ILLINOIS LIBRARY ,M URBANA-CHAMPAIGN BIOLOGY Botany NEW SERIES, NO. 34 PTERIDOPHYTA OF PERU Part VI 22. Marsileaceae-28. Isoetaceae Rolla M. Tryon Robert G. Stolze With the collaboration of: R. James Hickey Benjamin 011gaard December 30, 1994 Publication 1461 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY Information for Contributors to Fieldiana & 4, Wan Won t and "Literatu used after abbreviated mez ublished b; ind ild follow botanical p ;-s Information Service. Names of botanical authors sho s, Kew," 1984 edition 1 pp. nd floristics. Journal of Ecology, 51: 567- iltural patterns in visions, pp. 63-80. In V id Stars. Mouton Publishers, The Hague, Netherlan* ward, J. H., ed.. Handbook of Bulletin 143, Bureau of American Ethn<; Idiana: B< ions: II tatements in figure captions alone, sue: ic arrangement to be obtar 1 within tli ;o the made and u . THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER. FIELD1ANA Botany NEW SERIES, NO. 34 PTERIDOPHYTA OF PERU Part VI 22. Marsileaceae-28. Isoetaceae Rolla M. Tryon Robert G. Stolze Department of Biology Associate Curator University of South Florida Department of Botany Tampa, Florida 33620-5150 Field Museum of Natural History Roosevelt Road at Lake Shore Drive Chicago, Illinois 60605-2496 With the collaboration of: R. James Hickey Benjamin 011gaard Miami University, Oxford, Ohio Aarhus University, Risskov, Denmark Accepted March 30, 1994 MAD \ Q 1995 Published December 30, 1994 Publication 1461 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 1994 Field Museum of Natural History ISSN 00 15-0746 PRINTED IN THE UNITED STATES OF AMERICA Table of Contents List of Illustrations INTRODUCTION 1 ACKNOWLEDGMENTS 1 22. MARSILEACEAE 2 Marsilea 2 23. SALVINIACEAE 5 Salvinia 6 Azolla 8 24. PSILOTACEAE 11 Psilotum 11 25. EQUISETACEAE 12 Equisetum 12 26. LYCOPODIACEAE 16 Huperzia 19 Lycopodium 52 Lycopodiella 58 27. SELAGINELLACEAE 66 Selaginella 66 28. ISOETACEAE 88 Isoetes 89 ADDENDUM 97 1 . Species to Be Added to the Pterido- phyte Flora, Parts I-V 97 2. Consideration of Pteridophyte Diversity in Respect to Ecology and Geography ... 98 DEPARTMENTS OF PERU 103 COMPREHENSIVE INDEX TO NAMES . . 1 04 1. Marsilea: M. ancylopoda; M. vestita .... 3 2. Salvinia: S. minima; S. auriculata 7 3. Azolla: A. filiculoides\ A. microphylla .... 9 4. Psilotum: P. nudum . .13 14 18 53 5. Equisetum: E. giganteum; E. bogotense 6. Huperzia: H. eversa; H. binervia; H. linifolia var. tenuifolia; H. cuneifolia . 1. Lycopodium: L. clavatum; L. thyoides 8. Lycopodiella: L. caroliniana var. meridi- onale; L. cernua 59 9. Selaginella: S. peruviana; S. haema- todes; S. exaltata 67 10. Isoetes: I. andicola; I. lechleri; I. di- spora 90 List of Tables 1 . Data on the diversity of pteridophyte species in Peru 100 List of Maps 1 . The most species-rich departments in Peru 99 2. The main vegetational zones in Peru and their ferns . .101 111 Back cover: Selaginella haematodes PTERIDOPHYTA OF PERU Part VI 22. Marsileaceae-28. Isoetaceae Introduction This sixth and final part of the "Pteridophyta of Peru" contains the aquatic fern families Mar- sileaceae and Salviniaceae as well as the "fern al- lies" Psilotaceae, Equisetaceae, Lycopodiaceae, Selaginellaceae, and Isoetaceae. A brief section on the diversity and ecology are included, written by the senior author. The comprehensive index con- tains not only the names in this final part but those of the other five parts as well. 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, 1 989), and it is not necessary to repeat them here. Treatment of the Lycopodiaceae has been con- tributed by Benjamin 011gaard and that of Isoeta- ceae by R. James Hickey. Both are recognized spe- cialists in these two families. 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, 1976 et seq.). The acronyms for herbaria follow Index Herbariorum and are also provided below. Acknowledgments Benjamin 011gaard produced a recent Index to the names of Lycopodiaceae, as well as taxonomic treatments for the family in "The ferns and fern allies of Guatemala" and in the "Flora of Ecua- dor." R. James Hickey contributed Isoetaceae for "The ferns and fern allies of Guatemala" and is currently working on a monograph of the family. Consequently, both have a keen understanding of the problems in these pteridophyte families, and for their outstanding efforts in the production of this Flora the authors wish to express their deep appreciation. We would like to extend special thanks to Blanca Leon (USM) for her invaluable assistance in pre- paring loans and arranging for their packing and shipment from this important Peruvian herbari- um, as well as from the Universidad Nacional de Trujillo (HUT). Rolla M. Tryon appreciates the fa- cilities provided by the Department of Biology, University of South Florida, Tampa, and the aid of Alice F. Tryon in the preparation of the treat- ment of Selaginella. The illustrations were contributed by Field Mu- seum scientific illustrator Zorica Dabich, who cre- ated the original drawings and adapted the rest from those previously used in the Fieldiana: Bot- any publication "The ferns and fern allies of Gua- temala." Her art work, which now has appeared in all six parts of this Flora, is an invaluable com- plement to the descriptions. We are extremely grateful to her. Thanks also go to Bent Johnsen, free-lance artist of Copenhagen, Denmark, for his drawing ofHuperzia. We also appreciate the valu- able comments presented by reviewers of the manuscript. We are grateful to the officers of the following institutions for granting loans of their material or allowing us to examine specimens in their her- baria: Herbarium Jutlandicum, Aarhus Univer- sitet, Denmark (AAU); Field Museum of Natural History, Chicago (F); Harvard University, Cam- bridge, Massachusetts— most Gray Herbarium (GH), some Arnold Arboretum (A); Herbarium Truxillense, Universidad Nacional de Trujillo, Trujillo, Peru (HUT); Missouri Botanical Garden, St. Louis (MO); Miami University, Oxford, Ohio (MU); Museum National d'Histoire Naturelle, Par- is (P); University of California, Berkeley (uc); United States National Herbarium, Smithsonian Institution, Washington, D.C. (us); and Herbario FIELDIANA: BOTANY, N.S., NO. 34, DECEMBER 30, 1994, PP. 1-123 San Marcos, Universidad Nacional Mayor de San Marcos, Lima, Peru (USM). Materials, mostly types, have also been studied in the following herbaria (acronyms follow Index Herbariorum, ed. 8): AWH, B, BKL, BM, BONN, BR, C, CGE, CPUN, CR, DUKE, G, GB, GL, HB, HEID, K, KRA, L, LG, LIL, LINN, LP, MA, MICH, MOL, MSC, NO, NY, PR, Q, QCA, RB, S, SAPF, U, UPS, W, WIS, and Z. This project has been supported in part by grant #BSR-85-16358 from the National Science Foun- dation, Systematic Biology Program. The work would not have been possible without this assis- tance. However, any opinions and conclusions ex- pressed are those of the authors and do not nec- essarily reflect the views of the Foundation. Family 22. MARSILEACEAE Marsileaceae Mirb., Hist. nat. veg. (Lam. & Mirb.) 5: 126. 1802. TYPE: Marsilea L. Stem short- to long-creeping, slender, often branched, hardly indurated, bearing trichomes. Leaves ca. 1-40 cm long, with 4, 2, or no leaflets at the apex of the petiole, circinate in the bud. Sori borne within sporocarps, indusiate, with indehis- cent, short- to long-stalked, not annulate, mega- sporangia and microsporangia; heterosporous, spores without chlorophyll. The Marsileaceae are a family of three genera, all in tropical America. The genera (Marsilea, Reg- nellidiwn, and Pilularid) are clearly distinct and not very closely allied. There are numerous detailed studies on the Marsileaceae centering on the form and devel- opment of the elaborate reproductive structures and simplified leaves. These plants have also been useful in physiological and experimental work on factors influencing leaf form. However, some of the basic problems concerning the nature of the leaflets and the sporocarp are not resolved, and the phyletic relationships of the three genera are not wholly certain. Pilularia L. is not known from Peru. However, Pilularia americana A. Br. has been collected near- by in Bolivia (as P. mandonii) and it probably also grows in Peru. It is a small plant with no leaflets on the petiole; the filiform leaves (petioles) are 1- 1 0 cm long. A single subglobose sporocarp is borne at the base of a leaf. Unless sporocarps are present, the plant can easily be taken for a species of Cy- peraceae, which grows in wet habitats that are sim- ilar to those of Marsilea. I. Marsilea Marsilea L., Sp. pi. 1099. 1753; Gen. pi. ed. 5, 485. 1754. TYPE: Marsilea quadrifolia L. Figure 1. Plants palustral or aquatic. Stem usually long- creeping, commonly bearing long roots at the nodes of the stem or along the internodes. Leaves with the petiole terminated by 2 adjacent pairs of nar- rowly cuneate to broadly flabellate leaflets that are glabrous or pubescent. Veins more or less anas- tomosing, usually connected at the margin. Sori borne within 1 to several stalked, indurated spo- rocarps attached to the petiole or at its base, en- closed by a diaphanous indusium, with microspo- rangia and megasporangia. Megaspores somewhat ellipsoidal with an apical papillalike laesura, the surface papillate. Microspores spheroidal, trilete, the surface slightly rugulose. Marsilea is a nearly worldwide genus of perhaps 50 species, with about 1 2 in America. The tropical American species are poorly known for there are relatively few collections and these often lack spo- rocarps. The species of Marsilea are most diverse and common in regions that support vernal pools or shallow pools that dry out, at least along the borders. Mexico is one center of diversity, Aus- tralia and Africa are others. The following treatment is based directly on that of Johnson (1986). When necessary, specimens are indicated as cited by Johnson. The sporocarp is borne on a stalk that is con- nected to the petiole. If this stalk is attached by its end to the sporocarp, there is neither raphe nor inferior tooth. If the stalk is attached along its side (the raphe), there may be an inferior tooth (the end of the stalk). The superior tooth is an extension of the sporocarp near the inferior tooth (fig. Id). FIG. 1 . Marsilea ancylopoda: a, habit; b, leaflet; c, stem and roots, with sporocarp. Marsilea vestita: d, sporocarps. (a, b from Alston 6354, Venezuela, F, c from Alston 5874, Venezuela, F, d from Sandberg, United States (Idaho), F.) FIELDIANA: BOTANY TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. The sporocarps may be very long-lived (to 100 years; Johnson, 1986) and they presumably are dispersed by waterfowl. Species also may be dis- persed as a result of discarded aquaria. JOHNSON, D. M. 1986. Systematics of the New World species ofMarsilea (Marsileaceae). Syst. bot. monogr. 11: 1-87. References GUPTA, K. M. 1962. Marsilea, Bot. monogr. 2, pp. 1-113, Council Sci. India Res., New Delhi. Key to Species of Marsilea a. Roots borne only at the nodes of the stem (rarely a few to 1 cm distant from a node); raphe present b b. Superior tooth acute 3. M. vestita b. Superior tooth blunt or absent c c. Sporocarps borne below the ground or stem level; leaflets essentially glabrous 2. M. ancylopoda c. Sporocarps borne above the ground or stem level; leaflets pubescent 4. M. mollis a. Roots borne at the nodes of the stem and along the internodes d d. Sporocarps 1-4, borne on the basal 'A of the petiole 1. M. deflexa d. Sporocarps 5-20, borne on the basal % of the petiole M. crotophora (see Comments) 1. Marsilea deflexa A. Br., Monatsber. konigl. preuss. akad. wiss. Berlin. 1863: 421. 1864. TYPE: Piauhy, Brazil, Gardner 2760 (holo- type, G; isotypes, BM, G, K, P, all cited by John- son). Plants forming dense colonies. Roots at nodes and internodes. Sporocarps 1-4, on basal '/4 of the petiole, 3.5-6 mm long with conspicuous lateral ridges. In a pool, ca. 59 m; a single collection from San Martin. Mexico and Central America; Venezuela and Colombia, south to Brazil; Paraguay and Peru. San Martin: Morales, Tarapoto, Vie 6866 (G, K, P, all cited by Johnson). 2. Marsilea ancylopoda A. Br., Monatsber. ko- nigl. preuss. akad. wiss. Berlin. 1863: 434. 1864. TYPE: Guayaquil, Ecuador, Jameson 304 (holotype, G; isotypes, BM, G, K, all cited by Johnson). Originally as James 304, cor- rected later (A. Br., 1871, op. cit. 1870: 717). Figure la-c. Roots borne only at the nodes (rarely a few 2- 3 cm distant). Raphe present. Superior tooth ab- sent or slightly raised. Sporocarps 2.5-6 mm long, borne below the ground or stem level, 1 at the base of a petiole. Leaflets of non-floating leaves gla- brous or with a few trichomes. Northern Peru, Tumbes to La Libertad, near sea level. Mexico; Florida and Greater Antilles; Venezue- la, Colombia to Peru; Brazil; Argentina; Uruguay. Tumbes: Laguna de Salitoral Grande, Coronado 235 (GH, uc). Lambayeque: San Nicolas, entre Lambayeque et Chiclayo, Santos Quiroz 2401 (F, GH). Prov. Lam- bayeque, Yencala, Santos Quiroz 71 (HUT). Laguna Boro, Chiclayo, Leon 600 (USM). La Libertad: Prov. Trujillo, Huaman, Angulo & Lopez 348 (GH). 3. Marsilea vestita Hooker & Grev., Icon. fil. 2: t. 159. 1830. LECTOTYPE (designated by Johnson, Syst. bot. monogr. 11: 62. 1986): Columbia River, Scouler 338 (K; isolecto- types, GH, NY). Figure Id. FIELDIANA: BOTANY Marsilea uncinata A. Br., Amer. J. Sci. ser. 2, 3: 55. 1 847. TYPE: Arkansas, Englemann 33 (holotype, MO; isotypes, K, M, MO, all cited by Johnson). Marsilea mucronata A. Br., Amer. J. Sci. ser. 2, 3: 55. 1847. TYPE: Near Devil's Lake, North Dakota, Geyer 71 (holotype, MO; isotypes, K, NY, all cited by Johnson). Plants in diffuse or dense colonies. Roots only at the nodes (rarely a few to 10 mm distant). Raphe present. Superior tooth acute, often hooked at apex. Sporocarp 3.0-7.6 mm long with lateral ridges. Leaflets of non-floating leaves covered by over- lapping trichomes on both surfaces. All of the above pertains to ssp. vestita; ssp. tenuifolia (A. Br.) Johnson is endemic to central Texas. It differs from ssp. vestita in having very narrow leaflets with sparse trichomes or none abaxially. Known in Peru only from the Department of Lambayeque. Southern British Columbia east to western Min- nesota; south to Louisiana, Florida; Mexico; Peru. The Peru and Florida localities may be introduc- tions. Lambayeque: Chiclayo, Leon (MICH, det. Johnson). Prov. Lambayeque, Salazar HI (USM). 4. Marsilea mollis Rob. & Fern., Proc. Amer. Acad. 30: 123. 1895. TYPE: San Diego, Chi- huahua, Mexico, Hartman 604 (holotype, GH; isotypes, F, GH, MSC, NY, uc, all cited by John- son). Roots borne only at the nodes (occasionally to 10 mm distant). Raphe present. Superior tooth absent or to 0.2 mm long. Sporocarps at or near petiole base, 1.7-6.7 mm long, on recurved pe- duncles, borne above the ground level. Leaflets of non-floating leaves adaxially glabrous to sparingly pilose, abaxially densely pilose. 2000-3900 m, Cajamarca south to Puno. Arizona and Texas south to northern South America, south to Argentina, Chile and Brazil. The following sterile specimens are considered to represent this species. All have been seen by the present authors except the Hill collections from Puno. All have been placed here by Johnson except the Nunez et al. collection from Cuzco. Cajamarca: Cajamarca, Muller & Gutte 27322 (USM). Junin: Huancayo, Kunkel 422 (GH). Huancavelica: Prov. Angaraes, 4 km W of Huanta, Stork & Morton 10808 (F, uc). Cuzco: Prov. Espinar, Yauri, Nunez et al. 7810 (F, MO), Vargas 13524 (GH). Puno: Towards Juliaca, Hill 562 (K), 563 (K). Comments Marsilea crotophora D. M. Johnson, Syst. bot. monogr. 11: 46. 1986. TYPE: Mato Grosso, Brazil, Hatschbach & Scherer 30470 (holo- type, us; isotypes, c, LP, M, MICH, NY, uc, all cited by Johnson). This species may be found in the upper Amazon or its tributaries. It may be distinguished from Peruvian species by the roots being borne on the internodes as well as the nodes, the lack of a raphe and superior tooth, and the 5-20 sporocarps borne on the basal % of the petiole, the lowest one borne well above the petiole base. Family 23. SALVINIACEAE Salviniaceae Reichenb., Bot. damen kunst. freunde pflanzenw. 255. 1828. TYPE: Salvinia Se- guier. Azollaceae Wettst. Handb. syst. bot. 2: 77. 1903. TYPE: Azolla Lam. Plants floating in water or stranded in wet mud. Stem elongate, small, and slender, often branched, not indurated, usually bearing trichomes. Floating leaves ca. 0.5-2.5 cm long, not circinate in the bud. Sori borne on a lobe of a leaf or on a branched leaf, indusiate, with the sporangia stalked, not an- nulate, the sori either megasporangiate or micro- sporangiate; heterosporous, spores without chlo- rophyll. The structure bearing the sporangia is consid- ered to be a sorus, surrounded by an indusium, and accordingly the special term sporocarp is not used. The living Salviniaceae are represented by two genera, Salvinia and Azolla, floating aquatics of wide distribution, especially in the tropics. Al- though Azolla is sometimes included in a separate family, it has many basic similarities with Sal- vinia. TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. Key to Genera of Salviniaceae a. Leaves in whorls of 3, 2 floating, ca. 5-25 mm long, with anastomosing veins and not lobed, 1 submerged, highly branched I. Salvinia a. Leaves alternate, ca. 0.5-1.5 mm long, without veins, bilobed II. Azolla I. Salvinia Salvinia Seguier, Fl. veron. 3: 52. 1754. TYPE: Salvinia natans (L.) All. (Marsilea natans L.) Figure 2. Plants floating aquatics. Stem short or elongate, slender, bearing trichomes and no roots. Leaves borne in whorls of 3, dimorphic, 2 of them green, floating, entire, oblong to suborbicular, ca. 0.5-2.5 cm long, often pubescent, usually papillate on the upper surface, one of them submerged, pendent in the water, highly branched, bearing many tri- chomes. Veins anastomosing. Sporangia borne in stalked sori on the submerged leaf, enclosed by the indusium, with either megasporangia or mi- crosporangia. Spores trilete, enclosed within a vac- uolate tapetal formation, the megaspore surface perforate, the microspore surface somewhat ru- gulose. Salvinia is a widely distributed genus of about ten species, with seven of them in America. Shaparenko (1956) recognized four sections, and these were adopted by de la Sota (1962); they are not recognized here because the genus is a small one. The descriptions of the species are freely adapt- ed from Stolze(1983). References SHAPARENKO, K. K. 1956. History of the Sal- vinias (in Russian). Act. Inst. Bot. Komarov. Ser. VIII Paleobotania 2: 1-44. SOTA, E. R. DE LA. 1962-1964. Contribution al conocimenti de las "Salviniaceae" neotropi- cales. I-V. Darwiniana 12: 465-520, 612-623; 13: 529-536. STOLZE, R. G. 1983. Ferns and fern allies of Gua- temala, III. Fieldiana: Bot., n.s. 12: 1 1-13. Key to Species of Salvinia a. Trichomes at the apex of the papillae, on the upper surface of the floating leaves, separate (not joined) at their tip 1 . S. minima a. Trichomes at the apex of the papillae, on the upper surface of the floating leaves, joined at their tip . 2. S. auriculata 1. Salvinia minima Baker, J. Bot. 24: 98. 1886. TYPE: Santa Catarina, Brazil, Muller 479 (holotype, K; isotype, BM). Figure 2a-c. Stele of the stem more or less circular in cross- section. Floating leaves oblong-elliptic, the apex obtuse (rarely retuse), the larger ones 0.6-1.3 cm long, their upper surface papillate, each papilla with apical trichomes, these separate (not joined) at the apex. 130-160 m, Loreto and Madre de Dios. Southeastern United States; Mexico and Central America; South America, south to Argentina and Uruguay. This species has often been called S. rotundifolia but that name = S. auriculata. FIG. 2. Salvinia minima: a, habit; b, portion of leaf showing venation; c, papillae with free, spreading trichomes. Salvinia auriculata: d, habit; e, papillae with joined trichomes. (Adapted from Stolze, Ferns and fern allies of Guatemala, 1983.) FIELDIANA: BOTANY TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. Loreto: Iquitos, McDaniel 11776 (MO). Near Iquitos, Ellenberg 8301 (F). Rio Itaya, Iquitos, Revilla 467 (F, MO, us, USM). Lupuna Cocha, Iquitos, Tryon & Tryon 5188 (F, GH, us, USM). Madre de Dios: Prov. Tambopata, Tambopata Nature Reserve, 30 air km SSW of Puerto Maldonado, Barbour 5373 (MO). Prov. Manu, Parque Nacional Manu, Foster 12206 (F). 2. Salvinia auriculata Aublet, Hist. pi. Guiane Franc. 2: 969. 1775. TYPE: French Guiana, Terr. Caux, Aublet (p?). Figure 2d-e. Salvinia rotundifolia Willd., Sp. pi. ed. 4, 5: 5 1 7. 1 8 1 0. TYPE: Brazil, Hoffmannsegg(see Morton, Contr. U.S. Natl. Herb. 38: 75. 1967). Stele of the stem U-shaped in cross-section. Floating leaves oblong-elliptic and nearly circular, obtuse or retuse at the apex, larger floating leaves 1 .5-2.5 cm long, their upper surface papillate, each papilla with 4 apical trichomes, these joined at the apex. At ca. 100 m, Loreto and Madre de Dios. Cuba; Mexico south to Argentina. Loreto: Rio Amazonas, SE of Iquitos, Croat 19296 (MO). Prov. Maynas, mouth of Rio Nanay, Gentry el al. 21715 (MO, uc). Yanamono, Gentry et al. 42574 (F, MO). Madre de Dios: Prov. Tambopata, Tambopata Nature Reserve, 30 air km SSW of Puerto Maldonado, Barbour 4888 (MO). II. Azolla Lam., Encycl. 1: 343. 1783. TYPE: Azolla filiculoides Lam. Figure 3. Floating aquatics. Stem short, elongate, slender, sometimes bearing trichomes, and usually short roots. Leaves ca. 0.5-1.5 mm long, bilobed, the upper lobe usually minutely to strongly papillate, without veins. Sporangia borne in short-stalked sori enclosed by an indusium, with either 1 mega- sporangium or several microsporangia. Spores tri- lete, the megaspores irregularly marked, with a perforate surface, the microspores smooth, em- bedded in massulae. The megaspore has accessory structures usually referred to as a columella bearing floats. These are partially enclosed by a band of sporoderm, the collar, which is between the megaspore proper and the floats. In the living species the floats are either three or nine in number. The microsporangia con- tain a few to several massulae, within which the microspores are embedded. Projecting structures called glochidia are often on the outer surface of the massulae. Azolla is a widely distributed genus of six spe- cies, with four of them (sect. Azolla) in America and in Peru. It is notable in having the most com- plex reproductive structures of any plant and in having a colony of Anabaena azollae within the lobes of each leaf. This blue-green alga has the ability to fix atmospheric nitrogen, and accord- ingly Azolla is the center of wide interest as a green- manure (Lumpkin & Plucknett, 1980), especially for the rice paddies of Southeast Asia. Azolla is a very distinctive genus, but the species are poorly known due to the fact that sterile ma- terial is difficult to identify. Fertile material is un- common. Accordingly, the species in Peru and their characters are only provisionally known. In spite of evidence to the contrary, the treatment of Sven- son (1944) is largely followed. The key is adapted from his key to species of the Americas and the megaspore characters are taken from Perkins et al. (1985). Species names may be assigned to one of the four species adopted, although this does not mean that the material is accurately determined. The following Peru collections have not been iden- tified to species: Cajamarca: Dillon 2919. La Li- bertad: Killip & Smith 21511; D. Smith et al. 2225. Cuzco: Herrera 2616, 2618; Cook & Gilbert 237, 241, 1078; Cook 1963; Maldonado 17. Puno: Bar- clay 9273. Species of Azolla, as they are known, are not restricted to particular environments as differ- ences in range and habitat statements might imply. Accordingly, the range, habitat, and altitude are given for the genus as a whole in Peru, rather than for each species. In wet mud of riverbanks or stream banks, where stranded by higher water, usually floating in stand- ing water of lakeshores, ponds, or small streams, also in ditches, in stagnant or fast-flowing water, less often in swamps or wet parts of cultivated land, near sea level to 4100 m, Cajamarca, Ama- zonas, and Loreto, south to Puno and Moquegua. FIG. 3. Azolla filiculoides: a, habit; b, megasporangium; c, sporocarp indusia with microsporangia; d, massula with nonseptate trichomes. Azolla microphylla: e, megasporangium; f, massula with septate trichomes. (Adapted from Stolze, Ferns and fern allies of Guatemala, 1983.) FIELDIANA: BOTANY TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. References LUMPKIN, T. A., AND D. L. PLUCKNETT. 1980. Azolla, botany, physiology, and use as a green- manure. Econ. Bot. 34: 1 1 1-153. PERKINS, S. K., G. A. PETERS, T. A. LUMPKIN, AND H. E. CALVERT. 1985. Scanning electron mi- croscopy of perine architecture as a taxonomic tool in the genus Azolla Lamarck. Scanning Electron Microscopy 4: 1719-1734. SVENSON, H. K. 1944. The New World species of Azolla. Amer. Fern J. 34: 69-84. Key to Species of Azolla a. Plants dichotomously branched, 0.5-1.5 cm long; leaves less than 1 mm long; megaspore collar with or without filaments b b. Plants small, 0.5-1.0 cm in diameter; leaves small (0.5 mm long), nearly orbicular and nearly smooth; glochidia not septate; megaspore collar with dense filaments, the megaspore surface more or less smooth, obscured by dense filaments 1 . A. caroliniana b. Plants often larger, 1.0-1.5 cm in diameter; leaves with the upper lobe 0.7 mm long, the lower lobe much larger, more or less obovate; glochidia septate; megaspore collar lacking filaments, megaspore surface with large and small pits, but few filaments 2. A. mexicana a. Plants pinnately branched, 1-6 cm long; leaves 1 mm long; megaspore collar lacking filaments . .c c. Plants often 2-6 cm long; leaves oblong to ovate; glochidia not septate (or with 1-2 septa distally); megaspore surface with raised hexagonal markings 3. A. filiculoides c. Plants 1-2 cm long; leaves nearly orbicular; glochidia septate, some septa located below the middle; megaspore surface rugulate-verrucate 4. A. microphylla 1 . Azolla caroliniana in the sense of Svenson and other authors, not Willd., as usually thought. Plants dichotomously branched, 0.5-1.0 cm in diameter. Leaves divaricate (not closely imbri- cate), nearly orbicular, nearly smooth, small, 0.5 mm long. Glochidia not septate. Megaspore collar with dense filaments on its surface (magnification x200 or more), rugulate-verrucate beneath fila- ments. Eastern North America; Mexico and the Carib- bean; introduced in Europe and South America. The name is used in the sense of Svenson and most authors (not Willd.) until a definitive treat- ment is published. Jermy (Somerfeltia 6: ix-x. 1987) has pointed out that according to the Ph.D. thesis of D. Dun- ham (Portsmouth Polytechnic) the type of Azolla caroliniana is A. filiculoides. This leaves Azolla of eastern North America presumably without a name. Amazonas: Rio Santiago, S of La Poza, Berlin 3711 (MO, uc). La Libertad: Prov. Bolivar, Laguna de las Ichus, Lopez & Sagdstegui 3238 (GH, MO). San Martin: Prov. Mariscal Caceres, 2-4 km W of Tocache Nuevo, Plow- man 11437 (F). Loreto: Prov. Maynas, Rio Itaya, Revilla 632 (F, MO, uc.). Prov. Maynas, Yanuyaca, Gentry et al. 32119 (GH). Ayacucho: 44 km N of Tambo, Luteyn & Lebron-Luteyn 6369 (uc, us). Cuzco: Prov. Cuzco, Say- lla, Fernandez et al. 455 (USM). Puno: Santa Rosa, Staf- ford 495 (F). Ichu, Soukup 354 (F). 2. Azolla mexicana Presl, Abh. Bohm. Ges. Wiss. Ser. V, 3: 150. 1845. TYPE: Mexico, Schiede & Deppe (holotype, PR). Plants dichotomously branched, often 1.0-1.5 cm in diameter. Leaves divaricate (not closely im- bricate) with the upper lobe ca. 0.7 mm long, the lower one much larger. Glochidia septate. Mega- spore collar glabrous; megaspore surface (magni- fication x 200 or more) coarsely rugose, also with pits, and with many dense filaments. North America; Mexico; Central America; northern half of South America. Tumbes: Casa Fernandez, Haught 183 (us). Loreto: Prov. Maynas, Iquitos, Tryon & Tryon 5213 (GH, us). Ancash: Lago Santa Cruz Chico, Huascaran National Park, D. Smith et al. 9261 (F, HUT, MO). Junin: Huancayo, Kunkel 421-lh (GH). Acopalca, Kunkel 421 (GH). Cerca Carhuamayo, Ferreyra 5242 (USM). Cuzco: Pampa de Anta, Iltis & Ugent 785 (GH, uc, us, USM). 3.5 km NW of Saylla, Iltis et al. 1208 (GH, us). Prov. Paucartambo, Paucartambo, Plowman & Davis 4920 (F). Madre de Dios: Prov. Manu, Parque Nacional Manu, Foster 9872 10 FIELDIANA: BOTANY (MO). Puno: Pucara, Hunnewell 15874 (GH). Lake Titi- caca, Shepard 1511 (GH, us). Moquegua: Ilo, Stafford 926 (F). 3. Azolla filiculoides Lam., Encycl. 1: 343. 1783. TYPE: "Magellan Region," Herb. Lamarck (p). Figure 3a-d. Plants pinnately branched, often 2-6 cm long. Leaves closely appressed, imbricate, oblong to ovate, ca. 1 mm long. Glochidia not septate (rarely 1-2 "septa" toward the apex). Megaspore collar more or less glabrous; megaspore surface (mag- nification x 200 or more) prominently rugose with hexagonal markings, with few filaments. Southern South America north to western North America and to Alaska; introduced in Europe, Asia, and Australia. Azolla filiculoides var. rubra (R. Br.) Strasb., treated as a species A. rubra R. Br. by Perkins et al. (1985), is not considered to occur in Peru. Cajamarca: Prov. Contumaza, Rio San Benito, San Benito, Sagdstegui et al. 12509 (HUT, MO). Amazonas: Prov. Bongara, Laguna Pomacocha, Wurdack888 (F, GH, uc, us). La Libertad: Prov. Trujillo, Tschudi, Shimo- kowa 7294 (HUT). Playa de Chan Chan, Angulo 18 (us). Loreto: Paraiso, above Iquitos, Rio Amazonas, Fosberg 29076 (us). Huanuco: Chasqui, Macbride 3306 (F, us); 3307 (F, us). Lima: Prov. Lima, Laguna de Villa, Corona- do 5 (GH, MO, uc, us); Tryon & Tryon 5458 (GH); Cerrate 2766 (USM). Huancavelica: Prov. Tayacaja, Pampas, Stork & Morton 10243 (F, uc, us). Cuzco: Cook & Gilbert 197 (us). Prov. Paucartambo, Huilabamba, Balls B7632 (F, uc, us). Arequipa: Prov. Arequipa, Rio Yarabamba, Ponce 32 (USM). Family 24. PSILOTACEAE Psilotaceae Kanitz, Novenyrends. Attek. 43. 1887. TYPE: Psilotum Sw. Stem flaccid to somewhat indurated, lacking in- dument and roots. Leaves (or pinnae) small to minute, borne alternately, with a single vein or none. Two or 3 sporangia joined in a sessile, thick- walled synangium. Homosporous. Spores without chlorophyll. The Psilotaceae are a family of two genera, Tme- sipteris Bernh. of western Malesia, Australia, and the Pacific, and Psilotum, which is pantropical in distribution. After an extensive series of studies, D. W. Bier- horst concluded that the Psilotaceae were primi- tive elements in the Filicopsida. This interpreta- tion of the position of the family provoked further interest in its phylogeny and resulted in a sym- posium on the taxonomic and morphological re- lationships of the Psilotaceae (White et al., 1977). The Psilotaceae have a number of characters that occur in the Filicopsida, such as the details of early ontogeny of the leaf, the subterranean, cylindrical gametophytes, multiflagellate sperm, and several aspects of the rhizoids and gametangia. These and the evidence from the structure and wall formation of Psilotum spores are indicative of re- lationships with the ferns. However, Cooper- Driv- er (1977) has shown that the chemical evidence does not support this relationship. References 4. Azolla microphylla Kaulf, Enum. fil. 273. 1 824. TYPE: "California," Chamisso (LE?). Figure 3e-f. Plants pinnately branched, small, 1-2 cm long, many leaves orbicular, 1 mm long, most leaves closely imbricate. Glochidia septate. Megaspore collar glabrous; megaspore surface slightly fila- mentous, slightly pitted, rugulate (high magnifi- cation) to verrucate. Western and northern South America; southern North America; West Indies. Loreto: Yurimaguas, Killip & Smith 27707 (us); Prov. Requena, Rio Ucayali, Encarnacion E-1101 (us). Aya- cucho: Puquio, Gentry et al. 23296 (F, MO, us). Cuzco: Prov. Anta, Pampas de Anta, Tryon & Tryon 5364 (GH, us, USM). Yauri, Nunez et al. 7813 (F, MO). COOPER-DRIVER, G. 1977. Chemical evidence for separating Psilotaceae from the Filicales. Science 198: 1260-1261. WHITE, R. A., ET AL. 1977. Taxonomic and mor- phological relationships of the Psilotaceae. Brit- tonia 29: 1-68. I. Psilotum Psilotum Sw., J. Bot. (Schrader) 1800 (2): 8, 109. 1 80 1 , nom. nov. for Hoffmannia Willd. (not Sw.) and with the same type. Hoffmannia aphylla Willd. = Psilotum nudum (L.) Beauv. Figure 4. Plants terrestrial, rupestral or usually epiphytic. Stem compactly branched in the substrate, the ae- TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 11 rial stems green, erect to pendent, with several dichotomous divisions, ca. 20 cm to 1 m long, without roots. Leaves minute, alternate, leafy stem glabrous. Synangia large, borne singly in the axil of forked fertile leaves. Spores elongate-ellipsoid- al, monolete, the laesurae '/2 to % the spore length, the surface coarsely rugose to shallowly and com- pactly verrucate. Psilotum is a highly distinctive genus of wide distribution in the tropics and with some exten- sions to subtemperate regions. Notable characters of the genus are the dichotomously branched stem, the bifurcate sporophylls, and the large synan- gium. Psilotum has a simple morphology, proba- bly through reduction from more complex ances- tors. 1. Psilotum nudum (L.) Beauv., Prodr. Fam. Ae- theog. 112. 1805. Figure 4. Lycopodium nudum L., Sp. pi. 1100. 1753. TYPE: LINN 1257.1. Psilotum triquetrum Sw., J. Bot. (Schrader) 1800 (2): 109. 1801, nom. super/I, for Lycopodium nudum L. and with the same type. Plants pendent or usually erect. Branches 3-an- gled, to 1 .2 mm wide, leaves distant, 2-3 mm long, triangular-subulate. Epiphytic, especially on palms and tree ferns, 200-600 m, San Martin south to Madre de Dios. American and Old World tropics and subtrop- ics. P. complanatum Sw. may be a distinct species. It has flattened branches 2 mm or more wide and is especially distinctive in the Pacific islands. San Martin: Dist. Chazuta, Rimachi 7945 (MO). Dist. Tocache Nuevo, Schunke 4084 (F, us). Gramalote ad Saposoa, Woytkowski 7300 (GH, MO, us). Loreto: Lower Rio Huallaga, Killip & Smith 29244 (Coll. W. J. Dennis). Padre Isla, Iquitos, Encarnacion 26314 (MO, us). Prov. Requena, Rio Ucayali, Ayala et al. 3753 (MO). Huanuco: Tingo Maria, Moran 3687 (USM). Madre de Dios: Prov. Tambopata, Cuzco Amazonico Lodge, 15 km NE of Puerto Maldonado, Nunez 12214 (MO); Gentry et al. 68958 (MO). Prov. Manu, Parque Nacional Manu, Foster 11336(F). Family 25. EQUISETACEAE Equisetaceae A. P. DC., Fl. Franc. (Lam. & DC.) ed. 3, 2: 580. 1805. Type: Equisetum L. Stem jointed, more or less indurated, the aerial usually green, the subterranean with numerous wiry roots. Leaves each with a single vein, borne in whorls, joined in a sheath. Sporangia thin-walled, borne on stalked, peltate sporangiophores that form a strobilus. Homosporous. Spores with chloro- phyll and bearing elaters. I. Equisetum Equisetum L., Sp. pi. 1061. 1753; Gen. pi. ed. 5, 484. 1754. TYPE: Equisetum fluviatile L. Fig- ure 5. Terrestrial, palustral or aquatic. Stem subter- ranean, short- to long-creeping, freely branched, bearing erect, aerial stems that are jointed, lon- gitudinally ridged, and usually hollow, ca. 10 cm to 8 m long, often with whorls of branches. Leaves small, in whorls at nodes, the lower portion lat- erally fused in sheaths, the upper portion in more or less prolonged teeth. Sporangia large, several borne on each stalked, peltate, apically flattened sporangiophore, in a condensed terminal strobi- lus. Spores spheroidal with circular aperture, and 4 paddle-shaped elaters, the surface with small granulate and large spherical deposits. The Equisetum strobilus is composed of spo- rangiophores that are sometimes called sporo- phylls. A detailed study of these structures by Page (1972) shows they are partly modified from a leaf and partly from a cauline appendage; thus the term sporangiophore is more appropriate. The small leaves that surround each node are laterally fused below and form coarse teeth above. References HAUKE, R. L. 1961-1962. A resume of the tax- onomic reorganization of Equisetum subgenus FIG. 4. Psilotum nudum: a, habit; b, portion of fertile branch with sporangium. (Adapted from Stolze, Ferns and fern allies of Guatemala, 1983.) 12 FIELDIANA: BOTANY TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 13 5cm 14 FIELDIANA: BOTANY Hippochaete, I-IV. Amer. Fem J. 51: 131-137; Equisetum subgenus Equisetum. Nova Hedwig- 52: 29-35, 57-63, 122-130. ia 30: 385^55. HAUKE, R. L. 1963. A taxonomic monograph of PAGE, C. N. 1972. An interpretation of the mor- the genus Equisetum subgenus Hippochaete. phology and evolution of the cone and shoot of Beih. Nova Hedwigia 8: 1-123. Equisetum. J. Linn. Soc. Bot. 65: 359-397. HAUKE, R. L. 1973. A taxonomic monograph of Key to Species of Equisetum a. Main aerial stem large, usually 10 mm or more wide (pressed), rarely less (to 5 mm), usually 2-5 m tall, with a hollow center; strobilus sessile or nearly so 1 . E. giganteum a. Main aerial stem small, to 2 mm wide (pressed), usually 0.5m tall or less, with a solid center; strobilus stalked 2. E. bogotense 1 . Equisetum giganteum L., Syst. nat. ed. 10: 1318. 1759. TYPE: Plumier, PI. Amer. (ed. Bur- man) t. 125, f. 2 (1757 as to the plate). Figure 5a-b. Equisetum myriochaetum Schlect. & Cham., Linnaea 5: 623. 1 830. TYPE: Misantla, Veracruz, Mexico, Schiede and Deppe 833 (holotype, B?). Equisetum schaffneri Milde, Verb. Zool.-Bot. Ges. Wien 1 1: 345. 1861. TYPE: Orizaba, Mexico, W. Schaffner 315 (holotype, B?). Stem usually 2-4 m tall, often bearing an apical strobilus, usually 1 5-25 mm wide (10 mm or more, rarely 5 mm or less), with a central hollow. En- dodermis surrounding each of the vascular bun- dles. Sheaths of the main stem usually light brown, with often deciduous, rather indurated teeth. Branches in regular whorls, with 8-10 ridges. Strobilus sessile or nearly so. In wet, usually open areas, in sloughs, on the banks of streams, rivers, and irrigation ditches, in boggy areas, and in streambeds, 1900-3500 m, Piura, Cajamarca and Amazonas, south to Are- quipa and Puno. Greater Antilles; Mexico and Central America; Venezuela and Colombia, south to Chile, Argen- tina and Brazil. Equisetum myriochaetum may be a species, as Hauke and others have it, or it may be an extreme variation of E. giganteum, as treated here. Most likely it should be recognized as a subspecies, as is E. ramosissimum ssp. debile, but we do not wish to make a new combination at this time. Equisetum giganteum has the turbercles on the ridges of the stem square or flattened in profile while the turbercles of E. myriochaetum appear rather similar to the teeth of a saw. The former species has two to several lines of stomates on each side of a groove of the stem, while the latter species has a single line of stomates. Characters of the strobilus (apiculate or not) and of the sheath color (different from the stem or not) are not species- specific. Equisetum schaffneri has a variable mixture of the characters of E. giganteum and of E. schaffneri. It has a wider distribution than a sterile hybrid should have, and it occurs in regions in which only one of the putative parents is known (Venezuela and in Peru, for example). Accordingly, it may be a natural intermediate of the extremes of E. gi- ganteum. The abortive spores present in some col- lections (not all have strobili) are also present in some other species: in E. bogotense and E. var- iegatum (Hauke, 1963, 1978) where hybridization is not concerned. Hauke (Ph.D. dissertation, University of Mich- igan) cites the following collections of E. myri- ochaetum from Peru: Ayacucho, Killip & Smith 22757 (us); Cuzco, Cook & Gilbert 1372 (us). Also the following collections of E. x schaffneri from Peru: Lima, Rose 18762 (NY, us); Ucayali?, Herre- ra 1392 (us); Cuzco: Cook & Gilbert 1091 (us). FIG. 5. Equisetum giganteum: a, habit; b, apex of branch with strobilus. Equisetum bogotense: c, habit; d, apex of branch with strobilus. (a, b from Lowell 535, Ecuador, F; c, d from Cuatrecasas 1 1807, Colombia, F.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 15 Piura: Hills of Chiarnique, Barbour 2160 (MO). Lam- bayeque: Olmos to Jaen, Correll & Smith P792 (GH). Cajamarca: Prov. Cutervo, Llatun, Mostacero et al. 1740 (HUT). Prov. Celendin, alrededores de Celendin, Mos- tacero et al. 835 (MO, uc). Amazonas: Prov. Bagua, 3.9 km NE of Chiriaco, Barbour 4360 (F, MO). Utcabamba, Pennell 15586 (GH, us). La Libertad: Prov. Trujillo, valle de Rio Chicama, Miiller & Krebs 12137 (USM). San Mar- tin: Tarapoto, Martin & Plowman 1843 (F, GH). Prov. Mariscal Caceres, 45 km NE of Tingo Maria, Tryon & Tryon 5270 (GH). Ancash: Prov. Santa, alrededores de Jimbe, Mostacero et al. 1096 (F). Huanuco: Prov. Huanu- co 6 km S of Huanuco, Stork & Morton 9376 (F, GH, MO, uc). Lima: Villa Lagoon (Prov. Lima, Chorrillos), Coro- nado 6 (GH, uc, us). Pasco: Oxapampa, Soukup 2677 (us). Junin: 3 km N of La Merced, Tryon & Tryon 5442 (GH, uc). Ucayali: Prov. Coronel Portillo, Arboretum von Humboldt, Diaz et al. 687 (F, MO, us). Huancavelica: Prov. Tayacaja, Huari, Sounders 1148 (F, GH, us). Aya- cucho: Ayna, Killip & Smith 22757 (F). Apurimac: Prov. Abancay, Trancapata, Vargas 9617 (F, uc, us). Cuzco: Aguas Calientes, Solomon 3180 (F, MO); 39 km E of Urohuasi on Abancay-Cuzco road, Gentry et al. 23399 (MO, uc). Madre de Dios: Prov. Vlanu. Parque Nacional del Manu, Foster 9796 (MO); 9856 (MO). Arequipa: Yura, Solomon 2912 (F, MO, USM). Puno: Puno, Angulo 1760 (HUT). 2. Fquisetum bogotense HBK., Nov. gen. sp. pi. 1: 42. 1815. TYPE: "Prope Santa Fe de Bo- gota et Alto del Roble, Columbia," Humboldt & Bonpland (holotype, P?). Figure 5c-d. Steins usually 20-50 cm tall, commonly bearing an apical strobilus, 1-2 mm wide, solid in the center. Endodermis single, surrounding the ring of vascular bundles. Sheaths with short, brown, pa- pery teeth. Branches irregular, with 4 ridges. Strobilus stalked. In soil or among rocks, in wet open places, in- cluding banks of creeks, rivers, and irrigation ditches in moist to wet seeps and clay road banks, 100-4200 m, most often above 2000 m, Piura south to Puno and Tacna. Costa Rica and Panama; Galapagos Islands; the Andes of Venezuela and Colombia, south to Bo- livia; to southern Argentina and Chile. Piura: Prov. Huancabamba, Cerro Blanco, Stork 114 15 (GH, uc, us). Cajamarca: Prov. Contumaza, Guzmango to Santiago, Sagdstegui & Lopez 10584 (F, MO). San Pa- blo, Hutchison & Wright 5060 (F, MO, uc, us). Amazonas: Hills WNW of Pomacocha, Wurdack 9445 (GH, uc, us, USM). La Libertad: Above Yamobamba, Conrad 2728 (MO). San Martin: NW corner of Rio Abiseo National Park, Young & Leon 4534 (HUT, USM). Ancash: Prov. Recuay, Lopez et al. 7596 (MO, us). Huanuco: Prov. Huanuco, Cerro Carpish, Luteyn & Lebron-Luteyn 5483 (F, uc, us). Lima: Prov. Huarochiri, Sullivan et al. 1103 (F, MO); Prov. Canta, Dist. Huaros, Pennell 14715 (F, GH, us). Pasco: Prov. Oxapampa, Los Chacos, near Oxa- pampa, Smith & Pretel 1514(f, MO, USM). Junin: Above Conception, Correll & Smith P736 (GH). Huancavelica: Prov. Huancavelica, Yauli, Stork & Morton 10863 (F, uc). Ayacucho: Santiago, about 67 km from Nasca on road to Puquio, Correll & Smith PI 51 (GH). Cuzco: Prov. Urubamba, Pomatales, Nunez 7327 (F, MO). Arequipa: Chacra Pachacutac, Balls B5868 (F, uc, us). Puno: San Gabon (San Gaban) to Ollachea, Dillon et al. 1239 (MO). Tacna: Prov. Tarata, Chuvire, Metcalf 30407 (MO, uc, us). Family 26. LYCOPODIACEAE Contributed by Benjamin 011gaard Lycopodiaceae Mirbel, in Lam. & Mirbel, Hist, nat. veg. 4: 293. 1802. ("Lycopodia"). TYPE: Lycopodium L. Plants terrestrial or epiphytic, erect to pendu- lous herbs or climbers. Stems dichotomously branching, sometimes also with lateral branching, protostelic, with the xylem arranged radially or in parallel bands, or forming an incomplete cylinder (Phylloglossuni). Leaves simple, with 1 simple vein, arranged in low alternating spirals or irregular al- ternating whorls, or seemingly decussate, homo- phyllous or heterophyllous, isophyllous or aniso- phyllous. Sporophylls like the foliage leaves, or modified, in some groups specialized and aggre- gated into distinct strobili. Sporangia solitary, ax- illary or on the upper side of the sporophyll base, homosporous, unilocular, reniform to subglobu- lar, short-stalked, dehiscing by a transverse slit, which divides each sporangium into 2 nearly equal or strongly unequal valves. Spores without chlo- rophyll, tetrahedral, with a trilete scar. Gameto- phytes monoecious, tuberous, subterranean, and mycorrhizal and without chlorophyll or surface- living and green. The family occurs in almost all humid regions of the world, both warm and cold. It is here treated as consisting of four genera including Phylloglos- sum (Australia, New Zealand). Up to 14 genera have been recognized by other authors (Holub, 1975, 1983, 1985, 1 99 1 ; Wagner & Beitel, 1992), the genera of these authors generally correspond- ing to the infrageneric taxa of 011gaard ( 1 987, 1 989, 1992); for a review of the classification, an index to the family, and an overview of the Neotropical species, see these publications. Most of the Pe- ruvian species are illustrated in 011gaard (1988). 16 FIELDIANA: BOTANY The total number of species in the family probably exceeds 350, and approximately 185 of these are tropical American. The Andes have a particularly high diversity, and many species are conspicuous elements in the montane and alpine vegetation of these mountains. This treatment includes 62 species for Peru, but the real number is likely to be higher. Although largely all available material has been consulted for this treatment, it is evident that much collect- ing needs to be done before a complete presen- tation can be made. Many species are represented by a single or few collections, and intensive efforts in limited areas such as those of Blanca Leon and Kenneth Young in the Rio Abiseo National Park, of Abundio Sagastegui in the north of Peru, and of the late David N. Smith in several areas have turned up several species not formerly known from Peru and widely disjunct from other known oc- currences, as well as species new to science. References HOLUB, J. 1975. Diphasiastrum, a new genus in Lycopodiaceae. Preslia 47: 97-1 10. HOLUB, J. 1983. Validation of generic names in Lycopodiaceae: With a description of a new ge- nus Pseudolycopodiella. Folia Geobot. Phyto- tax. 18: 439-442. HOLUB, J. 1985. Transfers of Lycopodium spe- cies to Huperzia: With a note on generic clas- sification in Huperziaceae. Folia Geobot. Phy- totax. 20: 67-80. HOLUB, J. 1991. Some taxonomic changes with- in Lycopodiales. Folia Geobot. Phytotax. 26: 81-94. 0LLGAARD, B. 1987. A revised classification of the Lycopodiaceae sens. lat. Opera Bot. 92: 1 53- 178. 0LLGAARD, B. 1988. Lycopodiaceae, in G. Har- ling and L. Andersson, Flora of Ecuador 33: 1- 155. 0LLGAARD, B. 1989. Index of the Lycopodi- aceae. Biol. Skr. Dan. Vid. Selsk. 34: 1-135. 0LLGAARD, B. 1992. Neotropical Lycopodi- aceae—An overview. Ann. Missouri Bot. Gard. 79:687-717. ROTHMALER, W. 1 944. Pteridophytcn-Studien I, Feddes Repert. Spec. Nov. Regni Veg. 54: 55- 82. TRYON, A. F., AND B. LUGARDON. 1991. Spores of the Pteridophyta. Springer- Verlag, New York. TRYON, R. M., AND A. F. TRYON. 1982. Lyco- podiaceae, in Ferns and allied plants, with spe- cial reference to tropical America. Springer- Ver- lag, New York. WAGNER, W. H., AND J. M. BEITEL. 1992. Mod- ern North American Lycopodiaceae and their generic classification. Ann. Missouri Bot. Gard. 79: 676-686. Key to Genera of Lycopodiaceae a. Stem dichotomies equal (isotomous) throughout, plants without elongate, indeterminate main stems, but sometimes with equally thick, somewhat heteroblastic branches; roots usually forming 1 basal tuft; sporophylls and vegetative leaves alike, or the sporophylls, if smaller, persisting, not subpeltate, not ephemeral; spores foveolate or fossulate I. Huperzia a. Stem dichotomies unequal (anisotomous) throughout, the branches differentiated into elongate, in- determinate, rhizomatous, or creeping, or trailing, main stems, and usually determinate aerial branch- let systems; sporophylls strongly modified, ephemeral, unlike vegetative leaves, usually subpeltate, aggregated in compact, terminal strobili; spores reticulate or rugate b b. Strobili erect, sessile or pedunculate, borne on branchlet systems that arise in a dorsolateral position on the main stem; side walls of sporangium epidermis cells sinuate, lignified throughout; spores reticulate II. Lycopodium b. Strobili pendulous and sessile, or strobili erect and terminating simple erect branches that arise dorsally on the creeping stem, side walls of sporangium epidermis cells straight, not lignified, except for nodular or semiannular thickenings; spores rugate III. Lycopodiella TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 17 FIG. 6. Huperzia eversa: a, habit; b, portion of fertile division with sporangia. Huperzia binervia: c, habit; d, portion of fertile division with sporangia. Huperzia linifolia var. tenuifolia: e, habit; f, leaf from basal division; g, portion of fertile division with sporangia. Huperzia cuneifolia: h, habit; i, expanded leaves from basal division; j, fertile shoot tip. (a from Holm-Nielsen et al. 3873, Ecuador, AAU; b from Holmgren 674, Ecuador, s; c, d from Matthews, Peru, K; e, f, g from Holm-Nielsen et al. 4422, Ecuador, AAU; h, i, j from Hutchison & Wright 6943, Peru, p.) 18 FIELDIANA: BOTANY I. Huperzia Huperzia Bernh., J. Dot. (Schrader) 1800 (2): 126. 1801. LECTOTYPE (designated by Roth- maler, Feddes Repert. Spec. Nov. Regni Veg. 54: 59. 1944): Huperzia selago (L.) Martius & Schrank (Lycopodium selago L.). Figure 6. Plananthus Mirbel in Lamarck & Mirbel, Hist. nat. veg. 3: 476. 1802. TYPE: Plananthus selago (L.) Beauv. (Lycopodium selago L.). Lycopodium subgen. Selago Baker, Handb. Fern-Al- lies 8. 1887. TYPE: Lycopodium selago L. Lycopodium subgen. Urostachya Pritzel, Nat. Pflan- zenfam. 1 (4): 592. 1901. TYPE: Lycopodium se- lago L. Urostachys (Pritzel) Herter, Beih. Bot. Centralbl. 39: 249. 1922. TYPE: Urostachys selago (L.) Herter (Lycopodium selago L.). Phlegmariurus Holub, Preslia 36: 21. 1964. TYPE: Phlegmarius phlegmaria (L.) Sen & Sen (Lyco- podium phlegmaria L.). Plants epiphytic or terrestrial, pendent, re- curved, erect or ascending, isotomously branched throughout (except in connection with bulbil for- mation in the group of H. selago, and sprouting from the base of old plants), the branches all sim- ilar (homoblastic), or in some terrestrial species differentiated (heteroblastic) into prostrate (some- times subterranean) versus erect, aerial branches. Roots arising from the stem stele, descending through the cortex to the stem base, here emerging as 1 basal tuft, or in heteroblastic species some- times emerging directly along the underside of prostrate shoots. Shoots homophyllous or hetero- phyllous. Sporophylls and vegetative leaves alike, not peltate, without mucilage cavities, persisting and green after sporangium dehiscence, the grad- ual or abrupt constriction of distal divisions of heterophyllous species associated or not with pres- ence of sporangia. Sporangia axillary, reniform, isovalvate, with a short slender stalk; side walls of sporangium epidermis cells sinuate, thickened and lignified together with the inner walls. Spores fo- veolate or fossulate. Gametophytes (not known for any Peruvian species) usually subterranean (in ter- restrial plants), mycorrhizal, cylindrical with ra- dial or bilateral symmetry, with pluricellular, uni- seriate trichomes among the gametangia. The genus is virtually cosmopolitan, occurring in tropical, temperate, arctic, and alpine environ- ments. Species diversity is highest throughout the tropics in evergreen montane forests, and in the wet Andean grass and shrublands in South Amer- ica. There are approximately 300 species world- wide, 49 in Peru. Seven species are endemic to Peru, four are shared only with Bolivia and more southerly regions, 26 occur also in Ecuador or far- ther north in the Andes and the Neotropics, while 12 species occur both to the north and south of Peru. Species delimitation is problematic almost throughout the genus due to the simple morphol- ogy of the group and to plasticity of the characters (011gaard, 1992). Morphogenesis seems to be somewhat unstable in most species and may be modified by external factors. Most characters are plastic within a species (e.g., stem thickness, num- ber of leaf orthostichies, leaf crowding, leaf direc- tion, color, degree of heterophyllous differentia- tion). Hybridization seems to occur rather frequently, contributing to the blurring of species limits, and the putative hybrids often have nor- mally developed spores. As a consequence, species recognition is often based on some experience and comparison with identified material, rather than a set of definite characters. Measurements and terminology. The descrip- tions and key characters are based on dried ma- terial, unless otherwise indicated. Very often mea- surements and solid shape of organs in life deviate strongly from the dried condition. Branching is isotomous throughout the genus, except in connection with sprouting (lateral branching) from the base of old or injured indi- viduals. Isotomy results in the formation of equal- ly thick branches each with an equal amount of vascular tissue. However, many species of high Andean grasslands develop heteroblastic branch pairs; i.e., the isotomous branches differentiate into distinct aspects and functions due to different leaf development, as exemplified by the sporangiate, erect, aerial shoots and the sterile, prostrate or even subterranean, basal, rejuvenating shoots of Huperzia crassa and H. hypogaea. Homoblastic branches have the same aspect and function and similar leaf development. Phyllotaxis is poorly understood in the Lyco- podiaceae. It seems to be irregularly organized morphogenetically and quite variable within one species, sometimes even within the same individ- ual. Stevenson (Bot. J. Linn. Soc. 72: 8 1-1 00. 1 976) interpreted the phyllotaxis of Huperzia lucidula (Michx.) Trevisan (temperate North America) as consisting of low alternating spirals, and found a definite relation between the number of orthosti- chies and the number of protoxylem lobes in the stele. In many species this relation is different or TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 19 possibly absent, and transitions from spiral to ir- regularly whorled phyllotaxis is common, al- though whorled arrangement is the more com- mon. The number of longitudinal leaf ranks is readily observed in some cases but often must be estimated, or calculated, from the number of leaves in each leaf whorl, or from the number of leaves contained in one rotation of the leaf spiral. As the leaves of a whorl or a rotation of the spiral alter- nate with the adjacent whorls or spirals above and below, the number of leaves in two whorls or ro- tations makes up the number of longitudinal leaf ranks of a shoot. For the sake of brevity, I have described leaf arrangement in terms of leaf whorls, although this is not everywhere completely accu- rate. Reference ROLLERI, C. 1981. Sinopsis de las especies de Lycopodium L. (Lycopodiaceae-Pteridophyta) de la seccion Crassistachys Herter. Revista Mus. La Plata (n.s.) Bot. 13: 61-113. Key to Species of Huperzia a. Plants erect, or ascending to erect, terrestrial or epiphytic and shoot apices erect; leaf margins smooth, fimbriate, denticulate or erose, or leaves papillate b b. Leaf margins denticulate (at least of some leaves, sometimes minutely and remotely denticulate) by pointed teeth, ciliolate, fimbriate, or leaves papillate c c. Leaves densely papillate abaxially, at least at the apex 12. H. weberbaueri c. Leaves not papillate d d. Leaves of upper, sporangiate divisions loosely to closely appressed, abaxially strongly convex throughout; leaf margins fimbriate by irregularly shaped and directed pale and soft teeth; plants bright red or tinged with red e e. Erect shoots predominantly quadrangular 25. H. tetragona e. Erect shoots hexagonal or terete f f. Leaves of upper, sporangiate branches triangular-lanceolate to triangular-ovate, with smooth abaxial epidermis, arranged in alternating whorls of 3-4 24. H. attenuata f. Leaves of upper, sporangiate branches strongly uneven abaxially by protruding, blister- like epidermal cells, arranged in alternating whorls of 5 23. H. sagasteguiana d. Leaves of upper, sporangiate divisions patent to recurved (sometimes appressed and green in H. affinis), abaxially flat or convex; leaf margins ciliate or denticulate; plants green . . . g g. Leaves borne in alternating whorls of 4-5(-6), denticulate or long-ciliate, narrowly tri- angular-lanceolate, linear-lanceolate or nearly subulate, (2.5-)3-6 mm long, 0.5-1.5 mm wide h h. Sporangiate leaves usually with long-ciliate margins, 1-1.5 mm wide at the somewhat clasping base, 4-6 mm long, spreading to appressed 10. H. affinis h. Sporangiate leaves denticulate, to 1 mm wide at the base, 2.5-5 mm long, not clasping, spreading to strongly reflexed 9. H. eversa g. Leaves borne in alternating whorls of (6-)7-l 1, denticulate, subulate to linear-subulate, 4-10 mm long i i. Leaves borne in alternating whorls of (6-)7-8, ascending to spreading or sharply reflexed, straight to strongly recurved, usually evenly tapering, with very sparsely to densely and sharply denticulate to short-ciliolate margins throughout 7. H. reflexa i. Leaves borne in alternating whorls of 8-11, usually sharply bent upward from a perpendicular junction to the stem and then gently to strongly claw-like recurved, narrowed shortly above the base, with rather densely denticulate-ciliolate margins at base, usually sparsely denticulate or smooth at apex 8. H. acifolia b. Leaf margins entirely smooth, or uneven, rugose to erose-rugose, without pointed teeth or cilia, not fimbriate or papillate, or papillate only on leaf base margins j j. Epiphytes with spreading leaves; leaves filiform, up to 0.5 mm wide, (6-) 10-1 7 mm long, leaf bases often red . . . 36. H. wilsonii 20 FIELDIANA: BOTANY j. Terrestrial plants with spreading, reflexed or appressed leaves; leaves linear-filiform to nearly orbicular, not red k k. Leaves wide-spreading to reflexed or patent-ascending, the widest ones linear-filiform to lanceolate, without a basal swelling (air sac) 1 1. Broadest leaves of sporangiate divisions linear-filiform to subulate, 1.5 mm wide or less m m. Leaves softly herbaceous, usually straight or slightly recurved, 0.5-1.3 mm wide, usually flat abaxially n n. Leaves linear to linear-subulate, 0.8-1.3 mm wide at base, slightly convex to canaliculate adaxially 1 . H. hippuridea n. Leaves linear-filiform, 0.5-0.8 mm wide at the base, canaliculate abaxially .... 2. H. lechleri m. Leaves coriaceous, usually sigmoid, 1.2-1.5 mm wide, usually bisulcate abaxially . 4. H. binervia 1. Leaves lanceolate, the narrowest ones 1.5 mm wide or more o o. Leaves reflexed to patent-ascending, thickly coriaceous, adaxially evenly convex, or with slightly prominent vein, margins flat or involute, smooth 5. H. weddellii o. Leaves papery to thinly subcoriaceous, folded slightly down along the vein, margins revolute, minutely rugose 6. H. brongniartii k. Leaves, at least of sporangiate divisions, appressed, the narrowest ones linear-lanceolate to broadly triangular-cordate and appressed, with or without a basal swelling, or patent to reflexed and oblong, ovate or wider p p. Shoots strongly heteroblastic, with deeply subterranean, elongate, horizontal shoots bear- ing stiffly erect aerial branches 22. H. hypogaea p. Shoots homoblastic to heteroblastic, above the ground or shallowly subterranean, not deeply subterranean q q. Sporangiate divisions with shortest leaves 6 mm or longer, linear to widely lanceolate or widely triangular-ovate r r. Aerial shoots somewhat club-shaped, compactly caespitose, narrowed and some- what etiolated at the base and here with pale and irregularly appressed leaves; shoots strongly heteroblastic, the basal, prostrate shoots short and with much reduced leaves 21. H. saururus r. Aerial shoots usually equally thick throughout or tapering, loosely to densely caes- pitose, with all leaves nearly alike; shoots homoblastic or heteroblastic; prostrate shoots (if any) with leaves the same size or larger than those of aerial shoots . . s s. Shoots heteroblastic, with prostrate basal, rejuvenating, often rooting shoots bearing erect, fingerlike aerial shoots t t. Aerial shoots often short, unbranched, appearing narrower than creeping shoots; leaves of creeping shoots larger than in aerial shoots 20. H. andina t. Aerial shoots well developed, simple or branched, not appearing narrower than creeping shoots; leaves of creeping and aerial shoots of nearly equal size u u. Leaves dull, pruinose, usually red-tinged, abaxially rugose from protrud- ing, blisterlike epidermis cells (rarely smooth) v v. Leaves 5-9 x 1-2 mm, with a prominent basal air cavity causing a perpendicular appearance of the junction to the stem in sporangiate leaves 18. H. crassa v. Leaves 9-12 x 2-2.5 mm, without a prominent basal air cavity, ap- pressed from the base 19. H. nesselii u. Leaves dull to somewhat glossy, green, abaxially smooth w w. Leaves of aerial shoots lanceolate to widely lanceolate, slightly long- acuminate, (1.8-)2-2.5(-3) mm wide, strongly curved upward at apex . . 17. H. darwiniana TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 21 w. Leaves of aerial shoots lanceolate, evenly tapering, 1.5-2 mm wide, straight to slightly curved upward at apex 16. H. macbridei s. Shoots homoblastic, all essentially alike x x. Leaves of basal divisions linear to linear-subulate, up to 1 .5 mm wide, usually bisulcate abaxially, usually somewhat sigmoid 4. H. binervia x. Leaves of basal divisions linear-lanceolate or wider, 1.5—4 mm wide, not bisulcate abaxially, sigmoid or not y y. Plants rather fragile and slender; stems 2-3.5 mm thick excluding leaves; leaves 5-10 x 1.5-2 mm, distal divisions ca. 5-10 mm in diameter in- cluding leaves 15. H. capellae y. Plants large, robust; stems 3-8 mm thick excluding leaves; leaves 5-12 x 2-4 mm, distal divisions ca. (7-) 1 0-25 mm in diameter including leaves z z. Leaves of aerial shoots evenly tapering to short-acute, straight or curved apex aa aa. Leaves triangular-lanceolate to widely triangular-ovate, straight to slightly curved upward; leaf base margins in sporangiate leaves usually slightly revolute, sometimes subauriculate 11. H. kuesteri aa. Leaves lanceolate to widely lanceolate, usually strongly curved upward (rarely patent-ascending and somewhat sigmoid); leaf bas- es not revolute or auriculate 14. H. polylepidetorum z. Leaves of aerial shoots slightly long-acuminate, strongly curved upward at apex 1 7. H. darwiniana q. Sporangiate divisions with longest leaves up to 6 mm long, lanceolate or wider .... bb bb. Leaves (longest ones) up to 4 mm long, ovate, elliptic, or cordate cc cc. Broadest leaves ovate or elliptic, (1.8-)2-2.6(-3.5) mm long, (1. 2-) 1.5-2(2. 4) mm wide, sporangia 1-1.5 mm wide 26. H. sellifolia cc. Broadest leaves widely ovate to widely suborbicular-cordate or triangular- cordate, 2-4 mm long, 2— 4(-5) mm wide, sporangia 1.5-2.5 mm wide .... 28. H. brevifolia bb. Leaves, even shorter ones, 4 mm or longer, or if shorter, then lanceolate . . . dd dd. Stem, excluding leaves, 1-1.5 mm thick, sporangia 2-2.5 mm wide, subgl- obose 27. H. engleri dd. Stem, excluding leaves, usually more than 2-4 mm thick; sporangia 1.5-2 mm wide, flattened ee ee. Leaves triangular-ovate to widely ovate, usually red-tinged, leaf bases with a strongly prominent air cavity 29. H. hohenackeri ee. Leaves lanceolate, green, leaf bases without or with a slightly prominent air cavity 13. H. colanensis a. Plants pendulous, or initially erect with nodding to pendulous shoot apices, epiphytic or terrestrial; leaf margins entire ff ff. Leaves more or less sharply dimorphic: basal divisions with long, expanded leaves, apical divisions constricted, with appressed, reduced, decussate or subdecussate leaves; or the entire plant covered by appressed, short, broad, decussate leaves gg gg. Plants robust, stem excluding leaves at least 2 mm thick at the base, without red color- ation hh hh. Constricted divisions sharply quadrangular, 3-8 mm thick including leaves; sporangiate leaves 3-8 mm long, sharply carinate, often with a conduplicate apex 41. H. molongensis hh. Constricted divisions bluntly quadrangular to subterete, 2-4 mm thick; sporangiate leaves 3 mm long or less, rounded to bluntly carinate throughout abaxially, not conduplicate apically ii ii. Plants entirely pendulous, expanded leaves of proximal divisions in usually densely crowd- ed, alternating whorls of 3, the whorls 2.5-5 mm apart; sporangiate leaves 2-3 mm long 22 FIELDIANA: BOTANY 42. H. campiana ii. Plants erect to scandent, with recurved shoot tips and constricted divisions, expanded leaves of proximal divisions in distant, alternating whorls of 4, the whorls 6-9 mm apart; sporangiate leaves 1.2-1.6 mm long 9. H. pruinosa gg. Plants slender, stem excluding leaves usually 1 mm thick or less (rarely to 1 .7 mm) at the base; stem base with or without red coloration jj jj. Expanded leaves of basal divisions very uniform in size, shape, and position throughout, closely situated, with almost continuously overlapping leaf margins (pressed specimens), borne in alternating whorls of 3 at the base, broadly lanceolate to ovate, acute; stems not red 44. H. ericifolia jj . Expanded leaves uniform or variable in shape and position, close to distant and not continuously overlapping, decussate, subdecussate or in alternating whorls of 3 at the base, linear-subulate to ovate or oblanceolate; stems with or without red coloration kk kk. Expanded leaves decussate or subdecussate, lanceolate to ovate, the broadest ones 2-3.5 mm wide 11 11. Constricted terminal divisions usually not sharply distinct from expanded divisions, with irregularly sized and directed leaves; expanded divisions usually well developed and more extensive than constricted divisions 45. H. myrsinites 11. Constricted terminal divisions sharply distinct from expanded divisions, with reg- ularly sized and uniformly directed leaves throughout; expanded leaves restricted to a short zone at the base (rarely lacking), and sometimes inserted in short zones of constricted divisions 43. H. heteroclita kk. Expanded leaves decussate or in whorls of 3, linear-subulate to lanceolate or oblan- ceolate, the broadest ones 2 mm wide or less, or if wider then less than 6.5 mm long and oblanceolate mm mm. Expanded leaves decussate or in whorls of 3, linear-subulate to lanceolate, the broadest ones 1-2 mm broad, the longest ones 8-15 mm long nn nn. Expanded leaves linear, or linear-subulate to linear-lanceolate, 0.5-1 mm broad 47. H. subulata nn. Expanded leaves linear-lanceolate to lanceolate, 1.3-2 mm broad 46. H. phylicifolia mm. Expanded leaves decussate, lanceolate to oblanceolate, the broadest ones 1.5-3 mm broad, the longest ones less than 6.5 mm long 48. H. cuneifolia ff. Leaves uniform throughout, long and expanded, or gradually smaller and more appressed, but not predominantly decussate or subdecussate toward the shoot apices oo oo. Plants very delicate, the longest leaves less than 6(-8) mm long, acicular, less than 0.5 mm broad; stems less than 1 mm thick at base excluding leaves pp pp. Non-sporangiate leaves of narrow terminal divisions closely appressed to the stem, with cuneate to rounded base; leaves of basal divisions usually strongly curved upward and inward from a patent base, or sigmoid 39. H. curvifolia pp. Non-sporangiate leaves of terminal divisions with a patent, rounded to auriculate or subhastate leaf base; leaves of basal divisions spreading to ascending, often unilaterally curved 40. H. tenuis oo. Plants slender to robust, longest leaves of basal divisions more than 8 mm long; leaves filiform or linear to lanceolate; stems 0.5-5 mm thick at base, excluding leaves qq qq. Leaves of basal divisions alternate or paired, or in occasional whorls of 3, more than 13 mm long, 1—4 mm broad, with a narrowed, twisted, often petiolelike and approximately perpendicular lamina base, leaves of terminal divisions alternate to whorled; stem base usually 1 mm or less thick excluding leaves 35. H. linifolia qq. Leaves of basal divisions predominantly or entirely whorled, without petiolelike lamina bases, 8-23 mm long; stem base 0.5-5 mm or more thick rr rr. Leaves closely appressed throughout, linear-subulate to linear-lanceolate, apically con- vex to conduplicate abaxially ss ss. Leaves borne in whorls of 7-8, linear-subulate, falcate- appressed TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 23 33. H. funiformis ss. Leaves borne in whorls of 4-5, linear to linear-lanceolate, straight to slightly recurved at apex 34. H. buesii IT. Leaves, at least in basal divisions, ascending to patent, filiform to lanceolate, flat, concave, or convex tt tt. Leaves filiform to linear, or linear-subulate from an auriculate leaf base, up to 1 mm broad; the leaves, if linear and 1 mm broad, borne in whorls of 5 or more uu uu. Leaf bases of non-sporangiate leaves auriculate and often somewhat overlap- ping neighboring leaf bases of the same whorl; lamina usually less than 0.7 mm broad above the auriculate base 38. H. sarmentosa uu. Leaf bases of non-sporangiate leaves without auricles; lamina 0.2-1 mm broad vv w. Leaves filiform, 0.3-0.5 mm broad just above the base ww ww. Leaves not twisted at base, perpendicularly spreading to slightly upward curved, or ascending, (6-) 10- 17 mm long, with green or bright red bases; stems usually 1.5-2 mm thick . . . 36. H. wilsonii ww. Leaves twisted at base, obliquely falcate-ascending, 6- 1 0(- 1 2) mm long, with green leaf bases; stems usually 1-1.5 mm thick 37. H. polycarpos vv. Leaves linear, 0.7-1 mm broad, (10-)14-19 mm long ... 3. H. arcuata tt. Leaves linear- lanceolate to lanceolate, the broadest ones more than 1.5 mm broad; or the non-sporangiate leaves of basal divisions linear, 1 mm broad or more, and borne in whorls of 3-4 xx xx. Leaf margins minutely uneven by individually protruding epidermis cells, especially near the apex; leaves lanceolate, 2—4 mm broad 32. H. rosenstockiana xx. Leaf margins smooth; leaves linear to lanceolate, 1-3.5 mm broad yy yy. Stem base 2.5-5 mm thick excluding leaves; leaves, at least of basal di- visions, brightly shining, firmly coriaceous throughout, not twisted at base, 2.5-3.5 mm broad; sporangia 1.7-3 mm broad .... 30. H. hartwegiana yy. Stem base (1-) 1.5-2 mm thick; leaves dull or slightly shining, firmly her- baceous to subcoriaceous, twisted or straight at base, 2-2.5 mm broad; sporangia 1-2.2 mm broad 31. H. taxifolia 1. Huperzia hippuridea (Christ) Holub, Folia Geobot. Phytotax. 20: 73. 1985. Lycopodium hippurideum Christ in Pittier, Primit. Fl. Costar. 3 (1): 56. 1901. TYPE: Costa Rica, El Paramo, 3000 m, massif de Buena Vista, 1897, Pittier 10619 (holotype, P!; isotype, us!). Urostachys hippurideus (Christ) Nessel, Barlappge- wachse 88. 1939. Urostachys poseidonis Herter, Revista Sudamer. Bot. 10: 122-123. 195 3. TYPE: Ecuador, Prov.Chim- borazo, Penipe, 3400 m, Rose in Mi lie 35 (ho- lotype, us!; isotypes, GH!, NY!). Lycopodium poseidonis (Herter) Morton, Amer. Fern J. 54: 42. 1964. Shoots homophyllous, equally thick throughout, 10—35 mm in diameter including leaves. Stems excluding leaves 2.5-4 mm thick at the base, ta- pering to ca. 2-3 mm upward. Leaves borne in irregular alternating whorls of 5-8, spreading to reflexed, sometimes sharply reflexed and ap- pressed to the stem, linear to linear-subulate, even- ly tapering from the base or the middle, (10-)1 1- 19 mm long, 0.8-1.3 mm wide, not, or rarely, twisted at base, adaxially with a slightly prominent vein, with smooth, usually slightly revolute mar- gins, and indistinctly to prominently and widely decurrent bases. Sporangia 1.5-2 mm wide. Plants ascending to stiffly erect from a decum- bent base, up to 60 cm tall, sparsely branched. Upper montane forest, especially near the forest limit, usually on the forest floor, in semishade, alt. 24 FIELDIANA: BOTANY 2500-3500 m, Cajamarca, San Martin, Pasco, Cuzco. Central America; Andes from Venezuela to Bo- livia. Huperzia hippuridea belongs to a group of close- ly related taxa of high montane forests throughout tropical America, including also H. arcuata B. 011g. (Colombia, Ecuador), H. lechleri, H. montana (Underw. & Lloyd) Holub (Greater Antilles), H. nuda (Nessel) B. 011g. & Windisch (Brazil). Hutchison & Bismarck 6455 and Matthews 963 (presumably from Chachapoyas) have thicker stems and more crowded and coriaceous leaves than usual for the species, thus approaching Hu- perzia \veddellii (Herter) Holub and H. loxensis B. 011g. Matthews 963 is a mixed collection, sheets of the same number at BM pertaining to H. mac- bridei. Cajamarca: Prov. Hualgayoc, Hda. Taulis, 4.6 km be- yond Palmito junction on the road to La Playa, 2740 m, Hutchison & Bismarck 6455 (uc, USM). San Martin: Prov. Mariscal Caceres, NW corner of Rio Abiseo National Park, Chochos, 3500 m, Young & Leon 4660 (AAU). Prov. Mariscal Caceres, Puerta del Monte, 3100-3300 m, Young & Leon 4436 (AAU), 3450 m, Leon & Young 1305 (USM). Pasco: Prov. Oxapampa, Santa Barbara, 3200-3300 m, D. Smith 8176 (AAU, USM). Cuzco: Prov. Paucartambo, Cerro Macho Cruz, Parque Nacional Manu, 3400 m, Leon 2303 (USM). Department unknown: (possibly from Chachapoyas) Matthews 963 (G). 2. Huperzia lechleri (Hieron.) Holub, Folia Geo- bot. Phytotax. 20: 74. 1985. Lycopodium lechleri Hieron., Bot. Jahrb. Syst. 34: 57 1 . 1905. LECTOTYPE (designated by Rolleri, Re- vista Mus. La Plata (n.s.) Bot. 13 (78): 82. 1981): Peru (Dept. Puno), Tabina, Lechler, ed. Hohe- nacker 20/2 (B!; isotypes, G!, K!, P!, UPS!). Vrostachys lechleri (Hieron.) Nessel, Barlappgewachse 85. 1939. Vrostachys lechleri Hieron. var. lehmannii Nessel, Re- vista Sudamer. Bot. 6: 61. 1940. SYNTYPES: BONN, Herb. Nessel 148\, Ecuador, Spruce, Palla- tanga 1858, marked 7; Ecuador: unknown collec- tor "Aus dem Herbar Bonaparte, Paris," anno- tated "Ur. lehmannii Boge."). Urostachys lehmannii (Nessel) Herter, Index Lyco- podiorum 67. 1949. In most features resembling Huperzia hippuri- dea closely, but differing consistently in the nar- rower, linear to nearly filiform leaves 10-23 mm long and 0.5-0.8 mm wide at the base, adaxially usually canaliculate, with a slightly prominent vein at the base, and the margins not revolute, the spo- rangia 1-1.8 mm wide. Upper montane forest, especially near the forest limit, usually on the forest floor, in semishade, alt. 1 800-3600 m, Junin, Huancavelica, Cuzco, Puno. Peru and Bolivia. Closely related to Huperzia hippuridea (see the preceding species for discussion). Junin: Prov. Tarma, Carpapata, Soukup 3477 (F, GH). Huacapistana, 1800-2400 m, Killip & Smith 24503 (F, GH, us). Huancavelica: Prov. Tayacaja, Marcavalle, be- tween Huachocolpa and Tintay, 2600 m, Tovar 4085 (USM), Tovar 4089 (USM). Cuzco: Valle Occobamba [Oco- bamba], 1900 m, Bites 856 (us). Prov. Paucartambo, Valle de Pilcopata, near Pillahuata, 2500 m, Foster & Wachter 7501 (AAU). Pillahuata, 2850 m, Fitzpatrick & Willard (F). San Ignacio, Huadquina, Biies 1410 (us). Puno: Ollachea to San Gaban, 1000-2000 m, Dillon et al. 1156 (AAU, F, MO). Prov. Carabaya, road San Gaban (Lanlacuni Bajo) to Macusani, near Ollachea, 2600 m, Maas et al. 6111 (AAU, USM). Department unknown: Lechler 2023a (K). Vargas 16767 (GH). 3. Huperzia arcuata B. 011g. in Harling and An- dersson, Fl. Ecuador 33: 15, t. ID. 1988. TYPE: Ecuador, Prov. Carchi, Road El Angel to Tulcan, 3500 m, Holm-Nielsen et al. 5341 (holotype, AAU!; isotypes, GB!, QCA!, us!). Plants ascending to erect from a decumbent base, with nodding shoot apices, up to 30 cm tall (ter- restrial), or recurved to pendulous and up to 90 cm long (epiphytes), sparsely branched. Shoots homophyllous, equally thick throughout, 25-30 mm in diameter, or tapering to ca. 12 mm in- cluding leaves. Stem excluding leaves 2-3 mm thick at base, tapering to ca. 1.5 mm upward. Leaves borne in irregular alternating whorls of 6—7 near the base, upward of 5-6, spreading to falcately ascending, linear to subulate from a slightly wid- ened lamina base, ( 1 0-) 14-19 mm long, (0.5-)0.7- 1 mm wide above the widened base, gradually tapering in the distal half or so, adaxially flat to slightly concave, or with slightly revolute margins, with abaxially slightly prominent vein and smooth margins, with slightly widened lamina base, twist- ing the lamina to a vertical position. Vegetative leaves of basal divisions often spreading to re Hexed and not twisted. Sporangia 1 .3-2 mm in diameter. Upper montane forest, near timberline, on the forest floor, in semishade, or epiphytic, alt. 2700- 3500 m, Amazonas, San Martin. Southern Colombia to Peru. TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 25 Closely related to Huperzia hippuridea, from which it differs by its obliquely falcate-ascending, twisted leaves and the nodding to pendulous shoot tips. Huperzia arcuata also resembles H. dicho- toma (Jacq.) Trevisan (tropical America) but is larger in all parts and restricted to timberline forest habitats. The Peruvian collections generally have short leaves, 8-15 mm long. Cajamarca: Prov. Cutervo, Llama to Huambos, 2700 m, Lopez et al. 6591 (AAU, OH, MO). Cutervo, Raimondi 3126 (B, USM). Trail Socota to Tambillo, 3300 m, Stork & Morton 10174 (F, K, uc). Amazonas: Prov. Chacha- poyas, Cerros Calla Calla, W side, 45 km above Balsas, 3100 m, Hutchison & Wright 5784B (uc). Cerros Calla Calla, E side, 1 9 km above Leimebamba, 3 1 00 m, Hutch- ison & Wright 5559 (F, GH, M, MO, NY, p, uc). Cano Santa Lucia just E of Chachapoyas, 2000-2400 m, Wurdack 600 (F, GH, K, uc). Amazonas: Chachapoyas, Matthews (E). San Martin: Prov. Mariscal Caceres, Rio Abiseo National Park, Puer- ta del Monte, 3450 m, Young 1627 (AAU, USM). Prov. Mariscal Caceres, near La Playa Camp, 2700 m, Young & Leon 4954 (AAU). 4. Huperzia binervia (Herter) B. 011g., Opera Bot. 92: 169. 1987. Lycopodium binervium Herter, Bot. Jahrb. Syst. 43, Beibl. 98: 48. 1909. TYPE: Peru, Prov. Chacha- poyas, Matthews (holotype, P!; isotypes, BM!, GL!, K!). Urostachys binervius (Herter) Nessel, Barlappge- wachse 110. 1939. Plants ascending to erect from a decumbent base, up to 50 cm tall. Shoots homophyllous, equally thick throughout, 1 5-30 mm in diameter includ- ing leaves or with gradually shorter leaves and tapering to 5-8 mm. Stems excluding leaves 3-6 mm thick at the base, sometimes tapering to 2- 2.5 mm, ridged by decurrent leaf bases and mar- gins. Leaves borne in very close alternating whorls of 6-9, spreading or somewhat reflexed to ascend- ing or appressed upward, slightly recurved or sig- moid, linear to linear-subulate, 7-16 mm long, 1.2-1.5 mm wide, upwards sometimes reduced to 3-7 mm long, not twisted at base, thick, coria- ceous, adaxially convex and shallowly rounded or with a slightly to sharply prominent veinal ridge, abaxially convex and shallowly rounded, when dried usually with a prominent vein and irregular longitudinal ridges or wrinkles, or sometimes the vein sunken and hence the adjacent leaf tissue ap- pearing as 2 longitudinal veins, with smooth mar- gins, green. Stomates often forming 2 irregular, slightly sunken longitudinal bands along the vein. Sporangia 1.5-2 mm wide. Open shrubland and grassland (jalca), seepage areas, 2000-3100 m, Cajamarca, Amazonas. Endemic. Related to Huperzia loxensis B. 011g. (southern Ecuador) and H. hippuridea. 5. Huperzia weddellii (Herter) Holub, Folia Geo- bot. Phytotax. 20: 78. 1985. Lycopodium weddellii Herter, Bot. Jahrb. 43: Beibl. 98: 45. 1909. TYPE: Peru, Dept. Puno, Carabaya, 1847, Weddell 4684 (holotype, P!; isotype frag., BONN, Herb. Nessel 1701 in part). Urostachys weddellii (Herter) Nessel, Barlappge- wachse91. 1939. Plants erect, or erect from a decumbent base, very robust, up to 30 cm tall, or to 50(-70) cm long, sparsely branched. Shoots homophyllous, al- most equally thick throughout, (14-) 16-2 3 mm in diameter including leaves. Stems excluding leaves 3-6 mm thick at the base, tapering to 3—4 mm in diameter. Leaves almost uniform throughout, borne in alternating whorls of 6-7, ascending to spreading or reflexed, straight, curved upward or recurved, lanceolate, acute to slightly acuminate or short-acute, 6-ll(-12) mm long, 1.5-2.5 mm wide, thick and coriaceous, often glaucous, rarely with red tips, adaxially convex with a slightly prominent vein, usually irregularly concave abax- ially (dried), with vein abaxially obscure to dis- tinctly and widely prominent, sometimes in as- cending leaves abaxially convex and rounded with obscure vein, with usually smooth to minutely un- evenly rugose, narrowly sclerified, translucent margins, or rarely papillate near the base. Leaf bases prominently decurrent, with a small, some- times indistinct swelling (air sac), especially in sporangiate leaves. Sporangia 1.5-2.2 mm in di- ameter. Terrestrial in exposed habitats in timberline for- est and lower shrub paramo, 2600-3900 m, Ama- zonas, Puno. Ecuador to Bolivia. Amazonas: Prov. Bagua, Cordillera Colan, NE of La Peca, ca. 3 1 70 m, Barbour 3446 (MO). 6. Huperzia brongniartii (Spring) Trev., Atti Soc. Ital. Sci. Nat. 17:248. 1874. 26 FIELDIANA: BOTANY Lycopodium brongniartii Spring, Bull. Acad. roy. Sci. Bruxelles 8 (2): 515. 1841. TYPE: Bolivia, Yun- gas, D'Orbigny 227 (holotype, P!; isotypes, BONN, Herb. Nessel 169\ in part, BR!). Lycopodium taxifolium Sw. var. brongniartii (Spring) Baker, Handb. Fern Allies 16. 1887. Urostachys brongniartii (Spring) Nessel, Arch. Bot. Est. S. Paulo 1: 388. 1927. Plants erect from an ascending base, up to 50 cm tall, sparsely branched. Shoots homophyllous, almost equally thick throughout, 20-30 mm in diameter including leaves. Stems excluding leaves 2.5—4 mm thick near the base, sometimes tapering to 1.5-2 mm upward, somewhat ridged by decur- rent leaf bases. Leaves uniform throughout, borne in alternating, rather distant whorls of 4-5, wide- spreading to somewhat reflexed, usually straight, not twisted at the base, lanceolate, with long-acute apex, papery to subcoriaceous and opaque, 9-1 2(- 15) mm long, 2-3 mm wide, almost flat, with prominent vein adaxially, or folded slightly down along the vein, with slightly revolute, minutely rugose margins. Sporangia ca. 2.5 mm wide. Terrestrial in humid montane forest, ca. 2900- 3300 m, Cuzco. Colombia to Bolivia. Related to Huperzia rosenstockiana (Herter) Holub with which it shares the minutely rugose leaf margins. The specimens cited correspond to the Bolivian representatives of the species. Cuzco: Prov. Urubamba, Machu Picchu, hillside above Rio Mandor, 2920 m, Peyton 1350 (AAU). Alturas de Rio Calzada, Huadquina, 3300 m, Sues 756 (us). Prov. Paucartambo, Parque Nacional Manu, Road Acjanaco to Pilcopata, bridge between Pillahuata and La Esperan- za, 2750 m, Leon 2228 (USM); Prov. Paucartambo, Par- que Nacional Manu, below Acjanaco, 2950 m, Leon & Cano 2129 (USM). 7. Huperzia reflexa (Lam.) Trev., Atti Soc. Ital. Sci. Nat. 17: 248. 1874. Lycopodium reflexum Lam., Encycl. 3: 653. 1789. TYPE: Martinique, Comm. Joseph Martin (ho- lotype, P, Herb. Lam. 4421). Plananthus reflexus (Lam.) Beauv., Prodr. Aeth. 100. 1805. Urostachys reflexus (Lam.) Herter, Beih. Bot. Cen- tralbl. 39: 249. 1922. Plants erect or ascending from a decumbent base, soft, usually loosely caespitose, 10-30(-40) cm tall. Shoots homophyllous, almost equally thick throughout, 7-1 5 mm in diameter including leaves. Stems excluding leaves 1 .5-3(-4) mm thick at base, sometimes tapering to 1-1.5 mm in diameter, ridged by decurrent leaves or almost smooth. Leaves borne in alternating irregular whorls of (6-)7-8(-9), ascending to spreading or sharply re- flexed, straight to strongly recurved, linear-subu- late, widest just above the base, 4-8 mm long, 0.5- 1(-1.2) mm wide, softly herbaceous to subcoria- ceous, adaxially convex, or concave near the base, abaxially flat, or slightly concave to convex, with an obscure to somewhat prominent vein, with flat to revolute, very sparsely to densely denticulate to short-ciliolate margins. Leaf bases often some- what decurrent. Sporangia 1-1.5 mm in diameter. A frequent pioneer on permanently moist, dis- turbed ground, mainly in the montane forest zone, also in peat bogs, or rarely as a low epiphyte. Tropical America. Closely related to Huperzia eversa, H. acifolia, and H. affinis. Huperzia reflexa is variable with respect to leaf and stem size, direction and crowding of leaves, and leaf margin characters. Part of the variation undoubtedly reflects variable growth conditions, but very often several distinct forms can be found growing intermixed in the same habitat, indicating that genetic differences exist. The variation pat- terns are complex and in need of a detailed study. A variety with smaller dimensions stands some- what apart and is recognized taxonomically. The remaining material, referred to the type variety, is polymorphic. Key to Varieties a. Shoots including leaves 9-15(-20) mm in diameter, leaves (5-)6-8 mm long, 0.7-1 (-1.2) mm wide, with regularly denticulate to short-ciliolate margins, sporangia ca. 1.5 mm wide . . 7a. var. reflexa a. Shoots including leaves 7-10 mm in diameter, leaves 4-5(-6) mm long, 0.5-0.7 mm wide, with sparsely and remotely denticulate margins, sporangia ca. 1 mm in diameter 7b. var. minor TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 27 7a. Huperzia reflexa var. reflexa Lycopodium bifidum Willd., Sp. pi. ed. 4, 5: 53. 1810. TYPE: Venezuela, Cuchilla de Guajana Guajana, Humboldt & Bonpland 474 (holotype, B, Herb. Willd. 194211; isotype, p, Herb. Humb.!). Lycopodium reversum Presl, Reliq. haenk. 1:82. 1825. TYPE: Ecuador, Guayaquil, Haenke (holotype, PRC!). Lycopodium reflexum Lam. var. majus Spring, Mem. Acad. roy. Belg. 15 [Mon. Lye. 1]: 26. 1842. Huperzia reflexa (Lam.) Trev. var. bifida (Willd.) Trev., Atti Soc. Ital. Sci. Nat. 17: 248. 1874. Lycopodium reflexum Lam. var. densifolium Baker, Handb. Fern-Allies 11. 1887. SYNTYPES: Hart- weg 1480 (K!); Moritz 2266 (= 226bl K!); Brazil, Glaziou 15797 (K!). Lycopodium densifolium (Baker) Underw. & Lloyd, Bull. Torrey Bot. Club 33: 106. 1906. Lycopodium mexiae Copel., Univ. California Publ. Bot. 19: 294, t. 47. 1941. TYPE: Peru, Dept. Huanuco, Churubamba, trail Cotirarda to Mer- cedes, 1875 m, Mexia 8193a (holotype, uc!; iso- types, GH!, K!, MICH). Urostachys stellae-polaris Herter, Revista Sudamer. Bot. 10: 121. 1953. TYPE: Colombia, Cundina- marca, Guayabetal to Monte Redondo, SE of Quetame, 1 300-1 500 m, Pennell 1801 (holotype, us!). Lycopodium stellae-polaris (Herter) Morton, Amer. Fern J. 54: 72. 1964. Huperzia bifida (Willd.) Holub, Folia Geobot. Phy- totax. 20: 71. 1985. Huperzia mexiae (Copel.) Rolleri and Deferrari, No- tas Mus. La Plata, Bot. 21 (100): 156. 1988. cucho: Prov. La Mar, eastern Massif of the Cord. Central opposing the Cord. Vilcabamba between Tambo, San Miguel, Ayna, and Hda. Luisiana, 1 570 m, Dudley 11826 (GH). Ayna between Huanta and Rio Apurimac, 750- 1000 m, Killip & Smith 23198 (NY). Cuzco: Prov. Con- vencion, Hda. Guayanay, 1800 m, Vargas 13248 (GH). Quispicanchi, Mandor, Marcapata, 1000 m, Vargas 5220 (us). 7b. Huperzia reflexa var. minor (Spring) B. 011g. in Marling and Andersson, Fl. Ecuador 33: 26. 1988. Lycopodium reflexum Lam. var. minus Spring, Mem. Acad. roy. Belg. 15 [Mon. Lye. 1]: 26. 1842. SYN- TYPES: Brasil, Pr. Rio de Janeiro, Gaudichaud (P!); Langsdorff (M, Herb. Mart.l); Brazil, Serra dos Orgaos, fr. Majo, Guillemin (P!); In sylvis prov. Paraensis, Martins (M!); Brazil, in prov. Minarum, Claussen (P!). Lycopodium brutum Herter, Bot. Jahrb. 43: Beibl. 98: 47. 1909. TYPE: Trinidad, Hooker ded. 1845 (holotype, P!). Humid montane forest, road banks, alt. 1000- 2400 m, Huanuco, Cuzco, Puno. Brazil; Trinidad; Ecuador to Bolivia; probably more widespread. This variety seems identical to Huperzia par- vifolium (Nessel) Rolleri and Deferrari from SE Brazil. It often grows intermixed with individuals of var. reflexa. Landslides, road banks, and other open or dis- turbed habitats in montane forest, alt. 900-3400 m, Cajamarca, Amazonas, San Martin, Huanuco, Pasco, Junin, Ayacucho, Cuzco. Throughout humid mountainous regions of tropical America. Foster & Smith 9095 (USM) is very robust and tall, with thicker stems and slightly wider leaves than usual for this variety. Huanuco: Prov. Huanuco, Road Huanuco to Tingo Maria, N of Carpish Pass, 2350-2450 m, Plowman & Rury 11146 (F, GH). Cuzco: Prov. Urubamba, Machu Picchu, Soukup 167 (F). Near Machu Picchu, 2000 m, Tryon & Tryon 5409 in part (GH, us, USM). Puno: Prov. Sandia, below Cuyocuyo, 2900 m, Ferreyra 16624 (USM). 8. Huperzia acifolia (Rolleri) Rolleri and Defer- rari, Notas Mus. La Plata, Bot. 21 (100): 155. 1988. Cajamarca: Prov. Cutervo, El Pajonal, San Andres, 2200 m, Llatas & Suarez 2828 in part (F). Prov. Cutervo, La Pucarilla, between Socota and San Andres, 2500 m, Sanchez Vega el al. 5923 (AAU, F). Amazonas: Prov. Chachapoyas, roads to Molinopampa, 2700 m, Sanchez Vega et al. 2218 (AAU). San Martin: Prov. San Martin, Dist. Tarapoto, road Tarapoto to Yurimaguas, km 9- 1 3, ca. 700 m, Rimachi 4102 (F, GH, NY). Huanuco: Prov. Huanuco, Chinchao to Puente Duran, 1900 m, Ochoa 14597 (F, us). Carpish Pass, 2700 m, Hodge 6294 (GH, us). Pasco: Manto at Yaupi, Woytkowski 6534 (MO, us). Prov. Oxapampa, Ulcumanu SW of Oxapampa, road to Maria Teresa and Llaupi, 2150-2450 m, Foster et al. 7688 (AAU). Junin: Tarma, Chanchamayo, above La Merced, ca. 2000 m, Weberbauer 2006 (G, MOL). Aya- Lycopodium acifolium Rolleri, Revista Mus. La Plata (n.s.) 14: 2, /. 1-2. 1985. TYPE: Peru, Dept. Aya- cucho, Cearrapa, between Huanta and Rio Apu- rimac, 1 500 m, Killip & Smith 22368 (holotype, us!). Plants erect, or erect from a decumbent base, 1 5-30 cm tall, or up to 60 cm long. Shoots homo- phyllous, almost equally thick throughout, 10-20 mm in diameter including leaves. Stems excluding leaves 2-5(-6) mm thick at the base, sometimes tapering to 1-3 mm thick. Leaves uniform throughout, borne in alternating, usually densely 28 FIELDIANA: BOTANY crowded whorls of 8-1 1, usually sharply bent up- ward from a perpendicular junction to the stem and then gently to strongly clawlike recurved, lin- ear-subulate, widest just above the pale and soft- herbaceous base, 6-10 mm long, 0.7-1(-1.3) mm wide at the base, narrowed shortly above the base, subcoriaceous at apex, adaxially convex distally, abaxially flat to convex with obscure to somewhat prominent vein (sometimes sunken when dried), with rather densely denticulate-ciliolate margins at base, usually sparsely denticulate or smooth at apex. Sporangia 1-1.5 mm wide. Moist banks in montane forest at mid-altitudes, alt. 1350-2040 m, San Martin, Huanuco, Pasco, Junin, Ayacucho, Cuzco. Venezuela to Bolivia. Closely related to Huperzia rejlexa and H. un- guiculata B. 011g. (Colombia, Ecuador). Some ma- terial earlier tentatively referred to H. unguiculata from Ecuador and Peru belongs here, and thus H. unguiculata is yet unknown from Peru. San Martin: Prov. Rioja, Pedro Ruiz— Moyobamba road, km 390-394, Venceremos, 1910-2040 m, D. Smith 4522 (AAU, USM). Prov. Mariscal Caceres, 60 km NE of Tingo Maria, La Divisoria pass through Cerro Azul, 1 500 m, Tryon & Tryon 5269 (GH, us). Huanuco: Prov. Leon- cio Prado, Dist. Daniel Alomia Robles, Road Tingo Maria to Pucallpa, La Divisoria, 1600 m, J. Schunke V. 3082 (F, G, GH, NY, us); Ferreyra 1690 (USM). Pasco: Prov. Oxapampa, Rio Boqueria, ca. 26 km from Oxapampa via Rio Yamaquizu, 1870 m, D. Smith et al. 1772 (AAU, USM). Oxapampa [as Junin], 1600 m, Soukup 2675 (F, GH, us). Junin: Prov. Tarma, road to La Mina Pichita, 9 km from the Tarma-La Merced road, ca. 1 3 km from San Ramon, 1 600 m, Skog et al. 5055 (us). Chancha- mayo Valley, above La Merced at Cumbre Yacunay, 2000 m, Hutchison 11 94 A (F, M, NY, uc, us, USM). Cuzco: Prov. Paucartambo, Kosnipata, San Pedro, 1 350 m, Var- gas 10219 (MO, uc). 9. Huperzia eversa (Poiret) B. 011g. in Harling and Andersson, Fl. Ecuador 33: 28. 1988. Lycopodium reflexum Willd., Sp. pi. ed. 4, 5: 52. 1810, not Lam. 1789. TYPE: Ecuador: Tungurahua, (Humboldt) Nee D. D. (holotype, B, Herb. Willd. 19419\). Lycopodium eversum Poiret, in Lam., Encycl. 3: 556. 1814 [1813]. Lycopodium reflexum Lam. var. polycarpum Sodiro, Recens. crypt, vase. Quit. 90. 1883. TYPE: Ec- uador, valle de Nanegal, Sodiro (not located). Lycopodium polycarpum (Sodiro) Underw. & Lloyd, Bull. Torrey Bot. Club 33: 105. 1906, not L. poly- carpos Kunze 1835. Lycopodium ecuadoricum Herter, Bot. Jahrb. 43: Beibl. 98: 48. 1909. LECTOTYPE (designated by B. 011g. in Harling and Andersson, Fl. Ecuador 33: 28. 1988): Ecuador, Andium nemoribus humidis, Jameson 74 (P!). Huperzia ecuadorica (Herter) Holub, Folia Geobot. Phytotax. 20: 72. 1985. Plants erect or erect from a decumbent base, soft, often large, much-branched and caespitose, up to 30(-50) cm tall. Shoots homophyllous, equally thick throughout, (2.5-)3-6(-10) mm in diameter including leaves. Stems excluding leaves 1.5-2.5 mm thick at the base, sometimes tapering to 1-2 mm, prominently ridged by decurrent leaf bases (dried). Leaves uniform throughout, borne in irregular, alternating, subdistant to densely crowded whorls of (4-)5(-6), wide-spreading to sharply reflexed, usually strongly recurved, linear- lanceolate, widest in the basal half, (2.5-)3-5 mm long, 0.5-1 mm wide, softly herbaceous to sub- coriaceous, adaxially convex with obscure vein, abaxially irregularly concave (dried), with obscure to somewhat prominent vein, with slightly revo- lute, denticulate-ciliolate margins. Leaf base with prominently decurrent vein and margins. Sporan- gia 1-1.5 mm in diameter. Terrestrial, as a pioneer on landslides, road banks, and other open, moist habitats in upper montaine forest, alt. 2400-3400 m, Piura, Ama- zonas, San Martin, Huanuco, Pasco, Ayacucho, Cuzco. Costa Rica; Andes from Venezuela to Bolivia. Huperzia eversa resembles H. rejlexa, but has shorter and relatively wider leaves, which are usu- ally strongly recurved, so that that shoots appear much more slender. It also differs by usually form- ing very densely branched individuals. Piura: Prov. Huancabamba, Loma Redonda (Sapa- lache to Chinguela), 2400 m, Sagdstegui et al. 10174 (AAU, MO). Amazonas: Prov. Chachapoyas, middle east- ern Calla-Calla slopes, 3000-3200 m, Wurdack 1770 (F, GH, NY, uc, us, USM). San Martin: Prov. Mariscal Ca- ceres, Rio Abiseo National park, NW corner, Chochos, 3400 m, Young 3693a (AAU). Huanuco: Playapampa, 2700 m, Macbride 4486 (F). Cushi, trail to Tambo de Vaca, Bryan 622 (F, us). Pasco: Prov. Oxapampa, Dist. Oxapampa, Rio San Alberto, Abra Esperanza, 2400- 2700 m, Foster et al. 10300 (AAU). Ayacucho: Prov. La Mar, eastern Massif of the Cord. Central opposing the Cord. Vilcabamba between Tambo, San Miguel, Ayna, and Hda. Luisiana, 2920 m, Dudley 11972 (F, GH). Cuz- co: Prov. Paucartambo, Parque Nacional Manu, Camino Eriksson, Leon & Huapalla 2387 (AAU). Michehuanunca, TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 29 Huadquina, 2700 m, Biies 745 (us). Department un- known: Matthews 1082 (E). 10. Huperzia affinis Trev., Atti Soc. Ital. Sci. Nat. 17: 248. 1874. Lycopodium affineGrev. & Hooker, Bot. Misc, 2: 364. 1831, not Bory 1804. LECTOTYPE (designated by Nessel, Barlappgewachse 97. 1939): Ecuador, Pichincha, 1824, Herb. Greville, Jameson (K!; possible isotypes, E!, NY!, us!). Lycopodium blepharodes Maxon, Contr. U.S. Natl. Herb. 17: 423. 1914. Huperzia blepharodes (Maxon) Holub, Folia Geobot. Phytotax. 20: 71. 1985. Plants erect from a decumbent base, at least up to 25 cm tall. Shoots homophyllous, equally thick throughout, 5-10 mm in diameter including leaves. Stems excluding leaves 1.5-3 mm thick, upward sometimes slightly tapering. Leaves borne in rath- er close alternating whorls of 5, almost covering the stem, patent to loosely appressed, narrowly triangular-lanceolate to almost subulate, with slightly widened base, (3.5-)4-6 mm long, 1-1.5 mm wide, abaxially convex and evenly rounded or with prominent veinal ridge, at least at base, somewhat clasping the sporangia, with long, flac- cid marginal cilia and small teeth. Sporangia 1.5- 2 mm in diameter. Terrestrial on banks in upper montane forest and grassland, alt. 2700-3100 m, Amazonas, Huanuco. Colombia to Peru. The description above applies to the Peruvian specimens. These are generally smaller, are more compact, and have more appressed and abaxially more convex leaves than specimens from Colom- bia and Ecuador. Huperzia affinis is variable with respect to crowding and direction of the leaves. It is distinguished from H. reflexa var. reflexa, H. eversa, and H. pearcei (Baker) Holub (Bolivia) by its wider and more convex leaf bases and the slen- der cilia on the leaf margins. Amazonas: Prov. Chachapoyas, Cerros Calla Calla, W side, 45 km above Balsas, 3100 m, Hutchison & Wright 5830 (F, GH, NY, uc, USM). Huanuco: Mito, 2700 m, Bryan 389a (F). Prov. Huanuco, Punta de Panao, As- plund 13712 (s). Panao, 2700 m, Macbride 3602 (F). 1 1 . Huperzia kuesteri (Nessel) B. 011g., Opera Bot. 92: 169. 1987. Urostachys kuesteri Nessel, Repert. Spec. Nov. Regni Veg. 35: 182, t. 172. 1934. LECTOTYPE (des- ignated by 011gaard in Harling and Andersson, Fl. Ecuador 33: 37. 1988): Ecuador, Loja to Za- mora, 3500 m, 1875, "/. M. J. Mission" Quito 415 (BONN, Herb. Nessel 23 /!). Plants ascending to erect, massively robust and large caespitose plants, with almost completely homoblastic divisions, up to 40 cm tall. Shoots homophyllous, almost equally thick throughout, or slightly tapering upward and the leaves grad- ually shorter and wider, 12-25 mm in diameter including leaves near the base, sometimes tapering to (7-) 10-1 5 mm. Stems excluding leaves 3-8 mm thick at the base, tapering to 3-5 mm upward, almost completely concealed by leaves. Leaves uniform throughout or upward slightly reduced, densely crowded, borne in usually regular alter- nating whorls of 4-5, ascending to closely imbri- cate, rarely patent to recurved in basal divisions, straight to slightly upwardly curved, narrowly tri- angular-lanceolate near the base to widely trian- gular-ovate in terminal divisions, widest just above the leaf base, 8-12 mm long and 1.5—4 mm wide in basal divisions, upward (4-)5-10 mm long, 2.5- 4 mm wide, evenly tapering, with acute, upward- curved apex, thick, adaxially flat or slightly con- cave with flush or slightly prominent vein, abax- ially rounded with slightly to strongly prominent, narrow vein, with a short, flattened basal swelling (air sac), shining or pruinose, green or rarely with red-tinged margins, with smooth to uneven or ru- gulate, opaquely to transparently sclerified mar- gins. Leaf base margins usually slightly revolute, sometimes subauriculate in apical divisions. Spo- rangia 2-2.5 mm wide. Low, wet mossy and grassy paramos, alt. 2750- 3400 m, Piura, Lambayeque. Southern Ecuador and northernmost Peru. This species is related to Huperzia macbridei but can be distinguished from this by the homo- blastic, evenly spreading branching habit, wider leaves, and greater size. Plura: Purchased in the market of Huancabamba, Friedberg7631b(GH). Lambayeque: Prov. Ferranafe, Dist. Incahuasi, Laguna Tembladera to Cerro Negro, 3300 m, Sagdstegui et al. 12818 (AAU), 12848 (F in part). 12. Huperzia weberbaueri (Nessel) Holub, Folia Geobot. Phytotax. 20: 78. 1985. Urostachys weberbaueri Nessel, Revista Sudamer. Bot. 6: 162, /. 10, f. 41. 1940. TYPE: Peru, Dept. Amazonas, Ostlich Chachapoyas, Tambo Ven- 30 FIELDIANA: BOTANY tillas, 2400-2600 m, Rosen 672 (holotype, BONN, Herb. Nessel 1831). Lycopodium papillatum Rolled, Amer. Fern J. 65: 3. 1975. TYPE: Peru, Dept. Amazonas, Prov. Cha- chapoyas, Cerro Malcabal (Cerro Tumbe), 3-6 km SW of Molinopampa, 2900 m, Wurdack 1456 (holotype, us!; isotypes, GH!, K!, uc!, USM!). Huperzia papillata (Rolleri) Holub, Folia Geobot. Phytotax. 20: 75. 1985. Plants erect from an ascending base, large, caes- pitose, homoblastic, up to 30(-40) cm tall. Shoots homophyllous, 10-15(-20) mm in diameter in- cluding leaves, sometimes tapering to 7 mm in diameter, strikingly whitish papillate in life. Steins excluding leaves 2—4 mm thick. Leaves uniform throughout, or slightly smaller upward, borne in more or less regular, alternating whorls of 5, patent to ascending, upward curved, lanceolate, 7-10(- 15) mm long, 2-2.5 mm wide, not or slightly de- current, without a prominent basal swelling (air sac), abaxially convex, with an obscure veinal ridge, adaxially flat to concave (dried), densely long-pap- illate on margins and abaxially, smooth adaxially. Sporangia ca. 1.5 mm wide. Wet grassy and boggy paramos with some shel- ter, and lower puna 3000-3600 m, Lambayeque, Amazonas, San Martin, Cuzco. Southernmost Ecuador and Peru. The densely papillate, bright whitish leaves in the living plants are unique in the genus. The whit- ish color is often lost after drying, especially heat drying. Lambayeque: Prov. Ferranafe, Dist. Incahuasi, La- guna Tembladera, 3 1 50 m, Sagdstegui et al. 12790 (AAU). Amazonas: Prov. Chachapoyas, Leimebamba to Calla Calla, 3100 m, Sanchez V. 5 JO (AAU); Boeke 1809 (AAU). NE of Tambo de Ventilla, Cerro de Fraijaco (Huaui to Huni), 3450 m, Pennell 15883 (GH). Cerros Calla Calla, 3100 m, Hutchison & Wright 5564 (F, GH, MO, NY, uc, us). San Martin: Prov. Mariscal Caceres, NW corner of Rio Abiseo National Park, 3400 m, Young & Leon 4768 (AAU). Prov. Mariscal Caceres, Chochos Valley, 3300 m, Young & Leon 4877 (AAU). Huallaga, valley of Apison- cho, 30 km above Jucusbamba, 3600 m, Hamilton & Holligan 1216, 1217 (us). Cuzco: Prov. Paucartambo, Cord, de Tres Cruces, 3600 m, Vargas 12195 (GH). Prov. Paucartambo, Pavayoc, 2100 m, Woytkowski 555 (MOL, USM). 13. Huperzia colanensis B. 011g., sp. nov. Planta erecta vel ascendens; surculi homoblastici us- que ad 30 cm alti, homophylli 6-10(-12) mm diametro foliis inclusis, omnino crassitie aequali vel parum an- gustati. Caulis 2-4 mm crassus foliis exclusis. Folia om- nino uniformes vel sursum parum minores, 8-10-faria, patentes usque ad ascendentes vel perpendiculares usque ad parum reflexa, recta vel incurva, lanceolata, aequate angustata vel acuminata, (3-)4-6(-7) mm longa, 1.5-2 mm lata, parum decurrentes, abaxialiter convexa, ad basin vix vel parum ventricosa, plerumque angulo me- diali subacuto instructa, adaxialiter plana usque ad con- cava (siccata), subcoriacea, hypostomatica, plerumque polita, marginibus laevibus, scleroideis et translucidis. Sporangia ca. 1.5-2 mm lata. Plants erect from an ascending base, loosely caespitose, homoblastic, up to 30 cm tall. Shoots homophyllous, 6-10(-12) mm in diameter includ- ing leaves, equally thick throughout or slightly ta- pering. Stems, excluding leaves, 2-4 mm thick. Leaves uniform throughout, or slightly smaller up- ward, borne in more or less regular, alternating whorls of 4-5, patent to ascending or perpendic- ular to slightly reflexed in basal divisions, straight to upward curved, lanceolate, widest ca. '/4 of the leaf length above the base, evenly tapering in the distal % or slightly acuminate, straight to slightly upward curved, (3-)4-6(-7) mm long, 1.5-2 mm wide, somewhat decurrent, without or with a slightly prominent basal swelling (air sac), abaxi- ally convex, rounded or usually with a rather sharp medial ridge, adaxially flat to concave (dried), somewhat coriaceous, usually glossy on both sides, the margins smooth, sclerified and somewhat translucent. Sporangia ca. 1.5-2 mm wide. TYPE— Peru, Dept. Amazonas, Prov. Bagua, Cord. Colan NE of La Peca, 10,800 ft, 8 Sep 1978, Barbour 3384 (holotype, MO!; isotypes, AAU!, USM!). Andean grassland, open patches in shrub para- mo, alt. 3140-3600 m, Amazonas, San Martin. Endemic. The specimen cited from San Martin deviates from the other material by a softer texture and less glossy leaves with less sclerified margins and is included in the species with some doubt. Huperzia colanensis appears to be intermediate in position between the Huperzia brevifolia and H. saururus groups of 011gaard (1987, 1989). It does not appear to be intimately related to other Peruvian species. Amazonas: Prov. Bagua, Cord. Colan NE of La Peca, 3 1 70 m, Barbour 3446 (MO). Prov. Bagua, Cord. Colan NE of La Peca, ridge W of peaks, 3 140 m, Barbour 3197 (MO), 3198 (AAU). San Martin: Dist. Huallaga, Valley of Rio Apisoncho, 3600 m, Hamilton & Holligan 1218 (us). 14. Huperzia polylepidetorum B. 011g. in Harling and Andersson, Fl. Ecuador 33: 42,/ 6B. 1 988. TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 31 TYPE: Ecuador, Prov. Azuay, Laegaard 55117 (holotype, AAU!; isotype, QCA!). Plants ascending to erect, large, robust, caes- pitose, nearly homoblastic, up to 35 cm tall, or more than 50 cm long. Shoots homophyllous, equally thick throughout, or slightly tapering up- ward and the leaves gradually shorter, 10-25 mm in diameter including leaves. Stems excluding leaves 5-7 mm thick at the base, sometimes ta- pering to 2 mm thick, usually not completely con- cealed by leaves. Leaves uniform throughout, or gradually shorter upward, well spaced to densely crowded, borne in more or less regular, alternating whorls of 5-6, patent-ascending to arcuate-ap- pressed, strongly curved upward and inward, lan- ceolate to widely lanceolate, acute or short-acute, 6-12 mm long, (2-)2.5-3(-3.5) mm wide, some- what fleshy, abaxially convex or rarely flattened, with an obscure to prominent adaxial and abaxial, long decurrent median ridge, with an indistinct air sac, adaxially concave or flat, with smooth, slightly involute margins, green throughout or with red- tinged tips. Sporangia 2.5-3.5 mm wide. In open Polylepis forest among mossy rocks, 3900-4300 m, Ancash. Ecuador and Peru. The Peruvian collection was made in shaded Polylepis woods, and the specimen differs slightly from Ecuadorean plants by the more patent-as- cending leaves, which tend to be abaxially flat to slightly concave in the basal divisions, while the distal, more exposed divisions have abaxially con- vex leaves. It thus seems close to Huperzia wed- dellii. Ancash: Prov. Huari, Huascaran National Park, Que- brada de Yuraccocha, a lateral valley of Quebrada Ruri- chinchay, 3900-4300 m, D. Smith et al. 12716 (AAU, F). 1 5. Huperzia capellae (Herter) Holub, Folia Geo- bot. Phytotax. 20: 71. 1985. Urostachys capellae Herter, Revista Sudamer. Hot. 10: 114-115. 1953. LECTOTYPE (designated by Roller!, Revista Mus. La Plata, n.s., Bot. 13 (71): 78. 1981): Ecuador, prov. Napo [as Imbabura], E of Volcan de Cayambe, along trail between Rio Boqueron and Rio Arturo, 1 1,000 ft, Drew E 3 14 (us!; isotype, MSC!). Lycopodium capellae (Herter) Morton, Amer. Fem J. 54: 72. 1964. Plants ascending to erect, sparsely to densely branched, without prostrate-ascending, rejuvenat- ing shoots, up to ca. 35 cm tall. Shoots homo- blastic, homophyllous to gradually slightly hetero- phyllous, 12-20 mm in diameter at the base including leaves, usually tapering to 5-10 mm. Stems excluding leaves 2-3 mm thick at the base, often brown to reddish brown, terete to promi- nently ridged by decurrent leaf bases. Leaves of basal divisions (and shaded shoots) borne in ir- regular, rather distant, alternating whorls of 5-6, usually patent to ascending, not covering the stem, linear-lanceolate, 7-10 mm long, 1.5-2 mm wide, adaxially flat, shining, abaxially flat to slightly con- vex, dull to shining green, usually with long-de- current leaf bases. Leaves of distal divisions grad- ually shorter, closer, and more appressed, borne in alternating whorls of 4-5(-6), ascending to ar- cuate-appressed, straight to strongly upward or unilaterally curved or twisted, lanceolate to broad- ly lanceolate, often slightly cuspidate above the middle, (5-)6-8 mm long, 1.5-2 mm wide, adax- ially concave (dried) or flat to shallowly rounded in life, abaxially convex, rounded and irregularly wrinkled (dried), with slightly prominent, short to long, narrowly decurrent basal swelling (air sac), with smooth to densely irregularly rugulate, slight- ly to distinctly, translucently sclerified margins, shining to dull green. Sporangia ca. 2 mm wide. Grass paramos, usually growing in partial shade among grasses, 3600 m, Cajamarca, Cuzco. Andes from Venezuela to Peru. Known from two collections in Peru, matching the Ecuadorean plants perfectly. Cajamarca: Prov. Chota, Laguna Yahuarcocha, above Incahuasi, 3600 m, Sagdstegui et al. 12908 (F). Cuzco: Prov. Paucartambo, Parque Nacional Manu, vicinity of El Mirador, 3600 m, Leon 2270 (USM). 16. Huperzia macbridei (Herter) B. 011g., Opera Bot. 92: 169. 1987. Urostachys macbridei Herter, Revista Sudamer. Bot. 10: 115. 1953. TYPE: Peru, Dept. Huanuco, 3- 6 mi NW of Mito, 1 1,000 ft, Macbride & Feath- erstone 1922 (holotype, us!; isotypes, B!, F!, MA!, s!). Lycopodium macbridei (Herter) Morton, Amer. Fem J. 54: 72. 1964. Plants erect from a decumbent base, loosely to densely caespitose, with basal, prostrate-ascend- ing, rejuvenating shoots at the periphery, up to 32 cm tall. Shoots homophyllous or with leaves grad- ually slightly reduced upward, equally thick throughout or slightly tapering upward, 1 5-20 mm 32 FIELDIANA: BOTANY in diameter including leaves near the base, often tapering to 6-9 mm in diameter. Stems excluding leaves 2—4 mm thick at the base, upward tapering to ca. 2 mm, almost completely concealed by leaves. Leaves uniform throughout, or upward slightly reduced, borne in irregular, alternating whorls of 4-5(-6), densely crowded, spreading to ascending in basal divisions, upward often grad- ually becoming closely imbricate, straight or slightly upward curved, narrowly lanceolate to lanceolate, widest near the base, evenly tapering, 10-15 mm long and 1.5-2.5 mm wide in basal divisions, up- ward 6-11 mm long, 1.5-2 mm wide, without or with a slightly pronounced basal swelling (air sac), adaxially concave with flush or slightly prominent vein, abaxially rounded, with slightly prominent and often slightly darker and long decurrent vein, with smooth to slightly uneven, somewhat invo- lute, indistinctly sclerified margins, green to yel- lowish green. Sporangia ca. 2.5 mm wide. Terrestrial in low shrub paramos, at the edge of woods, 2500-3600 m, Amazonas, La Libertad, Huanuco, Ayacucho, Cuzco. Southern Ecuador and Peru. Individuals of shaded or sheltered habitats (e.g., Macbride 3487) tend to retain juvenile leaf mor- phology, being completely homophyllous with rel- atively wide and long, patent-ascending rather than appressed and gradually reduced leaves and often with a more loosely caespitose growth habit. Mat- thews 963 is a mixed collection, sheets of the same number at G pertaining to H. hippuridea. Amazonas: Chachapoyas (presumably), Matthews 963 (BM). La Libertad: Prov. Bolivar, Nevado del Cajamar- L] u ilia, puna, Ferreyra 1350 (USM). San Martin: Prov. Mariscal Caceres, Rio Abiseo National Park, Puerta del Monte, 3350 m, Leon & Young 1500 (AAU, USM). Prov. Mariscal Caceres, Rio Abiseo National Park, Valle de Chochos, 3450 m, Leon 1870 (AAU, USM). Huanuco: Mito, 2700 m, Bryan 381 (F). Cani, 7 mi NE of Mito, 2500 m, Macbride 3487 (F, us). Ayacucho: Prov. La Mar, east- ern Massif of the Cord. Central opposing the Cord. Vil- cabamba between Tambo, San Miguel, Ayna and Hda. Luisiana, 3200-3500 m, Dudley 11989 (F, GH). Cuzco: Paucartambo, Tres Cruces, 3600 m, Bikes 2214 (MO). Prov. Paucartambo, Parque Nacional Manu, vicinity of El Mirador, 3600 m, Leon 2269 (USM). 17. Huperzia darwiniana (Nessel) B. 011g., comb, nov. Urostachys darwinianus Nessel [Barlappgewachse 80, /. 3,f. 20. 1939, nom. nud.], Revista Sudamer. Bot. 6: 161. 1940. TYPE: "Columbien: Andinum montibus nemoribus, Ruiz" (holotype, BONN, Herb. Nessel 1331). Plants erect, or erect from a prostrate to as- cending base, sometimes loosely caespitose, with basal, prostrate-ascending, rejuvenating shoots at the periphery, and erect, fingerlike shoots in the center, at least up to 35 cm tall. Erect shoots homo- phyllous or almost so, equally thick throughout, 10-15 mm in diameter including leaves, or some- times tapering to ca. 8 mm upward. Stems ex- cluding leaves 3-5 mm thick at the base, some- times tapering to 2 mm, usually completely concealed by leaves. Leaves uniform throughout, or slightly reduced upward, borne in alternating whorls of 4-7, in distal fully sporangiate divisions usually 4—5, crowded, loosely to closely imbricate, slightly upward curved and usually with a strongly upward curved apex, lanceolate to widely lanceo- late, slightly long-acuminate from a wide and somewhat rounded base, 8-12 mm long, (1.8-)2- 2.5(-3) mm wide, upward sometimes reduced to 7 mm long, with an inconspicuous, narrow, long- decurrent basal swelling (air sac), adaxially con- cave to slightly convex at the base, abaxially con- vex and rounded, with a prominent, often darker colored (dried) ridge along the vein, smooth and often shining, with smooth, slightly sclerified mar- gins, green, not or scarcely pruinose. Sporangia 2- 3 mm wide. Boggy paramo and jalca, humid rocks, 2700- 3600 m, Huanuco, Pasco, Cuzco. Peru and Bolivia. The information on the type label is in Nessel's handwriting, and the geographic information is as dubious as on numerous other Nessel labels. Spec- imens in Geneva, "Peruvia, Herb. Pavon" (G!), are likely to be isotypes. See Comments under Hu- perzia polyclada. Huanuco: Cushi, trail to Tambo de Vaca, Bryan 677 (F, us). Playapampa, ca. 2700 m, Macbride 4475 (F, us). Pasco: Prov. Oxapampa, trail to summit of Cord. Yana- chaga via Rio San Daniel, 3 1 50-3300 m, D. Smith 7710 (AAU, USM). Cuzco: Cerro Puncuyoc, 4500 m, Biles 563 (us). Prov. Urubamba, near Wenner Gren ruins, 3400- 3600 m, Metcalf 30745 in part (uc). Trail Puyupata to Sayaccmarca, 3600 m, Vargas 2910 (us). Department unknown: Perou (1839-1840), Gay (p). Peru, Pearce (us 1431555). 18. Huperzia crassa (Willd.) Rothm., Feddes Re- pert. Spec. Nov. Regni Veg. 54: 60. 1944. Lycopodium crassum Willd., Sp. pi. ed. 4, 5: 50. 1810. TYPE: Ecuador, Antisana, Humboldt & Bon- pland 226 3 (holotype, B,Herb. mild. 194171; iso- types, BM!, P!). TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 33 Urostachys crassus (Willd.) Nessel, Barlappgewachse 75. 1939. Urostachys pilgerianus Nessel, Revista Sudamer. Bot. 6: 161, t. 9,f. 32. 1940. TYPE: "Peru, Cordillere, Rautenstock et Mann, in 1 90 1 " (holotype, BONN, Herb. Nessel 124\). Huperzia pilgeriana (Nessel) Holub, Folia Geobot. Phytotax. 20: 75. 1985. Plants erect from a prostrate to ascending base, slightly to strongly heteroblastic, loosely to densely caespitose with prostrate-ascending or shallowly subterranean basal shoots, these bearing often nu- merous erect, fingerlike shoots in the center, at least up to 40 cm tall. Erect shoots homophyllous or almost so, equally thick throughout, or slightly tapering upward, 5-10 mm in diameter including leaves. Stems excluding leaves 2-3 mm thick, usu- ally completely concealed by leaves. Leaves uni- form throughout, or slightly reduced upward, borne in irregular, alternating whorls (or low spirals) of 5-7, densely crowded, usually closely imbricate, straight (or slightly upward curved in shaded di- visions), linear-lanceolate to lanceolate, 5-9 mm long, (1-) 1.2-2 mm wide, upward sometimes re- duced to 4.5 mm long, with a prominent, short to long decurrent basal swelling (air sac) causing a perpendicular appearance of the very leaf base in sporangiate leaves, adaxially concave to slightly convex, with a raised veinal ridge near the base, abaxially convex and rounded or with a prominent veinal ridge, slightly to strongly rugose by pro- truding, blisterlike epidermal cells, with smooth to rugose margins, green to brick red or dark red, usually strongly pruinose. Sporangia 1-2 mm wide. Terrestrial in high paramo and superparamo, puna, 3600-4850 m, La Libertad, Ancash, Junin, Cuzco, Puno. As here delimited, Huperzia crassa is a widely distributed polymorphic species, occurring from Mexico to Panama, Hispaniola, and in the Andes from Venezuela to Bolivia. The Peruvian material represents a disjunct population, widely separated from the Ecuadorian populations, and it may be distinct at least at the variety level. Therefore, the synonyms included for the type variety in Ollgaard (1 988) are not cited here. The holotype of Urostachys pilgerianus most likely is a mislabeled duplicate of Weberbauer 6626 (F, GH, MOL, us) from Dept. Junin, Nevado Runa- tullu, SE of Jauja, on rocks, 4400-4500 m, as this collection matches it exactly. Mislabeled (or re- labeled?) specimens abound in Nessel's herbari- um. The type and the Weberbauer collection both have creeping basal shoots and are rather densely tufted. The leaves are twisted unilaterally, but oth- erwise this corresponds to most of the Peruvian material. La Libertad: Prov. Santiago de Chuco, Jalca de Ques- nada (Quiruvilca to Huamachuco), 4000 m, Sagdstegui & Fabris 7574 (MO, NY). Prov. Santiago de Chuco, Dist. Calchicadan, Escalerilla-Conzuzo road near Tamboras, 3960 m, Saunders 882 (F, GH). Ancash: Prov. Carhuaz, Huascaran National Park, Quebrada Ishinca, 4380-4500 m, D. Smith 9480 (AUU). Prov. Carhuaz, Huascaran Na- tional Park, Quebrada Honda, 4300-4750 m, D. Smith et al. 11643 (F). Cord. Blanca, above Vicos, toward Le- jiacocha, 3600 m, Hutchison & Wright 4323 (F, GH, NY, uc). Prov. Huaylas, Huascaran National Park, Quebrada Alpamayo, 4750 m, D. Smith et al. 9715 (AUU, F). Junin: Mount La Juntay, near Huancayo, 4700 m, Killip & Smith 22114 (F, NY, us). Cuzco: Cerro Salamanca, Valle Lucumayo, 2200 m, Biies 569 (us). Prov. Urubamba, Abra de Malaga, 4300 m, Chavez 2822 (MO); 4 1 50-4230 m, Molau & Ohman 1639 (GB). Puno: "Cord, jugis pr. Tabina," Lechler 2043 (E). Cord, near Agapata and Sa- chapata, Lechler 2028 (BR, G, s). 1 9. Huperzia nesselii (Nessel) Roller! & Deferrari, NotasMus.LaPlata,Bot.21 (100): 156. 1988. Urostachys nesselii Nessel, Revista Sudamer. Bot. 6: 161. 1940 [Barlappgewachse 7 8, / 12. 1939, nom. inval.]. TYPE: "Ost Peru, Cuzco, 1868, Rein- hardt" (holotype, BONN, Herb. Nessel 132, in part!). Plants erect from a prostrate to ascending base, slightly to strongly heteroblastic, loosely to densely caespitose with prostrate-ascending basal shoots, these bearing numerous erect, fingerlike shoots in the center, up to 20 cm tall. Erect shoots homo- phyllous or almost so, equally thick throughout, 10-15 mm in diameter including leaves. Stems excluding leaves 2-6 mm thick, usually completely concealed by leaves. Leaves uniform throughout, borne in irregular, alternating whorls (or low spi- rals) of 5-6, densely crowded, closely imbricate, straight to irregularly curved upward or twisted at the apex, linear-lanceolate to lanceolate or nar- rowly elliptic, 9-12 mm long, 2-2.5 mm wide, without a prominent basal swelling, adaxially con- cave to slightly convex, with a raised veinal ridge near the base, abaxially flat especially at the base to rounded especially at the apex, with a broadly prominent medial ridge especially at the base, slightly rugose because of protruding, blisterlike epidermal cells, with smooth to slightly uneven margins, pruinose. Sporangia 2-3 mm wide. 34 FIELDIANA: BOTANY High Andine grassland near the timber line, 3000-3500 m, Junin. Endemic. The material referred to this species including the type seems to represent a single gathering, re- ferable to Weberbauer. The information on the type label is in Nessel's handwriting, and the geo- graphic information is dubious as are numerous other Nessel labels. The species seems most closely related to Hu- perzia saururus with respect to size and growth habit; however, it differs by the open growth of basal portions of the plant, and the ridged leaf bases, and smooth margins. From Peruvian H. crassa it differs by the large, rather flat leaves with- out a prominent basal air sac. Junin: Tarma, mountains W of Huacapistana, 3400- 3500 m, Weberbauer 2222 (G, MOL). "Lagasca ex herb. Swartz, ex Tarma Provincia in Peruvia, n. 2222" (BONN, Herb. Nessel 132 in part). 20. Huperzia andina (Rosenst.) Holub, Folia Geobot. Phytotax. 20: 70. 1985. Lycopodium andinum Rosenst., Repert. Spec. Nov. Regni Veg. 5: 239. 1908, not Herter 1909. TYPE: Bolivia, La Paz, Murusata, 5000 m, Buchtien 173 (holotype, s!). Urostachys andinus (Rosenst.) Nessel, Barlappge- wachse 76. 1939. Plants with prostrate, rooting shoots from which 1 to several, usually short, erect, fingerlike shoots arise, sometimes loosely caespitose, up to 10(-15) cm tall. Prostrate shoots densely covered with 8- 15 mm long, unilaterally upward curved, linear, red-tinged leaves. Erect shoots homophyllous or almost so, equally thick throughout or slightly ta- pering upward, ca. 6-10 mm in diameter including leaves. Steins of erect shoots excluding leaves 2- 3 mm thick, usually completely concealed by leaves. Leaves of erect shoots uniform throughout, or somewhat reduced upward, borne in close, ir- regular, alternating whorls of 4-5, usually closely imbricate, straight to unilaterally or irregularly twisted and often somewhat deformed, linear-lan- ceolate to lanceolate or triangular-lanceolate, evenly to abruptly tapering from the slightly wid- ened base or slightly acuminate, 5-8(-l 1) mm long, l-2(-2.5) mm wide, upward sometimes reduced to 4-6 mm long, with a prominent basal swelling (air sac), adaxially concave to canaliculate apical- ly, abaxially convex and rounded, smooth without protruding, blisterlike epidermal cells, with smooth to irregularly rugose, often strongly sclerified mar- gins, green to reddish-tinged. Sporangia 2-2.5 mm wide. In boggy grassland, open turfy puna, wet ground at lakes, alt. (3300-)4000-4700(-5000) m, Caja- marca, La Libertad, Ancash, Junin, Huancavelica, Cuzco, Puno. Peru and Bolivia. Some collections, especially from lower alti- tudes, are referred to this species with doubt. Leon & Young 1566, 1621 (AAU), from San Martin, Rio Abiseo National Park, alt. 3650 m, and Leon & Young 1105 (USM), from La Libertad, Prov. Pataz, Rio Abiseo National Park, Cueva de Manachaqui, 3600-3800 m, are smaller, more slender, and en- tirely green but share the growth habit with rela- tively large leaves on the creeping shoots and rath- er slender erect shoots. Cajamarca: Prov. Contumaza, Jalca El Chuno (Pozo Chuno), 4500 m, Sagdstegui et al. 9380 (AAU, F). La Libertad: Prov. Santiago de Chuco, Laguna La Victoria (road to Conzuzo), 4000 m, Sagdstegui et al. 6180 (GH, MO). Huillilas, N of Cachicadan, 4000 m, Stork & Norton 10003 (F, uc). Ancash: Prov. Yungay, Dist. Yungay, Lake Llanganuco, ca. 3500 m, Sounders 503 (F). Junin: Prov. Huancayo, Dist. Huancayo, Huaytapallana, Huancayo- Pariahuanca road, ca. 4480 m. Sounders 1151 (GH). Dist. Huancayo, ca. 28 km E of Huancayo, 4500 m, Tyron & Tryon 5469 in part (F, GH, us). Huancavelica: Prov. Ta- yacaja, Millpu, puna de Tocas, between Colcabamba and Paucarbamba, 4000 m, Tovar 1945 (USM). Marcapata, 3600 m, Stafford 1007 (F, K). Puno: Carabaya, Aricoma Lake, 4400 m, Stafford 1118 (K). 21. Huperzia saururus (Lam.) Trev., Atti Soc. Ital. Sci. Nat. 17: 249. 1874. Lycopodium saururus Lam., Encycl. 3: 653. 1789. TYPE: He de Bourbon (Reunion), Commerson (holotype, P, Herb. Law.!). Lycopodium elongatum Sw., Syn. Fil. 175. 1806. TYPE: Peru, Prov. Junin, Tarma, collector un- known (holotype, s!). Urostachys saururus (Lam.) Herter, Repert. Spec. Nov. Regni Veg. 19: 162. 1923. Urostachys elongatus (Sw.) Herter, Index Lye. 60. 1 949. Lycopodium sanctae-barbarae Rolleri, Darwiniana 16: 129,/ 1 D-E, t. 1, 3B. 1970. TYPE: Argentina, Prov. Jujuy, de la Sola 2883 (holotype, LP). Huperzia sanctae-barbarae (Rolleri) Rolleri and De- ferrari, Notas. Mus. La Plata, Bot. 21 (100): 156. 1988. Plants erect, very compactly caespitose with basal, prostrate, rejuvenating shoots from which TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 35 stiffly erect, sparsely branched, fingerlike shoots arise, at least up to 50 cm tall. Erect shoots homo- phyllous or almost so, usually narrow and some- what etiolated at the base and gradually becoming wider and green upward, 5-10 mm in diameter including leaves at the base, upward usually 8-15 mm. Steins excluding leaves (2-)3-5 mm thick, usually completely concealed by leaves except at the etiolated base. Leaves uniform throughout or gradually shorter upward in old shoots, often smaller, narrower, pale and irregularly appressed at the etiolated base of shoots, borne in irregular, alternating whorls (or low spirals) of 6-8, densely crowded, closely appressed, the sporophylls ap- pressed from a nearly perpendicular leaf base, straight or somewhat secund, linear to linear-lan- ceolate, the uppermost sometimes lanceolate to widely lanceolate, 10-12(-15) mm long, l-2(-3) mm wide, upward sometimes reduced to 5-8 mm long, sporophylls with a prominent basal swelling (air sac), adaxially concave to slightly convex, abaxially convex and rounded throughout or at least at the apex, smooth, with minutely rugose, thickened, sclerified margins, especially in the bas- al leaves, green and shining to reddish-tinged or rusty-colored and sometimes strongly pruinose. Sporangia 2-2.5 mm wide. Moist shrubland and grassland (jalca), often among rocks, alt. 3300—4500 m, Cajamarca, La Libertad, Ancash, Pasco, Junin, Huancavelica, Cuzco. Peru to Argentina and Chile; Kerguelen; Tristan da Cunha; alpine regions of tropical and southern Africa; Madagascar; Reunion; Mauritius. Contrary to the view of Rolleri (Amer. Fern J. 67: 109-120. 1977), the present author finds the Neotropical distribution of Huperzia saururus re- stricted to Peru and more southern regions in America. The Central American and north Andine material referred to H. saururus by Rolleri belongs to various different species. Some plants of this species are distinctly prui- nose and reddish-tinged, similar to Huperzia cras- sa, but can usually be distinguished by the leaf shape, the etiolated character of the base of the erect shoots, and the lack of blisterlike, protruding epidermis cells on the abaxial leaf surface. Cajamarca: Contumaza, Jalca El Chuno, 4500 m, Sa- gdstegui et al. 9559 (AAU, F, NY). Pozo Kuan, 3790 m, Sagdstegui et al. 13089 (AAU, F). Pozo Kuan, Laguna el Toro, 4100 m, Sagdstegui 9452 (F). La Libertad: Prov. Santiago de Chuco, Pampas de la Julia, 3600 m, Sagds- tegui et al. 11123 (AAU, GH). Jalca de Quiruvilca, 4200 m, Lopez 1506 (us). Ancash: Prov. Huari, Huascaran National Park, Quebrada Rima Rima, a lateral valley of Quebrada Carhuazcancha, 4200-4440 m, D. Smith et al. 1231 1 (AAU, F). Prov. Yungay, Huascaran National Park, Quebrada Ranicuray, 4000-4300 m, D. Smith et al. 9142 (AAU, F). Pasco: Pasco, Huayllay, ex herb. Cruck- shanks in 1830 (GH). Junin: San Jose, ca. 3960 m, Mac- bride & Featherstone 1 1 1 1 (F, G, us). Huancavelica: Prov. Tayacaja, above Hda. Tocas, between Colcabamba and Paucarbamba, 3200 m, Tovar 1967 (USM). Cuzco: Neva- do Sallcantay, 3900-4200 m, Biies 744 (us). Prov. Uru- bamba, Dist Chinchero, Cuper, Hatun Wayk'o quebra- da, 3300 m, Sallo ex Franquemont 281 (AAU, F). Prov. Paucartambo, Pfuyucalla?, Huilcacunca, 4000 m, Var- gas 9840 in part (MO). Madre de Dios: Pinasniocj, Pan- tiacolla, Pass, 3600 m, Cook & Gilbert 1867 (us). De- partment unknown: Peru, Rusby 354A (us). 22. Huperzia hypogaea B. 011g. in Harling and Andersson, Fl. Ecuador 33: 58. 1988. TYPE: Ecuador, Prov. Carchi, 0llgaard & Balslev 8517 (holotype, AAU!; isotype, QCA!). Plants with short or up to 36 cm long, horizon- tal, usually subterranean, isotomously branching shoots, from which stiffly erect, aerial shoot sys- tems arise. Subterranean divisions with pale ap- pressed leaves, 2-4 mm thick including leaves. Aerial shoots up to 40 cm tall including erect sub- terranean divisions, stiffly erect, homophyllous, equally thick throughout, or slightly tapering, 4- 6(-8) mm thick including leaves. Stems of aerial shoots 1.5-3 mm thick excluding leaves, partly to completely concealed by leaves, brownish to bright red, with smooth to somewhat rugose epidermis. Leaves borne in alternating, irregular whorls of 4- 5, arcuate-appressed to closely appressed, abaxi- ally convex and rounded or with a prominent, long decurrent veinal ridge, with or without a slightly prominent basal swelling (air sac), adaxially con- cave to slightly convex, linear-lanceolate to lan- ceolate, 4-6(-7) mm long, l-1.8(-2) mm wide, upward often reduced to 3.5-4 mm long, red-tinged to entirely red, with smooth to unevenly rugose and sclerified margins. Sporangia 1-1.3(-1.5) mm wide. Wet and boggy paramo and jalca, alt. 3000- 4300 m, Lambayeque. Colombia to northern Peru. Usually a very easily recognizable species of marshes and wet depressions of the lower paramos with a soft substrate where the subterranean run- ner-shoots are well developed. Where the species grows on more solid substrates, e.g., open soil in landslides and road cuts, supraterranean creeping shoots with well-developed, patent-ascending 36 FIELDIANA: BOTANY leaves replace the runner shoots, and make the distinction from slender forms of Huperzia crassa more subtle. In such cases the difference of leaf curvature, leaf size, and the number of leaf or- thostichies are helpful characters. Lambayeque: Prov. Ferranafe, Dist. Incahuasi, La- guna Tembladera to Cerro Negro, 3300 m, Sagdstegui et al. 12847 (AAU, F, GH). 23. Huperzia sagasteguiana B. 011g., sp. nov. Species cum habitu Huperziae hypogaeae et H. atten- uatae. A H. hypogaea differt surculis horizontalibus epi- gaeis vel non profunde hypogaeis, foliis brevioribus, marginibus denticulato-fimbriatis, epidermide abaxiali rugulosa cellulis protrusis pustuliformibus. A H. atten- uata differt foliis 1 0-fariis, epidermide abaxiali foliorum rugulosa, basibus foliorum minus ventricosis. Plants with up to 20 cm long, horizontal, epi- terranean and creeping, or shallowly subterranean, isotomously branching shoots, rooting along the underside, bearing stiffly erect, up to 20 cm tall aerial shoot systems with short to long intervals between branches. Creeping shoots with loosely appressed to upward curved, secund leaves, 3-5 mm thick including leaves. Aerial shoots stiffly erect, homophyllous, terete, equally thick through- out, 4-6 mm thick including leaves. Stems of aerial shoots 1.5-3 mm thick excluding leaves, partly to completely concealed by leaves, brownish to bright red, with rugosely striate epidermis. Leaves borne in alternating, irregular whorls of 5, arcuate-ap- pressed to closely appressed, abaxially strongly convex and rounded to apically indistinctly cari- nate, with a prominent, short to long-decurrent veinal ridge, with a somewhat prominent basal swelling (air sac), adaxially concave (dried), lan- ceolate to triangularly ovate-lanceolate, 3-5 mm long, 1.5-2 mm wide, red-tinged to entirely red, abaxially wrinkled (dried) and strongly uneven due to the protruding blisterlike epidermal cells, with numerous irregularly shaped and directed, soft and pale marginal processes. Sporangia 1-1.5 mm wide. TYPE— Peru, Dept. La Libertad, Prov. Pataz, Paso de Alaska, carretera a Tayabamba, en ladera abierta de Gramineas, Jalca, 3900 m, Lopez & Sagdstegui 8177 (holotype, MO; isotypes, AAU!, GH!, NY!). Terrestrial in jalca vegetation, 3900-3950 m, La Libertad. Endemic. The growth habit resembles Huperzia hypogaea, but the trailing stems are usually on or just below the surface of the ground, and the leaves are short- er, strongly pruinose, with many blisterlike cells on the abaxial leaf epidermis, and have densely denticulate-fimbriate margins. Huperzia attenu- ata is rather similar but is less distinctly hetero- blastic, has a smooth abaxial leaf epidermis, and has a more sharply prominent basal abaxial air sac on the upper leaves, and the leaves are borne in whorls of three to four. La Libertad: Prov. Pataz, between Retama and La Paz, 3950 m, Lopez & Sagdstegui 3590 (GH). 24. Huperzia attenuata (Spring) Trev., Atti Soc. Ital. Sci. Nat. 17: 249. 1874. Lycopodium attenuatum Spring, Mem. Acad. roy. Belg. 24 [Mon. Lye. 2]: 8. 1849. LECTOTYPE (des- ignated by Lellinger, Proc. Biol. Soc. Wash. 89: 717. 1977): Ecuador (Prov. Pichincha), in declivi- tate montis Pichincha, Hartweg 1470 (us!; isolec- totypes, BM!, GL!, K!, NY!, P!). Urostachys attenuatus (Spring) Nessel, Barlappge- wachse 101. 1939. Plants ascending to erect, rather small, caespi- tose with erect shoots in the center, and prostrate- ascending, rejuvenating basal shoots in the pe- riphery, up to 20 cm tall, or up to 30 cm long. Shoots homophyllous to slightly gradually hetero- phyllous, terete to hexagonal, equally thick throughout, 4-7 mm in diameter including leaves. Steins excluding leaves 1 .5-2 mm thick at the base, sometimes tapering to 1-1.5 mm in diameter up- ward, completely concealed by leaves, tinged with red or brick-red. Leaves borne in alternating whorls of 3—4, forming 6 or 8 usually regular ranks, ap- pressed to the stem throughout, in basal divisions lanceolate and convex abaxially, upward gradually changing to narrowly triangular-ovate with acute or slightly acuminate apex, abaxially strongly con- vex to subcarinate near the apex, with a prominent basal swelling (air sac), (4-)5-6 mm long, in basal divisions 1-1.5 mm wide, distally 1.5-2.2 mm wide, with fimbriate margins, tinged with red. Spo- rangia 1.5-2 mm wide. Exposed cushion vegetation in paramo, pioneer vegetation on landslides, road banks, etc., in the paramo zone, alt. 3400-3500 m, Cajamarca, La Libertad, San Martin. Costa Rica; Ecuador and Peru to Bolivia. An easily recognizable species, characterized by fimbriate leaf margins. Huperzia attenuata is a TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 37 close relative of H. tetragona (see the following for discussion). C'ajamarca: Prov. Cajamarca, N of Canal Cumbe Mayo, 3400-3500 m, Sanchez. V. & Molau 3746 (AAU). La Libertad: Prov. Huamachuco, Pallar-Huaguil, road to Tayabamba, 3400 m, Lopez & Sagdstegui 8148 (AAU, F, MO). San Martin: Prov. Mariscal Caceres, Rio Abiseo National Park, NW sector, Valle de Chochos, 3450 m, Leon 1872 (AAU, USM). Prov. Mariscal Caceres, Rio Abi- seo National Park, Pampa de Cuy, 3450 m, Leon & Young 1401A (USM). 25. Huperzia tetragona (Hooker & Grev.) Trev., Atti Soc. Ital. Sci. Nat. 17: 248. 1874. Lycopodium tetragonum Hooker & Grev., Icon. Fil. 1: t. 109. 1829. TYPE: Ecuador (Prov. Pichin- cha), Prope verticem montis Pichincha, Jameson (E!, K!). Lycopodium catharticum Hooker, Ann. Nat. Hist. 1 : 430-^3 1 , /. 14. 1 838. SYNTYPES: Ecuador (Prov. Azuay), "Asuay of the Equator," W. Turner (K!); Ecuador (Prov. Canar), from the mountains of Pillzhum, Jameson (K!). Lycopodium myrsinites Lam. var. minus Spring, Mem. Acad. roy. Belg. 15 [Mon. Lye. 1]: 29. 1842, based on Lycopodium catharticum Hooker and with the same type. Lycopodium tetragonum Hooker & Grev. var. patu- lum Spring, Mem. Acad. roy. Belg. 24 [Mon. Lye. 2]: 12. 1849, based on Lycopodium catharticum Hooker and with the same type. Urostachys tetragonus (Hooker & Grev.) Nessel, Bar- lappgewachse 135. 1939. Urostachys catharticus (Hooker) Nessel, Barlappge- wachse 135. 1939. Plants ascending to erect, rather small, caespi- tose with erect shoots in the center, and usually prostrate to ascending rejuvenating basal shoots in the periphery, up to 20(-30) cm tall. Shoots homophyllous, or gradually slightly heterophyl- lous, quadrangular and equally thick throughout, 2.5-3.5(-4) mm in diameter including leaves. Steins excluding leaves 1-2 mm thick. Leaves dec- ussate throughout, imbricate, concealing the stem, in basal divisions widely lanceolate to triangular- ovate, abaxially rounded and evenly decurrent, upward triangular, carinate at least in the upper half, somewhat decurrent or abaxially with a prominent basal swelling (air sac), 3.5-5(-7) mm long, 2-4 mm wide at the base, evenly tapering into an acute or slightly acuminate apex, orange to bright red-tinged at the margins throughout, with fimbriate margins. An occasional branch may be hexagonal or terete, with leaves borne in alter- nating whorls of 3. Sporangia 1.3-2.5 mm in di- ameter. Exposed cushion vegetation, pioneer vegetation on landslides, road banks, etc., in the lower part of paramos and jalca, 2750-3600 m, Cajamarca, La Libertad, San Martin, Huanuco, Cuzco. Columbia to Bolivia. The red or reddish, quadrangular, erect shoots make this species an easily recognizable one. In- complete material of some epiphytic species with quadrangular constricted shoots have been fre- quently misidentified as this species. They are readily separated because of the fimbriate leaf margins in Huperzia tetragona. Huperzia tetragona is closely related to H. at- tenuata. The occasional aberrant hexagonal or te- rete shoots in some specimens of//, tetragona are virtually indistinguishable from slender shoots of H. attenuata. The two species are also ecologically rather similar and are often found to grow inter- mixed. There are numerous examples of species in which the number of leaves in each whorl is variable in the same population. The difference of terete and quadrangular shoots in Huperzia atten- uata and H. tetragona may be due to such simple and taxonomically overrated variation within populations of one species. Cajamarca: Prov. Chota, Laguna Yahuarcocha, above Incahuasi, 3600 m, Sagdstegui et al. 12901 (AAU). Prov. Cajamarca, N of canal Cumbe Mayo, 3400-3500 m, Sanchez Vega & Molau 3747 (AAU). La Libertad: Pataz, Puerta del Monte, ruta de Huaylillas, 3200 m, Lopez & Sagdstegui 3450 (GH). San Martin: Prov. Mariscal Ca- ceres, Rio Abiseo National Park, Chochos valley, 3425 m, Young 3507 (AAU); 3300 m, Young & Leon 4878 (AAU). Huanuco: 3-6 mi NW of Mito, ca. 3350 m, Mac- bride & Featherstone 1923 (F, s, us). Playapampa, ca. 2750 m, Macbride 4890 (F, o, us). Cuzco: Paucartambo, Huillcacunca to Huaisampilla, 3800 m, Vargas 9859 (MO, uc). 26. Huperzia sellifolia B. 011g. in Harling and An- dersson, Fl. Ecuador 33: 68, / 12C. 1988. TYPE: Ecuador, Prov. Carchi, B. 0llg. & Bal- slev 8432 (holotype, AAU!). Plants erect or ascending, usually forming large, caespitose, homoblastic plants, often more than 25 cm tall, with up to 45 cm long shoots. Shoots homophyllous, almost equally thick throughout, (5-)6-8(-10) mm in diameter including leaves. Stems excluding leaves (2.5-)3—4 mm thick, al- most completely concealed by leaf bases, except in basal divisions. Leaves almost uniform throughout, usually densely crowded throughout, borne in rather regular alternating whorls of 4-5, loosely imbricate or ascending to patent or sharply 38 FIELDIANA: BOTANY reflexed and appressed to the stem, usually dis- tinctly sigmoid, widely lanceolate to ovate or el- liptic, acute or obtuse, (1.8-)2-2.6(-3.5) mm long, (1.2-)1.5-2(-2.4) mm wide, coriaceous, usually brightly red-tinged, abaxially convex to concave with prominent vein in basal part of lamina, rounded in the apex, with prominent, short to long- decurrent basal swelling (air sac), with usually slightly involute, narrowly sclerified, darker, smooth to slightly uneven margins, adaxially with slightly to distinctly prominent vein. Sporangia 1- 1.5 mm in diameter. Terrestrial in exposed wet paramos, alt. 3 1 50- 3650 m, Piura, Lambayeque, Amazonas, San Martin. Andes of southern Colombia to northern Peru. Piura: Purchased in the market of Sullana, Friedberg 3933c (GH). Lambayeque: Prov. Ferranafe, Dist. Inca- huasi, Laguna Tembladera, 3150 m, Sagdstegui et al. 12799 (AAU, F). Amazonas: Prov. Bagua, Cord. Colan NE of La Peca, ca. 3300 m, Barbour 3382 (AAU, MO, USM). San Martin: Prov. Mariscal Caceres, Rio Abiseo National Park, Paredones, 3650 m, Leon & Young 1564 (AAU). 27. Huperzia engleri (Herter) B. 011g., Opera Bot. 92: 169. 1987. Lycopodium engleri Herter, Bot. Jahrb. Syst. 43, Beibl. 98: 45. 1909. TYPE: Peru, Dept. Huanuco, Prov. Huamalies, mountains SW of Monzon, 3200- 3300 m, Weberbauer 3359 (holotype, B!; isotype, BONN, Herb. Nessel 17 3\, MOL!). Urostachys engleri (Herter) Nessel, Barlappgewachse 94. 1939. Plants erect or ascending, up to 1 0 cm tall. Shoots homophyllous, equally thick throughout, 8-10 mm in diameter including leaves. Stems excluding leaves 1-1.5 mm thick, densely papillate. Leaves well spaced at base, crowded upward, borne in alternating whorls of 4, patent to somewhat re- flexed, widely lanceolate to oblong-lanceolate, acute to obtuse, 4-5.5 mm long, 1.5-2 mm wide, slightly convex abaxially with slightly prominent vein, with evenly decurrent base, without a basal swelling, adaxially with a shallow groove along the vein, with minutely rugose margins. Sporangia 2-2.5 mm wide, nearly globose. Habitat information according to protologue: Bog at grass steppe interrupting the shrub for- mation. Endemic, Huanuco. Possibly related to Huperzia rufescens (Hooker) Trev. (Ecuador) and H. brevifolia but apparently adapted to more sheltered habitats. The sporangia are disproportionately large, thick, and volumi- nous and do not open as is otherwise usual in dried specimens. This, plus the rarity of the species, sug- gests hybridity, but there is no other evidence of hybridity, because the spores seem normally de- veloped. The collection by Jelski from the Nessel Herbarium needs to be confirmed; it was not found in Krakow. Department unknown: "Ost-Cordilere, Jelskr [pos- sibly Dept. Cajamarca, Prov. Cutervo] (BONN, Herb. Nes- sel 1731). 28. Huperzia brevifolia (Grev. & Hooker) Holub, Folia Geobot. Phytotax. 20: 71. 1985. Lycopodium brevifolium Grev. & Hooker, Hooker Bot. Misc. 3: 104. 1832. TYPE: Peru, in Herb. Lam- bert, Ruiz and Pavon (BM?; possible isotypes, FI, Herb. Webb; AAU photo ex FI!, o!). Urostachys rufescens (Hooker) Nessel var. brevifolius (Grev. & Hooker) Nessel, Barlappgewachse 103. 1939. Urostachys brevifolius (Grev. & Hooker) Herter, Index Lye. 54. 1949. Plants erect or ascending, stiff and robust, form- ing small to large, caespitose, homoblastic plants, often more than 25 cm tall. Shoots homophyllous, almost equally thick throughout, (5-)7-12 mm in diameter including leaves. Stems excluding leaves (3-)4-7 mm thick at the base, slightly tapering upward, usually concealed by leaf bases. Leaves almost uniform throughout, densely crowded, or sometimes more spaced in basal divisions, borne in alternating whorls of 4— 5, ascending or perpen- dicular to sharply reflexed, with straight to upward curved apex, widely ovate or triangular-ovate to widely suborbicular-cordate or triangular-cordate, acute to almost obtuse or mucronulate, 2-4 mm long, 2-4(-5) mm wide, stiffly coriaceous, green- or red-tinged, abaxially concave, with sharply prominent vein, with sharply prominent, some- what flattened, short-decurrent basal swelling (air sac), with slightly revolute, strongly sclerified, darker and somewhat translucent, slightly erose margins. Sporangia 1.5-2.5 mm wide. Terrestrial in wet paramos, jalca, alt. 2700-4 1 20 m, Piura, Lambayeque, Cajamarca, Amazonas, Huanuco, Pasco, Cuzco. Costa Rica; Colombia to Peru. The total distribution range is uncertain, as there TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 39 is considerable regionally characteristic variation in the material referred to Huperzia brevifolia. Populations from Colombia, southern Ecuador and Peru are slightly different and usually recognizable; perhaps they are worth taxonomic recognition. Huperzia brevifolia differs from H. rufescens and H. sellifolia by its larger size, thick stems, and stiff and almost prickly leaves and by the frequent lack of red coloration. Dudley 11109 deviates by its slender shoots with very short leaves. Piura: Purchased in the market of Sullana, Friedberg 3933e (GH). Lambayeque: Prov. Ferranafe, Dist. Inca- huasi, Laguna Tembladera, 3150 m, Sagdstegui 12792 (F, GH), 12848 (GH in part). Cajamarca: From Chiclayo market, Leon (AAU, USM). Amazonas: Prov. Chachapo- yas, Leimebamba-Chilchos trail, Boeke 2130 (AAU). Prov. Bagua, Cord. Colan NE of La Peca, 3300 m, Barbour 3381 (AAU, MO). Huanuco: Playapampa, 2700 m, Mac- bride 4476 (F, us). Cushi, trail to Tambo de Vaca, Bryan 678 (F, us). Pasco: Prov. Oxapampa, trail to summit of Cord. Yanachaga via Rio San Daniel, 3350-3620 m, D. Smith 7742 (AAU, USM). Junin: Prov. Tarma, mountains W of Huacapistana, 3400-3500 m, Weberbauer 2225 (MOL). Cuzco: Prov. Paucartambo, Paso del Gallo, 2900 m, Vargas 4279 in part (uc, us). Urubamba, Machu Picchu, Runcuraccay-Sayamarca pass, 4120 m, Peyton 288 (AAU). Prov. La Convention, summit ridge of Vil- cabamba, ca. km 28 along trail from Hda. Luisiana and Rio Apurimac, 3330-3410 m, Dudley 11109 (GH, us). 29. Huperzia hohenackeri (Herter) Holub, Folia Geobot. Phytotax. 20: 73. 1985. Lycopodium hohenackeri Herter, Bot. Jahrb. 43: Beibl. 98: 46. 1 909. TYPE: Peru, Dept. Puno, in summis Cordiller. jugis pr. Tabina, Lechler 2043 (B!, BR!, K, NY!, p, s!). Urostachys hohenackeri (Herter) Nessel, Barlappge- wachse 105. 1939. Plants erect or ascending, forming small to large, caespitose, homoblastic plants, up to 40 cm tall. Shoots homophyllous or with gradually smaller leaves upward, equally thick throughout, 10-13(- 1 5) mm in diameter including leaves, or tapering to (5-) 7-10 mm in diameter upward. Stems ex- cluding leaves 5-6 mm thick at the base, upward tapering to (2-)3— 4 mm thick, partly to completely concealed by leaves. Leaves borne in more or less regular, alternating low spirals or whorls of 5-6, often well spaced in basal divisions, densely crowded upward, those of basal non-sporangiate divisions ascending to spreading, widely triangu- lar-lanceolate or ovate-lanceolate, 4.5-5.5 mm long, 2.5-3 mm wide, with short to long, evenly decurrent leaf bases. Leaves of upper, sporangiate divisions patent to loosely imbricate, triangular- ovate to ovate or widely ovate, acute, 3-5 mm long, 2-3.5 mm wide, softly to firmly coriaceous, green- to red-tinged, adaxially concave, or slightly convex and with a prominent veinal ridge at the base, abaxially usually slightly concave at the base, above the base convex and rounded or with a slightly to strongly prominent veinal ridge, with a strongly prominent, narrow basal swelling (air sac), with upward curved apex, with usually strongly sclerified, irregularly rugose to minutely erose margins. Sporangia 1.7-2 mm wide. Terrestrial on banks, steep, mossy slopes in wet paramos, jalca, alt. 3400-4000 m, Cuzco, Puno. Colombia to Peru. Huperzia hohenackeri resembles most the large individuals of H. brevifolia, with which it shares the general aspect of size and growth habit. It dif- fers in the longer, more acute and usually ap- pressed leaves. Metcalf 30745 deviates by having recurved leaves throughout. Cuzco: Prov. Urubamba, trail Piajupata to Sayacmar- ca, 3600 m, Vargas 2911 (us). Near Wenner Gren ruins, 3400-3600 m, Metcalf 307 45 (uc in part). Prov. La Con- vencion, Alturas de Rio Calzada, Huadquina, ca. 3660 m, Bties 755 (us). Puno: Tabina, in summis Cordilleras, julio 1854, Hohenacker (BONN, Herb. Nessel 199). Cor- dillera near Tabina, Hohenacker 4204 (BONN, Herb. Nes- sel 199). Cord, near Ayapata and Sachapata, Lechler 2028 (E). Department unknown: Cordillera, 1902, Lan- doni (BONN, Herb. Nessel 199). 30. Huperzia hartwegiana (Spring) Trev., Atti. Soc. Ital. Sci. Nat. 17: 248. 1874. Lycopodium hartwegianum Spring, Mem. Acad. roy. Belg. 24 [Mon. Lye. 2]: 14. 1849. LECTOTYPE (designated by Ollgaard in Harling and Anders- son, Fl. Ecuador 33: 76. 1988): Colombia, Prov. Popoyan, Hartweg 1466 (K! annotated by Spring; isolectotypes, BM!, G!, LG!, NY!). Lycopodium caracasicum Herter, Hedwigia 49: 88, /. 3a. 1909. TYPE: Venezuela, Caracas, Jan. 1856, Gollmer (holotype, B!). Urostachys hartwegianus (Spring) Nessel, Barlappge- wachse 90. 1939. Urostachys caracasicus (Herter) Nessel, Barlappge- wachse 90. 1939. Huperzia caracasica (Herter) Holub, Folia Geobot. Phytotax 20: 71. 1985. Plants robust, recurved or pendulous, at least up to 30 cm long. Shoots homophyllous to grad- ually heterophyllous, 15-30 mm in diameter in- cluding leaves in basal divisions, sometimes ta- pering to 10-15 mm in diameter. Stems excluding 40 FIELDIANA: BOTANY leaves 2.5-3.5(-5) mm thick at the base, tapering to 1.5-2 mm. Leaves of basal divisions borne in often regular alternating whorls of 4-5, falcately patent-ascending to imbricate, radially directed to somewhat secund, almost completely covering the stem and sporangia, lanceolate, coriaceous, usu- ally shining, 14-23 mm long, 2.5-3.5 mm wide, widest below the middle to near the base, abaxially convex at the base, usually slightly concave toward the tip, with smooth, flat to somewhat involute margins, the vein slightly prominent at the base and apically obscure. Leaves of terminal divisions conform, or usually smaller, 7-10(-13) mm long, 1 .5-3 mm wide, often somewhat clasping with the base widened, softer and less shining. Sporangia 1.7-3 mm in diameter. Rupestral in montane forest, alt. 2400-3100 m, Ancash, Cuzco. Southern Mexico; Guatemala; Andes from Ven- ezuela to Peru. Huperzia hartwegiana is widely distributed and variable. The species is often epiphytic outside Peru, but in Peru all records are from rocky sub- strates. The variation seems partly correlated with habitat type. The species frequently occurs in sites that are too dry for the other Huperzia species. In such exposed rupestral sites, the plants initially grow erect, then recurved, and may develop rather short and rigid shoots only. Such forms were rec- ognized as Lycopodium caracasicum by Herter and others. In moist, shaded, epiphytic situations long, lax, pendulous forms are developed. Huperzia hartwegiana is related to H. taxifolia. Ancash: Prov. Bolognesi, Quero E of Huasta, Cerrate 2471 (USM). Cuzco: Prov. Cuzco, Cadena de Pactatayan, Vargas 17044 (GH). Prov. Urubamba, Ollantaytambo, 3050 m, Plowman & Davis 4741 (GH). Cusuchaca, 2400 m, Vargas 22971 (GH). Valle de Urubamba, Herrera 3423 (c). Valle de Urubamba, ca. 1 0 km from the village, Leon 446 (USM). Quebrada Las Penas, 3100 m, Chavez 3493 (MO). 31. Huperzia taxifolia (Sw.) Trev., Atti Soc. Ital. Sci. Nat. 17: 248. 1874. Lycopodium taxifolium Sw., Prodr. 138.1788. TYPE: A remounted specimen, marked as type, without original annotation, in the Regnellian type her- barium in s!, is possibly the holotype. Lycopodium passerinoides Kunth, in HBK., Nov. gen. sp. 1: 41. 1816. TYPE: Peru, prope Olleras et Aipate, alt. 747 hexp., Humboldt (holotype, P, Herb. Humb.l; isotypes, B, Herb. Willd. 194101, BONN, Herb. Nessel 160\). Lycopodium nitens Schlecht. & Cham., Linnaea 5: 623. 1830. TYPE: Mexico, in arboris vetustis prope Jalapam, Schiede & Deppe (holotype, B!; isotype, BM!). Huperzia passerinoides (Kunth) Trev., Atti Soc. Ital. Sci. Nat. 17: 248. 1874. Huperzia passerinoides (Kunth) Trev. var. nitens (Schlecht. & Cham.) Trev., Atti Soc. Ital. Sci. Nat. 17: 248. 1874. Lycopodium schwendeneri Herter, Bot. Jahrb. 43: Beibl. 98: 50. 1909. TYPE: Not designated. Mexico in- dicated as locus classicus by Herter 1923. Urostachys taxifolius (Sw.) Herter, Repert. Spec. Nov. Regni Veg. 19: 162. 1923. Phlegmariurus taxifolius (Sw.) Love & Love, Taxon 26: 324. 1977. Plants lax and pendent, or sometimes recurved from an erect and somewhat rigid stem base, up to 50(-70) cm long. Shoots usually gradually ta- pering from ca. 20-30 mm in diameter including leaves at the base, to 4-10 mm in distal divisions of fully developed plants, sometimes not, or only slightly tapering, rarely abruptly constricted. Stems excluding leaves 1.5-2 mm thick at the base, ta- pering to 1-1.5 mm. Leaves usually reduced and modified upward, borne in irregular alternating whorls of 3-5. Leaves of basal divisions spreading to ascending or somewhat appressed, often twisted at the base, narrowly lanceolate, widest in the low- er half, firmly herbaceous to subcoriaceous, 14- 23 mm long, 2-3 mm wide, almost flat or some- what concave adaxially, with flat or slightly rev- olute, smooth margins, with evident to somewhat prominent vein abaxially. Leaves of middle and terminal divisions usually gradually shorter, nar- rower and more appressed, abaxially more convex, with involute margins. Leaves of fully sporangiate divisions often distinctly 6-( 1 0-) ranked, with wid- ened, clasping base, partly covering the sporangia, often abruptly contracted into a short to long, nar- row, involute apex, 3-8 mm long, l-1.5(-2) mm wide. Sporangia 1-1.5 mm wide. Usually epiphytic in lower to mid-altitude wet montane forest, alt. 1300-1400 m, Cajamarca, Amazonas. Central America; West Indies; northern tropical South America. Huperzia taxifolia is a variable species, the de- limitation of which is problematic, especially out- side Peru. The Peruvian specimens cited below are unusual in having the leaves arranged in al- ternating whorls of 5. Hutchison & Bismarck 6379, unusual in the nearly conform sporophylls and uniformly patent-ascending leaves throughout, is apparently juvenile. TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 41 Cajamarca: Prov. Hualgayoc, Hda. Taulis, between the Casa Hacienda and Palmito, 2000 m, Hutchison & Bismarck 6379 (uc, USM). Amazonas: Prov. Bongara, vicinity of Campamiento Ingenio 1-3 km up road to Pomacocha (and Rioja) from camp along the Rio In- genio, 1300-1400 m, Hutchison & Wright 3825 (F, GH, uc). 32. Huperzia rosenstockiana (Herter) Holub, Fo- lia Geobot. Phytotax. 20: 76. 1985. Lycopodium rosenstockianum Herter, Hedwigia 49: 90. 1909. TYPE: Ecuador, Ost-Abhang des Tun- gurahua, 3000 m, Rimbach 130 (holotype, Herb. Rosenstock, not located; isotypes, s!, uc!, us!). Urostachys rosenstockianus (Herter) Nessel, Bar- lappgewachse 174. 1939. Plants lax, pendulous, rarely recurved from an ascending base, at least up to 1 50 cm long. Shoots homophyllous, or with gradually slightly smaller leaves upward, 20-25(-30) mm in diameter in- cluding the leaves in basal divisions, sometimes tapering to 10-14 mm. Stems excluding leaves (1 .5-)2-3(-5) mm thick at the base, tapering to 1- 2 mm. Leaves of basal divisions borne in alter- nating irregular whorls of 3-5, irregularly spread- ing to ascending, usually somewhat twisted at the base, with oblique to vertical lamina, lanceolate to narrowly oblong, widest at or below the middle, with widely cuneate to somewhat rounded base, with short to long acute apex, 10-15 mm long, 2- 4 mm wide, subcoriaceous and opaque to herba- ceous and translucent, almost flat, with prominent vein adaxially, or sometimes folded down along the vein, with slightly revolute, minutely rugose margins with individually protruding epidermis cells, especially at the apex. Leaves of middle and terminal divisions conform, or gradually smaller and more ascending to loosely appressed, more densely crowded. Leaves of densely sporangiate divisions usually widest just above the base, lan- ceolate to narrowly triangular, with a rounded base, 6-9 mm long, 1.5-2.5 mm wide, partly covering the sporangia. Sporangia 1.5-2 mm wide. Usually epiphytic, rarely terrestrial, in cloud for- est and elfin forest above 3000 m, San Martin. Colombia to northern Peru. Huperzia rosenstockiana resembles H. taxifolia, but details of the leaf margin, color, and ecology are quite distinct. Closely related to H. brong- niartii with which it shares the rugulate leaf mar- gins. Huperzia socratis (Herter) Rolleri & Defer- rari (type from Colombia) seems to be a synonym of the present species. San Martin: Prov. Mariscal Caceres, Rio Abiseo Na- tional Park, Puerta del Monte, 3400 m, Leon & Young 1303 (AAU, USM). Prov. Mariscal Caceres, Rio Abiseo National Park, Laguna de Chochos, 3300 m, Young & Leon 4855 (AAU). 33. Huperzia funiformis (Spring) Trev., Atti Soc. Ital. Sci. Nat. 17: 248. 1874. Lycopodium funiforme Spring, Bull. Acad. roy. Sci. Bruxelles 8 (2): 516. 1841. LECTOTYPE (des- ignated by Spring, Mon. Lye. 2: 49. 1949): Gua- deloupe, L'Herminier (P!; isolectotypes, BR!, GH!, NY!, uc!). Urostachys funiformis (Spring) Herter in Urban, Symb. Antill. 9: 387. 1925. Plants robust, flaccidly pendulous, ropelike, at least up to 250 cm long. Shoots homophyllous, almost equally thick throughout, 5-10(-15) mm in diameter including leaves, or sometimes taper- ing from a thicker base with patent-ascending leaves. Stems excluding leaves 1 .5-3(-5) mm thick at the base, ridged by decurrent leaf bases. Leaves almost uniform throughout, or slightly shorter up- ward, borne in crowded alternating whorls of 7- 8, densely covering the stem, usually closely fal- cate-appressed throughout, not twisted, linear-su- bulate, widest just above the base, 6-10(-12) mm long, 1-1.5 mm wide, evenly tapering into a long pungent apex, firmly herbaceous to coriaceous, dull to shining, abaxially strongly convex, often api- cally conduplicate, with smooth margins. Sporan- gia 1-1.5 mm wide. Lower montane rain forest, up to alt. ca. 1 300 m, Huanuco. West Indies; southern Mexico to Panama; Guy- ana; Venezuela to Peru. Huperzia funiformis is a very distinct species, not easily confused with other species due to its smooth ropelike shoots. Huanuco: SW slope of the Rio Llullapichis watershed, on the ascent of Cerros del Sira, 1290 m, Wolfe 12330 (GH). 34. Huperzia buesii (Herter) B. 011g., comb. nov. Urostachys buesii Herter, Revista Sudamer. Bot. 10: 126. 1954 [as 1953]. TYPE: Peru, Tranca de Ya- racuaya, 1800 m, Bues 742 (holotype, us!). Lycopodium buesii (Herter) Morton, Amer. Fern J. 54: 72. 1964. Plants robust, pendent, up to 50 cm long. Shoots homophyllous, almost equally thick throughout, 42 FIELDIANA: BOTANY ca. 10-20 mm in diameter including leaves, or tapering to 10 mm. Steins excluding leaves 2-3 mm thick, prominently ridged by decurrent leaf bases. Leaves almost uniform throughout, or slightly shorter distally, borne in close alternating whorls of 4-5, densely covering the stem, usually closely appressed throughout, slightly recurved at the apex, not twisted, linear to linear-lanceolate, widest just above the slightly rounded base, 12- 19 mm long, 1-2 mm wide, evenly tapering into a long apex, firmly herbaceous to subcoriaceous, shining, abaxially convex, somewhat involute, with smooth margins. Leaves of sporangiate divisions sometimes reduced to 6-8 mm. Sporangia 1.5-2 mm wide. A rare endemic. Epiphytic in forest, 1800 m, Cuzco. Perhaps related to Huperzia funiformis and H. aristei (Nessel) Rolleri and Deferrari (?Colombia) with which it shares the high number of leaves in each whorl and the general leaf shape. However, the leaves are longer and more diverging. The sim- ilarity to the juvenile plants of H. sotae (Rolleri) Rolleri and Deferrari (NW Argentina to Bolivia, e.g., Venturi 2955, isotype in us!) is close. Cuzco: Prov. La Convention, Choquellohuanca, 1 800 m, Marin 2247 (F). 35. Huperzia linifolia (L.) Trev., Atti Soc. Ital. Sci. Nat. 17: 248. 1874. Lycopodium linifolium L., Sp. pi. 1 100. 1753. TYPE: Plumier, Traite foug. Amer. /. 166, f. C, from an unspecified locality, either Martinique or His- paniola (see Proctor, Fl. Lesser Antilles 2: 26. 1977). Urostachys linifolius (L.). Herter, Repert. Spec. Nov. Regni Veg. 19: 154. 1923. Plants pendulous, usually with flaccidly hanging divisions, the terminal divisions often aggregated in fasciculate clusters, up to 60(-80) cm long. Shoots homophyllous or gradually heterophyllous, equal- ly thick throughout, 20-30(-45) mm in diameter including leaves, or gradually tapering to (7-) 1 0- 15 mm in diameter in terminal, densely sporan- giate divisions. Stems excluding leaves 0.5-1 mm thick at the base, slightly tapering upward, almost straight to somewhat flexuous, pale greenish in life. Leaves of basal divisions spirally arranged, single, or in occasional pairs or whorls of 3, not predom- inantly whorled, forming about 6 indistinct lon- gitudinal ranks, subdistant, soft-herbaceous, wide- spreading to ascending, straight to slightly falcate, usually with the lamina vertical due to a twist of the lamina base, linear-lanceolate to lanceolate, widest in the basal '/3 or '/4, distinctly narrowed into a petiolelike, twisted, usually perpendicular to deflexed lamina base, (10-) 13-24 mm long, 1- 5 mm wide, flat, or with slightly revolute, smooth margins. Leaves of middle and terminal divisions spirally arranged, paired or borne in irregular to regular, alternating whorls of 3, conform, or usu- ally narrower, (5-) 10- 15 (-20) mm long, (1-)1.5- 2(-3) mm wide, often with a long narrow apex from a subauriculate, non-twisted lamina base. Sporangia 1-1.5 mm wide. Southern Mexico to Panama; West Indies; northern South America. Huperzia linifolia is a widespread and poly- morphic species. Its variation is especially com- plex in the northern Andes. The two varieties keyed out below are fairly well characterized morpho- logically and are geographically well defined. Key to Varieties a. Shortest sporophylls of densely sporangiate divisions with distinctly widened, usually not twisted lamina base, usually ascending to loosely appressed, opaquely green; stem usually opaque-stramin- eous, usually above 1000 m elevation 35a. var. tenuifolia a. Shortest sporophylls of densely sporangiate divisions usually conform to leaves of basal divisions, or shorter, usually with a twisted, narrow to slightly widened lamina base, perpendicular to spreading- ascending, pale, transparently green or brownish green; stem, at least in terminal divisions, usually transparently stramineous to reddish brown, the vascular tissue usually visible through the cortex, usually below 600 m elevation 35b. var. jenmanii TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 43 35a. Huperzia linifolia var. tenuifolia (Nessel) B. 011g. in Harling and Andersson, Fl. Ecuador 33: 90. 1988. Urostachys linifolius (L.) Herter var. tenuifolius Nes- sel, Revista Sudamer. Bot. 6: 164. 1940 [Bar- lappgewachse 162, Abb. 39, / 14. 1939, nom. nud.]. LECTOTYPE (designated by 011gaard, in Harling and Andersson, Fl. Ecuador 33: 90. 1 988): Peru, Prov. Huanuco, Pampayacu, Poeppig ["Moritz 210 und 1155"] (BONN, Herb. Nessel 38 1\; isotype, G!). Wet forests of the eastern Andine slopes, alt. 100-2700 m, Cajamarca, Amazonas, San Martin, Huanuco, Pasco, Junin, Cuzco. Colombia?, Ecuador and Peru. Cajamarca: Tabillo, Jelski 1022 (KRA). Huancabamba, Andre Kl 763 (F, GH, NY, us). Tambillo, Jelski 1022 (KRA). Amazonas: Prov. Bongara, 5 km N of end of Lake Po- macocha, on road to Rioja, 2000 m, Hutchison & Wright 6791 (F, GH, NY, uc). Prov. Chachapoyas, Cerros Calla Calla, 10 km above Leimebamba, 2700 m, Hutchison & Bennett 4743 (F, GH, M, MO, NY, uc, us). San Martin: Tarapoto, monte Campana, Spruce 4627 (BR). Prov. Rio- ja, Pedro Ruiz-Moyobamba road, km 390, Venceremos, 1750 m, D. Smith 4372 (AAU). Prov. Rioja, Pedro Ruiz- Moyobamba road, km 29 1 , 1 850 m, Gentry et al. 45502 (AAU). Pasco: Prov. Oxapampa, Quebrada San Alberto, on the borders of Parque Nacional Yanachaga-Chemi- llen, Leon et al. 944 (F, USM). Junin: La Merced, Chan- chamayo, Soukup 1007 (F); 1000 m, Esposto 11978 (USM). Cuzco: Prov. La Convention, Potrero, Sapan Sachayoc, 2300 m, Vargas 13643 (GH). Valley of Santa Ana, 1000- 1 500 m, Herrera 2628 (us). Department unknown: Peru, Ruiz & Pavon (E, G). 35b. Huperzia linifolia var. jenmanii (Underw. & Lloyd) B. 011g. & Windisch, Bradea 5: 13. 1987. Lycopodium jenmanii Underw. & Lloyd, Bull. Torrey Bot. Club 33: 1 12. 1906. TYPE: Guyana, Monica River, Jenman (holotype, NY!; isotypes, BONN, Herb. Nessel 388 in part, E!). Urostachys jenmanii (Underw. & Lloyd) Nessel, Bar- lappgewachse 158. 1939. Huperzia jenmanii (Underw. & Lloyd) Holub, Folia Geobot. Phytotax. 20: 74. 1985. Epiphytic in lowland rain forest up to ca. 600 m, San Martin. Throughout the Amazonian basin and the Guianas. San Martin: Prov. Mariscal Caceres, Palo Blanco, W of Puente, Tocache Nuevo, J. Schunke V. 5738 (F, NY, s, us). 36. Huperzia wilsonii (Underw. & Lloyd) B. 011g., Opera Bot. 92: 170. 1987. Lycopodium wilsonii Underw. & Lloyd, Bull. Torrey Bot. Club 33: 111. 1906. TYPE: Puerto Rico, Luquillo Mountains, Wilson 271 (holotype, NY!). Lycopodium trichodendron Herter, Bot. Jahrb. 43: Beibl. 98: 49. 1909. SYNTYPES: Guadeloupe, L'Herminier (P!); Bory 103 (P!). Lycopodium andinum Herter, Bot. Jahrb. 43, Beibl. 98: 49. 1909, not Rosenst. LECTOTYPE (des- ignated by 011gaard in Harling & Andersson, Fl. Ecuador 33: 97. 1988): Ecuador, Eraser, in Herb. Drake (P!; isolectotype, G!). Lycopodium lindavianum Herter, Hedwigia 49: 90. 1909, nom. nov. for L. andinum Herter, not Ro- senst., and with the same type. Lycopodium stamineum Maxon, Smithsonian Misc. Coll. 56 (29): 2, pi. 2. 1912. TYPE: Panama, Chi- riqui, above El Boquete, ca. 1 750 m, Maxon 5636 (holotype, us!). Urostachys wilsonii (Underw. & Lloyd) Herter, Re- pert. Spec. Nov. Regni Veg. 19: 163. 1923. Lycopodium arcanum Maxon in Yuncker, Field. Mus. Publ. Bot. 17: 310, t. 3. 1938. TYPE: Honduras, Comayagua, above El Achote, above plains of Siguatepeque, 1800 m, Yuncker et al. 6149 (ho- lotype, us!). Huperzia lindaviana (Herter) Holub, Folia Geobot. Phytotax. 20: 74. 1985. Plants erect, arcuate-spreading to pendulous, up to 20(-30) cm long. Shoots homophyllous, 15- 2 5 (-30) mm in diameter including leaves, equally thick throughout, or in some slender individuals gradually tapering to 1—1.5 cm. Stems excluding leaves (l-)1.5-2(-3) mm thick at the base, often tapering to (0.7-)1 mm, prominently ridged by decurrent leaf bases, pale green to stramineous, often with bright red spots on leaf bases. Leaves usually uniform throughout, borne in alternating, irregular whorls of 6-7, in basal divisions, upward often in whorls of 4-5, perpendicularly spreading to ascending, straight to upward curved, usually not twisted at the base, linear to filiform, (6-) 10- 1 7 mm long, 0.3-0.5 mm wide at the base, quickly narrowed to ca. 0.2 mm due to involution, grad- ually tapering, adaxially canaliculate to involute, often with a prominent vein abaxially near the base. Leaves of terminal divisions in old plants sometimes gradually reduced to 6(-4) mm long. Decurrent leaf bases usually not wider than the lamina base, often bright red. Sporangia 1-1.5 mm wide. Epiphytic in lower montane forests, up to alt. 2750 m, San Martin. West Indies; southern Mexico to Panama; An- dean South America south to Peru. 44 FIELDIANA: BOTANY Huperzia wilsonii is related to H. dichotoma and H. mandiocana (Raddi) Trev. (Brazil) and to H. polycarpos. With these it shares the bottlebrush- like growth habit, and with H. mandiocana the bright red coloration of the leaf bases in some individuals. The absence or presence of red color seems uncorrelated with other characters. The in- tensity and size of the coloration are variable. Pendent individuals of H. wilsonii are rather similar to H. polycarpos, but the latter has shorter, more flattened leaves and uniformly falcate leaves due to a basal twist, and this species apparently is always pendulous. The direction of leaves is cor- related with growth habit. In erect-growing plants the leaves are wide-spreading to almost reflexed, while in pendent plants the leaves are somewhat ascending. San Martin: Prov. Rioja, Pedro Ruiz-Moyobamba road, km 390, Venceremos, 1750 m, D. Smith 437 2 A (MO). Prov. Mariscal Caceres, Rio Abiseo National Park, trail La Playa camp-Papayas, 2650-2750 m, Young & Leon 4968 (AAU). Department unknown: Ex herb. Pavon (o, P). Peru, M. Matthews (G, K). Peru, Matthews 1082 (o). 37. Huperzia polycarpos (Kunze) B. 011g., Opera Bot. 92: 169. 1987. Lycopodium polycarpos Kunze, Linnaea 9: 5. 1834. TYPE: Peru, ad Cassapi [Casapi?, Huanuco], Poeppig (w!). Urostachys polycarpos (Kunze) Herter, Index Lyco- podiorum 76. 1949. Urostachys cuatrecasasii Herter, Revista Sudamer. Bot. 10: 123. 1953. TYPE: Colombia, Valle, Cord. Occidental, Vertiente Occidental, hoya del rio Di- gua, Piedra de Moler, 900-1 180 m, Cuatrecasas 15143A (holotype, us!). Lycopodium cuatrecasasii (Herter) Morton, Amer. Fem J. 54: 72. 1964. Plants pendulous, with flaccidly hanging divi- sions, up to 30 cm long. Shoots homophyllous, almost equally thick throughout, 10-15 mm in diameter including the leaves. Stems excluding leaves 1-1.5 mm thick near the base, tapering to 0.5-1 mm, somewhat ridged by decurrent leaf bas- es. Leaves almost uniform throughout, densely crowded, borne in alternating, irregular whorls of 6—7 in basal divisions, upward of (3-)4 whorls, wide-spreading to ascending, often subsecund, usually with vertical lamina due to a twist at the base, linear to filiform, 6-10(-12) mm long, 0.4- 0.5 mm wide, evenly tapering, usually oblique- falcately curved, adaxially flat or shallowly cana- liculate, often with an abaxially prominent vein near the leaf base, with green decurrent leaf bases. Sporangia ca. 1 mm wide. Pendulous epiphyte in montane forest, ca. 2000 m, Huanuco?, Junin. Costa Rica to Colombia; Peru. Huperzia polycarpos is close to H. wilsonii, which see for discussion. From H. sarmentosa it is dis- tinguished by the lack of leaf base auricles. From H. mollicoma (Spring) Holub (tropical America), it differs in the more patent, twisted leaves, with much less prominent veins. Junin: Chanchamayo Valley, 2000 m, C. Schunke 529 (F). 38. Huperzia sarmentosa (Spring) Trev., Atti Soc. Ital. Sci. Nat. 17: 248. 1874. Lycopodium sarmentosum Spring, Mem. Acad. roy. Belg. 24 [Mon. Lye. 2]: 13. 1849. LECTOTYPE (designated by 011gaard in Harling and Anders- son, Fl. Ecuador 33: 100. 1988): Ecuador, Qui- tonian Andes, Jameson 41 (K!). Urostachys sarmentosus (Spring) Herter, Repert. Spec. Nov. Regni Veg. 19: 165. 1923. Plants slender, pendulous, with flaccidly hang- ing divisions, up to 100(-200) cm long. Shoots homophyllous or almost so, equally thick through- out, 15-25 mm in diameter including leaves in basal divisions, or sometimes tapering to ca. 8 mm in diameter. Stems excluding leaves 1-1.5 mm thick at the base, often tapering to 0.4-0.8 mm, slightly to prominently ridged by decurrent leaf bases. Leaves borne in alternating irregular whorls of 4-7, spreading to ascending, or rarely loosely appressed in terminal divisions, usually twisted at the lamina base, straight to slightly upward curved, linear to linear-subulate, with distinctly widened, auriculate, usually strongly revolute, often over- lapping lamina bases, 7-15 mm long, 0.4-0.7 mm wide just above the auriculate base, usually ca. 0.3 mm wide in the middle, usually somewhat revo- lute, with obscure to prominent vein, with smooth margins, or the auricles irregularly dentate. Leaves of terminal divisions often densely crowded, con- form, or gradually reduced to 5-7 mm long. De- current leaf bases narrower than the lamina base. Sporangia 1-1.3 mm in diameter. Epiphytic or rarely hanging from banks, in up- per montane forest, alt. 2100-3450 m, Amazonas, San Martin. Ecuador and northern Peru. TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 45 Huperzia sarmentosa is quite variable with re- spect to leaf length. It resembles H. polycarpos, and pendulous forms of//, wilsonii but differs from these by its auriculate lamina base. Amazonas: Chachapoyas, Leimebamba-Lajasbamba trail, Boeke 2059 (MO, us, USM). San Martin: Prov. Maris- cal Caceres, Rio Abiseo National Park, NW sector, Valle de Chochos, 3450 m, Leon 2161 (AAU, USM). 39. Huperzia curvifolia (Kunze) Holub, Folia Geobot. Phytotax. 20: 72. 1985. Lycopodium curvifolium Kunze, Linnaea 9: 5. 1835. TYPE: Peru (Dept. Huanuco), sylvar. densarum arboribus prope Pampayaco (Pampayacu), 1829, Poeppig(LZ, holotype, destroyed; isotypes, AWH!, B!, BR!, E!, oi, K!, M!, NY!, wi). Lycopodium tenue Willd. var. tenuissimum Spring, Mem, Acad. roy. Belg. 24 [Mon. Lye. 2]: 2 1 . 1 849, based on L. curvifolium Kunze and with the same type. Urostachys curvifolius (Kunze) Nessel, Barlappge- wachse 129. 1939. Plants delicate to extremely delicate, flaccidly pendulous, up to 60 cm long. Shoots gradually (rarely abruptly) heterophyllous, 2.5-6 mm in di- ameter including the leaves in basal divisions, ta- pering to 0.3-1.5(-2) mm in diameter including leaves in terminal divisions. Stems excluding leaves 0.4-0.7 mm thick at the base, tapering to 0.2-0.4 mm upward, usually not concealed by leaves. Leaves of basal divisions borne in alternating whorls of 4-5, patently sigmoid to strongly falcately up- ward curved, acicular-nliform, ong and widely de- current, 2-4(-5) mm long, 0.2-0.3 mm wide at the base, extremely delicate, abaxially convex, flat or canaliculate adaxially. Vegetative leaves of ter- minal constricted divisions alternate, or borne in alternating irregular whorls of 3, closely appressed, lanceolate to linear-lanceolate, somewhat clasping with the base, 1.2-3.5 mm long, 0.2-0.6 mm wide at the base, usually strongly involute, often sharply carinate at the base, short to long decurrent. Spo- rophylls usually few and scattered, often unilat- eral, conform, or shorter and wider, lanceolate to widely ovate, or subcordate and short to long acu- minate, usually widely spreading, 1-3 mm long, 0.3-0.8 mm wide, sometimes scarcely exceeding the sporangia, abaxially with a prominent vein, or carinate. Sporangia 0.6-0.8 mm wide. Epiphytic in lower, wet montane forest, alt. 400- 1750 m, San Martin, Ucayali, Huanuco. Costa Rica to Peru. Huperzia curvifolia is variable in size and com- pactness, seemingly not only as a response to en- vironmental factors. It is easy to recognize by the strongly upward curved leaves of basal divisions and the appressed non-sporangiate leaves of the terminal divisions. The smallest forms with hair- like terminal shoots represent the most fragile and slender extreme known in the family. Only these are represented in Peru. They correspond to the type collection. Other, stronger forms to the north in the Andes and in Central America approach slender forms of H. acerosa and H. filiformis. San Martin: Boqueron pass, 92 km from Tingo Maria on road to Pucallpa, 400 m, Allard 22098 (us). Prov. Rioja, Pedro Ruiz-Moyobamba road, km 390, Vencere- mos, 1750 m, D. Smith 437 2A (USM). Ucayali (as Lo- reto): Prov. Coronel Portillo, Boqueron Padre Abad, 400 m, J. Schunke V. 3046 (F, GH, us), 480 m, Hutchison et al. 6054 (F, GH, uc, us, USM). 40. Huperzia tenuis (Willd.) Trev., Atti Soc. Ital. Sci. Nat. 17: 248. 1874. Lycopodium tenue Willd., Sp. pi. ed. 4, 5: 55. 1810. TYPE: Ecuador, Valley of Vinajacu, near Loja, Humboldt & Bonpland 3363 (holotype, B, Herb. mild. 194231; isotypes, BM!, LG!, P!). Urostachys tenuis (Willd.) Nessel, Arch. Bot. Est. S. Paulo 1: 401. 1927. Plants delicate to extremely delicate, flaccidly penduolous, at least up to 75 cm long. Shoots grad- ually heterophyllous, 3-8 mm in diameter includ- ing the leaves in the basal divisions, tapering to 1-3 mm. Stems excluding leaves 0.5-0.7 mm thick at the base, tapering to ca. 0.3 mm upward. Leaves of basal divisions borne in irregular alternating whorls of 3-5, or spirally arranged, forming 6-10 indistinct longitudinal ranks, usually densely crowded, spreading to ascending, straight to some- what curved, often secund, acicular, tapering from a subauriculate base, 3-5(-7) mm long, 0.3-0.5 mm wide, decurrent. Vegetative leaves and spo- rophylls of constricted terminal divisions almost conform, borne in alternating irregular whorls of 3, or irregularly alternate, patently diverging from the stem, with upward curved tips, widely ovate to subhastate, 1-3 mm long, 0.5-1.4 mm wide, clasping at base, tapering to abruptly narrowed into a short to long acuminate apex, bluntly to sharply carinate, usually unilaterally, discontinu- ously, but often densely sporangiate. Sporangia 0.8-1.2 mm wide. 46 FIELDIANA: BOTANY Usually epiphytic, in high montane forests, alt. 1350-3300 m, Amazonas, San Martin, Cuzco. Costa Rica; Andes from Venezuela to Peru. Huperzia tenuis exhibits the same type of size variation as mentioned for H. curvifolia. Some forms are as extremely fragile and slender but dif- fer by the spreading leaves in terminal divisions. The typical forms of the species are also ecologi- cally distinct, being restricted to the uppermost montane forests, while the extremely thin forms (LI. Williams 7348 and Spruce 4621) appear to occur only at lower altitudes. Amazonas: Prov. Chachapoyas, Cerros Calla Calla, 1 8 km above Leimebamba on road to Balsas, 3000-3100 m, Hutchison & Wright 6932 (F, GH, MO, NY, uc, us, USM); Wurdack 1762 (F, G, GH, K, NY, s, uc, USM). San Martin: Tarapoto, Monte Campana, Spruce 4621 (BM, BR, P). Prov. Mariscal Caceres, NW corner of Rio Abiseo National Park, Puerta del Monte, 3100-3300 m, Young & Leon 4458 (AAU). Prov. Mariscal Caceres, Chochos valley, 3100 m, Young 2667 (AAU, USM). San Roque, 1 350-1 500 m, LI. Williams 7348 (F, us). Cuzco: Camino a Huarcan, caqui Acobamba [Ocobamba], Valle Neapi- llo, 2400 m, Biies 1376 (NY). Department unknown: Poeppigin 1829 (MO). Lehmann 5021 (F, GH, us). 41. Huperzia molongensis (Herter) Holub, Folia Geobot. Phytotax. 20: 75. 1985. Lycopodium molongenseHerter, Bot. Jahrb. 43: Beibl. 98: 51. 1909. TYPE: "Molong," Pearce (holo- type, K!) [erroneously as "Ostaustralische Pro- vinz: Gebirgswalder Neu-Siid- Wales" in the pro- tologue]. Urostachys molongensis (Herter) Nessel, Barlappge- wachse240. 1939. Plants robust to very robust, pendulous, at least up to 1 50 cm long. Shoots heterophyllous, usually not sharply differentiated, 20-35 mm in diameter including the expanded leaves in basal divisions, distally gradually to abruptly constricted to 3-8 mm in diameter including the shorter, imbricate leaves in the terminal divisions. Stems excluding leaves 2-3.5 mm thick at the base, tapering to 1- 1.5 mm. Expanded leaves of basal divisions borne in alternating whorls of 3, patent to ascending, or slightly recurved, lanceolate to lanceolate-ovate, acute, 10-18 mm long, 3-6(-7) mm wide, flat, or with slightly revolute margins, coriaceous, often shining, usually twisted at the base. Leaves of con- stricted divisions decussate or subdecussate, im- bricate, discontinuously sporangiate, ovate to sub- cordate, acute, sharply carinate and clasping with the base, usually conduplicate in the apex, usually shining, (2.5-)3-10 mm long, 3-6 mm wide. Spo- rangia 1.5-2.5 mm in diameter. Usually epiphytic, in montane, wet cloud forest on the eastern Andean slopes, 3100-3550 m, Amazonas, San Martin. Andes of Venezuela, south to northern Peru. This species is easily distinguished because of its large size and the sharply quadrangular ter- minal divisions. The expanded leaves are some- times developed only along a very short portion of the stem base and are sometimes lacking in herbarium specimens. Amazonas: Prov. Chachapoyas, Leimebamba, road to Balsas, 3240 m, Luteyn 11370 (NY). Cerros Calla Calla, 3450-3550 m, Wurdack 1707 (F, GH, NY, us); 3360 m, Hutchison & Wright 6998 (c, E, F, G, GH, K, M, MO, NY, s, uc, us). San Martin: Prov. Mariscal Caceres, NW corner of Rio Abiseo National Park, trail to Mirador, 3 100-3300 m, Young& Leon 4457 (AAU). Prov. Mariscal Caceres, Rio Abiseo National Park, Puerta de Monte, 3400 m, Leon & Young 1308 (AAU). Prov. Mariscal Ca- ceres, Rio Abiseo National Park, Chochos Valley, 3100 m, Young 2620 (AAU). 42. Huperzia campiana B. 011g., in Harling and Andersson, Flora of Ecuador 33: 109. 1988. TYPE: Ecuador, Prov. Loja, E of Nudo de Cajanuma, 0llgaard et al. 57861 (holotype, AAU!; isotype, QCA!). Plants rather robust, pendulous, at least up to 1 m long. Shoots heterophyllous, the basal divi- sions ca. (15-)20-30 mm in diameter including the expanded leaves, the distal quadrangular di- visions abruptly constricted to 2-4 mm thick in- cluding the imbricate, reduced leaves. Stems ex- cluding leaves 2-4 mm thick at the base, tapering to 0.5-1.5 mm, pale greenish or brownish. Ex- panded leaves of basal divisions usually uniform throughout, borne in alternating whorls of 3, spreading to perpendicular, lanceolate to widely lanceolate-ovate, usually widest below the middle, (7.5-)10-15 mm long, 2.5-4(-4.5) mm wide, often somewhat shorter near the contraction of the shoot, twisted at the base, with flat to revolute margins, herbaceous to subcoriaceous, dull to somewhat shining. Leaves of terminal constricted divisions decussate, or rarely in alternating whorls of 3 near the base of constricted divisions, usually sporan- giate throughout, appressed and clasping with their bases, bluntly to sharply carinate, widely ovate to triangular-ovate, short-acuminate or mucronu- late, 2-3 mm long, 1.5-2 mm wide, the sporo- TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 47 phylls equalling or slightly exceeding the sporan- gia. Sporangia 1.2-2 mm in diameter. Epiphytic in upper cloud forest and elfin forest, alt. 2400-3500 m, San Martin, Huanuco. Ecuador; Peru. Bryan 779 deviates from typical Huperzia cam- piana by the relatively wide expanded leaves and slightly thicker constricted divisions. San Martin: Prov. Mariscal Caceres, Rio Abiseo Na- tional Park, NW sector, Valle de Chochos, 3450 m, Leon 2179 (AAU). Huanuco: Cushi, trail to Tambo de Vaca, Bryan 7 19 A (F, G, us). stricted divisions and is tentatively referred to this species. Amazonas: Leimebamba, 2100 m, Woytkowski 7749 (F, MO, NY, USM). 44. Huperzia ericifolia (Presl) Holub, Folia Geo- bot. Phytotax. 20: 72. 1985. Lycopodium ericaefolium Presl, Reliq. haenk. 1: 77. 1825. TYPE: "Peruvia, verosimiliter Luzon," Haenke (holotype, PRC!; isotype, BR!). Urostachys phlegmaria (L.) Herter var. ericaefolius (Presl) Nessel, Barlappgewachse 217. 1939. 43. Huperzia heteroclita (Poiret) Holub, Folia Geobot. Phytotax. 20: 73. 1985. Lycopodium heteroclitum Poiret, in Lam., Encycl. 3: 554. 1814 [1813]. LECTOTYPE (designated by 011gaard in Harling and Andersson, Fl. Ecuador 33: 1 12. 1988): America calidiore, Herbier de A. N. Desvaux (P!). Plants pendulous, lax, at least up to 80 cm (pos- sibly to 400 cm) long. Shoots heterophyllous, with expanded, perpendicular leaves, often only in the first 5-20 cm of the basal divisions, upward abruptly constricted, sharply to bluntly quadran- gular, 2-3 mm thick including the small, imbricate leaves, the constricted divisions often interrupted by short zones with expanded leaves. Stems ex- cluding leaves 1-1.7 mm thick at the base, tapering to 0.7-1 mm, often bright red. Expanded leaves decussate, or subdecussate, perpendicular to the stem, lanceolate to widely lanceolate, widest in the middle, acute, 6-9 mm long, 2-3.5 mm wide, soft- herbaceous, usually twisted at the base. Leaves of constricted divisions decussate, closely imbricate, usually discontinuously sporangiate, ovate-cor- date, acute, bluntly to sharply carinate to condu- plicate in the apex, 1.5-4 mm long, 2-2.5 mm wide, dull, sometimes with shallowly erose, or in- distinctly dentate margins. Sporangia ca. 1.5 mm wide. Epiphytic in upper wet montane forest, ca. 2 100 m, Amazonas. Southern Ecuador and northern Peru. The species is usually recognizable by its long, sharply quadrangular constricted shoots with ex- tensive non-sporangiate zones and sharply cari- nate, apically conduplicate leaves. However, the specimen cited below deviates from the typical in the terete rather than sharply quadrangular con- Plants slender, pendulous, at least up to 6 cm long. Shoots heterophyllous, in the basal divisions ca. 8-14 mm in diameter including the expanded leaves, then abruptly constricted to 1.5-2 mm in- cluding the imbricate, reduced leaves of the qua- drangular, terminal divisions. Stems excluding leaves ca. 1 mm thick, tapering to ca. 0.5 mm, pale greenish. Expanded leaves usually uniform in position, shape and size throughout, borne in al- ternating whorls of 3, or decussate upward, almost continuously overlapping throughout (pressed specimens), widely lanceolate to oblong-lanceo- late, acute to obtuse and mucronate, 7-9 mm long, 2-3 mm wide, with slightly revolute margins, the lamina usually twisted at the base to a vertical position. Leaves of terminal, constricted divisions decussate, discontinuously sporangiate, appressed and clasping with their bases, carinate to condu- plicate in the apex, lanceolate to ovate- or trian- gular-ovate, short to long acuminate, 2-4 mm long, 1.5-2 mm wide, the sporangiate leaves 1.5-2.5x longer than the sporangia. Sporangia 1-1.5 mm wide. Wet lower montane forest, alt. 625-2040 m, San Martin, Huanuco. Ecuador to Bolivia. Huperzia ericifolia is closely related to H. di- chaeoides (Maxon) Holub (Central America to Ec- uador) and H. aqualupiana (West Indies, N Ven- ezuela to NW Colombia). Its distinctness from the latter is doubtful, although H. ericifolia can usually be distinguished by its narrower, more acute ex- panded leaves in whorls of three, while H. aqualu- piana usually has ovate, subacute to obtuse, dec- ussate expanded leaves. Problems of typification were discussed by 011- gaard in Harling and Andersson, Fl. Ecuador 33: 113. 1988. 48 FIELDIANA: BOTANY San Martin: Prov. Rioja, Pedro Ruiz-Moyobamba road, km 390, Venceremos, 2040 m, Smith & Vdsquez 4589 (AAV, USM). Jingo Maria, 625-1 100 m, Allard20935 (us). Huanuco: SW slope of Rio Llullapichis watershed, ascent of Cerros del Sira, 1290 m, Dudley 13029 (GH, us). 45. Huperzia myrsinites (Lam.) Trev., Atti Soc. Ital. Sci. Nat. 17: 249. 1874. Lycopodium myrsinites Lam., Encycl. Meth. Bot. 3: 654. 1789. TYPE: Dominican Republic, "S. Do- mingue," Comm. Joseph Martin, (holotype, P, Herb. Lam.l). Urostachys myrsinites (Lam.) Herter, Repert. Spec. Nov. Regni Veg. 19: 166. 1923. Lycopodium skutchii Maxon, Proc. Biol. Soc. Wash. 46: 159 (1933). TYPE: Guatemala, Chimaltenan- go, Chichavac, alt. 2400-2700 m, Skutch 243 (ho- lotype, us!). Plants pendulous, slender, at least up to 65 cm long. Shoots heterophyllous, usually not all sharp- ly differentiated, the basal divisions 10-18 mm in diameter including the expanded leaves, distal di- visions gradually or abruptly constricted to 1.5-3 mm thick including the reduced, imbricate leaves. Stem excluding leaves ca. 1 mm thick at the base, tapering to ca. 0.5 mm, pale greenish or reddish. Expanded leaves of basal divisions decussate or subdecussate, often irregularly shaped, subdistant to densely crowded and somewhat overlapping, ascending to perpendicular, ovate-lanceolate or lanceolate, acute, usually the widest ones with a rounded base, 6-11 mm long, 1.5-3 mm wide, usually flat, straight to somewhat recurved. Leaves of terminal constricted divisions highly variable, often with complete reduction series, and with re- current series to expanded shape, decussate or sub- decussate, continuously or discontinuously spo- rangiate. Transitional leaves with widely ovate base, and short to long acuminate apex, appressed and clasping, with the wide base abaxially rounded to bluntly carinate, with straight to recurved apex, 2.5-5 mm long, 1.5-2 mm wide. Shortest leaves with base conform, but with straight to falcate apex, bluntly to sharply carinate, scarcely exceed- ing the sporangia, ca. 2 mm long. Sporangia 1-1.3 mm in diameter. Epiphytic in montane forest, ca. 2500 m, Ca- jamarca. Hispaniola; Central America; Trinidad; Guy- ana; Venezuela to Peru. For illustration of the species, see A. R. Smith (Pteridophytes, in Breedlove: Flora of Chiapas, part 2: 347,f.82a-b. 1981). Huperzia myrsinites is closely related to H. phy- licifolia. Species of the group of Huperzia quad- rifariata (Bory) Rothm. (SE Brazil) were errone- ously placed under H. myrsinites by several earlier authors. Cajamarca: Prov. Hualgayoc, Hda. Taulis, 2500 m, Hutchison & Bismarck 6391 (F, NY, us). Prov. Contu- maza, Bosque de Cachil, 2450 m, Sagdstegui 14873 (AAV). 46. Huperzia phylicifolia (Poiret) Holub, Folia Geobot. Phytotax. 20: 75. 1985. Lycopodium phylicifolium Poiret, in Lam., Encycl. 3: 546. 1814 [1813]. LECTOTYPE (designated by 011gaard in Harling and Andersson, Fl. Ecuador 33: 117. 1988): "Habitat in Chili," Herbier de A. N. Desvaux, Donnee par Mme Vve Lavallee en 1896(p!). Urostachys phylicifolius (Poiret) Nessel, Barlappge- wachse 246. 1939. Lycopodium congestifolium Spring, Mem. Acad. roy. Belg. 15 [Mon. Lye. 1]: 70. 1842. TYPE: Peruvia, Dombey, H. M. P., Herb. Deless. (P!). Lycopodium nubigenum Herzog, Meded. Rijks-Herb. 27: 2. 1915. TYPE: Bolivia, Comarapa, Herzog 1967 (holotype, L!; isotypes, B!, z!). Urostachys nubigenus (Herzog) Nessel, Barlappge- wachse 246. 1939. Plants slender, pendulous, up to 150 cm long. Shoots heterophyllous, in the basal divisions ca. 10-20(-25) mm in diameter including the ex- panded leaves, then abruptly (rarely gradually) constricted to (l-)1.5-2(-2.5) mm in diameter in- cluding the imbricate, reduced leaves. Stems ex- cluding leaves 0.7-1.2 mm thick at the base, up- ward tapering to ca. 0.5 mm, greenish to bright red. Expanded leaves of basal divisions borne in alternating whorls of 3, or decussate, subdecussate, or alternate, usually widely spaced in alternate- leaved stem portions, perpendicular to the stem to falcately ascending, lanceolate to linear-lanceo- late, widest at or below the middle, (4-)6-10(-13) mm long, (1-)1 .5-2 mm wide, softly to firmly her- baceous, with flat to slightly revolute margins, the lamina twisted to a vertical position. Expanded leaves near the constriction often sporangiate. Leaves of constricted terminal divisions decussate, or subdecussate, continuously or discontinuously sporangiate, appressed and clasping with their bas- es, abaxially rounded to carinate, widely lanceo- late to widely ovate or subcordate, acute to mu- cronate or cuspidate, rarely with an elongate flat apex, 1.7-2(-4) mm long (in gradually hetero- phyllous shoots sometimes longer), 1-1.5 mm wide, TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 49 equalling to more than twice as long as the spo- rangia. Sporangia 1.2-1.5 mm wide. Epiphytic or rarely rupestral in upper montane and elfin forest, 2300-3700 m, Cajamarca, Ama- zonas, Huanuco, Huancavelica, Apurimac, Cuzco, Puno. Due to uncertain delimitation of the species the total distribution of this species is unsettled. In an inclusive sense it is wide-ranging: from Costa Rica, along the Andes south to northern Argentina, and SE Brazil. Huperzia phylicifolia occurs in a wide range of epiphytic and rupestral habitats with highly vari- able exposure and humidity. Its growth habit (i.e., size, leaf size, texture and crowding, and stem col- or), responds strongly to the environmental con- ditions. It is closely related to H. myrsinites, H. subulata, H. biformis (Hooker) Holub (Brazil), and H. erythrocaulon (Fee) Holub (Brasil). Forms of the species corresponding to the type of Lycopo- dium nubigenum, with expanded leaves slightly longer and widest just above the base, are preva- lent in the southern part of the range (Peru, Bo- livia, Argentina). Cajamarca: Prov. Cutervo, Jelski 1013 (p). Amazonas: Chachapoyas, Matthews 1081 (G, K, p). Huanuco: Cani, NE of Mito, 2800 m, Macbride 3546 (F, us). Cushi, trail to Tambo de Vaca, Bryan 719B (F, GH, us). Huancaveli- ca: Prov. Tayacaja, Marcavalle, between Huachocolpa and Tintay, 2700 m, Tovar 4751 (GH, USM). Apurimac: Prov. Abancay, Ampay, 3400-3700 m, Vargas 8388 (MO). Cuzco: Prov. La Convencion, Dist. Vilcabamba, Yu- paanqui, 2700 m, Davis et al. 1210 (AAV, F, GH). Prov. Paucartambo, Paso de Aguila, Pillawata, 2450 m, Vargas 22995 (GH). Puno: Prov. Sandia, 2300 m, Weberbauer 633 (G, MOL). Prov. Carabaya, Ayapata-Kahuallayoc, 2600-3600 m, Vargas 10741 (GH). Tabina, Lechler2023 (us). 47. Huperzia subulata (Poiret) Holub, Folia Geo- bot. Phytotax. 20: 77. 1985. Lycopodium subulatum Poiret, in Lam., Encycl. 3: 544. 1814 [1813]. LECTOTYPE (designated by Ollgaard in Harling and Andersson, Fl. Ecuador 33: 1 19. 1988): HerbierdeA. N. Desvaux, Donnee par Mme Vve Lavallee en 1896 (P!). Urostachys subulatus (Poiret) Nessel, Arch. Hot. Est. S. Paulo 1: 420. 1927. Urostachys ewanii Herter, Revista Sudamer. Bot. 10: 126-127. 1953. TYPE: Colombia, drainage of upper Rio Pascual, above Cordoba, 2840 m, Ewan 16526 (holotype, us!; isotype, NO!). Lycopodium ewanii (Herter) Morton, Amer. Fern J. 54: 72. 1964. Plants slender, pendulous, up to 2 m long. Shoots heterophyllous, ca. 15-25 mm in diameter in- cluding the expanded leaves in the basal divisions, abruptly constricted to 1.5-2 mm in diameter in- cluding the reduced and appressed leaves of the terminal divisions. Stems excluding leaves ca. 1 mm thick at the base, upward tapering to less than 0.5 mm, pale greenish to bright red. Expanded leaves of basal divisions borne in alternating, often irregular whorls of 3, or often upward subdecus- sate, spreading to perpendicular to the stem, nar- rowly lanceolate to linear-subulate, straight to obliquely falcate-ascending, (6-) 10- 15 mm long, 0.5-1 mm wide, twisted at the base, flat, soft-her- baceous. Leaves of terminal constricted divisions subdecussate, or in alternating whorls of 3 just above the constriction of the shoots, sporangiate almost throughout, closely imbricate, widely ovate with obtuse to acute apex, abaxially rounded to carinate, 1.3-2 mm long, 1.3-1.5 mm wide, equalling or slightly exceeding the sporangia. Spo- rangia ca. 1 mm wide. Epiphytic, in cloud forest and elfin forest, alt. 1 500-3400 m, Amazonas, San Martin, Huanuco, Cuzco. Costa Rica; Colombia to Peru. Huperzia subulata is closely related to H. phy- licifolia but seems to tolerate a narrower range of growth conditions, being restricted to the most humid forests of the eastern slopes of the Andes. Amazonas: Prov. Bagua, Serrania de Bagua, ca. 12-18 trail km E of La Peca, 1 800-1 950 m, Gentry et al. 22904 (AAV, MO, USM). San Martin: Prov. Mariscal Caceres, Rio Abiseo National Park, Puerta del Monte, 3400 m, Leon & Young 1340 (AAU). Huanuco: Pampayacu, Hda. at mouth of Rio Chinchao, Bryan 725 (F, G, us). Cuzco: Prov. La Convencion, Cord. Vilcabamba, Hda. Lu- isiana-Rio Apurimac, 3250 m, Dudley 11 105 (F, GH, us, USM). Prov. La Convencion, Chontapampa, Valle San Miguel, 1500 m, Bties 2987 (GH). 48. Huperzia cuneifolia (Hieron.) Holub, Folia Geobot. Phytotax. 20: 72. 1985. Lycopodium cuneifolium Hieron., Bot. Jahrb. Syst. 34: 572. 1905. LECTOTYPE (designated by Nessel, Barlappgewachse 252. 1939): Costa Rica, Volcan Barba, Hoffman 50 (B!). Urostachys cuneifolius (Hieron.) Nessel, Barlappge- wachse 252. 1939. Plants pendulous, delicate, up to 60 cm long. Shoots heterophyllous, in the basal divisions (5-)7- 1 3 mm in diameter including the expanded leaves, 50 FIELDIANA: BOTANY then abruptly constricted to 1-2 mm in diameter including the reduced, imbricate leaves in the ter- minal, constricted, quadrangular divisions. Stems excluding leaves 1 mm thick, or less, at the base, tapering to ca. 0.5 mm, greenish to bright red. Expanded leaves of basal divisions decussate, spreading to perpendicular or slightly reflexed, spathulate to lanceolate or lanceolate-obovate, usually widest above the middle, obtuse or mu- cronulate, 2-6.5 mm long, 1.5-3 mm wide, flat, the lamina twisted to a vertical position, softly to firmly herbaceous. Leaves of terminal constricted divisions decussate, often sporangiate in rather short zones of the divisions, appressed and clasp- ing with their bases, widely ovate to subcordate, 1.5-2 mm long, 1-1.5 mm wide, carinate, with falcate to reflexed mucronulate tips, the sporo- phylls equalling or slightly exceeding the sporan- gia. Sporangia ca. 1 mm wide. Epiphytic or occasionally on banks in upper montane forest and elfin forest, ca. 3000-3100 m, Amazonas and San Martin. Costa Rica; Andean Venezuela; Colombia and Peru. Amazonas: Prov. Chachapoyas, Cerros Calla Calla, 19 km above Leimebamba, 3100 m, Hutchison & Wright 6943 (F, GH, K, M, P, us); Wurdack 1744 (F, GH, NY, us, USM). San Martin: Prov. Mariscal Caceres, NW sector of Rio Abiseo National Park, near Mirador, 3000-3100 m, Leon 27/9(AAu). 49. Huperzia pruinosa (Herter) Holub, Folia Geo- bot. Phytotax. 20: 76. 1985. Lycopodium pruinosum Herter, Bot. Jahrb. Syst. 43, Beibl. 98: 52. 1909. TYPE: Peru, Dept. Ama- zonas, Prov. Chachapoyas, Tambo Ventillas, 2400-2600 m, Weberbauer (err. as Ule in pro- tologue) 4410 (holotype, B!; isotypes, BONN, Herb. Nessel 62 l\,o\). Lycopodium durissimum Herter, Bot. Jahrb. Syst. 43, Beibl. 98: 52. 1909. TYPE: Peru, "Voyage a FEquateur et au Perou, 1876-77, Guayaba mars 1877" (err. "Colombia, Guayabal" in proto- logue), Vidal-Seneze ["Senege"] (holotype, P!; iso- type, BONN, Herb. Nessel 6221). Urostachys pruinosus (Herter) Nessel, Arch. Bot. Sao Paulo 1:420. 1927. Urostachys durissimus (Herter) Nessel, Barlappge- wachse 240. 1939. Huperzia durissima (Herter) Holub, Folia Geobot. Phytotax. 20: 72. 1985. Plants erect and distally recurved, unbranched or sparsely branched at base, distally with densely tassel like ramification, at least up to 70 cm long. Shoots heterophyllous, the basal divisions with expanded leaves ca. 10-18 mm in diameter in- cluding leaves, distally abruptly constricted to ca. 2-2.5 mm including the reduced, imbricate leaves. Stems excluding leaves up to 5 mm thick at the base, upward tapering to ca. 0.5-1.5 cm, rigid, dark brown to purplish brown. Expanded leaves of basal divisions borne in alternating, irregular, distant whorls of 4, the whorls 6-9 mm apart, ascending to spreading or sharply reflexed, lan- ceolate, straight or recurved, (6-)8-10 mm long, 2.5-3 mm wide, acute, flat, coriaceous, with smooth, revolute margins, often adaxially con- cave. Leaves of recurved, constricted divisions sub- decussate, densely crowded, sporangiate almost throughout, imbricate, widely ovate with obtuse to acute apex, abaxially rounded to carinate, 1.2- 1.6 mm long and wide, equalling or slightly ex- ceeding the sporangia. Sporangia 1-1.3 mm wide. Indicated as terrestrial scandent shrub, in ceja scrub (protologue) and as an epiphyte in high Swie- tenia trees (Nessel, in Hoehne, Fl. Bras. 11: 88. 1955). Endemic, Amazonas and San Martin. The two taxonomically synonymous basionyms were published simultaneously. The epithet prui- nosa is maintained due to the quality of its type material and greater certainty of its origin. San Martin: Mount Organero, ceja, 1900 m, Melin 96 (s). "Tarapoto, 1908, G. Huebner" (BONN, Herb Nessel 6211). Comments Huperzia polyclada (Sodiro) Rolleri and Deferrari, Notas Mus. La Plata, Bot. 21(1 00): 156.1988. Lycopodium polycladum Sodiro, Crypt, vase. Quit. 561.1893. TYPE: Ecuador, Mojanda, Sodiro (not located). The type locality in Ecuador contains several taxa of the Huperzia crassa group. Due to the ab- sence of authoritative material and the relatively scarce information of the protologue it is not pos- sible to certify to which of these taxa the name applies. However, none of the taxa correspond to the Peruvian material that was placed in this spe- cies by Rolleri (1980). This material is here re- ferred to H. darwiniana. TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 51 II. Lycopodium Lycopodium L., Sp. pi. 1100. 1753. TYPE: Ly- copodium clavatum L. Figure 7. Lepidotis Mirbel, in Lam. & Mirbel, Hist. nat. veg. 3: 477. 1802. TYPE: Lycopodium clavatum L. Lycopodium subgen. Lepidotis Baker, Handb. Fern- Allies 8. 1887. TYPE: Lycopodium clavatum L. Lycopodium subgen. Rhopalostachya Pritzel, Nat. Pflanzenfam. 1 (4): 601. 1901. TYPE: Lycopo- dium clavatum L. Diphasiastrum Holub, Preslia 47: 104. 1975. TYPE: Lycopodium complanatum L. Diphasium Rothm., Feddes Repert. Spec. Nov. Regni Veg. 54: 64. 1944. TYPE: Diphasium jussiaei (Poiret) Rothm. (= Lycopodium jussiaei Poiret). Austrolycopodium Holub, Folia Geobot. Phytotax. 26: 87, 9 1 . 1 99 1 . TYPE: Lepidotis magellanica Beauv. (= Lycopodium magellanicum (Beauv.) Sw.). Plants anisotomously branched, with elongate, indeterminate, subterranean, creeping, or scan- dent, plectostelic main stems (rhizomes), which, in a dorsolateral position, give rise to usually de- terminate, ascending to erect, dendroid or spread- ing, repeatedly inclinate-bilaterally branched, branchlet systems. Roots emerging directly along the underside of main stems, with plectostelic main roots. Branchlet leaves uniform or anisophyllous. Strobili erect, simple or forked, sessile or borne on simple or forked peduncles. Sporoph ylls peltate without a basal mucilage cavity, or subpeltate with a thin basal decurrent wing and with a basal mu- cilage cavity. Sporangia attached to the sporophyll base, reniform, with a short thick stalk, isovalvate or slightly anisovalvate, their epidermis cells with thin, lignified, sinuate side walls, without partial thickenings. Spores reticulate. Gametophytes ob- conic to convoluted disc-shaped, subterranean, mycoparasitic, lacking pluricellular uniseriate tri- chomes among the gametangia. The generic description and synonymy above includes only the Neotropical representatives. For a fuller general treatment of the genus, see 011gaard (1987). The genus Lycopodium as here circumscribed has perhaps 40 species mainly in temperate regions of both the Northern and Southern Hemispheres, with eight species in the Neotropics and five in Peru, all of these, except L. vestitum, with wide distributions. It includes several of the genera of Holub (1975, 1983, 1985, 1991), also advocated by Wagner and Beitel (1992). These genera cor- respond to the sections in 011gaard (1987, 1989). Four of the nine sections are represented in the Neotropics. These sections are very distinct groups of species. None of them are connected by inter- mediate species or by intersectional hybrids. Reference WILCE, J. H. 1965. Section Complanata of the genus Lycopodium. Beih. Nova Hedw. 19: 1-IX, 1-233, t. 1-40. Key to Species of Lycopodium a. Branchlets radially symmetrical, isophyllous b b. Branchlet leaves with a colorless hair tip, or a colorless membranous apex; sporophylls subpeltate, with a basiscopic, membranous wing on the stalk (sect. Lycopodium) c c. Branchlet leaves hair-tipped, or with a short membranous apex; branchlets green; strobili sessile or borne on short or long, simple or branched peduncles 1 . L. clavatum c. Apical half or more of branchlet leaves colorless, membranous; branchlets grayish to silvery white; strobili sessile or borne on short, simple, indistinct peduncles 2. L. vestitum b. Branchlet leaves green throughout, without membranous or hairlike tips; sporophylls peltate, with a slender, terete, wingless stalk (sect. Magellanica) 3. L. magellanicum a. Branchlets dorsiventral, flattened, anisophyllous, with dimorphic or trimorphic leaves d d. Branchlet leaves decussate, arranged in 4 ranks, with widened lateral leaves, and 1 rank of dorsal and 1 rank of ventral, median narrow leaves (sect. Complanata) 5. L. thyoides d. Branchlet leaves alternate, with 2 ranks of of wide, dorsolateral leaves, and ca. 3 indistinct ranks of narrow, ventral, apically membranous leaves (sect. Diphasium) 4. L. jussiaei FIG. 7. Lycopodium clavatum: a, aerial shoot, habit. Lycopodium thyoides: b, aerial shoot, habit; c, tip of branchlet, adaxial side; d, tip of branchlet, abaxial side. (Adapted from Stolze, Ferns and fern allies of Guatemala, 1983.) 52 FIELDIANA: BOTANY TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 53 1. Lycopodium clavatum L., Sp. pi. 1101. 1753. LECTOTYPE: Herb. Burser XX: 49 (UPS, des- ignated by Jonsell & Jarvis, Regnum Veg. 1 27: 63. 1993). Lepidotis clavata (L.) Beauv., Prod. Aetheogamie 108. 1805. Plants creeping, trailing, or hanging over banks. Main stem usually above ground with ascending to stiffly erect, up to at least 50 cm tall, repeatedly unequally branched aerial shoots, with strongly diverging to almost parallel branchlets. Ultimate branchlets terete. Leaves borne in low alternating spirals or whorls of 6-8(-10), patent to ascending or imbricate, linear-acicular, 6-8(-10) mm long and 0.5-0.8 mm wide, terminating in a long col- orless trichome or short membranous apex, with smooth to sparsely denticulate margins. Strobili sessile or pedunculate. Peduncles, if present, erect, up to 30 cm long, simple, or branched and bearing up to 6 pedicellate strobili. Strobili 1.5-6(-8) cm long, ca. 6 mm in diameter including sporophylls, sometimes forked. Sporophylls borne in alternat- ing whorls of 5-6, subpeltate, with triangular-ovate to rhombic-ovate, acuminate exterior face, with usually widely scarious, dentate to erose-laciniate margins. Sporangia 1.3-1.6 mm wide, their epi- dermal cells with strongly sinuate side walls with pocket-like in- and evaginations. Spores reticulate on all faces. Almost cosmopolitan, in humid temperate and boreal regions of the Northern Hemisphere and on tropical mountains in the Old and New World. The species is highly variable and adaptive to ex- ternal factors. The discovery of tetraploids in Ja- pan indicates that genetic variation is also in- volved. There is as yet no indication of polyploidy in the Neotropical representatives of the species. Lycopodium clavatum varies almost continu- ously from amply branched plants with diverging branches and spreading, soft leaves, and long- branched peduncles, growing in moist, warm, shel- tered habitats, to small, compact, parallel-branched plants with more imbricate and firm leaves, and lacking, or short, simple or once forked peduncles, belonging to cold, exposed habitats. The latter forms are here recognized as ssp. contiguum. Cor- responding monostachyous forms are found in the Arctic and high mountains of New Guinea. The two subspecies recognized here are often consid- ered to be distinct species. However, there are many intermediate forms that cannot be placed in one or the other taxon with certainty. These inter- mediates form normal spores and have normal meiosis. Also there are rare examples of single individuals showing characters of one subspecies in the growth of one year and of the other the following year. As the two forms are usually rec- ognizable and are ecologically rather distinct, I treat them as subspecies. Key to Subspecies of Lycopodium clavatum a. Strobili rarely forked, borne on elongate, simple or branched peduncles; aerial shoots amply branched, with more or less diverging branchlets; sporophylls short to long acuminate .... 1 a. ssp. clavatum a. Strobili sessile, or borne on short, simple, often indistinct peduncles, frequently forked; aerial shoots sparsely branched, usually stiffly erect, with parallel branchlets; sporophylls usually with a long, wide, scarious apex Ib. ssp. contiguum la. Lycopodium clavatum ssp. clavatum Lycopodium aristatum Willd., in L., Sp. pi. ed. 4, 5: 17. 1810. TYPE: Venezuela, Silla de Caracas, Humboldt & Bonpland (holotype, B, Herb. Willd. 79557!). Lycopodium piliferum Raddi, PI. Bras. nov. gen. 1 : 79, t. 3. 1825. Based on L. aristatum Willd. and with the same type. Lycopodium trichophyllum Desv., Mem. Soc. Linn. Paris 6: 184. 1827 [as trychophyllum]. LECTO- TYPE (designated by 011gaard in Harling and An- dersson, Fl. Ecuador 33: 125. 1988): "Habitat in Brasilia, no. 41 in herb. deA. N. Desvaux, Donnee par Mme Vve Lavallee en 1895" (P!). Lycopodium clavatum L. var. aristatum (Willd.) Spring, Flora 21 (1): 173. 1838. Lycopodium eriostachys Fee, Crypt. Vase. Bresil 1: 224. 1869. TYPE: Brasilia fluminensi, Serra dos Orgaos, Glaziou 1788 (BR!, c!, P!, RB!). Clearings exposed habitats in montane forest, 700-3200 m, Piura, Cajamarca, Amazonas, La Libertad, San Martin, Loreto, Huanuco, Pasco, Junin, Ucayali, Ayacucho, Cuzco, Puno. 54 FIELDIANA: BOTANY Humid temperate and boreal regions of the Northern Hemisphere, montane in the tropics. Absent from Australia. For more complete synonymy, see 011gaard (1988). Cajamarca: Prov. Cutervo, Cutervo-Socota, 2320 m, Lopez & Sagdstegui 5331 (F, MO). Prov. Hualgayoc, Culquirrumi, 2448 m, Sanchez Vega 2099 (AAU). Ama- zonas: Prov. Chachapoyas, Pipus, between Chachapoyas and Molinopampa, 2100 m, Sanchez Vega et al. 2180 (AAU). La Libertad: Prov. Bolivar, above Longotea, 2800 m, Sagdstegui 14189(r). San Martin: Prov. Rioja, Pedro Ruiz-Moyobamba road, km 390-394, Venceremos, 1910-2040 m, D. Smith 4524 (AAU). Zepelacio, near Moyobamba, 1 1 00-1 600 m, King 3360 (MO, s, us). Tara- poto, Spruce 4732 (BR, E, G, p, us). Huanuco: Prov. Leon- cio Prado, Dist. Rupa Rupa, near Tingo Maria, 700 m, King 337 (F). Prov. Huanuco, Dist. Churubamba, Santo Toribio, 1500 m, Mexia 8144 (F, MO, s, uc, us). Pasco: Prov. Oxapampa, Cord. San Gutardo, 2600 m, Leon 512 (AAU). Cord. Yanachaga, 1 2 km E of Oxapampa-Villa Rica road, 2100-2200 m, Gentry & Smith 35922 (AAU). Junin: Prov. Tarma, Chanchamayo Valley, above La Merced at Cumbre Yacunay, 2000 m, Hutchison 1 194 (F, M, s, uc, us). Huacapistana, Killip & Smith 24255 (us). Ucayali: Prov. Coronel Portillo, La Divisoria, ca. 20 km NNE of Tingo Maria on road to Pucallpa, 1 600 m, Dillon 2641 (F). Ayacucho: Prov. La Mar, eastern Massif of Cord. Central opposing the Cord. Vilcabamba, between Tambo, San Miguel, Ayna and Hda. Luisiana, 1 300 m, Dudley 1 1 953 (F). Cearapa, Killip & Smith 22459 (us). Cuzco: Valle del Urubamba, Machu Picchu, 2400 m, Herrera 3197 (c, F). Tres Cruces, Gentry et al. 23458 (us). Puno: Below Limbani, 3100-3200 m, Brandbyge 523 (AAU). At Lake Titicaca near Huancano, 2000 m, Charpin AC- 12704(0). Ib. Lycopodium clavatum ssp. contiguum (Klotzsch) B. 011g. in Harling & Andersson, Flora of Ecuador 33: 126. 1988. Lycopodium contiguum Klotzch, Linnaea 18: 519. 1844. LECTOTYPE (designated by 011gaard in Harling & Andersson, Flora of Ecuador 33: 126. 1988): Ecuador, Paramo de Tiopullo [between Cotopaxi and Illiniza], Hartweg 1474 (B!; isolec- totypes, BM!, GL!, K.!, NY!, P!). Lycopodium vestitum Poiret, var. herbaceum Spring, Mem. Acad. roy. Belg. 24 [Mon. Lye. 2]: 45. 1 849. TYPE: Colombia, Sierra Nevada, Moritz (holo- type, B!; isotype, LG!). Lycopodium clavatum L. var. pseudo-contiguum Christ, Primitiae Fl. Costar. 3 (1): 55. 1901. TYPE: Costa Rica, Cerro de las Vueltas, 3000 m, versant W du massif de Buena Vista, Pittier 10467 (ho- lotype?, BR!; isotype?, P!). Lycopodium herbaceum (Spring) Hieron., Bot. Jahrb. Syst. 34: 575. 1905. Clearings and exposed habitats in uppermost montane forest and in grass paramo, frequent in disturbed paramo, 2600-3800 m, Piura, Caja- marca, Amazonas, San Martin, Ancash, Pasco, Huancavelica, Apurimac, Cuzco. Costa Rica; Andes from Venezuela to Peru. For more complete synonymy, see 011gaard (1988). Piura: Prov. Huancabamba, Cuello del Indio, 2800 m, Lopez et al. 8883 (AAU). Cajamarca: Prov. San Miguel, near El Tingo (Agua Blanca), 3100 m, Mostacero et al. 1130 (AAU). Cutervo, Jelski 1016 (us). Amazonas: Prov. Chachapoyas, Saullamur-Calla Calla, 3250 m, Sanchez Vega et al. 2136 (AAU). San Martin: Prov. Mariscal Ca- ceres, NW corner of Rio Abiseo National Park, 3500 m, Young & Leon 4712 (AAU). Ancash: Prov. Carhuaz, Huascaran National Park, Quebrada Ishinca, 3900 m, D. Smith et al. 9588 A (AAU). Prov. Yungay, Huascaran National Park, Quebrada Ranincuray, 3850 m, D. Smith et al. 9067 (USM). Pasco: Prov. Oxapampa, Dist. Huan- cabamba; Sta. Barbara, above Lanturachi, 3300-3500 m, Foster et al. 10398 (AAU). Huancavelica: Prov. Tay- acaja, Montepungo, 5 km E of Surcubamba, 300 m, Stork & Horton 10386 (F). Prov. Tayacaja, vicinity of Huachocolpa, 3000-3100 m, Tovar 3950 (USM). Apu- rimac: Prov. Abancay, Cerro Turronmocco, NW of Nev- ado Ampay, 3500 m, Brandbyge 422 (AAU). Cuzco: Prov. Urubamba, Antakillqa, 3300 m, Franquemont 285 (F). 2. Lycopodium vestitum Poiret, in Lam., Encycl. 3: 546. 1814. LECTOTYPE (designated by 011gaard in Harling & Andersson, Flora of Ecuador 33: 128. 1988): Ecuador (Prov. Loja), prope Loxam, Humboldt (p-Humboldt!; iso- types, B, Herb. Willd. 194281, BR!, LG!, us!). Lycopodium scariosum Hooker, Icon. PI. 1 : /. 89. 1 836, not Forst. TYPE: Peru, Casapi, Matthews 1765 (holotype, K!; isotype, IVY!). Lycopodium albidum Baker, J. Bot. 25: 37. 1887. TYPE: Ecuador, Prov. Loja, Mataba 1883, Hu- beck (holotype, K!). Rhizome creeping, scrambling or hanging over banks, usually above ground, 3-5 mm in diameter including leaves. Aerial shoot systems stiffly erect, sparsely branched, with almost parallel branches, silvery whitish. Branchlet leaves appressed, linear- lanceolate, 5-9 mm long, 0.6-0.8 mm wide, herbaceous in less than half of their length, with colorless, translucent, membranous, slightly widened, coarsely erose-laciniate apex. Strobili sessile or borne on a short, simple, indistinct pe- duncle. Sporophylls borne in alternating whorls of 5-6, subpeltate, 5-7 mm long, 1.5-2 mm wide, narrowly triangular-ovate, with a long membra- nous apex, with widely membranous, erose-den- tate to laciniate margins. Sporangia 1.2-2 mm wide, their epidermis cells with strongly sinuate TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 55 side walls with pocketlike evaginations. Spores re- ticulate on all faces. Clearings, road banks, and exposed sites in up- per montane forest, and in the lower paramos, 2400-3500 m, Piura, Cajamarca, Amazonas, San Martin. Southern Ecuador and northern Peru. Lycopodium vestitwn is closely related to L. cla- vatum, especially to ssp. contiguum. In spite of the striking appearance of this species, it is only doubt- fully distinct from L. clavatum. Individuals with intermediate characters between L. vestitwn and both subspecies are frequent. These have normal meiosis and normal spores. Piura: Prov. Huancabamba, Cord. Chinguela (Sapa- lache-El Carmen), 2900 m, Sagdstegui et al. 10223 (AAU). Above Huancabamba, 3400-3500 m, Weberbauer 6145 (F, GH, us). Cajamarca: Prov. Cutervo, Fortaleza de Santa Cruz, San Andres, 2400 m, Llatas & Suarez 2866 (F). Cutervo, Jelski 990 (p), 994, 1007 (KRA). Amazonas: Prov. Bagua, Cord. Colan E of La Peca, 3200 m, Barbour 3261 (AAU). Prov. Chachapoyas, Cerros Calla Calla, E side, 19 km above Leimebamba on road to Balsas, 3100 m, Hutchison & Wright 5545 (F, GH, MO, P, uc, us, USM). San Martin: Rio Abiseo National Park, Mariscal Ca- ceres, NW corner of the Park, 3300 m, Young 27 54 (AAU, USM). Rio Abiseo National park, Mariscal Caceres, La- guna del Eco, Chochos, 3450 m, Young & Leon 4901 (USM). 3. Lycopodium magellanicum (Beauv.) Sw., Syn. Fil. 180. 1806. Lepidotis magellanica Beauv., Prod. Aetheogamie 102. 1805. TYPE: Fretum magellanicum, unknown collector (holotype presumably in G, Herb. De- lessen n.v.\ isotype, P!, Herb. Palisot de Beauvois, comm. M. de Jussieu). Lycopodium spurium Willd., Sp. pi. ed. 4, 5: 28. 1810. TYPE: Ecuador (Prov. Tungurahua), Quito, in vulcano Tungurahua, Humboldt (holotype, B, Herb. Willd. 193641). Lycopodium pichinchense Hooker, Icon. pi. 1: /. 55. 1837. LECTOTYPE (designated by B. 011gaard, Biol. Skr. Dan. Vid. Selsk. 34: 62. 1989): Ecuador, Pichincha, 10,000 ft, Col. Hall (38?) (K!). Austrolycopodium magellanicum (Beauv.) Holub, Fo- lia Geobot. Phytotax. 26: 91. 1991. Rhizome subterranean or occasionally above ground, 1-2 mm thick excluding leaves. Aerial shoots ca. 1 mm thick excluding leaves at origin, up to 25 cm long excluding peduncles and strobili, repeatedly unequally branched, with spreading to ascending branchlets. Branchlets 4-7 mm in di- ameter including leaves. Branchlet leaves acicular, flattened, smooth, with a long, pointed, non-pilif- erous apex, 3-5 mm long, 0.6-0.8 mm wide. Strobili sessile, or terminating somewhat indis- tinct, simple or up to twice-forked peduncles, 3- 4 mm thick, 2-10(-l 5) cm long, often forked. Spo- rophylls borne in alternating whorls of 4, peltate, with a slender, terete, wingless stalk, with widely ovate, short to long acuminate exterior face at- tached to the stalk near the center, with narrowly scarious, almost smooth to shallowly erose-den- ticulate margins. Sporangia ca. 2 mm wide; spor- anguium epidermis with evenly sinuate side walls. Spores with unornamented proximal faces, with an irregular, fine-meshed reticulum on distal faces. Creeping or hanging over banks in grass paramo, usually in relatively dry paramos, but also often in mossy bogs, 2750-4000 m, Lambayeque, Ca- jamarca, La Libertad, San Martin, Ancash, Huan- uco, Cuzco. Hispaniola; Costa Rica; Andes from Venezuela to Chile and Argentina. The subterranean rhizomes seem to enable this species to survive in vegetation that is burned fre- quently. Lycopodium magellanicum is possibly identical to certain forms included in L. fastigiatum R. Br. of New Zealand and Australia. Lambayeque: Ferranafe, Laguna Tembladera, Inca- huasi, 31 50 m, Sagdstegui et al. 12791 in part (F). Prov. Ferranafe, ca. 7 km NW of Incahuasi, near Cerro Puna- machay, 3300-3550 m, Dillon & Skillman 4125 (F). Ca- jamarca: Cajabamba-Luchubamba, 3800 m, Sagdstegui et al. 11202 (AAU). San Miguel, Millan (El Tingo-Taulis), 3000 m, Sagdstegui 9526 (NY). Cutervo, Jelski 993, 1012, 1017 (KRA). La Libertad: Bolivar, Laguna de los Ichus, 3600 m, Lopez & Sagdstegui 3239 (GH). San Martin: Prov. Mariscal Caceres, Parque Nacional Rio Abiseo, Paredones, 3650 m, Leon & Young 1606 (USM). Prov. Mariscal Caceres, Parque Nacional Rio Abiseo, Cho- chos, 3400 m, Young & Leon 4896B (USM). Ancash: Prov. Carhuaz, Huascaran National Park, Quebrada Ish- inca, 3900 m, D. Smith et al. 9588B (AAU). Prov. Huari, Huascaran National Park, Quebrada Pachachaca, 3700- 3860 m, D. Smith et al. 12547 (USM). Huanuco: Yanano, 2000 m, Macbride 4928 (F). Cuzco: Paucartambo, 3700 m, Vargas 13938 (GH). Paucartambo, Parque Nacional Manu, Cerro Macho Cruz, 3200 m, Leon & Aguilar 2345 (USM). 4. Lycopodium jussiaei Poiret, in Lam., Encycl. 3: 543. 1814. TYPE: Perou (holotype, P, Herb. Jussieu 6581). Lycopodium jussiaei Poiret var. microphyllum Poiret, in Lam., Encycl. 3: 543. 1814. TYPE: Amerique meridionale, Herb. Bonpland (holotype, P!). Lycopodium haenkei Presl, Reliq. haenk. 1: 78. 1825. 56 FIELDIANA: BOTANY TYPE: Peru (Dept. Huanuco), In montosis Pe- ruviae ad Huanocco, Haenke (holotype, PRC!; possible isotype, K!). Lycopodium heterophyllum Sprengel, Syst. Veg. ed. 16, 4: 13. 1827. TYPE: Peruvia, Humboldt (ho- lotype, B, Herb. Willd. 194251). Lycopodium scariosum Forst. var.jussieui(Poirei) Ba- ker, Handbook Fern Allies 29. 1887. Diphasium jussiaei (Poiret) [Presl ex] Rothm., Feddes Repert. Spec. Nov. Regni Veg. 54: 65. 1944 [as jussieui]. Rhizomes creeping, scrambling to scandent, rig- id, 2-4 mm thick, usually above ground, with leaves radially arranged, uniform, 3-5 mm long, ca. 1 mm wide, linear-lanceolate, with widely membra- nous, irregularly obtuse apex. Aerial shoot systems 5-75 cm tall, branched almost from the base, in large individuals with a main axis almost conform to the rhizome, upward gradually changing to an- isophyllous, bearing alternating, fan-shaped branchlet systems. Branchlets dorsiventral, ani- sophyllous, flattened, 4-6(-8) mm wide including leaves, with 2 dorsolateral ranks of wide leaves and 2-3 indistinct ventral ranks of narrow leaves. Dorsolateral leaves flattened in the plane of the branchlet system, or inclinate to it, obliquely el- liptic, with the acroscopic margin 2-3 mm long, 1—1.5 mm wide, forward and ventrally curved, mucronate to short hair-tipped, short to long de- current. Ventral leaves appressed, lanceolate-su- bulate, with membranous apices. Peduncles up to 50 cm long (rarely absent), simple or up to twice- forked, bearing 1-3 strobili, terete. Strobili (l-)3- 10 cm long, 4—6 mm in diameter including spo- rophylls. Sporophylls borne in alternating whorls of 4, subpeltate, with ovate, more or less acumi- nate exterior face, 4-6 mm long, ca. 2 mm wide, with narrowly membranous, shallowly erose-den- ticulate margins. Sporangia 1.5-2 mm wide, the side walls of epidermis cells evenly sinuate and finely curled. Spores reticulate, with large, regular meshes on distal faces, and unornamented prox- imal faces. Usually a vigorous scrambling to scandent plant, common in clearings and on road banks and open places in montane forest, 1700-3700 m, Piura, Cajamarca, Amazonas, San Martin, Huanuco, Pasco, Huancavelica, Ayacucho, Cuzco, Puno. Jamaica; Dominican Republic; Costa Rica; Venezuela; Andes south to Bolivia; Brazil (Ita- tiaia). A variable species, apparently due to environ- mental factors. Lycopodium jussiaei is closely re- lated to L. scariosum (Indonesia to New Zealand) and to plants from Chile and Juan Fernandez rec- ognized as L. gayanum Remy. The plants referred to Lycopodium jussiaei are generally larger and coarser than those species. A modern revision of the group is needed. For more complete synony- my, see 011gaard (1988). Piura: Huancabamba, Loma Redonda (Sapalache- Chinguela), 2400 m, Sagdstegui et al. 10203 (AAU). Ca- jamarca: Prov. Cutervo, 10 km NW of Socota, 3200 m, Stork & Horton 10143 (F, G, MO). Cutervo, Jelski 1025 (KRA). Amazonas: Prov. Bagua, Cord. Colan, E of La Peca, 3200 m, Barbour 3262 (AAU, MO, USM). Prov. Cha- chapoyas, Puma-urcu SE of Chachapoyas, 3100-3200 m, Wurdack 1158 (F, s, uc). San Martin: Prov. Mariscal Caceres, NW corner of Rio Abiseo National Park, 3300 m, Young & Leon 4882 (AAU). Huanuco: Prov. Huanuco, Carpish Pass, 3300 m, Hodge 6291 (AAU, F). Churubam- ba, Pampa Hermosa, 1400 m, Mexia 8146 (F, o, MO, s). Pasco: Cord. Yanachaga, 1 2 km E of main Oxapampa- Villa Rica Road, 2100-2200 m, Gentry & Smith 35924 (AAU, MO). Oxapampa, 1970 m, Smith & Pretel 1645 (AAU). Huancavelica: Prov. Tayacaja, Montepungo, 5 km E of Surcubamba, 3000 m, Stork & Horton 10384 (F). Ayacucho: Prov. La Mar, eastern Massif of Cord. Central opposing the Cord. Vilcabamba between Tambo San Miguel, Ayna and Hda. Luisiana, Dudley 1 1 973 (F). Cuz- co: Prov. Paucartambo, Valle de Pilcopata, Patria-Pil- lahuata, 2000 m, Foster & Wachter 7475 (AAU). Puno: Below Limbani, 3100-3200 m, Brandbyge 558 (AAU). 5. Lycopodium thyoides Willd., Sp. pi. ed. 4, 5: 18.1810. TYPE: Venezuela, Silla de Caracas, Humboldt (holotype, B, Herb. Willd. 193521). Lycopodium complanatum L. var. tropicum Spring, in Mart., Flora Bras. 1 (2): 1 16. 1840. Lycopodium complanatum L. var. thyoides (Willd.) Christ, in Schwacke, PI. Nov. Mineiras 2: 42. 1900 [as thujoides]. Lycopodium complanatum L. var. validum Weath- erby, Proc. Amer. Acad. Arts Sci. 45: 414. 1910, based on L. thyoides Willd. and with the same type. Diphasiastrum thyoides (Willd.) Holub, Preslia 47: 108. 1975. Plants with creeping, trailing to scandent rhi- zomes, usually above ground, or hanging over banks. Rhizomes terete, 1.2-2.5 mm in diameter excluding leaves. Rhizome leaves relatively dis- tant, borne in irregular spirals, or subverticillate, subulate, appressed to ascending. Aerial shoots as- cending to erect, 10-50 cm tall, with vegetative portions up to ca. 30 cm tall. Main upright axis terete to somewhat flattened, bearing lateral, flat- tened, fan-shaped branchlet systems. Ultimate branchlets flattened, dorsiventral, anisophyllous, 1.5-3 mm wide including leaves, with trimorphic, TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 57 decussate leaves. Upper, median branchlet leaves with pointed, subulate to acicular, appressed, 1-2 mm long, free blades, and a conspicuous, ca. 0.4- 0.6 mm wide, prominently decurrent base. Lateral branchlet leaves bilaterally compressed, long-de- current, 2.5-7 mm long including bases, the free blades 1-3 mm long, appressed to spreading, acu- minate to long-pointed, the leaf bases 0.6-1 .5 mm wide, with almost parallel to distinctly diverging margins, often curved down. Ventral leaves in- conspicuous, acicular, without decurrent base, 1- 2 mm long. Peduncles 10-25 cm long, terete, bear- ing 4-9 pedicellate strobili. Strobili 1 .5-5 cm long, 2-4 mm in diameter including sporophylls, often with protracted sterile tips. Sporophylls usually borne in alternating whorls of 3, subpeltate, widely deltoid-ovate, long-cuspidate, ca. 2-3 mm long, 1.5-2 mm wide, with almost entire, widely mem- branous margins. Sporangia 1.5-2 mm wide, with side walls of epidermal cells evenly sinuate. Spores densely reticulate on all faces. Clearings, road banks, open habitats, and sec- ondary scrub in upper montane forest, and in the lowest paramos, alt. 2000-3400 m, Piura, Lam- bayeque, Cajamarca, Amazonas, La Libertad, San Martin, Ancash, Huanuco, Pasco, Junin, Huanca- velica, Ayacucho, Cuzco, Puno. Throughout moist mountainous regions of trop- ical America, south to northern Argentina. The present application of the name Lycopo- dium thyoides corresponds to the "L. thyoides- complex" of Wilce (1965). I have not attempted to treat the infraspecific variation. The Peruvian material referred to this species is highly variable and may belong to more than one species, but also external factors apparently greatly affect the growth habit of the individuals. Ferreyra 15111 (USM), from Dept. Cajamarca, Prov. Celendin, Celedin, ca. 3100 m, has unusually wide branchlets with very wide lateral leaves. For more complete synonymy, see 011gaard (1988). Cajamarca: Prov. Chota, Chota Tacabamba road, km 14, 2800 m, Smith & Vdsquez 3549 (AAU, F). Prov. San Miguel, El Tingo, 3100 m, Mostacero et al. 1123 (AAU, F). Amazonas: Prov. Chachapoyas, Leimebamba-Calla Calla road, km 12-15, 2960-3100 m, Smith & Vdsquez 4992 (AAU). Donile-Cohechan, Soukup 4160 (F, us). La Libertad: Prov. Pataz, Cerro Colpar, above Yalen, 3300- 3700 m, Young 3046 (AAU, USM). Prov. Bolivar, above Longotea, 2800 m, Sagdstegui 14195 (F). San Martin: Rio Abiseo National Park, NW corner, 2800 m, Young 1480 (AAU); Rio Abiseo National Park, Laguna del Eco, Chochos, 3450 m, Young & Leon 4902 (USM). Ancash: Prov. Yungay, Huascaran National Park, Quebrada Ranincuray, 3650-3900 m, D. Smith et al. 10336 (AAU, USM). Huanuco: Prov. Huanuco, Huanuco-La Union road km 32, 2940-3100 m, Huapalla 2201 (AAU). Carpish Pass, 2850 m, Asplund 12748 (s). Pasco: Above Oxa- pampa (Cerro Corporation), ca. 2300 m, collector un- known (F). Junin: Huacapistana, 1 800-2400 m, Killip & Smith 24255 (F). Huancavelica: Prov. Tayacaja, Mar- cavalle, between Huachocolpa and Tintay, 2800 m, 70- var 4765 (USM). Ayacucho: Ccarrapa, between Huanta and Rio Apurimac, 2500 m, Killip & Smith 22293 (F, us). Cuzco: Prov. Calca, below Kachin, 2700-3800 m, Sallo ex Franquemont 244 (AAU, F). Prov. Paucartambo, Marcachea, Achirani, 2500-3000 m, Vargas 11133 (F, MO, uc). Paucartambo, Woyt kowski 6744 (MO, us). Puno: Below Limbani, 3100-3200 m, Brandbyge 546 (AAU). Prov. Sandia, between Sandia and Cuyo-Cuyo, 3100- 3300 m, Ferreyra 16813 (USM). Tabina, Lechler 2034 (E, G, s). III. Lycopodiella Lycopodiella Holub, Preslia 36: 22. 1964. TYPE: Lycopodiella inundata (L.) Holub. Figure 8. Lycopodium subgen. Cernuistachys Herter, Bot. Jahrb. 43: Beibl. 98: 29. 1909. TYPE: Lycopodium cern- uum L. Palhinhaea Vascon. & Franco, Bol. Soc. Brot. II, 41: 24 (1 967). TYPE: Palhinhaea cernua (L.) Vascon. & Franco (= Lycopodium cernuum L.). Lycopodium sect. Caroliniana Bruce, Amer. Fern J. 66: 136. 1976. TYPE: Lycopodium carolinianumL. Pseudolycopodiella Holub, Folia Geobot. Phytotax. 18:441. 1983. Lepidotis auct., not Mirbel 1802. Lycopodium subgen. Lepidotis auct., not Sprengel, Anleit. ed. 2, 2, 1: 108. 1817, nor Baker 1887. Piura: Prov. Huancabamba, Dist. Sondor, Cerro La Viuda, 2170 m, Sagdstegui et al. 8197 (AAU, MO). Piura: Loma Redonda, 2400 m, Sagdstegui 10175 (AAU). Lam- bayeque: Prov. Ferranafe, Incahuasi (Sinchigual-Laguna Tembladera, 3000 m, Sagdstegui et al. 12870 (AAU, F). Plants with prostrate, rooting, indeterminate, isophyllous or anisophyllous, horizontally branch- ing shoots, and dorsally arising, erect, simple strobiliferous branches (sect. Lycopodiella and sect. Caroliniana), or with trailing to arching or looping FIG. 8. Lycopodiella caroliniana var. meridionale: a, habit; b, portion of stem and branch. Lycopodiella cernua: c, habit; d, ultimate branchlet, showing sterile leaves and strobilus; e, apex of strobilus. (Adapted from Stolze, Ferns and fern allies of Guatemala, 1983.) 58 FIELDIANA: BOTANY TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 59 indeterminate runner shoots that root with usually long intervals, and branch occasionally in the hor- izontal plane giving off horizontal branchlet sys- tems, and usually bearing 1 vertical, dorsally aris- ing, dendroid branch system on each loop (sect. Campylostachys). Dendroid branch system with a series of subdecussate, spreading to hanging, fla- bellate branchlet systems, which in turn may ter- minate in sessile, nodding to pendulous strobili. Sporophylls subpeltate, with a median basiscopic wing, or with coalescent basal membranes that almost enclose the sporangia (sect. Campylosta- chys). Sporangia on the sporophyll base, or axil- lary (sect. Lycopodielld), strongly anisovalvate, or isovalvate (sect. Caroliniand), the epidermis cells with thin, straight, non-lignified side walls, but with lignified, nodular, or semiannular thicken- ings. Spores rugate. Gametophytes green, tuberous and lobed on the upper side, surface-living, with mycorrhiza in the base. Almost all moist temperate and tropical regions of the world. Perhaps 40 species, the majority of these in the Americas and probably nine in Peru, all fairly widely distributed. The generic description above covers only the Andean representatives of the genus. Although growth habit and morphological details are quite diverse, features of branching, sporangium anat- omy, spores, and gametophytes indicate that Ly- copodiella is a natural entity. In sect. Campylostachys, most species have erect, dendroid branch systems arising from the dorsal side of horizontal, looping main axes (e.g., Lyco- podiella cernud), but in some species the corre- sponding dendroid branch system is scandent, hanging, or creeping. In these cases the dendroid branch system can be distinguished from the hor- izontal branch system by the orientation of lateral branchlet systems on the main axes. In the hori- zontal system they are distichous, all arranged in the horizontal plane. In the dendroid branch sys- tem the lateral branchlets are polystichous, ar- ranged in an irregularly subdecussate manner. Zimmer, in a review in Bot. Jahrb. Syst. 112: 414, 1991, claimed that Lepidotis is the correct name for this genus as lectotypified by Rothmaler (1944). However, Rothmaler's lectotypification does not refer to the protologue of Lepidotis and is superceded by the choice of Lycopodium cla- vatum (Pichi Sermolli, Webbia 26: 145-149. 1971). Key to Species of Lycopodiella a. Strobili erect, terminating simple, erect branches that arise dorsally on the creeping stem b b. Creeping shoots isophyllous or nearly so; leaves of the erect branch crowded, borne in alternating whorls of 6 or more c c. Sporangia subglobular, strongly anisovalvate, ca. 1 mm wide; Sporophylls and vegetative leaves with the same color and texture; leaves of erect branch with flattened, appressed base (sect. Caroliniand) 1. L. alopecuroides c. Sporangia reniform, isovalvate, 1-1.5 mm wide; Sporophylls and vegetative leaves with different color and texture; leaves of erect, branch strongly upward curved from a subterete, diverging base (sect. Caroliniand) 3. L. contexta b. Creeping shoots strongly anisophyllous, with wide and long ventrolateral leaves and narrow dorsal leaves; leaves of the erect branch borne in distant, alternating whorls of 4 or 5 (sect. Caroliniand) 2. L. caroliniana var. meridionalis a. Strobili nodding to pendulous, terminating spreading to pendulous branchlets that are borne on dendroid, erect or scandent, amply branched shoot systems (sect. Campylostachys) d d. Branchlet leaves closely imbricate, flattened throughout, lanceolate, with fimbriate margins and densely hairy leaf bases 9. L. riofrioi d. Branchlet leaves spreading to ascending, subterete, angular or apically flattened, acicular to linear- lanceolate, glabrous or hairy, not fimbriate e e. Leaves of the main axes closely appressed, less than 4 mm long; stems usually densely hairy, dendroid shoot system flexible, usually bending to hanging from an erect base 6. L. descendens e. Leaves of main axes patent to ascending or reflexed or, if loosely appressed, more than 4 mm long; stems glabrous or hairy f f. Dendroid branch system usually indeterminate, up to 4 m long, scandent to creeping, or all 60 FIELDIANA: BOTANY axes scandent to creeping; main axes robust; leaves of main axes coriaceous, apically flat- tened, usually strongly upward curved and hook-shaped, with hairy leaf bases 7. L. glaucescens Dendroid branch system determinate, vertical, not scandent or climbing g g. Lateral branchlet systems of dendroid branch system recurved to long-pendulous; strobili 3-5 mm in diameter; sporophylls usually more than 3 mm long, with irregularly dentate margins 8. L. pendulina g. Lateral branchlet systems of dendroid branch system erect or spreading to horizontal and distally nodding; strobili 2.5-3 mm in diameter, sporophylls rarely more than 2 mm long, with coarsely dentate to erose-laciniate margins h h. Branchlet systems divaricate to somewhat aggregate, patent to horizontal, usually with gently recurved tips; strobilus-bearing branchlets softly nodding at tip 4. L. cernua h. Branchlet systems stiffly ascending or suberect, often densely aggregated, pointed up- ward, not recurved; strobilus-bearing branchlets usually sharply reflexed at tip, or only the strobilus reflexed 5. L. camporum 1 . Lycopodiella alopecuroides (L.) ( 'ran li 1 1 . Amer. Fern J. 71: 97. 1981. Lycopodium alopecuroides L., Sp. pi. 1102. 1753. TYPE: Eastern temperate North America, Vir- ginia, Kalm (LINN 1257.7). Lycopodium matthewsii Hooker, Icon. pi. 1: /. 26. 1836. TYPE: Peru, Bagasan, [Bagazan, Dept. Amazonas] Matthews 1778 (holotype, K.!; isotype, BM!). Lepidotis alopecuroides (L.) Rothmaler, Feddes Re- pert. Spec. Nov. Regni Veg. 54: 66. 1944. Lycopodiella matthewsii (Hooker) Holub, Folia Geo- bot. Phytotax. 18: 441. 1983. Horizontal shoots creeping and firmly rooted throughout, up to 25 cm long, sparsely and un- equally branched, densely covered on all sides by almost uniform, somewhat upwardly secund, or spreading to perpendicular leaves, 7-12 mm wide including leaves, bearing stiffly erect, dorsally aris- ing, simple, strobiliferous branches. Leaves linear- lanceolate to subulate, flat, with smooth to den- ticulate-ciliate margins, soft, light green, 5-8 mm long, 0.5-1 mm wide. Erect branches ca. 5-8 mm in diameter including leaves, up to 30 cm tall in- cluding the strobilus, with radially arranged leaves borne in alternating whorls of 6 or more, like leaves of horizontal shoots, or slightly narrower, usually more ascending. Strobili up to 12 cm long, 10-18 mm in diameter including sporophylls. Sporo- phylls arranged as peduncle leaves, narrowly lan- ceolate to lanceolate from a subpeltate base, not coalescent at base, with few to many spreading to hooked teeth on margins, 6-10 mm long, 1-1.7 mm wide at base. Sporangia axillary, subglobular, ca. 1 mm wide, strongly anisovalvate, almost com- pletely concealed by sporophyll bases, the epider- mal cells with straight, unlignified side walls with semiannular lignified thickenings. Spores rugate, with a distinct equatorial rim. On open, wet, peaty or sandy soil, in the jalca or montane forest zone, 2400-3450 m, Cajamarca, Amazonas, San Martin, Huanuco, Pasco. Eastern temperate North America; Cuba; throughout continental tropical America. As here delimited, in the wide sense, Lycopo- diella alopecuroides is a polymorphic species, con- sisting of several elements. In the narrow sense, the species is restricted to temperate North Amer- ica and Cuba, while other elements are present in South America. High altitude forms in the Andes are relatively compact, wide-leaved, and robust and perhaps deserve recognition as a species of which the correct name is Lycopodiella matthews- ii; the material cited below belongs to this form. Cajamarca: Prov. Cutervo, El Suro, 2400 m, Sanchez Vega et al. 6019 (AAU, F). Cumbemayo 10 km WSW of Cajamarca, 3550 m, Molau & Sanchez Vega 840 (GB). Amazonas: Prov. Chachapoyas, 1-5 km W of Molino- pampa, 2400 m, Wurdack 1381 (F, GH, uc, us). Lei- mebamba-Calla Calla, km 12-15, 2960-3100 m, Smith & Vdsquez 5000 (AAU). Chachapoyas, 2400-2600 m, Weberbauer 4397 (G). Prov. Bagua, Cord. Colan E of La Peca, 3500 m, Barbour 3197, 3200 A (AAU, MO). San Martin: Prov. Mariscal Caceres, Rio Abiseo National Park, NW corner of the park, 3425m, Young 3475 (AAU); 3450 m, Leon 1854 (AAU). Huanuco: Tingo Maria, Mor- row 11131 (GH). Pasco: Prov. Oxapampa, San Gotardo, 36 km W of Oxapampa, 2850 m, D. Smith 2753 (AAU). 2. Lycopodiella carol in iana (L.) Pichi-Sermolli var. meridionalis (Underw. & Lloyd) B. 011g. & Windisch, Bradea 5: 27. 1987. Lycopodium meridionale Underw. & Lloyd, Bull. Torr. TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 61 Bot. Club 33: 121. 1906. TYPE: Puerto Rico, dry savannahs, Luquillo Mountains, Wilson 94 (ho- lotype, NY!). Lycopodium carolinianum L. var. meridionalis Nes- sel. Arch. Bot. Est. S. Paulo 1: 431. 1927. Pseudolycopodiella meridionalis (Underw. & Lloyd) Holub, Folia Geobot. Phytotax. 18: 442. 1983. Horizontal steins creeping and firmly rooted throughout, to ca. 30 cm long, sparsely and un- equally branched, anisophyllous, with long and wide lateral leaves, and usually shorter and nar- rower dorsal leaves, 7-1 2(-l 5) mm wide including leaves, bearing 1 to few, dorsally arising, stiffly erect, simple, strobilus-bearing branches. Lateral leaves 3-5(-7) mm long, (l-)1.5-2.5(-3) mm wide at the base, triangular-ovate to lanceolate, oblique- ly spreading to falcately recurved, tapering into an acute to long pointed apex. Dorsal leaves arranged in (l-)2-4 longitudinal ranks (often on the same individual), widely lanceolate to subulate, diverg- ing to appressed, straight to upward curved, (1.5-)3-4(-5) mm long, l-1.3(-2) mm wide at the base. Erect branches up to 30 cm tall including the strobilus, 1-1.5 mm thick excluding leaves, bearing small, 3-5 mm long, acicular leaves in remote, alternating spirals or irregular whorls of 4-5. Strobili up to at least 13 cm long, 3-5 mm thick with appressed sporophylls. Sporophylls borne in alternating whorls of 4 or 5, subpeltate, rhombic or ovate-acuminate or ovate-cuspidate to triangular-lanceolate, 3.5-6 mm long, (l-)1.5-2(- 2.5) mm wide, with entire to erose-denticulate, minutely fimbriate-denticulate margins. Sporan- gia reniform, isovalvate, borne on the sporophyll stalk, ca. 1.5-2 mm wide. Spores rugate, with a distinct equatorial rim. Terrestrial or sometimes epilithic, usually on wet ground, on road banks, slopes, ledges, grass- lands in the jalca zone, on seepages, 1350-2400 m, Amazonas, San Martin. Widely distributed in tropical America, rare in the Andes. The distinctness of the present taxon from the type variety (temperate North America) has been doubted (e.g., Proctor, 1977, pp. 32-33), because the distinguishing characters usually indicated for them are widely overlapping, and seem to be based entirely on features related to the size of the plants. Amazonas: Chachapoyas, 1-5 km W of Molinopampa, 2400 m, Wurdack 1382 (F, GH, NY, us, USM). San Martin: San Roque, 1350-1500 m, LI. Williams 7763 (F, us). 3. Lycopodiella contexta (Mart.) Holub, Folia Geobot. Phytotax. 18: 441. 1983. Lycopodium contextum Mart., Icon. Crypt. Brasil. 38, t. 20, f. 1. 1834. TYPE: In campis graminosis apricis in summo monte Arara-Coara, fluvii Ja- pura cataractae imminente, alt. ca. 1200 p., Mar- tins (holotype, M!). Lycopodium alopecuroides L. var. contextum (Mart.) Baker, Handb. Fern Allies 19. 1887. Lycopodium sprucei Baker, Handb. Fern Allies 24. 1887. TYPE: Venezuela, San Carlos del Rio Ne- gro, Spruce 3151 (holotype, K!; isotype, BONN, frag. Herb. Nessel 200\ in part). Lycopodium alopecuroides L. subsp. contextum (Mart.) Hassler, Trab. Inst. Bot. Farm. Buenos Aires 45: 92. 1928. Lepidotis contexta (Mart.) Rothm., Feddes Repert. Spec. Nov. Regni Veg. 54: 66. 1944. Pseudolycopodiella contexta (Mart.) Holub, Folia Geobot. Phytotax. 26: 93. 1991. Horizontal shoots creeping and firmly rooted throughout, up to at least 30 cm long, sparsely and unequally branched, isophyllous, with somewhat upwardly secund, ascending, acicular leaves, 3-7 mm in diameter including leaves, bearing stiffly erect, dorsally arising, simple, or sometimes 1-2 x forked, up to at least 40 cm tall, vegetative or strobiliferous aerial shoots. Erect branches 4-6 mm in diameter including leaves, densely foliose, iso- phyllous. Leaves of erect branches borne in alter- nating irregular whorls of 6-7, subterete at the base (angular when dry), sometimes apically flattened, strongly upward curved from a strongly diverging to almost perpendicular base, 4-5(-7) mm long, ca. 0.5 mm wide, with conspicuously acroscopical- ly adnate and long decurrent leaf bases, with smooth margins. Strobili up to 8 cm long, 3-4 mm in diameter with appressed sporophylls. Sporo- phylls borne in alternating whorls of 4-5, subpel- tate, widely triangular-ovate to ovate-lanceolate at base, with a long, narrow apex, 4.5-7 mm long, 1-1.5 mm wide, with subentire to slightly erose- dentate margins at the base. Sporangia borne on the sporophyll base, reniform, isovalvate, 1-1.5 mm wide. Terrestrial, humid places on white sand, grass- lands, and river margins, ca. 1 20 m, Loreto. Circum-Amazonian, periphery of the Amazo- nian lowland of Brazil, Peru, Colombia, and Ven- ezuela. Loreto: Manfinfa, on the upper Rio Nanay, Ll. Wil- liams 1 106 (F, us); Vicinity of Iquitos, ca. 120 m, Revilla 4340 (F). 62 FIELDIANA: BOTANY 4. Lycopodiella cernua (L.) Pic.-Ser., Webbia 23: 165. 1968. Lycopodium cernuum L., Sp. pi. 1103. 1753. TYPE: LINN 1 257. 1 3, see Proctor, Ferns of Jamaica, Lon- don 29. 1985. Lepidotis cernua (L.) Beauv., Prodr. Aetheogamie 108. 1805. Lycopodium cernuum var. capillaceum Spring, Mem. Acad. roy. Belg. 15 [Mon. Lye. 1]: 80. 1842. TYPE: Venezuela, Edo. Monagas, Guanaguana, Hum- boldt 473 (holotype, B, Herb. Willd. 194291). Lycopodium capillaceum (Spring) Hieron., Bot. Jahrb. Syst. 34: 573. 1905. Palhinhaea cernua (L.) Vase. & Franco, Bol. Soc. Brot. 41:25. 1967. Plants with arching-looping runner shoots, with up to 1 m tall, erect, dendroid branch systems, these with several, subdecussate to alternate, high- ly compound, spreading to horizontal, 5-15(-20) cm long lateral branchlet systems. Ultimate branchlets nodding, 3— 4(-6) mm in diameter in- cluding leaves. Branchlet leaves usually borne in densely crowded, alternating whorls or low spirals of 3-5, usually 3-4 mm long, ca. 0.3 mm diameter, acicular, terete to angular (dried), with often con- spicuously acroscopically adnate and decurrent leaf bases, gradually changing from patent-reflexed and distant on main axes, to patent, upward curved and densely crowded in ultimate branchlets, oc- casionally with sparse, lax trichomes or minute spinules; leaf bases often with longer, irregularly crisped or branched trichomes, these rarely also on stern surfaces. Strobili usually numerous, ses- sile, terminating ultimate branchlets, 4-10(-20) mm long, 2.5-3 mm in diameter. Sporophylls usu- ally borne in alternating whorls of 5, ovate-deltoid, short to long cuspidate, ca. 2 mm long, ca. 1 mm wide, with membranous, coarsely erose-laciniate margins. Sporangia globose, 0.5-0.8 mm in di- ameter, strongly anisovalvate. Spores rugate, without a distinct equatorial rim. A common pioneer species on road cuts and moist disturbed soil, along rivers, in forest clear- ings, etc., from sea level up to 3000 m, Cajamarca, Amazonas, San Martin, Loreto, Huanuco, Pasco, Junin, Ayacucho, Cuzco, Puno. Pantropic. Where Lycopodiella cernua grows intermixed with L. descendens, intermediates may occur. Llanos & Chimouy 61 (USM), from Dept. Ca- jamarca: Prov. Jaen, San Patricio, Santa Rita, Chontali, 1 600 m, with the growth habit of L. cernua, deviates from typical forms of the latter by the densely crowded and strongly reflexed leaves on the main erect axis, by the somewhat flattened branchlet leaves, and by the densely hairy leaf bas- es and stem surfaces. It closely resembles forms from similar altitudes in Ecuador (Prov. Zamora- Chinchipe) and Bolivia and may represent a dis- tinct element, but more field observations and ma- terial are desirable for an assessment of its status. Cajamarca: Prov. Cutervo, La Pucarilla-San Andres, 2400 m, Sanchez Vega et al. 5999 (AAU, F). San Ignacio, San Jose de Lourdes, Villarica, 1650 m, Diaz 2053 (F). Amazonas: Prov. Bagua, road Chiriaco-Puente Vene- zuela, ca. 200-300 m, Barbour 4326 (AAU, MO). Prov. Chachapoyas, ca. 1800 m, Raimondi 350 (USM). San Martin: Prov. Mariscal Caceres, Dist. Tocache Nuevo, J. Schunke V. 3713 (F, us). La Divsioria, 59 km from Tingo Maria on road to Pucallpa, Allard 22175 (us). Loreto: Prov. Maynas, Dist. Iquitos, near mouth of Rio Nanay, Rimachi 449 (F, MO, USM). Balsapuerto, 220 m, Klug 2970 (F, MO, s, uc, us). Huanuco: Prov. Huanuco, Dist. Churubamba, Hda. Mercedes, 1875 m, Mexia 8193 (F, GH, s, us). Vilcabamba, 2000 m, Macbride 5014 (F, GH, MO, us). Pasco: Prov. Oxapampa, Palcazu valley, 300 m, D. Smith 3927 (AAU). Pichis Trail, Eneiias, 1700 m, Killip & Smith 25688 (F, us). Junin: Chanchamayo Valley, 1200 m, C. Schunke 188 (F). La Merced, Killip & Smith 23755 (us). Ayacucho: Prov. La Mar, eastern Massif of the Cord. Central opposing the Cord. Vilca- bamba between Tambo, San Miguel, Ayna and Hda. Luisiana, 1570 m, Dudley 11889 (GH, us). Cuzco: Pil- copata-Atalaya, 700 m, Vargas 13309 (GH). Puno: Olla- chea to San Gaban, 1000-2000 m, Dillon et al. 1166 (F, MO). 5. Lycopodiella camporum B. 011g. & P. G.Windisch, Bradea 5: 24, t. 3. 1987. TYPE: Brasil, Est. Minas Gerais, Mun. Santana do Riacho, Serra do Cipo, Prado et al. 69 (ho- lotype, HB!; isotypes, AAU!, RB!, SP!, SPF!). Palhinhaea camporum (B. 011g. and Windisch) Holub, Folia Geobot. Phytotax. 26: 93. 1991. Plants with arching-looping runner shoots, with up to 1 m tall, stiffly erect, dendroid branch sys- tems, these with several, subdecussate to alternate, often densely aggregated, stiffly ascending, 5-10 (-20) cm long lateral branchlet systems. Ultimate branchlets 2.5-7 mm in diameter including leaves, stiffly ascending to erect, only the strobiliferous ones rather sharply recurved at the tip. Branchlet leaves usually borne in densely crowded, alter- nating low spirals or oblique whorls of 5-7, 2.5- 4 mm long, ca. 0.3-0.5 mm in diameter, acicular, terete or angular (dried), sometimes flattened in the lower half, with often slightly acroscopically adnate and long decurrent leaf bases, patently ar- TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 63 cuate-ascending to arcuate-appressed, entirely smooth to densely hairy on leaf bases, rarely with soft trichomes on leaf margins. Strobili usually numerous, sessile, terminating tips of ultimate branchlets with a sharp bend, to 2.5 cm long, 2- 3 mm in diameter. Sporophylls borne in alternat- ing whorls of 5-7, widely ovate, short to long cus- pidate, 1.5-2 mm long, 0.6-0.9 mm wide, with scarious, coarsely erose-laciniate margins. Spo- rangia globose, ca. 0.6 mm in diameter, strongly anisovalvate. Swamps, moist campo-grasslands, on peaty or sandy soil, often in the light-open vegetation ad- jacent to gallery-vegetation at rivers, subject to flooding during the rainy season; alt. 1 20-1 800 m, San Martin, Junin, Loreto. Amazonian Colombia, Peru and Bolivia; sa- vanna region of Venezuela and Guyana, campos vegetation in Brazil. Lycopodiella camporum is a widespread species which has been confused with L. pendulina and synonyms thereof. The distinctive branching habit and characteristic ecology make L. camporum a rather easily recognizable and ecologically well- defined taxon, although detail characters of mor- phology do not seem strong. San Martin: Tarapoto, LI. Williams 5960 (F). Junin: Prov. Satipo, Gran Pajonal, 1000-1 200 m, Pena & Oven- tini 915 (USM). Loreto: Prov. Maynas, Quisto Cocha, Iquitos, 120 m, Sagdstegui & Aldave 5821 (GH, MO). 6. Lycopodiella descendens B. 011g., in Harling and Andersson, Fl. Ecuador 33: 143. 1988. TYPE: Ecuador, Prov. Pastaza, Mera, B. 0llg. & Balslev 9076 (holotype, AAU!; isotypes, F!, QCA!). Palhinhaea descendens (B. 011g.) Holub, Folia Geo- bot. Phytotax. 26: 93. 1991. Runner shoots arching-looping, usually densely hairy, with relatively distant, appressed, subulate leaves, with initially erect, then recurved and de- scending, amply branched, up to at least 75 cm long dendroid branch systems, these with several subdecussate to alternate, highly compound, spreading to hanging lateral branchlet systems, with usually densely hairy main axis with relatively dis- tant, appressed, subulate leaves. Ultimate branch- lets 3-4 mm in diameter including leaves. Branch- let leaves borne in alternating, densely crowded whorls or low spirals of 5-6, 2—4 mm long, ca. 0.3 mm thick, subterete, acicular to quadrangular, with long-decurrent and acroscopically adnate, hairy leaf bases, ascending and upward curved, glabrous, or with few lax trichomes above the base. Strobili usually numerous, sessile, terminating ultimate hanging branchlets, 5-20 mm long, ca. 3 mm in diameter. Sporophylls usually borne in alternating whorls of 5, ovate-deltoid, cuspidate, 1.5-2 mm long, ca. 1 mm wide, with membranous, coarsely erose-laciniate margins. Sporangia globose, ca. 0.5 mm in diameter, strongly anisovalvate. Spores ru- gate, without a distinct equatorial rim. Road banks, moist disturbed soil in lower, wet montane forest, alt. 1000-1 500 m, Amazonas, San Martin, Loreto, Huanuco. Lower eastern slopes of the Andes in Peru and Ecuador. Lycopodiella descendens is closely related to L. cernua, and is sometimes seen growing intermixed with it. In such situations intermediate individuals may be found. In addition to the key characters mentioned, L. descendens very often differs from L. cernua by its longer Strobili. Amazonas: Prov. Bagua, Mesones-Muro highway, 286 km E of Olmos, 8 km E of Montenegro, 650 m, Hutchi- son & Wright 3820 (F, uc, us). San Martin: Prov. Lamas, Dist. Lamas, Rio Curiyacu, affluent of Rio Cumbasa, ca. 450 m, Belshaw 3585 (GH, MO, NY, us, us). Prov. Maris- cal Caceres, Tocache Neuvo-Juanjui road, km 88, 850 m, D. Smith 2152 (AAU). Prov. Moyobamba, Jepelacio, Cerro Shallcahurco, 1320 m, Fernandez & Clemants 27 (USM). Loreto: Pumayacu, between Balsapuerto and Moyobamba, 600-1200 m, Klug 3232 (F, G, GH, K, MO, NY, s, us). Huanuco: Prov. Huanuco, Dist. Churubamba, Hda. Mercedes, 1640 m, Mexia 8195 (F, GH, K, s, uc, us). SW slope of the Rio Llullapichis watershed, on the ascent of Cerros del Sira, ca. 1000 m, Dudley 13125 (GH). 7. Lycopodiella glaucescens (Presl) B. 011g., Ope- ra Bot. 92: 176. 1987. Lycopodium glaucescens Presl, Reliq. haenk. 1: 81. 1825. TYPE: Peru (Huanuco), In montanis ad Huanocco, Haenke (holotype, PRC!). Palhinhaea glaucescens (Presl) Holub, Folia Geobot. Phytotax. 26: 93. 1991. Runner shoots long, robust, arching-looping to scandent, with initially erect, amply branched, up to several meters long, bending to long-scandent dendroid branch systems, these with several spreading or nodding to long pendulous, usually alternate, up to at least 40 cm long lateral branchlet systems. Leaves of main axes patent to reflexed, usually strongly upward curved to hook-shaped, 3-5 mm long, up to 1 mm wide, with terete to 64 FIELDIANA: BOTANY quadrangular, hairy leaf bases, apically flattened, usually coriaceous. Ultimate branchlets 3-5 mm in diameter including leaves, the sterile ones often tapering to less than 2 mm in diameter. Leaves of ultimate branchlets borne in densely crowded, al- ternating whorls or low spirals of 4-6, acicular to lanceolate, usually with flattened apex, 2.5-4 mm long, 0.3-0.8(-1.0) mm wide, usually strongly up- ward curved or hook-shaped from a spreading or perpendicular to reflexed base, soft herbaceous to rigidly coriaceous, hairy only on leaf base. Strobili variable, 5-30 mm long, 3-4.5(-5) mm in diam- eter. Sporophylls borne in alternating whorls of 5, lanceolate-ovate, apically acute-acuminate, 2-3 mm long, 0.8-1.2 mm wide, with shallowly den- tate-ciliate margins, of herbaceous, pale green tex- ture throughout. Sporangia globose, strongly an- isovalvate, 0.7 mm in diameter. Spores rugate, without a distinct equatorial rim. Clearings and open places in wet upper montane forest, alt. 1600-3700 m, Cajamarca, San Martin, Huanuco, Pasco, Cuzco. Costa Rica; Colombia to Peru; possibly farther north and south. A remarkable species with often very long scan- dent dorsally arising shoots. Under favorable con- ditions, these may reach a length of several meters. Lycopodiella glaucescens is highly variable with respect to strobilus size, size, coarseness, and di- rection of the branchlet systems. Plants of exposed habitats are usually relatively short and compact and with rigid, coriaceous leaves, approaching the growth habit of L. pendulina, while individuals from protected habitats are longer, more lax, and soft and with long pendulous branchlets. Cajamarca: Prov. Cutervo, La Pucarilla (San Andres- Socota), 2420 m, Lopez & Sagdstegui 5459 (GH). San Martin: Huallaga. Valley of Rio Apisoncho, 30 km above Jucusbamba, Hamilton & Holligan 1248 (K). Prov. Mariscal Caceres, Rio Abiseo National Park, hill past Las Palmas, 2650-2750 m, Young 4000 (USM). Huanuco: Prov. Huanuco, Carpish, 2850 m, Asplund 13076 (s). Playapampa, 3000 m, Macbride 4506 (F, G, us). Pasco: Prov. Oxapampa, Cord. San Gutardo, 2800 m, Leon 523a (AAU, USM). Cuzco: Prov. La Convencion, Cord. Vilcabamba, 2825 m, Dudley 10782 (USM). 8. Lycopodiella pendulina (Hooker) B. 011g., Ope- ra Dot. 92: 176. 1987. Lycopodium pendulinum Hooker, Icon. pi. 1: /. 90. 1837. TYPE: Peru, Casapi, Matthews 1776 (ho- lotype, K.!). Lycopodium cernuum L. var. pendulinum (Hooker) Baker, Handb. Fern Allies 23. 1887. Palhinhaea pendulina (Hooker) Holub, Folia Geobot. Phytotax. 26: 93. 1991. Runner shoots robust, shallowly arching, with up to ca. 40 cm tall, erect dendroid branch systems, these with several subdecussate to upward alter- nate, sparsely subequally branched, up to 1 2(-20) cm long, usually long pendulous lateral branchlet systems. Leaves of main axes loosely appressed, upward curved, 5-7 mm long, up to 1 mm wide. Ultimate branchlets (3-)5-6(-9) mm in diameter including leaves, rarely tapering to 2 mm in di- ameter. Branchlet leaves borne in densely crowded alternating whorls or low spirals of 5-7, acicular, terete to quadrangular, or sometimes apically flat- tened, (3-)4-6 mm long, up to 1 mm wide, patent- ascending, upward curved, softly to firmly her- baceous, glabrous, or with few marginal cilia. Leaf bases and stem surfaces usually glabrous, rarely with short crisped trichomes. Strobili 10-20 mm long, 5-6 mm in diameter. Sporophylls borne in alternating whorls of 5-6, lanceolate-ovate, short acuminate, (2.5-)3-3.5 mm long, ca. 1 .5 mm wide, with irregularly dentate, narrowly membranous margins, pale greenish, of herbaceous texture throughout. Sporangia globose, strongly aniso- valvate, ca. 1 mm in diameter. Spores rugate, without a distinct equatorial rim. Wet grassland, open spaces or clearings in the upper montane forest and, 2200-3600 m, Piura, Amazonas, Huanuco, Cuzco. Costa Rica; Andes from Venezuela to Bolivia, southeastern Brazil. Lycopodiella pendulina is most closely related to L. glaucescens, and distinguished from the latter by its stiffly erect main shoots with thick, sparsely ramified, weeping branchlet systems and large Strobili. Piura: Prov. Huancabamba, Loma Redonda (Sapa- lache-Chinguela), 2400 m, Sagdstegui et al. 10217 (AAU). Amazonas: Prov. Chachapoyas, Cerros Calla Calla, E side, 1 8 km above Leimebamba on road to Balsas, 3 100 m, Hutchison & Wright 5663 (BR, c, E, F, G, GH, M, MO, NY, s, uc, USM). Prov. Bagua, Cord, de Colan E of La Peca, 3400 m, Barbour 3199 (AAU, MO). Huanuco: Prov. Huanuco, Carpish, 2800 m, Asplund 13105 (s). Chushi, trail to Tambo de Vaca, Bryan 681 (F, us). Cuzco: Prov. La Convencion, Cord. Vilcabamba, 2825 m, Dudley 10707, 10782 (GH, MO). 9. Lycopodiella riofrioi (Sodiro) B. 011g., Opera Bot. 92: 176. 1987. Lycopodium riofrioi Sodiro, Crypt, vase. Quit. 582. 1893. TYPE: Ecuador (Prov. Pichincha), In silv. TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 65 occid. m. Pichincha ad Gualea, 9/888, Sodiro (ho- lotype, si 2 1230!). Lycopodium pensum Lell., Proc. Biol. Soc. Wash. 89: 7 1 7, t. 2. 1 977. TYPE: Costa Rica, Prov. Heredia, 6 mi from San Rafael de Heredia on slopes of Volcan Barba [Cerro Chompipe], McAlpin 216 (holotype, DUKE; isotype, GH!). Palhinhaea riofrioi (Sodiro) Holub, Folia Geobot. Phytotax., Praha 26: 93. 1991. Growth habit as in Lycopodiella pendulina, up to 2 m tall. Branchlet leaves imbricate, densely crowded, borne in alternating whorls of 4-6, lan- ceolate to ovate-lanceolate, acuminate, 2.5-4 mm long, ca. 1 mm wide, with densely ciliate-fimbriate margins, with a very short, patent, terete-suban- gular base. Stems and leaf bases densely hairy. Strobili 1-2.5 cm long, 3-5 mm thick including sporophylls. Sporophylls borne in alternating whorls of 5, ca. 2.5 mm long, ca. 1 mm wide. Sporangia 0.7-1 mm in diameter. Wet montane forest, in clearings and on road banks, 1700-2600 m. To be expected in north- ernmost Peru. Costa Rica; Panama; Venezuela; Colombia to southern Ecuador. A very distinctive species with its smooth-ap- pearing branchlets covered by closely imbricate, scalelike leaves. Family 27. SELAGINELLACEAE Selaginellaceae Milde, Hoher. Sporenpfl. Deutschl. Schweiz 136. 1865, as Selaginelleae. TYPE: Selaginella Beauv. Stem indurated or not, branched, bearing rather few, often long roots usually at a branch of the stem. Leaves simple, ca. 0.5-10 mm long, with 1 (very rarely 2) vein(s). Sporangia short-stalked, single, near the axil of a leaf. Heterosporous, spores without chlorophyll. The Selaginellaceae are a distinctive family in- cluding the single genus Selaginella, only distantly related to others such as the Lycopodiaceae and Isoetaceae. Heterospory, and the presence of ves- sels in some species of subgenus Selaginella, in- dicate the family is specialized. I. Selaginella Selaginella Beauv., Magasin Encycl. 5: 478. 1804; and Prod. Fam. Aetheog. 101. 1805, nom. conserv. TYPE: Selaginella spinosa Beauv., nom. nov. for Lycopodium selaginoides L. = Selaginella selaginoides (L.) Link. Figure 9. Bryodesma Sojak. Preslia 64: 154. 1992. Type: Bry- odesma rupestris (L.) Sojak (Lycopodium rupestre L.) = Selaginella rupestris (L.) Spring. Terrestrial, rupestral, or rarely epiphytic. Stem slender, branched, sometimes dichotomously, prostrate-creeping or with ascending branches, or pendent-epiphytic, or erect from a usually stolon- iferous base. Leaves ca. 1.0-10 mm long, with usually 1 vein, borne in a close spiral or usually alternate in 4 ranks. Sporangia large, borne in or near the axil of a well-differentiated sporophyll, in a quadrangular or (in S. deflexa and S1. selagi- noides) a cylindrical strobilus. Megasporangia commonly basal in the strobilus, usually with 4 megaspores, larger than the microsporangia and different in size and color. Microsporangia usually borne above the megasporangia and with many microspores. Megaspores tetrahedral-globose, tri- lete, the laesurae % to usually equaling the radius, often with a more or less prominent equatorial ridge, rugose-reticulate, rugose, papillate, tuber- culate, granulate, rarely nearly smooth on the proximal face. Microspores tetrahedral-globose, trilete, often compressed or the proximal face more or less depressed, the laesurae '/2 to equaling the radius, usually finely to coarsely echinate, rugose, papillate, perforate-cristate, or granulate. Selaginella is a large genus of perhaps 700 spe- cies, with about 40 of them in Peru, or expected there. The descriptions are drawn from Peru ma- terial and are intended to be parallel within related groups of species. The genus has not been ame- nable to segregation although five subgenera are FIG. 9. Selaginella peruviana: a, habit; b, tip of branch with aerial root. Selaginella haematodes: c, habit; d, portion of branch, abaxial side, showing axillary and lateral leaves. Selaginella exaltata: e, habit; f, portion of secondary branch, adaxial side, showing median leaves, (a, b from Solomon & Nee 17895, Bolivia, F; c, d from Taylor 5049, Brazil, F; e, f from Bohs 2215, Ecuador, F.) 66 FIELDIANA: BOTANY TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 67 currently recognized (Jermy, 1986), and Sojak has recently recognized a second genus, Bryodesma, for the S. rupestris group. The genus is probably more diverse in Peru than presently known. Species are difficult to identify, and many may appear similar to the field botanist unacquainted with special specific characters. At the same time, a better understanding of the vari- ation within species and the morphology of ju- venile plants will lead to an increase in synonyms. The species have been placed in a sequence that, presumably, extends from primitive to advanced. Species 1-2 are homophyllous, while the remain- der of the species are heterophyllous. Species 3-10 are prostrate and not articulate (both primitive characters), while species 11-12 are prostrate and articulate. These species relate to the former group by their small leaves and often intricate stems. Species 13-15 have erect, not ar- ticulate stems that are usually short and pinnately branched. Species 16-25 are also erect and not articulate but usually have a main stem with ap- pressed leaves toward the base and flabellate branches. Species 1 7 may be unbranched or with irregular branches. Species 26-30 are prostrate and articulate, as are species 11-12. However, these have large leaves and mostly separate stems. Spe- cies 31-34 are articulate and erect (both derived characters). Species 35 is a climbing species and 36 is a highly derived, xeric species. The following treatment is based to a large ex- tent upon the published work of A. H. G. Alston (1934), especially the (posthumous) publication with A. C. Jermy and J. M. Rankin (1981). The Koller and Scheckler (1986) paper is among the many that could be cited, but it is selected because of the great potential of their work to the classi- fication of the genus. References ALSTON, A. H. G. 1934. Notes on Selaginella, VI. J. Dot. 72: 223-226. ALSTON, A. H. G., A. C. JERMY, AND J. M. RANKIN. 1981. The genus Selaginella in tropical South America. Bull. Brit. Mus. (Nat. Hist.) Bot. 9: 233-330. BAKER, J. G. 1887. Handbook of the fern-allies. George Bell & Sons, London. JERMY, A. C. 1986. Subgeneric names in Selag- inella. Brit. Fern Gaz. 13: 117, 118. KOLLER, A. L., AND S. E. SCHECKLER. 1986. Vari- ations in microsporangia and microspore dis- persal in Selaginella. Amer. J. Bot. 773: 1274- 1288. SOJAK, J. 1992. Generische problematik der Se- laginellaceae. Preslia 64: 151-158. TRYON, R. M. 1 955. Selaginella rupestris and its allies. Ann. Mo. Bot. Gard. 42: 1-99. VALDESPFNO, I. A. 1993. Notes on Neotropical Selaginella (Selaginellaceae), including new spe- cies from Panama. Brittonia 45: 315-327. Key to Species of Selaginella a. Leaves of branches and stems more or less uniform, spirally arranged; main stem prostrate, creeping b b. Leafy stem radially symmetrical; leaves all alike; base of the lower leaves abruptly adnate, distinct from the stem 1 . S. sellowii b. Leafy stem somewhat dorsiventral; upper leaves differ from the lower leaves on the same portion of the stem; base of the lower leaves strongly decurrent on the stem 2. S. peruviana a. Leaves of the branches and often of the main stem dimorphic, in 4 ranks with 2 smaller median ranks on the upper side of the leafy stem and 2 larger lateral ranks; main stem erect, assurgent at the apex, to prostrate c c. Main stem very short, erect, bearing leafy branches that form a rosette; branches strongly involute when dry 36. S. convolute c. Main stem long, erect, assurgent at the apex, to prostrate; branches straight or nearly so when dry d d. Main stem erect, uniformly red or reddish, especially below the basal branch, not articulate e e. Acroscopic edge of lateral leaves ciliate or ciliolate, at least near the base; erect stem and branched portion ca. 10-25 cm, mostly 10-15 cm long 21. S. erythropus 68 FIELDIANA: BOTANY e. Acroscopic edge of lateral leaves minutely denticulate to essentially entire; erect stem and branched portion mostly 30-65 cm long 22. S. haematodes d. Main stem erect to prostrate, brownish, greenish, or stramineous, rarely red and then articulate (with a break, a dark ring, a swollen area, or usually a constriction) f f. Main stem climbing or scrambling to 8 m long, articulate 35. S. exaltata f. Main stem erect to prostrate, not exceeding 1 m long, articulate or not g g. Strobilus with sporophylls only on the upper side of the leafy stem 10. S. ramosissima g. Strobilus tetragonous, sporophylls spirally arranged on the stem h h. Branches pubescent, especially on the branch axils, trichomes sometimes sparse . . . 30. S. articulata h. Branches glabrous i i. Leaves tightly clasping the stems when dry 9. S. microphylla i. Leaves spreading, ascending or appressed, not clasping the stems when dry . . . . j j . Plants with an erect main stem and often a flabelliform arrangement of branches; basal part of the main stem usually with appressed leaves, these dimorphic or not; rhizophores usually confined to the basal portion of the main stem, or rarely above the first branch k k. Ultimate leafy branches 5-20 mm broad 1 1. Axis of primary branches fractiflex, branches sometimes flagelliform; main stem articulate 3 1 . S. parkeri 1. Axis of primary branches straight or nearly so, branches not flagelliform; main stem not articulate m m. Median leaves more or less acute to acuminate, not or hardly aristate n n. Median leaves inequilateral, the outer side larger, the costa acentric 24. S. praestans n. Median leaves equilateral, the sides nearly equal in size, the costa central 23. S. quadrifaria m. Median leaves aristate o o. Primary branches usually 1 -pinnate, sometimes 2-pinnate with branches nearly parallel from the first branch to the apex of the main stem 16. S. bombycina o. The main stem and branches irregularly or dichotomously divided P p. Branches usually few, erect, of irregular lengths, not forming a flabelliform arrangement 17. S. chrysoleuca p. Branches forming a broad, flabelliform arrangement 1 8. S. speciosa k. Ultimate leafy branches 5 mm broad or less q q. Axis of primary branches fractiflex r r. Median leaves oblong, with a basal auricle that is entire or nearly so 32. S. geniculata r. Median leaves ovate-lanceolate, with a ciliolate basal auricle 34. S. stellata q. Axis of primary branches straight or nearly so s s. Main stem articulate 33. S. asperula s. Main stem not articulate t t. Base of the main stem slender, 0.25-0.50 mm broad; leaves short, the longest 2 mm long . . 13. S. novae-hollandiae t. Base of the main stem stouter, 0.75-3.5 mm broad; longest leaves 2-5 mm long u u. Median leaves aristate 25. S. haenkeana u. Median leaves acute to acuminate, rarely a few aristate . v TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 69 v. Leaves of the main stem distal to the second branch, strongly ascending to spreading (patent); stem and leaves near the axil of the first branch usually 4-5 mm broad 19. S. lechleri v. Leaves of the main stem distal to the second branch, appressed; stem and leaves near the axil of the first branch 1-3 mm broad 20. S. anceps j. Plants with a prostrate or assurgent, sometimes intricate, or rarely erect stem; basal part of the main stem with dimorphic leaves; lateral leaves spreading; rhizophores usually borne throughout the main stem, or at least above the first branch w w. Main stem articulate x x. Leafy branches slender, 2-4 mm broad, often intricate y y. Lateral leaves with ciliolate margins at the base, without a prolonged auricle 12. S. lingulata y. Lateral leaves with entire margins, with 1 or 2 prolonged, ciliolate auricles 11. S. diffusa x. Leafy branches 4-20 mm broad, not intricate z z. Leafy branches 8-20 mm, often 10 mm broad aa aa. Base of the lateral leaves without prolonged auricles 28. S. trisulcata aa. Base of the lateral leaves with 1 or 2 prolonged auricles, the inner sometimes an acute lobe bb bb. Leafy main stem 12 mm or less broad, glabrous; branches forming an open, dichotomous arrangement 29. S. poeppigiana bb. Leafy main stem 1 5-20 mm broad, often pubescent beneath, especially on the axils, trichomes sometimes sparse; branches at the apex forming a broad, compact, flabellate arrangement 30. S. articulata z. Leafy branches 4-8 mm, usually 5 mm broad cc cc. Apical portion of the main stem, and often of the primary branches, forming a somewhat rhomboid arrangement, the apex acute to acuminate 26. S. kunzeana cc. Apical portion of the main stem, and often of the primary branches, forming a broadly ovate to obdeltate arrangement, the apex obtuse to truncate 27. S. silvestris w. Main stems not articulate dd dd. Median leaves with a short, acute apex often extended in a cusp 5. S. truncata dd. Median leaves aristate or subaristate ee ee. Edges of lateral leaves entire, denticulate, or ciliolate ff ff. Leafy branches, when present, 5-20 mm, mostly ca. 1 5 mm broad; main stem with no, or rarely few primary branches, these strongly ascending, mostly dichotomous 17. S. chrysoleuca ff. Leafy branches, 3-12 mm, mostly ca. 5 mm broad; main stem with many, spreading branches gg gg. Stems erect; rhizophores confined to the base of the main stem or not beyond the first branch hh hh. Ultimate branches short, the apex acute 1 5. S. flagellata hh. Ultimate branches longer, with the apex truncate to somewhat rounded 14. S. xiphophylla gg. Stems prostrate, creeping, sometimes intricate, forming a mat; 70 FIELDIANA: BOTANY rhizophores extending to the stem apex, or above the first branch to lh the length of the stem ii ii. Stems usually intricate, forming a mat; penultimate leafy branches 2-4 mm broad 6. S. t a ra pot ens is ii. Stems elongate, discrete; penultimate leafy branches 4-10 mm broad jj jj. Median leaves orbicular, the base truncate; lateral leaves oblong with an obtuse apex 8. S. product a jj. Median leaves ovate to oblong, the base with an ex- tended, round lobe; lateral leaves elongate-ovate, with an acute apex 7. S. seemannii ee. Edges of lateral leaves ciliate, the cilia most abundant from the base to the mid-part of the leaf kk kk. Median leaves without a prolonged auricle .... 3. S. revoluta kk. Median leaves with a prolonged auricle 11 1 1 . Main stem short, erect; rhizophores at base of the stem or mostly below the first branch ... 13. S. novae-hollandiae 1 1 . Main stems creeping, often intricate and forming a mat; rhizophores extending above the first branches . . 4. S. brevifolia 1 . Selaginella sellowii Hieron., Hedwigia 39: 306. 1900. LECTOTYPE (by R. Tryon, 1955): Brazil, Paria de San Diego Sellow, in 1821 (holotype, B; frag., NY). Selaginella rupestris f. amazonica Milde, Fil. Eur. At- lant. 263. 1867. TYPE: Peru, Cajamarca, (Rio) Maranon, Bonpland (holotype, B; isotype, BM). Selaginella amazonica (Milde) Hieron., Hedwigia 39: 310. 1900, not S. amazonica Spring, 1840. Selaginella mildei Hieron., in Engler & Prantl, Nat. Pflanzenfam. 1 (4): 671. 1902, nom. nov. for S. amazonica (Milde) Hieron. Main stem prostrate, creeping, assurgent at the tip, not articulate, pinnately branched throughout, green to brownish, glabrous. Primary branches mostly simple, 1 -pinnate, or often 2-pinnate, the ultimate branches as broad as the main stem or nearly so, 1-1.5 mm broad including the leaves. Rhizophores throughout the stems, mostly at the axils of the branches. Leaves similar (homophyl- lous), spirally arranged, subulate, long-triangular to broadly Ungulate to lanceolate, the edges whit- ish, denticulate, the apex setate, the seta opaque, milky-white, the base abruptly adnate, ciliate or ciliolate, rarely lacking cilia, abruptly distinct from the stem in form and color, glabrous. On exposed or wooded, rocky bluffs, or among stones, from ca. 300 to 2900 m, Lambayeque, Ca- jamarca, Ancash, and Cuzco. Central Mexico, Cuba, Venezuela, and Colom- bia south to Argentina, and eastern Brazil. The species belongs to the Sartorii series of the Selaginella rupestris group and appears most closely allied to S. sartorii Hieron. Geographically it is wide-ranging from Mexico and Cuba through South America. It is readily distinguished from the nine others of series S. sartorii and other Peruvian spe- cies by the glabrous leaf bases, especially by the opaque, milky-white setae on the leaves. This and S. peruviana have elongate stems that differ from the radially symmetrical plants of S. convoluta. Lambayeque: Hacienda Valor, Ellenberg 3613 (GH). Cajamarca: between Llaconora and Namora, Correll & Smith P893 (GH, MO). Between Jaen and Quemado, Fe- rreyra & Sanchez 19668 (USM). Ancash: Mancos, Yun- gay, Tryon & Tryon 6551 (GH). Cuzco: Biies 608 (us); Herrera 3009a (us). 2. Selaginella peruviana (Milde) Hieron., Hed- wigia 39: 307. 1900. Figure 9a-b. Selaginella rupestris f. peruviana Milde, Fil. Eur. At- lant. 263. 1891. TYPE: Peru, Huanuco, Ruiz 98 (holotype, B; isotype, NY). Selaginella peruviana var. dombeyana Hieron., Hed- wigia 39: 308. 1900. PARATYPE: Peru, Dombey 14 (B). Selaginella sheldonii Maxon, Proc. Biol. Soc. Wash. 31: 171. 1918. TYPE: United States, Quanah Mountains, Oklahoma, Sheldon 233 (holotype, us). Main stem prostrate, creeping, not articulate, pinnately branched throughout, glabrous, green to brownish. Leafy branches forming compact to dif- TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 71 fuse mats with discrete, simple or 1 -pinnate, some- times 2-pinnate stems. Rhizophores borne throughout the stems at the axils of branches. Leaves generally similar (homophyllous), spirally arranged, setate, the seta usually more or less lu- tescent, or often not well differentiated in color, the base adnate, glabrous, sometimes denticulate, the edges usually ciliate, or sometimes somewhat pubescent, leaves on the underside of the stem differ from those on the upper side, on the same part of the stem, those on the upper side linear, acuminate to long-triangular, the lower leaves su- bulate, acuminate, broadest at or near the base, to linear-lanceolate, gradually distinct from the stem. Mats on bluffs, dry rocky slopes, ledges or crev- ices of cliffs, exposed or in light shade, igneous, calcareous, or sandstone rock, 1750—4400 m, Ca- jamarca south to Puno. Southwestern United States south to Puebla, Mexico; Peru to Argentina. The species is recognized as the least specialized and basic species within the series Eremophilae, in the treatment of the Selaginella rupestris group (Tryon, 1955). Geographically it is the widest ranging species of that series. On the basis of the slender leaves bearing long setae, it is most closely related to S. arizonica Maxon, a species of north- west Mexico and the states of Texas and Arizona. However, it is distinguished from this as well as Peruvian selaginellas by the lower leaves that are strongly decurrent on the stem. Among the Pe- ruvian selaginellas, it is most closely allied with S. sellowii, another species with homophyllous leaves. Cajamarca: Between Cajamarca and San Juan, Mutter & Gutte 8869 (USM). Ancash: Near Mancos, Tryon & Tryon 6550 (GH). Huanuco: Prov. Huanuco, Plowman & Rury 11103 (F, GH, USM), Prov. Huanuco, Stork & Morton 9388 (F, uc). Lima: Prov. Canta, Punte San Jose, Lopez & Riccio 10080 (GH). Prov. Huarochiri, Leon 583 (USM). Junin: Above Conception, Correll & Smith P748 (GH, MO). Prov. Tarma, lit is et al. 108 (GH, wis). Apu- rimac: Prov. Andahuaylas, Stork & Morton 10719 (F, uc). Cuzco: Near Pisac, McDaniel & Gorski 11402 (GH, MO). Urubamba valley, Leon 443 (F, USM). Prov. Uru- bamba, Chicon, Vargas 11058 (F, uc). Puno: Near Puno, Solomon 2895 (F, MO). 3. Selaginella revoluta Baker, J. Bot. 21:141.1883. TYPE: Venezuela, Amazonas, near Mau- pures, Spruce 3621 (holotype, K; isotypes, CGE, us). Selaginella demissa Christ, Bull. Herb. Boissier 2, 1: 75. 1883. TYPE: Peru, Cerro Canchahuaya, Hu- ber 1421 (holotype, p). Main stem short, creeping, often forming com- pact mats, not articulate, stramineous, glabrous, the basal part with spreading leaves, and often smaller, appressed, ascending leaves. Primary branches short, distant, usually 1 -2-pinnate, ex- tending to the base of the main stem, 3-5 mm broad including the leaves, the terminal branches short. Rhizophores extending from the base of the main stem to the apex of the branches. Lateral leaves above the first branch not imbricate, spreading, elongate-ovate, broadest at the base, 0.5-0.2 mm long, ca. 3 x longer than broad, acute, with a round, ciliolate, scarcely prolonged basal lobe, without conspicuous whitish borders, the edges with long cilia, these denser at the base. Median leaves suborbicular, broadest at the cen- ter, aristate, the arista '/2 as long as the lamina, cuspidate, the auricles not prolonged, often with narrow, whitish borders, the edges denticulate. In wet, primary forests, 100-1100 m, Loreto and Huanuco. Costa Rica and Panama; in South America from the Guianas south to Peru and Brazil. The compact, creeping habit of the plants with short, lateral branches bearing discrete, wide- spread leaves along the main stem form a dis- tinctive growth pattern. The species was allied with Selaginella product a by Alston et al. (1981), but distinguished by the strongly tapering, ciliate, more acute lateral leaves, sometimes pubescent on the upper surface near the margin. However, that spe- cies also has broader, more delicate leafy stems. Loreto: Prov. Maynas, Mishuyacu, Klug 1220 (F, us). Prov. Maynas, below Iquitos, Moran 3651 (MO). Gami- tanacocha, Schunke 322 (F, GH, us, USM). Huanuco: Tin- go Maria, Allard 20485 (MO), 20846 (us). 4. Selaginella brevifolia Baker, J. Bot. 21:83.1883. TYPE: Brazil, Janarate Cochoeira, Rio Ne- gro, Spruce 2547 (holotype, K; isotypes, BM, CGE). Main stem prostrate, creeping, it and its branch- es forming a mat of stems, sometimes elongate to 15 cm, not articulate, stramineous, glabrous, the basal part with spreading leaves and often smaller, appressed, ascending leaves. Primary branches 2- 3 -pinnate, alternate, distant, extending to the base of the main stem, 2-3 mm broad including the leaves. Rhizophores above the first branch, up to 72 FIELDIANA: BOTANY '/3 the length of the stem. Lateral leaves ovate, acute, ca. 1-2 mm long, without prolonged auri- cles, ciliate especially on the acroscopic margin, the cilia usually long, discrete. Median leaves ovate- elliptic, acuminate to aristate, the arista '/4 the lam- ina length, the base with a broad, long, ciliolate auricle, without conspicuous whitish borders, the edges denticulate. On soil and among rocks, along roadsides, 400- 2400 m, Cajamarca, San Martin, and Huanuco. Venezuela, Colombia, Peru, and western Ama- zonas, Brazil. Although the plants are small and mosslike in habit, growing over soil or among rocks, they are well collected and known to be wide-ranging in South America and in Peru. The species is considered close to the Greater Antillean Selaginella cordifolia but with longer aristae on the median leaves (Alston et al., 1981). It also resembles S. novae- hollandiae, but the strongly prostrate habit with rhizophores above the median part of the main stem contrast with the erect habit and rhizophores, mainly at the base of the stem in S. novae- hollandiae. Cajamarca: Prov. Contumaza, Guzmango, Sagdstegui 3932 (GH). Prov. Cajamarca, El Molino, Sagdstegui 7995 (F). Contumaza, Lleden, Sagdstegui et al. 10887 (F). San Martin: Boqueron Pass, between Tingo Maria and Pu- calpa, Allard22124 (us). Huanuco: Cushi, trail to Tambo de Vaca, Bryan 718 (F, us). 5. Selaginella truncata A. Braun, Index Sem, Hort. Bot. Berol. Appendix 1857: 15. SYNTYPES: Colombia, Cundinamarca, Bogota, Andes of New Grenada, Karsten (B, BM); Susumuco, Triana 696 (BM), 238 (NY). Selaginella weberbaurei Knox, Trans. Bot. Soc. Ed- inburgh 35:282. 1950. Only spores are described. "TYPE": based on a specimen collected by We- berbauer, named by Hieronymus, B?. Main stem prostrate, assurgent, not articulate, stramineous, glabrous, the basal part with spread- ing leaves and often smaller appressed, ascending leaves. Primary branches 1-2-pinnate, pinnately branched, extending to the base of the main stem, the central branches distant on the main stem, strongly ascending, forming an open, dichotomous division of the branches, the ultimate branches as broad as the main stem, 4-6 mm broad including the leaves. Rhizophores slender, threadlike, throughout the main stem. Lateral leaves uniform in length, 2-3 mm long, oblong to broader near the base, the apex obtuse, the base truncate, with narrow, whitish borders and delicate, long cilia at the edges, especially dense at the base. Median leaves ovate, the apex acute, sometimes with a short cusp, the base truncate, with whitish borders, the edges regularly ciliate or ciliolate, more strong- ly so at the base. Epiphytic in dense jungle, or on rotted logs, in dense forests, 365-700 m, San Martin and Cuzco. Venezuela and Colombia south to Bolivia. The compact, deep green leaves, imbricate on the stems, give a distinctive aspect to the plants. Ecological information on collections indicates the plants are epiphytic on trees 15 ft above the ground. This is corroborated by the dense rhizo- phores on the branches and the stems that are free from soil. Selaginella applanata A. Braun, Ann. Sci. Nat. Bot. 5, 3: 274. 1865, may be a synonym; uc has a fragment of an isotype, Peru, Puno, San Gavan, (Gaban) Lechleri 2405. San Martin: Prov. Lamas, San Antonio, Belshaw 3569 (H, GH, uc, us). Guayrapurima, Tarapoto, Spruce 4024 (GH, us). Pasco: Prov. Oxapampa, Cordillera de San Ma- tias, D. Smith 2000 (F). Cuzco: Prov. Quispicanchis, entre Inombaris y Quincemil, Vargas 1 1690 (GH); entre Quin- cemil y San Lorenzo, Vargas 16481 (GH). 6. Selaginella tarapotensis Baker, J. Bot. 21: 98. 1883. TYPE: Peru, Mt. Guayrapurima, near Tarapoto, Spruce 4625 (holotype, K; isotypes, BM, CGE, us). Main stem prostrate or assurgent at the tip, often forming a mat of stems, not articulate, stramin- eous, glabrous, the basal part with spreading leaves, and often smaller, appressed, ascending leaves. Primary branches 3-4-pinnate, alternate, distant, extending to the base of the main stem, 2-4 mm broad including the leaves, the ultimate branches short. Rhizophores filamentous, dense at the base, extending nearly to the stem apex. Lateral leaves oblong-lanceolate, but not imbricate on the main stem, 0.5-2.0 mm long, approximately more than 3 x longer than broad, obtuse to acute, the auricles not prolonged, without conspicuous whitish bor- ders, the edges short-denticulate. Median leaves rhomboidal to ovate, aristate, the arista slender, usually whitish, '/2 as long to nearly equal the length of the lamina, the base not or hardly prolonged, often with narrow, whitish borders that extend into the whitish arista, the edges denticulate. TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 73 Rarely epiphytic, in deep cloud forests, and on shaded banks bordering roads, 500-1 200 m, Ama- zonas and San Martin south to Cuzco. Costa Rica south to Bolivia. The truncate leaf bases on both lateral and me- dian leaves, along with the rhomboid shape and prominent aristata on the median leaves, distin- guish the species. This is one of the slender- stemmed, mat-forming species most closely re- sembling Selaginella revoluta. Selaginella calosticha Spring, Nouv. Mem. Acad. roy Sci. Belg. 24: 206. 1849. TYPE: Venezuela, Caracas, Funck & Schlim 3321 (K), may be an earlier name for this species. Two Allard collec- tions from Peru— 20821 and 21206— are cited as S. calosticha in Alston et al. (1981). Amazonas: Prov. Bagua, Montenegro, Hutchison & Wright 3822 (F, GH, uc, us). Monte Campana, near Tara- poto, Spruce cf4625 (us). San Martin: Tocache Nuevo- Juanjui, Smith et al. 2132A (uc). Huanuco: Prov. Tingo Maria, Cueva de las Pavas, Aldave & Fernandez 5578 (GH). Pasco: Oxapampa, Cordillera San Matias, Leon et al. 321 (F, MO, USM). Junin: Satipo, Pichanaki, Leon 222 (F, USM). Cuzco: Prov. Paucartambo, Quebrada Quita Calzon, Leon et al. 2936 (USM). Achirani, Vargas 11147 (F). 7. Selaginella seemannii Baker, J. Bot. 21: 244. 1883. TYPE: Colombia, Choco, Cacaqual Is- land, Seemann 1006 (holotype, K; isotype, BM). Main stem assurgent at the tip, ca. 5-25 cm long, not articulate, stramineous, glabrous, the basal part with spreading leaves, and often smaller, ap- pressed, ascending leaves. Primary branches 2-3- pinnate, arising a short distance above the base of the main stem, sometimes gradually reduced, forming a flabelliform arrangement, the ultimate leafy branches short, unequal, 4-6 mm broad in- cluding the leaves. Rhizophores mostly near the base to the mid-portion of the main stem. Lateral leaves elongate-ovate, broadest at the base, spreading above the first branch, approximate, imbricate toward the apex, 2-3 mm long, mem- branous, acute, the base truncate or 1 side enlarged and rounded, with narrow, whitish, denticulate edges, or sparsely denticulate along the base. Me- dian leaves elongate-ovate, the apex acuminate to subaristate, the arista less than '/4 the lamina length, the base truncate or with an enlarged, rounded outer side, with narrow, whitish, denticulate edges, dentate at the base. On open floor of dense forest, in deep ravines, or dense, seasonally flooded forest, 130-625 m, Loreto and Huanuco. Costa Rica, Colombia, Suriname, Ecuador, and Peru. The flabelliform arrangement of the branches, with rhizophores concentrated at the base of the stem, distinguishes this from other slender- stemmed species such as S. producta and S. re- voluta that clearly are prostrate, creeping. Although the species is reported to be wide- ranging from Costa Rica to Peru, we have seen collections only from Loreto, and Huanuco in Peru. Loreto: Prov. Maynas, Calentura, Killip & Smith 29141 (us). Prov. Maynas, Iquitos, Tryon & Tryon 5195 (F, GH, us, USM). Prov. Maynas, Yanamono Tourist Camp, van der Werffet al. 9959 (uc). Rio Itaya, LI. Williams 238 (F, us). Huanuco: Tingo Maria, La Cueva de las Pavas, Allard 20514 (us). 8. Selaginella producta Baker, J. Bot. 21: 243. 1883. LECTOTYPE (chosen by Alston et al., 1981): Brazil, Amazonas, between Barcellos and San Gabriel, Spruce 2043 (holotype, BM; isotype, CGE). Main stem prostrate or assurgent, 1 0-20 cm long, not articulate, several stems usually arising at the base of the plants, stramineous to somewhat green- ish, glabrous, the basal part with spreading leaves and often smaller, appressed, ascending leaves. Primary branches 2-3 -pinnate, with distant sec- ondary divisions spreading, sometimes forming a broad, flabelliform arrangement, or with elongate, alternatively pinnate branches as long as the main stem or nearly so, 5-10 mm broad including the leaves, the ultimate leafy branches short, unequal in length. Rhizophores mostly at the stem base, or extending to the central part of the main stem and branches. Lateral leaves oblong to ovate, spread- ing, approximate to somewhat imbricate above the first branch, 2-5 mm, usually 4 mm long, broader at the base, obtuse to subacute, the base truncate, or with an enlarged, round basal lobe, without conspicuous whitish borders, the edges sparsely denticulate with the teeth somewhat dens- er at the base. Median leaves suborbicular, broad- est at the center, the apex acuminate to subaristate and cuspidate, the arista usually V* or less the lam- ina length, the base truncate, with or without nar- row, whitish margins, the edges evenly denticulate. Terrestrial, or a trunk epiphyte, on rotted logs, or commonly on white sand of the forest floor, 100-860 m, Loreto, Huanuco, and Pasco. 74 FIELDIANA: BOTANY Tobago and Trinidad, French Guiana west to Colombia and south to Peru and Brazil. The species is compared to Selaginella revoluta by Alston et al. (1981). However, it differs in broader lateral leaves that have a truncate base and lack cilia. The imbricate leaves covering most of the branches resemble those of S. truncata but are clearly broader and may have a whitish sur- face. Loreto: Prov. Maynas, Iquitos, Killip & Smith 27322 (GH, us). Iquitos, Klug 198 (F, us). Iquitos, Revilla 4285 (F, MO). Iquitos, McDaniel et al. 22107 (F). Mishana, Solomon 3595 (MO). Prov. Maynas, Alpahuayo, van der Werff et al. 10262 (F, MO). Huanuco: Rio Llullapichis, Cerros del Sira, Wolfe 12264 (GH). Pasco: Prov. Oxa- pampa, Palcazu, Foster 9532 (MO, USM). 9. Selaginella microphylla (HBK.) Spring, Bull. Acad. roy Sci. Bruxelles 10: 234. 1843. Lycopodium microphyllum HBK., Nov. gen. sp. 1: 37. 1816. TYPE: Colombia, Cauca, Qulcace, Bon- pland (holotype, p?, photo, GH; isotype, BM). Main stem short, creeping, usually it and its branches forming a compact mat, not articulate, stramineous or greenish, glabrous. Primary branches often intricate, 1-2-pinnate, short, dis- tant, extending to the base of the main stem, slen- der, ca. 0.5 mm broad, including the leaves. Rhi- zophores abundant at the base, extending nearly to the apex of the main stem and branches. Lateral leaves extending nearly to the base of the stems, minute, inserted obliquely, 0.25-0.50 mm long, ovate or ovate-elliptical, tightly clasping the stem and enveloping the median leaves when dry, im- bricate, less so on the ultimate branches, the apex acute with a more or less extended cusp, the bor- ders whitish, ciliolate to fimbriate along the edges, especially at the base. Median leaves ovate, about half as long as the lateral leaves, subacute, not auriculate, the edges with regular, dense cilia, the border broad, whitish. Terrestrial in open forest, on soil or among damp rocks, in deep shade, 800-2700 m, Cajamarca, La Libertad, Huancavelica, Apurimac, and Cuzco. Costa Rica and Panama, Venezuela along the Andes south to Argentina, east to southern Brazil, Paraguay and Uruguay. The abundant cilia along the borders of the leaves, and especially the lateral leaves that strong- ly clasp the dry stems, readily characterize the spe- cies. The slender, intricate stems have a growth form that resembles some species in the Selagi- nella rupestris group. The species is known from a broad geographic range through South America and has been widely collected in Peru. Specimens from southern Co- lombia collected by Bonpland on his travels in South America with Humboldt represent the type. Cajamarca: Prov. Contumaza, El Tunel, Sagdstegui et al. 12565 (MO). La Libertad: Prov. Pataz, Vista Florida, Leon & Young 1086 (USM). Huancavelica: Prov. Tuya- caja, Surcubamba, Tovar 3696 (GH). Apurimac: Prov. Abancay, Vargas 16584 (GH). Cuzco: Prov. Convention, Sahuayaco, Biles 834 (us). Urubamba Valley, San Mi- guel, Cook & Gilbert 1782 (us). 10. Selaginella ramosissima Baker, J. Bot. 23: 295. TYPE: Peru, San Martin, near Tarapoto, Spruce 4088 (holotype, K; isotypes, BM, GH, NY). Main stem erect, or assurgent at the tip, short, less than 10 cm long, not articulate, greenish, gla- brous, the basal part with spreading leaves and often smaller, appressed, ascending leaves. Leafy branches 2-pinnate, ascending, longest in the cen- tral part of the stem, the secondary branches straight, ascending, forming a short, deltate ar- rangement of the stems, ca. 3 mm broad. Rhizo- phores mostly at the base extending to the first branch. Lateral leaves slender, elongate, broadest at the center, closely placed to somewhat imbricate especially on the secondary branches, oblong, 0.5- 1 .0 mm long, membranous, acute, the base trun- cate, not prolonged, with narrow, whitish borders, the edges denticulate especially at the base. Me- dian leaves slender, elongate-ovate, or somewhat broader at the center, membranaceous, acuminate to aristate, the arista '/4 or more the length of the lamina, the base more or less equal, without a prolonged auricle, the borders narrow, whitish, the edges denticulate. Strobilus dorsiventral with 2 ranks of sporophylls on the upper surface of the leafy stem. Along roadside or bank of stream below the forest, 200-2700 m, Cajamarca, San Martin, Lima, and Cuzco. Ecuador and Peru. The strobilus is dorsiventral with two ranks of sporophylls on the upper part of the leafy stem. This readily distinguishes the species from all oth- ers in Peru. The thinner, membranaceous texture of the leaves also is a characteristic of the Peruvian plants. TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 75 Cajamarca: Prov. Hualgayoc, Soukup 3872 (us). Lima: Prov. Cantua, Acleto 758 (USM). Cuzco: Biies in Junio, 1930(F). 1 1 . Selaginella difTusa (Presl) Spring, Bull. Acad. roy. Sci. Bruxelles 10: 143. 1843. Lycopodium diffusum Presl, Reliq. haenk. 78. 1825. TYPE: Panama, Haenke (holotype, PR; isotype, B). Selaginella eggersii Sodiro, Crypt, vase. Quit. 605. 1893. TYPE: Ecuador, Valle Takatanga, Sodiro (holotype, not found; isotype, us). Selaginella atirrensis Hieron., in Engler & Prantl, Nat. Pflanzenfam. 1 (4), 71 1. 1 90 1 . LECTOTYPE: (by Alston, 1955), Costa Rica, Cartago, near Atirro, Donn. Smith 5103 (holotype, us; isotype, NY). Main stem elongate, prostrate, assurgent at the tips, sometimes compact and matlike, articulate, stramineous, glabrous, the basal part with spread- ing leaves, and often smaller, appressed, ascending leaves. Primary branches distant, ascending, un- equally forked, 2-3-pinnate, extending to or nearly to the base of the main stem, 2-4 mm broad in- cluding the leaves, the ultimate branches short. Rhizophores hairlike, extending nearly to the apex of the branches. Lateral leaves spaced or approx- imate, not imbricate, 1-2 mm long, elliptical-lan- ceolate, acute, short, with a prolonged auricle, of- ten densely ciliolate, sometimes with whitish borders, the edges sparsely ciliolate to nearly en- tire. Median leaves elongate-ovate, broadest near the middle, the apex long-acuminate, the base with a prolonged, Ungulate auricle that is usually cilio- late, with or without a whitish border, the edges finely denticulate. In moist places in deep forest, on steep banks, on logs or among rocks, sometimes forming large mats, 1100-2100 m. Huanuco, and Pasco south to Cuzco. Costa Rica; Trinidad; South America east to Suriname, and in the Andes south to Bolivia. Plants appear to have lateral leaves of more than one form. Most abundant are slender, elliptical leaves with entire margins. Other, larger leaves are broader with the base abundantly ciliolate, and the cilia sometimes extending along the margins to the mid-region of the leaf. The ciliolate base resembles that of the median leaves. Selaginella atirrensis Hieron. is included here although it has been recognized as a distinct spe- cies on the basis of its acute median leaves. It is not easily distinguished on leaf shape and possibly intergrades with S. diffusa. Huanuco: Tingo Maria, Croat 21245 (MO, uc). Churubamba, Mexia 8256 (F, GH, MO, uc, us). Tingo Maria, Tryon & Tryon 5227 (F, GH, us, USM). Junin: near La Merced, Killip & Smith 23949 (F, GH). Chanchamayo valley, Schunke 197 (F). Tarma, Pan de Azucar, Velarde 5490 (GH). Pasco: Prov. Oxapampa, between Oxapampa and Paucartambo, D. Smith 1466 (MO). Ucayali: Bo- queron, Ferreyra 8128 (USM). Cuzco: Prov. La Conven- cion, Rio Mapitunuari, Dudley 11352 (GH, MO). Kos- nipata, Sta. Isabel, Vargas 23011 (GH). 12. Selaginella lingulata Spring, Nouv. Mem. Acad. roy. Sci. Belg. 24: 224. 1849. TYPE: Ecuador, Pichincha, Jameson (holotype, K; isotype, BM). Selaginella intacta Baker, J. Bot. 21: 335. 1883. TYPE: Ecuador, San Nicolas, Sodiro (holotype, K). Main stem prostrate, intricate, usually compact, forming a dense mat, articulate, stramineous, gla- brous, the basal part with spreading leaves and often smaller, appressed, ascending leaves. Leafy branches 1-3-pinnate, usually pinnately branched, distant, extending to the base of the main stem, the ultimate branches short, 2-4 mm broad. Rhi- zophores extending nearly to the stem apex. Lat- eral leaves closely placed to distant, 1-3 mm long, spreading on the upper part of the stems and pen- ultimate branches, oblong, acute, the basal lobes more or less equal, not prolonged, with long, dense cilia, without whitish borders, the edges entire. Median leaves elongate-ovate, the apex usually long-acuminate, the base with 2 equally long au- ricles, ciliate at the base, without conspicuously whitish borders, the edges denticulate above. On soil or among rocks, in shade, sometimes forming dense mats on hillsides or on steep banks, sometimes in deep forests, 1000-1300 m, Pasco and Junin. Colombia south to Bolivia. The long marginal cilia at the base of the leaves form a small fringe especially on the lateral leaves. The median leaves are clearly more slender than the lateral ones and have a long-acuminate apex in the material examined. However, the apex of the median leaves was considered as acute in the treatment by Alston et al. (1981). The axillary leaves are distinctive, somewhat larger than the median and lateral leaves, and more or less ovate, acute, usually with one prolonged auricle and few long cilia. Pasco: Pichis trail, San Nicolas (as Junin), Killip & Smith 26056 (F, GH, us). Junin: Above San Ramon, 76 FIELDIANA: BOTANY Schunke A251 (us), Chanchamayo Valley, Schunke 230 (us), 755 (us). Prov. Tarma, San Ramon, Tryon & Tryon 5448 (F, GH, us). 13. Selaginella novae-hollandiae (Sw.) Spring, Bull. Acad. roy. Sci. Bruxelles 10: 234. 1843. Lycopodium novae-hollandiae Sw., Syn. fil. 184, 410. 1806. TYPE: "Nova Hollandiae," probably an error for Nova Granada, not located. Selaginella pearcei Baker, J. Bot. 22: 246. 1884. TYPE: Peru, Huanuco, Cordilleras of Pozuzo, Pearce 249 (holotype, K). Selaginella chionoloma Crabbe & Jermy, Amer. Fem J. 63: 137. 1973. TYPE: Peru, Cuzco, Cuquipata, Herrera 1636 (holotype, us; isotype, BM). Main stem short, erect or assurgent, usually less than 20 cm long, not articulate, stramineous, gla- brous, the basal part with spreading leaves and often smaller, appressed, ascending leaves, or the basal part with only appressed leaves. Primary branches 2-3-pinnate, often forming a complex system of branches broadest at the base, the pen- ultimate branches straight or nearly so, 1-3 mm broad including the leaves. Rhizophores at base of the main stem, mostly below the first branch. Lat- eral leaves 0.5-2.0 mm long, elliptical to some- what ovate, acute, not auriculate, the base cilio- late, sometimes with narrow whitish borders, the edges ciliolate, often densely so. Median leaves ovate, usually more or less elongate, the apex acu- minate to aristate, the arista '/4 the lamina length, the base with a prolonged, Ungulate, ciliolate au- ricle, or a somewhat enlarged, round, ciliate base, the borders narrow, whitish, the edges ciliolate or denticulate. On damp, shaded ledges or crevices of rock fac- es, or moist banks along streams, wooded hillsides, open woods, dense forests, sometimes carpeting the forest floor, rarely epiphytic, 400-3600 m, Lambayeque, south to Puno. Nicaragua south to Bolivia and Argentina. This differs from other slender-stemmed species in lacking articulations on the stems and having generally smaller leaves. It is one of the most com- mon species of tropical America and is wide-rang- ing in Peru. Lambayeque: Along Olmos-Jaen road, Correll & Smith P826 (GH, MO, us). Cajamarca: El Molino, Sagdstegui et al. 7995 (F, MO, uc). Amazonas: Prov. Bagua, Chacha- poyas, Hutchison & Wright 5782 (F, GH, MO, uc, us, USM). La Libertad: Prov. Otuzco, road to Paranday, Lopez 1044 (us). San Martin: Rio Sion, Schunke 3494 (F, us). Loreto: Balsapuerto, Killip & Smith 28488 (F, GH). Hua- nuco: Tingo Maria (as San Martin), Allard 21130 (MO, uc). La cueva de las Pavas, Schunke 3257 (F, GH, us). Lima: Matucana, Bryan 48 (F, MO). Junin: Prov. Tarma, San Ramon, H. Iltis & C. Iltis 18 (GH, wis). E of Quimiri Bridge, Killip & Smith 23945 (F, GH). Ucayali: Boqueron Padre Abad (as Loreto), Schunke 3065 (F, GH, us). Huan- cavelica: Surcubamba, Tovar 3682 (GH). Ayachucho: Ccarrapa, Killip & Smith 22331 (F, GH, us). Apurimac: Prov. Abancay, Vargas 11083 (GH). Cuzco: Chincheros, King et al. 293 (USM). Machu Picchu, Leon 457 (USM). Puno: Prov. Carabaya, Ollachea, Vargas 6914 (us). 14. Selaginella xiphophylla Baker, J. Bot. 22: 296. 1884. TYPE: Peru, San Martin, Mt. Guay- rapurima, near Tarapoto, Spruce 3990 (ho- lotype, K; isotypes, BM, CGE, GH, NY). Main stem erect, ascending, 10-14 cm long, not articulate, stramineous to greenish, glabrous, the basal part with appressed, ascending leaves. Pri- mary branches 1-2-pinnate, alternate, distant, ex- tending to, or nearly to the base of the main stem, the lower branches usually longest, the penulti- mate leafy branches 6-12 mm broad. Rhizophores confined to the lower Vs-'/z of the main stem, rarely above. Lateral leaves ascending to somewhat ap- pressed, those above the first branch approximate, 2-6 mm long, oblong-lanceolate, the apex acute, the base truncate, with denticulate edges especially at the base. Median leaves ovate-lanceolate, grad- ually attenuate, aristate, the arista as long as the lamina, the base truncate, the outer side ciliolate, with narrow, whitish denticulate margins, the teeth extending onto the arista. Panama, Colombia, and Peru. We have seen only a single collection, the type material from San Martin listed above. The spe- cies is also reported from Panama and Colombia by Valdespino (1993). The small size, erect habit of the plants, the rhizophores borne in the basal l/3-l/2 of the main stem, and median leaves with an arista as long as the lamina are characteristics by which the species can be recognized. 15. Selaginella flagellata Spring, Bull. Acad. roy. Sci. Bruxelles 10: 228. 1843. TYPE: French Guiana, Rio Inini, "source of the Rio Oya- pok," Leprieur (holotype, LG; isotype, P). Selaginella regularis Baker, J. Bot. 22: 277. 1884. TYPE: Peru, San Martin, near Tarapoto, Spruce 3977 (holotype, K; isotypes, BM, GCE). Main stem erect, short, 5-20 cm long, the base 2-3 cm below the leafy branches, not articulate, TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 77 stramineous, glabrous, the basal part with spread- ing leaves, and often smaller, appressed, ascending leaves. Primary branches 2-3-pinnate, sessile or subsessile, distant, spreading, the central ones longest, the lateral branches ascending in a broad, frondlike arrangement, the penultimate leafy branches straight, 3-5 mm broad. Rhizophores basal, or mostly below the lowest branches. Lat- eral leaves 1-3 mm long, elliptic, or broadest at the base, attenuate to acute, the base truncate, the borders not conspicuously whitish, the edges den- ticulate, Median leaves ovate to somewhat elon- gate, aristate, the arista l/4 as long as the lamina, the base not auriculate, the borders narrow, whit- ish or not, the edges denticulate, especially at the base. Terrestrial, or climbing in cloud forest, among moist rocks and on cliffs, 200-2000 m, San Mar- tin, Loreto, and Cuzco. Mexico south to French Guiana and Brazil, and in the Andes to Bolivia. The short, ultimate branches clearly differ from other slender, erect-stemmed species in Peru. There is a general resemblance to Selaginella xiphophylla in the erect habit and rhizophores confined to the basal part of the main stem. San Martin: Juan Jui, Klug 4230 (F, GH, us). Loreto: Pongo de Manseriche, Mexia 6333A (uc). Cuzco: Prov. Convention, Echarate, Sues 787 (us). 16. Selaginella bombycina Spring, Nouv. Mem. Acad. roy. Sci. Belg. 24: 191. 1849. TYPE. Peru, San Martin, Matthews (Mathews) 1781 (holotype, K). Main stem erect, 25-30 cm long, 1 -pinnate, not articulate, stramineous, glabrous, the basal part with appressed, ascending leaves. Primary branch- es alternate, mostly 1 -pinnate, extending in nearly parallel alignment from the first branch to the apex of the stem, 10-16 mm broad including the leaves, the axes straight or nearly so. Rhizophores largely at the base of the main stem, rarely with a few among the main branches. Lateral leaves imbri- cate on the lateral branches, oblong, slender 5-10 mm long, 4 or 5 x longer than broad, obtuse, the base with a slightly enlarged, round lobe, usually densely ciliate, without conspicuous whitish bor- ders. Median leaves suborbicular, asymmetric, the outer side larger, aristate, the arista '/2 to equal the length of the lamina, with a somewhat longer basal auricle, the base ciliolate, without conspicuous whitish borders, cilia often dense along the outer edges. On hillsides and deep ravines of wet forests, 700-1 100 m, San Martin and Huanuco. In Peru nearly restricted to the area around Tingo Maria. Costa Rica south to Peru. The plants usually have long primary branches that are 1 -pinnate. This feature and the ciliate lat- eral leaves distinguished the species. San Martin: Prov. Mariscal Caceres, 4 km de Puerto Pizana, Schunke 4893 (F, us). Huanuco: Tingo Maria, Tryon & Tryon 5255 (GH, us). Prov. Leoncio Prado, Tingo Maria, Plowman & Kennedy 5732 (F, GH, USM). Prov. Leoncio Prado, Rupa Rupa, Schunke 10172 (F, MO). 17. Selaginella chrysoleuca Spring, Bull. Acad. roy. Sci. Bruxelles 10: 226. 1843. TYPE: Bo- livia, D'Orbigny, (holotype, P). Selaginella sprucei Hooker, Sec. cent, ferns, t. 83. 1861. TYPE: Peru, San Martin, Mt. Campana, near Tarapoto, Spruce 4623 (holotype, K; isotypes, BM, CGE, us). Main stem below the first branch short, usually less than 5 cm long, erect, not articulate, greenish, glabrous, the basal part with spreading leaves and often smaller, appressed, ascending leaves, or the basal part only with appressed, ascending leaves. Leafy branches dichotomously divided 1 or 2 x , often with 1 longer branch, sometimes with tuft- like branches arising from a slender creeping run- ner, ultimate leafy branches 5-20 mm, mostly 1 5 mm broad, axes of primary branches straight. Rhi- zophores mostly at the base, sometimes extending to l/3 the length of the stem, but not among the broad, leafy branches. Lateral leaves spreading above the first branch, imbricate, 5-10 mm long, oblong to linear, 3-5 x longer than broad, the apex acute to obtuse, asymmetrical, with the upper (ac- roscopic) edge somewhat larger, round, and finely denticulate, with narrow whitish borders, the edg- es finely denticulate. Median leaves elongate-ovate to suborbicular, asymmetrical, acuminate to aris- tate, the arista sometimes Vz the lamina length, the base truncate with a slightly enlarged, round lobe, the borders narrow, whitish, the edges finely den- ticulate. Among mosses on rocks, in damp soil, on shad- ed slopes of ravines, on moist riverbanks, and in dense, dark cloud forests, 400-2000 m, Amazo- nas, south to Cuzco. 78 FIELDIANA: BOTANY Panama, Venezuela to Bolivia. Similarities in the general aspect of the broad, leafy branches, in the asymmetrical shape of the median leaves with a long arista and dentate mar- gins, suggest an alliance with Selaginella speciosa. Amazonas: Prov. Bagua, La Peca, Barbour 2839 (MO). Prov. Bagua, near Montenegro, Hutchison & Wright 3821 (F, GH, uc, us). Above Montenegro, Wurdack 1879 (F, GH, uc, us). San Martin: Road from Tarapoto to Yu- rimaguas, Kennedy 3544 (F, us). Between Tarapoto and Yurimaguas, Knapp & Mallet 8474 (F, MO). Tarapoto- Yurimaguas road, McDaniel 13805 (GH, MO). Near Tara- poto, Spruce 4628 (GH). Ucayali (as Loreto): Between Tingo Maria and Pucallpa, Ferreyra 1008 (USM). Huanu- co: Cerros del Sira, Rio Llullapichis Watershed, Dudley 13360 (GH). Cuzco: Prov. La Convention, Dudley 10355 (GH, MO). 18. Selaginella speciosa A. Braun, Ann. Sci. Nat. Bot. 5, 3: 274. 1865. TYPE: Colombia, Bo- gota, Triana (holotype, B?; isotype, BM). Selaginella huberi Christ, Bull. Herb. Boissier 2, 1: 73. 1901. TYPE: Peru, "entre Ucayali et Hualla- ga," Huber 1547 (holotype, P). Main stem erect, 30-70 cm long, not articulate, green to brownish, glabrous, the basal part with appressed, ascending leaves. Primary branches di- chotomous, long, 1 -pinnate, with 2 or more long divisions, the lower ones divided at short intervals forming a frondlike, somewhat flabelliform ar- rangement, 10-20 mm broad including the leaves, the axes straight. Rhizophores at the base of the main stem. Lateral leaves closely placed to im- bricate, linear-oblong, to broader above the base, 8-10 mm long, 5-6 x longer than broad, asym- metrical, the acroscopic side larger, the apex ob- tuse, the base truncate, the borders sometimes whitish at the base, the edges finely denticulate. Median leaves orbicular to somewhat ovate, asymmetric with the outer side larger, acuminate to aristate, the arista often nearly '/2 as long as the lamina, the auricles not prolonged, borders nar- row, whitish, the edges finely denticulate. In deep, shady forests, in rain forests, in moist ravines, rarely epiphytic, 100-600 m, Amazonas and Loreto. Colombia, Ecuador, and Peru. This and Selaginella chrysoleuca have excep- tionally broad leafy branches and asymmetrical median leaves that are long aristate. Both species occur in dense forest in the Andes, but S1. speciosa occurs mainly in the north of Peru while S. chry- soleuca extends south to Cuzco. Amazonas: Huampami, Kayap 1311 (uc). Loreto: Be- tween Yurimaguas and Balsapuerto, Killip & Smith 28340 (F, us). Balsapuerto, Klug 2921 (F, GH, MO, us). Prov. Maynas, Iquitos, McDaniel & Rimachi 17473 (GH). Si- erra del Pongo, Mexia 6283 (F, GH, MO, uc, us). Gamita- nacocha, Rio Mazan, Schunke 109 (F, GH, uc, us). 1 9. Selaginella lechleri Hieron., in Engler & Prantl, Nat. Pflanzenfam. 1 (4): 683. 1901. LEC- TOTYPE (chosen by Alston et al., 1981): Peru, Puno, near San Gavan (San Gaban), Lechler 2159 (holotype, B?; isotype, BM). Main stem erect, 1 2-40 cm long, not articulate, stramineous, glabrous, the basal part with ap- pressed, ascending leaves. Primary branches 3-pinnate, forming an extended, flabelliform ar- rangement, distant, the lower stalked, those above sessile or nearly so, penultimate leafy branches straight or nearly so, ultimate leafy branches slen- der, 3-5 mm broad. Rhizophores confined to the base of the main stem. Lateral leaves of the main stem, above the first branch, ascending, coarse, mostly strongly imbricate, 2-5 mm long, oblong, acute, the auricles not prolonged, without con- spicuous whitish borders, the edges denticulate to short ciliate, the cilia denser at the base. Median leaves elongate, ovate, acuminate, the base some- times unequal, with a slightly enlarged auricle, not or sparsely ciliolate, without conspicuous whitish borders, the edges slightly denticulate or entire. Terrestrial on clay or sandy soil, along trails or on stream banks in primary forest or rain forests, 1 30-1 200 m, Amazonas and Loreto south to Puno. Colombia and Peru. This taxon seems to intergrade with Selaginella anceps but is maintained here in its traditional sense. The broad, erect stems with appressed leaves, the primary branches forming a broad, flabelli- form arrangement, and small size of the leaves on the ultimate branches are similar to those of S. anceps. Amazonas: E of Cinkan, Berlin 254 (F, MO, uc). Loreto: Prov. Maynas, Puca Urquillo, Plowman et al. 6628 (F, us). La Victoria, LI. Williams 2919 (F, us). Pasco: Prov. Oxapampa, between Puerto Bermudez and San Matias, Leon et al. 333 (F, USM). Puerto Bermudez (as Junin), Killip & Smith 26467 (F, GH). Cuzco: between Mistiana and Keros, Scolnik 868 (uc). Madre de Dios: Alto Madre de Dios, Rauh & Hirsch PI 570 (GH). Puno: Prov. Caraba- ya, Qllachea a Quillabamba, Vargas 17539 (GH). 20. Selaginella anceps (Presl) Presl, Abh. Bonn. Ges. Wiss. 5, 3: 581. (Bot. Bemerk., 151). 1844. TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 79 Lycopodium anceps Presl, Reliq. haenk. 1: 80. 1825. TYPE: "Luzon," but probably from Peru, Haenke (holotype, PR, photo, GH). Lycopodium gracile Poiret, in Lam., Encycl. suppl. 3: 551. 1814. TYPE: Peru (Holotype, Herb. Des- vaux P, photo OH). Selaginella gracilis (Poiret) Hieron., Hedwigia 58: 293. 1917, not Moore, 1886. Main stem erect, 20-25 cm or longer, not artic- ulate, 2-3.5 mm broad at the base, stramineous, glabrous, the basal part with appressed, ascending leaves. Primary branches 3-4-pinnate, forming an elongate, flabelliform arrangement, distant, stalked, the penultimate branches straight or nearly so, the ultimate leafy branches slender, 1-2 mm broad. Rhizophores confined to the base of the main stem. Lateral leaves above the first branch, appressed, 2-3 mm long, elongate-ovate, broadest at the base, acute, with a somewhat prolonged, ciliate auricle, without conspicuous whitish borders, long ciliate at the base of the leaf. Median leaves ovate-elon- gate, acuminate with a somewhat prolonged au- ricle or an enlarged, ciliolate basal lobe, the bor- ders not conspicuously whitish, the edges usually prominently ciliolate. Disturbed forests, primary and secondary rain forests, steep hillsides in wet forests, 1 50-1 500 m, San Martin and Loreto to Cuzco and Madre de Dios. Costa Rica south to Venezuela and to Bolivia. On the basis of similarities in the small, imbri- cate leaves densely covering the ultimate branches Selaginella anceps appears to be allied to 5. lech- leri. However, the usually denticulate base of the median leaves in S. lechleri is distinct from the usually prominently ciliolate one in S. anceps. San Martin: Between Tocache Nuevo and Juanjui, Croat 58038 (F, MO). Loreto: Pongo Manseriche, Mexia 6371 (GH, MO, uc, us). Huanuco: Pampayacu, Kanehira 118 (GH, us). Pasco: Prov. Oxapampa, Palcazu, Foster & d'Achille 10177 (USM). Junin: Rio Perene, Killip & Smith 25104 (F, GH, us). Perene, Seibert 2193 (F, MO, us). Ayacucho: Estrella, between Huanta and Rio Apu- rimac, Killip & Smith 22671 (F, GH, us). Cuzco: Prov. Convencion, Dudley 11381 (GH, MO). Madre de Dios: Prov. Manu, Atalaya, Foster & Wachter 7456 (MO). 2 1 . Selaginella erythropus (Mart.) Spring, in Mar- tius, Fl. bras. 1 (2): 125. 1840. Lycopodium erythropus Martius, Icon. pi. crypt. 39. 1 834. TYPE: Brazil, Piaui, Martius (holotype, BR). Plants 10-25 cm, mostly 10-15 cm long. Main stem erect and simple below the lowest, well-de- veloped branch, not articulate, reddish, glabrous, the basal part with appressed, ascending leaves. Primary branches often irregular, mostly 2-pinnate, ultimate branches not exceeding 5 mm broad, in- cluding the leaves. Rhizophores usually confined to the base of the main stem, reddish. Lateral leaves strongly imbricate, reddish, especially the borders, the color extending into the marginal cil- ia, elongate-oblong, 0.25-0.5 mm long, the acro- scopic side somewhat larger, acuminate (not aris- tate), with a rounded, denticulate basal lobe, the upper edges ciliate or ciliolate. Median leaves elongate-ovate, strongly imbricate, somewhat asymmetrical, acute to aristate, the base more or less truncate, the borders whitish, the edges den- ticulate, somewhat denser at the base. On moist riverbanks, steep, wet road banks, thickets in dense forests, sometimes forming dark green mounds on the floor of lowland rain forests, 250-1800 m, Amazonas south to Cuzco. Costa Rica; Colombia to Bolivia; Brazil. Among the species with erect main stems, Se- laginella erythropus is distinguished by the reddish color of the main stem that often extends into the branches and rhizophores, and by the small size of the plants. It also differs from S. haematodes by the ciliate or ciliolate lateral leaves. Amazonas: Prov. Bagua, between Aramango and Montenegro, Lopez et al. 4212 (GH, MO). San Martin: Juan Jui, Klug 4267 (GH, MO, us). Tarapoto, Spruce 3989 (GH, us). Junin: Colonia Perene, Killip & Smith 24918 (F, GH, us). La Merced, Macbride 5549 (F, us). Ayacucho: Estrella, between Huanta and Rio Apurimac, Killip & Smith 22664 (F, us). Cuzco: Prov. La Convencion, Dud- ley 11438 (GH). 22. Selaginella haematodes (Kunze) Spring, in Martius, Fl. bras. 1 (2): 126. 1840. Figure 9c-d. Lycopodium haematodes Kunze, Linnaea 9: 9. 1834. TYPE: Peru, San Martin, Mission Tocache, Poep- pig, in June 1830 (isotype, K). Selaginella filicina Spring, Nouv. Mem. Acad. roy. Sci. Belg. 24: 189, 1849. SYNTYPE: Peru, Mat- thews (Mathews) 40 36 (K). Other collections from Venezuela are also cited. Plants 30-65 cm long. Main stem erect, not articulate, bright red below the branches, glabrous, the basal part with appressed, ascending leaves. Primary branches usually red, ultimate branches 3-8 mm broad including the leaves, the basal part 80 FIELDIANA: BOTANY with appressed, ascending leaves. Rhizophores near the base of the main stem, deep red. Lateral leaves often reddish, oblong to elongate-ovate, broader at the base, 0.25-2.0 mm long, acute, the base truncate, without conspicuous whitish borders, the upper (acroscopic) edge finely denticulate, the teeth longer toward the apex, or nearly entire. Median leaves slender, elongate-ovate, acuminate to aris- tate, the arista '/4 or less the lamina length, the base truncate, with narrow, whitish borders, the edges denticulate especially near the apex. Frequent along riverbanks, on steep, moist hill- sides, or scattered on floor of dense forests, pri- mary forests, rocky hillsides, or along trails, 135- 900 m, San Martin and Loreto, south to Madre de Dios. Panama, and Venezuela south to Bolivia. The large, deep green, branches borne on long, simple, red-colored stems, readily distinguish the species. The vivid red color extends into the base of the main branches and also to the rhizophores that are confined to the basal part of the main stem. San Martin: Tocache Nueve, Schunke 7077 (F, us). Prov. Mariscal Caceres, Uchiza, Schunke 3202 (F, GH, us). Tarapoto, Spruce 4036 (GH, us). Loreto: San An- tonio, Rio Samira, Ayala & Arevalo 4227 (F, MO). Puerto Arturo, Killip & Smith 27747 (F, GH, us). Cueva de las Pavas, Fukushima 6521 (F, GH, MO). Huanaco: Prov. Huanuco, Mexia 8302 (GH, MO, uc). Pasco: Pichis Trail, Killip & Smith 26214 (F). Junin: La Merced, Soukup 2369 (F, GH). Madre de Dios: Cocha Cashu, lake of Rio Manu, Foster et al. 3417 (F). 23. Selaginella quadrifaria Alston et al., Bull. Brit. Mus. (Nat. Hist.) Bot. 9: 261. 1981. TYPE: Peru, Prov. Loreto, Sierra del Pongo, Mexia 6282 (holotype, BM; isotypes, GH, uc, us). Main stem erect, 20-70 cm long, not articulate, stramineous, glabrous, the basal part with ap- pressed, ascending leaves. Primary branches 3-4- pinnate, distant, ascending, usually dichotomously divided, forming a broad, flabellate arrangement of ultimate branches, 8-20 mm broad including the leaves, the axes straight. Rhizophores confined to the base of the main stem. Lateral leaves coarse, rigid, imbricate, 4-8 mm long, oblong or broader near the base, equilateral, with a central costa, the apex acute, the base truncate, with an enlarged, rounded, ciliolate lobe, without conspicuous whit- ish borders, the edges denticulate. Median leaves ovate, equilateral, with a central costa, acute, the base truncate, the edges sometimes narrow, whit- ish, finely denticulate. In dense forest or rain forest, on slopes, on well- drained soils, 450-860 m, Amazonas to Pasco. Colombia and Peru. Among related species with an erect stem and appressed leaves Selaginella quadrifaria is gener- ally distinguished by the median leaves that are equilateral and not or hardly aristate. Differences from S. praestans are noted under that species. Amazonas: Prov. Bagua, near Montenegro, Hutchison & Wright 382 IB (uc). Prov. Bagua, near Campamento, Wurdack 1869 (GH, us). Huanuco: Cerro del Sira, Rio Llullapichis, Wolfl2263B (F, us). Pasco: (as Junin) Puer- to Yessup, Killip & Smith 26372 (F, us). Prov. Oxapam- pa, valle del Palcazu, Iscozacin, Foster 4556 (F). Cabeza de Mono, near Iscozacin, D. Smith 3752 (uc). 24. Selaginella praestans Alston et al., Bull. Brit. Mus. (Nat. Hist.) Bot. 9: 260. 1981. Selaginella sprucei A. Braun, Ann. Sci. Nat. Bot. 5, 3: 277. 1865, not Hooker 1861. TYPE: Peru, San Martin, near Tarapoto, Guayrapurina, Spruce 4788 (not 4708 as originally cited) (holotype, G; isotypes, BM, NY). Main stem erect, 20-70 cm long, not articulate, green to brownish, glabrous, the basal part with appressed, ascending leaves. Primary branches 1- 2-pinnate, distant, ascending, sparingly pinnate, the basal ones often longer, the axes straight, ul- timate branches 10-15 mm broad including the leaves. Rhizophores confined to the base of the main stem. Lateral leaves above the first branches, ascending, imbricate, those below coarse, elon- gate-ovate to oblong, broadest at the base, 7-8 mm long, 3 or 4 x longer than broad, inequilateral, the acroscopic side larger, the costa acentric, acute to obtuse, the base truncate, or with a slightly en- larged, ciliolate auricle, without conspicuous whit- ish borders, the edges delicately ciliolate, some- times denticulate, Median leaves suborbicular to somewhat ovate, asymmetric, inequilateral, the outer side larger, acute, the base truncate, without conspicuous whitish borders, the edges ciliolate or entire. In dense shade of deep woods or rain forests, and along stream sides, 630-1500 m, San Martin, Huanuco, and Pasco. Colombia to Peru. The species is similar to Selaginella quadrifaria in the robust size of the plants and coarse leaves TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 81 appressed to the main stem. It can be distinguished from that species by the inequilateral shape and acentric costa of the lateral and median leaves in contrast to the equilateral leaves and centric costa in 5. quadrifaria. San Martin: Valley of Huallaga, between Tocache Nuevo and Juanjui, Croat 58057 (MO). Huanuco: Prov. Huanuco, Cotirarda, Mexia 8196 (BM, GH, MO, uc, us). Pasco: Prov. Oxapampa, between Iscozacin and Villa America, D. Smith 2840 (F). 25. Selaginella haenkeana Spring. Bull. Acad. roy. Sci. Bruxelles 10: 225. 1843. TYPE: probably Peru, "cordilleris chilensibus," Hdenke (ho- lotype, PR; photo, GH; isotypes, B, us). Selaginella dimorpha Klotzsch, Linnaea 18: 523. 1844. TYPE: "Auf den Anden von Chile," Hdenke (ho- lotype, B; isotypes, PR, us). Probably the same Peru collection cited above. Main stem erect, up to 40 cm or more long, usually unbranched Vs or more above the base of the main stem, the base stout, 1-3 mm broad, not articulate, greenish to stramineous, glabrous, the basal part with appressed, ascending leaves. Pri- mary branches 1 -pinnate, sessile or subsessile, dis- tant, spreading, often arching upward, the central ones usually longest, slender, the axes straight, ul- timate branches not more than 5 mm broad in- cluding the leaves. Rhizophores basal, or below the lowest branch. Lateral leaves 2-3 mm long, oblong, acute, without prolonged auricules, with long, discrete cilia especially dense at the leaf base. Median leaves ovate to somewhat elongate, aris- tate, the arista slender, '/4 or less the lamina length, the base not auriculate, the borders narrow, whit- ish or not, the edges long-ciliate, especially long and dense at the base. Terrestrial, among rocks, or climbing, in cloud forest and dense forest, 1 700-2000 m, Amazonas, and Pasco. Colombia, south to Bolivia. Well-developed plants often have an unusual branching system with a long, slender central stem and widely spaced, 1 -pinnate lateral branches bearing short, ultimate divisions. Amazonas: Prov. Bagua, La Peca, Barbour 2820 (F, MO, uc), 2967 (F, MO). Pasco: Pichis Trail, Dos de Mayo (as Junin), Killip & Smith 25843 (F, GH). 26. Selaginella kunzeana A. Braun, Ann. Sci. Nat. Bot. 5, 3: 296. 1865. LECTOTYPE (chosen by Alston et al., 198 1): Peru, Huanuco, Pam- payacu, Poeppig 195 (holotype, B; isotype, BM, P). Main stem prostrate long-creeping to assurgent, articulate, stramineous, glabrous, the basal part with spreading leaves and often smaller, ap- pressed, ascending leaves. Primary branches 2-3- pinnate, ascending, mostly widely distant, these as well as the apex of the main stem forming a rhom- boid arrangement, with an acute to acuminate apex, ultimate branches 5-8 mm broad including the leaves. Rhizophores extending above the lowest branches, mostly in the lower half of the stem. Lateral leaves above the first branch distant, wide- spreading, imbricate at the stem apex, 2-3 mm long, oblong, elongate with somewhat parallel sides, acute, the base with a prolonged auricle forming a round or slender lobe, with narrow, whitish, fine- ly denticulate edges, the teeth especially dense at the base. Median leaves slender, elliptic, the apex long-acuminate to aristate, the arista to '/4 the lam- ina length, the base prolonged with one, some- times two, large, auricles, with narrow, whitish, finely denticulate edges. Trailing on steep banks, in dense woods, often in disturbed areas, 500-1950 m, Huanuco, south to Cuzco. Mexico to Panama; Venezuela, Andes of Co- lombia to Peru. A widely distributed species occurring from Mexico to Peru, with a broad range in Peru, often in disturbed places. The rhomboid arrangement of the terminal, leafy branches distinguishes this from other articulate, prostrate species. Huanuco: Prov. Huanuco (as San Martin), Tingo Ma- ria, Allard 20361 (us); Asplund 12140 (us). Hacienda Mercedes, Mexia 8197 (F, GH, MO, uc, USM). Pasco: south of Pozuzo, Teppner 79/269 (us). Ucayali: Divisoria, Ce- rro Azul, Tryon & Tryon 5272 (F, GH, us, USM). Aya- cucho: Prov. La Mar, southwest of Hacienda Luisiana, Dudley 11721 (GH). Estrella, between Huanta and Rio Apurimac, Killip & Smith 22663 (GH, us). Cuzco: Prov. Quispicanchis, Marcapata, Vargas 3178 (MO, us). Kos- nipata, Velarde 7987 (GH). 27. Selaginella silvestris Asplund, Ark. f. Bot. 20 A (7): 30. 1926. TYPE: Bolivia, Sur Yungas, El Chaco, Asplund 1140 (holotype, UPS; frag., BM; photo, GH). Main stem prostrate, long-creeping to decum- bent, articulate, stramineous, glabrous, the basal part with spreading leaves, and often smaller, ap- 82 FIELDIANA: BOTANY pressed, ascending leaves. Primary branches 2-3- pinnate, often dichotomous, forming a broadly ovate to obdeltate arrangement, the ultimate leafy branches short, with a round to truncate apex, 4- 8, usually 5 mm broad including the leaves. Rhi- zophores mostly at the base of the main stem, extending above the first branch. Lateral leaves distant, spreading above the first branch, 3-4 mm long, oblong, somewhat broader near the base, asymmetrical, the apex obtuse to acute, the auri- cles not prolonged, without conspicuous whitish borders, the edges denticulate, especially at the base. Median leaves elongate-ovate, usually acu- minate, or if aristate the arista less than 'A the lamina length, the base unequal with 1 prolonged, curved auricle that may be up to % the length of the lamina, the borders not conspicuously whitish, the edges prominently denticulate. In humid forests, tropical rain forests, wooded ravines or thickets, also among rocks, rarely on rock walls or epiphytic, 650-3000 m, Cajamarca, and Amazonas, south to Ayacucho and Cuzco. Mexico south to Panama; Venezuela and Co- lombia south to Bolivia. The long-creeping stems with abundant rhizo- phores and prolonged auricles on the median leaves suggest an alliance with Selaginella trisulcata. Cajamarca: Cueva San Andres, Velarde 6984 (GH). Prov. Cutervo, Guta de las Huacharos, San Andres, Lopez & Sagdstegui 5404 (GH). Amazonas: Prov. Chachapoyas, Quebrada Molino, Wurdack 636 (F, GH, uc, us); Prov. Bagua, 25 km E of La Peca, Barbour 2962 (F, MO). San Martin: Prov. Mariscal Caceresa, Rio Abiseo Nacional Park, Young & Leon 4957 (HUT, USM). Huanuco: Prov. Huanuco, Carpish, Asplund 12842 (us). Pasco: Oxapam- pa, Leon et al. 913 (F). Junin: Yucapata, Yaupi, Woyt- kowski 6627 (MO, us). Huancavelica: Prov. Tayacaja, Marcavalle, Tovar4754 (GH). Ayacucho: Ccarrapa, Killip & Smith 22359 (us). Cuzco: Machu Picchu, Peyton & Peyton 1320 (GH, MO); Nunez 7516 (MO, uc). 28. Selaginella trisulcata Asplund, Ark. f. Bot. 20A (7): 34. 1926. TYPE: Bolivia, La Paz, El Chaco, Asplund 1482 (holotype, UPS; photo, GH). Selaginella poeppigiana var. peruviana A. Braun, Ann. Sci. Nat. Bot. 5, 3: 295. 1865. TYPE: Peru, Puno, near Tabina, Lechler 2015 (holotype, B?). Main stem prostrate, elongate, creeping, only rarely assurgent at the apex, articulate, pale green or sometimes light brown, glabrous, the basal part with spreading leaves, and often smaller, ap- pressed, ascending leaves. Primary branches 3-4- pinnate, distant, extending to the base of the main stem, more or less equally divided, in a flabelliform arrangement, the terminal branches nearly the same length, 8-12 mm broad including the leaves. Rhi- zophores long, coarse, stramineous, throughout the stems. Lateral leaves above the first branch spread- ing, distant, imbricate only on the ultimate leafy branches, 6-8 mm long, oblong-lanceolate, the sides more or less parallel, symmetrical, acute, the auricles not prolonged, without whitish borders, the edges denticulate, sometimes sparingly. Median leaves elongate-ovate, long-acuminate to subaristate, the arista less than V* the lamina, the base with a pro- longed, Ungulate auricle, the borders narrow, whitish or not, denticulate along the edges. In moist ravines, in secondary forest, primary rain forests, in soil among rocks, 730-2700 m, Amazonas, south to Puno. Ecuador to Bolivia. The general aspect of the leaves and system of branches is similar to those of Selaginella poep- pigiana and S. articulata, but the lateral leaves in those species have prolonged auricles that are not found in 5". trisulcata. The stem also has two vas- cular bundles. Amazonas: Prov. Bongara, N of Pedro Ruiz, Smith & Vasquez 4896 (MO, uc, USM). Huanuco: Carpish Pass, Allard 20996 (us). Pasco: Prov. Oxapampa, Smith & Canne 5824 (MO). Prov. Oxapampa, Canyon de Huan- cabamba, Leon 670 (F, GH). Junin: Carpapata, above Huacapistana, Killip & Smith 24467 (us). Yaupe, Woyt- kowski 6331 (MO, us). Ayacucho: Between Huanta and Rio Apurimac, Killip & Smith 22751 (F, us). Cuzco: Ca. 5 km N of Aguas Calientes, Solomon 3162 (MO). Portero, 8 km W of Quillabamba, Tryon & Tryon 5394 (F, GH, us). 29. Selaginella poeppigiana (Spring) Splitg., Tijdschr. Naturl. Gesch. Physiol. 7: 443. 1840. Selaginella sulcata subsp. poeppigiana Spring, Flora (Jena), 21: 185. 18 38. TYPE: Ecuador, Pichincha, Jameson (holotype, K). Main stem prostrate, long-creeping, ca. 20-40 cm long, articulate, stramineous, sometimes light brown at the apex, glabrous, the basal part with spreading leaves, and often appressed, ascending leaves. Leafy branches 3-4-pinnate, dichotomous, in a subflabellate arrangement, extending to the base of the main stem, the ultimate branches sim- ilar in length or nearly so, 8-10 mm broad. Rhi- zophores at the base of the main stem, extending to the upper branches nearly to the apex. Lateral TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 83 leaves above the first branch distant, spreading to patent, 4-6 mm, usually 5 mm long, oblong, sym- metrical or nearly so, acute, the base narrowed with 1 or 2, prolonged, acute or Ungulate, ciliate auricles, without conspicuous whitish borders, the edges finely denticulate. Median leaves elongate- ovate, long-acuminate or subaristate, the arista less than 'A the lamina length, the base with a pro- longed, Ungulate, often ciliate auricle, without con- spicuous borders, the edges sparsely denticulate. Creeping, prostrate, forming dense mats on shaded floor of wet forest, trailing along banks and roadsides, or in rock crevices, 900-2800 m, Ama- zonas south to Puno. Ecuador and Peru. The long, creeping stems, with abundant rhi- zophores throughout and in a subflabelliform ar- rangement, suggest alliances with Selaginella tri- sulcata and S. articulata. However, the leafy branches are not as broad, and usually more open and dichotomous than in those species. Amazonas: Prov. Bagua, La Peca, Harbour 2493 (F, MO, uc), 281 9 (F, MO). La Peca, Serrania de Bagua, Gentry et al. 23065 (F, MO, uc). San Martin: Prov. Moyobamba, Ferreyra 18518 (USM). Tarapoto, Spruce 4626 (GH, us). Loreto: Upper Rio Macusari, below Ecuadorian border, McDaniel 10984 (GH, MO). Huanuco: Pampayacu, Ka- nehira 182 (GH, us). Pasco: Prov. Oxapampa, Rio San Alberto, Leon 645 (F, USM). Junin: Above San Ramon, Killip & Smith 24657 (F, GH); Chanchamayo Valley, Schunke 192 (F), 194 (F), 231 (F). Cuzco: Prov. Paucar- tambo, Suecia, Woytkowski 175 (USM). Puno: Churu- mayo, Soukup 434 (GH). 30. Selaginella articulata (Kunze) Spring, Flora (Jena), 21: 182. 1838. LycopodiumarticulatumKunze,Linnaea9: 10, 1834. TYPE: Peru, Tocache, Poeppig, in 1830 (not lo- cated). Main stem prostrate, long-creeping, articulate, stramineous or light brown near the apex, pubes- cent especially at the apex and branch axils, or the main stem glabrescent. Leafy branches usually 3-pinnate, the apex forming a broad, compact, fla- belliform arrangement, the ultimate branches 10- 20 mm broad. Rhizophores long, coarse, numer- ous, extending 3/4 the length of the stem, stramin- eous or golden-colored. Lateral leaves closely placed to somewhat imbricate at the apex, those on the leafy branches oblong, spreading, approx- imate, 8-10 mm long, oblong, somewhat broader near the base, nearly symmetrical, obtuse to acute, with a prolonged, round auricle, the edges narrow, whitish, denticulate to entire. Median leaves short, ovate, the apex attenuate, the base with 2 pro- longed, round auricles, the edges narrow, whitish, finely denticulate. Plants subrepent, on floor of rain forest, along streams and roadsides, 350-2000 m, Amazonas and Loreto south to Pasco and Ucayali. Panama; Suriname and French Guiana; Peru, Bolivia, Brazil, Paraguay, and Argentina. The large, flabellately aligned branches with im- bricate leaves on the terminal portion of the stems compose form-flattened sprays that distinguish this from other Peruvian species. The prolonged basal auricles on the outer side of the median leaves are also exceptional. Amazonas: Prov. Bagua, Montenegro, Hutchison & Wright 3749 (F, GH, MO, uc, us). Prov. Bagua, near Cam- pamento, Wurdack 1892 (F, GH, uc, USM). San Martin: Prov. Mariscal Caceres, Tocache Nuevo, Schunke 6916 (F, MO, uc). Tarapoto, Spruce 4627 (GH, us). Loreto: Prov. Maynas, Poeppig, in 1831 (MO). Pasco: Prov. Oxa- pampa, Smith & Salick 8366 (MO). Ucayali: Padre Abad, Schunke 3074 (F, GH). 3 1 . Selaginella parkeri (Hooker & Grev.) Spring, Bull. Acad. roy. Sci. Bruxelles 10: 146. 1843. Lycopodium parkeri Hooker & Grev., Bot. Misc. 2: 388. 1831. TYPE: Guyana, Demerara, Parker (holotype, K). Selaginella pedata Klotzsch, Linnaea 18: 521. 1844. TYPE: Guyana, Schomburgk 118 (holotype, B?; isotype, BM). Selaginella fragilis A. Braun, Ann. Sci. Nat. Bot. 5, 3: 305. 1865. TYPE: Brazil, Amazonas, Rio Vaupes, Spruce 2533 (holotype, B?; isotype, BM, CGE). Selaginella brachylepis Christ, Bull Herb. Boissier 2, 1: 74. 1901. TYPE: Peru, "entre Ucayali et Hua- llaga," Huber (holotype, P). Main stem erect, 20-25 cm long, articulate, stra- mineous, or rarely red, glabrous, the basal part with appressed, ascending leaves. Primary branch- es 3-4-pinnate, sessile, the axis fractiflex, the branches forming compact, frondlike, flabelliform sprays, ultimate branches 5-12 mm broad includ- ing the leaves. Rhizophores at or near the base of the main stem, well below the first branch. Lateral leaves above the first branch, coarse, spreading to somewhat ascending, usually approximate, 4-6 mm long, oblong, broader just above the base, acute, the base usually with a prolonged, dentic- ulate auricle, sometimes 2, 1 an acute lobe, with narrow whitish borders or not, the edges dentic- 84 F1ELDIANA: BOTANY ulate, the teeth somewhat longer and denser at the base. Median leaves lanceolate, the apex long-acu- minate, the base with a prolonged, peltate, Un- gulate auricle, or sometimes 2, without conspic- uous borders, the edges finely denticulate. Primary forests, sometimes in wet or swampy forests, on sand or clay soils, usually in deep shade, 140-650 m, Amazonas and Loreto, south to Pas- co. Colombia and Peru; the Guianas, Amazonian Venezuela; Brazil. The plants have stems with one vascular bundle, and there are short auricles on the axillary leaves similar to those on the median leaves. The long main stems bearing flagelliform branches that ap- pear somewhat frondlike, and the fractiflex axes of the primary branches clearly suggest alliances between this and S. geniculata. Some specimens with unusually attenuated, flagelliform branches have been recognized as a distinct species, Selag- inella fragilis. Amazonas: Prov. Bagua, Montenegro, Hutchison & Wright 3821 A (uc). San Martin: Alto Rio Huallaga, Juan Jui, King 3831 (F, GH, both mixed with S. anceps). Lo- reto: Prov. Requena, van der Werffet al. 9966 (MO, uc), 10108 (MO, uc). H nan urn: Prov. Pachitea, Bosque Na- tional de Iparia, Schunke 2258 (F, GH, us). Pasco: Prov. Oxapampa, Quebrada Castilla, Leon & Young 1036 (F, GH). Prov. Oxapampa, Iscozacin, D. Smith 3735 (MO, uc). 32. Selaginella geniculata (Presl) Spring, Bull. Acad. roy. Sci. Bruxelles 10: 230. 1843. Lycopodium geniculatum Presl, Reliq. haenk. 1 : 80. 1 825. TYPE: "Luzon," but doubtless Peru or Ec- uador, Haenke (holotype, PR; photo, BM, GH). Selaginella ferruminata Spring, Bull. Acad. roy. Sci. Bruxelles 10: 231. 1843, TYPE: Peru, Pangoa, Matthews (Mathews) 1083 (holotype, K.; isotypes, GH, us). Selaginella elongata Klotzsch, Linnaea 18: 522. 1844. TYPE: Peru, Cuchero, Ruiz 94 (holotype, B). Selaginella nodosa Presl, Abh. Bohm Ges. Wiss. 5 (2): 580 (Bot. Bermerk. 50). 1844. TYPE: Peru, Cas- sapi, Poeppig (not located). Selaginella tomentosa Spring, Nouv. Mem. Acad. roy. Sci. Belg. 24: 231. 1849. TYPE: Colombia, Cau- ca, Gorgona Isl., Hinds (holotype, K.). Main stem erect, 10-50 cm or longer, articulate, stramineous, glabrous, or sometimes pubescent, the basal part with appressed, ascending leaves. Primary branches 3-4-pinnate, sessile above the lowest branches, forming flat, imbricate sprays in an extended flabelliform, frondlike arrangement, axis of the primary branches fractiflex, the ulti- mate leafy branches 2-5 mm broad. Rhizophores borne at the base of the main stem, or below the first branch. Lateral leaves of the main stem, above the first branches, coarse, ascending or spreading, usually approximate, those below distinct, peltate, 2-5 mm long, elongate-ovate, broadest above the base, acute, the base with a short, acute lobe, with- out conspicuous whitish borders, the edges den- ticulate, especially at the base. Median leaves im- bricate, oblong, acute to acuminate with a long, Ungulate auricle, without conspicuous whitish bor- ders, the edges entire or nearly so. On white sand or clay soil in high forest or rain forest, usually terrestrial but sometimes on tree trunks, or on rocks, 120-1500 m, Amazonas and Loreto south to Cuzco and Madre de Dios. Colombia, Ecuador and Peru. The axillary leaves are truncate; there are two vascular bundles at the base of the main stem, and there are peltate leaves on the main stem, below the first branch. These characters distinguish the species from other erect, articulate species of Peru. An alliance with Selaginella parkeri is indicated by the long main stem, the alignment of branches, and the fractiflex axes of the primary branches; however, the leafy branches are not as broad as in S. parkeri. Amazonas: Rio Cenepa, vicinity of Huampami, Berlin 633 (F, MO, uc); Ancuash 1042 (MO, uc). Prov. Bagua, above Cascadas de Mayasi, Wurdack 1894 (F, GH, uc, us, USM). San Martin: Prov. Mariscal Caceres, Cam- panilla, Schunke 4118 (F, us). Loreto: Between Yuri- maguas and Balsapuerto, Killip & Smith 28250 (F, GH, us). Prov. Maynas, between Rio Momon and Rio Mo- monicillo, McDaniel 16121 (F, GH, MO). Huanuco: Prov. Huanuco, Vargas 5288 (MO). Pasco: Prov. Oxapampa, between Puerto Bermudez and Hito San Matias, Leon et al. 327 (F, USF, USM). Junin: Satipo, Ridoutt 11397 (USM). Ucayali: Boqueron del Padre Abad, Skog et al. 5128 (us). Cuzco: Marcapata, Quispicanchis, Vargas 3781 (MO, us). Madre de Dios: Prov. Manu, Vargas 16643 (GH). 33. Selaginella asperula Spring, in Martius, Fl. bras. 1 (2): 127. 1840. SYNTYPES: "ad flu- vium Amazonum," Martius (M), "et ad flu- men S. Francisco," Martius (M). Main stem slender, erect, sometimes arching, and rooting at the apex, usually 1 -pinnate, 1 5-30 cm long, articulate, stramineous, glabrous, the basal part with appressed, ascending leaves. Primary TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 85 branches distant, often forming tufts of slender, dichotomous branches along the main stem, sometimes rebranching into uniformly slender ul- timate leafy branches 1-3 mm broad, the axes straight or nearly so. Rhizophores mostly at or near the base of the main stem. Lateral leaves above the first branch ascending or wide-spreading, 0.5- 3 mm long, elongate-ovate, somewhat broader just above the base, acute, with an auricle sometimes forming a small, acute tooth, the edges narrow, whitish, ciliolate, these longer at the base. Median leaves ovate, asymmetrical, acute, with a pro- longed, ciliate auricle, the edges narrow, whitish, denticulate, the teeth longer and denser at the base. Steep slopes and road banks, open areas, thick- ets and moist areas in dense forests, 680-1 300 m, Amazonas and Loreto to Junin. Amazonian Venezuela and Brazil, south to Bo- livia. The diffuse branches that form a uniformly slen- der, dichotomous, leafy branch system, with small, strongly imbricate leaves, are distinctive among the Peruvian species. The species has a single stele at the base of the erect stem, a character that sep- arates it from Selaginella geniculata. Amazonas: Prov. Bagua, 43 km NE of Chiriaco, Bar- hour 4458 (MO, USM). San Martin: San Rogue, LI. Wil- liams 7415 (F, us). Prov. Rioja, Woytkowski 6036 (GH, MO, us). Loreto: Rio Huallaga. above Lagunas, Croat 18097 (F, GH, MO). Huanuco: Tingo Maria (as San Mar- tin) Allard 20823 (MO). Pasco: Prov. Oxapampa, Parque Nacional Yanachaga-Chemillen, Leon et al. 989 (F). Ju- nin: San Ramon, Killip & Smith 24785 (F, GH, us). Prov. Chanchamayo, La Merced, Macbride 5504 (F, us); La Merced, D. Smith 4063 (MO, uc). 34. Selaginella stellata Spring, in Martius, Fl. bras. 1 (2): 129. 1840. LECTOTYPE (designated here): Brazil, Prov. Para. Obidos, Martius (holotype, M). Selaginella conduplicata Spring, in Martius, Fl. bras. 1 (2): 129. 1840. TYPE: Brasil, Prov. Para, Mar- tius (holotype, M). Selaginella calcarata A. Braun, Ann. Sci. Nat. 5, 3: 305. 1865. TYPE: ?, several syntypes are cited. Main stem erect to sometimes creeping, 10-30 cm long, articulate, stramineous, glabrous, the basal part with appressed, ascending leaves. Primary branches 3-4-pinnate, usually with an extended, flabelliform, frondlike arrangement, branches dis- tant, the lowest often longest, or in creeping stems the branches of similar length, axes of the branches fractiflex, ultimate leafy branches slender, 2-5 mm broad. Rhizophores abundant on the lower part of the stem nearly to the first branch, or in creeping stems among the upper branches. Lateral leaves above the first branches distant, ascending, those below distant, peltate, 1—4 mm long, lanceolate, slightly broader above the base, acute, the base with a prolonged, membranaceous auricle on the acroscopic side and usually a sharp tooth on the basiscopic side, with narrow, whitish, denticulate edges. Median leaves very small, usually '/2 the length of the lateral leaves, ovate-lanceolate, acu- minate or subaristate, with a prolonged, Ungulate, ciliate basal auricle, with narrow, whitish, dentic- ulate edges. In dense stands on clay or often sandy soil in lowland woods or rain forests and secondary for- ests. Common in humid, disturbed areas, in chac- ras, especially old ones, ca. 100-250 m, Loreto and Junin. Guianas, Amazonian Brazil west to Peru. We follow Alston (Repert. Spec. Nov. Regni Veg. 40: 309. 1936) in accepting the name Selag- inella stellata over S. conduplicata, both published by Spring in the same work. The species is readily distinguished by the un- usually prolonged, ciliate auricles on axillary, me- dian, and lateral leaves. The bright stramineous color of the fractiflex axes contrasting with the deep green leaves resembles what is seen in Se- laginella geniculata, and there are peltate leaves on the lower part of the main stem and branches similar to that species. Loreto: Iquitos, Croat 18306 (MO, us), 20057 (GH, MO, uc). Iquitos, road to San Juan, Mexia 6496 (F, GH, MO,UC). Prov. Maynas, Picuruyacu, Revilla 123 (F, MO, uc). Ga- mitanacocha, Schunke 144 (F, GH, uc, us). Iquitos, Tryon & Tryon 5166 (F, GH, us). Junin: Chanchamayo Valley, La Merced, Soukup 1117 (F), 7775 (F). La Merced, Mac- bride 5503 (F). 35. Selaginella exaltata (Kunze) Spring, Bull. Acad. roy. Sci. Bruxelles 10: 234. 1843. Fig- ure 9e-f. Lycopodium exaltatum Kunze, Linnaea 9: 8. 1834. TYPE: Peru, San Martin: "ab Uchiza ad Toache," Poeppig, July 1830, Diar. 1953 (not located). Selaginella strobilifera Christ, Bull. Herb. Bossier 2, 1: 72. 1901. TYPE: Peru, "entre Rios Huallaga et Ucayali," Huber 1515 (holotype, P; isotype, MG). Main stem erect, scandent up to 8 m long, ar- ticulate, 2-6 mm broad including the leaves, stra- 86 FIELDIANA: BOTANY mineous, often pubescent. Primary branches widely spreading, 2-3-pinnate, often pubescent. Rhizo- phores few, borne in axils of branches. Lateral leaves distant, clasping on the main stem when dry, approximate, or imbricate on the ultimate branches, obtuse, elongate, 1-3 mm long, 3 or 4 x longer than broad, acute, or sometimes entire, somewhat acuminate, the base truncate, some- times with conspicuous whitish borders. Median leaves acentric, the outer side larger, ovate, acu- minate to aristate, the arista '/4 the lamina length, glabrous, the base entire. In lowland forests, rain forests, secondary for- ests, and disturbed areas, scandent or scrambling over shrubs, often forming dense thickets, 30-760 m, Amazonas south to Madre de Dios. Costa Rica south to Bolivia and western Brazil. The scandent habit of these plants, often form- ing dense thickets, is exceptional among species of Selaginella. The stems are noted to be up to 8 m long, but in most collections they are indicated as 3 m. The habit of these plants, with dense branches bearing closely placed leaves, forms a compact cover over other vegetation. Amazonas: Prov. Bagua, Montenegro, Sagdstegui et al. 7158 (F, GH, HUT, MO, uc). Prov. Bagua, above Cas- cadas de Mayasi, Wurdack 1800 (F, GH, uc, us). San Martin: between Tocache Nuevo and Juanjui, Croat 58035 (F, MO). Lamas, Naranjal, Knapp & Mallet 6958 (F, MO, uc). Loreto: Bosque Nacional von Humboldt, Gentry & Revilla 20418 (F, MO). Prov. Maynas, Plowman et al. 6502 (F, GH, us). Nanay, Woytkowski 5126 (GH, MO, us). Huanuco: Casa de Koepke, Rio Llullapichis, Dudley 12484 (GH). Bosque Nacional de Iparia, Schunke 1299 (F, GH, MO, us). Pasco: Prov. Oxapampa, Cabeza de Mono, D. Smith 3768 (MO, uc). Puerto Bermudez (as Junin), Killip & Smith 26498 (us). Madre de Dios: Prov. Tambopata, Tambopata Nature Reserve, Barbour 4881 (F, MO). Manu National Park, Gentry et al. 27209 (F, MO). 36. Selaginella convoluta (Arnott) Spring, in Mar- tius, Fl. bras. 1 (2): 131. 1840. Lycopodium convolutum Arnott, Mem. Wernerian Nat. Hist. Soc. 5: 199. 1824 TYPE: Brasil, Rio de Janeiro, Jameson (holotype, E). Main stem erect, very short, not articulate, green to brownish, glabrous. Primary branches forming an open rosette curling inward when dry, 1-2- pinnate, with the ultimate branches, including the leaves, 3-5 mm broad. Rhizophores few at the base of the main stem. Lateral leaves rigid, strongly imbricate, 1-2 mm long, ovate-lanceolate or somewhat elliptic, acuminate to acute, strongly imbricate, the base with a prolonged, densely cil- iate auricle, the acroscopic side with a broad, whit- ish border, the edges irregularly denticulate. Me- dian leaves broadly ovate to somewhat elliptical, obliquely oriented, acute or short aristate, the base not prolonged, the borders not conspicuously whitish, the edges regularly dentate. On dry desert slopes, 1250 m, Cajamarca. Guatemala; Greater Antilles; Guyana; Peru; Brazil and Paraguay. The habit of these plants forming a rosette of stems, with lateral branches curling inward, readi- ly characterizes the species. The leaves are di- morphic; both lateral and median leaves are strongly imbricate and have whitish margins and dentate edges. Selaginella convoluta is widely distributed in Central and South America, but only a single col- lection has been seen from Peru. The inrolled, brownish, leafy stems may be easily overlooked on the dry, desert slopes where the plants occur. Cajamarca: Prov. Celendin, Balsas-Celendin road, Rio Maranon valley, D. Smith 6179 (MO). Comments The following species are accepted by Alston et al. (1981) as growing in Peru. Adequate material of these has not been seen and a better evaluation of them is desired than is now possible. a. Selaginella acanthostachys Baker, J. Bot. 21: 99. 1 883. TYPE: Peru, San Martin, Spruce 4328 (holotype, K). b. Selaginella- asplundii Crabbe & Jermy, Fern Gaz. 11: 257. 1976. TYPE: Peru, Huanuco, Asplund 12822 (holotype, s; isotype, BM). c. Selaginella muscosa Spring, in Martius, Fl. bras. 1 (2): 120. 1840. Type: Brazil, Luschnatt (ho- lotype, M). d. Selaginella suavis (Spring) Spring, Bull. Acad. roy. Sci. Bruxelles 10: 229. 1843. Selaginella sulcata ssp. suavis Spring. Flora (Jena) 21: 185. 1838. TYPE: Brazil, Sellow (holotype, K; iso- type, B?). e. Selaginella wolfii Sodiro, Crypt, vase. Quit. 620. 1893. TYPE: Ecuador, "woods of the western region, 1800 m." Sodiro (holotype?, p). The following species was credited to Peru on the basis of the type, which is now considered to have been collected in Panama. TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 87 f. Selaginella horizontalis (Presl) Spring, Bull. Acad. roy. Sci. Bruxelles 10: 226. 1843. Lyco- podium horizontale Presl, Rel. haenk. 1: 78. 1825. TYPE: Panama, Haenke (holotype, PR), not from Peru as originally thought. Family 28. ISOETACEAE Isoetaceae Reichenb., Hot. Damen Kunst. Freunde Pflanzenw. 309. 1828. TYPE: Isoetes L. Stem erect, subterranean, not indurated, bearing numerous, fleshy to wiry, dichotomously branched roots. Leaves numerous, spirally arranged, ligu- late, acicular with a single vein and 4 rows of air chambers, each transversely partitioned by mul- ticellular septa. Sporangia foliar, large, single, ses- sile at or near the adaxial leaf base. Megaspores tetrahedral-globose and trilete, without chloro- phyll, germinating endosporically to form a small, short-lived megagametophyte. Microspores ellip- soidal and monolete, germinating endosporically to form a small megagametophyte producing 4 multiflagellate spermatozoa. An ancient genus with representatives known from the lower Cretaceous; represented today by a single, distinctive, and cosmopolitan genus ex- hibiting distant affinities with Selaginella and Ly- copodium. Species recognition is greatly hindered by tre- mendous phenotypic plasticity and active speci- ation, both through hybridization and polyploidy as well as through diploid divergence (Taylor & Hickey, 1992). Species identificatiop is also ham- pered by these phenomena but is made even more difficult by a suite of specialized characters not found elsewhere in the plant kingdom. Unique terminology, therefore, is an historic addendum to Isoetes systematics. Most of these novel terms were first introduced by Alexander Braun in 1 864. The use of them, although inconvenient at first, adds clarity and uniformity of understanding. Within the body of this treatment I employ them in the following contexts: corm— the "stem" of the plant, that part to which the leaves and roots are attached; fossa— the groove (or one of several) on the side of the corm (lateral fossa), or extending all the way around the corm (circumbasal fossa) and from which roots are derived; synchronous roots acropetal roots— in Isoetes, roots are initi- ated with a distinctive rhizotaxy (Paolillo, 1963) with root initiation simultaneous (synchronous) or not (acropetal); subula— the distal, non-laminate portion of the leaf; ala— the lateral, proximal lam- inate portion of the leaf, not including the ex- panded mid vein or sporangial regions; scales — leaf primordia that have been arrested early in ontogeny and have become sclerified and black- ened; these initially surround the corm apex during dormancy and, as new leaves are initiated, are displaced outward, and hence may be found sur- rounding the fully developed leaves on a corm; ligule— a multicellular, planar, and glandular tis- sue growing out of a small pit (foveola) in the proximal portions of the adaxial leaf surface, just distal to the sporangium; the ligule is composed of a thickened central region (the cushion) and peripheral, delicate margins; labium— a nonglan- dular, planar tissue that is apparently a modifi- cation of the proximal rim of the foveola; this tissue can be quite extensive and may be errone- ously identified as the ligule (it has also been called "pseudoligule"); velum— a tissue of leaf derivation that completely or partially covers the sporangi- um, the latter in most species somewhat impressed within the leaf tissue. References BRAUN, A. 1864. Les especes d'lle Sardaigne. Ann. Sci. Nat. 5: 306-377. FUCHS, H. P. 1982. Zur heutigen Kenntnis von Vorkommen und Verbreitung der siidameri- kanischen Isoetes-Arten. Proc. Kon. Ned. Akad. Wetensch. CSS: 205-260. HICKEY, R. J. 1984. Chromosome numbers of Neotropical Isoetes. Amer. Fern J. 74: 9-13. HICKEY, R. J. 1986. Isoetes megaspore surface morphology: Nomenclature, variation, and sys- tematic importance. Amer. Fern J. 76: 1-16. MARSDEN, C. R. 1976. Morphological variation and taxonomy of Isoetes muelleri A. Br., J. Ade- laide Bot. Gard. 1: 37-54. PAOLILLO, D. J. 1963. The developmental anat- omy of Isoetes. Illinois Biol. Monograph 31:1- 130. PFEIFFER, N. 1922. A monograph of the Isoe- taceae. Ann. Missouri Bot. Gard. 9: 79-232. TAYLOR, W. C., AND R. J. HICKEY. 1992. Hab- itat, evolution, and speciation in Isoetes. Ann. Missouri Bot. Gard. 79: 613-622. WEBER, U. 1922. Zur anatomic und systematik der gattung Isoetes L., Nova Hedwigia 63: 2 1 9- 262. 88 FIELDIANA: BOTANY I. Isoetes Isoetes L., Sp. pi. 2: 1 100. 1753. TYPE: /. lacus- tris L. Stylites Amstutz, Ann. Missouri Hot. Gard. 44: 121. 1957. TYPE: S. andicola Amstutz = /. andicola (Amstutz) Gomez. Figure 10. Aquatic, amphibious or terrestrial perennials. Stem globose, to vertically or horizontally elon- gate, buried in substrate. Leaves terete, trigonal (broadly 3-angled) or triquetrous (sharply 3-angled). Ligule delicate, to ca. 8 mm, glandular, auriculate. Labium variable in expression, minute to completely covering the ligule, generally thick and fleshy. Sporangium elliptic to ovate, embed- ded in the leaf or partially emergent, exposed or completely covered by the velum. Megaspores lae- vigate, reticulate, echinate, tuberculate or cristate, 250-1 ,200 Mm in diameter. Microspores laevigate, tuberculate, or echinate, 25-50 MHI long. 2n = 22- 132. The cosmopolitan genus Isoetes has been vari- ously divided into sections based on megaspore surface morphology. Recent studies, however, suggest that spore morphology is extremely vari- able and highly subject to convergence. Key to species of Isoetes a. Plants of lowland (less than 1 500 m) pools; subula triquetrous; scales present; megaspores baculate I.I. panamensis a. Plants of high altitude (more than 3000 m) pools, streams, or moist meadows; subula terete or trigonal; scales usually absent; megaspores laevigate, tuberculate, pustulate, cristate, echinate, rugulate, or reticulate, not baculate b b. Megaspores reticulate 2. I. novo-granadensis b. Megaspores not reticulate, surface ornamentation variable c c. Plants diminutive, the leaves of mature, fertile plants rarely longer than 30 mm in length, scale leaves around corm bases and surrounding shoot apex 3. I. parvula c. Plants larger, only juveniles with leaves less than 30 mm, fertile adult plants larger, lacking scale leaves d d. Velum covering, at most, '/4 of the sporangium, the sporangium essentially exposed . . . . e e. Corm vertically elongate, frequently producing unbranched roots only along a lateral groove (fossa); alae extending more than 50% of the total leaf length; sporangia elevated above the leaf base; megaspores laevigate, pustulate or rugulate— often variable within a single sporangium; mature plants deeply embedded in substrate, with only the distal (+/— ) quarter of the leaf tip exposed 4. I. andicola e. Corm globose to slightly elongate vertically and producing roots along a continuous circumbasal groove (fossa); alae extending less than 50% of the leaf length; sporangia basal; megaspores laevigate or with subtle, indistinct markings; mature plants not deeply embedded in substrate, typically with at least the distal half of the leaf exposed f f. Leaves broad, 3.5-6 mm wide at mid-length, 6-8 mm wide at the base, subula and alae dark brown to dark green, the apex subacute to acute; labium narrowly oblong, 40-60 Mm wide, 140-180 nm high; megaspores of 2 size and shape classes 5. I. dispora f. Leaves narrow, ca. 1.5-1.6 mm wide at mid-length, 3.6-4 mm wide at the base, subula and alae light green, the apex attenuate; labium depressed-ovate, 540 fim wide, 1 20 Mm high; megaspores similar in size and shape 6. I. hewitsonii d. Velum covering 3A or more of the sporangium g g. Plants terrestrial or amphibious; distal portions of subulae strongly trigonal; leaves often leathery or cartilaginous, the apex acute 7. I. saracochensis g. Plants aquatic or occasionally amphibious; distal portions of subulae terete to half-round, lax to erect, flaccid or stiffly turgid, the apex attenuate to subacute h h. Leaves stiffly erect, extremely turgid (to the point of being brittle), dark green, most often with dark brown to nearly sclerotic pigmentation basally; corm globose to extensively elongate horizontally; megaspores laevigate 8. I. lechleri TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 89 5cm 5mm 1mm 90 FIELDIANA: BOTANY h. Leaves flexuous to weakly erect, barely turgid, bright green and without dark pig- mentation basally; corm globose to slightly elongate horizontally; megaspores lae- vigate to minutely tuberculate or cristate 9. I. boliviensis 1. I soctes panamensis Maxon & Morton, Ann. Missouri Hot. Gard. 26: 272-273. 1939. TYPE: Panama, Prov. Panama, vicinity of Bejuco, Woodson et al. 1685 (holotype, us!; isotypes, GH!, NY!). Isoetes pacifica Svenson, Amer. Fern J. 34: 123. 1944. TYPE: Ecuador, Prov. Guayas, east of Chanduy, Svenson 1 1022 (holotype, BKL; isotypes, GH!, NY!). Isoetes savanarum Gomez, Phytologia 49: 339. 1891. TYPE: Costa Rica, Prov. Guanacaste, Gomez 17088, as 7055 (holotype, CR!). Corm globose, 7-34 mm in diameter, 2-3-lobed, dichotomous roots arising synchronously within the circumbasal fossa(e). Leaves to 100, erect, 160- 540(-760) mm long, 6-16 mm wide at the base, 0.5-1. 5(-2) mm wide at mid-length; alae colorless and hyaline proximally, stramineous to bright green or bluish green and chartaceous distally, 2-4 mm wide at the sporangium, 35-185(-230) mm long ([17-]22-30[-40]% of the total leaf length), apices attenuate; subula triquetrous (sharply 3-angled), stramineous to bright green to bluish green, the apex attenuate to subacute, with 3 groups of fi- brous bundles forming distinct longitudinal ridges, continuing to the apex; stomates present; scale leaves present. Sporangium obovate, tan, unspot- ted, 6-12 mm long, 3.5-7 mm wide, basal. Velum rudimentary, 0.2-0.5 mm wide along the lateral edges of the sporangium, absent along the upper edge of the sporangium. Ligule delicate, ephem- eral, occasionally represented by a whitish, deltate to triangular fragment, 3-4 mm long, 2.5-4.5 mm wide. Labium depressed-ovate, entire, laciniate or shallowly scalloped, 800-2,800 Mm high, 2,500- 4,500 Mm wide. Megaspores white, not shiny, 380- 580 (x = 496) Mm in diameter, baculate, the bacula occasionally laterally confluent; equatorial and proximal ridges narrowly triangular to deltoid in cross-section, equatorial ridges often undulate. Microspores ash-gray, 27—40 (x = 34) Mm long, 20-33 (x = 26) Mm wide, sparsely echinate, to papillate, to laevigate. 2n = 44. In ephemeral bodies of water such as rain pools, riverbanks, and marshes, from sea level to 1 500 m, Tumbes. Western Guatemala to Panama; coastal Ecua- dor and Peru; Brazil and Paraguay. Rarely col- lected. Throughout its range, this species exhibits con- siderable variation in the shape, size and distri- bution of the bacula on megaspores. Microspores also vary in surface morphology, but no parallel trends are evident. The single tetraploid count for this species is from Costa Rican material; it would not be surprising to find distinctive reproductively isolated elements within /. panamensis, with the Peruvian material presumably assigned to /. pa- cifica Svenson. Isoetes panamensis is the only Peruvian species with either triquetrous subulae or baculate mega- spores and as such is a distinctive element in the flora. Gomez's description of the ligule in /. sa- vannarum refers to the condition of the labium. Tumbes: South of Zarumilla, Ellenberg 1380 (GH, u). 2. Isoetes novo-granadensis Fuchs, Caldasia 8: 314-315. 1969. TYPE: Colombia, Comisaria del Putumayo, 3250 m, Cuatrecasas 11770 (us!). Isoetes dichotoma Mora-Osejo & Hagemann, Mutisia 43: 5. 1977. TYPE: Colombia, Dept. Narino, Volcan Galeras, 3900 m, Hagemann & Leist 1773 (COL, HEID!). Corm slightly elongate horizontally to globose, up to 15 mm tall and 30 mm in diameter, 2-lobed, roots arising synchronously within a continuous circumbasal fossa. Leaves to about 80, stiffly erect, up to 200 mm long, often with reflexed tips, 6-10 mm wide at the base, 2-7 mm wide at mid-length, dark green to brown proximally, bright to dark green distally; alae dark green to nearly black, membranaceous, 1.5—4 mm wide at the sporan- FIG. 10. Isoetes andicola: a, habit; b, proximal portion of leaf showing elevated sporangium; c, enlargement of "b" showing ligule (LI), labium (LA), sporangium (SP). Isoetes lechleri: d, habit; e, proximal portion of leaf showing ligule (LI), velum (VE), sporangium (SP). Isoetes dispora: f, proximal portion of leaf showing labium (LA), velum (VE), sporangium (SP). (a from Hutchison et al. 5890, F, b, c from Saunders 1154, MU; d from Sanchez Vega 1330, F, e from Leon 2074, MU; f from Skillman et al. 12850, MU.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 91 gium, 20-140 mm long ([44-]50-63[-70]% of the leaf length), parallel-sided, their apices attenuate; subula trigonal (broadly 3-angled in cross-section, erect to slightly reflexed, heavily cutinized, the apex sharply acute; fibrous bundles absent; stomates absent; scale leaves absent. Sporangium elliptic to ovate, diaphanous to light brown, concolorous, 4- 9 mm long, 3.5-5 mm wide, basal. Velum rudi- mentary, represented by a narrow ridge of tissue along the lateral portions of the sporangium, ab- sent along the upper edge of the sporangium. Lig- ule sagittate, brown, 7-9.5 mm high, 3.5-5 mm wide, delicate; the margins especially ephemeral, the cushion occasionally persistent as a widely ovate remnant, ca. 2 mm high x 2.5 mm wide. Labium obsolete. Megaspores white to light gray, not lustrous, 880-1,120 (;c = 1,015) »m in di- ameter, reticulate; equatorial and proximal ridges straight, distinct, as high or slightly higher than broad, generally obscured by the dense muri. Mi- crospores dark brown, 37-52 (x = 47) /urn long, 30-43 (x = 37) nm wide, smooth. 2n = 132. Wet boggy soil, 3500-3600 m, Cuzco. Colombia, Ecuador and Peru. Known from Peru only by this single collection. The species is common in moist paramos through- out Ecuador and Colombia where it grows either as a submerged aquatic or, more commonly, as a terrestrial species deeply embedded in surround- ing vegetation. The Peruvian material is typically terrestrial, with only the upper half to upper third of the leaves protruding above the substrate. Cuzco: Prov. Paucartambo, camino peatonal de El Mirador a Cerro Macho Cruz, 3500-3600 m, Leon 2246 (F). 3. Isoetes parvula Hickey, sp. nov. Ab aliis speciebus altitudis celsis statute parvulo, squamis scleroticis praesentibus, et megasporis subfari- nosis differt. Corm globose, 3-5 mm in diameter, bilobed; dichotomous roots arising synchronously within the circumbasal fossa. Leaves 8-15, stiffly reflexed, 20-30 mm long, 3-4 mm wide at the base, 0.3- 0.5 mm wide at mid-length; alae hyaline, 0.5-1 mm wide at the sporangium, 0.5-1 mm long (25- 50% of the total leaf length), each apex attenuate; subula half-terete (?), bright green, not highly cu- tinized, straight to slightly reflexed, the apex at- tenuate; fibrous bundles not evident; stomates not seen; scales present. Sporangia circular to elliptic, embedded, basal, hyaline, unspotted, 1-3 mm long, 1-2 mm wide. Velum incomplete, extending (10-) 50-75% down the sporangium. Ligule quickly ephemeral, not seen. Labium not evident. Mega- spores reddish brown en masse, 30-36.7 (x = 33.2) Mm long, 25-28.4 (x = 26.4) nm wide, shortly echinate. TYPE— Peru, Dept. Ayacucho, Prov. Ayacucho, Laguna Yaurihuiri, about 205 km from Nazca on the road to Abancay, rocky and stony slopes, dom- inant in site with seeping water, 4300 m, Brand- byge 321 (holotype, AAU!). High-altitude seeps, 4300 m, Ayacucho. Known only from the type specimen. 4. Isoetes andicola (Amstutz) Gomez, Brenesia 1 8: 4. 1980. Figure lOa-c. Stylites andicola Amstutz, Ann. Missouri Bot. Gard. 44: 121. 1957. TYPE: Peru, Dept. Lima, Prov. Huarochiri, above Casapalca, in alpine bogs, 4750 m, Amstutz 2000 (holotype, MO). Stylites gemmifera Rauh, in Rauh & Falk, Sitzungs- ber. Heidelberg Acad. Wiss. Math. Naturwiss. Kl. 1959: 11. 1959. TYPE: Peru, Dept. Lima, Prov. Huarochiri, above Casapalca, von Appen (holo- type, HEID). Isoetes andicola var. gemmifera (Rauh) Gomez, Bre- nesia 18: 4. 1980. Corm vertically elongate, 20-67 mm tall, 11-15 mm in diameter, 1 -lobed (2-3-lobed in juvenile plants), roots arising acropetally along a single (2- 3) lateral fossa(e) (in juveniles arising synchro- nously within a continuous circumbasal fossa). Leaves 200 or more, stiffly erect in terrestrial in- dividuals, weakly recurved in submerged individ- uals, 30-50(-75) mm long, 3.5-6 mm wide at the base, 4-7 mm wide at mid-length, dark brown to black proximally, bright to dark green distally; alae colorless and hyaline to dark brown or black, char- taceous proximally, membranaceous distally, 1- 2.8 mm wide at the sporangium, 20-37 mm long (55-85% of the leaf length), each apex obtuse; su- bula trigonal, highly cutinized, strongly reflexed in terrestrial plants, weakly reflexed to arcuate in sub- merged plants, the apex attenuate to acute; fibrous bundles absent; stomates absent, scale leaves ab- sent. Sporangium elliptic to obovate, superficial to emergent, chocolate-brown, concolorous, 2.5-5.5 mm long, 1.7-3 mm wide, positioned 3.5-9 mm above the leaf base. Velum rudimentary, repre- sented by a narrow ridge of tissue along the lateral portions of the sporangium, absent along the upper edge of the sporangium. Ligule triangular, weakly 92 FIELDIANA: BOTANY cordate, off- white, 3-4.5 mm high, 2-2.5 mm wide; the margin delicate and ephemeral, the cushion persistent, deltate, 2-3 mm high, 1 .8-2.3 mm wide. Labium deltate to depressed-ovate, entire, dark brown, 280-320 urn high, 180-240 Mm wide. Megaspores mottled, gray on brown, or uniformly brown, not lustrous, 460-720 (x = 559) pm in diameter, laevigate to obscurely pustulate, the pus- tules occasionally merging laterally to form short meandriform rugae; equatorial and proximal ridg- es straight, distinct, as high or slightly higher than broad. Microspores dark brown, 35-43.8 (x = 39) Mm long, 28.8-37.5 (x = 32) pm wide, scabrate. 2n = 44. Vegetative reproduction by cortical gem- mae. In wet, boggy soil or inundated plains of puna vegetation from 4100 to 4900 m, Lima, Pasco (numerous collections), Junin, Cuzco, Puno. Peru and Bolivia. The inclusion of Stylites as a generic synonym of Isoetes is well supported by the early develop- ment of the sporophyte and the internal anatomy of the plants (Hickey, unpubl.) as well as the con- firmation of the chromosome number as 2n = 44. Earlier reports of 2n = 48-52 are incorrect. Lima: Prov. Huarochiri, Laguna Caprichosa, 4800 m, Rauh 186 (M). Lago Aguascocha, near Mina Caprichosa, 4780 m, Hutchison & Tovar 4244 (c, E, F, o, L, MICH, MU, s, uc, us, wis). Pasco: Prov. Pasco, Cerro de Pasco, ca. 4300 m, Asplund 11830 (s). Junin: Prov. Jauja, Pa- chayo, 3800 m, Ameghino (MU). Prov. Huancayo, La- guna Condoray, 4700 m, Marshall, 6 Sept. 1961 (BM). Prov. Junin, 13-14 km N of Junin, Karrfalt & Hunter 22 (USM). Prov. Tarma, Lago Junin, east side near road, 13 km north of Junin, 4100 m, Hutchison et al. 5890 (F, NY, uc, us). Cuzco: In high Andes above Cuzco, toward Puno, Gomez AS2189 (CR). Prov. Chumbivilcas, Laguna Huarmicocha, 4600 m, Curlier 241 (USM). Puno: Prov. Carabaya, pampa de Lacka Macusani, 4360 m, Vargas 7128 (us). 5. Isoetes dispora Hickey, sp. nov. Figure lOf. Differt ab aliis speciebus Isoetes, megasporis dimor- phis necnon laevibus, sporangiis paene superficiaribus, subulis plus minusve triquetris et labiis ligulatis. Corm globose to slightly elongate vertically, ca. 20 mm wide, ca. 30 mm high, 2-lobed, dichoto- mous roots arising synchronously from a complete circumbasal fossa. Leaves ca. 30, stiffly erect, dark brown to dark green proximally, dark green dis- tally, to 65 mm long, 6-8 mm wide at the base, 3.5-6 mm wide at mid-length; alae dark green, chartaceous to sub-membranaceous, 2-3 mm wide at the sporangium, 10-20 mm long (15-34% of the total leaf length), each apex attenuate; subula trigonal, highly cutinized, straight to slightly re- flexed, the apex blunt, subacute; fibrous bundles not evident; stomates absent; scales absent. Spo- rangia ovate to obovate, essentially superficial, basal, light brown, concolorous or occasionally with scattered dark spots, 2.5-4.5 mm long, 2-3.5 mm wide. Velum completely absent. Ligule ephemeral, occasionally represented by a dark auriculate frag- ment, 2.5 mm long by 2 mm wide. Labium light to dark brown, narrowly oblong, often cryptic, 1 40- 1 80 Mm long, 40-60 fim wide, the apex asymmet- ric, erose. Megaspores gray, not lustrous, laevigate to minutely tuberculate, dimorphic, showing Types I and II spores (sensu Marsden 1976), 750-1,125 (x = 920) Mm wide, the proximal and distal ridges laevigate, straight, distinct. Microspores gray en masse, 37.5-42.5 (x = 40.3) urn long, 26.3-32.5 (x = 29.9) Mm wide, laevigate to slightly papillate. TYPE— Peru, Dept. Lambayeque, Prov. Ferre- fiafe, Dist. Incahuasi, Jalca, Laguna Tembladera, Cerro Negro, 3300 m, Skillman et al. 12850 (ho- lotype, MU!; isotypes, F!, HUT!). High-altitude paramo (aquatic?), 3300 m, Lam- bayeque. Known only from the type specimen. Isoetes dispora is, despite being known only from the type collection, a very distinctive species. The combination of dimorphic, laevigate megaspores, nearly superficial sporangia, more or less trigonal subulae, no velum, and a strap-shaped labium dif- ferentiate it from all other known collections of Isoetes. The presence of dimorphic megaspores is normally taken as an indication of hybridity, often involving unbalanced genomes. The materials of Skillman et al., however, have certain anomalies suggesting that alternative possibilities should be entertained. Notable among these is the absence of any other collected taxa from the Lambayeque region. Hybrids are typically found in lower num- bers than the sum of their parental taxa and it seems unlikely that three individual Fl hybrids would be the sole collections from a mixed pop- ulation of quillworts. Furthermore, there is no ev- idence of vegetative reproduction in these speci- mens, suggesting that all three have had separate origins from spore. In addition, there is a previ- ously undescribed regularity to the production of the two spore types in this material. Careful dis- section of individual sporangia showed that each sporangium contained 20 spores, of which 1 0 were large and fully formed and contained cytoplasm, whereas the other 10 were flattened in the equa- TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 93 tonal plane and contained no cytoplasm. Because of space limitations within the sporangia, the spores derived from each megaspore mother cell retained their tetrad orientation, and so it was possible to observe directly the end products of all meiotic events ( 1 0 total) in two separate sporangia. In all cases tetrads consisted of two large globose spores and two smaller flattened ones. Taken together, these data are inconsistent with previous data on Isoetes hybrids, leading us to believe that these plants are either sexual or apogamous. Apogamy in the genus is relatively rare, being confined to a few species in Australia and India. In virtually all known cases of apogamy in Isoetes, microspores are either rare or lacking and the megaspores are irregular in shape, often forming Type HI spores or having single contact faces, i.e., diplospores. Such is not the case in Isoetes dispora. The presence of normal microspores (less than 10% abortion) and the extreme regularity of megaspore production suggest meiotic consistency, albeit ir- regular. The presence of two types of megaspores in this species may, therefore, reflect differential resource allocation among meiotic products in a fashion analogous to that seen during megaspore development in Selaginella and angiosperms and during oogenesis in mammals. 6. Isoetes hewitsonii Hickey, sp. nov. Species nova L. dispora proxima, cujus megasporis laevibus, velo carenti et subulis infirme triquetris habet. Differt megasporis uniformibus et foliis angustatis ha- bens apicibus attenuatisque minus corneis. Corm globose, slightly elongate laterally, ca. 1 5 mm wide, 2-lobed, with fleshy dichotomous roots arising from a continuous circumbasal fossa. Leaves light green, 10-20, erect, 50-160 mm long, 3.5-6 mm wide at the base, 1.5-1.6 mm wide at mid- length; alae light green to hyaline, chartaceous, 0.9-1.5 mm wide at the sporangium, 23-27 mm long (36-46% of the total leaf length), each apex attenuate; subula weakly triquetrous, highly cutin- ized, straight, the apex attenuate, scales absent. Sporangia elliptic, embedded, although only slightly so for the megasporangia, light tan, con- colorous, 2.8-4 mm long, 1.5-2 mm wide, basal. Velum completely absent. Ligule widely ovate, au- riculate, ephemeral, ca. 1.1 mm high by 0.6 mm wide, the margins entire. Labium tan to pale green, depressed-ovate, ca. 12 Mm high by 54 (j.m wide. Megaspores white, laevigate, 650-750 (x = 715) Mm in diameter; proximal ridges straight, distinct, the equatorial ridge distinct or not. Microspores gray en masse, 28-35 (x = 31.7) nm long, 21-28 (x = 24.9) Mm wide, with acute to truncate echinae. Vegetative reproduction by cortical budding. TYPE— Peru, Dept. Cajamarca, Prov. Celendin, desvio a Guagal siguiendo la ruta a Celendin, 3700 m, Sanchez Vega et al. 2021 (holotype, MU!; iso- types, ASU!, F!, CPUN). High-altitude aquatic endemic, 3700 m, so far known only from Cajamarca. This species is dedicated to Dr. Walter Hewit- son, a student of the fern genus Osmunda and the man who first introduced me to the excitement of both botany and Isoetes. A delicate and distinctive species, with the gen- eral appearance of a rigid Isoetes boliviensis, but with stiffer leaves and no velum. The large mega- spores suggest polyploidy. Additional collections should be sought to determine the chromosome number of this species and to encompass its vari- ation. 7. Isoetes saracochensis Hickey, sp. nov. Species foliis latis, subulus valde trigonis, apicibus acutis corneisque, sporangiis basalibus, velis fere com- pletis et megasporis laevibus ad ieniter rugulatis a con- generibus diversa. Corm globose, 2-lobed, 17-23 mm in diameter, with dichotomous roots arising from a continuous circumbasal fossa. Leaves 13-14, stiffly erect, 55- 135 mm long, 6-10 mm wide basally, 5-7 mm wide at mid-length; alae chartaceous, light to dark brown, extending 1.2-3 mm wide at the sporan- gium, 35-75 mm long (46-73% of the leaf length), apices acute; subula highly cutinized and strongly trigonal, with 3 distinct, rounded ridges continuing to the apex; leaf apices acute, sharp, corneous; fibrous bundles absent; stomata absent; scale leaves absent. Sporangium elliptic to ovate, concolorous, 2.5-6 mm long, 2.5-3.5 mm wide, basal. Velum virtually complete, extending (%)% or more down from the upper edge of the sporangium. Ligule delicate, transparent, broadly to narrowly ovate with a weakly cordate base, 1 .9-2.7 mm high, 1 .7- 2 mm wide, ephemeral, frequently degraded in mature leaves. Labium light to dark brown, a transverse ridge along the lower border of the fove- ola, entire, 100-150 Mm high, 500-800 Mm wide. Megaspores white to gray, 400-620 (x = 515) Mm in diameter, laevigate or with indistinct to distinct, short, low meandriform rugae; equatorial and proximal ridges straight, distinct, with equatorial ridges as broad as high and proximal ridges fre- 94 FIELDIANA: BOTANY quently twice as broad as high. Microspores dark gray to dark brown, 30-36 (x = 33.5) t*m long, 22-28 (x = 24.3) ^m wide, with clavate or echinate projections. TYPE— Peru, Dept. Puno, Laguna Saracocha, 14,000 ft, Tutin 1424 (holotype, BM; isotype, BM). Submerged or emergent in high-altitude lakes, 3900-4400 m, Cuzco, Puno. Endemic. This species is fairly well described despite the few collections available. It is distinctive but ob- viously belongs in an alliance close to 7. lechleri. The extensive velum and the distinctive labium are shared characteristics that are quite unusual in the genus for this part of the world. The /. lechleri complex (below), however, is distinctive unto it- self, having a number of unique characters (hor- izontally elongate corm, turgid leaf condition) that set it apart from 7. saracochensis. Cuzco: Chectuyoc, near Sicuna, 3950 m, Tutin 1406 (BM). Prov. Cafias, Laguna Langui y Layo, Chavez 2313, 2321 (MO). Puno: Prov. Carabaya, Lake Chungara, 4400 m, de Macedo & Enriquez (GH). Prov. Lampa, Laguna Lagunilla, ca. 4300 m, Tutin 1407 a, 1420 (BM). 8. Isoetes lechleri Mett., Fil. Lechl. 2: 36. 1859. TYPE: Peru, "in laguna cacuminis, Cordler. pr. Agapata," Lechler 1937 (holotype, B!; iso- types, G! [2 sheets], UPS!, frag. s!). Figure lOd-e. Isoetes soda A. Br., Verb. Bot. Ver. Brandenb. 4: 332. 1862. TYPE: Peru, in laguna cacuminis (mixta cum pi an tula repente ignota), Lechler 1937 b (ho- lotype, B!). Isoetes glacialis Asplund, Ark. Bot. 20A: 34-35. 1926. TYPE: Bolivia, Dept. La Paz, Prov. Murillo, Jainvags stationem La Cumbre, 4700 m, Asplund 4041 (holotype, UPS!; isotypes, G!, M!, s!). Isoetes laevis Weber, Hedwigia 63: 252. 1922. TYPE: Peru, Dept. Ancash, Cordillera regia uber Caraz, auf dem Grunde eines Tumpels, vollig unterge- taucht, 4400 m, Weberbauer 3111 (holotype, B!). Isoetes peruviana Weber, Hedwigia 63: 246. 1922. TYPE: Peru, Dept. Junin, Prov. Tarma, Beige westl. von Huacapistana, 3500 m, Weberbauer 2228 (holotype, B!; isotype, B!; frag., UPS!; photos, s and UPS of B). Corm (globose) laterally elongate, distinctly bi- lobed [more so in damaged specimens], 1 5-44 mm wide, 2-5 mm high; dichotomous roots arising synchronously within the circumbasal fossa. Leaves to 20-40, stiffly turgid and erect, 88-160(-240) mm long, 8-19 mm wide at the base, 2-3 mm wide at mid-length; alae dark green to dark brown, nearly black, 2-3.5 mm wide at the sporangium, 20-55 mm long (46-75% of the total leaf length), each apex attenuate; subula terete, dark green, the apex attenuate to subacute, corneous; fibrous bun- dles absent; stomates absent; scale leaves absent. Sporangium obovate to elliptic, hyaline, unspot- ted, 4-19 mm long, 3.5-4 mm wide, basal. Velum complete to nearly complete, extending 75-100% down the sporangium. Ligule ephemeral, ovate to very widely ovate, with an auriculate base, 1.2- 1.8 mm high, 0.8-2 mm wide, the more persistent cushion ovate to widely ovate, ca. 0.8-1 mm high, 0.4-0.6 mm wide. Labium not evident. Mega- spores white, laevigate, typically shiny, showing variable abortion, 280-440 (x = 340) ^m wide, the equatorial ridges indistinct in larger spores, the proximal ridges sharp, distinct. Microspores gray in mass, 33.8-38.8 (x = 36.2) »m long, 26.3-30 (x = 28.1) Mm wide, with acute to truncate pro- jections. 2n = 44. In high-altitude ponds and streams, typically submerged, 3300—4750 m, Cajamarca, San Mar- tin, Ancash, Junin, Ayacucho, Cuzco, Puno. Peru and Bolivia. The high-altitude ponds, lagunas, and streams of south-central Bolivia and central Peru are re- plete with laevigate-megaspored plants that have the same overall form but that differ from one population to another in the extent of pigmenta- tion, the size and rigidity of the leaf, and the amount of lateral growth in the corms. Quite a few of these variants have been formally recognized as discrete species and even more are now published (Fuchs- Eckert, 1 982) as nom. nud. Nearly all populations show some degree of meiotic abnormality as ev- idenced by irregularly shaped or sized spores. To date, all of these populations have been found to be tetraploids, with 2n = 44. The publication (Hickey, 1 984) of 2n = 44 for 7. "ticlioensis" Fuchs ined., I. glacialis, and 7. herzogii (in part) are re- ferable to this assemblage. The distinctive color- ation, corm shape, velum structure, and ligule form strongly suggest a common origin for these plants. All of these plants undoubtedly have had a com- mon hybrid origin, and it would appear that rapid speciation via reciprocal gene silencing is occur- ring. Each population, or perhaps even subsets within a population, is differentially silencing the tetraploid genome down to the diploid level. As this phenomenon proceeds, genetic isolation slow- ly builds up and functional spores become rarer and rarer. We are left, then, with a mosaic of vari- able morphologies, established by drift or perhaps selection, which are partially isolated from each TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. other and perpetuated by vegetative reproduction (corm budding). An examination of only a few populations gives a false sense of discreteness, and only after a broad survey can one observe the com- plete intergradation in forms. All of these popu- lations are, therefore, best included within a single species concept, /. lechleri. Sagdstegui et al. 13098 and 14347, while of the general form of 7. boliviensis, have much longer, narrower, more slenderly tapering leaves, with some pigmentation along the bases. Provisionally, it has been identified as 7. lechleri off. Likewise, Leon & Young 2833 is provisionally assigned to 7. lechleri. This collection has the darkened leaf bases and alae, laevigate megaspores, and exten- sive velum coverage typical of 7. lechleri. It differs in having a large (up to 5 mm long), strongly au- riculate ligule more typical of 7. andina of Ecuador, Colombia, and Venezuela and in having emargin- ate ala apices reminiscent of those seen in 7. kar- stenii (s.l.) of Colombia and Venezuela. The species is best distinguished by its brownish (almost caramel) coloration and the distinctly te- rete leaves that are turgid to the point of being brittle. Herbarium material is best identified by coloration, the presence of a horizontally elongate corm (in most populations), the laevigate mega- spores, the more or less complete velum, and the widely ovate ligule which has a tendency toward an auriculate base. Isoetes lechleri is a plant of high-altitude (over 3500 m) lakes, streams, and pools. It grows in both shallow and deep water, with an individual plant's form being strongly influenced by water depth. Shallow plants have short, stiffly rigid leaves. Plants of deeper water "etiolate": the leaves are a bit more lax and considerably longer. Vegetative re- production, by corm budding, is very common and may be a primary means of reproduction. The plants often form dense mats in quiet water; in streams and shallows the plantlets are often broken off and transported by animals or wave/current action. In Bolivia one road, cutting between two lakes, was completely covered by washed-out plants and corms. Cajamarca: Pozo Kuan, 3790 m, Sagdstegui et al. 13098 & 14011 (F, HUT, MU). Prov. Contumaza, Pozo Kuan, 3900 m, Sagdstegui et al. 14347 (F, HUT, MU). San Martin: Prov. Mariscal Caceres, forest on the edge of Laguna de Chochos, NW corner of Rio Abiseo Na- tional Park, 3300 m, Young & Leon 4856 (MU). Ancash: Prov. Huari, Huascaran National Park, 4440-4490 m, D. Smith et al. 12355 (HUT). Prov. Yungay, Laguna de Llanganuco, 4100-4200 m, Enderlin (USM). Lima: Prov. Cajatambo, Raura, perimetro de Ada, 4800-4900 m, Chanco & Montoya 188 (MU). Pasco (as Junin): Huaron, lake, 4200 m, Asplund 11828 (c, G [2 sheets], NY, s, UPS, us.). Junin: Prov. Yauli, Ticlio, small lake, 4750 m, As- plund 11669 (s). Ayacucho: Prov. Huanta, Mt. Razu- huillca, 4000-4100 m, Weberbauer 7500 (F, NY, us). Cuzco: Prov. Canas, Laguna de Langui y Layo, 3900 m, Chavez 2323 (GH). Prov. Paucartambo, Leon & Young 2833 (F, USM). Puno: Cordillera Real, Laguna Rinconada, 4680 m, Thomasson (s). 9. Isoetes boliviensis Weber, Nova Hedwigia 63: 247. 1922. TYPE: Bolivia, circa La Paz, via ad Coroico, Lancha, 5000 m, Mandon 1532 (holotype, G!; isotypes, BM!, G!). Corm globose to slightly elongate laterally, 5- 1 6 mm in diameter, 2-lobed; dichotomous roots arising synchronously within the circumbasal fos- sa. Leaves to 40, delicate, flexuous, erect to laxly spreading, 70-1 20(-240) mm long, 6-1 6 mm wide at the base, 1-1.5 mm wide at mid-length; alae hyaline to light brown, 1-2 mm wide at the spo- rangium, 20-30 mm long (1 2-33% of the total leaf length), each apex attenuate (rarely truncate); su- bula terete, bright green, the apex attenuate; fi- brous bundles absent; stomates absent; scale leaves absent. Sporangium obovate to elliptic, hyaline, unspotted, 5-8 mm long, 3-5 mm wide, basal. Velum incomplete to complete, extending 6-100% down the sporangium. Ligule ephemeral, widely ovate, with an auriculate base, ca. 2 mm high, 1.7 mm wide, the more persistent cushion narrowly depressed-ovate, often bilobed, ca. 0.2 mm high, 0.6 mm wide. Labium not evident. Megaspores white to light gray, laevigate to minutely tuber- culate, shiny, 380-460 (x = 433) /nm wide, the equatorial and proximal ridges sharp, distinct. Mi- crospores gray in mass, 26.3-40 (x = 33) um long, 20-30 (x = 25.3) /mi wide, with acute to truncate projections. 2n = 22. In shallow water, edges of lakes, ponds, and ver- nal pools, 4100-5000 m, Cajamarca, San Martin, Ancash, Lima, Pasco, Junin, Ayacucho, Cuzco, Puno. Peru and Bolivia. A very distinctive plant due to its small and relatively delicate appearance; several collectors have even suggested that it is an annual. This is the only Peruvian species with lax, flexuous leaves. It has a partial to complete velum normally cov- ering about 75% of the sporangium, corms that are slightly elongate horizontally, and megaspores that are laevigate to minutely tuberculate. It is found in shallow (= ephemeral?) portions of ponds and lakes. The microspores are quite distinctive 96 FIELDIANA: BOTANY in appearance, having acute or club-shaped pro- jections which themselves are spiny. Microspores of some /. lechleri collections are similar. The only counts for this species, 2n = 22, are from Bolivian material. nearly complete velum, and has megaspores that vary from tuberculate to rugulate to nearly lae- vigate. Isoetes herzogii is a high-altitude, nearly emergent plant of streams and shallow pools. La Libertad: Prov. Huamachuco, east of Quiruvilca, 4100 m, Hutchison et al. 6143 (F, GH, MO, NY, uc, us). Ancash: Prov. Yungay, Huascaran National Park, 4100- 4200 m, D. Smith et al. 10432 (F, MO, MU). Huascaran National Park, 4500 m, D. Smith et al. 9210 (MO, MU). Lima: Prov. Huarochiri, Ticlio Pass, between Lima and Oroya, 4840 m, Hutchison et al. 6081 (c, E, F, G, GH, HEID, L, LIL, M, MICH, MO, NY, s, uc, wis). Junin: 5 km above Hacienda Cochas, 34 km W of la Oroya-Huan- cayo Route 3 on road to Pachacayo, 4425 m, Keeley & Keeley 11087, 11088 (MU, occ). Comments Isoetes triquetra A. Braun, Verh. Bot. Vereins Prov. Brandenburg 3 (4): 332-333. 1862. TYPE: Peru, Sachapata (am ostlichen Abhang der Cordillera von Peru), in pascuis humidis, Lechler 3337 (holotype, B!; photos, s and UPS ofe). Calamaria triquetra (A. Braun) Kuntze, Rev. gen. pi. 2:828. 1891-1893. Isoetes lechleri var. triquetra (A. Braun) Gomez, Bre- nesia 18: 5. 1980. The type material for this species is very frag- mentary. There are no megaspores and the micro- spores are of a very common form (echinate). The most distinctive features of the specimen are the trigonal subula, the moderate labium, and the large auriculate ligule. In these regards, the type of L. triquetra is nearly identical to /. andina of Ecua- dor, Colombia, and Venezuela but differs from all other known Peruvian collections. I have argued (Taxon 35: 243-246. 1986) for the conspecificity of these taxa. The major argument against the common identity is distribution (some 1 ,400 km separate the type locality of I. triquetra from the nearest "good" locality of /. andina). Isoetes herzogii Weber, Hedwigia 63: 250. 1922. TYPE: Bolivia, Tunari, 4300 m, Herzog2083 (holotype, M; isotypes, L!, M!, us!; frag., UPS!). Although not recorded from Peru, this species is geographically proximate in Bolivia and should be expected in Peru. The species has stiffly erect, narrow, subulate leaves with attenuate apices. It lacks the pigmentation typical of /. lechleri, has a Addendum 1. Species to Be Added to the Pteridophyte Flora, Parts I-V The following species have come to the atten- tion of the authors as growing in Peru, after their generic treatment had been published. Trichomanes arbuscula Desv., Mem. Soc. Linn. Paris 6: 326. 1827. TYPE: "Crescit in Guia- na," P, Herb. Desvaux. Trichomanes coriaceumKunze, Linnaea9: 105. 1834. TYPE: "Prope Collares, Brasiliae," Poeppig in 1832 (Diar. 2981), not located. This species has an alate rachis and petiole and is sometimes subdimorphic. Illustrations are Hooker & Greville, Icones Filicum, t. 204. 1831 (as T. bancroftii) and Vareschi, Flora de Vene- zuela, Vol. 1 (Helechos), t. 47. 1969. Loreto: Prov. Maynas, Puerto Almendras, van der Werff et al. 9845 (MO), 9795 (MO), det. K. U. Kramer. Trichomanes crispumL., Sp.pl. 1097. 1753. TYPE: "Martinica," Plumier, Traite foug. Amer., t. 86. 1705. This is a critical and polymorphic species (see Windisch, Bradea 6: 99. 1992 for discussion). Huanuco: Prov. Pachitea, region of Pucallpa, Wall- nofer 11-261088 (z), det. K. U. Kramer. Pterozonium paraphysatum (A. C. Sm.) Lell., Mem. New York Bot. Card. 17: 13. 1967. Syngramma paraphysata A. C. Sm., Bull. Torrey Bot. Club 58: 301. 1931. TYPE: Venezuela, Amazo- nas, Cerro Duida, Tate 441 (NY; frag., us). This species is distinguished by the scales (vs. trichomes) on the stem, the keeled lamina, and relatively distant veins. TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 97 Huanuco: Prov. Pachitea, region of Pucallpa, low, elfin forest, 1750 m, Wallnofer 110-13988 (z), det. K. U. Kramer. Lindsaea phassa Kramer, Bot. Helvetica 101: 207. 1991. TYPE: Peru, Loreto, Rio Ampiyaco, vicinity of Pucaurquillo, Davis et al. 849 (ho- lotype, uc). Paratypes: Loreto: Prov. Maynas, Yanamono Explorama tourist camp, van der Werffet al. 9869 (MO, z), 7994 (MO, z). Kramer provides a discussion of the relation- ship of this new species and an illustration (fig. 1 , p. 228) of it. 2. Consideration of Pteridophyte Diversity in Respect to Ecology and Geography This treatment of the pteridophytes of Peru in- cludes 28 families, 118 genera, and 1,060 native or adventive species. The account includes all fam- ilies of the Pteridophyta except for four that occur only in the Old World and the Hymenophyllopsi- daceae of northern South America. It includes most of the genera in the Americas and about one-third of the Neotropical species. This indicates that Peru ranks high in pteridophyte diversity. It is not sur- prising that among the 24 departments of Peru, those with the greatest number of species are also those that are ecologically diverse. The six most speciose departments (map 1 ; table 1 ) are ecolog- ically strongly diverse, and they include relatively large areas of montane rain forest and/or cloud forest. All of these have been relatively accessible to collectors since the late 1 8th century, and with the exception of Pasco they are all large depart- ments. About 90% of the species and 86% of the endemics are in one or more of the six most spe- ciose departments. It is doubtful whether any additional families will be added to the pteridophyte flora except by segregation of those already present. A few genera may be added, such as Microlepia in Ecuador and Bolivia, Pilularia, known to occur in Bolivia, and Maxonia, known in Ecuador. Additional species most likely will be found in the northern depart- ments of Amazonas and Cajamarca and the south- ern departments of Madre de Dios and Puno. The number also may be increased by addition to large genera such as Selaginella and Thelypteris. The 1 30 species endemic to Peru are included in Table 1 and some of them are also on Map 1 . The map shows that there appears to be no clear geographic center of endemism. Rather, the de- partments that have the most endemics are those with the most species. This suggests that disrup- tions of ranges during the Pleistocene may have occurred throughout the Andes of Peru. However, there is need for more detailed knowledge of the Pleistocene in the Andean region, such as that for Europe published by Lang (1992). There are few studies of local diversity, although the ecological work of Young and Leon (1989, 1991) on pteridophytes of Peru supplies critical new data. Their analysis of a low elevation forested area in central Peru, including 61 species within 2 hectares, suggests that edaphic and topographic gradients affect the diversity of species. Their work on a high elevation forest, on the east slope of the Peruvian Andes, including 1 09 species in about 5 square km, shows greatest pteridophyte diversity in the montane rain forest. They suggest that the high and nearly constant humidity is a factor help- ing to account for the species richness. The Andes are the major feature of Peru that has molded the vegetation and distribution pat- terns of the biota. The principle types of vegetation and species diversity are in large part based upon altitudinal zones affecting temperature and pre- cipitation. The species of Peruvian pteridophytes are treated here in four main zones, based upon their ecology. They include Montana Ferns, Ceja Ferns, Sierra Ferns and Loma Ferns. The ecolog- ical zones are shown on Map 2, along with a di- agrammatic section showing the zones in profile. Montana Ferns occur in the forest region that covers the largest part of Peru. On the east flank of the Andes, this extends up to 1 800 m. This region receives 1500-3500 mm of rain annually, usually between October and April. It is difficult to recognize floristic elements among several hun- dred species in the Montana, but they may be characterized by their large size or by their epi- phytic habit. The large ones have creeping stems, often forming large colonies with continued growth throughout the year. The leaves may be more than a meter long and are often modified for vegetative reproduction. There may be a relatively large number in a given locality, but many may be rare species. The large-leaved species include Nephro- lepis biserrata (leaves to 3.6 m), Lygodium volubile (to 12 m), Hemidictyum marginatum (to 3 m), Adiantum pectinatum (to 2.5 m), and the tree ferns, mostly ca. 3 m. Some of the large-leaved species that form conspicuous colonies are Dennstaedtia cicutaria, Gleichenia bifida, Dicranopteris pectin- ata, and Pteridium aquilinum var. arachnoideum. The last is the most aggressive and may occupy 98 FIELDIANA: BOTANY COLOMBIA BOLIVIA CHILE! MAP 1 . The most species-rich departments in Peru (Cuzco and northward) are hatched. The number of species is above the name, the number of endemics below. Puno is included to illustrate the transition to the less species- rich regions of Bolivia. TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 99 TABLE 1 . Data on the diversity of pteridophyte spe- cies in Peru. Department Size (ca. 1,000 km2) Number of Pteridophyta Species Endemic to Peru Loreto 207 252 8 Cuzco 152 559 45 Madre de Dios 144 169 8 Ucayali 106 105 6 Puno 68 188 9 Arequipa 57 32 3 Ayacucho 47 145 7 San Martin 45 490 32 Huanuco 40 541 39 Piura 39 38 2 Lima (incl. Callao) 39 72 4 Ancash 38 86 5 Amazonas 36 398 31 Cajamarca 32 225 19 Pasco 30 393 31 Junin 29 457 24 La Libertad 26 128 14 lea 25 1 0 Apurimac 21 57 3 Huancavelica 21 73 2 Moquegua 14 4 1 Tacna 13 1 0 Lambayeque 12 51 3 Tumbes 4 20 0 whole hillsides after clearing for agriculture. Fer- tile leaves may be present only during a brief pe- riod as in Bolbitis serratifolia, B. lindigii, Poly- botrya caudata, and P. osmundacea. Some species may be more common, or confined to more trop- ical areas, at altitudes below 300 m. This region may be distinctive, based on distributions of flow- ering plants, but it is clearly a part of the Montana Fern flora. The Montana Ferns have two main types of leaf modification for vegetative reproduction. Species that have buds toward the apex of the lamina in- clude Polystichum platyphyllum, Thelypteris mac- rotis, Bolbitis serratifolia, and Diplazium macro- phyllum. Doryopteris pedata var. palmata has buds at the base of the lamina, and Tectaria incisa has buds at the base of pinnae and along the pinna- rachises. In these species the buds are persistent and develop into plantlets, especially on old leaves, while still attached to the leaf. Deciduous buds are produced in the axils of pinnae in Dennstaedtia arborescens. Other species have leaves that root at the tip of the rachis.The rachis tip is elongated as in Adiantum deflectens, Trichomanes diversi- frons, and Asplenium radicans. All or nearly all of the leaves in Asplenium radicans become rooted. The epiphytic ferns that are primarily plants of the forest include Grammitis serrulata, Pecluma filicula, Campyloneurum angustifolium, Polypo- dium polypodioides var. burchellii, Pleopeltis per- cussa, Asplenium serrulatum, and Ophioglossum palmatum, as well as several species of Elapho- glossum. Some may grow on bare rock or may survive, for a time, on fallen branches. These spe- cies are biologically similar to the Sierra Ferns. They may become dormant during the relatively mild dry season, especially if growing on more exposed branches. Many Montana Ferns are not sufficiently common to use in defining the zone, but the presence of numerous species of Adiantum with large dimidiate segments, only in the Mon- tana, helps to characterize this zone. These include Adiantum petiolatum, A. latifolium, A. macro- phyllum, A. anceps, A. platyphyllum, A. pectina- tum, A. tetraphyllum, A. tomentosum, A. pulver- ulentum, and A. villosum. Ceja Ferns occur in the region extending along the higher eastern slopes or low summits of the Andes, at elevations of about 1800-3000 m. This is a moist, cool region where clouds and fog are present most of the year. It is a transitional cli- matic and vegetational band: the Ceja de la Mon- tana. Tree ferns and the bamboo Chusquea are the dominant elements in the Ceja Region. Under moist conditions where fog often occurs, frequent species are Hymenophyllum ruizianum, Eriosorus flexuosus, Lycopodium clavatum, L. jussiaei, and Lycopodiella pendulina. The absence of Adiantum species with large dimidiate segments, as noted above in the characterization of the Montana, helps to define the Ceja zone. In drier situations lacking mists, most of the ferns are terrestrial, the leaves less than 6 dm long. They are usually fertile and not modified for veg- etative reproduction. In these features the plants generally resemble those of the Sierra species. They grow more or less throughout the year or at least retain leaves in fresh condition. A mixture of Si- erra species such as Pellaea ovata, P. sagittata, and Woodsia montevidensis may occur here, as well as species more characteristic of the Montana such as Niphidium crassifolium or Nephrolepis pectin- ata. The Ceja Ferns are largely a mixture of species from the Montana and Sierra rather than species restricted to the Ceja. Sierra Ferns center in the high central part of the Sierra, or Altiplano, between 3000 and 4300 m elevation. The land is somewhat rolling or may be rather flat. There is considerable relief where valleys penetrate the mountains. The region is bor- dered, except in the north, by chains of high moun- 100 FIELDIANA: BOTANY COLOMBIA BRAZIL BOLIVIA CEJA FERNS SIERRA FERNS 1800m 3000m MAP 2. The main vegetational zones in Peru and their ferns (diagrammatic). TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 101 tains, many of which rise to 5500 m or more and support snow fields and glaciers. The temperature is relatively cool and the air is dry. The vegetation largely consists of grasses, sedges, semi-desert shrubs and cacti. Locally there are small woods, especially those dominaed by Polylepis. The puna, the land above the limits of agriculture, extends from 4000 m up to the limit of vascular plants, at about 5200 m. Here cold prevails and low cushion plants are the principal vegetational feature. The Sierra Ferns are seasonally dormant but the dormancy may be intermittent due to infrequent rains during the dry season. The available mois- ture is largely in the soil or rock crevices. Many species occur where there is local seepage or where conditions are suitable for condensation of at- mospheric moisture. The annual rainfall, between 500 and 1 1 00 mm, is by no means deficient but the dry air, winds, and strong insolation at the high altitudes combine to diminish the moisture. One of two main kinds of Sierra Ferns is the mesic type, typical of the locally moist habitats. Leaves of these plants die during the dry season unless they are in an unusually favorable place. Species characteristic of this habitat include Adiantum poiretii, A. digitatum, Cystopteris fra- gilis, Asplenium peruvianum, and Woodsia mon- tevidensis. Asplenium peruvianum and the closely related A. gilliesii are exceptional in this group in having proliferous buds on the leaves. The petioles of these plants elongate and act as stolons. The petiole persists after the lamina withers and a plantlet develops at what appears to be the tip of the rachis. Xeric species are the second characteristic type of Sierra Ferns. These plants retain their leaves but they become curled during the dry season, reviving during the brief moist periods. Species characteristic of this group include Cheilanthes in- carum, C. pruinata, C. bonariensis, C. scariosa, Polypodium pycnocarpum, Pellaea ternifolia, and Notholaena nivea. Loma Ferns occur on the Pacific side of the Andes, usually near the coast. This region becomes progressively drier toward the ocean and finally barren, or with very sparse vegetation. In the north of Peru, these desert conditions are less pro- nounced. At the northernmost tip of Peru there is a small forested area. The otherwise barren coastal zone is relieved, at intervals, by green, irrigated valleys, and by the naturally verdant Lomas. The ferns of this region are part of a unique vegetation that occurs along the coast of Peru and Chile, north to about 8 degrees south latitude. This vegetation develops in response to local physiographic con- ditions. In winter there is more or less constant fog and "guara" (a dense mist) on certain hills and upper parts of valleys. The summer months are continuously dry. A rather lush vegetation may develop where unusual moist conditions occur. The long, continuously dry periods evidently re- strict the flora. Loma Ferns include a selection of the Sierra species including Polypodium pycno- carpum, Adiantum subvolubile, and A. digitatum and, less commonly, Ophioglossum reticulatum, Anogramma leptophylla, Woodsia montevidensis, and Cheilanthes peruviana. It is of interest to note that while about 80% of the Loma seed plants are endemic, there are no endemic ferns. This may be due to slower evolution of the ferns but more likely reflects their superior means of dispersal. The spores undoubtedly are blown from the Sierra to the Lomas with sufficient frequency to prevent endemism. References LANG, G. 1992. Some aspects of European late- and post-glacial flora history. Acta Bot. Fennica 144: 1-17. YOUNG, K. R., AND B. LEON. 1 989. Pteridophyte diversity in the central Peruvian Amazon: Im- portance of edaphic specialization. Brittonia 41: 388-395. YOUNG, K. R., AND B. LEON. 1991. Diversity, ecology and distribution of high-elevation pte- ridophytes within Rio Abiseo National Park, north-central Peru. Fern Gaz. 14: 25-29. 102 FIELDIANA: BOTANY Colombia g-f i... 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 1 3. JunTn 14. Ucayali 15. lea 1 6. Huancavelica 1 7. A y acucho 1 8. Apurfmac 1 9. Cuzco 20. Madre de Dios 2 1 . Arequipa 22. Puno 23. Moquegua 24. Tacna Bolivia Chile DEPARTMENTS OF PERU TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 103 Comprehensive 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. Prefixes (1-6) indicate the number of the volume or part, followed by a hyphen and then the page number within that part. (Thus, the prefix "6" indicates entries for the present series number of Fieldiana, whereas lower numbers indicate entries for previously published parts, their series numbers as follows: 1 = New Series No. 20; 2 = No. 22; 3 = No. 29; 4 = No. 27; 5 = No. 32.) Entries for Parts I through V pertain only to names of species and genera. Listings for categories above and below these taxa may be found in the indices to those particular parts. Acrospermum maxonii 5-112 Acrostichum 2-81 acuminatum 4-154 adenolepis 4- 1 66 albescens 4-121 alcicorne 5-181 alienum 4- 1 00 apodum 4-157 aureonitens 2-48 aureum 2-83 auricomum 4-125 bakeri 4-125 barbatum 4-125 bonariense 2-28 calaguala 4-137 calomelanos 2-18 calophyllum 4-166 castaneum 4- 1 27 caudatum 4-64 cervinum 4-85 chrysoconium 2-20 chrysolepis 4-6 1 chrysophyllum 2-16 ciliatum 4-128 citrifolium 2-92 cladotrichum 4-125 curvans 4- 1 66 cuspidatum 4- 1 29 danaeifolium 2-83 decoration 4-129 denticulatum 4-131 dichotomum 1-33 digit at um 1-33 discolor 4- 1 30 dissimile 4- 1 66 diversifolium 4- 1 30 ebeneum 2- 1 8 elegans 1-34 elongatum 4- 1 62 engelii 4-131 erinaceum 4-132 erythrolepis 4- 1 32 eximium 4- \ 33 fendleri 3-68 flabellatum 4-169 flaccidum 4-133 fractiseriale 4-60 furfuraceum 4- 1 64 glabellum 4- 1 34 graminioides 5-72 guamanianum 4-135 hackelianum 4-6 1 hartwegii 4-135 hayesii 4-136 haynaldii 4-136 heterophyllum 5-63 hickenii 4-136 horridulum 4-137 huacsaro 4-137 /zysm';t4-138 ilvense 4-94 insigne 4-6 1 japurense 4-107 juglandifolium 4-6 1 laminarioides 4-139 lanuginosum 2-32 latifolium 4-140 lechlerianum 4-141 leprosum 4- 1 42 leptophyllum 4- 1 46 lindenii 4-142 /inrf/gii'4-103 lingua 4-142 litanum 4-143 luridum 4- 1 44 mathewsii 4-144 minutum 4-146 moorei 4-167 muscosum 4-147 nicotianifolium 4-102 nigrescens 4-148 nivosum 4-148 nodosum 1-15 oligarchicum 4-102 orbignyanum 4-151 osmundaceum 4-62 pachyphyllum 4-151 paleaceum 4-152 pandurifolium 4-105 papillosum 4-152 patinii 4-153 peltatum 4- 1 69 petiolosum 4-154 phlebodes 4-107 piloselloides 4- 1 54 plumbicaule 4-60 plumosum 4-155 poeppigianum 4-156 polybotryoides 4-6 1 polypodioides 5-137 preslianum 4-128 propinquum 4-156 quitense 4-157 reptans 5-153 rufum 2-47 scariosum 2-32 schomburgkii 4- 1 44 serratifolium 4-101 serratum 4-101 serrulatum 5-83 setigerum 4- 1 60 sinuatum 2-34 sorbifolium 4-105 sphenophyllum 4-169 squamipes 4-161 stenopyllum 4-161 suberectum 4-61 tambillense 4- 1 62 tartareum 2-19 tectum 4-162 tenuiculum 4-163 tereticaulon 2-40 thalictroides 2-50 thelypteris 3-6 trifoliatum 2-21 unitum 4-146 villosum 4-156 Actinostachys 1-33 digit ata 1-33 pennula 1-36 Adiantopsis 2-34 chlorophylla 2-36 paupercula 2-34 radiata 2-36 ternata 2-36 Adiantum 2-52 alarconianum 2-67 amabile 2-56 anceps 2-68 capillus-veneris 2-58 cayennense 2-64 ceciliae 2-63 chilense 2-57 concinnum 2-58 crenatum 2-57 cuneatum 2-56 decorum 2-56 deflectens 2-62 delicatulum 2-62 digitatum 2-61 dolabriforme 2-62 filiforme 2-62 flagellum 2-62 fructuosum 2-64 104 FIELDIANA: BOTANY fuliginosum 2-64 guianense 2-120 henslovianum 2-59 hirtum 2-64 humile 2-66 imbricatum 2-60 incisum 2-67 kalbreyeri 2-63 kaulfussii 2-66 killipii 2-66 laetum 2-59 lancea 2-121 latifolium 2-66 lobatum 2-60 lucidum 2-69 lunulatum Burm. 2-62 lunulatum Houtt. 4-40 macrocladum 2-64 macrophyllum 2-70 mathewsianum 2-67 mexiae 2-64 microsorium 2-62 moorei 2-56 obliquum 2-57 orbignyanum 2-60 palmatum 2-6 1 patens 2-6 1 pauperculum 2-34 pectinatum 2-62 pedatum 2-61 peruvianum 2-68 petiolatum 2-66 phillipense 2-62 phyllitidis 2-70 pilosum 2-63 platyphyllum 2-68 poeppigianum 2-69 poiretii 2-57 pulverulentum 2-65 raddianum 2-56 radiatum 2-36 rhizophyllum 2-62 ruizianum 2-62 scalare 2-69 serratodentaum 2-65 sessilifolium 2-59 speciosum 2-61 steerei 2-62 strictum 2-121 subaristatum 2-62 subvolubile 2-59 sulphureum 2-58 terminatum 2-65 tetraphyllum 2-64 thalictroides 2-57 tinctum 2-56 tomentosum 2-63 urophyllum 2-63 x variopinnatum 2-66 veitchianum 2-62 villosissimum 2-64 villosum 2-64 Aleuritopteris farinosa 2-23 peruviana 2-30 Allantodia asplenioides 4-75 Alsophila 1-116 armigera 1-123 aterrima 1-1 14 austral is 1-1 16 blechnoides 1-111 capensis 1-118 caracasana 1-133 conjugata 1-128 contracta 1-109 cuspidata 1-120 dombeyi 1-123 elongata 1-115 engelii 1-116 erinacea 1-118 floribunda 1-123 frigida 1-126 incana 1-120 infesta 1-123 kalbreyeri 1-124 ft/ftp// 1-125 lasiosora 1-125 latevagans 1-124 fcc/z/en 1-128 macrosora 1-115 melanopus 1-124 microdonta 1-127 H/gra 1-124 nigripes 1-124 pallescens 1-131 paucifolia 1-118 peruviana 1-123 phegopteroides 1-127 pilosissima 1-125 podophylla 1-124 poeppigii 1-115 procera 1-123 pruinata 1-109 pterorachis 1-123 pubescens 1-126 pycnocarpa 1-123 quadripinnata 1-109 rostrata 1-111 sprucei 1-115 swartziana 1-122 tarapotensis 1-125 tryonorum 1-128 w/e/ 1-128 Amauropelta 3-9 breutelii 3-9 cheilanthoides 3-34 concinna 3-29 5-22 ternatum 5-22 theciferum 5-48 tomentosum 2-46 trapezoides 5-48 tricholepis 5-30 trichomanes-dentatum 5-34 trilobatum 5-22 trilobum 5-48 triphyllum 5-22 tucumanense 5-26 tuerckheimii 5-29 tungurahuae 4-76 uniseriale 5-2 1 vargasii 5-17 vastum 4-73 venulosum 4-76 virens Desv. 5-3 1 virens Presl 5-30 106 FIELDIANA: BOTANY vomeriforme 5-28 wagneri 5-17 weberbaueri 5-49 Athyrium 4-88 achilleifolium 5-32 ambiguum 4-7 1 "bradearum" 4-88 celtidifolium 4-83 dombei 4-90 dombeyi 4-90 expansum 4-75 ferulaceum 4-90 filix-femina 4-88 flexuosum 4-77 fumaroides 4-92 haenkeanum 5-24 praestans 4-87 Austrolycopodium 6-52 magellanicum 6-56 Azolla 6-8 caroliniana 6-10 filiculoides 6-11 var. rw&ra 6- 1 1 mexicana 6-10 microphylla 6- 1 1 rubra 6- 1 1 Balantium karstenianum 1-105 Blechnum 5-56 acutum 5-64 andinum 5-61 angustifolium 5-64 arborescens 5-62 asplenioides 5-60 auratum 5-67 auriculatum 5-68 binervatum 5-63 blechnoides 5-60 brasiliense 5-6 1 buchtienii 5-67 caudatum 5-58 chilense 5-62 ciliatum 5-68 cognatum 5-68 columbiense 5-67 confluens 5-59 cordatum 5-62 delicatum 5-68 divergens 5-63 ensiforme 5-64 fragile 5-64 fraxineum 5-59 glandulosum 5-58 gracile 5-59 heterophyllum 5-63 kunthianum 5-64 1'herminieri 5-61 lanceola 5-60 lechleri 5-68 lehmannii 5-61 longifolium 5-59 loxense 5-65 magellanicum 5-68 malacothrix 5-68 maxonii 5-61 meridense 5-64 nigrosquamatum 5-61 nudum 5-56 obtusifolium 5-67 occidentale 5-58 orient ale 5-58 ornifolium 5-62 pectinatum 5-58 penna-marina 5-62 peruvianum 5-62 polypodioides Raddi 5-60 polypodioides (Sw.) Kuhn 5-64 pteropus 5-63 rubicundum 5-65 scandens 5-70 schomburgkii 5-66 serrulatum 5-60 sprucei 5-66 squamulosum 5-65 stenophyllum Presl 5-65 stenophyllum (Klotzsch) Mett. 5-65 stipitellatum 5-65 tr (angular e 5-59 trilobum 5-68 unilateral 5-60 volubile 5-70 Blotiella2-lll glabra2-lll lindeniana 2-113 Bolbitis 4-98 aliena4-100 bradeorum 4-103 crenata 4-101 guianensis 4-109 fci7/(p«4-102 Iindigii4-103 macrophylla 3-68 nicotianifolia 4-102 oligarchica 4-102 pandurifolia 4-105 portoricensis 4-101 serrata 4-101 serratifolia 4-101 stuebelii 4-103 Botrychium 1-6 cicutarium 1-8 lun an a 1-6 mirifica 1-58 schaffner 1-6 underwoodianum 1-6 virginianum 1-8 virginicum 1-8 Bryodisma 6-66 Byrsopteris 4-35 aristata 4-35 Caenopteris achilleifolia 5-32 myriophylla 5-25 Calamaria triquetra 6-97 Camptosorus 5-2 Campyloneurum 5-158 abruptum 5-172 aglaolepis 5-167 amphostenon 5-165 angustifolium 5-168 angustipaleatum 5-169 aphanophlebium 5-161 asplundii 5-167 brevifolium 5-170 caespitosum 5-162 chlorolepis 5-168 coarctatum 5-164 decurrens 5-132 densifolium 5-166 fasciale 5-163 fuscosquamatum 5-163 heterolepis 5-168 inflatum 5-164 irregular e 5-166 jamesonii 5-172 lapathifolium 5-162 /atara 5-170 lorentzii 5-167 magniftcum 5-132 nitidissimum 5-171 nodosum 5-172 occultum 5-161 ophiocaulon 5-162 pascoense 5-171 phyllitidis 5-169 repens 5-162 serpentinum 5-163 solutum 5-172 sphenodes 5-164 taeniosum 5-169 trichiatum 5-161 vulpinum 5-167 Ceradenia 5-72 capillaris 5-90 curvata 5-72 dendroxa 5-88 discolor 5-86 farinosa 5-89 herrerae 5-88 longipinnata 5-85 meridensis 5-85 mirabilis 5-91 pearcei 5-87 pilipes 5-90 praeclara 5-89 terrestris 5-87 Ceratopteris 2-50 pteridoides 2-50 richardii 2-50 thalictroides 2-50 Ceropteris adiantoides 2- 1 9 Ceterach aspidioides 3-13 Cheilanthes 2-23 andina 2-27 arequipensis 2-33 bonariensis 2-28 borsigniana 2-37 buchtienii 2-34 TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 107 cantangenste 2-32 chlorophylla 2-36 concolor 2-3 1 crenata 4-94 elegans 2-31 farinosa 2-30 fasciculata 2-27 fractifera 2-25 fraseri 2-28 glandulosa 2-106 glauca 2-30 hostilis 2-\\Q hypoleuca 2-34 incarum 2-33 intramarginalis 2-23 lonchophylla 2-33 macleanii 2-27 marginata 2-30 mathewsii 2-27 micropteris 2-23 mollis 2-29 moritziana 2-25 myriophylla 2-3 1 notholaenoides 2-25 obducta 2-29 obtusata 2-\\\ orbignyana 2-29 ornatissima 2-32 parallelogramma 2-110 peruviana 2-32 pilosa 2-27 pilosa x pruinata 2-27 poeppigiana 2-30 pruinata 2-27 radiata 2-36 rigida 2-34 rufopunctata 2-29 saundersii 2-25 scariosa 2-32 sinuata 2-34 squamosa 2-34 tripinnata 2-32 Cheiroglossa 1-8 palmata 1-9 Christella 3-39 dentata 3-42 hispidula 3-41 parasitica 3-39 Christensenia 1-13 Chrysodium lindigii 4-103 serratum 4-101 Cnemidaria 1-136 alatissima 1-138 cocleana 1-139 horrida 1-138 nervosa 1-139 speciosa 1-139 uleana 1-138 Cochlidium 5-72 graminioides 5-72 pumilum 5-82 serrulatum 5-82 Coptophyllum 1-24 buniifolium 1-24 Cryptosorus 5-72 blumei 5-72 Ctenitis 4-5 ampla 4-10 andicola 4- 1 9 biserialis 4-15 catocarpa 4-10 distans 4-5 hirsuto-setosa 4-16 honesta 4-14 karsteniana 4-17 microchlaena 4-9 mollicoma 4-17 nemophila 4-10 nigrovenia 4-9 protensa 4-2 1 pulverulenta 4- 1 7 refulgens 4-7 sloanei 4-10 subincisa 4-10 submarginalis 4-8 Ctenopteris 5-72 amylacea 5-87 anfractuosa 5-99 apiculata 5-94 asplenifolia 5-104 athyrioides 5-113 capillaris 5-90 congesta 5-89 contacta 5-107 crispata 5-72 cultrata 5-108 discolor 5-86 dolor ensis 5-104 ecuadorensis 5-110 farinosa 5-89 ./zrma 5-100 gracilis 5-96 herrerae 5-88 heteromorpha 5-105 lanigera 5-106 leucosticta 5-111 longipinnata 5-85 longiuscula 5-111 major 5-96 melanosticta 5-98 meridensis 5-85 moniliformis 5-98 obovata 5-95 peruviana 5-102 phlegmaria 5-97 pilosissima 5-101 pseudocapillaris 5-94 pseudonutans 5-103 pteropus 5-95 r/ge«5 5-101 rigescens 5-102 semihirsuta 5-112 sericeo-lanata 5-106 ste//a 5-106 subflabelliformis 5-107 subimpressa 5-94 subsessilis 5-95 taxifolia 5-113 tunguraguae 5-95 venulosa 5-72 yungensis 5-113 Culcita 1-103 coniifolia 1-103 macrocarpa 1-103 Cuspidaria 5-145 furcata 5-145 Cyathea 1-129 andina 1-130 arborea 1-129 aterrima 1-1 14 bipinnatifida 1-126 caracasana 1-133 castanea 1-135 cuspidata 1-120 delgadii 1-132 divergens 1-131 dudleyi 1-135 ebernina 1-135 elongata 1-117 equestris 1-131 erinacea 1-118 frigida 1-126 fulva 1-135 incana 1-120 kalbreyeh 1-124 lasiosora 1-125 latevagans 1-124 lechleri 1-135 macrosora 1-115 meridensis 1-135 mexicana 1-133 microdonta 1-127 microphylla 1-132 multiflora 1-130 multisegmenta 1-132 n/#ra 1-125 nigripes 1-124 oligocarpa 1-132 oreites 1-115 pallescens 1-131 panamensis 1-131 petiolata 1-131 phegopteroides 1-127 pilosa 1-132 pilosissima 1-125 poeppigii 1-115 polystichoides 1-118 primaeva 1-125 procera 1-123 pubens 1-127 pubescens 1-127 pungens 1-123 quindiuensis 1-116 rufescens 1-1 14 ruiziana 1-132 schanschin 1-132 sprucei 1-115 subtropica 1-128 wte/ 1-128 vilhelmii 1-130 willdenowiana 1-123 Cyclodium 4-47 guianense 4-48 meniscioides 4-48 trianae 4-47 Cyclopeltis 4-29 semicordata 4-29 108 FIELDIANA: BOTANY Cyclosorus 3-39 dentatus 3-42 gongy lodes 3-40 Cyrtomium 4-38 dubium 4-38 falcatum 4-40 nobile 4-38 Cyrtophlebium 5-158 phyllitidis 5-169 repens 5-162 Cystodium 1-103 Cystopteris 4-92 fragilis 4-92 translucens 4-92 Danaea 1-15 cuspidata 1-19 elliptica 1-18 grandifolia 1-17 humilis 1-20 jamaicensis 1-19 longifolia 1-17 moritziana 1-19 nodosa 1-17 oblanceolata 1-18 stenophylla 1-19 trichomanoides 1-20 wendlandii 1-20 Davallia arborescens 2-100 concinna Presl 2-100 concinna Schrader 5-4 glauca 2-99 inaequalis 2-103 multiflora 5-52 thecifera 5-48 Davalliopsis elegans 1-84 Dennstaedtia Bernh. 2-95 Dennstaedtia Moore 2-95 Dennstaedtia arborescens 2- 1 00 bipinnata 2-99 cicutaria 2-98 concinna 2-100 dissecta2-100 distenta 2-95 erosa 2-100 flaccida 2-95 glauca 2-99 globulifera 2-99 mathewsii 2-100 obtusifolia 2-100 pearcei 1-99 punctilobula 2-95 rubiginosa 2-98 sprucei 2-101 wercklei 2-101 Deparia mathewsii 2-100 Dicksonia 1-105 arborescens 1-105 berteriana 1-105 bipinnata 2-99 cicutaria 2-98 coniifolia 1-103 dissecta 2-100 erosa 2-100 gigantea 1-105 karsteniana 1-105 montevidensis 4-94 obtusifolia 2-100 pearcei 1-99 polypodioides 2-95 rubiginosa 2-98 sellowiana 1-105 spruceana 1-105 stuebelii 1-105 Dicranoglossum 5-49 desvauxii 5-147 furcatum 5-145 panamense 5-145 polypodioides 5-148 subnudum 5-147 Dicranopteris 1-46 o^nw 1-42 bancroftii 1-39 ftCfrfa 1-41 brittonii 1-44 dichotoma 1-46 flexuosa 1-47 linearis 1-47 longipinnata 1-45 nervosa 1-47 pectinata 1-49 pennigera 1-39 peruviana 1-45 pruinosa 1-42 remota 1-44 rubiginosa 1-46 schomburgkiana 1-47 seminuda 1-47 simplex 1-39 ve/ata 1-42 yungensis 1-44 Didymochlaena 4-40 lunulata 4-40 sinuosa 4-40 truncatula 4-40 Didymoglossum 1-76 angustifrons 1-86 hymenoides 1-86 krausii 1-87 membranaceum 1-88 muscoides 1-86 reptans 1-87 sphenoides 1-86 Diphasiastrum 6-52 thyoides 6-57 Diphasium 6-52 jussiaei 6-56 Diplazium 4-65 aberrans 4-87 alienum 4-72 altissimum 4-70 ambiguum 4-7 1 angelipolitanum 4-79 appolinaris 4-77 arboreum 4-80 asplenioides 4-75 bicolor 4-74 bogotense 4-70 bombonasae 4-8 1 bonapartii 4-75 bradeorum 4-88 brasiliense 4-79 buchtienii 4-72 callipteris 4-83 caracasanum 4-79 celtidifolium 4-83 centripetale 4-88 chimborazense 4-87 costale 4-77 crassifolium 4-76 cristatum 4-80 cuneifolium 4-82 delitescens 5-32 denticulosum 4-80 diplazioides 4-71 divergens 4-70 drepanolobium 4-8 1 eggersii 4-88 expansum 4-75 ferulaceum 4-90 flavescens 4-85 flexuosum 4-76 fraxinifolium 4-67 fuscopubescens 4-72 fuscum 4-87 gracilescens 4-72 grande 4-78 grandifolium 4-82 hians 4-70 induratum 4-78 lechleri 4-85 legalloi 4-84 lehmannii 4-72 lindbergii 4-78 lonchophyllum 4-80 macrodictyon 4-87 macrophyllum 4-77 melanopus 5-3 1 melanosorum 4-75 moritzianum 4-73 obscurum 4-85 obtusum 4-79 oxylobum 4-77 pactile 4-87 paucijugum 4-84 pedatum 4-74 pinnatifidum 4-86 plantagineum 4-86 plantaginifolium 4-86 praestans 4-87 preslianum 4-76 remotum 4-74 riedelianum 4-85 roemerianum 4-85 rostratum 4-75 sandwichianum 4-72 shepherdii 4-80 striatum 4-79 stuebelianum 4-80 subnudum 4-79 subobtusum 4-78 tabalosense 4-79 tarapotense 4-73 TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 109 tungurahuae 4-76 unilobum 4-81 vastum 4-73 venulosum 4-75 verapax 4-85 werckleanum 4-80 Diplopterygium 1-37 bancroftii 1-39 Doryopteris 2-43 concolor 2-3 1 crenulans 2-44 lomariacea 2-44 lorentzii 2-44 palmata 2-44 pedata 2-44 Drynaria acuminata 5-156 Dryopteris 4-35 ampla 4-10 anceps 3-68 ancyriothrix 3-63 andicola 4-19 andreana 3-76 angustifolia 3-74 arcana 3-76 aristata 4-35 aspidioides 3-13 assurgens 3-30 asterothrix 3-62 atrorubens 3-24 balbisii 3-3 1 bangii 3-42 biformata 3-60 biserialis 4- 1 5 boqueronensis 3-26 brachyodus 3-49 brachypus 3-18 brausei 3-28 canadasi 3-2 1 catocarpa 4-10 caucaensis 3-28 cheilanthoides 3-34 christii 4-45 chrysodioides 3-69 clypeata 3-65 coarctata 3-32 columbiana 3- 1 5 comosa 3-49 concinna 3-29 consobrina 3-73 contermina 3-32 corazonensis 3-27 ctenitis 4-5 decussata 3-49 deflexa 3-30 deltoidea 3-46 dmsa 3-34 densiloba 3-52 dent at a 3-42 denticulata 4-37 desvauxii 3-71 diplazioides 3-14 dumetorum 3-27 eggersii 3-59 engelii 3-26 ensiformis 3-75 euchlora 3-25 extensa 3-43 falcata 3-75 fibrillosa 4-13 filix-mas 4-35 /wrva 3-19 gigantea 3-68 glandulosa 3-49 glandulosolanosa 3-20 heterophlebia 4-45 hirsuto-setosa 4- 1 6 honesta 4-14 incana 3-52 jamesonii 3-56 jurgensenii 3-75 juruensis 3-61 karsteniana 4- 1 7 karstenii 4-9 killipii 4-33 laevigata 3-24 /ec/z/m' 3-69 leprieurii 3-5 1 leptosora 4-14 leucothrix 3-32 limaensis 3-20 lindigii 3-30 lingulata 3-75 linkiana 3-14 lomatosora 3-15 longicaudata 4-44 lugubriformis 3-58 macbridei 3-19 macrophylla 3-68 macrostegia 4-38 macrotis 3-57 mapiriensis 3-49 megalodus 3-64 mercurii 3-31 microchlaena 4-9 microsora 4-14 mi//« 3-28 mollicoma 4-11 mollis 3-42 multiformis 3-35 nemophila 4-10 nigrovenia 4-9 w'teHs 3-35 ochropteroides 4-38 oligocarpa 3-15 oligophylla 3-45 opaca 4-45 pachyrhachis 3-3 1 paleacea 4-36 paludosa 4-48 parallelogramma 4-36 patens 3-44 patula 4-36 paucinervata 4-45 pavoniana 3-23 pellucido-punctata 4-44 permollis 3-70 peruviana 3-23 phacelothrix 3-22 pilosohispida 3-27 pilosula 3-19 platyloba 4-15 poiteana 3-65 prasina 4-45 protensa 4-2 1 ptarmiciformis 3-17 pteroidea 3-25 pubescens 4-35 pulverulenta 4- 1 7 p«s/7/a 3-17 pyramidata 3-58 quadrangularis 3-41 refulgens 4-7 resinosofoetida 3-34 retrorsa 3-27 rimbachii 3-19 rivulorum 3-32 n&//.s 3-26 rw/a 3-20 ruiziana 3-35 saffordii 4-37 salzmannii 3-73 scalaris 3-16 sellensis 3-3 1 semihastata 3-57 serrata 3-71 simplicifrons 3-68 sprengelii 3-31 stuebelii 3-22 subandina 3-20 subincisa 4- 1 6 submarginalis 4-8 supina 3-26 supralineata 3-52 tarapotensis 4- 1 5 tetragona 3-64 thelypteris 3-6 thomsonii 3-22 tremula 3-9 m'sto 3-6 1 uliginosa 3-3 valdepilosa 3-50 vasta 4- 1 7 wallichiana 4-36 warmingii 3-56 yungensis 4-14 Elaphoglossum 4-111 albescens 4-121 alipes 4- 1 22 amazonicum 4-122 amphioxys 4-123 amplum 4-123 angustius 4- 1 24 apodum 4-123 atropunctatum 4- 1 24 atrosquamatum 4- 1 24 auricomum 4-125 bakeri 4-125 bangii 4-167 barbatum 4-125 blepharoglottis 4- 1 26 calaguala 4-137 cam pt oil-pis 4- 1 26 cardenasii 4-126 castaneum 4-127 110 FIELDIANA: BOTANY caudatum 4-154 chloodes4-127 ciliatum 4- 1 28 concinnum 4-128 conforme 4-111 craspedotum 4-128 crinipes 4-160 cuspidatum 4-129 decoratum 4- 1 29 denticulatum 4-131 dichroum4-129 discolor 4- 1 30 diversifrons 4- 1 30 eatonianum 4-127 elegantipes 4-131 elongatum 4-162 engelii 4-131 ensiforme 4-131 erinaceum 4-132 erythrolepis 4- 1 32 eximium 4-133 flaccidum 4-133 fortipes 4-133 glabellum 4-134 glossophyllum 4-134 glutinosum 4-154 gracillimum 4- 1 34 guamanianum 4-135 hartwegii 4-135 hayesii 4- 1 36 haynaldii 4-136 hickenii 4-136 hieracioides 4-137 "hikenii" 4-137 horridulum 4-137 huacsaro 4-137 hystrix 4-138 jucundum 4-138 killipii 4-13S laminarioides 4-139 Ianatum4-139 Iasioglottis4-139 latevagans 4- 1 40 latifolium 4-140 latum 4-123 lawyerae 4-141 laxisquama 4-141 lechlerianum 4-141 leprosum 4-142 leptophyllum 4-146 lindbergii 4- 1 26 lindenii 4-142 lingua 4-142 linguaeforme 4- 1 34 litanum 4-143 lloense 4- 1 32 longipes 4- 1 44 longius 4-143 luridum 4-144 macilentum 4- 1 44 mathewsii 4- 1 44 megalurum 4-145 megarhizon 4-146 meladenium 4-145 melancholicum 4-145 metallicum 4-146 minutum 4-146 molle4-152 moorei 4-167 moyeri 4-147 muscosum 4-147 nastukiae 4- 1 48 nidiformis4-148 nigriscens 4-148 nigrocostatum 4- 1 40 nivosum 4-148 obovatum 4-149 obtusum 4- 1 49 oculatum 4-150 odontolepis 4- 1 50 orbignyanum 4- 1 50 ornatum 4-149 oxyglossum 4-151 odphyllum 4- 1 50 pachyphyllum 4-151 pachyrrhizum 4-152 paleaceum 4-152 palorense 4-153 papillosum 4-153 pascoense 4-153 patinii 4-153 pattersoniae 4-166 peltatum 4-169 petiolosum 4-154 pichinchae 4-158 piloselloides 4-154 pilosius 4-155 plicatum 4-152 plumosum 4-155 poeppigianum 4-15 potomogeton 4-166 preslianum 4-128 propinquum 4-156 pseudohirtum 4-158 pumilio 4-156 punae 4-157 qui tense 4-157 raywaense 4-157 rimbachii 4-158 rosenstockii 4-158 rubellum 4-159 ruficomus 4-159 rufum4-159 russelliae 4- 1 60 schomburgkii 4-144 setigerum 4-160 siliquoides 4-136 simulans 4- 1 60 spatulatum 4-155 squamipes 4-161 stenophyllum 4-161 sty riacum 4-161 subciliatum 4-143 tambillense 4-162 tectum 4-162 tenue 4-163 tenuiculum 4-163 to men tell um 4-163 velongum 4- 1 64 vittarioides 4- 1 64 vulcanicum 4-164 wardiae 4-165 williamsiorum 4-165 williamsii 4-123 zebrinum 4-165 Enterosora 5-72 campbellii 5-72 parietina 5-84 trichosora 5-84 trifurcata 5-83 Equisetaceae 6- 1 2 Equisetum 6-12 bogotense 6-16 fluviatile 6- 1 2 giganteum 6- 1 5 myriochaetum 6- 1 5 ramosissimum ssp. debile 6- 1 5 schaffneri 6-15 x schaffneri 6- 1 5 variegatum 6- 1 5 Eriosorus 2-3 accrescens 2-6 aureonitens 2-6 cheilanthoides 2-6 elongatus 2-8 flabellatus 2-8 flexuosus 2-7 lechleri 2-8 orbignyanus 2-7 rufescens 2-4 ruizianus 2-20 scandens 2-7 stuebelii 2-6 wurdackii 2-7 Eschatogramme 5-145 desvauxii 5-147 furcata 5-147 polypodioides 5-148 subnuda 5-147 Eupodium 1-15 kaulfussii 1-15 Feea diversifrons 1-91 heterophylla 1-94 humboldtii 1-94 trollii 1-91 Glaphyropteris 3-48 decussata 3-49 Gleichenia 1-37 affinis 1-42 bancroftii 1-39 bifida 1-41 boliviensis 1-43 buchtienii 1-42 costaricensis 1-43 flexuosa 1-47 glauca 1-37 hypoleuca 1-43 lechleri 1-44 leucocarpa 1-44 longipinnata 1-45 mathewsii 1-41 mellifera 1-4 TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. Ill nervosa 1-47 nitidula 1-43 pectinata 1-49 pennigera 1-39 peruviana 1-45 polypodioides 1-37 pruinosa 1-42 remota 1-44 revoluta 1-42 rigida 1-47 rubiginosa 1-46 simplex 1-39 subandina 1-43 tomentosa 1-41 truncata 1-37 tuberculata 1-43 velata 1-42 yungensis 1-44 Glyphotaenium 5-72 trifurcatum 5-83 Goniophlebium 5-71 incanum 5-125 pectinatum 5-121 semipinnatum 5-140 subauriculatwn 5-72 Goniopteris 3-52 biolleyi 3-62 crenata 3-52 eggersii 3-59 juruensis 3-6 1 pennata 3-64 poiteana 3-65 pyramidata 3-58 tetragona 3-64 m'sto 3-6 1 Grammitis 5-72 albidula 5-87 alsopteris 5-110 amylacea 5-87 andicola 5-103 and/mz 5-109 anfractuosa 5-99 apiculata 5-94 aromatica 5-100 asplenifolia 5-104 assurgens 5-99 athyrioides 5-113 basalis 5-109 bipinnata 5-92 bishopii 5-86 blepharidea 5-112 blepharolepis 5-108 bryophila 5-81 Z>wes/7 5-112 campbellii 5-72 capillaris 5-90 cheilanthoides 2-6 chrysleri 5-104 congesta 5-89 crispata 5-72 cultrata 5-108 curvata 5-72 daguensis 5-109 david-smithii 5-109 dendroxa 5-88 dependens 5-104 discolor 5-86 dudleyi 5-92 elongata 5-143 erecta 5-98 farinosa 5-88 firma 5-100 flabelliformis 5-102 flexuosa 2-7 gracilis 5-96 graminioides 5-72 herrerae 5-88 heteromorpha 5-105 immixta 5-115 jamesonii 5-82 jamesonioides 5-93 lanceolata 5-143 lanigera 5-105 laxa 5-107 lehmanniana 5-104 leucosticta 5-111 limbata 5-8 1 linkiana 3-14 longipinnata 5-85 major 5-96 marginella 5-72 mathewsii 5-91 melanosticta 5-97 meridensis 5-85 mirabilis 5-91 moniliformis 5-98 myosuroides 5-82 myriophylla 5-114 nigrolimbata 5-81 obliquata 5-72 paramicola 5-8 1 parietina 5-84 peruviana 5-102 phlegmaria 5-97 pichinchae 5-110 pichinchensis 5-110 pilipes 5-90 pilosissima 5-100 praeclara 5-89 pseudocapillaris 5-94 pseudonutans 5-103 pumila 5-82 recondita 5-95 revoluta 5-143 rigens 5-101 rigescens 5-102 ruiziana 2-7 sectifrons 5-72 semihirsuta 5-111 senilis5-107 sericeo-lanata 5-106 serrulata 5-82 sprucei 5-84 squamulosa 5-143 subflabelliformis 5-106 subsessilis 5-95 taxifolia 5-113 terrestris 5-87 trichosora 5-83 trifurcata 5-83 truncicola 5-109 tunguraguae 5-94 variabilis 5-115 venulosa 5-72 werfii 5-92 xiphopteroides 5-101 youngii 5-97 Gymnogramma aureonitens 2-7 diplazioides 3-14 elongata 2-8 ferruginea 2-20 flabellata 2-8 flexuosa 2-7 goudotii2-\2 jamesonii 2-20 lechleri 2-8 mathewsii 2-4 mohriaeformis 2-4 ochracea 2- 1 9 orbignyana 2-1 pearcei 2-2 1 peruviana 2- 1 9 polypodioides 3-14 pumila 2-84 reniformis 2- 1 5 rufescens 2-4 stuebelii 2-6 Gymnopteris 2-46 aliena 4-100 nicotianifolia 4-102 pandurifolia 4-105 rufa 2-47 tomentosa 2-46 Hecistopteris 2-84 pumila 2-84 Hemidictyum 4-90 marginatum 4-90 Hemionitis 2-46 brasiliana 2-87 cajenensis 2-87 lanceolata 2-87 lineata 2-86 palmata 2-48 pinnata 2-48 plantaginea 2-84 rufa 2-47 tomentosa 2-46 Hemiphlebium kapplerianum 1-88 Hemitelia 1-129 andina 1-130 horrida 1-138 lechleri 1-131 multiflora 1-130 nervosa 1-139 petiolata 1-131 rufescens 1-1 14 speciosa 1-139 subincisa 1-139 uleana 1-138 Hicriopteris bancrofti 1-39 Histiopteris 2-115 incisa 2-115 vespertilionis 2-115 112 FIELDIANA: BOTANY Hoffmannia 6- 1 1 aphylla 6- 1 1 Holodictyum 5-2 Homoeotes 1-76 heterophylla 1-94 Huperzia 6-19 acerosa 6-46 acifolia 6-28 affinis 6-30 andina 6-35 aqualupiana 6-48 arcuata 6-25 aristei 6-43 attenuata 6-37 bifida 6-28 blepharodes 6-30 binervia 6-26 brevifolia 6-39 brongniartii 6-27 buesii 6-42 campiana 6-47 capellae 6-32 caracasica 6-40 colanensis 6-3 1 crassa 6-33 cuneifolia 6-50 curvifolia 6-46 darwiniana 6-33 dichaeoides 6-48 dichotoma 6-26 durissima 6-51 ecuadorica 6-29 engleri 6-39 ericifolia 6-48 e versa 6-29 filiformis 6-46 funiformis 6-42 hartwegiana 6-40 heteroclita 6-48 hippuridea 6-24 hohenackeri 6-40 hypogaea 6-36 kuesteri 6-30 jenmanii 6-44 lechleri 6-25 lindaviana 6-44 linifolia 6-43 var. jenmanii 6-44 var. tenuifolia 6-44 loxensis 6-26 lucidula 6-19 macbridei 6-32 mandiocana 6-45 mexiae 6-28 mollicoma 6-45 molongensis 6-47 montana 6-25 myrsinites 6-49 nesselii 6-34 nuda 6-25 papillata 6-3 1 parvifolium 6-28 passerinoides 6-4 1 var. nitens 6-4 1 pearcei 6-30 phylicifolia 6-49 pilgeriana 6-34 polycarpos 6-37 polyclada 6-5 1 polylepidetorum 6-3 1 pruinosa 6-5 1 quadrifariata 6-49 reflexa 6-27 var. bifida 6-28 var. minor 6-28 var. reflexa 6-28 rosenstockiana 6-42 sagasteguiana 6-37 sanctae-barbarae 6-35 sarmentosa 6-45 saururus 6-35 selago 6-19 sellifolia 6-38 socratis 6-42 sotae 6-43 subulata 6-50 taxifolia 6-41 tenuis 6-46 tetragona 6-38 unguiculata 6-29 weberbaueri 6-30 weddellii 6-26 wilsonii 6-44 Hydroglossum 1-33 oligostachyum 1-33 Hymenodium kunzeanum 4-151 Hymenophyllum 1-50 adiantoides 1-66 amabile 1-70 andinum 1-62 apiculatum 1-59 axillare 1-62 beyrichianum 1-68 calodictyon 1-56 ciliatum 1-67 contortum 1-64 crispatulum 1-68 crispum 1-67 cristatum 1-55 dendritis 1-59 dependens 1-72 dicranotrichum 1-50 ectocarpon 1-57 elegans 1-66 elegantulum 1-70 endiviifolium 1-63 fendlerianum 1-64 ferax 1-61 fragile 1-69 fucoides 1-55 fusagasugense 1-73 fusugasugense 1-73 hirsutum 1-66 interruptwn 1-72 karstenianum 1-71 lamella turn 1-58 latipes 1-59 lindenii 1-71 lineare 1-66 lobatoalatum 1-74 mathewsii 1-60 mexiae 1-59 microcarpum 1-68 mirincum 1-58 molle 1-65 nuil tialat urn 1-73 multiflorum 1-63 myriocarpum 1-62 nigrescens 1-63 nigricans 1-63 pedicellatum 1-57 peltatum 1-57 peruvianum 1-75 platylobum 1-68 plumieri 1-72 plumosum 1-72 poeppigianum 1-76 polyanthos 1-59 polycarpum 1-64 procerum 1-65 pulchellum Hooker 1-70 pulchellum Mett. 1-65 pyramidatum 1-74 reniforme 1-64 rimbachii 1-64 ruizianum 1-69 rupestre 1-82 simplex 1-69 speciosum 1-70 spectabile 1-70 sprucei 1-76 superbum 1-72 tarapotense 1-75 tenerrimum 1-66 tomentosum 1-73 tortuosum 1-57 trapezoidale 1-69 trianae 1-61 trichomanoides 1-61 trichophyllum 1-65 trifidum 1-76 tunbridgense 1-50 undulatum 1-63 valvatum 1-68 verecundum 1-75 Hymenostachys diversifrons 1-91 Hypoderris stuebelii 4-103 Hypolepis 2-106 bogotensis 2-1 10 flexuosa 2-110 hostilis 2-110 nigrescens 2-109 obtusata 2-110 parallelogramma 2-110 pteroides 2-111 stuebelii 2-109 tenuifolia 2- 106 Isoetaceae 6-88 Isoetes 6-89 andicola 6-92 var. gemmifera 6-92 andina 6-97 boliviensis 6-96 TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 113 dichotoma 6-9 1 dispora 6-93 glacialis 6-95 herzogii 6-97 hewitsonii 6-94 karstenii 6-96 lacustris 6-89 laevis 6-95 lechleri 6-95 var. triquetra 6-97 novo-granadensis 6-91 pacifica 6-9 1 panamensis 6-9 1 parvula 6-92 peruviana 6-95 saracochensis 6-94 savanarum 6-9 1 soda 6-95 "ticlioensis" 6-95 triquetra 6-97 Jamesonia 2-8 alstonii 2-12 blepharum 2-14 boliviensis 2-12 cinnamomea 2-14 glutinosa 2-13 goudotii 2-12 imbricata 2- 1 3 paleacea 4-54 peruviana 2- 1 1 pulchra 2-11 rotundifolia 2-10 scalaris 2- 1 2 scammanae 2-1 1 Lacostea tanaica 1-89 Lastrea cheilanthoides 3-34 nitens 3-35 poeppigiana 3-57 poiteana 3-65 pubescens 4-3 1 recedens 4-3 1 ntt/w 3-26 scabriuscula 3-44 Lastreopsis 4-3 1 amplissima 4-33 effusa 4-33 exculta 4-35 killipii 4-33 recedens 4-3 1 tenera 4-3 1 Lecanium membranaceum 1-88 Lellingeria 5-72 apiculata 5-94 major 5-96 myosuroides 5-82 phlegmaria 5-97 pseudocapillaris 5-94 subsessilis 5-95 tunguraguae 5-95 Lepicystis 5-125 incana 5-125 Lepidotus auct. 6-58 Lepidotus Mirbel 6-52 alopecuroides 6-6 1 cernua 6-63 clavata 6-54 contexta 6-62 magellanica 6-56 Lepisorus 5-140 Leptochilus alienus 4-100 bradeorum 4-103 crenatus 4-101 guianensis 4-109 killipii 4-102 lindigi 4-103 nicotianifolius 4-102 oligarchicus 4- 1 03 pandurifolium 4-105 serratifolius 4-101 serratus 4-101 stuebelii 4-103 Leptocionium 1-50 dicranotrichum 1-50 fucoides 1-56 pedicellatum 1-57 Leptopteris 1-20 Lindsaea 2-115 arcuata 2-117 divaricata 2-118 guianensis 2-120 hemiglossa 2-122 lancea 2-121 latifrons 2-122 portoricensis 2-120 phassa 6-98 schomburgkii 2-122 spruceana 2-118 sprucei 2-118 stricta2-121 taeniata 2-118 tarapotensis 2-118 trapeziformis 2-115 ulei2-122 Litobrochia horizontalis 2-77 Lomagramma 4-109 guianensis 4-109 Lomaria 5-56 acwta 5-64 andina 5-61 angustifolia 5-64 arborescens 5-62 aurata 5-67 caudata 5-66 chilensis 5-62 cordata 5-62 cuspidata 5-64 diver gens 5-63 ensiformis 5-64 euphlebia 1-101 fragile 5-64 fraxinea 5-59 heterophylla 5-63 linariaefolia 5-68 loxensis 5-65 meridensis 5-64 «M<&Z 5-56 obtusifolia 5-67 ornifolia 5-62 pteropus 5-63 schomburgkii 5-66 serrulosa 5-62 squamulosa 5-65 stenophylla 5-65 stipitellata 5-65 volubilis 5-70 Lomaridium semicordatum 1-101 Lomariopsis 4-105 erythrodes 4-107 fendleri4-106 japurensis 4-106 latipinna 4-107 nigropaleata 4-108 sorbifolia 4-105 Lonchitis 2-113 hirsuta 2-113 lindeniana 2-113 ;?«tea 2-80 tenuifolia 2-106 Lophidium 1-33 elegans 1-34 flabellum 1-34 latifolium 1-33 poeppigianum 1-36 Lophosoria 1-107 pruinata 1-109 quadripinnata 1-107 Loxoma 1-98 Loxoscaphe 5-4 concinna 5-4 thecifera 5-48 Loxsoma 1-98 Loxsomopsis 1-99 costaricensis 1-99 lehmannii 1-99 notabilis 1-99 pearcei 1-99 Lycopodiaceae 6-16 Lycopodiella 6-58 sect. Campylostachys 6-60 sect. Caroliniana 6-58 sect. Lycopodiella 6-60 alopecuroides 6-6 1 camporum 6-63 caroliniana var. meridionalis 6-6 1 cernua 6-63 contexta 6-62 descendens 6-64 glaucescens 6-64 inundata 6-58 matthewsii 6-6 1 pendulina 6-65 riofrioi 6-65 Lycopodium 6-52 sect. Caroliniana 6-60 sect. Complanata 6-52 114 FIELDIANA: BOTANY sect. Diphasium 6-52 sect. Lycopodium 6-52 sect. Magellanica 6-52 subg. Cernuistachys 6-58 subg. Lepidotis auct. 6-58 subg. Lepidotis Baker 6-52 subg. Rhopalostachya 6-52 subg. Selago 6-19 subg. Urostachya 6- 1 9 acifolium 6-28 affine 6-30 albidum 6-55 alopecuroides 6-6 1 var. contextum 6-62 ssp. contextum 6-62 anceps 6-80 andinum 6-35 arcanum 6-44 aristatum 6-54 articulatum 6-84 attenuatum 6-37 bifidum 6-28 binervium 6-26 blepharodes 6-30 brevifolium 6-39 brongniartii 6-27 brutum 6-28 Z>Mes» 6-42 capellae 6-32 capillaceum 6-63 caracasicum 6-40 carolinianum 6-58 var. meridionalis 6-62 catharticum 6-38 cernuum 6-64 var. capillaceum 6-64 var. pendulinum 6-65 clavatum 6-54 ssp. clavatum 6-54 ssp. contiguum 6-55 var. aristatum 6-54 var. pseudocontiguum 6-55 var. thyoides 6-57 var. validum 6-57 congestifolium 6-49 contextum 6-62 contiguum 6-55 convolutum 6-87 crassum 6-33 cuatrecasasii 6-45 cuneifolium 6-50 curvifolium 6-46 densifolium 6-28 diffusum 6-76 durissimum 6-5 1 ecuadoricum 6-29 elongatum 6-35 engleri 6-39 ericaefolium 6-48 eriostachys 6-54 erythropus 6-80 eversum 6-29 ewanii 6-50 exaltatum 6-86 fastigiatum 6-56 funiforme 6-42 gayanum 6-57 geniculatum 6-85 glaucescens 6-64 gracile 6-80 haematodes 6-80 haenkei 6-56 hartwegianum 6-40 herbaceum 6-55 heteroclitum 6-48 heterophyllum 6-57 hohenackeri 6-40 hypogaea 6-19 hippurideum 6-24 jenmanii 6-44 jussiaei 6-56 var. microphyllum 6-56 lechleri 6-25 linifolium 6-43 lucidula 6-19 macbridei 6-32 magellanicum 6-56 mathewsii 6-6 1 meridionale 6-6 1 mexiae 6-28 microphyllum 6-75 molongense 6-47 myrsinites 6-49 var. minus 6-38 -4 1 nubigenum 6-49 nova-hollandiae 6-77 nudum 6-12 papillatum 6-31 parkeri 6-84 parvifolium 6-28 passerinoides 6-4 1 pendulinum 6-65 pensum 6-68 phlegmaria 6-19 phylicifolium 6-49 pichinchense 6-56 piliferum 6-54 polycarpos 6-45 polycarpum 6-29 polycladum 6-5 1 poseidonis 6-24 pruinosum 6-5 1 reflexum Lam. 6-27 var. densifolium 6-28 var. majus 6-28 var. minus 6-28 var. polycarpum 6-29 reflexum Willd. 6-29 reversum 6-28 riofrioi 6-65 rosenstockianum 6-42 sanctae-barbarae 6-2 1 sarmentosum 6-45 saururus 6-35 scariosum 6-55 vsa.jussieui 6-57 schwendeneri 6-41 selaginoides 6-66 selago 6- 1 9 skutchii 6-49 sprucei 6-62 spurium 6-56 stamineum 6-44 stellae-polaris 6-28 subulatum 6-50 taxifolium 6-41 var. brongniartii 6-27 tenu? 6-46 var. tenuissimum 6-46 tetragonum 6-38 var. patulum 6-38 thujoides 6-57 thyoides 6-57 trichodendron 6-44 trichophyllum 6-54 trychopyllum 6-54 unguiculata 6-29 vestitum 6-55 weddellii 6-26 wilsonii 6-44 Lygodium 1-30 digitatum 1-32 mexicanum 1-33 micans 1-32 oligostachyum 1-33 polymorphyum 1-33 radiatum 1-32 scandens 1-30 venustum 1-30 volubile 1-32 Macrothelypteris 3-3 torresiana 3-3 Marattia 1-13 a/ara 1-15 kaulfussii 1-15 laevis 1-15 Marginaria 5-125 angustifolia 5-169 polypodioides 5-125 Marginariopsis 5-7 1 Marsilea 6-2 ancylopoda 6-4 crotophora 6-5 deflexa 6-4 mollis 6-5 natans 6-6 mucronata 6-5 uncinata 6-5 vestita 6-4 ssp. tenuifolia 6-5 ssp. vestita 6-5 Marsileaceae 6-2 Mecodium 1-50 apiculatum 1-59 contortum 1-64 dendritis 1-59 endiviifolium 1-63 fendlerianum 1-64 /mzx 1-61 mexiae 1-59 microcarpum 1-68 multiflorum 1-63 myriocarpum 1-62 polyanthos 1-59 trichomanoides 1-61 TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 115 undulatum 1-64 Megalastrum 4- 1 1 andicola 4- 1 8 biseriale 4-15 hirsutosetosum 4-16 honestum 4-13 leptosorum 4-14 microsorum 4-14 mollicomum 4- 1 7 pansamalense 4- 1 8 platylobum 4-15 pulverulentum 4-17 spectabile 4- 1 6 subincisum 4-16 vastum 4-17 villosum 4-1 1 yungense 4-14 Melpomene 5-96 Meniscium 3-66 andreanum 3-76 angustifolium 3-74 arborescens 3-70 cristatum 4-80 falcatum 3-75 giganteum 3-68 guyanense 3-68 jurgensenii 3-75 macrophyllum 3-68 opacum 4-45 salzmannii 3-73 serratum 3-71 Meringium fucoides 1-56 Mertensia 1-46 bancroftii 1-39 Z>(/zrfa 1-41 dichotoma 1-46 flexuosa 1-47 laevigata 1-37 longipinnata 1-45 mathewsii 1-41 nervosa 1-47 pectinata 1-49 pennigera 1-39 pruinosa 1-42 remota 1-44 revoluta 1-42 rigufa 1-47 simplex 1-37 tomentosa 1-41 ve/afa 1-42 Metaxya 1-109 rostrata 1-111 Microgramma 5-148 acatallela 5-152 acuminata 5-157 baldwinii 5-155 chrysolepis 5-151 ciVuzfcz 5-153 fuscopunctata 5-144 geminata 5-157 latevagans 5-151 Iindbergii5-157 lycopodioides 5- 1 54 megalophylla 5-157 percussa 5-144 persicariifolia 5-157 piloselloides 5-152 recreense 5-157 reptans 5-163 rosmarinifolia 5-154 squamulosa 5-156 tecta5-154 thurnii 5-156 ulei 5-156 vacciniifolia 5-152 Microlepia 2-95 flaccida 2-95 inaequalis 2-103 polypodioides 2-95 speluncae 2-95 Micropolypodium 5-72 pseudotrichomanoides 5-72 Microstaphyla bangii 4-167 moorei 4-167 Mildella 2-23 intramarginalis 2-23 Mohria 1-23 Nephelea 1-118 cuspidata 1-120 erinacea 1-118 incana 1-120 polystichoides 1-118 Nephrodium antioquoianum 4-26 brachypus 3-18 canadasii 3-21 "carazanense" 3-27 caucaense 3-28 conspersum 3-42 corazonense 3-27 crassipes 3-3 1 deflexum 3-30 eggersii 3-59 firmifolium 4-47 funestum 4-21 gardnerianum 3-52 jamesonii 3-56 kunzeanum 3-46 lagerheimii 4-8 fecA/m 3-69 leprieurii 3-51 lizarzaburui 4-25 longipilosum 3-14 macradenium 3-21 macrotis 3-57 microsorum 4-14 nemorale 3-62 nigrovenium 4-9 ochropteroides 4-38 pilosohispidum 3-27 polyphyllum 4-53 quadrangulare 3-4 1 resinosofoetidum 3-34 retrorsum 3-27 schizotis 3-44 sodiroi 4-26 supinum 3-26 tarapotense 4-8 trapezoides 4-53 valdepilosum 3-50 villosum 4- 1 8 Nephrolepis 5-49 biserrata 5-51 cordifolia 5-52 hirsutula 5-54 multiflora 5-52 occidental is 5-53 pectinata 5-53 pendula 5-52 rivularis 5-52 Neuromanes pinnatum 1-92 Neurophyllum hostmannianum 1-93 pinnatum 1-92 Niphidium 5-173 albopunctatissimum 5-174 americanum 5-173 anocarpos 5-177 carinatum 5-176 crassifolium 5-174 longifolium 5-173 macbridei 5-176 vittaria 5-177 Notholaena 2-37 arequipensis 2-33 awm* 2-28 bonariensis 2-28 brackenridgei 2-32 buchtienii 2-34 Candida 2-37 cantangensis 2-32 chrysophylla 2-40 /ra«?ri 2-28 lonchophylla 2-33 marantae 2-37 mo//w 2-29 nivea 2-38 obducta 2-29 peruviana 2-32 sinuata 2-34 stuebeliana 2-33 sulphurea 2-37 tectaria 2-34 tenera 2-40 tomentosa 2-34 trichomanoides 2-37 Odontomanes hostmannianum 1-93 Oleandra 4-96 articulata 4-96 distenta 4-96 hirta 4-98 lehmannii 4-97 micans 4-98 neriiformis 4-96 nodosa 4-96 pilosa 4-98 Olfersia 4-55 caudata 4-64 cervina 4-57 116 FIELDIANA: BOTANY ciliata 4-128 corcovadensis 4-57 Onoclea polypodioides 1-37 Ophioglossum 1-8 coriaceum 1-12 crotalophoroides 1-12 ellipticum 1-12 lusitanicum 1-12 nudicaule 1-11 opacum 1-12 palmatum 1-9 pendulum 1-8 peruvianum 1-9 petiolatum 1-9 reticulatum 1-9 scandens 1-30 scariosum 1-11 tenerum 1-11 vulgatum 1-8 ypanemense 1-11 Osmunda 1-21 adiantifolia 1-24 cervina 4-57 cicutaria 1-8 cinnamomea 1-21 flexuosa 1-27 hirsuta 1-28 humilis 1-29 lunaria 1-6 oblongifolia 1-29 palustris 1-23 phyllitidis 1-29 polypodioides 5-64 regalis 1-23 spectabilis 1-23 virginiana 1-8 Paesia2-106 amazonica 2-106 anfractuosa 2-106 glandulosa 2-106 viscosa 2-106 Palhinhaea 6-58 camporum 6-63 cernua 6-63 descendens 6-64 glaucescens 6-64 pendulina 6-65 riofrioi 6-66 Parablechnum ciliatum 5-68 Paraceterach marantae 2-37 Parkeria pteridoides 2-50 Pecluma 5-116 absidata 5-119 boliviano 5-120 camptophyllaria 5-122 choquetangensis 5-116 curvans 5-119 dispersa 5-125 divaricata 5-120 eury basis 5-121 filicula 5-119 funicula 5-116 hygrometrica 5-124 pectinata 5-121 plumula 5-118 ptilodon 5-123 venturii 5-122 Pellaea 2-40 atropurpurea 2-40 cordifolia 2-4 1 crenulans 2-44 dealbata 2-38 lorentzii 2-44 «/vea 2-38 ovata 2-4 peruviana 2-41 sagittata 2-41 tenera 2-40 ternifolia 2-41 weddeliana 2-4 1 wrightiana 2-41 Peltapteris4-167 moorei 4-167 peltata4-169 peruviana 4- 1 70 Peltochlaena 4-47 Phanerophlebia 4-38 nobilis 4-38 Phegopteris cochleata 4-55 dictyophylla 4-38 dwZ>/a 4-38 laevigata 3-24 lechleri 4-45 membranacea 3-69 mo/fo 3-70 pycnolepis 4-55 refulgens 4-7 Phlebodium 5-125 aureum 5-134 decumanum 5-135 Phlegmariurus 6- 1 9 taxifolius 6-4 1 Phyllitis 5-4 scolopendrium 5-4 Phylloglossum 6-16 Pilularia 6-2 americana 6-2 mandonii 6-2 Pityrogramma 2-16 austroamericana 2- 1 8 chrysoconia 2-20 chrysophylla 2-18 ebenea 2-18 ferruginea 2-20 ochracea 2- 1 9 pearcei 2-21 perelegans 2- 1 9 peruviana 2-19 presliana 2-20 tartarea 2- 1 9 trifoliata 2-21 Plagiogyria 1-101 costaricensis 1-101 denticulata 1-101 euphlebia 1-101 latifolia 1-101 semicordata 1-101 Plananthus 6-19 reflexus 6-27 selago 6- 1 9 Platycerium 5-181 alcorne 5-181 andinum 5-181 Plecosorus 4-49 mexicanus 4-49 peruvianus 2-111 speciosissimus 4-49 Pleopeltis5-140 angusta 5-140 astrolepis 5-143 fuscopunctata 5-144 lanceolata 5-143 macrocarpa 5-142 percussa 5-144 pinnatifida 5-135 revoluta 5-143 squamulosa 5-143 Pleuridium albopunctatissimum 5-174 Pleurosorus 5-2 Poecilopteris crenata 4-101 Polybotrya 4-57 aequatoriana 4-64 alfredii 4-65 altescandens 4-6 1 andina 4-6 1 appressa 4-64 caudata 4-64 cervina 4-57 crassirhizoma 4-60 decorata 4-62 fractiserialis 4-60 fulvastrigosa 4-65 glandulosa 4-63 hickeyi 4-65 juglandifolia 4-6 1 kalbreyeri 4-6 1 lechleriana 4-63 lomarioides 4-65 macbridei 4-60 nutans 4-65 osmundacea 4-62 plumbicaulis 4-60 polybotryoides 4-6 1 pubens 4-62 puberulenta 4-63 serratifolia 4-57 sorbifolia 4-57 subelliptica 4-63 suberecta 4-6 1 Polypodium 5-125 abitaguae 5-86 abruptum 3-58 absidatum 5-119 acrodontium 5-101 acrosorum 5-177 adiantiforme 4-3 1 adnatum 5-133 aglaolepis 5-168 TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 117 albopunctatissimum 5-174 alternifolium 5-108 americanum 5-173 amphostenon 5-166 amplum 4-10 andinum 5-109 anfractuosum 5-99 angustifolium 5-168 angustipaleatum 5-169 anocarpos 5-177 apiculatum 5-94 oppression 5-132 arboreum 1-129 aristatum 4-35 aromaticum 5-100 articulatum 5-133 asplenifolium 5-104 asplundii 5-167 astrolepis 5-143 athyrioides 5-113 aureum 5-134 azuyense 5-94 balaoense 5-138 fez/Mrii 3-31 biauriculatum 5-131 binervatum 5-63 biseriale 4- 1 5 blepharideum 5-112 blepharolepis 5-108 bolivianum 5-120 bombycinum 5-138 brachyodus 3-49 brevifolium 5-170 bryophilum 5-81 bryopodum 5-135 buchtienii 5-136 l>ues/7 5-112 caceresii 5-133 caespitosum 5-162 camptophyllarium 5-123 capillare 5-90 chacapoyense 5-131 chartaceum 5-129 chnoodes 5-129 chrysolepis 5-151 ciliatum 5-153 circinatum 5-119 coar datum 5-164 cochleatum 4-54 concinnum 3-29 cordatum 5-131 cordifolium 5-52 crassifolium 5-174 crenatodentatum 4-54 crenatum 3-65 crispatum 5-167 crossii 3-23 crystalloneuron 5-139 cultratum 5-108 curvans 5-1 19 curvatum 5-72 dasypleuron 5-131 decumanum 5-135 decurrens 5-132 decussatum 3-49 dentatum 3-41 denticulatum 4-37 dependens 5-104 dichotomum 1-46 discolor 5-86 dispersum 5-125 disectum 2-100 dissimile 5-129 divaricatum 5-120 dolor ense 5-104 5-83 ecostatum 5-85 ecuadorense 5-1 10 effusum 4-33 euchlorum 3-25 eurybasis 5-121 exaltatum 5-49 falcatum 4-40 farinosum 5-89 fasciale 5-163 fendleri 5-132 fibrillosum 4-13 fUicula5-H9 filix-femina 4-88 fiiix-mas 4-35 firmum 5-100 flabelliforme 5-102 flavopunctatum 4-42 fragile 4-92 fraseri 5-136 fraxinifolium 5-132 fulvescens 2-111 furfuraceum 5-136 fuscopunctatum 5-144 giganteum 5-133 gilliesii 5-130 glaucophyllum 5-134 glaucum 1-109 globuliferum 2-99 gracile 5-96 gracillimum 5-108 guianense 4-109 haynaldii 4-26 herzogii 5-100 heteromorphum 5-105 heterophlebium 4-45 hirsutulum 5-54 honestum 4-13 horridum 1-138 x huancayanum 5-139 hygrometricum 5-124 incanum 5-137 jamesonii 5-82 jamesonioides 5-93 karstenianum 4- 1 7 kunzeanum 5-131 lachniferum 5-123 /aftfwm 5-130 laevigatum 5-134 lanceolatum 5-142 lanigerum 5-106 lapathifolium 5-162 lasiopus 5-130 latevagans 5-151 5-129 5-170 5-107 leptophyllum 2-23 leucatomos 5-134 x leucosporum 5-139 leucosticton Fee 5-111 leucosticton Klotzsch 5-136 levigatum 5-134 limbatum 5-81 lomariiforme 5-125 lonchitis 4-49 longicaudatum 4-44 longifolium 5-132 longisetosum 5-114 longiusculum 5-111 longum 5-108 lor et ense 5-156 loriceum 5-129 lycopodioides 5-154 macrocarpum Presl 5-135 macrocarpum Willd. 5-143 macrophyllum 4-44 marginellum 5-72 mathewsii 5-91 medullare 1-112 megalodus 3-64 megalolepis 5-137 megalophyllum 5-157 melanostictum 5-97 meridense 5-85 microdontum 1-127 mo//e 3-41 mollendense 5-135 moniliforme 5-98 monosorum 5-138 montevidense 4-5 1 monticola 5-99 muricatum 4-55 murorum 5-139 myosuroides 5-82 myriophyllum 5-114 nigrolimbatum 5-81 /j/tefu 3-35 nitidissimum 5-172 nodosum 5-172 obliquatum 5-72 occultum 5-161 oligocarpum 3-15 oligophlebium 4-45 ophiocaulon 5-162 opposition 3-32 parietinum 5-84 patens 3-44 pavonianum 3-23 pearcei 5-87 pectinatum 5-121 pedicellata 1-15 pendulum 5-95 penna-marina 5-62 pennatum 3-64 percussum 5-144 persicariifolium 5-157 peruvianum 5-102 phlegmaria 5-97 118 FIELDIANA: BOTANY phyllitidis 5-169 pichinchae 5-1 10 pichinchense 5-110 pilipes 5-90 piloselloides 5-152 pilosissimum 5-100 plantagineum 4-27 platylobum 4- 1 5 plumula 5-118 polypodioides 5-137 pozuzoense 5-90 prasinum 4-45 preslianum 5-132 procerum 1-123 pruinatum 1-109 pseudoaureum 5-134 pseudocapillare 5-94 pseudonutans 5-103 pteroideum 3-25 pteropus 5-95 ptilodon 5-124 pubescens Hooker & Grev. 5- 1 30 pubescens L. 4-35 pulverulentum 4-17 punctatum 4-45 pungens 1-123 pycnocarpum 5-135 pycnolepis 4-55 quadripinnatum 1-109 ratibori 5-135 remotum 5-136 repens 5-162 reticulatum 3-66 richardii 5-133 rigens 5-101 rigescens 5-102 rigidum 4-53 rivulare 5-52 rivulorum 3-32 rosmarinifolium 5-154 rostratwn 1-111 n«fe 3-26 rufum 3-20 ruiz-lealii 5-135 ruizianum 3-35 rusbyi 5-135 salicifolium 3-74 saxatile 4-54 sectifrons 5-72 semicordatum 4-29 semihirsutum 5-111 semipinnatifidum 5-140 .wrote 5- 107 sericeo-lanatum 5-106 serpentinum 5-163 serrulatum 5-83 sessilifolium 5-131 sloanei 4-10 solutum 5-172 sororium 5-129 speluncae 2-95 sphenodes 5-164 spixianum 5-86 sprucei 5-84 squamulosum 5-156 subandinum 5-130 subauriculatwn 5-71 subflabelliforme 5-106 subincisum 4- 1 6 submarginale 4-8 subscabrum 5-1 10 subsessile 5-95 subvestitwn 5-135 surucuchense 5-131 taeniosum 5-168 tarapotense 4- 1 5 taxifolium 5-113 tectum 5-154 tenuiculum 5-96 tetragonum 3-64 thomsonii 3-22 thurnii 5-156 thyssanolepis 5-137 tottum 3-40 trichiatum 5-161 trichosorum 5-84 trifoliatum 4-21 trifurcatum 5-83 triseriale 5-132 m'ste 3-60 truncicola 5-109 tunguraguae 5-95 tweedianum 5-135 M/e/5-156 vacciniifolium 5-152 variabile 5-115 vastum 4- 1 7 venturii 5-122 venulosum 5-72 villosum 4-11 vittaria 5-177 vulgare 5-125 vulpinum 5-167 xantholepis 5-135 xiphopteroides 5-101 yungense 5-113 Polystichopsis 4-35 ochropteroides 4-38 Polystichum 4-49 amplissimum 4-33 boboense 4-54 bonapartii 4-38 cochleatum 4-54 dubium 4-38 gelidum 4-54 haenkeanum 4-54 lehmannii 4-51 lonchitis 4-49 mexiae 4-5>l montevidense 4-5 1 moritzianum 4-55 muricatum 4-55 nudicaule 4-52 orbiculare 4-53 orbiculatum 4-53 paleaceum 4-54 platyphyllum 4-52 polyphyllum 4-54 pycnolepis 4-54 sodiroi 4-54 speciosissimum 4-49 torresianum 3-3 trapezoides 4-54 VW//H 4-52 yungense 4-52 Polytaeniwn 2-84 brasilianum 2-87 cajenense 2-87 guayanense 2-87 lanceolatum 2-86 Pronephrium 3-66 Pseudolycopodiella 6-58 contexta 6-62 meridionalis 6-62 Psilogramme 2-8 Psilotaceae 6- 1 1 Psilotum 6-11 complanatum 6-12 nudum 6-12 Pteridanetium 2-92 citrifolium 2-92 Pteridium 2-105 aquilinum 2-105 arachnoideum 2-105 caudatum 2-105 Pteris 2-70 altissima 2-76 amazonica 2-106 amp/a 2-78 angustifolia 2-90 aquilina 2-105 arachnoidea 2-105 atropurpurea 2-40 awrea 2-28 bakeri 2-80 biaurita 2-77 concolor 2-3 1 consanguinea 2-77 coriacea 2-74 cretica 2-81 decomposita 2-80 decurrens 2-77 deflexa 2-74 edentula 2-74 farinosa 2-30 fraseri 2-78 fur cat a 5-145 grandifolia 2-79 haenkeana 2-78 horizontalis 2-77 imbricata 2-13 incisa 2-115 interrupta 3-40 intramarginalis 2-23 faV/i/Hi 2-78 kunzeana 2-76 lechleri 2-78 //'nea/a 2-90 livida 2-80 lonchitoides 2-113 longifolia 2-70 /wa'fito 2-69 muricata 2-74 «/v«2 2-38 notholaenoides 2-25 orbiculata 2- 1 3 ovate 2-43 palmata 2-44 TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 119 pedata 2-80 peruviana 2-41 petiolulata 2-79 podophylla 2-80 polita 2-74 propinqua 2-75 pungens 2-75 quadriaurita 2-75 reticulata 2-76 reticulatovenosa 2-76 rigida 2-34 ruffa 2-47 sagittata 2-41 speciosa 2-76 splendens 2-79 sulphured 2-37 ternifolia 2-41 transparens 2-76 trichomanoides 2-37 tripartita 2-8 1 vespertilionis 2-115 vert tftz 2-78 vittata 2-8 1 Pteropsis vittarioides 2-9 1 Pterozonium 2-14 brevifrons 2-15 paraphysatum 6-96 reniforme 2-15 Ptilophyllum bicorne 1-91 hostmannianum 1-93 lambertianum 1-95 martiusii 1-95 pellucens 1-96 Ragatelus crinitus 1-94 Regnellidium 6-2 Rhipidopteris 4-167 flabellata 4-\69 peltata 4-169 rusbyi 4-167 sphenophylla 4-169 Rumohra 4-3 1 adiantiformis 4-3 1 aspidioides 4-31 berteriana 4-3 1 Saccoloma 2-101 elegans 2-103 inaequale 2-103 wercklei 2-101 Saffordia 2-48 /rtrfwta 2-50 Salpichlaena 5-68 hookeriana 5-70 lomarioidea 5-70 volubilis 5-70 Salvinia 6-6 auriculata 6-8 minima 6-6 natans 6-6 rotundifolia 6-8 Salviniaceae 6-5 Schaffheria 5-2 Schizaea 1-33 dichotoma 1-33 digitata 1-33 elegans 1-34 nstulosa 1-34 flabellum 1-34 incurvata 1-36 pennula 1-36 poeppigiana 1-36 pusilla 1-34 Selaginella 6-66 acanthostachys 6-87 amazonica (Milde) Hieron. 6-7 1 amazonica Spring 6-7 1 anceps 6-79 applanata 6-73 arizonica 6-72 articulata 6-84 asperula 6-85 atirrensis 6-76 bombycina 6-78 brachylepis 6-84 brevifolia 6-72 calcarata 6-86 calosticha 6-74 chionoloma 6-77 chrysoleuca 6-78 conduplicata 6-86 convolute 6-87 cordifolia 6-73 demissa 6-72 diffusa 6-76 dimorpha 6-82 eggersii 6-76 elongata 6-85 erythropus 6-80 exaltata 6-86 ferruminata 6-85 filicina 6-80 flagellata 6-77 fragilis 6-84 geniculata 6-85 gracilis Moore 6-80 gracilis (Poiret) Hieron. 6-80 haematodes 6-80 haenkeana 6-82 horizontalis 6-87 huberi 6-79 intacta 6-76 kunzeana 6-82 lechleri 6-79 lingulata 7-76 microphylla 6-75 mildei 6-7 1 nodosa 6-85 nova-hollandiae 6-77 parkeri 6-84 pearcei 6-77 pedata 6-84 peruviana 6-7 1 var. dombeyana 6-71 poeppigiana 6-83 var. peruviana 6-83 praestans 6-8 1 producta 6-74 quadrifaria 6-81 ramosissima 6-75 regularis 6-77 revoluta 6-72 rupestris 6-66 f. amazonica 6-71 f. peruviana 6-7 1 sartorii 6-71 seemannii 6-74 selaginoides 6-66 sellowii 6-71 sheldonii 6-7 1 silvestris 6-82 speciosa 6-79 spinosa 6-66 sprucei A. Br. 6-8 1 sprucei Hooker 6-78 stellata 6-86 strobolifera 6-86 sulcata ssp. poeppigiana 6-83 ssp. suavis 6-87 tomentosa 6-85 trisulcata 6-83 truncata 6-73 wolfii 6-87 weberbaueri 6-73 xiphophylla 6-77 Selaginellaceae 6-66 Selaginelleae 6-66 Selenodesmium rigidum 1-85 Sitobolium 2-95 punctilobulum 2-95 Solanopteris 5-179 bifrons5-181 bismarckii 5-180 brunei 5-180 tuberosa 5-180 Soromanes 4-57 serratifolia 4-57 Sphaerocionium 1-50 adiantoides 1-66 ciliatum 1-67 crispum 1-67 elegans 1-66 elegantulum 1-70 fragile 1-69 hirsutum 1-67 interruptum 1-72 karstenianum 1-71 lindenii 1-71 lobatoalatum 1-74 microcarpum 1-68 /no//e 1-65 multialatum 1-73 nigricans 1-63 plumieri 1-72 plumosum 1-72 pyramidatum 1-74 ruizianum 1-69 simplex 1-69 spectabile 1-71 tenerrimum 1-66 tomentosum 1-73 120 FIELDIANA: BOTANY trichophyllum 1-65 valvatum 1-68 Sphaeropteris 1-112 u talma 11 pa 1-115 aterrima 1-1 14 bradei 1-115 elongata 1-115 hirsuta 1-114 horrida 1-1 12 macrosora 1-1 14 medullaris 1-112 quindiuensis 1-116 rufescens 1-114 Steiropteris 3-46 gardneriana 3-52 incana 3-52 valdepilosa 3-50 Stenochlaena angusta 4-106 fendleri 4-106 japurensis 4-107 vesfita 4-106 Sticherus 1-37 o#z/ii5 1-42 6//K/WS 1-41 buchtienii 1-42 laevigatus 1-37 lechleri 1-44 longipinnatus 1-45 mathewsii 1-41 nitidulus 1-43 penniger 1-39 pruinosus 1-42 revolutus 1-42 rubiginosus 1-46 simplex 1-39 tuberculatus 1-43 velatus 1-42 yungensis 1-44 Stigmatopteris 4-42 alloeoptera 4-45 ecuadorensis 4-45 guianense 4-48 heterophlebia 4-45 ichtiosma 4-44 lechleri 4-45 longicaudata 4-44 meniscioides 4-48 opaca 4-45 paludosa 4-48 pellucidopunctata 4-44 prasina 4-45 rotundata 4-42 Struthiopteris maxonii 5-61 Sr>>///es 6-89 andicola 6-92 gemmifera 6-92 Syngramma brevifrons 2- 1 5 paraphysata 6-97 Taenitis desvauxii 5-147 furcata 5-148 Tectaria 4-2 1 andina 4-25 antioquiana 4-26 brauniana 4-24 decurrens 4-29 draconoptera 4-26 fraxinea 5-51 haynaldii 4-26 heracleifolia 4-27 incisa 4-24 kunzei 4-29 lizarzaburui 4-25 martinicensis 4-24 plantaginea 4-27 poeppigii 4-29 sodiroi 4-26 transiens 4-25 trifoliata 4-27 vivipara 4-25 Thelypteris 3-5 abrupta 3-58 aequatorialis 3-45 ancyriothrix 3-63 andicola 3-16 andreana 3-76 angustifolia 3-74 arborea 3-24 arborescens 3-70 arcana 3-76 arenosa 3-33 argentina 3-21 arrecta 3-36 aspidioides 3-13 assurgens 3-30 atrorubens 3-24 balbisii 3-31 biformata 3-60 biolleyi 3-62 brachyodus 3-50 brachypus 3-18 brausei 3-28 canadasii 3-21 caucaensis 3-28 cheilanthoides 3-34 chrysodioides 3-69 clivalis 3-45 clypeata 3-65 coarctata 3-32 comosa 3-49 comptula 3-23 concinna 3-29 confluens 3-6 consobrina 3-73 conspersa 3-42 contermina 3-32 corazonensis 3-27 ctenitoides 3-37 curta 3-60 decussata 3-48 deflexa 3-30 deltoidea 3-46 demissa 3-21 densa 3-34 dentata 3-41 depilata 3-45 diplazioides 3-14 dudleyi 3-33 dumetorum 3-27 eggersii 3-59 enigmatica 3-16 ensiformis 3-75 erythrothrix 3-59 euchlora 3-25 extensa 3-43 exuta 3-37 falcata 3-75 frigida 3-18 funckii 3-18 furfuracea 3-34 furva 3-19 gardneriana 3-52 gigantca 3-68 glandulosa 3-50 glandulosolanosa 3-20 gongylodes 3-40 grandis 3-46 guyanensis 3-68 hispidula 3-41 hutchisonii 3-29 interrupta 3-40 invisa 3-46 jamesonii 3-56 juruensis 3-61 killipii 3-60 laevigata 3-24 lancea 3-74 leoniae 3-18 leprieurii 3-5 1 leucothrix 3-32 Hmaensis 3-20 limbata 3-9 lindigii 3-30 lingulata 3-75 linkiana 3-14 lomatosora 3- 1 5 longifolia 3-70 loretensis 3-30 lugubriformis 3-58 macbridei 3-19 macrophylla 3-68 macrotis 3-57 mapiriensis 3-49 maxoniana 3-71 megalodus 3-64 membranacea 3-69 mercurii 3-3 1 micula 3-33 millei 3-28 multiformis 3-35 navarrensis 3-15 nemoralis 3-62 nitens 3-35 oligocarpa 3-15 oligophlebia 3-3 opposita 3-32 opulenta 3-43 pachyrhachis 3-3 1 palustris 3-6 parasitica 3-39 patens 3-43 pavoniana 3-23 pennata 3-64 TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 121 pennellii 3-50 peripae 3-60 peruviana 3-23 phacelothrix 3-22 pilosohispida 3-27 pilosula 3-19 pinnatifida 3-56 poiteana 3-65 proboscidea 3-36 ptarmiciformis 3- 1 7 pteroidea 3-25 pusilla 3-17 quadrangularis 3-4 1 resinosofoetida 3-34 reticulata 3-66 retrorsa 3-27 rudis 3-25 rufa 3-20 ruiziana 3-35 salicifolia 3-74 salzmannii 3-73 scalaris 3-15 schunkei 3-63 semihastata 3-57 serrata 3-71 sodiroi 3-62 sprengelii 3-3 1 subandina 3-20 supina 3-26 tetragona 3-64 thomsonii 3-22 torresiana 3-3 totta 3-40 tristis 3-60 tryonorum 3-62 urbanii 3-64 valdepilosa 3-50 Tmesipteris 6-1 1 Todea 1-20 Trachypteris 2-48 aureonitens 2-48 induta 2-50 pinnata 2-48 Trichipteris 1-120 conjugata 1-128 corcovadensis 1-120 dombeyi 1-123 excelsa 1-120 flava 1-125 frigida 1-126 infest a 1-123 kalbreyeri 1-124 lasiosora 1-125 latevagans 1-124 lechleri 1-127 microdonta 1-127 nigra 1-124 nigripes 1-124 phegopteroides 1-127 pilosissima 1-125 procera 1-123 pubescens 1-126 serpens 1-126 tryonorum 1-128 Trichomanes 1-76 accedens 1-98 angustatum 1-83 angustifrons 1-86 ankersii 1-90 applerianum 1-88 arbuscula 6-97 bancroftii 6-97 bicorne 1-91 botryoides 1-92 brachyblastos 1-81 capillaceum 1-84 cellulosum 1-85 ciliatum 1-67 collariatum 1-81 coriaceum 6-97 crinitum 1-94 crispum 6-97 cristatum 1-97 delicatum 1-98 diaphanum 1-83 diversifrons 1-91 ekmanii 1-88 datum 1-96 elegans Rich. 1 -84 elegans Rudge 1 -9 1 flaccida 2-95 fragile 1-69 fucoides 1-56 haenkeanum 1-98 heterophyllum 1-94 hirsutum 1-67 hookeri 1-88 hostmannianum 1-93 humboldtii 1-94 hymenoides 1-86 hymenophylloides 1-83 krausii 1-87 kunzeanum 1-81 lambertianum 1-95 leptophyllum 1-83 lucens 1-95 martiusii 1-94 membranaceum 1-88 muscoides 1-86 opacum 1-84 pedicellatum 1-90 pellucens 1-96 pellucidum 1-96 peltatum 1-57 pennatum 1-92 pilosum 1-95 pinnatum 1-92 plumosum 1-96 plumula 1-95 poeppigii 1-89 polyanthos 1-59 polypodioides 1-89 prieurii 1-84 punctatum 1-86 pyxidiferum 1-82 radicans 1-81 reptans 1-87 rigidum 1-85 rupestre 1-82 scandens 1-76 sellowianum 1-97 sphenoides 1-86 spruceanum 1-91 sprucei 1-86 subsessile 1-90 tanaicum 1-89 tenerum 1-83 trollii 1-91 tuerckheimii 1-90 undulatum 1-97 vandenboschii 1-97 Trichopteris 1-122 Triplophyllum 4-19 acutilobum 4-2 1 dicksonioides 4-2 1 funestum 4-2 1 protensum 4-19 Trismeria 2-16 aurea 2-16 microphylla 2-2 1 trifoliata 2-21 Trogonospora 3-2 Urostachys 6-19 andinus 6-35 attenuatus 6-37 bifida 6-28 brevifolius 6-39 capellae 6-32 caracasicus 6-40 catharticus 6-38 cuatrecasasii 6-45 binervius 6-26 brogniarti 6-27 buesii 6-42 crassus 6-34 cuneifolius 6-50 curvifolius 6-46 darwinianus 6-33 durissimus 6-5 1 elongatus 6-35 engleri 6-39 ewanii 6-50 funiformis 6-42 hartwegianus 6-40 hippurideus 6-24 hohenackeri 6-40 jenmanii 6-44 kuesteri 6-30 lechleri 6-25 var. lehmannii 6-25 lehmannii 6-25 linifolius 6-43 var. tenuifolius 6-44 macbridei 6-32 mexiae 6-28 molongensis 6-47 myrsinites 6-49 nesselii 6-34 nubigenus 6-49 phlegmaria var. ericaefolius 6-48 phylicifolius 6-49 pilgerianus 6-34 polycarpos 6-45 poseidonis 6-24 pruinosus 6-5 1 122 FIELDIANA: BOTANY reflexus 6-27 pyx idifera 1-82 ere nata 4-94 rosenstockianus 6-42 radicans 1-81 ilvensis 4-94 rufescens 6-39 tenera 1-83 montevidensis 4-94 sarmentosus 6-45 Vittaria 2-89 peruviana 4-94 saururus 6-35 angustifolia 2-90 se/a#o 6- 1 9 costata 2-90 stellae-polaris 6-28 filifolia 2-90 Xiphopteris 5-72 subulatus 6-50 filiformis 2-90 blepharidea 5-112 taxifolius 6-4 1 gardneriana 2-91 blepharolepis 5-108 tenuis 6-46 graminifolia 2-90 £>u£H/5-112 tetragonus 6-38 lanceolata 2-86 jamesonii 5-82 weberbaueri 6-30 latifolia 2-91 myosuroides 5-82 weddellii 6-26 moritziana 2-9 1 serrulata 5-83 wilsonii6-44 remota2-9l truncicola 5-109 ruiziana 2-91 stipitata 2-9 1 Vandenboschia vittarioides 2-9 1 Zygophlebia 5-72 angustata 1-83 dudleyi 5-92 capillacea 1-84 mathewsii 5-91 diaphana 1-83 Woodia 4-94 sectifrons 5-72 hymenophylloides 1-83 Woodsia 4-94 wer# 5-92 TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 123 A Selected Listing of Other Fieldiana: Botany Titles Available D FERN ALLIES PTER1 )F PERI [RIDOPH »ublication 1397, S27.00 Pnhliratinn 1403 [RIDOPH ith a Publication 1433. $19.00 [RIDOPHYTA OF PERI Publi< 3.00 PTERIDOPH^ Robert < Publication 1447, $35.00 HECKMAN BINDERY INC. ^^••^••H FEB96