BIOLOGV Botany PTERIDOPHYTA OF PART V 18. Aspleniaceae-21. Polypodiacea< Rolla !V I Rob* Bill : •ation for Contributors to Fieldiana i he Uni iCAl -FREE PAPER FIELDIANA Botany NEW SERIES, NO. 32 PTERIDOPHYTA OF PERU PARTY 18. Aspleniaceae-21. Polypodiaceae Rolla M. Tryon Department of Biology University of South Florida Tampa, Florida 33620-5150 With the collaboration of: Blanca Leon Universidad Nacional Mayor de San Marcos Lima, Peru Robert G. Stolze Associate Curator Department of Botany Field Museum of Natural History Roosevelt Road at Lake Shore Drive Chicago, Illinois 60605-2496 Accepted January 6, 1993 Published June 30, 1993 Publication 1447 BIOLOGY LIBRARY 101 BURRILL HALL SEP 2 7 I993 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 1993 Field Museum of Natural History Library of Congress Catalog Card Number: 93-71887 ISSN 00 15-0746 PRINTED IN THE UNITED STATES OF AMERICA Table of Contents List of Illustrations INTRODUCTION 1 ACKNOWLEDGMENTS 1 1 8. ASPLENIACEAE 2 Asplenium 2 1 9. DAVALLIACEAE 49 Nephrolepis 49 20. BLECHNACEAE 54 Blechnum 56 Salpichlaena 68 2 1 . POLYPODIACEAE 70 Grammitis 72 Pecluma 116 Polypodium 125 Pleopeltis 140 Dicranoglossum 145 Microgramma 148 Campyloneurum 158 Niphidium 173 Solanopteris 1 79 Platycerium 181 MAP OF PERU 1 84 INDEX TO NAMES . .185 1. Asplenium: A. pseudoangustum; A. hallii; A. theciferum 3 2. Nephrolepis: N. pectinata; N. cordifo- lia; N. multiflora; N. rivularis 50 3. Blechnum: B. asplenioides; B.fraxi- neum; B. binervatum ssp. fragile 55 4. Salpichlaena: S. volubilis 69 5. Grammitis: G. myosuroides; G. limba- ta\ G. david-smithii; G. myriophylla ... 73 6. Pecluma: P. hygrometrica; P. filicula; P. eurybasis var. pilosa; P. curvans 117 7. Polypodium: P. decumanum; P. au- reum; P. loriceum; P. triseriale 126 8. Pleopeltis: P. macrocarpa var. macro- carpa; P. macrocarpa var. laciniata; P. percussa 141 9. Dicranoglossum: D. subnudum; D. des- vauxii; D. polypodioides 147 10. Microgramma: M. thurnii; M. reptans\ M. squamulosa 1 50 11. Campyloneurum: C. fasciale; C. phylli- tidis; C. angustipaleatum\ C. ophiocau- lon; C. amphostenon 159 12. Niphidium: N. crass ifolium; N. mac- bridei 175 13. Solanopteris: S. bifrons; S. bismarckii . . 178 14. Platycerium: P. andinum 182 in Back cover: Nephrolepis pectinata PTERIDOPHYTA OF PERU PartV 18. Aspleniaceae-21. Polypodiaceae Introduction This fifth part of the "Pteridophyta of Peru" contains the fern families Aspleniaceae, Blech- naceae, Davalliaceae, and Polypodiaceae. Al- though it was stated in the introduction to Part I that this pteridophyte flora would be contained in five parts, it has been deemed more practical to add a sixth part (now in progress), which will in- clude the "water ferns," "fern allies," a brief bio- geography of Peru, and a comprehensive index to the entire work. The general style, typography, form of citations, and so forth follow the previously published parts. 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. Campyloneurum of the Polypodiaceae has been contributed by Blanca Leon, and the other genera are a joint effort of Rolla M. Tryon and Robert G. Stolze, each critically reviewing the treatments prepared by the other. 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. Abbreviations of periodicals generally follow the system of Botanico-Periodicum-Huntianum (1968), while those of books and authors generally follow the system of Taxonomic Literature (TL- 2, 1 976 et seq.). A notable exception is Karel Dom- in's "The pteridophyta of the island of Dominica" (1929), which is cited throughout as "Pterid. Dominica," in the interest of brevity. According to TL-2, the full citation is Rozpr. Krai. Ceske Spolecn. Nauk, Tr. Mat. Prir. N.R. 2. The abbre- viations for herbaria follow Index Herbariorum. Acknowledgments Blanca Leon is nearing completion of a mono- graphic study of Campyloneurum (Polypodi- aceae), a genus in which there are very few diag- nostic characters to separate the species. She has seen nearly all of the type collections during sev- eral years of research and so has gained a thorough knowledge of a taxonomically difficult group of ferns. The authors are extremely fortunate to have her contribution to this flora. 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 their packing and shipment from those two important Peruvian herbaria. The illustrations were contrib- uted by Field Museum scientific illustrator Zorica Dabich, who created the original drawings and adapted the rest from those previously used in the Fieldiana: Botany publication, "The Ferns and Fern Allies of Guatemala." We are grateful to her, for her art has provided a valuable complement to the descriptions. We are indebted to Dr. Alan R. Smith and Dr. Robbin C. Moran for their crit- ical suggestions on the treatment of Grammitis (Polypodiaceae), and to Dr. David B. Lellinger for stimulating discussions and correspondence per- taining to a wide variety of taxonomic problems. To Dr. Brigitte Zimmer, of the Botanical Museum, Berlin-Dahlem, we extend special thanks for valu- FIELDIANA: BOTANY, N.S., NO. 32, JUNE 30, 1993, PP. 1-190 able comments on typification and nomenclature, particularly in regard to Pecluma and Microgram- ma. We also appreciate the valuable comments presented by reviewers of the manuscript. 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: Botanischer Garten und Botanisches Mu- seum, Berlin-Dahlem, Berlin (B); British Museum (Natural History), London (BM); Jardin Botanique National de Belgique, Meise (BR); Field Museum of Natural History, Chicago (F); Harvard Univer- sity, Cambridge, Mass. — most Gray Herbarium (GH), some Arnold Arboretum (A); Herbarium Truxillense, Universidad Nacional de Trujillo, Trujillo, Peru (HUT); Royal Botanic Gardens, Kew, England (K); Rijksherbarium, Leiden, The Neth- erlands (L); Missouri Botanical Garden, St. Louis (MO); New York Botanical Garden, New York (NY); Museum National d'Histoire Naturelle, Paris (P); National Museum, Prague, Czechoslovakia (PR); Botanical Institute, Charles University, Prague, Czechoslovakia (PRC); Herbario del Institute de Ciencias Naturales, Universidad Central, Quito, Ecuador (Q); Rijksmuseum, Stockholm, Sweden (s); University of California, Berkeley (uc); United States National Herbarium, Smithsonian Institu- tion, Washington, D.C. (us); Herbario San Mar- cos, Universidad Nacional Mayor de San Marcos, Lima, Peru (USM); and Naturhistorisches Museum, Vienna (w). This project has been supported in part by grant #BSR-85- 16358 from the National Science Foun- dation, Systematic Biology Program. The work would not have been possible without this assis- tance. However, any opinions and conclusions ex- pressed are those of the authors and do not nec- essarily reflect the views of the Foundation. Family 18. ASPLENIACEAE Aspleniaceae Newman, Hist. Brit, ferns, 6: 1840. TYPE: Asplenium L. Terrestrial, epiphytic, or epipetric. Stem com- monly erect, sometimes slender and long-creeping, bearing scales (and rarely also trichomes), the scales obviously clathrate, or occasionally only incon- spicuously so. Leaves circinate in vernation, pin- nate to decompound, or in several species entire, essentially monomorphic, of small to moderate size, or rarely to 1 m long, petiole continuous with the stem. Lamina essentially glabrous, or several species with minute trichomes or reduced scales, especially along the rachis, the rachis sometimes proliferous at the pinna axils or apex. Veins free, or in a few species casually to freely anastomosing. Sori linear to elliptic, borne along a vein, covered by a short to very long and narrow indusium, or rarely with a nearly marginal pouch formed by the indusium and opposed leaf tissue, paraphyses lacking. Sporangia usually long-stalked, the cells of the stalk uniseriate below its apex, the annulus interrupted by the stalk. Spores monolete, bilat- eral, lacking chlorophyll, usually with sharp folds in the perispore. The Aspleniaceae, as recognized here, are a fam- ily of about 700 species, included in seven genera, of which five are found in the New World: As- plenium, Camptosorus, Holodictyum, Pleurosorus, and Schaffneria. Only Asplenium occurs in Peru. The family is typically characterized by small to medium-sized leaves with clathrate stem scales and nonarticulate petioles, veins commonly free or, when anastomosing, the areoles without free included veinlets, and elongate sori with a narrow indusium borne along a vein. References MORTON, C. V., AND D. B. LELLINGER. 1986. The Polypodiaceae subfamily Asplenioideae in Ven- ezuela. Mem. New York Bot. Gard., 15: 1-49. STOLZE, R. G. 1986. Polypodiaceae— Asplenioi- deae. In Harling, G., and B. Sparre, eds., Flora of Ecuador 23: 1-83. Stockholm. TRYON, R. M., AND A. F. TRYON. 1982. Asplen- iaceae, pp. 627-654. In Ferns and Allied Plants. Springer- Verlag, New York. I. Asplenium Asplenium L., Sp. pi. 1079. 1753; Gen. pi., ed. 5, 485. 1754. TYPE: Asplenium marinum L. Figure 1. FIG. 1 . Asplenium pseudoangustum: a, habit. Asplenium hallii: b, habit; c, stem scale; d, base of pinna. Asplenium theciferum: e, apex of pinna, (a from Schunke V. 6951, F; b, c, d from Acosta Solis 13761, Ecuador, F; e from Holm- Nielsen et al. 4110, Ecudaor, F.) FIELDIANA: BOTANY TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. Phyllitis Hill, Brit. herb. 525. 1757. TYPE: Phyllitis scolopendrium Newman = Asplenium scolopen- drium L. Loxoscaphe Moore, Hooker's J. Bot. Kew Gard. Misc. 5: 227. 1853. TYPE: Loxoscaphe concinna (Schrader) Moore (Davallia concinna Schrader) = Asplenium concinnum (Schrader) Kuhn = As- plenium theciferum (HBK.) Mett. Leaves pinnate to 4-pinnate, in a few species entire, 3-60(-100) cm long. Lamina terminating in a conform apical segment, or more commonly tapering to a pinnatifid apex, or occasionally with a flagellate, proliferous tip. Pinnae, when present, several to many, subequilateral to (more com- monly) inequilateral at base. Veins simple or forked, free, or reticulate in A. purdieanum. Sori elliptic to (more commonly) linear, borne along the acroscopic side of veins or, in several species, a few of them also borne along the basiscopic side (thus back-to-back, i.e., diplazioid). Indusia nar- row, attached along the vein and opening toward the midrib, commonly persistent, rarely obsolete, or in A. theciferum cuplike or pouchlike, borne singly near segment tips, with sporangia deeply immersed. Sporangia on slender stalks composed of a single row of cells, annular cells commonly 20-25. There are more than 600 species in this nearly cosmopolitan genus, a majority of these occurring in wet forests of tropical regions. Unfortunately, although the numerous species are rather easily identified to genus, they are not readily (or at least obviously) classified into infrageneric groups. As in Diplazium, many can be separated on the basis of pinna dissection, and yet very closely related species (and even a species itself) can be 1-3-pin- nate. One exception to this is the group of species often placed in the genus Loxoscaphe. Most, in- cluding Asplenium theciferum, are quite distinc- tive in their indusia, which are generally pouch- or cup-shaped and occur near the segment margin. By means of this character, the latter species can- not be confused with any other in Peru. But there are a few Old World species in the group in which the sori intergrade to a more typical Asplenium sorus. For this reason Loxoscaphe is included with Asplenium for purposes of this treatment. A few species with back-to-back (diplazioid) sori are frequently determined as Diplazium. With these species, comparison of stem scales, and of spo- rangia stalks and annular cells, is necessary for positive identification to genus. (Also see Smith, 1976.) References MUTUI, K., N. MURIKAMI, AND K. IWATSUKI. 1989. Chromosomes and systematics of As- plenium sect. Hymenasplenium (Aspleniaceae). Amer. J. Bot., 76: 1689-1697. SMITH, A. R. 1976. Diplazium delitescens and the Neotropical species of Asplenium sect. Hy- menasplenium. Amer. Fern J., 66: 1 16-120. Key to Species of Asplenium a. Lamina simple, margins entire or serrate; petiole obsolete or VJ0 to V20 the length of the lamina . b b. Larger adult leaves 4.2-15 cm broad, 20-70(-100) cm long; veins usually spreading at 50-75° from the midrib 1 . A. serratum b. Larger mature leaves 0.5-3(-4) cm broad, 8-30(-40) cm long; veins spreading at 30-45° from the midrib c c. Midrib with scattered, minute, clathrate scales, at least abaxially; stem scales blackish, linear, attenuate, obscurely clathrate; veins commonly evident (if not prominent) ... 2. A. angustum c. Midrib lacking scales; stem scales brown, lanceolate to ovate, acute or acuminate; veins obscure 3. A. pseudoangustum a. Lamina lobed or pinnate to decompound or (in a few species) simple and unlobed, and then petiole '/4-'/2 the length of the lamina d d. Indusium cup- or pocket-shaped, borne along the acroscopic side of ultimate segments near their tips 62. A. theciferum d. Indusium linear to narrow-elliptic, attached along the vein and opening toward the midrib . . e e. Lamina simple, the margins entire to serrate or crenulate f f. Lamina base extremely long-decurrent; midrib abaxially with scattered, minute, blackish scales; lamina chartaceous, lacking glandular trichomes 4. A. stuebelianum f. Lamina base short-decurrent; midrib lacking scales; lamina thin- to firm-herbaceous, with scattered, minute (0.1-0.2 mm), appressed, often gland-tipped trichomes .. 5. A. pearcei FIELDIANA: BOTANY e. Lamina lobed to 1 -pinnate or decompound g g. Lamina shallowly lobed to pinnatifid (or some juvenile leaves with a pair of lobed basal pinnae); leaves diminutive, 2-6 cm long, 0.2-0.8 mm broad 6. A. escaleroense g. Lamina regularly 1 -pinnate or decompound (in a few species juvenile leaves sometimes simple and entire); leaves small to large h h. Basal pair of pinnae much more strongly produced basiscopically than acroscopically; lamina with few to many whitish, septate trichomes, especially on rachis, veins, and margins 7. A. pumilum h. Basal pair of pinnae subequilateral at base or more strongly produced acroscopically; lamina lacking whitish, septate trichomes i i. Petiole and base of rachis highly lustrous and dark brown, atropurpureous or blackish j j. Pinnae entire to sharply and deeply incised, 0.6-1.5 cm long k k. Pinnae deeply incised 8. A. formosum k. Pinnae entire to crenate, serrate, or shallowly lobed 1 1. Adaxial raised ribs of rachis conspicuously hirsute, the rigid, spreading, cas- taneous trichomes to 0.3 mm long; pinnae subequilateral ... 9. A. extensum 1. Adaxial ribs or wings of rachis entire, denticulate, or pustulate; pinnae inequi- lateral at base, cuneate basiscopically, truncate and often subauriculate ac- roscopically m m. Vein tips not dilated, inconspicuous, or obscured in leaf tissue; adaxial ribs of rachis each with a pronounced, foliaceous green wing throughout 10. A. resiliens m. Vein tips (most of them) somewhat to strongly dilated and conspicuous on adaxial side of pinnae; adaxial ribs commonly lacking foliaceous green wings 11. A. monanthes j. Pinnae 1-3 -pinnate, or if subentire to incised, then larger ones (1.8-)2-12 cm long n n. Lamina gradually reduced at base; petiole 1-6 cm long o o. Lamina 2-3-pinnate 12. A. rutaceum o. Lamina 1 -pinnate-pinnatisect 13. A. hallii n. Lamina truncate or nearly so p p. Rachis not prolonged or proliferous, the lamina apex pinnatifid; stem short- creeping (often inconspicuously so in A. laetum); pinnae subentire to shal- lowly pinnatifid q q. Pinnae 15-24 pairs, 2.5-4.5 cm long, most of them strongly inequilateral for nearly half their length 31. A. laetum q. Pinnae 7-1 1 pairs, larger ones 6-1 2 cm long, essentially equilateral except at the base 32. A. melanopus p. Rachis prolonged, often proliferous; stem erect or decumbent; pinnae sub- entire to 3-pinnate r r. Stem scales 0.6-1 mm broad, ovate or broadly lanceolate, acute, dark or grayish brown and often with lighter brown margins . . 14. A. macrurum r. Stem scales 0.1-0.3 mm broad, linear or narrowly lanceolate, attenuate, gray-brown to blackish, concolorous 15. A. radicans i. Petiole and base of rachis not or slightly lustrous, green, reddish or grayish brown (rarely darker, but then not highly lustrous) s s. Pinnae mostly trifoliolate or bifoliolate, occasionally with l(-2) additional pairs of pinnules t t. Leaves 1.5-5 cm long, remote, borne singly or in pairs from long-creeping, stoloniform roots; pinnae 4-8 pairs; stem scales few, 0.5 mm long 1 6. A. repens t. Leaves (4-)6-30 cm long, fasciculate from an erect stem, pinnae numerous; stem scales many, 1-2 mm long 17. A. triphyllum TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. s. Pinnae simple or regularly pinnate to decompound, not trifoliolate or bifoliolate u u. Lamina 2-pinnate or more, at least at pinna bases; basal segments (pinnules) of proximal pinnae obviously stalked v v. Lamina conspicuously reduced at base, proximal pinnae often rudimentary w w. Fertile ultimate segments mostly less than 2 mm long, conspicuously broader than adjacent sterile ones; stem scales bicolorous, blackish with narrow but conspicuously brown margins 18. A. divaricatum w. Fertile and sterile ultimate segments 2-4 mm long, subequal; stem scales concolorous, or occasionally faintly brown-margined x x. Lamina linear, 2-pinnate; pinnules 3-6 pairs on a pinna; plants epipetric (rarely terrestrial), growing above 2900 m 19. A. haenkeanum x. Lamina elliptic or elliptic-ovate, 2-pinnate-pinnatisect or 3-pinnate; pinnules of larger pinnae 7-14 pairs; plants terrestrial (rarely epipetric or epiphytic), at elevations from 250 to 2700 m y y. Pinnae commonly patent, margins parallel for much of their length; ultimate segments mostly narrow-elliptic, acute .20. A. cristatum y. Pinnae commonly ascending, broadest near the base, from there gradually to abruptly tapering to apex; ultimate segments mostly obovate, obtuse to subacute 21. A. myriophyllum v. Lamina not or scarcely reduced at base z z. Leaves 1.5-8(-10) cm long, borne directly on stoloniform (commonly long-creeping) roots; scales of petiole base 0.4-0.6 mm long a2 a2. Ultimate segments flabelliform, subdigitately incised at apex, veins commonly 1 -forked in each segment; petiole to 0.5 cm long 16. A. repens a2. Ultimate segments linear or narrowly oblanceolate, apex acute, veins simple in each segment; petiole 1.5—4 cm long . . 22. A. delicatulum z. Leaves (10-) 12- 100 cm long, borne on a stout, short-creeping to erect, stem; scales of stem and petiole base 1.5-20 mm long b2 b2. Rachis obviously alate throughout; most pinnae oblong, margins par- allel for much of their length, spreading at right angles, sessile, their basal pinnules strongly overlapping the rachis . . . 20. A. cristatum b2. Rachis nonalate, or narrowly so distally; pinnae ovate or deltate, tapering from base to apex, ascending, short-stalked, basal pinnules not or infrequently overlapping the rachis c2 c2. Lamina 1 5-20 cm broad; pinnules (2-)3-9 cm long; stem scales 10-20 mm long, 3-6 mm broad; indusia thin, commonly scarious 23. A. squamosum c2. Lamina 2-1 2(-l 5) cm broad; pinnules 0.5-2 cm long; stem scales 2—4 mm long, 0.3-0.8 mm broad; indusia firm or thick and fleshy, whitish to gray, or light brown d2 d2. Stem scales linear, attenuate, dark brown; lustrous; ultimate segments obovate to subflabellate, broadly and obtusely den- tate near apex; veins subdichotomously branched 56. A. cuneatum d2. Stem scales lanceolate to ovate, dull orange to gray-brown; ultimate segments linear-lanceolate or elliptic, acute or sub- acute; veins 1 -forked to pinnately branched 24. A. cuspidatum u. Lamina 1 -pinnate to 1 -pinnate-pinnatisect (in a few species nearly 2-pinnate at bases of proximal pinnae); basal secondary segments (if any) adnate, not freely stalked . . e2 FIELDIANA: BOTANY e2. Lamina terminating in a distinct, conform or subconform apical segment (this occasionally with a basal lobe or auricle) f2 f2. Veins anastomosing, especially toward pinna margin; petiole copiously scaly 61. A. purdieanum f2. Veins free; petiole naked or sparsely scaly (or sometimes in A. serra conspicuously scaly) g2 g2. Leaves approximate to remote on a short- to long-creeping stem h2 h2. Stem abundantly scaly; lamina firm-herbaceous to (more com- monly) chartaceous, the petiole and rachis scaly (scales at times caducous); pinnae subequilateral at base 57. A. serra h2. Stem essentially lacking scales; lamina membranaceous or thin- herbaceous, the petiole and rachis naked; pinnae conspicuously inequilateral at base 25. A. repandulum g2. Leaves crowded to caespitose on an erect to decumbent (rarely very short-creeping) stem i2 i2. Pinnae conspicuously inequilateral at base, there acroscopically truncate to cordate or auriculate, often overlapping the rachis J2 j2. Larger pinnae 2.5-4 cm broad; pinnae bases (at least prox- imal ones) mostly truncate to broadly rounded basiscopi- cally 26. A. vomeriforme j2. Larger pinnae 1-2 cm broad; pinna bases mostly narrowly cuneate basiscopically 53. A. sal id folium i2. Pinnae subequilateral at base, there broadly or narrowly cuneate to attenuate k2 k2. Stem scales filiform, 0. 1-0.3(-0.4) mm broad 27. A. juglandifolium k2. Stem scales lanceolate to ovate, 0.5-2 mm broad ... 12 12. Lamina with 1 (very rarely 2) pair of pinnae, these subopposite and narrowly adnate; stem scales 3-4 mm long 5. A. pearcei 12. Lamina with (l-)2-8 pairs of pinnae, these alternate, sessile or short-stalked, not or rarely adnate; stem scales mostly 5-1 5 mm long m2 m2. Stem scales 1-2.5 mm long, 0.5 mm broad, black- ish, often slightly vaulted; apical lamina segment 1-1.7 mm broad or occasionally obso- lete 28. A. tuerckheimii m2. Stem scales 5-15 mm long, mostly 1-2 mm broad, orange to brown, flat; apical segment 2.2-5.0 cm broad n2 n2. Pinna margins entire; lamina glabrous abaxially; plants epiphytic 29. A. tricholepis n2. Pinna margins crenate-serrate; lamina mod- erately to amply provided abaxially with minute, appressed, dark brown trichomes; plants terrestrial 30. A. davisii e2. Lamina tapering to a pinnatifid, caudate, or flagelliform apex or, if with a distinct terminal segment, this obviously differing in shape from lateral segments o2 o2. Petiole, rachis and tissue sparsely to copiously scaly, the scales dark and filiform, with prolonged, unicostate apices p2 TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. p2. Pinnae serrate to deeply and irregularly erose-lacerate, not cune- ately lobed 58. A. dimidiatum p2. Pinnae deeply lobed, the 3-6 segments cuneiform, their apices often lacerate 59. A. praemorsum o2. Petiole, rachis and tissue with scales lacking or scattered and incon- spicuous q2 q2. Stem short-creeping (sometimes not obviously so in A. laetum); leaves closely spaced or crowded along the stem r2 r2. Pinnae 14-24 pairs, larger ones 2.5-5 cm long; sori not or rarely diplazioid s2 s2. Stem scales orange; veins and sori subparallel, or diverging at 5-10°; pinnae deeply pinnatisect 60. A. lividum s2. Stem scales dark brown or blackish; veins and sori di- verging at 30-40°; pinnae bicrenate or biserrate 31. A. laetum r2. Pinnae 6-10(-12) pairs, larger ones 6-14 cm long; sori often diplazioid t2 t2. Pinnae (at least proximal ones) lobed to pinnatifid on both margins, the tip of each lobe with a few rounded teeth . 32. A. melanopus t2. Pinnae serrate to deeply crenate acroscopically, serrate to subentire basiscopically 33. A. delitescens q2. Stem erect, not creeping; leaves fasciculate to densely caespi- tose u2 u2. Pinnae incised nearly to the costa into 7-16 pairs of entire to crenate segments 34. A. achilleifolium u2. Pinnae subentire to crenate, or with a nearly free basal acro- scopic auricle, but not regularly pinnatisect v2 v2. Pinnae lacking a definite midrib, the veins few and com- monly subflabellately dichotomous w2 w2. Veins conspicuous, blackish, simple or 2-3-forked; pinnae 2-3 mm long, with short, blackish, stalks . . 35. A. projectum w2. Veins obscure, or if distinct only slightly darker than the laminar tissue, commonly subflabellate, 2-3 times dichotomous; pinnae 4-14 mm long, not black- stalked x2 x2. Larger pinnae 1 0- 1 4 mm long; plants terrestrial, growing below 1000 m 36. A. lorentzii x2. Larger pinnae 4-8(-10) mm long; plants epi- petric (rarely terrestrial), growing above 2700 m y 2 y2. Petiole 0.2-0.4 mm thick, this and rachis flexuous and greenish or stramineous; pin- nae broadly and deeply lobed and cuspidate 37. A. gilliesii y2. Petiole 0.5-1 mm thick, this and rachis rigid and commonly reddish or grayish brown; pinna margins entire to deeply dentate, rarely lobed 38. A. peruvianum v2. Pinnae, at least in the proximal half, with a definite, central (or basiscopic in subdimidiate pinnae) axis, this either a thick, raised, costa, or straight midvein bearing several to many pinnately arranged veins z2 FIELDIANA: BOTANY z2. Stem scales lacking; pinna midrib with 2(-3) acro- scopic, and usually 1 basiscopic, veins 39. A. quitense 2.2. Stem scales ample and conspicuous; pinnae, larger ones, with 3-many pairs of veins (sometimes less in A. pulchellum) a3 a3. Pinnae subdimidiate, i.e., conspicuously inequi- lateral for half their length or more; veins 1-3 (-4) on basiscopic side of the pinna, and the basi- scopic margin entire b3 b3. Lamina apex acuminate to subcaudate; pin- na apex acute; acroscopic margin deeply and acutely cleft 40. A. pulchellum b3. Lamina apex acute; pinna apex obtuse; ac- roscopic margin crenate-serrate 41. A. discrepans a3. Pinnae subequilateral, or inequilateral only to- ward the base; veins 5-many on basiscopic side (sometimes less in A. tabinense but then the basi- scopic margin narrowly and deeply crenate) c3 c3. Veins all simple, or proximal 1-3 veins forked on acroscopic side of pinnae d3 d3. Lamina gradually reduced to rudimen- tary basal pinnae; petiole very short, '/5- Vio the length of the lamina e3 e3. Rachis broadly alate, each wing of- ten as wide as the rachis; acroscopic pinna margin serrulate to deeply serrate; elevation 400-1 100 m ... 42. A. pteropus e3. Rachis narrow-alate; acroscopic pinna margin deeply and narrowly crenate to shallowly lobed, and of- ten with a discrete basal auricle; el- evation 1 500-3600 m f 3 f3. Basal acroscopic pinna auricle cut nearly to costa; sori com- monly 8-12 on larger pinnae . 43. A. sessilifolium f3. Basal acroscopic pinna auricle cut (if at all) only halfway to cos- ta; sori commonly 4-7 on larger pinnae 44. A. tabinense d3. Lamina scarcely reduced at base, or sev- eral pinnae reduced and the basal ones half the size of largest ones; petiole '/»- '/2 the length of the lamina g3 g3. Lamina apex with a subcaudate, proliferous tip ... 45. A. herzogii g3. Lamina apex pinnatifid, not prolif- erous h3 h3. Pinna margins subentire to crenulate; pinnae 1.2-2.2(-2.5) cm long; stem scales 1.5-3 mm TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. long, blackish, attenuate 46. A. bangii h3. Pinna margins deeply serrate to serrately lobed; pinnae (larger ones) mostly 2.5-9 cm long; stem scales 4-10 mm long, gray brown or, if dark brown to blackish (in A. harpeodes) then with a long, tortuous, unicos- tate tip i3 i3. Stem scales deep brown to blackish, linear, with long, tortuous, unicostate tips; pinnae (24-)30-50 pairs; rachis atropurpureous . . . 47. A. harpeodes i3. Stem scales gray-brown, linear- to ovate-lanceolate, acute to attenuate; pinnae (12-) 15-24 pairs; rachis gray- to red-brown 48. A. raddianum c3. Veins all forked on acroscopic side of pinnae, except 1-3 apical ones simple j3 j3. Lamina abruptly ending in a proliferous tip; rachis (and usually petiole) broadly alate, the wing 1-2 mm broad 49. A. alatum j3. Lamina tip not proliferous; rachis and petiole narrowly or vestigially alate, or occasionally the rachis wing to 1 mm broad k3 k3. Indusia thin and mostly scarious, often hyaline until mature .... 13 13. Lamina chartaceous, abruptly terminating in a broad-based, nonconform, apical segment; petiole 12-25 cm long; pinnae 4-12 pairs . 50. A. absciss urn 13. Lamina firm-herbaceous to membranaceous, gradually re- duced to a pinnatifid, usually acuminate, apex; petiole 4-9 cm long; pinnae 1 5—40 pairs 51. A. drepanoph yllum k3. Indusia (at least mature ones) firm and fleshy, opaque m3 m3. Lamina very gradually taper- ing to a pinnatifid, often sub- caudate apex; stem scales ecil- iate 52. A. auritum m3. Lamina abruptly reduced at apex, the apical section ser- rate-ligulate, broad-based and nonconform, or indefinite; 10 FIELDIANA: BOTANY stem scales with orange cil- ia n3 n3. Veins mostly 2-forked; lamina ending in a broad- based, nonconform, some- times hastate, terminal segment .... 53. A. salicifolium n3. Veins simple to 1 -forked; lamina ending in a nar- row, serrate-ligulate, or subcaudatc apex .... o3 o3. Stem scales 6-9 mm long; pinna apex commonly attenuate ... 54. A. hastatum o3. Stem scales 2^*(-5) mm long; pinna apex acute or subacute . . 55. A. auriculatum 1. Asplenium serratum L., Sp. pi. 1079. 1753. LECTOTYPE (designated by Proctor, in Howard, Flora Lesser Antilles 2: 313. 1977): Plumier, Descr. pi. Amer. /. 39. 1693. Plants epiphytic, occasionally terrestrial. Stem stout, provided with brown-tomentose roots and usually iridescent, gray-brown (rarely blackish) scales, these obviously clathrate, linear or linear- lanceolate, attenuate, (5-)7-18 mm long. Leaves densely caespitose, sessile or subsessile, simple and unlobed, larger mature ones 20-70(-100) cm long 4.2-15 cm broad. Petiole 0-4 cm long, '/i0-'/2o the length of the lamina. Lamina chartaceous, acute to acuminate or abruptly caudate at the apex, grad- ually reduced to a long-attenuate base, with entire to crenulate or bluntly serrate margins, glabrous and eglandular, but midrib with minute, scattered, blackish, clathrate scales, at least abaxially. Veins simple to 1 -forked, spreading at 50-75° from the midrib. Sori densely crowded, greatly elongated. Indusia elongated, thin, often scarious. In deep, wet forests, on tree trunks and branches, occasionally in alluvial soil on ravine or stream banks, 100-800(-1200) m, Amazonas and Loreto to Cuzco and Madre de Dios. United States (southern Florida); West Indies; southern Mexico to Argentina and Paraguay. This and similar species such as the Old World A. nidus L., have been given the common name "Bird's Nest Ferns," due to their growth habit. The large, densely caespitose leaves of older plants often form a tight circle, the center of which then becomes a receptacle for a mixture of organic mat- ter that falls, or is blown in, from above. With this perhaps should be included A. angustum and A. stuebelianum, which are probably only variants. The two taxa differ from A. serratum merely by subtle characters of leaf shape, vein angle, and stem scales, and a number of specimens examined during this study are clearly intermediates. Stem scales of A. serratum are typically 8-15 mm long, gray-brown, and conspicuously clathrate, but oc- casionally they are smaller, darker, and more nar- rowly clathrate like those of A. stuebelianum. Some specimens of A. pearcei have simple leaves and are often determined as A. serratum. How- ever, the former have long petioles like those of A. stuebelianum as well as several other differing characters, which are discussed under that species. Amazonas: Prov. Bagua, NE of Chiriaco, Harbour 45 13 (F, MO, USM). San Martin: Chazuta, Rio Huallaga, Klug 4140 (F, OH). Loreto: Gamitanacocha, Rio Mazan, J. Schunke 276 (F, GH, uc). Huanuco: Prov. Huanuco, near confluence of Rio Cayumba with Huallaga, Mexia 8276 (F, OH, MO, uc, us). Pasco: Prov. Oxapampa, Palcazu valley, Iscozacin, Foster 7973 (F, MO). Junin: Chancha- mayo, La Merced, Soukup 101 1 (F). Ucayali (as Lorelo): Prov. Coronel Portillo, between Divisoria and Boque- ron, Ferreyra 4273 (GH, USM). Cuzco: Prov. La Conven- tion, Rio Apurimac, below Puerto Capiro, Davis et al. 1300 (F, GH). Madre de Dios: Prov. Manu, Manu Na- tional Park, Cocha Cashu Station, Foster et al. 6629 (F), 69/6 (F). TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 11 2. Asplenium angustum Sw., Kongl. Vetensk. Acad. Handl. 38: 66, t. 4,f. 1. 1817. TYPE: "Habitat in Brasilia," Freyreis (holotype, s; photos, F, us). Plants epiphytic. Stem erect, provided with brown-tomentose roots and blackish scales, these linear, 4-5(-6) mm long including the attenuate, unicostate tip, obscurely clathrate. Leaves caes- pitose, sessile or subsessile, simple, 8-30(-40) cm long, 0.5-3(-4) cm broad. Petiole 0-2 cm long, '/io-'/2o the length of the lamina. Lamina firm-her- baceous to chartaceous, long-attenuate to apex and base, lacking proliferous buds, margins entire, gla- brous and eglandular, but midrib provided with scattered, minute, dark, clathrate scales, at least abaxially. Veins simple to 1 -forked, spreading at 30-45° from the midrib. Sori approximate to sub- distant, linear to elliptic. Indusia linear to elliptic, whitish to pale greenish, relatively thick, not trans- lucent. In wet forests, on tree trunks, 100-1 100 m, Lo- reto, Pasco, Madre de Dios. The Guianas to Colombia; Peru; Brazil. This is but provisionally maintained as distinct from A. serratum. Whereas the extreme forms of each appear to be discrete, a number of specimens toward the edge of the range tend to be interme- diate. Typical A. angustum, especially in Brazil, has leaves less than 2.5 cm broad and very grad- ually tapering to apex and base, the veins borne at an angle of less than 45° from the midrib, the indusia relatively broad and thick-textured, and whitish and obscure, not crowded, and the stem scales blackish and obscurely clathrate. This con- trasts greatly with typical A. serratum, with lamina 8-12 cm broad and abruptly acute or acuminate at apex, the veins at angles of 50-60°, the indusia very long and narrow, scarious and densely crowd- ed, and the stem scales brownish, iridescent, and obviously clathrate. However, a number of small- er specimens of A. serratum exhibit nearly the size and vein angle of A. angustum and have blackish and less conspicuously clathrate stem scales. Sev- eral of the specimens from Peru cited below are approximately intermediate but have features much closer to A. angustum than to A. serratum and therefore are so identified here. Monographic study may prove the former to be merely a variant of the latter. Loreto: Prov. Maynas, Dist. Iquitos, Rio Momon, be- low Balcon, Hickock 606 (GH). Lower Rio Momoncillo, lower Rio Momon, near Iquitos, Jones & Davidson 9515 (MO, us). Prov. Maynas, Mishana, van der Werff el al. 10181 (MO, uc). Pasco (as Junin): Pichis Trail, San Ni- colas, Killip & Smith 30698 (us). Madre de Dios: Tam- bopata Nature Reserve, SE bank of Rio Tambopata, Barbour 5185 (MO). Parque Nacional del Manu, Cocha Cashu Biological Station, M. Foster P-84-52 (uc). 3. Asplenium pseudoangustum Stolze, Amer. Fern J. 74: 49. 1984. TYPE: Peru, Huanuco, Tingo Maria, Tryon & Tryon 5257 (holotype, GH!; isotypes, F!, us!, USM!). Figure la. Plants epiphytic. Stem small, erect to decum- bent, provided with dark or light brown, clathrate scales, these lanceolate or ovate, acute to acumi- nate, 1.5-3 mm long. Leaves caespitose, sessile or subsessile, simple, 8-22 cm long, 1.3-2.5(-3) cm broad. Petiole essentially lacking. Lamina firm- herbaceous, subcarnose, elliptic, apex acute (sometimes somewhat attenuate), long-attenuate at base, lacking proliferous buds, margins entire, surfaces and midrib glabrous and lacking scales. Veins 1 (-2)-forked, obscure, spreading at 30^45° from the midrib. Sori well spaced, linear. Indusia linear, pale green or yellow-green, not translucent, becoming dark brown and thicker at maturity. On tree trunks in wet forests, 350-1400 m, San Martin to Cuzco and Madre de Dios. Ecuador; Peru. Although resembling A. angustum in size and shape, this is quite a distinct species, as evidenced by the key characters. In addition, A. pseudoan- gustum has somewhat succulent leaves that often commonly dry blackish green. Leaves of A. an- gustum dry gray- or yellow-green. San Martin: Prov. Mariscal Caceres, between Aucaya- cu and Uchiza, Ferreyra 1 7038 (GH, USM). Prov. Mariscal Caceres, Tocache Nuevo, /. Schunke V. 567 '1 (F, us). Huanuco (as San Martin): Tingo Maria, Allard 21596a (us), 22384 (us). Pasco: Paujil, near Puerto Bermudez, Leon et al. 292 (GH, USM). Prov. Oxapampa, Valle de Palcazu, near Iscozacin, Leon 715 (F, GH). Cuzco: Prov. La Convention, Tupitani, Bites 5450 (us). Madre de Dios: Prov. Manu, Cerro de Pantiacolla, Rio Palotoa, Foster et al. 10827 (F). 4. Asplenium stuebelianum Hieron., Hedwigia 47: 222. 1908. TYPE: Colombia, Prov. Cundi- namarca, near Villavicencio and Oca, Stiibel 659 (holotype, presumably B). Plants epiphytic, occasionally terrestrial. Stem stout, erect or decumbent, provided with brown- tomentose roots and gray-brown to blackish scales, 12 FIELDIANA: BOTANY these obscurely to narrowly clathrate, linear or lin- ear-lanceolate, attenuate, 2-4(-6) mm long. Leaves densely caespitose, usually conspicuously petio- late, simple and unlobed, larger mature ones 25- 50 cm long, 3.5-9 cm broad, proliferous buds oc- casionally borne at the apex. Petiole (3-)5-18 cm long, usually '/4-'/2 the length of the lamina. Lamina chartaceous, acute to acuminate or abruptly cau- date at the apex, at the base abruptly reduced and then long-attenuate, margins entire to crenulate or bluntly serrate, glabrous and eglandular, but mid- rib with minute, scattered, blackish, clathrate scales, at least abaxially. Veins simple to 1 -forked, spreading at 60-75° from the midrib. Sori densely crowded, greatly elongated. Indusia elongated, thin, often scarious. In deep, wet forests, on tree trunks and branches, occasionally in alluvial soil or ravine or stream banks, 100-1400 m, Amazonas and Loreto south to Junin, Madre de Dios. Venezuela; Colombia; Peru; Bolivia; Brazil. Asplenium stuebelianum is perhaps merely a va- riety of A. serratum, from which it differs princi- pally in the petiolate or subpetiolate lamina. In the latter the lamina tapers very gradually and regularly to the stem, with petioles (if any) only a few centimeters long. The lamina in A. stuebe- lianum is reduced suddenly and markedly and then is attenuate to a narrowly winged or naked petiole, which is typically 6-12 cm long. For further com- parison see treatment of A. serratum. The Wur- dack collection, cited below from Amazonas, is intermediate between the two taxa, in characters of both lamina and stem scales. Amazonas: Prov. Bagua, Rio Maranon above Casca- das de Mayasi, Wurdack 1971 (F, GH, uc, us). San Mar- tin: Prov. Mariscal Caceres, Dist. Tocache Nuevo, Puer- to Pizana, Schunke V. 6947 (F). Loreto: Mishiyacu, near Iquitos, Klug 1537 (F, us). Huanuco: Prov. Pachitea, Dist. Puerto Inca, Bosque Nacional de Iparia, Schunke V. 2968 (F, GH, us). Junin: Puente Perene, Coronado 254 (GH). Schunke Hacienda above San Ramon, Killip & Smith 24705 (F). Madre de Dios: Prov. Manu, Atalaya, Foster & Wachter 7461 (F). Prov. Manu, Cerro de Pan- tiacolla, Rio Palotoa, Foster et al. 10694 (F). 5. Asplenium pearcei Baker, Syn. fil. ed. 2: 483. 1874. TYPE: Peru (Pasco), banks of Rio Huancabamba, Pozuzo, Pearce 528 (holo- type, K.!; photo, F). Asplenium amazonicum Christ, Hedwigia 45: 191. 1906. LECTOTYPE (designated here): Brazil, Rio Jurua, Vie 5525 (P!; isolectotype, K!; photo, F of p). Vie 5525 (L!) is A. stuebelianum (mixed col- lection). Asplenium haplophyllum Domin, Pterid. Dominica 170, t. 29, f. 1 (Vie 5525, K.!). 1929. Plants low-epiphytic, rarely terrestrial. Stem stout, erect or decumbent, provided with clathrate, gray-brown scales, these lanceolate, acuminate, 2- 4 mm long and mostly 1 mm broad. Leaves fas- ciculate, 24-40 cm long, mostly long-petiolate, simple to trifoliolate or (very rarely) with 2 pairs of pinnae. Petiole commonly 5-18 cm long, dull reddish or greenish brown, sparsely clathrate-scaly at base. Lamina thin- to firm-herbaceous, in pin- nate leaves the apical segment conform but larger than lateral ones, sparsely to moderately provided with minute (0.1-0.2 mm), appressed, pluricel- lular trichomes, these often gland-tipped and vis- ible only under high magnification, rachis (in pin- nate specimens) and proximal part of petiole alate. Pinnae (when present) opposite or subopposite, narrowly and subequally cuneate at base, acute or acuminate at apex, narrowly adnate to the rachis, margins entire to crenulate. Veins 1-2-forked, in- distinct. Indusia linear, green or yellow-green, en- tire. In wet forests, low on tree trunks, on fallen logs, or rarely in wet, rocky soil, 1 00-500 m, San Martin and Loreto south to Puno. Venezuela; the Guianas; Ecuador; Peru; Brazil. Two strange occurrences have led to confusion attending the names of synonyms of A. pearcei. First, Ule 5525, one of the syntypes of A. ama- zonicum, represents a mixed collection, for the specimen at Leiden is A. stuebelianum, a distinct, albeit superficially similar, species. Second, Do- min, obviously unaware of the identity of A. ama- zonicum, later created A. haplophyllum, based on Ule 5525 at Kew (here designated isolectotype of the former name). Specimens of^4. pearcei with simple leaves have been confused with A. serratum, A. angustum, and A. pseudoangustum. It is distinguished from these three by the long petiole and thinner lamina and from the first two by lack of small, black scales along the costa abaxially. San Martin: Prov. San Martin, road from Tarapoto to Yurimaguas, Plowman 6001 (GH, USM). Prov. Maris- cal Caceres, Tocache Nuevo, Schunke V. 7069 (F, MO, USM). Loreto: Gamitanacocha, Rio Ma/an. J. Schunke 305 (F, GH, uc, us). Pasco (as Junin): Cahuapanas, on Rio Pichis, Killip & Smith 26736 (us). Ucayali (as Lo- reto): Rio Aguaytia above mouth of Yurac-Yacu, Croat 20905 (MO). Madre de Dios: Prov. Tambopata, Albergue TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 13 Cuzco Amazonico, Leon 890 (F, USM). Puno: Prov. Cara- baya, Hacienda Palmera, Inambari, Vargas 16142 (GH). 6. Asplenium escaleroense Christ, Hedwigia 44: 366. 1905. TYPE: Peru (Loreto), "Cairo de Escalero," Ule 6886 (holotype, P; isotype, B?). Plants epiphytic, occasionally epipetric. Stems minute, small and erect, often arising at subdistant intervals from long-creeping, stoloniform roots, scales lacking, or a few dark brown, clathrate ones at the petiole base. Leaves single, or several and fasciculate, 2-6(-9) cm long, 0.2-0.6(-1.0) cm broad. Petiole thin, 0.4-2 cm long, green or green- ish brown. Lamina thin-herbaceous, linear or lin- ear-lanceolate (or juvenile ones subflabellate to obovate), glabrous, shallowly lobed to pinnatifid, or rarely with a pair of lobed basal pinnae. Seg- ments or lobes 2-6 pairs, strongly ascending. Veins simple in each segment, often indistinct. Sori one per segment. Indusia to 3 mm long and 0.5 mm broad, yellowish or greenish. In rain forests, on trunks or branches of trees, or occasionally on wet rocks or cliffs near water- falls, 130-1200 m; in Peru thus far known from the type and two other collections. Colombia; Ecuador; Peru; Bolivia. Scattered collections have been made of this species from Colombia to Bolivia, but the leaves are tiny and inconspicuous and therefore easily overlooked, so it is likely to be far more common within its reported range. It should be searched for throughout Peru in wet forests, on trees, and on rocks around waterfalls. San Martin: Tarapoto-Yurimaguas Hwy., km 14-17, McDaniel 13802 (GH). Loreto: Prov. Maynas, Yana- mono Explorama Tourist Camp, van der Werff et al. 9923 (MO, uc). 7. Asplenium pumilum Sw., Prodr. 129. 1788. TYPE: Swartz, Jamaica (holotype, s). Plants terrestrial or epipetric. Stem small, erect, provided at apex with filiform, blackish scales to 3 mm long. Leaves 1 -pinnate or -pinnatisect, sub- caespitose, mature ones 4-20 cm long, 1.5-6 cm broad. Petiole thin, 1-1 Ocm long, green to lustrous and castaneous or atropurpureous at base abaxi- ally. Lamina firm-herbaceous to somewhat fleshy, not reduced at base, terminating in a nonconform, pinnatifid apex, provided, especially on rachis, veins, and margins, with few to many whitish, septate trichomes. Pinnae 1-4 pairs, sessile and adnate distally, the proximal ones often short- stalked, subentire to crenate or lobed, basal pair the largest, more strongly produced basiscopically than acroscopically. Veins 1 - or 2-forked, or pin- nately branched in basal lobes. Sori linear or el- liptic. Indusia thin, yellow to whitish. Rare, in wet forests, on rocks, or on the forest floor, 400-1800 m, Tumbes, Huanuco, Cuzco. United States (Florida); West Indies; Mexico to Panama; Venezuela and Colombia to Brazil and Argentina; Africa. This is one of a few species in the genus having basal pinnae more strongly produced on the basi- scopic side. This and the whitish, septate tri- chomes of the lamina easily separate it from other species in Peru. Although it is rarely collected, it is probably more common than thus far reported, as it is but another of the very inconspicuous spe- cies in Asplenium. Tumbes: Between Caucho and Cotrina, Coronado 221 (GH, uc). Between Tumbes and Caucho, Coronado 225 (GH, uc). Huanuco: Yanano, Macbride 3815 (F). Cuzco: Prov. La Convencion, Potrero, Bties A40 (us). Prov. La Convencion, Playa de Balsa, Mexia 8053b (uc). 8. Asplenium formosum Willd., in L. Sp. pi. ed. 4, 5: 329. 1810. TYPE: Venezuela, Caracas, Bredemeyer(ho\oiype,B,Herb. Willd. 1990S; photos, F, GH). Plants terrestrial, occasionally epipetric or epi- phytic. Stem small, erect, provided with rigid, lin- ear scales, these 1-3 mm long, obscurely clathrate, black with narrow, brown margins. Leaves 1 -pin- nate, caespitose, to 30 cm long and 2.5 cm broad. Petiole rather stout, 1.5-4 cm long, lustrous, black or atropurpureous, glabrous. Lamina firm-herba- ceous to chartaceous, tapering to a pinnatifid apex, gradually narrowed at base to mere auricles, gla- brous, lacking proliferous buds, rachis adaxially with narrow perpendicular wings. Pinnae numer- ous, 0.6-1.5 cm long, patent, sessile, narrow and subacute, subdimidiate, basiscopically cuneate at base and the margin entire to biserrate, acroscop- ically truncate and the margin biserrate to deeply and sharply incised. Veins simple or 1 -forked, in- distinct. Sori 1-3 to a pinna, elliptic to ovate, often confluent at maturity. Indusia pale green to whit- ish. In wet forests, usually in loose soil of banks, occasionally on tree trunks or in crevices of wet 14 FIELDIANA: BOTANY cliffs, 200-1 200 m, Loreto, Junin, Ayacucho, Cuz- co. West Indies; Mexico to Panama; the Guianas to Colombia, south to Brazil and Argentina. Loreto: Banks of Rio Santiago above Pongo de Man- seriche, Mexia 6318 (F, GH, MO, uc, us). Junin: Rio Pinedo N of La Merced, Killip & Smith 23566 (F). Puente Quimiri, Leon 469 (USM). La Merced, Macbride 5373 (F). Ayacucho: Estrella, between Huanta and Rio Apu- rimac, Killip & Smith 23077 (us). Cuzco: Prov. La Con- vencion. Potrero, W of Quillabamba, Tryon & Tryon 5382 (GH). Prov. La Convention, Rosalina, Vargas 12288 (GH). 9. Asplenium extensum Fee, Mem. foug. 7: 51, /. 13. 1857. TYPE: Colombia, Ocana, Schlim 629 (holotype, P; isotypes, L!, RB; frag., us! of L; photos, F, GH & us of L). Plants terrestrial or epipetric. Stem short, erect or decumbent, provided with linear-lanceolate scales, these 1-2 mm long, obscurely clathrate, black, with narrow brown margins. Leaves 1 -pin- nate, subcaespitose, to 40 cm long and 2 cm broad. Petiole 3-5 cm long, terete, lustrous, atropurpu- reous to blackish, essentially glabrous. Lamina chartaceous to subcoriaceous, linear, strongly and gradually tapering to apex, not or rarely reduced at base, occasionally with proliferous buds at apex or in pinna axils. Rachis adaxially provided with strong parallel, winglike ribs, these conspicuously hirsute with rigid, spreading, castaneous trichomes to 0.3 mm long. Pinnae numerous, to 1 cm long, patent, approximate to remote, sessile, oblong, subequilateral, margins entire or broadly and shal- lowly crenate. Veins obscure, but their tips com- monly enlarged and conspicuous near the pinna margin. Sori 2-7 to a pinna, sometimes confluent at maturity. Indusia large, persistent, pale green to whitish. In forests, in soil, or in crevices of wet, rocky, canyon walls, 1 800-3950 m, Cajamarca and Ama- zonas to Junin. Colombia; Peru. Cajamarca: Prov. Celendin, canyon of Rio Maranon above Balsas, Hutchison & Wright 5294 (F, GH, uc, us). Amazonas: Prov. Chachapoyas, Cerros Calla Calla above Leimebamba, Hutchison & Bennett 4546 (F, GH, uc, us). San Martin: Prov. Mariscal Caceres, Rio Abiseo Na- tional Park, Young 4224 (HUT). Huanuco: Yanano, Mac- bride 3831 (F, GH, us). Tambo de Vaca, Macbride 4380 (F, us). Junin: Prov. Chanchamayo, Mina Pichita, van der Werffet al. 8660 (MO, uc). 10. Asplenium resiliens Kunze, Linnaea 18: 331. "1844" (1845), nom. nov. for Asplenium par- vulum Mart. & Gal. and with the same type. Asplenium parvulum Mart. & Gal., Nouv. Mem. Acad. Roy. Sci. Bruxelles 15: 60. 1842, nom. illeg. (not Hooker, 1840). TYPE: Mexico, Oaxaca, Galeotti 6462 (holotype, BR, photos, BM, F, us; isotype, BR, photos, P, us). Plants epipetric, rarely terrestrial. Stem short, erect, provided at apex with linear to filamentous, blackish scales, these 3-4 mm long, often with narrow brown margins, obscurely clathrate. Leaves 1 -pinnate, densely caespitose, to 35 cm long and 2.5 cm broad. Petiole 1-5 cm long, lustrous, atro- purpureous to blackish, essentially glabrous. Lam- ina subcoriaceous, linear, gradually reduced to base and diminished abruptly to a pinnatifid apex, lack- ing proliferous buds. Rachis adaxially with parallel ribs, each of these with a narrow but pronounced, foliaceous, green wing nearly throughout. Pinnae numerous, large ones 0.5-1.2 cm long, patent to slightly deflexed, subsessile or short-stalked, ellip- tic or broadly oblong, inequilateral at base, cu- neate basiscopically, truncate and often subauricu- late acroscopically, margins entire to broadly crenulate. Veins obscure or indistinct, the tips not dilated. Sori several pairs to a pinna, mostly crowded at the pinna margins, usually confluent at maturity. Indusia broad, pale yellow to whitish, persistent, but often obscured at maturity by the numerous sporangia. In forests, on ledges and in crevices of rocky cliffs, or on boulders, rarely in rocky soil, 1200- 3500 m, Cajamarca and Amazonas south to Cuz- co. Southern United States; Jamaica; Hispaniola; Mexico; Guatemala; Colombia and Venezuela south to Argentina and Brazil. Cajamarca: About 40 km from Cajamarca on road to Chilete, Correll & Smith P844 (GH, us), P844A (GH). Amazonas: Prov. Chachapoyas, Jazan (Ingenio-Chacha- poyas), Lopez et al. 4262 (GH, HUT). La Libertad: Prov. Sanchez Carrion, between Sausagocha and Cajabamba, Smith & Vasquez 3382 (MO, uc). Ancash: Prov. Carhuas, Cordillera Blanca, Valley of Rio Marcara, Hutchison & Wright 4362 (F, GH, uc, us). Huanuco: Huacachi, esta- cion near Muna, Macbride 3871 (F, GH, us). Junin: South of Huancayo, between Viques and Ingahuasi, Killip & Smith 22157 (F, us). Apurimac: 5 km above Huamcara- ma, West 3802 (uc). Cuzco: Road from "Ccasapata" (Casapata) to Apurimac, Sues 1388 (us). TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 15 11. Asplenium monanthes L., Mant. pi. 130. 1767. TYPE: South Africa, Cape of Good Hope (ho- lotype, LINN 1250.17; photo, us). Plants epipetric, occasionally terrestrial, rarely epiphytic. Stem short, erect, provided at apex with filiform to narrow-deltate scales, these 1-5 mm long, gray-brown to blackish, scarcely to obviously clathrate. Leaves 1 -pinnate, caespitose, to 60 cm long and 3 cm broad. Petiole 1-15 cm long, lus- trous, dark brown, atropurpureous or blackish, es- sentially glabrous. Lamina firm-membranaceous to chartaceous, linear, tapering to a pinnatifid apex and strongly reduced at base. Rachis sparsely pro- vided with dark, fibrillose scales, adaxially with 2 parallel raised ribs, not proliferous at apex but, in var. wagneri, often with scale-covered gemmae in a distal pinna axil. Pinnae (larger ones) to 1.5 cm long, sessile to minutely stalked, patent, blunt and quadrangular to subacute and falcate, inequilateral at base, often subauriculate acroscopically. Veins often indistinct, but their tips conspicuous and somewhat to strongly dilated adaxially. Sori sol- itary near the basiscopic pinna margin, or borne in several pairs, often confluent at maturity. In- dusia broad, pale green to whitish, persistent, but often obscured at maturity by spreading sporangia. The Southern United States; Jamaica; Hispan- iola; Mexico south to Panama; Colombia and Ven- ezuela south to Argentina and Chile; Hawaii; Af- rica. Authors have recognized Asplenium monanthes and two closely related species separated by a few, mostly quantitative, features. Some of these char- acters are relatively constant in certain parts of the range, but variable in other areas, and interme- diate specimens are not uncommon. (See discus- sion of var. monanthes for detailed comparison of characters.) In this treatment the three taxa are recognized as geographical or elevational variants, a key to which is provided here: a. Ribs on adaxial side of rachis conspicuously pustulate or pustulate-dentate; stem scales l-2(-3) mm long; a proliferous bud sometimes borne in a distal pinna axil lie. var. wagneri a. Ribs on adaxial side of rachis entire to remotely and minutely denticulate; stem scales mostly 4-5 mm long; proliferous buds lacking b b. Stem scales commonly blackish, or with a narrow brown margin, not or obscurely clathrate; pinnae commonly 2-4 times as long as broad 1 la. var. monanthes b. Stem scales commonly gray-brown, clathrate; pinnae l-1.5(-2) times as long as broad 1 1 b. var. castaneum 1 la. Asplenium monanthes var. monanthes. Asplenium monanthemum Murray, in L., Syst. veg. ed. 14: 933. 1784. nom. superfl. for Asplenium monanthes and with the same type. Stem scales commonly 4-5 mm long, blackish, or with narrow brown margins, linear to filiform, not or obscurely clathrate, lumina narrow and their walls thick. Leaves to 60 cm long and 3 cm broad. Petiole atropurpureous (sometimes dark brown or blackish). Rachis with adaxial ribs entire to mi- nutely denticulate, lacking proliferous buds. Pin- nae (larger ones) mostly 2—4 times as long as broad, subentire to crenate or crenate-serrate. In forests and thickets, on and among rocks, in crevices of rock cliffs, and occasionally in rocky soil, (900-) 1300-4350 m, Lambayeque to Ama- zonas, south to Ayacucho and Madre de Dios. Southern United States; Jamaica; Hispaniola; Mexico to Panama; Colombia and Venezuela south to Argentina and Chile; Hawaii; Africa. The range of var. monanthes is very broad; it occurs at middle to very high elevations in Peru. Variety castaneum does not occur in the Old World, the West Indies, or middle Central America, is (with a few exceptions) confined to Andean regions in South America, and is found mostly over 3000 m in Peru. Variety wagneri thus far has been found only in Colombia and Ecuador, and in Peru com- monly between 3000 and 3800 m. The last has the smallest leaves of the three varieties and is easiest to recognize because of the pustulate rachis ribs, the much smaller stem scales, and the occasional rachis buds. Darker clathrate scales, darker peti- oles, and relatively narrower pinnae usually dis- tinguish var. monanthes from var. castaneum, but all these vary frequently enough so that compar- ison of a suite of characters is usually necessary for positive identification. Some characters seem to be more constant in certain geographic areas than in others; consequently, the above key is es- pecially designed to separate the varieties in Peru. Lambayeque: Prov. Ferrenafe, Quiros 2521 (F). Ca- 16 FIELDIANA: BOTANY jamarca: Prov. Celendin, Pumarrume, Mostacero el al. 1011 (F, GH, HUT, uc). Amazonas: Prov. Chachapoyas, Cerros Calla Calla, above Balsas, Hutchison & Wright 5824 (F, GH, uc, us, USM). La Libertad: Prov. Santiago de Chuco, 26 km from Santiago, D. Smith 2334 (F, MO). San Martin: Dist. Huallaga. Valley of Rio Aposoncho, Hamilton & Holligan 500 (us), 914 (us). Ancash: Prov. Corongo, Nueva Victoria, Mostacero el al. 2009 (F, HUT). Huanuco: Muna, Macbride 3939 (f, GH, us). Lima: Prov. Yauyos, Laguna Huacracocha, Cerrate 1255 (USM). Pas- co: Huariaca, Bryan 169 (F). Junin: Prov. Tarma, 5 km SW of Huacapistana, Tryon & Tryon 5425 (F, GH, us). Ayacucho: Prov. Huamanga, Totorabamba, Weberbauer 5469 (GH). Apurimac: Prov. Abanbcay, forests of Am- pay, Vargas 1068 (F, GH). Cuzco: Prov. Urubamba, near town of Machu Picchu, Tryon & Tryon 5405 (F, GH, us). Mad re de Dios: Pinasniocj, Panticalla [Pantiacolla?] Pass, Cook & Gilbert 1798 (us). lib. Asplenium monanthes var. cast am* urn (Schlecht. & Cham.) Stolze, Flora of Ecuador 23:45. 1986. Asplenium castaneum Schlecht. & Cham., Linnaea 5: 611. 1830. TYPE: Mexico, Veracruz, Mt. Ori- zaba, Schiede (holotype, HAL; frag., NY!). Stem scales commonly 4-5 mm long and gray- brown, linear to narrow-deltate, usually conspic- uously clathrate, the lumina broad and clear and the walls thin. Leaves to 35 cm long and 2 cm broad. Petiole dark brown, sometimes castaneous, rarely atropurpureous. Rachis with adaxial ribs subentire, lacking proliferous buds. Pinnae (larger ones) l-1.5(-2) times as long as broad, subentire to crenate. In forests, on rocks and in crevices of rock cliffs, and sometimes in rocky soil, (600-)2900-4700 m, Cajamarca and Amazonas south to Huancavelica and Puno. Southern Mexico; Guatemala; Costa Rica; Pan- ama; Venezuela and Colombia south to Bolivia. Authors often distinguish this from var. monan- thes by the petiole color. Petioles of var. casta- neum usually are dark brown and those of var. monanthes atropurpureous, but, at least in Peru, petioles on the same stem can be of both colors. Furthermore, Stolze (1981) pointed out that in Guatemala the short pinna stalks were usually green in "A. monanthes," but often partly brown in "A. castaneum." This character is useless in Peru, as pinna stalks are nearly always concolorous in both varieties. Cajamarca: Prov. Contumaza, Jalca El Chuno, Sa- gdstegui el al. 9368 (F, MO). Amazonas: Prov. Chacha- poyas, Cerro Campanario, Wurdack 1558 (GH, uc, us). La Libertad: Prov. Santiago de Chuco, near Santiago, Sagdstegui el al. 11708 (HUT). San Martin: Prov. Maris- cal Caceres, Rio Abiseo National Park, Young & Leon 4576 (USM). Ancash: Prov. Yungay, near Laguna Llan- ganuco, Mostacero el al. 1401 (F, HUT, MO). Lima: Prov. Huarochiri, Dist. San Mateo, Rio Blanco, Sounders 404 (F, uc). Pasco: Between La Quinua and Tambo, Soukup 5627 (us). Junin: Mt. La Juntay, near Huancayo, Killip & Smith 22058 (GH, us). Huancavelica: Prov. Castro- virreina, Choclococha, Tovar 2842 (USM). Cuzco: Prov. Espinar, Imantata, Vargas 10509 (GH). Puno: Granja Salcedo, Canon Viscachani, Mexia 4263 (uc). 1 Ic. Asplenium monanthes var. wagneri (Kuhn) Stolze, Flora of Ecuador 23: 46. 1986. Asplenium wagneri Kuhn, Linnaea 36: 96. 1 869. TYPE: Ecuador, Pichincha, near "Turtillas" between Quito and Esmeraldas, Wagner (holotype, M; iso- type, BM!; photos, BM & uc of M, GH of BM). Asplenium vargasii Abbiatti, Darwiniana 14: 61, /. 1. 1966. TYPE: Cuzco, "Sajasaihuaman" (Saxihua- man), Vargas 362 (holotype, LIL). Stem scales l-2(-3) mm long, blackish, linear or narrow-deltate, not or obscurely clathrate. Leaves to 22 cm long and 1.3 cm broad. Petiole castaneous or atropurpureous. Rachis with adaxial ribs conspicuously pustulate or pustulate-dentate, sometimes with a proliferous bud in one of the distal pinna axils. Pinnae 1-1.5 times as long as broad, subentire to crenate or crenate-serrate. In forests, on rocks or in crevices or rock cliffs, or in rocky soil, (2700-)3000-3800(-4350) m, An- cash, Junin, Cuzco. Venezuela; Colombia; Ecuador; Peru. In Ecuador, pinnae are sometimes deeply cre- nate to shallowly lobed, but in specimens from Peru they are commonly merely crenate-serrate. The type of A. vargasii has not been examined, but the diagnostic features of var. wagneri (deeply dentate rachis ribs and short stem scales) are clear- ly indicated in the former's description and illus- tration. Ancash: Prov. Bolognesi, Cochacutan, at the foot of "El Carnicero," Cerrate 2294 (GH). Junin: Prov. Huan- cayo, Dist. Huancayo, foot of Nevada Huaytapallana. Sounders 1165 (GH). Cuzco: Prov. Cuzco, Hills of Saxi- huaman, Herrera 1349 (us), 2370 (F), 2380 (F). Prov. Cuzco, Saxihuaman. Tryon & Tryon 5348, in part (GH, us). 12. Asplenium rutaceum (Willd.) Men.. Abh. Senckenberg Naturf. Ges. 3: 173. 1858. Aspidium rutaceum Willd., Sp. pi. ed. 4, 5: 266. 1810. TYPE: Plumier, Traite foug. Amer. /. 57. 1805. TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 17 Asplenium perkinsii Jenman, Gard. Chron. 3, 19: 8. 1896. TYPE: Guyana (as British Guiana), Kaie- teur Falls, Perkins (holotype, NY!). Asplenium conquisitum Christ, Bull. Herb. Boissier 2, 7: 270. 1907. LECTOTYPE (designated by Max- on, Contr. U.S. Natl. Herb. 10: 488. 1908): Ja- maica, Maxon 1558 (P; isolectotype, us!). Plants epiphytic or terrestrial. Stem erect, pro- vided with linear or lanceolate scales, these to 3 mm long, clathrate, castaneous or gray-brown. Leaves 2-3-pinnate, rigid, fasciculate, to 40 cm long and 10 cm broad. Petiole 0-2 cm long, lus- trous, castaneous to atropurpureous, glabrous. Lamina thin-herbaceous, gradually reduced to a naked and flagelliform apex, this often with a pro- liferous and radicant tip, strongly reduced at base, surface apparently glabrous but sparsely provided abaxially with minute, appressed trichomes. Ra- chis lustrous, dark brown or atropurpureous, nar- rowly alate adaxially. Pinnae 1 5-32 pairs, patent to slightly ascending, or proximal ones deflexed, sessile, the basal pinnules commonly overlapping the rachis. Pinnules 6-10 pairs, inequilateral at base, with several pairs of spatulate or obovate segments, their apices obtuse to acute (occasion- ally apiculate). Veins simple in most ultimate seg- ments. Sori solitary on the vein of each segment. Indusia thin, subentire, yellowish or greenish. In wet forests, on tree trunks, in humus, or on banks of ravines, 800-2600 m, Cajamarca and Amazonas south to Cuzco. Greater Antilles; southern Mexico to Panama; the Guianas to Colombia and south to Bolivia. This is often confused with other 2-3-pinnate species of Asplenium in Peru, such as A. myrio- phyllumandA. radicansvur. uniseriale. However, only A. rutaceum has the combination of such diagnostic features as: petiole dark, lustrous, and nearly obsolete; lamina strongly reduced at base and apex, with a naked, flagelliform, and usually radicant tip. Lellinger (Proc. Biol. Soc. Wash. 98: 372. 1985) separated A. maxonii Lell. from A. rutaceum in South America and southern Central America on a number of subtle characters and cited one para- type from Peru: Woytkowski 34518 (us). How- ever, upon reexamination, he has advised (pers. comm.) that the latter is actually A. rutaceum, thus A. maxonii has not yet been found in Peru. The latter is presumed to differ in having adnate ulti- mate segments and the basal acroscopic lobe of pinnules usually bifid. Ultimate segments of A. rutaceum are short-stalked, and basal lobes are never bifid. Other features cited in the original description of A. maxonii vary too greatly in Pe- ruvian specimens to be of significant value. Cajamarca: Prov. Jaen, above Tabaconas, Quebrada de Pajonal, Fosberg 27803 (us). Amazonas: Prov. Bagua, E of La Peca, Barbour 2821 (F, MO, us). San Martin: Prov. Rioja, Venceremos, D. Smith 4432 (F, MO, uc). Huanuco: Prov. Leoncio Prado, between Tingo Maria and Pucallpa, Sullivan & Young 1 159 (MO). Pasco: Prov. Oxapampa, trail to summit of Cordillera Yanachaga, D. Smith et al. 7815 (F, MO, USM). Junin: Prov. Tarma, Rio Palca Valley, Hodge 6299 (GH). Ucayali (as Huanuco): Divisoria, Woytkowski 34518 (uc, us). Cuzco: Prov. Paucartambo, Cosnipata Valley, Rio Tono, Wachter et al. 187 (F). 1 3. Asplenium hallii Hooker, Sp. fil. 3: 202. 1 860. TYPE: Ecuador, forest of Esmeraldas, Hall (holotype, K!, frag., B; photo, us of B). Figures Ib-d. Asplenium spruceanum Hieron., Hedwigia 60: 260. 1918. TYPE: Brazil, San Gabriel, Spruce 2357 (not 2375 as in protologue) (holotype, B!, frag, us!; isotype, GH!). Plants epiphytic. Stem rather stout, erect or de- cumbent, provided with linear-lanceolate scales, these 2-3 mm long, clathrate, dark brown to black- ish. Leaves 1 -pinnate-pinnatifid or -pinnatisect, erect, fasciculate, 18-40 cm long, (1.5-)2-9 cm broad. Petiole 1-6 cm long, lustrous, castaneous to atropurpureous, glabrous. Lamina thin- to firm- herbaceous, glabrous, gradually reduced to apex and base, the tip attenuate, usually flagelliform and/or proliferous, rachis dark and lustrous. Pin- nae 20-30 pairs, larger ones (1.5-)2—4 cm long, patent, or a few proximal ones deflexed, sessile, pinnatifid, or incised nearly to the costa at base, the basal acroscopic segment commonly overlap- ping the rachis. Veins simple in most segments, forked to pinnate in basal ones. Sori solitary on each vein. Indusia thin, yellowish or greenish. In deep forests, on stumps or tree trunks and branches, 100-200 m, San Martin and Loreto. Colombia; Ecuador; Peru; Amazonian Brazil. Some confusion has attended the nomenclature and descriptions of this species and its synonym. Hieronymus separated his A. spruceanum from A. hallii on the basis of several quantitative charac- ters and cited as its type a paratype of the latter (incidentally citing the number incorrectly as 2375). During the present study, comparison of the type material showed there is no reason to recognize 18 FIELDIANA: BOTANY two species. Furthermore, in the protologue, fol- lowing the description of A. hallii, Hooker made a curious statement: "My specimens from Col. Hall have the pinnae again pinnated, especially in the lower half . . . ." The Hall specimen at Kew exhibits nothing of the kind and, in fact, is iden- tical to Spruce 2357, except the pinnae are less deeply incised and the segments a little broader. Mexia 6184 (F, GH, MO, uc, us) from Loreto has dull, gray-green rachis and petiole, stem scales light brown and ovate, and roots somewhat golden-to- mentose (all characters similar to those of A. au- riculatum) but otherwise matches A. hallii. This may be a hybrid involving the two species, for many sporangia are barren or with some spores malformed. San Martin: Lamas, Santa Rosa de Davidcillo, near road to Tioyacu, Knapp & Mallet 7195 (F, MO, uc). Lo- reto: Mishuyacu, near Iquitos, Killip & Smith 29975 (F, GH, us). Prov. Maynas, Mishara, Lopez et al. 8650 (HUT). Prov. Requena, Jenaro Herrera, N of Requena, Tovar 6968 (USM). La Victoria on the Amazon River, LI. Wil- liams 2989 (F, us). 14. Asplenium macrurum Mickel & Stolze, in Stolze, Flora of Ecuador 23: 41. 1986, nom. nov. Asplenium longicaudatum Mickel & Stolze, Amer. Fem J. 74: 115. 1984, nom. illeg., not A. longicauda- tum Bonaparte. 1917. TYPE: Ecuador, Prov. Mo- rona-Santiago, Cordillera Cutucu, Camp E-1283 (holotype, NY!; isotype, s). Plants terrestrial or low-epiphytic. Stem stout, decumbent to erect, provided with ovate to lan- ceolate, acute scales, these 2-4 mm long, 0.6-1 mm broad, dark brown or gray-brown, or often bicolorous with narrow, lighter brown margins, subclathrate, the lumina isodiametric or slightly elongated. Leaves 1 -pinnate, fasciculate, to 1 mm long and 18 cm broad. Petiole 10-30 cm long, lustrous or sublustrous, dark brown to atropur- pureous, essentially glabrous. Lamina thin- to firm- herbaceous, glabrous, lanceolate or deltate-lanceo- late, not or scarcely reduced at base, gradually reduced to a long, flagelliform apex. Rachis gla- brous, not alate, dark reddish brown, the flagel- liform tip often proliferous but fragile and usually broken off in dried specimens. Pinnae 10-17 pairs, to 12 cm long and 1 .8 cm broad, sessile or short- stalked, equilateral except at base, there broadly cuneate to truncate acroscopically, narrow-cune- ate basiscopically, the apex narrowly acute to at- tenuate, margins subentire to broadly crenulate. Veins l(-2)-forked. Sori 5-8 mm long, linear or narrow-oblong, slightly arcuate. Indusia yellow- ish, firm, subentire. On the floor of deep, wet forests or at the base of tree trunks, 1000-1700 m, Amazonas, Ucayali. Venezuela; Ecuador; Peru. Heretofore known only by the type collection from Ecuador, several collections of the species now have been made in Peru. It is similar in ap- pearance to A. radicans var. cirrhatum but differs significantly in stem scales and in size and shape of pinnae. The Peruvian, especially Schunke, col- lections closely match the type, but pinnae of the two Harbour specimens are not attenuate as on the type; they are merely narrow-acute. Amazonas: Prov. Bagua, 12 km E of La Peca, Barbour 2404 (MO, uc), 2507 (uc). Ucayali (as Loreto): Prov. Coronel Portillo, Dist. Padre Abad, Rio Chino, J. Schunke V. 9197 (MO, uc). 15. Asplenium radicans L., Syst. nat. ed. 10, 2: 1323. 1759. LECTOTYPE (designated by Morton & Lellinger, Mem. New York Bot. Gard. 1 5: 38. 1966): Jamaica, Browne 92 (LINN 1250.16, right-hand specimen; photo, us). Plants terrestrial, rarely epiphytic or epipetric. Stem stout, erect, provided at apex with linear or lanceolate, attenuate, clathrate scales, these 2-5 mm long, gray-brown to blackish. Leaves 1 -pin- nate to 3-pinnate-pinnatisect, fasciculate, to 1 m long and 25 cm broad. Petiole 10-30 cm long, lustrous, dark brown to atropurpureous, essen- tially glabrous. Lamina thin- to firm-herbaceous, lanceolate to deltate-ovate, not or scarcely reduced at base, gradually tapering to a radicant apex. Ra- chis essentially glabrous, bisulcate adaxially, not or conspicuously green-alate, lustrous, terminat- ing in a naked and flagelliform apex. Pinnae nu- merous, extremely variable: simple, entire to cre- nate-serrate and commonly 2-5 cm long by 1 cm broad in var. cirrhatum, to 3-pinnate and 12-15 cm long by 4-5 cm broad in var. uniseriale; sessile or subsessile, patent to slightly ascending. Veins commonly 1-2-forked, or simple in ultimate seg- ments of decompound varieties. Sori elongated, or moderately so in decompound varieties. In- dusia thin, yellowish or greenish, oblong to elliptic, the margins subentire. The species occurs in the West Indies, and from Mexico to Bolivia and Brazil. TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 19 Few fern species are as highly variable in lamina architecture as Asplenium radicans. A combina- tion of features distinguish it from other species in the genus: lustrous, atropurpureous petiole and rachis; flagelliform, proliferous leaf tips; scarcely reduced lamina base. However, the lamina may be simply pinnate, or 3-pinnate, or varying in an infinite number of degrees between each extreme. This has prompted some authors to recognize four or more species, each based solely on very subtle differences. Obviously, questions posed by this and similar aggregate species cannot be satisfactorily resolved by comparison of herbarium specimens, within the limitations of a floristic treatment. Field and greenhouse observations coupled with cyto- logical studies are needed. A good beginning was made by Walker (Trans. Roy. Soc. Edinburgh 66: 169-237. 1966; and 69: 109-235. 1969), who re- vealed that some crossing is involved within and also out of the species complex. Tentatively, four varieties are recognized, which may or may not represent geographic or ecological taxa. Key to Varieties a. Lamina 1 -pinnate or 1 -pinnate-pinnatisect. Secondary segments (if any) adnate to a broadly alate costa b b. Pinnae lobed to deeply pinnatisect (but not dissected entirely to the costa) . . 1 5a. var. radicans b. Pinnae entire to deeply crenate-serrate, not lobed, not or rarely auriculate 1 5b. var. cirrhatum a. Lamina 2- to 3-pinnate, at least the larger secondary segments definitely stalked c c. Pinnae 1 -pinnate (at least in proximal half), the pinnules coarsely dentate to deeply pinnati- sect 1 5c. var. partitum c. Pinnae 1 -pinnate-pinnatisect to 2-pinnate, the pinnules (at least proximal one) with 1 -several free segments 1 5d. var. uniseriale 1 5a. Asplenium radicans var. radicans. Asplenium rhizophyllum L., Sp. pi. 1540. 1763, based on the same protologue as A. radicans. Not A. rhizophyllum L., 1753. (For full list of synonyms of A. radicans and varieties, see Morton and Lellinger, 1966.) Pinnae pinnatifid to deeply pinnatisect, basal segments often free, but not fully stalked, obtuse to subacute, subentire to dentate distally. Rare, on slopes or ravine banks in wet forests, 600-2300 m, Amazonas, San Martin, Huanuco. West Indies; southern Mexico south to Brazil and Bolivia. Degree of lamina dissection merges gradually into that of var. partitum, the subtle distinction here being that the latter has clearly stalked sec- ondary segments with coarsely dentate apices. Amazonas: Prov. Bongara, Shillac, D. Smith & Vas- quez 4839 (MO, uc). San Martin: Mt. Guayrapurima, near Tarapoto, Spruce 4021 (BR, GH, us). Huanuco (as San Martin): East of Tingo Maria, Allard 21562 (GH). 15b. Asplenium radicans var. cirrhatum (Willd.) Rosenst., Hedwigia 46: 102. 1906. Asplenium cirrhatum Willd., Sp. pi. ed. 4, 5: 321. 1810. TYPE: Guadeloupe, L. C. Richard (holo- type, B, Herb. Willd. 19894, frag., NY!; isotype, p; photos, F, GH, us of B). Pinnae entire to deeply crenate-serrate, acro- scopically truncate and sometimes subauriculate, basiscopically narrow-cuneate to excavate. In forests, on sandy or rocky soil, or on old logs or bases of tree trunks, 300-1700 m, Amazonas and Loreto to Cuzco and Puno. Guadeloupe; Greater Antilles; southern Mexi- co; Guatemala; Nicaragua to Brazil and Bolivia. Amazonas: Prov. Bagua, valley of Rio Maranon above Cascadas de Mayasi, Wurdack 1871 (GH, us). San Mar- tin: Prov. Mariscal Caceres, Dist. Uchiza, SE of Nuevo Progreso, Schunke V. 3159 (F, GH). Loreto: Valseca-Ru- dolpho, along Rio Corrientes, McDaniel & Marcos 11061 (GH, MO). Huanuco: Prov. Leoncio Prado, Castilo Alto, Plowman 5849 (F, GH, USM). Pasco: Prov. Oxapampa, Palcazu Valley, between Iscozacin and Villa America, D. Smith 3871 (MO, uc). Junin: Schunke Hacienda, above San Ramon, Killip & Smith 24577 (F, GH). Cuzco: Prov. Paucartambo, Cosnipata Valley, Rio Tono, Wachter et al. 201 (F, USM). Madre de Dios: Tambopata Nature Reserve, between the lodge and Rio la Torre, Funk et al. 8377 (uc, us). Puno: Prov. Carabaya, Palmera, Vargas 16150 (GH). 1 5c. Asplenium radicans var. partitum (Klotzsch) Hieron., Bot. Jahrb. Syst. 34: 464. 1904. 20 FIELDIANA: BOTANY Aspleniumflabellulatum var. dentatum Klotzsch, Lin- naea 9: 71. 1834. SYNTYPES: Columbia [Ven- ezuela], Moritz 44 (B?, P!; photos, F & us of P). Venezuela, prope Caracas (B?). Aspleniumflabellulatum Kunze var. partitum Klotzsch, Linnaea 20: 357. 1847. LECTOTYPE (designat- ed by Morton & Lellinger, Mem. New York Bot. Card. 15: 37. 1966): Venezuela, Caracas, Otto 651 (holotype, B; photo, us). Asplenium partitum (Klotzsch) C. Chr., Index fil. 1 25. 1905. Asplenium radicans var. dentatum (Klotzsch) Bonap., Notes Pteridologiques 7: 349. 1918. Pinnae 1 -pinnate, with at least several pairs of free pinnules, these coarsely dentate and often deeply incised. On slopes and ravine banks of wet forests, 650- 2400 m, Amazonas, Huanuco, Pasco, Junin, Cuz- co. West Indies; Mexico south to Peru and Bolivia. Stolze (1986) had not seen type material but cited A. flabellulatum as a synonym of var. par- titum, based on the original description. However, having now examined the lectotype (with fully tri- pinnate lamina), it can be seen that the former name is more properly placed under var. unise- riale. This is typical of the confusion surrounding the many names and variants of A. radicans, all of which have been distinguished merely on mi- nute degrees of variation in lamina dissection. Amazonas: Prov. Bongara, Shillac, Smith & Vdsquez 4893 (MO). Huanuco: La Divisoria, 25 km NE of Tingo Maria, Moran 3700 (MO). Pasco: Prov. Oxapampa, Dist. Oxapampa, Rio San Alberto, Leon 633 (F, GH). Junin: Chanchamayo Valley, C. Schunke 109 (F), 110 (F), 1452 (F). Above San Ramon, C. Schunke A2 11 (uc). Prov. Tarma, Agua Dulce, Woytkowski 3 5429 (MO, uc). Cuzco: Torontoy, Urubamba Valley, Cook & Gilbert 1097 (us). Prov. La Convention, Rio Apurimac, Davis et al. 1299 (F, GH). 15d. Asplenium radicans var. uniseriale (Raddi) Gomez, Brenesia 8: 53. 1975. Asplenium uniseriale Raddi, Opusc. Sci. 3: 29 1 . 1819. TYPE: Brazil, Raddi (holotype, n). Asplenium flabellulatum Kunze, Linnaea 9: 71. 1834. TYPE: Peru, Pampayacu (Huanuco), Poeppig (holotype, LZ, destroyed; lectotype, B!, frag., us!, photo, us). Pinnae 2-3-pinnate, with basal pinnules strong- ly overlapping the rachis, most pinnules with 1 to several free segments, these obovate to cuneiform, or once again pinnate. Terrestrial in and at edges of forests, 100-2700 m, Cajamarca to Loreto, south to Huancavelica and Cuzco. West Indies; southern Mexico to Brazil and Bo- livia. With this probably should be included A. bal- liviani Rosenst. of Bolivia, which Rosenstock dis- tinguished from A. uniseriale as having slightly less divided laminae (2-pinnate-pinnatifid). Asple- nium radicans var. uniseriale (including A. flabel- lulatum) has laminae 3-pinnate or more. Detailed analysis of the species complex is needed. Studies by Stolze in Guatemala and Ecuador, and now in Peru, have revealed no other morphological char- acters or ecological data to corroborate differences in degree of lamina dissection. Monographic study possibly may substantiate the premise that A. rad- icans is merely a species with highly varied leaf architecture, whether this conclusion is "conve- nient" to classification or not. Cajamarca: Prov. Cutervo, Grutas de San Andres, Lla- tas Quiroz & Suarez 2735 (F, uc). Amazonas: Prov. Ba- gua, Cordillera Colon SE of La Peca, Barbour 4003 (MO). San Martin: Zepelacio, near Moyobamba, Klug 3489 (F, GH, MO), 3490 (B). Loreto: Mishuyacu, near Iquitos. Klug 208 (F), 392 (F). Huanuco: Prov. Huanuco, Carpish, As- plund 12835 (s). Pasco: Prov. Oxapampa, Dist. Oxa- pampa, Rio San Alberto, Leon 651 (F, GH, USM). Junin: Chanchamayo Valley, C. Schunke 830 (F), 1399 (F). Huancavelica: Prov. Tayacaja, Marcavalle, Tovar 4755 (GH). Cuzco: Rio Tacate, Bues 1745 (us). 16. Asplenium repens Hooker, Sec. cent, ferns, /. 31. 1860. TYPE: Ecuador (Napo), growing in woods in Archedona, Jameson 787 (holotype, K.!). (Jameson 786 cited in the protologue.) Plants epiphytic, rarely terrestrial. Stems mi- nute, borne on long-creeping, stoloniform roots. Leaves remote, borne singly, or occasionally in pairs, 2-pinnate (at least proximal ones), 1 .5-5 cm long. Petiole to 0.5 cm long, yellow- or gray-green, at the base bearing a few, inconspicuous scales, these castaneous, clathrate, ovate, about 0.5 mm long. Lamina membranaceous, oblong or deltate- oblong, reduced to a pinnatifid apex, not or scarce- ly reduced at base. Rachis glabrous, lacking pro- liferous buds. Pinnae 4-8 pairs, approximate or subdistant, stalked, essentially pinnate, typically with only a single pinnule borne on the acroscopic side of the costa (thus nearly bifoliolate). ultimate segments about 3 mm long, cuneate, flabelliform, subdigitately incised at apex. Veins commonly 1 -forked in each ultimate segment. Sori solitary TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 21 on each ultimate segment. Indusia thin, broad, yellowish or greenish brown, entire. Rarely collected in Peru, on trunks or branches of trees, rarely on wet ground or mossy rocks, 1700-1800 m, Huanuco, Junin, Cuzco. Colombia; Ecuador; Peru; Bolivia. Jameson 786 was cited as the type by Hooker, and he also referred to it in Species Filicum (3: 194. 1860). However, the specimen in the holo- type folder is 787. A diligent search by Peter Ed- wards (K), as well as a canvass of other herbaria, failed to yield a collection numbered 786. Thus, it appears likely Hooker erred in citing the type number. Although shape of ultimate segments and lam- ina of A. repens are quite different from those of A. delicatulum, the two species are very closely related. Both have small, delicate and glabrous leaves borne on stoloniform roots, with a few clathrate petiole scales about 0.5 mm long. That neither species is well represented in herbaria is probably because they are so small and incon- spicuous. Huanuco: Prov. La Divisoria, 25 km NE of Tingo Maria, Moran & Fernandez 3704 (MO, uc, USM). Junin: La Merced, Chanchamayo, Soukup 1009 (F). Cuzco: Tablahuas, Lucumayo Valley, Biies 1034 (us). 17. Asplenium triphyllum Presl, Reliq. haenk. 1: 45. 1825. TYPE: "Hab. in Cord. Peru" (but possibly Ecuador), Haenke (holotype, PRC!; photo, F). Asplenium tenue Presl, Reliq. haenk. 1 : 44, t. 6, f. 5. 1825. TYPE: Ecuador (as Peru), monte Chim- borazo, Haenke (holotype, PR!; isotype, PRC!; probable isotype, NY!; photos, F & GH of PR, GH of NY). Asplenium ternatum Presl, Reliq. haenk. 1: 45. 1830. TYPE: in vallibus cordillerarum Peruviae, Haenke (PR!). Asplenium rhomboideum Brack., U.S. expl. exped. 1 6: 156, t. 21, f 2. 1854. TYPE: (Huanuco) Peru, Andes of Banos, Brackenridge 24 (holotype, us!). Plants epipetric, rarely terrestrial. Stem small to stout, provided with linear or linear-lanceolate scales, these obscurely clathrate, 1-2 mm long, blackish. Leaves fasciculate to densely caespitose, (4-)6-30 cm long, 0.5-1.8 cm broad. Petiole 1-10 cm long, dull, or rarely sublustrous, gray-brown to castaneous, sparsely provided with minute, glandular trichomes. Lamina thin-herbaceous to chartaceous, linear, gradually tapering to a pin- natifid apex, abruptly to gradually reduced at base. Rachis rigid to flexuous, sparsely to moderately provided with filiform scales and pluricellular tri- chomes, not or inconspicuously alate, often bear- ing a proliferous bud in a pinna axil. Pinnae nu- merous, subsessile to short-stalked, subdistant to densely crowded, most of them trifoliolate or bi- foliolate, or occasionally with another pair of lat- eral pinnules, ultimate segments obovate or ob- lanceolate, entire to bifid. Veins simple (rarely 1 -forked) in each ultimate segment. Sori solitary on the vein of each segment. Indusia thin, pale yellowish or translucent, subentire. In crevices and ledges of rock walls, at cave mouths, among rocks and boulders, rarely in wet, mossy, soil, (2800-)3 100-4800 m, from Caja- marca south to Arequipa and from Huanuco south to Puno. Colombia to Bolivia and Argentina. The taxonomy of this species and its near rel- atives has been very confused; consequently, a rather lengthy discussion is in order here. The type collection of A. tenue cited above contains leaves merely 2—4 cm long and 0.6-0.8 mm broad, and Stolze (1986) thought they closely resembled those of A. fragile (= A. peruvianum). Although pinnae are not conspicuously trifoliolate, close exami- nation reveals that many of them are deeply cleft; therefore, they resemble many of the smaller, bi- foliolate pinnae of A. triphyllum. The stem scales of the holotype of A. tenue are scarcely or very narrowly clathrate, like those of A. triphyllum, and not obviously clathrate like those of A. peruvia- num. Thus, it seems that A. tenue is nothing more than a depauperate form of A. triphyllum. The types of both A. tenue and A. triphyllum were collected by Haenke, and on the labels are marked "Peru." However, boundaries of the two countries were often confused, and the protologue of the former cited that the plants were actually collected in Monte Chimborazo, which is in Ec- uador. Presl based his descriptions of both species on the Haenke collections in hand, and it is quite conceivable that Haenke found both at the same collecting site. Despite the fact that these plants evidently represent extreme forms of the same spe- cies, Presl chose to describe them as different. Asplenium trilobatum C. Chr. of Bolivia is very closely related and also may be merely a depau- perate form. Other variability in leaves of Asplenium tri- phyllum similarly prompts their recognition as dif- ferent species. At one extreme, the stem is rather 22 FIELDIANA: BOTANY small, the rachis slender and flexuous, and the pinnae thin-herbaceous and subdistant; in other leaves (as in the type of A. ternatum), the stem is stout, the rachis stout and rigid, and the pinnae chartaceous and crowded (even imbricate). When proliferous buds are present, they tend to be found only on the thin and flexuous leaves. Nevertheless, it is doubtful that two distinct taxa are represented here, as too many intermediate conditions occur, and the extreme forms may simply indicate leaf development responding to different habitats. Field studies are needed to corroborate this. Besides being closely related to A. peruvianum, A. triphyllum also is part of a variable species com- plex involving A. myriophyllum, A. haenkeanum, and perhaps some others; in fact, several speci- mens in Peru are intermediate between A. tri- phyllum and A. haenkeanum. See treatment of the latter for further discussion. Cajamarca: Prov. Cajamarca, Cerro "Cumbe Mayo," Sanchez V. 39 (GH, HUT, us). La Libertad: Prov. Santiago de Chuco, Laguna La Victoria, Sagdstegui el al. 6189 (GH, HUT). Ancash: Prov. Huari, Huascaran National Park, D. Smith el al. 12411 (F, MO). Huanuco: Chasqui, Macbride & Featherstone 1757 (F, GH, us). Lima: Prov. Huarochiri, above Chumpicocha, Cerrate 2005 (F, GH, USM). Junin: Road from Tarma to La Oroya, Correll & Smith P786 (GH, us), P787 (GH, MO). Huancavelica: Prov. Tayacaja, Tocas, between Colcabamba and Paucarbam- ba, Tovar 2103 (GH, USM). Ayacucho: Between Quinua and road to Tambo, Soukup 6185 (us). Cuzco: Prov. Cuzco, near Saxihuaman, Tryon & Tryon 5361 (F, GH, us, USM). Arequipa: Pichu Pichu, Stafford 676. Puno: Near Puno, Soukup 302 (F, uc). 18. Asplenium divaricatum Kunze, Linnaea 9:71. 1834 (not Wallich, 1829, nom. nud.). TYPE: Peru, "Cerro San Christobal (sic), prope Cas- sapi," Poeppig (diar. 7757), 1829 (holotype, LZ, destroyed; isotypes, BM!, P!, w!; photos, F, GH, uc & us of BM; F of w). Plants epipetric, rarely terrestrial. Stem small to stout, erect, provided with linear or linear-lanceo- late scales, these obscurely clathrate, 1-2 mm long, bicolorous, blackish with narrow but conspicuous brown bands of marginal cells. Leaves caespitose, 2-pinnate-pinnatisect to 3-pinnate, 5-30 cm long, 1-7 cm broad. Petiole 0-6 cm long, dull, reddish to grayish brown. Lamina thin- to firm-herba- ceous, linear to elliptic, gradually reduced to apex and base. Rachis glabrous or with a few filiform scales, narrow-alate, lacking proliferous buds. Pin- nae numerous, sessile, crowded to imbricate, basal pinnules overlapping the rachis. Pinnules of larger pinnae 6-10 pairs, bifoliolate or with several pairs of ultimate segments, segments commonly less than 2 mm long, obtuse, fertile ones nearly twice as broad as adjacent sterile ones. Veins simple in each ultimate segment. Sori solitary on the vein of each segment, at maturity usually projecting beyond segment margin. Indusia thin, translucent, elliptic- oblong, margins entire. Commonly in crevices of dry rocky walls or in soil among rocks, 700-2700 m, generally along the Cordillera Central from Amazonas south to Cuz- co. Endemic to Peru. Superficially, this resembles A. rutaceum, but the latter differs, among other characters, in its darker, lustrous petiole and radiant leaf apex. A distinctive feature of most leaves of A. divaricatum is the relative size and shape of fertile and sterile segments, which is especially evident on pinnules that are not again pinnate but merely bifoliolate. The fertile segment is usually obovate, while the opposing sterile segment is often about half its width and nearly linear. Mickel and Beitel (Mem. New York Hot. Gard. 46: 55-56. 1988) included A. divaricatum in their Pteridophyte Flora of Oaxaca and cited its distri- bution from Mexico to Peru. They maintained that this name applies to specimens from Central and South America that traditionally have been in- cluded under A. myriophyllum and that true A. myriophyllum occurs only in the West Indies. However, the Oaxaca fern they described and il- lustrated as A. divaricatum and a supposed type fragment (NY) are substantially different from ac- tual type material (BM, P, w) of that species and from specimens of true A. divaricatum in Peru (cited below). References to neither the bicolorous stem scales nor the size and differences in fertile and sterile segments are seen in Mickel's descrip- tion and illustration. There is a fragmentary specimen at New York labeled simply "Asplenium rhizophyllum Kze., Peru, Poeppig," which is identified and filed as a type collection of A. divaricatum. Labels of all oth- er type specimens (cited above) contain not only the identification as A. divaricatum but also the full data of type locality. Asplenium rhizophyllum is a synonym of A. myriophyllum, and indeed the "type collection" at New York matches the type of the latter well. Perhaps it is this specimen that prompted the incorrect application of the name of A. divaricatum to the Mexican plants. This is also typical of the understandable confusion that has long attended the circumscription of species in this TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 23 highly variable complex. See discussion of A. haenkeanum for further details. Amazonas: Prov. Chachapoyas, Cerros Calla Calla, above Leimebamba, Hutchison & Bennett 4545 (F, GH, uc, us). Hacienda Limon, W of Balsas, Osgood & An- derson 57 (F), 63 (F, us). San Martin: Trip from Pacas- mayo to Moyobamba, Stiibel 1023 (B, us). Huanuco: Muna, Bryan 432 (F, us). Huacachi, estacion near Muna, Macbride 3870 (F, GH, us, w). Junin: Prov. Tarma, near San Felix Bridge, Ferreyra 14955 (GH, USM). Prov. Tar- ma, SW of Huacapistana, Tryon & Tryon 5433 (GH). Cuzco: Masias, high above Apurimac River, Sues 1383 (us). 1 9. Asplenium haenkeanum (Presl) Hieron., Hed- wigia47: 233. 1908. Athyrium haenkeanum Presl, Tent, pterid. 98. 1836. TYPE: Peru, near city of Huanuco, Haenke 52 (holotype, PRC!). Asplenium cladolepton Fee, Mem. foug. 7: 55. 1857. TYPE: Colombia, Prov. Ocana, Schlim 324 (ho- lotype, P; isotype, L; frag., us!: photos, F & us of L). Asplenium cladolepton var. angustifolium Hieron., loc. cit. SYNTYPES: Peru, (Loreto) prope Cuelapa (Loreto), Stiibel 1019b (B); supra Tambo Mayo, Amazonas, Stiibel 1043 (B). Asplenium cladolepton var. minor Hieron. loc. cit. TYPE: Peru (Loreto) prope Cuelapa, Stiibel 101 9a (B). Plants epipetric (rarely terrestrial). Stem small to rather stout, erect or suberect, provided with linear or deltate-lanceolate scales, these obscurely clathrate, 1.5-3 mm long, blackish, or occasionally faintly brown-margined. Leaves fasciculate, 2-pinnate, 12-38 cm long, 2.2-5(-7) cm broad. Petiole 3-1 2 cm long, dull reddish or grayish brown. Lamina thin-herbaceous or membranaceous, lin- ear, gradually reduced to apex and base. Rachis glabrous or with a few filiform scales, narrow-alate, lacking proliferous buds. Pinnae 20-28 pairs, ap- proximate to subdistant, mostly patent, or prox- imal ones deflexed, sessile, basal pinnules over- lapping the rachis. Pinnules of larger pinnae 3-6 pairs, shallowly to deeply bifid, fertile and sterile segments subequal, 2-3 mm long. Veins simple in each ultimate segment. Sori solitary on veins. In- dusia thin, yellowish or greenish, margins entire. In wet places, on or among rocks, or in crevices of rock walls, 2900-3800 m, Amazonas, La Liber- tad, Loreto, Huanuco, Pasco, Apurimac, Cuzco. Colombia; Venezuela; Ecuador; Peru. This is at the morphological center of a species complex involving A cristatum, A. myriophyllum, A. triphyllum, and, in a broader sense, A. divari- catum. These species are alike in many significant characters, although they have been separated tra- ditionally by some variable, quantitative, features: lamina dissection and shape, degree of reduction of lamina base, degree of pinna angle, and number of pinnules. Typical A. triphyllum has very long and narrow leaves with bi- or trifoliolate pinnae, in the axils of which are frequently borne proliferous buds, but not infrequently there are pinnae with an ad- ditional one or two lateral segments, whose con- dition thus approaches that of less robust forms of A. haenkeanum (3-6 pairs of pinnules). How- ever, no proliferous buds yet have been found in the latter species. Conversely, more robust leaves of A. haenkeanum (narrow-elliptic, with up to 6 pairs of pinnules) resemble narrower forms of A. myriophyllum (7-10 pairs of pinnules). The latter is characteristically terrestrial, growing at 1400- 2700 m in Peru, and A. haenkeanum is epipetric, at 2900 m or higher. Asplenium cristatum and A. myriophyllum have been distinguished by slight differences in scale color, pinna shape, and degree of reduction in the lamina base, which is seen under/!, cristatum. Some specimens of A. myriophyllum with un- usually small ultimate segments might be confused with A. divaricatum, but the latter commonly can be distinguished by its conspicuously bicolorous stem scales and the difference in size and shape of the fertile and sterile segments. Obviously the en- tire complex in the Neotropics is badly in need of detailed study, and it is with strong reservations that its various components are separated in this flora. La Libertad: Prov. Bolivar, Bolivar, Lopez & Sagds- tegui 3274 (GH, HUT). Huanuco: Chasqui, Macbride & Featherstone 1756 (F, us, in part). Pasco: Huariaca, Bry- an 168 (F). Road to Cerro de Pasco, Gentry et al. 37486 (F, MO). Apurimac: Prov. Andahuaylas, Huancarama, Vargas 8827 (uc). Cuzco: Ollantaytambo, Cook & Gilbert 395 (us). Prov. Urubamba, Chincheros, King et al. 141 (F, a variant). 20. Asplenium cristatum Lam., Encycl. 2: 310. 1786. TYPE: Locality and collector undesig- nated (holotype, P, Herb. Jussieu 1276; photo, us). Asplenium cicutarium Sw., Prodr. 130. 1788. TYPE: Jamaica, Swartz (holotype, s!; isotype, SET; pho- tos, F & us of s). 24 FIELDIANA: BOTANY Plants terrestrial, rarely epiphytic or epipetric. Stem stout, erect, provided with linear or deltate- lanceolate scales, these 2-3 mm long, subclathrate, gray-brown or blackish, occasionally faintly brown- margined. Leaves fasciculate, 2-pinnate-pinnati- sect to 3-pinnate, to 60 cm long and 1 5 cm broad. Petiole 5-25 cm long, dull gray or reddish brown to blackish. Lamina thin-herbaceous or membra- naceous, elliptic or oblong-lanceolate, gradually reduced to a pinnatifid apex, scarcely reduced at base, but occasionally with 1 or 2 proximal pairs of pinnae strongly reduced. Rachis glabrous or with a few, scattered, filiform scales, alate, lacking pro- liferous buds. Pinnae 1 5-20 pairs, approximate to crowded or imbricate, most of them patent, ob- long, the margins parallel for much of their length, sessile, basal pinnules overlapping the rachis. Pin- nules of larger pinnae 9-16 pairs, deeply pinnati- sect or pinnate, ultimate segments 2.5—4 mm long, narrow-elliptic, acute. Sori 3-8 on a pinnule. In- dusia thin, grayish or yellowish, margins subentire or slightly erose. On wet forest floor, in wooded ravines, rarely on or among rocks or on bases of tree trunks, 250- 2400 m, Cajamarca and Amazonas south to Aya- cucho and Madre de Dios. United States (Florida); West Indies; Mexico to the Guianas, south to Bolivia and Brazil. With this perhaps should be included A. myri- ophyllum, which shares nearly the same range and habitat and has been distinguished by subtle dif- ferences such as shape of lamina base, pinnae, and ultimate segments. See A. haenkeanum for further discussion. Cajamarca: Prov. Hualgayoc, Monte Seco, Soukup 381 1 (F, us). Amazonas: Prov. Bagua, Cerro Tapur near Rio Utcubamba, Hutchison 1480 (F, uc, us). San Martin: Near Tarapoto, Spruce 3975 (BR). Huanuco: Prov. Hua- nuco. Tingo Maria, Asplund 13201 (s). Mima, Bryan 424 (F). Pasco: Oxapampa, Soukup 2355 (F, GH). Junin: Chanchamayo Valley, C. Schunke 7 (F, us), 65 (F, us). Ayacucho: Aina [Ayna], between Huanta and Rio Apu- rimac, Killip & Smith 22838 (us). Cuzco: Prov. La Con- vention, Rio Apurimac below Puerto Capiro, Davis et al. 1293 (F, GH). Madre de Dios: Maldonado, Lopez et al. 4588 (GH). 21. Asplenium myriophyllum (Sw.) Presl, Reliq. haenk. 1:48. 1825. Caenopteris myriophylla Sw., J. Bot. (Schrader) 1800(2): 50. 1801. TYPE: Jamaica, Swam (ho- lotype, s!; photo, F). Plants terrestrial, rarely epipetric or epiphytic. Stem small to rather stout, erect or suberect, pro- vided with linear or deltate-lanceolate scales, these obscurely clathrate, 3-4 mm long, cast an ecus to gray-brown or blackish, occasionally faintly brown- margined. Leaves fasciculate, 2-pinnate-pinnati- sect or 3-pinnate, 12-35(-40) cm long, 3-7 cm broad. Petiole 3-15 cm long, dull reddish or gray- ish brown. Lamina thin-herbaceous or membra- naceous, ovate to elliptic, gradually reduced to a pinnatifid apex, abruptly reduced at base, with 2- 4(-6) proximal pairs of pinnae much shorter than central ones. Rachis glabrous or with a few, scat- tered, filiform scales, alate, lacking proliferous buds. Pinnae 15-28 pairs, approximate to subdistant, mostly ascending, broadest near base, tapering gradually or abruptly to apex, sessile, basal pin- nules overlapping rachis. Pinnules of larger pinnae (6-) 7-10 pairs, with one or more free ultimate segments, these 2-3 mm long, mostly obovate and obtuse to subacute. Veins simple in each ultimate segment. Sori solitary on the veins. Indusia thin, yellowish or greenish, margins entire. Infrequent and widely scattered in Peru, on moist forest floor, rarely on rocks, 1400-2700 m, Ca- jamarca, Amazonas, Huanuco, Pasco, Junin. United States (Florida); Greater Antilles; Mex- ico to Peru and Bolivia. This is the most highly dissected of several spe- cies in a confusing complex. See further discussion under A. haenkeanum and A. divaricatum. Cajamarca: Prov. Santa Cruz, Bosque de Monteseco, Sagdstegui et al. 12387 (F, HUT, MO, uc). Colasay, Woyt- kowski 7014 (GH, MO, us). Amazonas: Prov. Bagua, Cor- dillera Colan SE of La Peca, Barbour 3896 (F, MO). Hua- nuco: Mima, Woytkowski 5215 (GH, MO). Prov. Leoncio Prado, km 478 on Lima-Tingo road, Young & Sullivan 850 (F, MO). Pasco: Prov. Oxapampa, 2 km from Oxa- pampa, Leon 496 (USM). Junin: Huancayo, Soukup (F). 22. Asplenium delicatulum Presl, Reliq. haenk. 1: 47, /. 7,f. 5. 1825. TYPE: Ecuador, "in Cor- dilleris Quitensibus," Haenke (holotype, PRC!; isotype, HBG; photos, F, GH, us of HBO). Plants epiphytic or epipetric. Stems very small, crowded to subdistant, borne on (usually) long- creeping, stoloniform roots. Leaves borne singly or in small fascicles, 2-3-pinnate, 4— 7(-9) cm long. Petiole 1.5—4 cm long, yellow- or gray-green, with a few inconspicuous scales at the base, these dark brown, clathrate, ovate, 0.4-0.6 mm long. Lamina TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 25 membranaceous, deltate-ovate, reduced to a pin- natifid apex, not or scarcely reduced at base. Ra- chis glabrous, lacking proliferous buds. Pinnae 5- 8 pairs, approximate or subdistant, stalked, with 2-3 pinnules, these simple or again pinnate, ulti- mate segments 2—4 mm long, linear or narrow- oblanceolate, acute. Veins simple in each ultimate segment. Sori solitary on the veins. Indusia thin- textured, narrow-elliptic, yellowish or grayish green, entire. In wet forests and wooded ravines, on tree trunks, branches, and fallen logs, and on wet, moss-cov- ered rocks, 500-1500 m, San Martin, Huanuco, Pasco, Ucayali. Colombia; Venezuela; Ecuador; Peru; Chile. Although more than a dozen collections have been found in herbaria thus far from Peru (most of these in Huanuco), only a few specimens have been seen from Colombia and Venezuela, and ap- parently the species is represented to date in Ec- uador only by the type. As with the closely related A. repens, the paucity of existing collections is like- ly due to the inconspicuous nature of the plant rather than to its actual rarity. See A. repens for further discussion. San Martin: Prov. Mariscal Caceres, Cerro Verde, Leon 112 (USM). Near Tarapoto, Spruce 4035 (BR, GH, us). Huanuco: Prov. Huanuco, near confluence of Rio Ca- yumba with Huallaga, Mexia 8290 (F, GH, uc, us). Prov. Leoncio Prado, Dist. Rupa Rupa, 5 km S of Tingo Maria, Schunke V. 3252 (F, GH, us). Prov. Huanuco, Tingo Ma- ria, Tryon & Tryon 5238 (F, GH, us, USM). Pasco: Prov. Oxapampa, Valle del Palcazu, Rio Cacazu, Leon 682 (F). Ucayali: Prov. Coronel Portillo, Sinchono, near La Divi- soria, Aguilar 834 (GH). 23. Asplenium squamosum L., Sp. pi. 1082. 1753. TYPE: Pet. fil. \\2,t.5,f. 2, illustration based on a plant from Hispaniola. Asplenium jamesonii Hooker, Sp. fil. 3: 184. 1860. TYPE: Ecuador, Guayaquil, Jameson (holotype, K; isotype, BM; frag., NY!; photo, us of BM). Plants terrestrial. Stem stout, creeping or de- cumbent, provided with broad, flaccid, light brown scales, these 10-20 mm long, 3-6 mm broad, finely clathrate, often remotely fimbriate. Leaves sub- distant, 2-pinnate or 2-pinnate-pinnatifid, 40-100 cm long, 1 5-50 cm broad. Petiole to 40 cm long, dull, light or grayish brown. Lamina firm-herba- ceous to chartaceous, subdeltate, gradually re- duced to a pinnatifid apex, not reduced at base, most axes and veins sparsely to moderately pro- vided with minute, brown, appressed, 2-3-celled, articulate trichomes. Rachis light or gray-brown, glabrous, with a few, scattered, broad scales, non- alate, narrowly alate near apex, lacking proliferous buds. Pinnae 8-16 pairs, approximate, ascending, clearly stalked, deltate or oblong-deltate. Pinnules 2-9 cm long, ovate-lanceolate to subtrapeziform, acute or subacute, cuneate at base, crenate-dentate or deeply lobed. Veins several-forked, borne at acute angles. Sori borne in a single line along each side of costule. Indusia thin, often scarious, mar- gins entire to erose. In forests, thickets, or wooded ravines, in wet or rocky soil, 2100-3600(-3900) m, San Martin, Huanuco, Ayacucho, Apurimac, Cuzco. Hispaniola; Costa Rica; Colombia and Vene- zuela to Bolivia and Brazil. This species is distinguished by its huge leaves and the very large, nearly trapeziform, ultimate segments. With it probably should be included A. tucumanense Hieron. of Argentina. San Martin: Prov. Mariscal Caceres, Rio Abiseo, Young 4168 (USM). Huanuco: Mito, Bryan 379 (F). Muna, Bryan 627 (F). Tambo de Vaca, Macbride 4464 (F, us). Aya- cucho: Prov. La Mar, between El Tambo and Ayna, Plowman & Davis 4681 (GH, USM). Apurimac: Prov. Abancay, Sanchez 244 (USM). Cuzco: Prov. Paucartam- bo, Pillahuata, Aldave 5024 (HUT). Calca, Vargas 15645 (GH). 24. Asplenium cuspidatum Lam., Encycl. 2: 310. 1786. TYPE: Peru, Jussieu (holotype, P, Herb. Jussieu, photo, us; isotype, P, Herb. Lamarck; photo, us). Plants epiphytic or terrestrial, rarely epipetric. Stem stout, erect, provided with lanceolate to ovate, orange to gray-brown scales, these 2—4 mm long, 0.5-0.8 mm broad, acute, narrowly clathrate. Leaves caespitose, 2-pinnate (at least as to basal pinnae) to nearly 4-pinnate, 10-70 cm long, 2- 1 2(-l 5) cm broad. Petiole dull or sublustrous, green or gray to reddish brown or blackish. Lamina firm- herbaceous to chartaceous, lanceolate to ovate or subdeltate, tapered to a pinnatifid or serrate, often caudate, apex, not reduced at base, commonly gla- brous. Rachis glabrous, but with a few, minute, filiform scales, nonalate, or narrowly so distally, lacking proliferous buds. Pinnae 6 to many pairs, approximate to subdistant, most of them ascend- ing, short-stalked, ovate to linear-lanceolate. Pin- nules 0.5-2 cm long, lanceolate to ovate, acute or subacute, subentire to nearly 3 -pinnate, at least 26 FIELDIANA: BOTANY the basal ones of proximal pinnae obviously stalked. Veins forked to pinnately branched. Sori elliptic. Indusia relatively thick and fleshy, narrow, grayish or whitish, subentire. This and A. auritum are components of a highly varied species complex. Pending monographic study they are separated arbitrarily here as a mat- ter of convenience (see A. auritum for detailed discussion). In Central America and northern South America, A. cuspidatum is very common, but south of Ecuador it is encountered much less frequently. Two varieties are provisionally recognized in Peru, and var. tripinnatum is included in the following key, as it is to be expected here. The species occurs in the West Indies, from Mexico to Panama, and from Colombia and Ven- ezuela south to Bolivia and Brazil. a. Proximal pinnae 1 -pinnate (at least near their base), the pinnules deeply serrate or a few with basal acroscopic lobes 24a. var. cuspidatum a. Proximal pinnae 1-pinnate-pinnatifid to nearly 3-pinnate, most of the pinnules deeply pinnatifid or more b b. Proximal pinnae 1-pinnate-pinnatifid to nearly 2-pinnate, the pinnules deeply pinnatifid, some tertiary segments incised to the costule, but never freely stalked 24b. var. tripinnatum b. Proximal pinnae 2-pinnate to nearly 3-pinnate, tertiary segments commonly stalked 24c. var. foeniculaceum 24a. Asplenium cuspidatum var. cuspidatum. In forests, on wet slopes, rarely on mossy rocks or on tree trunks, 1 500-3 100 m, Cajamarca, Ama- zonas, La Libertad, San Martin, Cuzco. Mexico to Panama; Colombia and Venezuela to Bolivia and Brazil. Plants with less dissected pinnae merge with, and are often identified as, A. auritum. Cajamarca: Prov. Santa Cruz, Bosque de Montesco, Sagdstegui et al. 12382 (F, HUT, MO, uc). Amazonas: Prov. Chachapoyas, Cerros Calla Calla, 45 km above Balsas, Hutchison & Wright 5807 (GH, uc). La Libertad: Prov. Bolivar, between Unamen and Bolivar, Lopez & Sagdstegui 3331 (GH). San Martin: Prov. Mariscal Ca- ceres, Rio Abiseo National Park, Young & Leon 4945 (HUT), 4994 (HUT). Cuzco: Prov. Urubamba, Machu Pic- chu, above Rio Mandor, Peyton & Peyton 1286 (MO). Prov. La Convention, Huayopata, 10 km SW of Inca- tambo, Peyton & King 1428 (MO). 24b. Asplenium cuspidatum var. tripinnatum (Fourn.) Morton & Lell., Mem. New York Bot. Gard. 15: 29. 1966. Asplenium fragransSv/., Prodr. 180. 1788. TYPE: Ja- maica, Swartz (holotype, s; isotype, UPS; photo, us of s). Asplenium auritum Sw. var. tripinnatum Fourn., Mex- ic. pi. 1: 107. 1872. TYPE: Based onA.fragrans Sw. Not yet reported from Peru, but to be expected. Greater Antilles, except Puerto Rico; Mexico to Panama; Colombia and Venezuela to Bolivia and Brazil. 24c. Asplenium cuspidatum var. foeniculaceum (HBK.) Morton & Lell., Mem. New York Bot. Gard. 15: 29. 1966. Asplenium foeniculaceum HBK., Nov. gen. sp. 1:15. 1815. TYPE: "Novae Andalusiae," Humboldt & Bonpland (holotype, p; isotype, B, Herb. Willd. 19860; photos, us of p, F & GH of B). Asplenium abrotanoides Presl, Reliq. haenk. 1 : 47, /. 8,f. 2. 1825. TYPE: Peru, Huanuco, Haenke(ho- lotype, PRC!; frag., NY!). Asplenium cuspidatum var. abrotanoides ( Presl) Mor- ton & Lell., Mem. New York Bot. Gard. 15: 29. 1966. In wet forests and wooded canyons, in humus, on mossy logs, sometimes in rocky soil or on tree trunks, 2000-3950 m, Piura, Huanuco, Huanca- velica, Apurimac, Cuzco, Puno. Mexico to Panama; Colombia and Venezuela to Bolivia and Brazil. Of the two varieties reported from Peru, var. cuspidatum usually is found (with few exceptions) in the northwest and var. foeniculaceum in the central or (mostly) southern portions. With the absence, thus far, of the intermediate var. tripin- natum, the two others are easily separated, both geographically and morphologically. Piura: Prov. Ayabaca. near Ayabaca, Sagdstegui & Cabanidas 8700 (MO). Huanuco: Tambo de Vaca. Bryan 647 (F, GH). Macbride 4459 (F, us). Huancavelica: Prov. Tayacaja, Ampurco, between Salcabamba and Surcu- bamba, Tovar 3767 (GH). Apurimac: Prov. Abancay, for- ests of Ampay, Vargas 1066 (GH, us). Cuzco: Prov. Pau- cartambo. Dist. Marcachea, near Achirani. I'argas 1 1 140 TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 27 (F, GH, uc, us). Puno: Prov. Carabaya, between Ayapata and Kahualluyoc, Vargas 10770 (GH). 25. Asplenium repandulum Kunze, Linnaea 9: 65. 1834. TYPE: Peru, Huanuco, "Pampayaco" [Pampayacu], Poeppig'm July 1 829 (holotype, LZ, destroyed; frag., NY!). Plants epiphytic, at least in Peru. Stem short- to long-creeping, scales lacking or rare and small (less than 1 mm long). Leaves 1 -pinnate, subdis- tant to remote, 20-40 cm long, 6-12 cm broad. Petiole 3-1 5 cm long, fleshy, dull reddish or green- ish brown, naked or with a few small scales at base. Lamina thin-herbaceous or membranaceous, oblong-lanceolate to ovate, not or only slightly reduced at base, narrowing abruptly to a subcon- form apical segment, this often with a broad, basal lobe. Rachis scarcely to broadly alate, lacking pro- liferous buds. Pinnae 10-20 pairs, approximate to subdistant, sessile or short-stalked, apex acute to attenuate (rarely obtuse), strongly inequilateral at base, narrowly cuneate or excavate basiscopically, truncate and often auriculate acroscopically, mar- gins crenate to serrate. Veins 1-2-forked, ascend- ing from the costa at 30-40° angles. Sori mostly medial. Indusia linear, opaque, greenish or brown, subentire. In forests, on tree trunks, 1250-2000 m, Ama- zonas to Junin, Madre de Dios. Southern Mexico to Panama; Venezuela and Colombia to Brazil and Bolivia. With this perhaps should be included A. obtu- sifolium L. of northern South America and the West Indies, which supposedly differs from A., repandulum in having fewer pinnae with obtuse apices and a basal lobe incised nearly to the costa. Although A. repandulum is often epipetric or ter- restrial in Mexico and Central America, most col- lectors report it as epiphytic in Peru and Ecuador. Amazonas: East of La Peca in Serrania de Bagua, Gen- try et al. 23073 (MO, uc). Huanuco: Fundo Chela, Rio "Chin" (Chinchao?), Aguilar 911 (USM). Prov. Leoncio Prado, Dist. Hermilio Vladizan, La Divisoria, Plowman & Schunke 7363 (F, GH) Pasco: Prov. Oxapampa, Gran Pajonal, D. Smith 5168 (F, MO). Junin: Prov. Tarma, Chanchamayo Valley, above La Merced, Hutchison 1 196 (F, GH, uc, us). Chanchamayo Valley, C. Schunke 155 (F, us). Madre de Dios: Cocha Cashu, between Panagua and Tayakome, Foster et al. 3449 (F). 26. Asplenium vomeriforme Hooker, Sp. fil. 3: 109, /. 762. 1860. TYPE: Peru, Mathews in 1835 (K!). (Mathews 1851 was cited by Hooker in the protologue.) Asplenium mathewsii Moore. Index fil. 145. 1857. Nom. nud. Plants epiphytic or terrestrial. Stem decumbent, or rarely very short-creeping, provided with gold- en-tomentose roots and gray-brown scales, these narrowly clathrate, linear-lanceolate, attenuate, 4— 7 mm long, often ciliate toward base. Leaves crowded, long-petiolate, usually pendent, 1 -pinnate, to 90 cm long and 25 cm broad. Petiole 1 5-40 cm long, dull, dark brown to yellow-brown, glabrous. Lamina carnose or chartaceous, abruptly terminating in a subconform apical segment, or this often with a basal lobe or auricle, not or scarce- ly reduced at base, rachis nonalate. Pinnae 6-14 pairs, larger ones 9-13 cm long. 2.5-4 cm broad, patent, short-stalked, most of them narrowly to broadly deltate, the margins entire or inconspic- uously crenulate, conspicuously inequilateral at base, acroscopically truncate to cordate or auric- ulate, basiscopically rounded to broadly cuneate, but proximal ones mostly truncate. Veins once or twice dichotomously forked, indistinct or obscure. Indusia firm, entire, inrolled at maturity. In primary and secondary forests, pendent from tree trunks and branches, or on the forest floor, 1200-1800 m, Pasco and Junin. Endemic. This is very closely related to A. salicifolium and may be merely a very robust, local variant. Prin- cipal differences are size, shape and margins of pinnae. Pasco: Prov. Oxapampa, 2 km from Oxapampa, Leon et al. 500 (F, uc, us, USM). Prov. Oxapampa, in coffee plantation near Villa Rica, van der Werffet al. 8278 (F, MO, uc). Junin: Chanchamayo Valley, C. Schunke 695 (F), 696 (F, us). Prov. Satipo, Gran Pajonal, south of Chequitavo, D. Smith 5151 (F, MO, uc). 27. Asplenium juglandifolium Lam., Encycl. 2: 307. 1786. TYPE: "Antilles" (probably Puer- to Rico or Hispaniola), collector undesignated (holotype, P, Herb. Jussieu 128 3 A & B; pho- tos, F & us of 1283 A). Asplenium integerrimum Sprengel, Nova Acta Phys.- Med. Acad. Caes. Leop.-Carol. Nat. Cur. 10: 231. 1821. TYPE: Puerto Rico, Bertero (holotype, B; isotypes, L, P; photos, us of B, F, & us of P). Asplenium falcinellum Maxon, Contr. U.S. Natl. Herb. 28 FIELDIANA: BOTANY 13: 14. 1909. TYPE: Guatemala, Cubilguitz, Tuerckheim 1910 (holotype, us!). Plants epiphytic (rarely terrestrial outside Peru). Stem decumbent (rarely very short-creeping out- side Peru), provided with medium or dark brown scales, these narrowly clathrate, mostly filiform, 3-10 mm long, 0. 1-0.3(-0.4) mm broad. Leaves crowded, long-petiolate, 1 -pinnate (juvenile ones rarely simple), to 80 cm long and 25 cm broad. Petiole 10-30 cm long, dull reddish or greenish brown, with a few linear scales at very base. Lam- ina firm-herbaceous to chartaceous, abruptly ter- minating in a conform apical segment, not reduced at base, scarcely or moderately provided abaxially with minute (0.1-0.2 mm), appressed, trichomes, these often gland-tipped, rachis very narrowly alate (if at all) near the apex. Pinnae (0-)2-12 pairs, larger ones 7-20 cm long, 1.2-2.6(-4) cm broad, ascending, short-stalked, apices commonly atten- uate, subequilateral at the cunate base, margins entire. Veins 1-2-forked, indistinct. Indusia linear, firm and rather thick-textured, entire. On tree trunks in wet forests, 100-1500 m, Amazonas and Loreto to Cuzco. Southern Mexico to Panama; Greater Antilles; Trinidad; Colombia to Surinam, south to Bolivia and Brazil. This species is quite variable in number, size, and shape of pinnae and in length of stem scales, and these differences have encouraged earlier sep- aration into several species and varieties. At one extreme is A. integerrimum of the Greater Antilles, with 2-5 pairs of pinnae well over 2 cm broad, their bases quite inequilateral, and truncate to broadly rounded. Asplenium falcinellum of Cen- tral America typically has 6-12 pairs of pinnae 1.2-1.8 cm broad, their bases more narrowly and more equally cuneate. Pinnae in the type specimen of A. juglandifolium more closely resemble those of the former species. Length of stem scales fre- quently corresponds with pinna characters in that those of A. falcinellum tend to be less than 5 mm long and those of A. juglandifolium and A. inte- gerrimum 6-10 mm long. However, throughout South America, especially in Ecuador and Peru, a number of specimens with numerous and narrow- er pinnae have much longer scales, and in other specimens the converse is true, which renders the separation of these taxa untenable when examined throughout the entire range of distribution. Also included under A. juglandifolium should be A. kapplerianum Kunze of Surinam and probably A. neogranatense Fee of Colombia. Amazonas: Prov. Bagua, Dist. Cenepa, Tillett 672-67 (GH). San Martin: Zepelacio, near Moyobamba, Klug 3430 (F, GH, MO, us). Prov. Mariscal Caceres, Tocache Nuevo, Schunke V. 6948 (F, GH, MO). Loreto: Rio Ma- mon near Rio Nanay, Croat 19950 (GH, MO, uc). Rio Mazan, Salinas, J. Schunke 375 (F, GH, uc, us). Huanuco: Above Rio Huallaga at Tingo Maria, Croat 21033 (MO, us). Pasco (as Junin): Puerto Bermudez, Killip & Smith 26679 (us); Prov. Oxapampa, Cordillera San Matias, Leon 322 (F, USM). Cuzco: Prov. Paucartambo, Callanga, Woytkowski 392 (GH). 28. Asplenium tuerckheimii Maxon, Contr. U.S. Natl. Herb. 13: 15. 1909. TYPE: Guatemala, Baja Verapaz, Panzal, Tuerckheim 11-1677 (us!). Plants terrestrial. Stem erect or decumbent, pro- vided at apex with blackish brown, clathrate scales, these ovate, 1-2.5 mm long, often somewhat vaulted. Leaves crowded or fasciculate, to 50 cm long and 1 5 cm broad, long-petiolate, 1 -pinnate. Petiole to 25 cm long, dull grayish to reddish brown, glabrous. Lamina firm-herbaceous, abruptly ter- minating in a conform apical segment (this seg- ment sometimes obsolete), not reduced at base, glabrous except for scattered, minute (0. 1-0.2 mm), appressed, sometimes glandular trichomes, rachis narrowly alate. Pinnae typically 3-6 pairs, to 10 cm long and 1 .8 cm broad, alternate, at least prox- imal ones stalked, not or rarely adnate, narrowly and subequally cuneate at base, acute to attenuate at apex, margins broadly crenate-serrate. Veins 1- 2-forked, not distinct. Indusia thin, yellowish brown, entire. On the floor of deep forests, 600-1 500 m, Junin and Cuzco. Southern Mexico; Guatemala; Peru. The few collections made in Peru constitute a wide disjunction in range for this species, here- tofore known only from southern Mexico and Guatemala. It is sometimes confused with A. ab- scissum, or with specimens of A. juglandifolium with narrow pinnae. The latter species has pinna bases somewhat to strongly inequilateral, and long, filiform stem scales, whereas pinnae in A. tuerck- heimii are equilaterally and narrowly cuneate, and stem scales are minute, ovate, and often somewhat vaulted. Asplenium abscissum is much more close- ly related, for it has identical stem scales. The latter is usually distinguished by its pinnatifid or non- conform apical segments and by strongly inequi- lateral pinna bases. Junin: Chanchamayo Valley, C. Schunke 88 (F, us), TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 29 1023 (F). Cuzco: Prov. La Convencion, Rio Apurimac, below Puerto Capiro, Davis et al. 1311 (F, GH, uc). 29. Asplenium tricholepis Rosenst., Repert. Spec. Nov. Regni Veg. 12: 468. 1913. TYPE: Bo- livia, Yungas, Polo-Polo, near Coroico, Buch- tien 3330 (holotype, s!; isotype, us!). Asplenium rusbyanum Domin, Pterid. Dominica 171, /. 29, f. 2. 1929. TYPE: Bolivia, Yungas, Rusby 383 (holotype, presumably K; iso types, F!, GH!, us!). Plants epiphytic. Stem stout, erect or decum- bent, provided with orange to reddish brown, clathrate, scales, these lanceolate to ovate, flat, 5- 15 mm long, 1-2 mm broad, margins (especially at base) often bearing long, tortuous cilia. Leaves few, crowded, to 60 cm long and 20 cm broad, long-petiolate, 1 -pinnate. Petiole to 30 cm long, dull, gray-green to purplish brown, glabrous. Lam- ina firm-herbaceous to subcarnose, abruptly ter- minating in a conform apical segment, not reduced at base, glabrous, rachis not or scarcely alate. Pin- nae (l-)2-5 pairs, to 25 cm long and 4 cm broad, alternate, sessile or subsessile, not or rarely adnate, narrowly and subequilaterally cuneate at base, api- ces obtuse or acute, occasionally attenuate, mar- gins entire. Veins l-2(-3)-forked, distinct or in- distinct. Indusia thick, inrolled at maturity, entire. On tree trunks in dense, wet forests, 800-2 1 00 m, Amazonas to Cuzco. Ecuador; Peru; Bolivia. The long-cilate stem scales may be helpful in distinguishing this species, but they do occur, per- haps less frequently, in A. davisii, A. juglandifoli- um, and A. salicifolium and its relatives. However the cilia often go unobserved, either because they are readily broken off or are actually lacking. Asplenium tricholepis has been confused with A. oligophyllum Kaulf. (syn. A. escragnollei Fee) of Venezuela and Brazil, which is very similar; how- ever, the latter has crenulate pinna margins, and blackish, filiform scales along the costae abaxially. Amazonas: Prov. Bagua, E of La Peca, Barbour 2522 (MO), 2576A (MO). Huanuco: Prov. Huanuco, Tulumayo, near Tingo Maria, Ferreyra 2151 (GH). Pasco (as Junin): Pichis Trail, Yapas, Killip & Smith 25560A (us). Junin: Schunke Hacienda above San Ramon, Killip & Smith 24837 (F, us). Cuzco: Prov. La Convencion, NE of Ha- cienda Luisiana, Dudley 10398 (GH). Prov. La Conven- cion, Tinkuri, Vargas 5453 (uc). 30. Asplenium davisii Stolze, sp. nov. Plantae terrestres. Caulis erecta vel decumbens, pa- leis brunneolis, clathratis, ovato-lanceolatis, 5- 10 mm longis, 1.5 mm latis. Folium 50-100 cm longum, 16-28 cm latum. Petiolus 25-60 cm lon- gus, obscurus, flavidus vel purpurascens. Lamina 1-pinnata, segmento apicali conformi, in pagina abaxiali trichomatibus minutis, adpressis, brun- neis, glandulosibus. Pinnae 3-7 jugae, 14-24 cm longae, 3-4 cm latae, lanceolatae, attenuatae, ad basin cuneatae et subequilaterale, marginibus crenato-serratis. Plants terrestrial. Stem stout, erect or decum- bent, provided with brown or gray-brown, clath- rate scales, these ovate-lanceolate, flat, 5-10 mm long, and about 1.5 mm broad, the margins some- times ciliate. Leaves few, crowded, 50-100 cm long, 16-28 cm broad, long-petiolate, 1 -pinnate. Petiole 25-60 cm long, dull, yellowish to purplish brown, glabrous. Lamina chartaceous, abruptly terminat- ing in a conform apical segment, not reduced at base, amply to moderately provided abaxially with minute (0. 1-0.2 mm) appressed, dark brown, plu- ricellular, often gland-tipped trichomes, rachis nonalate. Pinnae (2-)3-7 pairs, larger ones 14-24 cm long and 3-4 cm broad, alternate, stalked, lan- ceolate, attenuate, broadly to narrowly cuneate and subequilateral at base, margins crenulate to cre- nate-serrate. Veins 1-2-forked, usually rather dis- tinct. Indusia linear, 1.5-2.5 cm long, greenish or yellowish, thin, often inrolled at maturity. TYPE— Peru, Cuzco, Prov. La Convencion, Rio Apurimac, above Boca del Tigre rapids, Davis et al. 1292 (holotype, F!; isotypes, GH!, uc!). Endemic. Terrestrial in wet forests, 100-1600 m, Amazonas and Loreto south to Cuzco and Ma- dre de Dios. This species has been most frequently identified as A. oligophyllum Kaulf. (syn. A. escragnollei Fee) of northern South America and Brazil, which dif- fers most significantly in the blackish, filiform, clathrate scales along the costa. Presence of these scales indicates a relationship with the simple- leaved A. serratum and A. angustum. Asplenium oligophyllum is further distinguished from A. da- visii by the absence of minute, appressed tri- chomes on the abaxial laminar surface. Asplenium davisii is probably more closely re- lated to A. canelense Rosenst., a rare species from Colombia and Ecuador (type: Ecuador, Spruce, Herb. Bonaparte no. 10010, P!). That species dif- fers by its much more numerous (12-14 pairs) pinnae and the absence of glandular trichomes. Another similar species from Ecuador is A. virens Presl (syn. A.flavidum Sodiro), which differs from 30 FIELDIANA: BOTANY A. davisii by its smaller (less than 40 cm) leaves, its much smaller (3-5 mm) stem scales, more deeply serrate, merely obtuse to acute, pinnae, and the absence of laminar trichomes. Amazonas: Prov. Bagua, Cerro Tapur on Rio Utcu- bamba, Hutchison 1485 (F, uc, us). Loreto: Puerto Ar- turo, lower Rio Huallaga below Yurimaguas, Killip & Smith 27773 (us). Huanuco: Prov. Leoncio Prado, Dist. Hermilio Valdizan, La Divisoria Plowman & Schunke 7377 (F). Junin: Chanchamayo Valley, C. Schunke 5 (F, us), 75 (F, us), 452 (F, us). Cuzco: Prov. Paucartambo, Kosnipata-Pilcopata, Vargas 11290 (GH). Madre de Dios: Prov. Mann. Atalaya, Foster & Wachter 7462 (F, MO). 3 1 . Asplenium laetum Sw., Syn. fil. 79: 271.1 806. TYPE: "Habitat in India occidentale," Swartz (holotype, s). Asplenium virens Desv., Mem. Soc. Linn. Paris 6: 273. 1827, not Presl, 1825. TYPE: "Habitat in Amer- ica," collector undesignated (holotype, p, Herb. Desv.; photo, us). with petioles quite crowded, so on some specimens it may appear to be decumbent. Besides the dif- ferences cited in the key, it differs from A. deli- tescens and A. melanopus in another subtle char- acter. The typical lamina in A. laetum becomes very gradually pinnatifid to a nearly caudate apex; i.e., each pair of distal segments becomes slightly shorter than the adjacent pair, the lamina finally terminating in a subcaudate tip. In the other two species, the distal segments are rather abruptly re- duced, the lamina commonly terminating in a nonconform apical segment. Amazonas: Bagua, 1 2 km S of La Peca, Barbour 2500 (F). San Martin: Juan Jui, Alto Rio Huallaga, Klug 4253 (F, GH, uc, us). Huanuco: Prov. Leoncio Prado, Dist. Rupa Rupa, 5 km S of Tingo Maria, J. Schunke V. 3261 (F, GH). Prov. Huanuco, Tingo Maria, Tryon & Tryon 5228 (F, GH, us, USM). Pasco: Prov. Oxapampa, Rio El Tunqui, D. Smith et al. 1710 (F, MO). Junin: La Merced, Chanchamayo, C. Schunke 20 (A). Cuzco: Prov. La Con- vention, below Rosalina, Vargas 12293 (GH). Plants terrestrial. Stem short-creeping (some- times not obviously so), provided with linear scales, these clathrate, 1-3 mm long, dark brown or black- ish. Leaves 1 -pinnate, crowded, to 40 cm long, 4- 8 cm broad. Petiole 6-18 cm long, dull grayish or greenish to lustrous and reddish brown or atro- purpureous, not alate, sparsely to amply scaly. Lamina thin-herbaceous, oblong-lanceolate, grad- ually diminishing to a pinnatifid, often subcau- date, apex, not or scarcely reduced at base. Rachis essentially glabrous, scarcely alate distally, lacking proliferous buds. Pinnae 1 5-24 pairs, larger ones 2.5-4.5 cm long, subsessile or short-stalked, most of them strongly inequilateral for nearly half their length, truncate acroscopically, narrow-cuneate or excavate basiscopically, obtuse to acute at apex, margins bicrenate or biserrate acroscopically, sub- entire basiscopically. Veins 1-2-forked. Sori spreading at 30—40° from costa, not or rarely di- plazioid. Indusia thin, pale green or yellowish, lin- ear, subentire. On floor of forests, on rocky slopes, or on mossy rocks or in soil pockets of rocky cliffs, 200-900 (-1600) m, Amazonas south to Cuzco. Southern Mexico to Panama; West Indies; Co- lombia to Surinam, south to Argentina and Par- aguay. This is one of a few species of Asplenium with the petiole varying from light to dark in color and from dull to lustrous, the variation observable on the same plant. The stem is very short-creeping, 32. Asplenium melanopus Sodiro, Crypt, vase. Quit. 189. 1893. TYPE: Ecuador, Chimbo- razo, Pallatanga Valley near Chimbo Bridge, Sodiro (holotype, QCA?). Diplazium melanopus (Sodiro) Hieron., Bot. Jahrb. Syst. 34: 456. 1904. Plants terrestrial or epipetric. Stem short-creep- ing, provided with linear to lanceolate scales, these clathrate, 0.5-1.5 mm long, dark brown. Leaves 1 -pinnate, approximate to crowded, 28-64 cm long, 10-1 8 cm broad. Petiole 1 2-32 cm long, dark red- dish brown to purplish, dull, or sometimes lus- trous, nonalate, sparsely scaly as on the stem. Lam- ina firm-herbaceous to membranaceous, deltate or ovate-deltate, rather abruptly reduced to a pin- natifid or nonconform apical segment, not or scarcely reduced at base. Rachis essentially gla- brous, not or scarcely alate, lacking proliferous buds. Pinnae 7-1 1 pairs, larger ones 6-12 cm long and 1.4-4 cm broad, short-stalked, equilateral ex- cept at base, truncate acroscopically, abruptly cu- neate or excavate basiscopically, acute to attenuate at apex, both margins lobed to pinnatifid, the tip of each lobe or segment with a few rounded teeth. Veins distinct, several times forked. Sori borne on the vein branches, often back-to-back (diplazioid). Indusia thin, yellow to reddish brown, linear, sub- entire. In forests, in soil or among rocks, sometimes on TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 31 rocky outcrops, 250-1700 m, Amazonas, Loreto, Junin, Madre de Dios. Colombia; Ecuador; Peru. This and A. delitescens have been included ear- lier in Diplaziwn because of the frequent occur- rence of back-to-back sori. However, each has the principal diagnostic characters ofAsplenium: ob- viously clathrate scales, single-rowed cells in spo- rangia stalks, annular cells of sporangia 20 or more, spores with folded perispore. The two species dif- fer from each other chiefly in the degree of pinna lobing and perhaps should be separated merely at an infraspecific level. Asplenium purpurascens Mett. of Ecuador, and A. melanopus may also be synonymous; however, the type of the former has not been examined dur- ing this study. The name of A. purpurascens would have priority. Amazonas: Prov. Bagua, E of La Peca, Barbour 2500 (F, a depauperate specimen). Loreto: Right bank of Rio Santiago, above Pongo de Manseriche (Amazonas), Mexia 6223 (GH, uc, us). Junin: Colonia Perene, Killip & Smith 24924 (F, us). Madre de Dios: Prov. Tambopata, Al- bergue Cuzco Amazonico, Leon 866 (USM). 33. Asplenium delitescens (Maxon) Gomez, Bre- nesia 8: 52. 1976. Diplazium delitescens Maxon, contr. U.S. Natl. Herb. 10: 497. 1908. TYPE: Cuba, Oriente, vicinity of San Luis, Pollard & Palmer 348 (holotype, us!). Plants terrestrial, occasionally epipetric or epi- phytic. Stem short-creeping, provided with linear or lanceolate, attenuate scales, these clathrate, 1- 2 mm long, dark brown to blackish. Leaves 1 -pinnate, approximate to crowded, 30-54 cm long, 1 2-24 cm broad. Petiole 1 4-25 cm long, dark red- dish brown to yellowish brown or greenish, dull to sublustrous, nonalate, sparsely scaly. Lamina thin- to firm-herbaceous, deltate or oblong-del- tate, nonalate, sparsely scaly. Lamina thin- to firm- herbaceous, deltate or oblong-deltate, gradually or abruptly reduced to a nonconform, sometimes at- tenuate, apical segment, not or scarcely reduced at base. Rachis glabrous, or with a few filiform scales, marginate or inconspicuously alate distally, lacking proliferous buds. Pinnae 6-10 pairs, larger ones 6-14 cm long and 1.5-2 cm broad, sessile, or proximal ones short-stalked, equilateral except at base, there truncate (and sometimes auriculate) acroscopically, narrowly cuneate to excavate basi- scopically, acute to attenuate at apex, margins ser- rate to deeply crenulate acroscopically, serrate or subentire basiscopically. Veins distinct or indis- tinct, 1-2-forked. Sori (some of them) diplazioid. Indusia yellow to light brown, linear, subentire. In forests, on tree trunks, on clay banks, or on the forest floor, occasionally on rocky cliffs, 135- 750 m, Loreto, Huanuco, Madre de Dios. Southern Mexico to Panama; Cuba; Colombia; Ecuador; Peru. This is probably only a less-dissected form of A. melanopus, under which see further discussion. Loreto: Yurimaguas, lower Rio Huallaga, Killip & Smith 29078 (us). Huanuco: Tingo Maria, Moran 3676 (MO, uc, USM). Prov. Pachitea, Dist. Puerto Inca, Bosque Na- tional de Iparia, /. Schunke V. 2967 (F, GH, us). Madre de Dios: Prov. Manu, Parque Nacional Manu, Pakitsa Station, Foster & Baldeon 12806 (F). 34. Asplenium achilleifolium (Mart. & Gal.) Liebm., Kongel. Danske Vidensk. Selsk. Skr., Naturv. Afd. V, 1: 249 (seors. 97). 1849, not (Lam.)C. Chr. 1905. Caenopteris achilleifolia Mart. & Gal., Nouv. Mem. Acad. Roy. Sci. Bruxelles 15: 63. 1842. SYN- TYPES: Mexico, Veracruz, vicinity of Mirador, Galeotti 6279 (BR), 6295 (BR). Athyrium achilleifolium (Mart. & Gal.) Fee (Mem. foug. 5) Gen. fil.: 186. 1852. Plants terrestrial. Stem erect or decumbent, am- ply provided with obscurely clathrate, ovate or lanceolate scales, these to 5 mm long and 3 mm broad, obtuse or acute, light to dark brown. Leaves 1 -pinnate-pinnatisect or nearly 2-pinnate, caes- pitose, 0.3-1.0 m long, 9-30 cm broad. Petiole somewhat shorter than the lamina, dull greenish or grayish brown, sparsely scaly, basal scales broad, the rest filiform. Lamina firm-herbaceous, ovate or lanceolate, tapering to an attenuate, pinnatifid apex, not or slightly reduced at base. Rachis broad- ly alate, lacking proliferous buds, sparsely filiform- scaly. Pinnae numerous, 5-15 cm long, crowded to subdistant, subsessile, truncate and narrowly decurrent at base, cut nearly to costa into 7-16 pairs of segments, these subentire to crenate or serrate. Veins 1 -several-forked, diverging from the midrib at a wide angle. Indusia thin, elliptic, yel- lowish to whitish. Thus far known in Peru from one collection, in wet forest, 2000 m, Amazonas. Southern Mexico to Nicaragua; Ecuador?; Peru. The collection cited below apparently is a de- pauperate form of A. achilleifolium. It matches 32 FIELDIANA: BOTANY typical plants in Central America except that the leaves and pinnae are significantly shorter. Stolze ( 1 986, p. 76) cited a single collection from Ecuador as another possible variant of this species. In any case, this represents a strikingly disjunct distri- bution. Amazonas: Prov. Bagua, 20 km E of La Peca, Harbour 2759 (F, MO, uc). 35. Asplenium projectum Kunze, Linnaea 9: 68. 1835. TYPE: Peru, "Pampayaco, in rupibus calidioribus" (Dept. Huanuco, Pampayacu), Poeppig (diar. 1156), Jul. 1829 (holotype, B!; isotypes, B!, p; photos, GH & uc of P). Plants epipetric. Stem very small, erect, provid- ed with linear-lanceolate scales, these 0.5-1 mm long, clathrate, blackish, sublustrous. Leaves fas- ciculate, to 7 cm long and 0.7 cm broad. Petiole 1-3 mm long, dull greenish or grayish brown, rare- ly blackish, marginate or narrowly alate. Lamina thin-herbaceous, linear, gradually tapering to a pinnatifid or flagelliform apex, somewhat reduced at base. Rachis dark brown or atropurpureous, moderately provided with minute, glandular tri- chomes, narrow-alate, often bearing a proliferous bud at apex or in a pinna axil. Pinnae 12-18 pairs, 2-3 mm long, subopposite, with short, blackish stalks, rotund to obovate, or shallowly and broadly bi- or trilobate, costa lacking. Veins distinct, black, simple or 2-3-forked in each pinna. Sori 1-2 on a pinna. Indusia thin, yellow-green, oblong-ellip- tic. In rock crevices, and among rocks, about 2500 m, Amazonas, Huanuco. Endemic. This rare species is one of the smallest of the genus in Peru and is closely related to A. peru- vianum, A. gilliesii, and A. triphyllum. The tiny, roundish pinnae and the black color of the stalks that continues onto the veins set it apart from its affiliates. The Lopez collection is the first for over a century, but this is not surprising, as the fern is so tiny and inconspicuous. The Poeppig collection cited below is incorrectly labeled as a type at Vienna. Amazonas: Prov. Chachapoyas, Pomacocha (Leime- hamha-Balsas). Lopez et al. 4398 (GH). Huanuco: "Cas- sapi" (Dist. Chinchao, Casapi), Poeppig, Oct. 1 829 (BM, w). 36. Asplenium lorentzii Hieron., Hot. Jahrb. Syst. 22: 375. 1896. LECTOTYPE (designated here): Argentina, Prov. Salta, Yacone, at foot of Nevada del Castillo, Hieronymus & Lo- rentz 304 (B!, photo, F, isolectotype, B!). SYN- TYPE: Argentina, Siambon, Sierra de Tucu- man, Hieronymus & Lorentz 896 (B!). Plants terrestrial. Stem erect or decumbent, pro- vided with linear, attenuate scales, these 1-3 mm long, clathrate, gray-brown to blackish. Leaves fas- ciculate, 1 -pinnate, 16-30 cm long, 1.7-2.7 cm broad. Petiole 4-10 cm long, 0.8-1 m thick, red- dish- or greenish brown, not or narrowly alate, glabrous, or with glandular trichomes. Lamina membranceous, linear, gradually tapering to a pin- natifid apex, somewhat reduced at base, glabrous, or with scattered, minute, glandular trichomes. Rachis straight or slightly flexuous, glabrous or with glandular trichomes, narrow-alate, prolifer- ous buds rare or lacking. Pinnae numerous, 10— 14 mm long, subopposite or alternate, subsessile, subdimidiate, most of them rhombic, deeply den- tate, patent, or proximal ones deflexed, lacking a distinct midrib. Veins distinct, slightly darker than the laminar tissue, 1-2 on basiscopic side of pinna, 4-6 on acroscopic side, 1-2-forked, the basal ac- roscopic ones subflabellately dichotomous. Sori 4- 6 on larger pinnae. Indusia thin, yellowish or greenish, oblong-elliptic. Thus far known in Peru by the lone collection cited below, on the ground, 700 m, Arequipa. Peru; Bolivia; Argentina. The sole collection from Peru differs slightly from the type in that pinnae are patent and moderately provided with minute, glandular trichomes on the abaxial surface. In the lectotypes and paratypes from Argentina laminae are glabrous, and pinnae are frequently deflexed. If these differences are substantiated by subsequent collections in Peru, it may warrant separation at an infraspecific level. This species is a low-elevation plant related to A. peruvianum and A. gilliesii. It is a more robust fern with stouter petiole, longer leaves, larger pin- nae, and more numerous sori. Arequipa: Lomas of Atequipa, Coronado 34 (GH, uc, us). 37. Asplenium gilliesii Hooker, Exot. fl. 3: 208 (overleaf). 1827. TYPE: Argentina, Cerro Grande Uspallata, Gillies, Jan. 31, 1825 (ho- lotype, K!; frag., GH!; ISOTYPE, GH!). TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 33 Asplenium gillesianum Hooker, Icon. fil. 1 : t. 73. 1 829, nom. nov. (an illegitimate renaming of A. gilliesii, and with the same type). Plants epipetric, rarely terrestrial. Stem small, erect, provided with linear, attenuate scales, these 1.5-3 mm long, narrow-clathrate, blackish, sub- lustrous. Leaves fasciculate, 1 -pinnate, 8-20 cm long, (0.8-)1-1.5 cm broad. Petiole 2-8 cm long, 0.2-0.4 mm thick, stramineous or greenish, mar- ginate or very narrowly alate. Lamina thin-her- baceous, linear, gradually tapering to a pinnatifid apex, somewhat reduced at base, provided abax- ially with minute, bacilliform glands. Rachis thin and flexuous, greenish, sparsely provided with minute, glandular trichomes, scarcely or narrowly alate, proliferous buds rare or lacking. Pinnae 7- 20 pairs, 4-8 mm long, subopposite or alternate, sessile or with short, greenish stalks, rhombic or subflabellate, lacking a costa, veins ending in acute, broad and deep, cuspidate lobes. Veins commonly distinct, slightly darker than the lamina tissue, subflabellately dichotomous. Sori 2-3 on a pinna. Indusia thin, yellowish or grayish green, oblong- elliptic. In rock crevices, or among rocks in dry mead- ows, 2750-4000 m, Ancash and Huanuco south to Ayacucho and Puno. Peru; Bolivia; Argentina. Very closely related are A. peruvianum and A. projectum, all three high-elevation ferns with lin- ear leaves and blackish, linear scales. See A. pe- ruvianum for further discussion. Ancash: Prov. Bolognesi, E of Huasta, Cerrata 2173, 2473 (GH, USM). Huanuco: Yanahuanca, Macbride & Featherstone 999 (F). Lima: Prov. Yauyos, Capia, below Tupe, Cerrate 1069 (GH, USM). Ayacucho: Prov. Pari- nacocha, Incuyo, Hutchison 1270 (F, NY, uc). Cuzco: Prov. Urubamba, road to Ollanta, Vargas 23280 (F, GH). Puno: Above Lake Titicaca, Mexia 7782 (F, GH, MO, uc, us). 38. Asplenium peruvianum Desv., Mem. Soc. Linn. Paris 6: 271. 1827. TYPE: Peru, col- lector undesignated, perhaps Dombey (holo- type, P!, Herb. Desv.; photos, GH, us). Asplenium fragilePres\,Tenl.pterid. 108. 1836. TYPE: "Montanis Peruviae," Haenke (holotype, PR!; frag., NY!; isotype, PRO!; photos, F, GH & us of PR, F of PRC). Plants epipetric, rarely terrestrial. Stem erect, provided with lanceolate or linear-lanceolate scales, these 1-1.5 mm long, clathrate, blackish, sublus- trous. Leaves densely caespitose, 1 -pinnate, 6-30 cm long, 0.6-1.5(-2) cm broad. Petiole 1-10 cm long, commonly 0.5-1 mm thick, dull grayish to reddish brown, scarcely alate, provided with a few filiform scales and minute, clavate trichomes, or glabrescent. Lamina firm-herbaceous or charta- ceous, linear, gradually tapering to a pinnatifid apex, somewhat to strongly reduced at base. Ra- chis typically straight, rigid, and reddish brown, occasionally thin, flexuous and greenish, with in- dument as on the petiole, narrow-alate to margin- ate, sometimes with a proliferous bud in a prox- imal pinna axil. Pinnae 1 2-30 pairs, 4-8(-10) mm long, subopposite, sessile or subsessile, rhombic, a distinct costa lacking (or rudimentary at pinna base), margins entire or shallowly to deeply den- tate, rarely lobed. Veins distinct to obscure, subfla- bellately dichotomous. Sori 4-6 on a pinna. In- dusia yellowish or greenish, narrow-elliptic. In open forests or thickets, in rock crevices, on and among rocks, or in rocky soil, 2750-4500 m, Cajamarca and La Libertad south to Ayacucho and Puno. Venezuela and Colombia south to Chile. The type specimen of Asplenium peruvianum consists merely of a single leaf, minus the stem, so the valuable character of stem scales is lacking. This matches the holotype of A. fragile, which also lacks stem scales, and the isotype (PRC), which has both stem and scales. The holotype label of A. peruvianum cites no data as to collector, number, or specific locality. There is another specimen at Paris, collected by Dombey at Tarma (Junin) also originally determined to be A. peruvianum. The label of a Morton photo (us) states that ". . . it may be the original plant from which the fragment which is the type was taken." This specimen is complete with a dozen densely caespitose leaves and small, blackish, clathrate, stem scales and is identical to the isotype of A. fragile in every way. It is unfortunate that this familiar name for a com- mon South American species of Asplenium must be subsumed under A. peruvianum. This is part of a species complex that includes A. gilliesii and A. projectum as well as A. tricho- manes-dentatum L. of Central America and the West Indies. They are all small ferns with linear, 1 -pinnate leaves, blackish, linear stem scales, and laminae with scattered filiform scales and glan- dular trichomes. Asplenium gilliesii and A. peru- vianum may be simply variants (witnessed by the intermediates cited separately below), and A. 34 FIELDIANA: BOTANY trichomanes-dentatum is a low-elevation form with smaller sterile leaves in a rosette pattern. (The name A. trichomanes-dentatum was later changed by Linnaeus to A. dentatum, and some authors still disagree on the application of the rules toward determination of the valid name. For further dis- cussion of the nomenclature problem, see Stolze [Fieldiana, Bot., n.s., 6: 92. 1981].) Monographic revision is needed. These species have close affinities with A. tri- phyllum and A. cladolepton, as well as with high- altitude plants with narrow leaves. The last two differ principally in their more dissected leaves. In addition, A. triphyllum has scattered, minute, glandular trichomes on the petiole. Cajamarca: Prov. Cajamarca, El Gavilan, road Caja- marca-Chilete, Ferreyra 3256 (USM). La Libertad: Prov. Otuzco, Cerro facing Rio Bado. Aguado 6741 (GH). An- cash: Prov. Recuay, Pativilea-Huaraz Hwy., D. Smith 8306 (MO). Huanuco: Mito, Bryan 207 (F). Lima: Rio Blanco. Killip & Smith 21707 (F, GH, us). Junin: NE of Huancayo, between Acopalca and Huari, Duncan et al. 2746 (MO, uc, us). Ayacucho: Between Huanta and Ha- cienda Pargora, Killip & Smith 23316 (us). Cuzco: Prov. Cuzco, near Saxihuaman, Tryon & Tryon 5350 (F, GH). Puno: Valley 10 mi NE of Macusani, Webster 7 (K). Intermediate between A. gilliesii and A. peru- vianum: Cajamarca: Near Hacienda Porcon, N of Cajamarca, Correll& Smith P875 (GH, us). Lima: Chicla, Ball (GH). Cuzco: Prov. Anta, below Chinchaypujio, Vargas 17186 (GH). 39. Asplenium quitense Hooker, Sec. cent, ferns, /. 20. 1860. TYPE: Ecuador, Andes of Quito, Archidona, Jameson 707 (holotype, K.!; prob- able isotype, K.!). Plants epiphytic. Stems minute, compact, bear- ing leaves singly or in small clusters, several to many of these commonly connected by long, creeping, stoloniform roots, scales lacking. Leaves 1-6 to a stem, 1 -pinnate, 3-1 1 cm long, 0.8-1.8 cm broad. Petiole 0.5-3.5 cm long, marginate or very narrow-alate to base, greenish to yellowish, glabrous but sometimes with a few dark brown, subclathrate scales at base, these 0. 1-0.4 mm long and nearly as broad. Lamina thin- to firm-her- baceous, glabrous, lanceolate or oblong-lanceo- late, rather abruptly reduced to a pinnatifid apex or with a nonconform terminal segment, not or scarcely reduced at base. Rachis marginate or nar- row-alate, lacking proliferous buds. Pinnae 5-10 pairs, to 10 mm long and 7 mm broad, stalked, irregularly obovate or rhombic, strongly inequi- lateral, the base acroscopically truncate and the margin with 1-3 shallow or deep, obtuse lobes, basiscopically narrow-cuneate and the margin en- tire or with a single lobe toward the apex. Veins simple or 1 -forked, mostly obscure, commonly 2 (3) on acroscopic side of midrib and only 1 basi- scopically. Sori usually 1-3 per pinna. Indusia yel- low to brownish, linear or narrow-oblong. In dark, wet forests, on trunks or branches or trees, or on fallen logs, 700-1200 m, Pasco and Cuzco. Colombia; Ecuador; Peru. As in A. escaleroense, the stem of this species lacks scales and often produces long, stoloniform roots, which at intervals in turn produce other stems. This root system may extend for more than 40 cm along a tree trunk or branch. However, the former species has merely lobed or pinnatifid leaves, whereas A. quitense has distinctly stalked and lobed pinnae. Pasco: Prov. Oxapampa, Palcazu Valley, Rio Cacazu, Foster & Smith 9427 (F, MO). Prov. Oxapampa, Rio Ca- cazu, 56 km from Villa Rica, D. Smith & Foster 2663 (F, MO). Cuzco: Altura de Tintininquiato, Bues 1 752 (us). Prov. Paucartambo, Cosnipata Hacienda Maria. Vargas 10206 (uc). Prov. Paucartambo. Cosnipata Valley, Rio Tono, Wachter et al. 189 (F). 40. Asplenium pulchellum Raddi, Opusc. Sci. 3: 291. 1819. TYPE: Brazil, Mandiocca, Raddi (holotype, FI; isotype, BR; photos, BM, F & us of BR). Asplenium poloenseRosensl., Repert. Spec. Nov. Reg- ni Veg. 12: 469. 1913. TYPE: Bolivia, North Yungas, Polo-Polo, near Coroico, Buchtien 3345 (holotype, s!; isotypes, B!, us!). Plants terrestrial or epipetric, very rarely epi- phytic. Stem erect, provided with linear or lan- ceolate, often attenuate, scales, these 1-3 mm long, narrow-clathrate, blackish, sublustrous. Leaves fasciculate, 1 -pinnate, 6-16 cm long, 1.3-3 cm broad. Petiole 1-2.5 cm long, reddish brown to greenish, narrow-alate to marginate, provided with a few, scattered, dark, filiform scales. Lamina membranceous, lanceolate or linear-lanceolate, gradually reduced to a pinnatifid, long-acuminate or subcaudate apex, somewhat reduced at base. Rachis reddish or greenish brown, with a few, dark, filiform scales, narrow-alate, proliferous buds TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 35 lacking. Pinnae 7-15 pairs, 8-14 mm long, short- stalked, somewhat to strongly ascending, subdi- midiate, conspicuously inequilateral for half their length or more, acroscopic margin deeply and acutely cleft, the basiscopic essentially entire, the apex acute. Veins usually distinct, 3-5 on the ac- roscopic side of pinnae, 1-3 on the basiscopic, all simple except forked at the base acroscopically, all borne along the principal vein running near and parallel to the basiscopic margin. Sori commonly 3-7 on a pinna. Indusia thin, yellowish or greenish, oblong-elliptic. In deep, wet forests, on the ground or on or among rocks, very rarely at the base of tree trunks, 400-1800 m, San Martin, Huanuco, Junin, Madre de Dios. Ecuador; Peru; Bolivia; Brazil. This has been confused with A. otites (see Com- ments below), but the latter is a more robust plant with a stouter, much longer petiole; also pinnae are larger and are equilateral except toward the base, and have more veins and serrations. Asple- nium otites has not been found in Peru, but it is apparently somewhat common in northern South America; A. pulchellum does not occur in northern South America. The Schunke collection from San Martin (cited below) is atypical in that pinnae are not strongly subdimidiate, but it matches A. pulchellum in ev- ery other way. San Martin: Prov. Mariscal Caceres, Dist. Tocache Nuevo, Fundo La Campina, J. Schunke V. 3428 (F, GH, us). Huanuco: Prov. Huanuco, Dist. Churubamba, Hda. San Carlos, Mexia 825 la (uc, us), 8256a (us). Junin: Chanchamayo Valley, C. Schunke 177, 180, 221, 773, 940, 1364 (all F). Madre de Dios: Prov. Mania, Manu Park, Cocha Cashu uplands, Nunez 5775 (MO). 4 1 . Asplenium discrepans Rosenst., Repert. Spec. Nov. Regni Veg. 12: 469. 1913. TYPE: Bo- livia, North Yungas, Polo-Polo, near Coroico, Buchtien 3341 (holotype, B?; isotypes, GH!, us!). Plants epipetric or terrestrial. Stem erect, pro- vided with linear, attenuate scales, these 2-3 mm long, narrow-clathrate, blackish, sublustrous. Leaves fasciculate, 1 -pinnate, 6-16(-20) cm long, 1-3 cm broad. Petiole 1-3.5 cm long, reddish brown to greenish, narrow-alate, provided with scattered, dark, filiform scales. Lamina thin-her- baceous, lanceolate, gradually tapering to an acute, pinnatifid apex, somewhat reduced at base. Rachis reddish brown or greenish, provided with scat- tered, dark, filiform scales, narrow-alate, prolif- erous buds lacking. Pinnae 10-20 pairs, 6-15 mm long, sessile or with short, greenish stalks, patent or slightly ascending, subdimidiate, conspicuously inequilateral for half their length or more, acro- scopic margin crenate-serrate, the basiscopic es- sentially entire, the apex obtuse. Veins 4-6 and mostly forked on acroscopic side of pinna, l-3(- 4) and unbranched basiscopically, all borne along a principal rib running near and parallel to the basiscopic margin. Sori commonly 4-8 on a pinna. Indusia thin, yellowish or greenish, oblong-ellip- tic. In forests, on rocky hillsides and ravine banks, or on and among rocks, 700-1400 m, San Martin, Junin, Cuzco. Peru; Bolivia; Paraguay. This is frequently identified as A. pulchellum. Besides the characters of the key, it also differs from the latter in its usually patent or slightly as- cending pinnae. Pinnae of A. pulchellum typically are borne on the rachis at a very acute angle, al- though occasionally they may be slightly ascend- ing. San Martin: Tarapoto, Spruce 3966 (BR, GH, us). Ju- nin: San Francisco de Satipo, along Rio Satipo, Solomon 3233 (MO). Yaupi, Woytkowski 6501 (MO, us), 6504 (MO, us). Cuzco: Santa Ana, Cook & Gilbert 1535 (us), 6501 (MO, us), 6504 (MO, us). Santa Rosa, Urubamba Valley, Cook & Gilbert 1726 (us). Prov. La Convencion, Potrero, Tryon & Tryon 5381 (GH). 42. Asplenium pteropus Kaulf, Enum. fil. 170. 1824. TYPE: Brazil, Chamisso (holotype, LE). Plants epiphytic, sometimes terrestrial. Stem erect or decumbent, provided with narrowly clath- rate, linear or lanceolate, acute to attenuate scales, these 3-5 mm long, gray-brown or castaneous. Leaves 1 -pinnate, fasciculate, 1 5^4 5 (-60) cm long, 2.5-6 cm broad. Petiole 1-5 cm long, dull grayish to reddish brown, with a few, scattered, linear to filiform scales, broadly alate distally or through- out. Lamina glabrous, thin- to firm-membrana- ceous, narrow-elliptic, gradually and conspicu- ously reduced at both ends, terminating in a pinnatifid, occasionally proliferous, apex. Rachis dull reddish or greenish brown, broadly alate, each wing often as broad as the rachis, naked, or with scattered filiform scales. Pinnae 1.5-3 cm long, numerous, approximate to subdistant, sessile, acute to obtuse, conspicuously inequilateral at base, ba- 36 FIELDIANA: BOTANY siscopically cuneate to excavate, acroscopically truncate and often subauriculate, the margins deeply serrate to serrulate. Veins distinct, 5-10 pairs, simple, except forked at or near the pinna base acroscopically. Sori commonly 7-1 2 on larger pinnae. Indusia yellowish to hyaline, narrowly el- liptic. Rare in Peru, in deep forests, on tree trunks or on the forest floor, 400-1950 m, San Martin, Huanuco, Junin, Ayacucho. Southern Mexico to Panama; West Indies; Co- lombia to Guyana, south to Bolivia and Brazil. The taxonomy of this species has been confused, treated by various authors in a strict to a broad sense, and it is in need of much closer scrutiny. Broadly delimited, it includes rather thin-textured plants with short or medium-length petioles, ta- pered gradually to a pinnatifid apex and with a somewhat or strongly reduced base, the rachis and petiole broadly alate, pinnae conspicuously in- equilateral at base, with veins mostly simple. In a strict sense, the species would be confined to those plants with petioles only a few centimeters in length and with laminae strongly and gradually reduced to auriclelike basal pinnae. Plants previously de- termined as A. pteropus Hieron., with somewhat longer petioles and with five or six proximal pin- nae somewhat, but not greatly, reduced are prob- ably other species: in Central America perhaps A. barbaense Hieron., in South America perhaps A. raddianum or (in Ecuador) A. rosenstockianum Brade. The first two have narrow rachis wings, the last strongly alate wings as in true A. pteropus. For purposes of this flora, A. pteropus is delimited in a strict sense and, as such, is not very common in Peru. San Martin: Prov. Mariscal Caceres, Dist. Tocache Nuevo, Quebrada de Saule Chico, J. Schunke V. 4358 (F, us). Huanuco (as San Martin): Tingo Maria, across the Huallaga on trail to "Monson" (Monzon), Allard 21864 (us). Junin: Prov. Chanchamayo, Rio Rondayacu, 45 km from San Ramon, D. Smith et al. 2614 (F, GH). La Merced, Chanchamayo, Soukup 1067, in part (F). Ayacucho: Rio Apurimac Valley, near Kimpitiriki, Killip & Smith 22918 (F, us). 43. Asplenium sessilifolium Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. 5: 322. 1811. TYPE: "Habitat in America ae- quinoctale," collector undesignated (holo- type, P, Herb. Jussieu 1265; isotype, B; photos, BM, GH, us of P). Plants terrestrial, at least in Peru. Stem stout, erect or decumbent, provided with clathrate, lin- ear or lanceolate, attenuate scales, these 2-5 mm long, gray-brown to blackish, often with narrow, lighter brown margins. Leaves 1 -pinnate, caespi- tose, 20-40 cm long, 2-5 cm broad. Petiole 3-16 cm long, gray-brown to atropurpureous or black- ish, dull or slightly lustrous, naked or with a few lanceolate scales, not or scarcely alate. Lamina gla- brous, thin- to firm-herbaceous, linear or narrow- elliptic, gradually and strongly reduced at both ends, terminating in a pinnatifid (rarely prolifer- ous) apex. Rachis castaneous to atropurpureous (occasionally grayish brown), narrow-alate to mar- ginate, naked or with a few dark, filiform scales. Pinnae 1-3 cm long, numerous, approximate to crowded, sessile, conspicuously inequilateral at base, basiscopically cuneate to excavate, acroscop- ically truncate and incised nearly to costa to form a nearly free basal auricle (occasionally opposed by a smaller basiscopic auricle), the rest of the pinna serrate, crenate, or shallowly lobed. Veins distinct, 4-7 pairs, simple, or subflabellate in the auricles. Sori 8-12 on larger pinnae. Indusia thin, linear or elliptic, yellowish or hyaline. On rocky slopes and wet ravine banks of cloud forests, 2500-3600 m, Cajamarca, Junin, Ayacu- cho, Apurimac, Cuzco. Southern Mexico to Costa Rica; Greater Antil- les; Ecuador; Peru; Bolivia. Pinnae of Asplenium triphyllum are typically bi- or trifoliolate, but in occasional plants they may vary in having extra lobes, crenations, or serra- tions, and such specimens sometimes have been identified as A. sessilifolium. However, the two differ significantly in indument. In the latter, the blackish stem scales are 2-5 mm long and often have narrow, brown margins, and petiole and ra- chis are naked or with a few, scattered, lanceolate or filiform scales. In A. triphyllum, the blackish stem scales are usually less than 2 mm long and lack lighter colored margins, and axes are sparsely to moderately provided with filiform scales and glandular trichomes. Cajamarca: Prov. Celendin, canyon of Rio Maranon above Balsas, Hutchison & Wright 5295 (uc). Junin: Huari, Kunkel, 1960 (B). Ayacucho: Prov. La Mar, above Jano, Plowman & Davis 4677 (GH). Between La Quinua and Tambo, Soukup 5625 (us). Apurimac: Forests of Ampay, Vargas 1069 (GH, MO). Cuzco: Prov. La Con- vention, Dist. Vilcabamba, between Rumichurco and Alcobamba, Davis et al. 1235 (F, GH). Prov. La Conven- tion, Choquequiran, Vargas 12999 (F). TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 37 44. Asplenium tabinense Hieron. Hedwigia 60: 224. 1918. TYPE: Peru, near Tabina (Puno), Lechler 2105 (holotype, B!; photo, F). Plants terrestrial, at least in Peru. Stem erect or decumbent, provided with narrowly clathrate, lan- ceolate or linear-lanceolate, attenuate scales, these 2.5-4 mm long, blackish, with hairlike tip and scattered marginal cilia (the latter processes usu- ally broken away in dried plants). Leaves 1 -pin- nate, fasciculate, 15-33 cm long, 2-4 cm broad. Petiole 1-5 cm long, atropurpureous, dull or slightly lustrous, with scattered, dark, filiform scales, nar- row-alate or marginate. Lamina glabrous, mem- branaceous, narrow-elliptic, gradually and con- spicuously reduced at both ends, terminating in a pinnatifid, usually attenuate, apex. Rachis atro- purpureous, narrow-alate, naked or with scattered, dark, filiform scales. Pinnae 1.2-2 cm long, nu- merous, approximate to crowded, subsessile, con- spicuously inequilateral at base, basiscopically cu- neate to excavate, acroscopically truncate and often auriculate, the auricle sometimes incised nearly halfway to midrib, the rest of the pinna narrowly and deeply crenate. Veins (3-)4-5 and simple on the basiscopic side of larger pinnae, 5-7 on the acroscopic side, simple, except subflabellate in the basal auricle and forked in the adjacent lobe. Sori commonly 4-7 on larger pinnae. Indusia thin, nar- row-elliptic, yellowish or hyaline. On densely wooded slopes or in thickets, 1500- 3100 m, Huancavelica, Ayacucho, Cuzco, Puno. Peru; Bolivia. All specimens seen from Bolivia are epiphytes, whereas plants in Peru grow on the forest floor. Besides the characters cited in the key, this also differs from A. sessilifolium in the more delicate lamina, usually firm-herbaceous in the latter, and membranaceous in A. tabinense. The rachis in A. sessilifolium varies from atropurpureous to cas- taneous (rarely lighter colored), but it is always atropurpureous in A. tabinense, at least on mature leaves. This perhaps might be considered merely a geographic variant of the widespread A. sessili- folium. Huancavelica: Prov. Tayacaja, Ampurco, between Sal- cabamba and Surcubamba, Tovar 3768 (GH). Ayacucho: Ccarrapa, between Huanta and Rio Apurimac, Killip & Smith 22385 (us). Cuzco: Quillabamba, Santa Teresa, 0.5 km W of La Playa, Peyton & Peyton 1236 (MO). Prov. La Convencion, Huayopata, Pistipata drainage, Peyton & King 1429 (MO). Prov. Calca, near Mantio, Vargas 15594 (GH). 45. Asplenium herzogii Rosenst., Meded. Rijks. Herb. 19: 12. 1913. TYPE: Bolivia, Incacor- ral, Herzog 2211 (holotype, B?; isotype, us!; photo, F of us). Plants terrestrial. Stem erect, provided with ob- scurely clathrate, lanceolate scales, these 2-3 mm long, gray-brown, acuminate. Leaves 1 -pinnate, fasciculate, 25-50 cm long, 3-5.5 cm broad. Pet- iole 6-12 cm long, gray-brown, narrowly to ves- tigially alate. Lamina linear, gradually reduced to a subcaudate, proliferous apex, slightly reduced at base, with a few proximal pairs of pinnae about half the length of the longest ones. Rachis dull red- or gray-brown, narrow-alate. Pinnae 2-2.8 cm long, 1 6-25 pairs, apex obtuse or subacute, inequilateral at base, basiscopically cuneate, acroscopically truncate and subauriculate, the margins crenate- serrate. Veins 5-8 pairs, simple, except forked near the pinna base acroscopically. Sori 9-15 on larger pinnae. Indusia thin, hyaline or yellow-brown, narrow-elliptic, entire. Thus far known in Peru from one collection, in thicket, about 2750 m, Huanuco. Peru; Bolivia. This may be confused with A. alatum because of the subcaudate, proliferous rachis but is easily separated from the latter by the narrow rachis wings and by the strongly inequilateral pinna bases. In A. alatum the wings on the rachis (and usually the petiole) are very broad (1-2 mm), and pinnae are mostly subequilateral. Asplenium herzogii is most closely related to A. raddianum, from which it can be distinguished by the proliferous rachis, the smaller (2-3 mm vs. 4-6 mm) stem scales and the linear lamina. Huanuco: Chaglla, Macbride 3639 (F). 46. Asplenium bangii Hieron., Hedwigia 60: 245. 1918. TYPE: Bolivia, Bang 2419 (holotype, B; isotypes, F!, NY!, us!). Plants terrestrial, rarely epiphytic or epipetric. Stem erect, provided with obscurely clathrate, lin- ear or linear-lanceolate, attenuate, scales, these 1.5- 3 mm long, blackish, sublustrous, their margins subentire. Leaves 1 -pinnate, caespitose, 30-65 cm long, 3.5-5 cm broad. Petiole (5-)6-15 cm long, dull purplish to reddish brown, narrowly to ves- tigially alate. Lamina linear or linear-lanceolate, gradually reduced to a pinnatifid apex scarcely reduced at base or several proximal pairs about 38 FIELDIANA: BOTANY half the length of the longest ones. Rachis dull purplish or reddish brown, narrow-alate (rarely the wing rather broad), lacking proliferous buds. Pin- nae (larger ones) 1.2-2.2(-2.5) cm long, (30-)35- 45 pairs, usually obtuse at apex, conspicuously inequilateral at base, basiscopically cuneate to ex- cavate, acroscopically truncate and often subauric- ulate, the margins subentire to crenulate. Veins 8- 12 on acroscopic side of costa, 5-8 basiscopically, simple, except forked at or near the pinna base acroscopically. Sori 12-18 on larger pinnae. In- dusia thin, yellowish or hyaline, margins entire. In damp forests, on wooded slopes and canyon walls, commonly on the forest floor, rarely on cliffs or tree trunks, 650-2150 m, Huanuco, Pasco, Ju- nin. Peru; Bolivia; Brazil. With this perhaps should be included A. claus- senii Hieron. of Brazil, northern South America, and the Greater Antilles, of which Hieronymus recognized three forms, based on minor differ- ences in length, margin and apex of pinnae. Mor- ton and Lellinger (1966, p. 16) designated Fendler 138, Venezuela (B; photo, us), the lectotype for this name, but unfortunately this specimen was unvailable for examination during our study. In the Field Museum herbarium, there is a sheet of Fendler 138, containing laminae of three different species. The center one is possibly A. claussenii and is very similar to A. bangii, but it lacks petiole and stem for definitive comparison. Given the ap- parent variability of A. claussenii, the entire com- plex (including A. drepanophyllum and A. pul- chellum) should be subjected to closer scrutiny. Therefore, A. claussenii is excluded from Peru for purposes of this treatment. Huanuco: Mima, Bryan 433 (F, GH). Mima, Macbride 4010 (F, GH, us). Pasco: Prov. Oxapampa, Valle del Pal- cazu, Rio Cacazu, Leon 678 (F, USM). Junin: La Merced, Macbride 5372 (F, GH, us). Chanchamayo Valley, C. Schunke 1371 (F in part, us). Prov. Tarma, Huacapis- tana, Tryon & Tryon 5453 (F, GH, uc, us, USM). 47. Asplenium harpeodes Kunze, Linnaea 18: 329. 1844. LECTOTYPE (designated by Morton & Lellinger, Mem. New York Bot. Gard. 15: 1 5. 1966): Mexico, Leibold26 (B!; photos, us). Plants terrestrial or epiphytic. Stem erect or de- cumbent, provided with clathrate, linear scales, these 4-10 mm long, deep brown or blackish, lus- trous, terminating in a long, tortuous, unicostate tip. Leaves 1 -pinnate, caespitose, 30-120 cm long 4-12 cm broad. Petiole 5-10 cm long, dull yel- lowish to reddish brown or purple-gray, wings lacking or vestigial. Lamina linear or linear-lan- ceolate, gradually reduced to a pinnatifid apex, scarcely reduced at base, or several proximal pairs about half the length of longest ones. Rachis atro- purpureous, narrow-alate, lacking proliferous buds. Pinnae (larger ones) 2.5-9 cm long, (24-)30-50 pairs, long-attenuate, inequilateral at base, basi- scopically excavate, acroscopically truncate, often subauriculate (but the basal lobe never incised more than halfway to the costa), the margins deeply, serrately lobed. Veins 1 2-20 pairs, mostly simple, except subdichotomously forked in the acroscopic auricle. Sori numerous. Indusia thin, yellowish, brownish or hyaline, margins subentire. Common, on the earth in wet forests and thick- ets, in humus, on wet banks, or rocky soil or on mossy rocks, often on tree trunks, 1 500-3600 m, Cajamarca to San Martin, south to Huancavelica and Puno. Southern Mexico to Panama; Greater Antilles; Colombia and Venezuela south to Bolivia and Brazil. In addition to the characters cited in the key, this usually can be recognized by the narrowly and deeply lobed pinnae that very gradually taper to long-attenuate tips. However, several depauperate specimens have been seen that have merely strong- ly serrate margins and acute tips. In these, the purple rachis and the stem scales with long, tor- tuous tips are diagnostic. This is one of the most common species of the genus in the Neotropics. Cajamarca: Colosay, Woytkowski 7010 (GH, MO, us). Amazonas: Prov. Chachapoyas, W of Molinopampa, Wurdack 1471 (GH, us, USM). San Martin: Prov. Maris- cal Caceres, Rio Abiseo National Park, Young 4 171 (USM), Young & Leon 501 1 (USM). Huanuco: Cani, 7 mi NE of Mito, Macbride 3390 (F, GH, us). Pasco: Prov. Oxapam- pa, 2-4 km N of Mallampampa, D. Smith & Canne 5865 (MO). Junin: Chanchamayo Valley, above La Merced, Hutchison 1197 (F, NY, uc, us). Huancavelica: Prov. Tay- acaja. Chuspi, near Tocas, Tovar 2063 (USM). Ayacucho: Prov. La Mar, between El Tambo and Ayna, Plowman & Davis 4675 (GH), 4680 (GH). Apurimac: Forests of Ampay, Vargas 1065 (GH)Cuzco: Prov. Urubamba, Cuz- co-Quillabamba Road, Skog & Skog 5173 (us). Puno: Prov. Carabaya, between Ayapata and K'ahualluyoc, Vargas 10763 (GH). 48. Asplenium raddianum Gaud., in Freycinet, Voy. Uranie, Bot., 316. 1828. Norn. nov. for A. brasiliense Raddi and based on the same type. TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 39 Asplenium brasiliense Raddi, PI. bras. nov. gen. 1 : 36, t. 51,f. 1. 1825, not Swartz, 1817. TYPE: Brazil, Prov. Rio de Janeiro, Raddi (holotype, n; isotype, BR; photo, BM of BR). Plants terrestrial, occasionally epiphytic. Stem erect, but often slender and elongate, provided with obscurely clathrate, ovate- to linear-lanceolate scales, these 4-6 mm long, gray-brown, acute to attenuate. Leaves 1 -pinnate, fasciculate, 25-50 cm long, 4-7 cm broad. Petiole 6-10 cm long, reddish or grayish brown, wings narrow or vestigial. Lam- ina lanceolate or elliptic-lanceolate, gradually re- duced to a pinnatifid apex, scarcely reduced at base, or several proximal pairs about half the length of the longest ones. Rachis dull reddish or grayish brown, narrow-alate, lacking proliferous buds. Pinnae (larger ones) 2.5-3.5 cm long, (12-) 15-24 pairs, apex acute to obtuse, inequilateral at base, basiscopically excavate, acroscopically truncate, often subauriculate (but the basal lobe never in- cised more than halfway to the costa), the margins deeply serrate or serrately lobed. Veins usually 8- 1 1 on acroscopic side of costa, 6-9 basiscopically, simple, except forked at or near the pinna base acroscopically. Sori 10-16 on larger pinnae, fre- quently diplazioid in the acroscopic auricle. In- dusia yellowish to light brown, margins subentire. On wet slopes or in litter on the forest floor, occasionally on trunks of trees or tree ferns, 1000- 2800 m, Cajamarca and Amazonas south to Cuzco and Puno. General distribution uncertain; probably Ven- ezuela and Colombia; Brazil; Peru; possibly Bo- livia. There is some question as to proper application of this name for the plants so circumscribed here. The type was not seen, but the specimens cited below closely match the description and most her- barium specimens identified as A. raddianum. As- plenium barbaense Hieron., from Costa Rica, may belong here, as the two species differ only quan- titatively and even share the distinctive feature of having relatively frequent diplazioid sori in the basal acroscopic pinna auricle. Otherwise, A. bar- baense essentially differs only in having narrower, more delicate laminae, proximal pinnae with short stalks, broader petiole and rachis wings. Perhaps plants from Costa Rica and northern South Amer- ica represent an infraspecific variant of those from Peru, Bolivia, and Brazil. Further study is re- quired. Cajamarca: Colasay, Woytkowski 701 1 (MO, us). Ama- zonas: Prov. Bongara, Shillac, N of Pedro Ruiz, D. Smith & Vdsquez 4885 (MO, uc). San Martin: San Roque, Li Williams 7189 (us). Ancash: Prov. Huaraz, Dist. Hua- raz, behind Hotel Monterrey, Sounders 1282, atypical (F, GH). Pasco (as Junin): Pichis Trail, "Yapas" [Yapaz], Killip & Smith 25520 (us). Junin: Schunke Hacienda above San Ramon, C. Schunke A208 (GH, us). Chan- chamayo Valley, C. Schunke 108 (us). Cuzco: Urubam- ba, Machu Picchu, Peyton & Peyton 372 (MO). Puno: Prov. Carabaya, San Gaban, Vargas 18913, atypical (GH). 49. Asplenium alatum Willd., Sp. pi. ed. 4, 5: 319. 1810. TYPE: Venezuela, Nova Andalusia, near Caripe, Humboldt & Bonpland (holo- type, B, Herb. Willd. 19889; isotype, p; pho- tos, F & GH of B). Plants terrestrial, rarely epipetric. Stem erect, provided with obscurely clathrate, ovate or ovate- lanceolate scales, these 2-3 mm long, gray-brown acute. Leaves 1 -pinnate, crowded to subfascicu- late, 20-50 cm long, 7-11 cm broad. Petiole 6-15 cm long, dull greenish or grayish brown, conspic- uously alate, often to base. Lamina broadly lan- ceolate to oblong, rather abruptly terminating in a short-ligulate, proliferous tip, not or scarcely re- duced at base. Rachis dull reddish or greenish brown, broadly alate, the wings each 1-2 mm broad. Pinnae to 7 cm long and 1.5 cm broad, 12- 20 pairs, obtuse to acute, broadly cuneate to trun- cate or scarcely auriculate at base, subequilateral, the margins broadly serrate or biserrate. Veins dis- tinct, mostly 1 -forked. Sori 9-12 pairs on larger pinnae. Indusia thin, linear, yellowish or greenish, margins entire. On the floor of wet forests or banks of wooded ravines, rarely on wet, mossy rocks, (700-) 1 200- 2400 m, along the Cordilleras Central and Ori- ental, Cajamarca and Amazonas south to Cuzco. Southern Mexico to Panama; Jamaica; Hispan- iola; Colombia and Venezuela south to Bolivia and Brazil. The very broad rachis wing and ligulate, prolif- erous rachis tip distinguish this from most other species of Asplenium in Peru. Cajamarca: Prov. Cutervo, Grutas de San Andres, Lla- tas Quiroz & Suarez 2734 (F, uc). Amazonas: Prov. Bon- gara, Quebrada Chachuaico, near Shilla, Young et al. 398 (MO, uc). San Martin: Between Moyobamba and Chachapoyas, Croat 58166 (F, MO). Prov. Mariscal Ca- ceres, Rio Abiseo National Park, Young 4221 (HUT, USM). Huanuco: Mima, Macbride 4032 (F, us). Pasco: Pozuzo, Hacienda Ballisteros, Bryan 692 (F, us). Junin: Chan- chamayo Valley, C. Schunke 57 (F, us), 727 (F, us). Cuz- co: Prov. Paucartambo, Kosnipata Valley, Plowman & Davis 4936 (GH). 40 FIELDIANA: BOTANY 50. Asplenium abscissum Willd., Sp. pi. ed. 4, 5: 321. 1810. LECTOTYPE (designated by Morton & Lellinger, Mem. New York. Hot. Gard. 15: 26. 1966): Venezuela, Caracas, Bre- demeyer (*, Herb. Willd. 19893-1, 19893-2; photos, F & GH). Plants terrestrial, occasionally epipetric, rarely epiphytic. Stem erect or decumbent, provided with obscurely clathrate, linear or lanceolate scales, these 1-2.5 mm long, blackish or with very narrow brown margins. Leaves 1 -pinnate, fasciculate, 30-50 cm long, 5-15 cm broad. Petiole 15-25 cm long, yel- lowish to reddish brown, not or scarcely alate. Lamina chartaceous, ovate to oblong, terminating abruptly in a broad-based, nonconform apical seg- ment, not reduced at base. Rachis dull, yellowish or reddish brown, narrow-alate, lacking prolifer- ous buds. Pinnae (larger ones) 1.5-12 cm long, 0.7-2 cm broad, 4-12 pairs, apex obtuse to atten- uate, scarcely to strongly inequilateral at base, the margins subentire to crenate-serrate. Veins few to many, 1-2-forked. Sori linear or narrow-elliptic. Indusia thin, yellow to brownish, scarious, often hyaline until mature, margins subentire. The species occurs in the United States (Flori- da); southern Mexico to Peru; West Indies; Co- lombia and Venezuela south to Bolivia and Uru- guay. Throughout most of its broad range the features of this species are rather constant. However, in Bolivia and southern Peru a number of specimens have been found with elongated and more equi- lateral pinnae. Also, throughout much of Peru and occasionally growing alongside typical A. abscis- sum, there occur plants with short, broad, and obtuse pinnae, and a few of these, including a syn- type of A. ruizianum, have been reported from Venezuela. Since these differences are merely quantitative, three taxa are recognized here as geo- graphical variants of A. abscissum. Another species, A. congestum C. Chr. of Ec- uador, is very closely related but should be con- sidered distinct, as its lamina is considerably smaller and membranaceous and tapers gradually to a narrow, pinnatifid apex. This contrasts with the chartaceous lamina of A. abscissum, which abruptly ends in a broad-based, nonconform apex. Key to Varieties a. Pinnae subequilateral at base, broadly cuneate acroscopically, narrowly so basiscopically, narrowly acute to attenuate at apex, commonly 6-10 times as long as broad; veins of larger pinnae 12-20 pairs 50b. var. subaequilaterale a. Pinnae conspicuously inequilateral at base, truncate acroscopically, cuneate to excavate basiscopically, obtuse to narrowly acute at apex, commonly 2-4.5 times as long as broad; veins 4-10(-12) pairs b b. Pinnae 3-4.5(-6) times as long as broad, subacute to narrowly acute at apex 50a. var. abscissum b. Pinnae 1.8-2.3 times as long as broad, broadly rounded at apex 50. var. ruizianum 50a. Asplenium abscissum var. abscissum. In wet forests or wooded canyons, on slopes or on the forest floor, occasionally on rocks or rocky cliffs, rarely on bases of tree trunks or fallen logs, 300-2200 m, Cajamarca and Amazonas south to Ayacucho and Madre de Dios. Range the same as the species. Cajamarca: Prov. Santa Cruz, 3.2 km ENE of Mon- teseco, Santisteban & Guevara 23 (F, GH, HUT, uc). Ama- zonas: Prov. Bagua, Cerro Tapur, near Rio Utcubamba, Hutchison 1473 in part (F, uc). San Martin: Juan Jui, Alto Rio Huallaga, Klug4312 (F, GH, MO, uc, us). Huanu- co: Cliffs W of Rio Huallaga, above bridge, Croat 57967 'a (MO in part, USM). Pasco: Cueva Grande, near Pozuzo, Macbride 4791 (F, us). Junin: Hacienda Perene, Coro- nado 247 (GH, uc). Ayacucho: Ayna, between Huanta and Rio Apurimac, Killip & Smith 22835 (F, us). Cuzco: Calca, Vargas 15595 (GH). Madre de Dios: Parque Na- cional del Manu, Cocha Cashu Biological Station, M. Foster P-84-50 (uc). 50b. Asplenium abscissum var. subaequilaterale Rosenst., Repert. Spec. Nov. Regni Veg. 12: 470. 1913. TYPE: Bolivia, southern Yungas, Polo-Polo, near Coroico, Buchtien 3349 (ho- lotype, presumably B). On the ground in dark, wet forests, 800-1600 m, Junin and Ayacucho. Peru; Bolivia. TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 41 This variety is very similar to A. tuerckheimii in many aspects, but it is primarily distinguished from the latter by the nonconform apical segment and the 7-12 pairs of pinnae. The lamina of A. tuerckheimii terminates in a conform apical seg- ment and bears only 3-6 pairs of pinnae. Junin: E of Quimiri Bridge, near La Merced, Killip & Smith 23900 (us). Chanchamayo Valley, C. Schunke 74 (F, us), 722 (us), 935 (F). Ayacucho: Ayna, between Huanta and Rio Apurimac, Killip & Smith 22782 (F, us). 50c. Asplenium abscissum var. ruizianum (Klotzsch) Stolze, comb, and stat. nov. As- plenium ruizianum Klotzsch, Linnaea 20: 354. 1847. LECTOTYPE (designated by Morton & Lellinger, Mem. New York Bot. Card. 15: 27. 1966): Andes of Peru, Ruiz (B!; fragment, NY!; photos, F & us). In wet forests and wooded canyons, in the earth or on or among wet rocks, very rarely on tree trunks, 800-2700 m, Tumbes to Cuzco. Venezuela; Peru. Tumbes: Prov. Zarumilla, Dist. Matapalo, Bosque Na- cional de Tumbes, Schunke V. 2408 (F). Cajamarca: Co- lasay, Woytkowski 7012 (MO, us). Amazonas: Prov. Ba- gua, 5 km above La Peca, Croat 58353. Prov. Bagua, Rio Utcubamba, Cerro Tapur, Hutchison 1473 (F in part, uc in part, us). Huanuco: Muna, Bryan 434 (F, GH, us). Junin: Chanchamayo Valley, C. Schunke 74 (F, us). Cuz- co: Prov. La Convencion, Quebrada Vilcabamba, Vargas 23397 (USM). 51. Asplenium drepanophyllum Kunze, Linnaea 9: 66. 1834, not Baker, 1874. TYPE: Peru, Pampayaco (Dept. Huanuco, Pampayacu), Poeppig July, 1829 (holotype, w; frag., BM!; photOS, BM Of W, F & GH of BM). Plants terrestrial, rarely epipetric. Stem erect or decumbent, provided with narrowly clathrate, lin- ear-lanceolate, attenuate, scales, these 2-3 mm long, dark brown to blackish. Leaves 1 -pinnate, fasciculate, 15-40 cm long, 3-4.5(-5) cm broad. Petiole 4-9(-12) cm long, yellowish to reddish brown, not or scarcely alate. Lamina membrana- ceous to firm-herbaceous, linear-lanceolate, grad- ually reduced to a narrow, pinnatifid, usually acu- minate apex, scarcely reduced at base or several proximal pairs about half the length of the longest ones. Rachis dull reddish brown, narrow-alate, lacking proliferous buds. Pinnae (larger ones) 1 .2- 3 cm long, 0.4-0.8 cm broad, 15—40 pairs, apex obtuse to attenuate, conspicuously inequilateral at base, basiscopically cuneate to excavate, acroscop- ically truncate and often subauriculate, the mar- gins crenulate to deeply biserrate. Veins 6-12 on acroscopic side of costa, (4-)5-10 basiscopically, 1-2-forked, except simple toward pinna base bas- iscopically. Sori 6-12 on larger pinnae, occasion- ally diplazioid. Indusia thin, yellowish or brown- ish, scarious, often hyaline until mature, margins entire. In dense, wet forests, on the forest floor, rarely on mossy rocks, 100-1600 m, San Martin and Loreto south to Ayacucho and Madre de Dios. Peru; Bolivia; Paraguay. Pinna apices vary greatly from obtuse to acu- minate, and extremes might be presumed different species, yet these differ from each other in no other way, and there are an infinite number of inter- mediate conditions. Leaves with obtuse pinnae are easily confused with A. bangii (under which see further discussion), but veins of the latter are all simple except at the acroscopic pinna base. As- plenium drepanophyllum is also similar to the enigmatic A. otites, discussed under Comments at the end of the genus treatment. San Martin: Along Rio Huallaga, NE of Shapaja on road to Chazuta, Knapp & Mallet 6915 (F, MO). Loreto: Puerto Arturo, lower Huallaga below Yurimaguas, Killip & Smith 27794 (us). Junin: E of Quimiri Bridge near La Merced, Killip & Smith 23909 (F, us). Between La Mer- ced and Satipo, Leon 188 (F, USM). Ayacucho: Estrella, between Huanta and Rio Apurimac, Killip & Smith 23078 (us). Cuzco: Prov. La Convencion, Rio Apurimac, be- tween Puerto Capiro and Boca de Tigre rapids, Davis el al. 1297 (F, GH). Prov. La Convencion, edges of Rio Mapitunuari, Dudley 10128 (GH, MO). Madre de Dios: Prov. Manu, Cocha Cashu uplands, Nunez 5766, in part (MO), 6092.4 (MO). 52. Asplenium auritum Sw., J. Bot. (Schrader) 1800(2): 52. 1801. TYPE: Jamaica, Swartz (s). (For detailed list of synonyms, see Morton &Lell., 1966, pp. 17-20.) Plants epiphytic or terrestrial, occasionally epi- petric. Stem erect, provided with clathrate, lan- ceolate to ovate scales, these 2—4 mm long, acute or acuminate, dark or grayish brown, eciliate. Leaves 1 -pinnate or 1 -pinnate-pinnatisect, rarely nearly 2-pinnate at base of proximal pinnae, fas- ciculate, 10-50 cm long. Petiole 5-20 cm long, dull green, gray, reddish brown, or blackish, not or scarcely alate. Lamina chartaceous to subcoria- ceous, deltate to oblong-lanceolate, very gradually tapering to a pinnatifid to serrate, often subcaudate 42 FIELDIANA: BOTANY apex, not reduced at base. Rachis green, gray, or reddish brown, scarcely alate, or sometimes broadly so distally, lacking proliferous buds. Pinnae 1-9 cm long, usually numerous, approximate to sub- distant, varying from obtuse and subentire and scarcely auriculate at base to deeply pinnatifid, subcaudate at apex, and with a nearly free basal acroscopic auricle. Veins indistinct or obscure, forked, or pinnately branched in larger segments. Son elliptic, frequently confluent at maturity. In- dusia firm and fleshy, opaque, commonly pale yel- low or whitish, subentire. In forests and thickets, on tree trunks and branches, on the forest floor, or occasionally on rocky soil or cliffs, 150-2900 m, Cajamarca to Loreto, south to Ayacucho and Puno. Southern United States (Florida); West Indies; southern Mexico to Bolivia and Argentina. This and A. cuspidatum are part of a highly vari- able and widely distributed complex that is char- acterized within the genus by the fleshy, light green lamina that contrasts with the rich brown sori (the latter often confluent at maturity) and by the thick, fleshy, opaque indusia. Because of the variable dissection of the lamina ( 1 -pinnate to 3-pinnate- pinnatifid) and the obtuse to subcaudate pinnae, authors have separated the species complex into as many as a dozen taxa. However, individual specimens essentially differ only in pinna shape and degree of lamina dissection; thus, for purposes of this treatment, only A. auritum and A. cuspi- datum (including three varieties) are recognized here. These are separated merely as a matter of convenience, by purely quantitative characters, as seen in the following key, and there are a number of intermediates even in these two components of the group. a. Pinnae simple and entire to pinnatifid (rarely nearly pinnate at base of proximal pinnae; basal segments (if any) adnate, never truly stalked A. auritum a. Pinnae (at least proximal ones) 1 -pinnate to nearly 3 -pinnate; basal segments (pinnules) of proximal pinnae obviously free and stalked A. cuspidatum Cajamarca: San Ignacio, San Juan de Piedra Bola, Diaz 2028 (F, MO). Amazonas: Kayamas Creek flowing into Rio Cenepa, Berlin 463 (MO, us, USM). San Martin: Tara- poto, Spruce 3956 (BR), 4677 (us). Loreto: Rio Mazan, J. Schunke 299 (F, GH, uc, us, USM). Huanuco: Prov. Huanuco, Dist. Churubamba, Hacienda Mercedes, Mex- ia 8179 (F, GH, uc). Pasco: Prov. Oxapampa, Dist. Ox- apampa, Rio Alberto, Leon 628 (USM). Junin: Prov. Tar- ma, Utcuyacu, Woytkowski 35363 (MO, uc). Ucayali: Prov. Coronel Portillo, Bosque von Humboldt, Young & Salazar 1026 (F, MO). Ayacucho: Ccarrapa, between Huanta and Rio Apurimac, Killip & Smith 22369 (us), 22477 (us). Cuzco: Prov. Paucartambo, road from Pilla- wata to Patria, Plowman & Davis 4941 (F, GH, USM). Madre de Dios: Prov. Manu. Rio Manu, Cocha Cashu Station, Foster 9703 (F, MO, USM). Puno: Prov. Carabaya, San Gaban, Vargas 18922 (GH). 53. Asplenium salicifolium L., Sp. pi. 1080. 1753. LECTOTYPE (designated by Morton & Lei- linger, Mem. New York Bot. Gard. 15: 23. 1966): Plumier, Traite foug. Amer. 18, t. 60. 1 705, based on a plant from the West Indies. Plants terrestrial (often epiphytic outside Peru), the roots densely golden-tomentose. Stem erect, provided with clathrate, lanceolate to ovate-lan- ceolate scales, these 2-6 mm long, acute to atten- uate, gray-brown, usually with long, orange cilia. Leaves 1 -pinnate, subfasciculate, 30-80 cm long, 9-20 cm broad. Petiole 8-30 cm long, dull yel- lowish to reddish brown, not alate, sparsely scaly and often golden-tomentose at base. Lamina thin- to firm-herbaceous, lanceolate to deltate-ovate, abruptly reduced at apex to a broad-based, non- conform or subconform segment, not or scarcely reduced at base. Rachis dull yellowish or reddish brown, narrow-alate, lacking proliferous buds. Pinnae 1 2-20 pairs, larger ones 4-10 cm long, 1 .2- 2.4 cm broad, subdistant to remote, narrowly acute to attenuate, inequilateral at base, commonly cor- date and auriculate, the auricles often overlapping the rachis acroscopically, narrow-cuneate basi- scopically, the margins obtusely crenulate to cre- nate or bicrenate. Veins distinct to obscure, 2-forked, at least in the proximal half of pinnae. Sori linear or narrow-elliptic, inframedial. Indusia firm and fleshy, opaque, yellowish to light brown. Dry woods or cloud forests, on the forest floor (but often on tree trunks and branches outside Peru), 2150-2700 m, Amazonas and Huanuco. Southern Mexico; Nicaragua; Costa Rica; West Indies; Colombia to the Guianas, south to Peru and Brazil. This, A. auriculatum, and A. hastatum have been combined by some authors. Their differences are merely quantitative, but they are maintained sep- arately here pending revision of the group. Morton TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 43 and Lellinger (1966) and Stolze (1986) partially based their separation of the species on width of rachis wing, texture of lamina, or acroscopic pinna bases overlapping the rachis. None of these char- acters are consistent or significant, whereas length of stem scales, forking of veins, and shape of pinna and lamina apex are far more reliable, over the broad range of distribution. Amazonas: Bongara, 4 km N of Pomacochas, Knapp et al. 7523 (F, MO). Prov. Chachapoyas, slopes of Puma- urcu SE of Chachapoyas, Wurdack 557 (F, GH, uc, us, USM). Huanuco: Muna, Bryan 528 (F). Cushi, trail to Tambo de Vaca, Bryan 618 (F). Huacachi, estacion near Muna, Macbride 4107 (F, us). 54. Asplenium hastatum Kunze, Linnaea 23: 235, 305. 1850. LECTOTYPE (designated by Morton & Lellinger, Mem. New York Bot. Gard. 15: 21. 1966): Hort. Lips., originally from Venezuela (B!, photos F, us; isolectotype, Plants epiphytic or terrestrial, the roots densely golden-tomentose. Stem erect, provided with clathrate, linear-lanceolate, attenuate scales, these commonly 6-9 mm long, attenuate, gray-brown, usually with long, orange cilia. Leaves 1 -pinnate, 30-80 cm long, 8-16 cm broad. Petiole 12-30 cm long, dull greenish to reddish brown, not or slightly alate, sparsely scaly and often golden-tomentose at base. Lamina firm-herbaceous to chartaceous, lance-ovate to oblong, abruptly reduced to a nar- row, serrate-ligulate or subcaudate apex, not or scarcely reduced at base. Rachis dull reddish or greenish brown, alate, lacking proliferous buds. Pinnae 12-25 pairs, larger ones 6-9 cm long, 1-2 cm broad, subdistant to remote, commonly atten- uate, inequilateral at base, truncate or subauricu- late acroscopically, cuneate basiscopically, the margins obtusely but prominently serrate or bi- serrate. Veins indistinct or obscure, simple to once- forked. Sori linear or narrow-elliptic, medial to inframedial. Indusia firm to subcartilaginous, yel- lowish to brown. Thus far known in Peru from one collection, on trees in mountains, San Martin. Colombia; Venezuela; Ecuador; Peru. The only collection seen from Peru perfectly matches the type specimens. The species seems to be intermediate between A. auriculatum and A. salicifolium, and all three could be synonymous. San Martin: Cerros de Campana, Spruce 4676 (us). 55. Asplenium auriculatum Sw., Kongl. Vetensk. Acad. Handl. 1817: 64. 1817, not (Thunb.) Kuhn, 1868. TYPE: Brazil, Minas Gerais, Freyreis (holotype, presumably s; isotype, BM; photos, F & us of BM). Plants epiphytic, sometimes terrestrial, rarely epipetric, the roots densely golden-tomentose. Stem erect, provided with clathrate, ovate to lanceolate scales, these 2-4(-5) mm long, acute, orange to light brown, usually with sparse, long, orange cilia. Leaves 1 -pinnate, 12-30(-50) cm long, 3.5-9 cm broad. Petiole 4-1 5(-l 8) cm long, dull green, gray, or reddish brown, rarely atropurpureous, non- alate, sparsely scaly and often golden-tomentose at base. Lamina thin- to firm-herbaceous, lanceo- late to oblong, abruptly reduced to a narrow, ser- rate-ligulate or subcaudate apex, not or scarcely reduced at base. Rachis dull reddish to greenish brown, marginate or narrow-alate, lacking prolif- erous buds. Pinnae 9-15 pairs, larger ones 3-6 cm long, 1-1.8 cm broad, approximate to subdistant, acute or subacute, inequilateral at base, truncate to subcordate and often overlapping the rachis ac- roscopically, cuneate to excavate basiscopically, the margins serrate or crenate- serrate. Veins sim- ple or 1 -forked, indistinct to obscure. Sori infra- medial, linear or elliptic. Indusia firm, fleshy, opaque, yellow, green, or light brown. In deep forests, commonly on tree trunks or branches, sometimes on wooded slopes or the for- est floor, rarely on mossy rocks, 500-2700 m, Ca- jamarca and Amazonas to Cuzco and Madre de Dios. Mexico to Panama; West Indies; Colombia to Trinidad, south to Bolivia and Brazil. This is closely related to, and possibly synon- ymous with, A. salicifolium and A. hastatum. See treatment of the former for further discussion. Cajamarca: Prov. Santa Cruz, ENE of Monteseco, Santisteban & Guevara 22 (F, GH, HUT, uc). Amazonas: Prov. Bagua, E of La Peca, Barbour 2492 (F, MO, uc). San Martin: Prov. Mariscal Caceres, Dist. Tocache Nue- vo, 4 km from Puerto Pizana, Schunke V. 4866 (F, us). Huanuco: Tingo Maria, Stork & Hort on 9469 (F, GH, uc, us). Pasco: Prov. Oxapampa, Dist. Oxapampa, Rio San Alberto, Leon 629 (F). Junin: Chanchamayo Valley, C. Schunke 178 (F), 1369(p). Cuzco: Quillabamba-Potrero, Coronado 1 19(GH, uc, us). Madre de Dios: Prov. Manu, Atalaya, near Hacienda Amazonia, Foster & Wachter 7414 (F, MO). 56. Asplenium cuneatum Lam., Encycl. 2: 309. 1786. TYPE: Locality and collector undesig- 44 FIELDIANA: BOTANY nated (holotype, P, Herb. Jussieu 1 725; photo, us). Plants epiphytic, rarely terrestrial, outside Peru sometimes epipetric. Stem stout, repent, provided with linear, attenuate, dark brown scales, these 2- 4 mm long, 0.3 mm broad, lustrous, broadly and conspicuously clathrate. Leaves fasciculate, 2-3- pinnate, 1 5-35(— 45) cm long, 4-1 2(-l 5) cm broad. Petiole 7-20 cm long, dull or sublustrous, reddish brown. Lamina chartaceous, subdeltate- to ob- long-lanceolate, reduced to a pinnatifid apex, not or scarcely reduced at base, glabrous. Rachis dark reddish brown, glabrous, but this and the costae sparsely to moderately provided with dark, tor- tuous, filiform scales, essentially nonalate, lacking proliferous buds. Pinnae 12-25 pairs, approxi- mate or subdistant, somewhat ascending, short- stalked, broad- or narrow-deltate. Pinnules 0.6-2 cm long, irregularly ovate to obovate, obtuse, lobed or with 1-2 pairs of segments, these obovate to subflabellate, broadly and obtusely dentate toward the apex. Veins indistinct, dichotomously branched, commonly forked. Sori 3-5 on the seg- ments, borne at a very acute angle to the costule. Indusia light brown or yellowish, firm. In dense forests, on trunks and branches of trees, rarely on forest floor, 100-900 m, San Martin and Loreto to Pasco, Puno. West Indies; Guatemala to Costa Rica; Colom- bia to the Guianas; Ecuador; Peru; Brazil; Africa; Polynesia. San Martin: Prov. Mariscal Caceres, Dist. Uchiza, E of Nuevo Progreso, Schunke V. 3195 (F, GH, us). Dist. Tocache Nuevo, Puerto Pizana, Schunke V. 6946 (F, MO, uc). Loreto: Bank of Rio Santiago, above Pongo de Man- seriche, Mexia 6222 (F, GH, MO, uc, us). Huanuco: Prov. Huanuco, Las Palmas, Asplund 12635 (s). Pasco (as Ju- nin): Puerto Bermudez, Killip & Smith 26518 (GH, us). Puno: Prov. Carabaya, Palmera, Vargas 16161 (GH). 57. Asplenium serra Langsd. & Fisch., Icon. fil. 16, t. 19. 1810. TYPE: Brazil, Santa Catarina, Langsdorff (holotype, LE; isotype, B, Herb, mild. 19880; photos, GH & us of B). Asplenium serra var. remotum Kunze, Linnaea 9: 64. 1834. TYPE: Peru, Pampayacu, Poeppigin 1829 (holotype, LZ, destroyed). Plants terrestrial, occasionally epiphytic or epi- petric. Stem short- to long-creeping, abundantly provided with lanceolate, attenuate, brown or gray- brown, iridescent, scales, these 3-5 mm long, con- spicuously clathrate. Leaves 1 -pinnate, approxi- mate to remote, to 1 m long and 30 cm broad. Petiole to 50 cm long, dull or slightly lustrous, dark brown to atropurpureous, sparsely (rarely co- piously) scaly. Lamina chartaceous (occasionally firm-herbaceous), oblong-lanceolate, terminating in a distinct, conform apical segment (this rarely with a basal lobe), not or scarcely reduced at base. Rachis nonalate, lacking proliferous buds. Pinnae 8-16 pairs, well spaced, spreading to ascending, stalked, long-attenuate, cuneate at base (often more broadly so acroscopically), margins serrate or bi- serrate (at least at apex). Veins 2-3-forked, as- cending at very acute (mostly 10-20°) angles, often subparallel to the costa. Sori inframedial, not ex- tending into the marginal V? of the pinna. Indusia yellowish or brown, long and narrow, often ob- scured by the sporangia at maturity. In forests, thickets, and edges of clearings, in Peru commonly on the forest floor, but occasion- ally low on tree trunks or among rocks, 100-3650 m, Cajamarca to Loreto, south to Ayacucho and Puno. West Indies; Mexico to Panama; Colombia and Venezuela south to Argentina and Paraguay. This species is commonly found as an epiphyte from Mexico to Ecuador, yet in Peru and the West Indies it is most often terrestrial. Also, the petiole and rachis commonly bear only a few, scattered, filiform scales, but several collections in Peru have rachises copiously beset with linear, often atten- uate, clathrate scales, to 5 mm long: C. Schunke 10, 137, and 655 (F) and Killip & Smith 24615 (F, GH, us). Curiously, all plants were collected be- tween 1927 and 1929 in Junin near Schunke's Ha- cienda, at about 1500 m. It is likely they come from the same site. Other than density of scales, they are typical A. serra. Cajamarca: Prov. Cutervo, "Petroglifos de Llipa," Mostacero et al. 1747 (F, GH, HUT, MO, uc). Amazonas: Prov. Chachapoyas, Calla-Calla slopes, near km 411- 4 1 6 of Leimebamba-Balsas road, Wurdack 1343 (F, GH). San Martin: Zepelacio, near Moyobamba, Klug 3671 (F, GH, MO, us). Loreto: Mishuyacu, near Iquitos, Klug 378 (F, us), 7509 (F, us). Huanuco: Prov. Huanuco, Mirador, road Acomayo to Chinchao, Mexia 7758 (GH, MO, uc, us). Pasco (as Junin): Pichis Trail, Enenas, Killip & Smith 25649 (F, us). Junin: Schunke Hacienda, above San Ra- mon, Killip & Smith 24615 (F, GH, us). Ayacucho: Prov. La Mar, between Tambo San Miguel, Ayna, and Haci- enda Luisiana, Dudley 11888 (GH). Cuzco: Prov. Pau- cartambo, Pillawata, Vargas 16714 (GH). Puno: Churu- mayo, Soukup 867 (F). TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 45 58. AspleniumdimidiatumSw.,Prodr. 129. 1788. TYPE: Jamaica, Swartz (holotype, s; isotypes, B, Herb. Willd. 19905, UPS, Herb. Thunb. 24798; photos, us of s, GH & us of B). Plants terrestrial, rarely epiphytic or epipetric. Stem erect or short-decumbent, abundantly pro- vided with narrowly clathrate, linear or filiform scales, these 5-8 mm long, with a tortuous, uni- costate tip, dark brown to blackish, often with a fine, lighter brown margin. Leaves 1 -pinnate, sub- caespitose, 1 8-50 cm long, 6-12 cm broad. Petiole to 20 cm long, dull, dark brown to blackish, abun- dantly clathrate-scaly at base, usually less so dis- tally. Lamina chartaceous or subcoriaceous, grad- ually reduced to a pinnatifid apex, or to a nonconform apical segment, not or scarcely re- duced at base. Rachis nonalate, lacking proliferous buds, moderately to amply provided with narrow, attenuate, blackish, clathrate scales. Pinnae 6-12 pairs, well-spaced, sessile or subsessile, lanceolate, subdimidiate, lacking a costa, apex attenuate, in- equilateral at base, cuneate acroscopically, excised basiscopically, margins serrate to deeply and ir- regularly erose-lacerate, abaxial surface and veins sparsely to moderately filiform-scaly. Veins re- peatedly subdichotomous, subparallel or diverg- ing from each other at 5-10° angles. Sori linear, crowded, at maturity usually covering pinnae nearly to margin. Indusia narrow, firm, light or yellow-brown, entire. On forest floor or in moist ravines, rarely on rocks or bases of tree trunks, 650-2000 m, Ama- zonas, San Martin, Junin, Cuzco. This has been confused with A. erosum L. of the Greater Antilles; however, pinnae of the latter are distinctly stalked and costate, and margins are more regularly and evenly serrate. Amazonas: Prov. Bagua, canyon of Rio Utcubamba on Chachapoyas road, Hutchison and Wright 3793 (GH, uc). San Martin: Near Tarapoto, Spruce 4753 (BR, GH, us). Junin: Prov. Tarma, Chanchamayo, Esposto (USM). Chanchamayo Valley, Puente Paucartambo to La Mer- ced, Gentry et al. 39823 (F, MO). La Merced, Soukup 3407 (GH, us). Cuzco: Sahuayacu, Biies 843 (us). Prov. La Convention, Rio Apurimac, 20 min. float below Puerto Capiro, Davis et al. 1291 (F, uc). 59. Asplenium praemorsum Sw., Prodr. 130. 1788. TYPE: Jamaica, Swartz (holotype, s; isotype, UPS, Herb. Thunb. 24834; photo, us of s). Asplenium nigricans Kunze, Linnaea 9: 69. 1834. TYPE: Peru, Huanuco, Sierra de Huanuco, "ad vicum Ambo," Poeppig, in Apr. 1830 (isotype, w; frag., BM; photos, BM, F & us of w). Plants epipetric, epiphytic, or sometimes ter- restrial. Stem erect or decumbent, abundantly pro- vided with narrowly clathrate, linear to filiform scales, these 3-7 mm long, usually with a tortuous, unicostate tip, dark brown to blackish, sometimes with a fine, lighter brown margin. Leaves 1-pin- nate-pinnatisect (in America), fasciculate, 12-60 cm long, 3-14 cm broad. Petiole somewhat shorter than the lamina, dull to slightly lustrous, dark brown to blackish, abundantly clathrate-scaly at base, usually less so distally. Lamina subcoria- ceous, tapering to a pinnatifid apex, not or scarcely reduced at base. Rachis nonalate, lacking prolif- erous buds, moderately to copiously scaly as on the petiole. Pinnae 7-18 pairs, approximate or subdistant, sessile, or proximal ones very short- stalked, deeply lobed (or sometimes fully pinnate at base), the segments cuneiform to obovate, their apices often lacerate, abaxial surface and veins fi- liform scaly. Veins indistinct, subparallel or di- verging from each other at a 5-10° angle. Sori linear, crowded. Indusia narrow, thin, light or yel- low-brown, entire. Common, in forests, usually on rocks and in crevices or rocky cliffs, often on trunks and branches of trees, or less frequently on the forest floor, 600-3700 m, Cajamarca to San Martin, south to Arequipa and Puno. West Indies; Mexico to Panama; Venezuela and Colombia to Brazil and Bolivia; Argentina; Ga- lapagos Islands. This variable species has been recognized for a long time under a number of names, from such diverse areas as Jamaica, Arabia, and South Af- rica. Recently, Braithwaite (J. Linn. Soc., Bot. 93: 349. 1986) proposed that all, including A. prae- morsum, should be included under the oldest name, A. aethiopicum (Burm.) Bech. African species have leaves no less than 2-pinnate, and two of the sub- species recognized by Braithwaite are 2-pinnate- pinnatifid. In the New World, and particularly in Peru, leaves are often only 1 -pinnate-pinnatisect, with the division of pinnae sometimes confined to one or two basal lobes; so perhaps this taxon should be recognized as an American variety. An exhaustive study of the American representatives of this species complex, such as Brathwaite's in Africa, is needed before the problem is resolved. For purposes of this treatment, however, A. prae- morsum is tentatively maintained as a distinct spe- cies. 46 FIELDIANA: BOTANY Cajamarca: Proc. Contumaza, near Guzmango, Sa- gdstegui 7703 (F, GH, MO). Amazonas: Prov. Chacha- poyas, slopes of Cano Santa Lucia, Wurdack 732 (F, GH, uc). La Libertad: Prov. Trujillo, Cerro Campana, Sa- gdstegui 6193 (GH, MO). San Martin: Dist. Huallaga, Valley of Rio Apisoncho, Hamilton & Holligan 940 (us). Ancash: Callejon de Huallas, 8 km SSE of Yungay, Tryon & Tryon 6548 (F, GH, us). Huanuco: Muna, Macbride 3931 (F, us). Lima: Prov. Canta, Muller & Lopez 12905 (GH). Junin: Satipo, Pichanaki, rodal del Proyecto Peru- ano-Aleman, Leon 236 (F, uc). Ayacucho: Prov. La Mar, Ayna, Ldpez & Soukup 15089 (GH). Apurimac: Prov. Andahuaylas, W of Pincos, Stork & Norton 10714 (F, uc, us). Cuzco: Machu Picchu, Leon 461 (F, USM). Are- quipa: Lomas de Arequipa, Coronado 32 (GH, uc, us). Puno: Prov. Carabaya, below Ollachea, Vargas 6906 (uc, us), 75997 (GH). 60. Asplenium lividum Kuhn, Linnaea 36: 100. 1 869. TYPE: Venezuela, Aragua, Colonia To- var, Fendler 156 (holotype, B; isotypes, F!, MO, us; photo, us of B). Plants terrestrial. Stem short-creeping, provided with ovate, narrowly clathrate scales, these 0.5-1 mm long, orange, acute to acuminate. Leaves 1-pinnate-pinnatisect, approximate, 25—45 cm long, 5-8 cm broad. Petiole somewhat shorter than the lamina, dull, reddish or grayish brown, non- alate, naked or with a few appressed scales like those on the stem. Lamina chartaceous, tapering to a pinnatifid apex, not or scarcely reduced at base. Rachis nonalate, lacking proliferous buds, this and the pinnae abaxially with scattered, ap- pressed, acuminate scales to 1 mm long. Pinnae 14-20 pairs, larger ones 3-5 cm long, subdistant, short-stalked, pinnatisect (to nearly pinnate at base), the 3-6 pairs of segments linear to cunei- form, their apices often lacerate. Veins indistinct, subparallel or diverging from each other at 5-10° angles. Sori linear, crowded. Indusia narrow, thin, light or yellowish brown, entire. Thus far known in Peru from one collection, in damp, shaded woods, about 2100 m, Huanuco. Venezuela; Peru; Africa. This has been confused with A. praemorsum, as the two species are nearly identical in lamina ar- chitecture, texture, veins, and sori; yet they differ significantly in habit and indument. The stem of A. lividum is short-creeping, although somtimes not obviously so (in the manner of A. laetum), and bears only a few approximate leaves. Minute, or- ange, mostly ovate-acuminate, scales are clustered at the stem apex and also are sparsely scattered along the axes and veins abaxially. Laminar scales are only microscopically evident. In marked con- trast are the erect (rarely decumbent) stem and densely fasciculate leaves of A. praemorsum, yet the latter differs even more conspicuously in the long, dark, filiform, usually copious scales that spread from the rachis, these quite evident to the naked eye. Although dozens of specimens of A. praemor- sum were examined during the course of this study, only the two sheets of A. lividum cited here have been found. However, Andean collections in var- ious herbaria should be searched for the latter. Any specimen of A. praemorsum is suspect on which dark, spreading lamina scales are not macroscop- ically evident. Huanuco: Muna, Bryan 417 (F, GH). 61. Asplenium purdieanum Hooker, Icon. PI. 10: t. 938. 1 854. TYPE: Colombia, Sierra de Santa Marta, La Fundacion (erroneously reported by Hooker as Jamaica). Purdie (holotype, K; isotype, BM; frag., NY). Plants epipetric or terrestrial. Stem stout, erect, provided with lanceolate, attenuate, gray-brown, iridescent scales, these 3-6 mm long, conspicu- ously clathrate. Leaves 1 -pinnate, caespitose, to 50 cm long. Petiole to 25 cm long, reddish brown to blackish, dull or slightly lustrous, densely scaly as on the stem. Lamina thin- to firm-herbaceous, with 1-2 pairs of pinnae and a conform apical segment, or in juvenile leaves simple and cordate to sagittate. Rachis not or scarcely alate, moder- ately scaly. Pinnae sessile or short-stalked, elliptic or broadly lanceolate, acute to short-attenuate, cu- neate at base, margins entire. Veins obscure or indistinct, anastomosing beyond the middle of the pinna to form elongate areoles, these lacking in- cluded veinlets. Sori in a single row on either side of the costa, spreading at acute angles. Indusia long and narrow, thin, often hyaline, shrivelling at ma- turity, the margin entire. Rare throughout the range, thus far known in Peru from one collection in San Martin. Elsewhere, in forests, on rocky soil or in crevices of limestone, 100-700 m, Honduras; Colombia; Venezuela; Peru. This is easily distinguished from other species in the genus by the few pinnae and conform apical segment, the densely scaly petiole, and the anas- tomosing veins. San Martin: Near Tarapoto, Spruce 4754 (BR, K). TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 47 62. Asplenium theciferum (HBK.) Mett., Ann. Sci. Nat. Bot. 2: 227. 1864. Figure le. Davallia thecifera HBK., Nov. Gen. Sp. 1: 23. 1815. TYPE: Venezuela, in monte Saraquen, Hwnboldt & Bonpland (holotype, P). Loxoscaphe thecifera (HBK.) Moore, Index fil. 302. 1861. Plants epiphytic, occasionally epipetric. Stem erect or decumbent, provided with ovate to lan- ceolate, bicolorous scales, these 2-3 mm long, gray- brown to blackish, with narrow lighter brown mar- gins. Leaves 2-pinnate to 2-pinnate-pinnatifid, fas- ciculate, 7-25 cm long. Petiole as long as or slightly longer than the lamina, stout and fleshy, dull greenish to yellowish, moderately to sparsely pro- vided with dark, conspicuously clathrate scales similar to those of the stem. Lamina subcoria- ceous, deltate- to elliptic-ovate, tapering to a pin- natifid apex. Rachis greenish, not or slightly alate, inconspicuously scaly. Pinnae 5-8 pairs, stalked, divided into linear to clavate segments or pin- nules, these sometimes incised deeply into tertiary segments. Veins completely obscured within the thickened tissue. Indusia cuplike or pocketlike, borne singly along the acroscopic side of ultimate segments, near their tips, with sporangia deeply immersed. Known thus far in Peru from a single specimen from Cuzco, 900 m (cited below). Hispaniola; southern Mexico; Costa Rica; Pan- ama; Colombia; Venezuela; Ecuador; Peru. Cuzco: Mercedes, Bties 1736 (us). Comments Asplenium denudatum Kuhn, Linnaea 36: 93. 1 869. TYPE: "Andes Peruvianae" (collector undes- ignated, B?). This was originally described by Mettenius, but not published until Kuhn's emendation of it in 1869. Judging from Kuhn's detailed description, it is probably A. monanthes. Stem scales are said to be concolorous, reddish brown, and clathrate, the lamina 6 in. long, and petioles lustrous and castaneous, so it is likely to be var. castaneum. Positive identification awaits study of the type. Asplenium discolor Kunze, Linnaea 9: 65. 1835. Kunze identified a collection by Poeppig as this species ("Peru, on the upper Huallaga, near Mission Tocache, Poeppig, July, 1830"). A search of several herbaria (B, BM, K, P, w) proved fruitless, and the original description is too incomplete to assign specimens to this name. The species was said to be a foot long, with petiole 4 in. long, rachis subalate, purplish and puberulent, and pinnae remote and obtusely serrate. These characters are common to many species of As- plenium. Asplenium otites Link, Fil. spec. 2: 60. 1 833. TYPE: Cultivated, presumably from a South Amer- ican plant (Link, in Fil. spec. 91. 1841, cited "Brasilia"), Hort. Mus. Bot. Berol., Cat. No. 053514 (holotype, B!; photos, F, us). It is difficult to place this name, because the type is similar to several other South American species, yet the specimen is too incomplete for definite comparison. It consists only of a single lamina with 6 cm of petiole, it lacks a stem, and the lamina is broken off near the tip. It has been compared to A. pulchellum, but its pinnae are inequilateral only at the base and bear 6-8 pairs of veins, which are mostly forked acroscopically. Pinnae of the latter are inequilateral throughout the proximal half, and veins are fewer and unbranched above the pinna base. It is probably more closely related to A. drepanophyllum, which does have more nu- merous and forked veins, but the latter has twice the number of pinnae and the lamina is more at- tenuate. The question is not likely to be resolved with certainty. Asplenium trapezoides Sw., Syn. fil. 76. 1806. TYPE: Peru (Junin), Tarma, Lagasca (holo- type, s; isotypes, B, Herb. Willd. 19869, P; photos, F & GH of B). The locality cited is undoubtedly incorrect. It certainly should be Chile, because the specimen is clearly the Chilean species A. trilobum Cav., and the Willdenow Herbarium sheet was so identified by Hieronymus. This rare fern has not been re- ported outside Chile for nearly 200 years, since the original citation of the type of A. trapezoides. The fasciculate leaves are subtrapezoidal, deeply crenate or with a pair of basal lobes. They lack a distinct midrib, and veins are flabellately branched from the lamina base. Admittedly, this is a small and inconspicuous fern, but its leaf shape is cu- rious and distinctive, and if it is indeed in Peru it certainly should have been rediscovered by now. 48 FIELDIANA: BOTANY Asplenium weberbaueri Hieron.. Hcdwigia 60: 2 1 1 . 1918. TYPE: Peru (Puno), Tambo Isilluma, on the way from Sandia to Chanchasmayo, Weberbauer 1198(B?). This plant was described as having simple, lin- ear leaves to 16 cm long and 1.5 cm broad that taper gradually at both ends, serrate or dentate margins, short (4 cm) petioles, veins spreading at 20-25° angles, and elongate-deltate stem scales. In characters of leaf size and shape, it is similar to Asplenium angustwn, but in lamina margin and stem scales it seems nearer to A. serratum. In Hieronymus's discussion, he refers to hook-shaped spines (stiff scales?) along the midrib. Both the above-named species have dark scales along the midrib, but these cannot be termed "hook-shaped." A canvass of European herbaria failed to locate the type, so the name cannot be precisely placed at this time. Family 19. DAVALLIACEAE Davalliaceae Frank, Syn. Pflanzensk. (Leunis), ed. 2, 3: 1474. 1877. TYPE: Davallia Sm. NephrolepidaceaePic.-Ser., Webbia 29: 8. 1974. TYPE: Nephrolepis Schott. Stem erect, decumbent, or long-creeping, mod- erately stout to slender, more or less indurated, bearing scales. Leaves ca. 1 5 cm to 3 m long, entire to usually pinnate, circinate in vernation, petiole without stipules. Sori roundish to somewhat elon- gate, rarely linear, abaxial, near or back of the margin, or on an intramarginal commissure, usu- ally not paraphysate, usually indusiate. Sporan- gium with a 2-rowed stalk below its apex, with a usually long stalk, the vertical annulus interrupted by the stalk. The Davalliaceae are a family of about 1 0 or fewer genera of the Old World, with Nephrolepis also in America. Most species are epiphytes with a dorsiventral stele and leaves that are articulate near the base of the petiole. Nephrolepis, the only genus in America, is in- cluded in the family with some uncertainty. The tuberculate spores and staurocytic stomata suggest a closer alliance with the davallioid ferns than with other groups. I. Nephrolepis Nephrolepis Schott, Gen. fil. no. 3. 1834. TYPE: Nephrolepis exaltata (L.) Schott (Polypodium exaltatum L.). Figure 2. Stem erect or decumbent, bearing scales, freely stoloniferous. Leaves monomorphic or slightly di- morphic with the fertile pinnae narrower than the sterile. Lamina 1 -pinnate, the pinnae articulate, entire or nearly so, glabrous, pubescent, and (or) scaly, the scales usually dissected, with hydathodes adaxially that usually have a thin, whitish, circular secretion, veins free. Sori roundish to lunate, borne on the vein ends, not paraphysate, covered by an orbicular to lunate indusium. Spores ellipsoidal, monolete, the surface irregularly tuberculate to ru- gose. Nephrolepis is a tropical and subtropical genus of perhaps 20 species, with five native species in America and four in Peru. The species of Nephrolepis are not well under- stood and, accordingly, current accounts of the genus in tropical America and other tropical regions often differ in the application of names and in the primary characters employed for identification. The following treatment is based primarily on Pe- ruvian materials, and differences with other mod- ern treatments do not necessarily imply alterations in the taxonomy of American species in other regions. Several species and their cultivated variations are commonly maintained in gardens and may escape to more or less native habitats, either through spore dispersal or by the growth of dis- carded cultivated plants. Some of the many cul- tivated variations (cultivars) have rarely been col- lected in Peru. These usually have pinnatifid to several times pinnate pinnae, forked pinnae or ra- chises, or nearly orbicular pinnae. They are not included in the treatment of the native and ad- ventive species. The key to species emphasizes the primary char- acters of the species, while uncommon variations are mostly mentioned in the descriptions. References NAUMAN, C. E. 1981. The genus Nephrolepis in Florida. Amer. Fern J., 71: 35-40. SMITH, A. R. 1981. Nephrolepis, pp. 151-154. In Flora of Chiapas, part 2, Pteridophytes. TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 49 50 FIELDIANA: BOTANY California Academy of Sciences, San Fran- cisco. TRYON, R. M., AND A. F. TRYON. 1982. Ne- phrolepis, pp. 656-662. In Ferns and Allied Plants. Springer- Verlag, New York. Key to Species of Nephrolepis \ . Indusia orbicular to orbicular-reniform, with a narrow to rather open sinus; mature sporangia pro- jecting all around the indusium b b. Pinnae short- or very short-stalked; costa definitely pubescent adaxially, or if glabrous then the base of the pinnae equilateral or nearly so c c. Base of the pinnae equilateral or nearly so, sometimes biauriculate, or slightly and broadly auriculate only on the acroscopic side; costa adaxially with long, often flexuous, trichomes and usually some dissected scales; stem scales light brown to brown, more or less spreading 1 . N. biserrata c. Base of the pinnae, or most of them, inequilateral, the acroscopic side narrowly auriculate; costa adaxially with short to rarely moderately long, stiff trichomes; stem scales dark brown to blackish, appressed 2. N. multiflora b. Pinnae sessile; costa glabrous or nearly so adaxially; base of the pinnae inequilateral, the acroscopic side broadly and acutely to subacutely auriculate 3. N. rivularis a. Indusia reniform, with a broad sinus, to lunate; mature sporangia projecting from the free portion of the indusium d d. Fertile pinnae with the basal sorus on the acroscopic side and the basal sorus on the basiscopic side ca. equally distant from, or close to, the base of the pinna, or many of them so; the distal pinnae, but not necessarily the apical ones, rounded, usually broadly so, at the base on the basiscopic side 4. N. cordifolia d. Fertile pinnae with the basal sorus on the acroscopic side much closer to the base of the pinna than the basal sorus on the basiscopic side, or most of them so; especially the distal pinnae cuneate at the base on the basiscopic side to sometimes slightly rounded 5. N. pectinata 1 . Nephrolepis biserrata (Sw.) Schott, Gen. fil. no. 3. 1834. Aspidium biserratum Sw., J. Bot. (Schrader) 1800(2): 32. 1802. TYPE: Mauritius, Grondahl (Grondal) (holotype, Herb. Sw., s!). Tectaria fraxinea Cav., Descr. pi. 250. 1802. TYPE: Peru. Obragillo, Nee, MA, seen by C. Chr. (Dansk Bot. Ark. 9(3): 15. 1937 and referred here). Stem scales light brown to brown, more or less spreading. Pinnae short-stalked, pubescent to rarely glabrous abaxially. the costa pubescent adaxially with long, often flexuous, trichomes and usually some dissected scales, rarely the trichomes short or the costa glabrous, the base equilateral or nearly so, sometimes biauriculate, or slightly and broadly auriculate only on the acroscopic side, broadly rounded to broadly cuneate on the basiscopic side. Lamina of fertile leaves ca. 10-30 cm broad. Basal sori on the acroscopic and basiscopic sides equally close to the base of the pinna. Indusium orbicular, with a narrow sinus, the mature sporangia pro- jecting all around the indusium. Epiphytic or usually terrestrial, along roadsides, in secondary growth, along forest borders, and in tall forests, 1 00-900 m, Cajamarca and Amazonas to Ayacucho. Tropical America; Old World. The lamina of this species is usually consider- ably broader than that of other Peruvian species. Even in rather small leaves it is rarely less than 10 cm broad, and in large leaves, which may be up to ca. 3 m long, it is commonly 30 cm broad or more. Small plants may have leaves similar to those of N. multiflora but that species has the pin- nae inequilateral at the base, dark, appressed stem FIG. 2. Nephrolepis pectinata: a, habit; b, fertile pinna. Nephrolepis cordifolia: c, fertile pinna. Nephrolepis mul- tiflora: d, pinna base. Nephrolepis rivularis: e, pinna base. (Adapted from Stolze, Ferns and Fern Allies of Guatemala, 1981.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 51 scales, and usually has the pinnae abaxially fibril- lose-scaly, especially at the base. Cajamarca: Prov. Jaen, Quebrada Tabaconas, Ferrey- ra & Sanchez 19646 (USM). Amazonas: Prov. Bagua, Chi- riaco-Puente Venezuela, Barbour 4331 (MO). La Poza, Rio Santiago, Berlin 3701 (MO). San Martin: NW of San Martin, Soukup 5218 (GH). Prov. San Martin, Vaca Pozo, Woytkowski 35136 (MO). Prov. San Martin, Ekin, Woyl- kowski 35230 (MO, uc). Loreto: Prov. Maynas, Rio Itaya, Tryon & Tryon 5176 (BM, F, GH, us, USM). Prov. Maynas, 3 km S of Iquitos, Tryon & Tryon 5163 (BM, F, GH, u, us, USM). Huanuco: Prov. Pachitea, Bosque Nacional de Iparia, J. Schunke V. 1735 (F, GH, us). Ayacucho: Kim- pitiriki, Rio Apurimac, Killip & Smith 22856 (F, us). 2. Nephrolepis multiflora (Roxb.) Morton, Contr. U.S. Natl. Herb. 38: 309. 1974. Figure 2d. Davallia multiflora Roxb., Calcutta J. Nat. Hist. 4: 515, t. 31 (left), 1844. LECTOTYPE: by Morton, he. cit., India, Roxburgh (BR; frag, and photo, us). Stem scales dark brown to blackish, appressed. Pinnae short- or very short-stalked, fibrillose-scaly abaxially, especially toward the base and (or) pu- bescent, the costa adaxially with usually short and stiff, to moderately long, trichomes, rarely with some dissected scales or long trichomes, the base inequilateral, narrow auriculate on the acroscopic side, broadly rounded to slightly auriculate on the basiscopic side. Lamina of fertile leaves ca. 8-20 cm broad. Basal sori on the acroscopic and basi- scopic sides equally close to the base of the pinna. Indusium orbicular to orbicular- reniform, with a narrow to rather open sinus, the mature sporangia projecting all around the indusium. Terrestrial, in primary forests and also disturbed areas, 120-2100 m, Cajamarca and Loreto. Paleotropics; sporadically adventive in tropical America. This species is probably more frequent in Peru than the few collections indicate. It is distin- guished by the abaxially fibrillose-scaly pinnae and the stem scales that are dark and appressed. It has perhaps been confused with another paleo tropical species, Nephrolepis hirsutula (Forster) Presl, which has the adaxial side of the costa moderately to densely invested with dissected scales, rather than pubescent as in N. multiflora. The species N. hir- sutula is not known to be adventive in Peru. Cajamarca: Provicutervo, San Andres, Quiroz 2840 (F). Loreto: Prov. Maynas, Puerto Almendras, Vasquez & Jaramillo 1415 (MO). Prov. Maynas, Quistococha, Vasquez & Jaramillo 4079 (F). Iquitos, Mitller & Miiller 2508 (GH). Quistococha, Gentry & Ayala 15531 (us). 3. Neprolepis rivularis (Vahl) Krug, Bot. Jahrb. Syst. 24: 122. 1897. Polypodium rivulare Vahl, Eclog. amer. 3: 51. 1807. TYPE: West Indies, Montserrat, Ryan (holotype, c). Stem scales light to dark brown, appressed to spreading. Pinnae sessile, glabrous to slightly pu- bescent abaxially, and (or) with a few dissected scales, rarely minutely pubescent, the costa gla- brous or nearly so adaxially, the base inequilateral, broadly acutely or subacutely auriculate on the acroscopic side, cuneate to rounded on the basi- scopic side. Lamina of fertile leaves ca. 8-15 cm broad. Basal sori on the acroscopic and basiscopic sides ca. equally close to the base of the pinna, or closer on one side than the other. Indusium or- bicular, with a narrow sinus, the mature sporangia projecting all around the indusium. Epiphytic, or usually terrestrial, in disturbed vegetation, secondary forests, or primary forests, 100-900 m, San Martin to Puno. Tropical America. This is a distinctive species, especially in its character of sessile pinnae. Contrary to some au- thors, it appears that the combination Nephrolepis rivularis was validly published by Krug, although the basionym was not cited. San Martin: Soritor, Woytkowski 6246 (GH, us). Lo- reto: Gamitanacocha, Rio Mazan, Schunke 279 (GH, uc). Prov. Maynas, Rio Nanay, McDaniel 13685 (GH), Hick- ok 662 (GH). Quista Cocha (Quistococha), near Iquitos, McDaniel 1 1739 (GH). Pasco: Prov. Oxapampa, Palcazu valley, Foster 9503 (MO). Cuzco: Prov. Paucartambo, Kosnipata-Pilcopata, Vargas 14750 (GH). Madre de Dios: Prov. Tambopata, Tambopata Nature Reserve, Barbour 4783 (MO), 5124 (F, MO). Puno: San Gavan (Gaban), Lechler 2515 (GH, K). 4. Nephrolepis cordifolia (L.) Presl, Tent. Pterid. 79. 1836. Figure 2c. Polypodium cordifolium L. Sp. pi. 2: 1089. 1 753. TYPE: Petiver, Pteri-graphia amer., t. l,f. 11 (not seen in Petiver, Hort. Sice, in Herb. Sloane, BM). Aspidium pendulum Raddi, Opusc. Sci 3: 289. 1819. TYPE: Brazil, Raddi (holotype, FI; isotype, K). Nephrolepis pendula (Raddi) J. Sm., J. Bot. (Hooker) 4: 197. 1841. 52 FIELDIANA: BOTANY Stem scales light to dark brown to blackish, more or less appressed. Pinnae short- to very short- stalked, essentially glabrous, the costa essentially glabrous adaxially, the base inequilateral or some- what inequilateral, slightly to definitely auriculate on the acroscopic side, rounded to broadly round- ed or sometimes slightly auriculate on the basi- scopic side. Lamina of fertile leaves ca. 3-10 cm broad. Basal sorus on the acroscopic and basi- scopic sides ca. equally distant from, or close to, the base of the pinna, or most of them so. Indusium reniform, with a broad sinus, to lunate, mature sporangia projecting from the free portion of the indusium. Epiphytic, rupestral, or terrestrial, rocky places, rarely on cliffs, cloud forests, forest borders, pri- mary forests, shrubby hillsides, or on fallen trunks and branches, 125-2800 m, Cajamarca and Ama- zonas south to Puno. Tropical America; Old World. The Petiver illustration is not adequate for the purposes of accurate identification. It is said to be copied from Plumier, Traite foug. amer., t. 71, from Hispaniola, which is more clearly, and usu- ally identified as, Nephrolepis cordifolia. Some epiphytic plants have very long, pendent leaves, up to 3 m and perhaps longer. However, in comparison to the long leaves of Nephrolepis biserrata, the lamina of those of N. cordifolia is narrow, about 8-10 cm broad. Sometimes there is a single basal basiscopic sorus closer to the base of the pinna than the basal basiscopic sorus. At times, Nephrolepis pendula is adopted for this species because its application is certain. At other times, N. pendula is considered to be a separate species from N. cordifolia. At least in Peru the two do not seem to be separable since there are inter- mediate specimens. This and the following species Nephrolepis pec- tinata are the most widely distributed and com- mon species of the genus in Peru. A discussion of the differences between them is provided under N. pectinata. Cajamarca: Prov. Santa Cruz, Bosque de Monteseco, Sagdstegui et al. 12402 (F, MO). Prov. Cutervo, Cutervo- Socota, Lopez et al. 5316 (GH). Amazonas: Prov. Bon- gara, Laguna Pomacocha, Wurdack 883 (GH, us). Prov. Chachapoyas, entre Chachapoyas y Caclic, Lopez et al. 4350 (GH, MO). San Martin: Zepelacio, near Moyobam- ba, Klug 3736 (F, GH, us). Prov. Mariscal Caceres, Dis- trito Tocache Nuevo, J. Schunke V. 4337 (F, GH, us). Loreto: Above Pongo de Manserische, Mexia 6163 (F, GH, uc, us). Huanuco: Prov. Pachitea, Bosque Nacional de Iparia, J. Schunke V. 2940 (F, GH, us). Hacienda Paty, abajo de Carpish, Ferreyra 9353 (GH, USM). Pasco: Prov. Oxapampa, Oxapampa-Cerro de Pasco, D. Smith 4083 (GH). Prov. Oxapampa, N of Chequitavo, D. Smith 5108 (GH). Junin: Huacapistana, Tryon & Tryon 5452 (BM, F, GH, u, us, USM). Ucayali: Prov. Coronel Portillo, Bo- queron del Padre Abad, Ridoutl (GH, USM). Ayacucho: Prov. La Mar, eastern massif of the Cordillera Central, Dudley 11724 (GH), 11880 (GH, us). Cuzco: Prov. La Convention, Rio Apurimac, Davis et al. 1328 (F, GH). Prov. Urubamba, Macchu Picchu station, Sounders 1230 (F, GH). Puno: Prov. Sandia, Asalaya, Vargas 1483 (F). 5. Nephrolepis pectinata (Willd.) Schott, Gen. fil. no. 3. 1834. Aspidium pectinatum Willd., Sp. pi. 5: 223. 1810. TYPE:"exAiton"(holotype,//er6. Willd. 19753. B!; photo, GH). Figures 2a-b. Stem scales brown to blackish, more or less ap- pressed. Pinnae sessile to usually short-stalked, essentially glabrous abaxially, or rarely minutely pubescent, the costa essentially glabrous adaxially, to rarely partly minutely pubescent, the base in- equilateral, auriculate on the acroscopic side, es- pecially the distal pinnae cuneate to slightly round- ed on the basiscopic side. Lamina of fertile leaves ca. 2-4 cm broad. Basal sorus on the acroscopic side much closer to the base of the pinna than the basal sorus on the basiscopic side. Indusium reni- form, with a broad sinus, to lunate, mature spo- rangia projecting from the free portion of the in- dusium. Epiphytic, rupestral, or terrestrial, rocky places in woods, in primary forests, cloud forests, on stream banks, forest borders, in moist and rough pastures, on fallen trunks and branches, 200-2900 m, Cajamarca and Amazonas to Puno. Tropical America. Nephrolepis pectinata is distinguished by hav- ing, especially in the distal portion of the lamina, the pinnae with a cuneate (evenly tapering) base on the basiscopic side and having the basal sorus on the acroscopic side much closer to the base of the pinna than the basal sorus on the basiscopic side. Sometimes N. pectinata has the base of the pinna on the basiscopic side slightly rounded, and N. cordifolia may have a single basal acroscopic sorus closer to the base of the pinna than the basal basiscopic sorus. Nephrolepis occidentalisKun7.e(L\nnaea 18: 343. 1844) is usually recognized as a species, with a poorly developed stem and thin, long-triangular, crenate to shallowly lobed pinnae with the veins clearly evident abaxially. However, at least in South TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 53 America, there is intergradation between speci- mens with the characters of N. occidentalis and those of N. pectinata and it is not possible to rec- ognize the former as a distinct species in that con- tinent. Plants referred to N. occidentalis usually grow in especially damp and shaded situations, and they may represent an ecological variation (shade form) or an ecotype of N. pectinata. Cajamarca: Prov. Santa Cruz, Monteseco, Sagdstegui et al. 12402 (HUT). Amazonas: Prov. Bagua, Chiriaco to Puente Venezuela, Barbour 4401 (MO). San Martin: Prov. Lamas, Lamas, Belshaw 3441 (F, GH). Loreto: Above Pongo de Manserische, Mexia 6219 (F, GH, uc, us). Pu- mayacu, between Balsapuerto and Moyobamba, Klug 3246 (F, GH, us). Huanuco: Muna, Bryan 415 (F, GH). Prov. Huanuco, above Cayumba, Rio Huallaga, Mexia 8324 (F, GH, uc, us). Prov. Huanuco, Hacienda Exito, Mexia 8135 (F, GH, uc, us). Pasco: Prov. Oxapampa, between Oxapampa and Paucartambo, Smith & Pretel 1485 (F, GH). Junin: Prov. Satipo, S of Chequitavo, D. Smith 5154 (GH). Above San Ramon, Killip & Smith 24613 (F, GH, us). Ucayali: Prov. Coronel Portillo, La Divisoria (as Loreto), Dillon 2630 (F, MO). Prov. Coronel Portillo, Boqueron del Padre Abad, Ferreyra 8109 (GH), 8129 (GH). Cuzco: Prov. Quispicanchi, San Pedro, Var- gas 9748 (GH). Prov. La Convention, Dudley 10432 (GH). Madre de Dios: Manu, Atalaya, Foster & Wackier 7415 (MO). Puno: Prov. Carabaya, Hacienda Palmera, Vargas 16123 (GH). Comments Nephrolepis hirsutula (Forst.) Presl, Tent. Pterid. 79. 1836. Polypodium hirsutulum Forst. Prod. 81. 1786. TYPE: South Sea Islands, probably Tahiti, Forster, pho- to, us of BM (see Morton, Contr. U.S. Natl. Herb. 38: 345. 1974). This species is native in the Malaysian and Pa- cific regions and is sparingly adventive in the American tropics. It may be collected in Peru. It is similar to N. biserrata but the adaxial surface of the costa bears dissected scales and lacks tri- chomes; also, a single, prominent, basal auricle is usually on the acroscopic side of the pinnae. Family 20. BLECHNACEAE Blechnaceae (Presl) Copel., Gen. fil. 155. 1947. Blechneae Presl, Epim. bot. 103. 1851, as Blechna- ceae. TYPE: Blechnum L. Stem erect and slender to very thick, to decum- bent and small to stout, or long-creeping, or scan- dent, hardly sclerotic to strongly indurated, bear- ing scales. Leaves ca. 10 cm to 1 5 m long, circinate in vernation, monomorphic to dimorphic, the lamina entire to usually pinnatisect or 1-2-pin- nate, glabrous, minutely glandular, pubescent, or scaly abaxially. Petiole lacking stipules, not artic- ulate to the stem. Veins wholly free or partly anas- tomosing. Sori abaxial, short to very long, borne on a vein (or sometimes the sporangia also borne on the leaf-tissue), usually covered by an indusium that opens on the side of the axis at the segment or exindusiate, not paraphysate. Sporangia with a 2-3-rowed stalk and a vertical or nearly vertical annulus, which is usually interrupted by the stalk. The Blechnaceae are a family of nine genera and about 175 species. There are two genera and 20 species in Peru. The spores are monolete, and the young leaves are usually tinged with red. Key to Genera of Blechnaceae a. Leaf to ca. 2 m long, determinate; lamina entire to 1 -pinnate I. Blechnum a. Leaf climbing to 15 m or more, indeterminate; lamina 2-pinnate II. Salpichlaena FIG. 3. Blechnum asplenioides: a, habit; b, fertile pinnae. Blechnum fraxineum: c, habit. Blechnum binervatum ssp. fragile: d, habit; e, portion of rachis with pinna base; f, stem scales. (Adapted from Stolze, Ferns and Fern Allies of Guatemala, 1981.) 54 FIELDIANA: BOTANY TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 55 I. Blechnum Blechnum L., Sp. pi. 1077. 1753; Gen. pi. ed. 5, 485, 1754. TYPE: Blechnum occidental L. (as B. orientale). Figure 3. Lomaria Willd., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 3: 160. 1809. TYPE: Lomaria nuda (Labill.) Willd. = Blechnum nudum (Labill.) Luers. Terrestrial, rupestral, or sometimes epiphytic, or scandent-epiphytic. Stem decumbent, or long- creeping, or scandent, or erect and small to mas- sive, often producing stolons. Leaves monomor- phic to dimorphic and the fertile usually longer than the sterile and with narrower segments. Lam- ina entire or usually pinnatisect to 1 -pinnate. Veins free or rarely partly anastomosing and without in- cluded veinlets. Sori short to long, on a vascular commissure close to the costa, not paraphysate, covered by an indusium of similar extent that opens toward the costa. Spores ellipsoidal, monolete, nearly smooth, slightly papillate, rugose, reticu- late, or echinate. Blechnum is a nearly worldwide genus of per- haps 150 species, 19 of them in Peru. Although infrageneric classifications are sometimes pre- sented (Smith, 1985), there is uncertainty con- cerning the relations of several species, and only two subgenera are mentioned here: subgenus Blechnum (species 1-7), with expanded green tis- sue beyond the attachment of the indusium and with monomorphic leaves, and subgenus Lomaria (Willd.) C. Chr. (Index fil. 1 50, 40 1 . 1 905) (species 8-19), with little or no green tissue beyond the attachment of the indusium and with usually di- morphic leaves. Species of Blechnum are often poorly defined, due in part to genetic and phenotypic variation and also to hybridization. Walker (1985) reports two hybrids in subgenus Blechnum and suggests that these probably represent but a few of those that occur in nature. For the purposes of identi- fication of the hybrid (Blechnum x caudatum (3 x ) B. fraxineum (2 x ) x occidental (4 x ) is placed under B. occidentale, and the hybrid B. asplenioi- des x fraxineum (4x) is treated under B. con- fluens. Much more cytological data are needed be- fore the taxonomy of the Andean Blechnum occidentale complex (species 1-5) is adequately known. Walker ( 1 985) and Mickel and Beitel (1988) discuss the characters of hybrids in this complex. Various characters, sometimes employed as species distinctions, have been found to be too variable in Peru to merit their use— for example, the presence or absence of aerophores on the lam- ina (at the base of the pinnae abaxially) or the shape of the aerophores, and pubescence and glan- dularity of the lamina. In this treatment, a pinna is considered to be a stalked or sessile segment or an adnate one that is longer than broad. A lobe is considered to be a reduced segment that is adnate and broader than long. While this treatment is centered on the spe- cies of Peru, it includes an assessment of all South American Blechnum materials. However, species tend to be more variable in the Andes than to the north, and we do not wish to imply that other species of Mexico, Central America, and the West Indies are not probably recognized. References BROADHURST, J. 1912. The genus Struthiopteris and its representatives in North America. I, II. Bull. Torrey Bot. Club, 39: 257-278, 357-385. LOOSER, G. 1947. Los Blechnum (Filices) de Chile. Rev. Universit. (Univ. Catol. Chile), 32(2): 7-106. MICKEL, J. T., AND J. M. BEITEL. 1988. Pteri- dophyta flora of Oaxaca, Mexico. Mem. New York Bot. Card., 46. MURILLO, M. T. 1968. Blechnum subgenus Blechnum en sur America. Nova Hedwigia, 16: 229-366. SMITH, A. R. 1981. Flora of Chiapas, 2, pp. 1- 370. In Breedlove, ed., Pteridophytes. Califor- nia Academy of Sciences, San Francisco. . 1985. Pteridophytes of Venezuela, an Annotated List. University of California, Berke- ley (not formally published). TRYON, R. M., AND A. F. TRYON. 1982. Blech- num, pp. 669-680. In Ferns and Allied Plants. Springer- Verlag, New York. WALKER, T. G. 1985. The cytological and tax- onomic implications. In A. C. Jermy and T. G. Walker, eds., Cytotaxonomic Studies of the Ferns of Trinidad. Bull. Brit. Mus. (Nat. Hist.) Bot., 13: 133-276. 56 FIELDIANA: BOTANY Key to Species of Blechnum a. Fertile segments with expanded green tissue beyond the attachment of the indusium; leaves mono- morphic, the sterile segments with essentially flat margins b b. Lamina entire, rarely with 1 or 2 small pinnae or lobes at the base 5. B. lanceola b. Lamina pi n na t i lid to 1 -pinnate c c. Lamina 1 -pinnate; pinnae with the short stalk articulate to the rachis 6. B. serrulatum c. Lamina often pinnatifid, pinnae of a 1 -pinnate lamina with the stalk continuous with the rachis d d. Lamina with the apex abruptly reduced to an apical segment that is longer than the longest pinna 3. B. fraxineum d. Lamina with the apex gradually reduced, the apex pinnatifid or with an entire tip that is usually shorter than the longest pinna e e. Lamina usually not or only slightly reduced at the base; basal pinnae truncate or semi- cordate to cordate, not fully adnate to the rachis 1 . B. occidentale e. Lamina reduced at the base; basal pinnae fully adnate to the rachis although an auricle may be partly free f f. Scales at the base of the petiole usually few, brown to blackish and usually nearly concolorous; the longest pinna ca. 0.5-4 cm long g g. Lamina hardly reduced at the base, or more or less abruptly reduced to basal pinnae that are as long as broad or longer than broad 2. B. confluens g. Lamina gradually reduced at the base to lobes that are broader than long 4. B. asplenioides f. Scales at the base of the petiole numerous, wholly blackish or with a blackish center and brown margins; the longest pinna (7-)15-20(-25) cm long . . 7. B. brasiliense a. Fertile segments contracted, with little or no green tissue extending beyond the attachment of the indusium; leaves dimorphic or if monomorphic then the sterile segments also contracted and with involute margins h h. Scales at the base of the petiole absent, few, or numerous, lanceolate to usually lanceolate-ovate; stem decument, erect or climbing, rarely arborescent i i. Leaves monomorphic; pinnae of the sterile lamina strongly involute, the margins usually reach- ing or nearly reaching the costa on the abaxial side 15. B. loxense i. Leaves dimorphic; pinnae of the sterile lamina flat or somewhat involute and the margins distant from the costa on the abaxial side j j. Pinnae of the sterile lamina narrowly or broadly adnate, or most of them so k k. Adult plants with a scandent stem, hemiepiphytic or epiphytic, its scales, especially the older ones, mostly bicolorous with a blackish center and lighter margins 13. B. binervatum k. Adult plants terrestrial, the stem scales concolorous 1 1. Sterile lamina abruptly reduced at the base, sometimes to a wing or to lobes, 1 2-30 cm broad, the petiole with some very small, inconspicuous lobes 12. B. divergens 1. Sterile lamina gradually reduced at the base, ca. 1.5-10 cm broad m m. Sterile lamina reduced at the base to small pinnae that are 2-4 mm long 9. B. andinum m. Sterile lamina reduced at the base to definite lobes n n. Pinnae of the sterile lamina mostly curved, ascending 8. B. lehmannii n. Pinnae of the sterile lamina straight, strictly patent or nearly so 10. B. penna-marina j. Pinnae of the sterile lamina sessile or short-stalked, or most of them so o o. Sterile lamina with often biauriculate pinnae, with a much prolonged, viviparous apex of reduced pinnae 16. B. sprucei o. Sterile lamina with cuneate, broadly rounded to cordate, not auriculate pinnae, the apex gradually to abruptly reduced, not viviparous p p. Apex of the sterile lamina gradually reduced to an entire tip; many or most (rarely TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 57 only some) of the scales of the costa and rachis abaxially denticulate to ciliate, more or less spreading, light brown to dark brown and then sometimes bicolorous 14. B. stipitellatum p. Apex of the sterile lamina abruptly reduced to a large, entire, terminal segment; scales of the costae and rachis abaxially entire, rarely denticulate or ciliate, mostly appressed, whitish to light brown 11. B. cordatum h. Scales at the base of the petiole numerous, long, acicular beyond the usually broader base, curved; stem usually arborescent q q. Sterile lamina with few or no fully adnate pinnae distally, with the base of the lamina abruptly and usually hardly reduced r r. Sterile lamina with the pinnae mostly patent or slightly ascending; pinnae with the apex acute to long-acuminate, the margins flat, or at least the tip flat or nearly so; stem mostly bearing few leaves 17. B. schomburgkii r. Sterile lamina with the pinnae ascending to strongly ascending; pinnae with the apex obtuse, the margins involute to strongly involute, the tip involute; stem bearing a dense crown of numerous leaves 18. B. aural um q. Sterile lamina with many fully adnate pinnae distally, with the lamina base gradually reduced .19. B. obtusifolium 1. Blechnum occidentale L., Sp. pi. 1077, as "ori- entale." 1753. TYPE: LINN 1247.1, photo A is this species. Blechnum caudatum Cav., Descr. pi. 262. 1802. TYPE (designated by C. Chr., Dansk Bot. Ark. 9(3): 20. 1937): Ecuador, San Antonio, Nee (holotype, MA). Blechnum glandulosum Link, Enum. hort. berol. alt. 2: 462. 1822. TYPE: Cultivated, "Hab in Brasi- lia" (holotype, Herb. Link, B; photo, GH). Blechnum pectinatum Hooker, Hooker's Icon. PI. 1 : t. 95. 1837, not Presl, 1825. TYPE: Peru, Casapi, Mathews (holotype, K). Blechnum x caudatum Cav. (pro sp.) (T. G. Walker, 1985). Stem small, erect to usually decumbent and short-creeping. Leaves monomorphic. Petiole with basal scales rather few, short, mostly lanceolate to ovate-lanceolate, stem scales wholly light brown to dark brown or often with lighter margins. Lam- ina pinnatisect to 1 -pinnate at the base, the apex gradually reduced, pinnatifid or with an entire tip shorter than the longest lateral pinna, not or slight- ly reduced at the base (rarely more), the pinnae essentially flat, the basal ones truncate, semicor- date to cordate. Fertile pinnae with green tissues extending well beyond the attachment of the in- dusium. Terrestrial and rupestral, lowland rain forests, montane forests, cloud forests, along streams, on banks, in thickets, among and on rocks, usually in moist and shaded sites, 400-3800 m, usually at 700-2400 m, Tumbes south to Arequipa and Puno. Tropical America. This is a common and variable species, part of the variation undoubtedly due to hybridization. Walker (1985) noted the hybrid B. fraxineum x occidentale, which is called Blechnum x caudatum Cav. While the morphology of the hybrid may agree with the holotype, the latter cannot be cy- tologically known and may represent a homoploid variation of B. occidentale. Walker also repeated his report of Blechnum asplenioides (as B. unila- terale) x occidentale. Mexia8216, Dept. Huanuco (MO), is clearly B. fraxineum x occidentale; an- other sheet of the same collection (GH) is B. oc- cidentale. Blechnum occidentale has a rachis that is gla- brous or nearly so, and in Peru it grows from 400 to 3850 m. It is much more common than Blech- num glandulosum (B. occidentale var. pubirhachis Rosenst.), which has a definitely glandular-pubes- cent rachis. The two variations are altitudinally distinct in Central America (Stolze, Fieldiana, Bot., n.s., 6: 110-111. 1981) but not in Peru where there is considerable overlap in the altitude. The glan- dular-pubescent variation occurs from 650 to 2700 m. Accordingly, the two are treated as one taxon in Peru. Tumbes: Between Caucho and Cotrina, Coronado 219 (GH). Prov. Zarumilla, near Campo Verde, Simpson & Schunke 457 (F, us, USM). Piura: Prov. Huancabamba, Sagdstegui & Cabanillas 8543 (MO, us). Lambayeque: Prov. Lambayeque, km 45 from Olmos, Plowman et al. 14282 (F, GH). Cajamarca: Prov. Contumaza, El Molino, Sagdstegui et al. 10895 (F). Amazonas: Prov. Bagua, entre Aramango y Montenegro, Lopez et al. 4217 (GH, us). Just E of Chachapoyas, Wurdack 730 (F). La Libertad: Prov. Otuzco, Dist. Huaranchal, Sagdstegui 95 (HUT). San Martin: Prope Tarapoto, Spruce 3950 (GH). San Roque, LI. Williams 7029, 7269, 7523 (F). Huanuco: 58 FIELDIANA: BOTANY Prov. Huanuco, Dist. Churubamba, Mexia 8137 (F, GH, MO). Prov. Pachitea, Panao, Ferreyra 1791 (GH, USM). Prov. Huanuco, Puente Durand, Stork & Morton 9425 (F, MO, us). Lima: Lomas de Atocongo, Coronado 291 (F, GH). Chosica, Macbride & Featherstone 489 (F, us). Prov. Huarochiri, Leon 177 (MO, USM). Pasco: 8 km N of Huancabamba, Skog et al. 5074 (us). Junin: Cerca a San Ramon, Cerrate 962 (GH). La Merced, Killip & Smith 23760 (F, us). Huancavelica: Prov. Tayacaja, SE of Tin- tay, Tovar 4704 (GH). Ayacucho: Estrella, between Huan- ta and Rio Apurimac, Killip & Smith 22659 (F, GH, us). Cuzco: Prov. Urubamba, Machu Picchu, Mexia 8084 (F, GH, MO). Prov. La Convention, Potrero, Tryon & Tryon 5395 (F, GH, us). Arequipa: Lomas de Atiquipa, Coro- nado 30 (GH, us). Puno: Prov. Sandia, Sandia, Vargas 14813 (F). 2. Blechnum confluens Schlect. & Cham., Linnaea 5: 613. 1830. TYPE: Mexico, Hacienda de Laguna, Schiede 779 (holotype, B, photo, GH; isotype, Schiede & Deppe 779, B, photo, GH). Blechnum triangulare Link. Hort. berol. 2: 76. 1833. TYPE: Cultivated (holotype, B, Herb. Link, photo GH; isotype,?, "e Mexico, Blechnum triangulare Spr." Sprengel script., B, photo, GH). Although Link published the name first, he says he received it from Sprengel. Stem small, erect to decumbent and short-creep- ing. Leaves monomorphic. Petiole with basal scales rather few, short, mostly lanceolate to ovate-lan- ceolate, stem scales wholly light brown to dark brown or often with lighter margins. Lamina pin- natisect, the apex gradually reduced, pinnatifid, reduced at the base to pinnae that are longer than broad or as long as broad, the pinnae essentially flat, the longest ca. 1—4 cm long. Fertile pinnae with green tissue extending well beyond the at- tachment of the indusium. Terrestrial or rupestral, in forests, on wet banks, among rocks, or in rock crevices, 670—2300 m, Amazonas south to Junin. Tropical America. This species is more or less intermediate be- tween B. occidentale and B. asplenioides; some (all?) specimens might represent hybrids between those species. The basal lobes in B. asplenioides are fully adnate and broader than long, while in B. confluens they are longer than broad or about as long as broad and only partly adnate to the rachis. Amazonas: E of Chachapoyas, Wurdack 730 (GH). San Martin: Monte Guayrapurima, prope Tarapoto, Spruce 4022 (GH, us). Prov. Lamas, Dist. Lamas, Belshaw 3424 (F, GH, us). Lamas, LI. Williams 6341 (F). Huanuco: Prov. Leoncio Prado, Dist. Rupa Rupa, J. Schunke V. 5081 (GH). Between Tingo Maria and Monzon, Croat 57909 (F). La Divisoria, Gentry et al. 18864 (F). Pasco: Prov. Oxapampa, Soukup 6810 (GH). Junin: Prov. Tar- ma, Utcuyacu, Woytkowski 35371 (GH). Near La Mer- ced, Killip & Smith 23994 (F). 3. Blechnum fraxineum Willd., Sp. pi. ed. 4, 5: 413. 1810. TYPE: Venezuela, Caracas, Bre- demeyer (holotype, B, Herb. Willd. 20044, photo, GH). Not Lomaria fraxinea Willd., op. cit. 1810. Figure 3c. Blechnum longifolium Willd., Sp. pi. ed. 4, 5: 413. 1810, not Cav. 1802. TYPE: Venezuela, Caripe, Humboldt 464 (holotype, B, Herb. Willd. 20045, photo, GH). Blechnum graciletiaulf., Enum. fil. 158. 1824. TYPE: "Brasilia, Otto comm." (not seen), ex char. Stem small, erect to decumbent and short-creep- ing. Leaves monomorphic. Petiole with basal scales usually rather few, rather short, mostly lanceolate to ovate-lanceolate, stem scales brown to dark brown, sometimes with narrow lighter margins. Lamina pinnatisect to 1 -pinnate, the apex abruptly reduced to an apical pinna that is longer than the longest lateral pinna, not or slightly reduced at the base, the pinnae essentially flat. Fertile pinnae with green tissue extending well beyond the attachment of the indusium. Terrestrial or rupestral, on clay banks, along streams, and among rocks, 450-2000 m, Ama- zonas, south to Cuzco. Tropical America. Walker (1985) reported a diploid and a tetra- ploid from Trinidad. The tetraploid, which is more dimorphic than the diploid and with additional (6-14) pinnae, is evidently more common in Peru than the diploid, with 2-4 pinnae. The identifi- cation of Peru material is uncertain pending cy- tological studies that have not been made. Blech- num gracile is a name for the tetraploid and B. meridense Kl. for the diploid. The type of B. fraxi- neum is considered to be a triploid hybrid of B. meridense and B. occidentale. The essential char- acter of the species and its hybrids is the long terminal segment. Amazonas: Prov. Bagua, valley of Rio Maranon, above Cascadas de Mayasi, Wurdack 1822 (GH. us. USM). San Martin: San Roque, LI. Williams 7519 (F). Tarapoto, Spruce 4026 (us). Junin: La Merced. Macbride 5500 (F. us). Chanchamayo valley. C. Schunke 3 1 (F). / 32 (F. us). 1352 (F). Above San Ramon. Killip & Smith 24731 (us). Ucayali: La Divisoria. Tryon & Tryon 5271 (GH. us). TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 59 Cuzco: Prov. Paucartambo, Pillawata, Vargas 16729 (GH). QUILLABAMBA, CORONADO 127 (US). 4. Blechnum asplenioides Sw., Kongl. Vetensk. Acad. Handl. 1817: 72, t. 3, f. 3. TYPE: Bra- zil, Villa Rica, Freyreis (holotype, s, Herb. Swartz, photo, GH, us). Figures 3a-b. Asplenium blechnoides Sw., Syn. fil. 76. 1806. TYPE: Peru, evidently Nee ex Lagasca (holotype, s; pho- to, GH, us). Blechnum unilaterale Sw., Ges. Naturf. Freunde Ber- lin Mag. Neuesten Entdeck. Gesammten Naturk. 4: 79. 1810, also Willd., Sp. pi. ed. 4, 5: 407. 1810. Both authors cite Asplenium blechnoides Sw. in synonymy; hence the name is superfluous and has the same type as Asplenium blechnoides. Blechnum polypodioides Raddi, Opusc. Sci. 3: 294. 1819, not (Sw.) Kuhn, 1 868. TYPE (from Raddi, PI. bras. nov. gen. 53, t. 60, f. 2. 1825): Brazil, Corcovado, Raddi (holotype, PI?). Blechnum blechnoides (Sw.) C. Chr., Index fil. 151. 1906, not Keyserl., 1873. Stem small, erect to decumbent and short-creep- ing. Leaves monomorphic. Petiole with basal scales rather few, mostly short, lanceolate to narrowly lanceolate, stem scales light brown to dark brown or with a dark brown center. Lamina pinnatifid to pinnatisect, the apex gradually reduced, pinnatifid, the base gradually reduced to lobes that are broad- er than long, pinnae essentially flat, the longest ca. 0.5-2.5 cm long. Fertile segments with green tissue extending well beyond the attachment of the in- dusium. Terrestrial or rupestral, wet forests, secondary forests, shaded stream banks, clay soil of road banks, or on rocks, 300-1 800 m, Amazonas south to Cuzco. Tropical America. This species is distinguished by the small basal lobes of the lamina. It is usually a small plant, although sometimes the leaf reaches a length of ca. 50 cm. Amazonas: Prov. Bagua, Rio Maranon, above Cas- cadas de Mayasi, Wurdack 2056 (F, GH, us). Prov. Bagua, NE of Chiriaco, Barbour 4509 (USM). San Martin: Ze- pelacio, near Moyobamba, Klug 3358 (F, GH, us). Prov. Lamas, Lamas, Ferreyra 4691 (GH, USM). San Roque, LI. Williams 7144 (F). Loreto: Pampayacu, Klug 3237 (F, GH, us). Huanuco: Villcabamba, Macbride 4991 (F, GH, us). Tingo Maria, Tryon & Tryon 5259 (F, GH, us). Pasco: Dos de Mayo, Pichis Trail, Killip & Smith 25861 (F, GH). Junin: La Merced, Soukup 2374 (GH). Near La Merced, Killip & Smith 23959 (F, us). Chanchamayo valley, C. Schunke2, 26, 182, 777, 1374(p). Ucayali: La Divisoria, Allard 22174 (GH). La Divisoria, Gentry et al. 18832 (F, USM). Cuzco: Prov. Quispicanchis, entre Quinze Mil y San Lorenzo, Vargas 11761 (GH). Prov. La Convention, valley of Rio Vilcanota, Mexia 8052 (F, GH, us). Prov. Paucartambo, Kosnipata valley, Plowman & Davis 5114 (GH). Madre de Dios: 39 km SW of Puerto Maldmado, Smith et al. 1 1 28 (us). 5. Blechnum lanceola Sw., Kongl. Vetensk. Acad. Handl. 1 8 1 7: 7 1 , t. 3, f. 2. TYPE: Brazil, Villa Rica, Freyreis (holotype, s, Herb. Swartz, photo, GH, us). Stem small, erect to decumbent and very short- creeping. Leaves monomorphic. Petiole with basal scales few to nearly lacking, short, mostly lanceo- late to ovate-lanceolate, stem scales light to dark brown, or with lighter narrow margins. Lamina entire or rarely with 1 or 2 small lobes at the base, mostly very narrowly elliptical, essentially flat. Fertile lamina with green tissue extending well be- yond the attachment of the indusium. A single collection from San Martin, Peru. Peru south to Argentina and east to Brazil. This species is represented by small plants 4— 1 5 cm, or occasionally to 30 cm, tall. The entire lamina is distinctive, although rarely there may be one or two lobes at the base. It is evidently rare in the Andes and more common in southeastern Brazil. San Martin: Monte Campana, prope Tarapoto, Spruce 4672 (GH, us). 6. Blechnum serrulatum Rich., Actes. Soc. Hist. Nat. Paris 1: 114. 1792. TYPE: "Cayenne" (French Guiana), Le Blond (holotype, p, Herb. Richard, photo, GH, us). Stem widely creeping (in deep sandy soil), its ascending branches bearing the leaves. Leaves monomorphic. Petiole with few or no scales at the base, stem scales wholly dark brown or with lighter margins or small and light brown only. Lamina 1 -pinnate, with an entire apical segment, not or hardly reduced at the base, pinnae essentially flat, articulate to the rachis, more or less serrulate, the largest ca. 5-1 5 cm long. Fertile pinnae with green tissue extending well beyond the attachment of the indusium. Sandy soil in secondary vegetation, 1 80 m, Lo- reto. Tropical America, south to Peru and north- eastern Argentina. 60 FIELDIANA: BOTANY The species has monomorphic leaves and differs from all species in South America in having ar- ticulate pinnae. In sandy deep soil, the main stem is deeply subterranean and has ascending branches that bear the leaves. Loreto: Jenaro Herrera, Mejia & Van der Well 221 (USM). 7. Blechnum brasiliense Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Ge- sammten Naturk. 5: 330. 1811. TYPE: Brazil, Dombey (holotype, P, Herb. Jussieu 1390, photo, GH, us). Blechnum nigrosquamatum Gilbert, Bull. Torrey Bot. Club 24: 258. 1897. TYPE: Bolivia, Colapampa, Bang 2314 (holotype, not located; isotype, GH!). Stem subarborescent to arborescent, stout, to 2 m tall. Leaves monomorphic. Petiole with scales at the base many, mostly 10-15 mm long, acicular, blackish or with a blackish center, stem scales dark brown to blackish or with a dark center and lighter (to brown) margins. Lamina pinnatifid to pinnati- sect, the apex gradually reduced, pinnatifid, or with a short entire tip, gradually reduced, pinnatifid, or with a short entire tip, gradually reduced at the base to small pinnae or lobes, pinnae essentially flat, the largest ca. (7-)15-20(-25) cm long. Fertile pinnae with green tissue extending well beyond the attachment of the indusium. Terrestrial, mountain forests, 1200-1600 m, San Martin. Guatemala; tropical South America. The petiole is usually short to nearly absent. The long, narrow scales at the base of the petiole com- bined with the monomorphic leaves are distinc- tive features of this species. San Martin: Zepelacio, near Moyobamba, Klug 3433 (F, GH, us). Murillo (1968) also cites: Monte Campana, Tarapoto, Spruce 4673 (BR). 8. Blechnum lehmannii Hieron., Bot. Jahrb. Syst. 34: 473. 1904. TYPE: Colombia, Rio Tim- biqui, Lehmann 8928 (holotype, B; isotype, us!). Struthiopteris maxonii Broadh., Bull. Torrey Bot. Club 39: 268. 1912. TYPE: Panama, Chiriqui, Cerro de la Horqueta, Maxon 5415 (holotype, us; iso- type, GH!, us!). Blechnum maxonii (Broadh.) C. Chr., Index fil. suppl. 16. 1913. Stem more or less erect, rather slender. Leaves dimorphic. Petiole with basal scales few to rather numerous, brownish, mostly large, concolorous, lanceolate-ovate or broader, stem scales acumi- nate, brown to dark brown, to blackish (some), concolorous. Sterile lamina pinnatisect, the apex gradually reduced or with a short to long entire tip, the base gradually reduced to lobes, pinnae curved-ascending, fully adnate to the rachis, the margins flat to somewhat involute, distant from the costa. Fertile pinnae with green tissue hardly extending beyond the attachment of the indusium. Terrestrial, cloud forests, 1750 m, Cuzco. Southern Mexico; Central America; Venezuela and Colombia, south to Peru. The stem may be erect and to 40 cm tall; the sterile lamina is 3-10 cm, usually 4-6 cm broad. This is closely related to, perhaps conspecific with, B. I'herminieri (Kunze) Mett. of the West Indies. That species has the petiole dark, and the lamina is abruptly reduced basally in contrast to B. lehmannii, which has a light-colored petiole and a lamina that is gradually reduced at the base. Lopez & Sagdstegui 5297 (GH, HUT), from Prov. Chota, Cajamarca, seems to be this species, but the petioles are mostly dark, rather than straw- colored as in typical B. lehmannii. The collection may represent an intermediate between the two taxa. Cuzco: Prov. La Convencion, Cordillera Vilcabamba, Knox's Cascade, Dudley 10463 (GH). Dudley 1 0489 from the same place (GH) is a sheet of juvenile plants, most probably of this species. 9. Blechnum andinum (Baker) C. Chr. Ind. fil. 1 50. 1905. Lomaria andina Baker, Syn. fil. ed. 2: 482. 1883. TYPE: Andes, "Unduari" (Unduavi), Bolivia, 10,000 ft, Pearce (holotype, K; photo, GH, us). Blechnum subtile Rosenst., Repert. Spec. Nov. Regni Veg. 11: 54. 1912. TYPE: Unduavi, Bolivia. Buchlien 2779 (holotype, not located; isotype, us!). Stem small, erect, stoloniferous, the stolons very slender and threadlike. Leaves small, dimorphic. The fertile and sterile about the same length. Pet- iole lacking scales or with a few, brownish, con- colorous ones, stem scales more or less small, brownish, sometimes with lighter margins. Sterile lamina 1 -pinnate, the apex and base gradually re- duced, the base reduced to small pinnae, the pin- nae obtuse, narrowly to broadly adnate. Fertile TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 61 pinnae with green tissue hardly extending beyond the attachment of the indusium. Wet forests, on trees and in a cave, 3050 m, Cuzco. Peru; Bolivia; and southern Brazil. This is a very small species, the leaves usually 6-12 cm long. Cuzco: Michihianuca, Bues 973 (us). Huadquina, Bues 980 (us). 10. Blechnum penna-marina (Poiret) Kuhn, Fil. Afr. 92. 1868. Polypodium penna-marina Poiret in Lam., Encycl. 5: 520. 1804. TYPE: Straits of Magellan, Commer- son (holotype, Herb. Desfontaine, p; photo, GH). Stem small, more or less erect, freely stolonif- erous and sometimes forming large clumps of plants. Leaves small, dimorphic, the sterile shorter and spreading, the fertile longer and erect. Petiole with a few brownish, concolorous scales, stem scales brown, more or less reddish brown, to light brown or dark brown, concolorous, more or less acute to acuminate. Sterile lamina mostly 5-10 (-15) cm long, narrow, pinnatisect, gradually re- duced to the apex, at the base reduced to small lobes. Fertile pinnae with green tissue hardly ex- tending beyond the attachment of the indusium. Terrestrial, 3390 m, Cuzco. Peru and southeastern Brazil; to southernmost Chile and Argentina; circumantarctic islands; southeastern Australia; New Zealand. Mettenius (Fil. Lech. 2: 15. 1859.) reported this species for Peru as Blechnum alpinum var. elongatum (Puno, Agapata, Lechler). I have not seen this specimen, and it is possible that it is some other species. In any event, the following is a recent collection. Cuzco: Prov. Paucartambo, Cerro Macho Cruz, Manu National Park, B. Leon et al. 2757 (cuz, USM). 11. Blechnum cordatum (Desv.) Hieron., Hed- wigia47: 239. 1908. Lomaria cordata Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 330. 1811. TYPE: omitted in the publication. Ho- lotype evidently "Perou-Concepcion," Dombey, p, Herb. Jussieu 1368; photo, GH, us. Lomaria chilensisKau\f.,Enum.fi\. 154. 1824. TYPE: Chile, Chamisso (holotype, LE?) Lomaria ornifolia Presl, Reliq. haenk. 51.1 825. TYPE: "Hab. in Peruviae?" Placed as a synonym of Blechnum cordatum by Hieron., Hedwigia 47: 239. 1908. Lomaria serrulosa Desv., Mem. Soc. Linn. Paris 6: 290. 1827. TYPE: "Peruvia" (holotype, P; photo, GH, us). Lomaria arborescens Klotzsch, Linnaea 20: 347. 1844. SYNTYPES (designated by Hieron., Hedwigia 47: 240. 1908): Venezuela, Merida, Moritz 299 (B) and Venezuela, Karsten Coll. II, 25 (B). Isosyn- type, Moritz 299, BM; photo, GH; frag., us!. Blechnum chilense (Kaulf.) Men., Fil. lechl. 1: 14. 1856. Blechnum ornifolium (Presl) Ettingsh., Denkschr. Kaiserl. Akad. Wiss. Wien. 23: 61. 1864. Blechnum arborescens (Klotzsch) Hieron., Hedwigia 47: 239. 1908. Blechnum peruvianum Hieron., Hedwigia 47: 245, /. 10. f. 19. 1908. TYPE: Peru, Rio Huallaga, Mo- yobamba, Stiibel 1 105 (holotype, B; photo, us). Stem erect and subarborescent to decumbent and short-creeping, moderately to very stout. Leaves dimorphic. Petiole with basal scales small to rather large, few to many, concolorous and mostly brown, linear to lanceolate to lanceolate- ovate or broader, stem scales broad, thin, concol- orous, light brown or reddish brown. Sterile lam- ina 1 -pinnate, the apex abruptly reduced to a ter- minal segment, the base not or only slightly reduced, pinnae sessile to definitely stalked, strict- ly patent to somewhat ascending, cuneate to broadly rounded to cordate, the margins flat or slightly involute, distant from the costa. Fertile pinnae with green tissue extending hardly beyond the attachment of the indusium. Terrestrial, wet montane forests, cloud forests, thickets, along streams, on clay road banks, and on old landslides, 700-3300 m, Piura south to Cuzco. Venezuela and Colombia, south to Peru; east- ward to southeastern Brazil; Chile; Juan Fernan- dez Islands, West Falkland Islands; entire range not certain. This is a variable species, primarily of the Andes from Venezuela and Colombia, south to Chile. The leaves are often large, to 1 m or more long, the apex is abruptly reduced to a large terminal segment, and the scales of the costae and rachis are mostly rather appressed and concolorous, whitish to light brown. The species is variable in many characters— for example, aerophores are absent, or present on some pinnae and then low to conical, the rachis may be smooth or strongly roughened and it varies from straw-colored to dark reddish. There seems to be 62 FIELDIANA: BOTANY no basis, at least in Peru, for the recognition of more than one taxon. A. R. Smith (1985) also treated Blechnum cordatum as a wide-ranging spe- cies with many synonyms. As in the case of other variable species, related ones recognized to the north of South America may be distinctive. A careful revision of this complex is much needed. The photograph of the type of Lomaria cordata clearly shows that the label reads "Perou-Concep- cion," rather than "Peru, Concepcion," as indi- cated by Morton on the covering label. Dombey never collected at or near a Concepcion in Peru, but he did travel from Lima to Concepcion, Chile, where he evidently collected the type specimen. Piura: Prov. Ayabaca, Sagdstegui & Cabanillas 8702 (F, HUT). Lambayeque: 4 km NW of Incahuasi, Dillon & Skill man 4155 (F, GH, USM). Arriba de Incahuasi, Sa- gdstegui el al. 12893 (F, GH). Amazonas: Mountain E of Balsas, Osgood & Anderson 72 (F). San Martin: Prov. San Martin, Dist. Tarapoto, Hickok 636 (GH). Prov. La- mas, Cerro Blanco, J. Schunke V. 6259 (F). Prov. Maris- cal Caceres, Rio Abiseo National Park, Young 1768 (USM). Loreto: Pumayacu, King 3243 (GH, us). Anacash: Above Yungay, Correll & Smith P964(GH). Prov. Huaylas, Lopez et al. 8378 (F, HUT). Prov. Yungay, Laguna Llanganuco, Mostacero et al. 1402 (F, GH). Huanuco: Carpish, Tryon & Tryon 5317 (GH). Prov. Huanuco, Dist. Churubamba, Mexia 8174 (F, GH, us). Mito, Macbride & Feat herst one 1522 (F, us). Lima: Prov. Canta, 4 km from Huaman- tanga, Sounders 1254 (GH). Pasco: Quillasu, Soukup 3289 (or 3284) (F, GH). Prov. Oxapampa, road Oxapampa- Paucartambo, Smith & Pretel 1631 (F). Prov. Oxapampa, "La Esperanza," Leon 613 (USM). Junin: Villa Amoretti, Kunkel 655 (GH). Chanchamayo valley, C. Schunke 162 (F). La Merced, Soukup 1087 (F). Huancavelica: Prov. Tayacaja, cerca a Tocas, Tovar 2044 (GH, USM). Aya- cucho: Prov. La Mar, 20-25 km from Tambo, Dudley 1 1942 (GH). Cuzco: Prov. La Convencion, San Luis, Var- gas 19858 (GH). Prov. La Convencion, Cordillera Vil- cabamba, Dudley 10329 (GH), 10584 (GH), 70609 (GH). 5 km N of Aguas Calientes, Solomon 3169 (F, USM). 12. Blechnum divergens (Kunze) Mett., Ann. Sci. Nat. Hot. 5, 2: 225. 1864. Lomaria divergens Kunze, Linnaea 9: 57. 1 834. TYPE: Peru, Dept. Huanuco, Pampayaco, Poeppig Diar. 1141 (not located). Lomaria pteropus Kunze, Farnkr. 1: 97. 1842, nom. nov. for Acrostichum heterophyllum Raddi, PI. bras. nov. gen. 5, /. 17. 1825. TYPE: Brazil, Man- diocca, Raddi (holotype, FI?), not Lomaria het- erophylla Desv., 1811, nor Blechnum hetero- phyllum (Desv.) Schlect., Adumbr. pi. 37. 1827. Blechnum pteropus (Kunze) Mett., Fil. hort. bot. Lips. 61. 1856. Stem erect to rather long-creeping, moderately stout. Leaves dimorphic. Petiole with basal scales few, rather large, concolorous, lanceolate to lan- ceolate-ovate, stem scales broad, dark brown to blackish, concolorous. Sterile lamina pinnatisect, the apex rather abruptly reduced to a short or long, entire tip, the base abruptly reduced, sometimes to a wing or to lobes that may be distant and inconspicuous, pinnae patent to somewhat as- cending, fully adnate or rarely partly adnate, the margins flat, distant from the costa. Fertile pinnae with green tissue extending hardly beyond the at- tachment of the indusium. Terrestrial or rarely epiphytic, cloud forests, dense, wet forests, 1500-2400 m, Cajamarca, Huanuco, Junin, and Cuzco. Tropical America. In South America, specimens of Blechnum di- vergens and B. binervatum ssp. acutum are not always readily distinguished, due to variation in both taxa. Blechnum divergens is a terrestrial spe- cies, rarely epiphytic in very wet cloud forest, and the stem has broad, concolorous scales. The sterile lamina is broad, 12-30 cm, and is abruptly re- duced at the base to a petiole that is either alate or has lobes, often small and inconspicuous, that are much broader than long. The basal or proximal pinnae are usually not or not much decurrent and hardly if at all narrowed toward the adnate base. Blechnum binervatum ssp. acutum often has a broad sterile lamina that is abruptly reduced at the base, and the basal or proximal pinnae are usually strongly decurrent or narrowed to the ex- panded and adnate base. Cajamarca: Prov. Cutervo, Parque Nacional de Cu- tervo, Diaz & Osores 259911 (MO). Huanuco: SW slope of Rio Llullapichis watershed, ascent of Cerros del Sira, Dudley 13407B (GH). Junin: Chanchamayo valley, C. Schunke 11, 159, 160, 1382 (F). Prov. Chanchamayo, Rio Rondayacu, Smith & Bokor 2104 (F). Cuzco: Prov. La Convencion, Quellouno to Chirumbia, Vargas 11396 (GH). Prov. Paucartambo, between Pillawata and Patria, Plowman & Davis 4935 (GH, us). 1 3. Blechnum binervatum (Poiret) Morton & Lell., Amer. Fern J. 57: 67. 1967. Polypodium binervatum Poiret in Lam., Encycl. 5:521. 1 804. TYPE: collector and locality not known but probably in the Lesser Antilles (holotype, p, Herb. Jussieu 1365; photo, GH). Stem scandent, somewhat slender to rather stout. Leaves dimorphic. Petiole with basal scales few or numerous, short to moderately long, rather lan- ceolate, mostly bicolorous (in Peru) with a black- ish center and lighter margins, or concolorous (ssp. TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 63 binervatum). Sterile lamina pinnatisect or 1 -pinnate proximally, the apex gradually or abruptly reduced to an entire tip, the base some- what to strongly reduced to lobes, pinnae patent to ascending, mostly fully adnate, the margins flat or nearly so, distant from the costa. Fertile pinnae with green tissue extending hardly or only slightly beyond the attachment of the indusium. This is a variable species, with three subspecies that formerly were usually recognized as species. The ssp. binervatum, with concolorous stem scales, is confined to the Lesser Antilles and Puerto Rico. This is the only scandent-epiphytic species in Peru. The stem may be rooted in the ground and climbing on tree trunks, especially on Cyathea- ceae, or it may (probably) be wholly epiphytic; very rarely the plant is rupestral, or juvenile plants may be terrestrial. Southern Mexico and Central America; Antilles; Venezuela to Colombia; south to Bolivia; south- eastern Brazil. Key to Subspecies of Blechnum binervatum The following key provides the typical charac- ters of the subspecies; these intergrade, and not all specimens have the characteristic complement of characters. For example, Peyton & King 1420, Dept. Cuzco, at MO is nearly ssp. fragile, while at GH it is more like ssp. acutum. a. Sterile lamina 3-10 cm broad, gradually reduced at the base; pinnae usually close and the sinus narrow; stem scales toothed, often freely so, especially near the apex of the stem 1 3a. ssp. fragile a. Sterile lamina 10-25 cm broad, more or less abruptly reduced at the base; pinnae often spaced and the sinus broad; stem scales with few and small teeth or none 1 3b. ssp. acutum 13a. Blechnum binervatum ssp. fragile (Liebm.) Tryon & Stolze, comb. nov. Figures 3d-f. Lomaria fragile Liebm., Kongel. Dankse Vid. Selsk. Skr. Naturvidensk. Afd. 5, 1: 232. 1849. TYPE: Mexico, Barranca de Huitamalco, Liebmann (Fl. Mex. 773) (lectotype, selected by A. R. Smith, 1981): C folio, frag., us!; isolectotype, Barranca de Huitamalco, L, photo, GH). Blechnum fragile (Liebm.) Morton & Lell., Amer. Fern J. 57: 68. 1967. OsmundapolypodioidesSv/., Prod. 127. 1788. TYPE: Jamaica, Swartz (holotype, s, photo, us; isotype, BM, photo, GH). Blechnum polypodioides(S'w.) Kuhn, Fil. afr. 92. 1 868, not Raddi, 1819. Other synonyms, mostly not currently used, are in the reference for Blechnum fragile. In dense rain forests and cloud forests, 1550- 3300 m, Cajamarca south to Cuzco. Southern Mexico and Central America; Greater Antilles; Venezuela and Colombia, south to Bo- livia; southeastern Brazil. Cajamarca: Prov. Cutervo, Parque Nacional de Cu- tervo, Diaz & Osores 2596 (MO). Amazonas: Upper slopes of Puma-urea, ESE of Chachapoyas, Wurdack 695 (GH). Prov. Bongara, Shillac, Smith & Vasquez 4898 (MO, USM). San Martin: Rio Abiseo National Park, Young & Watson 3350 (USM). Huanuco: SW slope of Rio Llullapichis wa- tershed, ascent of Cerros del Sira, Dudley 13284 (GH). Muna, Macbride 4292 (F, GH). Pasco: Oxapampa, Sou- kup 2337 (F, GH). Rio San Alberto, Foster et al. 10316 (USM). Junin: Pichita Caluga, Walden 37, 74 (GH). Cuzco: Prov. La Convencion, Huayopata, Peyton & King 1420 (MO). 13b. Blechnum binervatum ssp. acutum (Desv.) Tryon & Stolze, comb. nov. Lomaria angustifolia HBK., Nov. gen. sp. 1 : 18. 1816. TYPE: Ecuador, Villa de Ibarra, Humboldt & Bonpland (B or p?, not seen). Lomaria acuta Desv., Mem. Soc. Linn. Paris 6: 290. 1827. LECTOTYPE: Dombey (designated by Weatherby, Contr. Gray Herb. 114: 26. 1936): "Habitat in Peruvia" (holotype, P; photo, GH). Lomaria cuspidata Kunze, Linnaea 9: 59. 1 834. TYPE: Peru, Pampayaco, Poeppig, Diar. 1 120 (isotype, B?, not seen). Lomaria meridensis Klotzsch, Linnaea 20: 345. 1847. TYPE: Venezuela, Merida, Moritz 297 (holotype, B, not seen). Lomaria ensiformis Liebm., Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk. Afd. 5, 1: 234. 1849. TYPE (selected by A. R. Smith, 1981): Mexico, Oaxaca, Distr. Chinantla, Teotalcingo, Liebmann PL Mex. 2342 (lectotype, c, folio, sheet with both fertile and sterile leaves, frag., us!). Blechnum meridense (Klotzsch) Mett., Fil. hort. lips. 61. 1856, based on Lomaria meridensis, not Blechnum meridense Klotzsch, 1847. Blechnum acutum (Desv.) Mett., Ann. Sci. Nat. Bot. Ser. 5, 2: 225. 1864. Blechnum angustifolium (HBK.) Hieron., Bot. Jahrb. Syst. 34: 472. 1904, not Willd., 1810. Blechnum ensiforme (Liebm.) C. Chr., Index fil. 153. 1905. Blechnum kunthianum C. Chr., Index fil. suppl. 16: 1913, nom. nov. for Lomaria angustifolia HBK., not Blechnum angustifolium Willd., 1810. 64 FIELDIANA: BOTANY Southern Mexico and Central America; Vene- zuela and Colombia, south to Bolivia; southeast- ern Brazil. In humid forests, in rain forests, and in wet cloud forests, 1000-2700 m, Amazonas south to Cuzco. In South America this subspecies is especially variable in the characters of the sterile lamina. Usually the distal pinnae are fully adnate and broadest at the base, but neither excurrent nor decurrent, and the pinnae are close with a narrow sinus between them. Proximal pinnae may be sim- ilar or they may be somewhat spaced and excur- rently and decurrently broadened at the base. These characters, however, vary to the extreme (Lomaria acuta Desv.), with few and widely spaced pinnae that are narrowed toward the base where they are fully adnate. Amazonas: Prov. Bagua, 12 km E of La Peca, Barbour 2507 (F, MO). San Martin: Prov. Rioja, Pedro Ruiz-Mo- yobamba, D. Smith 4451 (F). Prov. Rioja, Pedro Ruiz- Moyobamba, Smith & Vasquez 4709 (MO, USM). Huanu- co: SW slope of Rio Llullapichis watershed, ascent of Cerros del Sira, Dudley 13253 (GH). Pasco: Prov. Oxa- pampa, Oxapampa, Soukup 2339 (GH). Prov. Oxapam- pa, Rio El Tunqui, Smith & Alban 5524 (F). Prov. Oxa- pampa, SW of Oxapampa, Foster 7624 (USM). Junin: Prov. Chanchamayo, Rio Rondayacu, Smith et al. 2616 (F, USM). Ucayali: Prov. Leoncio Prado, La Divisoria, Plowman & Schunke 7418 (F). Cuzco: Prov. La Con- vention, 3 km from Incatambo, Peyton & Peyton 851 (GH, MO). Quillabamba, Santa Teresa, Peyton & Peyton 1273 (GH, MO); Prov. Paucartambo, Parque Nacional Manu, Leon 2194 (F). 14. Blechnum stipitellatum (Sodiro) C. Chr., In- dex fil. 160. 1905. Lomaria stipitellata Sodiro, Recens. crypt, vase. Quit. 29. 1883. SYNTYPES: Ecuador, Corazon y Ata- cazo, 2500-3400 m. Sodiro (syntype or isosyn- type, Corazon, Sodiro, p; photo, GH). Blechnum rubicundum Hieron., Hedwigia 47. 242, /. 4,f. 16. 1908. TYPE: Colombia, Cumbal to La Ceja de Mayasquer, Stiibel 347 (holotype, B; pho- to, us). Stem moderately to rather stout, erect to de- cumbent and very short-creeping. Leaves dimor- phic. Petiole with basal scales mostly large, mostly concolorous, sometimes bicolorous, brown, few to many, linear to lanceolate to lanceolate-ovate, stem scales large, brown to dark brown or reddish brown often with lighter (broad) margins, acute more or less thin, sometimes concolorous. Sterile lamina 1 -pinnate, the apex gradually reduced to an entire tip, the base slightly or not reduced, pinnae usually patent to slightly ascending, sessile to short-stalked, more or less truncate, the margins flat to somewhat involute, distant from the costa. Fertile pinnae with green tissue extending only slightly beyond the attachment of the indusium. Terrestrial, in wet forests and among rocks, 2400-3700 m, Amazonas, south to Puno. Colombia, south to Bolivia. This seems to be a distinctive species, but it is little known and its variation is uncertain. Many or most of the scales on the costa and rachis abaxially are more or less spreading, light brown to dark brown and then sometimes bicolor- ous. The scales, or at least some of them, are den- ticulate to ciliate on the margins. The lamina has many, usually 20—40 pairs of close, patent pinnae. At least when dried, the pinnae are often promi- nently striate adaxially— the tissue between the veins being depressed. This species seems close to some variations of Blechnum cordatum, but the expanded sterile pin- nae suggest it may be a shade form of Blechnum loxense. Amazonas: Prov. Bagua, Cordillera Colan, Barbour 3467 (MO). Huanuco: Between Huanacoand Pampayacu, Kanehira 144 (GH, USM). Huanuco-Tingo Maria road, Gentry et al. 33851 (F). Pasco: Prov. Oxapampa, Cor- dillera San Gutardo, Leon et al. 517 (USM). Junin: Prov. Satipo, Dist. Pampa Hermosa, Sounders 1069 (GH). Cuz- co: Prov. Urubamba, above Palca, Peyton & Peyton 767 (GH, MO). Puno: Agapata, Lechler 2025, in part (GH). 15. Blechnum loxense (HBK.) Salomon, No- mencl. Gefaskrypt. 117. 1883. Lomaria loxensis HBK., Nov. gen. sp. 1: 18. 1816. TYPE: Ecuador, "Andibus Peruviae inter Gon- zanama et Loxa," Humboldt & Bonpland (holo- type, P; photo, GH). Lomaria squamulosa Desv., Mem. Soc. Linn. Paris 6: 290. 1827. "Habitat in Peruvia," Dombey (F, photo, us). Lomaria stenophylla Klotzsch, Linnaea 20: 346. 1847. TYPE: of alpha. Peru, Dombey (B; photo, us). This and also beta are both B. loxense according to Hieronymus, Hedwigia 47: 241. 1908. Not Blechnum stenophyllum Presl, 1851. Blechnum stenophyllum (Klotzsch) Men., Fil. hort. bot. lips. 64. 1856. Not Blechnum stenophvllum Presl. Blechnum squamulosum (Desv.) Men., Ann. Sci. Nat. Bot. Ser. 5,2: 225. 1864. Stem stout to massive, erect to somewhat de- cumbent and very short-creeping. Leaves mono- morphic. Petiole with basal scales usually numer- TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 65 ous, large, brown and concolorous, stem scales mostly broad, light brown to brown to more or less reddish brown with or without a darker center. Sterile lamina 1 -pinnate, the apex abruptly re- duced to a terminal segment, the base hardly or not reduced, the pinnae ascending, usually cor- date, the margins strongly involute, usually reach- ing or nearly reaching the costa on the abaxial side, the tip usually strongly involute. Fertile pinnae with green tissue extending only slightly beyond the attachment of the indusium. In open, often wet, sites, usually in the puna, grasslands, steep roadside banks, among rocks or in the crevices of rocks, 1750-4100 m, Lamba- yeque south to Puno. Costa Rica and Panama; Venezuela and Colom- bia, south to Bolivia. The stem is stout but only rarely (to ca. 2 m tall) arborescent. It is usually soft, mucilaginous and green inside, especially at the apex. Except for B. auratum var. columbiense, this species is distinc- tive among those of Peru in having very narrow sterile pinnae as well as fertile ones. Lambayeque: Dist. Incahuasi, Sagdstegui et al. 12781 (F, GH, HUT). Cajamarca: Prov. Hualgayoc, Carmona. Soukup 501 1 (us). Amazonas: Paso de Calla-Calla, San- chez 87 (GH). San Martin: Prov. Mariscal Caceres, Rio Abiseo National Park, Young & Watson 3499 (USM), 4400 (F, USM). La Libertad: Prov. Bolivar, Laguna de los Ichus, Lopez & Sagdstegui 3237 (GH). Prov. Santiago de Chuco, Dist. Mollepata, Sounders 875 (GH). Prov. Santiago de Chuco, Shoreyo-Trujillo, Smith 2344 (F). Ancash: NW slope of Nevada de Huascaran, Correll & Smith P958 (GH). Prov. Pallasca, Consuzo, Lopez 2394 (GH). Huascaran National Park, D. Smith 10613, Smith et al. 10380 (F), 77627 (F, HUT), 12395, 12457 (F). Huan- uco: Mito, Bryan 203 (F), Macbride & Featherstone 1790 (F, us), 2779 (F, us). Junin: Huancayo, Kunkel 425 (GH). Huancavelica: Prov. Tayacaja, entre Colcabamba y Pau- cartambo, Tovar 2005 (GH). Ayacucho: Between Huanta and Rio Apurimac, Killip & Smith 23258 (F, us). Apu- rimac: Prov. Andahuaylas, Dist. Chincheros, Saunders 736 (GH). Cuzco: Prov. La Convencion, Dist. Vilcabam- ba, Davis et al. 1217 (F, GH). Puno: Agapata, Lechler 2025, in part (GH). Near Puno, Soukup 218 (F). 16. Blechnum sprucei C. Chr., Index fil. 160. 1905, nom. nov. for Lomaria caudata Baker, not Blechnum caudatum Cav., 1802. Lomaria caudata Baker, in Hooker & Baker, Syn. fil. 1 79. 1 967. TYPE: Andes of Ecuador, Spruce 5329 (holotype, K, photo, Svensk Bot. Tidskr. 420. 1954). Stem rather stout, nearly erect to decumbent and very short-creeping. Leaves dimorphic. Pet- iole with basal scales few to many, rather large, concolorous, lanceolate to lanceolate-ovate or broader, stem scales light to dark brown often with lighter margins. Sterile lamina 1 -pinnate, the apex gradually reduced, much prolonged with small pinnae and with 1 or more proliferous buds (al- ways?), the base gradually reduced, pinnae strictly patent, sessile, truncate or often somewhat biau- riculate, the margins flat, distant from the costa. Fertile pinnae with green tissue extending hardly or only slightly beyond the attachment of the in- dusium. Undoubtedly in Peru. Costa Rica; Ecuador; Peru south to northwest Argentina and east to southeast Brazil. The much prolonged apex of the sterile lamina makes this a very distinctive species. 17. Blechnum schomburgkii (Klotzsch) C. Chr., Index fil. 157. 1905. Lomaria schomburgkii Klotzsch, Linnaea 20: 346. 1847. TYPE: "British Guiana," Schomburgk 1162 (B). Stem stout, arborescent to 3 m tall, to decum- bent. Leaves dimorphic. Petiole with basal scales very numerous, long (to 3.5 cm or more), acicular, the base usually broader, curved, mostly bicolor- ous with a blackish or dark center and lighter, brown margins, stem scales dark brown or with lighter margins or light brown or brown. Sterile lamina 1 -pinnate or pinnatisect distally, the apex abruptly reduced to a large, entire segment, the base hardly to somewhat reduced, the pinnae pa- tent to somewhat ascending, sessile to short-stalked or partly adnate distally, cuneate, truncate, broad- ly rounded to cordate, the margins flat to involute, distant from the costa. Fertile pinnae with green tissue extending hardly or only slightly beyond the attachment of the indusium. Usually terrestrial, rarely rupestral, cloud forests and elfin forests, or in open sites, seepage slopes, grassy hillsides, or at the edge of rocks, 1 450-3400 m, Cajamarca south to Cuzco. Costa Rica; Venezuela (Roraima) west to Co- lombia, south to Bolivia; southeastern Brazil. The leaves are often few and the pinnae are acute to long-acuminate, and the tip is flat or nearly so. Cajamarca: Namora, Correll & Smith P897 (GH). Amazonas: Prov. Chachapoyas, entre Leimebamba y Balsas, Lopez et al. 4446 (GH). Prov. Bongara, 3 km S 66 FIELDIANA: BOTANY of Pomacocha. Wurdack 998 (F. GH). La Libertad: Prov. Bolivar, Chomparen, Lopez & Sagdstegui 3207 (GH). Huanuco: SW slope of Rio Llullapichis watershed, ascent of Cerros del Sira. Dudlev 13417 A (GH). 13467 (GH), 13469 (GH). Mito. Macbride 3408 (F). Cuzco: Prov. La Convencion. Cordillera Vilcabamba, Dudlev 10975 (GH), 11118(GH), 11135(GH, MO). 18. Blechnum auratum (Fee) Tryon & Stolze, comb. nov. Stem arborescent, often to ca. 1 m tall, some- times multicipetal, stout. Leaves dimorphic or monomorphic. Petiole with basal scales usually many, to 3.5 cm long, curved, acicular beyond the usually broader base, bicolorous with a blackish center and lighter margins, or concolorous, brown, stem scales long, narrow, dark brown to blackish or center so but narrow margins lighter. Sterile lamina 1 -pinnate or pinnatisect distally, the apex abruptly reduced to a large, entire terminal seg- ment, the base not or hardly reduced, to gradually reduced to rather small pinnae, pinnae usually strongly ascending, a few distal ones partly to fully adnate, others sessile to short-stalked, cuneate to broadly rounded to cordate. Fertile pinnae with green tissue extending hardly or only slightly be- yond the attachment of the indusium. Costa Rica; Venezuela and Colombia; south to Bolivia. The stem is arborescent with a dense crown of stiffly straight leaves, the pinnae are strongly as- cending to rarely only slightly so, and their apex is more or less obtuse, with the tip involute or rarely only slightly involute. Key to Subspecies of Blechnum auratum a. Leaves dimorphic, the sterile pinnae broader than the fertile ones; margins of the sterile pinnae involute but not to the costa abaxially ssp. auratum a. Leaves monomorphic, the fertile and sterile pinnae of essentially the same width; margins of the sterile pinnae very strongly involute, reaching to or nearly to the costa abaxially . . . ssp. columbiense 1 8a. Blechnum auratum ssp. auratum. Lomaria aurata Fee. Mem. foug. 8: 71. 1857. TYPE: Colombia. Ocana. Schlim 394 (holotype, p?). Blechnum buchtienii Rosenst., Repert. Spec. Nov. Regni Veg. 5: 231. 1908. TYPE: Bolivia, Un- duavi, Buchtien 878 (holotype, not located; iso- type, us). In open sites, wet hillsides, grasslands, rocky ground, and boggy places, 1900-3400 m. Piura south to Puno. Range of the species. Piura: Prov. Huancabamba, above Huancabamba. Hutchison 1620 (GH). Amazonas: Prov. Bongara, Dist. Yambrasbamba, Tillet 673-340 (GH, USM). Balsas to Lei- mebamba, Barro Negro, E. E. Smith (habitat photo, GH). San Martin: Rio Abiseo National Park, Young 1981 (HUT, USM). Prov. Chachapoyas. south side of Molino- pampa-Disan pass, Wurdack 1614 (USM). Pasco: Prov. Oxapampa, Santa Barbara, D. Smith 8102 (F). Junin: Prov. Satipo, Dist. Pampa Hermosa, Sounders 1030 (GH), 7066 (GH). Cuzco: Prov. La Convencion, Cordillera Vil- cabamba, Dudley 11031 (GH). Prov. Urubamba. near Sayacmarco. Peyton & Peyton 290 (GH). Paucartambo. Soukup 375 (F). Tres Cruces, Gentry el al. 23440 (F). Puno: Prov. Sandia, Vargas 14856 (GH). Prov. Sandia. near Limbani. Metcalf 30519 (GH). 18b. Blechnum auratum ssp. columbiense (Hi- eron.) Tryon & Stolze, comb. nov. Blechnum columbiense Hieron., Hedwigia 47: 244, /. 5,f. 17. 1908. TYPE: Colombia, Huila. Paramo de Moras, Stubel 1268 (holotype, B). Blechnum columbiense var. bogotense Hieron.. Hed- wigia loc. cit. 244. 1908. SYNTYPES: Colombia. Bogota. Karsten (B), Lindig 27 (B): isosyntype, Lindig 27 (BM; photo. GH). Elfin forests and jalca, 2700-3300 m, Piura and Amazonas. Venezuela and Colombia, south to Peru. D. Smith 7720 (USM), Dept. Pasco, is this variety or near it. Piura: Prov. Huancabamba, Hutchison 1619 (F, GH). Amazonas: 20 km downstream from Balsas. Madison 1138 (GH). La Jalca. Chachapoyas. LI. Williams 7570 (F). 19. Blechnum obtusifolium Ettingsh.. Denkschr. Kaiserl. Akad. Wiss. Wien. 23: 59, /. 8. f. 4. 1864. Also Ettingsh., Farnkr. Jetzw. 122. 1865. Lomaria obtusifolia Presl, Tent. Pterid. 143. 1836. nom. nud. Since Ettingshausen uses Presl's name, the specimen cited by Presl. "Herb. bras. reg. be- rol. 101" may be considered to be the type (not located). / Stem more or less erect, massive, becoming ar- borescent. Leaves dimorphic. Petiole with basal TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 67 scales very numerous, long (to 3.5 cm or more), acicular beyond a usually broader base, usually bicolorous with a blackish center and lighter, brown margins, or mostly concolorous, brown, stem scales very long and narrow, often curved, attenuate, brownish often with dark brown to blackish cen- ter. Sterile lamina pinnatisect distally, 1 -pinnate proximally, the apex abruptly reduced to an entire, usually large terminal segment, the base gradually reduced to small pinnae or lobes, pinnae patent to ascending, sessile with a cuneate to rounded base proximally or broadly sessile or partly adnate there, margins flat to somewhat involute, distant from the costa. Fertile pinnae with green tissue extending hardly or only slightly beyond the at- tachment of the indusium. The status and nomenclature of this species are uncertain, the group of species related to Blech- num magellanicum being in need of revision. The description of Ettingshausen is not wholly satis- factory; additional characters are mentioned by Christ, Bull. Herb. Boisier 2, 4: 1091. 1904, and also by Hieronymus, Hedwigia 47: 243. 1908. Two collections, wet forest and forest border, 550 and 220 m, Cuzco, and Madre de Dios. Peru and northwestern Argentina; southeastern Brazil; range uncertain. Cuzco: Prov. Paucartambo, Pillawata, Vargas 16698 (GH). Madre de Dios: Prov. Manu, Shintuya, Chavez 834 (fertile), 835 (sterile) (MO). Comments One collection of what may be a new species of subgenus Blechnum has not been identified: Ama- zonas, Prov. Bagua, Cordillera Colan, Barbour 3589 (MO). The specimen is evidently the central portion of a large fertile leaf. The long and narrow fertile pinnae are short-stalked and bear costal sori with expanded green tissue well beyond the at- tachment of the indusium. The following names have not been associated with a Peruvian species of Blechnum for the rea- sons stated. Blechnum ciliatum Presl, Reliq. Haenk. 1: 50. 1825. "Hab. in Peruvia et in Cordilleris Chilen- sibus." Under Parablechnum ciliatum (Presl) Presl, Epim. bot. 109. 1851, only Chile is cited. Placed by Murillo (1968) as a synonym of Blechnum au- riculatum Cav. Blechnum cognatum Presl, Epim. bot. 107. 1851. "Habitat in vallibus Cordillerae Peruviae (Haenke)." Murillo (1968) places this name, ex char, as a synonym of Blechnum occidentale L. Blechnum delicatum Maxon & Morton, Bull. Torrey Bot. Club 66: 41. 1939. TYPE: Robore, Bolivia, Cardenas 3006 (holotype, us!) This seems to be an unusual variation of Blechnum asple- nioides. Blechnum lechleri Mett., Fil. Lechl. 2: 17. 1859. "Peruvia, Tatanara; Brasilia (Bongard)." I have not examined authentic material of this species. It may be Blechnum cordatum. Lomaria linariaefolia Presl, Reliq. Haenk. 1 : op. cit. 52. 1825. "Hab. in Peruviae?" Blechnum malacothrix Maxon & Morton, Bull. Torrey Bot. Club 66: 40. 1939. TYPE: Okara, Cor- dillera Real, Bolivia, Tate 951 (holotype, NY; iso- type, us!). This may be a valid species, or it may be Blechnum confluens. Blechnum trilobum Presl, loc. cit., t. 9,f.2.\ 825. "Hab. in Peruvia." The figure is Blechnum auri- culatum Cav. of Chila NE to southern Brazil. II. Salpichlaena Salpichlaena Hooker, Gen. fil., /. 93. 1842, as Sal- pichloena in text, corrected in index. TYPE: Salpichlaena volubilis (Kaulf.) Hooker (Blech- num volubile Kaulf.). Figure 4. Terrestrial. Stem long-creeping to short-creep- ing and moderately stout. Leaves dimorphic or partly dimorphic, the fertile pinnae with narrower, sometimes very narrow segments, scrambling or climbing to 15 m or more long, indeterminate. Lamina 2-pinnate, the pinnae imparipinnate. Veins free, connected by a marginal strand. Sori on a long vascular commissure close to the costa, not paraphysate, covered by an indusium of similar extent, arching completely over the sporangia, and opening toward the costa. Spores ellipsoidal, monolete, with spherical deposit over the papil- late-rugose surface. FIG. 4. Salpichlaena volubilis: a, sterile pinna; b, fertile pinnule; c, section of pinnule with sori. (From Plowman 2550, F.) 68 FIELDIANA: BOTANY 5cm TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 69 Salpichlaena is a widely distributed tropical American genus of one species. The leaves of ju- venile plants are entire or 1 -pinnate, and deter- minate, while those of adult plants are 2-pinnate, indeterminate, and scrambling or high-climbing in trees. 1. Salpichlaena volubilis (Kaulf.) Hooker, Gen. fil., t. 93. 1842. Figure 4. Blechnum volubile Kaulf., Enum. fil. 159. 1824. TYPE: perhaps Brazil, Chamisso (holotype, LZ, de- stroyed: no isotype located). Blechnum scandens Bory, in Duperrey, Voy. Coquille Bot. 272, t. 36. 1 829. TYPE: Brazil, St. Catharina, D'Urville (holotype, P; isotype, GH!). Lomaria volubilis Hooker, Sp. fil. 3: 39. 1860. TYPE: Brazil, Barra do Rio Negro, Spruce 1263 (holo- type, K; isotype, ?, GH!). Blechnum volubile var. lomarioideum Baker in Mar- tius, Fl. Brasil. 1(2): 428. 1870. Spicanta hookeriana O. Kuntze, Revis. gen. pi. 821. 1891. Nom. nov. for Lomaria volubilis Hook, and with the same type. Salpichlaena hookeriana (O. Kuntze) Alston, Bull. Misc. Inform 1432: 312. Salpichlaena lomarioidea (Baker) A. R. Sm., Ann. Missouri Bot. Gard. 77: 250. 1990. Stem scales small, rigid, dark brown to blackish. Pinnae usually subopposite. Segments acute, nar- rowly acuminate to caudate, entire or often serrate at the tip, sessile to rather long-stalked, cuneate to broadly rounded at the base, the margins cartilagi- nous, the costa prominently abaxially, the sterile ones ca. 1.5 mm to 3 cm broad and 10-30 cm long. Sori costal, very long. Terrestrial in rain forests, montane forests, in- undated thickets and forests near rivers, rarely in cloud forests, 100-1900 m, mostly below 700 m, Amazonas and especially Loreto, south to Cuzco. Central America; Lesser Antilles; south to Bo- livia and southeastern Brazil. The chromosome number of n = 40 is unique in the family. This is a very distinctive genus and species, with a long scrambling or climbing leaf. The habit is the same as that ofLygodium species. Rarely the fertile and sterile segments are of es- sentially the same breadth. It is probably more widely distributed than the collections indicated since the climbing leaves may be partly concealed among tree branches, or the plant may not be rec- ognized as a fern. In some specimens, the fertile segments are very narrow, with little or no green tissue extending beyond the attachment of the indusium. These have been named Salpichlaena hookeriana— for example, Plowman 2550 and McDaniel 11758, both Dept. Loreto (GH), are this narrow variant. Some specimens have narrow segments and only narrow green tissue beyond the indusium. Buds may be present in the axils of the pinnae (less often the axils of the pinnules), and these occur both in specimens with narrow and broad fertile segments. The narrow variant seems restricted to low ele- vations, while the specimens with broader seg- ments are also from higher elevations. Further study is required before another taxon is recog- nized with confidence. Amazonas: Rio Maranon, above Cascadas de Mayasi, Wurdack 1893 (F, GH). Prov. Bagua, 1 2 km E of La Peca, Barbour 2402 (F, MO). E of La Peca, Gentry et al. 22905 (F, USM). San Martin: Zepelacio near Moyobamba, Klug 3514 (F, GH, MO). Prov. Mariscal Caceres, Dist. Tocache Nuevo, J. Schunke V. 5541 (F, MO), 7079 (F, USM). Lo- reto: Sierra del Pongo, Mexia 6276 (F, GH, MO). Pumaya- cu, Klug 3 197 (F, GH, MO). Shiriara, Rio Nanay, Plowman 2550 (F, GH). Versailles, Iquitos, McDaniel 1 1758 (F, GH, MO). Huanuco: Tingo Maria, Allard 21386 (GH). Pasco: Prov. Oxapampa, Rio Pichis, 1 hr below Puerto Ber- mudez, Gentry et al. 42123 (F). Prov. Oxapampa, N of Chequitavo, D. Smith 5118 (MO). Junin: La Merced, Soukup 1 106 (F). Near La Merced, Killip & Smith 23923 (F). Above San Ramon, Killip & Smith 25565 (F). Cuzco: Prov. La Convencion, Cordillera Vilcabamba, Dudley 10451 (GH, MO, USM). Family 21. POLYPODIACEAE POLYPODIACEAE Bercht. & J. S. Presl, Priroz- en. Rostlin 1: 272. 1820. TYPE: Polypo- dium L. Stem erect to short-creeping, or very long-creep- ing, small to rather stout, or slender to filiform, with scales. Leaves ca. 3 cm to 2 m long, circinate in vernation, monomorphic or dimorphic, entire, pinnatifid, or variously forked, or pinnate, gla- brous, pubescent or scaly abaxially. Petiole lacking stipules, often articulate to the stem. Veins free to anastomosing. Sori abaxial, round to elongate, borne on a vein, at the tip of a vein, at the junction of veins, on a vascular commissure, or on a special series of fertile veins, exindusiate, paraphysate or not. Sporangia with a 1 - or 2-rowed stalk basally, the vertical annulus interrupted by the stalk. The Polypodiaceae are a large family of perhaps 40 genera and 1 000 species, most of these being 70 FIELDIANA: BOTANY epiphytic or epipetric. The classification of this group is difficult, and authors often disagree on the taxa to be recognized and their rank. Three tribes are adopted here, each of these sometimes being treated as a family. The tribe Polypodieae usually has the petiole sulcate and articulate to the stem, the petiole and lamina have short trichomes or none, the sporan- gium has two rows of cells at its base, and the spores are ellipsoidal, monolete and only rarely green. There are nine genera in Peru. The tribe Loxogrammeae is intermediate be- tween the two other tribes, having some characters of one and some of the other. It does not occur in Peru. The tribe Grammitideae usually has the petiole terete and continuous (not articulate) with the stem, the petiole and lamina often with long trichomes, the sporangium has a single row of cells at its base, and the spores are usually green and usually tet- rahedral-globose, trilete. It is represented in Peru by the single genus, Grammitis. References COPELAND, E. B. 1947. Polypodiaceae, pp. 174- 222. In Genera Filicum. Chronica Botanica, Waltham, Mass. TRYON, R. M., AND TRYON. 1982. Polypodi- aceae, pp. 684-758. In Ferns and Allied Plants. Springer- Verlag, New York. Key to Genera of Polypodiaceae a. Petiole and usually the lamina with long, multicellular, acuminate trichomes; sporangium stalk uniseriate at base; spores usually with chlorophyll (green) and tetrahedral-globose . . I. Grammitis a. Petiole and lamina without trichomes, or these present and usually unicellular, short; sporangium stalk bi- or triseriate; spores lacking chlorophyll (with rare exception), ellipsoidal, monolete . . . . b b. Lamina dichotomously or subdichotomously forked or veined c c. Lamina with scattered scales V. Dicranoglossum c. Lamina with stalked, stellate trichomes X. Platycerium b. Lamina entire, lobed, or pinnatifid to 3-pinnate d d. Stem bearing tubers; spores prominently echinate; epiphytic IX. Solanopteris d. Stem without tubers; spores nearly smooth, verrucate, rugose, tuberculate, papillate or with folds; terrestrial, rupestral, or epiphytic e e. Lamina pinnatifid, pinnatisect, or to 3-pinnate f f. Stem scales peltately attached III. Polypodium f. Stem scales basally attached II. Pecluma e. Lamina entire g g. Stem more or less flattened, its scales not clathrate, attached near the center VI. Microgramma g. Stem more or less terete, its scales usually clathrate h h. Sori in 2 rows between strong lateral veins, or lateral veins obscure or absent ... i i. Immature sori with peltate paraphyses; most sori served by 2 or more veins; lamina surface scaly (at least sparsely so) IV. Pleopeltis i. Paraphyses absent or minute; most sori borne back of a single vein tip; scales absent on the lamina or only a few on the costa VII. Campyloneurum h. Sori in 1 row between strongly developed lateral veins j j. Stem very long-creeping HI. Polypodium j. Stem more or less erect, decumbent or short-creeping VIII. Niphidium Marginariopsis C. Chr. (Dansk. Bot. Ark. 6(3): 42. 1 920) is an epiphyte that occurs from Ecuador north to Costa Rica and it may grow in Peru. Its stem is rather flattened and bears finely clathrate scales, the leaves are dimorphic, the fertile much narrower than the sterile, the lamina is entire, with small scales, and the sporangia are borne on anas- tomosing veins. Many American species of Polypodium have been placed in the genus Goniophlebium. That name, however, applies to an Old World genus and is typified by Polypodium subauriculatum TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 71 Blume (Goniophlebium subauriculatum (Blume) Presl). I. (.ram in it is Grammitis Sw., J. Bot. (Schrad.) 1800(2): 17. 1801. TYPE: Grammitis marginella (Sw.) Sw. (Polypodium marginellum Sw.). Figure 5. Xiphopteris Kaulf., Berl. Jahrb. Pharm. 21: 35. 1820. LECTOTYPE (designated by John Sm., Hist. fil. 179. 1875: Xiphopteris serrulata (Sw.) Kaulf. Grammitis section Xiphopteris (Kaulf.) Presl, Tent. Pterid. 208. 1836. Cochlidium Kaulf., Berl. Jahrb. Pharm. 21: 36. 1820. TYPE: Grammitis graminoides (Sw.) Sw. (Acros- tichum graminoides Sw., Cochlidium grami- noides (Sw.) Kaulf.) (section Pleurogramme). Ctenopteris Kunze, Bot. Zeit. (Berlin) 4: 425. 1846. TYPE: Ctenopteris venulosa (Blume) Kunze (Polypodium venulosum Blume = Grammitis ven- ulosa (Blume) R. & A. Tryon (section Cryptoso- rus). Cryptosorus Fee, Mem. foug. 5 (Gen. fil.): 231. 1852: TYPE: Cryptosorus blumei Fee, nom. super/], for Polypodium obliquatum Blume = Grammitis ob- liquata (Blume) Hassk. Polypodium section Cryp- tosorus (Fee) Fourn., Ann. Sci. Nat. Bot., ser. 5, 18: 282. 1873. Grammitis section Cryptosorus (Fee) R. & A. Tryon, Rhodora 84: 128. 1982. Glyphotaenium (John sm.) John Sm., Hist. fil. 187. 1875. Ctenopteris subgenus Glyphotaenium John Sm., Bot. Voy. Herald (Seemann) 227. 1854. TYPE: Ctenopteris crispata John Sm. (Glypho- taenium crispatum (John Sm.) John Sm.) = Grammitis crispata (John Sm.) Morton. Gram- mitis section Glyphotaenium (John Sm.) R. & A. Tryon, Rhodora 84: 128. 1982. Enterosora Baker, Timehri 5: 218. 1886. TYPE: En- terosora campbellii Baker = Grammitis camp- bellii (Baker) Proctor (section Glyphotaenium). Micropolypodium Hayata, Bot. Mag. Tokyo 42: 302, 341. 1928. TYPE: Micropolypodium pseudotri- chomanoides (Hayata) Hayata. Ceradenia Bishop, Amer. Fern J. 78: 2. 1988. TYPE: Polypodium curvatum Sw. (Ceradenia curvata) (Sw.) Bishop) = Grammitis curvata (Sw.) Ching. Zygophlebia Bishop, Amer. Fern J. 79: 107. 1989. TYPE: Polypodium sectifrons Mett. (Zygophlebia sectifrons (Mett.) Bishop) = Grammitis sectifrons (Mett.) Seymour. Lellingeria Smith & Moran, Amer. Fern J. 81: 76. 1991. TYPE: Polypodium apiculatum Klotzsch (Lellingeria apiculata (Klotzsch) Smith & Mor- an). Plants commonly epiphytic, occasionally epi- petric or terrestrial. Stem small, mostly short- creeping, sometimes erect, commonly scaly (scales clathrate or not), rarely naked, bearing slender, fibrous roots. Leaves monomorphic or rarely the apical fertile portion modified, 2-60 cm (rarely to 1 m) long, rarely articulate to the stem, scales lack- ing, or present only at petiole base, petiole long to obsolete. Lamina simple and entire to pinnatisect (very rarely to 3-pinnate), glabrous or variously pubescent (rarely densely so), the trichomes 1- few-celled. Veins commonly free, sometimes anas- tomosing and with or without free, included vein- lets. Sori round to elongate, sometimes confluent, borne on the vein or vein branch, sometimes deep- ly immersed in the laminar tissue, or sporangia in lines on each side of the costa, paraphysate or not, exindusiate. Spores trilete, tetrahedral-globose, greenish (chlorophyllous). Grammitis is a pantropical and austral genus of more than 400 species, with perhaps half of these in America. From the time of Copeland's (1947) "Genera Filicum" to the present it has been di- vided by various authors into as many as 20 genera or subgenera, and some of these divisions may be warranted. Alan R. Smith and Robbin C. Moran are currently attempting a revision of the Amer- ican species that will recognize nine genera, the concepts partially based on the recent work of Bishop (1977, 1978, 1988, 1989a). The authors have kindly provided provisional manuscripts of these genera, which have been very helpful in con- structing a key to the species in Peru. Nevertheless, pending the results of these new studies, Gram- mitis is recognized in the broadest sense for pur- poses of the present treatment. The key to species in Peru is based in part on the generic divisions ultimately to be proposed by Smith and Moran. For example, species 1-3 are Grammitis s.s., essentially defined in the Neo- tropics by the dark, sclerenchymatous border of the lamina, and species 7-9 share the characters of Enterosora (= Glyphotaenium): lamina subcar- FIG. 5. Grammitis myosuroides: a, habit. Grammitis limbata: b, habit; c, leaf section, with sori. Grammitis david- smithii: d, habit; e, pinna, adaxial side; f, stem scale. Grammitis myriophylla: g, section of rachis with fertile pinna base, (a from Camp E-5131, Ecuador, F; b from Steyermark & Wurdack 1 178, Venezuela, F; c from D. Smith 2038, F; d from Beck 1424 'A, Bolivia, F; e, f from Smith & Canne 5837, holotype, F; g from Leon 1005, F.) 72 FIELDIANA: BOTANY TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 73 nose and entire to lobed, with sori immersed in the tissue, hydathodes lacking. The character of hydathodes is effective in sep- arating some major species groups, in addition to Glyphotaenium. In many species, hydathodes are easily visible to the naked eye— usually seen as expanded vein tips differing in color from the ad- jacent tissue. In other species, closer scrutiny is necessary; i.e., hydathodes may be hidden by dense indument, or they may be located at the very seg- ment margin, the adaxial portion of which is strongly revolute, or they may be less conspicuous to the naked eye, yet quite evident with slight mag- nification (10-12x). Besides the group of Enterosora, two other an- hydathodous groups are to be recognized by Smith and Moran, these separated by a presumably good and consistent character: sori with conspicuous, pale, waxlike paraphyses (Ceradenid) vs. paraph- yses lacking, or dark and not waxlike (Zygophle- bid). Unfortunately, the presence and character of the paraphyses often cannot be ascertained in ma- ture sori nor in heat-dried specimens (Bishop, 1988, p. 2). Species of Zygophlebia are further charac- terized by strongly to casually anastomosing veins (free in Ceradenid), but in two of three species in Peru (G. dudleyi, G. werffii) the lamina texture is so thick that veins can only be examined by the lengthy process of clearing. Thus, the two most important distinguishing features of these two groups are in most cases functionally ineffective insofar as rapid field or herbarium identification is concerned. Species 29-66, with hydathodes, are much more difficult to separate into discrete groups; however, the characters used in the key that best seem to indicate species relations are those of stem scales, laminar trichomes, and sporangia (capsular setae present or lacking). Several species complexes in this treatment are still very poorly understood, and relationships of their components are sorely in need of study. The groups most in need of attention are those of G. cultrata/heteromorpha, G. semihirsuta, G. monil- iformis, and G. flabelliformis/pilosissima, each of these probably containing four to eight taxa. The species delineations herein seem to serve rather well insofar as Peruvian collections are concerned, but many of the taxa vary or intergrade consid- erably in Mexico and Central America, the West Indies, and other parts of South America. Most of the problems are pointed out in the respective species discussions that follow. References BISHOP, L. E. 1977. The American species of Grammitis section Grammitis. Amer. Fern J., 67: 101-106. . 1978. Revision of the genus Cochlidium (Grammitidaceae). Amer. Fern J., 68: 76-94. -. 1988. Ceradenia, a new genus of Gram- mitidaceae. Amer. Fern J., 78: 1-5. . 1989a. New species of Ceradenia. Amer. Fern J., 79: 14-25. . 1989b. Zygophlebia, a new genus of Grammitidaceae. Amer. Fern J., 79: 103-1 18. BISHOP, L. E., AND A. R. SMITH. 1992. Revision of the fern genus Enterosora (Grammitidaceae) in the New World. Syst. Bot., 17: 345-362. CHRISTENSEN, C. 1929. Taxonomic fern studies. I. Revision of the polypodioid genera with lon- gitudinal coenosori (Cochlidiinae and "Dry- moglossinae"; with a discussion of their phy- logeny. Dansk. Bot. Ark., 6(3): 1-93. COPELAND, E. B. 1952a. Grammitis. Phillip. J. Sci., 80:93-271. . 1952b. The American species of Xiph- opteris. Amer. Fern J., 42: 41-52, 93-1 10. . 1956. Ctenopteris in America. Philipp. J. Sci., 84:381-473. MAXON, W. R. 1914. Notes upon Polypodium duale and its allies. Contr. U.S. Natl. Herb., 17: 398-406. . 1916. Polypodium trichomanoides and its American allies. Contr. U.S. Natl. Herb., 17: 542-557. MORTON, C. V. 1967. The genus Grammitis in Ecuador. Contr. U.S. Natl. Herb., 38: 85-123. SMITH, A. R., R. C. MORAN, AND L. E. BISHOP. 1 99 1 . Lellingeria, a new genus of Grammiti- daceae. Amer. Fern J., 81: 76-88. Key to Species of Grammitis a. Lamina simple, entire, with a black or dark brown, sclerenchymatous border b b. Lamina less than 3 mm broad; stem scales 2-6 cells across at base 3. G. paramicola b. Lamina 3-8 mm broad; stem scales 8-12 cells across at base c c. Hydathodes present; sclerenchymatous border commonly less than 0. 1 mm broad; lamina firm- herbaceous to chartaceous . . 2. G. limbata 74 FIELDIANA: BOTANY c. Hydathodes lacking; border commonly 0.2 mm broad; lamina subcoriaceous 1 . G. bryophila a. Lamina lobed to decompound or, if simple, then lacking a dark, sclerenchymatous border d d. Lamina (at least fertile portions) simple and subentire to repand or serrate-dentate, 1-2 mm broad, the son confluent and commonly covering the costa at maturity e e. Leaves monomorphic, fertile and sterile portions entire throughout, ca. 1 mm broad 4. G. pumila e. Leaves partly dimorphic, fertile distal portion subentire, sterile leaf or portion serrate-dentate to pinnatisect, to 4 mm broad f f. Lamina trichomes castaneous, scattered but persistent at least on costa and veins; sporangia borne at base of veins, confluent as 2 subcostal lines until merging into a broad single line at full maturity; sterile portion of lamina pennatisect 5. G. myosuroides f. Lamina trichomes lacking (or pale and caducous on juvenile plants); sporangia borne on costa and veins, thus confluent as a single, costal line; sterile portion of lamina serrate- dentate or with short, acute lobes 6. G. serrulata d. Lamina pinnatisect to decompound throughout, or if simple to shallowly lobed then 4-20 mm broad and sori discrete, in 1 or more lines on each side of, and subdistant from, the costa . . . g g. Hydathodes lacking, or minute and discernible only with high (more than 20 x ) magnification h h. Lamina simple, subentire or lobed to pinnatifid (not cut nearly to the costa) i i. Leaves 1-3.5 cm broad; stem scales yellow or yellowish brown; sori in (l-)2-5 series between costa and margin j j. Leaves 2-3.5 cm broad, cut Vj-'/z to the costa; petiole lustrous black or atropurpureous; sori in 3-5 series between costa and margin 7. G. trifurcata j. Leaves 1-1.5 cm broad, subentire to very shallowly lobed; petiole dull yellowish or reddish brown; sori in (l-)2 series between costa and margin .... 8. G. trichosora i. Leaves 0.4-0.8 cm broad; stem scales deep orange to castaneous; sori in a single series between costa and margin k k. Stem scales deep orange, entire or remotely ciliolate; lamina base attenuate; petiole trichomes castaneous 9. G. parietina k. Stem scales castaneous, conspicuously setose; lamina base cuneate; petiole trichomes orange 10. G. phalacron h. Lamina pinnatisect (cut nearly to the costa) to 3-pinnate 1 1. Lamina 2-3-pinnate; ultimate segments 0.7-3 mm long m m. Stem scales clathrate, margins entire or with some short glands near apex; petiole and lamina puberulent, but lacking long, spreading, castaneous trichomes 24. G. in i rahi I is m. Stem scales nonclathrate, margins abundantly setose; petiole and lamina with spread- ing castaneous trichomes to 1.5 mm long (and puberulent as well) 28. G. bipinnata 1. Lamina 1 -pinnate or deeply pinnatisect; ultimate segments 5 mm long or (usually) much longer n n. Veins regularly merging to form usually quite evident areoles, at least when held to light 25. G. mathewsii n. Veins free, or, if casually anastomosing, then completely obscure, even when held to light o o. Lamina not or scarcely reduced at base, or basal pair of segments more than half as long as median ones; petiole about half as long as or longer than the lamina . P p. Pinnae, at least proximal ones, deeply serrate, the others crenulate 11. G. longipinnata p. Pinnae subentire q q. Stem scales rather flaccid, pale orange or yellow-brown, entire, lacking mar- ginal setae; plants epiphytic 26. G. dudleyi TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 75 q. Stem scales rigid, deep orange to castaneous, with conspicuous marginal setae; plants terrestrial r r. Rachis and costae (and usually segments) amply provided on both sides with long, spreading trichomes, these 0.5-2 mm long 12. G. meridensis r. Rachis (and usually costae) densely short-puberulent adaxially, the tri- chomes 0. 1-0.3(-0.4) mm long s s. Setae of stem scales dark, the same color as the scale body; stem long- creeping, the leaves mostly wide spaced 13. G. bishopii s. Setae of stem scales whitish, in strong contrast with the dark scale body; stem short-creeping to ascending, leaves approximate or crowded . . . 27. G. werffii o. Lamina somewhat to strongly reduced at base, several to many proximal segments much shorter than medial ones or reduced to mere auricles; petiole obsolete or much shorter than the lamina t t. Petiole lacking conspicuous, spreading trichomes u u. Petiole essentially obsolete; sclerenchyma of the rachis and costae obscured by the thick lamina tissue v v. Lamina segments 1-2 times as long as broad; lamina moderately to densely white-farinose, the farina mostly disposed in stellate patterns 14. G. discolor v. Lamina segments (larger ones) 3-7 times as long as broad; lamina mod- erately farinose to naked, the farina deposits solitary or in amorphous clusters 15. G. curvata ssp. pearcei u. Petiole (1-) 2- 10 cm long; sclerenchyma of the rachis (and sometimes costae) exposed, at least proximally w w. Lamina pinnatisect, segments closely spaced, joined by at least a narrow wing of tissue, gradually narrowing from longest medial pinnae to nearly auricular ones at base x x. Stem scales 3-5 mm long, setulose; segments ascending at a 30-50° angle; sori medial 16. G. terrestris x. Stem scales ca. 1 mm long, subentire; segments ascending at a 60-80° angle; sori supramedial 17. G. herrerae w. Lamina 1 -pinnate, pinnae adnate, but discrete and mostly subdistant, abruptly reduced at base, with only ca. 3-6 pinnae shorter than medial ones y y. Pinnae 1-3.5 cm long, 2.5-5 mm broad, ascending at a 60-80° angle, base rounded to truncate acroscopically; sori generally confined within pinna margins 19. G. farinosa y. Pinnae to 10 cm long, ca. 1.5 mm broad, ascending at a 20-35° angle, attenuate at base; sori at maturity extending well beyond pinna margins 18. G. dendrodoxa t. Petiole (and often the rachis and lamina) amply provided with orange to cas- taneous, spreading, unicellular trichomes, these 0.5-3 mm long z z. Lamina 0.6-1 cm broad; pinnae circular, obovate or broadly oblong, obtuse at the apex, 1-2 times as long as broad 20. G. congesta z. Lamina 2-10 cm broad; pinnae linear to deltate, acute to attenuate at the apex, several to many times longer than broad a2 a2. Pinna base rounded nearly to the costa acroscopically, short-decurrent basiscopically; sori not extending beyond the pinna margin 21. G. praeclara a2. Pinna base fully adnate and nearly perpendicular to the rachis acroscopi- cally, long-decurrent basiscopically; sori mostly extending beyond the pinna margin at maturity b2 76 FIELDIANA: BOTANY b2. Lamina thin- to firm-herbaceous; rachis evident and usually lus- trous, at least abaxially; petiole 0.2-0.4(-0.5) mm in diameter . . . 22. G. capillaris b2. Lamina subcarnose; sclerenchyma of the rachis usually obscured by laminar tissue, or rarely evident and then not lustrous; petiole (0.4-)0.6-0.9 mm in diameter 23. G. pilipes g. Hydathodes evident (or in a few species hidden by dense trichomes) c2 c2. Trichomes on petiole and/or lamina 0.1-0.3 mm long or lacking and the stem scales conspicuously setose d2 d2. Lamina subtruncate at base, or a few proximal pairs of pinnae about half as long as medial ones 30. G. apiculata d2. Lamina attenuate at base, 2-6 proximal pairs of pinnae rudimentary and auriculiform e2 e2. Rachis thin and conspicuously flexuous, sparsely puberulent adaxially; pinna base inequilateral, rounded or truncate acroscopically, decurrent basiscopically; pinnae ascending at a 20-45° angle 31. G. pseudocapillaris e2. Rachis stout and rigid, not or slightly flexuous, densely puberulent adaxially; pinna base subequilateral, either subtruncate or dilated; pinnae patent or slightly as- cending (often strongly ascending in G. subsessilis) f 2 f2. Mature leaves 0.8-1.5 cm broad; petiole 0.3-0.4 mm in diameter; pinnae oblong, 3—4 times longer than broad 32. G. tunguraguae f2. Mature leaves 2-7 cm broad; petiole (0.4-)0.5-1.2 mm in diameter; pinnae linear to deltate, 6-12 times longer than broad g2 g2. Pinnae often strongly ascending, linear from a strongly dilated base; stem scales 2-4 mm long, obviously clathrate, the proximal cells in 5-10 series across the scale and the lumina clear 33. G. subsessilis g2. Pinnae patent to slightly ascending, deltate or oblong-deltate; stem scales (4— )5-7 mm long, narrowly clathrate, the proximal cells in (12-) 15-20 series across the scale and the lumina mostly occluded . . 34. G. major c2. Trichomes on petiole and/or lamina (0.4— )0. 5-2 mm long or, if shorter or lacking, then stem scales entire h2 h2. Trichomes on petiole and/or lamina 0.1-0.3 mm long or lacking (occasionally a few, scattered, longer ones present, especially on the petiole in #40) i2 i2. Pinnae deeply lobed to pinnatifid, most of them ascending at a 30—45° angle .... 35. G. gracilis i2. Pinnae with margins entire to crenulate, patent or slightly ascending j2 j2. Pinnae conspicuously inequilateral at base (i.e., broadly rounded to subauricu- late acroscopically, strongly decurrent basiscopically); lamina thin-herbaceous 36. G. phlegmaria j2. Pinnae subequilateral, equally dilated at base, or basiscopically somewhat more spreading; lamina chartaceous to coriaceous k2 k2. Pinnae or segments ascending at a 60-70° angle, linear-deltate, 6-8 times as long as broad 37. G. youngii k2. Pinnae or segments patent, semicircular to oblong or deltate, l-3(— 4) times as long as broad 12 12. Laminar tissue amply provided with appressed, whitish, viscid tri- chomes, these often obscuring the hydathodes 29. G. jamesonioides 12. Laminar tissue essentially glabrous, the hydathodes conspicuous .... m2 m2. Stem scales l-2(-2.5) mm long, not iridescent, cell walls blackish; segments plane 38. G. melanosticta m2. Stem scales 3-8 mm long, iridescent, cell walls brown or reddish brown; segment margins (at least fertile ones) revolute n2 TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. n2. Leaves well spaced on the stem; segments 2—4 times as long as broad; plants not aromatic 39. G. erecta n2. Leaves commonly approximate to subfasciculate; segments 1-1.5 times as long as broad; plants (especially dried) with a resinous scent 40. G. moniliformis h2. Trichomes on petiole and/or lamina (0.4-)0.5-2 mm long o2 o2. Leaves 1 -pinnate and 1-2-forked 50. G. heteromorpha o2. Leaves pinnatisect to 2(-3)-pinnate, not (or very rarely) forked p2 p2. Pinnae crenate or lobed (at least near the base) to l(-2)-pinnate q2 q2. Pinnae crenate to deeply lobed; stem scales black or dark brown 64. G. athyrioides q2. Pinnae deeply pinnatisect to 1 (-2)-pinnate; stem scales orange or light brown r2 r2. Ultimate segments linear; margins of stem scales entire or with a few short glands; sporangia not setose 65. G. myriophylla r2. Ultimate segments oblong to nearly circular; margins of stem scales long-setose; sporangia long-setose s2 s2. Hydathodes commonly encrusted with white deposits; pale tri- chomes of rachis (and segments) mixed with castaneous, stel- lately branched ones 66. G. immixta s2. Hydathodes lacking white encrustations; pa> trichomes of rachis mixed with minute (0.1-0.3 mm), septate, gland-tipped ones . 67. G. variabilis p2. Pinnae (or segments) entire t2 t2. Sporangia with long setae; axes with most of the trichomes sericeous and hyaline to pale orange; stem scales setose u2 u2. Lamina deeply pinnatisect, not or scarcely reduced at base or only 1-2 basal pairs greatly reduced; petiole long, conspicuous 48. G. asplenifolia u2. Lamina fully 1 -pinnate, strongly and gradually reduced at base; pet- iole short or obsolete v2 v2. Rachis with stout, castaneous, sessile-stellate trichomes mixed among the simple, sericeous ones w2 w2. Pinnae 0.4-1 cm long, oblong-elliptic, obtuse, most of them narrowed at base; petiole 0.2-0.4 mm in diameter; sporan- gia each with 1 seta 49. G. dependens w2. Pinnae 1 . 1-4.5 cm long, deltate, acute or subacute, broadly adnate at base; petiole 0.4-1 mm in diameter; sporangia with 1-3 setae 5 Ib. G. lanigera var. stella v2. Rachis lacking stout, castaneous, sessile-stellate trichomes (al- though some thin, pale, stellate ones may be present) x2 x2. Pinnae 3-6 mm long; pinna base very constricted and in- equilateral, strongly rounded acroscopically, decurrent basi- scopically 52. G. subflabelliformis x2. Pinnae 10—45 mm long; pinna base broadly adnate and subequilateral, slightly rounded or truncate to somewhat dilated y2 y2. Pinnae 60-80 pairs, linear, 0.2-0.3 mm broad, 12-20 pairs along 5 cm of rachis; rachis trichomes all simple 53. G. laxa y2. Pinnae 20-50 pairs, oblong to deltate, (0.3-)0.4-1.2 mm broad, 4-10 pairs along 5 cm of rachis; rachis with few to many sessile-stellate trichomes mixed among the abundant simple ones z2 z2. Most pinnae slightly deflexed; petiole 0.2-0.3 mm 78 FIELDIANA: BOTANY in diameter; lamina sparsely to moderately pilose; sessile-stellate trichomes sparse on rachis and ab- axial surface 54. G. cultrata z2. Most pinnae patent to slightly ascending; petiole 0.4-1 mm in diameter; lamina densely pilose; sim- ple and sessile-stellate trichomes everywhere abun- dant 5 la. G. lanigera var. lanigera t2. Sporangia lacking long setae; axes with stout, rigid, trichomes (except sericeous in G. taxifolid), stem scales setose or not a3 a3. Stem scales with few to many rigid, marginal setae, these often as long as the width of the scale body b3 b3. Lamina of mature leaves to 1 cm broad; veins solitary in each segment and simple or with one acroscopic branch c3 c3. Petiole 0.5-1 .5 cm long; lamina chartaceous; segments often somewhat gibbous at base acroscopically 55. G. blepharolepis c3. Petiole obsolete or nearly so; lamina thin- to firm-herba- ceous; segments not gibbous 56. G. truncicola b3. Lamina of mature leaves ( 1 . 5-)2-8(- 1 2) cm broad; veins 3-many pairs in a pinna or segment d3 d3. Stem scales gray-brown or black, clathrate or not; sori me- dial to inframedial e3 e3. Stem scales gray-brown to blackish, essentially clath- rate; lamina trichomes mostly 0.4-0.5 mm long; veins 6-14 pairs per segment; hydathodes lacking white de- posits 57. G. david-smithii e3. Stem scales lustrous black, nonclathrate; lamina tri- chomes 1-2 mm long; veins 3-4 pairs per segment; hydathodes mostly encrusted with white deposits . . . 58. G. pichinchensis d3. Stem scales orange to castaneous, not or scarcely clathrate; sori supramedial to submarginal f3 f3. Pinna surface amply and minutely setulose abaxially; lamina not reduced at base, or with 1-3 pairs of pinnae somewhat shortened; rachis trichomes 0.4-0.7 mm long abaxially, shorter adaxially 59. G. alsopteris f 3. Pinna surface glabrous abaxially; lamina conspicuously reduced at base, 3-6 pairs of pinnae much shortened; rachis trichomes mostly 1-2 mm long (sometimes part- ly broken away on older leaves) g3 g3. Pinnae broad- or oblong-deltate, obtuse to sub- acute, 4-6 times as long as broad; veins 6-13 pairs per pinna 60. G. leucosticta g3. Pinnae linear or linear-deltate, acute, 7-10 times as long as broad; veins 1 6-24 pairs on larger pin- nae 61. G. semihirsuta a3. Stem scales with margins entire (scale apex setose in G. taxifolid) or minutely glandular or ciliolate h3 h3. Stem scales less than 1 mm long, few and inconspicuous; stems small (usually less than 1 mm), on long-creeping, stoloniform roots 41. G. anfractuosa h3. Stem scales (l-)2-8 mm long, conspicuous and usually in large masses at least at stem apex; stems 1-5 mm thick, roots not stoloniform i3 i3. Stem scales conspicuously clathrate, gray-brown, narrow- TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 79 delta te or -lanceolate, 8-15 cells across at scale base and the lumina large, clear and elongate j3 j3. Pinnae linear and many of them deflexed, conspicuously inequilateral at base, strongly dilated acroscopically, subtruncate basiscopically; lamina (2-)3-7 cm broad; trichomes on rachis adaxially 0.3-0.5 mm long and per- sistent 42. G. firma j3. Pinnae oblong to narrow- or oblong-deltate, not de- flexed, subequilaterally truncate or slightly dilated at base; lamina 0.4-2.5 cm broad; rachis trichomes 0.6-1 mm long (sometimes deciduous) k3 k3. Pinnae ascending, acute; lamina not or abruptly re- duced at base, proximal 0-3 pairs of pinnae some- what shortened 43. G. pilosissima k3. Pinnae patent, obtuse or subacute; lamina strongly and gradually reduced at base, with 3-many pairs of pinnae much shortened 13 13. Pinnae mostly oblong, 2.5-6 times as long as broad; petiole amply provided with spreading, dark, unicellular trichomes, but minute, tortu- ous, septate ones rare or lacking; dried speci- mens strongly aromatic 44. G. xiphopteroides 13. Pinnae deltate or oblong-deltate, 1.5-2 times as long as broad; petiole moderately provided with minute, septate trichomes, but spreading, uni- cellular ones sparse or lacking; specimens not or weakly aromatic . . . 45. G. flabelliformis i3. Stem scales not or obscurely clathrate, yellow to lustrous orange or reddish brown, or if gray-brown then ovate or broadly lanceolate and minutely clathrate, with 1 8-30 cells across at scale base and most lumina very small and nearly isodiametric m3 m3. Stem scales flaccid and dull yellow in mass, with scat- tered, minute, marginal glands; sori solitary at segment base; petiole 0.5-2.5 cm long . . . 62. G. blepharidea m3. Stem scales stout and bright orange to red-brown or thin and gray-brown, eglandular; sori 2-12 pairs on a pinna; petiole 3-10 cm long n3 n3. Stem scales orange to red-brown, stout, rigid and lustrous, nonclathrate, terminating in a pale, apical (often deciduous) seta; rachis trichomes pale and sericeous 63. G. taxifolia n3. Stem scales light or gray-brown, thin, minutely clathrate, lacking an apical seta; rachis trichomes rigid and castaneous or lacking o3 o3. Pinnae narrowly deltate or oblong-deltate, 4- 8 times as long as broad, margins essentially plane, apex acute or subacute 46. G. pseudonutans o3. Pinnae oblong or broadly oblong-deltate, 1.5- 2 times as long as broad, margins mostly rev- olute, apex obtuse (occasionally subacute) . . . 47. G. andicola 80 FIELDIANA: BOTANY 1 . Grammitis bryophila (Maxon) Seymour, Phy- tologia31: 172. 1975. Polypodium bryophilum Maxon, Amer. Fern J. 16: 7. 1926. TYPE: Costa Rica, Pcia. San Jose, vicinity of La Palma, Maxon & Harvey 7980 (holotype, us!; photo, F). Plants commonly epiphytic, rarely (and in Peru) terrestrial. Stem erect to ascending, provided with bright orange, lustrous, nonclathrate scales, these 1 .5-3 mm long, to 0.5 mm broad, 8-12 cells across at the base. Leaves simple, entire, 4-25 cm long, 3-5 mm broad. Petiole dark brown to castaneous, 0-3 mm long, 0.2-0.8 mm in diameter. Lamina linear to narrow-elliptic or -oblanceolate, erect, subcoriaceous, attenuate at base, the apex acute to attenuate, provided with a lustrous, black or castaneous, sclerenchymatous border about 0.2 mm broad, essentially glabrous, but with scat- tered, minute, subclavate, caducous trichomes, es- pecially on costa and margin. Veins indistinct, di- verging from the costa at 30-^tO(-50)° angles, sterile ones simple, fertile ones forked, hydathodes lack- ing. Sori elongate, often slightly immersed, usually confluent at maturity. In dwarf forest, in sphagnum, 2700 m, San Mar- tin and Pasco. Costa Rica; Panama; Venezuela; Colombia; Peru; Bolivia. The two specimens cited below from San Martin are mounted on the same sheet at Kew, each with the small, blue, handwritten label characteristic of Spruce, and both are numbered 4643. It is as- sumed the number is correct for the Spruce spec- imen, since other specimens of his, collected near this date, bear numbers in this sequence. Appar- ently mounters inadvertently copied this number onto the Nelson label; this specimen may have come from Ecuador (see Comments). San Martin: Mt. Picote near Moyobamba, Nelson in Dec. 1856 (K). Mt. Campana, near Tarapoto, Spruce 4643 in Aug. 1 856 (K). Pasco: Border between Provinces Pasco and Oxapampa, San "Cotardo" (Gutardo), van der Werffet al. 8584 (MO, uc). 2. Grammitis limbata Fee (Mem. foug. 5) Gen. fil.: 233. 1852. TYPE: Guadeloupe, Perrotet, in 1824 (holotype, P!). Figures 5b-c. Grammitis nigrolimbata Hooker, Sp. fil. 4: 164. 1862. in synon. Polypodium nigrolimbatum Jenman, Bull. Bot. Dept.. 'n.s. 4: 69. 1897. Polypodium limbatum (Fee) Maxon, Bull. Torrey Bot. Club 42: 222. 1915. Grammitis nigrolimbata (Jenman) Lell., Proc. Biol. Soc. Wash. 89:715. 1977. Plants terrestrial. Stem erect to ascending, pro- vided with bright orange or golden orange, lus- trous, rigid, nonclathrate scales, these 2-3 mm long, to 0.5 mm broad, 8-12 cells across at the base. Leaves simple, entire to weakly sinuate-undulate, 5-16 cm long, 3-8 mm broad. Petiole light brown, 1-5 mm long, 0.2-0.4 mm in diameter. Lamina linear to narrow-elliptic or -oblanceolate, erect, firm-herbaceous to chartaceous, attenuate at base, the apex acute or subacute, rarely apiculate, pro- vided with a lustrous black or castaneous, scler- enchymatous border usually less than 0.1 mm broad, essentially glabrous but with scattered, minute, caducous trichomes, especially on costa and margin, these simple and subclavate, rarely branched. Veins distinct or indistinct, diverging from the costa at 30—40° angles, sterile ones sim- ple, fertile ones commonly with a short spur bear- ing the sorus, hydathodes evident adaxially. Sori superficial, rounded to, usually, elongate, some- times confluent. In cloud forests, growing on trees, 750-910 m, Pasco and Cuzco. West Indies; Venezuela; Peru; Bolivia; Brazil. Pasco: Prov. Oxapampa, west divide of Cordillera de San Matias between Iscozacin and summit, D. Smith 2038 (F, MO). Cuzco: Prov. Paucartambo, Rio Tono Road, Foster et al. 10619(f). 3. Grammitis paramicola Bishop, Amer. Fern J. 67: 105. 1977. TYPE: Colombia, Depto. Hui- la, 25 km southeast of Gigante, Little 8663 (holotype, us!; photo, F). Plants terrestrial. Stem erect to ascending, pro- vided with deep orange, lustrous, rigid, nonclath- rate scales, these 1-2 mm long, ca. 0.2-0.3 mm broad, 2-6 cells across at the base. Leaves simple, entire to slightly sinuate, 2-7 cm long, 1.5-3 mm broad. Petiole light brown, 1-5 mm long, 0.2-0.4 mm in diameter. Lamina linear, or slightly broad- ened distally, rigid, subcoriaceous, attenuate at base, the apex obtuse to subacute, rarely cuspidate, provided with a black or dark brown, lustrous, sclerenchymatous border 0.1-0.2 mm broad, es- sentially glabrous, but with scattered, caducous, minute, subclavate trichomes especially on costa and margin. Veins obscure, diverging from the TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 81 costa at a very acute angle, sterile ones simple, fertile ones furcate, hydathodes lacking. Sori roundish to elongate, confluent at maturity and sometimes slightly immersed in the tissue. In open paramos or elfin forests, in sphagnum or on mossy hummocks, 2750-3500 m, San Mar- tin, Huanuco, Cuzco. Costa Rica; Colombia; Peru. This is one of the three species from Peru in the genus with dark, sclerenchymatous borders (Grammitis s.s., fide Smith and Moran). In addi- tion to the key characters, it differs also from G. bryophila and G. limbata in its smaller, more rig- idly erect leaves and by the more acute vein angle. In G. paramicola veins diverge from the costa at about 10-15°, whereas the angle is commonly SO- SO0 in the other two species. San Martin: Prov. Mariscal Caceres, Chochos, NW corner of Rio Abiseo National Park, Young & Leon 4760 (USM). Huanuco: Playapampa, Macbride 4520 (F, us). Cuzco: Prov. La Convention, Cordillera Vilcabamba, Dudley 11061 (GH). Huadquina, Biies 1271 (us). 4. Grammitis pumila (C. Chr.) Stolze, comb. nov. Cochlidium pumilum C. Chr., Dansk. Bot. Ark. 6(3): 19, t. 3,f.3. 1929. TYPE: Ecuador, Chuquiribam- ba, Massee, in 1868 (holotype, K). Plants epiphytic or terrestrial. Stem erect or as- cending, provided with bright orange, lanceolate, attenuate, nonclathrate scales, these 2-3 mm long. Leaves simple, entire to distally repand, 4-15 cm long, ca. 1 mm broad. Petiole brown to blackish, 2-5 mm long, 0.2-0.5 mm in diameter. Lamina linear, subcoriaceous, attenuate at base, fertile only in the distal portion, this often conduplicate, gla- brous. Veins obscure, reduced to short spurs, hydathodes indistinct to obscure. Sori elongate, arranged in interrupted or continuous lines (coe- nosori). In elfin forests or somewhat open places, on or at bases of trees, 2100-2600 m, Cajamarca and Huanuco. Venezuela; Colombia to Bolivia; Brazil. Cajamarca: Prov. Cutervo, La Pucarilla, Socota to San Andres, Lopez et al. 6717 (F, GH). Huanuco: SW slope of Rio Llullapichis watershed, ascent of Cerros del Sira, Dudley 134 54 A (GH), 13490 (GH). 5. Grammitis myosuroides (Sw.) Sw., J. Bot. (Schrader) 1800(2): 18. 1801. Figure 5a. Polypodium myosuroides Sw., Prodr. 131.1 788. TYPE: Jamaica, Swartz (holotype, s; photo, us). Xiphopteris myosuroides (Sw.) Kaulf., Enum. fil. 85. 1824. Xiphopteris jamesonii Hooker, Sec. cent, ferns, /. 14. 1860. TYPE: Ecuador, Andes of Quito, Jameson (holotype, K). Polypodium serrulatum var. strictissimum Hooker, Sp. fil. 4: 175. 1862, based on Xiphopteris jamesonii Hooker. Polypodium jamesonii (Hooker) Jenman, Bull. Bot. Dept. 4: 112. 1897, nom. illeg., not (Fee) Salo- mon, 1883. Polypodium strictissimum (Hooker) Hieron., Bot. Jahrb. Syst. 34: 501. 1904. Grammitis jamesonii (Hooker) Morton, Contr. U.S. Natl. Herb. 38: 96. 1967. Lellingeria myosuroides (Sw.) Smith & Moran, Amer. Fern J. 81: 85. 1991. Plants epiphytic. Stem small, erect, provided with grayish brown, clathrate, narrow-deltate, at- tenuate scales, these ca. 1 mm long. Leaves partly dimorphic, 3-10 cm long, 3-4 mm broad. Petiole dark brown to castaneous, 1-4 mm long, 0.2-0.4 mm in diameter. Lamina linear, chartaceous, at- tenuate at base, fertile only in the subentire distal portion, sterile portion deeply pinnatisect with 1 5- 20 pairs of segments, these deltate and acute to oblong and obtuse, provided with scattered, mi- nute, persistent, castaneous, setiform trichomes to 0.5 mm long. Veins simple and solitary in each sterile segment, black and extending at broad an- gles beyond the sporangia in the fertile portion, hydathodes indistinct or lacking. Sporangia borne at the base of veins, initially in discrete sori, but eventually confluent as 2 subcostal lines, merging into a broad, single, costal line at maturity. Known thus far in Peru by a single collection from Cajamarca, edge of the road, 2700 m. Mexico (Chiapas); Costa Rica; Jamaica; Cuba; Puerto Rico; Venezuela; Colombia to Peru. Besides the key characters this differs also from G. serrulata in the blackish veins of the fertile portion of the lamina, which spread at a very broad angle from the costa and contrast strongly with the laminar tissue. Fertile veins of G. serrulata spread at an acute angle and contrast little with the lam- inar tissue. The sole collection of G. myosuroides in Peru was found growing together with speci- mens of the latter and G. pumila. Cajamarca: Prov. Cutervo, La Pucarilla, between San Andres and Socota, Lopez et al. 6716 (GH, HUT). 6. Grammitis serrulata (Sw.) Sw., J. Bot. (Schrad- er) 1800(2): 18. 1801. 82 FIELDIANA: BOTANY AcrostichumserrulatumSw.,Prodr. 128. 1788. TYPE: Jamaica, Swart z (holotype, s; frag., us; isotypes, B, Herb. Willd. 19589-1 &-3; photos, GH of B). Xiphopteris serrulata (Sw.) Kaulf., Wes. Farrnkr. 87. 1827. Polypodium serrulatum (Sw.) Mett., Fil. hort. hot. Lips. 30. 1856, not. Sw. 1802. Polypodium duale Maxon, Contr. U.S. Natl. Herb. 1 6: 61. 1912, nom. nov. for Acrostichum serrulatum Sw. and with the same type. Cochlidium serrulatum (Sw.) Bishop, Amer. Fern J. 68: 80. 1978. Plants epiphytic, occasionally epipetric. Stem ascending, provided with brown, narrow-deltate, nonclathrate scales, these 2-3 mm long. Leaves partly dimorphic, 3-8 cm long, to 2.5 mm broad. Petiole very short or obsolete. Lamina linear, firm- herbaceous, attenuate at base, fertile only in the subentire, sometimes conduplicate, distal portion, sterile leaf or portion serrate-dentate or with short, acute lobes, glabrous, or with some pale and ca- ducous trichomes on juvenile plants. Veins indis- tinct to obscure, simple and solitary on each sterile segment, in fertile portion ascending at an acute angle and not contrasting strongly in color with the laminar tissue, hydathodes vestigial or lacking. Sporangia borne on costa and veins, becoming confluent in a single costal line (coenosorus). Polypodium trifurcatumL., Sp.pl. 1084. 1753. TYPE: Plumier t. 138, Traite foug. Amer., illustrating a Plumier specimen from Martinique. Glyphotaenium trifurcatum (L.) Lellinger, Amer. Fem J. 75: 31. 1985. Enterosora trifurcata (L.) Bishop, Syst. Bot. 17: 353. 1992. Plants epiphytic. Stem short-creeping to as- cending, provided at the apex with dense clusters of closely imbricate, yellow or yellow-brown scales, these narrow-deltate, not or scarcely clathrate, 2- 4 mm long, remotely ciliolate. Leaves monomor- phic, 10-27 cm long, 2-3.5 cm broad. Petiole 3- 8 cm long, commonly '/j-'/z the length of the lam- ina, lustrous, blackish or atropurpureous, amply to densely provided with spreading, castaneous, unicellular trichomes, these 1-1.5 mm long. Lam- ina subcarnose, lanceolate or elliptic-lanceolate, obtuse or subacute at apex, cuneate at base, cut '/3-'/2 to the costa, the lobes obtuse (rarely sub- acute), the margins sparsely provided with tri- chomes like those of the petiole, or glabrate. Veins obscure, free or rarely anastomosing, pinnately branched in each lobe, hydathodes lacking. Sori round to elongate, borne at the tips of acroscopic vein branches, in 3-5 irregular series on each side of the costa, somewhat immersed in the tissue. In wet forests or clearings, on trees or fallen logs, 325-1 800(-2700) m, Cajamarca, Loreto, and Huanuco south to Cuzco. Southern Mexico to Panama; West Indies; Co- lombia to the Guianas, south to Bolivia and Brazil; wet tropical Africa. This is one of the most widespread species of Grammitis in the Neotropics and one of the easiest to distinguish. For further comparisons, see dis- cussion under G. myosuroides. Cajamarca: Prov. Cutervo, La Pucarilla, between San Andres and Socota, Ldpez et al. 6715 (GH). San Martin: Prov. Mariscal Caceres, Dist. Tocache Nuevo, Schunke y. 7807 (MO). Loreto: Sierra del Pongo, Mexia 6287 (GH, MO, us). Huanuco: Prov. Huanuco, Dist. Churubamba, Pampa Hermosa, Mexia 8258a (F, GH). Pasco: Prov. Oxapampa, west side of Cordillera de San Matias, be- tween Iscozacin and summit, D. Smith 2041 (F, MO). Junin: East of Quimiri Bridge near La Merced, Killip & Smith 23886 (F, us). Cuzco: Valle de Cosnipata, between Yaramayo and Santa Isabel, Scolnik 863 (uc, us). Uncommon, in dense forests, usually arched or pendent from tree trunks, 700-1900 m, Huanuco, Pasco, Madre de Dios. West Indies; Costa Rica and Panama; the Guianas to Colombia; Ecuador to Bolivia. Specimens of G. trifurcata from the West Indies, especially Jamaica, commonly bear two different kinds of indument: intermixed with the long, spreading, unicellular, trichomes can be found some scattered, minute, tortuous ones with 3—4 cells, the apical cell often glandular. No such tri- chomes have been observed in the few specimens found thus far in Peru. The character merits fur- ther examination in future monographic study. Huanuco: SW slope of Rio Llullapichis watershed, as- cent of Cerros del Sira, Dudley 13084b (GH). Pasco (as Junin): Pichis Trail, Dos de Mayo, Killip & Smith 25803 (F, us). Pichis Trail, Porvenir, Killip & Smith 25894 (NY, us). Madre de Dios: Prov. Manu, Cerro de Pantiacolla, Rio Palotoa, Foster et al. 10847 (F, USM). 7. Grammitis trifurcata (L.) Copel., Gen. fil. 21 1. 1947. 8. Grammitis trichosora (Hooker) Ching, Bull. Fan. Mem. Instil. Biol., Bot. 10: 241. 1941. TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 83 Polypodium trichosorum Hooker, Sec. cent, ferns, /. 12. 1860. TYPE: Ecuador, Archidona, Andes of Quito, Jameson 349 (holotype, K.!; photos, F, us). Enterosora trichosora (Hooker) Bishop, Syst. Bot. 1 7: 357. 1992. Plants epiphytic. Stem short-creeping or repent, provided at the apex with dense clusters of closely imbricate, pale yellow scales, these narrow-deltate, nonclathrate, 1-2 mm long, entire to remotely denticulate or ciliolate. Leaves monomorphic, 8- 18 cm long, 1-1.5 cm broad. Petiole 1-5 cm long, 0.4-0.8 mm in diameter, yellowish to reddish brown, densely covered with spreading, casta- neous, unicellular trichomes 2 mm long. Lamina firm-herbaceous, elliptic to oblanceolate, apex subacute, base cuneate, margins subentire to re- mand or broadly and shallowly lobed, abundantly provided with persistent trichomes like those of the petiole. Veins obscure, pinnately branched, free to irregularly anastomosing especially away from the costa, hydathodes lacking. Sori round or slight- ly elongate, in 1-2 irregular series on each side of the costa, not or scarcely immersed in the tissue. Thus far known in Peru by a single collection from San Martin, epiphyte in montane rain forest, 2100 m. Ecuador and Peru. Very similar is Polypodium ecostatum Sodiro (Central America; Antilles; northern South Amer- ica). However, the latter is a much longer fern of lower elevations (under 1 000 m), with sori rather deeply immersed in the subcarnose tissue. Also, the veins frequently anastomose along the costa, whereas areoles of G. trichosora are quite irregular, occurring infrequently along the costa, and are much more common about midway between costa and margin. San Martin: Prov. Rioja, Venceremos, Pedro Ruiz- Moyobamba Road, D. Smith & Vdsquez 4727 (MO, USM). 9. Gram mil is parietina (Klotzsch) Fee (Mem. foug. 5) Gen. fil. 233. 1852. Polypodium parietinum Klotzsch, Linnaea 20: 373. 1847. TYPE: Venezuela (as "Columbia"), Colo- nia Tovar, Moritz 253 (holotype, B!; isotype, BM; photo, F of B). Polypodium sprucei Hooker, Sec. cent, ferns, t. 10. 1 860. TYPE: Peru, San Martin, Tarapoto, Spruce 4746 (holotpye, K!; isotype, us!). Grammitis sprucei (Hooker) John Sm., Hist. fil. 181. 1875. Enterosora parietina (Klotzsch) Bishop, Syst. Bot. 17: 357. 1992. Plants epiphytic. Stem small, ascending, pro- vided at apex with a few inconspicuous, deep or- ange scales, these entire or ciliolate, obscurely clathrate, 0.5-1 mm long. Leaves monomorphic, 4-9 cm long, 0.4-0.8 cm broad. Petiole to 0.7 cm long or obsolete, 0.4-0.6 mm in diameter, brown to blackish, amply provided with spreading, cas- taneous, unicellular trichomes about 1 mm long. Lamina subcarnose to chartaceous, oblong-ellip- tic, apex obtuse or subacute, base attenuate, mar- gins subentire to sinuate or crenate, amply pro- vided with trichomes like those of the petiole. Veins obscure, free, commonly 1 -forked, hydathodes lacking. Sori roundish, in a single series on each side of the costa, superficial or slightly immersed in the tissue. In cloud forests, on tree trunks, 1900-2400 m, Amazonas, Pasco, San Martin and Cuzco. Southern Mexico; Guatemala; Costa Rica and Panama; Jamaica; Hispaniola; Venezuela; Colom- bia to Peru. The species thus far is represented in Peru by only three widely scattered collections. Because of its small size and inconspicuous leaves it has likely been overlooked, and a more diligent search should reveal it to be in intervening localities. Amazonas: Prov. Bagua, 12-20 km E of La Peca, Bar- hour 2723 (F). Pasco: Prov. Oxapampa, San Alberto, Cordillera de Yanachaga, van der Werffel al. 8496 (MO). Cuzco: Prov. La Convencion, Cordillera Vilcabamba, Dudley 11 302 (GH). 10. Grammitis phalacron Stolze, sp. nov. Caulis breve gracilisque, paleis 1-1.5 mm longis, castaneis, setosis, angusti-deltatis, non clathratis. Folia 7-12 cm longa, 0.6-0.8 cm lata. Petiolus 1-2 cm longus, trichomatibus circa 2mm longis, aurantiacis, unicellularibus. Lamina anguste el- liptica, subintegra vel repanda, glabrata vel tri- chomatibus 0.2-0.4 cm longis, pallescentibus, tortuosis, pluricellularibus. Hydathodi nulli. Sori rotundi, discreti, praphysibus glandulis. Plants probably epiphytic. Stem short, slender, provided with castaneous, narrow-deltate, setose, nonclathrate scales, these 1-1.5 mm long. Leaves monomorphic, 7-12 cm long, 0.6-0.8 cm broad. Petiole 1-2 cm long, 0.2-0.4 mm in diameter, dark brown to blackish, amply provided with spread- ing, orange, unicellular trichomes, these to 2 mm 84 FIELDIANA: BOTANY long. Lamina firm-herbaceous to chartaceous, nar- row-elliptic, apex obtuse to subacute, cuneate at base, margins subentire or repand to very broadly and shallowly lobed, abaxially provided with scat- tered, pale, tortuous, pluricellular trichomes, these 0.2-0.4 mm long, caducous. Veins indistinct, 1- 2-forked to pinnate, hydathodes lacking. Sori round, borne on basal vein branches and forming a single series on each side of the costa, paraphyses glandular. TYPE— Peru, Cuzco, Valley of Rio Urubamba, Bues A29 (holotype, us!). Known only from the type. In general aspect, this appears to be interme- diate between G. parietina and Polypodium (Grammitis) ecostatum Sodiro (Ecuador). How- ever, the completely different indument, the glan- dular paraphyses and the thinner textured lamina (vs. laminae with spongy parenchyma) argue against such affinity. The specimen was annotated by Bishop and Smith as Ceradenia phalacron, with intent to publish at a later date. Their kind per- mission for publication as Grammitis in this flora is greatly appreciated. 1 1. Grammitis longipinnata (Copel.) Loll.. Amer. Fern J. 74: 58. 1984. Ctenopteris longipinnata Copel., Philipp. J. Sci. 84: 459. 1956. TYPE: Peru, Cuzco, Cabeceradel Ko- ribeni, Bues 1952 (holotype, us!). Ceradenia longipinnata (Copel.) Bishop, Amer. Fern J. 78: 5. 1988. Plants probably epiphytic. Stem unknown. Leaves to 30 cm long and 10 cm broad. Petiole 8-10 cm long, 1 mm in diameter, blackish, sub- lustrous, glabrescent. Lamina chartaceous to sub- coriaceous, subdeltate, not or scarcely reduced at base, deeply pinnatisect to fully 1 -pinnate toward the base, rachis amply provided with spreading, castaneous trichomes to 1 mm long; pinnae falcate and somewhat ascending, linear-deltate, acute to attenuate, margins with scattered, castaneous tri- chomes and deeply serrate, but distal ones merely crenulate or entire. Veins obscure, free, 1 -forked, hydathodes lacking. Sori round to slightly elon- gate, in a single supramedial to submarginal series on each side of the costa. Endemic. Thus far known from the type and two other collections from Cuzco, 2750-3250 m. The Dudley specimen was pendulous from trees in a dark, wet, cloud forest. Nothing is known of the stem characteristics of this species, since leaves of the only known spec- imens are broken off at the petiole base. No habit was mentioned on labels of the type and the other Biles collection. Cuzco: Valle de Lares, Bues 1793 (us). Prov. La Con- vention, 25 km from Hacienda Luisiana and the Apu- rimac River, Dudley 1 1 106 (us). 12. Grammitis meridensis (Klotzsch) Seymour, Phytologia 31: 179. 1975. Polypodium meridense Klotzsch, Linnaea 20: 380. 1847. TYPE: Venezuela, Edo. Merida, Merida, Moritz 335 (holotype, B!; photo, F). Ctenopteris meridensis (Klotzsch) Copel., Philipp. J. Sci. 84: 464. 1956. Ceradenia meridensis (Klotzsch) Bishop, Amer. Fern J. 78: 5. 1988. Plants terrestrial, at least in Peru. Stem short- creeping to ascending, abundantly provided with linear, rigid, lustrous, deep orange to castaneous scales, these (2-)3-6 mm long, nonclathrate, am- ply setulose, the setae whitish. Leaves monomor- phic, articulate and approximate on the stem, 20- 40(-50) cm long, 3-7 cm broad. Petiole 8-30 cm long (usually as long as the lamina), 0.4-1 mm in diameter, lustrous, castaneous to atropurpureous or blackish, sparsely to amply pilose, or glabres- cent at maturity, the trichomes castaneous. spreading, unicellular, 0.4-1 mm long. Lamina firm-herbaceous to chartaceous. oblong or oblong- deltate, truncate at base, deeply pinnatisect, or rarely 1 -pinnate at base, rachis moderately pro- vided on both sides with spreading, castaneous, trichomes to 1 mm long. Segments patent or slightly ascending, linear-deltate, acute or subacute, tri- chomes like those of the rachis. sparse along the margins and sometimes on the abaxial surface, especially among the sori. Veins completely ob- scure, commonly once-forked, hydathodes lack- ing. Sori superficial, round, supramedial to sub- marginal, immature sporangia intermixed with pale, waxy paraphyses. In dense, wet forests or wooded rav ines. on clay banks or in humus, 1500-3600 m. Huanuco to Cuzco. Costa Rica; Panama; Venezuela and Colombia south to Brazil and Bolivia. This species is especially distinguished by the TRYON & STOLZE. PTERIDOPHYTA OF PERU. V. 85 long (to 1 mm), spreading, dark, unicellular tri- chomes on both sides of the rachis and by the truncate lamina base with basal pinnae longer or nearly as long as medial ones. Very similar, and probably conspecific, are Polypodium spixianum Mett. of Brazil and P. abitaguae Hooker of Ec- uador. Both apparently differ from G. meridensis in the lighter orange color and greater length (to 9 mm) of stem scales, and sori of P. abitaguae are medial rather than supramedial or submarginal. Some specimens have been determined as P. spixi- anum on the basis of slightly thinner lamina tex- ture and nearly obtuse segment apices, but both characters are too variable to be significant. This condition seems to pertain to juvenile and/or de- pauperate plants, while thicker texture and longer, acute segments are common to fully mature leaves. Huanuco: Cani, pueblo 7 mi NE of Mito, Macbride 3491 (F, us). Vilcabamba, Macbride 5145 (F in part, us). Pasco: Pichis Trail, Porvenir (as Junin), Killip & Smith 25939 (F, NY, us). Prov. Oxapampa, Dist. Chontabamba, near the summit of San Gutardo, Leon et al. 524 (USM, juvenile specimens), 526 (F, USM). Junin: Chanchamayo Valley, C. Schunke 853 (F), 1380 (F). Cuzco: Prov. La Convencion, Valle San Miguel, Biles 2133 (us). Prov. Paucartambo, near El Mirador, Manu National Park, Leon 2265 (F, USM). Prov. Paucartambo, Cordillera de Tres Cruces, Vargas 12191 (GH). 13. Grammitis bishopii Stolze, sp. nov. Plantae terrestres. Caulis repens, 1.5-3 mm crassus, paleisabundantibus, linearibus, rigentibus, niten- tibus, castaneis, 2-4 mm longis, 0.2-0.4 mm latis, setulosis, setulis concoloribus. Folia 18-34 cm longa, 4-10 cm lata, subdistantia, ad phyllopodia bulbosa affixa. Petiolus 10-24 cm longus, nitidus, castaneus vel atropurpureus, glaber. Lamina 1-pinnata, deltata, basi non reducta. Rhachis et costa trichomatibus minutis, castaneis, unicel- lularibus, abundantibus praeditae. Venae ob- scurae, 1-2-furcatae, raro anastomosantes. Hy- dathodi nulli. Plants terrestrial. Stem long-creeping, 1.5-3 mm in diameter, densely clothed with linear, rigid, lus- trous, castaneous scales, these 2-4 mm long, 0.2- 0.4 mm broad, nonclathrate, sparingly to amply setulose, the setae concolorous with the scale body. Leaves monomorphic, borne on short (1-2 mm), often bulbous phyllopodia, usually 1-2 cm apart, 18-34 cm long, 4-10 cm broad. Petiole 10-24 cm long, 1-2 mm in diameter, lustrous, castaneous to atropurpureous, essentially glabrous. Lamina sub- coriaceous, narrowly or broadly deltate, not or scarcely reduced at base, 1 -pinnate, the segments adnate, slightly ascending to slightly reflexed, lin- ear, acute, their margins often with minute, scat- tered, glandular trichomes, the rachis adaxially with dense, minute, castaneous, rigid, unicellular tri- chomes, these 0.1-0.3 mm long, costae with sim- ilar, but shorter and fewer, trichomes. Veins completely obscured (even when held to light), 1- 2-forked, often connivent distally, on some spec- imens frequently anastomosing, hydathodes lack- ing. Sori superficial, commonly round, essentially medial between (and at maturity often crowding both) costa and margin, paraphyses not seen. TYPE— Peru, Pasco, Prov. Oxapampa, border Prov. Oxapampa and Pasco, van der Werff et al. 8569 (holotype, uc!; isotype, MO!). In elfin or cloud forests, in clay or sphagnum, 2700-3000 m, Amazonas, Pasco and Cuzco. Peru; Bolivia (Tate 349 [NY]). This may be confused with G. werffii. In addition to the characters used in the key, this differs in the rarely (vs. frequently in the latter) anastomosing veins and in the short, rather swollen, phyllopo- dium that joins the petiole base to the stem. In G. werffii the petiole base is articulate and the line of abscission very evident. The species is named in honor of the late L. Earl Bishop, who recognized this as a new species and tentatively labeled several of the specimens as "Ceradenia vaga," but until now it has remained undescribed. Amazonas: Prov. Bagua, Cordillera Colan, E of La Peca, Barbour 3264 (MO, USM). Pasco: Prov. Oxapampa, Dist. Chontabamba, near the summit of San Gutardo, Leon et al. 526 (USM). Prov. Oxapampa, Cerro Pajonal, 29 km from Oxapampa, D. Smith & Foster 2504 (uc). Border, Prov. Oxapampa and Pasco, below San Gutardo (as "Cotardo"), van der Werff et al. 8550 (MO). Cuzco: Prov. La Convencion, Cordillera Vilcabamba, Dudley 10892 (F, GH), 11145 (GH, US). 14. Grammitis discolor (Hooker) Morton, Amer. Fern J. 60: 66. 1970. Polypodium discolor Hooker, Icon. pi. 4: /. 386. 1841. TYPE: "British Guiana" (Guyana), Schomburgk 1031 (holotype, K; photo, F). Ctenopteris discolor (Hooker) John Sm., Hist. fil. 185. 1875. Ceradenia discolor (Hooker) Bishop, Amer. Fern J. 78:4. 1988. Plants probably terrestrial. Stem stout, erect, provided with linear, brown scales, these 3-6 mm long, nonclathrate, entire to sparsely and remotely setulose. Leaves not articulate, 9-20 cm long, 1- 2 cm broad. Petiole essentially obsolete. Lamina 86 FIELDIANA: BOTANY subcarnose, linear to narrow-elliptic, strongly and gradually reduced at base, deeply pinnatisect (about 3/4 to the rachis) the segments 1-2 times as long as broad, rachis rarely evident (usually immersed in the tissue), rachis and tissue lacking trichomes, moderately to densely farinose, the farina white, mostly disposed in stellate patterns. Veins ob- scure, free, pinnately branched in the segments, hydathodes lacking. Sori round, submarginal, with farina deposited among the sporangia. In dense forests, apparently terrestrial (habit not cited on any of the labels seen, but clay particles evident among the roots), 1 300-2000 m, thus far found only in Junin. Guyana; Venezuela; Colombia; Peru; Bolivia. This is very similar to G. curvata ssp. pearcei, differing only in the farina pattern and the shape of segments and may represent merely another infraspecific variant of the latter. Junin: Schunke Hacienda above San Ramon, C. SchunkeA152(GH, us). Chanchamayo Valley, C. Schunke 470 (F, us), 513 (F, us). 15. Grammitis curvata (Sw.) Ching, ssp. pearcei (Baker) Stolze, stat. & comb. nov. Polypodiutn pearcei Baker, in Hooker and Baker, Syn. fil., ed. 2: 508. 1874. LECTOTYPE (designated here): Ecuador, Cuesta de Puente Grande, Pearce in June 1865 (K!, photo, F, isolectotype, 2 laminae on the right, K!) Ctenopteris amylacea Copel., Philipp. J. Sci. 84: 469. 1956. TYPE: Bolivia, Cargadira, R. S. Williams 1134 (holotype, us!; isotypes, GH!, uc!). Grammitis amylacea (Copel.) Morton, Contr. U.S. Natl. Herb. 38: 233. 1973. Ceradenia pearcei (Baker) Bishop, Amer. Fern J. 78: 4. 1988. Plants epiphytic or epipetric. Stem stout, erect, provided with linear, orange-brown to ferruginous scales, these 3-6 mm long, nonclathrate, sparsely to amply setose, the setae pale orange or whitish. Leaves not articulate, 12-30(-35) cm long, (2.5-) 3-8 cm broad. Petiole essentially obsolete. Lamina subcarnose, rachis and costae rarely evident (usu- ally immersed in the tissue), narrowly to broadly elliptic, strongly and gradually reduced at base, deeply pinnatisect, the segments 3-7 times as long as broad, rachis and tissue lacking trichomes, moderately farinose, the farina particles white (of- ten brown on dried specimens), solitary or dis- posed in small, amorphous clusters. Veins ob- scure, free, once-forked from the costa, hydathodes lacking. Sori round, borne at the tip of the acros- copic vein branch, submarginal, at maturity ex- tending beyond the segment margin, with farina deposited among the sporangia. In scrub and cloud forests, on tree trunks or in crevices of rocky cliffs, 2000-2750 m, Amazonas, Huanuco, Cuzco. Peru; Bolivia. This differs from ssp. curvata (West Indies) in the longer and more abundant setae on the stem scales. Furthermore, ssp. curvata has minute, ap- pressed (sometimes glandular?) trichomes scat- tered on the lamina, in addition to the farina de- posits. These trichomes are 0.1-0.2 mm long, cylindrical, castaneous, and are often quite abun- dant, though very inconspicuous. Such processes have not been seen in specimens of ssp. pearcei. Also very similar (and likely another subspecies) is Grammitis albidula (Baker) Morton, from southeastern Brazil, but the latter has smaller leaves and supramedial (vs. submarginal) sori. There are two sheets of Pearce specimens in the type folder at Kew. One has the stem and several leaves and bears a printed label marked "Type Specimen." The other sheet has two laminae with- out a stem, and a third lamina from another Pearce collection with the number 279 and "holotype" handwritten in blue ink. The two stemless laminae were annotated by Bishop (1988) as lectotype, but that typification has not been published. The other sheet is a much preferable lectotype, since the plant is complete and not mixed, and is so designated here. Amazonas: Prov. Bagua, E of La Peca, Barbour 2767 (F, uc). Prov. Chachapoyas, west of Molinopampa, Wur- dack 1499 (us), 1543 (us). Huanuco: Cushi, trail to Tam- bo de Vaca, Bryan 686 (F, us). Cuzco: Prov. Paucartam- bo, Manu Park, between Pillahuata and Pilcopata, Skog &Skog5198(vs). 16. Grammitis terrestris (Bishop) Stolze, comb, nov. Ceradenia terrestris Bishop, Amer. Fern J. 79: 23. 1989. TYPE: Peru, Amazonas, Prov. Chacha- poyas, south of Molinopampa-Diosan Pass, Wurdack 1643 (holotype, us!; isotype, GH!). Plants terrestrial. Stem stout, erect, provided with linear-deltate, deep orange to castaneous, nonclathrate scales, these 3-5 mm long, moder- ately to amply setulose. Leaves not or scarcely articulate, 8-18 cm long, 1-2 cm broad, lacking long, spreading unicellular trichomes. Petiole 1-2 cm long, 0.6-0.9 mm in diameter, minutely glan- TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 87 dulose, often white-farinose. Lamina subcarnose, linear-elliptic, gradually reduced at base and apex, copiously white-farinose and with some minute, glandular trichomes (especially on segment mar- gins), pinnatisect, the segments closely spaced, joined by a narrow wing of tissue and ascending at a 30-50° angle, sclerenchyma of the rachis abax- ially exposed, or sometimes concealed by laminar tissue. Veins obscure, free, usually once-forked, hydathodes lacking. Sori roundish, essentially me- dial, at maturity often extending from costa to margin, with farina deposited among the sporan- gia. Thus far known only from the type collection, on moist bank in scrub forest, 2700-3100 m, Amazonas. In Peru, this is most closely related to G. her- rerae. In addition to the key characters, it differs also by the narrower, more rigid lamina and by the petiole not or scarcely articulate to the stem. While the base of the petiole in G. herrerae is not swollen or noticeably discolored, in age it does fracture clearly at the base, leaving a clear and somewhat concave scar at the stem. 17. Grammitis herrerae (Copel.) Morton, Contr. U.S. Natl. Herb. 38: 233. 1973. Ctenopteris herrerae Copel., Philipp. J. Sci. 84: 467. 1956. LECTOTYPE (designated by Morton, 1973): Peru, Cuzco, Huadquina, Bues 1269 (us!). Ceradenia herrerae (Copel.) Bishop, Amer. Fern J. 78: 4. 1988. Plants possibly epiphytic. Stem small, erect, sparsely provided with narrow-deltate, orange- brown, nonclathrate, subentire scales ca. 1 mm long. Leaves cleanly but inconspicuously articulate at the stem, 10-30 cm long, 1.7-4 cm broad, lack- ing long, spreading, unicellular trichomes. Petiole ( l-)2.5-7 cm long, 0.5-0.7 mm in diameter, densely glandulose. Lamina subcarnose, elliptic, gradually reduced at apex and base, white-farinose and with some minute, glandular trichomes, pinnatisect, the segments closely spaced, joined by a narrow wing of tissue and ascending at a 60-80° angle, scleren- chyma of the rachis mostly exposed. Veins ob- scure, free, simple to once-forked, hydathodes lacking. Sori round, supramedial, at maturity rare- ly extending beyond the segment margin, with fa- rina deposited among the sporangia. In forests, 2750-3000 m, Cuzco. No habit was recorded on labels of specimens seen, but the one specimen with a stem (Sues 1268) has only ad- herent organic matter, so the species is apparently epiphytic. Endemic. The species most closely related to this is G. terrestris, under which see further comparison. Cuzco: Huadquina, Bites 1268 (paratype, us). Prov. La Convention, Abra Mirador, Bues 2073 (us). 1 8. Grammitis dendrodoxa (Bishop) Stolze, comb, nov. Ceradenia dendrodoxa Bishop, Amer. Fern J. 79: 15. 1989. TYPE: Peru, Amazonas, Prov. Chacha- poyas, Cerros de Calla Calla, Wurdack 1715 (ho- lotype, uc!; isotypes, F!, GH!, NY!, us!). Plants epiphytic (in Peru) or epipetric. Stem erect, provided with linear or narrow-deltate, lustrous, castaneous, nonclathrate scales, these 1-3 mm long, the margins remotely white-setulose. Leaves 1 -pinnate, cleanly but inconspicuously articulate at the stem, pendent, 1 2-75 cm long, 4-8 cm broad, lacking long, spreading, unicellular trichomes. Petiole 2-6 cm long, 0.3-0.6 mm in diameter, glandulose. Lamina firm-herbaceous to charta- ceous, elliptic, abruptly reduced at apex, less abruptly so at base, several proximal segments much shorter than the medial ones, glandulose and with some pale farina scattered on both sides, ra- chis castaneous to atropurpureous, glandulose, nonalate. Pinnae to 10 cm long, ca. 1.5 mm broad, subdistant to remote, ascending at a 20-35° angle, linear, attenuate to apex and base. Veins obscure, free, simple, hydathodes lacking. Sori round, at maturity extending well beyond the pinna mar- gins. In forests or wooded ravines, on tree trunks or branches, 3400-3950 m, Amazonas and Huanuco. Ecuador; Peru. Of the Grammitis species lacking hydathodes and long, dark, spreading trichomes, this is the most easily identified. It can be clearly distin- guished by strongly ascending, nearly filiform pin- nae whose mature sori extend so far beyond the margins that the pinnae at first appear to be lobed or dentate. Huanuco: Tambo de Vaca, Bryan 626 (F, us). 19. Grammitis farinosa (Hooker) Morton, Contr. U.S. Natl. Herb. 38: 99. 1967. 88 FIELDIANA: BOTANY Polypodium farinosum Hooker, Icon. pi. 10: /. 947. 1854; Cent, ferns /. 47. 1854. TYPE: Ecuador, Cordillera of Quito, Jameson (holotype, K; frag., us!). Ctenopteris farinosa (Hooker) Copel., Philipp. J. Sci. 84: 470. 1956. Ceradenia farinosa (Hooker) Bishop, Amer. Fern J. 78:4. 1988. Plants terrestrial or epiphytic. Stem erect, pro- vided with elliptic-lanceolate, orange to casta- neous, nonclathrate scales, these 1-2.5 mm long, minutely glandular-ciliolate. Leaves 1 -pinnate, clearly but inconspicuously articulate at the stem, 12—40 cm long, 1.8-5 cm broad, lacking long, spreading, unicellular trichomes. Petiole 5-10 cm long, 0.5-0.9 mm in diameter, densely glandulose. Lamina chartaceous, elliptic-lanceolate or narrow- ly oblanceolate, abruptly reduced at apex and base, at least several pairs of proximal segments less than half as long as medial ones, densely white- farinose, the farina turning brown at maturity, ra- chis dark brown, glandulose, nonalate. Pinnae 1- 3.5 cm long, 2.5-5 mm broad, approximate to subdistant on the rachis, ascending at a 60-80° angle, linear-deltate, obtuse to acute, base rounded to truncate acroscopically, decurrent basiscopi- cally. Veins obscure, free, commonly once-forked, hydathodes lacking. Sori round, supramedial, not or rarely extending beyond pinna margins. In forests or clearings, terrestrial or on tree trunks, thus far known by one collection from Peru, ca. 2800 m, Huanuco. Colombia; Ecuador; Peru. In the only collection found thus far in Peru, the lamina is not densely white-farinose as in some other specimens seen from Ecuador. In this species the farina apparently often turns brownish or falls away as the leaf reaches maturity. Huanuco: Cushi, trail to Tambo de Vaca, Bryan 569B (F, us). 20. Grammitis congests (Copel.) Lell., Amer. Fern J. 74: 58. 1984. Ctenopteris congesta Copel., Philipp. J. Sci. 84: 397. 1956. TYPE: Peru, Cuzco, Prov. La Convencion, Lorn a Grande, Biies 2172 (holotype, us!). Habit and stem unknown. Leaves 10-15 cm long, 0.6-1 cm broad. Petiole 2—4 cm long, 0.2-0.5 mm in diameter, sparsely and deciduously pilose with castaneous trichomes. Lamina coriaceous, linear, 1 -pinnate, rachis and pinnae abundantly provided with spreading, castaneous, unicellular trichomes to 2 mm long, rachis dark brown or blackish. Pin- nae crowded, slightly ascending, to 5 mm long and 2.5 mm broad, circular, obovate or broadly ob- long, the apex obtuse. Veins obscure, free, simple, hydathodes lacking or very indistinct. Sori round, 2-5 on a pinna, discrete, or confluent at maturity. Thus far known only from the type, 4000-^4300 m, Cuzco. In general aspect, this resembles G. monilifor- mis, but it is distinguished from the latter by its lack of (or very indistinct) hydathodes, and by the abundant, spreading unicellular trichomes on the rachis and pinnae. In G. moniliformis hydathodes are present (although sometimes conspicuous); furthermore, the long, unicellular trichomes are very sparsely scattered on the lamina and the ra- chis is beset with numerous, minute, pluricellular, glandular trichomes. The leaves on the type specimen lack stems, and there is no habit or habitat data on the label, so these vital characters are still open to speculation. 21. Grammitis praeclara (Bishop) Stolze, comb, nov. Ceradenia praeclara Bishop, Amer. Fern J. 79: 16. 1989. TYPE: Peru, Cuzco, Prov. La Convencion. Valle de San Miguel, Biies 2179 (holotype, us!). Plants probably epiphytic. Stem small, provided with narrow-deltate, lustrous, deep orange to cas- taneous, nonclathrate scales, these 1-2 mm long, with a few marginal glands and short setae. Leaves lax (pendent?), 1 -pinnate, not or scarcely articu- late, 1 5-30 cm long, 2.5-5 cm broad. Petiole brown to blackish, 1-3 cm long, 0.2-0.3 mm in diameter, amply provided with spreading, orange, unicel- lular trichomes ca. 1.5 mm long, and rather abun- dantly with minute glandular trichomes and sessile glands. Lamina firm-herbaceous to chartaceous, narrow-elliptic, rather abruptly narrowed at apex and base, indument as on the petiole but the long, spreading trichomes rather sparse, rachis and cos- tae blackish. Pinnae well spaced, ascending at a 50-70° angle, 1.5-3.5 cm long, 2-3.5 mm broad, narrow-deltate, subacute at apex, the base rounded nearly to costa acroscopically, short-decurrent basiscopically. Veins obscure, free, simple, or oc- casionally once-forked, hydathodes lacking. Sori round, medial or inframedial, the sporangia na- ked. TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 89 Known only from the type, 3 1 50 m, Cuzco. The type label cites no habit or habitat data, but organic material clings to the stem and the leaves are rather lax, which implies a pendent habit, so the plant was probably epiphytic. 22. Grammitis capillaris (Desv.) Proctor, Brit. FernGaz. 9:218. 1965. Polypodium capillare Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck., Gesammten Na- turk. 5: 316: 181 1. TYPE: Jamaica, Tussacl (ho- lotype, P, Herb. Desvaux; photo, us). Ctenopteris capillaris (Desv.) Copel., Philipp. J. Sci. 84: 402. 1956. Ceradenia capillaris (Desv.) Bishop, Amer. Fern J. 78: 4. 1988. Plants apparently epiphytic. Stem erect or as- cending, provided with narrow-deltate, lustrous, deep orange to castaneous, nonclathrate scales, these 1-2 mm long, with a few marginal glands and short setae. Leaves 1 -pinnate (at least proxi- mally), not or scarcely articulate, 6-15(-18) cm long, 1.5-4 cm broad. Petiole 1-2.5 cm long, 0.2- 0.4(-0.5) mm in diameter, sparsely glandular and moderately to densely provided with spreading, deep orange, unicellular trichomes ca. 1 mm long. Lamina thin- to firm-herbaceous, narrowly or broadly elliptic, rather abruptly reduced at apex and base, indument as on the petiole, but the spreading trichomes sparse to lacking, rachis and (sometimes) costae dark brown to blackish and usually lustrous, especially abaxially. Pinnae well spaced, ascending at a 25-40° angle, 0.8-2.5 cm long, 0.8-2 mm broad, linear to narrow-deltate, acute or subacute at apex, the base fully adnate and nearly perpendicular to the rachis acroscopi- cally, long-decurrent basiscopically. Veins free, in- distinct or obscure, simple, hydathodes lacking. Sori round, supramedial, most of them extending beyond the pinna margin at maturity. In forests, probably on tree trunks (although habit is not cited on any of the specimen labels from Peru), 1600-2750 m, Huanuco to Cuzco. West Indies; Venezuela; Colombia to Peru; Bo- livia; southern Brazil. Copeland did not see the type of Polypodium capillare, and his concept of the species (Copeland, 1956, p. 402) was actually that of P. pilipes, under which see further discussion. Huanuco: Pampayacu, Kanehira 185 (GH, us). Playa- pampa, Macbride 4522 (F, Os). Pasco: Oxapampa, Sou- kup 2356 (us). Junin: Schunke Hacienda, above San Ra- mon, C. Schunke A153 (us). Chanchamayo Valley, C. Schunke 469 (F). Cuzco: Prov. La Convention, Valle de Santa Ana, Herrera 3011 (us). Department unknown: "Perou," Lechler 153 (F), Mathews 921 (NY). 23. Grammitis pilipes (Hooker) Morton, Phyto- Iogia22: 77. 1971. Polypodium pilipes Hooker, Icon. pi. 3: /. 227. 1840. TYPE: Peru (Amazonas), Chachapoyas, Mathews in 1838 (holotype, 2 sheets, K; probable isotype, F!; photo, us of K). Polypodium pozuzoense Baker, Icon. pi. 17: t. 7672. 1 886. TYPE: Peru (Pasco), Pozuzo, Pearce in 1 863 (holotype, 2 sheets, K; photo, us). Ceradenia pilipes (Hooker) Bishop, Amer. Fern J. 78: 4. 1988. Plants epiphytic. Stem erect or ascending, pro- vided with narrow-deltate, lustrous, deep orange to castaneous, nonclathrate scales, these 1-2 mm long, with a few marginal glands and short setae. Leaves 1 -pinnate (at least proximally), not or scarcely articulate, (12-) 17-4 5 cm long, 3-7(-10) cm broad. Petiole 1—4 cm long, (0.4-)0.6-0.9 mm in diameter, sparsely to abundantly glandular and amply to densely provided with spreading, deep orange, unicellular trichomes ca. 2 mm long. Lam- ina subcarnose, narrowly or broadly elliptic, rather abruptly reduced at apex and base, indument as on the petiole, but the spreading trichomes sparse to lacking, sclerenchyma of the rachis usually ob- scured by the laminar tissue or, if rarely evident, then not lustrous. Pinnae well spaced, ascending at a 25-40° angle, (2-)3-4(-l 2) cm long, 1.5-2 mm broad, linear, acute to attenuate at apex, the base fully adnate and nearly perpendicular to the rachis acroscopically, long-decurrent basiscopically, proximal pinnae rarely bearing a few irregular short and narrow segments. Veins free, obscure, simple, hydathodes lacking. Sori round, most of them ex- tending beyond the pinna margin at maturity. In elfin or cloud forests or thickets, on tree trunks or branches, 1900-3700 m, Amazonas, San Mar- tin, Pasco, and Cuzco. Costa Rica; Venezuela; Colombia to Peru. In addition to the key characters, this species can also be distinguished from G. capillaris by the usually longer leaves and petiole trichomes. Fur- thermore, there is a strong tendency for the lamina of G. pilipes to have greatly elongate pinnae (to 1 2 cm!) scattered irregularly among the normal ones. Also, very rarely a few proximal pinnae may be partially pinnatifid, with a few short, irregular seg- ments. These were the characters on which Poly- 90 FIELDIANA: BOTANY podium pozuzoense was based. Such leaves seem to be monstrous forms, but those with elongate pinnae are not uncommon. Neither do these spec- imens appear to be hybrids, for no abortive spores or sporangia have been detected. Despite all these differences, G. capillaris and G. pilipes are very closely related and there are frequent intermediates, which can only be iden- tified by using the larger combination of a suite of characters. Amazonas: Prov. Bagua, Cordillera Colan east of La Peca, Barbour 3263 (MO, USM). San Martin: Moyobam- ba, Mathews (K). Pasco: Prov. Oxapampa, Parque Na- cional Yanachaga, Foster & d'Achille 12256 (F). Cuzco: Prov. La Convention, Hacienda Pintobamba above Huaillayoc, Biies 1956 (GH, us). Prov. La Convention, Cordillera Vilcabamba, Dudley 10893 (GH), 70996 (GH, MO). Prov. Paucartambo, Pillahuata, Puente de Aguila, Vargas 16746 (GH). 24. Grammitis mirabilis (Bishop) Stolze, comb, nov. Ceradenia mirabilis Bishop, Amer. Fern J. 79: 22. 1989. TYPE: Bolivia, Cochabamba, Carmen, Brooke 6134, corrected from 6134a (holotype, BM!; isotypes, NY, u, us). Plants epiphytic or terrestrial. Stem short-creep- ing, provided with narrow-deltate, clathrate scales, these 1-3 mm long, gray-brown, iridescent, the margins entire, or with glands near the tips. Leaves lax, not or scarcely articulate to the stem, approx- imate or crowded, 2-pinnate, 1 2-30 cm long, 3-8 cm broad. Petiole 4-8 cm long, 0.5-0.8 mm in diameter, sparsely puberulent with simple or branched, septate trichomes 0.1-0.3 mm long, or glabrescent. Lamina chartaceous, indeterminate, abruptly reduced at base, elliptic, with 10-15 pairs of strongly ascending pinnae. Rachis flexuous, moderately puberulent as on the petiole. Pinnae sessile, 3-8 cm long, linear, 1 -pinnate, costa pu- berulent, the pinnules adnate, 2-3 mm long, spath- ulate. Veins obscure, simple or 1 -forked in the pinnules, hydathodes lacking or very indistinct. Sori round, one to a pinnule, sporangia glabrous. Thus far known in Peru from a single collection, growing between boulders in woods, 4090 m, An- cash. Peru and Bolivia. Ancash: Prov. Huaraz, Huascaran National Park, Quebrada Llaca, D. Smith & Buddensiek 11142 (MO). 25. Grammitis mathewsii (Mett.) Morton, Amer. Fern J. 60: 66. 1970. Polypodium mathewsii Mett., Abh. Senckenberg Na- turf. Ges. 2: 74 (iiber farngat. I. Polypodium). 1856. LECTOTYPE (designated by Bishop, Amer. Fern J. 79: 108. 1989): Peru. Mathews 1811 (B!, photo, F; isolectotype, B!, BM! K, p). Zygophlebia mathewsii (Mett.) Bishop, Amer. Fern J. 79: 108. 1989. Plants epiphytic, rarely epipetric or terrestrial. Stem stout, suberect, provided with dense clusters of linear, rigid, lustrous, deep reddish brown scales, these 4-6 mm long, 0.2-0.4 mm broad, nonclath- rate, entire. Leaves monomorphic, 25-60 cm long, 5-12 cm broad. Petiole 10-28 cm long, 0.7-2 mm in diameter, dull yellowish or grayish brown, am- ply provided with spreading, castaneous, unicel- lular trichomes, these to 3 mm long and inter- mixed with minute, pluricellular, gland-tipped ones. Lamina chartaceous, lanceolate or elliptic, somewhat or scarcely reduced at base, pinnatisect nearly to the rachis, the segments strongly ascend- ing, narrow-deltate, obtuse to acute, amply to abundantly provided with trichomes like those of the petiole. Veins distinct or indistinct, 1-3-forked, the branches frequently merging distally to form areoles, hydathodes lacking. Sori superficial, roundish, forming a single series on each side of the costa, medial to inframedial, borne on the ac- roscopic vein branch, sporangia intermixed with brownish, glandular, viscid paraphyses (these rare- ly evident in mature sori). In wet forests, pendent from tree trunks or oc- casionally on stone walls, very rarely terrestrial on grassy paramos, 1800-3400 m, Amazonas, San Martin, and Cuzco. Costa Rica and Panama; Colombia to Bolivia. This species is easily recognized within the ge- nus by its large leaves, which lack hydathodes and have frequently anastomosing veins that are com- monly distinct enough to be seen when held to light. Grammitis dudleyi and G. wcrffii are two other species in Peru that Bishop ( 1 989b) included in his new genus Zygophlebia. Both have veins that form areoles (at least occasionally), but their laminae are so thick that the veins are impossible to discern without clearing the tissue. See treat- ment of these species for further discussions. Amazonas: Prov. Chachapoyas. Cerros Calla Calla above Leimcbamba. Hutchison & Wright 5554 (F, GH, NY, us), Wurdack 1743 (us). \\'urdack 1765 (GH, NY, uc, us. USM). Prov. Chachapoyas, Mathews "3281 '?(B). San TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 91 Martin: Prov. Mariscal Caceres, Chochos, NW corner of Rio Abiseo Park, Young & Leon 4744 (USM). Prov. Chanchamayo, Mina Pichita, above San Ramon, van der Werffel al. 8663 (MO). Cuzco: Altura de Sicre, Biies 1538 (us), 7556 (us). Prov. Urubamba, trail to Intipata, Leon & Cano 2100 (F, USM). 26. Grammitis dudleyi (Bishop) Stolze, comb. nov. Zygophlebia dudleyi Bishop, Amer. Fern J. 79: 1 13. 1989. TYPE: Peru, Cuzco, Prov. La Convencion, Cordillera Vilcabamba, NE of Rio Apurimac, Dudley 1 1144 (holotype, NA; isotypes, GH!, us!). Plants epiphytic. Stem short-creeping or as- cending, provided with linear or lanceolate, rather flaccid orange or yellow-brown scales, these 1.5- 4 mm long, 0.4-0.8 mm broad, subclathrate, with margins entire. Leaves monomorphic, articulate to the stem, 35-60 cm long, 6-8 cm broad. Petiole 18-30 cm long, 1-2 mm in diameter, dull yellow- ish or reddish brown, sparsely to densely provided with spreading, castaneous, unicellular trichomes, these to 5 mm long. Lamina coriaceous, deltate, not or scarcely reduced at base, deeply pinnatisect to nearly pinnate, the segments rather strongly as- cending and subdistant, linear-deltate, acute, the rachis and (sometimes) segments provided with widely scattered trichomes as on the petiole, or glabrate. Veins completely obscure (even when held to light), 1-2-forked, the branches often connivent distally and occasionally merging to form irregular areoles, hydathodes lacking. Sori slightly im- pressed, round to slightly elongate, in a single se- ries between costa and margin, at maturity often reaching both costa and margin. On trees, in thickets and cloud forests, 1550- 3400 m, Huanuco and Cuzco. Apparently endemic. Although veins were completely obscured with- in the thick segment tissue, careful clearing will reveal them to be essentially free. However, there is a tendency for vein branches to curve distally toward adjacent ones and occasionally even to merge. This tendency toward areolate venation, along with other characters, indicates a relation- ship with G. mathewsi and G. werffii, under which may be seen further discussion. Huanuco: SW slope of Rio Llullapichis watershed, as- cent of Cerros del Sira, Dudley 13249 (GH), 13406 (GH). Cuzco: Prov. La Convencion, Cordillera Vilcabamba, facing Apurimac Valley, Dudley 11218 (GH, us). 27. Grammitis werffii (Bishop) Stolze, comb. nov. Zygophlebia werffii Bishop, Amer. Fern J. 79: 115. 1989. TYPE: Peru, Pasco, border between Prov. Oxapampa and Prov. Pasco, van der Werffel al. 8570 (holotype, uc!; isotype, MO!). Plants terrestrial. Stem stout, short-creeping to ascending, densely provided with linear, rigid, lus- trous, deep orange to brown scales, these 5-1 0 mm long, 0.4-0.7 mm broad, nonclathrate, sparingly to amply setulose, the setulae whitish and spread- ing. Leaves monomorphic, articulate and approx- imate on the stem, 25-50 cm long, 3.5-7 cm broad. Petiole 6-26 cm long, 1-2 mm in diameter, lus- trous, deep castaneous to blackish, often glabres- cent near the base, but distally rather densely pro- vided with castaneous, rigid, unicellular trichomes, these 0.1-0.4 mm long. Lamina subcoriaceous, linear to narrow-elliptic or -oblong, not or scarcely reduced at base, deeply pinnatisect or fully 1 -pinnate proximally, the segments patent, linear, acute, the rachis with abundant trichomes like those of the petiole, the segments with scattered, minute, glandular, simple or branched pluricellular tri- chomes on surface and margins. Veins completely obscure (even when held to light), commonly once- forked, many of them merging distally to form areoles, hydathodes lacking. Sori superficial, com- monly round, in a single series on each side of the costa, essentially medial between costa and mar- gin, sporangia intermixed with brownish, glan- dular, viscid paraphyses (these mostly evident in immature sori). In elfin and cloud forests, on clay banks or in sphagnum, 1900-2700 m, Huanuco and Pasco. Apparently endemic. This is included by Bishop in his new genus Zygophlebia, related to G. mathewsii and G. dud- leyi in Peru by anastomosing veins, among other characters. However, veins are so completely ob- scure in G. werffii and G. dudleyi that the character cannot be used effectively in the key. Consequent- ly, more practical means have been employed to key out these species. Huanuco: Vilcabamba, Hacienda on Rio Chinchao, Macbride 5145 (F, in part). 28. Grammitis bipinnata Stolze, sp. nov. Caulis erectus vel repens, paleis lanceolatis, non clathratis, castaneis, 1-2 mm longis, setulosis, se- tulis niveis vel hyalinis. Folia 1 0-30 cm longa, 2- 92 FIELDIANA: BOTANY 6 cm lata, subfasciculata. Petiolus 3-20 cm longus, 0.3-0.5 mm latus, trichomatibus 0. 1-0.3 mm lon- gis, simplicibus vel ramosis, septatis, et trichoma- tibus 1.5 mm longis, castaneis, unicellularis inter- mixtis. Lamina bipinnata, basis abrupte reducta, paleis iis petioli equalibus. Pinnae 10-20 jugae, ascendentes. Venae simplices vel ramosae. Hy- dathodi nulli. Plants pendent, probably epiphytic. Stem sub- erect or very short-creeping, provided with lan- ceolate, nonclathrate scales, these 1-2 mm long, lustrous, castaneous, the margins with rigid, whit- ish or hyaline setae. Leaves lax, articulate to the stem, crowded to subfasciculate, 2-pinnate (rarely 3-pinnate), 10-30 cm long, 2-6 cm broad. Petiole 3-20 cm long, 0.3-0.5 mm in diameter, sparsely puberulent with simple or branched, septate tri- chomes 0.1-0.3 mm long, and with simple, spreading, unicellular, castaneous trichomes to 1.5 mm long, or glabrescent. Lamina firm-herbaceous, abruptly reduced at base, ovate or broadly lan- ceolate, with 10-20 pairs of ascending pinnae. Ra- chis thin and flexuous, amply provided with spreading, castaneous unicellular trichomes to 1.5 mm long, and also sparsely provided with simple or branched septate trichomes 0.1-0.3 mm long. Pinnae stalked, 1.5-7 cm long, linear, 1 -pinnate (rarely 2-pinnate), costa and segment margins pi- lose as on the rachis, and costa also with minute septate trichomes, ultimate segments sessile, 0.7- 3 mm long, 0.6-2 mm broad, obovate or spatulate. Veins indistinct or obscure, simple or 1 -forked in the segments, hydathodes lacking. Sori round, usually 1 to a segment, sporangia glabrous. TYPE— Peru, Cuzco, Prov. La Convencion, Loma Grande, Biies 2167 (holotype, us!; isotype, F!; pho- to, F of us). In wet forests, 3150-4100 m, Cuzco. Although no data are provided on any of the labels, the long and delicate, flexuous, pendent leaves suggest that this species may be epiphytic. Endemic. The type specimen and paratypes have all been determined previously as Polypodium variabile(= Grammitis variabilis), and there is certainly a su- perficial resemblance to the latter as well as to G. heteromorpha. However, hydathodes are present adaxially on the vein tips of these two species (al- though sometimes obscured by indument), where- as hydathodes are lacking in G. bipinnata. There are two kinds of long, unicellular laminar tri- chomes in G. heteromorpha: yellow or orange and unbranched, and these mixed with others that are castaneous and stellately branched. The long, uni- cellular trichomes are yellowish and simple in G. variabilis, and castaneous and simple in G. bipin- nata, but they are never stellate in these two spe- cies. Leaves of G. bipinnata and G. variabilis are at least 2-pinnate, but in G. heteromorpha they are 1 -pinnate with lamina once or twice forked. There has been much confusion attending the relationship of this species complex (see Copeland, 1956; Morton, 1967), and the entire group is in great need of revision. Further comparison of characters can be found in the treatment of G. heteromorpha. Cuzco: Altura de Sicre, Biies 1553 (us). Raccaypata, Vilcabamba, Biies 1606 (us). Prov. La Convencion, Loma Grande, Biies 2164 (F, GH, us), 2765 (us, monstrous form, with extremely long, pinnate pinnae), 2766 (us). Paucartambo, Acjanaco, subida a Quellhua Ccocha, Cano & Baldeon 4980 (F, USM). 29. Grammitis jamesonioides (Fee) Morton, Contr. U.S. Natl. Herb. 38: 108. 1967. Polypodium jamesonioides Fee, Mem. foug. 7: 59, t. 21, f. 4. 1857. TYPE: Colombia, Dept. Santan- der, Ocana, Schlim 399 (holotype, L!; photos, F, uc, us). Plants epiphytic. Stem erect, small, sparsely pro- vided with ovate, brown, nonclathrate scales, these less than 1 mm long, viscid, the margins entire or glandular-ciliolate. Leaves 1 -pinnate, inconspic- uously articulate, 12-30 cm long, 1-1.7 cm broad, axes and tissue moderately provided with ap- pressed, septate, viscid trichomes, these 0.1-0.2 mm long, simple, or occasionally branched, whit- ish or hyaline. Petiole 0.5-5 cm long, 0.2-0.4 mm in diameter. Lamina firm-herbaceous to charta- ceous, abruptly reduced at apex, gradually reduced at base, the basalmost pinnae rudimentary. Pinnae 0.5-1.2 cm long, 2-2.5 mm broad, approximate but discrete, ascending at a 70-80° angle, linear- to narrow-deltate, mostly acute, margins entire to crenulate, base rounded to truncate acroscopically, slightly decurrent basiscopically. Veins indistinct or obscure, free, short and unbranched, hyda- thodes present, but usually partially obscured by the broad, appressed laminar trichomes. Sori round, mostly inframedial, sometimes confluent at maturity. On tree trunks in montane forests, ca. 3350 m, Cuzco. Costa Rica; Colombia to Peru. TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 93 This may be confused with G. farinosa, from which it differs chiefly in its narrower leaves and viscid-puberulent, vs. farinose, lamina. With it probably should be included Polypodium azuay- ense Sodiro of Ecuador; however, this is not cer- tain, for the type of the latter was not examined during this study. Cuzco: Mollepata, Hacienda Pincopata, Bishop 2520 (uc). Prov. Calca, valley of Rio Calca, Biies 1805 (us). 30. Grammitis apiculata (Klotzsch) Seymour, Phytologia31: 176. 1975. Polypodium apiculatum Klotzsch, Linnaea 20: 378. 1 847. LECTOTYPE (designated here); Venezuela (as Colombia), Colonia Tovar, Moritz 247 (B!; photo, F; isolectotype, us!). Syntype: British Gui- ana (Guyana) (Rich. Schomburgk 1213 (B!, cat. #091628); another sheet of no. 1213 (B!, cat. #088656) is G. capillaris. Ctenopteris apiculata (Klotzsch) Copel., Philipp. J. Sci. 84:417. 1956. Ctenopteris subimpressa Copel., Philipp. J. Sci. 84: 463. 1 956. TYPE: Peru, Junin, E of Quimiri Bridge near La Merced, Killip & Smith 23842 (holotype, us!; photo, F). Lellingeria apiculata (Klotzsch) Smith & Moran, Amer. Fern J. 81: 83. 1991. Plants epiphytic. Stem short-creeping or decum- bent, provided with lanceolate or ovate, blackish or castaneous scales, these 1-3 mm long, clathrate, with margins amply setose, the setae hyaline or whitish. Leaves 10-28 cm long, 2-5 cm broad. Petiole 3-9 cm long, 0.5-0.8 mm in diameter, light or reddish brown, densely puberulent, the tri- chomes orange or whitish, 0. 1-0.2 mm long. Lam- ina firm-herbaceous to chartaceous, oblong-del- tate, subtruncate at base, or with a few proximal pairs of somewhat reduced pinnae, usually abrupt- ly reduced at apex to a pinnalike apex, pectinate, incised nearly or quite to the rachis, the segments (or pinnae) patent or slightly ascending, linear, ob- tuse or subacute, the rachis sparsely to amply pu- berulent on both sides as on the petiole. Veins indistinct to obscure, simple, hydathodes distinct. Sori round, superficial or slightly impressed, me- dial. In cloud forests or clearings, on trunks or branches of trees, 700-2600 m, Cajamarca, San Martin (?), Pasco, and Junin. Hispaniola; Mexico; Honduras; Costa Rica; Guyana to Colombia, south to Bolivia. Cajamarca: Prov. Jaen, Quebrada de Pajonal, above Tabaconas, Fosberg 27804 (us). Pasco: Prov. Oxapam- pa, trail between Santa Rosa de Chives and the San Matias signpost, Leon 328 (F, USM). Junin: Prov. Satipo, Gran Pajonal S of Chequitavo, D. Smith 5147 (MO). Department unknown: "Cumbasamma Mts." (Cumbasa, San Martin?), Steere (GH). 31. Grammitis pseudocapillaris (Rosenst.) Mor- ton, Contr. U.S. Natl. Herb. 38: 108. 1967. Polypodium pseudocapillare Rosenst., Meded. Rijks- Herb. 19: 17. 19 13. TYPE: Bolivia, above Tablas, Herzog 2190a (holotype, L; isotypes, uc!, us). Ctenopteris pseudocapillaris (Rosenst.) Copel., Phil- ipp. J. Sci. 84: 407. 1956. Lellingeria pseudocapillaris (Rosenst.) Smith & Mor- an, Amer. Fern J. 81: 86. 1991. Plants epiphytic. Stem short-creeping or decum- bent, provided with narrow-deltate, castaneous or blackish scales, these subclathrate, 1.5-3 mm long, with setose margins, the setae hyaline or whitish. Leaves fasciculate, usually pendulous, 8-50 cm long, 1.5-4 cm broad. Petiole 0.5-2 cm long, 0.2- 0.6 mm in diameter, brown or reddish brown, sparsely to moderately puberulent, the trichomes simple, branched at base or stellate, orange or whitish, 0.1-0.3 mm long. Lamina firm-herba- ceous, 1 -pinnate, narrow-elliptic, strongly reduced at apex and base, the rachis somewhat to (com- monly) strongly flexuous, sparsely puberulent as on the petiole. Pinnae ascending at 20-45° angles, narrow-deltate to linear (proximal ones mere au- ricles), approximate to well spaced, inequilateral at base, subtruncate or rounded acroscopically, de- current basiscopically. Veins indistinct or obscure, free, simple, hydathodes evident. Sori round, su- perficial or slightly impressed, inframedial. On tree trunks in montane forests, 3050-3740 m, Cuzco. Colombia to Bolivia. In addition to the key characters, this can often be distinguished from the closely related G. sub- sessilis by the occurrence of some branched and stellate trichomes mixed among the simple ones, especially on the petiole. Petiole trichomes of G. subsessilis are always simple. Cuzco: Huadquina, Biies 1010 (us). 15 km from Qui- llabamba, Ellenberg 4774a (GH). Prov. Paucartambo, forest on Cerro Macho Cruz, Leon et al. 2784 (F, USM). Department unknown: Mat hews 1102 (BM). 32. Grammitis tunguraguae (Rosenst.) Morton, Contr. U.S. Natl. Herb. 38: 109. 1967. 94 FIELDIANA: BOTANY Polypodium tunguraguae Rosenst., Repert. Spec. Nov. Regni Veg. 7: 307. 1909. TYPE: Ecuador, Prov. Tungurahua, Mt. Tungurahua, Spruce (holotype, p, Herb. Bonap.; isotype, BM, frag., us!). Ctenopteris tunguraguae (Rosenst.) Copel., Philipp. J. Sci. 84: 394. 1956. Lellingeria tunguraguae (Rosenst.) Smith & Moran, Amer. Fem J: 81: 88. 1991. Plants terrestrial or epipetric. Stem short -creep- ing or decumbent, provided with narrow-lanceo- late, gray-brown scales, these 1-2 mm long, clath- rate, margins densely setose, the setae hyaline or pale orange. Leaves 5-15 cm long, 0.8-1.5 cm broad. Petiole 0.5-4 cm long, 0.3-0.4 mm in di- ameter, light or reddish brown, densely puberu- lent, the trichomes orange or whitish, 0. 1-0.2 mm long. Lamina firm-herbaceous to chartaceous, 1 -pinnate, narrow-elliptic, strongly reduced at both ends, the rachis atropurpureous or blackish, sparsely puberulent abaxially, densely so adaxi- ally. Pinnae patent or slightly ascending, oblong, obtuse at apex, adnate and subequilateral at base, subtruncate to slightly dilated. Veins indistinct to obscure, simple, hydathodes evident. Sori round, superficial or slightly immersed, medial to infra- medial. In forests or at edges of roads, on the ground or in rock crevices, 2000-3000 m, Cajamarca and Cuzco. Ecuador; Peru. This is one of the most diminutive of the fully pinnate species of Grammitis, with which smaller specimens of G. apiculata might be confused. However, the latter is epiphytic, with lamina sub- truncate at the base and segments relatively much longer and narrower. Cajamarca: Prov. Cutervo, La Pucarillo, Lopez et al. 6718 (F, HUT). Cuzco: Machu Picchu, Bishop 1999 (uc), 2576 (uc). Prov. Urubamba, summit of Huayna Picchu, Hutchison 1755 (F, K., NY, uc). Prov. Paucartambo, bor- der of Manu National Park, road to Pilcopata, Leon 2175 (USM). 33. Grammitis subsessilis (Baker) Morton, Contr. U.S. Natl. Herb. 38: 107. 1967. Polypodium pteropus Hooker, Sp. fil. 4: 192, /. 2755. 1863, not Blume, 1828. LECTOTYPE (desig- nated here): Ecuador, Mt. Abitagua. Spruce 5276 (K!). Polypodium subsessile Baker, in Hooker & Baker, Syn. fil. 329. 1 867, nom. nov. for P. pteropus Hooker and with the same type. Ctenopteris pteropus John Sm., Hist. fil. 185. 1875. Ctenopteris subsessilis (Baker) Copel., Philipp. J. Sci. 84:411. 1956. Lellingeria subsessilis (Baker) Smith & Moran, Amer. FernJ. 81: 87. 1991. Plants epiphytic, occasionally epipetric. Stem short-creeping or decumbent, provided with del- tate, gray-brown to blackish scales, these 2-4 mm long, obviously clathrate, the proximal lumina in 5-10 series across the scale and clear, margins densely setose, the setae hyaline or whitish. Leaves often pendulous, (8-) 12-40 cm long, (2-)3.5-7 cm broad. Petiole 0.5-3 cm long, (0.4-)0. 5-0.9 mm in diameter, brown or blackish, moderately to densely puberulent, the trichomes unbranched, or- ange or whitish, 0.1-0.2 mm long. Lamina firm- herbaceous to chartaceous, 1 -pinnate or deeply pinnatisect, elliptic, strongly reduced at apex and base, the rachis straight or slightly flexuous, sparsely puberulent abaxially, densely so adaxially. Pinnae spreading or (more commonly) strongly ascend- ing, narrow-deltate to linear (proximal ones mere auricles), usually well spaced and joined by a nar- row herbaceous wing at a broadly rounded sinus, segment base subequilaterally and conspicuously dilated on each side. Veins obscure, free, simple, hydathodes evident. Sori round, slightly to rather deeply impressed, inframedial. In cloud or elfin forests, usually pendulous on trees, or occasionally growing on mossy rocks or rocky cliffs, 1300-3750 m, Amazonas, Huanuco, Pasco, and Cuzco. Costa Rica; Panama; Venezuela; Colombia to Bolivia. Of the five syntypes (K) cited from Colombia and Ecuador, Spruce 5276 is selected as lectotype because it best displays the typical variation of the lamina shape in this species and also was appar- ently the specimen from which Hooker's original illustration was drawn. Leaves of this species can be highly variable; those of some specimens are greatly reduced in size but more erect, but this probably is due to their growing on rocks. Eiphytes are invariably pendulous, much longer, and rather flexuous. Ctenopteris obovata Copel. (syn. Polypodium pendulum, var. boliviense Rosenst.), from Bolivia, probably should be included here, at least as a low- elevation variant. It differs from G. subsessilis only in its very short and broad lamina. Grammitis recondita Morton (Ecuador) is very similar in gen- eral aspect, for the rachis is thin and flexuous and pinnae are strongly ascending and have inequilat- eral bases. However, hydathodes are lacking in G. TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 95 recondita, lamina texture is very thin, and the few reduced basal pinnae are neither auriculiform nor broader than long. Amazonas: Mendoza, Woytkowski 8216 (GH, MO). Huanuco: SW slope of Rio Llullapichis watershed, slopes of Cerros del Sira, Dudley 13409 (GH). Cani, 7 mi NE of Mito, Macbride 3396 (F, us). Pasco: Prov. Oxapampa, summit of San Matias, Leon et al. 318 (F, USM). Prov. Oxapampa, San Alberto, Cordillera de Yanachaga, van der Werffet al. 8492 (MO). Cuzco: Cerro Chuyapi, Bites A39 (us). Prov. La Convencion, Cordillera Vilcabamba, Dudley 11106 (GH). Prov. La Convencion, Canchyoj?, Vargas 23314 (GH). these segments are relatively broad and thin in texture. In others, including the Peruvian speci- mens, segments are thicker, more constricted, and sori occur throughout their length. No other char- acteristics have been observed to support sepa- ration into two separate taxa, and the condition is possibly due to individual plants' responses to dif- ferent habitats. Cuzco: Prov. La Convencion, Huayapata, 7 km from village of Incatambo, Peyton & Peyton 918 (GH, MO). Urubamba, Macchu Picchu, above Rio Mandor, Peyton & Peyton 1314 (MO). 34. Grammitis major (Copel.) Morton, Contr. U.S. Natl. Herb. 38: 109. 1967. Polypodium tenuiculum var. acrosorum Hieron., Bot. Jahrb. Syst. 34: 510. 1904, as "acrosora," not P. acrosorum Kunze, 1834. TYPE: Ecuador, Azuay, near Yerbabuena, Lehmann 5727 (holotype, B?; isotypes, F!, us). Ctenopteris major Copel., Philipp. J. Sci. 84: 455. 1956. Type same as for Polypodium tenuiculum var. acrosora. Lellingeria major (Copel.) Smith & Moran, Amer. Fern J. 81: 84. 1991. Plants epiphytic or (outside Peru) terrestrial or epipetric. Stem short-creeping or decumbent, pro- vided with deltate or oblong-deltate, gray- or light brown, iridescent scales, these (4-)5-7 mm long, narrow-clathrate, the proximal lumina ( 1 2-) 1 5-20 across and mostly occluded, margins densely se- tose, the setae hyaline or whitish. Leaves 15-38 cm long, 2-5 cm broad. Petiole 2-9 cm long, 0.6- 1.2 mm in diameter, reddish brown to atropur- pureous, moderately to densely puberulent, the trichomes pale orange to white, 0. 1-0.3 mm long. Lamina chartaceous, deeply pinnatisect to 1 -pinnate, narrow-elliptic, strongly reduced at both ends. Rachis castaneous to atropurpureous, adax- ially densely puberulent like the petiole, glabrous or sparsely puberulent abaxially. Pinnae patent or slightly ascending, deltate, obtuse to subacute at apex, adnate and subequally dilated at base. Veins indistinct to obscure, simple, hydathodes evident. Sori round or oblong, superficial or slightly im- mersed, medial, often confined to distal half of pinnae. On tree trunks in humid forests and wooded ravines, 2560-3430 m, Cuzco. Ecuador; Peru. On the type and most other Ecuador specimens, sori are confined to the distal half of segments; 35. Grammitis gracilis (Hooker) Stolze, comb. nov. Polypodium gracile Hooker, Bot. misc. 2: 239. 1831. TYPE: Peru, Pasco, Huayllay, Cruckshanks, in 1830 (holotype, K; isotype. GH!; photo, F of GH!). Ctenopteris gracilis (Hooker) John Sm., Hist. fil. 185. 1875. Habit unknown, dried plants with a faint aroma like that of pine leaves. Stem short-creeping, pro- vided with narrow-deltate, gray-brown scales, these subclathrate, 2-3 mm long, the margins entire. Leaves subfasciculate, articulate to very short phyllopodia, pendulous, 1 8-26 cm long, to 2.2 cm broad. Petiole 5-7 cm long, 0.5-0.7 mm in di- ameter, light brown or castaneous, sparsely to am- ply puberulent, the trichomes simple or branched, septate, tortuous, 0.2-0.3 mm long. Lamina char- taceous, 1 -pinnate-pinnatifid, narrow-elliptic, re- duced at apex and base, the rachis slightly flexu- ous, puberulent as on the petiole. Pinnae strongly ascending at a 30—45° angle, sessile, linear or lan- ceolate, well spaced, incised about halfway to the costa into 4-6 pairs of broad, obtuse lobes. Veins obscure, simple, one in each segment and termi- nating in a hydathode adaxially. Known thus far only from the type, ca. 4500 m, Pasco. It is interesting to discover that the fragrance of the isotype still persists 1 60 years after it was col- lected. To the author it is much like the aroma of some species of Pinus, and the odor can be de- tected also in several related species, such as G. andicola, G.firma, G. melanosticta, G. monilifor- mis, and G. xiphopteroides. Most of these are to be included by Smith et al. in a new genus to be named Melpomene. The species is quite distinc- tive in its strongly ascending, pinnatifid pinnae, and it is curious that it has not been collected since 1830. 96 FIELDIANA: BOTANY 36. Grammitis phlegmaria (John Sm.) Proctor, Rhodora 68: 467. 1966. Polypodium phlegmaria John Sm., London J. Bot. 1: 194. 1842. TYPE: Venezuela, Edo. Bolivar, near Mt. Roraima, Schomburgk 161 (holotype, K.!; photos, F, us). Ctenopteris phlegmaria (John Sm.) Copel., Philipp. J. Sci. 84: 435. 1956. Lellingeria phlegmaria (John Sm.) Smith & Moran, Amer. Fern J. 81: 86. 1991. Plants epiphytic. Stem erect, provided with ovate, brownish scales, these clathrate, 1-2 mm long, cordate at base, margins entire. Leaves es- sentially glabrous, fasciculate, articulate to the stem, pendent, 10-25 cm long, 1-1.7 cm broad. Petiole 0.2-0.5 cm long, 0.2-0.3 mm in diameter, brown or black. Lamina thin-herbaceous, pinnatisect nearly to rachis, linear to narrow-elliptic, reduced at apex and base, the rachis blackish or obscured by the laminar tissue. Pinnae sessile, broadly and obtusely deltate, entire, or (at least on the acro- scopic margin) broadly crenulate, inequilateral at base, broadly rounded to subauriculate acroscop- ically, strongly decurrent basiscopically. Veins dis- tinct or indistinct, free, short, pinnately branched in the segments, hydathodes evident. Sori round, 3-5 to a segment. Thus far known in Peru from a single specimen, on trees, 2200 m, Cuzco. Honduras; Costa Rica; Guyana; Venezuela; Co- lombia to Peru. Proctor (Rhodora 68: 467. 1966) separated ma- terial from the Lesser Antilles as var. antillana, which differs from var. phlegmaria in the narrower and subentire segments. Cuzco: Sahuayacu, Cerro Sombreruyoc, Biies 825 (us). 37. Grammitis youngii Stolze, sp. nov. Plantae epiphyticae. Caulis repens, 1.0-1.4 mm crassus, paleis abundantibus, 5-7 mm longis, 0.4- 0.7 mm latis, clathratis, lanceolatis, attenuatis, basi cordatis. Folia 15-30 mm longa, 1.5-3 cm lata. Petiolus 3-9 cm longus, 0.6-1.2 mm latus, minute puberulus, trichomatibus 0. 1-0.3 mm longis, sep- tatis, simplicibus vel ramosis. Lamina coriacea, 1 -pinnata vel profunde pinnatisecta, basis et apicis valde reducta. Pinnae lineato-deltatae, ascen- dentes, basi subaequilateris, marginibus integris et revolutis. Venae liberae, obscurae, hydathodis ad- sunt. Sori superficial!, mediali vel inframediali. Plants epiphytic. Stem short- to long-creeping, 1-1.4 mm in diameter, pruinose, abundantly pro- vided with iridescent brown or gray-brown scales, these 5-7 mm long, 0.4-0.7 mm broad, lanceolate, attenuate, clathrate, cordate at base, margins en- tire. Leaves approximate to well spaced, neither articulate nor aromatic, 15-30 cm long, 1.5-3 cm broad. Petiole 3-9 cm long, 0.6-1.2 mm in di- ameter, brown or blackish, sparsely to moderately puberulent, the trichomes 0.1-0.3 mm long, sep- tate, simple or branched. Lamina coriaceous, in- cised nearly or quite to the rachis, narrow-elliptic, strongly reduced to apex and base, the rachis and adaxial laminar tissue sparsely and minutely pu- berulent as on the petiole, or glabrescent. Pinnae sessile, linear-deltate, 6-8 times as long as broad, ascending at a 60-70° angle, the margins entire and usually revolute, subequilateral at base, sub- truncate acroscopically, slightly dilated basiscop- ically. Veins obscure, free, short and ascending, 6- 1 2 on each side of the costa, each terminating in a hydathode adaxially. Sori superficial, round, me- dial or inframedial, lacking paraphyses. TYPE— Peru, San Martin, Prov. Mariscal Ca- ceres, Puerta del Monte, forest patch above tim- berline, Young 1684 (holotype, USM!; isotype, F!). Apparently endemic. On trees in forests or forest patches, 3000-3400 m, Piura, Amazonas, San Martin. This is named for Dr. Ken Young, who not only gathered half of the known specimens but has been collecting in Peru for several years. The species is part of the group of G. moniliformis, which is dis- tinguished by large, clathrate, stem scales with en- tire margins and a cordate base, coriaceous pinnae with revolute margins, and laminae having scat- tered, septate, often branched trichomes on the petiole and rachis instead of long, spreading tri- chomes. Pinnae of G. youngii are linear-deltate and somewhat ascending. Piura: Road from Piura to Huancabamba, El Tambo, Scolnik 1413 (us). Amazonas: E of Balsas, Osgood & Anderson 70 (F). San Martin: Prov. Mariscal Caceres, Rio Abiseo National Park, near Mirador, Ledn 2132 in part (uc). Prov. Mariscal Caceres, NW corner of Rio Abiseo National Park, Young & Leon 4559 (USM), 4609a (USM). 38. Grammitis melanosticta (Kunze) Seymour, Phytologia31: 179. 1975. Polypodium melanostictum Kunze, Linnaea 9: 44. 1834. TYPE: Peru, Cuesta de Carpis, Poeppig, in 1 829 (holotype, LZ, destroyed; isotype, w). TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 97 Ctenopteris melanosticta (Kunze) Copel., Philipp. J. Sci. 84: 385. 1956. Plants epipetric or epiphytic, scarcely to strongly aromatic. Stem short-creeping to erect, provided with a few ovate or lanceolate, clathrate scales, these l-2(-2.5) mm long, 0.3-0.5 mm broad, mar- gins entire, cordate at base, the cells 6-10 across at widest point, their walls blackish, not or scarcely lustrous. Leaves glabrous, or with a few scattered, septate trichomes ca. 0. 1 mm long on the petiole and rachis abaxially, approximate to fasciculate, not articulate, 6-17 cm long, 0.7-1.2 cm broad. Petiole ca. 1 cm long, 0.4-0.6 mm in diameter, brown or blackish. Lamina subcoriaceous, pin- natisect nearly to the rachis, linear to narrow-el- liptic, reduced at apex and base, the rachis black- ish. Segments about twice as long as broad, patent, joined near the rachis by a narrow wing, oblong, obtuse, margins entire and essentially plane, sub- equilateral at base, subtruncate acroscopically, slightly dilated basiscopically, dotted with reddish stomates abaxially. Veins indistinct or obscure, free, 2-3 pairs ascending from the costa and each terminating in a hydathode adaxially. Sori round, 1-4 to a segment. On dry rock faces or on tree trunks in forests, 600-1800 m, San Martin, Huanuco, Junin. Southern Mexico; Guatemala; Costa Rica; Cuba; Hispaniola; Venezuela; Colombia to Bolivia. Although rather widespread in its distribution, collections of this species are nowhere very com- mon, but perhaps it has been overlooked because of its small and inconspicuous leaves. Stolze (Fieldiana, Hot., n.s., 6: 26 1 . 1 98 1) included it with G. moniliformis in Guatemala, and indeed the two are very similar. The principal difference is in the stem scales, which are large and iridescent in G. moniliformis, usually with brown cell walls, whereas scales of G. melanosticta are small and inconspicuous, scarcely lustrous, and with black- ish cell walls. Also, lamina segments of the latter are consistently about twice as long as broad and dotted with red stomates; segments of G. monili- formis are commonly (but not always) about as broad as long and are eglandular. It is not certain if the description has been prop- erly applied to the name of this species, for no type material was available for this study. However, in the original description the segments are described as "subtus punctulatis," which apparently refers to the red stomates. Nearest allies of this species, at least in Peru, have normal stomates. San Martin: Prov. Mariscal Caceres, Tocache Nuevo, J. Schunke V. 5723 (F, NY). Huanuco (as San Martin): E of Tingo Maria, Allard 20603, 20731, 21363, 21469 (all us). Junin: La Merced, Hacienda Schunke, Macbride 5638 (F). Chanchamayo Valley, C. Schunke 315 (F), 1548 (F). 39. Grammitis erecta Morton, Phytologia 22: 72. 1971. TYPE; Ecuador, Mt. Tunguragua, Spruce 5279a (holotype, K). Plants terrestrial or epipetric, not aromatic. Stem long-creeping, usually decumbent, sometimes pru- inose, 1-3 mm in diameter, abundantly provided with lanceolate, iridescent, clathrate scales, these 4-8 mm long, 0.5-1 mm broad, margins entire, cordate at base, the cells 1 5-40 across at the widest point, their walls light or reddish brown. Leaves with a few, scattered, minute, appressed, septate trichomes ca. 0.1 mm long, or glabrescent, well spaced, erect and continuous on the stem, 10-18 cm long, 0.7-1.4 cm broad. Petiole 1—4 cm long, 0.7-1 mm in diameter, brown. Lamina coriaceous, deeply pinnatisect, linear to narrow-elliptic, re- duced at apex and base, the rachis reddish brown. Segments 2—4 times as long as broad, patent, me- dial ones narrowly or broadly oblong, obtuse, mar- gins entire and revolute, subequilaterally dilated at base. Veins obscure, free, 3-4 pairs to a segment, each terminating in a hydathode adaxially. On wet hummocks or bryophytes or on wet rocks, 3100-3500 m, Cuzco. Venezuela; Ecuador; Peru. The habit of G. erecta is distinctive. The stem is stout and very long-creeping, eventually as- cending, and upon it are borne well-spaced, rigid, very narrow, ascending leaves. The 30-60 pairs of rigid segments are perpendicular to the rachis. The species is closely allied with G. moniliformis and, thus far, is known in Peru from but one collection. Cuzco: Prov. La Conversion, Cordillera Vilcabamba, Dudley 11 062 (GH). 40. Grammitis moniliformis (Sw.) Proctor, Brit. FernGaz. 9: 219. 1965. Polypodium moniliformeSw., Syn. fil. 33. 1806. TYPE: Peru, ex Lagasca (holotype, s; photos, GH, us). Ctenopteris moniliformis (Sw.) John Sm., Hist. fil. 1 84. 1875. Plants terrestrial, epipetric, or epiphytic, aro- matic. Stem short- to long-creeping, 1-3 mm in diameter, abundantly provided with ovate to lan- 98 FIELDIANA: BOTANY ceolate, clathrate, iridescent scales, these 3-8 mm long, 0. 1-1 .2 mm broad, margins entire, base cor- date, the cells ( 1 2-) 1 6-30 across at the widest point, their walls reddish brown. Leaves approximate to subfasciculate (rarely to 1 cm apart), not articulate, 4-25 cm long, 0.4-1.5 cm broad. Petiole 0.7-6 cm long, 0.4-1 mm in diameter, brown (rarely black- ish at base), sparsely to moderately provided with septate, simple or branched trichomes about 0. 1 mm long, but occasionally with some dark, rigid ones to 1 mm long. Lamina coriaceous, cut nearly or quite to the rachis, linear, reduced at apex and base, the rachis brown, glabrous, or puberulent as on the petiole. Pinnae (or segments) 1-1.5 times as long as broad, patent, semicircular to broadly oblong, obtuse, margins entire and mostly revo- lute, subequilateral at base, glabrous. Veins ob- scure, free, with 1-2 short branches that terminate in hydathodes adaxially. Sori round, several to a pinna, slightly impressed. In forests on tree trunks, humus or rocky soil, or on open road cuts or exposed rock cliffs, (1800-) 2150-4600 m, Piura to Amazonas, south to Aya- cucho, Cuzco, and Puno. Southern Mexico to Panama; Greater Antilles; Colombia and Venezuela south to Bolivia and Brazil. This is perhaps the most common species of Grammitis in Peru, and it is found in many hab- itats. Its leaves vary considerably depending upon where the plant grows. Those on tree trunks, es- pecially in elevations below 2500 m, tend to be more robust, with longer and broader laminae, whereas the plants found on rocks or crevices of cliffs and in higher elevations are apt to be narrow and stunted. The robust plants are sometimes sep- arated as G. assurgens (Maxon) Morton (Ecuador), but they differ in no other way than size, and even this is not wholly consistent with their habitat or elevation. Several higher elevation specimens from Peru are quite robust. Leaves consistently have the pungent odor discussed under G. gracilis. Besides G. assurgens, there are several other taxa that form a natural group with G. moniliformis, but these definitely merit distinction. Grammitis erecta and G. melanosticta (under which see fur- ther comparisons) are similar in leaf shape but have other clear but subtle differences. Although G. youngii surely belongs in this group, it can be more quickly distinguished by its broad lamina and linear segments. Perhaps G. moniliformis is most easily confused with G. flabelliformis (under which see further discussion). Piura: Prov. Huancabamba, above Canchaque on road to Huancabamba, Hutchison 1638 (uc). Lambayeque: Prov. Ferrenafe, near Cerro Punamachay on trail to La- guna Hualtaco, Dillon & Skillman 4117 (F, GH). Caja- marca: Prov. Chota, Laguna Yahuarcocha above Inca- huasi, Sagdstegui et al. 12899 (F, GH). Amazonas: Prov. Chachapoyas, Cerros Calla Calla above Leimebamba, Hutchison & Wright 5665 (F, GH, uc). La Libertad: Prov. Bolivar, Las Quinuas, Ldpez & Sagdstegui 3347 (F, GH, MO,UC). San Martin: Prov. Mariscal Caceres, NW corner of Rio Abiseo National Park, Young & Leon 4542 (USM), 4750 (USM). Ancash: Above Yungay near Laguna Llan- ganuco, Tryon & Tryon 6560 (GH). Huanuco: Carpish, Coronado 63 (GH, uc, us). Lima: Near Antaicocha, Cerro Colorado, E of Canta, Pennell 14692 (F, GH, us). Pasco: Prov. Oxapampa, Dist. Chontabamba, road to top of San Gutardo, Leon et al. 525 (USM). Huancavelica: Prov. Huanuco, Machajhuay, Tovar877 (GH, USM). Ayacucho: Pampalca, between Huanta and Rio Apurimac, Killip & Smith 23295 (NY, us). Apurimac: Forests of Ampay, Vargas 1063 (GH). Cuzco: Prov. La Convencion, NE of Hacienda Luisiana and Rio Apurimac, Dudley 11 138 (GH, us). Madre de Dios: Pinasniocj, Cook 1926 (us). Puno: Prov. Sandia, between Sandia and Cuyocuyo, Fe- rreyra 16812 (GH, USM). 41. Grammitis anfractuosa (Klotzsch) Proctor, Rhodora63: 35. 1961. Polypodium anfractuosum Klotzsch, Linnaea 20: 375. 1 847. TYPE: Venezuela, Merida, Moritz 330 (ho- lotype, B!, photo, F; isotypes, B!, us). Polypodium monticola Klotzsch, Linnaea 20: 377. 1 847; reduced to P. anfractuosum by Hieronymus (Hedwigia 48: 251. 1909). LECTOTYPE (desig- nated here); Peru, Huanuco, Muna, Ruiz 58 (B!, photo, F; frag., B!). Ctenopteris anfractuosa (Klotzsch) Copel., Philipp. J. Sci 84: 431. 1956. Plants epiphytic, rarely epipetric. Stem small, usually ca. 1 mm thick, bearing 1-few leaves at intervals along long-creeping, stoloniform roots, sparsely provided with narrow-deltate, gray-brown to blackish scales, these 1 mm or less, subclathrate (lumina much constricted), the margins naked. Leaves 5-14 cm long, 0.4-1.3 cm broad. Petiole 0.2-2 cm long, 0.2-0.5 mm in diameter, usually marginate throughout, sparsely or moderately pro- vided with spreading, reddish to castaneous, uni- cellular trichomes, these 0.4-1 mm long. Lamina firm-herbaceous to chartaceous, linear-elliptic, deeply pinnatisect nearly to the rachis, strongly and gradually reduced at both ends, pilose as on the petiole, especially long and abundant among the sori. Segments 2-6 mm long, 1.5-2.5 mm broad, oblong or oblong-deltate, obtuse or sub- acute, occasionally host to black, clavate fungi, margins entire. Veins simple, obscure, the hyda- thodes distinct and sometimes encrusted with TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 99 small, calcareous deposits. Sori 1-2 pairs on a seg- ment, sporangia glabrous. In wet forests or wooded ravines, on stumps, logs, or trunks and branches of trees, rarely on moss-covered rocks, 1400-2750 m, along the Cor- dillera Central from Cajamarca and Amazonas to Cuzco. Southern Mexico to Panama; Guadeloupe and the Greater Antilles; Guianas; Venezuela and Co- lombia to Peru. This little fern is distinguished especially by the very small, erect stems that bear only one or a few leaves and that proliferate by long, slender, sto- loniform roots. This habit has not been observed in any other species of Grammitis in Peru, al- though there are several species ofAsplenium with the stems connected in this manner. The black, clavate fungi that so commonly infest species in the G. semihirsuta complex are found but occa- sionally on the lamina of G. anfractuosa, even though they have been reported as quite prevalent on specimens from Mexico and the Antilles. Cajamarca: Prov. Jaen, Rio Manchara valley E of Ta- baconas, Hodge 6102 (us). Amazonas: Prov. Chacha- poyas, Cerros Calla Calla above Leimebamba, Hutchison & Bennett 4754 (F), 4754 in part (GH, uc). Prov. Bongara, Dist. Yambrasbamba, Tillett 673-231 (F, GH). San Mar- tin: Rioja, trail to Venceremos and Rio Serranoyacu, Knapp & Alcorn 7776 (F, MO). Near Tarapoto, Spruce 4642 (GH, p, us). Huanuco: Prov. Huanuco, 5 km from Carpish Pass, between Acomayo and Chinchao, Tryon & Tryon 5310 (USM), 5310-1/2 (F, GH, us), Tryon 5310 at GH is G. moniliformis. 42. Grammitis firma (John Sm.) Morton, Contr. U.S. Natl. Herb. 38: 110. 1967. Polypodiumfirmum Klotzsch, Linnaea 20: 378. 1847, not Kaulf., 1827. LECTOTYPE (designated by Looser, Rev. Univ. Chile 36: 75. 1951): Guyana, Rich. Schomburgk 1 170 (B; isolectotype, K.!, pho- tos, F & us of K). Ctenopteris firma JohnSm., Hist. fil. 184. 1875. Nom. nov. for Polypodium firmum Klotzsch and with the same type. Polypodium aromaticum Maxon, Proc. U.S. Natl. Mus. 27: 743. 1904. TYPE: Jamaica, Blue Mt. Peak, Underwood 1449 (holotype, NY!; isotype,us!; pho- to, F of us). Polypodium herzogii Rosenst., Repert. Spec. Nov. Regni Veg. 6: 176. 1908. TYPE: Bolivia, Prov. Cochabamba, Inca Corral, Herzog 783 (holotype, s; isotype, us!; photo, F of us). Grammitis aromatica (Maxon) Proctor, Brit. Fern Gaz. 9: 218. 1965. Plants epiphytic or epipetric, strongly aromatic. Stem 1.5-4 mm in diameter, short-creeping to de- cumbent, provided with narrow-deltate or -lan- ceolate, thin, iridescent gray-brown scales, these 3-5 mm long, clathrate, 10-15 cells across at scale base and the lumina large, clear and elongate, mar- gins naked. Leaves 8-28 cm long, (2-)3-7 cm broad. Petiole 1.5-7 cm long, amply provided with cas- taneous, rigid, unicellular trichomes to 1 mm long. Lamina 1 -pinnate, chartaceous to subcoriaceous, elliptic or oblong-elliptic, slightly and abruptly re- duced at base, dark unicellular trichomes 0.5-1 mm long on tissue and rachis abaxially, 0.3-0.5 mm long on rachis adaxially. Pinnae (0.6-)l-4 cm long, linear, many proximal ones deflexed, (4-)6- 14 times longer than broad, adnate, acute or sub- acute, conspicuously inequilateral at base, strongly dilated acroscopically, subtruncate basiscopically, margins entire and usually strongly revolute. Veins simple, 8-20 pairs on a pinna, hydathodes distinct. Sori 6-12 pairs on larger pinnae, sporangia gla- brous. In wet forests, on tree trunks, on boulders, and among rocks, 1900-3050 m, along the Cordillera Central from Cajamarca and Amazonas to Cuzco. Jamaica; southern Mexico; Guatemala; Costa Rica; Venezuela and Guyana; Colombia to Boliv- ia. The character most clearly distinguishing this species from its nearest allies is the many rigid, linear pinnae that are strongly deflexed in the prox- imal half of the lamina. It is also distinctive in the pungent odor of its leaves, persistent even on older herbarium specimens, as discussed under G. graci- lis. Cajamarca: Colasay, Woytkowski 7009 (MO, us). Ama- zonas: Leimebamba, Woytkowski 7839 (MO, us). Huanu- co: Carpish, Coronado 79 (us). Prov. Huanuco, Mirador along road from Acomayo to Chinchao, Mexia 4142 (uc). Pasco: Prov. Oxapampa, "Palamazu" (Palcazu), van der Werffet al. 8411 (MO). Junin: Prov. Tarma, Agua Dulce, Woytkowski 35480 (uc). Cuzco: Prov. Urubam- ba, Machu Picchu, Ferreyra 9889 (GH). Prov. Urubam- ba, Machu Picchu near Winay Wayna, Leon et al. 2108 (F). 43. Grammitis pilosissima (Mart. & Gal.) Mor- ton, Contr. U.S. Natl. Herb. 38: 1 14. 1967. Polypodium pilosissimum Mart. & Gal. Nouv. Mem. Acad. Roy. Sci. Bruxelles 15: 39, t. 9,f. 2. 1842. LECTOTYPE (designated by A. R. Smith, Fl. Chiapas 2: 124. 1981): Mexico, Veracruz, Zacu- apan, Galeotti 6379 (BR, frag., us; isolectotype, K; photo, us of BR). 100 FIELDIANA: BOTANY Ctenopteris pilosissima (Mart. & Gal.) Copel., Philipp. J. Sci. 84: 390. 1955. Plants epiphytic, rarely epipetric, not or slightly aromatic. Stem 1-3 mm in diameter, short- to long-creeping, provided with narrow-deltate or lanceolate, thin, iridescent, gray-brown scales, these 3-6 cm long, clathrate, 8-15 cells across at scale base and the lumina large, clear and elongate, scale margins naked. Leaves 8-27 cm long, 0.8-1.5 cm broad. Petiole 1.5-9 cm long, amply provided with castaneous, unicellular trichomes 1-2.5 mm long. Lamina 1 -pinnate, chartaceous to subcoriaceous, elliptic, not or abruptly reduced at base, proximal 0-3 pairs somewhat shorter than longest ones, ra- chis on both sides with dark, unicellular trichomes about 1 mm long or glabrescent. Pinnae (0.5-)1- 4 cm long, narrowly oblong-deltate to linear-del- tate, mostly ascending at 60-80°, 4-10(-16) times longer than broad, adnate, acute, slightly and sub- equilaterally dilated at base, margins entire and usually strongly revolute. Veins simple, 4—1 6 pairs on a pinna, hydathodes distinct. Sori (3-)4-l 2 pairs on larger pinnae, sporangia glabrous. In wet forests or thickets, on trunks and branch- es of trees, rarely on rocks, (2000-)2450-4000 m, Piura, La Libertad, San Martin, Huanuco, and Cuzco. Southern Mexico; Guatemala; Costa Rica; Ven- ezuela and Colombia to Brazil and Bolivia. This species varies greatly in size, especially in South America, and specimens at the extreme size limits understandably may be determined as dif- ferent species. Leaf width in Mexico and Central America rarely exceeds 2.5 cm, and pinnae are deltate or oblong-deltate; but in South America the lamina is often 3—4 cm broad, and many of the pinnae are linear beyond the dilated base. Grammitis pilosissima is especially distinguished from related species by the acute, ascending pin- nae, with none or only a few of them shortened at the base of the lamina, whereas laminae of most allied species are strongly and gradually reduced to often rudimentary auricles. Long, dark, spread- ing, unicellular trichomes are usually ample on petiole, rachis, and lamina tissue (although some- times broken away on dried specimens). With this probably should be included Polypodium acro- dontium Fee of Brazil. Further comparison of al- lied species can be found under G. xiphopteroides. Piura: Prov. Huancabamba, environs of Ayabaca, Sa- gdstegui & Cabanillas 8701 (F, MO, uc). La Libertad: Prov. Pataz, Puerta del Monte, Lopez & Sagdstegui 3452 (GH). San Martin: Dist. Huallaga, Valley of Rio Apison- cho, Hamilton & Holligan 708, 736 (K). Prov. Mariscal Caceres, Chochos, Young 2509 (USM), Young & Leon 4716 (USM). Huanuco: Tambo de Vaca, Bryan 629 (F, us). Macbride 4410 (F, us). Cuzco: Prov. Urubamba, Machu Picchu, Nunez & Arque 8339 (MO, uc), Peyton & Peyton 1103 (GH, MO). 44. Grammitis xiphopteroides (Liebm.) A. R. Smith, Amer. Fern J. 70: 26. 1980. Polypodium xiphopteroides Liebm., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Afd. ser. 5, 1: 196. 1849. LECTOTYPE (designated by A. R. Smith, Fl. Chiapas 2: 125. 1981): Mexico, Ve- racruz. "Nac. de Mirador," Liebmann PI. Mex. 2548 (c). Polypodium rigens Maxon, Proc. U.S. Natl. Mus. 27: 741. 1904. TYPE: Jamaica, John Crow Peak, Maxon 1346 (us). Ctenopteris rigens (Maxon) Copel., Philipp. J. Sci. 84: 422. 1956. Grammitis rigens (Maxon) Proctor, Brit. Fern Gaz. 9: 219. 1965. Plants epiphytic, rarely terrestrial, strongly ar- omatic. Stem 1.5-2 mm in diameter, short-creep- ing, provided with narrow-deltate to -lanceolate, thin, iridescent, gray-brown scales, these 2-5 mm long, clathrate, 8-12 cells across at base and the lumina large, clear and elongate, scale margins na- ked. Leaves 6-30 cm long, (0.7-)0.9-2.3 cm broad. Petiole (0.5-)5-8 cm long, amply provided with reddish brown, spreading, unicellular trichomes l-2(-3) mm long, but minute pluricellular ones rare or lacking. Lamina cut nearly or quite to ra- chis, chartaceous to subcoriaceous, linear or nar- row-lanceolate, gradually reduced at base to 6- many short pinnae or auricles, trichomes ample, 0.6-1 mm long on both sides of the rachis and longer on the segments, especially among the sori. Pinnae (or segments) 0.5-1 .2 cm long, 2.5-6 times longer than broad, patent or slightly ascending, adnate, mostly oblong, obtuse, subequilateral at base, either subtruncate or slightly dilated, mar- gins plane to somewhat revolute. Veins simple, 4- 10 pairs on a pinna, hydathodes evident. Sori 3— 7 pairs, sporangia glabrous. In wet forests, on trunks or branches of trees, rarely on wet clay banks, 800-2300(-2700) m, Ca- jamarca and Amazonas south to Cuzco. Mexico to Panama; Greater Antilles; Venezuela and Colombia to Brazil and Bolivia. This, G. pilosissima, and G. flabelliformis form a poorly understood species complex that is in great need of revision. Essentially the taxa are sep- arated here on the basis of some subtle and not TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 101 too consistent features: apex, shape, and orienta- tion of pinnae, degree of reduction of lamina base, and abundance or paucity of laminar trichomes. Most specimens can be separated by the key char- acters, but enough intermediates have been found to cast doubts as to whether there are one, two, or three discrete species. Grammitis xiphopteroides, as dilimited here, is basically a lower elevation epiphyte, occurring mostly at 2000 m and below. The other two are generally found at 2500-4000 m, but G. flabelliformis is an epipetric species of exposed habitats, whereas G. pilosissima is an epi- phyte in deep woods. Cajamarca: Colasay, Woytkowski 7016 (MO, us). Ama- zonas: Prov. Bagua, 12 km E of La Peca, Barbour 2397 (MO). Loreto: Sierra del Pongo, Mexia 6292a. Pasco: Prov. Oxapampa, Abra Los Mellizos, 4 km from Enenas, Skog et al. 5041 (us). Junin: Schunke Hacienda above San Ramon, Killip & Smith 24847 (NY, us). 24848 (NY). Ucayali (as Loreto): La Divisoria, between Tingo Maria and Pucallpa, Ferreyra 1689 (USM). Cuzco: Machu Pic- chu, Bishop 2513 (uc). 45. Grammitis flabelliformis (Poiret) Morton, Contr. U.S. Natl. Herb. 38: 57. 1967. Polypodium flabelliforme Poiret, in Lam., Encycl. 5: 519.1 804. LECTOTYPE (designated by Morton, 1967): Bourbon (Reunion), Commerson (P, Herb. Juss. 1098-C; photo, us, also Pichi-Sermolli pho- to in Webbia, 37: 121. 1983). Polypodium rigescens Willd. Sp. PI. ed. 4, pi. 5: 183. 1810. TYPE: Bourbon, Bory (holotype, B, Herb. Willd. 19668; isotype?, K, figured by Hooker & Grev. Icon. fil. t. 216). Polypodium peruvianum Desv., Mem. Soc. Linn. Paris 6: 231. 1827. TYPE: Peru, Dombey (holotype, P, Herb. Desv.; possible isotypes, L, P!; photo, us of L). Ctenopteris peruviana (Desv.) John Sm., Hist. fil. 1 84. 1875. Ctenopteris rigescens (Desv.) John Sm., Hist. fil. 184. 1875. Grammitis peruviana (Desv.) Morton, Contr. U.S. Natl. Herb. 38: 115. 1967. Grammitis rigescens (Willd.) Lell., Proc. Biol. Soc. Wash. 89: 383. 1985. Plants epipetric or terrestrial, often slightly ar- omatic. Stem 1-2.5 mm in diameter, short- or long-creeping, provided with narrow-deltate to -lanceolate, thin, iridescent, gray-brown scales, these 2.5-5 mm long, clathrate, 8-12 cells across at scale base and the lumina large, clear and elon- gate, margins naked. Leaves 5-20 cm long, 0.4- 1 (-1 .4) cm broad. Petiole 1-7 cm long, moderately provided with minute (0.1-0.2 mm), simple or branched, septate trichomes, but dark, spreading, unicellular ones sparse or lacking. Lamina cut nearly or quite to the rachis, chartaceous to sub- coriaceous, linear, gradually reduced at base to 3- 8 pairs of short and broad pinnae or auricles, cas- taneous, spreading, unicellular trichomes sparse or lacking on rachis, to 1.5 mm long on segments abaxially, especially among the sori. Pinnae 0. 1 5- 0.7 cm long, 1.5-2 times as long as broad, patent, adnate, deltate to oblong-deltate, obtuse or sub- acute, slightly and equally dilated or truncate at base, margins usually strongly revolute. Veins sim- ple, 2-4 pairs on a pinna, hydathodes evident. Sori (l-)2-4 pairs on a pinna, sporangia glabrous. On paramos, exposed slopes, high grasslands or open woods, on rocks, crevices of cliffs or on rocky ground, 2500-4800 m, Cajamarca south to Huan- cavelica and Puno. Costa Rica; Hispaniola; Venezuela and Colom- bia to Brazil and Bolivia; Africa. Long-standing confusion on the nomenclature and lectotypification of this and G. rigescens was resolved by Morton (Contr. U.S. Natl. Herb. 38: 57-59. 1967) and further substantiated by Bishop (Taxon 38: 91-95. 1989). However, Morton (loc. cit. 1 1 5) elected to separate G. peruviana from G. flabelliformis on the basis of the latter's sometimes longer stems and more distant leaves. Neither of these characters is significant nor consistent. This is easily confused with G. moniliformis, another species with stout, creeping stems, linear leaves, and broad, obtuse pinnae that are often nearly as broad as long. However, in G. flabelli- formis the lamina abaxially is amply provided with stout, castaneous trichomes, especially among the sori, and stem scales are narrow, with large, elon- gate lumina, these only 8-12 cells across at the base of the scale. Leaves of G. moniliformis are glabrous, and stem scales are broader, these with lumina commonly 1 6-30 across at scale base and most of the central ones nearly isodiametric. Grammitis flabelliformis is most closely related to G. pilosissima and G. xiphopteroides. See treat- ment of the latter for further discussion of the relationship. Cajamarca: Cerro Cumbe Mayo, Sanchez V. 34 (GH, us). La Libertad: Prov. Huamachuco, 5 km S of Por- tachuelo Pass, West 8147 (GH, uc, us). Ancash: NW slope of Nevada de Huascaran, below Llanganuco, Cor- rell & Smith P961 (GH, us). Prov. Yungay, Huascaran National Park, D. Smith & Goodwin 8896 (MO), 5597 (MO). Huanuco: Chavinillo, Coronado 177 (uc), 775 (uc), 506 (uc). Pasco: Huayllay, Cruckshanks (GH). Junin: San Jose, Macbride & Featherstone 1 1 10 (F, us). Huancav- elica: Prov. Tayacaja, Huacracocha, 20 km from Huan- 102 FIELDIANA: BOTANY cayo, Tovar 2184 (GH, USM). Apurimac: Abancay region, Sanlander (uc). Cuzco: Cerro de Colquipata, Pennell 13736 (F, GH, us). Puno: Prov. Sandia, between Sandia and Cuyocuyo, Ferreyra 16753 (GH). 46. Grammitis pseudonutans (Christ & Rosenst.) Morton, Contr. U.S. Natl. Herb. 38: 1 14. 1 967. Polypodium pseudonutans Christ & Rosenst., Repert. Spec. Nov. Regni Veg. 5: 15. 1908. TYPE: Ec- uador, Mt. Tungurahua. Rimbach (Rosenst. Exsicc. 39) (holotype, B?; isotype, us!). Ctenopteris pseudonutans (Christ & Rosenst.) Copel., Philipp. J. Sci. 84: 389. 1956. Plants epiphytic, not or scarcely aromatic. Stem 1.5-2.5 mm in diameter, short- or long-creeping, provided with ovate or ovate-lanceolate, thin, ir- idescent, gray-brown scales, these 5-8 mm long, clathrate, 20-30 cells across at scale base and most lumina small and central ones often nearly iso- diametric, scale margins naked. Leaves 18-36 cm long, 1.5-4.5 cm broad. Petiole 3-6 cm long, sparsely provided with minute (0. 1-0.2 mm) sim- ple or branched, pluricellular trichomes, long, spreading unicellular ones lacking. Lamina 1 -pin- nate, chartaceous, narrow-elliptic, gradually ta- pering at both ends, 5-10 proximal pinnae grad- ually reduced to mere auricles, rachis and lamina tissue with castaneous, unicellular trichomes to 1 mm long, glabrous adaxially. Pinnae 1-3 cm long, 4-8 times longer than broad, narrow- to oblong- deltate, patent or slightly ascending, adnate, acute or subacute, subequilaterally dilated at base, or occasionally subtruncate acroscopically, margins entire and plane. Veins simple, 6-12 pairs on a pinna, hydathodes evident. Sori 5-9 pairs on a pinna, sporangia glabrous. In forests or thickets, on tree trunks or branches, 3000-3425 m, San Martin. Colombia; Ecuador; Peru. Very similar to this is G. sodiroi (Christ & Ro- senst.) Morton of Ecuador, whose type was col- lected by Rimbach at the type location of G. pseu- donutans. The former differs in the abruptly reduced lamina, the completely glabrous rachis, and the immersed son. San Martin: Prov. Manscal Caceres, Chochos, Rio Abiseo National Park, Leon 2132. in part (uc). Prov. Mariscal Caceres, Mirador, Rio Abiseo National Park, Young 2154 (USM). 47. Grammitis andicola Stolze, sp. nov. Caulis 1-1.5 mm crassus, repens. Squamae cau- lis 4-6 mm longae, fuliginosac. clathratae, margine integra, luminibus 1 8-25 horizontaliter et pro par- tern maximam parvis et isodiametris. Folia ap- proximata vel distantia, 12-30 cm longa, 0.6-2.2 cm lata. Lamina 1 -pinnata vel pinnatisecta, basis gradatim reductis. Rachis glabra. Segmenta 0.4- 1.2 cm longa, oblonga vel oblonge deltata, obtusa vel subacuta. trichomatibus circa 1 mm longis, rigidis, castaneis. Venae simplices, hydathodis evidentibus. Sori 2-5(-7) jugi per segmento. Spo- rangia glabra. Plants terrestrial, rarely epiphytic or epipetric, somewhat aromatic. Stem 1-1.5 mm in diam- eter, long-creeping, provided with ovate or broadly lanceolate, thin, iridescent, gray-brown scales, these 4-6 mm long, clathrate, 18-25 cells across at scale base and at least the lumina of central cells small and nearly isodiametric, scale margins naked. Leaves approximate to well spaced, 12-30 cm long, 0.6-2.2 cm broad. Pet- iole 3-9 cm long, scarcely to moderately pro- vided with orange to red-brown, spreading, uni- cellular trichomes to 1 mm long, moderately to amply provided with simple to 1 -forked pluri- cellular trichomes 0. 1-0.2 mm long. Lamina cut nearly or quite to the rachis, chartaceous to sub- coriaceous, linear or narrow-elliptic, tapering at both ends, 5-many proximal pinnae gradually reduced to mere auricles, rachis glabrous, seg- ments amply provided with rigid, castaneous trichomes to 1 mm long, especially among the sori. Pinnae 0.4-1 .2 cm long, 1.5-2 times as long as broad, patent, adnate, oblong or oblong-del- tate, obtuse or subacute, slightly dilated and subequilateral at base, the margins entire and strongly re volute. Veins simple, 3-8(-10) pairs on a pinna, hydathodes evident. Sori 2-5 (-7) pairs on a pinna, sporangia glabrous. TYPE— Peru, Amazonas, Prov. Chachapoyas, Cerros Calla Calla, Hutchison & Wright 5826 (ho- lotype, F!; isotypes, GH!, uc!). In forests or thickets, on the forest floor, or rarely on rock cliffs or low on tree trunks, 2500-4250 m, along the Cordillera Central from Amazonas to Cuzco. Thus far known from Peru and one specimen from Colombia: Cauca, Parque Nacional de Pu- race, Lozano et al. 4464 (F). This is very similar to G. moniliformis and G. flabelliformis in the linear leaves cut nearly or quite to the rachis into many short and broad, obtuse TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 103 to subacute pinnae, many of which are as broad as long. Leaves of G. moniliformis are approxi- mate to subfasciculate and glabrous, whereas those of G. andicola are usually well spaced, and seg- ments and often the petiole are provided with rig- id, castaneous, unicellular trichomes. From G.fla- belliformis, G. andicola differs especially in the broader stem scales, cells with lumina about twice as many across at scale base, and those at the middle of the scale mostly isodiametric. Scales of G. flabelliformis are larger and mostly elongate throughout. In G. andicola, pinnae are usually lon- ger (mostly over 5 mm long) and petioles thicker (mostly over 0.6 mm). Pinnae of G. flabelliformis are commonly less than 5 mm long and petioles less than 0.6 mm thick. Amazonas: Prov. Bagua, Cordillera Colan SE of La Peca (aberrant form), Barbour 3556 (MO, uc). Prov. Cha- chapoyas, Cerros Calla Calla, Hutchison 4754 (GH, uc, in part). Prov. Chachapoyas, slopes of Puma-Urcu ESE of Chachapoyas, Wurdack 702 (us). San Martin: Dist. Huallaga, Valley of Rio Apisoncho, Hamilton & Holli- gan 1223 (us). Prov. Mariscal Caceres, NW corner of Rio Abiseo National Park, Young & Leon 4423 (USM), 4826 (USM). Prov. Mariscal Caceres, Rio Abiseo Na- tional Park, Puerta del Monte, Young 1983 (HUT). Pasco: Prov. Oxapampa, Santa Barbara, D. Smith 81 17 (F, MO). Cuzco: Cabecera del Rio Korebeni, Biies 1948 (us), 7955 (us). Prov. La Convention, Huayopata, Peyton & Peyton 974 (GH, MO). 48. Grammitis asplenifolia (L.) Proctor, Brit. Fern Gaz. 9: 76. 1962. Polypodium asplenifolium L., Sp. pi. 2: 1084. 1753. TYPE: Petiver, Pter. Amer., t. 7,f. 16, from Plu- mier, Traite foug. Amer., t. 102a, based on a specimen from Martinique. Polypodium dolorense Hieron., Bot. Jahrb., Syst. 34: 512. 1904. TYPE: Colombia, Prov. Antioquia, Rio Dolores, Lehmann 7380 (holotype, B?; iso- type, us!). Ctenopteris asplenifolia (L.) Copel., Gen. fil. 2 1 9. 1 947. Ctenopterisdolorensis (Hieron.) Copel., Philipp. J. Sci. 84:449. 1956. Plants epiphytic. Stem short-creeping or decum- bent, provided with abundant, lustrous, orange scales, these not or scarcely clathrate, 1-3 mm long, the margins with whitish or hyaline setae. Leaves deeply pinnatisect, 25-60 cm long, 3-7 cm broad. Petiole 10-30 cm long, 0.4-1.2 mm in di- ameter, sparsely to abundantly provided with se- riceous, spreading, unicellular trichomes, these 1- 3 mm long, tawny to orange. Lamina thin-her- baceous, lanceolate, cut nearly to the rachis, grad- ually reduced to apex, scarcely reduced at base (or only 1-2 basal pairs of segments strongly reduced), rachis and tissue amply provided with orange or reddish brown, spreading trichomes. Segments 1- 4 cm long, patent, narrow-deltate, acute, dilated subequally at base, or more strongly decurrent ba- siscopically, the margins entire and with simple, spreading trichomes. Veins simple, but fertile ones with a rudimentary spur on which the sorus is borne, hydathodes dark brown, small but usually distinct. Son inframedial, sporangia with long, pale setae. On tree trunks in rain forests, 1500-2400 m, Huanuco, Pasco, and Junin. Mexico and Central America; West Indies; Trinidad; Venezuela; Colombia to Bolivia. This differs from G. lehmanniana (Hieron.) Morton, of Central America and northern South America, in a few subtle characters: the petiole is shorter, the stem scales have longer setae, and the sporangia have 2-3 (vs. 1) setae. Grammitis chrysleri (Copel.) Proctor, of Jamai- ca, probably should be included with G. aspleni- folia. The former has been separated on the basis of its larger leaves with forked veins. Veins of G. asplenifolia are indeed forked, but the fork is re- duced to a short spur on which the sorus is borne. At best, G. chrysleri should be recognized only as a variety. Huanuco: Prov. Huanuco, Mirador, on road from Acomayo to Chinchao, Mexia 4141, in part (uc). Pasco: Prov. Oxapampa, Oxapampa, van der Werffet al. 8608 (MO, uc). Junin: Chanchamayo Valley, C. Schunke 23 (F, us). Prov. Tarma, Agua Dulce, Woytkowski 35485 (uc). 49. Grammitis dependens (Baker) Morton, Contr. U.S. Natl. Herb. 38: 104. 1967. Polypodium dependens Baker, in Hooker & Baker, Syn. fil. 335. 1867. TYPE: Ecuador, Mt. Pichin- cha, Spruce 5637 (as 563) (holotype, K!; photos, F, us). Plants epiphytic. Stem small, suberect, sparsely provided with minute, deltate, sublustrous, cas- taneous scales, these 1 mm long, nonclathrate, the margins with rigid, whitish setae. Leaves 12-30 cm long, 1-1.5 cm broad. Petiole 1 cm long or less, 0.2-0.4 mm in diameter, amply provided with spreading, yellowish, unicellular trichomes to 1.5 mm long. Lamina thin-herbaceous, linear, 1 -pinnate, strongly and gradually reduced to apex and base, pilose as on the petiole, but also inter- 104 FIELDIANA: BOTANY mixed with stouter, castaneous, sessile-stellate tri- chomes. Pinnae 0.4-1 cm long, oblong-elliptic, ob- tuse at apex, somewhat narrowed at base, the margins entire and the trichomes mostly in fas- cicles of 2 or 3. Veins 1-2 pairs in each pinna, hydathodes small, brown, sometimes obscured by the indument. Sori several to a pinna, sporangia each with 1 long seta. On tree trunks in wet forests, 3000-3100 m, Cuzco. Colombia to Peru. Copeland (1956) combined this with G. hetero- morpha, suggesting that it is merely an unbranched form of the latter. The supposition may be correct, for the two taxa are remarkably similar in such critical features as stem scales and lamina indu- ment. However, pending monograhic revision, G. dependens is separated here by the unbranched leaves and the oblong-elliptic (vs. obovate) pinnae. Cuzco: Michihuanunca, Bites 731 (us). Huadquina, Bites 975 (us). 50. Grammitis heteromorpha (Hooker & Grev.) Morton, Contr. U.S. Natl. Herb. 38: 102. 1967. Polypodium heteromorphum Hooker & Grev., Icon, fil. 1, t. 108. 1829. TYPE: Ecuador, Prov. Pi- chincha, between Mt. Chimborazo and Pichin- cha, Jameson (holotype, K.!; isotype.n; photos, F of K, us of FI). Ctenopteris heteromorpha (Hooker & Grev.) Copel., Philipp. J. Sci. 84:412. 1956. Plants terrestrial (in Peru), epiphytic, or epipet- ric. Stem small, short-creeping or decumbent, pro- vided with ovate or deltate, nonclathrate scales, these 0.5-1.5 mm long, lustrous, castaneous, the margins with rigid, whitish setae. Leaves lax, swol- len and articulate at the stem, crowded to subfas- ciculate, 1 -pinnate and 1-2-forked, to 60 cm long and 8 cm broad. Petiole 1-5 cm long, 0.2-0.4 mm in diameter, amply provided with thin, spreading trichomes, these tawny to orange, 1-2 mm long and often intermixed with a few pluricellular, sim- ple or branched trichomes about 0.2 mm long. Lamina thin- to firm-herbaceous, indeterminate, reduced at base. Rachis brown to blackish, indu- ment as on the petiole, except occasionally some long, castaneous, stellate trichomes intermixed with the simple, light-colored ones. Pinnae sessile, 4— 7 mm long, 2—4 mm broad, obovate or spatulate, amply pilose as on the rachis. Veins distinct to obscure, pinnately branched in each pinna and the branches terminating in hydathodes adaxially, the latter (outside Peru) sometimes covered by cal- careous deposits. Sori round, 3-5 per segment, sporangia (in Peru) glabrous. On shaded, rocky slopes, among boulders (but often epiphytic outside Peru), 4400-4500 m, An- cash. Mexico; Guatemala; Colombia to Bolivia. This is a variable species that may include some distinct forms or varieties. Plants in Mexico and Guatemala are thinner in texture and sometimes bear white deposits on the hydathodes. Some spec- imens from Colombia have larger, oblong pinnae nearly 1 cm long, with white hydathodes and se- tose sporangia. Copeland ( 1956) included with this G. dependens and G. variabilis; see treatment of the latter for detailed discussion of taxonomy and typification. The entire complex needs mono- graphic revision. Ancash: Prov. Huaraz, Huascaran National Park, Quebrada Llaca, D. Smith et al. 8987 (F, MO). Prov. Huaraz, Huascaran National Park, Quebrada Ishinca, D. Smith et al. 9544 (F, HUT, MO). 51. Grammitis lanigera (Desv.) Morton, Contr. U.S. Natl. Herb. 38: 105. 1967. Plants epiphytic, rarely terrestrial in var. lani- gera. Stem suberect, provided with ligulate to nar- row-deltate, lustrous, castaneous scales, these 1-3 mm long, nonclathrate, the margins with rigid or- ange to hyaline setae. Leaves indeterminate, pen- dulous, 25-70 cm long, 2.5-8 cm broad. Petiole 0.5-4 cm long, 0.4-1 mm in diameter, copiously provided with spreading, sericeous, yellowish or hyaline trichomes to 2 mm long. Lamina firm- herbaceous to chartaceous, narrow-elliptic, 1 -pin- nate, strongly reduced at apex and base, densely and simply pilose as on the petiole (often nearly obscuring the lamina surface), but also with abun- dant sessile-stellate trichomes, these (in var. Stella) stouter and castaneous. Pinnae commonly 20-50 pairs, 4-10 pairs along 5 cm of rachis, 1.1-4.5 cm long, 0.4-1.2 cm broad, patent to slightly ascend- ing, broadly to narrowly deltate, apex acute or subacute, subequilateral at base, there truncate to slightly dilated acroscopically, rather strongly di- lated basiscopically, the margins entire and with trichomes simple to sessile-stellate. Veins obscure, simple, hydathodes brown or occasionally with circular white deposits, often completely obscured by the dense indument. Sori 6-12 pairs on a pinna, sporangia with 1-3 long setae. TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 105 The species occurs in Costa Rica, Hispaniola, the Lesser Antilles, and from Colombia to Bolivia. This is part of the confusing species complex involving G. cultrata (Willd.) Proctor, G. depen- dens, G. heteromorpha, G. laxa, G. subflabellifor- mis, and probably a few more taxa outside Peru. Diagnostic features of many of these are highly variable and species lines are not clearly drawn. The entire group is in need of monographic re- vision. From the other species in the complex, G. lanigera can best be distinguished by the type and density of lamina trichomes. All the species have long, pale, spreading, sericeous trichomes, and most have these on the margins as well as some in fas- cicles of 2-4. Other species have a few fascicled trichomes along the rachis with up to 6-7 arms (in the keys and descriptions these are called "ses- sile-stellate"). However, G. lanigera is the only species that has these sessile-stellate trichomes in such dense masses as to often obscure the abaxial laminar surface. It is separated here into two va- rieties. Key to Varieties a. Sessile-stellate trichomes on lamina sericeous and yellowish to hyaline (occasionally pale orange) . 5 la. var. lanigera a. Sessile-stellate trichomes on rachis (and often the laminar surface) rigid and castaneous . 51b. var. stella 5 la. Grammitis lanigera var. lanigera. Polypodium lanigerum Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck, Gesammten Na- turk. 5: 316. 1811. TYPE: Peru, Dombey, Herb. Jussieu 1099 (holotype, p; isotype, P!; photos, F, GH, us). Polypodium sericeolanatum Hooker, Sp. fil. 4: 221. 1864. LECTOTYPE (designated by Morton, Contr. U.S. Natl. Herb. 38: 105. 1967): Ecuador, Ravines of Pichincha, Jameson 235 (K; frag., us). Ctenopteris lanigera (Desv.) Copel., Philipp. J. Sci. 84: 420. 1956. Ctenopteris sericeolanata (Hooker) Copel., Philipp. J. Sci. 84: 453. 1956. Grammitis sericeolanata (Hooker) Proctor, Rhodora 63: 35. 1961. In wet forests, pendent from trunks and branch- es of trees, rarely on wet, moss-covered banks or boulders, 2200-3900 m, Central Peru, from Ama- zonas to Cuzco. Hispaniola; Martinique; Costa Rica to Bolivia; possibly Venezuela. Amazonas: Prov. Chachapoyas, Calla Calla slopes near km 4 1 5-4 1 8 of Leimebamba-Balsas road, Wurdack 1 742 (F, GH, uc, us, USM). San Martin: Prov. Mariscal Caceres, NW corner of Rio Abiseo National Park, Young & Leon 4604 (USM). Huanuco: Muna, trail to Tambo de Vaca, Macbride 4303 (F, GH, us). Pasco: Prov. Oxapampa, San Alberto, Cordillera de Yanachaga, van der Werff et al. 8464 (MO, us). Cuzco: Montana de Calca, Valle de Lares, Bites 1930 (us). 5 Ib. Grammitis lanigera var. stella (Copel.) Mor- ton, Contr. U.S. Natl. Herb. 38: 106. 1967. Ctenopteris stella (Copel, Philipp. J. Sci. 84: 452. 1956. TYPE: Peru, Cuzco, Valley of Rio Urubamba, Biies A3 2 (holotype, us!). In wet forests and wooded ravines, ( 1 600-)2400- 3740 m, Central Peru, Amazonas to Cuzco. Colombia to Bolivia. The distinctive feature of rigid, castaneous, ses- sile-stellate trichomes on the lamina is shared by only one other species in the G. cultrata complex, G. dependens. Variety stella is normally epiphytic at higher elevations in Peru, usually above 2500 m. However, the Woytkowski specimen cited from Amazonas was found at 1 600 m on a rocky slope. The lamina pubescence is especially dense, the pinnae are large (to 4 cm long and 1 .2 cm broad) and very crowded, and the hydathodes are white- encrusted. (These calcareous deposits are infre- quent in the species.) This is certainly an aberrant specimen, and perhaps its unusual features are merely products of the different elevation and hab- itat. Amazonas: Mendoza, Woytkowski 8254 (MO). San Martin: Prov. Mariscal Caceres, Rio Abiseo National Park, Leon 1908 (uc), Young & Leon 4723 (USM), Young & Leon 4816 (USM). Huanuco: Huanuco-Tingo Maria road near Carpish Divide, Gentry & Smith 44857, in part (MO). Pasco: Prov. Oxapampa, Rio San Alberto Val- ley E of Oxapampa, D. Smith & Pretel 7976 (F, MO, USM). Cuzco: Huadquina, Biies 967 (us), 7007 (us). 52. Grammitis subflabelliformis (Rosenst.) Mor- ton, Contr. U.S. Natl. Herb. 38: 104. 1967. Polypodium subflabelliforme Rosenst., Repert. Spec. 106 FIELDIANA: BOTANY Nov. Regni Veg. 7: 306. 1909. TYPE: Ecuador, Mt. Abitagua, Spruce 5271 (holotype, K.; isotypes, NY!, us). Polypodium subflabelliforme var. minor Rosenst., Re- pert. Spec. Nov. Regni Veg. 7: 307. 1909. TYPE: Ecuador, Mt. Tungurahua, Spruce 5272 (holo- type, K). Polypodium senile var. minor Rosenst., Meded. Rijks- Herb. 19: 19. 1913. TYPE: Bolivia, Yungas de San Mateo, Herzog 1985 (holotype, L; isotype, us!). Ctenopteris subflabelliformis (Rosenst.) Copel., Phil- ipp. J. Sci. 84: 400. 1956. Plants epiphytic, epipetric or terrestrial. Stem small, suberect, sparsely provided with minute, ligulate to oblong, sublustrous, deep orange scales, these 0.5-0.8 mm long, nonclathrate, the margins with whitish or hyaline setae. Leaves 7-20 cm long, 0.7-1.2 cm broad. Petiole to 1 cm long, 0.1-0.2 mm in diameter, provided with spreading, whitish or yellowish, unicellular trichomes to 1.5 mm long. Lamina thin-herbaceous, linear, 1 -pinnate, strongly reduced at apex and base, thin-pilose as on the petiole. Pinnae 15-35 pairs, 3-6 mm long, 2-3 mm broad, obovate or (occasionally) oblong, ob- tuse at apex, rounded acroscopically to a very nar- row base, often strongly decurrent basiscopically, the margins entire (or slightly gibbous acroscopi- cally) the long, marginal trichomes simple or paired from the base. Veins evident, simple to pinnate with 2 pairs of veinlets, hydathodes evident. Sori several to a pinna, sporangia with 2-3 long setae. In wet forests, on tree trunks, humus, or rocky cliffs, 1800-2800 m, Amazonas, Huanuco, Pasco, and Cuzco. Panama; Ecuador to Bolivia. Perhaps this should be combined with G. senilis (Fee) Morton var. senilis, which supposedly differs only in its slightly longer, more oblong pinnae. (The type of the latter was not located for this study, so a comparison could not be made.) Based on specimens examined, the distribution in Peru (as well as through the entire range) ap- pears to be rather disjunct. However, since this is a very inconspicuous fern, it can be easily over- looked, and it is likely future collecting efforts will prove it to be much more widely distributed. Amazonas: Prov. Bongara, road to La Rioja, 5 km N of Lake Pomacocha, Hutchison & Wright 6805 (uc). Huanuco: Vilcabamba, hacienda on Rio Chinchao, Bry- an 730 (F). Pasco: Prov. Oxapampa, Dist. Oxapampa, from Oxapampa to Villa Rica, Ledn 661 (USM). Cuzco: Valle de Lares, above Rio Lachac, Sues 1812 (us). Prov. La Convention, 1 0 km SW of Incatambo, Peyton & King 1433 (MO). Prov. Urubamba, Winawaina, Vargas 22373 (GH). 53. Grammitis laxa (Presl) Morton, Contr. U.S. Natl. Herb. 38: 105. 1967. Polypodium laxum Presl, Reliq. haenk. 1 : 23, /. 4, f. 1. 1825. TYPE: "In vallibus cordillerum Peru- viae," Haenke (holotype, PRO!; isotype, PR!). Ctenopteris contacta Copel., Philipp. J. Sci. 84: 447. 1956. TYPE: Bolivia, Yungas, Bang 483 (holo- type, us!; isotype, F!). Plants epiphytic or occasionally terrestrial. Stem suberect, sparsely provided with ligulate to nar- row-deltate, sublustrous, deep orange to casta- neous scales, these 0.5-1 .5 mm long, nonclathrate, the margins with rigid, whitish or hyaline setae. Leaves 14-40 cm long, 1-4 cm broad. Petiole 1 cm long or less, 0.2-0.3 mm in diameter, provided with spreading, sericeous, whitish or hyaline, uni- cellular trichomes to 2 mm long. Lamina thin-to firm -herbaceous, narrow-elliptic, 1 -pinnate, pec- tinate, strongly reduced at apex and base, sparsely to moderately sericeous-pilose as on the petiole. Pinnae patent, commonly 60-80 pairs, very crowded (12-20 pairs along 5 cm of rachis), 1-2.5 cm long, 2-3 mm broad, linear, obtuse at apex, broadly adnate and subequilateral at base, there truncate to slightly rounded acroscopically, trun- cate to slightly dilated basiscopically, the margins entire and subparallel most of their length, the long, marginal trichomes simple, paired or in fas- cicles of 3. Veins indistinct or obscure, simple, hydathodes evident. Sori 6-12 pairs on a pinna, sporangia with 1-3 long setae. In wet forests, on tree trunks or moist, shaded bluffs, 1500-2400 m, Amazonas, Huanuco, and Cuzco. Ecuador to Bolivia. It is interesting that Copeland (1956, pp. 420- 421) combined G. laxa with G. lanigera, and in so doing he cited Bang 483 from Bolivia as a typ- ical example. Yet on page 427 he cited this same collection as type of his new Ctenopteris contacta (= G. laxa), placed in another group. This is typical of the confusion in the complex of Grammitis spe- cies with sericeous trichomes and setose sporangia, because the other characters separating these spe- cies are rather subtle. For further discussion see treatment of G. lanigera. TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 107 Amazonas: Prov. Bagua, Cordillera Colan SE of La Peca, Harbour 3601 (F, MO, USM). Huanuco: Pampayacu, Kanehira 122 (GH, us). Cuzco: Trail from Machu Picchu ruins to Inca Bridge, Bishop 2509, 2510 (both uc). Prov. La Convention, Valle San Miguel, Sues 2130 (us). De- partment unknown: Poeppig 170 (p), Poeppig in 1829 (MO). 54. Grammitis cultrata (Willd.) Proctor, Rhodora 63:35. 1961. Polypodium cultratum Willd., Sp. pi. ed. 4, 5: 187. 1810. TYPE: Jamaica, Swartz (holotype, B, Herb. Willd. 19674; photos, GH, us). Ctenopteris cultrata (Willd.) Copel., Gen. fil. 2 1 9. 1 947. Plants epiphytic. Stem erect, sparsely provided with ligulate to narrow-deltate, sublustrous, cas- taneous or deep orange scales, these 0.5-2 mm long, nonclathrate, the margins with rigid, orange or hyaline setae. Leaves pendulous, 30-50 cm long, 1.5-5 cm broad. Petiole 0.5-3 cm long, 0.2-0.3 mm in diameter, amply provided with spreading, sericeous, yellowish or hyaline trichomes to 2 mm long. Lamina thin- to firm-herbaceous, narrow- elliptic, 1 -pinnate, strongly reduced at apex and base, sparsely to moderately pilose as on the pet- iole, and with scattered sessile-stellate ones on the rachis. Pinnae commonly 20-50 pairs, 5-10 pairs along 5 cm of rachis, l-2(-2.5) cm long, (0.3-)0.4- 1.2 cm broad, patent or often somewhat deflexed, broadly to narrowly deltate, apex obtuse to acute, mostly subequilateral at base, there slightly round- ed or dilated acroscopically, usually obviously di- lated basiscopically, the margins entire and with trichomes simple, paired, or in fascicles of 3. Veins indistinct or obscure, simple, hydathodes evident. Sori 4-8 pairs on a pinna, sporangia with 1-3 long setae. flexed, with bases subequilateral, i.e., slightly rounded or dilated on each side. In Central Amer- ica and northern South America, both shapes seem to be rather equally common. The most extreme form is represented by the type (K) of Polypodium alternifolium Hooker (= P. longum C. Chr.) from Ecuador, with acute, broadly deltate, subequilater- al pinnae to 3 cm long and 1 cm broad. This spec- imen resembles larger specimens of G. lanigera and seems to be intermediate between the two species. The entire species group of G. cultrata is greatly in need of revision. For further discussion of prob- lems in this complex, along with a list of some of the other species, see treatment of G. lanigera. Amazonas: Prov. Chachapoyas, Cerros Calla Calla, 19 km above Leimebamba, Hutchison & Wright 5582 (GH, uc). San Martin: Prov. Rioja, Venceremos, D. Smith 4472 (F, MO, uc). Huanuco: Prov. Huanuco, Mirador, road from Acomayo to Chinchao, Mexia 4141 (GH, MO, uc in part). Junin: Carpapata, above Huacapistana, Kil- lip & Smith 24458 (F, NY, us). Pichis Trail, San Nicolas, Killip & Smith 25988 (F, NY, us). Pasco: Prov. Oxapam- pa, surroundings of Oxapampa, van der Werffet al. 8349 (MO, uc). Cuzco: Cerro Chuyapi, Bues A43 (us), A47, (us). Puno: Near San Gaban (as Gavan), Lechler 2166 (GH). 55. Grammitis blepharolepis (C. Chr.) Morton, Contr. U.S. Natl. Herb. 38: 98. 1967. Polypodium gracillimum Hieron., Hedwigia 48: 250, t. 12, f. 18. 1909, not Copel. 1905. TYPE: Ec- uador, Quito to Mindo, Stiibel 747 (holotype, B). Polypodium blepharolepis C. Chr., Index fil. suppl. 1: 58. 1913, nom. nov. for P. gracillimum Hieron. and with the same type. Xiphopteris blepharolepis (C. Chr.) Copel., Amer. Fern J. 42: 109. 1952. In dense forests or thickets, on trunks or high branches of trees, 1100-3100 m, in central Peru from Amazonas to Puno. Southern Mexico to Panama; Greater Antilles, Colombia and Venezuela to Bolivia and Brazil. Pinna shape varies widely in this species, and monographic study may reveal that there are sev- eral varieties or forms. In the Greater Antilles pin- nae are patent, small, obtuse, nearly oblong, with a somewhat equilateral base, i.e., rounded acro- scopically and dilated basiscopically (approaching pinna shape of G. subflabelliformis). Most speci- mens from Peru and Bolivia have deltate, acute or subacute pinnae, many of them somewhat re- Plants epiphytic. Stem small, suberect, provided with lanceolate or ovate, lustrous, deep orange or reddish brown scales, these 1-2 mm long, sub- clathrate, the margins with long, rigid, concolorous setae. Leaves 5-15 cm long, 0.4-0.7 cm broad. Petiole 0.5-1.5 cm long, 0.3-0.4 mm in diameter, abundantly provided with spreading, rigid, red- dish brown, unicellular trichomes about 1 mm long. Lamina chartaceous, linear, divided nearly or quite to the rachis into numerous, crowded seg- ments, abruptly reduced at apex and base, pilose as on the petiole. Segments 2-3 mm long, oblong or oblong-elliptic, the apex obtuse, the base trun- cate or somewhat dilated, the margins entire, often gibbous near the acroscopic base. Veins obscure, 108 FIELDIANA: BOTANY simple in each segment, or with one short acro- scopic branch, hydathodes evident. Sori solitary and at or near the base in each segment, sporangia glabrous. In forests on tree trunks, 300-1400 m, Pasco, Junin, and Puno. Venezuela; Colombia to Peru. With this probably should be included Gram- mitis basalts (Morton) Lell. (Ecuador) and G. da- guensis (Hieron.) Morton (Colombia). The former seems to be identical. The latter was distinguished by Morton (1967) solely on the basis of its non- gibbous segments; however, an isotype (B!) has gibbous and nongibbous segments alike, and no other different features were noted. Pasco: Prov. Oxapampa, Pichis Valley, San Matias Ridge, Foster 8616 (F, MO), Leon 315 (USM). Junin: Schunke Hacienda above San Ramon, Killip & Smith 24648 (us), C. Schunke Al 57 (us). La Merced, Hacienda Schunke, Macbride 5639 (F, us). Puno: La Pampa, Rio Tavara, Watkins 1916 (us). 56. Grammitis truncicola (Klotzsch) Morton, Contr. U.S. Natl. Herb. 38: 98. 1967. Polypodium truncicola Klotzsch, Linnaea 20: 374. 1847. TYPE: Venezuela, Colonia Tovar, Moritz 252 (isotype, us). Polypodium andinum Hooker, Sec. cent, ferns, /. 6. 1847. LECTOTYPE (designated by Morton, Contr. U.S. Natl. Herb. 38: 98. 1967): Ecuador, Banks of Rio Hondacha, Jameson 780 (K). Grammitis andina (Hooker) Ching, Bull. Fan. Mem. Inst. Biol., Bot. 10: 240. 1941. Xiphopteris truncicola (Klotzsch) Copel., Amer. Fem J. 42: 101. 1952. Plants epiphytic. Stem small, suberect, provided with lanceolate or ovate, lustrous, deep orange or reddish brown scales, these 1-2 mm long, sub- clathrate, the margins with rigid, concolorous se- tae. Leaves 5-10 cm long, 0.6-1 cm broad. Petiole obsolete or nearly so, abundantly provided with spreading, rigid, reddish brown, unicellular tri- chomes to 1 mm long. Lamina thin- to firm-her- baceous, linear to narrow-elliptic, pinnatisect, re- duced at apex and base, pilose as on the petiole. Segments 2-3 mm long, broadly oblong or oblong- elliptic, the apex obtuse, the margins entire. Veins obscure, simple in each segment, or with a short, acroscopic branch, hydathodes evident. Sori sol- itary and borne at or near the base in each segment, sporangia glabrous. Thus far known in Peru only from San Martin. The collection represents one of the syntypes of Polypodium andinum. Costa Rica; Venezuela; Colombia to Peru. This scarcely differs from G. blepharolepis. Monographic study is needed. San Martin: Mount Picote, near Moyobamba, Nelson (Spruce 4780) (K). 57. Grammitis david-smithii Stolze, sp. nov. Fig- ures 5d-f. Caulis erectus, paleis 2—4 mm longis, lineari- deltatis, attenuatis, fuliginosis, clathratis, setosis. Folia 12-35 cm longa, 2-5 cm lata, pectinata, 1-pinnata vel pinnatisecta. Petiolus 1-8 cm lon- gus, (0.4-)0.5-1 .2 mm latus, trichomatibus sparsis vel abundantibus, 0.4-0.9 mm longis, unicellulari- bus, aurantiacis vel rubiginosis. Segmenta 0.9-2.8 cm longa, 0.2-0.4 cm lata, linearia vel lineari-del- tata, glabra vel hirsuta. Venae simplices, 6-14 jugae. Hydathodi brunneae. Plants epiphytic, rarely epipetric. Stem erect, provided with linear-deltate, attenuate, iridescent gray-brown scales, these 2-4 mm long, essentially clathrate, the margins with rigid orange or reddish brown setae. Leaves 1 2-35 cm long, 2-5 cm broad. Petiole 1-8 cm long, (0.4-)0.5-1.2 mm in diam- eter, sparsely to amply provided with rigid, orange or reddish brown trichomes 0.4-0.9 mm long. Lamina chartaceous, elliptic, pectinate, divided nearly or quite to the rachis, apex gradually re- duced, the base abruptly reduced, the rachis amply hirsute like the petiole, but trichomes shorter (0.3- 0.5 mm), surface glabrous adaxially, moderately hirsute or glabrescent abaxially. Segments 0.9-2.8 cm long, 0.2-0.4 cm broad beyond the dilated base, linear or linear-deltate, subacute, the mar- gins entire, usually host to clavate, black fungi (Ascomycetes) on the abaxial surface. Veins sim- ple, distinct or indistinct, 6-14 pairs in each seg- ment, hydathodes distinct but lacking white en- crustations. Sori medial to supramedial, 6-12 pairs on a segment, sporangia glabrous. TYPE— Peru, Prov. Oxapampa, 2—4 km N of Mallampampa, D. Smith & Canne 5837 (hololype, F!; isotype, MO!). In forests, on tree trunks and branches, rarely on mossy boulders, 1800-2800 m, Huanuco, Pas- co, Junin, and Cuzco. TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 109 Peru and Bolivia. This is another of the species with setose scales very closely related to G. semihirsuta (under which may be seen further discussion), but the only one in Peru with scales iridescent, gray-brown, and clathrate; i.e., the cell walls are relatively thin, with lumina transparent (although often quite narrow). Scales of the other species in the complex are black, orange, or reddish brown and have thick, crowded cell walls and the lumina occluded. The species is named in honor of the late David N. Smith, a friend and indefatigable collector of Andean plants who had an especially keen eye for ferns. His excellent collections are well represented throughout this flora. Huanuco: Huanuco-Tingo Maria road near Carpish Divide, Gentry & Smith 44857, in part (MO). Pasco: Prov. Oxapampa, San Alberto, Cordillera de Yanachaga, van der Werff et al. 8451 (MO, uc), 8452 (MO, uc). Junin: Prov. Chanchamayo, Rio Rondayacu, 45 km from San Ramon, D. Smith et al. 2607 (MO). Cuzco: Machu Picchu, trail to Winay Waina, Bishop 2506 (uc). Huadquina, Biies 1252 (us). Valle de Lares, Bues 1929 (us). Additional paratypes follow. Ecuador: Prov. Pichincha, Paramo de Guamani, road from Pifo to Papallacta, just west of the pass, 4000 m, Stolze & Stolze 1663 (F). Bolivia: Cochabamba, Prov. Chapare, Cochabamba, Beck 1424a (F). La Paz: Prov. Nor Yungas, Chuspipata, Beck 7600 (F). 58. Grammitis pichinchensis (Hieron.) Morton, Contr. U.S. Natl. Herb. 38: 111. 1967. Polypodium subscabrum sensu Hooker, Sp. fil. 4: 183, /. 274A. 1864 (based on Moritz 332, K), not Klotzsch, 1 847 (based on Moritz 332, B). Polypodium pichinchense Hieron., Bot. Jahrb. Syst. 34: 506. 1904. LECTOTYPE (designated by Morton, 1 967): Ecuador, western side of Pichin- cha, Jameson in 1862 (B!, photo, F; probable iso- lectotypes, B!, BM!, us). Polypodium ecuadorense C. Chr., Index fil. 524. 1906. Nom. superfl., an illegitimate renaming of P. pi- chinchense Hieron. 1904. Ctenopteris ecuadorensis (C. Chr.) Copel., Philipp. J. Sci. 84: 434. 1956. Plants epiphytic, rarely (in Ecuador) epipetric. Stem suberect, provided with lanceolate or nar- row-deltate, attenuate, lustrous, blackish scales, these 1-3 mm long, nonclathrate, the margins with dark, rigid setae. Leaves 10-20 cm long, 0.7-2 cm broad. Petiole 1-4 cm long, 0.3-0.6 mm in di- ameter, abundantly provided with spreading, rig- id, castaneous, unicellular trichomes 1-2 mm long. Lamina firm-herbaceous to chartaceous, narrow- elliptic, deeply pinnatisect nearly to rachis, grad- ually reduced to apex and base, rachis and at least the abaxial surface sparsely to moderately pilose as on the petiole. Segments 0.3-1 cm long, 0.15- 0.25 cm broad at base, deltate or oblong-deltate, obtuse or subacute, the margins entire, usually host to clavate, black fungi (Ascomycetes) on the ab- axial surface. Veins simple, indistinct or obscure, 3-4 pairs in each segment, hydathodes distinct and each usually encrusted with a calcareous deposit. Sori medial to inframedial, 2-4 pairs per segment, sporangia glabrous. In forests on tree trunks, rarely (in Ecuador) in rock crevices, known thus far in Peru by a single specimen, 2600 m, Huanuco. Ecuador; Peru. Nomenclature has been badly confused and ap- plication of the name incorrectly applied because of a spurious type collection of Polypodium sub- scabrum Klotzsch, Moritz 332 (Venezuela). Ap- parently a label of the latter was inadvertently af- fixed to an Ecuadorean specimen (K) of what is now recognized as Grammitis pichinchensis. Hooker (1862), unaware of a true isotype of P. subscabrum (BM), applied this name to the speci- men he thought was type material at Kew. He was followed by subsequent authors, including Morton (1967), who made the combination of Grammitis subscabra. Polypodium subscabrum Klotzsch is a true Polypodium, not Grammitis, and is likely con- fined to Venezuela. Further confusion resulted when Polypodium pichinchense Hieron. was published. Despite its similar name, Polypodium pichinchae Sodiro (Grammitis pichinchae (Sodiro) Morton) is a dif- ferent species. Grammitis pichinchae (Ecuador) has leaves up to twice as long and broad, pinnae most- ly acute, and glabrous adaxially, and hydathodes lacking white deposits. For full clarification of no- menclature and correct application of names, see Stolze (Amer. Fern J. 81: 139-140. 1991). Huanuco: Cani, 7 mi NE of Mito, Bryan 383 (F). 59. Grammitis alsopteris Morton, Contr. U.S. Natl. Herb. 38: 112. 1967. TYPE: Ecuador, Napo-Pastaza, Canton Mera, Mexia 7013 (holotype, us!; photo, F). 110 FIELDIANA: BOTANY Plants epiphytic, very rarely epipetric. Stem erect, provided with deltate or linear-deltate, lus- trous orange or reddish brown scales, these 2—4 mm long, not or scarcely clathrate, the margins and apex with hyaline setae. Leaves 15-45 cm long, (3.5-)4-10(-12) cm broad. Petiole 7-16 cm long, 0.8-1.5 mm in diameter, amply provided with pale to deep orange, spreading, unicellular trichomes to 2 mm long. Lamina chartaceous, del- tate or lanceolate-deltate, pectinate, 1 -pinnate, gradually reduced to apex, base not or scarcely reduced, proximal pinnae as long as the rest, or 1-3 pairs of them slightly reduced, the rachis tri- chomes 0.4-0.7 mm long abaxially, 0.2-0.3 mm long adaxially, surface amply but minutely cas- taneous-puberulent abaxially, glabrous adaxially. Pinnae 2-6 cm long, 0.2-0.5 cm broad beyond the dilated base, 6-15 times as long as broad, linear or linear-deltate, apex acute or subacute, adnate, usually host to clavate, black fungi on the abaxial surface, margins entire. Veins simple, 10-26 pairs on a pinna, hydathodes distinct and commonly encrusted with calcareous deposits. Sori supra- medial to submarginal, sporangia glabrous. In rain forests, on tree trunks, or, very rarely, on wet, moss-covered rocks, 2300-2850 m, Pasco and Cuzco. Costa Rica; Panama; Colombia to Peru. The species occurs at lower elevations (900-2 1 00 m) in Colombia and Ecuador. It is easily distin- guished from most of the related taxa by the mi- nute, rigid (0. 1 mm), castaneous trichomes on the abaxial laminar surface, while the others have gla- brous surfaces. Grammitis taxifolia has pale, usu- ally longer trichomes on the abaxial pinna surface, but the lamina is conspicuously reduced at base, and the margins of the stem scales are naked, not setose. Hieronymus described two varieties of G. semihirsuta from Colombia: Polypodium semihir- sutum var.fuscosetosum and var. hirtopuberulwn. Both these varieties supposedly have longer, spreading trichomes on the pinnae. Pasco: Prov. Oxapampa, Mallampampa, D. Smith & Canne 5800, in part (MO, uc). Cuzco: Trail from Machu Picchu to Winay Waina, Bishop 2507 (uc). Prov. Uru- bamba, Machu Picchu, trail to Intipata, Leon 2107 (F). 60. Grammitis leucosticta (John Sm.) Morton, Contr. U.S. Natl. Herb. 38: 112. 1967. Polypodium leucosticton Fee, (Mem. foug. 5) Gen. fil. 240. 1852, not Klotzsch, 1847. TYPE: Ecuador, Quito, Jameson (holotype, p?; isotype, us!, photo, F of us). Ctenopte ris leucosticta John Sm., Hist. fil. 185. 1875. Type same as for P. leucosticton Fee. Polypodium longiusculum C. Chr., Index fil. 54 1 . 1 906. Ctenopteris longiuscula (C. Chr.) Copel., Philipp. J. Sci. 84: 456. 1956, nom. super/], for C leucosticta John Sm. Plants epiphytic, very rarely terrestrial. Stem erect, provided with narrow-deltate, lustrous, deep orange to castaneous scales, these 2-4 mm long, not or scarcely clathrate, the margins with con- colorous setae. Leaves 25-60 cm long, 2.5—4 cm broad. Petiole 6-16 cm long, 0.8-1.2 mm in di- ameter, amply provided with deep orange to cas- taneous, spreading, unicellular trichomes to 2 mm long. Lamina chartaceous, narrow-elliptic or ob- long-elliptic, 1 -pinnate, gradually reduced to apex and base, the rachis trichomes 1-2 mm long (at least proximally), sometimes partially broken off on dried specimens, surfaces glabrous, although costae adaxially often with a few, long, deciduous trichomes. Pinnae 1.2-2 cm long, 0.3-0.45 cm broad beyond the often broadened base, 4.5-6 times as long as broad, deltate or oblong-deltate, obtuse to subacute, adnate, usually host to clavate, black fungi on the abaxial surface, margins entire. Veins simple, 6-17 pairs on a pinna, hydathodes distinct and commonly encrusted with calcareous deposits. Sori supramedial or submarginal, spo- rangia glabrous. In wet forests, on tree trunks or rarely on the forest floor, 1900-3100 m, Amazonas and Pasco. Ecuador; Peru. The most obvious distinction between this and its nearest relatives, G. semihirsuta and G. alsop- teris, is the shorter and broader, usually obtuse, pinnae. The lamina of the other two species is pectinate, with crowded, linear or linear-deltate, usually acute pinnae. Amazonas: Prov. Bagua, Cordillera Colan SE of La Peca, Barbour 4137, 4177 (both MO). Prov. Chachapo- yas, Cerros Calla Calla above Balsas, Hutchison A Wright 5827 (F, GH, uc). 40 mi E of Chachapoyas, Osgood & Anderson 51 (F, us). Pasco: Prov. Oxapampa, 4-5 km N of Mallampampa, D. Smith & Canne 5800, in part (MO, uc). 61. Grammitis semihirsuta (Klotzsch) Morton, Contr. U.S. Natl. Herb. 38: 113. 1967. Polypodium semihirsutum Klotzsch, Linnaea 20: 379. TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. Ill 1847. TYPE: Peru, Ruiz 9 (holotype, B?; isotype, us!, photo, F of us). Ctenopteris semihirsuta (Klotzsch) Copel., Philipp. J. Sci. 84: 450. 1956. Plants epiphytic. Stem erect, provided with nar- row-deltate, lustrous, castaneous scales, these 2-3 mm long, nonclathrate, the margins with orange or castaneous setae. Leaves 1 5-50 cm long, 3.5-8 cm broad. Petiole 5-14 cm long, 0.8-1.5 mm in diameter, sparsely to amply provided with deep orange to castaneous, spreading, unicellular tri- chomes to 2 mm long. Lamina firm-herbaceous to chartaceous, lanceolate or elliptic, 1 -pinnate, gradually reduced to apex and base, the rachis tri- chomes scattered to ample, to 1 mm long abaxially often with a few long, deciduous trichomes. Pinnae 1.2-3.5 cm long, 0.3-0.5 cm broad beyond the often dilated base, 7-9 times as long as broad, linear-deltate, acute, adnate, usually host to cla- vate, black fungi on the abaxial surface, margins entire. Veins simple, 1 6-24 pairs on a pinna, hy- dathodes distinct and often encrusted with cal- careous deposits. Sori supramedial or submargin- al, sporangia glabrous. In wet forests, on tree trunks, 2450-3100 m, Amazonas, San Martin, and Huanuco. Southern Mexico to Panama; Jamaica; Hispan- iola; Venezuela and Colombia to Bolivia and Bra- zil. This is part of a large complex of species in- cluding, among others, G. alsopteris, G. leucostic- ta, G. pichinchensis, and G. david-smithii and characterized by conspicuously setose stem scales and axes provided with orange to castaneous, spreading, unicellular trichomes 0.4-2 mm long. Most species have pectinate laminae, with linear pinnae or segments, white-encrusted hydathodes, and are host to a black, clavate, ascomycete, Ac- rospermum maxonii Riddle, which grows es- pecially on the costae and among the sori. Because of the very subtle differences and misapplied spe- cies names, great confusion has attended both tax- onomy and nomenclature. Furthermore, speci- mens of G. taxifolia and a few of its closest relatives are often identified as species within this complex, as they share some of these same diagnostic fea- tures, and are also hosts to Acrospermum. How- ever, G. taxifolia can be separated by the stem scales, which lack marginal setae, and the pale, sericeous trichomes on the petiole and lamina. Also, it is likely that some of the species are hy- bridizing, and monographic study is sorely needed of the entire group. Amazonas: Cerros Calla Calla, between Leimebamba and Balsas, Hutchison & Wright 5827a (GH, uc). San Martin: Prov. Mariscal Caceres, Rio Abiseo National Park, Leon 2159 (uc), Young 2659 (USM), 2728 (USM), 3187 (USM). Huanuco: Muna, trail to Tambo de Vaca, Macbride 4294 (F, us). 62. Grammitis blepharidea (Copel.) Stolze, comb, nov. Polypodium blepharideum Copel., Univ. Calif. Publ. Hot. 19: 304. 1941. TYPE: Peru, Huanuco, Dist. Churubamba, Cresta Santa Toribio, Mexia 8 147 a (holotype, uc; isotype, us!; photos, F & GH of us). Polypodium buesii Maxon, Contr. Gray Herb. 165: 72. 1947. TYPE: Peru, Cuzco, Cerro Chuyapi, Bikes A45 (holotype, us!). Xiphopteris blepharidea (Copel.) Copel., Amer. Fern J. 42: 99. 1952. Xiphopteris buesii (Maxon) Copel., Amer. Fern J. 42: 105. 1952. Grammitis buesii (Maxon) Lell., Amer. Fern J. 74: 58. 1984. Plants epiphytic, rarely epipetric. Stem small, erect, provided with broad, flaccid, dull yellow scales, these 2-3 mm long, minutely clathrate, the margins entire or with scattered, minute, rotund glands. Leaves 8-20 cm long, 0.6-0.9 cm broad. Petiole 0.5-2.5 cm long, 0.4-0.7 mm in diameter, abundantly provided with stout, spreading reddish to castaneous, unicellular trichomes to 2 mm long, and with a few, scattered simple or branched sep- tate trichomes 0.1-0.2 mm long. Lamina thin- to firm-herbaceous, linear-elliptic, divided nearly or quite to the rachis, strongly and gradually reduced at both ends, unicellular trichomes on rachis, sur- face and margins as on the petiole, but less abun- dant. Pinnae (or segments) 3-5 mm long, 1.5-2.5 mm broad, deltate or oblong-deltate, obtuse or subacute, margins entire. Veins usually simple, but fertile ones with a short acroscopic branch near the segment base on which the sorus is borne, hydathodes distinct, lacking calcareous deposits. Sori solitary at the base of each segment, sporangia glabrous. In wet forests, on tree trunks, or very rarely at the base of rocky cliffs, 2000-2750 m, along the Cordillera Central from Cajamarca and Amazonas to Puno. Endemic. 112 FIELDIANA: BOTANY Copeland (1952b) separated this from G. buesii by only two characters: the latter with "toothed" stem scales and 1 -forked veins; the former with entire scales and simple veins. The type of Poly- podium blepharideum consists of two sterile, im- mature leaves, each pinna bearing a simple vein, and the stem scales apparently with no marginal processes. The type of Polypodium buesii contains mostly fertile leaves, their pinnae with a single vein and a short, basal, acroscopic spur that bears the sorus; however, on the few sterile leaves the veins are unbranched. Stem scales commonly have a few, nearly undetectable, rotund glands scattered along the margin, but at least a few scales have naked margins. The scale glands are delicate and apparently are easily dislodged. Therefore, the sep- aration of G. buesii is unwarranted, since all other characters are identical. Cajamarca: Prov. Cutervo, Chontacruz, San Andres, Quiroz & Suarez 2894, in part (F). Amazonas: Prov. Cha- chapoyas, along Rio Ventanilla W of Molinopampa, Wurdack 1511 (F, GH, uc, us). Pasco: Prov. Oxapampa, trail to Cordillera Yanachaga via Rio San Daniel, D. Smith el al. 7853 (MO), 8490 (MO). Cuzco: Cabecera del Koribeni, Sues 1963 (GH, us). Cerro Chuyapi, Biies A38 (us). Puno: Prov. Sandia, slopes of Oconeque, E of Lim- bani, Hodge 6087 (F, GH, us). 63. Grammitis taxifolia (L.) Proctor, Rhodora 63: 35. 1961. Polypodium taxifolium L., Sp. pi. 2: 1086. 1 753. TYPE: Plumier t. 89, Traite foug. Amer., based on a Plumier specimen from Martinique. Ctenopteris taxifolia (L.) Copel., Philipp. J. Sci. 84: 447. 1956. Plants epiphytic. Stem short-creeping to erect, provided with narrow-deltate, rigid, lustrous, or- ange to red-brown scales, these 1-2.5 mm long, nonclathrate, the margins naked but the tip setose. Leaves 12-35 cm long, 3-6 cm broad. Petiole 3- 10 cm long, 0.7-1.2 mm in diameter, abundantly provided with yellow or pale orange, sericeous, unicellular trichomes to 2 mm long. Lamina pec- tinate, 1 -pinnate, thin- to firm-herbaceous, ellip- tic, gradually or abruptly reduced at both ends, trichomes on axes, surface and margins of seg- ments as on the petiole, but less abundant and a little shorter. Pinnae 1.5-3.5 cm long, 3-5 mm broad, linear, adnate, obtuse or subacute, truncate at base, usually host to clavate, black fungi on the abaxial surface, margins entire. Veins simple, 6— 18 pairs on a pinna, hydathodes distinct, rarely (but not in Peru) with calcareous deposits. Sori medial, sporangia glabrous. In wet forests, on tree trunks, 1950-2500 m, along the Cordillera Central from Amazonas to Pasco. Costa Rica; Hispaniola; Puerto Rico; Lesser An- tilles; Trinidad; Venezuela and Colombia to Brazil and Bolivia. This is often confused with some of the species in the complex of G. semihirsuta (q.v.). There is a close resemblance in the size and shape of leaves, the pectinate lamina, and all are hosts to the dis- tinctive black ascomycete Acrospermum, yet there are subtle but distinct diagnostic characters. Spe- cies in the G. semihirsuta complex have conspic- uous setae on margins of stem scales, and petiole and rachis trichomes are stout, rigid, and casta- neous. In G. taxifolia stem scales have an apical seta, but none on the margins, and axes trichomes are sericeous and yellow or orange. In addition, most species in the semihirsuta group have sub- marginal son, whereas those in G. taxifolia are medial. South American specimens seem to be more robust than West Indian ones, with thicker and broader laminae. Also, Proctor (Flora Lesser An- tilles, 1 977) contended that hydathodes of G. taxi- folia are encrusted with the circular, calcareous deposits common to the semihirsuta group. How- ever, these white dots are lacking on the Antillean specimens at Field Museum and are not present on any of the Peruvian material. Amazonas: Prov. Bagua, 20 km E of La Peca, Barbour 2835 (F, MO). San Martin: Prov. Rioja, Pedro Ruiz- Moyobamba Road, Venceremos, D. Smith & Vdsquez 4577 (MO, uc). Huanuco: Prov. Huanuco, road from Acomayo to Chinchao, Mexia 4142a (uc). Pasco: Prov. Oxapampa, trail to Cordillera Yanachaga via Rio San Daniel, D. Smith et al. 7855 (F, MO, USM). 64. Grammitis athyrioides (Hooker) Morton, Contr. U.S. Natl. Herb. 38: 255. 1973. Polypodium athyrioides Hooker, Sp. fil. 4: 224, /. 277b. 1863. TYPE: Peru, Junin, Pangoa (San Martin de Pangoa), Mathews 1 103 (holotype, K!; photo, F). Polypodium yungense Rosenst., Repert. Spec. Nov. Regni Veg. 5: 236. 1908. TYPE: Bolivia, North Yungas, Unduavi, Buchtien 891 (holotype, B?; isotype, us!). Ctenopteris athyrioides (Hooker) Copel., Philipp. J. Sci. 84: 406. 1956. TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 113 Ctenopterisyungensis(Rosensl.) Copel. Philipp. J. Sci. 84: 407. 1956. Plants epiphytic. Stem short-creeping or decum- bent, sparsely provided with narrow-deltate, non- clathrate scales, these 2-3 mm long, dark brown to black, with rigid brown or orange setae. Leaves lax, usually pendent, swollen and irregularly ar- ticulate at the stem, crowded to subfasciculate, 1 -pinnate, 20-50 cm long, 2.5-6 cm broad. Petiole 6-10 cm long, 0.6-1.2 mm in diameter, amply provided with spreading trichomes, these orange to castaneous, 0.5-2 mm long. Lamina firm-her- baceous to chartaceous, reduced at apex and base. Rachis brown to blackish, amply provided with spreading, castaneous trichomes to 1 mm long. Pinnae slightly ascending, adnate, narrow deltate, deeply lobed to broadly crenate, at least near the base, commonly bearing some black clavate fungi on the abaxial surface. Veins commonly simple, hydathodes often covered by circular, calcareous deposits. Sori round, supramedial. Sporangia gla- brous. In forests, on tree trunks, 2100-3750 m, from Amazonas to Cuzco. Peru and Bolivia. A number of specimens have been determined as the Bolivian Polypodium yungense, the latter supposedly differing from G. athyrioides in the cre- nate (vs. deeply lobed) pinna margins. However, examination of type and other specimens from Peru and Bolivia identified as either species re- vealed no other differences. Throughout a large number of specimens there can be observed a pro- gressive gradation from subentire to broadly cre- nate pinnae to those lobed about halfway to the costa; thus, it is presumed that the deeply lobed pinnae are simply manifestations of a more robust grown habit. Furthermore, the types of both ex- hibit two very distinctive features: short, blackish stem scales with brownish setae, and the (usual) presence of black, clavate fungi on the pinnae. Amazonas: Prov. Chachapoyas, Cerros Calla Calla above Leimebamba, Hutchison & Wright 6988 (us). Prov. Chachapoyas, slopes of Puma-Urcu, Wurdack 703 (GH, us). San Martin: Prov. Mariscal Caceres, Rio Abiseo National Park, Leon 1898 (uc), Young & Leon 4649 (USM). Huanuco: Mufia, Bryan 550B (F). Mima, trail to Tambo de Vaca, Macbride 4305 (F), 4329 (F). Cuzco: Valle de Lares, above Rio Lachae, Sues 1822 (us). Prov. La Convencion, Dist. Vilcabamba, Bues 2116 (us). Be- low Abra de Malaga, 1 5 km from Quillabamba, Ellen- berg 4776 (GH, in part). 65. Grammitis myriophylla (Baker) Morton, Contr. U.S. Natl. Herb. 38: 108. 1967. Figure 5g. Polypodium myriophyllum Baker, in Hooker and Ba- ker, Syn. fil. 338. 1868. TYPE: Peru, Puno, Ta- tanara, Lechler 2567 (holotype, 2 sheets, K!; iso- type, L; frag., us of L; photos, F of K, us of K & L). Plants epiphytic. Stem suberect, very sparsely provided with minute, nonclathrate, ovate or del- tate, orange scales, these 0.5-0.8 mm long, their margins subentire or with short, scattered glands. Leaves lax, pendent, not or scarcely articulate, densely fasciculate, 1 -pinnate-pinnatisect to (rare- ly) 2-pinnate, 1 5-30 cm long, 2-5 cm broad. Pet- iole 3-9 cm long, 0.4-0.8 mm in diameter, copi- ously provided with orange, spreading, unicellular trichomes to 2 mm long. Lamina firm-herbaceous, gradually reduced at apex and base, rachis and segments moderately provided with deep-casta- neous, rigid, spreading, unicellular trichomes. Pin- nae ascending at a 40-60° angle, 1.5-5 cm long, 0.4—1 cm broad, linear to narrow-lanceolate, di- vided nearly or quite to the costa into well-spaced, linear segments, these rarely with a single, basal, linear segment. Veins simple, one to a pinna or ultimate segment, each terminating in a brownish hydathode. Sori round, solitary at the very base of each ultimate segment and much broader than the segment at maturity, sporangia glabrous. In forests, on tree trunks and on thick layers of mosses on the forest floor, 2800-3400 m, Pasco, Cuzco, and Puno. Ecuador to Bolivia. Very similar to this (and perhaps to be com- bined with it) is Polypodium longisetosum Hooker of Ecuador (type not seen). The latter differs in the broadly oblong (vs. linear) ultimate segments. Grammitis myriophylla is not apt to be confused with any other species in the genus. It is easily distinguished by its nearly 2-pinnate leaves with strongly ascending pinnae, and the widely spaced ultimate segments which are so narrow that the mature sori extend beyond the margins. Pasco: Prov. Oxapampa, Yanachaga National Park, path from Abra La Esperanza to Rio Pescado, Leon 1005 (F). Cuzco: Altura de Sicre, Bues 1539 (us), 7554 (us). Prov. La Convencion, Valle de San Miguel, Bues 2191 (us). Prov. La Convencion, Cordillera Vilcabamba, Dud- ley 11 134 A (GH). Prov. Paucartambo, Cerro Macho Cruz, Leon 2301 (F). 114 FIELDIANA: BOTANY 66. Grammitis immixta Stolze, sp. nov. Species haec ab Grammite variabile (Kuhn) Morton differt pubibus densioribus, hydathodibus calcareis, et trichomibus longis, stellatis castane- isque vel simplicibus pallidisque immixtis. Plants epiphytic, possibly also epipetric. Stem suberect, provided with a few orange, ovate or deltate, nonclathrate scales, these 1-1.5 mm long, the margins with long, pale setae. Leaves lax, pen- dent, not or scarcely articulate, fasciculate, 2-pin- nate, 15-50 cm long, 2.5-7 cm broad. Petiole 2- 6 cm long, 0.2-0.5 mm in diameter, sparsely to amply provided with thin, yellow, spreading, uni- cellular trichomes 1-2 mm long. Lamina firm-her- baceous, gradually reduced to apex and base, ra- chis and segments densely covered with (and often obscured by) trichomes like those of the petiole, and intermixed with castaneous ones, these stel- lately branched at base into 5-6 long arms. Pinnae ascending at a 20-60° angle, 2.5-7 cm long, 0.4- 0.6 cm broad, linear, bearing many crowded, ob- long to nearly circular segments. Veins simple in each ultimate segment, hydathodes usually en- crusted with a round, calcareous deposit. Sori one to a segment, sporangia provided with rigid setae, these often longer than the sporangia. TYPE— Peru, Cuzco, Prov. La Convention, Distr. Vilcabamba, Ruina Idma-huasi, Bues 2103 (ho- lotype, us!; isotypes, F!, GH!, MO!). In forests, on tree trunks (or, though not stated on labels, possibly pendent from rocky walls of ruins), 3200-3650 m, Cuzco. Endemic. This is closely related to G. variabilis, but in addition to the characters used in the key, it can be distinguished further by the density of the pu- bescence. Although trichomes on the lamina of G. variabilis are abundant, the tissue of the segments is always evident beneath them. However, in G. immixta the segments are so copiously villous that their surfaces are often completely obscured, es- pecially abaxially. Cuzco: Ccarcco, Bues 1393 (us, in part). Prov. La Con- vencion, Abra Mirador, Bues 2071 (us). Prov. Urubam- ba, Machu Picchu, Peyton & Peyton 1 102 (MO). 67. Grammitis variabilis (Kuhn) Morton, Contr. U.S. Natl. Herb. 38: 102. 1967. Polypodium variabile Kuhn, Linnaea 36: 133. 1869. SYNTYPES: Ecuador, Andes of Quito, Jameson (B?, K.!; photo, F of K); New Granada (Colombia), Purdie (B?). Plants epiphytic, rarely epipetric. Stem suberect, provided with a few orange or brown, ovate or deltate, nonclathrate scales, these 1-1.5 mm long, the margins with pale setae. Leaves lax, pendent, not or scarcely articulate, fasciculate, 2-pinnate, not or very rarely forked, 1 5-30 cm long, 2-4 cm broad. Petiole 2-5 cm long (often appearing longer because of deciduous proximal pinnae), 0.2-0.4 mm in diameter, amply provided with yellow or orange, spreading, unicellular trichomes to 2 mm long. Lamina membranaceous to firm-herbaceous, reduced to apex and base, rachis and segments amply pilose like the petiole, but the long tri- chomes of the rachis sparsely to moderately in- termixed with minute (0. 1-0.3 mm), septate, sim- ple or branched, gland-tipped ones. Pinnae ascending at a 40-60° angle, 1—4 cm long, 0.3-0.7 cm broad, linear to narrow-lanceolate, bearing several to many, approximate to crowded, oblong to circular segments. Veins simple in each ultimate segment, each terminating in a brownish hyda- thode. Sori round, one to a segment, sporangia provided with rigid setae, these sometimes longer than the sporangia. In forests, on tree trunks, or rarely in crevices of rocky cliffs, 3350-3950 m, Junin and Cuzco. Colombia to Peru. Copeland (1956) included this with G. hetero- morpha, suggesting that the two differed only in the dissection and branching of the lamina, but there are other distinctions, particularly the rachis indument and, less obviously, the stem base. Nei- ther of the type specimens (and few extant speci- mens) seen have stems, but those few seen are swollen and articulate at base in G. heteromorpha, but not or scarcely so in G. variabilis. Although rachises of both species are abundantly provided with the same abundant, long-spreading, light-col- ored, unicellular trichomes, each has another kind of trichome intermixed: in G. heteromorpha, some scattered, long castaneous, sessile-stellate ones; in G. variabilis, some minute, septate, gland-tipped ones. There are two sheets in the type folder of Polypo- dium heteromorphum at Kew, one with 2-pinnate leaves and the other with 1 -pinnate but forking leaves. Both are Jameson specimens from the "Andes of Quito." Hooker obviously considered TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 115 these the same collection, and even the original drawing (Icon. fil. /. 108) was based on the two different lamina patterns, hence the epithet "het- eromorpha." On the sheet with the 2-pinnate leaf is pencilled, in a different hand (Mettenius'?) "P. variabile Mett." It is not certain if Mettenius saw this as well as another Jameson syntype (B?), but it differs from the holotype off. heteromorpha in the characters mentioned above. Obviously, there are a number of problems con- cerning these two species and G. dependens, and the entire group is in need of further study. Junin: San Jose, Macbride 1108 (F, us). Cuzco: Sucra- cucha (?), Bikes 1585 (us). Ccarcco, Biies 1393 (us, in part). Prov. La Convention, NE of Hacienda Luisiana, Dudley 11 189 (GH, us). II. Pecluma Pecluma Price, Amer. Fern J. 73: 109. 1983. TYPE: Pecluma pectinata (L.) Price (Polypodium pectinatum L.). Figure 6. Plants epiphytic or epipetric, occasionally ter- restrial. Stem short, creeping to decumbent or sub- erect, unbranched, not glaucous, bearing often- proliferous roots and abundant scales, the latter basally attached, nonclathrate, and with essen- tially undifferentiated margins. Leaves monomor- phic, fasciculate to crowded (not over 1 .5 cm apart), short- or long-petiolate, the petiole terete, dark brown or black, articulate to short phyllopodia. Lamina pectinate, deeply incised nearly or quite to the rachis into numerous, narrow, usually crowded segments, sparsely to amply provided with pluricellular trichomes on laminar surface and/or rachis. Veins free or casually anastomosing, forked (or simple in 3 species). Sori terminal on the sim- ple, basal acroscopic vein branch, exindusiate, the receptacle paraphysate, sporangia glabrous or se- tose, the stalks with 2-3 rows of cells. Spores mon- olete, bilateral, reniform to subglobose, yellow (not chlorophyllous). The genus contains about 27 species from trop- ical or subtropical regions of the New World. It is especially distinguished by the pectinate lamina with pluricellular trichomes, the dark, terete pet- ioles articulate to short phyllopodia, the short, un- branched stems bearing basally attached, non- clathrate scales, and the exindusiate sori borne on the tips of usually free veinlets. The pectinate as- pect of the lamina is imparted by the numerous, crowded, spreading, linear segments, which are en- tire, or occasionally crenate. Only in P. funicula (Fee) Price of Cuba and P. c!~oquetangensis (Ro- senst.) Price of Bolivia are the segments conspic- uously pinnatifid. According to Price (1983) Pe- cluma shares ancestry with Polypodium and Ctenopteris (Grammitis subg. Cryptosorus), with whose species it is commonly confused. This treat- ment essentially follows that of Evans (1969). Eleven species are known from Peru. References EVANS, A. M. 1969. Interspecific relationships in the Polypodium pectinatum- plumula com- plex. Ann. Missouri Bot. Gard., 55: 193-293. PRICE, M. G. 1983. Pecluma, a new tropical American fern genus. Amer. Fern J., 73: 109- 116. Key to Species of Pecluma a. Rachis scales lanceolate to ovate, conspicuous and persistent, at least abaxially; rachis and petiole black (or brown in P. filiculd) b b. Leaf 6-18 cm long: petiole and rachis brown; stem scales ovate, to 1-2 mm long 2. P. ti lieu la b. Leaf 20—60 cm long; petiole and rachis black; stem scales linear or narrow-deltate, 2-5 mm long 1 . P. plumula a. Rachis scales lacking, or sparse, filiform, and inconspicuous; rachis and petiole brown c c. Proximal segments not reduced, or basal pair at least half as long as the longest ones d d. Costae perpendicular to rachis; sori mostly supramedial; sporangia glabrous 5. P. divaricata d. Costae decurrent on rachis; sori medial; sporangia setose 11. P. hygrometrica c. Proximal segments strongly reduced, often to only wings or auricles e e. Segments, especially distal ones, usually ascending at a 45-50° angle; rachis abaxially with scattered trichomes to 1 .5 mm long 4. P. curvans 116 FIELDIANA: BOTANY FIG. 6. Pecluma hygrometrica: a, habit. Peduma filicula: b, habit. Pecluma eurybasis var. pilosa: c, portion of rachis with pinna base. Pecluma curvans: d, rachis and pinnae, (a from Moran 3665. F, b from Tryon & Tryon 5373. F, c from Young 4938, USM; d from Soukup 3642, us.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 117 e. Segments patent to slightly ascending, or sometimes basal ones deflexed; rachis trichomes of various lengths f f. Base of segments asymmetrical, mostly perpendicular or rounded acroscopically, decurrent basiscopically; trichomes on segment margins lacking or scattered and inconspicuous .... 3. P. absidata f. Base of segments essentially symmetrical, or if asymmetrical then decurrent acroscopically and perpendicular basiscopically; trichomes conspicuous on segment margins (sometimes caducous) g g. Lamina base with a few segments ^ -'/2 the length of larger ones; sporangia glabrous h h. Trichomes on rachis 0.2-0.5 mm long, on costa 0.2-0.3 mm long; veins glabrous . . 6. P. eurybasis h. Trichomes on rachis (at least abaxially) 0.6-1 mm long, on costa 0.4-0.8 mm long; veins sparsely to moderately provided with acicular trichomes 0.4-0.8 mm long . . . 8. P. venturii g. Lamina base gradually or abruptly reduced to rudimentary segments or wings; sporangia with 1-3 capsular (often deciduous) setae j i. Laminar tissue glabrous except for a small, oblong patch of minute trichomes sur- rounding the sorus; costa perpendicular to the rachis; veins free .... 10. P. ptilodon i. Laminar tissue regularly pilose or minutely puberulent; costa slightly to strongly de- current on the rachis; veins free to partly anastomosing k j. Rachis abaxially puberulent with trichomes 0.1-0.2 mm long; lamina abruptly reduced to a few small lobes or auricles; trichomes on costa and laminar tissue inconspicuous, 0. 1 mm long 7. P. pectinata j. Rachis abaxially pilose with trichomes 0.4-0.8 mm long; lamina cuneate to atten- uate at base, the segments (often many) gradually reduced to small lobes or auricles; trichomes on costa and laminar tissue usually conspicuous, 0.3-0.5 mm long . . . 9. P. camptophyllaria 1 . Pecluma plumula (Willd.) Price, Amer. Fern J. 73: 115. 1983. PolypodiumplumulaWi\\d.,Sp.pl.ed.4,5: 178. 1810. LECTOTYPE (designated by Evans, 1 969, p. 230): Venezuela, Caracas, Bredemeyer (B, Herb. Willd. 19655-1; frag., NY; photos, F, GH). Plants epiphytic, epipetric or terrestrial. Stem short-creeping, provided with linear or narrow- deltate, acuminate or attenuate, reddish brown scales, these 2-5 mm long, subentire, often with a filiform tip. Leaves 20-60 cm long, crowded to subfasciculate. Petiole 2-10(-12) cm long, black. Lamina elliptic or elliptic-lanceolate, 3-7.5 cm broad, gradually or abruptly reduced at base. Ra- chis black, abaxially provided with orange to red- dish brown scales, these cordate to basifixed, plane to sub-bullate, often ciliate, to 1 mm long, and with scattered to abundant, acicular, pluricellular trichomes. Segments perpendicular to the rachis, or distal ones slightly ascending, the costae, mar- gins, and (occasionally) the laminar surface pu- bescent like the rachis. Veins obscure, 1 (-2)-forked, free. Sori medial to supramedial. Sporangia se- tose. In forests and wooded canyons, on trees or fallen logs, on rocks or in rock crevices, occasionally on the forest floor, 100-2000 m, Amazonas and Lo- reto, south to Ayacucho and Madre de Dios. United States (Florida); Mexico to Panama; West Indies; Colombia to the Guianas, south to Bolivia and Brazil. Practically indistinguishable from this is P. dis- persa (Evans) Price, which shares the same range, but which heretofore has not been found in Peru. Evans ( 1 969) described the latter as a triploid, with 32 spores per sporangium, whereas P. plumula is a tetraploid, with 64 spores per sporangium. The only other distinctions given between the two taxa were that the stem scale margins of P. plumula are somewhat papillose and the veins are rarely twice forked, whereas scale margins of P. dispersa are somewhat papillose and the veins are often twice forked. Amazonas: Prov. Bongara, Shipasbamba-Pomacocha trail, Wurdack 1093 (F, GH, us). San Martin: Alto Rio Huallaga, LI. Williams 5586 (F). Loreto: Mishuyacu, near Iquitos, King 1382 (F, us). Prov. Maynas, Lupuna Cocha, Tryon & Tryon 5186 (F, GH, us). Huanuco: Piedra Gran- de Estacion, near Santo Domingo, Macbride 3703 (F, 118 FIELDIANA: BOTANY OH, us). Junin: La Merced, Killip & Smith 23796 (F, us). Yaupe: Woytkowski 6393 (GH, MO, us). Ayacucho: "Aina" (Ayna), Tidschack 89 (B). Cuzco: Prov. La Convention, Vilcanota Valley, Mexia 8090 (F, GH, MO, NY, uc, us). Madre de Dios: Prov. Manu, Parque National del Manu, Cocha Cashu, Foster et al. 3454 (F). 2. 1 Yd u ma filicula (Kaulf.) Price, Amer. Fern J. 73: 114. 1983. Figure 6b. Polypodiumfilicula Kaulf., Enum. fil. 275. 1 824. TYPE: Brazil, Chamisso (holotype, LE?). Plants epiphytic or epipetric. Stem very short- creeping, provided with ovate, acute, brown or reddish brown scales, these about 1-2 mm long, subentire or finely serrulate. Leaves 6-1 8 cm long, fasciculate. Petiole 0.5—4 cm long, reddish brown. Lamina elliptic, 1 .5-3 cm broad, tapered gradually at both ends. Rachis reddish brown, amply and conspicuously provided abaxially with reddish brown, cordate, sub-bullate, subentire to dentic- ulate scales to 1 mm long and abundantly pubes- cent on both sides with acicular, pluricellular tri- chomes to 0.3 mm long. Segments perpendicular to the rachis, or distal ones slightly ascending, the costae and margins pubescent like the rachis. Veins simple, free. Sori supramedial. Sporangia lacking setae. In wet forests, on trees or fallen logs, or on rocks or in crevices of rock walls, 800-2400 m, Caja- marca, Amazonas, San Martin, Huanuco, and Cuzco. Colombia to Argentina and southern Brazil. This is the smallest species of Pecluma in Peru and is also notable for the simple veins, broad, cordate, conspicuous rachis scales, and suprame- dial son. Cajamarca: Prov. Jaen, Chontali, Chimoy 261 (USM). Amazonas: Prov. Bagua, on Cerro Tapur ca. 40 km S of Bagua Grande, Hutchison 1472 (uc, us). San Martin: San Martin, km 28 of Tarapoto-Yurimaguas road, Knapp & Mallet 8402 (F, MO). Huanuco: Muna, Bryan 431 (F). Yanano, Macbride 3820 (F, us). Muna, Woytkowski 5216 (GH, MO, us). Cuzco: Prov. La Convention, Rio Chau- pimayo, Soukup 807 (F). Prov. La Convention, Potrero, 8 km W of Quillabamba, Tryon & Tryon 5373 (F, GH, us). 3. Pecluma absidata (Evans) Price, Amer. Fern J. 73: 113. 1983. Polypodium absidatum Evans, Ann. Missouri Bot. Card. 55: 238. 1 969. TYPE: Columbia, Dist. San- tander, Paramo de Romeral, Killip & Smith 18518 (holotype, us!; isotype, GH!). Plants epiphytic or epipetric. Stem rather long- creeping, the scales linear to narrow-deltate, at- tenuate, light or dark brown, sublustruous, sub- entire, 3-6 mm long. Leaves 20-40(-50) cm long, approximate. Petiole 3-8 cm long, reddish brown, Lamina subcoriaceous, 4-6 cm broad, elliptic, rather abruptly reduced at base. Rachis reddish brown, scales lacking or rare and filiform, sparsely provided abaxially with acicular trichomes about 0.4 mm long, and moderately to amply adaxially with trichomes 0.7-1 mm long. Segments (at least distal ones) somewhat ascending, at a 60-70° an- gle, most of these nearly perpendicular to or rounded at the rachis acroscopically, decurrent basiscopically, the costae strongly decurrent, gla- brous or with scattered trichomes, the margins usually sinuate and glabrous or with scattered, in- conspicuous trichomes. Veins free, obscure, 1 -forked. Sporangia with 1 or 2 long setae. Sori medial. In forests or at edges of clearings, on rocks or trees, 2900-3700 m, Cajamarca, La Libertad, and Ancash. Greater Antilles; Venezuela; Colombia to Bo- livia. This is easily confused with those specimens of P. curvans whose pinnae are not strongly ascend- ing. See treatment of the latter for further discus- sion. Cajamarca: Prov. Celendin, Canyon of Rio Maranon above Balsas, Hutchison & Wright 5324, in part (F, GH). La Libertad: Prov. Otuzco, near Agallpampa, Lopez 1024 (HUT, us). Ancash: Prov. Yungay, near Laguna Llangan- uco, Mostacero et al. 1417 (HUT, MO). 4. Pecluma curvans (Mett.) Price, Amer. Fern J. 73: 114. 1983. Figure 6d. Polypodium curvans Mett., Ann. Sci. Nat. Bot. 5, 2: 253. 1864. TYPE: Peru, Agapata, Lechler 2006 (holotype, B!; isotypes, B!, GH!, L, possible isotype, F!; frag., us!; photos, F of B, us oft). Polypodium circinatum Sodiro, Crypt, vase. Quit. 333. 1 893. TYPE: Ecuador, Azuay, near Cuenca, Rim- bach 35 (holotype, Q or QCA). Plants epiphytic or epipetric, rarely terrestrial. Stem short- to long-creeping, the scales narrow- deltate, attenuate, castaneous, lustrous, entire, 2- 4 mm long. Leaves 15-35(-40) cm long, approx- imate or fasciculate, with apex often circinate at apex even at maturity. Petiole 2-8 cm long, dark reddish brown. Lamina firm-herbaceous to char- taceous, elliptic, 3-10 cm broad, gradually nar- TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 119 rowed to apex and base. Rachis reddish brown, scales lacking or rare and filiform, abaxially with scattered acicular (sometimes caducous) tri- chomes 1-1.5 mm long, trichomes scattered adax- ially, 0.4-1 mm long. Segments (at least distal ones) strongly ascending, at a 45-50° angle, most of them with base nearly perpendicular to or rounded at the rachis acroscopically, decurrent basiscopically, the costae and laminar tissue subglabrous or with scattered, acicular trichomes 0.4-0.8 mm long, the margins sinuate and essentially glabrous. Veins indistinct or obscure, 1 -forked, free. Sori medial. Sporangia with 1 or 2 setae about the length of the capsule, these usually deciduous on mature sporangia. In forests and wooded canyons, on tree trunks and branches, or among rocks or on rocky cliffs, rarely on mossy earth, 1600-4000 m, Cajamarca to Apurimac and Puno. Ecuador to Bolivia. Some specimens of Pecluma curvans from Peru, atypical in their scarcely ascending pinnae, may be confused with P. absidata or P. camptophyllaria var. abbreviata. These can be distinguished from the latter by the virtually glabrous pinna margins (vs. margins with conspicuous trichomes) and from both by the much longer (nearly 2 mm) flexuous, abaxial rachis trichomes and by the often circinate leaves (even at maturity). Cajamarca: Cerro Cumbre Mayo, Sanchez V. 35 (GH, us). La Libertad: Prov. Bolivar, Nevado de Cajamar- quilla, Lopez & Sagdstegui 3235 (GH, HUT, MO). Ancash: NW slope of Nevada de Huascaran, below Llanganuco, Correll & Smith P955 (us). Prov. and Dist. Yungay, Llanganuco, Sounders 516 (F). Huanuco: Chasqui, Mac- bride & Featherstone 1949 (F, us). Junin: Huancayo, Quebrada de Occopilla, Soukup 3642 (F, GH, us), 3646 (F), 5884 (GH). Huancavelica: Prov. Tayacaja, Ampurco, Tovar 3763 (GH). Apurimac: Prov. Abancay, Ampay, Tamburco, Nunez & Vargas 7213 (MO, uc). Cuzco: Mol- lepata, above Pincopata, Bishop 2530 (us). Prov. Pau- cartambo, Hacienda Churi, Herrera 271 (GH, us), 7655 (us). Puno: Granja Salcedo, near Puno, Soukup 64 (F, uc). 5. Pecluma divaricata (Fourn.) Mickel & Beitel, Mem. New York Bot. Gard. (Pterid. fl. Oa- xaca) 46: 269. 1988. Polypodium divaricatum Fourn., Mexic. pi. 1: 180. 1872. TYPE: Mexico, Veracruz, Zacuapan, Gal- eotti 6287 (holotype, P; isotypes, B, BR, G). Polypodium bolivianum Rosenst., Repert. Spec. Nov. Regni Veg. 5: 236. 1908. TYPE: Bolivia, South Yungas, Sirupaya, near Yanacachi, Buchtien 481 (holotype, s; isotype, us!). Pecluma boliviano (Rosenst.) Price, Amer. Fern J. 73: 113. 1983. Plants epiphytic (at least in Peru). Stem short- to long-creeping, the scales linear to narrow-del- tate, acuminate or attenuate, orange to dark brown, 2-5 mm long, often remotely ciliate or ciliolate. Leaves (35-)42-130 cm long, approximate to 2 cm apart. Petiole 8-25 cm long, reddish brown. Lamina narrow-delta te, (8-) 1 0-25 cm broad, trun- cate or occasionally with a few proximal segments slightly reduced. Rachis reddish brown, scales lacking or sparse, filiform and inconspicuous, sparsely and minutely pubescent on the abaxial side. Segments perpendicular to the rachis, gla- brous or sparsely and minutely pubescent abaxi- ally, tissue subglabrous adaxially, the costa per- pendicular to the rachis, not or scarcely decurrent. Veins usually obscure, 2-3-forked. Sori mostly su- pramedial, receptacle surrounded by a small clus- ter of setiform trichomes, these obscured when sporangia are mature. Sporangia glabrous. In forests or thickets, on tree trunks, 1000-2400 m, along the Cordillera Central from Amazonas to Cuzco. Southern Mexico to Panama; Venezuela; Co- lumbia to Bolivia. Some specimens of P. eurybasis may be con- fused with P. divaricata in that the laminae are very large and subtruncate and the costae are not or scarcely decurrent on the rachis. However, in the former species the rachis is moderately to densely villous, whereas in P. divaricata it is gla- brous or sparsely pubescent. Bryan 517 (F), from Muna, Dept. Huanuco, is an aberrant form: the lamina base is truncate but with a few greatly reduced pinnae and with costae somewhat decurrent on the rachis. Many abortive sporangia appear among the normal ones, so this is apparently a hybrid. Some characters, including indument, suggest P. pectinata as a putative par- ent, but the specimen was collected at 2150 m, and P. pectinata does not occur in Peru above 790 m. Barbour4179, from Amazonas (MO), is another aberrant form, also with abortive sporangia. Amazonas: Trail E of La Peca in Serrania de Bagua, Gentry et al. 22972 (MO, us). San Martin: Prov. Rioja, Venceremos, D. Smith 4462 (MO). Huanuco: Muna, Bry- an 546 (F), 556 (F). Junin: Huacapistana, Coronado 272 (GH, uc, us). Prov. Satipo, road to La Merced, Leon et al. 184 (MO, USM). Chanchamayo Valley, C. Schunke 118 (us). Pasco: Oxapampa (as Junin), Soukup 2347 (GH). Villa Rica, Woytkowski 7346 (GH, MO, us). Cuzco: San 120 FIELDIANA: BOTANY Miguel, Urubamba Valley, Cook & Gilbert 1752 (us), 7 762 (us). 6. Pecluma eurybasis (C. Chr.) Price, Amer. Fern J. 73: 114. 1983. Plants terrestrial or epiphytic, occasionally epi- petric. Stem usually long-creeping, the scales linear to narrow-deltate, attenuate, light to dark brown, 2-5 mm long. Leaves 25-120 cm long, approxi- mate. Petiole 7-35 cm long, reddish brown. Lam- ina herbaceous to subcoriaceous, 5-22 cm broad, narrow-ovate, abruptly reduced at base, with a few segments V6-'/2 the length of the longest ones. Ra- chis reddish brown, scales lacking or rare and fi- liform, moderately to densely villous, the mostly acicular trichomes 0.2-0.4(-0.5) mm long. Seg- ments perpendicular to the rachis, the base ex- panded symmetrically, or sometimes perpendic- ular to the rachis acroscopically, costae mostly perpendicular to the rachis and sparsely to densely puberulent with acicular trichomes 0.2-0.3 mm long. Veins usually distinct, 1-2-forked, free, gla- brous. Sori medial. Sporangia glabrous. Evans (1969) recognized three varieties of this species: var. eurybasis from the Greater Antilles and Venezuela, and vars. glabrescens and villosa, both from southern Central America to Bolivia. He separated var. glabrescens from the first of these on the basis of its once- (vs. twice-) forked veins. Key to Varieties a. Rachis sparsely to moderately provided with acicular trichomes 0.2-0.3 mm long; stem scales often inconspicuously ciliolate 6a. var. glabrescens a. Rachis densely villous with acicular trichomes 0.2-0.5 mm long; stem scales entire . 6b. var. villosa 6a. Pecluma eurybasis var. glabrescens (Rosenst.) Lell., Amer. Fern J. 74:59. 1984. Polypodium lachniferum var. glabrescens Rosenst., Repert. Spec. Nov. Regni Veg. 11: 57. 1912. TYPE: Bolivia, North Yungas, Unduavi, Buch- tien 2770 (holotype, s; isotype, us!). Polypodium eurybasis var. glabrescens (Rosenst.) Evans, Ann/Missouri Bot. Card. 55: 244. 1969. In wet and rocky soil, 2000-2800 m, Cajamarca. Costa Rica; Venezuela; Colombia to Bolivia. Cajamarca: Prov. Contumaza, Cruz del Hueco, Sa- gdstegui et al. 9926 (HUT, MO, uc). Prov. and Dist. Ca- jamarca, Lluscapampa, Sanchez 373 (GH). 6b. Pecluma eurybasis var. villosa (Evans) Lell., Amer. Fern J. 74: 59. 1984. Figure 6c. Polypodium eurybasis var. villosum Evans, Ann. Mis- souri Bot. Gard. 55: 245. 1969. TYPE: Colombia, Cundinamarca, above Bogota, Fosberg 19688 (holotype, us!; isotype, us!). In forests, thickets, or wooded canyons, on tree trunks, in humus or among rocks, 2000-2750 m, Amazonas to Cuzco. Panama; Venezuela; Colombia to Bolivia. Amazonas: Prov. Chachapoyas, Puma-urcu SE of Cha- chapoyas, Wurdack 555 (us). San Martin: Prov. Mar- iscal Caceres, Rio Abiseo National Park, Young & Leon 4938 (USM). Loreto: Leticia, on Amazon River, LI. Wil- liams 3040 (F). Huanuco: Cani, 7 mi SE of Mito, Mac- bride 3407 (F, us). Muna, trail to Tambo de Vaca, Mac- bride 4280 (F, us). Mito, Macbride & Featherstone 1621 (F, us), 7757 (F, us). Pasco: Prov. Oxapampa, trail to Cordillera Yanachaga via Rio San Daniel, D. Smith et al. 7854 (MO). Cuzco: Quillabamba. Santa Teresa, 0.5 km W of La Playa, Peyton & Peyton 1234 (GH, MO). 7. Pecluma pectinata (L.) Price, Amer. Fern J. 73: 115. 1983. Polypodium pectinatum L., Sp. pi. 1085. 1753. LEC- TOTYPE (designated by Evans, 1969, p. 246): Plumier t. 37, Descr. pi. Amer., based on a spec- imen from Martinique. Goniophlebium pectinatum (L.) John Sm., London J. Bot. 4: 57. 1842. Plants epiphytic or epipetric, occasionally ter- restrial. Stem short- or long-creeping, the scales 3-6 mm long, orange to reddish brown, linear, with an attenuate, often filiform, tip. Leaves 20- 100 cm long, crowded to approximate. Petiole (3-) 4—15 cm long, reddish brown. Lamina firm-her- baceous to chartaceous, 3-9 cm broad, abruptly reduced to a few small lobes, wings, or auricles. Rachis reddish brown, scales lacking or rare and filiform, abaxially minutely puberulent, the tri- chomes 0.5-2 mm long. Segments perpendicular to the rachis, the base expanded symmetrically, TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 121 the margins with ample, trichomes, the costae slightly decurrent on the rachis, the costae, veins and lamina tissue regularly but minutely and in- conspicuously puberulent with trichomes 0.05-0. 1 mm long. Veins indistinct or obscure, 1-2-forked, mostly anastomosing. Sori medial. Sporangia cap- sules each usually bearing a single seta. In deep forests or wooded ravines, on tree trunks or branches, fallen logs, rocks, or occasionally on the forest floor, 150-750 m, Huanuco and Loreto to Cuzco and Madre de Dios. Costa Rica and Panama; West Indies; Vene- zuela; Columbia to Peru. For a long time this name was a catchall for many species ofPolypodium in the Pecluma group, so that it was perceived to be far more common than it actually is. Evans (1969) found it to be a plant of lower elevations, usually under 1 000 m (or lower in Peru). It is characterized by the strong- ly and abruptly reduced lamina, which is regularly though minutely and inconspicuously puberulent. Loreto: Bank of Rio Santiago above Pongo de Man- seriche, Mexia 6212 (F, GH, MO, uc, us). Huanuco (as San Martin): Ridge east of Tingo Maria, Allard 21442 (us). Ucayali (as San Martin): Boqueron Pass, 92 km from Tingo Maria, Allard 21734 (us). Pasco: Prov. Ox- apampa, Palcazu, Rio Alto Iscozacin, Foster et al. 10115 (F). Cuzco: Prov. La Convencion, Rio Apurimac, above Boca de Tigre rapids, Davis et al. 1301 (F, CH). Madre de Dios: Parque Nacional de Manu, Cocha Cashu Sta- tion, Foster & Terborgh 6632 (F). 8. Pecluma venturii (de la Sola) Price, Amer. Fern J. 73: 115. 1983. Polypodium venturii de la Sola, Opera Lilloana 5: 1 86. 1960. TYPE: Argentina, Tucuman, Tafi, Venturi 1232 (holotype, LIL). Plants epiphytic or terrestrial. Stem short- to long-creeping, the scales broad- or narrow-delta te, acuminate or attenuate, sublustrous, reddish brown, 1-2 mm long. Leaves 28-60 cm long, ap- proximate. Petiole 5-12 cm long, reddish brown. Lamina firm-herbaceous to chartaceous, 6-10 cm broad, narrow-ovate, abruptly reduced at base, with a few segments 'A-'/z the length of the longest ones. Rachis reddish brown, scales lacking or rare and filiform, abundantly pilose, the acicular trichomes 0.6-1 mm long abaxially, a little shorter adaxially Segments perpendicular to the rachis, the base ex- panded symmetrically, the margins with ample, minute, trichomes, the costae perpendicular to or somewhat decurrent on the rachis and amply pro- vided on both sides with acicular trichomes 0.4- 0.8 mm long. Veins distinct or indistinct, 1-2- forked, free, sparsely to moderately provided with acicular trichomes 0.4-0.8 mm long. Sori medial or inframedial. Sporangia glabrous. In and at edges of forests, on trunks or branches of trees, on fallen logs or in rocky soil, 2300-3600 m, Lambayeque, Cajamarca, San Martin, Huan- uco, and Cuzco. Ecuador; Peru; Argentina. Lambayeque: Prov. Ferrenafe, 7 km NW of Incahuasi, Dillon & Skillman 41 1 1 (F, GH, uc). Cajamarca: Colasay, Woytkowski 7013 (MO, us). San Martin: District Hual- laga, Valley of Rio Apisoncho 30 km above Jucusbamba, Hamilton & Holligan 516 (us). Huanuco: Mito, Bryan 191 (F). Pampayacu, Kanehira 183 (us). Cuzco: Prov. La Convencion, Garavito, Hacienda Potrero, Vargas 13214 (GH). 9. Pecluma camptophyllaria (Fee) Price, Amer. FernJ. 73. 113. 1983. Plants epipetric, epiphytic, or terrestrial. Stem short- to long-creeping, the scales 2-5 mm long, light to dark reddish brown, linear to narrow-del- tate, the apex attenuate and often with a filiform tip. Leaves 16-150 cm long, crowded to approx- imate. Petiole 2-20 cm long, reddish brown. Lam- ina firm-herbaceous to subcoriaceous, attenuate at base, segments (often many of them) gradually re- duced to lobes or auricles. Rachis reddish brown, scales essentially lacking, abaxially densely pilose with trichomes 0.4-0.8 mm long, these adaxially a little smaller. Segments perpendicular to the ra- chis, the base usually expanded symmetrically or, if sometimes asymmetrical then the basiscopic edge perpendicular to the rachis, the costae decurrent on the rachis, the costae, veins and laminar tissue conspicuously pilose with trichomes 0.2-0.5 mm long. Veins indistinct to obscure, 1-2-forked, free to partly anastomosing. Sori medial to suprame- dial. Sporangia with 1-3, often deciduous, cap- sular setae. Of the four varieties recognized by Evans ( 1 969) only var. macedoi (Bolivia and southern Brazil) does not occur in Peru. The latter is distinguished by forked laminar trichomes and fully anasto- mosing veins. The other varieties have simple tri- chomes, and veins are free to only partly anasto- mosing. 122 FIELDIANA: BOTANY Key to Varieties a. Leaves 1 6-30 cm long; petiole 2-5(-7) cm long; veins 1 -forked, free 9c. var. abbreviate a. Leaves 30-1 50 cm long; petiole 6-20 cm long; veins 2-forked, partly anastomosing b b. Trichomes of rachis, costae, and laminar surface copious, all about the same density and length (ca. 0.5 mm) 9a. var. camptophyllaria b. Trichomes of rachis and costae moderate and to 0.8 mm long, noticeably longer and much more abundant than those of the laminar tissue 9b. var. lachnifera 9a. Pecluma camptophyllaria var. camptophyllar- ia. Cuba and Jamaica: Venezuela: Colombia to Bo- livia. Polypodium camptophyllarium Fee, Mem. foug. 8: 86. 1857. TYPE: Colombia, north of Santander, near Ocana. Schlim 128 (holotype, P!; isotypes, BR, G, K.!; photos, F of P, us of K). In forests, on trees or rocks, or on the wet forest floor, 1800-3200 m, San Martin, Huanuco, Junin, Apurimac to Puno. Costa Rica and Panama; Greater Antilles; Co- lumbia and Venezuela to Bolivia and Brazil. This variety is distinguished by the dense and evenly disposed trichomes of the axes and lamina. It and the conspicuously smaller var. abbreviata are sparsely represented in scattered locations in Peru, whereas var. lachnifera is far more common and widespread. Trichomes of the latter are coarse, of moderate abundance and of irregular length, often to 0.8 mm on rachis and costa, but half as long and much more scattered on the laminar tis- sue. San Martin: Dist. Huallaga. valley of Rio Apisoncho, Hamilton & Holligan 910 (us). Huanuco: Hwy. from Chinchao to Durand, Coronado 91 (uc, us). Mima, Woytkowski 5238 (GH, MO). Junin: Yaupe, Woytkowski 6489 (MO). Apurimac: Chirhuai, Quebrada Matara, Var- gas 2299 (GH). Cuzco: Urubamba River, Machu Picchu, Heller 2206 (us), Herrera 3479 (us). Puno: Prov. Sandia, near Sandia, Vargas 14806 (GH). 9b. Pecluma camptophyllaria var. lachnifera (Hi- eron.) Lell., Amer. Fern J. 74: 59. 1984. Polypodium camptophyllarium var. lachniferum (Hi- eron.) Evans, Ann. Missouri Bot. Gard. 55: 254. 1969. Polypodium lachniferum Hieron., Bot. Jahrb. Syst. 34: 515. 1904. LECTOTYPE (designated by Evans, 1969): Ecuador, Mt. Tungurahua, Lehmann 458 (B; isolectotypes, LE, us!). In forests, clearings, and wooded canyons, on trees, rocks, or rocky slopes, 1000-2900 m, Ca- jamarca and Amazonas south to Ayacucho and Puno. Cajamarca: Prov. Cutervo, Cutervo-Socota, Lopez & Sagdstegui 5333 (GH, HUT, MO); Prov. Celendin, Can- tange, Sagdstegui et al. 8448 (F, HUT, MO, uc). Amazonas: Prov. Chachapoyas, road Palmira-Leimebamba, Fer- nandez 78 (NY, USM). Prov. Chachapoyas, Cerros Calla Calla, San Miguel, Hutchison & Bennett 4544 (F, GH, uc, us). Huanuco: Muna, Macbride3968 (F, us), Woytkowski 5217 (GH, MO). Pasco: Prov. and Dist. Oxapampa, Rio San Alberto, Foster et al. 10284 (F). Junin: Prov. Tarma, 5 km SW of Huacapistana, Tryon & Tryon 5426 (GH, us). Ayacucho: Ccarrapa, between Huanta and Rio Apu- rimac, Killip & Smith 22473 (us). Cuzco: Quillabamba. Santa Teresa, Mandornilloc, Peyton & Peyton 1276 (GH). Puno: Prov. Sandia near Sandia, Soukup & Lopez 14762 (GH). 9c. Pecluma camptophyllaria var. abbreviata (Evans) Stolze, comb. nov. Polypodium camptophyllarium var. abbreviatum Evans, Ann. Missouri Bot. Gard. 55: 256. 1969. TYPE: Peru, Cuzco, Prov. La Convencion, Dist. Santa Ana, Hacienda Echarabi, Herrera 872 (ho- lotype, us!). In forests or in clearings, or rocks or rocky slopes, 1300-2900 m, Cajamarca, Huancavelica, and Cuzco. Endemic. Cajamarca: Prov. Celendin. Canyon of Rio Maranon above Balsas, Hutchison & Wright 5324 (F in part, GH in part, MO, us). Huancavelica: Prov. Tayacaja, Tovar 2015 (GH), 3701 (GH). Cuzco: "Pampakjahue," Coronado 105 (uc, us). 20 km N of Ollantaytambo, Hitchcock 22516 (us). Urubamba Valley, Leon 442 (F, USM). 10. Pecluma ptilodon (Kunze) Price, Amer. Fern J. 73: 115. 1983. Plants epiphytic or terrestrial. Stem long- or short-creeping, the scales narrow-deltate, attenu- ate, light to reddish brown. 2—4 mm long. Leaves 25-150 cm long, crowded to fasciculate. Petiole 1-35 cm long, reddish brown. Lamina firm-her- baceous to subcoriaceous. to 20 cm broad, elliptic TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 123 to narrow-ovate, abruptly or very gradually re- duced at base to mere lobes or auricles. Rachis reddish brown, scales lacking or rare and filiform, sparsely to abundantly pilose with acicular tri- chomes 0.2-1.5 mm long, or these a little shorter adaxially. Segments perpendicular to the rachis, the base (in varieties occurring in Peru) expanded symmetrically, the margins with ample, minute trichomes, the costae (in Peru) perpendicular to the rachis and glabrous or sparsely to amply pu- berulent or pilose. Veins distinct or obscure, 1-3- forked, free, glabrous or with a few, scattered, acic- ular trichomes. Sori medial. Sporangia with 1-3 setae. The species occurs in the United States (Flori- da); Greater Antilles; Mexico to Honduras; South America. Four varieties were recognized by Evans (1969). Variety robusta (Fee) Lell. (southern South Amer- ica) has the largest leaves, with the basiscopic base of segments perpendicular to the rachis at least in the proximal half of the lamina. In the other va- rieties, segment bases are subequally expanded on both edges. Variety caespitosa (Jenm.) Lell. (Flor- ida, West Indies, Mexico to Honduras) is suppos- edly distinguished from var. ptilodon in subtle dif- ferences in length of leaf, petiole, and spores. The other two varieties occur in Peru. Key to Varieties a. Petiole 8-20 cm long; lamina subtruncate at base, abruptly reduced to a few small lobes or auricles; rachis trichomes 0.2-0.3 mm long lOa. var. ptilodon a. Petiole l-3(-8) cm long; lamina attenuate at base, many segments very gradually reduced to auricles; rachis trichomes (at least abaxially) to 1 .5 mm long lOb. var. pilosa lOa. Pecluma ptilodon var. ptilodon. Polypodium ptilodon Kunze, Linnaea 9: 42. 1834 (as "ptiloton"). TYPE: Peru, San Martin Pampaya- cu, Poeppig in July 1 829 (holotype, LZ, destroyed; isotype, B!, designated as lectotype by Evans, 1 969; photo, F of B). In dense forests, 1000-1700 m, San Martin, Huanuco, and Junin. Peru; Bolivia. Huanuco: Prov. Huanuco, Dist. Churubamba, Cotir- arda, Mexia 8220a (GH, uc). Prov. Huanuco, Puente Durand, Stork & Norton 9571 (uc). Junin: Schunke Ha- cienda, above San Ramon, C. Schunke A-l 51 (us). La Merced, Chanchamayo, C. Schunke 23 (GH, uc, us). Chanchamayo Valley, C. Schunke 126 (us). 1 Ob. Pecluma ptilodon var. pilosa (Evans) Stolze, comb. nov. Polypodium ptilodon var. pilosum Evans, Ann. Mis- souri Bot. Gard. 55: 259. 1969. TYPE: British Guiana (Guyana), Demerara, Essequibo River, Jenman (NY). On trees in forests, 130-1 100 m. Amazonas to Cuzco and Madre de Dios. Trinidad: Guyana: Venezuela; Peru; Bolivia; Brazil. Amazonas: Prov. Bagua, Cerro Tapur on Rio Utcu- bamba, Hutchison 1470 (GH, uc). San Martin: San Mar- tin, km 28 of Tarapota-Yurimaguas Road, Knapp & Mallet 8373 (MO). Loreto: Prov. Maynas, Yanamono Tourist Camp, van der Werffet al. 9890 (MO, uc). Huan- uco (as San Martin): 10 km from Tingo Maria on road to Lima, Allard 21541 (us). Prov. Huanuco, Tingo Ma- ria, along Rio Huallaga, Tryon & Tryon 5334 (GH, us). Pasco (as Junin): Pichis Trail, Santa Rosa, Killip & Smith 26201 (us). Prov. Oxapampa, Quebrada Castilla, on Rio Omaiz, Leon & Young 1066 (USM). Ucayali (as Loreto): Coronel Portillo, Pucallpa-Lima Hy., McDaniel 13963 (GH). Cuzco: Prov. Paucartambo, Cosnipata Valley, Rio Tono, Wachteret al. 181 (F). Madre de Dios: Prov. Manu, Cerro de Pantiacolla, Rio Palotoa, Foster et al. 10716 (F). 11. Pecluma hygrometrica (Splitg.) Price, Amer. Fern J. 73: 115. 1983. Figure 6a. Polypodium hygrometricum Splitg., Tijdschr. Natuurl. Gesch. Physiol. 7: 409. 1840. TYPE: Surinam, Para, Splitgerber 1069 (holotype, L, isotype, us; photo, us of L). Plants epiphytic or epipetric. Stem short- to long- creeping, the scales linear to narrow-deltate, acu- minate or attenuate, orange to reddish brown, 2- 4 mm long, entire to remotely and inconspicuously papillate or denticulate. Leaves 27-40(-50) cm long, approximate. Petiole 5-12 cm long, reddish brown. Lamina narrow- or ovate-deltate, 6-10 (-12) cm broad, truncate or occasionally a few proximal segments slightly reduced. Rachis red- dish brown, scales lacking or inconspicuous and filiform, amply but minutely pilose with silvery 124 FIELDIANA: BOTANY trichomes abaxially. Segments perpendicular to the rachis, lamina tissue silver-pilose on both sides, the costa decurrent on the rachis. Veins usually obscure, l(-2)-forked. Sori mostly medial. Spo- rangia setose, the setae often deciduous at matu- rity. In montane or lowland forests, on trees or rocks, 100-3200 m, Loreto, Junin, Cuzco, and Madre de Dios. Mexico to Panama: Guyana to Colombia, south to Bolivia. In addition to the characters used in the key, this can be further distinguished from P. divaricata by the smaller leaves that are more crowded on the stem and by the amply but minutely silver- pilose lamina tissue abaxially. The laminar tissue in P. divaricata is glabrous or sparsely provided with minute, brownish trichomes. Loreto: Prov. Maynas, 50 mi downriver from Iquitos, Moran 3655 (F, MO). Lower Rio Huallaga basin between Yurimaguas and Balsapuerto, Killip & Smith 28164 (F, us). Junin: Rio Pinedo, N of La Merced, Killip & Smith 23608 (us). Prov. Satipo, road to La Merced, Leon 184 (F). Cuzco: Prov. Calca, Lares, Marin 2309 (F). Madre de Dios: Parque Nacional de Manu, Cocha Cashu Sta- tion, M. Foster P-84-2 (uc). Comments Pecluma dispersa (Evans) Price, Amer. Fern J. 73: 114. 1983. Polypodium dispersum Evans, Ann. Missouri Bot. Gard. 55: 235. 1969. TYPE: United States, Flor- ida, Citrus County, Pineola Grottoes, Evans 2008 (holotype, MICH; isotypes, TENN, us!). Although this species has been cited by Evans as occurring throughout the Neotropics, including Ecuador, Brazil, and Bolivia, it has not yet been found in Peru. It scarcely differs from P. plumula, under which may be seen further discussion. Polypodium lomariiforme Kunze, Linnaea 9: 42. 1834. TYPE: Peru (Huanuco), dry mountains of Cassapi, Poeppigin 1829, Diar. 1152 (not located). Kunze's description is insufficient to place this name, and the type has not been located. The de- scription, insofar as it goes, seems to agree with that of Pecluma camptophyllaria var. lachnifera. If this is correct, the name of Polypodium loma- riiformewou\d have priority as a species, but with- out examining the type, nothing further can be determined. For further discussion of the question, see Evans (1969, p. 265). III. Polypodium Polypodium L., Sp. pi. 1082. 1753; Gen. PI. ed. 5, 485. 1 754. TYPE: Polypodium vulgare L. Fig- ure 7. Marginaria Bory, Diet. Class. d'Hist. Nat. 6: 587. 1824. Species listed in 10: 176. 1826. TYPE: Marginaria polypodioides (L.) Tidestr. (Acrosti- chum polypodioides L.) = Polypodium polypo- dioides (L.) Watt. Phlebodium (R. Br.) John Sm., J. Bot. (Hooker) 4: 58. July 1 84 1 . Polypodium section Phlebodium R. Br., PL Jav. Rar. (Bennett & Brown; Horsefield) 4. 1838. TYPE: Polypodium aureum L. (Phlebodi- um aureum (L.) John Sm.). Lepicystis (John Sm.) John Sm., London J. Bot. 1: 195. 1842. Goniophlebium section Lepicystis John Sm., J. Bot. (Hooker) 4: 56. 1841. TYPE: Gon- iophlebium incanum John Sm. Polypodium in- canum Sw., nom. superjl. ) (Lepicystis incana John Sm., nom. nud.) = Polypodium polypodioides (L.) Watt. Terrestrial, rupestral or epiphytic. Stem short- to long-creeping (to 5 m or more), moderately stout to slender, bearing peltate-based scales. Petiole ar- ticulate. Leaves monomorphic to somewhat di- morphic (the fertile longer than the sterile), borne in a cluster to widely spaced. Lamina pinnatifid, pinnatisect, or 1 -pinnate, rarely to 2-pinnate-pin- natified or 3-pinnate, or (in P. laevigatum) entire, glabrous, pubescent, or sparsely to densely scaly. Veins free to anastomosing with or without free included veinlets. Sori round or roundish, borne at the tip of a vein, rarely back of the vein tip, or at the junction of veins, on a usually slightly to moderately raised receptacle, paraphysate or not, exindusiate. Spores ellipsoidal, monolete, the lae- sura '/2-2/3 the length, often with a low, verrucate, tuberculate, or papillate surface, sometimes with more or less winglike folds. The leaves of Polypodium are articulate, leaving a clean scar above the base of the petiole. The petiole below the scar is sometimes called a phyl- lopodium. The veins may be free or variously anastomosing and the sori are borne at the tip of a veinlet, rarely back of the vein tip, or at the junction of veins. Paraphyses are not conspicuous in Polypodium. They are usually filamentous, TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 125 126 FIELDIANA: BOTANY sometimes branched, or they may be expanded at the apex, or they may resemble sporangia. Some species such as Polypodium pycnocarpum have the immature son covered by scales. These are not considered to be paraphyses because they are at- tached at the periphery of the receptacle rather than on the receptacle. Nectaries may be present in several species of Polypodium— for example, in P. thyssanolepis (Koptur et al., 1982). A dark-col- ored sheath around each meristele is reported for P. loriceum, P. subandinum, P. dasypleuron, P. fraxinifolium, P. adnatum, P. caceresii, and P. lae- vigatum and sometimes in P. lasioipus (Hensen, 1990). In the key and descriptions, the term pinna is used for a primary segment that is sessile or stalked. The term pinna-segment is used for a primary segment that is adnate to the rachis or basally joined to other primary segments (the lamina pin- natifid). There are many problems in this genus that can only be resolved by an adequate monographic treatment of the Andean species. References HENSEN, R. V. 1990. Revision of the Polypodium loriceum-comp\e\ (Filicales, Polypodiaceae). Nova Hedwigia, 50: 279-336. KOPTUR, S., A. R. SMITH, AND I. BAKER. 1982. Nectaries in some Neotropical species of Poly- podium. Biotropica, 14: 108-113. MAXON, W. R. 1916a. Polypodium furfur aceum and related species. Contr. U.S. Natl. Herb., 17: 557-579. . 1 9 1 6b. Polypodium squamatum and its allies. Contr. U.S. Natl. Herb., 17: 579-596. SOTA, E. R. DE LA. 1 966. Las especias escomosas del genero " 'Polypodium squamatum.'''' Mus. La Plata Sec. Bot., 10: 60-186. Key to Species of Polypodium a. Lamina irregularly lobed or pinnatifid, at least in part two hybrids, see Comments. a. Lamina simple and entire, the margins even or essentially so; stem very slender and long-creeping; scales at the apex of the stem obtuse 12. P. laevigatum a. Lamina regularly deeply pinnatifid or 1-3-pinnate b b. Sorus often at the junction of 2 or 3 veins; scales of the stem apex reddish brown to light brown, elongate, concolorous, not clathrate; lamina margin cartilaginous, usually notched c c. Sori in 1 or 2 rows between the costa and the margin; lamina often glaucous abaxially; primary lateral veins absent or not prominent abaxially where all veins are equally prominent or nearly so 13. P. aureum c. Sori in 4 to 6 rows between the costa and the margin; lamina light green abaxially; primary lateral veins prominently raised or abaxially rather so 14. P. decuman u m b. Most or all sori on 1 vein; scales of the stem apex brown, or darker d d. Lamina 2-pinnate-pinnatifid to 3-pinnate at the base 23. P. monosorum d. Lamina deeply pinnatifid to 1 -pinnate-pinnatifid e e. Sori in 3 or more rows between the costa and the margin or predominantly so f f. Primary lateral veins connected by obvious transverse veins, these forming several are- olae between the costa and the margin; areolae mostly with 2-4 sori between adjacent transverse veins 9. P. fendleri f. Transverse secondary veins evident, obscure, or absent; areolae with 1 sorus between transverse veins g g. Scales of the stem apex dull or shining, not iridescent h h. Scales of the stem apex obtuse to acute, rarely acuminate; scales mostly appressed to the stem; stem long-creeping 8. P. fraxinifolium h. Scales of the stem apex acuminate to attenuate, sometimes with a filiform tip i FIG. 7. Polypodium decumanum: a, portion of fertile pinna. Polypodium aureum: b, portion of fertile pinna. Polypodium loriceum: c, habit; d, portion of fertile pinna; e, petiole base and part of stem. Polypodium triseriale: f, portion of fertile pinna. (Adapted from Stolze, Ferns and Fern Allies of Guatemala, 1981.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 127 i. Pinnae, pinna-segments, and rachis glabrous; scales of the stem apex acuminate to long-acuminate; stem short-creeping, proximal pinnae usually sessile to slightly adnate 7. P. triseriale i. Pinna-segments and rachis pubescent (may be slightly so) adaxially and abax- ially, or at least the segments pubescent on the costa and margins; scales of the stem apex with a filiform tip; stem long-creeping and proximal pinnae somewhat adnate 10. P. adnatum g. Scales of the stem apex iridescent, spreading, light brown to brown, or rarely blackish, the cell walls thin, many of the scales, especially of the stem apex, with a long, filiform tip; distal pinnae sessile or slightly adnate 11. P. caceresii e. Sori in 1 or 2 rows between the costa and the margin, or predominantly so j j. Pinnae and pinna-segments with scales few or lacking abaxially, veins readily visible because they are raised, or the tissue thin and the veins readily visible with transmitted light k k. Scales of the stem apex concolorous, uniformly colored 1 1. Veins free or a few casual anastomoses present 1. P. sororium 1. Veins regularly anastomosing, at least along the costa m m. Stem very slender, lamina pinnatisect, proximal pinna-segments fully adnate; stem apex scales dark brown, hardly clathrate, narrow ... 4. P. subandinum m. Stem rather stout, lamina rarely pinnatisect, usually 1 -pinnate; proximal pinna- segments short-stalked to sessile or slightly adnate; stem apex scales brownish, clathrate, broad 6. P. sessilifolium k. Scales of the stem apex more or less bicolorous, with a darker center and lighter, thinner margins; the stem and apex sometimes nearly devoid of scales n n. Pinnae mostly or all sessile, or if pinna-segments adnate, then these broadest at about the center o o. Distal pinna-segments adnate; scales of the stem apex acuminate to attenuate; stem short-creeping 7. P. triseriale o. Distal pinnae sessile or only slightly adnate; scales of the stem apex obtuse to acute; scales mostly appressed to the stem; stem long-creeping 8. P. fraxinifolium n. Pinna-segments mostly or all fully adnate to the rachis and usually broadest there P p. Scales of the stem apex and the stem roundish, usually few except at the apex; stem very long-creeping; rachis and costae long-pubescent, especially abaxially 5. P. dasypleuron p. Scales of the stem apex and stem obtuse to acuminate; stem short- to long- creeping; rachis and costae glabrous to short-pubescent q q. Stem very long-creeping, often with deciduous scales; lamina rarely short- pubescent and slightly so; veins regularly anastomosing; 1 or 2 rows of sori between the costa and the margin; not on lomas 2. P. loriceum q. Stem short- to rather long-creeping, densely covered by persistent scales; lamina, especially the rachis, glandular-short-pubescent; veins often free dis- tally, or all veins free; 1 row of sori between the costa and the margin; sometimes on lomas 3. P. lasiopus j. Pinnae and pinna-segments densely covered with scales, or the tissue thick and the veins not visible with transmitted light r r. Lamina gradually reduced at the base to small, often auriculiform lobes s s. Pinna-segments abundantly scaly abaxially, the scales reddish brown 21. P. bombycinum s. Pinna-segments sparingly scaly abaxially, the scales whitish . . 22. P. balaonense r. Lamina not or hardly reduced at the base t t. Scales of the stem apex concolorous u 128 FIELDIANA: BOTANY u. Stem short-creeping, the leaves and phyllopodia approximate; pinna-segments abaxially densely scaly with whitish scales; veins free ... 17. P. furfuraceum u. Stem long-creeping, leaves and phyllopodia distant; pinna- segments abaxially more or less scaly, with dark brown scales or the scales with a dark brown center; veins anastomosing 19. P. megalolepis Scales of the stem bicolorous, with a dark center and lighter margins v v. Scales of the stem apex in mass appearing woolly with projecting dark spines, many scales with a narrow, thick, blackish projecting center and thin, broad margins; leaves usually dimorphic, the fertile often taller than the sterile .... 16. P. remotum v. Scales of the stem apex closely appressed to spreading, the tip attenuate to filiform, the darker center not projecting, the tips brown or lighter, the lighter margins narrow to moderately broad w w. Stem very slender, ca. 1 mm wide dry; sori immersed in the leaf tissue; veins anastomosing, at least in part 18. P. polypodioides w. Stem stouter, ca. 2 mm or more wide dry; sori superficial; veins free or anastomosing x x. Margin of the pinna-segments notched, especially distally; adaxial side of the pinna-segments glabrous; veins free 15. P. pycnocarpum x. Margin of the pinna-segments entire, not notched; adaxial side of the pinna-segments scaly, rarely glabrous; veins anastomosing 20. P. thyssanolepis 1. Polypodium sororium Willd., Sp. pi. ed. 4, 5: 191. 1910. TYPE: Venezuela, "Prope Car- ipe," Humboldt 424 (holotype, B!, Herb. Willd. 19684, sheets 1 and 2; photos, GH). Stem short- to long-creeping, leaves approxi- mate to distant. Scales of stem apex light brown to reddish brown, not iridescent, not clathrate, appressed, the apex acuminate to mostly filiform. Lamina pinnatisect to 1 -pinnate, to ca. 100 cm long, monomorphic, usually only somewhat re- duced at the base. Pinnae or pinna-segments gla- brous or somewhat scaly or short-pubescent abax- ially, the margins notched or not, the distal ones adnate at the base and margins notched or not, the distal ones adnate at the base and broadest there, the proximal ones sessile to adnate. Veins free, 2-3-forked, visible. Sorus at the end of a vein, 1 row of sori between the costa and margin. In forests, terrestrial (one collection epiphytic), 350-1500 m, San Martin, Huanuco, and Pasco. Mexico and Central America; West Indies; northern South America; Colombia south to Peru. This free-veined species has commonly been called P. dissimile L. However, according to Lei- linger (Amer. Fern J: 70: 30. 1980) that species has anastomosing veins and is the correct name for what has usually been called P. chnoodes Sprengel. The earliest name for the present species is P. sororium Willd. San Martin: Prov. Mariscal Caceres, Tocache Nuevo, J. Schunke V. 6944 (MO). Huanuco: Sinchono, Aquilar 879 (USM). Pasco: San Nicolas, Pichis Trail, Killip & Smith 260122 (us, USM). 2. Polypodium loriceum L., Sp. pi. 2: 1086. 1753. LECTOTYPE (by Proctor, Ferns of Jamaica, 529. 1985): Petiver, Pterigraph. Amer., /. 7, f. 10 (redrawn from Plumier, Traite Foug. Amer., /. 78), from Martinique. Figures 7c- e. Polypodium latipes Langsd. & Fisch., PI. voy. Russes monde(Icon. fil.) 1: 10, 1. 10. 18 10. TYPE: Brazil, "Insula St. Catharina," Langsdorff(ho\olype, LE). Polypodium laetum Raddi, Opusc. Sci. 3: 287. 1819. TYPE: Brazil, Raddi (holotype, F), not Salisb. Prod. 403. 1796. Polypodium chartaceum Baker, J. Bot. 15: 166. 1877. TYPE: Ecuador, Andes of Quito, Volcan el Cor- azon, 3200-3400 m, Sodiro (holotype, K; frag, and photo, us!). Stem long-creeping, the leaves mostly distant, scales appressed or sometimes spreading, or the stem naked. Scales of the stem apex bicolorous, brown, dark in the center where clathrate (the cell walls thicker), the margins thin, lighter in color, not iridescent, apex obtuse to acuminate. Lamina pinnatisect, to ca. 80 cm long, monomorphic, not reduced at the base or slightly so. Pinna-segments adnate, broadest at the base, the margin smooth (not notched), or slightly so (crenulate), especially TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 129 toward the apex, glabrous abaxially or slightly scaly, or short- to long-pubescent. Veins visible, anas- tomosing, at least along the costa or a few free. Sorus at the end of a vein, 1-2 rows of sori between the costa and margin. Epiphytic or rarely terrestrial, 300-3300 m, usu- ally ca. 2000 m, Amazonas to Cuzco and Madre de Dios. Mexico and West Indies; Central America; Ven- ezuela and Colombia; south to Argentina; east to southeastern Brazil. Polypodium laetum Raddi of southeastern Bra- zil is close to or the same as P. loriceum, its char- acters falling within the variation of that species at least when interpreted broadly. Polypodium la- tipes is a less certain synonym. Hensen (1990) treated it as a widespread species (including Peru), whereas Sehnem (Fl. Illust. Catar. Polypodiaceae) restricted its range to southeast Brazil and British Guiana. Amazonas: Serrania de Bagua, Gentry el al. 23060 (F, uc). Prov. Chachapoyas, Quebrada Molino, Wurdack 765 (GH, us). San Martin: Prov. Rioja, Pedro Ruiz- Moyobamba, D. Smith 4414 (MO, USM). Prov. Mariscal Caceres, Dist. Tocache Nuevo, J. Schenke V. 4656 (us). Huanuco: Tingo Maria (as San Martin), Allard 22300 (GH). Rio Llullapichis watershed, ascent of Cerros del Sira, Dudley 13372 (GH, MO, us), 13560 (GH). Prov. Huanuco, Chinachao, Ferreyra 16981 (GH, us, USM). Pasco: Oxapampa (as Junin), Soukup 2365 (GH, us), 3358 (GH, us). Junin: Huacapistana, Killip & Smith 24499 (F, GH, us). La Merced, Soukup 1086 (F). Ucayali: Prov. Coronel Portillo, Aguaytia, J. Schunke V. 5457 (F, us). Ayacucho: Prov. La Mar, Cordillera Central, Dudley 12029 (GH). Cuzco: Prov. Urubamba, Machu Picchu, Peyton & Peyton 1300 (GH, MO); Prov. La Convention, Huayopata, Peyton & Peyton 1389 (GH, MO). Madre de Dios: Prov. Manu, Cerro de Pantiacolla, Foster et al. 10685 (F), 10753 (F). 3. Polypodium lasiopus Klotzsch, Linnaea 20: 393. 1 847. TYPE: Venezuela, Colonia Tovar, Mo- ritz 256 (holotype, B; isotypes, BM, p.). Polypodium pubescens Hooker & Grev., Icon, fil., t. 182. 1830, not L. 1759. TYPE: Argentina, Cerro del Morro, San Luis, Gillies (holotype, K). Polypodium gilliesii C. Chr., Ind. fil. 529. 1906. nom. nov. for Polypodium pubescens Hooker & Grev., not L., 1759, and with the same type. Polypodium lasiopus f. bipinnatifidum Kunkel, Nova Hedwigia 3: 379. 1961. TYPE: Chamiseria, "bei Huancayo," Peru, Kunkel (holotype, Herb. Kun- kel). Polypodium lasiopus var. weberbauerianum Kunkel, Nova Hedwigia 3: 380. 1961. Chamiseria, "bei Huancayo," Peru, Kunkel (holotype, herb, not given). Stem somewhat short- to moderately long- creeping, leaves distant or usually rather approx- imate, the scales appressed or the tips spreading. Scales of the stem apex brown, bicolorous, dark brown in the center where clathrate, and cell walls thick, apex of scales acuminate to somewhat fili- form, not iridescent. Lamina pinnatisect or rarely the pinna-segments pinnatifid, to ca. 20 cm long, monomorphic, not reduced at the base or rarely so. Pinna-segments adnate or the proximal ones sessile or slightly adnate, broadest at the base, scales few or none, pubescent abaxially or not, rachis the same, usually short, glandular-pubescent, rarely long-pubescent. Veins visible, free or anastomos- ing. Sorus at the tip of a vein, sori in 1 row between costa and margin. Terrestrial or rarely (especially on lomas) epi- phytic, often among rocks or on cliffs and ledges in forests on stream banks, sometimes on lime- stone, 400-3600 m, usually above 2000 m except lower on lomas; Lambayeque to Cuzco. Andean South America; range uncertain. Lambayeque: Prov. Chiclayo, Cerro Reque, Llatas 340 (GH, HUT, MO). Prov. Lambayeque, Entre Beatita de Hu- may y km 38, Lopez et al. 4043 (GH, HUT). Cajamarca: Between Llacanora and Namora, Correll & Smith P882 (GH, MO). Prov. Contumaza, Pampa de Guzmango, Sa- gdstegui 1 1440 (F, GH, MO). La Libertad: Prov. Trujillo, Cerro Campana, Sagdstegui & Kobats 9213 (F, MO). Sa- gdstegui et al. 12956 (F, GH, MO). Ancash: Lomas de Lupin, Velarde 4441 (GH). Huanuco: Mito, Macbride & Featherstone 1707 (F), Bryan 194 (F). Muna, Bryan 418 (F). Lima: Loma de Lachay (Luchay), Tryon & Tryon 5418 (F, GH, us), Ferreyra 197 (USM). Atacongo Lomas, Stork et al. 9283 (GH, us). Lomas of Granados, Stork & Vargas 9338 (GH, us). Junin: Huancayo, Soukup 2966 (F). Prov. Tarma, Huacapistana, Tryon & Tryon 5455 (GH, us). Mato ad Yaupi, Woytkowski 6543 (MO, us). Ayacucho: Pampalca, between Huanta and Rio Apuri- mac, Killip & Smith 22263 (GH, us). Cuzco: Urubamba Valley, Leon 447 (F, USM). 4. Polypodium subandinum Sodiro, Crypt, vase. Quit. 348. 1893. TYPE: "Corazon, Pasochoa, Pichincha, etc.," 3000-3500 m, Ecuador, So- diro (holotype, ?). Stem very slender and long-creeping, more or less naked or somewhat scaly, the scales mostly appressed with their tips spreading. Scales of the stem apex hardly clathrate, dark brown, concol- orous, not iridescent, the apex attenuate to fili- form. Lamina pinnatisect, to ca. 20 cm long, monomorphic, not reduced at the base, or only slightly so. Pinna-segments adnate, broadest at the 130 FIELDIANA: BOTANY base, margins smooth, rather sparingly to densely long-pubescent abaxially, the rachis similar. Veins visible, anastomosing at least along the costa. So- rus at the end of a vein, 1 row of sori between costa and margin. Terrestrial, in forests, 200-3000 m, Amazonas, San Martin, and Pasco. Andes; range uncertain. The pubescence on the abaxial surface of the pinna-segments may be similar to that of P. la- siopus and P. dasypleuron. However, the stem is very slender and the scales of the stem apex are narrow and concolorous. Amazonas: Prov. Chachapoyas, Cerros Calla Calla, 1 8 km above Leimebamaba on road to Balsas, Hutchison & Wright 4872 (F, GH). San Martin: Dist. Huallaga, Hamilton & Holligan 1202 (us). Pasco: Pichis Trail, San Nicholas to Azupizu, Killip & Smith 26098 (us). 5. Polypodium dasypleuron Kunze, Linnaea 9: 43. 1834. TYPE: Peru, Pampayaco (Pampayacu), Poeppig, Diar. 1111 (holotype, LZ, destroyed; isotype, ? Poeppig in 1829, K; frag. us!). Stem long-creeping, leaves distant, scales usu- ally lacking, or a few appressed ones present. Scales of the stem apex mostly few, clathrate in the center where the cell walls are thicker, margins lighter, narrow, not iridescent, apex obtuse. Lamina to 50 cm long, pinnatisect, monomorphic, slightly re- duced at the base, or not. Pinna-segments adnate, broadest at the base, margins smooth (not notched), sparsely to rather densely long-pubescent, es- pecially on the costa abaxially, hardly scaly. Veins readily visible, anastomosing at least along the costa, some veins may be free. Sorus at tip of a vein, 1 row of sori between the costa and the mar- gin. Epiphytic, rarely on rotten wood, in dense for- ests, rarely in inundated forests, 200-1800 m, Amazonas and Loreto to Madre de Dios and Cuz- co. Costa Rica south to Peru. San Martin: Prov. Mariscal Caceres, Tocache Nuevo, J. Schunke V. 4656 (F, MO, USM); 6942 (F, MO). Loreto: Above Pongo de Manseriche, Mexia 6142 (F, GH, MO, us). Valencia, Rio Corrientes near Platanoyacu, Mc- Daniel and Marcos 11162 (F, GH, MO). Huanuco: Tingo Maria, Tryon & Tryon 5284 (F, GH, us, USM). Pasco: Prov. Oxapampa, vicinity of Chequitavo, D. Smith 5167 (MO). Junin: Chanchamayo Valley, C. Schunke 127 (F). Cuzco: Prov. La Convencion, above Hacienda Luisiana, Davis et al. 1330 (F, GH). Madre de Dios: Prov. Manu, Cerro de Pantiacolla, Foster et al. 10715 (F). 6. Polypodium sessilifolium Desv., Mem. Soc. Linn. Paris 6: 238. 1827 TYPE: Herb. Des- vaux, P; photo, us of p. (spelling corrected from on the label of the type, see Weath., Contr. Gray Herb. 114: 33. 1936). Polypodium cordatum Kunze, Linnaea 9: 44. 1834. TYPE: "ad Cassapi," July 1829. Poeppig (Diar. 1184), not Desv. 1827. Polypodium surucuchense Hooker, Hooker's Icon. 1 : /. 69. 1837. TYPE: Surucucho, near Cuenca "on trunks, Colombia" (Ecuador) Jameson (holotype, K). Polypodium biauriculatum Hooker, Hooker's Icon. 2: t. 121. 1837. TYPE: Chachapoyas, Peru, Ma- thews in 1836 (holotype, K). Polypodium chacapoyense Hooker, Sp. fil. 5: 29, t. 281. 1863. TYPE: Sesuja, Chachapoyas, Peru, Mathews 3278 (holotype, K). Polypodium kunzeanum C. Chr., Index fil. 536. 1906. Nom. nov. for Polypodium cordatum Kunze, not Desv. Stem rather stout, short-creeping, and the leaves approximate to moderately long-creeping, leaves distant, the scales mostly spreading. Scales of the stem apex brownish, clathrate, concolorous, rarely iridescent, usually not so, the tip acute to some- what filiform. Lamina pinnatisect to usually 1 -pinnate, to ca. 100 cm long, monomorphic, not or hardly reduced at the base. The proximal pinnae or pinna-segments, sessile to short-stalked, the dis- tal ones often slightly adnate to adnate, margins smooth, no scales abaxially or very few, essentially glabrous to usually short- to long-pubescent abax- ially, and sparsely to definitely so. Veins visible, anastomosing at least along the costa. Sorus at the tip of a vein in 1 or 2 (rarely 3) rows between costa and margin. Terrestrial or sometimes epiphytic, in rock crev- ices, on rocks or at the edge of rocks, in moist ravines, in thickets, at the edge of streams, in for- ests, 1300-4270 m, Piura south to Puno. Tropical America; especially northern South America; range uncertain. Piura: Prov. Huancabamba, Cerro La Viuda, Sagds- tegui et al. 8208 (F, MO). Cajamarca: Prov. Cajamarca, Pampa de Guzmango, Sagdstegui 1440A (F, MO). Prov. Celendin, Pumarrume, Mostacero et al. 1030 (HUT, MO). Amazonas: Prov. Chachapoyas, Calla Calla slopes, Wur- dack 1755 (GH, us). La Libertad: Prov. Otuzco, Cerro Ragache, Sagdstegui et al. 1 1609 (F, MO). Ancash: Prov. Yungay, Laguna Llanganuco, Sagastergui et al. 12318 TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 131 (GH, HUT, F). Prov. Recuay, Laguna de Querococha, Lopez el al. 75/9(GH, HUT). Huanaco: Muna, Woytkowski 5180 (GH, MO, us). Junin: Prov. Huancayo, 1 1 km from Huan- cayo toward Chameseria, Sounders 655 (F, GH, us). Cuz- co: Prov. Urubamba, Machu Picchu, Peyton & Peyton 1041 (GH, MO). Prov. Quispicanchi, 10 km northeast of Marcapata, Wasshausen & Encarnacidn 767 (MO, USM). Puno: Prov. Caranaya, Ollachea, Vargas 6910 (MO, us). 1. Polypodium triseriale Sw., J. Bot. (Schrader) 1800(2): 126. 1 80 1 . TYPE: nothing cited. Fig- ure If. Polypodium longifolium Presl, Delic. prag. 1:167.1 822. TYPE: "ad Rio Janeiro" (PR or PRC, not seen). Polypodium preslianum Spreng., Syst. Veg. ed. 16, 5: 556 (index). 1828. Nom nov. for P. longifolium Presl, 1822, not Cav. 1802. Stem rather stout, short- to long-creeping, the leaves approximate to distant, scales more or less appressed. Scales of the stem apex large, with a brown to dark brown, clathrate center, and lighter brown, narrow margins, the apex acute to atten- uate, slightly iridescent or not. Lamina pinnatisect distally or 1 -pinnate proximally, to 90 cm long, monomorphic, not or hardly reduced at the base. Pinna-segments adnate to slightly so, the pinnae sessile, margins not notched, smooth, hardly pu- bescent or scaly abaxially. Veins visible, anasto- mosing. Sorus at the tip of a vein, son in (l-)2-3 (rarely 4) rows between costa and margins. Terrestrial or epiphytic, on or among rocks, in forests, 325-1900 m, probably higher; Tumbes and Amazonas to Cuzco. Tropical America. This is a variable species, especially in the num- ber of rows of sori between the costa and margin. Some specimens seem to be intermediate with P. fraxinifolum. Tumbes: Prov. Zarumilla, Arro de Tres Picos, Simp- son & Schunke 379 (F). Amazonas: West of Molinopam- pa, Wurdack 1407 (F, GH, us). Cajamarca: Prov. Santa Cruz, Monteseco, Sagdstegui et al. 12365 (F, MO). San Martin: Prov. Mariscal Caceres, Pueblo Viejo de To- chache, /. Schunke V. 3718 (F, GH, us). Prov. Lamas, San Juan de Pacaizapa, J. Schunke 9543 (F, MO). Loreto: Rio Tacsha Curaray, Croat 20458 (MO). Huanuco: East of Tingo Maria, Croat 21128 (MO). Pasco: Prov. Oxa- pampa, Palcazu Valley, D. Smith 3723 (MO). Junin: Hu- acapistana, Coronado 252 (GH). La Merced, Soukup 1094 (F). Cuzco: Prov. Paucartambo, Kosnipata, Vargas 1 1323 (GH). Prov. La Convention, Urusaiwa, Vargas 22347 (GH). 8. Polypodium fraxinifolium Jacq., Collectanea 3: 187. 1789. TYPE: "Cult, ex Caracas" (holo- type, w) not Hooker, Sp. fil. 5: 4. 1863. Polypodium appressum Copel., Univ. Calif., Publ. Bot. 10: 305. 1941. TYPE: Bolivia, Dept. La Paz, Prov. Sur Yungas, Colaya, Mexia 7812 (holotype, uc; isotype, GH!). Stem long-creeping or rarely short-creeping, leaves distant, the scales appressed, roundish, or sometimes nearly naked. Scales of the stem apex bicolorous, with a dark brown to blackish, clath- rate center and lighter, thinner margins, obtuse, rarely acuminate. Lamina 1 -pinnate, to 75 cm long, monomorphic, not or hardly reduced at the base. Pinnae sessile or slightly adnate distally, sessile proximally, margins smooth, essentially glabrous abaxially, or rarely moderately pubescent. Veins visible, anastomosing. Sorus at tip of a vein; sori in 1 or 2, usually 3 (rarely 4 or 5), rows between costa and margin. Terrestrial, epiphytic, rarely climbing on a tree trunk, or on rocks, in moist, shady places, in for- ests, 650-2700 m, Cajamarca to Puno. Tropical America. Cajamarca: Colasy, Woytkowski 7015 (GH, MO, us). Prov. Cutervo, San Andres, Quiroz 2720 (F). Amazonas: Prov. Chachapoyas, 5 km below Chachapoyas, Wurdack 629 (GH, us). Prov. Bagua, 1 2 km east of La Peca, Bar- hour 2589 (F, MO). San Martin: Prov. Rioja, Pedro Ruiz- Moyobamba, D. Smith 4362 (MO). Prov. Lamas, 4 km de San Juan de Pacayzapa, J. Schunke V. 6116 (F). Lo- reto: Sierra del Pongo, Mexia 6281 (GH, MO, us). Huanu- co: Tingo Maria, Aguilar 310 (F, GH, us). Pasco: Oxa- pampa, Soukup 3353 (F, GH, us). Junin: Chanchamayo Valley, C. Schunke 77 (F), 794 (F). Above San Ramon, Killip & Smith 24704 (F). Ucayali: Prov. Coronel Potillo (as Loreto), entre Previsto y Boqueron, Aguilar 958 (F, GH). Ayacucho: Ayna, between Huanta and Rio Apuri- mac, Killip & Smith 23180 (F, us). Cuzco: Prov. Uru- bamba, Machu Picchu, Peyton & Peyton 1394 (GH, MO). Madre de Dios: Prov. Manu, Carbon a Salvation, Vargas 16939 (GH, us). Puno: Prov. Carabaya, San Gaban (Ga- ban), Vargas 18915 (GH, us). 9. Polypodium fendleri D. C. Eaton, Mem. Amer. Acad., n.s., 8: 199. 1860. TYPE: Venezuela, Fendler 410 (holotype, YU; isotype, GH!). Campyloneurum magnificum Moore, Index fil. 226. March 1 86 1 . TYPE: Venezuela, Fendler 410 (ho- lotype, K.?; isotype, GH!), nom nov. for Polypodium fendleri D. C. Eaton, not Campyloneurum fendleri Moore. Campyloneurum fendleri (D. C. Eaton) John Sm., Hist, fil. 97. 1 875, not Moore, Index fil. 224, Oct. 1 860. 132 FIELDIANA: BOTANY Stem short- to long-creeping, leaves approxi- mate or distant, scales appressed or spreading. Scales of the apex concolorous, brownish, clath- rate, not iridescent, the apex acute to acuminate. Lamina to 80 cm long (or more), 1 -pinnate gla- brous or with a few scales abaxially, monomor- phic, not reduced at the base, or slightly so. Pinnae 6-10 cm wide, sessile to slightly adnate, the mar- gins smooth, not pubescent abaxially. Veins readi- ly visible, anastomosing. Sorus at the end of a vein; sori in 11-15 rows between the costa and the mar- gin, 2 or usually 3^4 between the main lateral veins. Terrestrial or epiphytic, in dense forest, 1 500- 1 800 m, San Martin, Junin and Pasco. Panama; Venezuela and Columbia south to Bo- livia. This species is closely related to Polypodium decurrens Raddi but seems to be distinct due to a lack of intermediates. It is a larger species than P. decurrens, especially in characters of the stem di- ameter, leaf length, petiole diameter, and pinna width. Polypodium fendleri and P. decurrens are both 1 -pinnate species and have the sorus at the tip of a vein. These characters place the two species in the genus Polypodium. However, some authors prefer to place them in the genus Campyloneurum (Lellinger, Amer. Fern J. 78: 14-35. 1988). Moore (Index fil. Ixxiv-lxxv) recognized that the terminal sori are exceptional in the genus Campyloneurm but considered that the venation was a more im- portant character. San Martin: Tarapoto, 4 mi E of Tarapoto, Woyt- kowski 35218 (MO, uc). Pasco: Oxapampa, Gran Pajonal, trail to S tin m uhuuni from Chequitavo, D. Smith 5212 (MO). Junin: Above San Ramon, C. Schunke Al 70 (GH, us). Chanchamayo Valley, C. Schunke 22 (F), 683 (F), 913 F). La Merced, Soukup 1089 (F). 10. Polypodium adnatum Klotzsch, Linnaea 20: 395. 1847. TYPE: "Merida Colombiae" (Venezuela), Moritz 353 (holotype, B; isotype, BM). Stem rather stout, long-creeping, the leaves dis- tant, scales appressed or at least the tips spreading. Scales at the stem apex narrow, light to dark brown, concolorous, clathrate, the apex acuminate to fi- liform, not or slightly iridescent. Lamina to 45 cm long (or more), 1 -pinnate, monomorphic, not re- duced at the base, or hardly so. Pinnae and pinna- segments sessile to adnate distally, sessile to slight- ly adnate proximally, margin smooth, scales none or very few abaxially, sparsely to rather densely short- to long-pubescent abaxially, especially along the costa. Veins not readily visible, anastomosing. Sorus at the end of a vein, sori in 5-7 rows between costa and margin. Terrestrial or epiphytic, in forests, 250-930 m, Huanuco, Ucayali, Cuzco, and Madre de Dios. Tropical South America; distribution uncertain. This species is a member of a complex that needs careful monographic study. Polypodium richardii Klotzsch and Polypodium giganteum Desv. are members of this complex and may merit recog- nition. Huanuco (as San Martin): east of Tingo Maria, Allard 22233 (GH, us), 223 30 (GH, us), Croat 21158 (MO). Tingo Maria, Tryon & Tryon 5222 (F, GH, us). Ucayali (as Loreto): Vicinity of Aguaytia, Croat 20937 (MO). Cuzco: Prov. Paucartambo, Skog & Skog 5208 (us). Prov. Quis- picanchis, entre 15 mil y San Lorenzo, Vargas 11758 (GH). Prov. La Convention, Cordilla Vilcabamba, Dud- ley 10140 (GH). Madre de Dios: Prov. Tambopata, DistTambopata, Young 160 (F, MO, USM). 11. Polypodium caceresii Sodiro, Crypt., vase. Quit. 360. 1893. TYPE: "Bosques de Oriente en la orilla del Rio Napo, Cdceres" (holotype, ?; isotype, s; photo, GH of s). Stem rather stout, long-creeping, leaves distant, scales appressed. Scales of the stem apex brown- ish, concolorous, clathrate, moderately broad, or brown to dark brown, narrow, attenuate to filiform at the apex, iridescent. Lamina to ca. 75 cm long, 1 -pinnate, monomorphic, not reduced at the base or hardly so. Pinnae or pinna-segments sessile to slightly adnate distally, the proximal pinnae ses- sile, margins smooth, glabrous abaxially or rarely very sparsely short-pubescent not scaly abaxially. Veins not readily visible, anastomosing. Sorus at tip of a vein; sori in 3-7 rows between costa and margin. Terrestrial or usually epiphytic, rarely climbing on a tree trunk, border of woods, riverbanks, cloud forests, primary forests, 100-1 100 m, San Martin to Puno. Colombia to Bolivia; Venezuela east to French Guiana; adjacent Brazil. Polypodium articulatum Desv. (Mem. Soc. Linn. Paris 6: 236. 1827, not Poiret, 1804) is sometimes considered to be a synonym of this species. How- ever, if the published locality "Brasilia" is correct, TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 133 it is more likely a synonym of Polypodium fraxi- nifolium Jacq., as Sehnem (Fl. Illust. Catar. Pol- ypod. 87. 1970) has it. A photograph of the sheet in the Desvaux Herbarium P (GH, us) was consid- ered to be P. caceresii by Weatherby (Contr. Gray Herb. 114: 29. 1936; and 124: 17. 1939). This is evidently correct. San Martin: Prov. Lamas, Lamas, Belshaw 3430 (GH, MO, us). Tarapoto-Yurimaguas, Knapp & Alcorn 7793 (MO). Loreto: Salinas, Rio Mazan, J. M. Schunke 376 (F, GH, us). Mishuyacu, near Iquitos, Klug 2572 (F, us). Huanuco: Prov. Huanuco, Rio Ysabel, Mexia 8 161 a (F, GH, MO, us). Prov. Huanuco, 5 km of Carpish, Tryon & Tryon 5304 (GH). Santa Rosa, Pichis Trail, Killip & Smith 26177 (us). Pasco: Oxapampa, Soukup 2363 (GH). Junin: La Merced, Soukup 1093 (F). Cuzco: Prov. La Conven- cion, Cordillera Vilcabamba, Dudley 10180 (GH). Prov. Paucartambo, Kosnipata-Pilcopata, Vargas 1 1322(GH). Madre de Dies: Prov. Manu, Cerro de Pantiacolla, Foster et al. 10820 (F). Puno: Prov. Carabaya, Inambari, Vargas 16139 (GH). 12. Polypodium laevigatum Cav., Descr. PI. 244. 1802. TYPE: San Antonio, Ecuador, Nee(ho- lotype, MA seen by C. Chr., Dansk Bot. Ark. 9(3): 11. 1937; isotype, s, Herb. Swartz). Orig- inally as levigatum, altered to laevigatum by Swartz, Syn. fil. 28. 1806. Polypodium glaucophyllum Klotzsch, Linnaea 20: 393. August 1847. LECTOTYPE: (here designated): Merida, Venezuela, Moritz 305 (lectotype, B; iso- lectotypes, BM, BR, K). The same name was pub- lished by Kunze, Farrnkrauter 1: 227, t. 93, in September 1847. Stem very slender, long-creeping, leaves mostly distant, naked or a few appressed, roundish scales present. Scales of stem apex roundish to acute, clathrate in the dark brown center, nearly concol- orous, not iridescent. Lamina to ca. 25 cm long, simple, entire, gradually reduced at the apex and the base, or the base abruptly reduced, or rounded, monomorphic, margins not notched, glabrous abaxially or rarely sparingly pubescent or with a few scales. Veins visible, anastomosing. Sorus at the tip of a vein; sori in 2-9 rows between costa and margin. Terrestrial, rupestral or usually epiphytic, in open woods, cloud or fog forests or dense, wet, montane rain forests, 100-2740 m, Piura south to Cuzco. Costa Rica; Guadeloupe; Guianas to Colombia south to Boliva. Campyloneurum repens, which may be similar to this species, has two rows of sori between the primary lateral veins while this species has a single row. The altered spelling, laevigatum, has been ac- cepted since its adoption in 1806. Piura: Prov. Huancabamba, Canchaque-Minas Tur- malina, Sagdstegui et al. 8275 (GH, MO, us). Cajamarca: Prov. Hualgoyoc, Palmito and vicinity, Hutchison & Bis- mark 6399 (F, GH, MO, us, USM); Prov. San Miguel, Taulis Recorco, Mostacero et al. 1174 (F, GH, HUT, MO). Ama- zonas: Serrania de Bagua, Gentry et al. 22906 (F, MO). San Martin: Prov. Mariscal Caceres, Cerro Azul, Tryon & Tryon 5264 (GH, us). Huanuco: Chinchao, entre Hua- nuco y Tingo Maria, Ferreyra 16944 (GH, USM). Yanano, Macbride 3817 (F, GH, us). Pasco: Prov. Oxapampa, km 53 Oxapampa-Paucartambo, Smith & Pretel 1486 (F, MO). Yapas (as Junin), Killip & Smith 25541 (F, us). Junin: Above San Ramon, Killip & Smith 24592 (F). Chanchamayo Valley, C. Shunke 133 (us). Ucayali: Prov. Coronel Portillo, La Divisoria, Dillon 2629A (MO). Aya- cucho: Ayna, between Huanta and Rio Apurimac, Killip & Smith 23140 (F, GH, us). Cuzco: Prov. La Convention, Cordillera Vilcabamba, Dudley 10449 (GH, MO). 13. Polypodium aureum L., Sp. pi. 1087. 1753. TYPE: LINN 1251. 10. Figure 7b. Polypodium leucatomos Poiret, in Lam., Encycl. 5: 516. 1804. TYPE: Cayenne, (French Guiana) Le Blond (holotype, Herb. Lam.) Phlebodium aureum (L.) John Sm., J. Bot. (Hooker) 4: 59. 1841. Stem stout, rather short-creeping, leaves mostly approximate, densely covered with spreading scales. Scales of the stem apex reddish brown to light brown, concolorous, not clathrate, apex fili- form, not iridescent, apex filiform. Lamina to 50 cm long (or more), deeply pinnatifid, monomor- phic, not or hardly reduced at the base. Pinna- segments adnate, margins very slightly notched or not, glabrous abaxially, or hardly scaly. Veins vis- ible, anastomosing. Sorus at the junction of 2 or 3 (rarely 4 or 1 ) veins, sori in 1 or 2 rows between costa and margin. Terrestrial, especially among rocks, or epiphyt- ic, 900-2600 m, Amazonas south to Puno. Tropical America. Phlebodium is often recognized as a genus, but if this is done then it is not obvious where to stop the segregation of Polypodium s.l. Polypodium leu- catomos is a synonym of P. aureum, not an earlier name for P. decumanum as previously thought (Morton, Contr. U.S. Natl. Herb. 38: 63. 1967). Polypodium pseudoaureum Cav. is diploid com- pared to P. aureum, which is tetraploid, and the former is sometimes treated as a separate species. 134 FIELDIANA: BOTANY Cajamarca: Prov. Cutervo, Socota, Mostecero el al. 1751 (HUT). Prov. Cajamarca, Matara-San Marcos, Lopez & Sagastegui 5530 (GH). Ama/onas: Cano Santa Lucia just east of Chachapoyas, Wurdack 717 (GH, us). La Libertad: Prov. Otuzco, Huaranchal, Sagastegui 0204 (GH). San Martin: Zepelacio, Klus 3537 (GH, us). Huanu- co: Prov. Huanuco. road to Gasa, Stork & Morton 9875 (GH, us). Junin: Huacapistana, Aguilar 514 (GH). Aya- cucho: Between Huanta and Rio Apurimac, Killip & Smith 22433 us). Cuzco: Quillabamba-Potrero, Coronado 126 (GH, us). Puno: Prov. Carabaya, San Gaban (Gaban), Vargas 18981 (GH). 14. Polypodium decumanum Willd., Sp. pi. ed. 4, 5: 1 70. 1 8 10. TYPE: "Brasilia," Sieber, comm. Hoffmansegg (B, Herb. Willd.). Figure 7a. Phlebodium decumanum (Willd.) John Sm., J. Bot (Hooker) 4: 59. 1841. Stem short-creeping, rather stout, the leaves more or less approximate, with mostly spreading scales. Scales of the stem apex light brown to red- dish brown, the tip filiform, not clathrate, not iridescent. Lamina to ca. 1 m long (or more), pin- natisect, more or less monomorphic. Pinna-seg- ments adnate, the margins notched, glabrous (not scaly nor pubescent), not glaucous, abaxially, fully adnate but broadest toward the middle. Veins vis- ible, anastomosing. Sorus at the junction of 2 or 3 veins, or son in 4-6 rows between the costa and margin. Epiphytic, rarely terrestrial, on steep banks, in forests, 1 00-1 300 m; Amazonas to Madre de Dios. Central America; West Indies; tropical South America. This is the largest of our species of Polypodium. It was erroneously called P. leucatomos for many years, but that name is now a synonym for Poly- podium aureum. Amazonas: Bagua. Ellenberg 3571 (GH). San Martin: Chazuta, Klug 3983 (GH). Rioja, northwest of San Mar- tin, Soukup 5217 (GH). Loreto: Prov. Maynas, Huaya- bamba, cerca de Iquitos, Ferreyra 3391 (USM). Prov. Maynas, 3 km S of Iquitos, Tryon & Tryon 5210 (GH). Huanuco: Bosque Nacional de Iparia,/. Schunke V. 1799 (GH, us). Ucayali: Pucallpa, Woytkowski 5751 (GH). Ju- nin: Above San Ramon, C. Schunke A 165 (us). Cuzco: Prov. Quispicanchis, Inamburi, Vargas 15441 (GH). Prov. La Convencion, below San Martin, Davis et al. 1326 (GH). Madre de Dios: Maldonado, Ldpez & Soukup 4663 (GH). 15. Polypodium pycnocarpum C. Chr., Index fil. 326. 1905. 557. 1906, nom. nov. for Polypo- dium macrocarpum Presl, not Willd. Polypodium macrocarpum Presl, Reliq. haenk. 1 : 23, /. l,f. 4. 1825, not Willd., Sp. pi. ed. 4, 5: 147. 1810. TYPE: "Hab. in Montanis Peruviae," Haenke (holotype, PR). Stem short- to long-creeping, leaves distant or approximate, scales appressed or spreading. Scales of the stem apex clathrate, dark brown to nearly blackish, with light brown borders, the apex obtuse to filiform, not iridescent. Lamina pinnatifid to pinnatisect, to ca. 15 cm long, scaly abaxially, monomorphic to dimorphic, not reduced at the base. Pinna-segments adnate distally, and adnate to slightly adnate proximally, sometimes pinnat- ifid or partly lobed, the margins notched, scaly abaxially, not pubescent usually fully adnate at the base and broadest there. Veins not readily visible, free. Sorus at the tip of a vein; sori in 1 row be- tween costa and margin. The correct name and classification for this spe- cies depends on a careful review of the consider- able material now present in the larger herbaria. For example, among the species Maxon recog- nized—P. bryopodum Maxon, P. mollendense Maxon, P. rusbyi Maxon, P. subvestitum Maxon, P. tweedianum Hook, and P. xantholepis Harr.— all may be synonyms of P. pycnocarpum. Among other names that require assessment are Pleopeltis pinnatifida Hook. & Grev. (now valid), Polypo- dium ratibori Copel., and P. ruiz-lealii Sola. Key to Varieties of Polypodium pycnocarpum a. Pinna-segments entire, obtuse, leaves monomorphic, lamina usually 5—8 cm long 1 5a. var. pycnocarpum a. Pinna-segments, especially the proximal or basal ones, partly lobed or pinnatifid, acute to acuminate; leaves dimorphic, the fertile taller than the sterile; lamina usually 10-15 cm long . 15b. var. bucht it-nil TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 135 15a. Polypodium pycnocarpum var. pycnocar- pum. Epiphytic, terrestrial or usually rupestral, in rock crevices, on rocky slopes, partly underneath rocks, 400-4100 m, on lomas at 400-800 m, Cajamarca to Puno. Peru; Bolivia; Argentina; Chile; range uncertain. Cajamarca: Prov. Celendin, cerca a Celendin, Ferreyra 15034 (USM). Prov. Contumaza, Cruz Grande Contu- maza, Sagdstegui et at 6477 (F). La Libertad: Prov. Otuzco, Cerro Ragache, Sagdstegui et al. 11611 (F, MO). San Martin: Prov. Mariscal Caceres, Chochos, Young & Leon 4583 (USM). Ancash: Prov. Carhuas, Cordillera Blanca, Chancos, Hutchison & Wright 4370 (F, GH, MO, us). Huanuco: Mito, Bryan 186 (F). Lima: Lomas de Luchay, Tryon & Tryon 5415 (F, GH, us). Lomas de Iquanil (Granado), Dillon et al. 4728 (HUT, USM). Pasco: Prov. Pasco, Entre Salcachupan y Cerro de Pasco, Ferr- eyra 6593 (GH). Junin: Tarma to San Ramon, Croat 57703 (F). lea: Los Cerrillos, 52 km south of Nasca, Rauh (USM). Ayacucho: 67 km from Nasca on road to Puquio, Correll & Smith PI 50 (GH). Apurimac: Prov. Aymaraes, toward Caraybamba, Saunders 77 1 (GH). Cuzco: Saxi- huaman, Cuzco, Tryon & Tryon 5346 (F, GH, us). Prov. Urubamba, Machu Picchu, Peyton & Peyton 1057 (GH, MO). Arequipa: Lomas de Atiquipa, Coronado 29 (GH, us). Puno: Puno, above Lake Titicaca, Mexia 7786 (F, GH, MO, us). 15b. Polypodium pycnocarpum var. buchtienii (Rosenst.) Tryon and Stolze, comb. nov. Polypodium buchtienii Rosenst., Repert. Spec. Nov. Regni Veg. 5: 237. 1908. Terrestrial, rupestral, rarely epiphytic, on steep rocky slopes, on rocks, in dense woods, 2000-4400 m, Cajamarca south to Cuzco. Venezuela, Ecuador, Peru, Bolivia; range un- certain. Cajamarca: Prov. Cajamarca, Lacia San Juan, San- chez 4046 (F). Prov. Celendin, Canyon of Rio Maranon above Balsas, Hutchison & Wright 5335 (GH, us). Ama- zonas: ESE of Chachapoyas, Wurdack 693 (us). La Lib- ertad: Pataz, Young 2889 (USM). 3 km west of Huana- chaco, Correll & Smith P934 (GH). San Martin: Prov. Mariscal Caceres, Puerta del Monte, Young & Leon 4916 (USM). Ancash: Muna, Woytkowski 5290 (GH, MO); Prov. Huaraz, lower slopes of Huascaran Norte, Saunders 1334 (F, GH). Huanuco: Mito, Macbride & Featherstone 1420 (F). 7520 (F, us). Lima: Prov. Yauyos, cerca a Tupe, Cerrate 1075 (GH). Junin: Huancayo, Soukup 2757 (F, GH). Huancavelica: Prov. Tayacaja, Tocas, Tovar 2017 (GH). Ayacucho: between La Quinua and Tambo, Soukup 5652 (us). Apurimac: Bosques de Ampay, Vargas 1067 (GH). Cuzco: Prov. Urubamaba, Chincheros, Davis et al. 1486 (F, USM). 1 6. Polypodium remotum Desv., Mem. Soc. Linn. Paris 6: 232. 1827. TYPE: "Habitat in Amer- ica Calidiori" (holotype; P, Herb. Desvaux), seen by Weatherby, Contr. Gray Herb. 114: 32. 1936. Polypodium leucosticton KJotzsch, Linnaea 20: 380. 1847. SYNTYPES: Colombia, Hartweg 1499 (B); (Venezuela) Merida, Moritz 336 (B). Polypodium fraseri Kuhn, Linnaea 36: 137. 1869. TYPE: Ecuador, Fraser (B). Stem short-creeping, leaves approximate, the scales appressed or somewhat spreading. Scales of the stem apex brownish, concolorous, or strongly bicolorous with a blackish center where clathrate, and broad, thin, crispate margins, the center often projecting so that the mass of scales appears as if it had blackish spines, not iridescent. Lamina to ca. 1 3 cm long, pinnatisect to 1 -pinnate, very spar- ingly scaly abaxially or not so, usually dimorphic, with the fertile leaf taller than the sterile. Distal pinnae or pinna-segments adnate, the proximal ones sessile to adnate, margins often notched, es- pecially toward the apex, trichomes absent. Veins not readily visible, free. Sorus at the tip of a vein; sori in 1 row between costa and margin. A. R. Smith (Pterid. Venezuela, 185. 1985) re- ports that the spores are chlorophyllous (green). Epiphytic, usually rupestral, or terrestrial, in forests, in cacao groves, in wet forests, on exposed cliffs, and in rocky soil, ca. 700-2600 m; Tumbes south to Cuzco. Mexico; Central America; Greater Antilles; Venezuela and Colombia south to Bolivia. Tumbes: Prov. Zarumillo, Campoverde, J. Schunke V. 2409 (F, GH). Lambayeque: 31 km from Olmos on road to Jaen, Correll & Smith P180 (GH). Cajamarca: Prov. Chota, Llama, Lopez & Sagdstequi 5309(GH, HUT). Prov. Santa Cruz, Sagdstegui & Leiva 14092 (F, GH, uc). Amazonas: Leimebamba, Woytkowski 7751 (GH, MO). Middle eastern Calla Calla slopes, Wurdack 1342 (us). La Libertad: Prov. Trujillo, Cerro Negro, Sagdstegui et al. 11015 (GH). Huanuco: Muna, Bryan 425 (F, GH), Mac- bride 3933 (us). Pasco: Quillasu, Soukup 3287 (GH, us). Prov. Oxapampa, Canyon de Huancabamba, Leon 609 (F), 620 (F). Junin: Chanchamayo Valley, C. Schunke 33 (F). Carpapata, Leon 248 (GH, USM). Ayacucho: Ayna, between Huanta and Rio Apurimac, Killip & Smith 23 1 74 (us). Cuzco: Prov. La Convention, Potrero, Tyron & Tryon 5389 (F, GH, us). 17. Polypodium furfuraceum Schlect. & Cham., Linnaea 5: 607. 1830. TYPE: Near Jalapa, Mexico, Schiede & Deppe (holotype, B). 136 FIELDIANA: BOTANY Stem short-creeping, leaves approximate, scales more or less appressed. Scales of the stem apex not clathrate, very light brown to whitish, con- colorous, elongate, the apex acuminate not irides- cent. Lamina to ca. 1 5 cm long, pinnatisect, dense- ly scaly abaxially with whitish or nearly whitish scales, not or hardly reduced at the base. Pinna- segments adnate, margins not notched, trichomes absent, fully adnate at the base and broadest there. Veins not readily visible, free. Sorus at the end of a vein; sori in 1 row between costa and margin. Epiphytic, 850 m, Cuzco. Rarely collected in southern Peru. Mexico and Central America; Peru and Bolivia. Cuzco: Santa Ana, Cook & Gilbert 1510 (us). Prov. La Convencion, Ichiquita, Vargas 22334 (GH). Prov. La Convention, Sahuayaco, Vargas 6273 (GH). Prov. La Convencion. Echarate, Vargas 22771 (GH). Quillabam- ba, Soukup 149 (F). 18. Polypodium polypodioides (L.) Watt, Canad. Nat. Ser. 2, 3: 158. 1867. Acrostichum polypodioidesL.Sp.pl. 1068. 1753. LEC- TOTYPE (by Weatherby, Contr. Gray Herb. 1 24: 28. 1939): Jamaica, Herb. Plukenet (BM). Polypodium incanum Sw., Prod. 131. 1788, nom. su- perfl. illegit. Stem long-creeping, slender, the leaves mostly distant, scales appressed. Scales of the stem apex bicolorous, dark brown to blackish with lighter margins, clathrate in the darker center, the apex acute to acuminate not iridescent. Lamina to ca. 1 5 cm long, usually less, deeply pinnatifid to pin- natisect, densely scale abaxially, monomorphic, not or only slightly reduced at the base. Pinna- segments adnate distally, the proximal ones sessile to adnate, margins not notched, the scales abaxi- ally broadly to narrowly triangular, trichomes ab- sent. Veins not readily visible; some free or all anastomosing at least along the costa. Sorus at the end of a vein; sori in 1 row between the costa and margin. All collections from Peru are evidently var. bur- chellii ; (Baker) Weath., Contr. Gray Herb. 124: 29. 1939 (Polypodium incanum var. burchellii Baker in Mart. Fl. Bras. 1(2): 526. 1870. TYPE: Goyaz, Brazil, Burchell 6998, holotype, K; isotype, GH!). The five varieties of P. polypodioides are distin- guished and discussed by C. A. Weatherby in the reference cited above. Variety burchellii has some, or most, veins anastomosing, the scales of the lam- ina, abaxially, strongly fimbriate, ovate and dilat- ed at the base and with a pronounced subulate apex (scales gomphoid or subgomphoid). Epiphytic or rarely rupestral, in dense woods, in cacao groves, on fallen trees in forests, in pas- tures, on rock walls, 250-1300 m, Cajamarca to Madre de Dios. Guianas, Venezuela, and Colombia to Bolivia; Brazil. Cajamarca: San Ignacio, Diaz 2056 (F, MO). Amazonas: Prov. Bagua, entre Aramango y Montenegro, Lopez el al 4259 (GH). San Martin: Prov. Lamas, Lamas to San Antonio, Belshaw 3455 (F, GH, MO, us). 5 km abajo de Tocache Nuevo, J. Schunke V. 3883 (F, MO, us, USM). Loreto: Prov. Maynas, Lupuna Cocha. across the Am- azon from Iquitos, Tryon & Tryon 5190(f, GH, us). Prov. Maynas, island in front of Iquitos, Simpson, & Schunke 691 (F, GH, us). Junin: Yaupe, \Voytkowski 6411 (GH, MO, us). Ayacucho: Estrella, between Huanta and Rio Apurimac, Killip & Smith 23060 (F, GH, us). Cuzco: 2 km east from Colpe de Guacomayo, Nunez 6518 (MO). Madre de Dios: Cocha Cashu, Parque National de Manu, Foster et al. 3451 (F). 19. Polypodium megalolepis Maxon & Morton, Bull. Torrey Bot. Club 66: 39. 1939. TYPE: Dept. Cuzco, Lucumayo Valley, Peru, Cook & Gilbert 1283 (holotype, us!). Stem long-creeping, leaves mostly distant, scales more or less appressed. Scales of the stem apex brown to mostly light brown, concolorous, or slightly bicolorous, not clathrate, apex acuminate to filiform not iridescent. Lamina pinnatifid to pinnatisect, to ca. 12 cm long, monomorphic, broadest at the base. Pinna-segments fully adnate or slightly joined to others, broadest at the base, not pubescent, definitely to moderately scaly abax- ially, the scales roundish to ovate-lanceolate. Veins not readily visible anastomosing. Sorus at the tip of a vein; sori in 1 row between the costa and margin. Rupestral, among scattered shrubs, 3900-4 1 00 m, Cuzco. Peru; Bolivia. Only known from Peru by the type collection. 20. Polypodium thyssanolepis KJotzsch, Linnaea 20: 392. 1847. SYNTYPES: Colombia, Otto 896 (B); Venezuela?, Moritz 22 (B). Stem short- or moderately long-creeping, the leaves mostly approximate, scales more or less TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 137 appressed. Scales of the stem apex light brown to reddish brown, slightly to definitely clathrate, slightly to definitely bicolorous, the apex acute to acuminate, not iridescent. Lamina pinnatisect or 1 -pinnate-pinnatifid proximally, to 35 cm long, monomorphic to dimorphic, not reduced at the base or only slightly so. Pinna-segments slightly adnate to adnate, the margins not notched, mod- erately to densely scaly abaxially, the scales lan- ceolate to ovate-lanceolate, the apex acuminate to filiform, not pubescent, often fully adnate at the base and broadest there. Veins not readily visible, partly free or anastomosing along the costa. Sorus at the tip of a vein; son in 1 row between costa and margin. Terrestrial to usually rupestral or rarely epi- phytic, at the base of rocks, on rocky banks, on stone walls, among open sunny rocks, 1600-3200 m, Cajamarca south to Cuzco. Southwestern United States; Mexico; Central America; Greater Antilles; Venezuela to Bolivia. Cajamarca: Prov. Celendin, Conga de Uriquis, Sa- gdstequi et al. 8417 (F, MO). Amazonas: Leimebamba, Woytkowski 7790 (GH, MO), 7843 (GH, MO). La Libertad: Prov. Otuzco, Chaullacocha, Sounders 899 (F, GH). Prov. Otuzco, Cerro Chologday, Sagstegui 0076 (GH, HUT). Huanuco: Pachachupan, cerca de Acomayo, Ridoutt (GH, USM). Mito, Macbride & Featherstone 1576 (F, GH, us), 1774 (F, GH, us). Junin: Canyon of Rio Huasahuasi, Hutchison 1060 (F, GH). Apurimac: Prov. Andahuallas, Quebrada de Toxoma, Velarde 4947 (GH). Cuzco: Prov. Urubamba, Hacienda Huarochari, Vargas 14968 (GH). 2 1 . Polypodium bombycinum Maxon, Contr. U.S. Natl. Herb. 17: 592. 1916. TYPE: Boqueron del Rio Dagua, Prov. Cali, Colombia, Leh- mann 7666 (holotype, us!; isotype, GH!; pho- to, GH of us). Stem short-creeping, leaves approximate, scales appressed. Scales of the stem apex light brown to brown, slightly bicolorous, the darker center hard- ly or not clathrate, apex acuminate to filiform, not iridescent. Lamina pinnatisect, to 60 cm long, monomorphic, gradually reduced at the base of auriculiform lobes. Pinna-segments adnate, the margins not notched, abundantly scaly abaxially, the scales reddish brown, not pubescent, fully ad- nate at base and broadest there. Veins not readily visible, anastomosing. Sorus at the end of a vein; sori in 1 row between costa and margin. Epiphytic or rarely terrestrial, swamp forests, on palms or on tree ferns, 100-580 m, Loreto, Cuzco, and Madre de Dios. Panama; northern South America; Amazonian Brazil; Peru; Bolivia. Polypodium balaoense Hieron. has the scales on the lamina abaxially ovate-triangular with a long tip in contrast to the gomphoid scales of P. bom- bycinum, which are mostly orbicular, with a fili- form tip. Loreto: Prov. Maynas, Rio Santa Maria, King 458 (F). Cuzco: 2 km east from Colpa de Guacamayos, Nunez 6522 (MO). Puente de Inambari, Vargas 18464 (GH). Ma- dre de Dios: Prov. Tambopata, Tambopata Reserve, Young 39 (MO), Funk et al. 8198 (us). 39 km southwest of Puerto Maldonado, Smith et al 1130 (F). 22. Polypodium balaoense Hieron. Bot. Jahrb. Syst. 34: 529. 1905. TYPE: "prope Balao (Ec- uador)," Eggers 14286 (holotype, B). Stem short-creeping, leaves approximate, scales appressed. Scales of the stem apex brown, bicolor- ous, the darker center clathrate, apex acuminate to filiform, not iridescent. Lamina pinnatisect to ca. 60 cm long, monomorphic, gradually reduced at the base to auricliform lobes. Pinna-segments adnate, the margins not notched, somewhat scaly abaxially, the scales whitish, not pubescent, fully adnate at base and broadest there. Veins not readi- ly visible, anastomosing. Sorus at the end of a vein; sori in 1 row between costa and margin. This species seems to intergrade with P. bom- bycinum. A single collection from northern Peru. Panama south to Peru; Galapagos Islands. Tumbes: (Prov. Tumbes), hwy. to El Caucho, Coro- nado 234 (uc). 23. Polypodium monosorum Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck Ge- samten Naturk 5: 319. 1811. TYPE: "Peru- via" (holotype, P), Herb. Desvaux, a single pinna). A label on the sheet refers to Herb. Jussieu, where sheet 1 1 24, Jos. Jussieu, "Pe- rou," P, photo, GH is undoubtedly the source of Desvaux's pinna. The specimen could have been collected in what is now Ecuador. Stem long-creeping, leaves mostly distant, scales appressed, or sometimes the stem partially naked. Scales of the stem apex linear, usually minutely denticulate, bicolorous with very narrow, lighter 138 FIELDIANA: BOTANY margins and a clathrate, brown or darker center, not iridescent or slightly so, the apex attenuate. Lamina to 20 cm long, 2-pinnate-pinnatind to 3-pinnate, slightly dimorphic, slightly reduced at the base, or not. Pinnae stalked, mostly triangular, not pubescent abaxially, where the scales are del- toid-acuminate. Veins not readily visible, free. So- rus at the end of a vein; sori individual on the ultimate segments. A single collection from Huanuco at 2745 m, in addition to the type. Colombia to Peru. The somewhat similar Polypodium murorum Hooker is common in Ecuador and may also grow in Peru. It has the lamina to 1 -pinnate-pinnatifid, most or many of the adaxial scales are rounded, and the scales or the stem apex are definitely and strongly denticulate. Huanuco: Mito, Bryan 208 (F). Comments Following are two names based on Peru material or credited to Peru. Polypodium crystalloneuron Rosenst., Repert. Spec. Nov. Regni Veg. 11: 57. 1912. SYN- TYPES: Unduavi, Bolivia, Buchtien "2773- 75" (holotype, not located). Hensen (1990) has selected Buchtien s.n., B, as lectotype; iso- syntype, Buchtien 2775 (uc!, us!). Material assigned to this species occurs in Bo- livia and southern and central Peru. However, it is quite possible that it is a southern variation of Polypodium loriceum and should be treated as a synonym or as an infraspecific taxon under that species. Because of doubt about its taxonomic sta- tus, it is dealt with here rather than in the text proper. The taxon has a stem that is usually thicker than that of P. loriceum, and the stem scales are more persistent. The stem also more often has a chalky white deposit (dry), which is rare in P. loriceum. Hensen ( 1 990) said that P. loriceum has the vas- cular bundles of the stem (meristeles) with a dark sclerenchymatous sheath, while this is lacking in P. crystalloneuron. The taxon grows in Peru and Bolivia. The Peru material seen is as follows. Ancash: Prov. Yungay, Huascaran National Park, D. Smith et al. 9091 (MO), D. Smith & K. Goodwin 8828 (MO). Cuzco: Prov. Paucartambo, Nunez et al. 8491 (MO, uc). Prov. Paurcartambo, Tupayachi 55 (uc), 56 (MO). Prov. Calca, Nunez 6724 (MO). Prov. Urubamba, Peyton & Peyton 1091 (MO). Polypodium mutabile Kunze, Linnaea 9: 46. 1 834. TYPE: Peru, Poeppig (holotype, LZ, de- stroyed; isotype, ?, w). On the basis of an authentic specimen at w, Hensen ( 1 990) placed this name as a synonym of Polypodium fraxinifolium. The following three hybrids have been proposed for Peru Polypodium. Others will probably be de- tected as the species are studied closely. Polypodium lasiopus • molledense. Polypodium Xhuancayanum Kunkel, Biota 6: 157, figs. C-D. 1966. TYPE: Huancayo, Peru, Kunkel 6569 (holotype, Herb. Kunkel ?). In the present treat- ment, P. mollendense is a synonym of P. pyc- nocarpum var. pycnocarpum. Polypodium thyssanolepis • Pleopeltis macrocar- pa. Polypodium leucosporum KJotzsch, Linnaea 20: 404. 1847. TYPE: Mucuchies, Merida (Venezuela), Moritz 306 (holotype, B). This is a putative hybrid of Pleopeltis macro- carpa and Polypodium thyssanolepis. Other spe- cies of Pleopeltis may be involved as parents of other hybrids in Mexico and Central America (see Mickell & Beitel, Amer. Fern J. 77: 16-27. 1987). The hybrid plants are variously intermediate be- tween a species with an entire lamina and one with a deeply pinnatifid or pinnatisect lamina. The spores are abortive and the lamina is definitely scaly abaxially. Plants of this hybrid occur sporadically from Piura and Amazonas to Cuzco, at 1800-2800 m. Piura: Prov. Huancabamba. 10 km N of Huancabam- ba, Hutchison & Wright 6601 (F, GH, us). Amazonas: Prov. Chachapoyas, Calla Calla Slopes, Wurdack 1181 (F, GH, us). Prov. Chachapoyas, Pomachocha, Lopez et al. 4395 (GH), 4396 (GH). Huanuco: La Molina, near Panao, Asplund 13662 (us). Junin: Between Tarma and San Ramon, Croat 57700 (F). Apurimac: Andahuailas, Herrera 1496 (GH). Cuzco: Prov. Anta. Vargas 17639 (GH). TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 139 Polypodium laevigatum x lasiopus. Goniophlebium semipinnatifidum Fee (Mem. foug. 5) Gen. fil. 256. 1852. TYPE: Pamplona, Columbia, Funck & Schlim 1363 (holotype, P?). Polypodium semipinnatifidum (Fee) Mett., Abh. Senckenberg Naturf. Ges. 2: 80. 1856. A single collection of this hybrid is from Peru, Amazonas, 1-5 km west of Molinopampa, Wur- dack 1363 (MO, us). Other specimens of this col- lection at F and GH are Polypodium laevigatum. The parents of this hybrid are uncertain. In Peru they are probably P. laevigatum x lasiopus and elsewhere another parent with P. laevigatum is likely involved. The lamina has few or no scales on the abaxial side. At the present time the epithet semipinnatifidum is used to include all similar hy- brids. Terrestrial in the jalca zone, 2400-2500 m, Amazonas. Columbia; Ecuador; Peru. Polypodium dulce Poir. in Lam., Encycl. 5: 523. 1804. TYPE: Morne de la Calebasse, Marti- nique. Plumier (Traite foug. Amer., t. 80. 1705) has been cited as the lectotype by Proctor and Lourteig (Bradea V (40): 387. 1990). However, the speci- men may take precedence over the figure as a type and this (Jussieu Herb. Cat. 1085) has been iden- tified in microfiche as Polypodium loriceum by Lellinger (pers. comm.). Because there are obvious problems in the typification of the name and the identification of the type, the name has neither been accepted nor placed in synonymy. IV. Pleopeltis Pleopeltis Willd., Sp. pi. 5: 2 1 1 . 1 8 1 0. TYPE: Pleo- peltis angusta Willd. Figure 8. Plants epiphytic or epipetric, rarely terrestrial. Stem long- or short-creeping, bearing peltate or peltately attached, clathrate or nonclathrate, usu- ally comose, scales. Leaves ca. 5—40 cm long, monomorphic or slightly dimorphic, articulate to the stem or to short phyllopodia, approximate to widely spaced. Lamina entire (or pinnatisect in P. angusta, not in Peru), moderately to abundantly scaly, especially abaxially, the scales peltately at- tached, most of them circular (or with stellate arms from the circular base in P. fuscopunctata). Veins anatomosing, with or without included free vein- lets. Sori round to somewhat elongate, usually borne at the junction of 2 or more veins, on a somewhat raised receptacle, exindusiate, inter- mixed with peltate or filiform paraphyses. Spores monolete, ellipsoid, with long laesura. Circumscription of this genus is still actively debated, especially in relationship to Microgram- ma. To some taxonomists it consists of a dozen Neotropical species, plus one other found in Af- rica, India, and Sri Lanka. Others include the Asian genus Lepisorus, with about 25 species. Even with Pleopeltis, s.s. some authors prefer to exclude P. fuscopunctata and P. percussa, transferring these two to Microgramma (Sola, 1986). For purposes of this flora, P. percussa is included in Pleopeltis on the basis of its circular lamina scales, subclath- rate stem scales, and the peltate paraphyses that cover the immature sorus (although the latter are rarely seen on dried specimens). Stem scales are not or scarcely clathrate in Pleo- peltis fuscopunctata or P. percussa, but they do have deciduous soral paraphyses. Lamina scales, though very small and appearing at first to be merely glandular dots, actually consists of stellate arms spreading from a minute, circular base. Ex- cept for P.fuscopunctatum, leaves of Pleopeltis are mostly coriaceous, whereas leaves of Microgram- ma are commonly chartaceous or thinner in tex- ture. There has been hybridization between some species of Pleopeltis and Polypodium, in one case involving Pleopeltis macrocarpa. See treatment of Polypodium thyssanolepis x Pleopeltis macrocar- pa in Comments following the genus Polypodium. References SOTA, E. R. DE LA. 1986. Sobre la position sis- tematica de Polypodium fuscopunctatum Hook. FIG. 8. Pleopeltis macrocarpa var. macrocarpa: a, habit; b, lamina scales. Pleopeltis macrocarpa var. laciniata: c, sterile leaf; d, lamina scale. Pleopeltis percussa: e, section of sterile lamina, (a, b from Hutchison & Wright 5370, F, c, d from Wurdack 2034, USM; e from Dudley 11418, GH.) 140 FIELDIANA: BOTANY TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 141 y Polypodium percussum Cav. Physis, C44: 1 9- 28. WEATHERBY, C. A. 1922. The group of Polypo- dium lanceolatum in North America. Contr. Gray Herb., 65: 3-14. Key to Species of Pleopeltis a. Lamina firm-herbaceous; veins distinct on both surfaces; lamina dark-punctate with minute, stellate scales usually less than 0. 1 mm in diameter (these, without high magnification, appearing as glandular dots) 4. P. fuscopunctata a. Lamina chartaceous to subcoriaceous; veins obscure or indistinct, at least abaxially; lamina with circular scales mostly 0.2-0.5 mm in diameter b b. Sporangia intermixed with dense masses of persistent, filiform paraphyses 3. P. percussa b. Sporangia intermixed with few to many, commonly fugacious, peltate paraphyses c c. Stem scales lanceolate to ovate, bicolorous, each not obscured by a mass of trichomes; sori circular to ovoid, 1-1.5 times as long as broad 1. P. macrocarpa c. Stem scales mostly circular, blackish, each usually completely obscured by a dense mass of trichomes; sori elongate, commonly 2-4 times as long as broad 2. P. astrolepis 1. Pleopeltis macrocarpa (Willd.) Kaulf, Berlin Jahrb. Pharm. 21:41. 1820. Plants epiphytic or epipetric, rarely terrestrial. Stem long-creeping, densely covered with lanceo- late to ovate scales, these 1-2 mm long, bicolorous (tawny to orange with blackish centers), and with erose-ciliolate margins. Leaves 6—40 cm long, 0.8- 4 cm broad, petiolate, articulate to the stem, es- sentially monomorphic. Petiole 0.5-7 cm long, te- rete (in Peru). Lamina simple, margins entire or rarely slightly sinuate, chartaceous to subcoria- ceous, narrow-to oblong-elliptic, with acute to at- tentuate apex, and cuneate or attenuate base, abundantly scaly, the scales peltate, scattered, of 2 kinds, most of them oblong, 0.2-0.5 mm in di- ameter, concolorous or bicolorous, these often in- termixed with larger, ovate-acuminate ones. Costa yellowish (or rarely blackish at base) abaxially. Veins distinct to obscure. Sori somewhat im- mersed, ovoid, discrete (or rarely a few of them merging), the sporangia intermixed with few to many circular, peltate, commonly fugacious, pa- raphyses. The species occurs in the Greater Antilles, from Mexico to Chile and Argentina, and in Africa to India and Sri Lanka. There are five varieties of P. macrocarpa, three of them confined to Mexico and/or Central Amer- ica. One of these, var. trichophora (Weath.) Pic.- Ser., is distinguished by the conspicuous trichomes borne at the bases of stem scales. Another, var. complanata (Weath.) Lell., has a conspicuously flattened petiole. A third, var. crassinervata (Fee) Moore, is distinguished by the stem scales, which are pale and concolorous (except for the dark point of attachment). The other two varieties occur in Peru. Key to Varieties a. Lamina scales distinctly bicolorous, circular ones 0.3-0.5 mm in diameter, the margins erose or erose-ciliolate; lamina attenuate at base, coriaceous or subcoriaceous and the veins obscure la. var. macrocarpa a. Lamina scales concolorous, circular ones 0.2-0.3 mm in diameter, the margins deeply laciniate; lamina broadly or narrowly cuneate at base, chartaceous and the veins distinct adaxially . Ib. var. laciniata 142 FIELDIANA: BOTANY la. Pleopeltis macrocarpa var. macrocarpa. Fig- ures S.i I). Polypodium lanceolatum L., Sp. pi. 2: 1082. 1 753, not Pleopeltis lanceolata Kaulf. 1 824. TYPE: Petiver, Pteri-graphia Americana, t. 6,f. 2. 1712. Polypodium macrocarpum Willd., Sp., pi. ed. 4, 5: 147. 1810. TYPE: Mauritius, Bory (holotype, B, Herb. Willd. 19629; isotype, p). Leaves 6-35 cm long, 0.8-2.4(-3) cm broad. Petiole 0.5-7 cm long. Lamina narrow-elliptic, at- tenuate at base, coriaceous or subcoriaceous, scales castaneous, with margin whitish to tawny, erose or erose-ciliolate. rieties the scales are bicolorous, i.e., with a much darker center, and the margins are subentire to, at best, erose-fimbriate. In var. laciniata, scales are rather uniformly castaneous, and both the circular and the elongated ones are deeply dissected with long and acute laciniae, much like those ofP.fus- copunctata. Even the abundant peltate scales in- termixed with maturing sporangia are similarly laciniate. Although the new variety is known thus far only from the type and two other specimens, its rela- tively large leaves should be conspicuous enough to enable collectors to locate more specimens in the future. In wet forests and wooded ravines, or in lomas, on tree trunks and branches, on cliffs or among rocks, occasionally in rocky soil, 500-2900 m, Piu- ra to Amazonas, south to Arequipa and Puno. General distribution is the same as the species. Piura: Prov. Huancabamba, Dist. Sapalacha, Sagds- tegui & Cabanillas 8596 (F, HUT). Lambayeque: Prov. Ferrenafe, Tute, Llatas Q. 2513 (F). Cajamarca: Prov. Celendin, Canyon of Rio Maranon above Balsas, Hutch- ison & Wright 5370 (F, GH). Amazonas: Prov. Chacha- poyas, Quebrada Molino below Chachapoyas, Wurdack 633 (F, GH, us). Huanuco: Mito, Macbride & Featherstone 1775 (F, GH, us). San Martin: San Martin, Boca Toma del Shilcayo, N of Tarapoto, Alcorn & Mallet 25 (MO). Junin: Prov. Tarma, 5 km SW of Huacapistana, Tryon & Tryon 5427 (GH, us). lea: San Gallan Island, Murphy 3475 (us). Apurimac: Prov. Caraveli, Lomas de Pongo, Ferreyra 13434 (GH, USM). Cuzco: Quillabamba, Potrero, Coronado 125 (GH, us). Arequipa: Lomas de Atiquipa, Coronado 36 (GH, us). Puno: Ollachea to San Gaban, Dillon et al. 1155 (F, USM). Ib. Pleopeltis macrocarpa var. laciniata Stolze, var. nov. Figures 8c-d. Folia 2.5—4 cm latae. Lamina chartacea, oblon- go-elliptica, ad basin cuneata, paleis concoloribus, pro parte maxima circular! bus, 0.2-0.3 mm dia- metris, ad marginem profunde laciniatis. TYPE— Peru, Dept. Huanuco, Tingo Maria, hills above river, Moran & Fernandez 3681 (holotype, USM!; isotypes, F!, MO; photos, F, MO, uc of USM). In forests, on rocky slopes or on trunks of trees, 400-700 m, Amazonas, Huanucu, and Pasco. Endemic. This is the most distinctive variety of P. macro- carpa. It is readily distinguished from all the others by the larger lamina with broad- to narrow-cu- neate (rather than attenuate) base and by the mar- gin and color of the laminar scales. In other va- Amazonas: Prov. Bagua, Rio Maranon, above Cas- cadas de Mayasi, Wurdack 2034 (USM). Pasco: Prov. Oxapampa, Palcazu, Rio Alto Iscozacin, Ozuz to Pes- cado, Foster & d'Achille 10121 (F). 2. Pleopeltis astrolepis (Liebm.) Fourn., Mexic. pi. 1: 87. 1872. Grammitis lanceolata Schkuhr, Vier Zwangiste Kl. Linn. Pfl.-Syst. 1 : 9, t. 7 in part. 1 804, not Polypo- dium lanceolatum L. 1 753, nor Pleopeltis lanceo- lata Kaulf. 1824. TYPE: Jamaica, probably Swart z (holotype, HAL). Grammitis elongata Sw., Syn. fil. 22, 213. 1806, not Pleopeltis elongata Kaulf., 1824. Nom. nov. for G. lanceolata Schkuhr, not Swartz. Grammitis revoluta Willd., Sp. pi. ed. 4: 139. 1810, not Pleopeltis revoluta Alderwerelt. 1909. TYPE: Locality and collector unknown (holotype, B, Herb. Willd. 19584). Grammitis squamulosa Splitg., Tijdschr. Nat. Gesch. Physiol. 7: 398. 1 840, not Pleopeltis squamulosa (Kaulf.) Presl, 1836, nom. illeg. TYPE: Surinam, collector unknown (holotype, L). Polypodium astrolepis Liebm., Kongel. Danske Vi- densk. Selsk, Skr., Naturvidensk. Afd., ser. 5, 1: 185. 1849. LECTOTYPE (designated by A. R. Smith, Fl. Chiapas 2: 177. 1981): Mexico, Oa- xaca, Trapiche de la Conception, Leibmann Fl. Mex. 87, lower specimen (c). Pleopeltis revoluta (Willd.) A. R. Smith, Proc. Calif. Acad. Sci. 40: 230. 1975, not Alderwerelt, 1909. Plants epiphytic, occasionally epipetric. Stem usually long-creeping, amply provided with black- ish peltate, circular, or elongate scales, these 0.3- 0.5 mm long, bearing, and usually obscured by, abundant, castaneous, unicellular trichomes to 1 mm long. Leaves 4-1 5 cm long, 0.4-1 .8 cm broad, subsessile or very short-petiolate, borne on short (0.5-1.5 mm) phyllopodia, monomorphic or sub- dimorphic (sterile one frequently shorter and/or broader than the fertile); petiole 0.4-1.5 cm long, conspicuously flattened. Lamina simple, entire, TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 143 chartaceous to coriaceous, narrow-elliptic to lin- ear-lanceolate, attenuate at base, bearing scat- tered, circular, lacerate-stellate scales to 0.3 mm in diameter, and some longer, ovate-lanceolate ones near the costa and margin. Costa castaneous to blackish abaxially. Veins indistinct or obscure. Sori oblong or elliptic, 2—4 times as long as broad, rare- ly merging to form longer lines, a few, deciduous, peltate scales intermixed with sporangia, but rarely seen. In forests, on trees or rarely on mossy rocks, 900-1500 m, Junin and Cuzco. Mexico to Panama; West Indies; Columbia to the Guianas, south to Bolivia and Brazil. Junin: 20 km from San Ramon, road to Tarma, Leon et al. 339 (F, USM). Chanchamayo, San Ramon near La Merced, Teppner 206 (us). Cuzco: Chaupimayo, Valle del Puerto, Biies 20 J 2 (us). Quillabamba, Soukup 179 (F). 3. Pleopeltis percussa (Cav.) Hooker & Grev., Icon, fil., /. 67. 1828. Figure 8e. Polypodium percussum Cav., Descr. pi. 243. 1802. TYPE: Peru (as Mariannas), Nee (holotype, MA; photo, F). Microgramma percussa (Cav.) Sola, Physis C44: 1 9. 1986. Plants epiphytic, occasionally terrestrial, rarely epipetric. Stem long-creeping, 0.2-2 mm thick, densely scaly, the scales of two kinds: minute, cas- taneous to blackish, appressed, circular ones, these often obscured by larger to (3 mm), and flaccid, subappressed or spreading ones, the latter lanceo- late, subentire, nonclathrate, tawny to orange, usu- ally with a castaneous point of attachment, peltate above the base. Leaves widely spaced, petiolate, articulate to the stem, essentially monomorphic. Petiole 0.5-3 cm long. Lamina simple, entire, 7- 27 cm long, 1-3.5 cm broad, coriaceous, opaque, narrow-elliptic, attenuate at both ends, essentially naked adaxially, minutely peltate-scaly abaxially, the scales abundant, scattered, mostly 0.2-0.3 mm broad, circular, entire, some much larger, ovate- lanceolate ones along the costa. Veins immersed and usually obscure, copiously anastomosing, only a few areoles containing free veinlets, these ori- ented in all directions. Sori discrete, circular to oval, rather deeply impressed, paraphyses copi- ous, filiform, ferruginous to castaneous, forming dense masses among the sporangia. Very common, in or at edges of wet forests and thickets, on stumps, tree trunks, and high branch- es, occasionally terrestrial or clambering over rocks, 100-2250 m, Cajamarca to Loreto, south to Aya- cucho and Madre de Dios. Southern Mexico to Panama; Columbia to the Guianas, south to Bolivia and Brazil. Cajamarca: Prov. Cutervo, 1 km from San Andres, Sanchez Vega 4475 (F, MO). Amazonas: Prov. Bagua, on Cerro Tapur, 40 km S of Bagua Grande, Hutchison 1477 (F, GH, uc). San Martin: Prov. Mariscal Caceres, Dist. Campanilla, Muyuna de Murga, Schunke V. 4139 (f, uc, us). Loreto: Dist. Iquitos, Rio Napa near Mazan, Mexia 6473 (F, GH, uc, us). Huanuco: Prov. Huanuco, Dist. Churubamba, Hacienda Exito, Mexia 8164 (BM, F, GH, uc, us). Pasco: 45 km from Pte. Rio Paucartambo on road to Oxapampa, Leon et al. 483 (USM). Junin: Prov. Tarma, Huacapistana, between Tarma and San Ramon, Ferreyra 11260 (GH, USM). Ucayali: Prov. Coronel Por- tillo, La Divisoria, 20 km NNE of Tingo Maria, Dillon 2629 (F, USM). Ayacucho: Ccarrapa, between Huanta and Rio Apurimac, Killip & Smith 22374 (F, us). Cuzco: 40 km W of Pilcopata on road to Paucartambo, Gentry et al. 23671 (F, MO, us). Madre de Dios: Cocha Cashu, Manu National Forest, between Paragua and Tayakome, Foster et al. 3419(f). 4. Pleopeltis fuscopunctata (Hooker) R. & A. Tryon, Rhodora 84: 129. 1982. Polypodium fuscopunctatum Hooker, Sp. fil. 5: 69. 1 863. TYPE: Ecuador, Chimborazo, Spruce 5734 (holotype, K; photo, GH). Microgramma fuscopunctata (Hooker) Vareschi, Flo- ra Venezuela 1: 893. 1969. Plants epiphytic. Stem long-creeping, 0.8-1.5 mm in diameter, densely (sometimes completely) covered with appressed to (at the tip) spreading scales, these 2-3 mm long, flaccid, ovate, acute, peltately attached above the base, orange or tawny, the margins subentire. Leaves ( 1 0-) 1 2-25(-30) cm long, (1.3-)1.8-2.5(-3) cm broad, subsessile to short-petiolate and articulate to the stem or to very short phyllopodia, essentially monomorphic. Pet- iole 0-10 mm long. Lamina simple, entire, firm- herbaceous, lance-elliptic, attenuate at both ends, surfaces punctate with dark brown, minute (0.05- 0.1 mm), peltate, circular- stellate scales, costa abaxially with a few, scattered, light brown, ovate scales to 2 mm long. Veins distinct, somewhat prominulous, primary and other ones equally prominent, areolate, the costal areoles frequently containing one, short, recurrent, free veinlet. Sori circular, borne in a single medial to supramedial row between costa and each margin. Climbing on tree trunks and branches, or along 144 FIELDIANA: BOTANY lianas, in and at edges of dense forests, 100-1250 m, Tumbes, Amazonas and Loreto south to Cuzco and Puno. Trinidad; the Guianas to Columbia, south to Brazil and Bolivia. The relationship of this species is not clear. Some authors place it in Microgramma, partly because of its thinner lamina texture, distinct veins, and nonclathrate stem scales. It also apparently lacks the peltate paraphyses and circular lamina scales so typical ofPleopeltis. However, there are stalked, peltate paraphyses in immature son, although these are rarely seen in dried specimens. Furthermore, the laminar surface is finely punctate with what appear to be glandular dots. High magnification (80- 1 00 x ) reveals these to be peltate, circular scales ca. 0.05 mm in diameter, each with 6-8 short to long, marginal arms in a stellate pattern. In this respect, they are nearer to the typical lamina scales of Pleopeltis than to the noncircular ones of Mi- crogramma. This species apparently occupies an interme- diate position between the two genera. For pur- poses of this treatment, it is maintained in Pleo- peltis, since it keys out most effectively with the characters of lamina scales and paraphyses. Tumbes: Prov. Zarumilla, Dist. Matapalo, Bosque Na- cional de Tumbes, Simpson & Schunke 381 (F, GH, us, USM). Amazonas: Prov. Bagua, forest behind Parcelacion Monterrico, Knapp & Alcorn 7624 (F, MO). San Martin: Prov. Mariscal Caceres, Tocache Nuevo, Schunke V. 6950 (F, uc). Loreto: Left bank of Rio Santiago above Pongo de Manseriche, Mexia 6144a (GH, uc, us). Huanu- co: Near Divisoria, 80 km E of Tingo Maria, Gentry et al. 16053 (F, MO, uc). Pasco: Prov. Oxapampa, Gran Pajonal, near Chequitavo, D. Smith 5187 (MO, uc, USM). Cuzco: Prov. La Convention, Rio Klause near conflu- ence with Rio Mapitunuari, Dudley 10130 (GH). Madre de Dios: Confluence of Rios Tambopata and La Torre, SW of Puerto Maldonado, S. F. Smith et al. 544 (F, us). Puno: Prov. Carabaya, near San Gaban, Vargas 17559, 18914 (GH). Plants epiphytic. Stem small, short-creeping, bearing clathrate scales and fibrous roots. Leaves monomorphic, loosely fasciculate, to 40 cm long, articulate to the stem, with petioles obsolete. Lam- ina attenuate at base, subdichotomously forked into 2 to several segments, or deeply pinnatisect and the 3-4 segments on either side strongly as- cending, sparsely to copiously scaly. Veins free, or anastomosing without included free veinlets. Sori round and terminal on vein tips, or elongate, or in a long, inframarginal line the receptacle slightly raised, not paraphysate, exindusiate. Spores mon- olete, ellipsoid, with long laesura. Dicranoglossum is a Neotropical genus of three to five species that are weakly distinguished by patterns of venation and size and shape of stem and laminar scales, and their taxonomy is still open to question. Some taxa (e.g., D. subnudum) have laminae so thick that the venation pattern can only be seen by the laborious process of clear- ing. In other taxa, leaves must be held to strong light to observe the veins. Authors have separated other species according to apparent differences in size, shape, and frequency of laminar scales, which, however, tend to be highly variable. These scales typically are minute and circular, sometimes sparse, sometimes copious. Some have short cusps, others have a long, attentuate apex, and there are inter- mediate gradations. Sporangia typically are borne on a long, narrow receptacle served by a vascular commissure, but in D. polypodioides the receptacle is usually con- fined to the vein tips, with sori more or less round and discrete. Sporangia of D. panamense (C. Chr.) Gomez (Central America and northern South America) are generally in long lines, but these sometimes grade into separate sori and the two species may be synonymous. See D. polypodioides for further discussion. V. Dicranoglossum Dicranoglossum John Sm., Bot. voy. Herald (See- mann) 232. 1854, nom. nov. for Cuspidaria Fee and with the same type. Figure 9. Cuspidaria Fee, Mem. Soc. Mus. Nat. Hist. Strasbourg 4: 201. 1850, not DC. 1838. TYPE: Cuspidaria furcata (L.) Fee (Pteris furcata L.) = Dicranoglos- sum furcatum (L.) John Sm. Eschatogramme C. Chr., Bot. Tidsskr. 26(2): 285. 1904. References CHRISTENSEN, C. 1929. Taxonomic fern studies. I. Revision of the polypodioid genera with lon- gitudinal coenosori (Cochlidiinae and "Dry- moglossinae"); with a discussion of their phy- logeny. Dansk. Bot. Ark., 6(3): 1-93. TRYON, R. M., AND A. F. TRYON. 1982. Dicran- oglossum, pp. 708-7 1 1 . In Ferns and Allied Plants. Springer- Verlag, New York. TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 145 146 FIELDIANA: BOTANY Key to Species of Dicranoglossum a. Stem scales obviously bicolorous, reddish-brown with a conspicuous, black, central band; laminar scales few and scattered or lacking; lamina subcoriaceous and the veins obscure . . 1 . D. subnudum a. Stem scales essentially concolorous, dark gray- or red-brown; laminar scales abundant; lamina firm- herbaceous to chartaceous and veins mostly visible when backlighted b b. Veins anastomosing, forming costal areoles; sporangia in continuous lines (coenosori); rachis and costae darker than adjacent tissue, at least in the proximal half of the lamina ... 2. D. desvauxii b. Veins of sterile portion of lamina free; sporangia round or oblong, mostly discrete (some confluent at maturity); rachis and costae lighter in color than laminar tissue, stramineous nearly throughout 3.D. polypodioides 1. Dicranoglossum subnudum (C. Chr.) Stolze, comb. nov. Figures 9a-c. Eschatogramme furcata var. subnuda C. Chr., Dansk Hot. Ark. 6(3): 36. 1929. TYPE: Bolivia, Nor- dyungas, Polo Polo near Coroico, Buchtien 3586 (holotype, us!; isotypes, F!, K). Eschatogramme subnuda (C. Chr.) Copel., Univ. Calif. Publ. Bot. 19: 305. 1941. Stem scales 1-2 mm long, lanceolate to ovate, acuminate, conspicuously bicolorous, reddish- brown, with a narrow to broad, black, often lus- trous, central band of cells. Leaves 1 5-40 cm long, 10-20 cm broad. Lamina subdichotomously di- vided into 2—4 segments, these often once again forked, subcoriaceous, naked, or with a few scales widely scattered on the abaxial side, these 0.4-0.8 mm long, lanceolate. Rachis and costae red- or gray-brown, abaxially, obviously darker than the adjacent laminar tissue, rarely lighter distally. Ul- timate segments 3-7 mm broad, fertile portions contracted and the margins usually revolute. Veins obscure, 1-2-forked, essentially free. Sporangia in submarginal. linear coenosori on the distal portion of segments. Pendent from tree trunks or branches, in deep forests, 700-1550 m, Amazonas, Huanuco, and Junin. Peru; Bolivia. Mr. J. K. Sommer, University of Utrecht, The Netherlands, had proposed the new combinations of D. subnudum as part of his monograph of the genus. Sadly, due to his untimely death in 1989, this work was not published, so the new combi- nation is published here. Christensen (1929) separated D. furcatum into several varieties, including vars. bicolor and sub- nudum. However, the bicolorous stem scales of the last two more clearly separate them from D. furcatum than the highly variable laminar scales distinguish D. furcatum from D. polypodioides or D. panamense. Hence, D. subnudum is here ele- vated to the level of species. Specimens from Co- lumbia, designated by Christensen as var. bicolor, apparently differ from D. subnudum only in the numerous laminar scales and probably should be treated as a form of the latter. Amazonas: Prov. Bagua. Cerro Tapur on Rio Utcu- bamba. Hutchison 1488 (uc). Huanuco: Tingo Maria to Pucallpa, Ellenberg 3895 (GH). Prov. Huanuco, Cotir- arda, Mexia 8220 (F, GH, us). Junin: Prov. Tarma, Chan- chamayo, Esposto, USM cat. no. 11023 (USM). La Mer- ced, Jardin el Perezoso, Leon 267 (GH, USM). La Merced, Hacienda Schunke, Macbride 5658 (F, us). Chancha- mayo Valley, C. Schunke 113 (F). 2. Dicranoglossum desvauxii (Klotzsch) Proctor. Rhodora 63: 35. 1961. Figure 9d. Taenitis desvauxii Klotzsch, Linnaea 20: 431. 1847. LECTOTYPE (designated by Proctor, Flora Less- er Antilles 348. 1977): Hooker & Greville, Icon, fil. \, t. 7. 1827, based on a Guilding specimen from St. Vincent. Eschatogramme desvauxii (Klotzsch) C. Chr., Dansk Bot. Ark. 6(3): 37. 1929. Stem scales about 1 mm long, ovate, essentially concolorous, dark brown or blackish, rarely slight- ly lighter in color just at the margin. Leaves 8-38 cm long, 5-14 cm broad. Lamina firm-herbaceous to (occasionally) chartaceous, abundantly scaly FIG. 9. Dicranoglossum subnudum: a, habit; b, portion of fertile plinna; c, stem scale. Dicranoglossum desvauxii: d, portion of fertile pinna; Dicranoglossum polypodioides: e, portion of fertile pinna, (a, b, c from Mexia 8220. F, d from Hutchison & Wright 3726, GH; e from Simpson 482, F.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 147 abaxially, scales on the laminar tissue circular and 0. 1 mm in diameter, or with a cusp as long as the scale body. Rachis and costae red- or gray-brown, obviously darker than the adjacent laminar tissue, sometimes lighter distally, provided with a few scattered, lanceolate, clathrate scales to 1 mm long. Ultimate segments 5-12 mm broad, fertile por- tions sometimes slightly contracted, the margins plane, rarely revolute. Veins indistinct, but com- monly evident when held to light, merging to form elongated costal areoles. Sporangia in continuous, narrow, inframarginal lines (coenosori) in the dis- tal portions of segments. In wet forests and wooded ravines, on tree trunks and branches, 375-850 m, Amazonas and Loreto, south to Pasco, Madre de Dios. St. Vincent; Trinidad; Columbia to the Guianas, south to Bolivia and Brazil. In Dicranoglossum desvauxii segments tend to be more numerous than in other species— often three or four on each side. Yet even the basal ones are so long and strongly ascending as to reach near the tip of the apical segment. Thus, the general aspect of the lamina is still of a subdichotomous pattern. This is the only species in the genus having veins of sterile portions of segments merging to form costal areoles. Amazonas: Prov. Bagua, 5.5 km NE of Montenegro, Hutchison & Wright 3726 (GH, uc). Prov. Bagua, Rio Maranon above Cascades de Mayasi, Wurdack 1855 (us). San Martin: Prov. Mariscal Caceres, Dist. Tocache Nue- vo, Quebrada de Yacu Sisa, Schunke V. 7554 (F, uc). Loreto: Prov. Maynas, Quebrada Sucusari, Vdsquez & Jaramillo 11838 (F, MO). Huanuco: 5 km NE of Tingo Maria, Stork & Morton 9577 (F, GH, uc, us). Prov. Huan- uco, Tingo Maria, Tryon & Tryon 5226 (GH, us, USM). Pasco: Puerto Bermudez (as Junin), Killip & Smith 26417 (F, us). Prov. Oxapampa, Valle de Palcazu, near Iscoz- acin, Leon 709 (F, USM). Madre de Dios: Prov. Mania, Rio Alto Madre de Dios, near Atalaya, Foster & Baldeon 12854 (F). 3. Dicranoglossum poly pod ioides (Hooker) Lei- linger, Selbyana 2: 283. 1978. Figure 9e. Taenitis furcata var. polypodioides Hooker, Sp. fil. 5: 1 88. 1 864. TYPE: Ecuador, base of the Andes on road to Guayaquil, Jameson (holotype, K). Eschatogramme polypodioides (Hooker) C. Chr., Dansk Bot. Ark. 6(3): 38. 1929. Stem scales 1-2 mm long, lanceolate to ovate, dark brown or blackish, rarely slightly lighter in color just at the margin, Leaves 1 2-30 cm long, 4-8 cm broad. Lamina subdichotomously divided into 2-4 segments, these often once again forked, firm-herbaceous to chartaceous, abundantly scaly abaxially, scales on the laminar tissue circular and 0. 1 mm in diameter, or with a cusp as long as the scale body. Rachis and costae lighter in color than the laminar tissue, stramineous throughout (or ra- chis darker near the base), provided with a few, scattered, lanceolate, clathrate scales to 1 mm long. Ultimate segments 4-12 mm broad, fertile por- tions sometimes slightly contracted, the margins plane, not revolute. Veins indistinct, but com- monly evident when held to light, 1-2-forked, free. Sporangia grouped in round or oblong, discrete sori (occasionally some of these confluent on very mature leaves), borne on distal portions of seg- ments. Pendent on trees and shrubs in thickets and for- ests, 600-800 m, Tumbes. Western Ecuador; northern Peru. Dicranoglossum polypodioides is very similar to D.furcatum (L.) John Sm., s.s. (West Indies, north- ern South America) and D. panamense (C. Chr.) Gomez (Central America, northern South Amer- ica). The diagnostic characters separating these are mostly variable: D. polypodioides has discrete sori, mostly stramineous midribs, and abundant lam- inar scales; D. panamense has linear coenosori, mostly stramineous midribs, and abundant lam- inar scales; and D. furcatum has linear coenosori, mostly dark midribs, and more scattered laminar scales. Some gradation toward either discrete or confluent sori has been observed in the former two species, and relative frequency of scales is not a strong diagnostic character in the latter two. Thus, D. panamense seems to be intermediate between the others, but further study is needed to deter- mine whether the three taxa are distinct or merely variable forms of one species. Tumbes: Prov. Zarumilla, Dist. Matapalo, Bosque Na- tional de Tumbes, Schunke V. 2373 (F, GH), Simpson & Schunke V. 482 (F, GH). VI. Microgramma Microgramma Presl, Tent, pterid. 213. 1836. TYPE: Microgramma persicariifolia (Schrad- er) Presl (Polypodium persicariifolium Schrader). Figure 10. Plants epiphytic or epipetric. Stem long-creep- ing, usually somewhat flattened, bearing peltately 148 FIELDIANA: BOTANY attached, nonclathrate scales and few to many fi- brous roots. Leaves monomorphic to dimorphic, articulate on short phyllopodia, widely spaced. Lamina simple and essentially entire, glabrous to slightly pubescent, sparsely to abundantly scaly, the scales not or scarcely clathrate, peltately at- tached, variously shaped, but not circular. Veins free to anastomosing, areoles lacking or (usually) with free, included veinlets. Sori round, or elon- gate in a few species, borne in a single row on each side of the costa on or usually at the tip of a vein or junction of veins, on a scarcely to somewhat raised receptacle, paraphyses (if present) not pel- tate. Spores monolete, ellipsoid, with long laesura. This ill-defined genus contains about 20 Amer- ican species, plus one or two more in Africa, Mad- agascar, and the Mascarenes. There are 1 4 known from Peru. A number of species have goniophle- bioid venation, as in Polypodium, and there is also a close relationship with Pleopeltis. Monographic study is sorely needed to ascertain proper rela- tionships. For further elaboration of the problems, see discussion under the genus Pleopeltis, and treatment of P. fuscopunctata. Key to Species of Microgramma a. Scales ample and conspicuous on costa and laminar tissue, at least abaxially, sometimes long and hairlike from an expanded base b b. Leaves not or only slightly dimorphic c c. Veins, especially of sterile leaves, free; laminar scales ovate to lanceolate, acuminate to atten- uate, but not abruptly filiform-tipped; lamina base attenuate 1 . M. chrysolepis c. Veins anastomosing, forming l(-2) series of areoles between costa and margin; laminar scales mostly filiform from an abruptly expanded base; lamina base obtuse or cuneate 2. M. latevagans b. Leaves conspicuously dimorphic, the sterile 2-6 times broader than fertile ones d d. Scales on abaxial lamina surface whitish, filiform and tortuous, most of them stellately branched 3. M. vacciniifolia d. Scales on abaxial lamina surface orange to castaneous, tips rigid and setiform, or filiform and rather straight, not tortuous e e. Fertile lamina 1.5-2 mm broad, mature sori commonly projecting beyond the margin; stem scales somewhat appressed to spreading 5. M. reptans e. Fertile lamina 3-6 mm broad, sori well contained within the margins; stem scales either tightly appressed or obviously spreading f f. Sterile lamina 4-8.5 cm long; laminar scales linear-deltate from a scarcely expanded base; stem scales tightly appressed, the tips not filiform, or if slightly filiform, then not or rarely spreading 4. M. piloselloides f. Sterile lamina 0.8-2(-2.5) cm long; lamina scales setiform from an abruptly expanded base; stem scales spreading, the tips filiform 6. M. tecta a. Scales essentially lacking on laminar tissue between costa and margin g g. Fertile leaves (1.3-)2-9 cm broad, broadly lanceolate or elliptic, the apex acuminate to attenuate h h. Stem 10—40 mm broad; sori mostly inframedial; leaves 24—40 cm long 13. M. megalophylla h. Stem 1-3 mm broad; sori mostly medial; leaves to 2 1 cm long . . . i i. Sori elongate to linear; primary veins much thicker and more prominent than the others 14. M. persicariifolia i. Sori round; primary and other veins equally prominent .12. M. thurnii g. Fertile leaves 0.2-1(-1.5) cm broad, linear to linear-lanceolate, the apex obtuse to subacute, rarely acute J j. Leaves not or only slightly dimorphic; included free veinlets both recurrent and excurrent within areoles • ^ k. Petiole 5-15 mm long; stem commonly more than 2 mm (to 5 mm) in diameter, the scales TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 149 150 FIELDIANA: BOTANY J. whitish from a castaneous base, with an attenuate tip; sori medial to (mostly) inframedial 7. M. rosmarinifolia k. Petiole obsolete; stem commonly less than 2 mm in diameter, the scales orange to castaneous (sometimes aging whitish), with a filiform tip; sori medial to (mostly) supramedial 1 1. Veins (at least primary ones) distinct, more or less prominulous; sori not or slightly immersed; lamina chartaceous to subcoriaceous, plane 8. M. lycopodioides 1. Veins all obscure (at least abaxially); sori deeply immersed; lamina coriaceous, the margins mostly strongly revolute 9. M. baldwinii Leaves conspicuously dimorphic, sterile ones 2-5 times as broad as the fertile and often some- times shorter; included free veinlets lacking or few and only excurrent within areoles .... m m. Sterile leaves 1— 4(-6) cm long, circular to broadly elliptic, the base cuneate 3. M. vacciniifolia m. Sterile leaves (4-)5-l 1 cm long, lanceolate to narrow-elliptic, the base attenuate n n. Costa scales broad; sori medial; stem scales ciliate 10. M. squamulosa n. Costa scales mostly filiform; sori subcostal to inframedial; stem scales denticulate .... 11. M. ulei 1. Microgramma chrysolepis (Hooker) Crabbe, Brit. Fern Gaz. 9: 316. 1967. Polypodium chrysolepis Hooker, Icon. pi. 8: /. 727. 1845. TYPE: Ecuador, Andes of Quito, Jameson 37 (in error in the protologue as "73") (holotype, K!; photo, us). Plants epiphytic, or sometimes epipetric. Stem (excluding scales) to 1 mm in diameter, provided with dense, spreading scales, these 4-6 mm long, linear-lanceolate and attenuate to a filiform tip, peltately attached above the base, orange, or whit- ish in age, with a castaneous point where attached, margins entire to denticulate. Leaves chartaceous to subcoriaceous, abundantly scaly throughout, es- sentially monomorphic, subsessile to conspicu- ously stalked. Petiole 2-12 mm long. Lamina lan- ceolate or elliptic-lanceolate, 3-10(-14) cm long, 0.6-1.7 cm broad, apex acute, or more commonly subacute, base attenuate, margin entire to slightly sinuate, scales ovate to lanceolate, acuminate to attenuate. Veins (especially of sterile leaves) free, 1-2-forked, indistinct or obscure. Sori round or slightly elongated, more or less medial, sporangia intermixed with, and sometimes covered by, ovate, acuminate, orange scales. In and at edges of dense or open forests, on tree trunks and branches, sometimes on mossy boul- ders, (1800-)2600-4100 m, Amazonas, La Lib- ertad, Ancash, Huanuco, and Cuzco. Ecuador; Peru; Bolivia; Argentina. This is the only species of Microgramma in Peru with predominantly free veins. The character of venation is the principal reason it is maintained in the genus Polypodium by Sola and Perez-Garcia (Biotica 7: 45-64. 1982). It occurs mostly above 2500 m, although Macbride 3826 Huanuco (cited below) was collected at 1800 m. Amazonas: Approaches to Cerro Campanario NNE of Diosan, Wurdack 1592 (GH, us). La Libertad: Prov. Hua- machuco, between Pallar and Huaguil, Lopez & Sagds- tegui 8129 (F, HUT). Ancash: Prov. Huari, Huascaran National Park, D. Smith et al. 12415 (F, HUT, MO), 12648 (F, MO). Huanuco: Yanano, Macbride 3826 (F, us). Cuzco: Huayoccari to Yanococha, Urubamba, Nunez et al. 6988 (F, MO). Prov. Urubamba, Hacienda Huayockayi, Vargas 14983 (GH). 2. Microgramma latevagans (Maxon) Lell., Amer. FernJ. 74: 59. 1984. Polypodium latevagans Maxon, Proc. Biol. Soc. Wash. 52: 120. 1939. TYPE: Bolivia, Unduavi, Rusby 361 (holotype, us!; isotype, NY; photo, F of us). Plants epiphytic, rarely epipetric. Stem (exclud- ing scales) less than 1 mm in diameter, provided with dense, spreading scales, these 4-6 mm long, linear-lanceolate and attenuate to a filiform tip, peltately attached above the base, orange, often castaneous at base, or whitish in age, margins en- tire to denticulate. Leaves chartaceous, abundant- FIG. 10. Microgramma thurnii; a, habit. Microgramma reptans: b, habit, fertile and sterile leaves; c, lamina scales. Microgramma squamulosa: d, base of sterile lamina, (a from S. F. Smith et al. 981. GH; b from Klug 366, F, c from Killip & Smith 22951, GH; d from Soukup & Lopez 14751. GH.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 151 ly scaly throughout, essentially monomorphic, or fertile ones slightly smaller, short- to long-stalked. Petiole 3-15 mm long. Lamina ovate to narrow- lanceolate, 1-5 cm long, 0.5-1.5 cm broad, apex acute or subacute, base obtuse or cuneate, margin entire, scales mostly filiform from an abruptly ex- panded base. Veins merging to form 1 (-2) rows of areoles on each side of the costa, indistinct or ob- scure. Sori round or elongated, often confluent at maturity, inframedial or subcostal, sporangia in- termixed with filiform orange scales. In forests, growing on tree trunks and branches, very rarely on rocks, (200-)650-2250(-2420) m, Cajamarca, San Martin, Loreto, Pasco, Puno. Besides the characters in the key, this also can be distinguished from M. chrysolepis by the son, which usually crowd the costa, whereas sori of the latter are essentially medial. Also, M. chrysolepis grows at much higher elevations, usually above 2500 m, while M. latevagans usually occurs below 2300 m. Cajamarca: Prov. Cutervo, Parque Nacional San Andres, Lopez & Sagdstegui 5551 (GH, HUT, uc). Prov. Santa Cruz, above Charro Blanco, Sagdstegui et al. 13008 (F, HUT). San Martin: 23 km S of Nuevo San Martin, Gentry et al. 37604 (F, MO). Loreto: Prov. Maynas, near Brilla Nueva on upper Rio Yaguasyacu, Gentry & Revilla 20472 (MO, USM). Pasco: Palcazu Valley, Iscozacin, Sal- ick 7135 (F). Puno: Prov. Sandia, near Sandia, Vargas 14801 (GH). 3. Microgramma vacciniifolia (Langsd. & Fisch.) Copel., Gen. fil. 185. 1947. Polypodium vacciniifolium Langsd. & Fisch., Icon. fil. 8. 1810. TYPE: Brazil, Santa Catarina Island, Langsdorff 6/7 (holotype, LE; isotype, B, Herb. Willd. 19600; photos, F & GH of B). Plants epiphytic, rarely epipetric. Stem 3-5 mm in diameter, completely covered by appressed, densely imbricate scales, or the tips often spread- ing, the scales 3-7 mm long, lanceolate, attenuate to a filiform tip, peltately attached above the base, castaneous, usually with a blackish point where attached, tips orange to tawny, or whitish in age, margins denticulate. Leaves chartaceous to sub- coriaceous, moderately scaly along costa abaxially, laminar surface abaxially naked or often with scat- tered to abundant, tortuous, filiform scales, con- spicuously dimorphic, sessile or subsessile. Petiole 0-2 mm long. Lamina obtuse to subacute, fertile ones 3-10 cm long, 0.2-0.4(-0.6) cm broad, linear, with attenuate base, sterile ones l-4(-6) cm long, 0.8-1.7 cm broad, circular to broadly elliptic or oblong-elliptic, cuneate at base. Veins areolate, in- cluded free veinlets lacking, or in sterile leaves these present and excurrent, indistinct, not or scarcely prominulous. Sori round, medial or in- framedial, sporangia usually intermixed with long, pale, filiform paraphyses. Rare in Peru, in forests, on tree trunks or on rocks, 480-1100 m, Cajamarca, Amazonas, Huancavelica. Jamaica; Grenada; Trinidad & Tobago; Vene- zuela and Columbia south to Paraguay and Ar- gentina. On the lamina are usually found what appear to be pale trichomes but which are actually filiform scales. These are attached by a nearly circular, peltate base and are abruptly reduced to a tortuous, hairlike scale body. They are commonly on the costa and intermixed among the sporangia, but on a few specimens they are abundant over the ab- axial surface of the lamina. Those on the laminar surface are often stellately branched from the base, with several greatly elongated and tortuous arms appearing like very thin tentacles. Apparently they are easily abraded, for on some mature specimens they are virtually lacking. Cajamarca: Colasay, Woytkowski 6938 (MO, us). Ama- zonas: Prov. Bongara, between Bongara Grande and El Ingenio, Sagdstegui 5949 (GH, HUT). Prov. Bongara, Ja- zan, above Rio Utcubamba, Young et al. 500 (MO, uc). Huancavelica: Prov. Tayacaja, above Virgen-pampa, E of Tintay, Tovar 4601 (GH). Prov. Tayacaja, mouth of Imaybamba, SE of Tintay, Tovar 4613 (GH). 4. Microgramma piloselloides (L.) Copel., Gen. fil. 185. 1947. Polypodium piloselloides L., Sp. pi. 2: 1083. 1753. TYPE: "Habitat in America Meridionali" (prob- ably from Martinque) (holotype, LINN 1251.3). Microgramma acatallela Alston, J. Wash. Acad. Sci. 48: 232. 1958. TYPE: Columbia, Putumayo, Rio Mocoa drainage, Ewan 16706 (holotype, BM; is- otypes, GH!, MO, uc!, us!). Plants epiphytic. Stem 0.6-1 mm in diameter, amply provided with and sometimes obscured by tightly appressed scales, these 2-4 mm long, pel- tately attached near the base, broadly lanceolate and often attenuate but the tips not filiform or spreading, orange or tawny, with a castaneous point where attached, margins subentire. Leaves char- taceous, amply scaly, conspicuously dimorphic (at least in Peru), short-petiolate. Petiole 0.3-0.6 cm long. Sterile lamina 4-8.5 cm long, 1-1.8 cm broad, 152 FIELDIANA: BOTANY lanceolate or oblong-lanceolate, apex acute or sub- acute, base cuneate, provided throughout with minute, appressed scales, these orange to tawny, often whitish on the adaxial surface, linear-deltate from a slightly expanded peltate base to a setiform tip; veins indistinct to obscure, merging to form 1 (-2) rows of areoles on each side of the costa, costal areoles each with a free, excurrent veinlet. Fertile lamina linear to very narrow-lanceolate, attenuate at base, (2.5-)4-10 cm long, 0.3-0.6 cm broad, son round, inframedial, sporangia inter- mixed with abundant, orange or ferruginous, fili- form scales. In wet forests, on tree trunks and branches, 1 00- 700 m, Amazonas, San Martin, and Loreto. Southern Mexico (Chiapas); Guatemala; West Indies; Columbia to Guyana, south to Bolivia and Brazil. In addition to the characters of the key, this usually can be distinguished from M. tecta and M. reptans by the often whitish scales on the adaxial lamina surface. Adaxial scales of the other two are orange to castaneous. Stem scales of M. pilosel- loides are also diagnostic in this species complex. Most of them are broad and tightly appressed to the stem from base to tip, whereas scales of M. reptans are only loosely appressed, with widely spreading tips, and those of M. tecta are filiform and strongly spreading. The shape of fertile leaves of M piloselloides is variable in the West Indies and Central America, in some plants conspicuously longer and narrower than the sterile ones, but in others less obviously so. Alston separated M. acatallela from M. pilo- selloides based on strong vs. slight dimorphism, but he pointed out no other substantiating char- acters. Mexia 6144b (Loreto) was cited as a para- type of M. acatallela. Amazonas: Prov. Bagua, Rio Maranon opposite Que- brada Mirana, Wurdack 2035 (us). San Martin: Prov. Lamas, trail to Rio Tiriyacu and Rio Cashiyacu, Knapp & Mallet 7209 (F, MO). Loreto: San Antonio, on Rio Ataya, Killip & Smith 29356 (F, us). Mishayacu, near Iquitos, Klug 2539 (F, GH, us). Pongo de Manseriche, bank of Rio Santiago, Mexia 6144b (BM, GH, us). 5. Microgramma reptans (Cav.) A. R. Smith, Proc. Calif. Acad. Sci. IV, 40: 230. 1975. Figures lOb-c. Acrostichum reptans Cav., Anales Hist. Nat. 1: 104. 1 799. TYPE: Ecuador, Prov. Guayas, Guayaquil, Nee (holotype, MA). Polypodium ciliatum Willd., Sp. pi. ed. 4, 5: 144. 1 8 10. TYPE: Brazil, Para, Hoffmannsegg (holotype, B, Herb. Willd. 19601- photo, GH). Microgramma ciliata (Willd.) Alston, Bull. Jard. Bot. Etat. 27: 56. 1957. Plants epiphytic, rarely epipetric. Stem 0.4-0.7 mm in diameter, amply to moderately provided with (but not obscured by) weakly appressed to spreading scales, these 2-4 mm long, linear-lan- ceolate and attenuate to a filiform tip, peltately attached above the base, orange to castaneous, margins essentially entire. Leaves chartaceous, amply scaly, conspicuously dimorphic, subsessile to short-petiolate. Petiole 0.2-0.7 cm long. Sterile lamina 1.5-5 cm long, 0.8-1.5 cm broad, ovate, elliptic, oblanceolate or oblong, apex acute or sub- acute, base narrowly to broadly cuneate, rarely obtuse, provided throughout with minute, ap- pressed scales, these mostly orange to castaneous, setiform to abruptly attentuate from an expanded, peltate base; veins obscure, merging to form l(-2) rows of areoles on each side of the costa, costal ones with a free, excurrent veinlet. Fertile lamina linear, attenuate at base, 2-5(-7) cm long, 0. 1 5- 0.22 cm broad, sori round, at maturity filling the lamina between costa and margin or, more com- monly, projecting well beyond the margin, spo- rangia amply intermixed with orange, acicular scales. In or at edges of forests, on trunks or branches of trees, once reported (Macbride 5582, Junin) growing on a rocky cliff, 100-650(-1200) m, San Martin and Loreto south to Ayacucho and Puno. Southern Mexico to Panama; Trinidad and To- bago; Columbia to the Guianas, south to Bolivia and Brazil. The large sori projecting over the margins of the vary narrow fertile lamina easily distinguishes this from the other species of Microgramma in Peru, but specimens containing only sterile laminae could be confused with those of M. tecta. However, in the latter the sterile lamina is rarely more than 2 cm long and the base is usually truncate, or even subcordate, and the stem scales are mostly con- spicuously spreading. In M. reptans the sterile lamina is commonly over 2 cm long and with a cuneate base, and stem scales are usually some- what appressed in the proximal half. San Martin: Prov. Lamas, km 2 of road from Pongo de Cainarache to Barranquita, Knapp 8150 (F, MO). Lo- reto: Bank of Rio Santiago above Pongo de Manseriche, Mexia 6144 (F, GH, uc, us). Huanuco: Prov. Pachitea, TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 153 Dist. Honoria, Bosque Nacional de Iparia, Schunke V. 2010 (F, OH). Pasco: Prov. Oxapampa, Quebrada Castilla on Rio Omaiz, Leon & Young 101 9a (MO, USM). Junin: La Merced, Macbride 5582 (F, GH, us). Ayacucho: Rio Apurimac Valley, near Kimpitiriki, Killip & Smith 22951 (F, GH, us). Cuzco: Rio Tambopata, 2 km E of Colpa de Guacamayos, Nunez 6516 (MO). Madre de Dios: Prov. Tambopata, Albergue Cuzco Amazonico, Leon 859 (USM). Puno: Below San Gaban on Rio San Gabon, Dillon et al. 1223 (F, MO, USM). 6. Microgramma tecta (Kaulf.) Alston, J. Wash. Acad. Sci. 48: 232. 1958. Polypodium tectum Kaulf., Enum. fil. 87. 1824. TYPE: Brazil, Ilha de Santa Catarina, Chamisso (LE). Plants epiphytic. Stem 0.3-0.5 mm in diameter, amply provided with (but not obscured by) spread- ing scales, these 2-3 mm long, peltately attached near the base, linear and gradually attenuate to a filiform tip, orange to castaneous, with a blackish point of attachment, margins entire. Leaves firm- herbaceous to chartaceous, amply scaly, conspic- uously dimorphic, short- to long-petiolate. Petiole 0.2-2 cm long. Sterile lamina 0.8-2(-2.5) cm long, 0.5-1 .3 cm broad, ovate to oblong-elliptic, to near- ly circular, apex obtuse or subacute, base broadly cuneate, truncate or subcordate, provided throughout with minute, appressed scales, these orange, and setiform from an abruptly expanded, nearly circular base; veins obscure, merging to form 1 (-2) row of areoles on each side of the costa, costal areoles with a free, excurrent veinlet. Fertile lam- ina linear to very narrow-elliptic or -oblanceolate, attenuate at base, 1 .6-5 cm long, 0.3-0.5 cm broad, sori round, inframedial, sporangia intermixed with abundant, orange, long, filiform scales. In forests, on tree trunks and branches, or twin- ing among lianas, 100-1350 m, Amazonas, San Martin, Loreto, Huanuco, and Junin. Columbia to the Guianas, south to Bolivia and Brazil. Some specimens were separated by Mickel and Beitel (Pterid. Fl. Oaxaca 251.1 988) as var. nana, said to differ from var. tecta by somewhat nar- rower lamina scales. This, M. piloselloides, and M. reptans form a closely knit complex of species with scaly, strongly dimorphic laminae. See treatments of the others for further comparisons. Amazonas: Prov. Bagua, between Aramango and Montenegro, Rio Maranon, Lopez et al. 4164 (GH, HUT), 4222 (GH, HUT). Prov. Bagua, Rio Maranon opposite Quebrada Mirana, Wurdack 2037 (F, GH, uc, us). San Martin: San Roque, LI. Williams 7531 (F, us). Loreto: Soledad, on Rio Itaya, Killip & Smith 29806 (F, GH, us). Prov. Maynas, Dist. Pebas, Rio Ampiyacu, Revilla 864 (F, MO, us, USM). Huanuco (as San Martin): Tingo Maria, Allard 22569 (GH, us). Junin: Colonia Perene, Killip & Smith 25086 (us). 7. Microgramma rosmarinifolia (Kunth) R. & A. Tryon, Rhodora 84: 129. 1982. Polypodium rosmarinifolium Kunth, Syn. pi. 1: 71. 1822. TYPE: Peru?, "Crescit in crepidinibus et ad arbores locis frigidis, alt. 1300 hex.," Kunth (holotype, p; photos, GH, us). Polypodium kunthii Desv., Prodr. 226. 1827. TYPE: type citation the same as for Polypodium ros- marinifolium, collector unspecified, but probably Kunth (holotype, P; photos, GH, us). This frag- ment very likely was cut from the Kunth type of P. rosmarinifolium. See Weatherby for discussion (Contr. Gray Herb. 124: 17. 1939). Plants epipetric, occasionally epiphytic. Stem 2- 5 mm broad, completely covered by appressed, imbricate scales, these with tips spreading, 3-6 mm long, lanceolate, attenuate at apex, dull white, but castaneous at base, margins entire, or distally remotely ciliolate. Leaves 5-14(-20) cm long, 0.2- 0.7(-1.0) cm broad, short-stalked, essentially monomorphic. Petiole 5-15 mm long. Lamina na- ked, subcoriaceous (rarely chartaceous), linear (rarely narrow-elliptic), obtuse to subacute, occa- sionally acute, the base attenuate. Veins obscure, areolate, the supracostal areoles each usually with a free, excurrent, veinlet, this bearing the sorus. Sori round, often slightly impressed, medial to inframedial, paraphyses sparse or lacking. In forest, growing on or among rocks, or on tree trunks, Cajamarca, La Libertad, Cuzco. Ecuador; Peru. This species thus far is represented in herbaria by only a few collections. All but one were growing on rocks. The only epiphyte (Sagdstegui et al. 10892) is much larger, with leaves to 20 cm long and 1 cm broad, and of thinner texture than the typical specimens. Cajamarca: Prov. Contumaza, El Molino, Sagdstegui et al. 10892 (F, HUT). La Libertad: Prov. Otuzco, road Casmiche to Otuzco, Sagdstegui 6268 (GH, HUT). Cuzco: Valle de Paucartambo, Herrera 3001 (us). Prov. Pau- cartambo, Chacclabamba, Vargas 9453 (GH). 8. Microgramma lycopodioides (L.) Copel., Gen. fil. 185. 1947. Polypodium lycopodioides L., Sp. pi. 2: 1082. 1753. TYPE: Probably West Indies, collector unknown 154 FIELDIANA: BOTANY (holotype, LINN 1251.2). For a comprehensive list of basionyms, see Lellinger, Pteridologia 2 A: 250. 1989. Plants epiphytic. Stem 1.5-2 mm in diameter, completely covered by appressed, densely imbri- cate scales, these 3-6 mm long ovate to lanceolate- attenuate, with a filiform, often deciduous tip, pel- tately attached above the base, castaneous to deep orange, sometimes whitish in age, usually with a black point where attached, margins setulose or sometimes ciliate. Leaves chartaceous to subcor- iaceous, naked, monomorphic to slightly dimor- phic (sterile ones a little broader), sessile, the pet- iole essentially obsolete. Lamina obtuse to subacute, rarely acute, (4.5-)6-14 cm long 0.6-1.3 cm broad, linear or linear-lanceolate, with atten- uate base, the margins plane. Veins areolate, pri- mary ones (at least) distinct and more or less prominulous, included free veinlets spreading in all directions. Sori round, medial or slightly su- pramedial, sporangia usually intermixed with oc- casional orange paraphyses. In rain forests, climbing among shrubs or on branches and trunks of trees, 700-2000 m, Ama- zonas to Loreto, south to Cuzco and Puno. West Indies; southern Mexico to Panama; Co- lumbia to the Guianas, south to Bolivia and Brazil; Africa?. Microgramma baldwinii is very similar to this and might be considered a geographic and eleva- tional variant. Throughout the range of distribu- tion the species complex varies markedly in lam- ina width and texture, which apparently affects the ways in which sori and veins are immersed in the tissue and the way the margins are inrolled. In the West Indies the lamina is often firm-herbaceous and 2 cm broad, with margins perfectly plane and the veins conspicuous, and sori are clearly super- ficial. At the other extreme, in northern South America and at lower elevations in Peru (M. bald- winii), the lamina is much narrower, nearly cori- aceous, with margins revolute, and veins and sori deeply immersed in the tissue. In central American specimens of M. lycopodioides, the lamina more closely resembles that in the West Indies, whereas at higher elevations in South America it approach- es the character of M. baldwinii. Further study is needed, but the two taxa are treated here as distinct species. Amazonas: Prov. Bagua, between Montenegro and Chiriaco, Sagdstegui 5926 (GH, HUT). San Martin: Prov. Lamas, Dist. Lamas, below English Evangelical Mission, Belshaw 3427 (F, GH, MO, uc, us). Loreto: Sierra del Pongo, Mexia 6276a (GH, uc, us). Huanuco: Prov. Hua- nuco, Dist. Churubamba, Hacienda Exito, Mexia 8227 (F, GH, uc, us). Pasco: Prov. Oxapampa, Abra los Mel- lizos, 4-8 km from Enenas, Skoget al. 5045, in part (us). Junin: Chanchamayo Valley, C. Schunke 1545 (F, us). Cuzco: Paucartambo, between Chontachaca and Pilla- huata, Nunez 8107 (F, MO). Puno: Below San Gaban on Rio San Gaban, Dillon el al. 1211 (F, USM). 9. Microgramma baldwinii Brade, Arch. Jard. Bot. Rio Janeiro 18: 30, t. 1. 1965. TYPE: Brazil, Amazonas, Marabitantas, Rio Negro, Bald- win 3229 (holotype, IAN; isotype, us). Plants epiphytic, very rarely terrestrial. Stem long-creeping, 1-2 mm in diameter, completely covered by appressed, densely imbricate scales, these 3— 4(-5) mm long, ovate to lanceolate-atten- uate, with a filiform, often deciduous tip, peltately attached above the base, castaneous to deep or- ange, sometimes whitish in age, usually with a blackish point where attached, margins entire to setulose. Leaves simple, entire, coriaceous, naked, monomorphic to slightly dimorphic (sterile ones a little broader), sessile, the petiole essentially ob- solete. Lamina obtuse to subacute, rarely acute, (4-)6-12 cm long, 0.4-1 cm broad, linear to nar- row-oblong, with attenuate base, the margins strongly revolute. Veins areolate, obscure, im- mersed in the tissue, included free veinlets both recurrent and excurrent within areoles. Sori round, borne in a single supramedial row on each side of the costa, sporangia usually intermixed with short, orange to castaneous paraphyses. In forests, climbing on trunks and branches of trees, in a single collection terrestrial (Wurdack 1943), 100-850 m, Amazonas, San Martin, Lo- reto, Huanuco, Junin, Ucayali, and Cuzco. Venezuela; Ecuador; Peru; Amazonian Brazil. This is very similar to M. lycopodioides, and it is questionable if they are truly distinct species. See treatment of the latter for further discussion. Amazonas: Prov. Bagua. between Aramango and Montenegro, Lope: et al. 4213 (GH, HUT). Prov. Bagua, valley of Rio Maranon near Cascadas de Mayasi, Wur- dack 1943 (F, us). San Martin: Lamas. Santa Rosa de Davidcillo. Knapp & Mallet 8467 (F, MO). Mariscal Ca- ceres, Tocache Nuevo, Schunke V. 5744 (F, us). Loreto: Mishuyacu. near Iquitos, Killip & Smith 29901 (us). Klug 11 94 (F, us). Huanuco: Prov. Leoncio Prado, Dist. Rupa Rupa, E of Jingo Maria. Schunke, I'. 10591 (F). Junin: Satipo. Ledn 276 (USM). Ucayali: Prov. Requena, Jenaro Herrera, V&squez et al. 2099 (F, MO). Cuzco: Prov. Pau- cartambo, Cosnipata Valley, N of Patria. Foster et al. 10617 (F). TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 155 10. Microgramma squamulosa (Kaulf.) Sola, Opera Lilloana 5: 59. 1961. Figure lOd. Polypodium squamulosum Kaulf., Enum. fil. 89. 1824. TYPE: Brazil, Ilha de Santa Catarina, Chamisso (LE). Plants epiphytic or epipetric. Stem 2.5-4 mm in diameter, completely covered by appressed, im- bricate scales, or the tips of these often spreading, the scales 3-6 mm long, lanceolate, attenuate to a filiform tip, peltately attached above the base, cas- taneous, with a blackish point where attached, tips usually whitish, often delicate and worn away, margins usually ciliate. Leaves chartaceous to sub- coriaceous, naked except for scattered, broad, cil- iate scales on costa, conspicuously dimorphic, sub- sessile to short-stalked. Petiole 1-10 mm long. Lamina obtuse to subacute, the base attenuate, fertile ones 5-12 cm long, 0.2-1 cm broad, linear, sterile ones 4-10 cm long, 1-2 cm broad, lanceo- late to narrow-elliptic. Veins areolate, included free veinlets few and excurrent or lacking, primary veins distinct and prominulous, the rest indistinct. Sori round, medial, sporangia intermixed with abun- dant, filiform paraphyses. In forests, climbing on tree trunks or scrambling among rocks, 850-2500 m, Apurimac, Cuzco, and Puno. Peru; Bolivia; SE Brazil; Argentina; Paraguay; Uruguay. Stem scales of this species are varied. At their based they are deep castaneous to nearly black, lustrous, rigid and with entire margins. But toward the center they become dull, whitish and nearly translucent, with margins long- and laxly ciliate. On older parts of the stem the delicate, whitish tips often are worn away, leaving only the firm, appressed darker portions. Thus, if a particular dried specimen contains only an older section of the stem, and another specimen only the growing tip, the two might be interpreted as representing different species. Apurimac: Rio Huacchac, Hacienda Sondor, Bites 1390 (us). Cuzco: San Miguel, Urubamba Valley, Cook & Gil- bert 907 (us), 1753 (us). Prov. La Convention, Rio Ma- pillo Above Pomobamba, Davis et al 1259 (GH). Quil- labamba, Soukup 161 (F). Prov. La Convencion, Huadquina, Vargas 13909 (GH). Puno: Sandia, Soukup & Lopez 14751 (GH), Vargas 11867 (GH), 74792 (GH). 1 1 . Microgramma ulei (Ule) Stolze, comb. nov. Polypodium ulei Ule, in Karsten & Schenk, Vegeta- tionsbilder 2(1), t. 3. 1 904. TYPE: Peru, San Mar- tin (as Loreto), Tarapoto, Spruce 4127 (holotype, B!; isotype, K!; photo, F of B, GH of K). Plants epiphytic. Stem 1.5-3 mm in diameter, completely covered by appressed, imbricate scales, or the tips often spreading 3-6 mm long, lanceo- late, attentuate to a filiform tip, peltately attached above the base, castaneous, usually with a blackish point where attached, tips orange to tawny or whit- ish, delicate and sometimes worn away, margins denticulate. Leaves chartaceous, naked, or more often with scattered, white, hairlike scales on the costa, conspicuously dimorphic, subsessile. Peti- ole 0-3 mm long. Lamina obtuse to subacute, the base attenuate, fertile ones 8-1 5 cm long, 0.2-0.7(- 0.9) cm broad, linear, sterile ones 7-1 1 cm long, 1-1.8 cm broad, lanceolate or narrow-elliptic. Veins areolate, primary ones distinct but not prominu- lous, included free veinlets lacking, or in sterile leaves few and excurrent, indistinct. Sori round, inframedial to subcostal, sporangia intermixed with long, pale, filiform paraphyses. In forests, climbing on shrubs or tree trunks, 360-900 m, San Martin. Endemic and rare. This was originally described and typified in an unpublished manuscript by Hieronymus and later published by Christ (Hedwigia 44: 363. 1905). However, Ule's publication of it in 1 904, although poor and short, is nevertheless valid. We are in- debted to Dr. Brigitte Zimmer at Berlin-Dahlem for bringing this to our attention (in litt.). San Martin: Tarapoto, Alto Puca Yacu, Monies 53 (F). Alto Rio Huallaga, LI. Williams 6642 (F, us). Juan Guerra, LI. Williams 6857 (F, us). Spruce 4127 and Ule 6600, both from Tarapoto and cited by Christ, are doubt- less this species. 1 2. Microgramma thurnii (Baker) Tryon & Stolze, comb. nov. Figure lOa. Drynaria acuminata Fee, Crypt, vase. Bresil 1: 122, t. 137,f. 3. 1869 (not Brack. 1854). TYPE: Brazil, Amazonas, Prov. Rio Negro, vicinity of Barra, Spruce in 1850-1851 (holotype, RB). Polypodium thurnii Baker, Ann. Bot. (London) 5: 476. 1891. LECTOTYPE (designated here): Guyana (as British Guiana), Ankers, in 1829 (K!); PARA- TYPE: Jenman, in June " 1 895" (K!). Jenman date obviously incorrect! Polypodium loretense Maxon, Amer. Fern J. 23: 105. 1933. TYPE: Peru, Loreto, Balsapuerto, Killip & 156 FIELDIANA: BOTANY Smith 28602 (holotype, us!; paratypes, us, nu- merous). Microgramma acuminata Lellinger, Amer. Fern J. 74: 59. 1984. Plants epiphytic. Stem 1 .2-2.5 mm in diameter, completely covered with scales, these appressed or with spreading tips, 4—6 mm long, lanceolate, acu- minate or attenuate to a filiform tip, peltately at- tached well above the base, orange to reddish brown, with a blackish or castaneous point of at- tachment, the margins entire. Leaves 10-21 cm long, (1.8-)2-4.5 cm broad, subsessile, essentially monomorphic. Petiole 0-3 mm long. Lamina firm- herbaceous to chartaceous, elliptic, attenuate at apex and base, glabrous. Veins distinct, primary ones no more prominent than the others, areolate, often with a recurrent, free veinlet in barren are- oles. Sori superficial, round, medial. Climbing on trees in dense forests, 100-450 m, San Martin, Loreto, Pasco, Ucayali, and Madre de Dios. Guyana; Surinam; Venezuela; Peru; Amazonian Brazil. This is often confused with three other species: M. recreense(Hieron.) Lell. (Ecuador) has a thicker stem (to 4 mm), with bicolorous, erose-ciliolate scales; the lamina of M. lindbergii (Mett.) Sota (Brazil) is rather amply pubescent; and M. gemi- nata (Schrader) R. & A. Tryon (Brazil) has nar- rower leaves, impressed sori, and abundant free included veinlets spreading in various directions. San Martin: Lamas, trail to Tioyacu and Nuevo La- mas, Knapp & Mallet 7119 (F, MO, uc). Loreto: Balsa- puerto, lower Rio Huallaga, Killip & Smith 28392 (us), 28500 (us); Mishuyacu, near Iquitos, Killip & Smith 29890 (GH, us), Klug 553 (F, us). Prov. Maynas, Iquitos, Sagdstegui & Aldave 5796 (HUT). Pasco: Puerto Ber- mudez (as Junin), Killip & Smith 26556 (us). Prov. Ox- apampa, Valle del Palcazu, Iscozacin, Leon 696 (F, GH, USM). Ucayali: Prov. Coronel Portillo, Bosque von Hum- boldt, Young & Grandez 1041 (F, MO). Madre de Dios: Tambopata Nature Reserve, Funk et al. 8155 (F). Main trail from Explorer's Inn, near confluence of Rios Tam- bopata and La Torre, S. F. Smith et al. 186 (us), 957 (GH, uc). 13. Microgramma megalophylla (Desv.) Sota, Bol. Soc. Argent. Bot. 10: 158. 1963. Polypodium megalophyllum Desv., Prodr. 227. 1827. LECTOTYPE (designated by Weatherby, Contr. Gray Herb. 114: 32. 1939): Brazil, Rio Negro, collector unknown (p, Herb. Desvaux). Plants epiphytic. Stem commonly flattened and appressed to tree trunks, 10-35 mm broad, com- pletely covered with appressed scales, these 4—7 mm long, lanceolate, attenuate to a filiform tip, peltately attached near the base, bicolorous, tawny to whitish with a castaneous center stripe, the mar- gins conspicuously erose. Leaves 24-50 cm long, (2.5-)3.5-9 cm broad, short-petiolate, essentially monomorphic. Petiole 0.5-4 cm long. Lamina subcoriaceous, oblong or elliptic, acuminate to caudate at apex, cuneate at base, glabrous. Veins indistinct or, more commonly, obscure, copiously areolate, with numerous secondary areoles and with free veinlets spreading in various directions. Sori 4-8 mm long, round or elliptic, inframarginal, pa- raphyses abundant, orange to castaneous, filiform. In lowland, often primary forests, climbing on, and usually appressed to, tree trunks, 100-200 m, Amazonas, San Martin, and Loreto. Venezuela; Guyana; Columbia to Peru; Brazil. The great size of the leaves and the scale-cov- ered, broad, flattened stem easily distinguish this from all other species in the genus. Amazonas: Mobil Oil Company trail, 3 km from La Poza, E bank of Santiago River, Berlin 3508 (MO, us). San Martin: Lamas, Santa Rosa de Davidcillo, Knapp & Mallet 7171 (MO). Loreto: Prov. Maynas, Rio Ampi- yacu, vicinity of Puca Urquillo, Plowman et al. 6966 (F, GH, USM). Prov. Maynas, Dist. Nanay, 3 km from Santa Maria de Nanay, Schunke K 2503 (F, GH, us). Prov. Maynas, Saboya, Vasquez & Jaramillo 7418 (MO, uc). Prov. Requena, Pan, Jenaro Herrera, Vasquez & Jar- amillo 1057 (F, USM). 14. Microgramma persicariifolia (Schrader) Presl, Tent, pterid. 214. 1836. Polypodium persicariifolium Schrader, Gott, Gel. Anz. 1824: 867. 1824. TYPE: Brazil, Maximilian von Wied-Neuwied (BR?). Plants epiphytic. Stem 1-2 mm in diameter, completely covered with appressed scales, these 4-5 mm long, lanceolate, attenuate to a filiform tip, peltately attached well above the base, orange or tawny to whitish, with a castaneous point where attached, the margins subentire. Leaves 10-20 cm long, (1.5-)2-3.5 cm broad, subsessile to short- petiolate, essentially monomorphic. Petiole 0.2- 1.5 cm long. Lamina firm-herbaceous to charta- ceous, broadly lanceolate or elliptic, acuminate at apex, attenuate at base, glabrous, but with a few, scattered, filiform scales along the costa abaxially. Veins rather distinct, primary ones much thicker TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 157 and more prominent than the rest, areolate, with many free, included veinlets spreading in various directions. Sori oblong to linear, approximately midway between and parallel to primary veins, medial. In or at edges of forests, climbing on shrubs or small trees, 100-350 m, Loreto, Huanuco, and Madre de Dios. Costa Rica; Panama; Trinidad; Columbia to the Guianas, south to Paraguay and Brazil. Loreto: Mishuyacu, near Iquitos, Klug 1594 (F, GH, us). Prov. Maynas, Iquitos, Sagdstegui & Aldave 5824 (F, GH, HUT). Rio Mazan, Quebrada Salinas, J. Schunke 359 (F, GH, uc, us, USM). Huanuco: Prov. Pachitea, Dist. Honoria, Bosque nacional de Iparia, Schunke V. 1973 (F, GH, us). Madre de Dios: Parque Nacional del Manu, Cocha Cashu Station, Rio Manu, Foster et al. 6947 (F), M. Foster P84- 13 (uc). Prov. Tambopata, Albergue Cuz- co Amazonico, Leon 863 (USM). VII. Carnpyloneurum Contributed by Blanca Leon Carnpyloneurum Presl, Tent, pterid. 189. 1836. LECTOTYPE: Carnpyloneurum repens (Au- blet) Presl (chosen by John Sm., Hist. fil. 95. 1875). Figure 11. Polypodium subg. Cyrtophlebium R. Br., in Benn., PI. Jav. Rar. 1:4.1838. TYPE: Cyrtophlebium repens (Aublet) John Sm. Cyrtophlebium (R. Br.) John Sm., J. Bot. (Hooker) 4: 58. 1841. Plants epiphytic, hemiepiphytic or terrestrial. Stem short or long-creeping, sometimes branched, green, green-stramineous, brown-black, some- times pruinose, indument of clathrate or nonclath- rate scales, these brown, ferrugineous, dark-ochra- ceous, or rarely whitish. Leaves simple, monomorphic, ca. 20-100(-200) cm long, glabrate or rarely pubescent; phyllopodia present; petiole usually present and articulate. Costa usually prom- inent on both sides of the lamina, venation are- olate, primary veins usually parallel, 45-75° di- vergent from the costa, secondary veins transverse to primary veins, anastomosing to form 2 or more primary areoles, these with 1-6 included veinlets, veinlets simple or furcate, excurrent, rarely recur- rent, free or anastomosed sometimes to secondary veins to form secondary areoles, apex of free vein- lets with hydathodes. Sori without indusia, usually at the medial to apical zone of the free veinlet, rarely at the junction of 2 veinlets, in (l-)2— 4(-6) series between primary veins. Carnpyloneurum is a Neotropical genus con- taining 47 species, a few of which extend into sub- tropical regions of America, a majority of these occurring in forests. The generic limits are estab- lished based on the pattern of venation, which is cyrtophlebioid, rather than goniophlebioid; i.e., secondary veins that form the areoles are curved rather than straight. Polypodium magnificum is sometimes placed in Carnpyloneurum; however, for the purposes of this flora, the 1 -pinnate species are treated under Polypodium. The characters in the key to species apply to mature leaves. The prominence of veins should be examined at the middle of the leaf, and this can often be determined with the naked eye. However, examination of trichomes and stem scales requires the use of a stereomicroscope. In the past, leaf characters such as lamina width and shape were extensively used in distinguishing species, but these are only partially reliable for this purpose. Vari- ation within several species (Carnpyloneurum am- phostenon, C. angustifolium, C. chlorolepis, C. coarctatum) shows that leaf characters cannot be used alone. For purpose of identification, a com- plete specimen, including stem scales, should be available, since these are the most important char- acters for recognizing species, as has been shown by Lellinger (1988). However, there are still some problems within some species (C. amphostenon, C. angustifolium, and C. densifolium), such as the presence of different types of scales within indi- viduals. Also, the presence of many individuals with intermediate characteristics suggests that more studies are needed to evaluate the variation of stem scale characteristics within and among pop- ulations. In certain species there are differences at the stem scale margins in cell shape, size, and dis- position. These differences are expressed in this treatment as scales with margins differentiated FIG. 1 1 . Carnpyloneurum fasciale: a, habit. Carnpyloneurum phyllitidis: b, portion of fertile lamina. Carnpylo- neurum. angustipaleatum: c, stem scale. Carnpyloneurum ophiocaulon: d, stem scale. Carnpyloneurum amphostenon: e, stem scale (Adapted in part from Stolze, Ferns and Fern Allies of Guatemala, 1981.) 158 FIELDIANA: BOTANY TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 159 (such as in fig. 1 Id) or not differentiated (figs. 1 Ic, Reference 1 le). Such differences have been referred to by some other authors as "scales with margins non- LELLINGER, D. B. 1988. Some new species of conform or conform." Campyloneurum and a provisional key to the genus. Amer. Fern J., 78: 14-34. Key to Species of Campyloneurum a. Primary veins not well defined, obscure, or prominulous (rarely prominent) in different degree on either side of the lamina; primary areoles divided and stem less than 4 mm wide b b. Stem scales whitish in mass, not clathrate 13. C. chlorolepis b. Stem scales light or dark brown, not whitish in mass c c. Lamina always puberulent, with minute, inconspicuous, furcate trichomes on both sides of the lamina 1. C. aphanophlebium c. Lamina glabrous adaxially, or with indument of simple trichomes only abaxially d d. Leaves usually more than 5 mm apart e e. Stem scales not clathrate f f. Stem scales light brown, not shining, the cell lumina usually not occluded; stem scale bases peltate 1 2. C. vulpinum f. Stem scales dark brown, usually shining, the cell lumina mostly occluded; stem scale bases short-biauriculate 1 1 . C. asplundii e. Stem scales clathrate or slightly clathrate g g. Stem scales usually 2-3 times as long as broad; scale bases usually more than 2.5 mm wide, with margins at the central and basal portion slightly differentiated 10. C. densifolium g. Stem scales usually 3-5 times as long as broad; scale bases usually 1.5-2(-2.5) mm wide, margins sometimes differentiated at the basal portion ... 9. C. amphostenon d. Leaves usually less than 5 mm apart h h. Stem scales 6-10 times longer than broad, linear from a broad base, less than 1 mm wide 16. C. angustipaleatum h. Stem scales 5 times as long as broad or less, linear-lanceolate, ovate to triangular-lan- ceolate, most stem scale bases usually 1 mm or more wide i i. Stem scales not clathrate, dark brown, often shining, the cell lumina mostly occluded 1 1 . C. asplundii i. Stem scales clathrate or slightly clathrate, the cell lumina translucent or slightly yellow J j. Stem scales gray-brown in mass, the cell lumina slightly iridescent; stem not pru- inose 14. C. aglaolepis j. Stem scales brown in mass, the cell lumina not iridescent; stem usually pruinose k k. Stem scale bases less than 1.5 mm wide; stem scales narrowly triangular-lan- ceolate 15. C. angustifolium k. Stem scale bases more than 1.5 mm wide; stem scales broadly triangular-lan- ceolate to ovate-lanceolate 1 1. Stem scales strongly clathrate, brown in mass, margins not differentiated; most scales 3-5 times as long as broad 9. C. amphostenon 1. Stem scales slightly clathrate, light brown in mass, margins slightly differ- entiated; most of scales 2-3 times longer than broad ... 1 0. C. densifolium a. Primary veins well defined, prominent or prominulous to the same degree on both sides of the lamina; primary areoles divided and stem more than 4 mm wide or primary areoles not divided and stem less than 4 mm wide m m. Most primary areoles divided (transverse veinlets connected by a 1 or more veinlets), often with more than 2 excurrent veinlets in each primary areole; stem usually more than 5 mm wide . . . n n. Stem scales linear, black or dark brown 20. C. nitidissimum 160 FIELDIANA: BOTANY n. Stem scales triangular-lanceolate or ovate-lanceolate, brown o o. Secondary areoles essentially isodiametric; sori usually in 2 rows of son between primary veins; stem scales triangular-lanceolate at the growing tip 17. C. phyllitidis o. Secondary areoles irregularly shaped; sori usually in more than 2 rows between primary veins; stem scales ovate-lanceolate p p. Secondary veins obscure 18. C. brevifolium p. Secondary veins prominulous 19. C. pascoense m. Most primary areoles not divided (i.e., transverse veinlets not connected by 1 or more excurrent veinlets), usually with 2 excurrent veinlets in each; stem usually less than 4 mm wide (except C. abruptum) q q. Petiole usually more than '/3 the length of the lamina r r. Lamina subcoriaceous; stem scales usually persistent 6. C. inflatum r. Lamina chartaceous or herbaceous; stem scales usually caducous s s. Stem scales linear-lanceolate, brown or light brown, not strongly clathrate 7. C. sphenodes s. Stem scales linear to subulate, dark brown, strongly clathrate 8. C. coarctatum q. Petiole less than '/3 the length of the lamina t t. Stem scales usually more than 5 mm long; stem diameter (4-)5-10 mm; leaves usually less than 5 mm apart 21. C. abruptum t. Stem scales usually less than 5 mm long; stem diameter 2-4 mm; leaves usually more than 5 mm apart u u. Cell lumina of the stem scales always narrow, dark yellow or almost occluded; stem scales dark brown, linear-lanceolate 5. C. fuscosquamatum u. Cell lumina of the stem scales transparent or yellowish, not occluded; stem scales brown, linear-lanceolate or ovate-lanceolate v v. Apex of stem scales obtuse, stem diameter usually more than 3 mm 3. C. ophiocaulon v. Apex of stem scales always acuminate, stem diameter usually less than 3 mm . . . w w. Stem scales with differentiated margins 2. C. repens w. Stem scales without differentiated margins x x. Lamina narrowly lanceolate, usually less than 5 cm wide, the base long-de- current; stem scales narrowly lanceolate 4. C. fasciale x. Lamina broadly lanceolate, usually more than 5 cm wide, the base broad- cuneate or short-decurrent; stem scales linear 8. C. coarctatum 1. Campyloneurum aphanophlebium (Kunze) Moore, Index fil. 223. 1861. Polypodium aphanophlebium Kunze, Bot. Zeit. (Ber- lin) 288. 1845. TYPE: Venezuela, Caracas, Mo- ritz 17 (holotype. B!). Polypodium occultum Christ, Bull. Herb. Boissier II, 5: 7. 1905. TYPE: Costa Rica, Prov. Cartage, Rio de las Vueltas, Tonduz 12752 (holotype, P!; iso- types B!, BM!). Polypodium trichiatum Rosenst., Repert. Spec. Nov. Regni Veg. 7: 148. 1909. TYPE: Ecuador, Rio- bamba, Cordillera Occidental, Rimbach 87 (ho- lotype, s!; isotypes, us!; photo, BM of s). Campyloneurum trichiatum (Rosenst.) Ching, Suny- atsenia 5: 263. 1940. Campyloneurum occultum (Christ) Gomez, Brenesia 8:46. 1976. Epiphytic, rarely terrestrial. Stem creeping, 1- 3(-4) mm wide, not pruinose, with scales brown, linear-lanceolate. Leaves (10-)20-40(-50) cm long, 2-4(-6) mm apart, petiole (0.3-)0.7-3(-6) cm long, stramineous or brown-stramineous, indument of minute furcate trichomes. Lamina oblanceolate, rarely narrowly lanceolate, chartaceous, (1.2-)2- 4(-4.7) cm wide, with attenuate base, with plane or slightly revolute margins, with acuminate or subcaudate apex, puberulent, indument of minute and very unequally furcate trichomes on both sides of the lamina, relatively dense abaxially. Costa prominent, plane or slightly canaliculate adaxially, convex or anuglate abaxially; primary veins slight- ly prominulous or obscure, more or less concol- orous with the adjacent tissue, slightly flexuous, (3-)4-7(-9) mm apart, diverging 50-60(-65)° from the costa, areoles between costa and margin 3-6, with 2(-3) excurrent veinlets in each areole. Sori medial or subterminal on the excurrent veinlets, paraphyses dendritic. TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 161 On tree trunks, sometimes in rock crevices, in shady places of lowland forests, 100-1000 m, Amazonas, San Martin, Loreto, Huanuco, Pasco, and Junin. Belize; Nicaragua to Venezuela, south to Bolivia and Brazil. Dry specimens of Campyloneurum aphano- phlebium sometimes are dull green or brownish on both sides of the lamina. Because of the ve- nation pattern, it is here allied with C. repens and related species. Amazonas: Prov. Bagua, ca. 1 km NE of Quebrada Chinganza, 10 km NE of Mayo, Knapp & Alcorn 7733 (NY). Prov. Bagua, Montenegro-Chiriaco, Sagdstegui 5924 (HUT). San Martin: Prov. Mariscal Caceres, Campanilla, Cajon Pericote, J. Schunke V. 4288 (uc, USM). Rio Ma- ranon, Spruce 3912 (K). Loreto: Prov. Alto Amazonas, Balsapuerto, lower Rio Huallaga, Killip & Smith 28642 (NY, us). Prov. Loreto, San Jose de Parinari, Vdsquez et al. 2274 (NY). Huanuco: Sinchono, Fundo Chela, Aguilar 946 (USM). Prov. Pachitea, Puerto Inca, Agua Dulce, al este de Puerto Inca, J. Schunke V. 2818 (GH, us). Tingo Maria, Tryon & Tryon 5335 (GH, us, USM). Cachicoto, Woytkowski 7868 (uc). Pasco (as Junin): Prov. Oxapam- pa, Puerto Bermudez, Killip & Smith 26647 (us). Junin: Prov. Chanchamayo, La Merced, Killip & Smith 23781 (GH, NY, us). Rio Paucartambo, near Perene bridge, Killip & Smith 25268 (s). 2. Campyloneurum repens (Aublet) Presl, Tent, pterid. 190. 1836. Polypodium repens Aublet, Hist. pi. Guiane 2: 962. 1775. TYPE: Plumier, Taite foug. Amer., t. 134, 1705. Polypodium lapathifolium Poiret, in Lam., Encycl. 5: 514. 1804. TYPE: America meridional, Herb. Jussieu 1071-E (holotype, P; photos, BM, s). Polypodium caespitosum Link, Hort. berol. 2: 9 1 . 1 833. TYPE: Cultivated, ex Hort. Loddiges (holotype, B!). Campyloneurum caespitosum (Link) Link, Fil. spec. 125. 1841. Cyrtophlebium repens (Aublet) John Sm., J. Bot. (Hooker) 4: 58. 1841. Campyloneurum lapathifolium (Poiret) Ching, Suny- atsenia 5: 263. 1940. Epiphytic or hemiepiphytic. Stem long-creep- ing, l-3(-4) mm wide, not pruinose, stem scales brown or light brown, lanceolate, clathrate, with center and margins differentiated in cell wall color and cell structure. Leaves 20-60 cm long, (5-) 10- 15 mm apart, petiole 0.5-7(-13) cm long, stra- mineous. Lamina oblong-lanceolate or obovate- lanceolate, sometimes narrowly lanceolate, her- baceous-chartaceous, (2.5-)3-8 cm wide, with at- tenuate, sometimes narrowly cuneate, base, with slight sinuate, cartilaginous, plane or very slightly revolute margins, with acuminate or caudate apex, indument scarce, of simple and sometimes furcate trichomes, scarce abaxially. Costa prominent, slightly sulcate adaxially; primary veins promi- nent or prominulous on both sides of the lamina, slightly flexuous or straight, often lighter in color than the adjacent tissue, (3-)5-7 mm apart, di- verging (65-)70-75° from the costa, areoles be- tween costa and margin (4-)6-12, with 2(-4) free excurrent veinlets in each areole. Sori medial or subterminal on the excurrent veinlets, paraphyses not seen. Usually in shady and humid places of lowland and montane forests, 100-2000 m, Amazonas, San Martin, Loreto, Huanuco, Pasco, Ayacucho, Cuz- co, and Madre de Dios. Mexico; Central America; Lesser Antilles; Guianas; Venezuela; Columbia to Bolivia and cen- tral Brazil. Amazonas: Prov. Bagua, Cordillera Colan, SE of La Peca, Barbour 4188 (F). San Martin: Prov. Mariscal Ca- ceres, Tocache Nuevo, J. Schunke V. 6952 (F, MO). Par- que Nacional Rio Abiseo, hill pass Las Palmas, Young 4324 (USM). Loreto: Prov. Maynas, Gamitanacocha, Rio Mazan, /. Schunke 205 (F, s, uc, us). Huanuco: Prov. Leoncio Prado, Dist. Hermilio Valdizan, from Puma- huasi to La Cumbre, Plowman & Schunke 7414 (F). Pas- co: Prov. Oxapampa, Gran Pajonal, vicinity of Chequi- tavo, D. Smith 5265 (uc). Ayachuco: Prov. La Mar, E massif of Cordillera Central, between Tambo San Mi- guel, Ayna and Hacienda Luisiana, Dudley 1 1908 (GH). Cuzco: Prov. Paucartambo, Hacienda Villa Carmen, Vargas 14697 (GH). Madre de Dios: Prov. Tambopata, Tambopata Nature Reserve, SSW of Puerto Maldonado, Barbour 4853 (F, MO). 3. Campyloneurum ophiocaulon (Klotzsch) Fee (Mem. foug. 5) Gen. fil. 258. 1852. Figure lid. Polypodium ophiocaulon Klotzsch, Linnaea 20: 401. 1847. TYPE: Peru, Junin, Dombey 41 (holotype, B!; photo, BM). Epiphytic. Stem long-creeping, brown or dark stramineous, (2-)2.5-4 mm wide, scales brown, clathrate, ovate-lanceolate, apex obtuse, margins differentiated. Leaves 30-50 cm long, 1-2 cm apart, petiole dark stramineous, 2.5-4.5 cm long. Lam- ina oblanceolate or broadly lanceolate, charta- ceous, 3.5-7 cm wide, with attenuate base, with sinuate, cartilaginous margins, with acuminate, apex, indument of sparse, simple trichomes, spreading abaxially. Costa prominent; primary veins prominulous, lighter in color than the ad- 162 FIELDIANA: BOTANY jacent tissue, 6-7 mm apart, diverging 6 5-70(-80)° from the costa, with 8-1 1 areoles between costa and margin, with 2(-3) free excurrent veinlets in each areole. Sori medial on the excurrent veinlets, paraphyses inconspicuous or lacking. In shady places, rarely in open light gaps, from lowland forests to montane forests, (1500-)2000- 2500 m, Cajamarca, Amazonas, San Martin, Uca- yali, Huanuco, Pasco, Junin, and Cuzco. Columbia to Bolivia. Cajamarca: Prov. Cutervo, San Andres de Cutervo, arriba de Saucedal, pasando por Chorro Blanco, Diaz & Osores 2963 (MO). Amazonas: Prov. Bagua, E of La Peca, Barbour 2812 (AAU, F, uc). San Martin: Prov. Rioja, Pedro Ruiz-Moyobamba road, km 390 Venceremos, D. Smith 4498 (F, NY, uc). Ucayali: La Divisoria, ca. 25 km NE of Tingo Maria, Moran & Fernandez 3693 (MO, uc, USM). Huanuco: Prov. Huanuco, Pampayacu, Ka- nehira 1 79(GH, us). Prov. Leoncio Prado, road between Tingo Maria and Pucallpa, Sullivan & Young 1 147 (F). Pasco: Oxapampa, Ulcumanu SW of Oxapampa, road to Maria Teresa and Llaupi, Foster et al. 7682 (USM). Prov. Oxapampa, rodal del Sr. Miiller, cerca a Oxapam- pa, Leon 495 (F, USM). Junin: Villa Amoretti, Kunkel 524 (GH). In the area of Pichita Caluga, Walden 26 (BM). Cuzco: Prov. La Convencion, entre Quillabamba y Abra de Malaga, Ellenberg 4739 (LPB). Prov. Paucartambo, km 1 36 carretera Acjanaco-Pilcopata, abajo de Buenos Aires, Ledn 2183 (F, USM). Prov. Urubamba, Machu Picchu Sanctuary, Puncuyo, Peyton & Tilney-Peyton 1365 (GH). Prov. La Convencion, Amaibamba, Vargas 9804 (uc). 4. Campyloneurum fasciale (Willd.) Presl, Tent, pterid. 190. 1836. Figure lla. Polypodium fasciale Willd., Sp. pi. 5. 156. 1810. TYPE: Venezuela, Caripe, Humboldt 426 (holotype, B!, Herb. Willd. 19632; photo, BM, GH). Polypodium serpentinum Christ, Bull. Herb. Boissier II, 6: 51. 1906. TYPE: Costa Rica, Navarro, Werckle (holotype, P!; photo, BM). Campyloneurum serpentinum (Christ) Ching, Suny- atsenia 5: 263. 1940. Epiphytic. Stem long-creeping, 1-3 mm wide, not pruinose, dark green or green, the scales brown or dark brown, narrowly lanceolate. Leaves erect or subpendent, 20-40(-50) cm long, 5-10 mm apart, petiole 1-2 cm long, stramineous or brown- ish. Lamina lanceolate or narrowly lanceolate, rarely linear-lanceolate, herbaceous-chartaceous, (0.8-)2.5-5 cm wide, with attenuate, rarely nar- rowly cuneate base, with slightly revolute or plane, slightly sinuate, cartilaginous margins, with acu- minate apex, indument of simple trichomes, these scarce abaxially. Costa prominent, with caducous scales; primary veins prominulous, 3.5-5 mm apart, diverging 60-70(-75)° from the costa, with (4-)5-8 areoles between costa and margin. Sori medial or subterminal, paraphyses inconspicuous or lacking. In mountain forests, 300-2400 m, Amazonas, San Martin, Loreto, Pasco, Junin, and Madre de Dios. Southern Mexico; Central America; Venezuela and French Guiana; Columbia to Bolivia. Campyloneurum fasciale can be distinguished from the related species C. fuscosquamatum and C. repens by the stem scale characters, as noted in the key. Campyloneurum fasciale differs from C. coarctatum on leaf shape and stem scale features; in the former, the lamina base is attenuate, rarely narrowly cuneate, with slightly rigid, adpressed, stem scales, while in the latter the lamina base in usually acuminate or narrowly cuneate with flex- uous stem scales. Amazonas: Prov. Bongara, SW of Pomacocha, Wur- dack 844 (F, NY, uc, us, USM). Sipabamba, Shillac, Young & Eisenberg 352a (MO, NY, uc). Prov. Bagua, 1 2 km E of La Peca, Barbour 2487 (MO, uc), Barbour 2495 (F, uc), Gentry et al. 22930 (F, MO, uc). San Martin: Cam- pana, Tarapoto, Spruce 4647 (BM). Loreto: Santa Rosa, lower Rio Huallaga. below Yurimaguas, Killip & Smith 28927 (us). Pasco: Prov. Oxapampa, Paujil, near Puerto Bermudez, Leon 287 (GH, USM). Junin: Pichis trail, Ya- pas, Killip & Smith 25606 (F, us). Madre de Dios: Prov. Manu, Manu National Park, Cocha Cashu, Nunez 6070 (NY). 5. Campyloneurum fuscosquamatum Loll.. Amer. Fern J. 78: 21. 1988. TYPE: Peru, Huanuco, Tingo Maria, Stork & Morton 9452 (holotype, us; isotypes, F!, GH!, uc!; photo, USM of us). Epiphytic. Stem long-creeping, 2-3 mm wide, green or green-stramineous to black, not pruinose, with scales dark brown, linear-lanceolate, the cell lumina occluded. Leaves erect or partially pen- dent, 20-40 cm long, 4-8 mm apart, petiole 0.5- 5 cm long, stramineous, sometimes darker abax- ially. Lamina narrowly oblanceolate or oblong, herbaceous-chartaceous, 2-5(-8) cm wide, with at- tenuate or narrowly cuneate base, with slightly rev- olute or plane, cartilaginous margins, with acute or acuminate apex, indument of scarce, simple, inconspicuous trichomes. Costa prominent, some- times with scales similar to those of the stem; pri- mary veins prominulous, slightly flexuous, di- verging (60-)70-75(-80)° from the costa, with 6- 10 areoles between costa and margin, with 2(-3) TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 163 excurrent veinlets in each noncostal areole. Sori medial or subterminal on the free excurrent vein- let, paraphyses inconspicuous or lacking. In humid and shady places, humid lowland for- ests or premontane forests, 100-1 250 m, San Mar- tin and Loreto, south to Ayacucho and Madre de Dios. Columbia to Bolivia. This species is closely related to C. fasciale and C. repens. San Martin: Prov. Mariscal Caceres, Campanilla, sud- oeste del Caserio de Sion, J. Schunke V. 3556 (F, USM). Prov. San Martin, 4 mi E of Tarapoto, Woytkowski 35235 (MO, s, uc). Loreto: Prov. Maynas, Quebrada Tamshi- yacu, E of Tamshiyacu, Gentry et al. 25837 (F, MO). Huanuco: Tingo Maria, Ferreyra 10276 (GH, USM). Prov. Pachitae, Codo de Pozuzo, Foster 9399 (F). Prov. Huan- uco, Dist. Churubamba, Hacienda Mercedes, Balsa-pla- ya, Mexia 8176 (BM, F, GH, MO, s, uc, USM). Pasco: Prov. Oxapampa, Pozuzo, Macbride 4585 (F, us). Junin: Chan- chamayo valley, Soukup 1103 (F). Ayacucho: Rio Apu- rimac, near Kimpitiriki, Killip & Smith 22861 (F, us). Prov. La Mar, 2 km NW of Santa Rosa, Wasshausen & Encarnacion 621 (MO, USM). Cuzco: Prov. Convencion, NE from Hacienda Luisiana, Dudley 11501 (GH, us). Madre de Dios: Prov. Manu, Parque Nacional Manu, Cocha Cashu Biological Station, M. Foster P-84- 17 (MO). Prov. Tambopata, alrededores de "Cuzco Amazonico," Leon 836 (USM). 6. Campyloneurum inflatum Lell., Amer. Fern. J. 78: 22. 1988. TYPE: Columbia, Cauca, W slope of Cerro Munchique, Perez Arbelaez & Cuatrecasas 6244 (holotype, us!; isotypes, COL, F!; photo, USM of us). Epiphytic. Stem long creeping, 2 mm wide, not pruinose, green, stramineous or black, the scales bright brown, lanceolate. Leaves 30-55 cm long, 3-4 cm apart, petiole 9-21 cm long, stramineous to dark stramineous. Lamina elliptic, cartilagi- nous, subcoriaceous, 6-10.5 cm wide, with acute base, with strongly cartilaginous, slightly revolute margins, with caudate apex, indument lacking. Costa prominent, slightly sulcate, indument of ca- ducous scales; primary veins slightly prominulous adaxially, prominent abaxially, slightly flexuo- sous, lighter in color than the adjacent tissue, 7- 8 mm apart, diverging 65-70° from the costa, with 6-7 areoles between costa and margin, with 2(-4) free excurrent veinlets per areole, obsolete, some- times secondary areoles close to the margin. Sori subterminal on the excurrent veinlets, paraphyses not seen. In Peru only known from Amazonas, 2200-2400 m. Columbia; Peru. Amazonas: Prov. Bongara, WSW of Pomacocha, Wur- dack 868 (us). 7. Campyloneurum sphenodes (Klotzsch) Fee, (Mem. foug. 5) Gen. fil. 258. 1852. Polypodlum sphenodes Klotzsch, Linnaea 20: 242. 1 847. TYPE: Venezuela, Merida, Moritz 304 (ho- lotype, B!; isotypes, BM!, K!). Epiphytic, hemiepiphytic, rarely terrestrial. Stem long-creeping, ( l-)2-3(-4) mm wide, not pruinose, with scales brownish, light brown, lanceolate or narrowly lanceolate, adpressed, slightly clathrate. Leaves erect, 25-50 cm long, 0.7-2.5 cm apart; petiole 5-15 cm long, stramineous or dark-stra- mineous. Lamina narrowly elliptic or lanceolate, chartaceous, (2-)4-6 cm wide, with narrowly cu- neate, short attenuate base, with slightly undulate, cartilaginous margins, with acuminate apex, in- dument inconspicuous or lacking. Costa promi- nent, primary veins priminulous on both sides of the lamina, straight, 5-7 mm apart, (60-)65-70° divergent from the costa, with 6-12 areoles be- tween costa and margin, with 2(-3) free excurrent veinlets in each areole. Sori medial or subterminal on the excurrent veinlets, paraphyses inconspic- uous or lacking. Uncommon in Peru, on tree trunks, in montane or lowland forests, 200-2500 m, Piura, Cajamar- ca, Amazonas, and Junin. Costa Rica; Panama; Venezuela and Columbia, south to Bolivia. Piura: Prov. Huancabamba, Canchaque, cerca a Cho- rro Blanco, Ramirez & Lamas (F, USM). Cajamarca: Prov. Santa Cruz, Dist. Catache, ca. 5 km above Monte Seco, Dillon et al. 4901 (F). Amazonas: Prov. Bagua, Cordillera Colan, SE of La Peca, Barbour 3893 (MO). Junin: Prov. Chanchamayo, E of Quimiri bridge, Killip & Smith 23896 (F, GH, us). 8. Campyloneurum coarctatum (Kunze) Fee, (Mem, foug. 5) Gen. fil. 258. 1852. Polypodium coarctatum Kunze, Linnaea 9: 39. 1834. TYPE: Peru, Huanuco, Cucheros, Poeppigin July 1829 (holotype, LZ, destroyed; isotypes, P!, w!; photo, BM of w). Epiphytic, rarely terrestrial. Stem long-creeping, 164 FIELDIANA: BOTANY (1.5-)2.5-3.5 mm wide, green, turning black with age, rarely stramineous, the scales narrowly lan- ceolate to linear-lanceolate. Leaves erect, 45-85 cm long, 5-9 mm apart, petiole 10-16 cm long, stramineous to dark stramineous. Lamina usually broadly elliptic to ovate-elliptic, rarely narrow- elliptic, chartaceous, (6-)8-13 cm wide, with nar- rowly cuneate or acuminate, rarely short-attenuate base, with slightly sinuate, cartilaginous, margins, with attenuate apex, indument of inconspicuous, simple trichomes, spreading abaxially. Costa prominent; primary veins priminulous on both sides of the lamina, diverging 60-70° from the costa, with 10-19 undivided areoles between costa and margin, with usually 2 free veinlets in each areole. Sori subterminal or medial, on the excur- rent veinlets, paraphyses inconspicuous or lacking. Usually in humid and shady places of humid forests, 1 00-1 000(-2000) m, Cajamarca, San Mar- tin, Loreto, Ucayali, Huanuco, Junin, and Aya- cucho. Costa Rica; Panama; French Guiana; Colum- bia; Bolivia; central Brazil. Campyloneurum coarctatum can be recognized by having nearly linear, strongly clathrate stem scales. It is closely related to C. sphenodes. Cajamarca: Prov. Santa Cruz, ENE Monteseco, San- tisteban & Guevara 31 (F, HUT). San Martin: Prov. San Martin, Tarapoto, Spruce 4646 (BM, NY, w). Loreto: Prov. Loreto, Rio Maranon, arriba del caserio Suramuro, Diaz & Ruiz 875 (MO). Santa Rosa, lower Rio Huallaga, below Yurimaguas, Killip & Smith 28854 (F, NY), 27665 (F, NY). Ucayali: Prov. Coronel Portillo, Bosque von Hum- boldt, Young & Salazar 1015 (F, MO). Huanuco: Huan- uco, gorge of Rio Chinchao, Tryon & Tryon 5302 (BM, F, GH, NY, uc, us). Junin: Tarma, Agua Dulce, Woyl- kowski 37025 (MO, uc). Ayacucho: Prov. La Mar, Aina (Ayna), between Huanta and Rio Apurimac, Killip & Smith 22720 (us). 9. Campyloneurum amphostenon (KJotzsch) Fee (Mem. foug. 5) Gen. fil. 258. 1852. Terrestrial or epiphytic. Stem long-creeping, (2-) 3-5 mm wide, usually pruinose, the scales brown, lanceolate. Leaves erect, 1 5-60(-70) cm long, (2-)- 5- 1 0 mm apart, petiole 4-30 cm long, stramineous or brown-stramineous. Lamina linear-lanceolate or lanceolate, chartaceous or subcoriaceous, (0.7-) 2-5 cm wide, with attenuate base, with slightly revolute or plane, cartilaginous margins, with acu- minate apex, indument of simple trichomes, scarce abaxially. Costa prominent, indument of caducous scales; primary veins obscure or prominulous adaxially, slightly prominulous abaxially, often concolorous with the adjacent tissue, straight or slightly flexuosous, 5-7 mm apart, diverging 45- 60° from the costa, with 2-4(-5) areoles between costa and margin, with 1-2 free excurrent veinlets in each noncostal areole. Sori medial or subter- minal on the excurrent veinlets, paraphyses in- conspicuous or lacking. I recognize Campyloneurum amphostenon as a complex of different populations, especially in the Andean region, probably reflecting incomplete speciation. Leaf and stem scale characteristics are extremely variable, as reflected in the description, sometimes at the individual level. Therefore, more studies, especially using cytogenetic techniques, are needed to evaluate this complex. Based on the currently available information, I recognize two varieties in this species. Key to Varieties a. Most of stem scales with elongate cells parallel to the main axis 9a. var. amphostenon a. Most of stem scales with cells irregularly ordered 9b. var. irregulare Campyloneurum amphostenon is closely related to C. asplundii and C. densifolium. Besides the characters provided in the key, C. amphostenon can be distinguished by the stem scales having an acuminate apex. Campyloneurum densifolium has the apex acute or broad and short-acuminate. Campyloneurum amphostenon differs from C. as- plundii by having clathrate stem scales, with bases usually more than 1.5 mm wide and cell lumina translucent or yellowish, while the latter has non- clathrate stem scales, with bases usually less than 1.5 mm wide and cell lumina brownish. However there are some intermediate specimens such as Nunez 8422 (F, NY), Vargas 1 1864 (GH), and Young & Leon 4519 (USM), with scales strongly clathrate but shaped like those of C. asplundii. TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 165 9a. Campyloneurum amphostenon var. atnphos- tenon. Figure lie. Polypodium amphostenon Klotzsch, Linnaea 20: 399. 1847. TYPE: Venezuela, Merida, Moritz 120b (holotype, B!; isotypes, BM!, K). Campyloneurum angustifolium var. amphostenon (Klotzsch) Farw., Amer. Midi. Naturalist 1 2: 296. 1931. This variety is recognized by the character pro- vided in the key. Leaves, sometimes in rather dense tufts, on tree trunks in remnant forested areas, or in shady rock crevices, in montane areas, (1500-)2000-4000 m, Piura, Cajamarca, Amazonas, La Libertad, San Martin, Ancash, Huanuco, Lima, Pasco, Junin, Huancavelica, Ayacucho, Cuzco, and Puno. Mexico; Central America; West Indies; Vene- zuela; Columbia to Bolivia. Piura: Prov. Huancabamba, Cuello del Indio, Lopez et al. 8894 (GH, MO, uc). Cajamarca: Prov. Santa Cruz, Dist. Catache, upper Rio Zana valley, ca. 5 km above Monte Seco, Dillon et al. 4896 (F, uc). Prov. Cajamarca, arriba de La Encanada, ruta a Celendin, Sagdstegui et al. 9404 (F, NY). Amazonas: Prov. Chachapoyas, Cano Santa Lucia, E of Chachapoyas, Wurdack 736 (us, USM). La Libertad: Prov. Sanchez Carrion, Huamachuco-Ca- jabamba road, D. Smith & Vdsquez 3378 (F, uc). San Martin: Prov. Mariscal Caceres, Parque Nacional Rio Abiseo, Chochos, Young & Leon 4562 (USM, HUT). An- cash: Prov. Santa, Jalca de Ultu-Cruz (Jimbe), Mostacero et al. 1869) (F, NY). Prov. Huari, Parque Nacional Huas- caran, Quebrada Carhuaz-Cancha, D. Smith et al. 12243 (F). Huanuco: Mito, Macbride & Featherstone 1919 (us), Huacachi, estacion near Muna, Macbride 4 127 (F). Lima: Prov. Huarochiri, Infiernillo, entre Lima y La Oroya, Ferreyra 6236 (BM, USM). Viso, Goodspeed et al. 11547 (GH, uc, us). Pasco: Prov. Pasco, entre Huariaca y Cerro de Pasco, Ferreyra 9502 (GH). Prov. Oxapampa, trail to summit of Cordillera Yanachaga, via Rio San Daniel, D. Smith et al. 7856a (USM). Junin: Prov. Tarma, In- catacuna, entre Tarmatambo y Acolla, Constance & To- var 2348 (uc). Prov. Satipo, Dist. Pampa Hermosa, road Concepcion-Satipo, Sounders 1068 (GH). Huancavelica: Prov. Tayacaja, Huaribamba, 1 km before Huari, Saun- ders 1147 (GH). Chuspi, cerca a Tocas, Tovar 2039 (GH, USM). Ayacucho: Prov. La Mar, E massif of the Cordillera Central, between Tambo San Miguel, Ayna and the Ha- cienda Luisiana, Dudley 12029 (GH). Road from Tambo to Ayna, Plowman & Davis 4676 (GH). Cuzco: Prov. Urubamba, Quebrada above Pojpoj, Davis et al. 1477 (F, NY, USM). Prov. Paucartambo, Parque Nacional Manu, Leon 2290 (F, USM). Prov. Urubamba, Machu Picchu Sanctuary, in Urcoscancha, above the village of Palcay, Peyton & Tilney- Peyton 761 (GH). Puno: Granja Salcedo, near Puno, Soukup 10 (F). 9b. Campyloneurum amphostenon var. irregulare (Lell.) B. Leon, comb. nov. Campyloneurum irregulare Lellinger, Amer. Fern J. 78: 24. 1988. TYPE: Ecuador, Pichincha, Hold- ridge 1580 (holotype, us!). This variety grows between 2700 and 4000 m, from Cajamarca to Puno. It seems to occur com- monly in open, rocky areas. Venezuela to Bolivia. Cajamarca: Prov. Hualgayoc, Cajamarca-Bambamar- ca road, pass above Hualgayoc, D. Smith & R. Vdsquez 3519 (F, MO, uc). Amazonas: Prov. Chachapoyas, upper slopes and summit of Cerro Campanario, Wurdack 1574 (GH, NY, us). La Libertad: Prov. Otuzco, Llaguen, Shilte, Lopez 1560 (GH). San Martin: Prov. Mariscal Caceres Parque Nacional Rio Abiseo, Puerta del Monte, P5 patch of forest, Young 1986 (F, USM). Huanuco: Mito, Bryan 204 (F). Pasco (as Junin): Prov. Pasco, between Cerro de Pasco and La Quinua, Asplund 11869 (us). Junin: Prov. Huancayo, Huancayo, Soukup 3646 (BM, F). Cuzco: Prov. Paucartambo, Tres Cruces, upper edge of Parque Na- cional Manu, Gentry et al. 23450 (MO, us). Prov. Cuzco, near Saxaihuaman, Tryon & Tryon 5345 (BM, F, GH, us). Puno: Baja Isla, Lake Titicaca, Mexia 7787 (BM, F, GB, GH, MO, s, uc). Prov. Huancane, Moho, Shepard 54 (GH). 10. Campyloneurum densifolium (Hieron.) Lell., Amer. Fern J. 78: 19. 1988. Polypodium angustifolium f. densifolium Hieron., Bot. Jahrb. Syst. 34: 532. 1904. LECTOTYPE (des- ignated by Lellinger, Amer. Fern J. 78: 1919. 1988): Ecuador, Azuay, near Las Yerbas Buenas, Lehmann 5723 (us; isolectotypes, B!, F!). Epiphytic or terrestrial. Stem long-creeping, 3- 6 mm wide, usually pruinose, the scales ovate- lanceolate, light brown or yellow-brown. Leaves erect, 30-70 cm long, 3-10 mm apart, petiole 10- 30 cm long, brown-stramineous. Lamina linear- lanceolate or lanceolate, chartaceous or subcor- iaceous, (l-)2-4(-5) cm wide, with attenuate or narrowly cuneate base, with slightly or strongly revolute margins, with acuminate apex, indument inconspicuous or lacking. Costa prominent, in- dument of caducous scales; primary veins obscure or prominulous and more or less lighter in color than the adjacent tissue adaxially, 5-7 mm apart, diverging (4 5-) 5 0-6 5 (-70)° from the costa, with 2-4 areoles between costa and margin. Sori medial or subterminal on the excurrent veinlet, paraph- yses inconspicuous or lacking. In open exposed areas or in open and humid places of montane forests, (2000-)2500-3800 (-^4200) m, Piura, Cajamarca, La Libertad, San Martin, and Ancash. 166 FIELDIANA: BOTANY Southern Mexico; Central America; Greater Antilles; Colombia and Venezuela to Bolivia. I recognize Campyloneurum densifolium as dis- tinct from C. amphostenon based on the differ- ences in dimensions of the stem scales, which in the former are mostly ovate-lanceolate with acute or short-acuminate apex. Also, C. densifolium has usually adpressed, rather persistent stem scales, while the latter has slightly spreading, semipersis- tent scales. I have seen both species growing in the same area and same type of habitat. In rocky places C. densifolium seems to predominate, but it is scarcer in closed mountain forests. Campyloneu- rum lorentzii (Hieron.) Ching, which occurs in Bo- livia and Argentina, differs by having broader stem scales with an obtuse apex. Piura: Prov. Huancabamba, above Canchaque, Hutchison 1646 (uc). Cajamarca: Prov. Contumaza, Pampa de la Sal. Sagdstegui el al. 10747 (F). La Libertad: Prov. Santiago de Chuco, 26 km from Santiago. D. Smith 2329(vsM). San Martin: Prov. Mariscal Caceres, Parque Nacional Rio Abiseo, Chochos valley, forest patch C9 above timberline, Young 2586 (F, USM). Ancash: Prov. Bolognesi, Tinya, valle del Rio Fortaleza, Cerrate 2585 (GH. USM). Prov. Yungay, Parque Nacional Huascaran, Quebrada Llanganuco, Perez 100 (USM). 1 1 . Campyloneurum asplundii (C. Chr.) Ching, Sunyatsenia 5: 263. 1940. Polypodium asplundii C. Chr., Ark. Bot. 20A(7): 24. 1926. TYPE: Bolivia, Prov. Sur Yungas, La Sir- ena, ca. 2300 m. Asplund 282 (holotype, s!: iso- type, BM). Epiphytic or terrestrial. Stem creeping, some- times branched, (2-)3-4(— 4.5) mm wide, some- times pruinose, black, with spreading, usually ca- ducous, persistent only at growing tips, linear, nonclathrate, dark brown, shining scales, with cell lumina occluded, scale base short-biauriculate. Leaves erect, 35-60 cm long, 2-10 mm apart, pet- iole 0.8-3.8(-20) cm long, stramineous or casta- neous. Lamina linear to narrowly lanceolate, char- taceous, (0.6-)1.5-3.5(-5.8) cm wide, with decurrent base, with cartilaginous slightly revolute margins, with acuminate apex, indument abaxially of simple trichomes, these sometimes with dark apical cell. Costa prominent, indument of often persistent scales; primary veins obscure or slightly prominulous to different degrees on either side of the lamina, sometimes to the same degree, slightly flexuous, 4-5 mm apart, diverging 50-60° from the costa, with 2-4 areoles between costa and mar- gin, with 1 (-2) free excurrent veinlet in each areole. Sori subterminal on the excurrent veinlets, pa- raphyses inconspicuous or lacking. Usually epiphytic, or when terrestrial growing in abundant organic matter, in montane forests between (1500-)! 800 and 2700(-3200) m, Ama- zonas, San Martin, Huanuco, Lima, Pasco, Junin, Ayacucho, Cuzco, and Puno. Ecuador; Peru; Bolivia. Amazonas: Prov. Chachapoyas, along Rio Ventilla, 1- 2 km W of Molinopampa, Wurdack 1490 (us, USM). San Martin: Prov. Mariscal Caceres, Parque Nacional Rio Abiseo, trail between La Playa camp and Papayas camp, Young & Leon 4995 (F, USM). Huanuco: Carpish, Cor- onado 62 (GH, uc, us). Yanano, Macbride 3818 (F, GH, us). Muna, Woytkowski 5181 (GH, us). Lima: Prov. Chancay, Huaccana, cerro al este de Supe, Cerrate 1062 (USM). Pasco: Prov. Oxapampa, Canyon of Huancabam- ba, Fundo La Esperanza, Leon 617 (F, GH, USM). Prov. Oxapampa-Pasco, below San Gotardo, Werffel al. 8521 (NY, uc). Junin: Carpapata, above Huacapistana. Killip & Smith 24429 (NY, us). Chanchamayo, C. Schunke(BM, F). Prov. Chanchamayo, Rio Rondayacu, D. Smith 21 17 (USM). Ayacucho: Aina, between Huanta and Rio Apu- rimac, Killip & Smith 22588 (BM). Cuzco: Prov. La Con- vencion, Dist. Vilcabamba, trail from Yupanqui to Apu- rimac, Dudley 1231 (GH). Prov. Urubamba, Machu Picchu, Ferreyra 2707 (BM, USM). Prov. Paucartambo, Parque Nacional Manu, Leon 2130 (F, USM). Puno: Prov. Sandia. Limbani a Chacani, Vargas 14936 (GH). 12. Campyloneurum vulpinum (Lindman) Ching, Sunyatsenia 5: 263. 1940. Polypodium vulpinum Lindman, Ark. Bot. 1: 245. 1903. Nom. nov. for Polypodium laevigatum var. crispatum C. Chr., and with the same type, not Polypodium crispatum L. 1753. Polypodium laevigatum var. crispatum C. Chr., Bot. Tiddskr. 25: 79. 1902. TYPE: Brazil, Minas Ger- ais, Serra de Caldas, Mosen 2220 (holotype, c; isotypes, BM!, s!, us!; photo, BM of s). Epiphytic. Stem long-creeping, 1-2 mm wide, with reddish-brown, linear-lanceolate scales, less than 1 mm wide, with cell walls conspicuously thinner toward the margin. Leaves 1 5-35 cm long, 7-15 mm apart, petiole 2.5-8(-10) cm long, stra- mineous or greenish stramineous. Lamina linear- lanceolate or narrowly lanceolate, herbaceous, slightly chartaceous, 1.5-3 cm wide, with attenu- ate base, with slightly cartilaginous, sinuate mar- gins, with acuminate apex, indument scarce, of simple trichomes. Costa prominent, primary veins obscure or slightly prominulous, more or less con- colorous with the adjacent tissue, 4-8 mm apart, diverging 50-60° from the costa, with areoles be- tween costa and margin 2-4, with 1-3 veinlets in TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 167 each areole. Sori subterminal on the excurrent veinlets, paraphyses dendritic. Growing in shady and humid places on tree trunks or in rock crevices, often in montane for- ests, (600-) 1200-2 100 m, Amazonas, San Martin, Huanuco, Pasco, Junin, Ayacucho. Haiti; Dominican Republic; Ecuador to Bolivia; Brazil. Amazonas: Prov. Bagua, 12 km E of La Peca, Barbour 2585 (F, MO, NY, uc). San Martin: Prov. Rioja, D. Smith 4364 (MO). Huanuco: Mima, Bryan 421 (F, GH), Woyt- kowski 5218 (GH). Rio Huallaga, below Santo Domingo, Macbride 4208 (F). Pasco: Prov. Oxapampa, 5 km SE of Oxapampa, Leon 506 (USM). Junin: Yucapata, Woyt- kowksi 6658 (us). Ayacucho: Prov. Huanta, Ccarrapa, between Huanta and Rio Apurimac, Killip & Smith 22386 (NY), 22401 (NY). 1 3. Campyloneunim chlorolepis Alston, Bull. Jard. Bot. Etat 27: 56. 1 957. TYPE: Colombia, Cal- das, Chinchina (Pereira-Manizales), Koie 5208 (holotype, not found; photo, BM of c). Polypodium angustifolium var. heterolepis Rosenst, Mem. Soc. Neuchatel 5: 54. 1 9 1 2. TYPE: Colom- bia, Antioquia, near Angelopolis, Mayor 140 (ho- lotype, P!; isotypes, s!, uc!, us!). Campyloneurum heterolepis (Rosenst.) Lell., Amer. Fem J. 67: 58. 1977. Epiphytic, or rarely terrestrial. Stem creeping, 3— 4(-5) mm wide, rarely pruinose, the scales whit- ish, linear-lanceolate. Leaves pendulous or rarely erect, 40-85 cm long, 1-2 mm apart, petiole 4— 8(-10) cm long, stramineous or brown. Lamina linear or narrowly lanceolate, herbaceous-charta- ceous, l-3(-5) cm wide, with narrow cuneate or attenuate base, with slightly revolute, cartilaginous margins, with acute or acuminate apex, indument of inconspicuous simple trichomes, scarce abaxi- ally. Costa prominent; primary veins inconspic- uous or when dry the color darker than the leaf tissue, 5-8 mm apart, diverging 50-60° from the costa, with 2-4 areoles between costa and margin, with 2-4 veinlets in each noncostal areole, anas- tomosing veinlets forming secondary areoles. Sori subterminal on the free excurrent veinlet, paraph- yses inconspicuous or lacking. In low and premontane forests, 200-1000(-1 800) m, Amazonas, San Martin, Loreto, Huanuco, Ju- nin, Ayacucho, and Madre de Dios. Venezuela, Colombia to Bolivia and Brazil. Amazonas: Al lado de Huampani, Kayap 1206 (MO). San Martin: Km 21-22 of Tarapoto-Yurimaguas road, Knapp et al. 7883 (MO, NY, USM). Prov. Mariscal Caceres, NW del caserio Nuevo Progreso, /. Schunke V. 3241 (GH, NY, us, USM). Prov. Tocache, Dist. Tocache Nuevo, Quebrada de Almendras, /. Schunke V. 4453 (F, GH, us). Loreto: Rio Corrientes, between Teniente Lopez and Puesto Avanzado, Gentry et al. 19057 (F, MO, USM). Huanuco: Tingo Maria, Solomon 3389 (MO). Junin: Prov. Satipo, Colonia del Perene, Killip & Smith 25089 (F). Chanchamayo valley, C. Schunke 124 (us), Soukup 1102 (F). Ayacucho: Rio Apurimac valley, near Kimpitiriki, Killip & Smith 22890 (NY, us). Madre de Dios: Prov. Manu, Atalaya, vicinity of Hacienda Amazonia, Foster & Wachter 7417 (USM). Atalya, Nunez 6866 (MO). 14. Campyloneurum aglaolepis (Alston) Sota, Opera Lilloana 5: 96. 1960. Polypodium aglaolepis Alston, J. Bot. 77: 346. 1939. TYPE: Argentina, Tucuman, Siambon, Lorentz & Hieronymus 948 (holotype, BM!; isotype, CORD). Epiphytic. Stem short creeping, 2-3(-4) mm wide, green or green stramineous, with narrowly lanceolate or triangular-lanceolate, brown or gray- ish-brown in mass scales. Leaves pendent, 30-50(- 60) cm long, 2-4 mm apart, petiole 2-4(-8) cm long, stramineous. Lamina linear-lanceolate, 0.5- 1.5 cm wide, with attenuate base, with revolute or plane, cartilaginous margins, with acuminate apex. Costa prominent, primary veins obscure on both sides, or prominulous adaxially and more or less obscure abaxially, concolorous with the adjacent tissue, diverging 40-60° from the costa, 3-5 mm apart, with 2(-3) areoles between costa and mar- gin, with 1 veinlet in each areole. Sori medial or subterminal on the excurrent veinlets, paraphyses inconspicuous or lacking. In dense tufts on tree trunks or in shady crevices of rocks, in humid montane forest, 1200-3600 m, Cuzco and Puno. Peru; Bolivia; Brazil; Argentina. Cuzco: Pinasniocj, "Panticalla" (Pantiacolla) pass,Coo/c & Gilbert 1826 (us). Prov. Quispicanchis, Marcapata, Cadena, Vargas 1 1665 (F). Puno: Prov. Carabaya, abajo de Ollachea, Vargas 6885 (uc, us). 15. Campyloneurum angustifolium (Sw.) Fee (Mem. foug. 5), Gen. fil. 257. 1852. Polypodium angustifolium Sw., Prodr. 130. 1788. TYPE: Jamaica, Swart z (holotype, s!; isotypes, BM!, LD!, s!). Polypodium taeniosumWi\ld.,Sp. 5, 155. 1810. TYPE: 168 FIELDIANA: BOTANY Venezuela, Caripe, Humboldt (holotype, B!, Herb. Willd. 19631; photos, BM, USM). Marginaria angustifolia (Sw.) Presl, Tent, pterid. 1 88. 1836. Campyloneurum taeniosum (Willd.) Fee (Mem. foug. 5), Gen. fil. 257. 1852. Polypodium angustifolium var. gramineum Sodiro, Crypt, vase. Quit. 366. 1893. TYPE: Ecuador, Pichincha-Manabi, a lado del Rio Suma [as Zuma], Cdceres (holotype, P; photo, BM). Epiphytic or epipetric. Stem creeping, (1.5-)2- 3 mm wide, pruinose or not, the scales brown, lanceolate or narrowly lanceolate. Leaves partially erect or pendent, 25-60(-70) cm long, 1^4 mm apart, petiole 1-5 cm long, brown-stramineous. Lamina linear or narrowly lanceolate, chartaceous or subcoriaceous, 0.5-1.5(-2.5) cm wide, with at- tenuate base, with revolute or plane, cartilaginous margins, with acuminate apex. Costa prominent, with caducous scales; primary veins obscure or usually of different color than the lamina tissue abaxially, 4-6 mm apart, with 1-2 areoles between costa and margin, with 1 free veinlet in each areole. Sori medial or subterminal on the excurrent vein- lets, paraphyses scarce, simple. Forming clumps on tree trunks or in crevices of rocks, in open or shady places from low to mon- tane forests, 100-2500(-3600) m, Lambayeque, Cajamarca, Amazonas, La Libertad, San Martin, Loreto, Huanuco, Pasco, Junin, Ucayali, Ayacu- cho, and Cuzco. Southern Florida; Central America; Greater An- tilles; Colombia to Bolivia and Brazil. This species is closely related to C. aglaolepis and C. angustipaleatum. Lambayeque: Prov. Ferrenafe, Tute, Llatas Quiroz2514 (F). Cajamarca: Prov. Jaen, alrededor de Chontali, Lla- nos & Chimoy s.n. (USM). Amazonas: Prov. Bagua, ca. 1 km NE of Quebrada Chingunza, Knapp & Alcorn 7732 (F, NY). La Libertad: Prov. Otuzco, Huaranchal, Lopez et al. 2693 (GH). San Martin: Prov. Mariscal Caceres, 6 km NE of Tingo Maria, Tryon & Tryon 5273 (GH, us). Loreto: Prov. Maynas, Lupuna Cocha, Tryon & Tryon 5187 (BM, F, GH, us, USM). Huanuco: Prov. Pachitea, Dist. Honoria, Bosque Nacional Iparia. Isla del Pacanasi, J. Schunke V. 2322 (F, GH). Tingo Maria, at confluence of Huallaga and Monzon rivers, Stork & Morton 9493 (uc, us). Pasco: Prov. Oxapampa, 5 km SE of Oxapampa, D. Smith 3653 (F, MO, NY, USM). Junin: Rio Pinedo, N of La Merced, Killip & Smith 23647 (F, GH, NY, us). Ucayali: Prov. Coronel Portillo, Bosque Nacional Alexander von Humboldt, Gentry et al. 41417 (MO, USM). Ayacucho: Prov. La Mar, Ayna, between Huanta and Rio Apur- imac, Killip & Smith 22588 (F). Cuzco: Prov. Paucar- tambo, entre Mistiana y Keros, valle de Cosnipata, Scol- nik 882 (us). 16. Campyloneurum angustipaleatum (Alston) Lellinger, Amer. Fern J. 74: 56. 1984. Figure lie. Polypodium angustipaleatum Alston, J. Bot. 77: 346. 1939. TYPE: Bolivia, Prov. Cochabamba, vie. Cochabamba, Bang 1288 (holotype, BM!; iso- types, B!, F!, us). Epiphytic. Stem creeping, 1-2 mm wide, dark green not pruinose, the scales dark brown, clath- rate, cell walls of uniform thickness, linear from a round base, less than 1 mm wide. Leaves pen- dent, 40-80 cm long, 2-4 mm apart, petiole 0.5- 4 cm long, stramineous, glabrate. Lamina linear, herbaceous-chartaceous, 0.5-1(-1 .5) cm wide, with attenuate base, with plane or slightly revolute, slightly cartilaginous margins, with acuminate apex, indument inconspicuous or lacking. Costa prominent; primary veins obscure, diverging 60- 65° from the costa, with l-2(-3) areoles between costa and margin, with 1 veinlet in each areole. Sori medial or submedial, paraphyses inconspic- uous or lacking. Usually in shady places in forests, rarely in areas of remnant forests, 650-2900 m, Cajamarca, Amazonas, San Martin, Huanuco, Pasco, Junin, Cuzco, and Puno. Colombia to Bolivia. Cajamarca: Prov. Jaen, Colasay, Woytkowski 6941 (GH, MO). Amazonas: Prov. Chachapoyas, along Rio Ventilla, 1-2 km W of Molinopampa, Wurdack 1503 (F, GH, NY, uc, us, USM). San Martin: Prov. Rioja, Pedro Ruiz- Moyobamba road, km 390, Venceremos, D. Smith & Vdsquez 4630 (MO, NY). Huanuco: Prov. Pachitea, near bridge Oxapampa-Pozuzo road, Skog et al. 5094 (NY). Pasco: Prov. Oxapampa, ca. 3 km NE del Puente Pau- cartambo, camino a Oxapampa, Leon 624 (F, USM). Ju- nin: Prov. Tarma, Utcuyacu, Woytkowski 37006 (MO, s, uc). Cuzco: Prov. Convention, valle de Santa Ana, Plow- man & Davis 4800 (F, GH). Puno: Prov. Carabaya, km 62 road Macusani-San Gaban, Maas et al. 6051 (USM). 17. Campyloneurum phyllitidis (L.) Presl, Tent, pterid. 190. 1836. Figure lib. Polypodium phyllitidis L., Sp. pi. 2: 1083. 1753. LEC- TOTYPE (chosen by Proctor, Pteridophyta, p. 341, in Howard, Flora Lesser Antilles, vol. 2. 1977): Plumier, Descr. pi. Amer. /. 38. 1705. Cyrtophlebium phyllitidis (L.) John Sm., J. Bot. (Hooker) 4: 58. 1841. Polypodium laevigatum var. rigidum Harr., J. Linn. Soc. Bot. 16: 36. 1877. LECTOTYPE (designated by Price, Contr. Univ. Michigan Herb. 15: 203. 1982): Peru, Amazonas, Rio Utcubamba, Jul. 1872, Steere s.n. (MICH!). TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 169 Polypodium phyllitidis var. elongata Hieron., Bot. Jahrb. Syst. 34: 534. 1904. SYNTYPES: Colom- bia, Prov. Tolima, Rio Paez, Lehmann 5721 (B); Ecuador, Cerro Yanghuan, Lehmann 7679 (B!). Epiphytic, hemiepiphytic, or terrestrial. Stem (4-)6-15 mm wide, not pruinose, scales brown, triangular-lanceolate, clathrate. Leaves (45-)60- 150 cm long, 2-6(-7) mm apart; petiole almost absent to 10 cm long, sulcate or slightly sulcate adaxially, stramineous or brownish. Lamina ob- lanceolate or lanceolate, chartaceous or subcor- iaceous, (2.7-)5-16 cm wide, with attenuate base, with slightly sinuate, plane or slightly revolute, cartilaginous margins, with acuminate or subcau- date apex, indument of scarce, simple trichomes, spreading abaxially. Costa prominent, sometimes with scales similar to those of the stem; primary veins prominent or prominulous on both sides of the lamina, straight, usually lighter in color than the adjacent tissue, (5-)6-10 mm apart, diverging (55-)60-70° from the costa, secondary veins ob- solete, with (6-)8-16 areoles between costa and margin, with 3(— 4) free excurrent veinlets in each noncostal primary areole, the central excurrent veinlet connected with the transverse veinlets forming isodiametric secondary areoles. Sori sub- terminal or terminal on the excurrent veinlets, pa- raphyses inconspicuous or lacking. In shady or open places, in montane or lowland forests, 100-2500 m, Piura, La Libertad, San Mar- tin, Loreto, Huanuco, Junin, Cuzco, and Madre de Dios. Southern United States (Florida); Central America; West Indies; Colombia and Venezuela to Bolivia and central Brazil. I recognize Campyloneurum phyllitidis as dis- tinct from C. brevifolium based on venation char- acters; the former has isodiametric areoles at the middle of the lamina and two rows of sori between the primary veins. Lellinger (1988) pointed out the inconstant venation in specimens of both spe- cies from Central and South America and the need for studies on the development of the leaves to clarify the confusion. Piura: Prov. Huancabamba, carretera entre Canc- haque y Huancabamba, Diaz & Baldeon 2394 (MO, NY, USM). La Libertad: Prov. Otuzco, Chuquizongo, Lopez et al. 2630 (GH). San Martin: Tarapoto, Woytkowski 35070 (MO, uc). Loreto: Prov. Alto Amazonas, above Pongo de Manseriche, left bank of Rio Santiago, Mexia 6324 (F, GH, MO, NY, s, uc). Huanuco: Prov. Pachitea, Honoria, Bosque Nacional Iparia. 1 km arriba de Tour- navista, J. Schunke K 2181 (F, GH, NY). Junin: Prov. Satipo, Alto Quimiriqui, Leon 278 (AAU, F, USM). Cuzco: Prov. Convencion, Quillabamba, 1 50 km NW from Cuz- co, Nunez et al. 6772 (MO). Madre de Dios: Prov. Manu, Parque Nacional Manu, Cocha Cashu Biological Station, M. Foster P-84-68 (MO, uc). 18. Campyloneurum brevifolium (Link) Link, Fil. spec. 124. 1841. Polypodium brevifolium Link, Hort. berol. 90. 1833. TYPE: Based on cultivated plants (holotype, B!). Campyloneurum latum Moore, Index fil. 225. 1861. LECTOTYPE (designated by Proctor, Pterido- phyta, p. 342, in Howard, Flora Lesser Antilles, vol. 1977: Windward Islands, St. Vincent, Guild- ing (K!). Polypodium phyllitidis var. latum (Moore) Hook., Sp. fil. 5: 38. 1864. Polypodium latum (Moore) Sodiro, Crypt, vase. Quit. 371. 1893. Terrestrial, hemiepiphytic or epiphytic. Stem creeping, not pruinose, (4— )6— 17(-24) mm wide, the scales usually light brown, ovate-lanceolate. Leaves erect, (35-)40-120 cm long, 1— 4(-10) mm apart, petiole (2-)5-28 cm long, stramineous or brownish, indument of rather persistent broad scales similar to those of the stem. Lamina elliptic- lanceolate, chartaceous or subcoriaceous, 6-13.5 cm wide, with attenuate or subcuneate base, with cartilaginous margins, with usually acuminate, sometimes subcaudate apex, indument of incon- spicuous, entire trichomes, spreading abaxially. Costa prominent, slightly sulcate abaxially, con- vex or slightly angulate abaxially, sometimes with spreading ovate-lanceolate scales; primary veins prominent on both surfaces, straight or slightly flexuous, diverging 70-75° from the costa, (4-)6- 9 (-1 3) mm apart, with 8-1 8 areoles between costa and margin, 2-5 veinlets in each noncostal areole, free or anastomosing forming usually irregular sec- ondary areoles. Sori medial or subterminal, pa- raphyses inconspicuous or lacking. In shady and humid places, in disturbed humid forests, sometimes in rocky places, 100-1300 m, San Martin, Loreto, Huanuco, Pasco, Junin, Cuz- co, Madre de Dios, and Puno. Central America; West Indies; Colombia and Venezuela to Bolivia and Central Brazil. Campyloneurum brevifolium is the correct name for the species commonly known as C. latum. The type of the former is a specimen with one sterile leaf that displays the venation character that is distinctive in this species: the noncostal primary areoles are irregularly divided. One additional specimen of Link's species is found at B, which 170 FIELDIANA: BOTANY has a short -creeping stem and three closely spaced leaves, plus one irregular apically branched leaf. A photograph of this specimen at BM is erroneously labeled as the type. Campyloneurum brevifolium usually occurs in the same areas as C. phyllitidis, from which it can be distinguished by the characters provided in the key. See discussion under C. phyllitidis. Campyloneurum brevifolium is also closely re- lated to C. pascoense. Both species have a similar pattern of venation; however, in the former species secondary veins are not prominulous. San Martin: Prov. Mariscal Caceres, Dist. Campani- lla, Mashuyacu, J. Schunke V. 4225 (GH, us). Prov. Rio- ja, Rio Negro, Soukup 5146 (GH). Loreto: Prov. Maynas, 12 km SW of Iquitos, Croat 18257 (F, uc, USM). Prov. Alto Amazonas, Yurimaguas, lower Rio Huallaga. Killip & Smith 27668 (F, s, us). Huanuco: Prov. Huanuco, near confluence of Rio Cayumba with Huallaga, Mexia 8272 (BM, F, uc, s). Pasco: Prov. Oxapampa, Palcazu valley, D. Smith 3874 (MO, uc). Junin: Prov. Satipo, Puente Perene, Coronado 253 (GH, uc, us). Cuzco: Prov. Con- vencion, Potrero, 8 km W of Quillabamba, Tryon & Tryon 5393 (BM, F, GH, us). Madre de Dios: Prov. Manu, Rio Palotoa, tributary of Alto Madre de Dios, NW of Shintuya, Foster & Terborgh 6759 (F). Puno: Prov. Car- abaya, Hacienda Palmera-Inambari, Vargas 16149 (GH). 19. Campyloneurum pascoense R. & A. Tryon, Rhodora 84: 125. 1982. TYPE: Peru, Pasco (as Junin), Oxapampa, Soukup 2340 (holo- type, GH!; isotype, F!). Terrestrial. Stem long-creeping, 10-20 mm wide, not pruinose, the scales brown, ovate-lanceolate. Leaves 1.5-2 m long, 15-25 mm apart, petiole 8- 14 cm long, brown or red-brown. Lamina lanceo- late, chartaceous, 10-20 cm wide, with attenuate or cuneate base, with cartilaginous, slightly sinuate margins, with acuminate or subcaudate apex, in- dument inconspicuous or lacking. Costa promi- nent, indument of sparse scales similar to those of the stem; primary veins prominent on both sur- faces, stramineous, straight, (6-)9-l 1 mm apart, diverging 70-75° from costa, veinlets prominulous to prominent, with 1 2-20 primary areoles between costa and margin, with 3-7 veinlets in each non- costal areole, free or anastomosing, excurrent or sometimes decurrent. Sori subterminal or com- pital, paraphyses threadlike. In shady, very disturbed areas, usually in rem- nant or patches of montane forests, 1600-2500 m, Cajamarca, Amazonas, San Martin, Huanuco, Pasco, Ayacucho, and Cuzco. Ecuador to Bolivia. Cajamarca: Prov. Chota, Huambos, Soukup 4488 (us). Amazonas: Prov. Bagua, Cordillera Colan, SE of La Peca, Barbour 4187 (F). San Martin: Prov. Mariscal Caceres, Parque Nacional Rio Abiseo, Rio Montecristo water- shed, hill E from Pajaten ruins, Young 4 32 3 (NY). Huanu- co: Prov. Pachitea, Huacachi, near Muna, Macbride 4082 (F, us). Prov. Huanuco, km 468 on Lima-Tingo Maria road, above San Miguel Chinchao, Young & Sullivan 619 (MO, NY, uc). Pasco: Prov. Oxapampa, Cordillera Yanachaga, road over shoulder of Cerro Pajonal to Villa Rica, Foster & Smith 907 1 (F, MO). Ayacucho: Prov. La Mar, San Miguel, Urubamba valley, Cook & Gilbert 1 756 (us). Cuzco: Prov. Urubamba, Machu Picchu station, Hutchison 1760a (uc). 20. Campyloneurum nitidissimum (Mett.) Ching, Sunyatsenia 5: 263. 1940. Terrestrial, rarely epiphytic. Stem long-creep- ing, not pruinose, 8-10 mm wide, the scales dark brown, linear-lanceolate. Leaves 60-90 cm long, 4-9 mm apart, petiole 17-25 cm, dark strami- neous, glabrate. Lamina narrowly lanceolate, co- riaceous, 4-8 cm wide, the base attenuate or slight- ly cuneate, then decurrent, with plane or scarcely revolute, cartilaginous margins, with acute or acu- minate apex, glabrate. Costa prominent, slightly sulcate adaxially, plane or angulate abaxially; pri- mary veins prominent on both sides of the lamina, diverging 60-65° from the costa, with 7-12 areoles between costa and margin, with 3—4 veinlets in each areole, simple or furcate, free or anastomos- ing, usually slightly prominulous. Sori subtermi- nal on the excurrent veinlets, paraphyses incon- spicuous or lacking. Leaves in C. nitidissimum are characterized by the coriaceous texture, petiole length variable from approximately half the size of the lamina to less than '/4, and venation with secondary areoles sim- ilar to those in C. brevifolium. I recognize Campyloneurum nitidissimum with two varieties. Key to Varieties a. Lamina coriaceous, usually less than 5 cm wide 20a. var. nitidissimum a. Lamina herbaceous to chartaceous, usually more than 5 cm wide 20b. var. latior TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 171 20a. Campyloneurum nitidissimum var. nitidis- simum. Polypodium nitidissimum Mett., Ann. Sci. Nat. V, 2: 258. 1864. TYPE: Colombia, San Antonio, Jan. 1861, Lindig 363 (holotype B!; isotypes, BM!, K!; frag., us! of BM). Variety nitidissimum is only known from the type specimen from Colombia and two Peruvian specimens. Huanuco: Mufia, Macbride 4040 (F, GH, NY, us). Dept. unknown: Ruiz 12 (B). 20b. Campyloneurum nitidissimum var. latior (Rosenst.) B. Leon, comb. nov. Polypodium nitidissimum var. latior Rosenst., Repert. ' Spec. Nov. Regni Veg. 12: 474. 1913. TYPE: Bo- livia, Yungas Septentrionalis, Coroico, Buchtien 3526 (holotype, not located; isotypes, F!, s!, us!). In open and disturbed areas of montane forest, ( 1 000-) 1 200-2 1 00 m, Huanuco, Pasco, and Junin. Colombia to Bolivia. Huanuco: Rio Cayumba with Huallaga. Mexia 827 2a (uc). Pasco: Prov. Oxapampa, Canon del Huancabamba, entre Quebrada Honda y Tunqui, Leon 667 (F, USM). Junin: Prov. Chanchamayo, aprox. 12 km de San Ra- mon, camino a Tarma, antes de Carpapata, Leon 340 (F, USM). 2 1 . Campyloneurum abruptum (Lindm.) B. Leon, comb. nov. Polypodium repens var. abruptum Lindm., Ark. Bot. 1: 245. 1903. TYPE: Brazil, Matto Grosso, Serra do Itapirapuam, Lindman (Regnell Exped. I) 3345 (holotype, s!; isotype, K!). Campyloneurum nitidissimum var. abruptum (Lindm.) Leon, Ann. Missouri Bot. Gard 77: 212, 865. 1990. Terrestrial or hemiepiphytic. Stem creeping, not pruinose, 5-10 mm wide, scales blackish, linear- lanceolate. Leaves erect, 60-1 10 cm long, 2-5 mm apart, petiole 2-7 cm long, dark stramineous, gla- brate. Lamina lanceolate or elliptic-lanceolate, chartaceous or herbaceous-chartaceous, (4.5-)6.5- 13.5 cm wide, with usually abruptly cuneate or attenuate, then decurrent base, with slightly sin- uate, cartilaginous margins, with usually acumi- nate apex, indument of inconspicuous, unicellular trichomes, spreading abaxially. Costa prominent, slightly sulcate on the adaxial side, indument of caducous scales; primary veins prominent, straight, lighter in color than the adjacent tissue, diverging (60-)65-70° from the costa, (3-)5-7(-l 0) mm apart, with 9-15 areoles between costa and margin, with 2-4 excurrent veinlets in each noncoastal areole, veinlets entire or furcate, usually free, rarely form- ing irregular secondary areoles. Sori subterminal on the excurrent veinlet, paraphyses inconspicu- ous or lacking. In shady and humid places, with mosses or abundant organic debris, sometimes on rocks, rarely in exposed areas, in ravines of humid for- ests, 100-1900 m, San Martin, Junin, and Madre de Dios. Colombia and Venezuela, south to Bolivia and Brazil. Campyloneurum abruptum was misidentified as C. coarctatum because of leaf morphology; how- ever, stem diameter and stem scale characters are very different in these species. Campyloneurum abruptum has primary areoles mostly undivided, with 2 excurrent veinlets; how- ever, more veinlets may occur in each areole, and one of these can be connected to the transverse veins. Campyloneurum abruptum appears to be closely related to C. phyllitidis. In Peru, C. abrup- tum is commonly found at low elevations, and the base of leaves is usually abruptly attenuate, with the base of the lamina decurrent along the petiole. San Martin: Chazuta, Rio Huallaga, Klug 4080 (BM, F, GH, NY, s, uc, us); 1 5 km E of Shapaja, along Quebrada Chumia, Knapp 7867 (MO, USM). Prov. Mariscal Caceres, Campanilla, Mashuyacu, J. Schunke V. 4226 (F, us). Junin: E of Quimiri bridge, near La Merced, Killip & Smith 24004 (NY, us). Madre de Dios: Prov. Tambopata, SE de Puerto Maldonado, bosque del Albergue "Cuzco Amazonico," Leon 884 (F, USM). Comments Campyloneurum solutum (Klotzsch) Fee (Mem. foug. 5) Gen. fil. 258. 1852. Polypodium solutum Klotzsch, Linnaea 20: 399. 1 847. LECTOTYPES (designated here): Without exact locality, Hartweg 1493 (B!; isolectotype, BM!, LD!). Polypodium nodosum Klotzsch, Linnaea 20: 400. 1 847. TYPE: Colombia, Paramo de la Culata, Moritz 310 (holotype, B!; isotype, BM!). Campyloneurum nodosum (Klotzsch) Fee (Mem. foug. 5) Gen. fil. 258. 1852. Campyloneurum jamesonii Fee (Mem. foug. 5) Gen. fil. 259. 1852. TYPE: Ecuador, Pichincha, Quito, Jameson (holotype, not found; isotype, BM!). 172 FIELDIANA: BOTANY Polypodium angustifolium var. solutum (Klotzsch) Christ, Denkschr. Kaiserl. Akad. Wiss. Math. Na- turwiss. Kl. 79: 42. 1908. Campyloneurum angustifolium var. solutum (Klotzsch) Farw., Amer. Midi. Naturalist 12: 296. 1931. This species has been recorded from Colombia and Ecuador where it grows above 3000 m ele- vation in open rocky areas and may occur in north- ern Peru. It is closely related to Campyloneurum amphostenon, especially in pattern of venation, pruinose condition, and diameter of the stem. However, it differs from the latter species in the character of the stem scales. In C. solutum the stem scales are slightly clathrate, 5-7 mm long, 1.5-2 mm wide, base auriculate. The leaf is long-peti- olate and lanceolate, but smaller than those found in C. amphostenon. VIII. Niphidium Niphidium John Sm., Hist. fil. 99. 1875. TYPE: Niphidium americanum (Hooker) John Sm. (Polypodium americanum Hooker) = Niphi- dium longifolium (Cav.) Morton & Lell. Fig- ure 12. Plants terrestrial, epiphytic, or sometimes epi- petric. Stem slender to rather stout, short- to long- creeping, provided with clathrate scales, when epi- phytic usually with a dense mat of soft roots, when terrestrial with many long fibrous roots. Leaves monomorphic, mature ones 25-125 cm long, ar- ticulate, phyllopodia short or obsolete. Lamina simple and entire, oblong to linear-lanceolate, na- ked to slightly scaly (rarely densely so), the adaxial surface provided with dark or white-encrusted hy- dathodes. Veins anastomosing, with free, included veinlets. Sori round to sometimes oblong, borne at the juncture of veins, in a single series between the principal lateral veins and in two or more rows on each side of the midrib, paraphysate or not. Receptacle somewhat raised. Indusium lacking, the sporangia sometimes bearing trichomes. Spores monolete, bilateral, smooth, hyaline. Niphidium is confined to the Neotropics and 7 of its 10 species are Andean. It is distinguished by long, simple leaves, anastomosing veins with in- cluded, free veinlets, and roundish, exindusiate sori borne in a single series between the prominent, straight, main lateral veins. Although the genus is easily recognized, the task of separating species is not so simple. One of the principal diagnostic char- acters deals with subtle differences in the stem scales, and another with the thickness and length of the stems. Therefore, the scales must be very carefully examined and, furthermore, specimens lacking stems usually cannot be accurately iden- tified. Of nearly 300 specimens examined during this study, nearly 30 could not be identified with certainty. The treatment that follows is based principally on the recent revision of Lellinger (1972). Reference LELLINGER, D. B. 1972. A revision of the fern genus Niphidium. Amer. Fern J., 62: 101-120. Key to Species of Niphidium a. Stems 3-5(-6) mm in diameter; midrib scales persistent; primary veins abaxially not or slightly raised, most of them darker than the lamina b b. Stem scales usually appressed throughout, attached well above the base, less than 3 mm long, with erose, often fimbriate, margins; plants most commonly epiphytic 6. N. anocarpos b. Stem scales with tips usually spreading, attached near the base, at least 4 mm long, with erose or often ciliolate margins; plants most commonly epipetric 5. N. vittaria a. Stems (6-)8-l 5 mm in diameter; midrib scales deciduous or lacking, rarely persistent; primary veins abaxially raised, often lighter in color than the lamina c c. Sporangia naked; apex of stem scales filiform and often contorted 4. N. macbridei c. Sporangia capsules bearing few to 12 stout, 1-2-celled trichomes; apex of stem scales acute to attenuate d d. Stem scales not or scarcely clathrate (the lumina small and occluded), longitudinally somewhat involute toward the apex, especially on those scales near the petiole base . . 3. N. carinatum d. tem scales strongly and obviously clathrate (their lumina translucent), essentially plane through- out TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 173 e. Lumina of the central portion of stem scales obviously elongate, often 3-5 x as long as broad; plants epiphytic, very rarely epipetric 1 . N. crassifolium e. Lumina of the central portion of stem scales essentially isodiametric, or toward the apex slightly elongate (l-2x as long as broad); plants terrestrial or epipetric, rarely epiphytic . 2. N. albopunctatissimum 1. Niphidium crassifolium (L.) Lell., Amer. Fern J. 62: 106. 1972. Figures 12a-b. Polypodium crassifolium L., Sp. pi. 1083. 1753. LEC- TOTYPE (designated by Lellinger, 1972): Petiv- er, "Pteri-graphia Americana," /. 6, f. 1. 1712. For additional synonymy, see Lellinger (1972, p. 106). Plants epiphytic, very rarely epipetric. Stem creeping, 6-12 mm in diameter, rarely pruinose, provided with lanceolate-acuminate scales, these 8-12 mm long, plane throughout, conspicuously clathrate, usually strongly bicolorous, the central portion blackish, the margins pale brown, entire or repand, lumina of the central portion conspic- uously elongate, the distal ones often 3-5 times longer than broad. Leaves crowded, to 1 .5 cm apart, on phyllopodia 5 mm long. Petiole 3-16(-30) cm long. Lamina narrowly lanceolate or oblanceolate, attenuate or acuminate at base, acute or rounded at apex, 40-80 cm long, (3.5-)5-l 2 cm broad, pro- vided adaxially with brown hydathodes, these rarely with a white encrustation. Midrib scales de- ciduous or lacking, rarely seen. Primary veins abaxially raised, often lighter in color than the laminar tissue. Sori in (6-) 7- 14 rows on each side of the midrib (at widest part of lamina). Sporangia with a few to 10, stout, 1-2-celled trichomes on the capsule. In or at edges of forests, on trunks and branches of trees, very rarely among rocks, 100— 2 100(-2600) m, Tumbes to Loreto, south to Puno. Neotropics. This is the only South American species of M- phidium that also occurs elsewhere. Scarcely dis- tinct from it is N. albopunctatissimum (see further discussion below). Herrera 3284 (us) from Cuzco, at 2200 m, at first appears to be N. macbridei as it has very small, narrow leaves and stem scales typical of the latter. But sporangia have conspicuous trichomes, so it is perhaps a depauperate or precociously fertile specimen of N. crossifolium. Tumbes: Prov. Zarumilla, Bosque Nacional de Tumbes, Schunke V. 2381 (F, GH). Cajamarca: Prov. Contumaza, between Cascas and Contumaza, Sagdstegui et al. 9108 (GH, MO). Amazonas: Prov. Bongara, El Ingenio, Sagds- tegui 5959 (GH, HUT). Prov. Chachapoyas, slopes of Puma- urcu, Wurdack 553 (F, uc, us, USM). San Martin: Prov. Mariscal Caceres, Dist. Tocache Nuevo, left bank of Rio Huallaga, Schunke V. 3636 (F, GH). Loreto: Above Pongo de Manseriche, Mexia 6 195 a (F, GH, MO, uc). Pasco: Prov. Oxapampa, km 36-37, road to Oxapampa, Leon 479 (USM). Junin: Chanchamayo, San Ramon, near La Merced, Teppner 73/207 (us). Cuzco: Machu Picchu, Cook & Gilbert 864 (GH, us). Puno: La Pampa, Rio Ta- vara, Watkins (us). 2. Niphidium albopunctatissimum Lell., Amer. Fern J. 62: 109. 1972. TYPE: Bolivia, Prov. La Paz, Apolo, R. S. Williams 1062 (holo- type, NY; isotypes, GH!, us!). Polypodium albopunctatissimum Linden, Cat. 1860, nom. nud. Pleuridium albopunctatissimum John Sm., Ferns Brit. for., ed. 1: 95. 1866, nom. nud. Plants terrestrial, occasionally epipetric, very rarely epiphytic. Stem creeping, 8-10 mm in di- ameter, often pruinose, provided with broadly lan- ceolate, acuminate scales, these 7-10 mm long, plane throughout, conspicuously clathrate, usually strongly bicolorous, the central portion dark brown, the margins pale brown, entire or repand, lumina of the central portion essentially isodiametric or, toward the apex, slightly longer than broad. Leaves crowded, to 1.5 cm apart, on phyllopodia 2-10 mm long. Petiole 4—30 cm long. Lamina lanceo- late, attenuate or acuminate at base, acuminate or acute at apex (20-)35-80 cm long, (3-)5-12 cm broad, provided adaxially with (usually) white-en- crusted hydathodes. Midrib scales deciduous or lacking, rarely seen. Primary veins abaxially raised, often lighter in color than the laminar tissue. Sori in (5— )6— 12 rows on each side of the midrib (at widest part of lamina). Sporangia with a few to 10 stout, (2-)3-celled trichomes on the capsule. In or at the edges of forests or thickets, on the FIG. 1 2. Niphidium crassifolium: a, habit; b, section of sterile lamina. Niphidium macbridei: c, stem scale. (Adapted in part from Stolze, Ferns and Fern Allies of Guatemala, 1981.) 174 FIELDIANA: BOTANY TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 175 forest floor or on road cuts or riverbanks, occa- sionally in rock crevices, very rarely on tree trunks, (300-) 1600-3 500 m, Piura to San Martin, south- ward to Huancavelica and Puno. Southern Colombia to Bolivia. This is scarcely distinct from N. crassifolium and perhaps might better be considered an elevational or geographic variant. The principal differences cited by Lellinger are the relative shape of lumina in the central, darker, portion of stem scales, and the relative width of the lighter colored distal mar- gins. These are very subtle and often somewhat variable features that are usually supported by habit and elevation: N. crassifolium 95% of the time is an epiphyte, usually growing at lower elevation, and N. albopunctatissimum is usually terrestrial at higher elevation. Additionally, the latter often has pruinose stems and white encrustations on the hy- dathodes, whereas N. crassifolium seldom has pru- inose stems and white hydathodes. Unfortunately, none of these supporting features are constant, so the two taxa must frequently be identified by using a majority of the suite of characters. Piura: Prov. Huancabamba, Dist. Sondor, Cerro La Viuda, Sagdstegui et al. 8201 (GH, HUT, MO, us). Caja- marca: Prov. Celendin, road to Gelig, Mostacero et al. 852 (F, HUT, MO). Amazonas: Prov. Chachapoyas, slopes of Puma-urcu SE of Chachapoyas, Wurdack 553 (F, us, USM). La Libertad: Prov. & Dist. Otuzco, near Chaul- lacocha, Sounders 897 (F, GH). San Martin: Prov. Mar- iscal Caceres, Rio Abiseo National Park, Young 1381 (HUT, USM). Huanuco: Muna, Bryan 676 (F, us). Pasco: Prov. Oxapampa, road to Maria Teresa and Llaupi, Fos- ter 7611 (F, MO). Junin: Prov. Tarma, E of Huasahuasi, Hutchison 1124 (F, MO, us). Huancavelica: Prov. Taya- caja, Chuspi-Tocas, between Colcabamba and Paucar- bamba, Tovar 2123 (GH, USM). Cuzco: Prov. La Con- vencion, below San Martin on Rio Apurimac, Davis et al. 1327 (F, GH). Puno: Prov. Sandia, between Sandia and Cuyocuyo, Ferreyra 16804 (GH, USM). 3. Niphidium carinatum Lell., Amer. Fern J. 62: 113. 1972. TYPE: Peru, Huanuco, Hacienda Mercedes, Cayumba, Mexia 8223 (holotype, uc; isotypes, BM!, F!, GH!, MICH, MO; photos, F, MO & us of uc). Plants epiphytic or terrestrial, rarely epipetric. Stem short-creeping, 6-10 mm in diameter, not pruinose, provided with lanceolate, acute to at- tenuate scales, these 8-10 mm long, longitudinally somewhat involute toward apex, not or scarcely clathrate (lumina small and occluded), bicolorous, the central portion dark brown to castaneous, the margins pale brown, entire or repand. Leaves crowded, to 1 cm apart, on phyllopodia 5-6 mm long. Petiole commonly obsolete, rarely to 8 cm long. Lamina linear-lanceolate, attenuate at base, acute to rounded at apex, 30-65 cm long, 2—4.5 (-5) cm broad, provided adaxially with incon- spicuous, light brown hydathodes, or these some- times with a white incrustation. Midrib scales lacking. Primary veins abaxially raised, often light- er in color than the laminar tissue. Sori in 4-7 rows on each side of the midrib (at widest part of lamina). Sporangia with a few, to 1 2, stout, linear, unicellular trichomes on the capsule. In forests or open places, commonly on trees, occasionally on road banks or in grassland, very rarely among rocks, 700-2200 m, along the Cor- dillera Central from Amazonas to Cuzco. Colombia; Peru; Bolivia. This species has rather narrow leaves crowded along a thick, short-creeping rhizome, but the scale characters that finally separate it from its nearest relatives are very subtle ones and often difficult to perceive. As Lellinger (1972, p. 113) observed in his revision of the genus, "it is the only species of the genus with involute— not plane— rhizome scales; the cells of the central band are only weakly clathrate and have occluded lumina." However, the scales are involute only toward the apex, and then not strongly so. Therefore, careful study un- der high magnification is necessary to assess this character. Amazonas: Prov. Bongara, trail to Shillac north of Pedro Ruiz, D. Smith & Vdsquez 4951 (F, MO, uc). San Martin: Prov. Lamas, Alonso de Alvarado, Fundo las "Flores," Schunke V. 6239 (F). Huanuco: Confluence of Monzon and Huallaga Rivers, near Tingo Maria, Stork & Norton 9502 (F, uc, us). Pasco: Prov. Oxapampa, Gran Pajonal, vicinity of Chequitavo, D. Smith 5274 (F, MO, uc). Junin: Huacapistana, Coronado 239 (GH, uc). Aya- cucho: Prov. La Mar, 25 km from Tambo San Miguel, Dudley 1 1881 (GH). Cuzco: Aina (Ayna), between Huanta and Rio Apurimac, Killip & Smith 22775 (us). 4. Niphidium macbridei Lell., Amer. Fern J. 62: 117. 1972. TYPE: Peru, Junin, Yanahuanca, Macbride & Featherstone 1234 (holotype, us!; isotypes, F!, GH!; photo, GH & uc of us). Figure 12c. Plants epipetric. Stem creeping, 6-10 mm in diameter, pruinose, provided with lanceolate, long- attenuate scales, these 6-8(-10) mm long, obvi- ously clathrate (the lumina large, clear, and con- spicuous), bicolorous, the central portion blackish, the margins pale brown, irregularly ciliolate-den- tate (at least distally), the apex filiform and often 176 FIELDIANA: BOTANY contorted, the cells at this point uniseriate. Leaves commonly 0.5-2 mm apart, on phyllopodia 2-5 mm long. Petiole (3-)6-18 cm long. Lamina nar- row-lanceolate, attenuate at base, rounded or acute at apex, 35-60 cm long, (2.5-)3.5-6.5 cm broad, provided adaxially with inconspicuous, whitish hydathodes. Midrib scales deciduous or lacking, rarely persistent. Primary veins abaxially raised, often lighter in color than the lamina tissue. Sori in 4-7(-9) rows on each side of the midrib (at widest part of lamina). Sporangia lacking tri- chomes. In forests or open places, among rocks or in crevices of rock cliffs, 2500-3600 m, La Libertad, Huanuco, Junin, Apurimac, and Cuzco. Endemic. Besides the naked sporangia capsules, N. mac- bridei can also be distinguished by the apices of stem scales. The long and filiform tips are often strongly contorted and consist of a single series of cells. Scales of similar species may be very nar- rowly acute to somewhat attenuate, but never with a filiform, uniseriate tip. La Libertad: Between Huamachuco and Cajabamba, Correll & Smith P917 (GH). Huanuco: Prov. Pachitea, Panao, Asplund 13619 (us). Junin: Main road below Palca, Correll & Smith P760 (GH). Apurimac: Trail be- tween Huancarama and Abancay, West 3765, (uc), 7765 (uc). Cuzco: Ollantaytambo, Cook & Gilbert 544 (us). Prov. Urubamba, Chincheros, Davis et al. 1751 (F), 7777 (F). Urubamba, Soukup 158 (F, GH). 5. Niphidium vittaria (Mett.) Lell., Amer. Fern J. 62: 119. 1972. Polypodium vittaria Mett., Fil. Lechl. 1 : 8. 1 856. TYPE: Peru, Puno, Tabina, Lechler 2039 (holotype, B!, Herb. Men.; isotypes, L, w; photos, F of B, us of B&L). Plants epipetric, rarely epiphytic. Stem long- creeping, 3-5.5 mm in diameter, amply provided with lanceolate, long-attenuate scales, these 4-6 mm long, attached near the broadened base, their filiform tips rather conspicuously spreading, bi- colorous, the central portion castaneous to black- ish, the margins orange or pale brown, erose or often irregularly ciliolate. Leaves commonly 1.5- 3 cm apart, on phyllopodia 3-5 mm long. Petiole 3-10 cm long. Lamina linear-lanceolate, attenuate at both ends, rarely rounded at apex, 30-50 cm long, 0.8-2.2(-3) cm broad, provided adaxially with blackish or white-encrusted hydathodes. Midrib sparsely provided abaxially with (usually) persis- tent scales like those of the stem, the base rounded, or occasionally sucordate with auricles approxi- mate or overlapping. Primary veins abaxially not or slightly raised, most of them darker in color than the lamina. Sori in 2-4 rows on each side of the midrib (at widest part of lamina). Sporangia rarely with several 1- or 2-celled trichomes on the capsule. In forests, on and among rocks, or on rocky cliffs, rarely on trees, 1200-3200 m, La Libertad, Huanuco, Junin, Cuzco, and Puno. Endemic. This and 7V. anocarpos are usually easily distin- guished from other species in Peru by the slender, long-creeping stems and the very narrow, well- spaced leaves. Both also have subpersistent scales scattered along the midrib abaxially, whereas mid- rib scales in other species of Niphidium are lacking or caducous. See TV. anocarpos for further discus- sion. La Libertad: Prov. Santiago de Chuco, Motil-Shorey, Sagdstegui et al. 11699 (MO) (F specimen is Campylo- neurum). Huanuco: Prov. Pachitea, Chaglla, Huapalla 2028 (USM). Muna, Macbride 3964 (us). Junin: Yaupe, Woytkowski 6496 (MO, us). Cuzco: Prov. Cuzco, banks of Rio Urubamba, Angulo 1755 (GH, HUT). Puno: Olla- chea to San Gabon (Gaban), Dillon et al. 1151 (F, MO). Prov. Carabaya, Ollachea to Quillabamba, Vargas 6905 (uc, us), 77574 (GH). 6. Niphidium anocarpos (Kunze) Lell., Amer. Fern J. 62: 120. 1972. Polypodium anocarpos Kunze, Linnaea 9: 40, 1834. TYPE: Peru, Huanuco, near Pampayaco (Pam- payacu), Poeppig 1 128 (holotype, LZ, destroyed; isotype, designated as lectotype by Lellinger, B, Herb. Mett; fragment, us!; photo, us of B). Polypodium acrosorum Kunze, Linnaea 9: 39. 1834. TYPE: Peru, Huanuco, near Pampayaco (Pam- payacu), Poeppig 1112 (holotype, LZ, destroyed; isotype, designated as lectotype by Lellinger, HBG; frag., us!; isolectotypes, BR, HBG, L, LE, MO!; pho- tos, F, GH, MO, UC & US of HBG). Plants epiphytic or terrestrial, occasionally epi- petric. Stem long-creeping, 3-5.5(-6) mm in di- ameter, copiously provided with ovate to ovate- lanceolate, acute to acuminate scales, these 2-3 mm long, tightly apressed, attached well above the base, bicolorous, the central portion black, the margins castaneous to pale brown, erose to short- fimbriate (the fringe often abraded in age). Leaves commonly 1-3 cm apart, on phyllopodia 2—4 mm long. Petiole 1-14 cm long. Lamina linear- to nar- TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 177 178 FIELDIANA: BOTANY rowly elliptic-lanceolate, attenuate at both ends, 40-70 cm long, 1-3 cm broad, provided adaxially with brown hydathodes. Midrib sparsely provided abaxially with (usually) persistent scales similar to those of the stem, the base rounded, or occasion- ally cordate with an open sinus between the au- ricles. Primary veins abaxially not or slightly raised, most of them darker in color than the lamina. Sori in 3—4 rows on each side of the midrib (at widest part of lamina). Sporangia with several unicellular trichomes persistent on the capsule. In forests, on trees and logs, sometimes on the forest floor, rarely on rocks, 1200-3950 m, Ca- jamarca, San Martin, Huanuco, Pasco, Cuzco, and Puno. Peru; Bolivia. This and N. vittaria are very similar and are maintained here, with some reluctance, as distinct. They are distinguished, at least in Peru, principally by subtle differences in their stem scales and hab- itat. Those of the latter species are commonly lon- ger and narrower, with filiform tips spreading con- spicuously from the stem. Their point of attachment is somewhat above the base but ap- pears to be relatively nearer the base because of the long-attenuate scale apices. Stem scales of N. anocarpos are shorter and broader, with subacute or short-acuminate tips and are attached relatively near the scale center. Scale margins of both species are eorse, but sometimes irregularly ciliolate along the attenuate tips in AT. vittaria, or sometimes short- fimbriate in N. anocarpos. Tryon & Tryon 5392 (F, GH, us), from Prov. La Convention, Cuzco, is intermediate; scales are short (3 mm) and the base fimbriate, but the somewhat attenuate tips are ir- regularly tiliolate. Caiamarca: La Palma, 10 k NW of Chirinos, Gentry et al. 61190 (MO, uc). San Martin: Prov. Rioja, Ven- ceremos, D. Smith 4514 (MO, uc, USM), 46 36 (MO, uc, USM). Huanuco: Tambo de Vaca, Bryan 675 (F, us). Muna, Macbride 3975 (F, GH, us), 4041 (F, us). Pasco: Prov. Oxapampa, vicinity of Huanacabamba, D. Smith et al. 1679 (F, MO). Cuzco: Prov. La Convension, Dist. Vil- cabamba. trail from Yupanqui to Apurimac, Davis et al. 1231 (GH). Urubamba, Machu Picchu, Peyton & Peyton 595 (GH), 1458 (GH, MO). Puno: Prov. Carabaya, below Ollachea, Vargas 6905 (us). IX. Solanopteris Solanopteris Copel. Amer. Fern J. 41: 75, 128, 1951. TYPE: Solanopteris bifrons (Hooker) Copel. (Polypodium bifrons Hooker). Figure 13. Microgramma subg. Solanopteris (Copel.) Lell., Amer. FemJ. 67: 59. 1977. Plants epiphytic. Stem usually pruinose, long- creeping, commonly with highly modified, swol- len, hollow (in age) tubers borne on short, lateral branches, with mostly circular scales and usually few roots. Leaves strongly to weakly dimorphic (the fertile commonly longer and narrower than the sterile), 3-1 8 cm long, well spaced on the stem, occasionally borne on a tuber, articulate to short phyllopodia on the stem. Lamina entire to pin- nately lobed, naked or sparsely scaly, attenuate to base, the petiole essentially obsolete. Veins anas- tomosing, with included, free veinlets spreading in all directions. Sori discrete and round to oval, or merged into continuous or interrupted lines, on a raised receptacle, served by few to many veins, exindusiate, sporangia intermixed with abundant paraphyses. Spores monolete, ellipsoid, echinate with a dense glandular deposit on the spines. Solanopteris contains three or four species, all in South America, with one, S. brunei (Christ) Wagner, also occurring in Costa Rica. This and Lecanopteris are unique in the family in the swol- len tubers which commonly are inhabited by ants, and the long-spiny perispore of the spores (Hen- nipman, 1990, p. 423; Tryon & Tryon, 1972, p. 737). It further differs from Microgramma, with which it was included by Lellinger (1977), by the small, mostly circular, stem scales. The plants are mostly high epiphytes, often growing in upper branches of trees, so they are easily overlooked; thus species may be far more widespread than currently supposed. Two species thus far have been found in Peru, but the other two (which may be synonymous) eventually may be collected here as well. See treatment of S. bismarckii for further discussion. FIG. 13. Solanopteris bifrons: a, habit, with sterile leaves and tuber; b, fertile leaf; c, section of sterile lamina. Solanopteris bismarckii: d, habit, with fertile leaf, (a, b, c from King 2643, GH; d from Rauh 35685, USM.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 179 The genus was initially published as Solenop- teris. However Copeland immediately corrected it to Solanopteris in the index and in errata in the same publication. References HENNIPMAN, E. 1 990. The significance of the SEM for character analysis of spores of Polypodi- aceae. Systematics Association, 41: 23—44. Clar- endon Press, Oxford. LELLINGER, D. B. 1977. Nomenclatural notes on some ferns of Costa Rica, Panama, and Colom- bia. Amer. Fern J., 67: 58-60. RAUH, W. 1955. Botanische Mitteilungen aus den Anden, I. Morphologische und anatomische Betrachtungen an Polypodium bifrons Hooker. Abh. Akad. Wiss. Abh. Lit. Mainz, Math.-Na- turwiss. KL, 3: 45-57. . 1973. Solanopteris bismarckii Rauh, Abh. Akad. Wiss. Abh. Lit. Mainz, Math.-Na- turwiss. Kl., 3: 223-256; and Tropische und subtropische Planzenwelt, 5: 5-38. TRYON, R. M., AND A. F. TRYON. 1982. Solan- opteris, pp. 735-738. In Ferns and Allied Plants. Springer- Verlag, New York. Key to Species of Solanopteris a. Sterile leaf entire to (rarely) slightly repand, chartaceous to subcoriaceous; stem scales copious, often obscuring the surface of the stem; sori merged into a continuous or interrupted line, the discrete portions (if any) more than twice as long as broad 1 . S. bismarckii a. Sterile leaf pinnately lobed, thin-herbaceous; stem scales sparsely scattered; sori usually discrete, round or oval . . 2. S. bifrons 1. Solanopteris bismarckii Rauh, Akad. Wiss. Abh. Math.-Naturwiss. Kl. 5: 232. 1973. TYPE: Peru, Pasco, Chanchamayo Valley near Oxapampa, Ceja de Montana, Rauh 25790 (holotype, HEID). Figure 13d. Plants epiphytic. Stems to 3 mm thick, bearing globular, swollen, often ant-inhabited tubers to 2.5 cm in diameter, stem scales abundant, often so crowded as to obscure the stem surface, most of them round, 0.2-0.5 mm in diameter, but oc- casionally short- and broad-ovate, conspicuously bicolorous, castaneous at point of attachment and with narrow or broad, white or ash-gray margins. Leaves weakly dimorphic, sterile ones 3-5 cm long, 0.6-1 .4 cm broad, fertile ones relatively longer and narrower. Lamina lanceolate to oblanceolate, chartaceous to subcoriaceous, the margins entire or weakly repand, sparsely dotted with small, ovate, castaneous scales, these especially along the costa abaxially. Veins indistinct, secondary and teritary veinlets about the same thickness and prominence as the primary ones which enclose them. Sporan- gia merged into a continuous, long, line which, at maturity, covers the lamina between costa and margin. In forests, creeping along the branches of trees, 1 100-2000 m, San Martin, Pasco, and Cuzco. Apparently endemic. This might be confused with S1. tuberosa (Max- on) Rauh, of Ecuador, which has similar (although slightly larger) leaves and continuous to somewhat interrupted sori. But the latter differs significantly in type of venation and stem scales. In S. tuberosa primary veins are thicker and more prominent than the subsidiary veinlets they enclose. Stem scales, though often round, are predominantly lan- ceolate-attenuate and castaneous, with very nar- row and tawny margins. In S. bismarckii (as well as in S. bifrons) all veins are of the same thickness, and stem scales are predominantly circular and with mostly broad, whitish margins. Solanopteris tuberosa scarcely differs from S. brunei (Christ) Wagner (Costa Rica, Panama, Co- lombia). Both have similar leaf shape, venation, and stem scales, and have been separated essen- tially by the discrete, round sori of the latter. How- ever, there is enough variability in this character to render it of questionable value. Distribution of the 5. brunei/tuberosa complex may extend from Costa Rica to Ecuador, and perhaps even into Peru (pending more careful exploration in the canopies of tropical forests). San Martin: Prov. San Martin, Rioja-Pomacochas Road, below Venceremos, Gentry & Smith 45184 (MO). Pasco: Prov. Oxapampa, 10 km from Oxapampa, Rauh 180 FIELDIANA: BOTANY 35685 (USM). Cuzco: "Cuyulichayoc" Hacienda, Cco- chayoc, Bites 1732 (us). 2. Solanopteris bifrons (Hooker) Copel., Amer. Fern J. 41: 75, 128. 1951. Figures 13a-c. Polypodium bifrons Hooker, Fil. exot., /. 52. 1859. TYPE: Ecuador, riverside near Archidona, Jame- son 789 (holotype, K; frag., us!). Microgramma bifrons (Hooker) Lell., Amer. Fern J. 67: 59. 1977. Plants epiphytic. Stems 1-2 mm thick, bearing spherical, swollen, ant-infested tubers to 2.5 cm in diameter, stem scales sparsely scattered, cir- cular, 0.2-0.4 mm in diameter, somewhat bico- lorous, castaneous, with a rather narrow, orange (rarely whitish) margin. Leaves strongly dimor- phic; sterile ones 4-10 cm long, 1-3 cm broad; lamina elliptic, thin-herbaceous, pinnately lobed '/3-% to the costa, scales rare or lacking; veins dis- tinct, secondary and tertiary veinlets the same thickness and prominence as the primary ones which enclose them. Fertile leaves like the sterile ones, but 6-18 cm long and 0.3-1 cm broad, mar- gins sinuate to weakly lobed. Sporangia grouped into large (4-5 mm), discrete, round or oval sori, in a single series between costa and margin. In wet forests, on shrubs, small trees, or on branches in the canopy, 180-900 m, Amazonas south to Cuzco and Madre de Dios. Colombia; Ecuador; Peru. In addition to the differences noted in the key, the stem scales of S1. bismarckii are strongly bi- colorous, the margins mostly broad and whitish; whereas scale margins of S. bifrons are narrow and orange-brown. Furthermore, S. bifrons occurs at lower elevations than S. bismarckii, the former under and the latter over 1000 m. Amazonas: Rio Santiago, 700 m past Caterpiza, Hua- shikat 754 (MO). San Martin: Pongo de Cainarachi, Rio Cainarachi, Klug 2643 (F, GH, us). Km 28, road to Yu- rimaguas, Maguire61522 (uc). Tarapoto, El Pongo, Rauh (USM). Pasco: Prov. Oxapampa, Palcazu, Rio Alto Is- cozacin, Ozuz, Foster & d'Achille 9957 (F). Drainage of Rio Palcazu on road NW of Villa Rica, Gentry & Smith 36054 (F, MO). Ucayali (as Loreto): Prov. Coronel Por- tillo, lower Boqueron of Padre Abad, Hutchison et al. 6039 (F, uc, us). Cuzco: Prov. Paucartambo, Cosnipata Valley, Rio Tono Road, Foster et al. 10578 (F). Prov. Quispicanchi, Inambari, Vargas 13497 (GH). Madre de Dios: Manu, Rio Salvation, Nunez et al. 8067 (F, MO). X. Platycerium Platycerium Desv., Mem. Linn. Soc. Paris 6: 213. 1827. TYPE: Platycerium alcicorne Desv., nom. nov. for Acrostichum alcicorne Sw. 1801, not Willemet, 1796. Figure 14. Plants epiphytic. Stem ascending, stout, provid- ed with scales and many fibrous roots. Leaves sol- itary or fasciculate, dimorphic (or in one African species monomorphic), the sterile, "nest" leaves 25-100 cm long, appressed at least basally, with petiole continuous, the fertile ones to 3 m long, erect-spreading to (more commonly) pendent, with petiole articulate. Lamina entire or lobed to di- chotomously forked, closely and finely pubescent, with stellate trichomes. Veins anastomosing, with or without included, free veinlets. Sporangia borne in soral patches in certain discrete, extensive areas of the fertile lamina, along the nearly unmodified veins, with or without paraphyses. Indusia lacking. Spores monolete, ellipsoid, the surface nearly smooth, but with a dense, spherical deposition. The genus consists of 14 species in Asia and Africa and only one in America. The species, all epiphytic, occur in lower to middle elevations in tropical or subtropical forests. They grow on the trunks or lowest forks of trees or in the higher branches. The broad, overlapping "nest" leaves are modified for collection of humus. Sporangia, in all but two species, are borne along the veins, but at maturity they are so abundant and crowded they appear acrostichoid. References HENNIPMAN, E., AND M. C. Roos. 1982. A monograph of the fern genus Platycerium. Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk., Tweede Sect., 80: 1-126. HOSHIZAKI, B. J. 1972. Morphology and phy- logeny of Platycerium species. Biotropica, 4: 93- 117. 1. Platycerium andinum Baker, Ann. Bot. (Lon- don) 5: 496. 1891. LECTOTYPE (designated by Hennipman and Roos, 1982): Peru, San Martin (as Loreto), Tarapoto, Spruce 4739 (K; isolectotype, P). Figures 14a-c. Plants epiphytic. Stem abundantly provided with linear or linear-lanceolate, attenuate, basally at- TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 181 FIG. 14. Platycerium andinum: a, habit, sterile leaf; b, distal portion of fertile leaf; c, venation of sterile leaf near margin, (a from LI. Williams 5734, F; b, c from King 3872, F.) 182 FIELDIANA: BOTANY tached scales, these 0.5-1.5 cm long, light or red- dish brown. Leaves fasciculate, densely pubescent with stellate trichomes, these often obscuring the fertile lamina abaxially. Nest leaves sessile, wedge- shaped, entire, or lobed apically, soon turning brown: veins copiously reticulate, primary and secondary ones prominulous and conspicuous, with frequent, usually obscure, free, included veinlets. Fertile leaves short-stalked, 1-3 m long, 3-5 times subdichotomously forked; primary veins promi- nent, the rest obscure; soral patches usually ex- tending from near the first fork of the lamina to beyond the ultimate fork. On tree trunks in wet forests, 210-800 m, San Martin and Junin. Peru; Bolivia. The dichotomously branching, pendent, fertile leaves and the sterile, clasping, nest leaves are so distinctive that this cannot be confused with any species of fern in Peru. It apparently is confined to the low, wet forests of Peru and Bolivia, al- though Hennipman and Roos (1982) saw a spec- imen with a label that was illegible except for the word "Ecuador." San Martin: Juan Jui, Alto Rio Huallaga, Klug 3872 (F, OH, MO, us). Along Rio Huallaga, NE of Shapaja on road to Cazuta, Knapp & Mallet 6921 (F, MO). Prov. Mariscal Caceres, Dist. Pajarillo, Schunke V. J3493(usM). Alto Rio Huallaga, LI. Williams 5734 (F, us), 66 38 (F). Junin: Prov. Satipo, 1 km from Puerto Ocopa, road from Mazamari to Puerto Ocopa, Fernandez & Callegari 683 (USM). TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 183 -^••v Colombia 1. Tumbes 2. Piura 3. Lambayeque 4. Cajamarca 5. Amazonas 6. La Libertad 7. San Martin 8. Loreto 9. Ancash 10. Huanuco 11. Lima 12. Pasco 13. Junin 14. Ucayali 15. lea 16. Huancavelica 1 7. Ayacucho 1 8. Apurfinac 1 9. Cuzco 20. Madre de Dios 21. Arequlpa 22. Puno 23. Moquegua 24. Tacna Chile DEPARTMENTS OF PERU 184 FIELDIANA: BOTANY Index to Names Accepted names are in roman type, synonyms are in italics, and new names are in boldface. A page number is provided for the principal place, or the only place, where the name occurs. Acrospermum maxonii 1 12 Acrostichum alcicorne 181 graminoides 72 heterophyllum 63 polypodioides 137 reptans 153 serrulatum 83 Aspidium biserratum 51 pectinatum 53 rutaceum 17 Aspleniaceae 2 Asplenium 2 abrotanoides 27 abscissum 41 var. abscissum 41 var. ruizianum 42 var. subaequilaterale 4 1 achilleifolium 32 aethiopicum 46 alatum 40 amazonicum 13 angustum 12 auriculatum 44 auritum 42 var. tripinnatum 27 ballivianii 21 bangii 38 barbaense 37 blechnoides 60 brasiliense 40 canelense 30 castaneum 17 cicutarium 24 cirrhatum 20 cladolepton 24 var. anguslifolium 24 var. minor 24 claussenii 39 concinnum 4 congestum 41 conquisitum 18 cristatum 24 cuneatum 44 cuspidatum 26 var. abrotanoides 27 var. cuspidatum 27 var. foeniculaceum 27 var. tripinnatum 27 davisii 30 delicatulum 25 delitescens 32 dentation 35 denudatum 48 dimidiatum 46 discolor 48 discrepans 36 divaricatum 23 drepanophyllum 42 erosum 46 escaleroense 14 escragnollei 30 extensum 15 falcinellum 28 flabellulatum 21 var. dentatum 21 var. partitum 2 1 flavidum 30 foeniculaceum 27 formosum 14 fragile 34 fragrans 27 gillesianum 34 gilliesii 33 haenkeanum 24 hallii 18 haplophyllum 13 harpeodes 39 hastatum 44 herzogii 38 integerrimum 28 jamesonii 26 juglandifolium 28 kapplerianum 29 laetum 31 lividum 47 longicaudatum 19 lorentzii 33 macrurum 19 marinum 2 mathewsii 28 maxonii 18 melanopus 31 monanthemum 16 monanthes 16 var. castaneum 17 var. monanthes 16 var. wagneri 17 myriophyllum 25 neogranatense 29 nidus 1 1 nigricans 46 oligophyllum 30 otites 48 partitum 21 parvulum 15 pearcei 13 perkinsii 18 peruvianum 34 poloense 35 praemorsum 46 projectum 33 pseudoangustum 12 pteropus 36 pulchellum 35 pumilum 14 purdieanum 47 purpurascens 32 quitense 35 raddianum 39 rad leans 19 var. cirrhatum 20 var. dentatum 21 var. partitum 20 var. radicans 20 var. uniseriale 21 repandulum 28 repens 2 1 resiliens 15 rhizophyllum 20 rhomboideum 22 rosenstockianum 37 ruizianum 43 rusbyanum 30 rutaceum 17 salicifolium 43 scolopendrium 4 serra 45 var. remotum 45 serratum 1 1 sessilifolium 37 spruceanum 18 squamosum 26 stuebelianum 12 tabinense 38 tenue 22 ternatum 22 theciferum 48 trapezoides 48 tricholepis 30 trichomanes-dentatum 34 trilobatum 22 trilobum 48 triphyllum 22 tucumanense 26 tuerckheimii 29 uniseriale 2 1 vargasii 17 virens Desv. 31 virens Presl 30 vomeri forme 28 wagneri 1 7 weberbaueri 49 Athyrium achilleifolium 32 haenkeanum 24 Blechnaceae 54 Blcchncae 54 ttlechnum 56 subg. Blechnum 56 subg. Lomaria 56 acutum 64 TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 185 alpinum var. elongatum 62 andinum 61 angustifolium (HBK.) Hieron. 64 angustifolium Willd. 64 arborescens 62 asplenioides 60 auratum 67 ssp. auratum 67 ssp. columbiense 67 auriculatum 68 binervatum 63 ssp. acutum 64 ssp. binervatum 64 ssp. fragile 64 blechnoides 60 brasiliense 6 1 buchtienii 67 caudatum 58 chile nse 62 ciliatum 68 cognatum 68 columbiense 67 var. bogotense 67 confluens 59 cordatum 62 delicatum 68 divergens 63 ensiforme 64 fragile 64 fraxineum 59 glandulosum 58 gracile 59 heterophyllum 63 kunthianum 64 lanceola 60 lechleri 68 lehmannii 61 1'herminieri 61 longifolium 59 loxense 65 magellanicum 68 malacothrix 68 maxonii 6 1 meridense 64 nigrosquamatum 61 nudum 56 obtusifolium 67 occidentale 58 var. pubirhachis 58 x fraxineum 58 x unilaterale 58 orientate 58 ornifolium 62 pectinatum 58 penna-marina 62 peruvianum 62 polypodioides Raddi 60 polypodioides (Sw.) Kuhn 64 pteropus 63 rubicundum 65 scandens 70 schomburgkii 66 serrulatum 60 sprucei 66 squamulosum 65 stenophyllum Presl 65 stenophyllum (KJotzsch) Mett. 65 stipitellatum 65 subtile 6 1 triangulare 59 trilobum 68 unilaterale 60 volubile 70 var. lomarioideum 70 Caenoptehs achilleifolia 32 myriophylla 25 Camptosorus 2 Campyloneurum 158 abrupt um 172 aglaolepis 167 amphostenon 165 var. amphostenon 166 var. irregulare 166 angustifolium 168 var. amphostenon 166 var. solutum 173 angustipaleatum 169 aphanophlebium 161 asplundii 167 brevifolium 170 caespitosum 162 chlorolepis 168 coarctatum 164 decurrens 132 densifolium 166 fasciale 163 fendleri 132 fuscosquamatum 163 heterolepis 168 inflatum 164 irregulare 166 jamesonii 172 lapathifolium 162 /a/wra 170 lorentzii 167 magnificum 132 nitidissimum 171 var. abruptum 172 var. latior 172 var. nitidissimum 172 nodosum 172 occultum 161 ophiocaulon 162 pascoense 171 phyllitidis 169 repens 162 serpentinum 163 solutum 172 sphenodes 164 taeniosum 169 trichiatum 161 vulpinum 167 Ceradenia 72 capillaris 90 curvata 72 dendrodoxa 88 discolor 86 farinosa 89 herrerae 88 longipinnata 85 meridensis 85 mirabilis 9 1 pearcei 87 pilipes 90 praeclara 89 terrestris 87 Cochlidium 72 graminoides 72 pumilum 82 serrulatum 82 Cryptosorus 72 blumei 72 Ctenopteris 72 subg. Glyphotaenium 72 amylacea 87 anfractuosa 99 apiculata 94 asplenifolia 104 athyrioides 1 1 3 capillaris 90 congest a 89 contacta 107 crispata 72 cultrata 108 discolor 86 dolorensis 104 ecuadorensis 1 1 0 farinosa 89 ./zrma 100 gracilis 96 herrerae 88 heteromorpha 105 lanigera 106 leucosticta 1 1 1 longipinnata 85 longiuscula 1 1 1 major 96 melanosticta 98 meridensis 85 moniliformis 98 obovata 95 peruviana 102 phlegmaria 97 pilosissima 101 pseudocapillaris 94 pseudonutans 103 pteropus 95 r/gens 101 rigescens 102 semihirsuta 1 1 2 sericeolanata 106 ste//a 106 subflabelliformis 107 subimpressa 94 subsessilis 95 taxifolia 1 1 3 tunguraguae 95 venulosa 72 yungensis 1 1 3 Cuspidaria 145 furcata 145 Cyrtophlebium 158 phyllitidis 169 repens 162 186 FIELDIANA: BOTANY Davallia concinna 4 multiflora 52 thecifera 48 Davalliaceae 49 Dicranoglossum 145 desvauxii 147 furcatum 145 var. bicolor 147 var. subnudum 147 panamense 145 polypodioides 148 subnudum 147 Diplazium delitescens 32 melanopus 31 Drynaria acuminata 156 Enterosora 72 campbellii 72 parietina 84 trichosora 84 trifurcata 83 Eschatogramme 145 desvauxii 147 furcata var. subnuda 147 polypdioides 148 subnuda 147 Glyphotaenium 72 trifurcatum 83 Goniophlebium 71 sect. Lepicystis 125 incanum 125 pectinatum 121 semipinnatum 140 subauriculatum 72 Grammitis 72 albidula 87 alsopteris 1 10 amylacea 87 andicola 103 andina 109 anfractuosa 99 apiculata 94 aromatica 100 asplenifolia 104 assurgens 99 athyrioides 1 1 3 basalis 109 bipinnata 92 bishopii 86 blepharida 1 1 2 blepharolepis 108 bryophila 8 1 £>ues/t 1 1 2 campbellii 72 capiilaris 90 chrysleri 104 conge sta 89 crispata 72 cultrata 108 curvaia 72 ssp. curvata 87 ssp. pearcei 87 daguensis 109 da N id-sin ithii 109 dendrodoxa 88 dependens 104 discolor 86 dudleyi 92 elongata 143 erecla 98 farinosa 88 firma 100 flabelliformis 102 gracilis 96 graminoides 72 herrerae 88 heteromorpha 105 immixta 1 1 5 jamesonii 82 jamesonioides 93 lanceolata 143 lanigera 105 var. lanigera 106 var. Stella 106 laxa 107 lehmanniana 104 leucosticta 1 1 1 limbata 8 1 longipinnata 85 major 96 marginella 72 mathewsii 91 melanosticta 97 meridensis 85 mirabilis 91 moniliformis 98 myosuroides 82 myriophylla 1 14 nigrolimbala 81 obliquata 72 param icola 8 1 parientina 84 peruviana 102 phalacron 84 phlegmaria 97 var. antillana 97 var. phlegmaria 97 pichinchae 1 10 pichinchensis 1 10 pilipes 90 pilosissima 100 praeclara 89 pseudocapillaris 94 pseudonutans 103 pumila 82 recondita 95 revoluta 143 rigens 101 rigescens 102 sectifrons 72 semihirsuta 1 1 1 senilis var. senilis 107 sericeolanata 106 semilata 82 sprucei 84 squamulosa 143 subtlabelliformis 106 subsessilis 95 taxifolia 1 1 3 terrestris 87 trichosora 83 trifurcata 83 truncicola 109 tunguraguae 94 variabilis 1 15 venulosa 72 werfii 92 xiphopteroides 101 youngii 97 Holodictyum 2 Lellingeria 72 apiculata 94 major 96 myosuroides 82 phlegmaria 97 pseudocapillaris 94 subsessilis 95 tunguraguae 95 Lepicystis 125 incana 125 Lepisorus 140 Lomaria 56 acuta 64 andina 6 1 angustifolia 64 arborescens 62 aurata 67 caudata 66 chilensis 62 cor dot a 62 cuspidata 64 diver gens 63 ensiformis 64 fragile 64 fraxinea 59 heterophylla 63 linariaefolia 68 loxensis 65 meridensis 64 nuda 56 obtusifolia 67 ornifolia 62 pier opus 63 schomburgkii 66 serrulosa 62 squamulosa 65 stenophylla 65 stipitellata 65 volubilis 70 Loxoscaphe 4 concinna 4 thecifera 48 Marginaria 125 angustifolia 169 polypodioides 125 TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 187 Marginariopsis 71 Melpomene 96 Microgramma 148 acatallela 152 acuminata 157 baldwinii 155 bifrons 181 chrysolepis 151 ciliata 153 fuscopunctata 144 geminata 157 latevagans 151 lindbergii 157 lycopodioides 154 megalophylla 157 percussa 144 persicariifolia 157 piloselloides 152 recreense 157 reptans 153 rosmarinifolia 154 squamulosa 156 tecta 154 var. nana 154 var. tecta 154 thurnii 156 ulei 156 vacciniifolia 152 Micropoly podium 72 pseudotrichomanoides 72 Nephrolepidaceae 49 Nephrolepis 49 biserrata 5 1 cordifolia 52 exaltata 49 hirsutula 54 multiflora 52 accident alls 53 pectinata 53 pendula 52 rivularis 52 Niphidium 173 albopunctatissimum 174 americanum 173 anocarpos 177 carinatum 176 crassifolium 174 longi folium 173 macbridei 176 vittaria 177 Osmunda polypodioides 64 Parablechnum ciliatum 68 Pecluma 1 16 absidata 1 1 9 boliviano 120 camptophyllaria 122 var. abbreviata 123 var. camptophyllaria 123 var. lachnifera 123 choquetangensis 116 curvans 119 dispersa 125 divaricata 120 eurybasis 121 var. glabrescens 1 2 1 var. villosa 121 filicula 119 funicula 1 16 hygrometrica 124 pectinata 121 plumula 1 18 ptilodon 123 var. caespitosa 124 var. pilosa 124 var. ptilodon 124 venturii 122 Phlebodium 125 aureum 134 decumanum 135 Phyllitis 4 scolopendrium 4 Platycerium 181 alcorne 181 andinum 181 Pleopeltis 140 angusta 140 astrolepis 143 fuscopunctata 144 lanceolata 143 macrocarpa 142 var. complanata 142 var. crassinervata 142 var. laciniata 143 var. macrocarpa 143 var. trichophora 142 percussa 144 pinnatifida 135 revoluta 143 squamulosa 143 Pleuridium albopunctatissimum 1 74 Pleurosorus 2 Polypodiaceae 70 Tribe Loxogrammeae 7 1 Tribe Grammitideae 7 1 Polypodium 125 subg. Cyrtophlebium 158 sect. Cryptosorus 72 sect. Phlebodium 125 abitaguae 86 absidatum 1 19 acrodontium 101 acrosorum 177 adnatum 133 aglaolepis 168 albopunctatissimum 1 74 alternifolium 108 americanum 173 amphostenon 166 andinum 109 anfractuosum 99 angustifolium 168 var. gramineum 169 var. heterolepis 168 var. solutum 173 f. densifolium 166 angustipaleatum 169 anocarpos 177 aphanophlebium 161 apiculatum 94 appressum 132 aromaticum 100 articulatum 133 asplenifolium 104 asplundii 167 astrolepis 143 athyrioides 1 1 3 aureum 134 azuayense 94 balaoense 138 biauriculatum 131 bifrons 181 binervatum 63 blepharideum 1 1 2 blepharolepis 108 bolivianum 120 bombycinum 138 brevifolium 170 bryophilum 81 bryopodum 135 buchtienii 136 buesii 1 1 2 caceresii 133 caespitosum 162 camptophyllarium 123 var. abreviatum 123 var. lachnifera 123 var. macedoi 122 capillare 90 chacopoyense 131 chartaceum 129 chnoodes 129 chrysolepis 151 ciliatum 153 circinatum 119 coarctatum 164 cordatum 131 cordifolium 52 crassifolium 174 crispatum 167 crystalloneuron 139 cultratum 108 curvans 1 1 9 curvatum 72 dasypleuron 131 decumanum 135 decurrens 132 dependens 104 discolor 86 dispersum 125 dissimile 129 divaricatum 120 dolorense 104 83 140 ecostatum 85 ecuadorense 1 10 188 FIELDIANA: BOTANY eurybasis var. glabrescens 1 2 1 var. villosum 121 exaltatum 49 farinosum 89 fasciale 163 Yendleri 132 yz//CM/a 119 firmum 100 flabelliforme 102 fraseri 136 fraxinifolium 132 furfuraceum 136 fuscopunctatum 144 giganteum 133 gilliesii 130 glaucophyllum 134 gracile 96 gracillimum 108 herzogii 100 heteromorphum 105 hirsutulum 54 x. huancayanum 139 hygrometricum 124 incanum 137 var. burchellii 137 jamesonii 82 jamesonioides 93 kunthii 154 kunzeanum 131 lachniferum 123 var. glabrescens 1 2 1 laetum 130 laevigatum 134 var. crispatum 167 var. rigidum 169 x lasiopus 140 lanceolatum 142 lanigerum 106 lapathifolium 162 lasiopus 130 var. weberbauerianum 1 30 f. bipinnatifidum 130 x laevigatum 140 x mollendense 139 latevagans 151 129 170 laxum 107 leucatomos 134 leucosporum 1 39 leucosticton Fee 1 1 1 leucosticton KJotzsch 136 levigatum 134 limbatum 81 lomariiforme 125 longifolium 132 longisetosum 1 1 4 longiusculum 1 1 1 longum 108 loretense 156 loriceum 129 lycopodioides \ 54 macrocarpum Presl 135 macrocarpum Willd. 143 marginellum 72 mathewsii 91 megalolepis 137 megalophyllum 157 melanostictum 97 meridense 85 mollendense 135 moniliforme 98 monosorum 138 monlicola 99 murorurn 139 mutabile 139 myosuroides 82 myriophyllum 1 1 4 nigrolimbatum 81 nitidissimum 172 var. /af/or 172 nodosum 172 obliquatum 72 occulturn 161 ophiocaulon 162 parietinum 84 pearcei 87 pectinatum 121 pendulum var. boliviense 95 penna- marina 62 percussum 144 persicariifolium 157 peruvianum 102 phlegmaria 97 phyllitidis 169 var. elongata 170 var. /a/wm 170 pichinchae 1 10 pichinchense 1 10 pilipes 90 piloselloides 152 pilosissimum 100 plumula 1 1 8 polypodioides 137 var. burchellii 137 var. polypopdioides 137 pozuzoense 90 preslianum 132 pseudoaureum 134 pseudocapillare 94 pseudonutans 103 pteropus 95 ptilodon 124 var. pilosum 1 24 pubescens 130 pycnocarpum 135 var. buchtienii 136 var. pycnocarpum 136 ratibori 135 remotum 136 repens 162 var. abrupt um 172 richardii 133 rigens 101 rigescens 102 rivulare 52 rosmarinifolium \ 54 ruiz-lealii 135 rusbyi 135 sectifrons 72 semihirsutum 1 1 1 var. fuscosetosum 1 1 1 var. hirtopuberulum 1 1 1 semipinnatifidum 140 var. minor 107 sericeolanatum 106 serpentinum 163 serrulatum 83 var. strictissimum 82 sessilifolium 131 solutum 172 sororium 129 sphenodes 164 spixianum 86 sprucei 84 squamulosum 156 strictisissimum 82 subandinum 130 subauriculatum 71 subflabelliforme 106 var. minor 107 subscabrum KJotzsch 110 subscabrum sensu Hooker 1 10 subsessile 95 subvestitum 135 surucuchense 131 taeniosum 168 taxifolium 1 1 3 tectum 154 tenuiculum var. "acrosora" 96 var. acrosorum 96 thurnii 156 thyssanolepis 137 x Pleopeltis macrocarpa 1 39 trichiatum 161 trichosorum 84 trifurcatum 83 triseriale 132 trundcola 109 tunguraguae 95 tweedianum 135 M/« 156 vacciniifolium 152 variabile 1 1 5 venturii 122 venulosum 12 vittaria 177 vulgare 125 vulpinum 167 xantholepis 135 xiphopteroides 101 yungense 1 1 3 loretense 156 Pteris fur cat a 145 Salpichlaena 68 hooker iana 70 lomarioidea 70 volubilis 70 TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 189 Schaffneria 2 furcata serrulata 83 Solanopteris 179 \ar.polypodioides\48 truncicola 109 bifrons 181 Tectaria bismarckii 180 fraxinea 51 brunei 180 Zygophlebia 12 tuberosa 180 dudleyi 92 Strut hiopteris Xiphopteris 12 mathewsii 9 1 maxonii 6 1 blepharidea 1 1 2 sectifrons 12 blepharolepis 108 wer#z/ 92 l>ues/f 1 1 2 Taenitis jamesonii 82 desvauxii 147 myosuroides 82 190 FIELDIANA: BOTANY - elected <>f Otlu HECKMAN BINDERY INC. FEB96 N. MANCHESTER. INDIANA 46962