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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

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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 1. Marsilea: M. ancylopoda; M. vestita .... 3 ACKNOWLEDGMENTS 1 2. Salvinia: S. minima; S. auriculata 7 22. MARSILEACEAE 2 3. Azolla: A. filiculoides\ A. microphylla .... 9

Marsilea 2 4. Psilotum: P. nudum . .13 23. SALVINIACEAE 5 5. Equisetum: E. giganteum; E. bogotense 14 Salvinia 6 6. Huperzia: H. eversa; H. binervia; H.

Azolla 8 linifolia var. tenuifolia; H. cuneifolia . 18 24. PSILOTACEAE 11 1. Lycopodium: L. clavatum; L. thyoides 53 Psilotum 11 8. Lycopodiella: L. caroliniana var. meridi- 25. EQUISETACEAE 12 onale; L. cernua 59 Equisetum 12 9. Selaginella: S. peruviana; S. haema- 26. LYCOPODIACEAE 16 todes; S. exaltata 67 Huperzia 19 10. Isoetes: I. andicola; I. lechleri; I. di- Lycopodium 52 spora 90 Lycopodiella 58 27. SELAGINELLACEAE 66 Selaginella 66 28. ISOETACEAE 88 List of Tables Isoetes 89 ADDENDUM 97

1 . Species to Be Added to the Pterido- 1 . Data on the diversity of pteridophyte phyte Flora, Parts I-V 97 species in Peru 100 2. Consideration of Pteridophyte Diversity in Respect to Ecology and Geography ... 98 DEPARTMENTS OF PERU 103

COMPREHENSIVE INDEX TO NAMES . . 1 04 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 "The ferns and fern allies of Guatemala" and is currently working on a monograph of the family. both have a keen of This sixth and final part of the "Pteridophyta Consequently, understanding of Peru" contains the aquatic fern families Mar- the problems in these pteridophyte families, and sileaceae and Salviniaceae as well as the "fern al- for their outstanding efforts in the production of this Flora the authors wish to their lies" Psilotaceae, Equisetaceae, Lycopodiaceae, express deep Selaginellaceae, and Isoetaceae. A brief section on appreciation. would like to extend thanks to Blanca the diversity and ecology are included, written by We special the senior author. The comprehensive index con- Leon (USM) for her invaluable assistance in pre- loans for their and tains not only the names in this final part but those paring and arranging packing from this Peruvian herbari- of the other five parts as well. The general style, shipment important as well as from the Universidad Nacional de typography, form of citations, and so forth follow um, Rolla M. the fa- the previously published parts. These matters are Trujillo (HUT). Tryon appreciates cilities the of adequately dealt with in Part I (Fieldiana: Botany, provided by Department Biology, of South and the aid n.s., No. 20, 1 989), and it is not necessary to repeat University Florida, Tampa, them here. of Alice F. Tryon in the preparation of the treat- Treatment of the Lycopodiaceae has been con- ment of Selaginella. tributed by Benjamin 011gaard and that of Isoeta- The illustrations were contributed by Field Mu- ceae by R. James Hickey. Both are recognized spe- seum scientific illustrator Zorica Dabich, who cre- cialists in these two families. The other genera are ated the original drawings and adapted the rest Bot- a joint effort of Rolla M. Tryon and Robert G. from those previously used in the Fieldiana: ferns fern allies of Gua- Stolze, each critically reviewing the treatments any publication "The and prepared by the other. temala." Her art work, which now has appeared Type collections from Peru are mentioned in in all six parts of this Flora, is an invaluable com- the nomenclature but are not repeated in the spec- plement to the descriptions. We are extremely also to Bent imen citations. They are, however, included in the grateful to her. Thanks go Johnsen, for his Peru range and ecology. The nomenclature of the free-lance artist of Copenhagen, Denmark, valu- genera and species is not intended to be complete. drawing ofHuperzia. We also appreciate the It includes all names based on Peru material and able comments presented by reviewers of the other names that are considered useful to mention. manuscript. the officers of the Abbreviations of periodicals generally follow the We are grateful to following system of Botanico-Periodicum-Huntianum institutions for granting loans of their material or in their her- (1968), while those of books and authors generally allowing us to examine specimens Univer- follow the system of Taxonomic Literature (TL- baria: Herbarium Jutlandicum, Aarhus Field Museum of Natural 2, 1976 et seq.). The acronyms for herbaria follow sitet, Denmark (AAU); Index Herbariorum and are also provided below. History, Chicago (F); Harvard University, Cam- bridge, Massachusetts most Gray Herbarium (GH), some Arnold Arboretum (A); Herbarium Acknowledgments Truxillense, Universidad Nacional de Trujillo, Trujillo, Peru (HUT); Missouri Botanical Garden, Benjamin 011gaard produced a recent Index to St. Louis (MO); Miami University, Oxford, Ohio the names of Lycopodiaceae, as well as taxonomic (MU); Museum National d'Histoire Naturelle, Par-

treatments for the family in "The ferns and fern is (P); University of California, Berkeley (uc); allies of Guatemala" and in the "Flora of Ecua- United States National Herbarium, Smithsonian dor." R. James Hickey contributed Isoetaceae for 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 grows in Peru. It is a small plant with no leaflets Marcos, Lima, Peru (USM). on the petiole; the filiform leaves (petioles) are 1- Materials, mostly types, have also been studied 1 cm long. A single subglobose sporocarp is borne in the following herbaria (acronyms follow Index at the base of a leaf. Unless sporocarps are present, Herbariorum, ed. 8): AWH, B, BKL, BM, BONN, BR, the plant can easily be taken for a species of Cy- C, CGE, CPUN, CR, DUKE, G, GB, GL, HB, HEID, K, peraceae, which grows in wet habitats that are sim-

KRA, L, LG, LIL, LINN, LP, MA, MICH, MOL, MSC, NO, ilar to those of Marsilea.

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- I. Marsilea dation, Systematic Biology Program. The work would not have been possible without this assis- Marsilea L., Sp. pi. 1099. 1753; Gen. pi. ed. 5, tance. However, any opinions and conclusions ex- 485. 1754. TYPE: Marsilea quadrifolia L. pressed are those of the authors and do not nec- Figure 1. essarily reflect the views of the Foundation. Plants palustral or aquatic. Stem usually long- creeping, commonly bearing long roots at the nodes of the stem or along the internodes. Leaves with Family 22. MARSILEACEAE the petiole terminated by 2 adjacent pairs of narrowly cuneate to broadly flabellate leaflets that are Marsileaceae Mirb., Hist. nat. veg. (Lam. & Mirb.) glabrous or pubescent. Veins more or less anas- 5: 126. 1802. TYPE: Marsilea L. tomosing, usually connected at the margin. Sori borne within 1 to several stalked, indurated spo- Stem short- to long-creeping, slender, often rocarps attached to the petiole or at its base, en- branched, hardly indurated, bearing trichomes. closed by a diaphanous indusium, with microspo- Leaves ca. 1-40 cm long, with 4, 2, or no leaflets rangia and megasporangia. Megaspores somewhat at the apex of the petiole, circinate in the bud. Sori ellipsoidal with an apical papillalike laesura, the borne within sporocarps, indusiate, with indehis- surface papillate. Microspores spheroidal, trilete, cent, short- to long-stalked, not annulate, mega- the surface slightly rugulose. sporangia and microsporangia; heterosporous, spores without chlorophyll. Marsilea is a nearly worldwide genus of perhaps 50 species, with about 1 2 in America. The tropical The Marsileaceae are a family of three genera, American species are poorly known for there are all in tropical America. The genera (Marsilea, Reg- relatively few collections and these often lack spo- nellidiwn, and Pilularid) are clearly distinct and rocarps. The species of Marsilea are most diverse not very closely allied. and common in regions that support vernal pools There are numerous detailed studies on the or shallow pools that dry out, at least along the Marsileaceae centering on the form and devel- borders. Mexico is one center of diversity, Aus- opment of the elaborate reproductive structures tralia and Africa are others. and simplified leaves. These plants have also been The following treatment is based directly on that useful in physiological and experimental work on ofJohnson (1986). When necessary, specimens are factors influencing leaf form. However, some of indicated as cited by Johnson. the basic problems concerning the nature of the The sporocarp is borne on a stalk that is con- leaflets and the sporocarp are not resolved, and nected to the petiole. If this stalk is attached by the phyletic relationships of the three genera are its end to the sporocarp, there is neither raphe nor not wholly certain. inferior tooth. If the stalk is attached along its side Pilularia L. is not known from Peru. However, (the raphe), there may be an inferior tooth (the Pilularia americana A. Br. has been collected near- end ofthe stalk). The superior tooth is an extension by in Bolivia (as P. mandonii) and it probably also of the sporocarp near the inferior tooth (fig. Id).

vestita: FIG. 1 . Marsilea ancylopoda: a, habit; b, leaflet; c, stem and roots, with sporocarp. Marsilea 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 JOHNSON, D. M. 1986. Systematics of the New years; Johnson, 1986) and they presumably are World species ofMarsilea (Marsileaceae). Syst. dispersed by waterfowl. Species also may be dis- bot. monogr. 11: 1-87. persed as a result of discarded aquaria.

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. Roots borne only at the nodes (rarely a few 2- preuss. akad. wiss. Berlin. 1863: 421. 1864. 3 cm distant). Raphe present. Superior tooth ab- TYPE: Piauhy, Brazil, Gardner 2760 (holo- sent or slightly raised. Sporocarps 2.5-6 mm long, type, G; isotypes, BM, G, K, P, all cited by John- borne below the ground or stem level, 1 at the base son). of a petiole. Leaflets of non-floating leaves glabrous or with a few trichomes. 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 Northern Peru, Tumbes to La Libertad, near sea ridges. level. Mexico; Florida and Greater Antilles; Venezue- In a pool, ca. 59 m; a single collection from San la, Colombia to Peru; Brazil; Argentina; Uruguay. Martin. Mexico and Central America; Venezuela and Colombia, south to Brazil; Paraguay and Peru. Tumbes: Laguna de Salitoral Grande, Coronado 235 (GH, uc). Lambayeque: San Nicolas, entre Lambayeque et Chiclayo, Santos Quiroz 2401 (F, GH). Prov. Lam- San Martin: Morales, Tarapoto, Vie 6866 (G, K, P, all bayeque, Yencala, Santos Quiroz 71 (HUT). Laguna Boro, cited by Johnson). Chiclayo, Leon 600 (USM). La Libertad: Prov. Trujillo, Huaman, Angulo & Lopez 348 (GH). 2. Marsilea ancylopoda A. Br., Monatsber. konigl. preuss. akad. wiss. Berlin. 1863: 434. 1864. TYPE: Guayaquil, Ecuador, Jameson 3. Marsilea vestita Hooker & Grev., Icon. fil. 2: 304 (holotype, G; isotypes, BM, G, K, all cited t. 159. 1830. LECTOTYPE (designated by by Johnson). Originally as James 304, cor- Johnson, Syst. bot. monogr. 11: 62. 1986): rected later (A. Br., 1871, op. cit. 1870: 717). Columbia River, Scouler 338 (K; isolecto- Figure la-c. types, GH, NY). Figure Id.

FIELDIANA: BOTANY Marsilea uncinata A. Br., Amer. J. Sci. ser. 2, 3: 55. Angaraes, 4 km W of Huanta, Stork & Morton 10808 (F, et al. 7810 1 847. TYPE: Arkansas, Englemann 33 (holotype, uc). Cuzco: Prov. Espinar, Yauri, Nunez (F, Hill MO; isotypes, K, M, MO, all cited by Johnson). MO), Vargas 13524 (GH). Puno: Towards Juliaca, Marsilea mucronata A. Br., Amer. J. Sci. ser. 2, 3: 55. 562 (K), 563 (K). 1847. TYPE: Near Devil's Lake, North Dakota, Geyer 71 (holotype, MO; isotypes, K, NY, all cited by Johnson). Comments

Plants in diffuse or dense colonies. Roots only Marsilea crotophora D. M. Johnson, Syst. bot. at the nodes (rarely a few to 10 mm distant). Raphe monogr. 11: 46. 1986. TYPE: Mato Grosso, present. Superior tooth acute, often hooked at apex. Brazil, Hatschbach & Scherer 30470 (holo- Sporocarp 3.0-7.6 mm long with lateral ridges. type, us; isotypes, c, LP, M, MICH, NY, uc, all Leaflets of non-floating leaves covered by over- cited by Johnson). lapping trichomes on both surfaces.

This species may be found in the upper Amazon All of the above pertains to ssp. vestita; ssp. or its tributaries. It may be distinguished from tenuifolia (A. Br.) Johnson is endemic to central Peruvian species by the roots being borne on the Texas. It differs from ssp. vestita in having very internodes as well as the nodes, the lack of a raphe narrow leaflets with sparse trichomes or none and superior tooth, and the 5-20 sporocarps borne abaxially. on the basal % of the petiole, the lowest one borne Known in Peru only from the Department of well above the petiole base. Lambayeque. Southern British Columbia east to western Min- nesota; south to Louisiana, Florida; Mexico; Peru. The Peru and Florida localities may be introduc- tions.

23. SALVINIACEAE Lambayeque: Chiclayo, Leon (MICH, det. Johnson). Family Prov. Lambayeque, Salazar HI (USM). Salviniaceae Reichenb., Bot. damen kunst. freunde 4. Marsilea mollis Rob. & Fern., Proc. Amer. pflanzenw. 255. 1828. TYPE: Salvinia Se- Acad. 30: 123. 1895. TYPE: San Diego, Chi- guier. huahua, Mexico, Hartman 604 (holotype, GH; isotypes, F, GH, MSC, NY, uc, all cited by John- Azollaceae Wettst. Handb. syst. bot. 2: 77. 1903. TYPE: Azolla Lam. son).

Plants in water or stranded in wet mud. Roots borne only at the nodes (occasionally to floating Stem and often 10 mm distant). Raphe present. Superior tooth elongate, small, slender, branched, not trichomes. absent or to 0.2 mm long. Sporocarps at or near indurated, usually bearing Floating leaves ca. 0.5-2.5 cm not circinate in the petiole base, 1.7-6.7 mm long, on recurved pe- long, bud. Sori borne on a lobe of a leaf or on a branched duncles, borne above the ground level. Leaflets of with the not an- non-floating leaves adaxially glabrous to sparingly leaf, indusiate, sporangia stalked, the sori either or micro- pilose, abaxially densely pilose. nulate, megasporangiate sporangiate; heterosporous, spores without chlo- 2000-3900 m, Cajamarca south to Puno. rophyll. Arizona and Texas south to northern South The structure bearing the sporangia is considered to be a surrounded an America, south to Argentina, Chile and Brazil. sorus, by indusium, and the term is not The following sterile specimens are considered accordingly special sporocarp used. to represent this species. All have been seen by the The Salviniaceae are two present authors except the Hill collections from living represented by Salvinia and of Puno. All have been placed here by Johnson except genera, Azolla, floating aquatics in the Al- the Nunez et al. collection from Cuzco. wide distribution, especially tropics. though Azolla is sometimes included in a separate it has basic similarities with Sal- Cajamarca: Cajamarca, Muller & Gutte 27322 (USM). family, many Junin: Huancayo, Kunkel 422 (GH). Huancavelica: Prov. 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 is a widely distributed genus of about ten species, with seven of them in America. Salvinia Seguier, Fl. veron. 3: 52. 1754. TYPE: Shaparenko (1956) recognized four sections, and Salvinia natans (L.) All. (Marsilea natans L.) these were adopted by de la Sota (1962); they are Figure 2. not recognized here because the genus is a small one.

The descriptions of the species are freely adapt- Plants floating aquatics. Stem short or elongate, ed from Stolze(1983). 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 References cm long, often pubescent, usually papillate on the upper surface, one of them submerged, pendent in SHAPARENKO, K. K. 1956. History of the Sal- the water, highly branched, bearing many tri- vinias (in Russian). Act. Inst. Bot. Komarov. chomes. Veins anastomosing. Sporangia borne in Ser. VIII Paleobotania 2: 1-44. stalked sori on the submerged leaf, enclosed by SOTA, E. R. DE LA. 1962-1964. Contribution al the indusium, with either megasporangia or mi- conocimenti de las "Salviniaceae" neotropi- crosporangia. Spores trilete, enclosed within a vac- cales. I-V. Darwiniana 12: 465-520, 612-623; uolate tapetal formation, the megaspore surface 13: 529-536. perforate, the microspore surface somewhat ru- STOLZE, R. G. 1983. Ferns and fern allies of Gua- gulose. 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. with apical trichomes, these separate (not joined) TYPE: Santa Catarina, Brazil, Muller 479 at the apex. (holotype, K; isotype, BM). Figure 2a-c. 130-160 m, Loreto and Madre de Dios. Southeastern United States; Mexico and Central Stele of the stem more or less circular in cross- America; South America, south to Argentina and section. Floating leaves oblong-elliptic, the apex Uruguay. obtuse (rarely retuse), the larger ones 0.6-1.3 cm This species has often been called S. rotundifolia = long, their upper surface papillate, each papilla 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, collar, which is between the megaspore proper and Rio Revilla Ellenberg 8301 (F). Itaya, Iquitos, 467 (F, the floats. In the living species the floats are either MO, us, USM). Lupuna Cocha, Iquitos, Tryon & Tryon three or nine in number. The microsporangia con- 5188 (F, GH, us, USM). Madre de Dios: Prov. Tambopata, tain a few to several massulae, within which the Tambopata Nature Reserve, 30 air km SSW of Puerto Maldonado, Barbour 5373 (MO). Prov. Manu, Parque microspores are embedded. Projecting structures Nacional Manu, Foster 12206 (F). called glochidia are often on the outer surface of the massulae. Azolla is a widely distributed genus of six spe- 2. Salvinia auriculata Aublet, Hist. pi. Guiane cies, with four of them (sect. Azolla) in America Franc. 2: 969. 1775. TYPE: French Guiana, and in Peru. It is notable in having the most com- Terr. Caux, Aublet (p?). Figure 2d-e. plex reproductive structures of any plant and in having a colony of Anabaena azollae within the Salvinia rotundifolia Willd., Sp. pi. ed. 4, 5: 5 1 7. 1 8 1 0. lobes of each leaf. This blue-green alga has the TYPE: Brazil, Hoffmannsegg(see Morton, Contr. U.S. Natl. Herb. 38: 75. 1967). ability to fix atmospheric nitrogen, and accordingly Azolla is the center ofwide interest as a green- Stele of the stem U-shaped in cross-section. manure (Lumpkin & Plucknett, 1980), especially Asia. Floating leaves oblong-elliptic and nearly circular, for the rice paddies of Southeast obtuse or retuse at the apex, larger floating leaves Azolla is a very distinctive genus, but the species 1 .5-2.5 cm long, their upper surface papillate, each are poorly known due to the fact that sterile ma- papilla with 4 apical trichomes, these joined at the terial is difficult to identify. Fertile material is un- apex. common. Accordingly, the species in Peru and their At ca. 100 m, Loreto and Madre de Dios. characters are only provisionally known. In spite Cuba; Mexico south to Argentina. ofevidence to the contrary, the treatment of Sven- son (1944) is largely followed. The key is adapted

Loreto: Rio Amazonas, SE of Iquitos, Croat 19296 from his key to species of the Americas and the (MO). Prov. Maynas, mouth of Rio Nanay, Gentry el al. megaspore characters are taken from Perkins et al. 21715 (MO, uc). Yanamono, et al. 42574 (F, Gentry MO). (1985). Species names may be assigned to one of Madre de Dios: Prov. Tambopata, Tambopata Nature the four species adopted, although this does not Reserve, 30 air km SSW of Puerto Maldonado, Barbour that the material is determined. 4888 (MO). mean accurately The following Peru collections have not been identified to species: Cajamarca: Dillon 2919. La Li- II. Azolla Lam., Encycl. 1: 343. 1783. TYPE: bertad: Killip & Smith 21511; D. Smith et al. 2225. Azolla filiculoides Lam. Figure 3. Cuzco: Herrera 2616, 2618; Cook & Gilbert 237, 241, 1078; Cook 1963; Maldonado 17. Puno: Bar- Floating aquatics. Stem short, elongate, slender, clay 9273. sometimes bearing trichomes, and usually short Species of Azolla, as they are known, are not roots. Leaves ca. 0.5-1.5 mm long, bilobed, the restricted to particular environments as differ- upper lobe usually minutely to strongly papillate, ences in range and habitat statements might imply. without veins. Sporangia borne in short-stalked Accordingly, the range, habitat, and altitude are sori enclosed by an indusium, with either 1 mega- given for the genus as a whole in Peru, rather than sporangium or several microsporangia. Spores tri- for each species. lete, the megaspores irregularly marked, with a In wet mud of riverbanks or stream banks, where perforate surface, the microspores smooth, em- stranded by higher water, usually floating in stand- bedded in massulae. ing water of lakeshores, ponds, or small streams, also in ditches, in stagnant or fast-flowing water, The megaspore has accessory structures usually less often in swamps or wet parts of cultivated referred to as a columella bearing floats. These are land, near sea level to 4100 m, Cajamarca, Ama- partially enclosed by a band of sporoderm, the 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 H. E. CALVERT. 1985. Scanning electron microscopy of perine architecture as a taxonomic LUMPKIN, T. A., AND D. L. PLUCKNETT. 1980. tool in the genus Azolla Lamarck. Scanning Azolla, botany, physiology, and use as a green- Electron Microscopy 4: 1719-1734. manure. Econ. Bot. 34: 1 1 1-153. SVENSON, H. K. 1944. The New World species PERKINS, S. K., G. A. PETERS, T. A. LUMPKIN, AND 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 32119 (GH). Ayacucho: 44 km N of Tambo, Luteyn & 6369 Cuzco: Prov. other authors, not Willd., as usually thought. Lebron-Luteyn (uc, us). Cuzco, Say- lla, Fernandez et al. 455 (USM). Puno: Santa Rosa, Staf- ford 495 (F). Ichu, Soukup 354 (F). Plants dichotomously branched, 0.5-1.0 cm in diameter. Leaves divaricate (not closely imbri- 2. Azolla mexicana Presl, Abh. Bohm. Ges. Wiss. cate), nearly orbicular, nearly smooth, small, 0.5 Ser. 3: 150. 1845. TYPE: Schiede mm long. Glochidia not septate. Megaspore collar V, Mexico, & with dense filaments on its surface (magnification Deppe (holotype, PR). x200 or more), rugulate-verrucate beneath filaments. Plants dichotomously branched, often 1.0-1.5 cm in diameter. Leaves divaricate (not closely im- with the ca. Eastern North America; Mexico and the Carib- bricate) upper lobe 0.7 mm long, the lower one much Glochidia bean; introduced in Europe and South America. larger. septate. Mega- collar surface The name is used in the sense of Svenson and spore glabrous; megaspore (magnification x 200 or also with most authors (not Willd.) until a definitive treat- more) coarsely rugose, and with dense filaments. ment is published. pits, many Jermy (Somerfeltia 6: ix-x. 1987) has pointed North Central out that according to the Ph.D. thesis of D. Dun- America; Mexico; America; northern half of South America. ham (Portsmouth Polytechnic) the type of Azolla caroliniana is A. filiculoides. This leaves Azolla of Tumbes: Casa Loreto: eastern North America presumably without a Fernandez, Haught 183 (us). Prov. & 5213 name. Maynas, Iquitos, Tryon Tryon (GH, us). Ancash: Lago Santa Cruz Chico, Huascaran National Park, D. Smith et al. 9261 (F, HUT, MO). Junin: Huancayo, l Amazonas: Rio Santiago, S of La Poza, Berlin 3711 Kunkel 421- h (GH). Acopalca, Kunkel 421 (GH). Cerca (MO, uc). La Libertad: Prov. Bolivar, Laguna de las Ichus, Carhuamayo, Ferreyra 5242 (USM). Cuzco: Pampa de Lopez & Sagdstegui 3238 (GH, MO). San Martin: Prov. Anta, Iltis & Ugent 785 (GH, uc, us, USM). 3.5 km NW Mariscal Caceres, 2-4 km W of Tocache Nuevo, Plow- of Saylla, Iltis et al. 1208 (GH, us). Prov. Paucartambo, man 11437 (F). Loreto: Prov. Maynas, Rio Itaya, Revilla Paucartambo, Plowman & Davis 4920 (F). Madre de 632 (F, MO, uc.). Prov. Maynas, Yanuyaca, Gentry et al. Dios: Prov. Manu, Parque Nacional Manu, Foster 9872

10 FIELDIANA: BOTANY (MO). Puno: Pucara, Hunnewell 15874 (GH). Lake Titi- Family 24. PSILOTACEAE caca, Shepard 1511 (GH, us). Moquegua: Ilo, Stafford 926 (F). Psilotaceae Kanitz, Novenyrends. Attek. 43. 1887. TYPE: Psilotum Sw. 3. Azolla filiculoides Lam., Encycl. 1: 343. 1783. TYPE: "Magellan Region," Herb. Lamarck Stem flaccid to somewhat indurated, lacking in- (p). Figure 3a-d. dument and roots. Leaves (or pinnae) small to minute, borne alternately, with a single vein or Plants pinnately branched, often 2-6 cm long. none. Two or 3 sporangia joined in a sessile, thick- Leaves closely appressed, imbricate, oblong to walled synangium. Homosporous. Spores without ovate, ca. 1 mm long. Glochidia not septate (rarely chlorophyll. 1-2 "septa" toward the apex). Megaspore collar more or less glabrous; megaspore surface (mag- The Psilotaceae are a family oftwo genera, Tme- nification x 200 or more) prominently rugose with sipteris Bernh. of western Malesia, Australia, and hexagonal markings, with few filaments. the Pacific, and Psilotum, which is pantropical in distribution. Southern South America north to western North After an extensive series of studies, D. W. Bier- America and to Alaska; introduced in Europe, Asia, horst concluded that the Psilotaceae were primi- and Australia. Azollafiliculoides var. rubra (R. Br.) tive elements in the Filicopsida. This interpreta- Strasb., treated as a species A. rubra R. Br. by tion of the position of the family provoked further Perkins et al. (1985), is not considered to occur in interest in its phylogeny and resulted in a sym- Peru. posium on the taxonomic and morphological re-

lationships of the Psilotaceae (White et al., 1977). Cajamarca: Prov. Contumaza, Rio San Benito, San The Psilotaceae have a number of characters Benito, Sagdstegui et al. 12509 (HUT, MO). Amazonas: that occur in the such as the details Prov. Bongara, Laguna Pomacocha, Wurdack888 (F, GH, Filicopsida, uc, us). La Libertad: Prov. Trujillo, Tschudi, Shimo- of early ontogeny of the leaf, the subterranean, kowa 7294 de Chan Chan, 18 (us). (HUT). Playa Angulo cylindrical gametophytes, multiflagellate sperm, Loreto: Paraiso, above Iquitos, Rio Amazonas, Fosberg and several aspects ofthe rhizoids and gametangia. 29076 (us). Huanuco: Chasqui, Macbride 3306 (F, us); These and the evidence from the structure and wall 3307 (F, us). Lima: Prov. Lima, Laguna de Villa, Corona- do 5 (GH, MO, uc, us); Tryon & Tryon 5458 (GH); Cerrate formation of Psilotum spores are indicative of re- 2766 (USM). Huancavelica: Prov. Tayacaja, Pampas, Stork lationships with the ferns. However, Cooper-Driv- & Morton 10243 uc, Cuzco: Cook & Gilbert 197 (F, us). er (1977) has shown that the chemical evidence (us). Prov. Paucartambo, Huilabamba, Balls B7632 (F, does not support this relationship. uc, us). Arequipa: Prov. Arequipa, Rio Yarabamba, Ponce 32 (USM). References

4. Azolla microphylla Kaulf, Enum. fil. 273. 1 824. COOPER-DRIVER, G. 1977. Chemical evidence TYPE: "California," Chamisso (LE?). Figure for separating Psilotaceae from the Filicales. 3e-f. Science 198: 1260-1261. WHITE, R. A., ET AL. 1977. Taxonomic and mor- Plants pinnately branched, small, 1-2 cm long, phological relationships of the Psilotaceae. Brit- many leaves orbicular, 1 mm long, most leaves tonia 29: 1-68. closely imbricate. Glochidia septate. Megaspore collar glabrous; megaspore surface slightly fila- I. Psilotum mentous, slightly pitted, rugulate (high magnification) to verrucate. Psilotum Sw., J. Bot. (Schrader) 1800 (2): 8, 109.

1 1 80 , nom. nov. for Hoffmannia Willd. (not Western and northern South America; southern Sw.) and with the same type. North America; West Indies. = Hoffmannia aphylla Willd. Psilotum nudum (L.) Loreto: & Smith 27707 Prov. Yurimaguas, Killip (us); Beauv. Figure 4. Requena, Rio Ucayali, Encarnacion E-1101 (us). Aya- cucho: Puquio, Gentry et al. 23296 (F, MO, us). Cuzco: Plants or Prov. Anta, Pampas de Anta, Tryon & Tryon 5364 (GH, terrestrial, rupestral usually epiphytic. us, USM). Yauri, Nunez et al. 7813 (F, MO). Stem compactly branched in the substrate, the ae-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 11 rial stems green, erect to pendent, with several Family 25. EQUISETACEAE dichotomous divisions, ca. 20 cm to 1 m long, without roots. Leaves minute, alternate, leafy stem Equisetaceae A. P. DC., Fl. Franc. (Lam. & DC.) glabrous. Synangia large, borne singly in the axil ed. 3, 2: 580. 1805. Type: Equisetum L. of forked fertile leaves. Spores elongate-ellipsoidal, monolete, the laesurae '/2 to % the spore length, Stem jointed, more or less indurated, the aerial the surface coarsely rugose to shallowly and com- usually green, the subterranean with numerous wiry pactly verrucate. roots. Leaves each with a single vein, borne in whorls, joined in a sheath. Sporangia thin-walled, Psilotum is a highly distinctive genus of wide borne on stalked, peltate sporangiophores that form distribution in the tropics and with some exten- a strobilus. Homosporous. Spores with chloro- sions to subtemperate regions. Notable characters phyll and bearing elaters. of the genus are the dichotomously branched stem, the bifurcate sporophylls, and the large synangium. Psilotum has a simple morphology, proba- I. Equisetum bly through reduction from more complex ances- tors. Equisetum L., Sp. pi. 1061. 1753; Gen. pi. ed. 5, 484. 1754. TYPE: Equisetumfluviatile L. Fig- ure 5. 1. Psilotum nudum (L.) Beauv., Prodr. Fam. Ae- theog. 112. 1805. Figure 4. Terrestrial, palustral or aquatic. Stem subterranean, short- to long-creeping, freely branched, Lycopodium nudum L., Sp. pi. 1100. 1753. TYPE: aerial stems that are lon- LINN 1257.1. bearing erect, jointed, and ca. 10 Psilotum triquetrum Sw., J. Bot. (Schrader) 1800 (2): gitudinally ridged, usually hollow, cm 109. 1801, nom. super/I, for Lycopodium nudum to 8 m long, often with whorls of branches. Leaves L. and with the same type. small, in whorls at nodes, the lower portion laterally fused in sheaths, the upper portion in more Plants or erect. Branches 3-an- pendent usually or less prolonged teeth. Sporangia large, several to 1 .2 leaves 2-3 mm gled, mm wide, distant, long, borne on each stalked, peltate, apically flattened triangular-subulate. sporangiophore, in a condensed terminal strobilus. Spores spheroidal with circular aperture, and on and tree Epiphytic, especially palms ferns, 4 paddle-shaped elaters, the surface with small 200-600 San Martin south to Madre de Dios. m, granulate and large spherical deposits. American and Old World tropics and subtrop- ics. The Equisetum strobilus is composed of spo- P. Sw. be a distinct complanatum may species. rangiophores that are sometimes called sporo- It has flattened branches 2 mm or more wide and phylls. A detailed study of these structures by Page is distinctive in the Pacific islands. especially (1972) shows they are partly modified from a leaf and partly from a cauline appendage; thus the term San Martin: Dist. Rimachi 7945 Dist. Chazuta, (MO). sporangiophore is more appropriate. The small Tocache Nuevo, Schunke 4084 (F, us). Gramalote ad leaves that surround each node are laterally fused Saposoa, Woytkowski 7300 (GH, MO, us). Loreto: Lower below and form coarse teeth above. 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: 3687 de Dios: Prov. Tingo Maria, Moran (USM). Madre References Tambopata, Cuzco Amazonico Lodge, 15 km NE of Puerto Maldonado, Nunez 12214 (MO); Gentry et al. tax- 68958 (MO). Prov. Manu, Parque Nacional Manu, Foster HAUKE, R. L. 1961-1962. A resume of the 11336(F). 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 a tall, with 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. likely it should be recognized as a subspecies, as 1759. TYPE: Plumier, PI. Amer. (ed. Bur- is E. ramosissimum ssp. debile, but we do not wish man) t. 125, f. 2 (1757 as to the plate). Figure to make a new combination at this time. 5a-b. Equisetum giganteum has the turbercles on the ridges of the stem square or flattened in profile Equisetum myriochaetum Schlect. & Cham., Linnaea while the turbercles of E. myriochaetum appear 5: 623. 1 830. TYPE: Misantla, Veracruz, Mexico, rather similar to the teeth of a saw. The former Schiede and Deppe 833 (holotype, B?). species has two to several lines ofstomates on each Equisetum schaffneri Milde, Verb. Zool.-Bot. Ges. side of a ofthe stem, while the latter species Wien 1 1: 345. 1861. TYPE: Orizaba, Mexico, W. groove Schaffner 315 (holotype, B?). 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- Stem usually 2-4 m tall, often bearing an apical specific. strobilus, usually 1 5-25 mm wide (10 mm or more, Equisetum schaffneri has a variable mixture of rarely 5 mm or less), with a central hollow. En- the characters ofE. giganteum and ofE. schaffneri. dodermis surrounding each of the vascular bun- It has a wider distribution than a sterile hybrid dles. Sheaths ofthe main stem usually light brown, should have, and it occurs in regions in which only with often deciduous, rather indurated teeth. one of the putative parents is known (Venezuela Branches in regular whorls, with 8-10 ridges. and in Peru, for example). Accordingly, it may be Strobilus sessile or nearly so. a natural intermediate of the extremes of E. giganteum. The abortive spores present in some col- In wet, usually open areas, in sloughs, on the lections (not all have strobili) are also present in banks of streams, rivers, and irrigation ditches, in some other species: in E. bogotense and E. var- boggy areas, and in streambeds, 1900-3500 m, iegatum (Hauke, 1963, 1978) where hybridization Piura, Cajamarca and Amazonas, south to Are- is not concerned. quipa and Puno. Hauke (Ph.D. dissertation, University of Mich- Greater Antilles; Mexico and Central America; igan) cites the following collections of E. myri- Venezuela and Colombia, south to Chile, Argen- ochaetum from Peru: Ayacucho, Killip & Smith tina and Brazil. 22757 (us); Cuzco, Cook & Gilbert 1372 (us). Also Equisetum myriochaetum may be a species, as the following collections of E. x schaffneri from

Hauke and others have it, or it may be an extreme Peru: Lima, Rose 18762 (NY, us); Ucayali?, Herre- variation of E. giganteum, as treated here. Most 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- GH, us). Pasco: Prov. Oxapampa, Los Chacos, near Oxa- bayeque: Olmos to Jaen, Correll & Smith P792 (GH). pampa, Smith & Pretel 1514(f, MO, USM). Junin: Above Cajamarca: Prov. Cutervo, Llatun, Mostacero et al. 1740 Conception, Correll & Smith P736 (GH). Huancavelica: Stork 10863 (HUT). Prov. Celendin, alrededores de Celendin, Mos- Prov. Huancavelica, Yauli, & Morton (F, tacero et al. 835 (MO, uc). Amazonas: Prov. Bagua, 3.9 uc). Ayacucho: Santiago, about 67 km from Nasca on Correll Smith PI 51 Cuzco: Prov. km NE of Chiriaco, Barbour 4360 (F, MO). Utcabamba, road to Puquio, & (GH). Pennell 15586 (GH, us). La Libertad: Prov. Trujillo, valle Urubamba, Pomatales, Nunez 7327 (F, MO). Arequipa: de Rio Chicama, Miiller & Krebs 12137 (USM). San Mar- Chacra Pachacutac, Balls B5868 (F, uc, us). Puno: San to Dillon et al. 1239 tin: Tarapoto, Martin & Plowman 1843 (F, GH). Prov. Gabon (San Gaban) Ollachea, (MO). Mariscal Caceres, 45 km NE of Tingo Maria, Tryon & Tacna: Prov. Tarata, Chuvire, Metcalf 30407 (MO, uc, Tryon 5270 (GH). Ancash: Prov. Santa, alrededores de us). 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 Family 26. LYCOPODIACEAE (GH, uc). Ucayali: Prov. Coronel Portillo, Arboretum von Humboldt, Diaz et al. 687 (F, MO, us). Huancavelica: Contributed by Benjamin 011gaard Prov. Tayacaja, Huari, Sounders 1148 (F, GH, us). Aya- cucho: Ayna, Killip & Smith 22757 (F). Apurimac: Prov. in Hist, Abancay, Trancapata, Vargas 9617 (F, uc, us). Cuzco: Lycopodiaceae Mirbel, Lam. & Mirbel, of Aguas Calientes, Solomon 3180 (F, MO); 39 km E nat. veg. 4: 293. 1802. ("Lycopodia"). TYPE: on et al. 23399 Urohuasi Abancay-Cuzco road, Gentry Lycopodium L. (MO, uc). Madre de Dios: Prov. Vlanu. Parque Nacional del Manu, Foster 9796 (MO); 9856 (MO). Arequipa: Yura, Plants terrestrial or erect to Solomon 2912 (F, MO, USM). Puno: Puno, Angulo 1760 epiphytic, pendu- (HUT). lous herbs or climbers. Stems dichotomously branching, sometimes also with lateral branching, 2. Fquisetum bogotense HBK., Nov. gen. sp. pi. protostelic, with the xylem arranged radially or in 1: 42. 1815. TYPE: "Prope Santa Fe de Bo- parallel bands, or forming an incomplete cylinder gota et Alto del Roble, Columbia," Humboldt (Phylloglossuni). Leaves simple, with 1 simple vein, & Bonpland (holotype, P?). Figure 5c-d. arranged in low alternating spirals or irregular alternating whorls, or seemingly decussate, homo- Steins usually 20-50 cm tall, commonly bearing phyllous or heterophyllous, isophyllous or aniso- an apical strobilus, 1-2 mm wide, solid in the phyllous. Sporophylls like the foliage leaves, or center. Endodermis single, surrounding the ring of modified, in some groups specialized and aggre- vascular bundles. Sheaths with short, brown, pa- gated into distinct strobili. Sporangia solitary, ax- pery teeth. Branches irregular, with 4 ridges. illary or on the upper side of the sporophyll base, Strobilus stalked. homosporous, unilocular, reniform to subglobu-

lar, short-stalked, dehiscing by a transverse slit, In soil or among rocks, in wet open places, in- which divides each sporangium into 2 nearly equal cluding banks of creeks, rivers, and irrigation or strongly unequal valves. Spores without chlo- ditches in moist to wet seeps and clay road banks, rophyll, tetrahedral, with a trilete scar. Gameto- 100-4200 m, most often above 2000 m, Piura phytes monoecious, tuberous, subterranean, and south to Puno and Tacna. mycorrhizal and without chlorophyll or surface- Costa Rica and Panama; Galapagos Islands; the living and green. Andes of Venezuela and Colombia, south to Bo- livia; to southern Argentina and Chile. The family occurs in almost all humid regions of the world, both warm and cold. It is here treated Piura: Prov. Huancabamba, Cerro Blanco, Stork 114 15 as consisting of four genera including Phylloglos- Prov. (GH, uc, us). Cajamarca: Contumaza, Guzmango sum (Australia, New Zealand). Up to 14 genera to Santiago, Sagdstegui & Lopez 10584 (F, MO). San Pa- have been recognized by other authors (Holub, blo, Hutchison & Wright 5060 (F, MO, uc, us). Amazonas: 1 99 1 & Beitel, ; 1992), Hills WNW of Pomacocha, Wurdack 9445 (GH, uc, us, 1975, 1983, 1985, Wagner USM). La Libertad: Above Yamobamba, Conrad 2728 the genera of these authors generally correspond-

Martin: corner of Rio Abiseo National 1 1 (MO). San NW ing to the infrageneric taxa of 011gaard ( 987, 989, Park, & Leon 4534 (HUT, USM). Ancash: Prov. Young 1992); for a review of the classification, an index Recuay, Lopez et al. 7596 (MO, us). Huanuco: Prov. to the family, and an overview of the Neotropical Huanuco, Cerro Carpish, Luteyn & Lebron-Luteyn 5483 see these Most of the Pe- (F, uc, us). Lima: Prov. Huarochiri, Sullivan et al. 1103 species, publications. (F, MO); Prov. Canta, Dist. Huaros, Pennell 14715 (F, ruvian species are illustrated in 011gaard (1988).

16 FIELDIANA: BOTANY The total number of species in the family probably cies to Huperzia: With a note on generic clas- exceeds 350, and approximately 185 of these are sification in Huperziaceae. Folia Geobot. Phy- tropical American. The Andes have a particularly totax. 20: 67-80. high diversity, and many species are conspicuous HOLUB, J. 1991. Some taxonomic changes with- elements in the montane and alpine vegetation of in Lycopodiales. Folia Geobot. Phytotax. 26: these mountains. 81-94. This treatment includes 62 species for Peru, but 0LLGAARD, B. 1987. A revised classification of the real number is likely to be higher. Although the Lycopodiaceae sens. lat. Opera Bot. 92: 1 53- largely all available material has been consulted 178. for this treatment, it is evident that much collect- 0LLGAARD, B. 1988. Lycopodiaceae, in G. Har- ing needs to be done before a complete presen- ling and L. Andersson, Flora of Ecuador 33: 1- tation can be made. Many species are represented 155. by a single or few collections, and intensive efforts 0LLGAARD, B. 1989. Index of the Lycopodi- in limited areas such as those of Blanca Leon and aceae. Biol. Skr. Dan. Vid. Selsk. 34: 1-135. Kenneth Young in the Rio Abiseo National Park, 0LLGAARD, B. 1992. Neotropical Lycopodi- of Abundio Sagastegui in the north of Peru, and aceae An overview. Ann. Missouri Bot. Gard. of the late David N. Smith in several areas have 79:687-717. turned up several species not formerly known from ROTHMALER, W. 1 944. Pteridophytcn-Studien I, Peru and widely disjunct from other known oc- Feddes Repert. Spec. Nov. Regni Veg. 54: 55- currences, as well as species new to science. 82. TRYON, A. F., AND B. LUGARDON. 1991. Spores of the Pteridophyta. Springer-Verlag, New York. References TRYON, R. M., AND A. F. TRYON. 1982. Lyco- podiaceae, in Ferns and allied plants, with spe- HOLUB, J. 1975. Diphasiastrum, a new genus in cial reference to tropical America. Springer-Ver- Lycopodiaceae. Preslia 47: 97-1 10. lag, New York. HOLUB, J. 1983. Validation of generic names in WAGNER, W. H., AND J. M. BEITEL. 1992. Mod- Lycopodiaceae: With a description of a new ge- ern North American Lycopodiaceae and their nus Pseudolycopodiella. Folia Geobot. Phyto- generic classification. Ann. Missouri Bot. Gard. tax. 18: 439-442. 79: 676-686. HOLUB, J. 1985. Transfers of Lycopodium spe-

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, indeterminate, rhizomatous, or creeping, or trailing, main stems, and usually determinate aerial branchlet 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 et al. Holm-Nielsen 4422, Ecuador, AAU; h, i, j from Hutchison & Wright 6943, Peru, p.)

18 FIELDIANA: BOTANY I. Huperzia There are approximately 300 species worldwide, 49 in Peru. Seven species are endemic to

Huperzia Bernh., J. Dot. (Schrader) 1800 (2): 126. Peru, four are shared only with Bolivia and more 1801. LECTOTYPE (designated by Roth- southerly regions, 26 occur also in Ecuador or far- maler, Feddes Repert. Spec. Nov. Regni Veg. ther north in the Andes and the Neotropics, while 54: 59. 1944): Huperzia selago (L.) Martius 12 species occur both to the north and south of & Schrank (Lycopodium selago L.). Figure 6. Peru. Species delimitation is problematic almost Plananthus Mirbel in Lamarck & Mirbel, Hist. nat. throughout the genus due to the simple morphol- 3: 476. 1802. TYPE: Plananthus veg. selago (L.) ogy of the group and to plasticity of the characters Beauv. (Lycopodium selago L.). (011gaard, 1992). Morphogenesis seems to be Lycopodium subgen. Selago Baker, Handb. Fern-Al- somewhat unstable in most and be lies 8. 1887. TYPE: Lycopodium selago L. species may Lycopodium subgen. Urostachya Pritzel, Nat. Pflan- modified by external factors. Most characters are zenfam. 1 592. 1901. se- (4): TYPE: Lycopodium plastic within a species (e.g., stem thickness, num- lago L. ber of leaf orthostichies, leaf crowding, leaf direc- Urostachys (Pritzel) Herter, Beih. Bot. Centralbl. 39: tion, color, degree of heterophyllous differentia- 249. 1922. TYPE: Urostachys selago (L.) Herter seems to occur rather (Lycopodium selago L.). tion). Hybridization Phlegmariurus Holub, Preslia 36: 21. 1964. TYPE: frequently, contributing to the blurring of species Sen & Sen Phlegmarius phlegmaria (L.) (Lyco- limits, and the putative hybrids often have nor- podium phlegmaria L.). mally developed spores. As a consequence, species Plants epiphytic or terrestrial, pendent, re- recognition is often based on some experience and curved, erect or ascending, isotomously branched comparison with identified material, rather than throughout (except in connection with bulbil for- a set of definite characters. mation in the group of H. selago, and sprouting Measurements and terminology. The descrip- from the base of old plants), the branches all sim- tions and key characters are based on dried ma- ilar (homoblastic), or in some terrestrial species terial, unless otherwise indicated. Very often mea- differentiated (heteroblastic) into prostrate (some- surements and solid shape of organs in life deviate times subterranean) versus erect, aerial branches. strongly from the dried condition. Roots arising from the stem stele, descending Branching is isotomous throughout the genus, through the cortex to the stem base, here emerging except in connection with sprouting (lateral as 1 basal tuft, or in heteroblastic species some- branching) from the base of old or injured indi- times emerging directly along the underside of viduals. Isotomy results in the formation of equal- prostrate shoots. Shoots homophyllous or hetero- ly thick branches each with an equal amount of phyllous. Sporophylls and vegetative leaves alike, vascular tissue. However, many species of high not peltate, without mucilage cavities, persisting Andean grasslands develop heteroblastic branch and green after sporangium dehiscence, the grad- pairs; i.e., the isotomous branches differentiate into ual or abrupt constriction of distal divisions of distinct aspects and functions due to different leaf heterophyllous species associated or not with pres- development, as exemplified by the sporangiate, ence of sporangia. Sporangia axillary, reniform, erect, aerial shoots and the sterile, prostrate or isovalvate, with a short slender stalk; side walls of even subterranean, basal, rejuvenating shoots of sporangium epidermis cells sinuate, thickened and Huperzia crassa and H. hypogaea. Homoblastic lignified together with the inner walls. Spores fo- branches have the same aspect and function and veolate or fossulate. Gametophytes (not known for similar leaf development. any Peruvian species) usually subterranean (in ter- Phyllotaxis is poorly understood in the Lyco- restrial plants), mycorrhizal, cylindrical with ra- podiaceae. It seems to be irregularly organized dial or bilateral symmetry, with pluricellular, uni- morphogenetically and quite variable within one seriate trichomes among the gametangia. species, sometimes even within the same individual. Stevenson (Bot. J. Linn. Soc. 72: 8 1-1 00. 1 976) The genus is virtually cosmopolitan, occurring interpreted the phyllotaxis of Huperzia lucidula in tropical, temperate, arctic, and alpine environ- (Michx.) Trevisan (temperate North America) as ments. Species diversity is highest throughout the consisting of low alternating spirals, and found a tropics in evergreen montane forests, and in the definite relation between the number of orthosti- wet Andean grass and shrublands in South Amer- chies and the number of protoxylem lobes in the ica. stele. In many species this relation is different or

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 19 possibly absent, and transitions from spiral to ir- ranks of a shoot. For the sake of brevity, I have regularly whorled phyllotaxis is common, al- described leaf arrangement in terms of leaf whorls, though whorled arrangement is the more com- although this is not everywhere completely accu- mon. The number of longitudinal leaf ranks is rate. readily observed in some cases but often must be estimated, or calculated, from the number of leaves Reference in each leaf whorl, or from the number of leaves contained in one rotation of the leaf spiral. As the ROLLERI, C. 1981. Sinopsis de las especies de leaves of a whorl or a rotation of the spiral alter- Lycopodium L. (Lycopodiaceae-Pteridophyta) nate with the adjacent whorls or spirals above and de la seccion Crassistachys Herter. Revista Mus. below, the number of leaves in two whorls or ro- La Plata (n.s.) Bot. 13: 61-113. tations makes up the number of longitudinal leaf

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, blisterlike 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 triangular-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 somewhat 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 caespitose, 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 protruding, 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, appressed 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 ofbasal 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, 1.54 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 including 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 ofaerial 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 bases 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, subglobose 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 coloration 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. Plants entirely pendulous, expanded leaves ofproximal divisions in usually densely crowded, 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 ofbasal 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 regularly 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 oblanceolate, 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, 14 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 convex 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 overlapping 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, 24 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 divisions, 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 herbaceous 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 Shoots homophyllous, equally thick throughout, Geobot. Phytotax. 20: 73. 1985. 1035 mm in diameter including leaves. Stems excluding leaves 2.5-4 mm thick at the base, ta- Lycopodium hippurideum Christ in Pittier, Primit. Fl. pering to ca. 2-3 mm upward. Leaves borne in Costar. 3 56. 1901. TYPE: Costa El (1): Rica, irregular alternating whorls of 5-8, spreading to Paramo, 3000 m, massif de Buena Vista, 1897, reflexed, sometimes sharply reflexed and ap- Pittier 10619 (holotype, P!; isotype, us!). to the stem, linear to linear-subulate, even- Urostachys hippurideus (Christ) Nessel, Barlappge- pressed wachse 88. 1939. ly tapering from the base or the middle, (10-)1 1- Urostachys poseidonis Herter, Revista Sudamer. Bot. 19 mm long, 0.8-1.3 mm wide, not, or rarely, 10: 122-123. 195 3. TYPE: Ecuador, Prov.Chim- twisted at base, adaxially with a slightly prominent borazo, Penipe, 3400 m, Rose in Milie 35 (ho- vein, with smooth, usually slightly revolute mar- lotype, us!; isotypes, GH!, NY!). to and Lycopodium poseidonis (Herter) Morton, Amer. Fern gins, and indistinctly prominently widely J. 54: 42. 1964. decurrent bases. Sporangia 1.5-2 mm wide.

Plants ascending to stiffly erect from a decum- Upper montane forest, especially near the forest bent base, up to 60 cm tall, sparsely branched. limit, usually on the forest floor, in semishade, alt.

24 FIELDIANA: BOTANY 2500-3500 m, Cajamarca, San Martin, Pasco, at the base, and the margins not revolute, the spo- Cuzco. rangia 1-1.8 mm wide. Central America; Andes from Venezuela to Bo- Upper montane forest, especially near the forest livia. limit, usually on the forest floor, in semishade, alt.

1 800-3600 m, Junin, Huancavelica, Cuzco, Puno. Huperzia hippuridea belongs to a group ofclose- Peru and Bolivia. related to the ly related taxa of high montane forests throughout Closely Huperzia hippuridea (see for tropical America, including also H. arcuata B. 011g. preceding species discussion). (Colombia, Ecuador), H. lechleri, H. montana (Underw. & Lloyd) Holub (Greater Antilles), H. Junin: Prov. Tarma, Carpapata, Soukup 3477 (F, GH). Huacapistana, 1800-2400 m, & Smith 24503 (F, nuda (Nessel) B. 011g. & Windisch (Brazil). Killip GH, us). Huancavelica: Prov. be- Hutchison & Bismarck 6455 and Matthews 963 Tayacaja, Marcavalle, tween Huachocolpa and Tintay, 2600 m, Tovar 4085 from have thicker (presumably Chachapoyas) (USM), Tovar 4089 (USM). Cuzco: Valle Occobamba [Oco- stems and more crowded and coriaceous leaves bamba], 1900 m, Bites 856 (us). Prov. Paucartambo, Valle de near than usual for the species, thus approaching Hu- Pilcopata, Pillahuata, 2500 m, Foster & Wachter 7501 (AAU). Pillahuata, 2850 m, Fitzpatrick & perzia \veddellii (Herter) Holub and H. loxensis Willard (F). San Ignacio, Huadquina, Biies 1410 (us). B. Matthews 963 is a mixed collection, sheets 011g. Puno: Ollachea to San Gaban, 1000-2000 m, Dillon et at to H. mac- of the same number BM pertaining al. 1156 (AAU, F, MO). Prov. Carabaya, road San Gaban bridei. (Lanlacuni Bajo) to Macusani, near Ollachea, 2600 m, Maas et al. 6111 (AAU, USM). Department unknown: Lechler 2023a (K). Vargas 16767 (GH). Cajamarca: Prov. Hualgayoc, Hda. Taulis, 4.6 km beyond Palmito junction on the road to La Playa, 2740 m, 3. Huperzia arcuata B. 011g. in Harling and An- Hutchison & Bismarck 6455 (uc, USM). San Martin: Prov. dersson, Fl. Ecuador 33: 15, t. ID. 1988. Mariscal Caceres, NW corner of Rio Abiseo National Park, Chochos, 3500 m, Young & Leon 4660 (AAU). TYPE: Ecuador, Prov. Carchi, Road El Angel Prov. Mariscal Caceres, Puerta del Monte, 3100-3300 to Tulcan, 3500 m, Holm-Nielsen et al. 5341 m, & Leon 4436 (AAU), 3450 m, Leon & Young Young (holotype, AAU!; isotypes, GB!, QCA!, us!). 1305 (USM). Pasco: Prov. Oxapampa, Santa Barbara, 3200-3300 m, D. Smith 8176 (AAU, USM). Cuzco: Prov. Nacional Paucartambo, Cerro Macho Cruz, Parque Plants ascending to erect from a decumbent base, Manu, 3400 m, Leon 2303 (USM). Department unknown: with nodding shoot apices, up to 30 cm tall (ter- (possibly from Chachapoyas) Matthews 963 (G). restrial), or recurved to pendulous and up to 90 cm long (epiphytes), sparsely branched. Shoots 2. Huperzia lechleri (Hieron.) Holub, Folia Geo- homophyllous, equally thick throughout, 25-30 bot. Phytotax. 20: 74. 1985. mm in diameter, or tapering to ca. 12 mm including leaves. Stem excluding leaves 2-3 mm thick

Lycopodium lechleri Hieron., Bot. Jahrb. Syst. 34: 57 1 . at base, tapering to ca. 1.5 mm upward. Leaves 1905. LECTOTYPE Re- (designated by Rolleri, borne in irregular alternating whorls of 67 near vista Mus. La Plata (n.s.) Bot. 13 (78): 82. 1981): the base, upward of 5-6, spreading to falcately Peru (Dept. Puno), Tabina, Lechler, ed. Hohe- ascending, linear to subulate from a slightly wid- nacker 20/2 (B!; isotypes, G!, K!, P!, UPS!). Vrostachys lechleri (Hieron.) Nessel, Barlappgewachse ened lamina base, ( 1 0-) 14-19 mm long, (0.5-)0.7- 85. 1939. 1 mm wide above the widened base, gradually Vrostachys lechleri Hieron. var. lehmannii Nessel, Re- tapering in the distal half or so, adaxially flat to vista Sudamer. Bot. 6: 61. 1940. SYNTYPES: slightly concave, or with slightly revolute margins, BONN, Herb. Nessel 148\, Ecuador, Spruce, Palla- with vein and smooth tanga 1858, marked 7; Ecuador: unknown collec- abaxially slightly prominent tor "Aus dem Herbar Bonaparte, Paris," anno- margins, with slightly widened lamina base, twist- tated "Ur. lehmannii Boge."). ing the lamina to a vertical position. Vegetative Urostachys lehmannii (Nessel) Herter, Index Lyco- leaves of basal divisions often spreading to re Hexed podiorum 67. 1949. and not twisted. Sporangia 1 .3-2 mm in diameter.

In most features resembling Huperzia hippuridea closely, but differing consistently in the nar- Upper montane forest, near timberline, on the rower, linear to nearly filiform leaves 10-23 mm forest floor, in semishade, or epiphytic, alt. 2700- long and 0.5-0.8 mm wide at the base, adaxially 3500 m, Amazonas, San Martin. usually canaliculate, with a slightly prominent vein Southern Colombia to Peru.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 25 Closely related to Huperzia hippuridea, from Cajamarca: Prov. Cutervo, Llama to Huambos, 2700 m, et al. 6591 (AAU, OH, MO). Cutervo, Raimondi which it differs by its obliquely falcate-ascending, Lopez 3126 (B, USM). Trail Socota to Tambillo, 3300 m, Stork twisted leaves and the nodding to pendulous shoot & Morton 10174 (F, K, uc). Amazonas: Prov. Chacha- arcuata also resembles H. dicho- tips. Huperzia poyas, Cerros Calla Calla, W side, 45 km above Balsas, toma (Jacq.) Trevisan (tropical America) but is 3100 m, Hutchison & Wright 5784B (uc). Cerros Calla E 1 9 km above 3 1 00 Hutch- larger in all parts and restricted to timberline forest Calla, side, Leimebamba, m, ison & Wright 5559 (F, GH, M, MO, NY, p, uc). Cano Santa habitats. The Peruvian collections generally have Lucia just E of Chachapoyas, 2000-2400 m, Wurdack short leaves, 8-15 mm long. 600 (F, GH, K, uc).

Amazonas: Chachapoyas, Matthews (E). San Martin: 5. Huperzia weddellii (Herter) Holub, Folia Geo- Prov. Mariscal Rio Abiseo National Puer- Caceres, Park, bot. Phytotax. 20: 78. 1985. ta del Monte, 3450 m, Young 1627 (AAU, USM). Prov. Mariscal Caceres, near La Playa Camp, 2700 m, Young Bot. Beibl. & Leon 4954 (AAU). Lycopodium weddellii Herter, Jahrb. 43: 98: 45. 1909. TYPE: Peru, Dept. Puno, Carabaya, 1847, Weddell 4684 (holotype, P!; isotype frag., BONN, Herb. Nessel 1701 in part). 4. Huperzia binervia (Herter) B. 011g., Opera Bot. Urostachys weddellii (Herter) Nessel, Barlappge- 92: 169. 1987. wachse91. 1939.

binervium Herter, Bot. Jahrb. 43, Lycopodium Syst. Plants erect, or erect from a decumbent base, Beibl. 98: 48. 1909. TYPE: Peru, Prov. Chacha- very robust, up to 30 cm tall, or to 50(-70) cm poyas, Matthews (holotype, P!; isotypes, BM!, GL!, branched. Shoots al- K!). long, sparsely homophyllous, Urostachys binervius (Herter) Nessel, Barlappge- most equally thick throughout, (14-) 16-2 3 mm in wachse 110. 1939. diameter including leaves. Stems excluding leaves 3-6 mm thick at the base, tapering to 34 mm in Plants to erect a ascending from decumbent base, diameter. Leaves almost uniform throughout, to 50 cm tall. Shoots up homophyllous, equally borne in alternating whorls of 6-7, ascending to thick 1 5-30 in diameter includ- throughout, mm spreading or reflexed, straight, curved upward or leaves or with shorter leaves ing gradually and recurved, lanceolate, acute to slightly acuminate to 5-8 mm. Stems leaves tapering excluding 3-6 or short-acute, 6-ll(-12) mm long, 1.5-2.5 mm thick at the sometimes to 2- mm base, tapering wide, thick and coriaceous, often glaucous, rarely 2.5 decurrent leaf bases and mar- mm, ridged by with red tips, adaxially convex with a slightly Leaves borne in close whorls gins. very alternating prominent vein, usually irregularly concave abax- of or somewhat reflexed to ascend- 6-9, spreading ially (dried), with vein abaxially obscure to dis- or recurved or ing appressed upward, slightly sig- tinctly and widely prominent, sometimes in as- linear to 7-16 moid, linear-subulate, mm long, cending leaves abaxially convex and rounded with 1.2-1.5 sometimes to mm wide, upwards reduced obscure vein, with usually smooth to minutely un- 3-7 mm not twisted at coria- long, base, thick, evenly rugose, narrowly sclerified, translucent convex and rounded or ceous, adaxially shallowly margins, or rarely papillate near the base. Leaf with a to slightly sharply prominent veinal ridge, bases prominently decurrent, with a small, some- convex and when abaxially shallowly rounded, times indistinct swelling (air sac), especially in dried with a vein and usually prominent irregular sporangiate leaves. Sporangia 1.5-2.2 mm in di- longitudinal ridges or wrinkles, or sometimes the ameter. vein sunken and hence the adjacent leaf tissue appearing as 2 longitudinal veins, with smooth mar- Terrestrial in exposed habitats in timberline for- Stomates gins, green. often forming 2 irregular, est and lower shrub paramo, 2600-3900 m, Ama- sunken slightly longitudinal bands along the vein. zonas, Puno. Sporangia 1.5-2 mm wide. Ecuador to Bolivia.

shrubland Open and grassland (jalca), seepage Amazonas: Prov. Bagua, Cordillera Colan, NE of La areas, 2000-3100 m, Cajamarca, Amazonas. Peca, ca. 3 1 70 m, Barbour 3446 (MO). Endemic. Related to Huperzia loxensis B. 011g. (southern 6. Huperzia brongniartii (Spring) Trev., Atti Soc. Ecuador) and H. hippuridea. Ital. Sci. Nat. 17:248. 1874.

26 FIELDIANA: BOTANY Lycopodium brongniartii Spring, Bull. Acad. roy. Sci. Lycopodium reflexum Lam., Encycl. 3: 653. 1789. Bruxelles 8 (2): 515. 1841. TYPE: Bolivia, Yun- TYPE: Martinique, Comm. Joseph Martin (ho- gas, D'Orbigny 227 (holotype, P!; isotypes, BONN, lotype, P, Herb. Lam. 4421). Herb. Nessel 169\ in part, BR!). Plananthus reflexus (Lam.) Beauv., Prodr. Aeth. 100. Lycopodium taxifolium Sw. var. brongniartii (Spring) 1805. Baker, Handb. Fern Allies 16. 1887. Urostachys reflexus (Lam.) Herter, Beih. Bot. Cen- Urostachys brongniartii (Spring) Nessel, Arch. Bot. tralbl. 39: 249. 1922. Est. S. Paulo 1: 388. 1927.

Plants erect or ascending from a decumbent base, soft, usually loosely caespitose, 10-30(-40) cm tall. Plants erect from an ascending base, up to 50 Shoots homophyllous, almost equally thick cm tall, sparsely branched. Shoots homophyllous, throughout, 7-1 5 mm in diameter including leaves. almost equally thick throughout, 20-30 mm in Stems excluding leaves 1 .5-3(-4) mm thick at base, diameter including leaves. Stems excluding leaves sometimes tapering to 1-1.5 mm in diameter, 2.54 mm thick near the base, sometimes tapering ridged by decurrent leaves or almost smooth. to 1.5-2 mm upward, somewhat ridged by decur- Leaves borne in alternating irregular whorls of rent leaf bases. Leaves uniform throughout, borne (6-)7-8(-9), ascending to spreading or sharply rein alternating, rather distant whorls of 4-5, wide- flexed, straight to strongly recurved, linear-subu- spreading to somewhat reflexed, usually straight, late, widest just above the base, 4-8 mm long, 0.5- not twisted at the base, lanceolate, with long-acute 1(-1.2) mm wide, softly herbaceous to subcoria- apex, papery to subcoriaceous and opaque, 9-1 2(- ceous, adaxially convex, or concave near the base, 15) mm long, 2-3 mm wide, almost flat, with abaxially flat, or slightly concave to convex, with prominent vein adaxially, or folded slightly down an obscure to somewhat prominent vein, with flat along the vein, with slightly revolute, minutely to revolute, very sparsely to densely denticulate rugose margins. Sporangia ca. 2.5 mm wide. to short-ciliolate margins. Leaf bases often somewhat decurrent. Sporangia 1-1.5 mm in diameter. Terrestrial in humid montane forest, ca. 2900- 3300 m, Cuzco. A frequent pioneer on permanently moist, dis- Colombia to Bolivia. turbed ground, mainly in the montane forest zone, Related to Huperzia rosenstockiana (Herter) also in peat bogs, or rarely as a low epiphyte. Holub with which it shares the minutely rugose Tropical America. leaf margins. The specimens cited correspond to Closely related to Huperzia eversa, H. acifolia, the Bolivian representatives of the species. and H. affinis. Huperzia reflexa is variable with respect to leaf and stem size, direction and crowding of leaves, Cuzco: Prov. Urubamba, Machu Picchu, hillside above and leaf margin characters. Part of the variation Rio Mandor, 2920 m, 1350 Alturas de Peyton (AAU). undoubtedly reflects variable growth conditions, Rio Calzada, Huadquina, 3300 m, Sues 756 (us). Prov. but often several distinct forms can be found Paucartambo, Parque Nacional Manu, Road Acjanaco very to Pilcopata, bridge between Pillahuata and La Esperan- growing intermixed in the same habitat, indicating za, 2750 m, Leon 2228 (USM); Prov. Paucartambo, Par- that genetic differences exist. The variation pat- Nacional below 2950 Leon & que Manu, Acjanaco, m, terns are complex and in need of a detailed study. Cano 2129 (USM). A variety with smaller dimensions stands somewhat apart and is recognized taxonomically. The 7. Huperzia reflexa (Lam.) Trev., Atti Soc. Ital. remaining material, referred to the type variety, is Sci. Nat. 17: 248. 1874. 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 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 Lycopodium bifidum Willd., Sp. pi. ed. 4, 5: 53. 1810. (GH). Ayna between Huanta and Rio Apurimac, 750- TYPE: Venezuela, Cuchilla de Guajana Guajana, 1000 m, Killip & Smith 23198 (NY). Cuzco: Prov. Con- Humboldt & Bonpland 474 (holotype, B, Herb. vencion, Hda. Guayanay, 1800 m, Vargas 13248 (GH). Willd. 194211; isotype, p, Herb. Humb.!). Quispicanchi, Mandor, Marcapata, 1000 m, Vargas 5220 Lycopodium reversum Presl, Reliq. haenk. 1:82. 1825. (us). TYPE: Ecuador, Guayaquil, Haenke (holotype, PRC!). Lycopodium reflexum Lam. var. majus Spring, Mem. 7b. Huperzia reflexa var. minor (Spring) B. 011g. Acad. roy. Belg. 15 [Mon. Lye. 1]: 26. 1842. in and Fl. Ecuador 33: Huperzia reflexa (Lam.) Trev. var. bifida (Willd.) Trev., Marling Andersson, Atti Soc. Ital. Sci. Nat. 17: 248. 1874. 26. 1988. Lycopodium reflexum Lam. var. densifolium Baker, Handb. Fern-Allies 11. 1887. SYNTYPES: Hart- Lycopodium reflexum Lam. var. minus Spring, Mem. weg 1480 (K!); Moritz 2266 (= 226bl K!); Brazil, Acad. roy. Belg. 15 [Mon. Lye. 1]: 26. 1842. SYN- Glaziou 15797 (K!). TYPES: Brasil, Pr. Rio de Janeiro, Gaudichaud Lycopodium densifolium (Baker) Underw. & Lloyd, (P!); Langsdorff (M, Herb. Mart.l); Brazil, Serra Bull. Torrey Bot. Club 33: 106. 1906. dos Orgaos, fr. Majo, Guillemin (P!); In sylvis Lycopodium mexiae Copel., Univ. California Publ. prov. Paraensis, Martins (M!); Brazil, in prov. Bot. 19: 294, t. 47. 1941. TYPE: Peru, Dept. Minarum, Claussen (P!). Huanuco, Churubamba, trail Cotirarda to Mer- Lycopodium brutum Herter, Bot. Jahrb. 43: Beibl. 98: cedes, 1875 m, Mexia 8193a (holotype, uc!; iso- 47. 1909. TYPE: Trinidad, Hooker ded. 1845 types, GH!, K!, MICH). (holotype, P!). Urostachys stellae-polaris Herter, Revista Sudamer. Bot. 10: 121. 1953. TYPE: Colombia, Cundina- marca, Guayabetal to Monte Redondo, SE of Humid montane forest, road banks, alt. 1000- Quetame, 1 300-1 500 m, Pennell 1801 (holotype, 2400 m, Huanuco, Cuzco, Puno. Brazil; Trinidad; us!). Ecuador to Bolivia; probably more widespread. Lycopodium stellae-polaris (Herter) Morton, Amer. This variety seems identical to Huperzia par- Fern J. 54: 72. 1964. Rolleri and Deferrari from SE Huperzia bifida (Willd.) Holub, Folia Geobot. Phy- vifolium (Nessel) totax. 20: 71. 1985. Brazil. It often grows intermixed with individuals mexiae Rolleri and No- Huperzia (Copel.) Deferrari, of var. reflexa. tas Mus. La Plata, Bot. 21 (100): 156. 1988.

Landslides, road banks, and other open or dis- Huanuco: Prov. Huanuco, Road Huanuco to Tingo N of Pass, 2350-2450 Plowman & turbed habitats in montane forest, alt. 900-3400 Maria, Carpish m, Rury 11146 (F, GH). Cuzco: Prov. Urubamba, Machu m, Cajamarca, Amazonas, San Martin, Huanuco, Picchu, Soukup 167 (F). Near Machu Picchu, 2000 m, Cuzco. Pasco, Junin, Ayacucho, Tryon & Tryon 5409 in part (GH, us, USM). Puno: Prov. Throughout humid mountainous regions of Sandia, below Cuyocuyo, 2900 m, Ferreyra 16624 (USM). tropical America. Foster & Smith 9095 (USM) is very robust and 8. Huperzia acifolia (Rolleri) Rolleri and Defer- tall, with thicker stems and slightly wider leaves rari, Notas Mus. La Plata, Bot. 21 (100): 155. than usual for this variety. 1988.

Cajamarca: Prov. Cutervo, El Pajonal, San Andres, Lycopodium acifolium Rolleri, Revista Mus. La Plata 2200 m, Llatas & Suarez 2828 in part (F). Prov. Cutervo, (n.s.) 14: 2, /. 1-2. 1985. TYPE: Peru, Dept. Aya- La Pucarilla, between Socota and San Andres, 2500 m, cucho, Cearrapa, between Huanta and Rio Apu- Sanchez Vega el al. 5923 (AAU, F). Amazonas: Prov. rimac, 1 500 m, Killip & Smith 22368 (holotype, Chachapoyas, roads to Molinopampa, 2700 m, Sanchez us!). Vega et al. 2218 (AAU). San Martin: Prov. San Martin, Dist. Tarapoto, road Tarapoto to Yurimaguas, km 9- Plants erect, or erect from a decumbent base, 1 3, ca. 700 m, Rimachi 4102 (F, GH, NY). Huanuco: Prov. Huanuco, Chinchao to Puente Duran, 1900 m, Ochoa 1 5-30 cm tall, or up to 60 cm long. Shoots homo- 14597 (F, us). Carpish Pass, 2700 m, Hodge 6294 (GH, phyllous, almost equally thick throughout, 10-20 us). Pasco: Manto at Yaupi, Woytkowski 6534 (MO, us). mm in diameter including leaves. Stems excluding Prov. Oxapampa, Ulcumanu SW of Oxapampa, road to leaves 2-5(-6) mm thick at the base, sometimes Maria Teresa and Llaupi, 2150-2450 m, Foster et al. to 1-3 thick. Leaves uniform 7688 (AAU). Junin: Tarma, Chanchamayo, above La tapering mm Merced, ca. 2000 m, Weberbauer 2006 (G, MOL). Aya- throughout, borne in alternating, usually densely

28 FIELDIANA: BOTANY 98: 48. 1909. LECTOTYPE B. crowded whorls of 8-1 1, usually sharply bent up- (designated by in Harling and Andersson, Fl. Ecuador 33: ward from a perpendicular junction to the stem 011g. 28. 1988): Ecuador, Andium nemoribus humidis, and then gently to strongly clawlike recurved, lin- Jameson 74 (P!). soft- ear-subulate, widest just above the pale and Huperzia ecuadorica (Herter) Holub, Folia Geobot. herbaceous base, 6-10 mm long, 0.7-1(-1.3) mm Phytotax. 20: 72. 1985. wide at the base, narrowed shortly above the base, subcoriaceous at apex, adaxially convex distally, Plants erect or erect from a decumbent abaxially flat to convex with obscure to somewhat base, often much-branched and prominent vein (sometimes sunken when dried), soft, large, caespitose, to cm tall. Shoots with rather densely denticulate-ciliolate margins up 30(-50) homophyllous, thick mm in at base, usually sparsely denticulate or smooth at equally throughout, (2.5-)3-6(-10) diameter leaves. Stems leaves apex. Sporangia 1-1.5 mm wide. including excluding 1.5-2.5 mm thick at the base, sometimes tapering to 1-2 decurrent leaf Moist banks in montane forest at mid-altitudes, mm, prominently ridged by bases Leaves uniform borne alt. 1350-2040 m, San Martin, Huanuco, Pasco, (dried). throughout, in subdistant to Junin, Ayacucho, Cuzco. irregular, alternating, densely Venezuela to Bolivia. crowded whorls of (4-)5(-6), wide-spreading to linear- Closely related to Huperzia rejlexa and H. un- sharply reflexed, usually strongly recurved, lanceolate, widest in the basal half, mm guiculata B. 011g. (Colombia, Ecuador). Some ma- (2.5-)3-5 0.5-1 mm herbaceous to sub- terial earlier tentatively referred to H. unguiculata long, wide, softly convex with obscure from Ecuador and Peru belongs here, and thus H. coriaceous, adaxially vein, concave with obscure unguiculata is yet unknown from Peru. abaxially irregularly (dried), to somewhat prominent vein, with slightly revolute, denticulate-ciliolate margins. Leaf base with San Martin: Prov. Pedro Ruiz Rioja, Moyobamba prominently decurrent vein and margins. Sporan- road, km 390-394, Venceremos, 1910-2040 m, D. Smith gia 1-1.5 mm in diameter. 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 Terrestrial, as a pioneer on landslides, road to Pucallpa, La Divisoria, 1600 m, J. Schunke V. 3082 banks, and other open, moist habitats in upper (F, G, GH, NY, us); Ferreyra 1690 (USM). Pasco: Prov. montaine forest, alt. 2400-3400 m, Piura, Ama- Oxapampa, Rio Boqueria, ca. 26 km from Oxapampa zonas, San Martin, Huanuco, Pasco, Ayacucho, via Rio Yamaquizu, 1870 m, D. Smith et al. 1772 (AAU, Cuzco. USM). Oxapampa [as Junin], 1600 m, Soukup 2675 (F, GH, us). Junin: Prov. Tarma, road to La Mina Pichita, Costa Rica; Andes from Venezuela to Bolivia. 9 km from the Tarma-La Merced ca. 1 3 km from road, Huperzia eversa resembles H. rejlexa, but has San Ramon, 1 600 m, Skog et al. 5055 (us). Chancha- shorter and relatively wider leaves, which are usu- mayo Valley, above La Merced at Cumbre Yacunay, ally strongly recurved, so that that shoots appear 2000 m, Hutchison 11 94A (F, M, NY, uc, us, USM). Cuzco: differs Prov. Paucartambo, Kosnipata, San Pedro, 1 350 m, Var- much more slender. It also by usually form- gas 10219 (MO, uc). ing very densely branched individuals.

9. Huperzia eversa (Poiret) B. 011g. in Harling and Piura: Prov. Huancabamba, Loma Redonda (Sapa- Andersson, Fl. Ecuador 33: 28. 1988. lache to Chinguela), 2400 m, Sagdstegui et al. 10174 (AAU, MO). Amazonas: Prov. Chachapoyas, middle east- Lycopodium reflexum Willd., Sp. pi. ed. 4, 5: 52. 1810, ern Calla-Calla slopes, 3000-3200 m, Wurdack 1770 (F, not Lam. 1789. TYPE: Ecuador: Tungurahua, GH, NY, uc, us, USM). San Martin: Prov. Mariscal Ca- (Humboldt) Nee D. D. (holotype, B, Herb. Willd. ceres, Rio Abiseo National park, NW corner, Chochos, 19419\). 3400 m, Young 3693a (AAU). Huanuco: Playapampa, Lycopodium eversum Poiret, in Lam., Encycl. 3: 556. 2700 m, Macbride 4486 (F). Cushi, trail to Tambo de 1814 [1813]. Vaca, Bryan 622 (F, us). Pasco: Prov. Oxapampa, Dist. Lycopodium reflexum Lam. var. polycarpum Sodiro, Oxapampa, Rio San Alberto, Abra Esperanza, 2400- Recens. crypt, vase. Quit. 90. 1883. TYPE: Ec- 2700 m, Foster et al. 10300 (AAU). Ayacucho: Prov. La uador, valle de Nanegal, Sodiro (not located). Mar, eastern Massif of the Cord. Central opposing the Lycopodium polycarpum (Sodiro) Underw. & Lloyd, Cord. Vilcabamba between Tambo, San Miguel, Ayna, Bull. Torrey Bot. Club 33: 105. 1906, not L. poly- and Hda. Luisiana, 2920 m, Dudley 11972 (F, GH). Cuz- carpos Kunze 1835. co: Prov. Paucartambo, Parque Nacional Manu, Camino Lycopodium ecuadoricum Herter, Bot. Jahrb. 43: Beibl. Eriksson, Leon & Huapalla 2387 (AAU). Michehuanunca,

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 29 Huadquina, 2700 m, Biies 745 (us). Department un- ignated by 011gaard in Harling and Andersson, known: Matthews 1082 (E). Fl. Ecuador 33: 37. 1988): Ecuador, Loja to Za- mora, 3500 m, 1875, "/. M. J. Mission" Quito 415 (BONN, Herb. Nessel 23 /!). 10. Huperzia affinis Trev., Atti Soc. Ital. Sci. Nat. 17: 248. 1874. Plants ascending to erect, massively robust and large caespitose plants, with almost completely Lycopodium affineGrev. & Hooker, Bot. Misc, 2: 364. homoblastic to 40 cm tall. Shoots 1831, not Bory 1804. LECTOTYPE (designated divisions, up by Nessel, Barlappgewachse 97. 1939): Ecuador, homophyllous, almost equally thick throughout, Herb. Jameson Pichincha, 1824, Greville, (K!; or slightly tapering upward and the leaves grad- possible isotypes, E!, NY!, us!). ually shorter and wider, 12-25 mm in diameter Lycopodium blepharodes Maxon, Contr. U.S. Natl. leaves near the sometimes Herb. 17: 423. 1914. including base, tapering Huperzia blepharodes (Maxon) Holub, Folia Geobot. to (7-) 10-1 5 mm. Stems excluding leaves 3-8 mm Phytotax. 20: 71. 1985. thick at the base, tapering to 3-5 mm upward, almost completely concealed by leaves. Leaves Plants erect from a decumbent base, at least up uniform throughout or upward slightly reduced, to 25 cm tall. Shoots homophyllous, equally thick densely crowded, borne in usually regular alter- throughout, 5-10 mm in diameter including leaves. nating whorls of 4-5, ascending to closely imbri- Stems excluding leaves 1.5-3 mm thick, upward cate, rarely patent to recurved in basal divisions, sometimes slightly tapering. Leaves borne in rath- straight to slightly upwardly curved, narrowly tri- er close alternating whorls of 5, almost covering angular-lanceolate near the base to widely trian- the stem, patent to loosely appressed, narrowly gular-ovate in terminal divisions, widest just above triangular-lanceolate to almost subulate, with the leaf base, 8-12 mm long and 1.54 mm wide slightly widened base, (3.5-)4-6 mm long, 1-1.5 in basal divisions, upward (4-)5-10 mm long, 2.5- mm wide, abaxially convex and evenly rounded 4 mm wide, evenly tapering, with acute, upward- or with prominent veinal ridge, at least at base, curved apex, thick, adaxially flat or slightly con- somewhat clasping the sporangia, with long, flac- cave with flush or slightly prominent vein, abax- cid marginal cilia and small teeth. Sporangia 1.5- ially rounded with slightly to strongly prominent, 2 mm in diameter. narrow vein, with a short, flattened basal swelling (air sac), shining or pruinose, green or rarely with Terrestrial on banks in upper montane forest red-tinged margins, with smooth to uneven or ru- and grassland, alt. 2700-3100 m, Amazonas, gulate, opaquely to transparently sclerified mar- Huanuco. gins. Leaf base margins usually slightly revolute, Colombia to Peru. sometimes subauriculate in apical divisions. Spo- The description above applies to the Peruvian rangia 2-2.5 mm wide. specimens. These are generally smaller, are more compact, and have more appressed and abaxially Low, wet mossy and grassy paramos, alt. 2750- more convex leaves than specimens from Colom- 3400 m, Piura, Lambayeque. Southern Ecuador bia and Ecuador. Huperzia affinis is variable with and northernmost Peru. respect to crowding and direction of the leaves. It This species is related to Huperzia macbridei is distinguished from H. reflexa var. reflexa, H. but can be distinguished from this by the homo- eversa, and H. pearcei (Baker) Holub (Bolivia) by blastic, evenly spreading branching habit, wider its wider and more convex leaf bases and the slen- leaves, and greater size. der cilia on the leaf margins. Plura: Purchased in the market of Huancabamba, Amazonas: Prov. Chachapoyas, Cerros Calla Calla, W Friedberg7631b(GH). Lambayeque: Prov. Ferranafe, Dist. side, 45 km above Balsas, 3100 m, Hutchison & Wright Incahuasi, Laguna Tembladera to Cerro Negro, 3300 m, et al. 12818 12848 in 5830 (F, GH, NY, uc, USM). Huanuco: Mito, 2700 m, Sagdstegui (AAU), (F part). Bryan 389a (F). Prov. Huanuco, Punta de Panao, As- plund 13712 (s). Panao, 2700 m, Macbride 3602 (F). 12. Huperzia weberbaueri (Nessel) Holub, Folia

1 1 . Bot. Huperzia kuesteri (Nessel) B. 011g., Opera Geobot. Phytotax. 20: 78. 1985. 92: 169. 1987.

Urostachys weberbaueri Nessel, Revista Sudamer. Bot. Urostachys kuesteri Nessel, Repert. Spec. Nov. Regni 6: 162, /. 10, f. 41. 1940. TYPE: Peru, Dept. Veg. 35: 182, t. 172. 1934. LECTOTYPE (des- Amazonas, Ostlich Chachapoyas, Tambo Ven-

30 FIELDIANA: BOTANY tillas, 2400-2600 m, Rosen 672 (holotype, BONN, patentes usque ad ascendentes vel perpendiculares usque Herb. Nessel 1831). ad parum reflexa, recta vel incurva, lanceolata, aequate Lycopodium papillatum Rolled, Amer. Fern J. 65: 3. angustata vel acuminata, (3-)4-6(-7) mm longa, 1.5-2 1975. TYPE: Peru, Dept. Amazonas, Prov. Cha- mm lata, parum decurrentes, abaxialiter convexa, ad chapoyas, Cerro Malcabal (Cerro Tumbe), 3-6 basin vix vel parum ventricosa, plerumque angulo me- km SW of Molinopampa, 2900 m, Wurdack 1456 diali subacuto instructa, adaxialiter plana usque ad con- (holotype, us!; isotypes, GH!, K!, uc!, USM!). cava (siccata), subcoriacea, hypostomatica, plerumque Huperzia papillata (Rolleri) Holub, Folia Geobot. polita, marginibus laevibus, scleroideis et translucidis. Phytotax. 20: 75. 1985. Sporangia ca. 1.5-2 mm lata.

Plants erect from an ascending base, large, caes- Plants erect from an ascending base, loosely pitose, homoblastic, up to 30(-40) cm tall. Shoots caespitose, homoblastic, up to 30 cm tall. Shoots homophyllous, 10-15(-20) mm in diameter in- homophyllous, 6-10(-12) mm in diameter includ- cluding leaves, sometimes tapering to 7 mm in ing leaves, equally thick throughout or slightly ta- diameter, strikingly whitish papillate in life. Steins pering. Stems, excluding leaves, 2-4 mm thick. excluding leaves 24 mm thick. Leaves uniform Leaves uniform throughout, or slightly smaller up- throughout, or slightly smaller upward, borne in ward, borne in more or less regular, alternating more or less regular, alternating whorls of 5, patent whorls of 4-5, patent to ascending or perpendic- to ascending, upward curved, lanceolate, 7-10(- ular to slightly reflexed in basal divisions, straight

15) mm long, 2-2.5 mm wide, not or slightly de- to upward curved, lanceolate, widest ca. '/4 of the current, without a prominent basal swelling (air leaf length above the base, evenly tapering in the sac), abaxially convex, with an obscure veinal ridge, distal % or slightly acuminate, straight to slightly adaxially flat to concave (dried), densely long-pap- upward curved, (3-)4-6(-7) mm long, 1.5-2 mm illate on margins and abaxially, smooth adaxially. wide, somewhat decurrent, without or with a Sporangia ca. 1.5 mm wide. slightly prominent basal swelling (air sac), abaxially convex, rounded or usually with a rather sharp Wet grassy and boggy paramos with some shel- medial ridge, adaxially flat to concave (dried), ter, and lower puna 3000-3600 m, Lambayeque, somewhat coriaceous, usually glossy on both sides, Amazonas, San Martin, Cuzco. the margins smooth, sclerified and somewhat Southernmost Ecuador and Peru. translucent. Sporangia ca. 1.5-2 mm wide. The densely papillate, bright whitish leaves in the living plants are unique in the genus. The whit- TYPE Peru, Dept. Amazonas, Prov. Bagua, ish color is often lost after drying, especially heat Cord. Colan NE of La Peca, 10,800 ft, 8 Sep 1978, drying. Barbour 3384 (holotype, MO!; isotypes, AAU!, USM!). Andean grassland, open patches in shrub para- Lambayeque: Prov. Ferranafe, Dist. Incahuasi, La- mo, alt. 3140-3600 m, Amazonas, San Martin. guna Tembladera, 3 1 50 m, Sagdstegui et al. 12790 (AAU). Endemic. Amazonas: Prov. Leimebamba to Calla Chachapoyas, The specimen cited from San Martin deviates Calla, 3100 m, Sanchez V. 5JO (AAU); Boeke 1809 (AAU). from the other material a softer texture and less NE of Tambo de Ventilla, Cerro de Fraijaco (Huaui to by Huni), 3450 m, Pennell 15883 (GH). Cerros Calla Calla, glossy leaves with less sclerified margins and is 3100 m, Hutchison & Wright 5564 (F, GH, MO, NY, uc, included in the species with some doubt. us). San Martin: Prov. Mariscal NW corner of Caceres, Huperzia colanensis appears to be intermediate Rio Abiseo National Park, 3400 m, Young & Leon 4768 in position between the Huperzia brevifolia and (AAU). Prov. Mariscal Caceres, Chochos Valley, 3300 m, H. saururus of It Young & Leon 4877 (AAU). Huallaga, valley of Apison- groups 011gaard (1987, 1989). cho, 30 km above Jucusbamba, 3600 m, Hamilton & does not appear to be intimately related to other 1216, 1217 Cuzco: Prov. Holligan (us). Paucartambo, Peruvian species. Cord, de Tres Cruces, 3600 m, Vargas 12195 (GH). Prov. Paucartambo, Pavayoc, 2100 m, Woytkowski 555 (MOL, USM). 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 13. colanensis B. nov. Huperzia 011g., sp. (MO), 3198 (AAU). San Martin: Dist. Huallaga, Valley of Rio Apisoncho, 3600 m, Hamilton & Holligan 1218 Planta erecta vel ascendens; surculi homoblastici us- (us). que ad 30 cm alti, homophylli 6-10(-12) mm diametro foliis inclusis, omnino crassitie aequali vel parum an- 14. B. in gustati. Caulis 2-4 mm crassus foliis exclusis. Folia om- Huperzia polylepidetorum 011g. Harling nino uniformes vel sursum parum minores, 8-10-faria, and Andersson, Fl. Ecuador 33: 42,/ 6B. 1 988.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 31 TYPE: Ecuador, Prov. Azuay, Laegaard ing shoots, up to ca. 35 cm tall. Shoots homo- 55117 (holotype, AAU!; isotype, QCA!). blastic, homophyllous to gradually slightly heterophyllous, 12-20 mm in diameter at the base Plants ascending to erect, large, robust, caes- including leaves, usually tapering to 5-10 mm. pitose, nearly homoblastic, up to 35 cm tall, or Stems excluding leaves 2-3 mm thick at the base, more than 50 cm long. Shoots homophyllous, often brown to reddish brown, terete to promi- equally thick throughout, or slightly tapering up- nently ridged by decurrent leaf bases. Leaves of ward and the leaves gradually shorter, 10-25 mm basal divisions (and shaded shoots) borne in ir- in diameter including leaves. Stems excluding regular, rather distant, alternating whorls of 5-6, leaves 5-7 mm thick at the base, sometimes ta- usually patent to ascending, not covering the stem, pering to 2 mm thick, usually not completely con- linear-lanceolate, 7-10 mm long, 1.5-2 mm wide, cealed by leaves. Leaves uniform throughout, or adaxially flat, shining, abaxially flat to slightly con- gradually shorter upward, well spaced to densely vex, dull to shining green, usually with long-de- crowded, borne in more or less regular, alternating current leaf bases. Leaves of distal divisions grad- whorls of 5-6, patent-ascending to arcuate-ap- ually shorter, closer, and more appressed, borne pressed, strongly curved upward and inward, lan- in alternating whorls of 4-5(-6), ascending to ar- ceolate to widely lanceolate, acute or short-acute, cuate-appressed, straight to strongly upward or 6-12 mm long, (2-)2.5-3(-3.5) mm wide, some- unilaterally curved or twisted, lanceolate to broad- what fleshy, abaxially convex or rarely flattened, ly lanceolate, often slightly cuspidate above the with an obscure to prominent adaxial and abaxial, middle, (5-)6-8 mm long, 1.5-2 mm wide, adax- long decurrent median ridge, with an indistinct air ially concave (dried) or flat to shallowly rounded sac, adaxially concave or flat, with smooth, slightly in life, abaxially convex, rounded and irregularly involute margins, green throughout or with red- wrinkled (dried), with slightly prominent, short to tinged tips. Sporangia 2.5-3.5 mm wide. long, narrowly decurrent basal swelling (air sac), with smooth to densely irregularly rugulate, slight- In open Polylepis forest among mossy rocks, ly to distinctly, translucently sclerified margins, 3900-4300 m, Ancash. shining to dull green. Sporangia ca. 2 mm wide. Ecuador and Peru. The Peruvian collection was made in shaded Grass paramos, usually growing in partial shade Polylepis woods, and the specimen differs slightly among grasses, 3600 m, Cajamarca, Cuzco. from Ecuadorean plants by the more patent-as- Andes from Venezuela to Peru. cending leaves, which tend to be abaxially flat to Known from two collections in Peru, matching slightly concave in the basal divisions, while the the Ecuadorean plants perfectly. distal, more exposed divisions have abaxially convex leaves. It thus seems close to Huperzia wed- Cajamarca: Prov. Chota, Laguna Yahuarcocha, above dellii. Incahuasi, 3600 m, Sagdstegui et al. 12908 (F). Cuzco: Prov. Paucartambo, Parque Nacional Manu, vicinity of El Mirador, 3600 m, Leon 2270 (USM). Ancash: Prov. Huari, Huascaran National Park, Que- brada de Yuraccocha, a lateral valley of Quebrada Ruri- 16. macbridei B. chinchay, 3900-4300 m, D. Smith et al. 12716 (AAU, F). Huperzia (Herter) 011g., Opera Bot. 92: 169. 1987.

1 5. Huperzia capellae (Herter) Holub, Folia Geo- Urostachys macbridei Herter, Revista Sudamer. Bot. bot. Phytotax. 20: 71. 1985. 10: 115. 1953. TYPE: Peru, Dept. Huanuco, 3-

6 mi NW of Mito, 1 1,000 ft, Macbride & Feath- Revista Sudamer. Hot. 10: Urostachys capellae Herter, erstone 1922 (holotype, us!; isotypes, B!, F!, MA!, 114-115. 1953. LECTOTYPE (designated by s!). Roller!, Revista Mus. La Plata, n.s., Bot. 13 (71): Lycopodium macbridei (Herter) Morton, Amer. Fem 78. 1981): Ecuador, prov. Napo [as Imbabura], J. 54: 72. 1964. E of Volcan de Cayambe, along trail between Rio Boqueron and Rio Arturo, 1 1,000 ft, Drew E 3 14 Plants erect from a decumbent base, loosely to (us!; isotype, MSC!). Lycopodium capellae (Herter) Morton, Amer. Fem J. densely caespitose, with basal, prostrate-ascend- 54: 72. 1964. ing, rejuvenating shoots at the periphery, up to 32 cm tall. Shoots homophyllous or with leaves grad- Plants ascending to erect, sparsely to densely ually slightly reduced upward, equally thick branched, without prostrate-ascending, rejuvenat- throughout or slightly tapering upward, 1 5-20 mm

32 FIELDIANA: BOTANY in diameter including leaves near the base, often Plants erect, or erect from a prostrate to as- tapering to 6-9 mm in diameter. Stems excluding cending base, sometimes loosely caespitose, with leaves 24 mm thick at the base, upward tapering basal, prostrate-ascending, rejuvenating shoots at to ca. 2 mm, almost completely concealed by the periphery, and erect, fingerlike shoots in the leaves. Leaves uniform throughout, or upward center, at least up to 35 cm tall. Erect shoots homo- slightly reduced, borne in irregular, alternating phyllous or almost so, equally thick throughout, whorls of 4-5(-6), densely crowded, spreading to 10-15 mm in diameter including leaves, or some- ascending in basal divisions, upward often grad- times tapering to ca. 8 mm upward. Stems ex- ually becoming closely imbricate, straight or slightly cluding leaves 3-5 mm thick at the base, some- upward curved, narrowly lanceolate to lanceolate, times tapering to 2 mm, usually completely widest near the base, evenly tapering, 10-15 mm concealed by leaves. Leaves uniform throughout, long and 1.5-2.5 mm wide in basal divisions, up- or slightly reduced upward, borne in alternating ward 6-11 mm long, 1.5-2 mm wide, without or whorls of 4-7, in distal fully sporangiate divisions with a slightly pronounced basal swelling (air sac), usually 45, crowded, loosely to closely imbricate, adaxially concave with flush or slightly prominent slightly upward curved and usually with a strongly vein, abaxially rounded, with slightly prominent upward curved apex, lanceolate to widely lanceo- and often slightly darker and long decurrent vein, late, slightly long-acuminate from a wide and with smooth to slightly uneven, somewhat invo- somewhat rounded base, 8-12 mm long, (1.8-)2- lute, indistinctly sclerified margins, green to yel- 2.5(-3) mm wide, upward sometimes reduced to lowish green. Sporangia ca. 2.5 mm wide. 7 mm long, with an inconspicuous, narrow, long- decurrent basal swelling (air sac), adaxially con- Terrestrial in low shrub paramos, at the edge of cave to slightly convex at the base, abaxially con- woods, 2500-3600 m, Amazonas, La Libertad, vex and rounded, with a prominent, often darker Huanuco, Ayacucho, Cuzco. colored (dried) ridge along the vein, smooth and Southern Ecuador and Peru. often shining, with smooth, slightly sclerified mar-

Individuals of shaded or sheltered habitats (e.g., gins, green, not or scarcely pruinose. Sporangia 2- Macbride 3487) tend to retain juvenile leaf mor- 3 mm wide. phology, being completely homophyllous with relatively wide and long, patent-ascending rather than Boggy paramo and jalca, humid rocks, 2700- appressed and gradually reduced leaves and often 3600 m, Huanuco, Pasco, Cuzco. with a more loosely caespitose growth habit. Mat- Peru and Bolivia. thews 963 is a mixed collection, sheets of the same The information on the type label is in Nessel's number at G pertaining to H. hippuridea. handwriting, and the geographic information is as dubious as on numerous other Nessel labels. Spec- Amazonas: Chachapoyas (presumably), Matthews 963 imens in Geneva, "Peruvia, Herb. Pavon" (G!), Libertad: Prov. Nevado del (BM). La Bolivar, Cajamar- are likely to be isotypes. See Comments under Hu- ilia, L] u puna, Ferreyra 1350 (USM). San Martin: Prov. perzia polyclada. 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 Huanuco: Cushi, trail to Tambo de Vaca, Bryan 677 Chochos, 3450 m, Leon 1870 (AAU, USM). Huanuco: Mito, (F, us). Playapampa, ca. 2700 m, Macbride 4475 (F, us). Pasco: Prov. trail to summit of Cord. Yana- 2700 m, Bryan 381 (F). Cani, 7 mi NE of Mito, 2500 Oxapampa, via Rio San 3 1 50-3300 D. Smith 7710 m, Macbride 3487 (F, us). Ayacucho: Prov. La Mar, east- chaga Daniel, m, ern Massif of the Cord. Central opposing the Cord. Vil- (AAU, USM). Cuzco: Cerro Puncuyoc, 4500 m, Biles 563 3400- cabamba between Tambo, San Miguel, Ayna and Hda. (us). Prov. Urubamba, near Wenner Gren ruins, 3600 30745 in Trail to Luisiana, 3200-3500 m, Dudley 11989 (F, GH). Cuzco: m, Metcalf part (uc). Puyupata Paucartambo, Tres Cruces, 3600 m, Bikes 2214 (MO). Sayaccmarca, 3600 m, Vargas 2910 (us). Department Pearce Prov. Paucartambo, Parque Nacional Manu, vicinity of unknown: Perou (1839-1840), Gay (p). Peru, (us El Mirador, 3600 m, Leon 2269 (USM). 1431555).

17. Huperzia darwiniana (Nessel) B. 011g., comb, 18. Huperzia crassa (Willd.) Rothm., Feddes Re- nov. pert. Spec. Nov. Regni Veg. 54: 60. 1944.

Urostachys darwinianus Nessel [Barlappgewachse 80, ed. 5: 50. 1810. /. 3,f. 20. 1939, nom. nud.], Revista Sudamer. Lycopodium crassum Willd., Sp. pi. 4, Bot. 6: 161. 1940. TYPE: "Columbien: Andinum TYPE: Ecuador, Antisana, Humboldt & Bon- montibus nemoribus, Ruiz" (holotype, BONN, pland 2263 (holotype, B,Herb. mild. 194171; iso- Herb. Nessel 1331). types, BM!, P!).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 33 Urostachys crassus (Willd.) Nessel, Barlappgewachse um. The type and the Weberbauer collection both 75. 1939. have creeping basal shoots and are rather densely Urostachys pilgerianus Nessel, Revista Sudamer. Bot. tufted. The leaves are twisted unilaterally, but oth- 6: 161, t. 9,f. 32. 1940. TYPE: "Peru, Cordillere, " erwise this corresponds to most of the Peruvian Rautenstock et Mann, in 1 90 1 (holotype, BONN, Herb. Nessel 124\). material. Huperzia pilgeriana (Nessel) Holub, Folia Geobot. 20: 75. 1985. Phytotax. La Libertad: Prov. Santiago de Chuco, Jalca de Ques- nada (Quiruvilca to Huamachuco), 4000 m, Sagdstegui Plants erect from a prostrate to ascending base, & Fabris 7574 (MO, NY). Prov. Santiago de Chuco, Dist. Calchicadan, Escalerilla-Conzuzo road near Tamboras, slightly to strongly heteroblastic, loosely to densely 3960 m, Saunders 882 (F, GH). Ancash: Prov. Carhuaz, caespitose with prostrate-ascending or shallowly Huascaran National Park, Quebrada Ishinca, 4380-4500 basal often nu- subterranean shoots, these bearing m, D. Smith 9480 (AUU). Prov. Carhuaz, Huascaran Na- merous erect, fingerlike shoots in the center, at tional Park, Quebrada Honda, 4300-4750 m, D. Smith et al. 11643 Cord. above toward Le- least up to 40 cm tall. Erect shoots homophyllous (F). Blanca, Vicos, jiacocha, 3600 m, Hutchison & Wright 4323 (F, GH, NY, or almost so, equally thick throughout, or slightly uc). Prov. Huaylas, Huascaran National Park, Quebrada tapering upward, 5-10 mm in diameter including Alpamayo, 4750 m, D. Smith et al. 9715 (AUU, F). Junin: leaves. Stems excluding leaves 2-3 mm thick, usu- Mount La Juntay, near Huancayo, 4700 m, Killip & ally completely concealed by leaves. Leaves uni- Smith 22114 (F, NY, us). Cuzco: Cerro Salamanca, Valle Lucumayo, 2200 m, Biies 569 (us). Prov. Urubamba, form throughout, or slightly reduced upward, borne Abra de Malaga, 4300 m, Chavez 2822 (MO); 4 1 50-4230 in irregular, alternating whorls (or low spirals) of m, Molau & Ohman 1639 (GB). Puno: "Cord, jugis pr. imbricate, 5-7, densely crowded, usually closely Tabina," Lechler 2043 (E). Cord, near Agapata and Sa- straight (or slightly upward curved in shaded di- chapata, Lechler 2028 (BR, G, s). visions), linear-lanceolate to lanceolate, 5-9 mm long, (1-) 1.2-2 mm wide, upward sometimes reduced to 4.5 mm long, with a prominent, short to 1 9. Huperzia nesselii (Nessel) Roller! & Deferrari, long decurrent basal swelling (air sac) causing a NotasMus.LaPlata,Bot.21 (100): 156. 1988. perpendicular appearance of the very leaf base in sporangiate leaves, adaxially concave to slightly Urostachys nesselii Nessel, Revista Sudamer. Bot. 6: convex, with a raised veinal ridge near the base, 161. 1940 [Barlappgewachse 7 8, / 12. 1939, nom. convex and rounded or with a abaxially prominent inval.]. TYPE: "Ost Peru, Cuzco, 1868, Rein- veinal ridge, slightly to strongly rugose by pro- hardt" (holotype, BONN, Herb. Nessel 132, in part!). truding, blisterlike epidermal cells, with smooth to rugose margins, green to brick red or dark red, Plants erect from a prostrate to ascending base, usually strongly pruinose. Sporangia 1-2 mm wide. slightly to strongly heteroblastic, loosely to densely caespitose with prostrate-ascending basal shoots, Terrestrial in high paramo and superparamo, these bearing numerous erect, fingerlike shoots in puna, 3600-4850 m, La Libertad, Ancash, Junin, the center, up to 20 cm tall. Erect shoots homo- Cuzco, Puno. phyllous or almost so, equally thick throughout, As here delimited, Huperzia crassa is a widely 10-15 mm in diameter including leaves. Stems distributed polymorphic species, occurring from excluding leaves 2-6 mm thick, usually completely Mexico to Panama, Hispaniola, and in the Andes concealed by leaves. Leaves uniform throughout, from Venezuela to Bolivia. borne in irregular, alternating whorls (or low spi- The Peruvian material represents a disjunct rals) of 5-6, densely crowded, closely imbricate, population, widely separated from the Ecuadorian straight to irregularly curved upward or twisted at populations, and it may be distinct at least at the the apex, linear-lanceolate to lanceolate or nar- variety level. Therefore, the synonyms included rowly elliptic, 9-12 mm long, 2-2.5 mm wide, for the type variety in Ollgaard (1 988) are not cited without a prominent basal swelling, adaxially con- here. cave to slightly convex, with a raised veinal ridge The holotype of Urostachys pilgerianus most near the base, abaxially flat especially at the base likely is a mislabeled duplicate of Weberbauer 6626 to rounded especially at the apex, with a broadly (F, GH, MOL, us) from Dept. Junin, Nevado Runa- prominent medial ridge especially at the base, tullu, SE of Jauja, on rocks, 4400-4500 m, as this slightly rugose because of protruding, blisterlike collection matches it exactly. Mislabeled (or re- epidermal cells, with smooth to slightly uneven labeled?) specimens abound in Nessel's herbari- margins, pruinose. Sporangia 2-3 mm wide.

34 FIELDIANA: BOTANY High Andine grassland near the timber line, protruding, blisterlike epidermal cells, with smooth 3000-3500 m, Junin. to irregularly rugose, often strongly sclerified mar- Endemic. gins, green to reddish-tinged. Sporangia 2-2.5 mm The material referred to this species including wide. the type seems to represent a single gathering, re- ferable to Weberbauer. The information on the In boggy grassland, open turfy puna, wet ground type label is in Nessel's handwriting, and the geo- at lakes, alt. (3300-)4000-4700(-5000) m, Caja- graphic information is dubious as are numerous marca, La Libertad, Ancash, Junin, Huancavelica, other Nessel labels. Cuzco, Puno. The species seems most closely related to Hu- Peru and Bolivia. perzia saururus with respect to size and growth Some collections, especially from lower alti- habit; however, it differs by the open growth of tudes, are referred to this species with doubt. Leon basal portions of the plant, and the ridged leaf & Young 1566, 1621 (AAU), from San Martin, Rio bases, and smooth margins. From Peruvian H. Abiseo National Park, alt. 3650 m, and Leon & crassa it differs by the large, rather flat leaves with- Young 1105 (USM), from La Libertad, Prov. Pataz, out a prominent basal air sac. Rio Abiseo National Park, Cueva de Manachaqui, 3600-3800 m, are smaller, more slender, and en- Junin: Tarma, mountains W of Huacapistana, 3400- tirely green but share the growth habit with rela- 3500 Weberbauer 2222 ex herb. m, (G, MOL). "Lagasca tively large leaves on the creeping shoots and rath- ex Tarma Provincia in n. 2222" Swartz, Peruvia, (BONN, er slender erect shoots. Herb. Nessel 132 in part).

Cajamarca: Prov. Contumaza, Jalca El Chuno (Pozo Chuno), 4500 m, Sagdstegui et al. 9380 (AAU, F). La 20. Huperzia andina (Rosenst.) Holub, Folia Libertad: Prov. Santiago de Chuco, Laguna La Victoria to 4000 et al. 6180 Geobot. Phytotax. 20: 70. 1985. (road Conzuzo), m, Sagdstegui (GH, MO). Huillilas, N ofCachicadan, 4000 m, Stork & Norton 10003 (F, uc). Ancash: Prov. Yungay, Dist. Yungay, Lake Lycopodium andinum Rosenst., Repert. Spec. Nov. Llanganuco, ca. 3500 m, Sounders 503 (F). Junin: Prov. 5: 239. not Herter 1909. TYPE: Regni Veg. 1908, Huancayo, Dist. Huancayo, Huaytapallana, Huancayo- Bolivia, La Paz, Murusata, 5000 m, Buchtien 173 Pariahuanca road, ca. 4480 m. Sounders 1151 (GH). Dist. (holotype, s!). Huancayo, ca. 28 km E of Huancayo, 4500 m, Tyron & Urostachys andinus (Rosenst.) Nessel, Barlappge- Tryon 5469 in part (F, GH, us). Huancavelica: Prov. Ta- wachse 76. 1939. yacaja, Millpu, puna de Tocas, between Colcabamba and Paucarbamba, 4000 m, Tovar 1945 (USM). Marcapata, Plants with prostrate, rooting shoots from which 3600 m, Stafford 1007 (F, K). Puno: Carabaya, Aricoma Lake, 4400 m, Stafford 1118 (K). 1 to several, usually short, erect, fingerlike shoots arise, sometimes loosely caespitose, up to 10(-15) cm tall. Prostrate shoots densely covered with 8- 21. Huperzia saururus (Lam.) Trev., Atti Soc. Ital. 15 mm long, unilaterally upward curved, linear, Sci. Nat. 17: 249. 1874. red-tinged leaves. Erect shoots homophyllous or almost so, equally thick throughout or slightly ta- Lycopodium saururus Lam., Encycl. 3: 653. 1789. ca. 6-10 mm in diameter pering upward, including TYPE: He de Bourbon (Reunion), Commerson leaves. Steins of erect shoots leaves 2- excluding (holotype, P, Herb. Law.!). 3 mm thick, usually completely concealed by Lycopodium elongatum Sw., Syn. Fil. 175. 1806. TYPE: Prov. collector un- leaves. Leaves of erect shoots uniform throughout, Peru, Junin, Tarma, known (holotype, s!). or somewhat reduced upward, borne in close, ir- Urostachys saururus (Lam.) Herter, Repert. Spec. Nov. whorls of regular, alternating 4-5, usually closely Regni Veg. 19: 162. 1923. imbricate, straight to unilaterally or irregularly Urostachys elongatus (Sw.) Herter, Index Lye. 60. 1 949. twisted and often somewhat deformed, linear-lan- Lycopodium sanctae-barbarae Rolleri, Darwiniana 16: 129,/ 1 D-E, t. 1, 3B. 1970. TYPE: Argentina, ceolate to lanceolate or triangular-lanceolate, Prov. Jujuy, de la Sola 2883 (holotype, LP). evenly to abruptly tapering from the slightly wid- Huperzia sanctae-barbarae (Rolleri) Rolleri and De- ened base or slightly acuminate, 5-8(-l 1) mm long, ferrari, Notas. Mus. La Plata, Bot. 21 (100): 156. l-2(-2.5) mm wide, upward sometimes reduced 1988. to 4-6 mm long, with a prominent basal swelling (air sac), adaxially concave to canaliculate apical- Plants erect, very compactly caespitose with

ly, abaxially convex and rounded, smooth without basal, prostrate, rejuvenating shoots from which

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 35 1506 Ancash: Prov. Huascaran stiffly erect, sparsely branched, fingerlike shoots m, Lopez (us). Huari, National Park, Quebrada Rima Rima, a lateral valley of arise, at least up to 50 cm tall. Erect shoots homo- Quebrada Carhuazcancha, 4200-4440 m, D. Smith et phyllous or almost so, usually narrow and some- al. 1231 1 (AAU, F). Prov. Yungay, Huascaran National etiolated at the base and what gradually becoming Park, Quebrada Ranicuray, 4000-4300 m, D. Smith et wider and green upward, 5-10 mm in diameter al. 9142 (AAU, F). Pasco: Pasco, Huayllay, ex herb. Cruck- in 1830 Junin: San ca. 3960 Mac- including leaves at the base, upward usually 8-15 shanks (GH). Jose, m, bride & Featherstone 1 1 1 1 (F, G, us). Huancavelica: Prov. mm. Steins excluding leaves (2-)3-5 mm thick, Tayacaja, above Hda. Tocas, between Colcabamba and concealed leaves at usually completely by except Paucarbamba, 3200 m, Tovar 1967 (USM). Cuzco: Neva- the etiolated base. Leaves uniform throughout or do Sallcantay, 3900-4200 m, Biies 744 (us). Prov. Uru- gradually shorter upward in old shoots, often bamba, Dist Chinchero, Cuper, Hatun Wayk'o quebrada, 3300 m, Sallo ex Franquemont 281 (AAU, F). Prov. smaller, narrower, pale and irregularly appressed Paucartambo, Pfuyucalla?, Huilcacunca, 4000 m, Var- at the etiolated base of shoots, borne in irregular, gas 9840 in part (MO). Madre de Dios: Pinasniocj, Pan- whorls low of alternating (or spirals) 6-8, densely tiacolla, Pass, 3600 m, Cook & Gilbert 1867 (us). De- crowded, closely appressed, the sporophylls ap- partment unknown: Peru, Rusby 354A (us). pressed from a nearly perpendicular leaf base, straight or somewhat secund, linear to linear-lan- 22. Huperzia hypogaea B. 011g. in Harling and ceolate, the uppermost sometimes lanceolate to Andersson, Fl. Ecuador 33: 58. 1988. TYPE: widely lanceolate, 10-12(-15) mm long, l-2(-3) Ecuador, Prov. Carchi, 0llgaard & Balslev mm wide, upward sometimes reduced to 5-8 mm 8517 (holotype, AAU!; isotype, QCA!). long, sporophylls with a prominent basal swelling (air sac), adaxially concave to slightly convex, Plants with short or up to 36 cm long, horizon- abaxially convex and rounded throughout or at tal, usually subterranean, isotomously branching least at the apex, smooth, with minutely rugose, shoots, from which stiffly erect, aerial shoot sys- thickened, sclerified margins, especially in the bas- tems arise. Subterranean divisions with pale ap- al leaves, green and shining to reddish-tinged or pressed leaves, 2-4 mm thick including leaves. rusty-colored and sometimes strongly pruinose. Aerial shoots up to 40 cm tall including erect sub- Sporangia 2-2.5 mm wide. terranean divisions, stiffly erect, homophyllous, equally thick throughout, or slightly tapering, 4- Moist shrubland and grassland (jalca), often 6(-8) mm thick including leaves. Stems of aerial among rocks, alt. 33004500 m, Cajamarca, La shoots 1.5-3 mm thick excluding leaves, partly to Libertad, Ancash, Pasco, Junin, Huancavelica, completely concealed by leaves, brownish to bright Cuzco. red, with smooth to somewhat rugose epidermis. Peru to Argentina and Chile; Kerguelen; Tristan Leaves borne in alternating, irregular whorls of 4-

da Cunha; alpine regions of tropical and southern 5, arcuate-appressed to closely appressed, abaxi- Africa; Madagascar; Reunion; Mauritius. ally convex and rounded or with a prominent, long Contrary to the view of Rolleri (Amer. Fern J. decurrent veinal ridge, with or without a slightly 67: 109-120. 1977), the present author finds the prominent basal swelling (air sac), adaxially con- Neotropical distribution of Huperzia saururus re- cave to slightly convex, linear-lanceolate to lan- stricted to Peru and more southern regions in ceolate, 4-6(-7) mm long, l-1.8(-2) mm wide, America. The Central American and north Andine upward often reduced to 3.5-4 mm long, red-tinged material referred to H. saururus by Rolleri belongs to entirely red, with smooth to unevenly rugose to various different species. and sclerified margins. Sporangia 1-1.3(-1.5) mm Some plants of this species are distinctly prui- wide. nose and reddish-tinged, similar to Huperzia crassa, but can usually be distinguished by the leaf Wet and boggy paramo and jalca, alt. 3000- shape, the etiolated character of the base of the 4300 m, Lambayeque. erect shoots, and the lack of blisterlike, protruding Colombia to northern Peru. epidermis cells on the abaxial leaf surface. Usually a very easily recognizable species of marshes and wet depressions ofthe lower paramos Cajamarca: Contumaza, Jalca El Chuno, 4500 m, Sa- with a soft substrate where the subterranean run- et al. 9559 Pozo 3790 gdstegui (AAU, F, NY). Kuan, m, ner-shoots are well developed. Where the species Sagdstegui et al. 13089 (AAU, F). Pozo Kuan, Laguna el grows on more solid substrates, e.g., open soil in Toro, 4100 m, Sagdstegui 9452 (F). La Libertad: Prov. landslides and road creeping Santiago de Chuco, Pampas de la Julia, 3600 m, Sagds- cuts, supraterranean tegui et al. 11123 (AAU, GH). Jalca de Quiruvilca, 4200 shoots with well-developed, patent-ascending

36 FIELDIANA: BOTANY leaves replace the runner shoots, and make the but the trailing stems are usually on or just below distinction from slender forms of Huperzia crassa the surface of the ground, and the leaves are short- more subtle. In such cases the difference of leaf er, strongly pruinose, with many blisterlike cells curvature, leaf size, and the number of leaf or- on the abaxial leaf epidermis, and have densely thostichies are helpful characters. denticulate-fimbriate margins. Huperzia attenuata is rather similar but is less distinctly hetero- Lambayeque: Prov. Ferranafe, Dist. Incahuasi, La- blastic, has a smooth abaxial leaf epidermis, and guna Tembladera to Cerro Negro, 3300 m, Sagdstegui has a more sharply prominent basal abaxial air et al. 12847 (AAU, F, GH). sac on the upper leaves, and the leaves are borne in whorls of three to four. 23. Huperzia sagasteguiana B. 011g., sp. nov.

La Libertad: Prov. Pataz, between Retama and La Paz, et H. atten- Species cum habitu Huperziae hypogaeae 3950 m, Lopez & Sagdstegui 3590 (GH). uatae. A H. hypogaea differt surculis horizontalibus epi- gaeis vel non profunde hypogaeis, foliis brevioribus, abaxiali marginibus denticulato-fimbriatis, epidermide 24. Huperzia attenuata (Spring) Trev., Atti Soc. rugulosa cellulis protrusis pustuliformibus. A H. atten- Ital. Sci. Nat. 17: 249. 1874. uata differt foliis 1 0-fariis, epidermide abaxiali foliorum rugulosa, basibus foliorum minus ventricosis. Lycopodium attenuatum Spring, Mem. Acad. roy. Belg. 24 [Mon. Lye. 2]: 8. 1849. LECTOTYPE (des- Plants with up to 20 cm long, horizontal, epi- ignated by Lellinger, Proc. Biol. Soc. Wash. 89: terranean and or subterranean, creeping, shallowly 717. 1977): Ecuador (Prov. Pichincha), in declivi- isotomously branching shoots, rooting along the tate montis Pichincha, Hartweg 1470 (us!; isolec- underside, bearing stiffly erect, up to 20 cm tall totypes, BM!, GL!, K!, NY!, P!). attenuatus aerial shoot systems with short to long intervals Urostachys (Spring) Nessel, Barlappge- wachse 101. 1939. between branches. Creeping shoots with loosely appressed to upward curved, secund leaves, 3-5 Plants ascending to erect, rather small, caespi- mm thick including leaves. Aerial shoots stiffly tose with erect shoots in the center, and prostrate- erect, homophyllous, terete, equally thick through- ascending, rejuvenating basal shoots in the pe- out, 4-6 mm thick including leaves. Stems ofaerial riphery, up to 20 cm tall, or up to 30 cm long. shoots 1.5-3 mm thick excluding leaves, partly to Shoots homophyllous to slightly gradually hetero- completely concealed by leaves, brownish to bright phyllous, terete to hexagonal, equally thick red, with rugosely striate epidermis. Leaves borne throughout, 4-7 mm in diameter including leaves. in alternating, irregular whorls of 5, arcuate-ap- Steins excluding leaves 1 .5-2 mm thick at the base, pressed to closely appressed, abaxially strongly sometimes tapering to 1-1.5 mm in diameter up- convex and rounded to apically indistinctly cari- ward, completely concealed by leaves, tinged with nate, with a prominent, short to long-decurrent red or brick-red. Leaves borne in alternating whorls veinal ridge, with a somewhat prominent basal of 34, forming 6 or 8 usually regular ranks, ap- swelling (air sac), adaxially concave (dried), lan- pressed to the stem throughout, in basal divisions ceolate to triangularly ovate-lanceolate, 3-5 mm lanceolate and convex abaxially, upward gradually long, 1.5-2 mm wide, red-tinged to entirely red, changing to narrowly triangular-ovate with acute abaxially wrinkled (dried) and strongly uneven due or slightly acuminate apex, abaxially strongly con- to the protruding blisterlike epidermal cells, with vex to subcarinate near the apex, with a prominent numerous irregularly shaped and directed, soft and basal swelling (air sac), (4-)5-6 mm long, in basal pale marginal processes. Sporangia 1-1.5 mm wide. divisions 1-1.5 mm wide, distally 1.5-2.2 mm wide, with fimbriate margins, tinged with red. Spo- TYPE Peru, Dept. La Libertad, Prov. Pataz, rangia 1.5-2 mm wide. Paso de Alaska, carretera a Tayabamba, en ladera abierta de Gramineas, Jalca, 3900 m, Lopez & Exposed cushion vegetation in paramo, pioneer Sagdstegui 8177 (holotype, MO; isotypes, AAU!, GH!, vegetation on landslides, road banks, etc., in the NY!). paramo zone, alt. 3400-3500 m, Cajamarca, La Terrestrial in jalca vegetation, 3900-3950 m, La Libertad, San Martin. Libertad. Costa Rica; Ecuador and Peru to Bolivia. Endemic. An easily recognizable species, characterized by The growth habit resembles Huperzia hypogaea, fimbriate leaf margins. Huperzia attenuata is a

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 37 close relative ofH. tetragona (see the following for Exposed cushion vegetation, pioneer vegetation discussion). on landslides, road banks, etc., in the lower part of paramos and jalca, 2750-3600 m, Cajamarca, C'ajamarca: Prov. Cajamarca, N ofCanal Cumbe Mayo, La Libertad, San Martin, Huanuco, Cuzco. 3400-3500 m, Sanchez. V. & Molau 3746 (AAU). La Columbia to Bolivia. Libertad: Prov. Huamachuco, road to Pallar-Huaguil, The red or reddish, quadrangular, erect shoots Tayabamba, 3400 m, Lopez & Sagdstegui 8148 (AAU, F, make this species an easily recognizable one. In- MO). San Martin: Prov. Mariscal Caceres, Rio Abiseo National Park, NW sector, Valle de Chochos, 3450 m, complete material of some epiphytic species with Leon 1872 (AAU, USM). Prov. Mariscal Caceres, Rio Abi- quadrangular constricted shoots have been fre- seo National de 3450 Leon & Park, Pampa Cuy, m, quently misidentified as this species. They are Young 1401A (USM). readily separated because of the fimbriate leaf margins in Huperzia tetragona. 25. Huperzia tetragona (Hooker & Grev.) Trev., Huperzia tetragona is closely related to H. at- Atti Soc. Ital. Sci. Nat. 17: 248. 1874. tenuata. The occasional aberrant hexagonal or terete shoots in some specimens of//, tetragona are Lycopodium tetragonum Hooker & Grev., Icon. Fil. virtually indistinguishable from slender shoots of 1: t. 109. 1829. TYPE: Ecuador (Prov. Pichin- H. attenuata. The two are also cha), Prope verticem montis Pichincha, Jameson species ecologically (E!, K!). rather similar and are often found to grow inter- Lycopodium catharticum Hooker, Ann. Nat. Hist. 1 : mixed. There are numerous examples of species 1 /. 1 430-^3 , 14. 838. SYNTYPES: Ecuador (Prov. in which the number of leaves in each whorl is Azuay), "Asuay of the Equator," W. Turner (K!); variable in the same population. The difference of Ecuador (Prov. Canar), from the mountains of terete in Pillzhum, Jameson (K!). and quadrangular shoots Huperzia atten- Lycopodium myrsinites Lam. var. minus Spring, Mem. uata and H. tetragona may be due to such simple Acad. 15 29. based roy. Belg. [Mon. Lye. 1]: 1842, and taxonomically overrated variation within on Lycopodium catharticum Hooker and with the populations of one species. same type. Lycopodium tetragonum Hooker & Grev. var. patulum Spring, Mem. Acad. roy. Belg. 24 [Mon. Lye. Cajamarca: Prov. Chota, Laguna Yahuarcocha, above 2]: 12. 1849, based on Lycopodium catharticum Incahuasi, 3600 m, Sagdstegui et al. 12901 (AAU). Prov. Hooker and with the same type. Cajamarca, N of canal Cumbe Mayo, 3400-3500 m, Urostachys tetragonus (Hooker & Grev.) Nessel, Bar- Sanchez Vega & Molau 3747 (AAU). La Libertad: Pataz, lappgewachse 135. 1939. Puerta del Monte, ruta de Huaylillas, 3200 m, Lopez & Urostachys catharticus (Hooker) Nessel, Barlappge- Sagdstegui 3450 (GH). San Martin: Prov. Mariscal Ca- wachse 135. 1939. 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- Plants ascending to erect, rather small, caespi- bride & Featherstone 1923 (F, s, us). Playapampa, ca. tose with erect shoots in the center, and usually 2750 m, Macbride 4890 (F, o, us). Cuzco: Paucartambo, prostrate to ascending rejuvenating basal shoots Huillcacunca to Huaisampilla, 3800 m, Vargas 9859 in the periphery, up to 20(-30) cm tall. Shoots (MO, uc). homophyllous, or gradually slightly heterophyllous, quadrangular and equally thick throughout, 26. Huperzia sellifolia B. 011g. in Harling and An- 2.5-3.5(-4) mm in diameter including leaves. dersson, Fl. Ecuador 33: 68, / 12C. 1988. Steins excluding leaves 1-2 mm thick. Leaves dec- TYPE: Ecuador, Prov. Carchi, B. 0llg. & Bal- ussate throughout, imbricate, concealing the stem, slev 8432 (holotype, AAU!). in basal divisions widely lanceolate to triangular- ovate, abaxially rounded and evenly decurrent, Plants erect or ascending, usually forming large, upward triangular, carinate at least in the upper caespitose, homoblastic plants, often more than

half, somewhat decurrent or abaxially with a 25 cm tall, with up to 45 cm long shoots. Shoots prominent basal swelling (air sac), 3.5-5(-7) mm homophyllous, almost equally thick throughout, long, 2-4 mm wide at the base, evenly tapering (5-)6-8(-10) mm in diameter including leaves. into an acute or slightly acuminate apex, orange Stems excluding leaves (2.5-)34 mm thick, al- to bright red-tinged at the margins throughout, most completely concealed by leaf bases, except with fimbriate margins. An occasional branch may in basal divisions. Leaves almost uniform be hexagonal or terete, with leaves borne in alter- throughout, usually densely crowded throughout, nating whorls of 3. Sporangia 1.3-2.5 mm in di- borne in rather regular alternating whorls of 4-5, ameter. loosely imbricate or ascending to patent or sharply

38 FIELDIANA: BOTANY reflexed and appressed to the stem, usually dis- Trev. (Ecuador) and H. brevifolia but apparently tinctly sigmoid, widely lanceolate to ovate or el- adapted to more sheltered habitats. The sporangia liptic, acute or obtuse, (1.8-)2-2.6(-3.5) mm long, are disproportionately large, thick, and volumi- (1.2-)1.5-2(-2.4) mm wide, coriaceous, usually nous and do not open as is otherwise usual in dried brightly red-tinged, abaxially convex to concave specimens. This, plus the rarity of the species, sug- with prominent vein in basal part of lamina, gests hybridity, but there is no other evidence of rounded in the apex, with prominent, short to long- hybridity, because the spores seem normally de- decurrent basal swelling (air sac), with usually veloped. The collection by Jelski from the Nessel slightly involute, narrowly sclerified, darker, Herbarium needs to be confirmed; it was not found smooth to slightly uneven margins, adaxially with in Krakow. slightly to distinctly prominent vein. Sporangia 1- 1.5 mm in diameter. Department unknown: "Ost-Cordilere, Jelskr [possibly Dept. Cajamarca, Prov. Cutervo] (BONN, Herb. Nes- sel 1731). Terrestrial in exposed wet paramos, alt. 3 1 50- 3650 m, Piura, Lambayeque, Amazonas, San Martin. 28. Huperzia brevifolia (Grev. & Hooker) Holub, Andes of southern Colombia to northern Peru. Folia Geobot. Phytotax. 20: 71. 1985.

Piura: Purchased in the market of Sullana, Friedberg Lycopodium brevifolium Grev. & Hooker, Hooker Bot. 3933c (GH). Lambayeque: Prov. Ferranafe, Dist. Inca- Misc. 3: 104. 1832. TYPE: Peru, in Herb. Lam- huasi, Laguna Tembladera, 3150 m, Sagdstegui et al. bert, Ruiz and Pavon (BM?; possible isotypes, FI, 12799 (AAU, F). Amazonas: Prov. Bagua, Cord. Colan Herb. Webb; AAU photo ex FI!, o!). NE of La Peca, ca. 3300 m, Barbour 3382 (AAU, MO, Urostachys rufescens (Hooker) Nessel var. brevifolius USM). San Martin: Prov. Mariscal Caceres, Rio Abiseo (Grev. & Hooker) Nessel, Barlappgewachse 103. National 3650 Leon & 1564 Park, Paredones, m, Young 1939. (AAU). Urostachys brevifolius (Grev. & Hooker) Herter, Index Lye. 54. 1949. 27. Huperzia engleri (Herter) B. 011g., Opera Bot. 92: 169. 1987. Plants erect or ascending, stiff and robust, forming small to large, caespitose, homoblastic plants, Lycopodium engleri Herter, Bot. Jahrb. Syst. 43, Beibl. often more than 25 cm tall. Shoots homophyllous, 98: 45. 1909. TYPE: Peru, Dept. Huanuco, Prov. almost equally thick throughout, (5-)7-12 mm in Huamalies, mountains SW of Monzon, 3200- diameter including leaves. Stems excluding leaves 3300 m, Weberbauer 3359 (holotype, B!; isotype, BONN, Herb. Nessel 17 3\, MOL!). (3-)4-7 mm thick at the base, slightly tapering Urostachys engleri (Herter) Nessel, Barlappgewachse upward, usually concealed by leaf bases. Leaves 94. 1939. almost uniform throughout, densely crowded, or sometimes more spaced in basal divisions, borne Plants erect or to 1 cm tall. Shoots ascending, up in alternating whorls of 4 5, ascending or perpen- thick 8-10 mm homophyllous, equally throughout, dicular to sharply reflexed, with straight to upward in diameter leaves. Stems including excluding curved apex, widely ovate or triangular-ovate to leaves 1-1.5 Leaves mm thick, densely papillate. widely suborbicular-cordate or triangular-cordate, well spaced at base, crowded upward, borne in acute to almost obtuse or mucronulate, 2-4 mm whorls of 4, to somewhat re- alternating patent long, 2-4(-5) mm wide, stiffly coriaceous, green- lanceolate to acute flexed, widely oblong-lanceolate, or red-tinged, abaxially concave, with sharply to 4-5.5 1.5-2 obtuse, mm long, mm wide, slightly prominent vein, with sharply prominent, some- convex with with abaxially slightly prominent vein, what flattened, short-decurrent basal swelling (air decurrent without a basal evenly base, swelling, sac), with slightly revolute, strongly sclerified, with a shallow the adaxially groove along vein, darker and somewhat translucent, slightly erose with 2-2.5 minutely rugose margins. Sporangia margins. Sporangia 1.5-2.5 mm wide. mm wide, nearly globose.

Habitat information according to protologue: Terrestrial in wet paramos, jalca, alt. 2700-4 1 20 Bog at grass steppe interrupting the shrub for- m, Piura, Lambayeque, Cajamarca, Amazonas, mation. Huanuco, Pasco, Cuzco. Endemic, Huanuco. Costa Rica; Colombia to Peru. Possibly related to Huperzia rufescens (Hooker) The total distribution range is uncertain, as there

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 39 is considerable regionally characteristic variation divisions patent to loosely imbricate, triangular- in the material referred to Huperzia brevifolia. ovate to ovate or widely ovate, acute, 3-5 mm Populations from Colombia, southern Ecuador and long, 2-3.5 mm wide, softly to firmly coriaceous, Peru are slightly different and usually recognizable; green- to red-tinged, adaxially concave, or slightly perhaps they are worth taxonomic recognition. convex and with a prominent veinal ridge at the Huperzia brevifolia differs from H. rufescens and base, abaxially usually slightly concave at the base, H. sellifolia by its larger size, thick stems, and stiff above the base convex and rounded or with a and almost prickly leaves and by the frequent lack slightly to strongly prominent veinal ridge, with a of red coloration. strongly prominent, narrow basal swelling (air sac), Dudley 11109 deviates by its slender shoots with with upward curved apex, with usually strongly very short leaves. sclerified, irregularly rugose to minutely erose margins. Sporangia 1.7-2 mm wide. Piura: Purchased in the market of Sullana, Friedberg Prov. Dist. Inca- 3933e (GH). Lambayeque: Ferranafe, Terrestrial on banks, steep, mossy slopes in wet huasi, Laguna Tembladera, 3150 m, Sagdstegui 12792 paramos, jalca, alt. 3400-4000 m, Cuzco, Puno. (F, GH), 12848 (GH in part). Cajamarca: From Chiclayo Colombia to Peru. market, Leon (AAU, USM). Amazonas: Prov. Chachapo- hohenackeri resembles the yas, Leimebamba-Chilchos trail, Boeke 2130 (AAU). Prov. Huperzia most large Bagua, Cord. Colan NE of La Peca, 3300 m, Barbour individuals of H. brevifolia, with which it shares 3381 Huanuco: 2700 Mac- (AAU, MO). Playapampa, m, the general aspect of size and growth habit. It dif- bride 4476 (F, us). Cushi, trail to Tambo de Vaca, Bryan fers in the longer, more acute and usually ap- 678 (F, us). Pasco: Prov. Oxapampa, trail to summit of leaves. 30745 deviates Cord. Yanachaga via Rio San Daniel, 3350-3620 m, D. pressed Metcalf by having Smith 7742 (AAU, USM). Junin: Prov. Tarma, mountains recurved leaves throughout. W of Huacapistana, 3400-3500 m, Weberbauer 2225 (MOL). Cuzco: Prov. Paucartambo, Paso del Gallo, 2900 Cuzco: Prov. Urubamba, trail Piajupata to Sayacmar- m, Vargas 4279 in part (uc, us). Urubamba, Machu ca, 3600 m, Vargas 2911 (us). Near Wenner Gren ruins, Picchu, Runcuraccay-Sayamarca pass, 4120 m, Peyton 3400-3600 m, Metcalf30745 (uc in part). Prov. La Con- 288 (AAU). Prov. La Convention, summit ridge of Vil- vencion, Alturas de Rio Calzada, Huadquina, ca. 3660 cabamba, ca. km 28 along trail from Hda. Luisiana and m, Bties 755 (us). Puno: Tabina, in summis Cordilleras, Rio Apurimac, 3330-3410 m, Dudley 11109 (GH, us). julio 1854, Hohenacker (BONN, Herb. Nessel 199). Cor- dillera near Tabina, Hohenacker 4204 (BONN, Herb. Nes- 29. Huperzia hohenackeri (Herter) Holub, Folia sel 199). Cord, near Ayapata and Sachapata, Lechler 2028 unknown: Lan- Geobot. Phytotax. 20: 73. 1985. (E). Department Cordillera, 1902, doni (BONN, Herb. Nessel 199).

Lycopodium hohenackeri Herter, Bot. Jahrb. 43: Beibl. 30. Atti. 98: 46. 1 909. TYPE: Peru, Dept. Puno, in summis Huperzia hartwegiana (Spring) Trev., Cordiller. jugis pr. Tabina, Lechler 2043 (B!, BR!, Soc. Ital. Sci. Nat. 17: 248. 1874.

K, NY!, p, s!). Urostachys hohenackeri (Herter) Nessel, Barlappge- Lycopodium hartwegianum Spring, Mem. Acad. roy. wachse 105. 1939. Belg. 24 [Mon. Lye. 2]: 14. 1849. LECTOTYPE (designated by Ollgaard in Harling and Anders- Plants erect or ascending, forming small to large, son, Fl. Ecuador 33: 76. 1988): Colombia, Prov. 1466 annotated caespitose, homoblastic plants, up to 40 cm tall. Popoyan, Hartweg (K! by Spring; isolectotypes, BM!, G!, LG!, NY!). Shoots homophyllous or with gradually smaller Lycopodium caracasicum Herter, Hedwigia 49: 88, /. leaves thick upward, equally throughout, 10-13(- 3a. 1909. TYPE: Venezuela, Caracas, Jan. 1856,

1 5) mm in diameter including leaves, or tapering Gollmer (holotype, B!). to (5-)7-10 mm in diameter upward. Stems ex- Urostachys hartwegianus (Spring) Nessel, Barlappge- wachse 90. 1939. cluding leaves 5-6 mm thick at the base, upward Urostachys caracasicus (Herter) Nessel, Barlappge- to 4 to tapering (2-)3 mm thick, partly completely wachse 90. 1939. leaves. in concealed by Leaves borne more or less Huperzia caracasica (Herter) Holub, Folia Geobot. regular, alternating low spirals or whorls of 5-6, Phytotax 20: 71. 1985. often well spaced in basal divisions, densely crowded upward, those of basal non-sporangiate Plants robust, recurved or pendulous, at least divisions ascending to spreading, widely triangu- up to 30 cm long. Shoots homophyllous to grad- lar-lanceolate or ovate-lanceolate, 4.5-5.5 mm ually heterophyllous, 15-30 mm in diameter in- long, 2.5-3 mm wide, with short to long, evenly cluding leaves in basal divisions, sometimes ta- decurrent leaf bases. Leaves of upper, sporangiate pering to 10-15 mm in diameter. Stems excluding

40 FIELDIANA: BOTANY leaves 2.5-3.5(-5) mm thick at the base, tapering Lycopodium nitens Schlecht. & Cham., Linnaea 5: to 1.5-2 mm. Leaves of basal divisions borne in 623. 1830. TYPE: Mexico, in arboris vetustis prope Jalapam, Schiede & Deppe (holotype, B!; often regular alternating whorls of 4-5, falcately isotype, BM!). to directed to patent-ascending imbricate, radially Huperzia passerinoides (Kunth) Trev., Atti Soc. Ital. somewhat secund, almost completely covering the Sci. Nat. 17: 248. 1874. stem and sporangia, lanceolate, coriaceous, usu- Huperzia passerinoides (Kunth) Trev. var. nitens (Schlecht. & Cham.) Trev., Atti Soc. Ital. Sci. Nat. ally shining, 14-23 mm long, 2.5-3.5 mm wide, 17: 248. 1874. widest below the middle to near the base, abaxially Lycopodium schwendeneri Herter, Bot. Jahrb. 43: Beibl. convex at the base, usually slightly concave toward 98: 50. 1909. TYPE: Not designated. Mexico in- the tip, with smooth, flat to somewhat involute dicated as locus classicus by Herter 1923. Urostachys taxifolius (Sw.) Herter, Repert. Spec. Nov. margins, the vein slightly prominent at the base Regni Veg. 19: 162. 1923. and apically obscure. Leaves of terminal divisions Phlegmariurus taxifolius (Sw.) Love & Love, Taxon conform, or usually smaller, 7-10(-13) mm long, 26: 324. 1977. 1 .5-3 mm wide, often somewhat clasping with the base widened, softer and less shining. Sporangia Plants lax and pendent, or sometimes recurved 1.7-3 in diameter. mm from an erect and somewhat rigid stem base, up to 50(-70) cm long. Shoots usually gradually tain alt. 2400-3100 Rupestral montane forest, m, pering from ca. 20-30 mm in diameter including Ancash, Cuzco. leaves at the base, to 4-10 mm in distal divisions Southern Andes from Ven- Mexico; Guatemala; of fully developed plants, sometimes not, or only ezuela to Peru. slightly tapering, rarely abruptly constricted. Stems Huperzia hartwegiana is widely distributed and excluding leaves 1.5-2 mm thick at the base, ta- is outside variable. The species often epiphytic pering to 1-1.5 mm. Leaves usually reduced and but in Peru all records are from sub- Peru, rocky modified upward, borne in irregular alternating strates. The variation seems partly correlated with whorls of 3-5. Leaves of basal divisions spreading in habitat type. The species frequently occurs sites to ascending or somewhat appressed, often twisted that are too for the other In dry Huperzia species. at the base, narrowly lanceolate, widest in the low- such the exposed rupestral sites, plants initially er half, firmly herbaceous to subcoriaceous, 14- then and rather grow erect, recurved, may develop 23 mm long, 2-3 mm wide, almost flat or some- short and shoots Such forms were rec- rigid only. what concave adaxially, with flat or slightly rev- as caracasicum Herter and ognized Lycopodium by olute, smooth margins, with evident to somewhat others. In situations moist, shaded, epiphytic long, prominent vein abaxially. Leaves of middle and are lax, pendulous forms developed. Huperzia terminal divisions usually gradually shorter, nar- is related to H. hartwegiana taxifolia. rower and more appressed, abaxially more convex, with involute margins. Leaves of fully sporangiate Ancash: Prov. E of Cerrate Bolognesi, Quero Huasta, divisions often distinctly 6-( 1 0-) ranked, with wid- 2471 (USM). Cuzco: Prov. Cuzco, Cadena de Pactatayan, ened, clasping base, partly covering the sporangia, Vargas 17044 (GH). Prov. Urubamba, Ollantaytambo, often contracted into a short to long, nar- 3050 m, Plowman & Davis 4741 (GH). Cusuchaca, 2400 abruptly m, Vargas 22971 (GH). Valle de Urubamba, Herrera row, involute apex, 3-8 mm long, l-1.5(-2) mm 3423 (c). Valle de Urubamba, ca. 1 km from the village, wide. Sporangia 1-1.5 mm wide. Leon 446 (USM). Quebrada Las Penas, 3100 m, Chavez 3493 (MO). Usually epiphytic in lower to mid-altitude wet montane forest, alt. 1300-1400 m, Cajamarca, Amazonas. 31. Huperzia taxifolia (Sw.) Trev., Atti Soc. Ital. Central America; West Indies; northern tropical Sci. Nat. 17: 248. 1874. South America. Huperzia taxifolia is a variable species, the de- Lycopodium taxifolium Sw., Prodr. 138.1788. TYPE: limitation of which is problematic, especially out- A remounted specimen, marked as type, without original annotation, in the Regnellian type her- side Peru. The Peruvian specimens cited below barium in s!, is possibly the holotype. are unusual in having the leaves arranged in al- in Nov. Lycopodium passerinoides Kunth, HBK., gen. ternating whorls of 5. Hutchison & Bismarck 6379, sp. 1: 41. 1816. TYPE: Peru, prope Olleras et unusual in the nearly conform sporophylls and Aipate, alt. 747 hexp., Humboldt (holotype, P, leaves is Herb. Humb.l; isotypes, B, Herb. Willd. 194101, uniformly patent-ascending throughout, BONN, Herb. Nessel 160\). apparently juvenile.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 41 Cajamarca: Prov. Hualgayoc, Hda. Taulis, between San Martin: Prov. Mariscal Caceres, Rio Abiseo Na- the Casa Hacienda and Palmito, 2000 m, Hutchison & tional Park, Puerta del Monte, 3400 m, Leon & Young Bismarck 6379 (uc, USM). Amazonas: Prov. Bongara, 1303 (AAU, USM). Prov. Mariscal Caceres, Rio Abiseo vicinity of Campamiento Ingenio 1-3 km up road to National Park, Laguna de Chochos, 3300 m, Young & Pomacocha (and Rioja) from camp along the Rio In- Leon 4855 (AAU). genio, 1300-1400 m, Hutchison & Wright 3825 (F, GH, uc). 33. Huperzia funiformis (Spring) Trev., Atti Soc. Ital. Sci. Nat. 17: 248. 1874. 32. Huperzia rosenstockiana (Herter) Holub, Fo- lia Geobot. Phytotax. 20: 76. 1985. Lycopodium funiforme Spring, Bull. Acad. roy. Sci. Bruxelles 8 (2): 516. 1841. LECTOTYPE (des- Lycopodium rosenstockianum Herter, Hedwigia 49: ignated by Spring, Mon. Lye. 2: 49. 1949): Gua- 90. 1909. TYPE: Ecuador, Ost-Abhang des Tun- deloupe, L'Herminier (P!; isolectotypes, BR!, GH!, gurahua, 3000 m, Rimbach 130 (holotype, Herb. NY!, uc!). Rosenstock, not located; isotypes, s!, uc!, us!). Urostachysfuniformis (Spring) Herter in Urban, Symb. Urostachys rosenstockianus (Herter) Nessel, Bar- Antill. 9: 387. 1925. lappgewachse 174. 1939.

Plants robust, flaccidly pendulous, ropelike, at Plants recurved from an lax, pendulous, rarely least up to 250 cm long. Shoots homophyllous, ascending base, at least up to 1 50 cm long. Shoots almost equally thick throughout, 5-10(-15) mm or with smaller homophyllous, gradually slightly in diameter including leaves, or sometimes taper- leaves in diameter in- upward, 20-25(-30) mm ing from a thicker base with patent-ascending the leaves in basal sometimes cluding divisions, leaves. Stems excluding leaves 1 .5-3(-5) mm thick to 10-14 Stems leaves tapering mm. excluding at the base, ridged by decurrent leaf bases. Leaves thick at the to 1- (1 .5-)2-3(-5) mm base, tapering almost uniform throughout, or slightly shorter up- 2 mm. Leaves of basal divisions borne in alter- ward, borne in crowded alternating whorls of 7- whorls of nating irregular 3-5, irregularly spread- 8, densely covering the stem, usually closely fal- to somewhat twisted at the ing ascending, usually cate-appressed throughout, not twisted, linear-su- with to vertical lanceolate base, oblique lamina, bulate, widest just above the base, 6-10(-12) mm to widest at or below the narrowly oblong, middle, long, 1-1.5 mm wide, evenly tapering into a long with cuneate to somewhat rounded widely base, pungent apex, firmly herbaceous to coriaceous, dull with short to acute 10-15 2- long apex, mm long, to shining, abaxially strongly convex, often api- 4 subcoriaceous and to herba- mm wide, opaque cally conduplicate, with smooth margins. Sporan- ceous and almost with translucent, flat, prominent gia 1-1.5 mm wide. vein adaxially, or sometimes folded down along the with vein, slightly revolute, minutely rugose Lower montane rain forest, up to alt. ca. 1 300 margins with individually protruding epidermis m, Huanuco. at the Leaves of middle and cells, especially apex. West Indies; southern Mexico to Panama; Guy- terminal divisions conform, or gradually smaller ana; Venezuela to Peru. and more to more ascending loosely appressed, Huperzia funiformis is a very distinct species, crowded. Leaves of densely densely sporangiate not easily confused with other species due to its divisions widest the lan- usually just above base, smooth ropelike shoots. ceolate to narrowly triangular, with a rounded base, 6-9 mm 1.5-2.5 long, mm wide, partly covering Huanuco: SW slope of the Rio Llullapichis watershed, the sporangia. Sporangia 1.5-2 mm wide. on the ascent of Cerros del Sira, 1290 m, Wolfe 12330 (GH).

Usually epiphytic, rarely terrestrial, in cloud forest and elfin forest above 3000 m, San Martin. 34. Huperzia buesii (Herter) B. 011g., comb. nov. Colombia to northern Peru. buesii Revista Sudamer. Bot. 10: Huperzia rosenstockiana resembles H. taxifolia, Urostachys Herter, 126. 1954 [as 1953]. TYPE: Peru, Tranca de Ya- but details of the leaf margin, color, and ecology racuaya, 1800 m, Bues 742 (holotype, us!). are quite distinct. Closely related to H. brong- Lycopodium buesii (Herter) Morton, Amer. Fern J. niartii with which it shares the rugulate leaf mar- 54: 72. 1964. gins. Huperzia socratis (Herter) Rolleri & Defer- rari (type from Colombia) seems to be a synonym Plants robust, pendent, up to 50 cm long. Shoots of the present species. homophyllous, almost equally thick throughout,

42 FIELDIANA: BOTANY ca. 10-20 mm in diameter including leaves, or Plants pendulous, usually with flaccidly hanging tapering to 10 mm. Steins excluding leaves 2-3 divisions, the terminal divisions often aggregated mm thick, prominently ridged by decurrent leaf in fasciculate clusters, up to 60(-80) cm long. Shoots bases. Leaves almost uniform throughout, or homophyllous or gradually heterophyllous, equal- slightly shorter distally, borne in close alternating ly thick throughout, 20-30(-45) mm in diameter whorls of 4-5, densely covering the stem, usually including leaves, or gradually tapering to (7-) 1 0- closely appressed throughout, slightly recurved at 15 mm in diameter in terminal, densely sporan- the apex, not twisted, linear to linear-lanceolate, giate divisions. Stems excluding leaves 0.5-1 mm widest just above the slightly rounded base, 12- thick at the base, slightly tapering upward, almost 19 mm long, 1-2 mm wide, evenly tapering into straight to somewhat flexuous, pale greenish in life. a long apex, firmly herbaceous to subcoriaceous, Leaves of basal divisions spirally arranged, single, shining, abaxially convex, somewhat involute, with or in occasional pairs or whorls of 3, not predom- smooth margins. Leaves of sporangiate divisions inantly whorled, forming about 6 indistinct lon- sometimes reduced to 6-8 mm. Sporangia 1.5-2 gitudinal ranks, subdistant, soft-herbaceous, wide- mm wide. spreading to ascending, straight to slightly falcate, usually with the lamina vertical due to a twist of A rare endemic. Epiphytic in forest, 1800 m, the lamina base, linear-lanceolate to lanceolate,

Cuzco. widest in the basal '/3 or '/4, distinctly narrowed Perhaps related to Huperzia funiformis and H. into a petiolelike, twisted, usually perpendicular aristei (Nessel) Rolleri and Deferrari (?Colombia) to deflexed lamina base, (10-) 13-24 mm long, 1- with which it shares the high number of leaves in 5 mm wide, flat, or with slightly revolute, smooth each whorl and the general leaf shape. However, margins. Leaves of middle and terminal divisions the leaves are longer and more diverging. The sim- spirally arranged, paired or borne in irregular to ilarity to the juvenile plants of H. sotae (Rolleri) regular, alternating whorls of 3, conform, or usu- Rolleri and Deferrari (NW Argentina to Bolivia, ally narrower, (5-) 10- 15 (-20) mm long, (1-)1.5- e.g., Venturi 2955, isotype in us!) is close. 2(-3) mm wide, often with a long narrow apex from a subauriculate, non-twisted lamina base.

Cuzco: Prov. La Convention, Choquellohuanca, 1 800 Sporangia 1-1.5 mm wide. m, Marin 2247 (F). Southern Mexico to Panama; West Indies; 35. Huperzia linifolia (L.) Trev., Atti Soc. Ital. northern South America. Sci. Nat. 17: 248. 1874. Huperzia linifolia is a widespread and polymorphic species. Its variation is especially com- 1 100. 1753. TYPE: Lycopodium linifolium L., Sp. pi. plex in the northern Andes. The two varieties keyed Plumier, Traite foug. Amer. /. 166, f. C, from an out below are fairly well characterized morpho- unspecified locality, either Martinique or His- and are well defined. paniola (see Proctor, Fl. Lesser Antilles 2: 26. logically geographically 1977). Urostachys linifolius (L.). Herter, Repert. Spec. Nov. Regni Veg. 19: 154. 1923.

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-stramineous, 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. 36. Huperzia wilsonii (Underw. & Lloyd) B. 011g., 011g. in Harling and Andersson, Fl. Ecuador Opera Bot. 92: 170. 1987. 33: 90. 1988. Lycopodium wilsonii Underw. & Lloyd, Bull. Torrey Bot. Club 33: 111. 1906. TYPE: Puerto Urostachys linifolius (L.) Herter var. tenuifolius Nes- Rico, Wilson 271 sel, Revista Sudamer. Bot. 6: 164. 1940 [Bar- Luquillo Mountains, (holotype, NY!). trichodendron Bot. lappgewachse 162, Abb. 39, / 14. 1939, nom. Lycopodium Herter, Jahrb. 43: Beibl. 98: 49. 1909. SYNTYPES: nud.]. LECTOTYPE (designated by 011gaard, in Guadeloupe, L'Herminier 103 Harling and Andersson, Fl. Ecuador 33: 90. 1 988): (P!); Bory (P!). Peru, Prov. Huanuco, Pampayacu, Poeppig Lycopodium andinum Herter, Bot. Jahrb. 43, Beibl. 98: 49. not ["Moritz 210 und 1155"] (BONN, Herb. Nessel 1909, Rosenst. LECTOTYPE (des- in & Fl. 38 1\; isotype, G!). ignated by 011gaard Harling Andersson, Ecuador 33: 97. 1988): Ecuador, Eraser, in Herb. Drake (P!; isolectotype, G!). Wet forests of the eastern Andine alt. slopes, Lycopodium lindavianum Herter, Hedwigia 49: 90. 100-2700 m, Cajamarca, Amazonas, San Martin, 1909, nom. nov. for L. andinum Herter, not Ro- Huanuco, Pasco, Junin, Cuzco. senst., and with the same type. stamineum Smithsonian Colombia?, Ecuador and Peru. Lycopodium Maxon, Misc. Coll. 56 (29): 2, pi. 2. 1912. TYPE: Panama, Chi- riqui, above El Boquete, ca. 1 750 m, Maxon 5636 Jelski 1022 Cajamarca: Tabillo, (KRA). Huancabamba, (holotype, us!). Andre 763 Jelski 1022 Kl (F, GH, NY, us). Tambillo, (KRA). Urostachys wilsonii (Underw. & Lloyd) Herter, Re- Amazonas: Prov. 5 km N of end of Lake Po- Bongara, pert. Spec. Nov. Regni Veg. 19: 163. 1923. macocha, on road to Rioja, 2000 m, Hutchison & Wright Lycopodium arcanum Maxon in Yuncker, Field. Mus. 6791 Prov. Cerros Calla (F, GH, NY, uc). Chachapoyas, Publ. Bot. 17: 310, t. 3. 1938. TYPE: Honduras, 10 km above 2700 Hutchison Calla, Leimebamba, m, & Comayagua, above El Achote, above plains of Bennett 4743 San Martin: (F, GH, M, MO, NY, uc, us). Siguatepeque, 1800 m, Yuncker et al. 6149 (ho- monte 4627 Prov. Rio- Tarapoto, Campana, Spruce (BR). lotype, us!). Pedro ja, Ruiz-Moyobamba road, km 390, Venceremos, Huperzia lindaviana (Herter) Holub, Folia Geobot. 1750 D. Smith 4372 Prov. Pedro Ruiz- m, (AAU). Rioja, Phytotax. 20: 74. 1985. 1 1 et al. Moyobamba road, km 29 , 850 m, Gentry 45502 (AAU). Pasco: Prov. Oxapampa, Quebrada San Alberto, Plants to on the borders of Parque Nacional Yanachaga-Chemi- erect, arcuate-spreading pendulous, up llen, Leon et al. 944 (F, USM). Junin: La Merced, Chan- to 20(-30) cm long. Shoots homophyllous, 15- chamayo, Soukup 1007 (F); 1000 m, Esposto 11978 (USM). 2 5 (-30) mm in diameter including leaves, equally Cuzco: Prov. La Convention, Potrero, Sapan Sachayoc, thick throughout, or in some slender individuals 2300 m, Vargas 13643 (GH). Valley of Santa Ana, 1000- gradually tapering to 11.5 cm. Stems excluding 1 500 m, Herrera 2628 (us). Department unknown: Peru, leaves thick at the often Ruiz & Pavon (E, G). (l-)1.5-2(-3) mm base, tapering to (0.7-)1 mm, prominently ridged by decurrent leaf bases, pale green to stramineous, 35b. Huperzia linifolia var. jenmanii (Underw. & often with bright red spots on leaf bases. Leaves uniform borne in Lloyd) B. 011g. & Windisch, Bradea 5: 13. usually throughout, alternating, 1987. irregular whorls of 6-7, in basal divisions, upward often in whorls of 4-5, perpendicularly spreading to to Lycopodium jenmanii Underw. & Lloyd, Bull. Torrey ascending, straight upward curved, usually Bot. Club 33: 1 12. 1906. TYPE: Guyana, Monica not twisted at the base, linear to filiform, (6-) 10- Jenman River, (holotype, NY!; isotypes, BONN, 1 7 mm long, 0.3-0.5 mm wide at the base, quickly Herb. Nessel 388 in part, E!). narrowed to ca. 0.2 mm due to involution, grad- Urostachys jenmanii (Underw. & Lloyd) Nessel, Bar- canaliculate to involute, lappgewachse 158. 1939. ually tapering, adaxially Huperzia jenmanii (Underw. & Lloyd) Holub, Folia often with a prominent vein abaxially near the Geobot. Phytotax. 20: 74. 1985. base. Leaves of terminal divisions in old plants sometimes gradually reduced to 6(-4) mm long. Decurrent leaf bases not wider than the Epiphytic in lowland rain forest up to ca. 600 usually lamina often red. 1-1.5 mm m, San Martin. base, bright Sporangia wide. Throughout the Amazonian basin and the Guianas. Epiphytic in lower montane forests, up to alt. 2750 m, San Martin. San Martin: Prov. Mariscal Caceres, Palo Blanco, W West southern Mexico to An- of Puente, Tocache Nuevo, J. Schunke V. 5738 (F, NY, Indies; Panama; s, us). dean South America south to Peru.

44 FIELDIANA: BOTANY Huperzia wilsonii is related to H. dichotoma and near the leaf base, with green decurrent leaf bases. H. mandiocana (Raddi) Trev. (Brazil) and to H. Sporangia ca. 1 mm wide. polycarpos. With these it shares the bottlebrush- like growth habit, and with H. mandiocana the Pendulous epiphyte in montane forest, ca. 2000 bright red coloration of the leaf bases in some m, Huanuco?, Junin. individuals. The absence or presence of red color Costa Rica to Colombia; Peru. seems uncorrelated with other characters. The in- Huperzia polycarpos is close to H. wilsonii, which tensity and size of the coloration are variable. see for discussion. From H. sarmentosa it is dis- Pendent individuals of H. wilsonii are rather tinguished by the lack of leaf base auricles. From similar to H. polycarpos, but the latter has shorter, H. mollicoma (Spring) Holub (tropical America), more flattened leaves and uniformly falcate leaves it differs in the more patent, twisted leaves, with due to a basal twist, and this species apparently is much less prominent veins. always pendulous. The direction of leaves is correlated with growth habit. In erect-growing plants Junin: Chanchamayo Valley, 2000 m, C. Schunke 529 the leaves are wide-spreading to almost reflexed, (F). while in pendent plants the leaves are somewhat ascending. 38. Huperzia sarmentosa (Spring) Trev., Atti Soc. Ital. Sci. Nat. 17: 248. 1874. San Martin: Prov. Rioja, Pedro Ruiz-Moyobamba road, km 390, Venceremos, 1750 m, D. Smith 4372A sarmentosum Mem. Acad. (MO). Prov. Mariscal Caceres, Rio Abiseo National Park, Lycopodium Spring, roy. 24 13. 1849. trail La Playa camp-Papayas, 2650-2750 m, Young & Belg. [Mon. Lye. 2]: LECTOTYPE in and Anders- Leon 4968 (AAU). Department unknown: Ex herb. Pavon (designated by 011gaard Harling son, Fl. Ecuador 33: 100. (o, P). Peru, M. Matthews (G, K). Peru, Matthews 1082 1988): Ecuador, Qui- tonian Jameson 41 (o). Andes, (K!). Urostachys sarmentosus (Spring) Herter, Repert. Spec. Nov. Regni Veg. 19: 165. 1923. 37. Huperzia polycarpos (Kunze) B. 011g., Opera Bot. 92: 169. 1987. Plants slender, pendulous, with flaccidly hanging divisions, up to 100(-200) cm long. Shoots Lycopodium polycarpos Kunze, Linnaea 9: 5. 1834. homophyllous or almost so, equally thick through- TYPE: ad Peru, Cassapi [Casapi?, Huanuco], out, 15-25 mm in diameter including leaves in Poeppig (w!). basal divisions, or sometimes tapering to ca. 8 mm Urostachys polycarpos (Kunze) Herter, Index Lyco- in diameter. Stems leaves 1-1.5 podiorum 76. 1949. excluding mm Urostachys cuatrecasasii Herter, Revista Sudamer. Bot. thick at the base, often tapering to 0.4-0.8 mm, 10: 123. 1953. TYPE: Colombia, Valle, Cord. slightly to prominently ridged by decurrent leaf Occidental, Vertiente Occidental, del rio Di- hoya bases. Leaves borne in alternating irregular whorls gua, Piedra de Moler, 900-1 180 m, Cuatrecasas of 4-7, spreading to ascending, or rarely loosely 15143A (holotype, us!). in terminal twisted at Lycopodium cuatrecasasii (Herter) Morton, Amer. Fem appressed divisions, usually J. 54: 72. 1964. the lamina base, straight to slightly upward curved, linear to linear-subulate, with distinctly widened, Plants pendulous, with flaccidly hanging divi- auriculate, usually strongly revolute, often over- sions, up to 30 cm long. Shoots homophyllous, lapping lamina bases, 7-15 mm long, 0.4-0.7 mm almost equally thick throughout, 10-15 mm in wide just above the auriculate base, usually ca. 0.3 diameter including the leaves. Stems excluding mm wide in the middle, usually somewhat revo- leaves 1-1.5 mm thick near the base, tapering to lute, with obscure to prominent vein, with smooth 0.5-1 mm, somewhat ridged by decurrent leafbas- margins, or the auricles irregularly dentate. Leaves es. Leaves almost uniform throughout, densely of terminal divisions often densely crowded, con- crowded, borne in alternating, irregular whorls of form, or gradually reduced to 5-7 mm long. De- 67 in basal divisions, upward of (3-)4 whorls, current leaf bases narrower than the lamina base. wide-spreading to ascending, often subsecund, Sporangia 1-1.3 mm in diameter. usually with vertical lamina due to a twist at the base, linear to filiform, 6-10(-12) mm long, 0.4- Epiphytic or rarely hanging from banks, in up- 0.5 mm wide, evenly tapering, usually oblique- per montane forest, alt. 2100-3450 m, Amazonas, falcately curved, adaxially flat or shallowly cana- San Martin. liculate, often with an abaxially prominent vein Ecuador and northern Peru.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 45 Huperzia sarmentosa is quite variable with re- Huperzia curvifolia is variable in size and com- spect to leaf length. It resembles H. polycarpos, pactness, seemingly not only as a response to en- and pendulous forms of//, wilsonii but differs from vironmental factors. It is easy to recognize by the these by its auriculate lamina base. strongly upward curved leaves of basal divisions and the appressed non-sporangiate leaves of the Amazonas: Chachapoyas, Leimebamba-Lajasbamba terminal divisions. The smallest forms with hair- Martin: trail, Boeke 2059 (MO, us, USM). San Prov. Maris- like terminal shoots represent the most fragile and cal Caceres, Rio Abiseo National Park, NW sector, Valle slender extreme known in the family. Only these de Chochos, 3450 m, Leon 2161 (AAU, USM). are represented in Peru. They correspond to the type collection. Other, stronger forms to the north 39. Huperzia curvifolia (Kunze) Holub, Folia in the Andes and in Central America approach Geobot. Phytotax. 20: 72. 1985. slender forms of H. acerosa and H. filiformis.

Lycopodium curvifolium Kunze, Linnaea 9: 5. 1835. San Martin: Boqueron pass, 92 km from Tingo Maria TYPE: Peru (Dept. Huanuco), sylvar. densarum on road to Pucallpa, 400 m, Allard 22098 (us). Prov. arboribus prope Pampayaco (Pampayacu), 1829, Rioja, Pedro Ruiz-Moyobamba road, km 390, Vencere- Poeppig(LZ, holotype, destroyed; isotypes, AWH!, mos, 1750 m, D. Smith 4372A (USM). Ucayali (as Lo- B!, BR!, E!, oi, K!, M!, NY!, wi). reto): Prov. Coronel Portillo, Boqueron Padre Abad, 400 Lycopodium tenue Willd. var. tenuissimum Spring, m, J. Schunke V. 3046 (F, GH, us), 480 m, Hutchison et Mem, Acad. roy. Belg. 24 [Mon. Lye. 2]: 2 1 . 1 849, al. 6054 GH, uc, us, USM). based on L. curvifolium Kunze and with the same (F, type. Urostachys curvifolius (Kunze) Nessel, Barlappge- wachse 129. 1939. 40. Huperzia tenuis (Willd.) Trev., Atti Soc. Ital. Plants delicate to extremely delicate, flaccidly Sci. Nat. 17: 248. 1874. pendulous, up to 60 cm long. Shoots gradually (rarely abruptly) heterophyllous, 2.5-6 mm in di- Lycopodium tenue Willd., Sp. pi. ed. 4, 5: 55. 1810. TYPE: Ecuador, of near ameter including the leaves in basal divisions, ta- Valley Vinajacu, Loja, Humboldt & Bonpland 3363 (holotype, B, Herb. pering to 0.3-1.5(-2) mm in diameter including mild. 194231; isotypes, BM!, LG!, P!). leaves in terminal divisions. leaves Stems excluding Urostachys tenuis (Willd.) Nessel, Arch. Bot. Est. S. 0.4-0.7 mm thick at the base, tapering to 0.2-0.4 Paulo 1: 401. 1927. mm upward, usually not concealed by leaves. Leaves of basal divisions borne in alternating whorls Plants delicate to extremely delicate, flaccidly of 4-5, patently sigmoid to strongly falcately up- penduolous, at least up to 75 cm long. Shoots grad- ward curved, acicular-nliform, ong and widely de- ually heterophyllous, 3-8 mm in diameter includ- current, 2-4(-5) mm long, 0.2-0.3 mm wide at the ing the leaves in the basal divisions, tapering to base, extremely delicate, abaxially convex, flat or 1-3 mm. Stems excluding leaves 0.5-0.7 mm thick canaliculate adaxially. Vegetative leaves of ter- at the base, tapering to ca. 0.3 mm upward. Leaves minal constricted divisions alternate, or borne in of basal divisions borne in irregular alternating alternating irregular whorls of 3, closely appressed, whorls of 3-5, or spirally arranged, forming 6-10 lanceolate to linear-lanceolate, somewhat clasping indistinct longitudinal ranks, usually densely with the base, 1.2-3.5 mm long, 0.2-0.6 mm wide crowded, spreading to ascending, straight to some- at the base, usually strongly involute, often sharply what curved, often secund, acicular, tapering from carinate at the base, short to long decurrent. Spo- a subauriculate base, 3-5(-7) mm long, 0.3-0.5 rophylls usually few and scattered, often unilat- mm wide, decurrent. Vegetative leaves and spo- eral, conform, or shorter and wider, lanceolate to rophylls of constricted terminal divisions almost widely ovate, or subcordate and short to long acu- conform, borne in alternating irregular whorls of minate, usually widely spreading, 1-3 mm long, 3, or irregularly alternate, patently diverging from 0.3-0.8 mm wide, sometimes scarcely exceeding the stem, with upward curved tips, widely ovate the sporangia, abaxially with a prominent vein, or to subhastate, 1-3 mm long, 0.5-1.4 mm wide, carinate. Sporangia 0.6-0.8 mm wide. clasping at base, tapering to abruptly narrowed into a short to long acuminate apex, bluntly to Epiphytic in lower, wet montane forest, alt. 400- sharply carinate, usually unilaterally, discontinu- 1750 m, San Martin, Ucayali, Huanuco. ously, but often densely sporangiate. Sporangia Costa Rica to Peru. 0.8-1.2 mm wide.

46 FIELDIANA: BOTANY Usually epiphytic, in high montane forests, alt. shining, (2.5-)3-10 mm long, 3-6 mm wide. Spo- 1350-3300 m, Amazonas, San Martin, Cuzco. rangia 1.5-2.5 mm in diameter. Costa Rica; Andes from Venezuela to Peru. Huperzia tenuis exhibits the same type of size Usually epiphytic, in montane, wet cloud forest variation as mentioned for H. curvifolia. Some on the eastern Andean slopes, 3100-3550 m, forms are as extremely fragile and slender but dif- Amazonas, San Martin. fer by the spreading leaves in terminal divisions. Andes of Venezuela, south to northern Peru. The typical forms of the species are also ecologi- This species is easily distinguished because of cally distinct, being restricted to the uppermost its large size and the sharply quadrangular ter- montane forests, while the extremely thin forms minal divisions. The expanded leaves are some-

(LI. Williams 7348 and Spruce 4621) appear to times developed only along a very short portion occur only at lower altitudes. of the stem base and are sometimes lacking in herbarium specimens.

Amazonas: Prov. Chachapoyas, Cerros Calla Calla, 1 8 Amazonas: Prov. road to km above Leimebamba on road to Balsas, 3000-3100 Chachapoyas, Leimebamba, Balsas, 3240 m, Luteyn 11370 (NY). Cerros Calla Calla, m, Hutchison & Wright 6932 (F, GH, MO, NY, uc, us, 3450-3550 m, Wurdack 1707 (F, GH, NY, us); 3360 m, USM); Wurdack 1762 (F, G, GH, K, NY, s, uc, USM). San Hutchison & 6998 Martin: Tarapoto, Monte Campana, Spruce 4621 (BM, Wright (c, E, F, G, GH, K, M, MO, NY, s, uc, us). San Martin: Prov. Mariscal Caceres, NW BR, P). Prov. Mariscal Caceres, NW corner of Rio Abiseo corner of Rio Abiseo National trail to National Park, Puerta del Monte, 3100-3300 m, Young Park, Mirador, 3 100-3300 Leon 4457 Prov. Mariscal & Leon 4458 (AAU). Prov. Mariscal Caceres, Chochos m, Young& (AAU). Caceres, Rio Abiseo National Park, Puerta de Monte, valley, 3100 m, Young 2667 (AAU, USM). San Roque, 3400 m, Leon & 1308 Prov. Mariscal Ca- 1 350-1 500 m, LI. Williams 7348 (F, us). Cuzco: Camino Young (AAU). Rio Abiseo National Chochos 3100 a Huarcan, caqui Acobamba [Ocobamba], Valle Neapi- ceres, Park, Valley, m, 2620 (AAU). llo, 2400 m, Biies 1376 (NY). Department unknown: Young Poeppigin 1829 (MO). Lehmann 5021 (F, GH, us).

42. Huperzia campiana B. 011g., in Harling and 41. Huperzia molongensis (Herter) Holub, Folia Andersson, Flora of Ecuador 33: 109. 1988. Geobot. 20: 75. 1985. Phytotax. TYPE: Ecuador, Prov. Loja, E of Nudo de Cajanuma, 0llgaard et al. 57861 (holotype, Bot. Jahrb. 43: Beibl. Lycopodium molongenseHerter, AAU!; isotype, QCA!). 98: 51. 1909. TYPE: "Molong," Pearce (holotype, K!) [erroneously as "Ostaustralische Pro- Plants rather at least to vinz: Gebirgswalder Neu-Siid-Wales" in the pro- robust, pendulous, up tologue]. 1 m long. Shoots heterophyllous, the basal divi- Urostachys molongensis (Herter) Nessel, Barlappge- sions ca. (15-)20-30 mm in diameter including wachse240. 1939. the expanded leaves, the distal quadrangular divisions abruptly constricted to 2-4 mm thick in- Plants robust to very robust, pendulous, at least cluding the imbricate, reduced leaves. Stems ex- up to 1 50 cm long. Shoots heterophyllous, usually cluding leaves 2-4 mm thick at the base, tapering not sharply differentiated, 20-35 mm in diameter to 0.5-1.5 mm, pale greenish or brownish. Ex- including the expanded leaves in basal divisions, panded leaves of basal divisions usually uniform distally gradually to abruptly constricted to 3-8 throughout, borne in alternating whorls of 3, mm in diameter including the shorter, imbricate spreading to perpendicular, lanceolate to widely leaves in the terminal divisions. Stems excluding lanceolate-ovate, usually widest below the middle, leaves 2-3.5 mm thick at the base, tapering to 1- (7.5-)10-15 mm long, 2.5-4(-4.5) mm wide, often 1.5 mm. Expanded leaves of basal divisions borne somewhat shorter near the contraction of the shoot, in alternating whorls of 3, patent to ascending, or twisted at the base, with flat to revolute margins, slightly recurved, lanceolate to lanceolate-ovate, herbaceous to subcoriaceous, dull to somewhat acute, 10-18 mm long, 3-6(-7) mm wide, flat, or shining. Leaves of terminal constricted divisions with slightly revolute margins, coriaceous, often decussate, or rarely in alternating whorls of 3 near shining, usually twisted at the base. Leaves of con- the base of constricted divisions, usually sporan- stricted divisions decussate or subdecussate, im- giate throughout, appressed and clasping with their bricate, discontinuously sporangiate, ovate to sub- bases, bluntly to sharply carinate, widely ovate to cordate, acute, sharply carinate and clasping with triangular-ovate, short-acuminate or mucronu- the base, usually conduplicate in the apex, usually 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- stricted divisions and is tentatively referred to this gia. Sporangia 1.2-2 mm in diameter. species.

Epiphytic in upper cloud forest and elfin forest, Amazonas: Leimebamba, 2100 m, Woytkowski 7749 alt. 2400-3500 m, San Martin, Huanuco. (F, MO, NY, USM). Ecuador; Peru. Bryan 779 deviates from typical Huperzia cam- 44. Huperzia ericifolia (Presl) Holub, Folia Geo- piana by the relatively wide expanded leaves and bot. Phytotax. 20: 72. 1985. slightly thicker constricted divisions.

Lycopodium ericaefolium Presl, Reliq. haenk. 1: 77. San Martin: Prov. Mariscal Caceres, Rio Abiseo Na- 1825. TYPE: "Peruvia, verosimiliter Luzon," tional Park, NW sector, Valle de Chochos, 3450 m, Leon Haenke (holotype, PRC!; isotype, BR!). 2179 (AAU). Huanuco: Cushi, trail to Tambo de Vaca, Urostachys phlegmaria (L.) Herter var. ericaefolius Bryan 7 19A (F, G, us). (Presl) Nessel, Barlappgewachse 217. 1939.

43. heteroclita Holub, Folia Huperzia (Poiret) Plants slender, pendulous, at least up to 6 cm Geobot. 20: 73. 1985. Phytotax. long. Shoots heterophyllous, in the basal divisions ca. 8-14 mm in diameter including the expanded Lycopodium heteroclitum Poiret, in Lam., Encycl. 3: leaves, then abruptly constricted to 1.5-2 mm in- 554. 1814 [1813]. LECTOTYPE (designated by cluding the imbricate, reduced leaves of the qua- 011gaard in Harling and Andersson, Fl. Ecuador terminal divisions. Stems 33: 1 12. 1988): America calidiore, Herbier de A. drangular, excluding N. Desvaux (P!). leaves ca. 1 mm thick, tapering to ca. 0.5 mm, pale greenish. Expanded leaves usually uniform in Plants pendulous, lax, at least up to 80 cm (pos- position, shape and size throughout, borne in al- sibly to 400 cm) long. Shoots heterophyllous, with ternating whorls of 3, or decussate upward, almost expanded, perpendicular leaves, often only in the continuously overlapping throughout (pressed first 5-20 cm of the basal divisions, upward specimens), widely lanceolate to oblong-lanceo- abruptly constricted, sharply to bluntly quadran- late, acute to obtuse and mucronate, 7-9 mm long, gular, 2-3 mm thick including the small, imbricate 2-3 mm wide, with slightly revolute margins, the leaves, the constricted divisions often interrupted lamina usually twisted at the base to a vertical by short zones with expanded leaves. Stems ex- position. Leaves of terminal, constricted divisions cluding leaves 1-1.7 mm thick at the base, tapering decussate, discontinuously sporangiate, appressed to 0.7-1 mm, often bright red. Expanded leaves and clasping with their bases, carinate to condu- decussate, or subdecussate, perpendicular to the plicate in the apex, lanceolate to ovate- or trian- stem, lanceolate to widely lanceolate, widest in the gular-ovate, short to long acuminate, 2-4 mm long, middle, acute, 6-9 mm long, 2-3.5 mm wide, soft- 1.5-2 mm wide, the sporangiate leaves 1.5-2.5x herbaceous, usually twisted at the base. Leaves of longer than the sporangia. Sporangia 1-1.5 mm constricted divisions decussate, closely imbricate, wide. usually discontinuously sporangiate, ovate-cordate, acute, bluntly to sharply carinate to condu- Wet lower montane forest, alt. 625-2040 m, San plicate in the apex, 1.5-4 mm long, 2-2.5 mm Martin, Huanuco. wide, dull, sometimes with shallowly erose, or in- Ecuador to Bolivia. distinctly dentate margins. Sporangia ca. 1.5 mm Huperzia ericifolia is closely related to H. di- wide. chaeoides (Maxon) Holub (Central America to Ec- uador) and H. aqualupiana (West Indies, N Ven- Epiphytic in upper wet montane forest, ca. 2 100 ezuela to NW Colombia). Its distinctness from the m, Amazonas. latter is doubtful, although H. ericifolia can usually Southern Ecuador and northern Peru. be distinguished by its narrower, more acute ex- The species is usually recognizable by its long, panded leaves in whorls of three, while H. aqualu- sharply quadrangular constricted shoots with ex- piana usually has ovate, subacute to obtuse, dec- tensive non-sporangiate zones and sharply cari- ussate expanded leaves. nate, apically conduplicate leaves. However, the Problems of typification were discussed by 011- specimen cited below deviates from the typical in gaard in Harling and Andersson, Fl. Ecuador 33: the terete rather than sharply quadrangular con- 113. 1988.

48 FIELDIANA: BOTANY San Martin: Prov. Rioja, Pedro Ruiz-Moyobamba Huperzia myrsinites is closely related to H. phy- road, km 390, 2040 Smith & Venceremos, m, Vdsquez licifolia. Species of the group of Huperzia quad- 4589 (AAV, USM). Jingo Maria, 625-1 100 m, Allard20935 rifariata (Bory) Rothm. (SE Brazil) were errone- (us). Huanuco: SW slope of Rio Llullapichis watershed, placed under H. several earlier ascent of Cerros del Sira, 1290 m, Dudley 13029 (GH, ously myrsinites by us). authors.

45. Huperzia myrsinites (Lam.) Trev., Atti Soc. Cajamarca: Prov. Hualgayoc, Hda. Taulis, 2500 m, Hutchison Ital. Sci. Nat. 17: 249. 1874. & Bismarck 6391 (F, NY, us). Prov. Contu- maza, Bosque de Cachil, 2450 m, Sagdstegui 14873 (AAV).

Lycopodium myrsinites Lam., Encycl. Meth. Bot. 3: 654. 1789. TYPE: Dominican Republic, "S. Do- mingue," Comm. Joseph Martin, (holotype, P, 46. Huperzia phylicifolia (Poiret) Holub, Folia Herb. Lam.l). Geobot. 20: 75. 1985. Urostachys myrsinites (Lam.) Herter, Repert. Spec. Phytotax. Nov. Regni Veg. 19: 166. 1923. skutchii Proc. Biol. Soc. Wash. Lycopodium Maxon, Lycopodium phylicifolium Poiret, in Lam., Encycl. 3: 46: 159 TYPE: Chimaltenan- (1933). Guatemala, 546. 1814 [1813]. LECTOTYPE (designated by alt. Skutch go, Chichavac, 2400-2700 m, 243 (ho- 011gaard in Harling and Andersson, Fl. Ecuador lotype, us!). 33: 117. 1988): "Habitat in Chili," Herbier de A. N. Desvaux, Donnee par Mme Vve Lavallee en Plants pendulous, slender, at least up to 65 cm 1896(p!). Urostachys phylicifolius (Poiret) Nessel, Barlappge- long. Shoots heterophyllous, usually not all sharp- wachse 246. 1939. ly differentiated, the basal divisions 10-18 mm in Lycopodium congestifolium Spring, Mem. Acad. roy. diameter distal including the expanded leaves, di- Belg. 15 [Mon. Lye. 1]: 70. 1842. TYPE: Peruvia, visions gradually or abruptly constricted to 1.5-3 Dombey, H. M. P., Herb. Deless. (P!). Meded. mm thick including the reduced, imbricate leaves. Lycopodium nubigenum Herzog, Rijks-Herb. 27: 2. 1915. TYPE: Bolivia, Comarapa, Herzog Stem excluding leaves ca. 1 mm thick at the base, 1967 (holotype, L!; isotypes, B!, z!). to ca. 0.5 tapering mm, pale greenish or reddish. Urostachys nubigenus (Herzog) Nessel, Barlappge- Expanded leaves of basal divisions decussate or wachse 246. 1939. subdecussate, often irregularly shaped, subdistant to densely crowded and somewhat overlapping, Plants slender, pendulous, up to 150 cm long. ascending to perpendicular, ovate-lanceolate or Shoots heterophyllous, in the basal divisions ca. lanceolate, acute, usually the widest ones with a 10-20(-25) mm in diameter including the ex- rounded base, 6-11 mm long, 1.5-3 mm wide, panded leaves, then abruptly (rarely gradually) usually flat, straight to somewhat recurved. Leaves constricted to (l-)1.5-2(-2.5) mm in diameter in- of terminal constricted divisions highly variable, cluding the imbricate, reduced leaves. Stems ex- often with complete reduction series, and with re- cluding leaves 0.7-1.2 mm thick at the base, up- current series to expanded shape, decussate or sub- ward tapering to ca. 0.5 mm, greenish to bright decussate, continuously or discontinuously spo- red. Expanded leaves of basal divisions borne in rangiate. Transitional leaves with widely ovate alternating whorls of 3, or decussate, subdecussate, base, and short to long acuminate apex, appressed or alternate, usually widely spaced in alternate- and clasping, with the wide base abaxially rounded leaved stem portions, perpendicular to the stem to bluntly carinate, with straight to recurved apex, to falcately ascending, lanceolate to linear-lanceo- 2.5-5 mm long, 1.5-2 mm wide. Shortest leaves late, widest at or below the middle, (4-)6-10(-13) with base conform, but with straight to falcate mm long, (1-)1 .5-2 mm wide, softly to firmly her- apex, bluntly to sharply carinate, scarcely exceed- baceous, with flat to slightly revolute margins, the ing the sporangia, ca. 2 mm long. Sporangia 1-1.3 lamina twisted to a vertical position. Expanded mm in diameter. leaves near the constriction often sporangiate. Leaves of constricted terminal divisions decussate, Epiphytic in montane forest, ca. 2500 m, Ca- or subdecussate, continuously or discontinuously jamarca. sporangiate, appressed and clasping with their bas- Hispaniola; Central America; Trinidad; Guy- es, abaxially rounded to carinate, widely lanceo- ana; Venezuela to Peru. late to widely ovate or subcordate, acute to mu- For illustration of the species, see A. R. Smith cronate or cuspidate, rarely with an elongate flat (Pteridophytes, in Breedlove: Flora of Chiapas, apex, 1.7-2(-4) mm long (in gradually hetero- part 2: 347,f.82a-b. 1981). 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- Plants slender, pendulous, up to 2 m long. Shoots rangia. Sporangia 1.2-1.5 mm wide. heterophyllous, ca. 15-25 mm in diameter including the expanded leaves in the basal divisions, Epiphytic or rarely rupestral in upper montane abruptly constricted to 1.5-2 mm in diameter in- and elfin forest, 2300-3700 m, Cajamarca, Ama- cluding the reduced and appressed leaves of the zonas, Huanuco, Huancavelica, Apurimac, Cuzco, terminal divisions. Stems excluding leaves ca. 1 Puno. mm thick at the base, upward tapering to less than Due to uncertain delimitation of the species the 0.5 mm, pale greenish to bright red. Expanded total distribution of this species is unsettled. In an leaves of basal divisions borne in alternating, often inclusive sense it is wide-ranging: from Costa Rica, irregular whorls of 3, or often upward subdecus- along the Andes south to northern Argentina, and sate, spreading to perpendicular to the stem, nar- SE Brazil. rowly lanceolate to linear-subulate, straight to Huperzia phylicifolia occurs in a wide range of obliquely falcate-ascending, (6-) 10- 15 mm long, epiphytic and rupestral habitats with highly vari- 0.5-1 mm wide, twisted at the base, flat, soft-her- able exposure and humidity. Its growth habit (i.e., baceous. Leaves of terminal constricted divisions size, leaf size, texture and crowding, and stem col- subdecussate, or in alternating whorls of 3 just or), responds strongly to the environmental con- above the constriction of the shoots, sporangiate ditions. It is closely related to H. myrsinites, H. almost throughout, closely imbricate, widely ovate subulata, H. biformis (Hooker) Holub (Brazil), and with obtuse to acute apex, abaxially rounded to H. erythrocaulon (Fee) Holub (Brasil). Forms of carinate, 1.3-2 mm long, 1.3-1.5 mm wide, the species corresponding to the type of Lycopo- equalling or slightly exceeding the sporangia. Spo- dium nubigenum, with expanded leaves slightly rangia ca. 1 mm wide. longer and widest just above the base, are preva- lent in the southern part of the range (Peru, Bo- Epiphytic, in cloud forest and elfin forest, alt. livia, Argentina). 1 500-3400 m, Amazonas, San Martin, Huanuco, Cuzco.

Cajamarca: Prov. Cutervo, Jelski 1013 (p). Amazonas: Costa Rica; Colombia to Peru. Matthews 1081 Huanuco: Chachapoyas, (G, K, p). Cani, Huperzia subulata is closely related to H. phy- NE of Mito, 2800 m, Macbride 3546 (F, us). Cushi, trail licifolia but seems to tolerate a narrower range of to Tambo de Vaca, Bryan 719B (F, GH, us). Huancaveli- restricted to the most ca: Prov. Tayacaja, Marcavalle, between Huachocolpa growth conditions, being and Tintay, 2700 m, Tovar 4751 (GH, USM). Apurimac: humid forests of the eastern slopes of the Andes. Prov. Abancay, Ampay, 3400-3700 m, Vargas 8388 (MO). Cuzco: Prov. La Convencion, Dist. Vilcabamba, Yu- Amazonas: Prov. Bagua, Serrania de Bagua, ca. 12-18 paanqui, 2700 m, Davis et al. 1210 (AAV, F, GH). Prov. trail km E of La Peca, 1 800-1 950 m, Gentry et al. 22904 Paucartambo, Paso de 2450 Aguila, Pillawata, m, Vargas (AAV, MO, USM). San Martin: Prov. Mariscal Caceres, 22995 Puno: Prov. 2300 Weberbauer (GH). Sandia, m, Rio Abiseo National Park, Puerta del Monte, 3400 m, 633 (G, MOL). Prov. Carabaya, Ayapata-Kahuallayoc, Leon & Young 1340 (AAU). Huanuco: Pampayacu, Hda. 2600-3600 m, Vargas 10741 (GH). Tabina, Lechler2023 at mouth of Rio Chinchao, Bryan 725 (F, G, us). Cuzco: (us). 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). 47. Huperzia subulata (Poiret) Holub, Folia Geo- bot. Phytotax. 20: 77. 1985. 48. Huperzia cuneifolia (Hieron.) Holub, Folia Geobot. 20: 72. 1985. Lycopodium subulatum Poiret, in Lam., Encycl. 3: Phytotax. 544. 1814 [1813]. LECTOTYPE (designated by Ollgaard in Harling and Andersson, Fl. Ecuador Lycopodium cuneifolium Hieron., Bot. Jahrb. Syst. 34: 33: 1 19. 1988): HerbierdeA. N. Desvaux, Donnee 572. 1905. LECTOTYPE (designated by Nessel, par Mme Vve Lavallee en 1896 (P!). Barlappgewachse 252. 1939): Costa Rica, Volcan subulatus Arch. Hot. Est. Urostachys (Poiret) Nessel, Barba, Hoffman 50 (B!). S. Paulo 1: 420. 1927. Urostachys cuneifolius (Hieron.) Nessel, Barlappge- Urostachys ewanii Herter, Revista Sudamer. Bot. 10: wachse 252. 1939. 126-127. 1953. TYPE: Colombia, drainage of upper Rio Pascual, above Cordoba, 2840 m, Ewan Plants pendulous, delicate, up to 60 cm long. 16526 (holotype, us!; isotype, NO!). Shoots in the basal divisions Lycopodium ewanii (Herter) Morton, Amer. Fern J. heterophyllous, (5-)7- 54: 72. 1964. 1 3 mm in diameter including the expanded leaves,

50 FIELDIANA: BOTANY then abruptly constricted to 1-2 mm in diameter Shoots heterophyllous, the basal divisions with including the reduced, imbricate leaves in the ter- expanded leaves ca. 10-18 mm in diameter in- minal, constricted, quadrangular divisions. Stems cluding leaves, distally abruptly constricted to ca. excluding leaves 1 mm thick, or less, at the base, 2-2.5 mm including the reduced, imbricate leaves. tapering to ca. 0.5 mm, greenish to bright red. Stems excluding leaves up to 5 mm thick at the Expanded leaves of basal divisions decussate, base, upward tapering to ca. 0.5-1.5 cm, rigid, spreading to perpendicular or slightly reflexed, dark brown to purplish brown. Expanded leaves spathulate to lanceolate or lanceolate-obovate, of basal divisions borne in alternating, irregular, usually widest above the middle, obtuse or mu- distant whorls of 4, the whorls 6-9 mm apart, cronulate, 2-6.5 mm long, 1.5-3 mm wide, flat, ascending to spreading or sharply reflexed, lan- the lamina twisted to a vertical position, softly to ceolate, straight or recurved, (6-)8-10 mm long, firmly herbaceous. Leaves of terminal constricted 2.5-3 mm wide, acute, flat, coriaceous, with divisions decussate, often sporangiate in rather smooth, revolute margins, often adaxially con- short zones of the divisions, appressed and clasp- cave. Leaves of recurved, constricted divisions sub- ing with their bases, widely ovate to subcordate, decussate, densely crowded, sporangiate almost 1.5-2 mm long, 1-1.5 mm wide, carinate, with throughout, imbricate, widely ovate with obtuse falcate to reflexed mucronulate tips, the sporo- to acute apex, abaxially rounded to carinate, 1.2- phylls equalling or slightly exceeding the sporan- 1.6 mm long and wide, equalling or slightly ex- gia. Sporangia ca. 1 mm wide. ceeding the sporangia. Sporangia 1-1.3 mm wide.

Epiphytic or occasionally on banks in upper Indicated as terrestrial scandent shrub, in ceja montane forest and elfin forest, ca. 3000-3100 m, scrub (protologue) and as an epiphyte in high Swie- Amazonas and San Martin. tenia trees (Nessel, in Hoehne, Fl. Bras. 11: 88. Costa Rica; Andean Venezuela; Colombia and 1955). Peru. Endemic, Amazonas and San Martin. The two taxonomically synonymous basionyms Amazonas: Prov. Chachapoyas, Cerros Calla Calla, 19 were published simultaneously. The epithet prui- above 3100 Hutchison km Leimebamba, m, & Wright nosa is maintained due to the quality of its type 6943 (F, GH, K, M, P, us); Wurdack 1744 (F, GH, NY, us, material and greater certainty of its origin. USM). San Martin: Prov. Mariscal Caceres, NW sector of Rio Abiseo National Park, near Mirador, 3000-3100 m, Leon 27/9(AAu). San Martin: Mount Organero, ceja, 1900 m, Melin 96 (s). "Tarapoto, 1908, G. Huebner" (BONN, Herb Nessel 6211). 49. Huperzia pruinosa (Herter) Holub, Folia Geo- bot. Phytotax. 20: 76. 1985.

Lycopodium pruinosum Herter, Bot. Jahrb. Syst. 43, Beibl. 98: 52. 1909. TYPE: Ama- Peru, Dept. Comments zonas, Prov. Chachapoyas, Tambo Ventillas, 2400-2600 m, Weberbauer (err. as Ule in protologue) 4410 (holotype, B!; isotypes, BONN, Herb. Huperzia polyclada (Sodiro) Rolleri and Deferrari, Nessel 62 l\,o\). Notas Mus. La Plata, Bot. 21(1 00): 156.1988. Lycopodium durissimum Herter, Bot. Jahrb. Syst. 43, Beibl. 98: 52. 1909. TYPE: Peru, "Voyage a vase. FEquateur et au Perou, 1876-77, Guayaba mars Lycopodium polycladum Sodiro, Crypt, Quit. 561.1893. TYPE: Ecuador, Mojanda, Sodiro (not 1877" (err. "Colombia, Guayabal" in proto- located). logue), Vidal-Seneze ["Senege"] (holotype, P!; isotype, BONN, Herb. Nessel 6221). Urostachys pruinosus (Herter) Nessel, Arch. Bot. Sao The type locality in Ecuador contains several Paulo 1:420. 1927. taxa of the Huperzia crassa group. Due to the ab- Urostachys durissimus (Herter) Nessel, Barlappge- sence of authoritative material and the wachse 240. 1939. relatively scarce information of the it is Huperzia durissima (Herter) Holub, Folia Geobot. protologue not pos- Phytotax. 20: 72. 1985. sible to certify to which of these taxa the name applies. However, none of the taxa correspond to Plants erect and distally recurved, unbranched the Peruvian material that was placed in this spe- or sparsely branched at base, distally with densely cies by Rolleri (1980). This material is here re- tassel like ramification, at least up to 70 cm long. ferred to H. darwiniana.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 51 II. Lycopodium base, reniform, with a short thick stalk, isovalvate or slightly anisovalvate, their epidermis cells with side Lycopodium L., Sp. pi. 1100. 1753. TYPE: Ly- thin, lignified, sinuate walls, without partial copodium clavatum L. Figure 7. thickenings. Spores reticulate. Gametophytes ob- conic to convoluted disc-shaped, subterranean,

Lepidotis Mirbel, in Lam. & Mirbel, Hist. nat. veg. 3: mycoparasitic, lacking pluricellular uniseriate tri- 477. 1802. TYPE: Lycopodium clavatum L. chomes among the gametangia. Lycopodium subgen. Lepidotis Baker, Handb. Fern- Allies 8. 1887. TYPE: Lycopodium clavatum L. The generic description and synonymy above Lycopodium subgen. Rhopalostachya Pritzel, Nat. includes the For Pflanzenfam. 1 (4): 601. 1901. TYPE: Lycopo- only Neotropical representatives. dium clavatum L. a fuller general treatment ofthe genus, see 011gaard Diphasiastrum Holub, Preslia 47: 104. 1975. TYPE: (1987). Lycopodium complanatum L. The genus Lycopodium as here circumscribed Diphasium Rothm., Feddes Repert. Spec. Nov. Regni has perhaps 40 species mainly in temperate regions Veg. 54: 64. 1944. TYPE: Diphasium jussiaei (Poiret) Rothm. (= Lycopodium jussiaei Poiret). of both the Northern and Southern Hemispheres, Austrolycopodium Holub, Folia Geobot. Phytotax. 26: with eight species in the Neotropics and five in

9 1 . 1 99 1 . TYPE: Beauv. 87, Lepidotis magellanica Peru, all of these, except L. vestitum, with wide (= Lycopodium magellanicum (Beauv.) Sw.). distributions. It includes several of the genera of Holub (1975, 1983, 1985, 1991), also advocated Plants anisotomously branched, with elongate, by Wagner and Beitel (1992). These genera cor- indeterminate, subterranean, creeping, or scan- respond to the sections in 011gaard (1987, 1989). dent, plectostelic main stems (rhizomes), which, Four of the nine sections are represented in the in a dorsolateral position, give rise to usually de- Neotropics. These sections are very distinct groups terminate, ascending to erect, dendroid or spread- of species. None of them are connected by inter- ing, repeatedly inclinate-bilaterally branched, mediate species or by intersectional hybrids. branchlet systems. Roots emerging directly along the underside of main stems, with plectostelic main roots. Branchlet leaves uniform or anisophyllous. Reference Strobili erect, simple or forked, sessile or borne on simple or forked peduncles. Sporoph ylls peltate WILCE, J. H. 1965. Section Complanata of the without a basal mucilage cavity, or subpeltate with genus Lycopodium. Beih. Nova Hedw. 19: 1-IX, a thin basal decurrent wing and with a basal mu- 1-233, t. 1-40. cilage cavity. Sporangia attached to the sporophyll

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 ofbranchlet, 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. Almost cosmopolitan, in humid temperate and LECTOTYPE: Herb. Burser XX: 49 (UPS, des- boreal regions of the Northern Hemisphere and ignated by Jonsell & Jarvis, Regnum Veg. 1 27: on tropical mountains in the Old and New World. 63. 1993). The species is highly variable and adaptive to external factors. The discovery of tetraploids in Ja- Lepidotis clavata (L.) Beauv., Prod. Aetheogamie 108. pan indicates that genetic variation is also in- 1805. volved. There is as yet no indication of polyploidy in the Neotropical representatives of the species. Plants creeping, trailing, or hanging over banks. Lycopodium clavatum varies almost continu- Main stem usually above ground with ascending ously from amply branched plants with diverging to stiffly erect, up to at least 50 cm tall, repeatedly branches and spreading, soft leaves, and long- unequally branched aerial shoots, with strongly branched peduncles, growing in moist, warm, shel- diverging to almost parallel branchlets. Ultimate tered habitats, to small, compact, parallel-branched branchlets terete. Leaves borne in low alternating plants with more imbricate and firm leaves, and spirals or whorls of 6-8(-10), patent to ascending lacking, or short, simple or once forked peduncles, or imbricate, linear-acicular, 6-8(-10) mm long belonging to cold, exposed habitats. The latter and 0.5-0.8 in a col- mm wide, terminating long forms are here recognized as ssp. contiguum. Cor- orless trichome or short membranous apex, with responding monostachyous forms are found in the smooth to sparsely denticulate margins. Strobili Arctic and high mountains of New Guinea. The sessile or pedunculate. Peduncles, if present, erect, two subspecies recognized here are often consid- to 30 or branched and up cm long, simple, bearing ered to be distinct species. However, there are many up to 6 pedicellate strobili. Strobili 1.5-6(-8) cm intermediate forms that cannot be placed in one ca. 6 in diameter long, mm including sporophylls, or the other taxon with certainty. These inter- sometimes forked. Sporophylls borne in alternat- mediates form normal spores and have normal whorls of with ing 5-6, subpeltate, triangular-ovate meiosis. Also there are rare examples of single to rhombic-ovate, acuminate exterior face, with individuals showing characters of one subspecies usually widely scarious, dentate to erose-laciniate in the growth of one year and of the other the 1.3-1.6 their margins. Sporangia mm wide, epi- following year. As the two forms are usually rec- cells side walls dermal with strongly sinuate with ognizable and are ecologically rather distinct, I pocket-like in- and evaginations. Spores reticulate treat them as subspecies. on all faces.

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 Brasilia, no. 41 in herb. deA. N. Desvaux, Donnee par Mme Vve Lavallee en 1895" (P!). Lycopodium clavatum L. var. aristatum (Willd.) Spring, Lycopodium aristatum Willd., in L., Sp. pi. ed. 4, 5: Flora 21 (1): 173. 1838. 17. 1810. TYPE: Venezuela, Silla de Caracas, Lycopodium eriostachys Fee, Crypt. Vase. Bresil 1: Humboldt & Bonpland (holotype, B, Herb. Willd. 224. 1869. TYPE: Brasilia fluminensi, Serra dos 79557!). Orgaos, Glaziou 1788 (BR!, c!, P!, RB!). Lycopodium piliferum Raddi, PI. Bras. nov. gen. 1 : 79, t. 3. 1825. Based on L. aristatum Willd. and with the same type. Clearings exposed habitats in montane forest, Lycopodium trichophyllum Desv., Mem. Soc. Linn. 700-3200 m, Piura, Cajamarca, Amazonas, La Paris 6: 184. 1827 [as trychophyllum]. LECTO- TYPE (designated by 011gaard in Harling and An- Libertad, San Martin, Loreto, Huanuco, Pasco, dersson, Fl. Ecuador 33: 125. 1988): "Habitat in Junin, Ucayali, Ayacucho, Cuzco, Puno.

54 FIELDIANA: BOTANY Humid temperate and boreal regions of the disturbed paramo, 2600-3800 m, Piura, Caja- Northern Hemisphere, montane in the tropics. marca, Amazonas, San Martin, Ancash, Pasco, Absent from Australia. Huancavelica, Apurimac, Cuzco. For more complete synonymy, see 011gaard Costa Rica; Andes from Venezuela to Peru. (1988). For more complete synonymy, see 011gaard (1988). Cajamarca: Prov. Cutervo, Cutervo-Socota, 2320 m, & 5331 Prov. Lopez Sagdstegui (F, MO). Hualgayoc, Piura: Prov. Huancabamba, Cuello del Indio, 2800 m, 2448 Sanchez 2099 Ama- Culquirrumi, m, Vega (AAU). Lopez et al. 8883 (AAU). Cajamarca: Prov. San Miguel, zonas: Prov. between Chachapoyas, Pipus, Chachapoyas near El Tingo (Agua Blanca), 3100 m, Mostacero et al. and 2100 Sanchez et al. 2180 Molinopampa, m, Vega 1130 (AAU). Cutervo, Jelski 1016 (us). Amazonas: Prov. La Libertad: Prov. above 2800 (AAU). Bolivar, Longotea, Chachapoyas, Saullamur-Calla Calla, 3250 m, Sanchez San Martin: Prov. Pedro m, Sagdstegui 14189(r). Rioja, Vega et al. 2136 (AAU). San Martin: Prov. Mariscal Ca- km Ruiz-Moyobamba road, 390-394, Venceremos, ceres, NW corner of Rio Abiseo National Park, 3500 m, 1910-2040 D. Smith 4524 near m, (AAU). Zepelacio, Young & Leon 4712 (AAU). Ancash: Prov. Carhuaz, 1 1 00-1 600 3360 Tara- Moyobamba, m, King (MO, s, us). Huascaran National Park, Quebrada Ishinca, 3900 m, 4732 Huanuco: Prov. Leon- poto, Spruce (BR, E, G, p, us). D. Smith et al. 9588 A (AAU). Prov. Yungay, Huascaran cio Dist. near 700 Prado, Rupa Rupa, Tingo Maria, m, National Park, Quebrada Ranincuray, 3850 m, D. Smith 337 Prov. Dist. Santo King (F). Huanuco, Churubamba, et al. 9067 (USM). Pasco: Prov. Oxapampa, Dist. Huan- Toribio, 1500 m, Mexia 8144 (F, MO, s, uc, us). Pasco: cabamba; Sta. Barbara, above Lanturachi, 3300-3500 Prov. Cord. San 2600 Leon 512 Oxapampa, Gutardo, m, m, Foster et al. 10398 (AAU). Huancavelica: Prov. Tay- Cord. 1 2 E of (AAU). Yanachaga, km Oxapampa-Villa acaja, Montepungo, 5 km E of Surcubamba, 300 m, Rica road, 2100-2200 & Smith 35922 m, Gentry (AAU). Stork & Horton 10386 (F). Prov. Tayacaja, vicinity of Junin: Prov. above La Tarma, Chanchamayo Valley, Huachocolpa, 3000-3100 m, Tovar 3950 (USM). Apu- at Hutchison 1 194 Merced Cumbre Yacunay, 2000 m, rimac: Prov. Abancay, Cerro Turronmocco, NW ofNev- & Smith 24255 (F, M, s, uc, us). Huacapistana, Killip ado Ampay, 3500 m, Brandbyge 422 (AAU). Cuzco: Prov. (us). Ucayali: Prov. Coronel Portillo, La Divisoria, ca. Urubamba, Antakillqa, 3300 m, Franquemont 285 (F). 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, 2. Lycopodium vestitum Poiret, in Lam., Encycl. between Tambo, San Miguel, Ayna and Hda. Luisiana, 3: 546. 1814. LECTOTYPE 1 300 m, Dudley 1 1 953 (F). Cearapa, Killip & Smith 22459 (designated by (us). Cuzco: Valle del Urubamba, Machu Picchu, 2400 011gaard in Harling & Andersson, Flora of Herrera Tres et al. m, 3197 (c, F). Cruces, Gentry 23458 Ecuador 33: 128. 1988): Ecuador (Prov. Loja), (us). Puno: Below Limbani, 3100-3200 m, Brandbyge prope Loxam, Humboldt (p-Humboldt!; iso- 523 (AAU). At Lake Titicaca near Huancano, 2000 m, B, Herb. Willd. 194281, BR!, LG!, us!). Charpin AC- 12704(0). types,

Lycopodium scariosum Hooker, Icon. PI. 1 : /. 89. 1 836, not Forst. TYPE: Matthews 1765 Ib. Lycopodium clavatum ssp. contiguum Peru, Casapi, (holotype, K!; isotype, IVY!). (Klotzsch) B. 011g. in Harling & Andersson, Lycopodium albidum Baker, J. Bot. 25: 37. 1887. Flora of Ecuador 33: 126. 1988. TYPE: Ecuador, Prov. Loja, Mataba 1883, Hu- beck (holotype, K!). Lycopodium contiguum Klotzch, Linnaea 18: 519. 1844. LECTOTYPE in (designated by 011gaard Rhizome creeping, scrambling or hanging over Harling & Andersson, Flora of Ecuador 33: 126. banks, usually above ground, 3-5 mm in diameter 1988): Ecuador, Paramo de Tiopullo [between leaves. Aerial shoot stiffly erect, Cotopaxi and Illiniza], Hartweg 1474 (B!; isolec- including systems totypes, BM!, GL!, K.!, NY!, P!). sparsely branched, with almost parallel branches, vestitum var. herbaceum Lycopodium Poiret, Spring, silvery whitish. Branchlet leaves appressed, linear- Mem. Acad. roy. Belg. 24 [Mon. Lye. 2]: 45. 1 849. lanceolate, 5-9 mm long, 0.6-0.8 mm wide, TYPE: Colombia, Sierra Nevada, Moritz (holo- herbaceous in less than half of their length, with type, B!; isotype, LG!). Lycopodium clavatum L. var. pseudo-contiguum colorless, translucent, membranous, slightly Christ, Primitiae Fl. Costar. 3 (1): 55. 1901. TYPE: widened, coarsely erose-laciniate apex. Strobili Costa Rica, Cerro de las 3000 versant Vueltas, m, sessile or borne on a short, simple, indistinct pe- W du massif de Buena Vista, Pittier 10467 (ho- duncle. Sporophylls borne in alternating whorls of lotype?, BR!; isotype?, P!). 5-7 1.5-2 Lycopodium herbaceum (Spring) Hieron., Bot. Jahrb. 5-6, subpeltate, mm long, mm wide, Syst. 34: 575. 1905. narrowly triangular-ovate, with a long membranous apex, with widely membranous, erose-den- Clearings and exposed habitats in uppermost tate to laciniate margins. Sporangia 1.2-2 mm montane forest and in grass paramo, frequent in wide, their epidermis cells with strongly sinuate

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 55 side walls with pocketlike evaginations. Spores re- erous apex, 3-5 mm long, 0.6-0.8 mm wide. ticulate on all faces. Strobili sessile, or terminating somewhat indistinct, simple or up to twice-forked peduncles, 3- Clearings, road banks, and exposed sites in up- 4 mm thick, 2-10(-l 5) cm long, often forked. Spo- per montane forest, and in the lower paramos, rophylls borne in alternating whorls of 4, peltate, 2400-3500 m, Piura, Cajamarca, Amazonas, San with a slender, terete, wingless stalk, with widely Martin. ovate, short to long acuminate exterior face at- Southern Ecuador and northern Peru. tached to the stalk near the center, with narrowly Lycopodium vestitwn is closely related to L. cla- scarious, almost smooth to shallowly erose-den- vatum, especially to ssp. contiguum. In spite of the ticulate margins. Sporangia ca. 2 mm wide; spor- striking appearance of this species, it is only doubt- anguium epidermis with evenly sinuate side walls. fully distinct from L. clavatum. Individuals with Spores with unornamented proximal faces, with intermediate characters between L. vestitwn and an irregular, fine-meshed reticulum on distal faces. both subspecies are frequent. These have normal meiosis and normal spores. Creeping or hanging over banks in grass paramo, usually in relatively dry paramos, but also often Piura: Prov. Huancabamba, Cord. Chinguela (Sapa- in mossy bogs, 2750-4000 m, Lambayeque, Ca- lache-El Carmen), 2900 m, Sagdstegui et al. 10223 (AAU). jamarca, La Libertad, San Martin, Ancash, Huan- Above Huancabamba, 3400-3500 m, Weberbauer 6145 uco, Cuzco. (F, GH, us). Cajamarca: Prov. Cutervo, Fortaleza de Santa Hispaniola; Costa Rica; Andes from Venezuela Cruz, San Andres, 2400 m, Llatas & Suarez 2866 (F). to Chile Cutervo, Jelski 990 (p), 994, 1007 (KRA). Amazonas: Prov. and Argentina. Bagua, Cord. Colan E of La Peca, 3200 m, Barbour 3261 The subterranean rhizomes seem to enable this Prov. Cerros Calla E (AAU). Chachapoyas, Calla, side, species to survive in vegetation that is burned fre- 19 km above Leimebamba on road to Balsas, 3100 m, quently. Hutchison & Wright 5545 (F, GH, MO, P, uc, us, USM). San Martin: Rio Abiseo National Park, Mariscal Ca- Lycopodium magellanicum is possibly identical ceres, NW corner ofthe Park, 3300 m, Young 2754 (AAU, to certain forms included in L. fastigiatum R. Br. USM). Rio Abiseo National park, Mariscal Caceres, La- of New Zealand and Australia. guna del Eco, Chochos, 3450 m, Young & Leon 4901 (USM). Lambayeque: Ferranafe, Laguna Tembladera, Inca- huasi, 31 50 m, Sagdstegui et al. 12791 in part (F). Prov. 3. Lycopodium magellanicum (Beauv.) Sw., Syn. Ferranafe, ca. 7 km NW of Incahuasi, near Cerro Puna- Fil. 180. 1806. 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), Lepidotis Beauv., Prod. 102. magellanica Aetheogamie 3000 m, Sagdstegui 9526 (NY). Cutervo, Jelski 993, 1012, 1805. TYPE: Fretum unknown magellanicum, 1017 (KRA). La Libertad: Bolivar, Laguna de los Ichus, collector (holotype in G, Herb. De- presumably 3600 m, Lopez & Sagdstegui 3239 (GH). San Martin: lessen n.v.\ P!, Herb. Palisot de Beauvois, isotype, Prov. Mariscal Caceres, Parque Nacional Rio Abiseo, comm. M. de Jussieu). Paredones, 3650 m, Leon & Young 1606 (USM). Prov. ed. 5: 28. 1810. Lycopodium spurium Willd., Sp. pi. 4, Mariscal Caceres, Parque Nacional Rio Abiseo, Cho- TYPE: Ecuador (Prov. Quito, in Tungurahua), chos, 3400 m, Young & Leon 4896B (USM). Ancash: vulcano Humboldt Tungurahua, (holotype, B, Prov. Carhuaz, Huascaran National Park, Quebrada Ish- Herb. Willd. 193641). inca, 3900 m, D. Smith et al. 9588B (AAU). Prov. Huari, Hooker, Icon. pi. 1: /. 55. Lycopodium pichinchense Huascaran National Park, Quebrada Pachachaca, 3700- 1837. LECTOTYPE B. (designated by 011gaard, 3860 m, D. Smith et al. 12547 (USM). Huanuco: Yanano, Biol. Skr. Dan. Vid. Selsk. 34: 62. 1989): Ecuador, 2000 m, Macbride 4928 (F). Cuzco: Paucartambo, 3700 Pichincha, 10,000 ft, Col. Hall (38?) (K!). m, Vargas 13938 (GH). Paucartambo, Parque Nacional Austrolycopodium Holub, Fo- magellanicum (Beauv.) Manu, Cerro Macho Cruz, 3200 m, Leon & Aguilar 2345 lia Geobot. 26: 91. 1991. Phytotax. (USM).

Rhizome subterranean or above occasionally 4. Lycopodium jussiaei Poiret, in Lam., Encycl. ground, 1-2 mm thick leaves. Aerial excluding 3: 543. 1814. TYPE: Perou (holotype, P, Herb. shoots ca. 1 mm thick leaves at excluding origin, Jussieu 6581). up to 25 cm long excluding peduncles and strobili, repeatedly unequally branched, with spreading to Lycopodium jussiaei Poiret var. microphyllum Poiret, ascending branchlets. Branchlets 4-7 mm in di- in Lam., Encycl. 3: 543. 1814. TYPE: Amerique ameter leaves. Branchlet leaves acicular, including meridionale, Herb. Bonpland (holotype, P!). flattened, smooth, with a long, pointed, non-pilif- Lycopodium haenkei Presl, Reliq. haenk. 1: 78. 1825.

56 FIELDIANA: BOTANY TYPE: Peru (Dept. Huanuco), In montosis Pe- and to plants from Chile and Juan Fernandez rec- ruviae ad Haenke Huanocco, (holotype, PRC!; ognized as L. gayanum Remy. The plants referred possible isotype, K!). to Lycopodium jussiaei are generally larger and Lycopodium heterophyllum Sprengel, Syst. Veg. ed. coarser than those A modern revision of 16, 4: 13. 1827. TYPE: Peruvia, Humboldt (ho- species. lotype, B, Herb. Willd. 194251). the group is needed. For more complete synony- Lycopodium scariosum Forst. var.jussieui(Poirei) Ba- my, see 011gaard (1988). ker, Handbook Fern Allies 29. 1887. Diphasium jussiaei (Poiret) [Presl ex] Rothm., Feddes Piura: Huancabamba, Loma Redonda (Sapalache- Repert. Spec. Nov. Regni Veg. 54: 65. 1944 [as Chinguela), 2400 m, Sagdstegui et al. 10203 (AAU). Ca- jussieui]. jamarca: Prov. Cutervo, 10 km NW of Socota, 3200 m, Stork & Horton 10143 (F, G, MO). Cutervo, Jelski 1025 Rhizomes creeping, scrambling to scandent, rig- (KRA). Amazonas: Prov. Bagua, Cord. Colan, E of La id, 2-4 mm thick, usually above ground, with leaves Peca, 3200 m, Barbour 3262 (AAU, MO, USM). Prov. Cha- chapoyas, Puma-urcu SE of Chachapoyas, 3100-3200 radially arranged, uniform, 3-5 mm long, ca. 1 m, Wurdack 1158 (F, s, uc). San Martin: Prov. Mariscal mm wide, linear-lanceolate, with widely membra- Caceres, NW corner of Rio Abiseo National Park, 3300 obtuse Aerial shoot nous, irregularly apex. systems m, Young & Leon 4882 (AAU). Huanuco: Prov. Huanuco, 5-75 cm tall, branched almost from the base, in Carpish Pass, 3300 m, Hodge 6291 (AAU, F). Churubam- 1400 Mexia 8146 large individuals with a main axis almost conform ba, Pampa Hermosa, m, (F, o, MO, s). Pasco: Cord. Yanachaga, 1 2 km E of main Oxapampa- to the rhizome, upward gradually changing to an- Villa Rica Road, 2100-2200 m, Gentry & Smith 35924 isophyllous, bearing alternating, fan-shaped (AAU, MO). Oxapampa, 1970 m, Smith & Pretel 1645 branchlet systems. Branchlets dorsiventral, ani- (AAU). Huancavelica: Prov. Tayacaja, Montepungo, 5 km sophyllous, flattened, 4-6(-8) mm wide including E of Surcubamba, 3000 m, Stork & Horton 10384 (F). Prov. La eastern MassifofCord. Central leaves, with 2 dorsolateral ranks of wide leaves Ayacucho: Mar, opposing the Cord. Vilcabamba between Tambo San and 2-3 indistinct ventral ranks of narrow leaves. Miguel, Ayna and Hda. Luisiana, Dudley 1 1 973 (F). Cuz- Dorsolateral leaves flattened in the of the plane co: Prov. Paucartambo, Valle de Pilcopata, Patria-Pil- branchlet system, or inclinate to it, obliquely el- lahuata, 2000 m, Foster & Wachter 7475 (AAU). Puno: Below 3100-3200 558 liptic, with the acroscopic margin 2-3 mm long, Limbani, m, Brandbyge (AAU). 11.5 mm wide, forward and ventrally curved, mucronate to short hair-tipped, short to long de- 5. Lycopodium thyoides Willd., Sp. pi. ed. 4, 5: current. Ventral leaves appressed, lanceolate-su- 18.1810. TYPE: Venezuela, Silla de Caracas, bulate, with membranous apices. Peduncles up to Humboldt (holotype, B, Herb. Willd. 193521). 50 cm long (rarely absent), simple or up to twice- forked, bearing 1-3 strobili, terete. Strobili (l-)3- L. var. 10 cm long, 46 mm in diameter including spo- Lycopodium complanatum tropicum Spring, in Mart., Flora Bras. 1 (2): 1 16. 1840. rophylls. Sporophylls borne in alternating whorls Lycopodium complanatum L. var. thyoides (Willd.) of with more or less acumi- 4, subpeltate, ovate, Christ, in Schwacke, PI. Nov. Mineiras 2: 42. nate exterior face, 4-6 mm long, ca. 2 mm wide, 1900 [as thujoides]. with narrowly membranous, shallowly erose-den- Lycopodium complanatum L. var. validum Weath- Proc. Amer. Acad. Arts Sci. 45: 414. 1910, ticulate margins. Sporangia 1.5-2 mm wide, the erby, based on L. thyoides Willd. and with the same side walls of epidermis cells evenly sinuate and type. curled. finely Spores reticulate, with large, regular Diphasiastrum thyoides (Willd.) Holub, Preslia 47: 108. meshes on distal faces, and unornamented prox- 1975. imal faces.

Plants with creeping, trailing to scandent rhi- Usually a vigorous scrambling to scandent plant, zomes, usually above ground, or hanging over common in clearings and on road banks and open banks. Rhizomes terete, 1.2-2.5 mm in diameter places in montane forest, 1700-3700 m, Piura, excluding leaves. Rhizome leaves relatively dis- Cajamarca, Amazonas, San Martin, Huanuco, tant, borne in irregular spirals, or subverticillate, Pasco, Huancavelica, Ayacucho, Cuzco, Puno. subulate, appressed to ascending. Aerial shoots as- Jamaica; Dominican Republic; Costa Rica; cending to erect, 10-50 cm tall, with vegetative Venezuela; Andes south to Bolivia; Brazil (Ita- portions up to ca. 30 cm tall. Main upright axis tiaia). terete to somewhat flattened, bearing lateral, flat- A variable species, apparently due to environ- tened, fan-shaped branchlet systems. Ultimate mental factors. Lycopodium jussiaei is closely re- branchlets flattened, dorsiventral, anisophyllous, lated to L. scariosum (Indonesia to New Zealand) 1.5-3 mm wide including leaves, with trimorphic,

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 57 decussate leaves. Upper, median branchlet leaves Cajamarca: Prov. Chota, Chota Tacabamba road, km 2800 Smith & 3549 Prov. San with pointed, subulate to acicular, appressed, 1-2 14, m, Vdsquez (AAU, F). Miguel, El Tingo, 3100 m, Mostacero et al. 1123 (AAU, mm long, free blades, and a conspicuous, ca. 0.4- F). Amazonas: Prov. Chachapoyas, Leimebamba-Calla 0.6 decurrent base. Lateral mm wide, prominently Calla road, km 12-15, 2960-3100 m, Smith & Vdsquez branchlet leaves bilaterally compressed, long-de- 4992 (AAU). Donile-Cohechan, Soukup 4160 (F, us). La current, 2.5-7 mm long including bases, the free Libertad: Prov. Pataz, Cerro Colpar, above Yalen, 3300- 3700 m, Young 3046 (AAU, USM). Prov. Bolivar, above blades 1-3 mm long, appressed to spreading, acu- Longotea, 2800 m, Sagdstegui 14195 (F). San Martin: minate to long-pointed, the leaf bases 0.6-1 .5 mm Rio Abiseo National Park, NW corner, 2800 m, Young wide, with almost parallel to distinctly diverging 1480 (AAU); Rio Abiseo National Park, Laguna del Eco, margins, often curved down. Ventral leaves in- Chochos, 3450 m, Young & Leon 4902 (USM). Ancash: Prov. Huascaran National conspicuous, acicular, without decurrent base, 1- Yungay, Park, Quebrada Ranincuray, 3650-3900 m, D. Smith et al. 10336 (AAU, 2 mm long. Peduncles 10-25 cm long, terete, bear- USM). Huanuco: Prov. Huanuco, Huanuco-La Union road 4-9 strobili. Strobili 1 .5-5 cm ing pedicellate long, km 32, 2940-3100 m, Huapalla 2201 (AAU). Carpish 2-4 mm in diameter including sporophylls, often Pass, 2850 m, Asplund 12748 (s). Pasco: Above Oxa- with protracted sterile tips. Sporophylls usually pampa (Cerro Corporation), ca. 2300 m, collector unknown (F). Junin: Huacapistana, 1 800-2400 m, Killip & borne in alternating whorls of 3, subpeltate, widely Smith 24255 (F). Huancavelica: Prov. Tayacaja, Mar- deltoid-ovate, long-cuspidate, ca. 2-3 mm long, cavalle, between Huachocolpa and Tintay, 2800 m, 70- 1.5-2 mm wide, with almost entire, widely mem- var 4765 (USM). Ayacucho: Ccarrapa, between Huanta branous margins. Sporangia 1.5-2 mm wide, with and Rio Apurimac, 2500 m, Killip & Smith 22293 (F, Cuzco: Prov. below side walls ofepidermal cells evenly sinuate. Spores us). Calca, Kachin, 2700-3800 m, Sallo ex Franquemont 244 (AAU, F). Prov. Paucartambo, densely reticulate on all faces. Marcachea, Achirani, 2500-3000 m, Vargas 11133 (F, MO, uc). Paucartambo, Woytkowski 6744 (MO, us). Puno: Clearings, road banks, open habitats, and sec- Below Limbani, 3100-3200 m, Brandbyge 546 (AAU). Prov. Sandia, between Sandia and Cuyo-Cuyo, 3100- ondary scrub in upper montane forest, and in the 3300 m, Ferreyra 16813 (USM). Tabina, Lechler 2034 (E, lowest paramos, alt. 2000-3400 m, Piura, Lam- G, s). bayeque, Cajamarca, Amazonas, La Libertad, San Martin, Ancash, Huanuco, Pasco, Junin, Huanca- velica, Ayacucho, Cuzco, Puno. III. Lycopodiella Throughout moist mountainous regions oftrop- ical America, south to northern Argentina. Lycopodiella Holub, Preslia 36: 22. 1964. TYPE: The present application of the name Lycopo- Lycopodiella inundata (L.) Holub. Figure 8. dium thyoides corresponds to the "L. thyoides- complex" of Wilce (1965). I have not attempted Lycopodium subgen. Cernuistachys Herter, Bot. Jahrb. to treat the infraspecific variation. The Peruvian 43: Beibl. 98: 29. 1909. TYPE: Lycopodium cern- material referred to this species is highly variable uum L. Palhinhaea Vascon. & Bol. Soc. Brot. 41: and may belong to more than one species, but also Franco, II, 24 (1 967). TYPE: Palhinhaea cernua (L.) Vascon. external factors apparently greatly affect the growth & Franco (= Lycopodium cernuum L.). habit of the individuals. 15111 Ferreyra (USM), Lycopodium sect. Caroliniana Bruce, Amer. Fern J. from Dept. Cajamarca, Prov. Celendin, Celedin, 66: 136. 1976. TYPE: Lycopodium carolinianumL. ca. 3100 m, has unusually wide branchlets with Pseudolycopodiella Holub, Folia Geobot. Phytotax. 18:441. 1983. very wide lateral leaves. Lepidotis auct., not Mirbel 1802. For more complete synonymy, see 011gaard Lycopodium subgen. Lepidotis auct., not Sprengel, (1988). Anleit. ed. 2, 2, 1: 108. 1817, nor Baker 1887.

Plants with prostrate, rooting, indeterminate, Piura: Prov. Huancabamba, Dist. Sondor, Cerro La isophyllous or anisophyllous, horizontally branch- Viuda, 2170 m, Sagdstegui et al. 8197 (AAU, MO). Piura: shoots, and arising, erect, simple Loma Redonda, 2400 m, Sagdstegui 10175 (AAU). Lam- ing dorsally bayeque: Prov. Ferranafe, Incahuasi (Sinchigual-Laguna strobiliferous branches (sect. Lycopodiella and sect. Tembladera, 3000 m, Sagdstegui et al. 12870 (AAU, F). 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 Andean representatives of the genus. Although long intervals, and branch occasionally in the hor- growth habit and morphological details are quite izontal plane giving off horizontal branchlet sys- diverse, features of branching, sporangium anat- tems, and usually bearing 1 vertical, dorsally aris- omy, spores, and gametophytes indicate that Ly- ing, dendroid branch system on each loop (sect. copodiella is a natural entity. Campylostachys). Dendroid branch system with a In sect. Campylostachys, most species have erect, series of subdecussate, spreading to hanging, fla- dendroid branch systems arising from the dorsal bellate branchlet systems, which in turn may ter- side of horizontal, looping main axes (e.g., Lyco- minate in sessile, nodding to pendulous strobili. podiella cernud), but in some species the corre- Sporophylls subpeltate, with a median basiscopic sponding dendroid branch system is scandent, wing, or with coalescent basal membranes that hanging, or creeping. In these cases the dendroid almost enclose the sporangia (sect. Campylosta- branch system can be distinguished from the hor- chys). Sporangia on the sporophyll base, or axil- izontal branch system by the orientation of lateral lary (sect. Lycopodielld), strongly anisovalvate, or branchlet systems on the main axes. In the hori- isovalvate (sect. Caroliniand), the epidermis cells zontal system they are distichous, all arranged in with thin, straight, non-lignified side walls, but the horizontal plane. In the dendroid branch sys- with lignified, nodular, or semiannular thicken- tem the lateral branchlets are polystichous, ar- ings. Spores rugate. Gametophytes green, tuberous ranged in an irregularly subdecussate manner. and lobed on the upper side, surface-living, with Zimmer, in a review in Bot. Jahrb. Syst. 112: mycorrhiza in the base. 414, 1991, claimed that Lepidotis is the correct name for this genus as lectotypified by Rothmaler Almost all moist temperate and tropical regions (1944). However, Rothmaler's lectotypification of the world. Perhaps 40 species, the majority of does not refer to the protologue of Lepidotis and these in the Americas and probably nine in Peru, is superceded by the choice of Lycopodium cla- all fairly widely distributed. vatum (Pichi Sermolli, Webbia 26: 145-149. 1971). The generic description above covers only the

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 flattened, 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 upward, not recurved; strobilus-bearing branchlets usually sharply reflexed at tip, or only the strobilus reflexed 5. L. camporum

( li 1 1 . 1 . Lycopodiella alopecuroides (L.) 'ran Amer. semiannular lignified thickenings. Spores rugate, Fern J. 71: 97. 1981. with a distinct equatorial rim.

Lycopodium alopecuroides L., Sp. pi. 1102. 1753. On open, wet, peaty or sandy soil, in the jalca North Vir- TYPE: Eastern temperate America, or montane forest zone, 2400-3450 m, Cajamarca, Kalm (LINN 1257.7). ginia, Amazonas, San Martin, Huanuco, Pasco. Lycopodium matthewsii Hooker, Icon. pi. 1: /. 26. Eastern North 1836. TYPE: Peru, Bagasan, [Bagazan, Dept. temperate America; Cuba; Amazonas] Matthews 1778 (holotype, K.!; isotype, throughout continental tropical America. BM!). As here delimited, in the wide sense, Lycopo- Lepidotis alopecuroides (L.) Rothmaler, Feddes Re- diella alopecuroides is a polymorphic species, con- pert. Spec. Nov. Regni Veg. 54: 66. 1944. sisting of several elements. In the narrow sense, Lycopodiella matthewsii (Hooker) Holub, Folia Geo- the is bot. Phytotax. 18: 441. 1983. species restricted to temperate North Amer- ica and Cuba, while other elements are present in Horizontal shoots creeping and firmly rooted South America. High altitude forms in the Andes are and robust throughout, up to 25 cm long, sparsely and un- relatively compact, wide-leaved, of equally branched, densely covered on all sides by and perhaps deserve recognition as a species almost uniform, somewhat upwardly secund, or which the correct name is Lycopodiella matthews- the material cited below to this form. spreading to perpendicular leaves, 7-12 mm wide ii; belongs including leaves, bearing stiffly erect, dorsally arising, simple, strobiliferous branches. Leaves linear- Cajamarca: Prov. Cutervo, El Suro, 2400 m, Sanchez lanceolate to subulate, flat, with smooth to den- Vega et al. 6019 (AAU, F). Cumbemayo 10 km WSW of Cajamarca, 3550 m, Molau & Sanchez Vega 840 (GB). ticulate-ciliate margins, soft, light green, 5-8 mm Amazonas: Prov. Chachapoyas, 1-5 km W of Molino- long, 0.5-1 mm wide. Erect branches ca. 5-8 mm pampa, 2400 m, Wurdack 1381 (F, GH, uc, us). Lei- in to tall in- diameter including leaves, up 30 cm mebamba-Calla Calla, km 12-15, 2960-3100 m, Smith cluding the strobilus, with radially arranged leaves & Vdsquez 5000 (AAU). Chachapoyas, 2400-2600 m, Weberbauer 4397 Prov. Cord. Colan E of La borne in alternating whorls of 6 or more, like leaves (G). Bagua, Peca, 3500 m, Barbour 3197, 3200A (AAU, MO). San of horizontal shoots, or slightly narrower, usually Martin: Prov. Mariscal Caceres, Rio Abiseo National more Strobili to 12 cm 10-18 ascending. up long, Park, NW corner ofthe park, 3425m, Young 3475 (AAU); mm in diameter including sporophylls. Sporo- 3450 m, Leon 1854 (AAU). Huanuco: Tingo Maria, Mor- phylls arranged as peduncle leaves, narrowly lan- row 11131 (GH). Pasco: Prov. Oxapampa, San Gotardo, 36 km W of 2850 m, D. Smith 2753 ceolate to lanceolate from a subpeltate base, not Oxapampa, (AAU). coalescent at base, with few to many spreading to hooked teeth on margins, 6-10 mm long, 1-1.7 2. Lycopodiella carol in iana (L.) Pichi-Sermolli var. mm wide at base. Sporangia axillary, subglobular, meridionalis (Underw. & Lloyd) B. 011g. & ca. 1 mm wide, strongly anisovalvate, almost com- Windisch, Bradea 5: 27. 1987. pletely concealed by sporophyll bases, the epidermal cells with straight, unlignified side walls with Lycopodium meridionale Underw. & Lloyd, Bull. Torr.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 61 Bot. Club 33: 121. 1906. TYPE: Puerto Rico, dry 3. Lycopodiella contexta (Mart.) Holub, Folia Wilson 94 savannahs, Luquillo Mountains, (ho- Geobot. Phytotax. 18: 441. 1983. lotype, NY!). Lycopodium carolinianum L. var. meridionalis Nes- contextum Icon. Brasil. sel. Arch. Bot. Est. S. Paulo 1: 431. 1927. Lycopodium Mart., Crypt. 38, t. 1. 1834. TYPE: In Pseudolycopodiella meridionalis (Underw. & Lloyd) 20, f. campis graminosis in summo monte fluvii Holub, Folia Geobot. Phytotax. 18: 442. 1983. apricis Arara-Coara, Ja- pura cataractae imminente, alt. ca. 1200 p., Mar- tins (holotype, M!). Horizontal steins and rooted creeping firmly Lycopodium alopecuroides L. var. contextum (Mart.) throughout, to ca. 30 cm long, sparsely and un- Baker, Handb. Fern Allies 19. 1887. equally branched, anisophyllous, with long and Lycopodium sprucei Baker, Handb. Fern Allies 24. 1887. TYPE: Venezuela, San Carlos del Rio Ne- wide lateral leaves, and usually shorter and nar- gro, Spruce 3151 (holotype, K!; isotype, BONN, rower dorsal leaves, 7-1 2(-l 5) mm wide including frag. Herb. Nessel 200\ in part). 1 to leaves, bearing few, dorsally arising, stiffly Lycopodium alopecuroides L. subsp. contextum (Mart.) erect, simple, strobilus-bearing branches. Lateral Hassler, Trab. Inst. Bot. Farm. Buenos Aires 45: 92. 1928. leaves 3-5(-7) mm long, (l-)1.5-2.5(-3) mm wide Lepidotis contexta (Mart.) Rothm., Feddes Repert. at the base, triangular-ovate to lanceolate, oblique- Spec. Nov. Regni Veg. 54: 66. 1944. to into an ly spreading falcately recurved, tapering Pseudolycopodiella contexta (Mart.) Holub, Folia acute to long pointed apex. Dorsal leaves arranged Geobot. Phytotax. 26: 93. 1991. in (l-)2-4 longitudinal ranks (often on the same individual), widely lanceolate to subulate, diverg- Horizontal shoots creeping and firmly rooted to to upward curved, ing appressed, straight throughout, up to at least 30 cm long, sparsely and mm mm wide at the (1.5-)3-4(-5) long, l-1.3(-2) unequally branched, isophyllous, with somewhat base. Erect branches to 30 cm tall including up upwardly secund, ascending, acicular leaves, 3-7 the strobilus, 1-1.5 mm thick excluding leaves, mm in diameter including leaves, bearing stiffly small, 3-5 mm long, acicular leaves in bearing erect, dorsally arising, simple, or sometimes 1-2 x remote, alternating spirals or irregular whorls of forked, up to at least 40 cm tall, vegetative or 4-5. Strobili to at least 13 cm 3-5 mm up long, strobiliferous aerial shoots. Erect branches 4-6 mm thick with appressed sporophylls. Sporophylls in diameter including leaves, densely foliose, iso- borne in alternating whorls of 4 or 5, subpeltate, phyllous. Leaves of erect branches borne in alter- rhombic or ovate-acuminate or ovate-cuspidate to nating irregular whorls of 6-7, subterete at the base triangular-lanceolate, 3.5-6 mm long, (l-)1.5-2(- (angular when dry), sometimes apically flattened, 2.5) mm wide, with entire to erose-denticulate, strongly upward curved from a strongly diverging minutely fimbriate-denticulate margins. Sporan- to almost perpendicular base, 4-5(-7) mm long, gia reniform, isovalvate, borne on the sporophyll ca. 0.5 mm wide, with conspicuously acroscopical- stalk, ca. 1.5-2 mm wide. Spores rugate, with a ly adnate and long decurrent leaf bases, with distinct equatorial rim. 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- Terrestrial or sometimes epilithic, usually on tate, widely triangular-ovate to ovate-lanceolate wet ground, on road banks, slopes, ledges, grass- at base, with a long, narrow apex, 4.5-7 mm long, lands in the jalca zone, on seepages, 1350-2400 1-1.5 with subentire to erose- m, Amazonas, San Martin. mm wide, slightly dentate margins at the base. Sporangia borne on Widely distributed in tropical America, rare in 1-1.5 the Andes. the sporophyll base, reniform, isovalvate, mm wide. The distinctness of the present taxon from the type variety (temperate North America) has been Terrestrial, humid places on white sand, grass- doubted (e.g., Proctor, 1977, pp. 32-33), because lands, and river margins, ca. 1 20 m, Loreto. the distinguishing characters usually indicated for Circum-Amazonian, periphery of the Amazo- them are widely overlapping, and seem to be based nian lowland of Brazil, Peru, Colombia, and Ven- entirely on features related to the size of the plants. ezuela.

Amazonas: Chachapoyas, 1-5 km W of Molinopampa, Loreto: Manfinfa, on the upper Rio Nanay, Ll. Wil- 2400 m, Wurdack 1382 (F, GH, NY, us, USM). San Martin: liams 1 106 (F, us); Vicinity of Iquitos, ca. 120 m, Revilla San LI. Roque, 1350-1500 m, Williams 7763 (F, us). 4340 (F).

62 FIELDIANA: BOTANY 4. Lycopodiella cernua (L.) Pic.-Ser., Webbia 23: by the densely crowded and strongly reflexed leaves 165. 1968. on the main erect axis, by the somewhat flattened branchlet leaves, and by the densely hairy leaf bas-

Lycopodium cernuum L., Sp. pi. 1103. 1753. TYPE: es and stem surfaces. It closely resembles forms

LINN 1 257. 1 3, see Proctor, Ferns ofJamaica, Lon- from similar altitudes in Ecuador (Prov. Zamora- don 29. 1985. Chinchipe) and Bolivia and may represent a dis- Lepidotis cernua (L.) Beauv., Prodr. Aetheogamie 108. 1805. tinct element, but more field observations and ma- Lycopodium cernuum var. capillaceum Spring, Mem. terial are desirable for an assessment of its status.

Acad. roy. Belg. 15 [Mon. Lye. 1]: 80. 1842. TYPE: Venezuela, Edo. Hum- Monagas, Guanaguana, Cajamarca: Prov. Cutervo, La Pucarilla-San Andres, boldt 473 (holotype, B, Herb. Willd. 194291). 2400 m, Sanchez Vega et al. 5999 (AAU, F). San Ignacio, Lycopodium capillaceum (Spring) Hieron., Bot. Jahrb. San Jose de Lourdes, Villarica, 1650 m, Diaz 2053 (F). 34: 573. 1905. Syst. Amazonas: Prov. Bagua, road Chiriaco-Puente Vene- Palhinhaea cernua (L.) Vase. & Franco, Bol. Soc. Brot. zuela, ca. 200-300 m, Barbour 4326 (AAU, MO). Prov. 41:25. 1967. Chachapoyas, ca. 1800 m, Raimondi 350 (USM). San Martin: Prov. Mariscal Caceres, Dist. Tocache Nuevo, Plants with arching-looping runner shoots, with J. Schunke V. 3713 (F, us). La Divsioria, 59 km from Tingo Maria on road to Pucallpa, Allard 22175 (us). up to 1 m tall, erect, dendroid branch systems, Loreto: Prov. Maynas, Dist. Iquitos, near mouth of Rio these with several, subdecussate to alternate, high- Nanay, Rimachi 449 (F, MO, USM). Balsapuerto, 220 m, to ly compound, spreading horizontal, 5-15(-20) Klug 2970 (F, MO, s, uc, us). Huanuco: Prov. Huanuco, cm long lateral branchlet systems. Ultimate Dist. Churubamba, Hda. Mercedes, 1875 m, Mexia 8193 2000 Macbride 5014 branchlets nodding, 3 4(-6) mm in diameter in- (F, GH, s, us). Vilcabamba, m, (F, GH, MO, us). Pasco: Prov. Oxapampa, Palcazu valley, cluding leaves. Branchlet leaves usually borne in 300 m, D. Smith 3927 (AAU). Pichis Trail, Eneiias, 1700 densely crowded, alternating whorls or low spirals m, Killip & Smith 25688 (F, us). Junin: Chanchamayo of 3-5, usually 3-4 mm long, ca. 0.3 mm diameter, Valley, 1200 m, C. Schunke 188 (F). La Merced, Killip acicular, terete to angular (dried), with often con- & Smith 23755 (us). Ayacucho: Prov. La Mar, eastern Massif of the Cord. Central the Cord. Vilca- spicuously acroscopically adnate and decurrent leaf opposing bamba between Tambo, San Miguel, Ayna and Hda. bases, gradually changing from patent-reflexed and Luisiana, 1570 m, Dudley 11889 (GH, us). Cuzco: Pil- distant on main to curved axes, patent, upward copata-Atalaya, 700 m, Vargas 13309 (GH). Puno: Olla- and densely crowded in ultimate branchlets, oc- chea to San Gaban, 1000-2000 m, Dillon et al. 1166 (F, casionally with sparse, lax trichomes or minute MO). spinules; leaf bases often with longer, irregularly or branched these also crisped trichomes, rarely 5. Lycopodiella camporum B. 011g. & P. on stern surfaces. Strobili ses- usually numerous, G.Windisch, Bradea 5: 24, t. 3. 1987. TYPE: ultimate sile, terminating branchlets, 4-10(-20) Brasil, Est. Minas Gerais, Mun. Santana do 2.5-3 in diameter. usu- mm long, mm Sporophylls Riacho, Serra do Cipo, Prado et al. 69 (ho- borne in whorls of ally alternating 5, ovate-deltoid, lotype, HB!; isotypes, AAU!, RB!, SP!, SPF!). short to long cuspidate, ca. 2 mm long, ca. 1 mm with erose-laciniate wide, membranous, coarsely Palhinhaea camporum (B. 011g. and Windisch) Holub, margins. Sporangia globose, 0.5-0.8 mm in di- Folia Geobot. Phytotax. 26: 93. 1991. ameter, strongly anisovalvate. Spores rugate, without a distinct equatorial rim. Plants with arching-looping runner shoots, with

up to 1 m tall, stiffly erect, dendroid branch sys- A common pioneer species on road cuts and tems, these with several, subdecussate to alternate, moist disturbed soil, along rivers, in forest clear- often densely aggregated, stiffly ascending, 5-10 ings, etc., from sea level up to 3000 m, Cajamarca, (-20) cm long lateral branchlet systems. Ultimate Amazonas, San Martin, Loreto, Huanuco, Pasco, branchlets 2.5-7 mm in diameter including leaves,

Junin, Ayacucho, Cuzco, Puno. stiffly ascending to erect, only the strobiliferous Pantropic. ones rather sharply recurved at the tip. Branchlet Where Lycopodiella cernua grows intermixed leaves usually borne in densely crowded, alter- with L. descendens, intermediates may occur. nating low spirals or oblique whorls of 5-7, 2.5- Llanos & Chimouy 61 (USM), from Dept. Ca- 4 mm long, ca. 0.3-0.5 mm in diameter, acicular, jamarca: Prov. Jaen, San Patricio, Santa Rita, terete or angular (dried), sometimes flattened in Chontali, 1 600 m, with the growth habit of L. the lower half, with often slightly acroscopically cernua, deviates from typical forms of the latter adnate and long decurrent leaf bases, patently ar-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 63 cuate-ascending to arcuate-appressed, entirely long-decurrent and acroscopically adnate, hairy leaf smooth to densely hairy on leaf bases, rarely with bases, ascending and upward curved, glabrous, or soft trichomes on leaf margins. Strobili usually with few lax trichomes above the base. Strobili numerous, sessile, terminating tips of ultimate usually numerous, sessile, terminating ultimate branchlets with a sharp bend, to 2.5 cm long, 2- hanging branchlets, 5-20 mm long, ca. 3 mm in 3 mm in diameter. Sporophylls borne in alternat- diameter. Sporophylls usually borne in alternating ing whorls of 5-7, widely ovate, short to long cus- whorls of 5, ovate-deltoid, cuspidate, 1.5-2 mm pidate, 1.5-2 mm long, 0.6-0.9 mm wide, with long, ca. 1 mm wide, with membranous, coarsely scarious, coarsely erose-laciniate margins. Spo- erose-laciniate margins. Sporangia globose, ca. 0.5 rangia globose, ca. 0.6 mm in diameter, strongly mm in diameter, strongly anisovalvate. Spores ru- anisovalvate. gate, without a distinct equatorial rim.

Swamps, moist campo-grasslands, on peaty or Road banks, moist disturbed soil in lower, wet sandy soil, often in the light-open vegetation ad- montane forest, alt. 1000-1 500 m, Amazonas, San jacent to gallery-vegetation at rivers, subject to Martin, Loreto, Huanuco. flooding during the rainy season; alt. 1 20-1 800 m, Lower eastern slopes of the Andes in Peru and San Martin, Junin, Loreto. Ecuador. Amazonian Colombia, Peru and Bolivia; sa- Lycopodiella descendens is closely related to L. vanna region of Venezuela and Guyana, campos cernua, and is sometimes seen growing intermixed vegetation in Brazil. with it. In such situations intermediate individuals Lycopodiella camporum is a widespread species may be found. In addition to the key characters which has been confused with L. pendulina and mentioned, L. descendens very often differs from synonyms thereof. The distinctive branching habit L. cernua by its longer Strobili. and characteristic ecology make L. camporum a rather easily recognizable and ecologically well- Amazonas: Prov. Bagua, Mesones-Muro highway, 286 km E of 8 km E of 650 Hutchi- defined taxon, although detail characters of mor- Olmos, Montenegro, m, son & Wright 3820 (F, uc, us). San Martin: Prov. Lamas, phology do not seem strong. Dist. Lamas, Rio Curiyacu, affluent of Rio Cumbasa, ca. 450 m, Belshaw 3585 (GH, MO, NY, us, us). Prov. Maris- San Martin: Tarapoto, LI. Williams 5960 (F). Junin: cal Caceres, Tocache Neuvo-Juanjui road, km 88, 850 Prov. Satipo, Gran Pajonal, 1000-1 200 m, Pena & Oven- m, D. Smith 2152 (AAU). Prov. Moyobamba, Jepelacio, tini 915 (USM). Loreto: Prov. Maynas, Quisto Cocha, Cerro Shallcahurco, 1320 m, Fernandez & Clemants 27 Iquitos, 120 m, Sagdstegui & Aldave 5821 (GH, MO). (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, 6. Lycopodiella descendens B. 011g., in Harling Hda. Mercedes, 1640 m, Mexia 8195 (F, GH, K, s, uc, and Andersson, Fl. Ecuador 33: 143. 1988. us). SW slope of the Rio Llullapichis watershed, on the TYPE: Prov. B. Ecuador, Pastaza, Mera, 0llg. ascent ofCerros del Sira, ca. 1000 m, Dudley 13125 (GH). & Balslev 9076 (holotype, AAU!; isotypes, F!, QCA!). 7. Lycopodiella glaucescens (Presl) B. 011g., Ope- ra Bot. 92: 176. 1987. Palhinhaea descendens (B. 011g.) Holub, Folia Geo- bot. Phytotax. 26: 93. 1991. Lycopodium glaucescens Presl, Reliq. haenk. 1: 81. 1825. TYPE: Peru (Huanuco), In montanis ad Runner shoots arching-looping, usually densely Huanocco, Haenke (holotype, PRC!). with subulate hairy, relatively distant, appressed, Palhinhaea glaucescens (Presl) Holub, Folia Geobot. leaves, with initially erect, then recurved and de- Phytotax. 26: 93. 1991. scending, amply branched, up to at least 75 cm long dendroid branch systems, these with several Runner shoots long, robust, arching-looping to subdecussate to alternate, highly compound, scandent, with initially erect, amply branched, up spreading to hanging lateral branchlet systems, with to several meters long, bending to long-scandent usually densely hairy main axis with relatively dis- dendroid branch systems, these with several tant, appressed, subulate leaves. Ultimate branch- spreading or nodding to long pendulous, usually lets 3-4 mm in diameter including leaves. Branch- alternate, up to at least 40 cm long lateral branchlet let leaves borne in alternating, densely crowded systems. Leaves of main axes patent to reflexed, whorls or low spirals of 5-6, 24 mm long, ca. 0.3 usually strongly upward curved to hook-shaped, mm thick, subterete, acicular to quadrangular, with 3-5 mm long, up to 1 mm wide, with terete to

64 FIELDIANA: BOTANY quadrangular, hairy leaf bases, apically flattened, Palhinhaea pendulina (Hooker) Holub, Folia Geobot. 26: 93. 1991. usually coriaceous. Ultimate branchlets 3-5 mm Phytotax. in diameter including leaves, the sterile ones often Runner shoots with tapering to less than 2 mm in diameter. Leaves of robust, shallowly arching, to ca. 40 cm erect dendroid branch ultimate branchlets borne in densely crowded, al- up tall, systems, these with several subdecussate to alter- ternating whorls or low spirals of 4-6, acicular to upward to 1 lanceolate, usually with flattened apex, 2.5-4 mm nate, sparsely subequally branched, up 2(-20) cm long, lateral branchlet long, 0.3-0.8(-1.0) mm wide, usually strongly up- usually long pendulous Leaves of main axes ward curved or hook-shaped from a spreading or systems. loosely appressed, 5-7 mm to 1 mm wide. perpendicular to reflexed base, soft herbaceous to upward curved, long, up Ultimate branchlets (3-)5-6(-9) mm in diameter rigidly coriaceous, hairy only on leaf base. Strobili including leaves, to 2 mm in di- variable, 5-30 mm long, 3-4.5(-5) mm in diam- rarely tapering ameter. Branchlet leaves borne in crowded eter. Sporophylls borne in alternating whorls of 5, densely alternating whorls or low spirals of 5-7, acicular, lanceolate-ovate, apically acute-acuminate, 2-3 terete to quadrangular, or sometimes flat- mm long, 0.8-1.2 mm wide, with shallowly den- apically mm to 1 mm tate-ciliate margins, of herbaceous, pale green tex- tened, (3-)4-6 long, up wide, patent- curved, to her- ture throughout. Sporangia globose, strongly an- ascending, upward softly firmly baceous, or with few cilia. Leaf isovalvate, 0.7 mm in diameter. Spores rugate, glabrous, marginal bases and stem surfaces without a distinct equatorial rim. usually glabrous, rarely with short crisped trichomes. Strobili 10-20 mm 5-6 mm in diameter. borne in Clearings and open places in wet upper montane long, Sporophylls whorls of 5-6, lanceolate-ovate, short forest, alt. 1600-3700 m, Cajamarca, San Martin, alternating ca. 1 .5 Huanuco, Pasco, Cuzco. acuminate, (2.5-)3-3.5 mm long, mm wide, with membranous Costa Rica; Colombia to Peru; possibly farther irregularly dentate, narrowly north and south. margins, pale greenish, of herbaceous texture aniso- A remarkable species with often very long scan- throughout. Sporangia globose, strongly ca. 1 mm in diameter. dent dorsally arising shoots. Under favorable con- valvate, Spores rugate, without a distinct rim. ditions, these may reach a length ofseveral meters. equatorial Wet or in the Lycopodiella glaucescens is highly variable with grassland, open spaces clearings montane forest 2200-3600 Piura, respect to strobilus size, size, coarseness, and di- upper and, m, Cuzco. rection of the branchlet systems. Plants ofexposed Amazonas, Huanuco, Costa Rica; Andes from Venezuela to Bolivia, habitats are usually relatively short and compact southeastern Brazil. and with rigid, coriaceous leaves, approaching the is most related growth habit of L. pendulina, while individuals Lycopodiella pendulina closely to L. and from the latter from protected habitats are longer, more lax, and glaucescens, distinguished its erect main shoots with soft and with long pendulous branchlets. by stiffly thick, sparsely ramified, weeping branchlet systems and large Strobili. Cajamarca: Prov. Cutervo, La Pucarilla (San Andres- Socota), 2420 m, Lopez & Sagdstegui 5459 (GH). San Martin: Huallaga. Valley of Rio Apisoncho, 30 km above Piura: Prov. Huancabamba, Loma Redonda (Sapa- Jucusbamba, Hamilton & Holligan 1248 (K). Prov. lache-Chinguela), 2400 m, Sagdstegui et al. 10217 (AAU). Mariscal Caceres, Rio Abiseo National Park, hill past Amazonas: Prov. Chachapoyas, Cerros Calla Calla, E Las Palmas, 2650-2750 m, Young 4000 (USM). Huanuco: side, 1 8 km above Leimebamba on road to Balsas, 3 100 Prov. Huanuco, Carpish, 2850 m, Asplund 13076 (s). m, Hutchison & Wright 5663 (BR, c, E, F, G, GH, M, MO, Pasco: Playapampa, 3000 m, Macbride 4506 (F, G, us). NY, s, uc, USM). Prov. Bagua, Cord, de Colan E of La Prov. Oxapampa, Cord. San Gutardo, 2800 m, Leon Peca, 3400 m, Barbour 3199 (AAU, MO). Huanuco: Prov. 523a (AAU, USM). Cuzco: Prov. La Convencion, Cord. Huanuco, Carpish, 2800 m, Asplund 13105 (s). Chushi, Vilcabamba, 2825 m, Dudley 10782 (USM). trail to Tambo de Vaca, Bryan 681 (F, us). Cuzco: Prov. La Convencion, Cord. Vilcabamba, 2825 m, Dudley 10707, 10782 (GH, MO). 8. Lycopodiella pendulina (Hooker) B. 011g., Ope- ra Dot. 92: 176. 1987.

9. Lycopodiella riofrioi (Sodiro) B. 011g., Opera Lycopodium pendulinum Hooker, Icon. pi. 1: /. 90. Bot. 92: 176. 1987. 1837. TYPE: Peru, Casapi, Matthews 1776 (ho-

lotype, K.!). Lycopodium cernuum L. var. pendulinum (Hooker) Lycopodium riofrioi Sodiro, Crypt, vase. Quit. 582. Baker, Handb. Fern Allies 23. 1887. 1893. TYPE: Ecuador (Prov. Pichincha), In silv.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 65 occid. m. Pichincha ad Gualea, 9/888, Sodiro (ho- Isoetaceae. Heterospory, and the presence of ves- lotype, si 2 1230!). sels in some species of subgenus Selaginella, in- Proc. Biol. Soc. Wash. 89: Lycopodium pensum Lell., dicate the family is specialized. 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!). I. Selaginella Palhinhaea riofrioi (Sodiro) Holub, Folia Geobot. Phytotax., Praha 26: 93. 1991. Selaginella Beauv., Magasin Encycl. 5: 478. 1804; and Prod. Fam. Aetheog. 101. 1805, nom. Growth habit as in Lycopodiella pendulina, up conserv. TYPE: Selaginella spinosa Beauv., to 2 m tall. Branchlet leaves imbricate, densely = nom. nov. for Lycopodium selaginoides L. crowded, borne in alternating whorls of 4-6, lan- Selaginella selaginoides (L.) Link. Figure 9. ceolate to ovate-lanceolate, acuminate, 2.5-4 mm long, ca. 1 mm wide, with densely ciliate-fimbriate Bryodesma Sojak. Preslia 64: 154. 1992. Type: Bry- a terete-suban- margins, with very short, patent, odesma rupestris (L.) Sojak (Lycopodium rupestre = gular base. Stems and leaf bases densely hairy. L.) Selaginella rupestris (L.) Spring. Strobili 1-2.5 cm long, 3-5 mm thick including sporophylls. Sporophylls borne in alternating Terrestrial, rupestral, or rarely epiphytic. Stem whorls of 5, ca. 2.5 mm long, ca. 1 mm wide. slender, branched, sometimes dichotomously, Sporangia 0.7-1 mm in diameter. prostrate-creeping or with ascending branches, or pendent-epiphytic, or erect from a usually stolon- Wet montane forest, in clearings and on road iferous base. Leaves ca. 1.0-10 mm long, with banks, 1700-2600 m. To be expected in north- usually 1 vein, borne in a close spiral or usually ernmost Peru. alternate in 4 ranks. Sporangia large, borne in or Costa Rica; Panama; Venezuela; Colombia to near the axil of a well-differentiated sporophyll, in

1 southern Ecuador. a quadrangular or (in S. deflexa and S . selagi- A very distinctive species with its smooth-ap- noides) a cylindrical strobilus. Megasporangia pearing branchlets covered by closely imbricate, commonly basal in the strobilus, usually with 4 scalelike leaves. 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- Family 27. SELAGINELLACEAE lete, the laesurae % to usually equaling the radius, often with a more or less prominent equatorial Selaginellaceae Milde, Hoher. Sporenpfl. Deutschl. ridge, rugose-reticulate, rugose, papillate, tuber- Schweiz 136. 1865, as Selaginelleae. TYPE: culate, granulate, rarely nearly smooth on the Selaginella Beauv. proximal face. Microspores tetrahedral-globose, trilete, often compressed or the proximal face more Stem indurated or not, branched, bearing rather or less depressed, the laesurae '/2 to equaling the few, often long roots usually at a branch of the radius, usually finely to coarsely echinate, rugose, stem. Leaves simple, ca. 0.5-10 mm long, with 1 papillate, perforate-cristate, or granulate. (very rarely 2) vein(s). Sporangia short-stalked, single, near the axil of a leaf. Heterosporous, spores Selaginella is a large genus of perhaps 700 spe- without chlorophyll. cies, with about 40 of them in Peru, or expected there. The descriptions are drawn from Peru ma- The Selaginellaceae are a distinctive family in- terial and are intended to be parallel within related cluding the single genus Selaginella, only distantly groups of species. The genus has not been ame- related to others such as the Lycopodiaceae and 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, of portion ofbranch, abaxial side, showing axillary and lateral leaves. Selaginella exaltata: e, habit; f, portion 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 (1934), especially the (posthumous) publication recently recognized a second genus, Bryodesma, with A. C. Jermy and J. M. Rankin (1981). The for the S. rupestris group. Koller and Scheckler (1986) paper is among the The genus is probably more diverse in Peru than many that could be cited, but it is selected because presently known. Species are difficult to identify, of the great potential of their work to the classi- and many may appear similar to the field botanist fication of the genus. unacquainted with special specific characters. At the same time, a better understanding of the variation within species and the morphology of ju- References venile plants will lead to an increase in synonyms. The species have been placed in a sequence that, ALSTON, A. H. G. 1934. Notes on Selaginella, presumably, extends from primitive to advanced. VI. J. Dot. 72: 223-226. Species 1-2 are homophyllous, while the remain- ALSTON, A. H. G., A. C. JERMY, AND J. M. RANKIN. der of the species are heterophyllous. 1981. The genus Selaginella in tropical South Species 3-10 are prostrate and not articulate America. Bull. Brit. Mus. (Nat. Hist.) Bot. 9: (both primitive characters), while species 11-12 233-330. are prostrate and articulate. These species relate BAKER, J. G. 1887. Handbook of the fern-allies. to the former group by their small leaves and often George Bell & Sons, London. intricate stems. Species 13-15 have erect, not ar- JERMY, A. C. 1986. Subgeneric names in Selag- ticulate stems that are usually short and pinnately inella. Brit. Fern Gaz. 13: 117, 118. branched. Species 16-25 are also erect and not KOLLER, A. L., AND S. E. SCHECKLER. 1986. Vari- articulate but usually have a main stem with ap- ations in microsporangia and microspore dis- pressed leaves toward the base and flabellate persal in Selaginella. Amer. J. Bot. 773: 1274- branches. Species 1 7 may be unbranched or with 1288. irregular branches. Species 26-30 are prostrate and SOJAK, J. 1992. Generische problematik der Se- articulate, as are species 11-12. However, these laginellaceae. Preslia 64: 151-158. have large leaves and mostly separate stems. Spe- TRYON, R. M. 1 955. Selaginella rupestris and its cies 31-34 are articulate and erect (both derived allies. Ann. Mo. Bot. Gard. 42: 1-99. characters). Species 35 is a climbing species and VALDESPFNO, I. A. 1993. Notes on Neotropical 36 is a highly derived, xeric species. Selaginella (Selaginellaceae), including new spe- The following treatment is based to a large ex- cies from Panama. Brittonia 45: 315-327. tent upon the published work of A. H. G. Alston

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 ofprimary 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 ofthe main stem slender, 0.25-0.50 mm broad; leaves

short, the longest 2 mm long . . 13. S. novae-hollandiae t. Base ofthe 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 ofthe primary branches, forming a somewhat rhomboid arrangement, the apex acute to acuminate 26. S. kunzeana cc. Apical portion of the main stem, and often ofthe 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 extended, 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. The species belongs to the Sartorii series of the 1900. LECTOTYPE (by R. Tryon, 1955): Selaginella rupestris group and appears most closely Brazil, Paria de San Diego Sellow, in 1821 allied to S. sartorii Hieron. Geographically it is (holotype, B; frag., NY). wide-ranging from Mexico and Cuba through South America. It is readily distinguished from the nine

Selaginella rupestris f. amazonica Milde, Fil. Eur. At- others of series S. sartorii and other Peruvian spe- lant. 263. 1867. TYPE: Peru, Cajamarca, (Rio) cies by the glabrous leaf bases, especially by the Maranon, Bonpland (holotype, B; isotype, BM). opaque, milky-white setae on the leaves. This and Selaginella amazonica (Milde) Hieron., Hedwigia 39: S. peruviana have stems that differ from 310. 1900, not S. amazonica Spring, 1840. elongate Selaginella mildei Hieron., in Engler & Prantl, Nat. the radially symmetrical plants of S. convoluta. Pflanzenfam. 1 (4): 671. 1902, nom. nov. for S. amazonica (Milde) Hieron. Lambayeque: Hacienda Valor, Ellenberg 3613 (GH). Cajamarca: between Llaconora and Namora, Correll & Main stem prostrate, creeping, assurgent at the Smith P893 (GH, MO). Between Jaen and Quemado, Fe- rreyra & Sanchez 19668 (USM). Ancash: Mancos, Yun- tip, not articulate, pinnately branched throughout, gay, Tryon & Tryon 6551 (GH). Cuzco: Biies 608 (us); green to brownish, glabrous. Primary branches Herrera 3009a (us). mostly simple, 1 -pinnate, or often 2-pinnate, the ultimate branches as broad as the main stem or 2. Selaginella peruviana (Milde) Hieron., Hed- nearly so, 1-1.5 mm broad including the leaves. wigia 39: 307. 1900. Figure 9a-b. Rhizophores throughout the stems, mostly at the axils of the branches. Leaves similar (homophyl- Selaginella rupestris f. peruviana Milde, Fil. Eur. At- lous), spirally arranged, subulate, long-triangular lant. 263. 1891. TYPE: Peru, Huanuco, Ruiz 98 to broadly to lanceolate, the whit- Ungulate edges (holotype, B; isotype, NY). ish, denticulate, the apex setate, the seta opaque, Selaginella peruviana var. dombeyana Hieron., Hed- milky-white, the base abruptly adnate, ciliate or wigia 39: 308. 1900. PARATYPE: Peru, Dombey 14 (B). ciliolate, rarely lacking cilia, abruptly distinct from Selaginella sheldonii Maxon, Proc. Biol. Soc. Wash. the stem in form and color, glabrous. 31: 171. 1918. TYPE: United States, Quanah Mountains, Oklahoma, Sheldon 233 (holotype, On exposed or wooded, rocky bluffs, or among us). stones, from ca. 300 to 2900 m, Lambayeque, Ca- jamarca, Ancash, and Cuzco. Main stem prostrate, creeping, not articulate, Central Mexico, Cuba, Venezuela, and Colom- pinnately branched throughout, glabrous, green to bia south to Argentina, and eastern Brazil. brownish. Leafy branches forming compact to dif-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 71 fuse mats with discrete, simple or 1 -pinnate, some- 75. 1883. TYPE: Peru, Cerro Canchahuaya, Hu- ber 1421 times 2-pinnate stems. Rhizophores borne (holotype, p). throughout the stems at the axils of branches. Main stem short, often com- Leaves generally similar (homophyllous), spirally creeping, forming pact mats, not articulate, stramineous, arranged, setate, the seta usually more or less lu- glabrous, the basal with and often tescent, or often not well differentiated in color, part spreading leaves, smaller, leaves. the base adnate, glabrous, sometimes denticulate, appressed, ascending Primary branches short, distant, 1 ex- the edges usually ciliate, or sometimes somewhat usually -2-pinnate, to the base of the main 3-5 pubescent, leaves on the underside of the stem tending stem, mm broad the the terminal branches differ from those on the upper side, on the same including leaves, short. from the base of the part of the stem, those on the upper side linear, Rhizophores extending main stem to the of the branches. Lateral acuminate to long-triangular, the lower leaves su- apex leaves above the first branch not bulate, acuminate, broadest at or near the base, to imbricate, broadest at the linear-lanceolate, gradually distinct from the stem. spreading, elongate-ovate, base, 0.5-0.2 mm long, ca. 3 x longer than broad, acute, with a round, ciliolate, basal Mats on bluffs, dry rocky slopes, ledges or crev- scarcely prolonged lobe, without conspicuous whitish borders, the ices of cliffs, exposed or in light shade, igneous, with these denser at the base. calcareous, or sandstone rock, 17504400 m, Ca- edges long cilia, Median leaves broadest at the cen- jamarca south to Puno. suborbicular, the arista '/2 as as the Southwestern United States south to Puebla, ter, aristate, long lamina, the auricles not often with Mexico; Peru to Argentina. cuspidate, prolonged, narrow, whitish borders, the denticulate. The species is recognized as the least specialized edges and basic species within the series Eremophilae, In wet, forests, 100-1100 Loreto in the treatment of the Selaginella rupestris group primary m, and Huanuco. (Tryon, 1955). Geographically it is the widest Costa Rica and Panama; in South America from ranging species of that series. On the basis of the the Guianas south to Peru and Brazil. slender leaves bearing long setae, it is most closely The habit of the with related to S. arizonica Maxon, a species of north- compact, creeping plants lateral branches wide- west Mexico and the states of Texas and Arizona. short, bearing discrete, spread leaves along the main stem form a dis- However, it is distinguished from this as well as tinctive growth pattern. The species was allied with Peruvian selaginellas by the lower leaves that are Selaginella producta Alston et al. (1981), but strongly decurrent on the stem. Among the Pe- by distinguished the strongly tapering, ciliate, more ruvian selaginellas, it is most closely allied with by acute lateral leaves, sometimes on the S. sellowii, another species with homophyllous pubescent surface the that leaves. upper near margin. However, species also has broader, more delicate leafy stems.

Cajamarca: Between Cajamarca and San Juan, Mutter & Gutte 8869 (USM). Ancash: Near Mancos, Tryon & Loreto: Prov. Maynas, Mishuyacu, Klug 1220 (F, us). Tryon 6550 (GH). Huanuco: Prov. Huanuco, Plowman Prov. Maynas, below Iquitos, Moran 3651 (MO). Gami- Schunke 322 Huanuco: Tin- & Rury 11103 (F, GH, USM), Prov. Huanuco, Stork & tanacocha, (F, GH, us, USM). Allard Morton 9388 (F, uc). Lima: Prov. Canta, Punte San Jose, go Maria, 20485 (MO), 20846 (us). Lopez & Riccio 10080 (GH). Prov. Huarochiri, Leon 583 Junin: Above Correll & Smith P748 (USM). Conception, 4. Selaginella brevifolia Baker, J. Bot. 21:83.1883. (GH, MO). Prov. Tarma, litis et al. 108 (GH, wis). Apu- TYPE: Brazil, Janarate Cochoeira, Rio Ne- rimac: Prov. Andahuaylas, Stork & Morton 10719 (F, uc). Cuzco: Near Pisac, McDaniel & Gorski 11402 (GH, gro, Spruce 2547 (holotype, K; isotypes, BM, MO). Urubamba valley, Leon 443 (F, USM). Prov. Uru- CGE). bamba, Chicon, Vargas 11058 (F, uc). Puno: Near Puno, Solomon 2895 (F, MO). Main stem prostrate, creeping, it and its branches forming a mat of stems, sometimes elongate to 3. Selaginella revoluta Baker, J. Bot. 21:141.1883. 15 cm, not articulate, stramineous, glabrous, the TYPE: Venezuela, Amazonas, near Mau- basal part with spreading leaves and often smaller, pures, Spruce 3621 (holotype, K; isotypes, CGE, appressed, ascending leaves. Primary branches 2- us). 3 -pinnate, alternate, distant, extending to the base of the main stem, 2-3 mm broad including the Selaginella demissa Christ, Bull. Herb. Boissier 2, 1: leaves. Rhizophores above the first branch, up to

72 FIELDIANA: BOTANY '/3 the length of the stem. Lateral leaves ovate, in length, 2-3 mm long, oblong to broader near acute, ca. 1-2 mm long, without prolonged auri- the base, the apex obtuse, the base truncate, with cles, ciliate especially on the acroscopic margin, narrow, whitish borders and delicate, long cilia at the cilia usually long, discrete. Median leaves ovate- the edges, especially dense at the base. Median elliptic, acuminate to aristate, the arista '/4 the lam- leaves ovate, the apex acute, sometimes with a ina length, the base with a broad, long, ciliolate short cusp, the base truncate, with whitish borders, auricle, without conspicuous whitish borders, the the edges regularly ciliate or ciliolate, more strong- edges denticulate. ly so at the base.

On soil and among rocks, along roadsides, 400- Epiphytic in dense jungle, or on rotted logs, in 2400 m, Cajamarca, San Martin, and Huanuco. dense forests, 365-700 m, San Martin and Cuzco. Venezuela, Colombia, Peru, and western Ama- Venezuela and Colombia south to Bolivia. zonas, Brazil. The compact, deep green leaves, imbricate on Although the plants are small and mosslike in the stems, give a distinctive aspect to the plants. habit, growing over soil or among rocks, they are Ecological information on collections indicates well collected and known to be wide-ranging in the plants are epiphytic on trees 15 ft above the South America and in Peru. ground. This is corroborated by the dense rhizo- The species is considered close to the Greater phores on the branches and the stems that are free Antillean Selaginella cordifolia but with longer from soil. aristae on the median leaves (Alston et al., 1981). Selaginella applanata A. Braun, Ann. Sci. Nat. It also resembles S. novae- hollandiae, but the Bot. 5, 3: 274. 1865, may be a synonym; uc has strongly prostrate habit with rhizophores above a fragment of an isotype, Peru, Puno, San Gavan, the median part of the main stem contrast with (Gaban) Lechleri 2405. the erect habit and rhizophores, mainly at the base of the stem in S. novae- hollandiae. 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- Cajamarca: Prov. Contumaza, Guzmango, Sagdstegui tias, D. Smith 2000 (F). Cuzco: Prov. Quispicanchis, entre 3932 (GH). Prov. Cajamarca, El Molino, Sagdstegui 7995 Inombaris y Quincemil, Vargas 1 1690 (GH); entre Quin- (F). Contumaza, Lleden, Sagdstegui et al. 10887 (F). San cemil y San Lorenzo, Vargas 16481 (GH). Martin: Boqueron Pass, between Tingo Maria and Pu- calpa, Allard22124 (us). Huanuco: Cushi, trail to Tambo de Vaca, Bryan 718 (F, us). 6. Selaginella tarapotensis Baker, J. Bot. 21: 98. 1883. TYPE: Peru, Mt. Guayrapurima, near Tarapoto, Spruce 4625 (holotype, K; isotypes, 5. Selaginella truncata A. Braun, Index Sem, Hort. BM, CGE, us). Bot. Berol. Appendix 1857: 15. SYNTYPES: Colombia, Cundinamarca, Bogota, Andes of Main stem prostrate or assurgent at the tip, often New Grenada, Karsten (B, BM); Susumuco, forming a mat of stems, not articulate, stramin- Triana 696 (BM), 238 (NY). eous, glabrous, the basal part with spreading leaves, and often smaller, appressed, ascending leaves. Selaginella weberbaurei Knox, Trans. Bot. Soc. Ed- Primary branches 3-4-pinnate, alternate, distant, inburgh 35:282. 1950. Only spores are described. to the base of the main 2-4 mm "TYPE": based on a specimen collected by We- extending stem, berbauer, named by Hieronymus, B?. broad including the leaves, the ultimate branches short. Rhizophores filamentous, dense at the base, Main stem prostrate, assurgent, not articulate, extending nearly to the stem apex. Lateral leaves stramineous, glabrous, the basal part with spread- oblong-lanceolate, but not imbricate on the main ing leaves and often smaller appressed, ascending stem, 0.5-2.0 mm long, approximately more than leaves. Primary branches 1-2-pinnate, pinnately 3 x longer than broad, obtuse to acute, the auricles branched, extending to the base of the main stem, not prolonged, without conspicuous whitish bor- the central branches distant on the main stem, ders, the edges short-denticulate. Median leaves strongly ascending, forming an open, dichotomous rhomboidal to ovate, aristate, the arista slender, division of the branches, the ultimate branches as usually whitish, '/2 as long to nearly equal the length broad as the main stem, 4-6 mm broad including of the lamina, the base not or hardly prolonged, the leaves. Rhizophores slender, threadlike, often with narrow, whitish borders that extend throughout the main stem. Lateral leaves uniform into the whitish arista, the edges denticulate.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 73 Rarely epiphytic, in deep cloud forests, and on or dense, seasonally flooded forest, 130-625 m, shaded banks bordering roads, 500-1 200 m, Ama- Loreto and Huanuco. zonas and San Martin south to Cuzco. Costa Rica, Colombia, Suriname, Ecuador, and Costa Rica south to Bolivia. Peru. The truncate leaf bases on both lateral and me- The flabelliform arrangement of the branches, dian leaves, along with the rhomboid shape and with rhizophores concentrated at the base of the prominent aristata on the median leaves, distin- stem, distinguishes this from other slender- guish the species. This is one of the slender- stemmed species such as S. producta and S. re- stemmed, mat-forming species most closely revoluta that clearly are prostrate, creeping. sembling Selaginella revoluta. Although the species is reported to be wide- Selaginella calosticha Spring, Nouv. Mem. Acad. ranging from Costa Rica to Peru, we have seen roy Sci. Belg. 24: 206. 1849. TYPE: Venezuela, collections only from Loreto, and Huanuco in Peru. Caracas, Funck & Schlim 3321 (K), may be an earlier name for this species. Two Allard collec- Loreto: Prov. Maynas, Calentura, Killip & Smith 29141 tions from Peru 20821 and 21206 are cited as (us). Prov. Maynas, Iquitos, Tryon & Tryon 5195 (F, GH, us, USM). Prov. Maynas, Yanamono Tourist Camp, van S. calosticha in Alston et al. (1981). der Werffet al. 9959 (uc). Rio Itaya, LI. Williams 238 (F, us). Huanuco: Tingo Maria, La Cueva de las Pavas, Amazonas: Prov. Bagua, Montenegro, Hutchison & Allard 20514 (us). Wright 3822 (F, GH, uc, us). Monte Campana, near Tara- poto, Spruce cf4625 (us). San Martin: Tocache Nuevo- 8. Selaginella producta Baker, J. Bot. 21: 243. Juanjui, Smith et al. 2132A (uc). Huanuco: Prov. Tingo 1883. et Maria, Cueva de las Pavas, Aldave & Fernandez 5578 LECTOTYPE (chosen by Alston al., (GH). Pasco: Oxapampa, Cordillera San Matias, Leon et 1981): Brazil, Amazonas, between Barcellos al. 321 (F, MO, USM). Junin: Satipo, Pichanaki, Leon 222 and San Gabriel, Spruce 2043 (holotype, BM; (F, USM). Cuzco: Prov. Paucartambo, Quebrada Quita isotype, CGE). Calzon, Leon et al. 2936 (USM). Achirani, Vargas 11147 (F). Main stem prostrate or assurgent, 1 0-20 cm long, not articulate, several stems usually arising at the 7. Selaginella seemannii Baker, J. Bot. 21: 244. base ofthe plants, stramineous to somewhat green- 1883. TYPE: Colombia, Choco, Cacaqual Is- ish, glabrous, the basal part with spreading leaves land, Seemann 1006 (holotype, K; isotype, BM). and often smaller, appressed, ascending leaves. Primary branches 2-3 -pinnate, with distant sec- Main stem assurgent at the tip, ca. 5-25 cm long, ondary divisions spreading, sometimes forming a not articulate, stramineous, glabrous, the basal part broad, flabelliform arrangement, or with elongate, with spreading leaves, and often smaller, ap- alternatively pinnate branches as long as the main pressed, ascending leaves. Primary branches 2-3- stem or nearly so, 5-10 mm broad including the pinnate, arising a short distance above the base of leaves, the ultimate leafy branches short, unequal the main stem, sometimes gradually reduced, in length. Rhizophores mostly at the stem base, or forming a flabelliform arrangement, the ultimate extending to the central part of the main stem and leafy branches short, unequal, 4-6 mm broad in- branches. Lateral leaves oblong to ovate, spread- cluding the leaves. Rhizophores mostly near the ing, approximate to somewhat imbricate above base to the mid-portion of the main stem. Lateral the first branch, 2-5 mm, usually 4 mm long, leaves elongate-ovate, broadest at the base, broader at the base, obtuse to subacute, the base spreading above the first branch, approximate, truncate, or with an enlarged, round basal lobe, imbricate toward the apex, 2-3 mm long, mem- without conspicuous whitish borders, the edges branous, acute, the base truncate or 1 side enlarged sparsely denticulate with the teeth somewhat dens- and rounded, with narrow, whitish, denticulate er at the base. Median leaves suborbicular, broad- edges, or sparsely denticulate along the base. Me- est at the center, the apex acuminate to subaristate dian leaves elongate-ovate, the apex acuminate to and cuspidate, the arista usually V* or less the lam- subaristate, the arista less than '/4 the lamina length, ina length, the base truncate, with or without nar- the base truncate or with an enlarged, rounded row, whitish margins, the edges evenly denticulate. outer side, with narrow, whitish, denticulate edges, dentate at the base. Terrestrial, or a trunk epiphyte, on rotted logs, or commonly on white sand of the forest floor, On open floor of dense forest, in deep ravines, 100-860 m, Loreto, Huanuco, and Pasco.

74 FIELDIANA: BOTANY Tobago and Trinidad, French Guiana west to form that resembles some species in the Selagi- Colombia and south to Peru and Brazil. nella rupestris group. The species is compared to Selaginella revoluta The species is known from a broad geographic by Alston et al. (1981). However, it differs in range through South America and has been widely broader lateral leaves that have a truncate base collected in Peru. Specimens from southern Co- and lack cilia. The imbricate leaves covering most lombia collected by Bonpland on his travels in of the branches resemble those of S. truncata but South America with Humboldt represent the type. are clearly broader and may have a whitish surface. Cajamarca: Prov. Contumaza, El Tunel, Sagdstegui et al. 12565 (MO). La Libertad: Prov. Pataz, Vista Florida, Leon & Young 1086 (USM). Huancavelica: Prov. Tuya- Loreto: Prov. Maynas, Iquitos, Killip & Smith 27322 caja, Surcubamba, Tovar 3696 (GH). Apurimac: Prov. (GH, us). Iquitos, Klug 198 (F, us). Iquitos, Revilla 4285 Abancay, Vargas 16584 (GH). Cuzco: Prov. Convention, (F, MO). Iquitos, McDaniel et al. 22107 (F). Mishana, Sahuayaco, Biles 834 (us). Urubamba Valley, San Mi- Solomon 3595 (MO). Prov. Maynas, Alpahuayo, van der guel, Cook & Gilbert 1782 (us). Werff et al. 10262 (F, MO). Huanuco: Rio Llullapichis, Cerros del Sira, Wolfe 12264 (GH). Pasco: Prov. Oxa- Foster 9532 pampa, Palcazu, (MO, USM). 10. Selaginella ramosissima Baker, J. Bot. 23: 295. TYPE: Peru, San Martin, near Tarapoto, Spruce 4088 (holotype, K; isotypes, BM, GH, 9. Selaginella microphylla (HBK.) Spring, Bull. NY). Acad. roy Sci. Bruxelles 10: 234. 1843.

Main stem erect, or assurgent at the tip, short, Lycopodium microphyllum HBK., Nov. gen. sp. 1: 37. less 10 not 1816. TYPE: Colombia, Cauca, Qulcace, Bon- than cm long, articulate, greenish, gla- pland (holotype, p?, photo, GH; isotype, BM). brous, the basal part with spreading leaves and often smaller, appressed, ascending leaves. Leafy Main stem short, creeping, usually it and its branches 2-pinnate, ascending, longest in the cen- branches forming a compact mat, not articulate, tral part of the stem, the secondary branches stramineous or greenish, glabrous. Primary straight, ascending, forming a short, deltate ar- branches often intricate, 1-2-pinnate, short, dis- rangement of the stems, ca. 3 mm broad. Rhizo- tant, extending to the base of the main stem, slen- phores mostly at the base extending to the first der, ca. 0.5 mm broad, including the leaves. Rhi- branch. Lateral leaves slender, elongate, broadest zophores abundant at the base, extending nearly at the center, closely placed to somewhat imbricate to the apex of the main stem and branches. Lateral especially on the secondary branches, oblong, 0.5- leaves extending nearly to the base of the stems, 1 .0 mm long, membranous, acute, the base trun- minute, inserted obliquely, 0.25-0.50 mm long, cate, not prolonged, with narrow, whitish borders, ovate or ovate-elliptical, tightly clasping the stem the edges denticulate especially at the base. Me- and enveloping the median leaves when dry, im- dian leaves slender, elongate-ovate, or somewhat bricate, less so on the ultimate branches, the apex broader at the center, membranaceous, acuminate acute with a more or less extended cusp, the bor- to aristate, the arista '/4 or more the length of the ders whitish, ciliolate to fimbriate along the edges, lamina, the base more or less equal, without a especially at the base. Median leaves ovate, about prolonged auricle, the borders narrow, whitish, the half as long as the lateral leaves, subacute, not edges denticulate. Strobilus dorsiventral with 2 auriculate, the edges with regular, dense cilia, the ranks of sporophylls on the upper surface of the border broad, whitish. leafy stem.

Terrestrial in open forest, on soil or among damp Along roadside or bank of stream below the rocks, in deep shade, 800-2700 m, Cajamarca, La forest, 200-2700 m, Cajamarca, San Martin, Lima, Libertad, Huancavelica, Apurimac, and Cuzco. and Cuzco. Costa Rica and Panama, Venezuela along the Ecuador and Peru. Andes south to Argentina, east to southern Brazil, The strobilus is dorsiventral with two ranks of Paraguay and Uruguay. sporophylls on the upper part of the leafy stem. The abundant cilia along the borders of the This readily distinguishes the species from all oth- leaves, and especially the lateral leaves that strong- ers in Peru. The thinner, membranaceous texture ly clasp the dry stems, readily characterize the spe- of the leaves also is a characteristic of the Peruvian cies. The slender, intricate stems have a growth plants.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 75 Cajamarca: Prov. Hualgayoc, Soukup 3872 (us). Lima: Huanuco: Tingo Maria, Croat 21245 (MO, uc). Prov. Cantua, Acleto 758 (USM). Cuzco: Biies in Junio, Churubamba, Mexia 8256 (F, GH, MO, uc, us). Tingo 1930(F). 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 1 1 . Selaginella difTusa (Presl) Spring, Bull. Acad. 5490 (GH). Pasco: Prov. Oxapampa, between Oxapampa Sci. Bruxelles 10: 143. 1843. roy. and Paucartambo, D. Smith 1466 (MO). Ucayali: Bo- queron, Ferreyra 8128 (USM). Cuzco: Prov. La Conven- Rio Lycopodium diffusum Presl, Reliq. haenk. 78. 1825. cion, Mapitunuari, Dudley 11352 (GH, MO). Kos- Sta. TYPE: Panama, Haenke (holotype, PR; isotype, nipata, Isabel, Vargas 23011 (GH). B). vase. 605. Selaginella eggersii Sodiro, Crypt, Quit. 12. Selaginella lingulata Spring, Nouv. Mem. 1893. TYPE: Ecuador, Valle Takatanga, Sodiro Acad. roy. Sci. Belg. 24: 224. 1849. TYPE: (holotype, not found; isotype, us). Jameson Selaginella atirrensis Hieron., in Engler & Prantl, Nat. Ecuador, Pichincha, (holotype, K;

Pflanzenfam. 1 (4), 71 1. 1 90 1 . LECTOTYPE: (by isotype, BM). Alston, 1955), Costa Rica, Cartago, near Atirro, Donn. Smith 5103 (holotype, us; isotype, NY). Selaginella intacta Baker, J. Bot. 21: 335. 1883. TYPE: Ecuador, San Nicolas, Sodiro (holotype, K). Main stem elongate, prostrate, assurgent at the sometimes and tips, compact matlike, articulate, Main stem prostrate, intricate, usually compact, the basal with stramineous, glabrous, part spread- forming a dense mat, articulate, stramineous, gla- often ing leaves, and smaller, appressed, ascending brous, the basal part with spreading leaves and leaves. Primary branches distant, ascending, un- often smaller, appressed, ascending leaves. Leafy to or equally forked, 2-3-pinnate, extending nearly branches 1-3-pinnate, usually pinnately branched, to the base of the 2-4 in- main stem, mm broad distant, extending to the base of the main stem, the the ultimate short. cluding leaves, branches the ultimate branches short, 2-4 mm broad. Rhi- to the Rhizophores hairlike, extending nearly apex zophores extending nearly to the stem apex. Lat- of the branches. Lateral leaves or spaced approx- eral leaves closely placed to distant, 1-3 mm long, not 1-2 imate, imbricate, mm long, elliptical-lan- spreading on the upper part of the stems and pen- ceolate, acute, short, with a prolonged auricle, of- ultimate branches, oblong, acute, the basal lobes ten sometimes with whitish densely ciliolate, more or less equal, not prolonged, with long, dense the ciliolate to en- borders, edges sparsely nearly cilia, without whitish borders, the edges entire. tire. leaves Median elongate-ovate, broadest near Median leaves elongate-ovate, the apex usually the middle, the apex long-acuminate, the base with long-acuminate, the base with 2 equally long au- a auricle that is cilio- prolonged, Ungulate usually ricles, ciliate at the base, without conspicuously with or without a whitish the late, border, edges whitish borders, the edges denticulate above. finely denticulate. On soil or among rocks, in shade, sometimes In moist in on places deep forest, steep banks, forming dense mats on hillsides or on steep banks, on or sometimes logs among rocks, forming large sometimes in deep forests, 1000-1300 m, Pasco mats, 1100-2100 m. Huanuco, and Pasco south and Junin. to Cuzco. Colombia south to Bolivia. Costa Rica; Trinidad; South America east to The long marginal cilia at the base of the leaves and in the Andes south to Bolivia. Suriname, form a small fringe especially on the lateral leaves. Plants to have lateral leaves of than appear more The median leaves are clearly more slender than one form. Most abundant are slender, elliptical the lateral ones and have a long-acuminate apex leaves with entire margins. Other, larger leaves in the material examined. However, the apex of are broader with the base abundantly ciliolate, and the median leaves was considered as acute in the the cilia sometimes extending along the margins treatment by Alston et al. (1981). to the of the leaf. mid-region The ciliolate base The axillary leaves are distinctive, somewhat resembles that of the leaves. median larger than the median and lateral leaves, and more Selaginella atirrensis Hieron. is included here or less ovate, acute, usually with one prolonged although it has been recognized as a distinct spe- auricle and few long cilia. cies on the basis of its acute median leaves. It is not on leaf easily distinguished shape and possibly Pasco: Pichis trail, San Nicolas (as Junin), Killip & intergrades with S. diffusa. Smith 26056 (F, GH, us). Junin: Above San Ramon,

76 FIELDIANA: BOTANY Schunke A251 (us), Chanchamayo Valley, Schunke 230 nuco: Tingo Maria (as San Martin), Allard 21130 (MO, (us), 755 (us). Prov. Tarma, San Ramon, Tryon & Tryon uc). La cueva de las Pavas, Schunke 3257 (F, GH, us). 5448 (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 13. Selaginella novae-hollandiae (Sw.) Spring, Padre Abad (as Loreto), Schunke 3065 (F, GH, us). Huan- Bull. Acad. roy. Sci. Bruxelles 10: 234. 1843. cavelica: Surcubamba, Tovar 3682 (GH). Ayachucho: Ccarrapa, Killip & Smith 22331 (F, GH, us). Apurimac: Prov. Lycopodium novae-hollandiae Sw., Syn. fil. 184, 410. Abancay, Vargas 11083 (GH). Cuzco: Chincheros, 1806. TYPE: "Nova Hollandiae," probably an King et al. 293 (USM). Machu Picchu, Leon 457 (USM). error for Nova Granada, not located. Puno: Prov. Carabaya, Ollachea, Vargas 6914 (us). Selaginella pearcei Baker, J. Bot. 22: 246. 1884. TYPE: Peru, Huanuco, Cordilleras of Pozuzo, Pearce 249 14. Selaginella xiphophylla Baker, J. Bot. 22: 296. (holotype, K). 1884. TYPE: Peru, San Martin, Mt. Selaginella chionoloma Crabbe & Jermy, Amer. Fem Guay- near J. 63: 137. 1973. TYPE: Peru, Cuzco, Cuquipata, rapurima, Tarapoto, Spruce 3990 (ho- Herrera 1636 (holotype, us; isotype, BM). lotype, K; isotypes, BM, CGE, GH, NY).

Main stem short, erect or assurgent, usually less Main stem erect, ascending, 10-14 cm long, not than 20 cm long, not articulate, stramineous, gla- articulate, stramineous to greenish, glabrous, the brous, the basal part with spreading leaves and basal part with appressed, ascending leaves. Pri- often smaller, appressed, ascending leaves, or the mary branches 1-2-pinnate, alternate, distant, ex- basal part with only appressed leaves. Primary tending to, or nearly to the base of the main stem, branches 2-3-pinnate, often forming a complex the lower branches usually longest, the penulti- system of branches broadest at the base, the pen- mate leafy branches 6-12 mm broad. Rhizophores ultimate branches straight or nearly so, 1-3 mm confined to the lower Vs-'/z of the main stem, rarely broad including the leaves. Rhizophores at base of above. Lateral leaves ascending to somewhat ap- the main stem, mostly below the first branch. Lat- pressed, those above the first branch approximate, eral leaves 0.5-2.0 mm long, elliptical to some- 2-6 mm long, oblong-lanceolate, the apex acute, what ovate, acute, not auriculate, the base cilio- the base truncate, with denticulate edges especially late, sometimes with narrow whitish borders, the at the base. Median leaves ovate-lanceolate, grad- edges ciliolate, often densely so. Median leaves ually attenuate, aristate, the arista as long as the ovate, usually more or less elongate, the apex acu- lamina, the base truncate, the outer side ciliolate, minate to aristate, the arista '/4 the lamina length, with narrow, whitish denticulate margins, the teeth the base with a prolonged, Ungulate, ciliolate au- extending onto the arista. ricle, or a somewhat enlarged, round, ciliate base, the borders narrow, whitish, the edges ciliolate or Panama, Colombia, and Peru. denticulate. We have seen only a single collection, the type material from San Martin listed above. The spe- On damp, shaded ledges or crevices of rock fac- cies is also reported from Panama and Colombia es, or moist banks along streams, wooded hillsides, by Valdespino (1993). The small size, erect habit open woods, dense forests, sometimes carpeting of the plants, the rhizophores borne in the basal

l l the forest floor, rarely epiphytic, 400-3600 m, /3- /2 of the main stem, and median leaves with Lambayeque, south to Puno. an arista as long as the lamina are characteristics Nicaragua south to Bolivia and Argentina. by which the species can be recognized. This differs from other slender-stemmed species in articulations on the stems and lacking having 15. Selaginella flagellata Spring, Bull. Acad. roy. generally smaller leaves. It is one of the most com- Sci. Bruxelles 10: 228. 1843. TYPE: French mon of America and is species tropical wide-rang- Guiana, Rio Inini, "source of the Rio Oya- in Peru. ing pok," Leprieur (holotype, LG; isotype, P).

Olmos-Jaen Correll & Smith Lambayeque: Along road, Selaginella regularis Baker, J. Bot. 22: 277. 1884. P826 MO, El et (GH, us). Cajamarca: Molino, Sagdstegui TYPE: Peru, San Martin, near Tarapoto, Spruce al. 7995 (F, MO, uc). Amazonas: Prov. Chacha- Bagua, 3977 (holotype, K; isotypes, BM, GCE). poyas, Hutchison & Wright 5782 (F, GH, MO, uc, us, USM). La Libertad: Prov. Otuzco, road to Paranday, Lopez Main stem erect, short, 5-20 cm the base 1044 (us). San Martin: Rio Sion, Schunke 3494 (F, us). long, Loreto: Balsapuerto, Killip & Smith 28488 (F, GH). Hua- 2-3 cm below the leafy branches, not articulate,

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 77 stramineous, glabrous, the basal part with spread- whitish borders, cilia often dense along the outer ing leaves, and often smaller, appressed, ascending edges. leaves. Primary branches 2-3-pinnate, sessile or subsessile, distant, spreading, the central ones On hillsides and deep ravines of wet forests, longest, the lateral branches ascending in a broad, 700-1 100 m, San Martin and Huanuco. In Peru frondlike arrangement, the penultimate leafy nearly restricted to the area around Tingo Maria. branches straight, 3-5 mm broad. Rhizophores Costa Rica south to Peru. basal, or mostly below the lowest branches. Lat- The plants usually have long primary branches eral leaves 1-3 mm long, elliptic, or broadest at that are 1 -pinnate. This feature and the ciliate lat- the base, attenuate to acute, the base truncate, the eral leaves distinguished the species. borders not conspicuously whitish, the edges denticulate, Median leaves ovate to somewhat elon- San Martin: Prov. Mariscal Caceres, 4 km de Puerto l Schunke 4893 Huanuco: gate, aristate, the arista /4 as long as the lamina, Pizana, (F, us). Tingo Maria, Tryon & Tryon 5255 (GH, us). Prov. Leoncio Prado, the base not auriculate, the borders narrow, whit- Tingo Maria, Plowman & Kennedy 5732 (F, GH, USM). ish or not, the edges denticulate, especially at the Prov. Leoncio Prado, Rupa Rupa, Schunke 10172 (F, base. MO).

Terrestrial, or climbing in cloud forest, among 17. Selaginella chrysoleuca Spring, Bull. Acad. moist rocks and on cliffs, 200-2000 m, San Mar- roy. Sci. Bruxelles 10: 226. 1843. TYPE: Bo- tin, Loreto, and Cuzco. livia, D'Orbigny, (holotype, P). Mexico south to French Guiana and Brazil, and in the Andes to Bolivia. Selaginella sprucei Hooker, Sec. cent, ferns, t. 83. 1861. The short, ultimate branches clearly differ from TYPE: Peru, San Martin, Mt. Campana, near Tarapoto, Spruce 4623 K; BM, other slender, erect-stemmed species in Peru. There (holotype, isotypes, CGE, us). is a general resemblance to Selaginella xiphophylla in the erect habit and rhizophores confined to the Main stem below the first branch short, usually basal part of the main stem. less than 5 cm long, erect, not articulate, greenish, glabrous, the basal part with spreading leaves and San Martin: Juan Jui, Klug 4230 (F, GH, us). Loreto: often smaller, appressed, ascending leaves, or the Pongo de Manseriche, Mexia 6333A (uc). Cuzco: Prov. basal part only with appressed, ascending leaves. Convention, Echarate, Sues 787 (us). 1 x Leafy branches dichotomously divided or 2 , often with 1 longer branch, sometimes with tuft- 16. Selaginella bombycina Spring, Nouv. Mem. like branches arising from a slender creeping run- Acad. roy. Sci. Belg. 24: 191. 1849. TYPE. ner, ultimate leafy branches 5-20 mm, mostly 1 5 Peru, San Martin, Matthews (Mathews) 1781 mm broad, axes ofprimary branches straight. Rhi- (holotype, K). zophores mostly at the base, sometimes extending to l/3 the length of the stem, but not among the Main stem erect, 25-30 cm long, 1 -pinnate, not broad, leafy branches. Lateral leaves spreading articulate, stramineous, glabrous, the basal part above the first branch, imbricate, 5-10 mm long, with appressed, ascending leaves. Primary branch- oblong to linear, 3-5 x longer than broad, the apex es alternate, mostly 1 -pinnate, extending in nearly acute to obtuse, asymmetrical, with the upper (ac- parallel alignment from the first branch to the apex roscopic) edge somewhat larger, round, and finely of the stem, 10-16 mm broad including the leaves, denticulate, with narrow whitish borders, the edg- the axes straight or nearly so. Rhizophores largely es finely denticulate. Median leaves elongate-ovate at the base of the main stem, rarely with a few to suborbicular, asymmetrical, acuminate to aris- among the main branches. Lateral leaves imbri- tate, the arista sometimes Vz the lamina length, the cate on the lateral branches, oblong, slender 5-10 base truncate with a slightly enlarged, round lobe, mm long, 4 or 5 x longer than broad, obtuse, the the borders narrow, whitish, the edges finely den- base with a slightly enlarged, round lobe, usually ticulate. densely ciliate, without conspicuous whitish borders. Median leaves suborbicular, asymmetric, the Among mosses on rocks, in damp soil, on shad- outer side larger, aristate, the arista '/2 to equal the ed slopes of ravines, on moist riverbanks, and in length of the lamina, with a somewhat longer basal dense, dark cloud forests, 400-2000 m, Amazo- auricle, the base ciliolate, without conspicuous nas, south to Cuzco.

78 FIELDIANA: BOTANY Panama, Venezuela to Bolivia. Amazonas: Huampami, Kayap 1311 (uc). Loreto: Be- tween and & Smith 28340 Similarities in the general aspect of the broad, Yurimaguas Balsapuerto, Killip (F, us). Balsapuerto, Klug 2921 (F, GH, MO, us). Prov. leafy branches, in the asymmetrical shape of the Maynas, Iquitos, McDaniel & Rimachi 17473 (GH). Si- median leaves with a arista and dentate mar- long erra del Pongo, Mexia 6283 (F, GH, MO, uc, us). Gamita- gins, suggest an alliance with Selaginella speciosa. nacocha, Rio Mazan, Schunke 109 (F, GH, uc, us).

Amazonas: Prov. Bagua, La Peca, Barbour 2839 (MO). 1 9. Selaginella lechleri Hieron., in Engler & Prantl, Prov. near Hutchison & 3821 Bagua, Montenegro, Wright Nat. Pflanzenfam. 1 (4): 683. 1901. LEC- (F, GH, uc, us). Above Montenegro, Wurdack 1879 (F, TOTYPE (chosen by Alston et al., 1981): GH, uc, us). San Martin: Road from Tarapoto to Yu- Peru, Puno, near San Gavan (San Gaban), rimaguas, Kennedy 3544 (F, us). Between Tarapoto and Lechler 2159 Yurimaguas, Knapp & Mallet 8474 (F, MO). Tarapoto- (holotype, B?; isotype, BM). Yurimaguas road, McDaniel 13805 (GH, MO). Near Tara- 4628 (as Loreto): Between poto, Spruce (GH). Ucayali Main stem erect, 1 2-40 cm long, not articulate, Tingo Maria and Pucallpa, Ferreyra 1008 (USM). Huanu- stramineous, glabrous, the basal part with ap- co: Cerros del Sira, Rio Llullapichis Watershed, Dudley leaves. branches 13360 (GH). Cuzco: Prov. La Convention, Dudley 10355 pressed, ascending Primary (GH, MO). 3-pinnate, forming an extended, flabelliform arrangement, distant, the lower stalked, those above 18. Selaginella speciosa A. Braun, Ann. Sci. Nat. sessile or nearly so, penultimate leafy branches Bot. 5, 3: 274. 1865. TYPE: Colombia, Bo- straight or nearly so, ultimate leafy branches slen- gota, Triana (holotype, B?; isotype, BM). der, 3-5 mm broad. Rhizophores confined to the base of the main stem. Lateral leaves of the main

Selaginella huberi Christ, Bull. Herb. Boissier 2, 1: stem, above the first branch, ascending, coarse, 1901. "entre et Hualla- 73. TYPE: Peru, Ucayali mostly strongly imbricate, 2-5 mm long, oblong, ga," Huber 1547 (holotype, P). acute, the auricles not prolonged, without conspicuous whitish borders, the denticulate to Main stem erect, 30-70 cm long, not articulate, edges short the cilia denser at the base. Median green to brownish, glabrous, the basal part with ciliate, leaves ovate, acuminate, the base some- appressed, ascending leaves. Primary branches di- elongate, times unequal, with a slightly enlarged auricle, not chotomous, long, 1 -pinnate, with 2 or more long or ciliolate, without whitish divisions, the lower ones divided at short intervals sparsely conspicuous borders, the denticulate or entire. forming a frondlike, somewhat flabelliform ar- edges slightly rangement, 10-20 mm broad including the leaves, Terrestrial on clay or sandy soil, along trails or the axes straight. Rhizophores at the base of the on stream banks in forest or rain forests, main stem. Lateral leaves closely placed to im- primary 1 30-1 200 m, Amazonas and Loreto south to Puno. bricate, linear-oblong, to broader above the base, Colombia and Peru. 8-10 mm long, 5-6 x longer than broad, asym- This taxon seems to intergrade with Selaginella metrical, the acroscopic side larger, the apex ob- anceps but is maintained here in its traditional tuse, the base truncate, the borders sometimes sense. The broad, erect stems with appressed leaves, whitish at the base, the edges finely denticulate. the branches a flabelli- Median leaves orbicular to somewhat ovate, primary forming broad, form arrangement, and small size of the leaves on asymmetric with the outer side larger, acuminate the ultimate branches are similar to those of S. to aristate, the arista often nearly '/2 as long as the lamina, the auricles not prolonged, borders nar- anceps. row, whitish, the edges finely denticulate. Amazonas: E ofCinkan, Berlin 254 (F, MO, uc). Loreto: Prov. Maynas, Puca Urquillo, Plowman et al. 6628 (F, In deep, shady forests, in rain forests, in moist us). La Victoria, LI. Williams 2919 (F, us). Pasco: Prov. ravines, rarely epiphytic, 100-600 m, Amazonas Oxapampa, between Puerto Bermudez and San Matias, and Loreto. Leon et al. 333 (F, USM). Puerto Bermudez (as Junin), Killip & Smith 26467 (F, GH). Cuzco: between Mistiana Colombia, Ecuador, and Peru. and Keros, Scolnik 868 (uc). Madre de Dios: Alto Madre This and Selaginella chrysoleuca have excep- de Dios, Rauh & Hirsch PI 570 (GH). Puno: Prov. Caraba- tionally broad leafy branches and asymmetrical ya, Qllachea a Quillabamba, Vargas 17539 (GH). median leaves that are long aristate. Both species

1 occur in dense forest in the Andes, but S . speciosa 20. Selaginella anceps (Presl) Presl, Abh. Bonn. occurs mainly in the north of Peru while S. chry- Ges. Wiss. 5, 3: 581. (Bot. Bemerk., 151). soleuca extends south to Cuzco. 1844.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 79 Lycopodium anceps Presl, Reliq. haenk. 1: 80. 1825. stem erect and simple below the lowest, well-de- TYPE: "Luzon," but probably from Peru, Haenke veloped branch, not articulate, reddish, glabrous, (holotype, PR, photo, GH). the basal part with appressed, ascending leaves. Lycopodium gracile Poiret, in Lam., Encycl. suppl. 3: Primary branches often irregular, mostly 2-pinnate, 551. 1814. TYPE: Peru (Holotype, Herb. Des- ultimate branches not 5 in- vaux P, photo OH). exceeding mm broad, Selaginella gracilis (Poiret) Hieron., Hedwigia 58: 293. cluding the leaves. Rhizophores usually confined 1917, not Moore, 1886. to the base of the main stem, reddish. Lateral leaves strongly imbricate, reddish, especially the stem 20-25 cm or not artic- Main erect, longer, borders, the color extending into the marginal cil- 2-3.5 mm broad at the stramineous, ulate, base, ia, elongate-oblong, 0.25-0.5 mm long, the acro- the basal with glabrous, part appressed, ascending scopic side somewhat larger, acuminate (not aris- leaves. branches an Primary 3-4-pinnate, forming tate), with a rounded, denticulate basal lobe, the elongate, flabelliform arrangement, distant, stalked, upper edges ciliate or ciliolate. Median leaves the branches or the penultimate straight nearly so, elongate-ovate, strongly imbricate, somewhat ultimate branches 1-2 mm broad. leafy slender, asymmetrical, acute to aristate, the base more or confined to the base ofthe main stem. Rhizophores less truncate, the borders whitish, the edges den- Lateral leaves above the first branch, appressed, ticulate, somewhat denser at the base. 2-3 mm long, elongate-ovate, broadest at the base, with a somewhat ciliate acute, prolonged, auricle, On moist riverbanks, steep, wet road banks, without whitish ciliate conspicuous borders, long thickets in dense forests, sometimes forming dark at the base of the leaf. Median leaves ovate-elon- green mounds on the floor of lowland rain forests, gate, acuminate with a somewhat prolonged au- 250-1800 m, Amazonas south to Cuzco. ricle or an ciliolate basal the bor- enlarged, lobe, Costa Rica; Colombia to Bolivia; Brazil. ders not the conspicuously whitish, edges usually Among the species with erect main stems, Se- ciliolate. prominently laginella erythropus is distinguished by the reddish color of the main stem that often extends into the Disturbed and rain forests, primary secondary branches and rhizophores, and by the small size hillsides in wet 1 50-1 500 forests, steep forests, m, of the plants. It also differs from S. haematodes San Martin and Loreto to Cuzco and Madre de by the ciliate or ciliolate lateral leaves. Dios. Costa Rica south to Venezuela and to Bolivia. Amazonas: Prov. Bagua, between Aramango and On the basis of similarities in the small, imbri- Montenegro, Lopez et al. 4212 (GH, MO). San Martin: cate leaves densely covering the ultimate branches Juan Jui, Klug 4267 (GH, MO, us). Tarapoto, Spruce 3989 Junin: Colonia Perene, & Smith 24918 Selaginella anceps appears to be allied to 5. lech- (GH, us). Killip (F, GH, us). La Merced, Macbride 5549 (F, us). Ayacucho: leri. However, the usually denticulate base of the Estrella, between Huanta and Rio Apurimac, Killip & median leaves in S. lechleri is distinct from the Smith 22664 (F, us). Cuzco: Prov. La Convencion, Dud- usually prominently ciliolate one in S. anceps. ley 11438 (GH).

San Martin: Between Tocache Nuevo and Juanjui, 22. haematodes (Kunze) Spring, in Croat 58038 (F, MO). Loreto: Pongo Manseriche, Mexia Selaginella 6371 (GH, MO, uc, us). Huanuco: Pampayacu, Kanehira Martius, Fl. bras. 1 (2): 126. 1840. Figure 118 (GH, us). Pasco: Prov. Oxapampa, Palcazu, Foster 9c-d. & d'Achille 10177 (USM). Junin: Rio Perene, Killip & Smith 25104 (F, GH, us). Perene, Seibert 2193 (F, MO, Lycopodium haematodes Kunze, Linnaea 9: 9. 1834. us). Ayacucho: Estrella, between Huanta and Rio Apu- TYPE: Peru, San Martin, Mission Tocache, Poep- rimac, Killip & Smith 22671 (F, GH, us). Cuzco: Prov. in June 1830 (isotype, K). Convencion, Dudley 11381 (GH, MO). Madre de Dios: pig, Selaginella filicina Spring, Nouv. Mem. Acad. roy. Prov. Manu, Atalaya, Foster & Wachter 7456 (MO). Sci. Belg. 24: 189, 1849. SYNTYPE: Peru, Mat- thews (Mathews) 4036 (K). Other collections from Venezuela are also cited. 2 1 . Selaginella erythropus (Mart.) Spring, in Mar-

tius, Fl. bras. 1 (2): 125. 1840. Plants 30-65 cm long. Main stem erect, not red below the branches, Lycopodium erythropus Martius, Icon. pi. crypt. 39. articulate, bright glabrous, 1 834. TYPE: Brazil, Piaui, Martius (holotype, BR). the basal part with appressed, ascending leaves. Primary branches usually red, ultimate branches Plants 10-25 cm, mostly 10-15 cm long. Main 3-8 mm broad including the leaves, the basal part

80 FIELDIANA: BOTANY with appressed, ascending leaves. Rhizophores near base truncate, the edges sometimes narrow, whit- the base of the main stem, deep red. Lateral leaves ish, finely denticulate. often reddish, oblong to elongate-ovate, broader at the base, 0.25-2.0 mm long, acute, the base In dense forest or rain forest, on slopes, on well- truncate, without conspicuous whitish borders, the drained soils, 450-860 m, Amazonas to Pasco. upper (acroscopic) edge finely denticulate, the teeth Colombia and Peru. longer toward the apex, or nearly entire. Median Among related species with an erect stem and leaves slender, elongate-ovate, acuminate to aris- appressed leaves Selaginella quadrifaria is gener- tate, the arista '/4 or less the lamina length, the base ally distinguished by the median leaves that are truncate, with narrow, whitish borders, the edges equilateral and not or hardly aristate. Differences denticulate especially near the apex. from S. praestans are noted under that species.

Frequent along riverbanks, on steep, moist hill- Amazonas: Prov. Bagua, near Montenegro, Hutchison IB Prov. sides, or scattered on floor of dense forests, pri- & Wright 382 (uc). Bagua, near Campamento, Wurdack 1869 (GH, us). Huanuco: Cerro del Sira, Rio mary forests, rocky hillsides, or along trails, 135- Llullapichis, Wolfl2263B (F, us). Pasco: (as Junin) Puer- 900 m, San Martin and Loreto, south to Madre to Yessup, Killip & Smith 26372 (F, us). Prov. Oxapam- de Dios. pa, valle del Palcazu, Iscozacin, Foster 4556 (F). Cabeza Panama, and Venezuela south to Bolivia. de Mono, near Iscozacin, D. Smith 3752 (uc). The large, deep green, branches borne on long, simple, red-colored stems, readily distinguish the 24. Selaginella praestans Alston et al., Bull. Brit. species. The vivid red color extends into the base Mus. (Nat. Hist.) Bot. 9: 260. 1981. of the main branches and also to the rhizophores that are confined to the basal part of the main Selaginella sprucei A. Braun, Ann. Sci. Nat. Bot. 5, stem. 3: 277. 1865, not Hooker 1861. TYPE: Peru, San Martin, near Tarapoto, Guayrapurina, Spruce 4788 (not 4708 as originally cited) (holotype, G; San Martin: Tocache Schunke 7077 Nueve, (F, us). isotypes, BM, NY). Prov. Mariscal Caceres, Uchiza, Schunke 3202 (F, GH, us). Tarapoto, Spruce 4036 (GH, us). Loreto: San An- Main stem erect, 20-70 cm not articulate, tonio, Rio Samira, Ayala & Arevalo 4227 (F, MO). Puerto long, Arturo, Killip & Smith 27747 (F, GH, us). Cueva de las green to brownish, glabrous, the basal part with Pavas, Fukushima 6521 (F, GH, MO). Huanaco: Prov. appressed, ascending leaves. Primary branches 1- Huanuco, Mexia 8302 (GH, MO, uc). Pasco: Pichis Trail, 2-pinnate, distant, ascending, sparingly pinnate, Killip & Smith 26214 (F). Junin: La Merced, Soukup the basal ones often longer, the axes straight, ul- 2369 (F, GH). Madre de Dios: Cocha Cashu, lake of Rio timate branches 10-15 broad the Manu, Foster et al. 3417 (F). mm including leaves. Rhizophores confined to the base of the main stem. Lateral leaves above the first branches,

23. Selaginella quadrifaria Alston et al., Bull. Brit. ascending, imbricate, those below coarse, elon- Mus. (Nat. Hist.) Bot. 9: 261. 1981. TYPE: gate-ovate to oblong, broadest at the base, 7-8 mm Peru, Prov. Loreto, Sierra del Pongo, Mexia long, 3 or 4 x longer than broad, inequilateral, the 6282 (holotype, BM; isotypes, GH, uc, us). acroscopic side larger, the costa acentric, acute to obtuse, the base truncate, or with a slightly en- Main stem erect, 20-70 cm long, not articulate, larged, ciliolate auricle, without conspicuous whit- stramineous, glabrous, the basal part with ap- ish borders, the edges delicately ciliolate, some- pressed, ascending leaves. Primary branches 3-4- times denticulate, Median leaves suborbicular to pinnate, distant, ascending, usually dichotomously somewhat ovate, asymmetric, inequilateral, the divided, forming a broad, flabellate arrangement of outer side larger, acute, the base truncate, without ultimate branches, 8-20 mm broad including the conspicuous whitish borders, the edges ciliolate or leaves, the axes straight. Rhizophores confined to entire. the base of the main stem. Lateral leaves coarse, rigid, imbricate, 4-8 mm long, oblong or broader In dense shade of deep woods or rain forests, near the base, equilateral, with a central costa, the and along stream sides, 630-1500 m, San Martin, apex acute, the base truncate, with an enlarged, Huanuco, and Pasco. rounded, ciliolate lobe, without conspicuous whit- Colombia to Peru. ish borders, the edges denticulate. Median leaves The species is similar to Selaginella quadrifaria ovate, equilateral, with a central costa, acute, the 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 by Alston et al., 198 1): Peru, Huanuco, Pam- from that species by the inequilateral shape and payacu, Poeppig 195 (holotype, B; isotype, BM, acentric costa of the lateral and median leaves in P). contrast to the equilateral leaves and centric costa in 5. quadrifaria. Main stem prostrate long-creeping to assurgent, articulate, stramineous, glabrous, the basal part San Martin: Valley of Huallaga, between Tocache with spreading leaves and often smaller, ap- Nuevo and Croat 58057 Huanuco: Prov. Juanjui, (MO). pressed, ascending leaves. Primary branches 2-3- Huanuco, Cotirarda, Mexia 8196 (BM, GH, MO, uc, us). pinnate, ascending, mostly widely distant, these as Pasco: Prov. Oxapampa, between Iscozacin and Villa well as the apex of the main stem a rhom- America, D. Smith 2840 (F). forming boid arrangement, with an acute to acuminate apex, ultimate branches 5-8 mm broad the 25. Selaginella haenkeana Spring. Bull. Acad. roy. including leaves. above the lowest Sci. Bruxelles 10: 225. 1843. TYPE: probably Rhizophores extending in the lower half of the stem. Peru, "cordilleris chilensibus," Hdenke (ho- branches, mostly Lateral leaves above the first branch distant, wide- lotype, PR; photo, GH; isotypes, B, us). spreading, imbricate at the stem apex, 2-3 mm long, with somewhat parallel sides, Selaginella dimorpha Klotzsch, Linnaea 18: 523. 1844. oblong, elongate TYPE: "Aufden Anden von Chile," Hdenke (ho- acute, the base with a prolonged auricle forming lotype, B; isotypes, PR, us). Probably the same a round or slender lobe, with narrow, whitish, fine- Peru collection cited above. ly denticulate edges, the teeth especially dense at the base. Median leaves slender, elliptic, the apex Main stem erect, up to 40 cm or more long, long-acuminate to aristate, the arista to '/4 the lam- usually unbranched Vs or more above the base of ina length, the base prolonged with one, some- the main stem, the base stout, 1-3 mm broad, not times two, large, auricles, with narrow, whitish, articulate, to stramineous, glabrous, the greenish finely denticulate edges. basal part with appressed, ascending leaves. Pri- branches 1 -pinnate, sessile or subsessile, dis- mary Trailing on steep banks, in dense woods, often tant, spreading, often arching upward, the central in disturbed areas, 500-1950 m, Huanuco, south ones the ul- usually longest, slender, axes straight, to Cuzco. timate branches not more than 5 mm broad in- Mexico to Panama; Venezuela, Andes of Co- the leaves. or cluding Rhizophores basal, below lombia to Peru. the lowest branch. Lateral leaves 2-3 mm long, A widely distributed species occurring from acute, without with oblong, prolonged auricules, Mexico to Peru, with a broad range in Peru, often long, discrete cilia especially dense at the leaf base. in disturbed places. The rhomboid arrangement Median leaves ovate to somewhat aris- elongate, of the terminal, leafy branches distinguishes this tate, the arista slender, '/4 or less the lamina length, from other articulate, prostrate species. the base not auriculate, the borders narrow, whitish or not, the edges long-ciliate, especially long Huanuco: Prov. Huanuco (as San Martin), Tingo Ma- and dense at the base. ria, Allard 20361 (us); Asplund 12140 (us). Hacienda Mercedes, Mexia 8197 (F, GH, MO, uc, USM). Pasco: south of Pozuzo, 79/269 Divisoria, Ce- Terrestrial, among rocks, or climbing, in cloud Teppner (us). Ucayali: rro Azul, Tryon & Tryon 5272 (F, GH, us, USM). Aya- forest and dense forest, 1 700-2000 m, Amazonas, cucho: Prov. La Mar, southwest of Hacienda Luisiana, and Pasco. Dudley 11721 (GH). Estrella, between Huanta and Rio Colombia, south to Bolivia. Apurimac, Killip & Smith 22663 (GH, us). Cuzco: Prov. Well-developed plants often have an unusual Quispicanchis, Marcapata, Vargas 3178 (MO, us). Kos- nipata, Velarde 7987 (GH). branching system with a long, slender central stem and widely spaced, 1 -pinnate lateral branches 27. Selaginella silvestris Asplund, Ark. f. Bot. 20A bearing short, ultimate divisions. (7): 30. 1926. TYPE: Bolivia, Sur Yungas, El Chaco, Asplund 1140 (holotype, UPS; frag., BM; Amazonas: Prov. Bagua, La Peca, Barbour 2820 (F, MO, uc), 2967 (F, MO). Pasco: Pichis Trail, Dos de Mayo photo, GH). (as Junin), Killip & Smith 25843 (F, GH). Main stem prostrate, long-creeping to decum- 26. Selaginella kunzeana A. Braun, Ann. Sci. Nat. bent, articulate, stramineous, glabrous, the basal Bot. 5, 3: 296. 1865. LECTOTYPE (chosen part with spreading leaves, and often smaller, ap-

82 FIELDIANA: BOTANY pressed, ascending leaves. Primary branches 2-3- pinnate, distant, extending to the base of the main pinnate, often dichotomous, forming a broadly stem, more or less equally divided, in a flabelliform ovate to obdeltate arrangement, the ultimate leafy arrangement, the terminal branches nearly the same branches short, with a round to truncate apex, 4- length, 8-12 mm broad including the leaves. Rhi- 8, usually 5 mm broad including the leaves. Rhi- zophores long, coarse, stramineous, throughout the zophores mostly at the base of the main stem, stems. Lateral leaves above the first branch spread- extending above the first branch. Lateral leaves ing, distant, imbricate only on the ultimate leafy distant, spreading above the first branch, 3-4 mm branches, 6-8 mm long, oblong-lanceolate, the sides long, oblong, somewhat broader near the base, more or less parallel, symmetrical, acute, the auricles asymmetrical, the apex obtuse to acute, the auri- not prolonged, without whitish borders, the edges cles not prolonged, without conspicuous whitish denticulate, sometimes sparingly. Median leaves borders, the edges denticulate, especially at the elongate-ovate, long-acuminate to subaristate, the base. Median leaves elongate-ovate, usually acu- arista less than V* the lamina, the base with a pro- minate, or if aristate the arista less than 'A the longed, Ungulate auricle, the borders narrow, whitish lamina length, the base unequal with 1 prolonged, or not, denticulate along the edges. curved auricle that may be up to % the length of the lamina, the borders not conspicuously whitish, In moist ravines, in secondary forest, primary the edges prominently denticulate. rain forests, in soil among rocks, 730-2700 m, Amazonas, south to Puno. In humid forests, tropical rain forests, wooded Ecuador to Bolivia. ravines or thickets, also among rocks, rarely on The general aspect of the leaves and system of rock walls or epiphytic, 650-3000 m, Cajamarca, branches is similar to those of Selaginella poep- and Amazonas, south to Ayacucho and Cuzco. pigiana and S. articulata, but the lateral leaves in Mexico south to Panama; Venezuela and Co- those species have prolonged auricles that are not lombia south to Bolivia. found in 5". trisulcata. The stem also has two vas- The long-creeping stems with abundant rhizo- cular bundles. phores and prolonged auricles on the median leaves suggest an alliance with Selaginella trisulcata. Amazonas: Prov. Bongara, N of Pedro Ruiz, Smith & Vasquez 4896 (MO, uc, USM). Huanuco: Carpish Pass, Allard 20996 (us). Pasco: Prov. Oxapampa, Smith & Cajamarca: Cueva San Andres, Velarde 6984 (GH). Canne 5824 (MO). Prov. Oxapampa, Canyon de Huan- Prov. Cutervo, Guta de las Huacharos, San Andres, Lopez cabamba, Leon 670 (F, GH). Junin: Carpapata, above & Sagdstegui 5404 (GH). Amazonas: Prov. Chachapoyas, Huacapistana, Killip & Smith 24467 (us). Yaupe, Woyt- Quebrada Molino, Wurdack 636 (F, GH, uc, us); Prov. kowski 6331 (MO, us). Ayacucho: Between Huanta and Bagua, 25 km E of La Peca, Barbour 2962 (F, MO). San Rio Apurimac, Killip & Smith 22751 (F, us). Cuzco: Ca. Martin: Prov. Mariscal Caceresa, Rio Abiseo Nacional 5 km N of Aguas Calientes, Solomon 3162 (MO). Portero, Park, Young & Leon 4957 (HUT, USM). Huanuco: Prov. 8 km W of Quillabamba, Tryon & Tryon 5394 (F, GH, Huanuco, Carpish, Asplund 12842 (us). Pasco: Oxapam- us). pa, Leon et al. 913 (F). Junin: Yucapata, Yaupi, Woyt- kowski 6627 (MO, us). Huancavelica: Prov. Tayacaja, Marcavalle, Tovar4754 (GH). Ayacucho: Ccarrapa, Killip 29. (Spring) Splitg., & Smith 22359 (us). Cuzco: Machu Picchu, Peyton & Selaginella poeppigiana Peyton 1320 (GH, MO); Nunez 7516 (MO, uc). Tijdschr. Naturl. Gesch. Physiol. 7: 443. 1840.

28. Selaginella trisulcata Asplund, Ark. f. Bot. Selaginella sulcata subsp. poeppigiana Spring, Flora 21: 185. 18 38. TYPE: 20A (7): 34. 1926. TYPE: Bolivia, La Paz, El (Jena), Ecuador, Pichincha, Jameson (holotype, K). Chaco, Asplund 1482 (holotype, UPS; photo, GH). Main stem prostrate, long-creeping, ca. 20-40 cm long, articulate, stramineous, sometimes light Selaginella poeppigiana var. peruviana A. Braun, Ann. brown at the the basal with Sci. Nat. Bot. 5, 3: 295. 1865. TYPE: Peru, Puno, apex, glabrous, part near Tabina, Lechler 2015 (holotype, B?). spreading leaves, and often appressed, ascending leaves. Leafy branches 3-4-pinnate, dichotomous, Main stem prostrate, elongate, creeping, only in a subflabellate arrangement, extending to the rarely assurgent at the apex, articulate, pale green base of the main stem, the ultimate branches sim- or sometimes light brown, glabrous, the basal part ilar in length or nearly so, 8-10 mm broad. Rhi- with spreading leaves, and often smaller, ap- zophores at the base of the main stem, extending pressed, ascending leaves. Primary branches 3-4- to the upper branches nearly to the apex. Lateral

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 83 leaves above the first branch distant, spreading to with a prolonged, round auricle, the edges narrow, patent, 4-6 mm, usually 5 mm long, oblong, sym- whitish, denticulate to entire. Median leaves short, metrical or nearly so, acute, the base narrowed ovate, the apex attenuate, the base with 2 pro- with 1 or 2, prolonged, acute or Ungulate, ciliate longed, round auricles, the edges narrow, whitish, auricles, without conspicuous whitish borders, the finely denticulate. edges finely denticulate. Median leaves elongate- ovate, long-acuminate or subaristate, the arista less Plants subrepent, on floor of rain forest, along than 'A the lamina length, the base with a pro- streams and roadsides, 350-2000 m, Amazonas longed, Ungulate, often ciliate auricle, without con- and Loreto south to Pasco and Ucayali. spicuous borders, the edges sparsely denticulate. Panama; Suriname and French Guiana; Peru, Bolivia, Brazil, Paraguay, and Argentina. Creeping, prostrate, forming dense mats on The large, flabellately aligned branches with im- shaded floor ofwet forest, trailing along banks and bricate leaves on the terminal portion of the stems roadsides, or in rock crevices, 900-2800 m, Ama- compose form-flattened sprays that distinguish this zonas south to Puno. from other Peruvian species. The prolonged basal Ecuador and Peru. auricles on the outer side of the median leaves are The long, creeping stems, with abundant rhi- also exceptional. zophores throughout and in a subflabelliform arrangement, suggest alliances with Selaginella tri- Amazonas: Prov. Bagua, Montenegro, Hutchison & 3749 Prov. near sulcata and S. articulata. However, the leafy Wright (F, GH, MO, uc, us). Bagua, Cam- pamento, Wurdack 1892 (F, GH, uc, USM). San Martin: branches are not as broad, and usually more open Prov. Mariscal Caceres, Tocache Nuevo, Schunke 6916 and dichotomous than in those species. (F, MO, uc). Tarapoto, Spruce 4627 (GH, us). Loreto: Prov. Maynas, Poeppig, in 1831 (MO). Pasco: Prov. Oxa- Smith & Salick 8366 Padre Amazonas: Prov. Bagua, La Peca, Harbour 2493 (F, pampa, (MO). Ucayali: Abad, Schunke 3074 MO, uc), 281 9 (F, MO). La Peca, Serrania de Bagua, Gentry (F, GH). 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, 3 1 . Selaginella parkeri (Hooker & Grev.) Spring, McDaniel 10984 Huanuco: Ka- (GH, MO). Pampayacu, Bull. Acad. roy. Sci. Bruxelles 10: 146. 1843. nehira 182 (GH, us). Pasco: Prov. Oxapampa, Rio San Alberto, Leon 645 (F, USM). Junin: Above San Ramon, Lycopodium parkeri Hooker & Grev., Bot. Misc. 2: Killip & Smith 24657 (F, GH); Chanchamayo Valley, 388. 1831. TYPE: Guyana, Demerara, Parker Schunke 192 (F), 194 (F), 231 (F). Cuzco: Prov. Paucar- tambo, Suecia, Woytkowski 175 (USM). Puno: Churu- (holotype, K). pedata Klotzsch, Linnaea 18: 521. 1844. mayo, Soukup 434 (GH). Selaginella TYPE: Guyana, Schomburgk 118 (holotype, B?; isotype, BM). Selaginella fragilis A. Braun, Ann. Sci. Nat. Bot. 5, 3: 30. Selaginella articulata (Kunze) Spring, Flora 305. 1865. TYPE: Brazil, Amazonas, Rio Vaupes, 21: 182. 1838. (Jena), Spruce 2533 (holotype, B?; isotype, BM, CGE). Selaginella brachylepis Christ, Bull Herb. Boissier 2, 1: 74. 1901. "entre et Hua- LycopodiumarticulatumKunze,Linnaea9: 10, 1834. TYPE: Peru, Ucayali TYPE: Peru, Tocache, Poeppig, in 1830 (not lo- llaga," Huber (holotype, P). cated). Main stem erect, 20-25 cm long, articulate, stra- Main stem prostrate, long-creeping, articulate, mineous, or rarely red, glabrous, the basal part stramineous or light brown near the apex, pubes- with appressed, ascending leaves. Primary branch- cent especially at the apex and branch axils, or the es 3-4-pinnate, sessile, the axis fractiflex, the main stem glabrescent. Leafy branches usually branches forming compact, frondlike, flabelliform 3-pinnate, the apex forming a broad, compact, fla- sprays, ultimate branches 5-12 mm broad includ- belliform arrangement, the ultimate branches 10- ing the leaves. Rhizophores at or near the base of 20 mm broad. Rhizophores long, coarse, numer- the main stem, well below the first branch. Lateral 3 ous, extending /4 the length of the stem, stramin- leaves above the first branch, coarse, spreading to eous or golden-colored. Lateral leaves closely somewhat ascending, usually approximate, 4-6 placed to somewhat imbricate at the apex, those mm long, oblong, broader just above the base, on the leafy branches oblong, spreading, approx- acute, the base usually with a prolonged, dentic- imate, 8-10 mm long, oblong, somewhat broader ulate auricle, sometimes 2, 1 an acute lobe, with near the base, nearly symmetrical, obtuse to acute, narrow whitish borders or not, the edges dentic-

84 F1ELDIANA: BOTANY ulate, the teeth somewhat longer and denser at the an extended flabelliform, frondlike arrangement, base. Median leaves lanceolate, the apex long-acu- axis of the primary branches fractiflex, the ulti- minate, the base with a prolonged, peltate, Un- mate leafy branches 2-5 mm broad. Rhizophores gulate auricle, or sometimes 2, without conspic- borne at the base of the main stem, or below the uous borders, the edges finely denticulate. first branch. Lateral leaves of the main stem, above the first branches, coarse, ascending or spreading, Primary forests, sometimes in wet or swampy usually approximate, those below distinct, peltate, forests, on sand or clay soils, usually in deep shade, 2-5 mm long, elongate-ovate, broadest above the 140-650 m, Amazonas and Loreto, south to Pas- base, acute, the base with a short, acute lobe, with- co. out conspicuous whitish borders, the edges den- Colombia and Peru; the Guianas, Amazonian ticulate, especially at the base. Median leaves im- Venezuela; Brazil. bricate, oblong, acute to acuminate with a long, The plants have stems with one vascular bundle, Ungulate auricle, without conspicuous whitish bor- and there are short auricles on the axillary leaves ders, the edges entire or nearly so. similar to those on the median leaves. The long main stems bearing flagelliform branches that ap- On white sand or clay soil in high forest or rain pear somewhat frondlike, and the fractiflex axes forest, usually terrestrial but sometimes on tree of the primary branches clearly suggest alliances trunks, or on rocks, 120-1500 m, Amazonas and between this and S. geniculata. Some specimens Loreto south to Cuzco and Madre de Dios. with unusually attenuated, flagelliform branches Colombia, Ecuador and Peru. have been recognized as a distinct species, Selag- The axillary leaves are truncate; there are two inella fragilis. vascular bundles at the base of the main stem, and there are peltate leaves on the main stem, below Amazonas: Prov. Bagua, Montenegro, Hutchison & the first branch. These characters distinguish the 3821A San Martin: Alto Rio Wright (uc). Huallaga, Juan species from other erect, articulate species of Peru. Jui, King 3831 (F, GH, both mixed with S. anceps). Lo- An alliance with Selaginella parkeri is indicated reto: Prov. Requena, van der Werffet al. 9966 (MO, uc), by the main stem, the of branches, 10108 (MO, uc). H nan urn: Prov. Pachitea, Bosque Na- long alignment fractiflex tional de Iparia, Schunke 2258 (F, GH, us). Pasco: Prov. and the axes of the primary branches; Oxapampa, Quebrada Castilla, Leon & Young 1036 (F, however, the leafy branches are not as broad as in Prov. D. Smith 3735 GH). Oxapampa, Iscozacin, (MO, S. parkeri. uc).

Amazonas: Rio Cenepa, vicinity of Huampami, Berlin 633 (F, MO, uc); Ancuash 1042 (MO, uc). Prov. Bagua, above Cascadas de Wurdack 1894 (F, GH, uc, 32. Selaginella geniculata (Presl) Spring, Bull. Mayasi, us, USM). San Martin: Prov. Mariscal Caceres, Cam- Acad. roy. Sci. Bruxelles 10: 230. 1843. panilla, Schunke 4118 (F, us). Loreto: Between Yuri- maguas and Balsapuerto, Killip & Smith 28250 (F, GH,

Lycopodium geniculatum Presl, Reliq. haenk. 1 : 80. us). Prov. Maynas, between Rio Momon and Rio Mo- 1 825. TYPE: "Luzon," but doubtless Peru or Ec- monicillo, McDaniel 16121 (F, GH, MO). Huanuco: Prov. uador, Haenke (holotype, PR; photo, BM, GH). Huanuco, Vargas 5288 (MO). Pasco: Prov. Oxapampa, Selaginella ferruminata Spring, Bull. Acad. roy. Sci. between Puerto Bermudez and Hito San Matias, Leon Bruxelles 10: 231. 1843, TYPE: Peru, Pangoa, et al. 327 (F, USF, USM). Junin: Satipo, Ridoutt 11397

Matthews (Mathews) 1083 (holotype, K.; isotypes, (USM). Ucayali: Boqueron del Padre Abad, Skog et al. GH, us). 5128 (us). Cuzco: Marcapata, Quispicanchis, Vargas 3781 Selaginella elongata Klotzsch, Linnaea 18: 522. 1844. (MO, us). Madre de Dios: Prov. Manu, Vargas 16643 TYPE: Peru, Cuchero, Ruiz 94 (holotype, B). (GH). Selaginella nodosa Presl, Abh. Bohm Ges. Wiss. 5 (2): 580 (Bot. Bermerk. 50). 1844. TYPE: Peru, Cas- sapi, Poeppig (not located). 33. Selaginella asperula Spring, in Martius, Fl. Selaginella tomentosa Spring, Nouv. Mem. Acad. roy. bras. 1 127. 1840. SYNTYPES: "ad flu- Sci. Belg. 24: 231. 1849. TYPE: Colombia, Cau- (2):

ca, Gorgona Isl., Hinds (holotype, K.). vium Amazonum," Martius (M), "et ad flu- men S. Francisco," Martius (M). Main stem erect, 10-50 cm or longer, articulate, stramineous, glabrous, or sometimes pubescent, Main stem slender, erect, sometimes arching, the basal part with appressed, ascending leaves. and rooting at the apex, usually 1 -pinnate, 1 5-30 Primary branches 3-4-pinnate, sessile above the cm long, articulate, stramineous, glabrous, the basal

lowest branches, forming flat, imbricate sprays in part with appressed, ascending leaves. Primary

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 85 branches distant, often forming tufts of slender, fractiflex, ultimate leafy branches slender, 2-5 mm dichotomous branches along the main stem, broad. Rhizophores abundant on the lower part of sometimes rebranching into uniformly slender ul- the stem nearly to the first branch, or in creeping timate leafy branches 1-3 mm broad, the axes stems among the upper branches. Lateral leaves straight or nearly so. Rhizophores mostly at or near above the first branches distant, ascending, those the base of the main stem. Lateral leaves above below distant, peltate, 14 mm long, lanceolate, the first branch ascending or wide-spreading, 0.5- slightly broader above the base, acute, the base 3 mm long, elongate-ovate, somewhat broaderjust with a prolonged, membranaceous auricle on the above the base, acute, with an auricle sometimes acroscopic side and usually a sharp tooth on the forming a small, acute tooth, the edges narrow, basiscopic side, with narrow, whitish, denticulate whitish, ciliolate, these longer at the base. Median edges. Median leaves very small, usually '/2 the leaves ovate, asymmetrical, acute, with a pro- length of the lateral leaves, ovate-lanceolate, acu- longed, ciliate auricle, the edges narrow, whitish, minate or subaristate, with a prolonged, Ungulate, denticulate, the teeth longer and denser at the base. ciliate basal auricle, with narrow, whitish, denticulate edges. Steep slopes and road banks, open areas, thickets and moist areas in dense forests, 680-1 300 m, In dense stands on clay or often sandy soil in Amazonas and Loreto to Junin. lowland woods or rain forests and secondary for- Amazonian Venezuela and Brazil, south to Bo- ests. Common in humid, disturbed areas, in chac- livia. ras, especially old ones, ca. 100-250 m, Loreto The diffuse branches that form a uniformly slen- and Junin. der, dichotomous, leafy branch system, with small, Guianas, Amazonian Brazil west to Peru. strongly imbricate leaves, are distinctive among We follow Alston (Repert. Spec. Nov. Regni the Peruvian species. The species has a single stele Veg. 40: 309. 1936) in accepting the name Selag- at the base of the erect stem, a character that sep- inella stellata over S. conduplicata, both published arates it from Selaginella geniculata. by Spring in the same work. The species is readily distinguished by the un- Amazonas: Prov. Bagua, 43 km NE of Chiriaco, Bar- usually prolonged, ciliate auricles on axillary, me- hour 4458 San Martin: San LI. Wil- (MO, USM). Rogue, dian, and lateral leaves. The bright stramineous liams 7415 (F, us). Prov. Rioja, Woytkowski 6036 (GH, color of the fractiflex axes contrasting with the MO, us). Loreto: Rio Huallaga. above Lagunas, Croat deep green leaves resembles what is seen in Se- 18097 (F, GH, MO). Huanuco: Tingo Maria (as San Mar- tin) Allard 20823 (MO). Pasco: Prov. Oxapampa, Parque laginella geniculata, and there are peltate leaves Nacional Yanachaga-Chemillen, Leon et al. 989 (F). Ju- on the lower part of the main stem and branches nin: San Ramon, Killip & Smith 24785 (F, GH, us). Prov. similar to that species. Chanchamayo, La Merced, Macbride 5504 (F, us); La Merced, D. Smith 4063 (MO, uc). 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- 34. stellata in Fl. bras. Selaginella Spring, Martius, mitanacocha, Schunke 144 (F, GH, uc, us). Iquitos, Tryon 1 (2): 129. 1840. LECTOTYPE (designated & Tryon 5166 (F, GH, us). Junin: Chanchamayo Valley, La 1117 7775 La Mac- here): Brazil, Prov. Para. Obidos, Martius Merced, Soukup (F), (F). Merced, bride 5503 (F). (holotype, M).

35. Selaginella exaltata (Kunze) Spring, Bull. Selaginella conduplicata Spring, in Martius, Fl. bras. Acad. Sci. Bruxelles 10: 234. 1843. 1 (2): 129. 1840. TYPE: Brasil, Prov. Para, Mar- roy. Fig- tius (holotype, M). ure 9e-f. Selaginella calcarata A. Braun, Ann. Sci. Nat. 5, 3: 305. 1865. TYPE: ?, several syntypes are cited. Lycopodium exaltatum Kunze, Linnaea 9: 8. 1834. TYPE: Peru, San Martin: "ab Uchiza ad Toache," Poeppig, July 1830, Diar. 1953 (not located). Main stem erect to sometimes creeping, 10-30 Selaginella strobilifera Christ, Bull. Herb. Bossier 2, cm long, articulate, stramineous, glabrous, the basal 1: 72. 1901. TYPE: Peru, "entre Rios Huallaga with leaves. part appressed, ascending Primary et Ucayali," Huber 1515 (holotype, P; isotype, branches 3-4-pinnate, usually with an extended, MG). flabelliform, frondlike arrangement, branches distant, the lowest often longest, or in creeping stems Main stem erect, scandent up to 8 m long, ar- the branches of similar length, axes ofthe branches ticulate, 2-6 mm broad including the leaves, stra-

86 FIELDIANA: BOTANY mineous, often pubescent. Primary branches widely imbricate, the base with a prolonged, densely cil- spreading, 2-3-pinnate, often pubescent. Rhizo- iate auricle, the acroscopic side with a broad, whit- phores few, borne in axils of branches. Lateral ish border, the edges irregularly denticulate. Me- leaves distant, clasping on the main stem when dian leaves broadly ovate to somewhat elliptical, dry, approximate, or imbricate on the ultimate obliquely oriented, acute or short aristate, the base branches, obtuse, elongate, 1-3 mm long, 3 or 4 x not prolonged, the borders not conspicuously longer than broad, acute, or sometimes entire, whitish, the edges regularly dentate. somewhat acuminate, the base truncate, sometimes with conspicuous whitish borders. Median On dry desert slopes, 1250 m, Cajamarca. leaves acentric, the outer side larger, ovate, acu- Guatemala; Greater Antilles; Guyana; Peru; minate to aristate, the arista '/4 the lamina length, Brazil and Paraguay. glabrous, the base entire. The habit of these plants forming a rosette of stems, with lateral branches curling inward, readi- In lowland forests, rain forests, secondary for- ly characterizes the species. The leaves are di- ests, and disturbed areas, scandent or scrambling morphic; both lateral and median leaves are over shrubs, often forming dense thickets, 30-760 strongly imbricate and have whitish margins and m, Amazonas south to Madre de Dios. dentate edges. Costa Rica south to Bolivia and western Brazil. Selaginella convoluta is widely distributed in The scandent habit of these plants, often form- Central and South America, but only a single col- ing dense thickets, is exceptional among species lection has been seen from Peru. The inrolled, of Selaginella. The stems are noted to be up to 8 brownish, leafy stems may be easily overlooked m long, but in most collections they are indicated on the dry, desert slopes where the plants occur. as 3 m. The habit of these plants, with dense branches bearing closely placed leaves, forms a Cajamarca: Prov. Celendin, Balsas-Celendin road, Rio Maranon D. Smith 6179 compact cover over other vegetation. valley, (MO).

Amazonas: Prov. Bagua, Montenegro, Sagdstegui et al. 7158 (F, GH, HUT, MO, uc). Prov. Bagua, above Cas- Comments cadas de Mayasi, Wurdack 1800 (F, GH, uc, us). San Martin: between Tocache Nuevo and Juanjui, Croat The following species are accepted by Alston et 58035 (F, MO). Lamas, Naranjal, Knapp & Mallet 6958 al. (1981) as growing in Peru. Adequate material (F, MO, uc). Loreto: Bosque Nacional von Humboldt, Gentry & Revilla 20418 (F, MO). Prov. Maynas, Plowman of these has not been seen and a better evaluation et al. 6502 (F, GH, us). Nanay, Woytkowski 5126 (GH, of them is desired than is now possible. MO, us). Huanuco: Casa de Koepke, Rio Llullapichis, Dudley 12484 (GH). Bosque Nacional de Iparia, Schunke a. Selaginella acanthostachys Baker, J. Bot. 21: 1299 Pasco: Prov. Cabeza (F, GH, MO, us). Oxapampa, 99. 1 883. TYPE: Peru, San Martin, Spruce 4328 de Mono, D. Smith 3768 (MO, uc). Puerto Bermudez (as (holotype, K). Junin), Killip & Smith 26498 (us). Madre de Dios: Prov. b. asplundii Crabbe & Fern Tambopata, Tambopata Nature Reserve, Barbour 4881 Selaginella- Jermy, Gaz. 11: 257. 1976. TYPE: (F, MO). Manu National Park, Gentry et al. 27209 (F, MO). Peru, Huanuco, Asplund 12822 (holotype, s; isotype, BM). c. Selaginella muscosa Spring, in Martius, Fl. bras. 36. Selaginella convoluta (Arnott) Spring, in Mar- 1 (2): 120. 1840. Type: Brazil, Luschnatt (ho- tius, Fl. bras. 1 (2): 131. 1840. lotype, M). d. Selaginella suavis (Spring) Spring, Bull. Acad. Lycopodium convolutum Arnott, Mem. Wernerian Nat. Hist. Soc. 5: 199. 1824 TYPE: Brasil, Rio de roy. Sci. Bruxelles 10: 229. 1843. Selaginella Janeiro, Jameson (holotype, E). sulcata ssp. suavis Spring. Flora (Jena) 21: 185. 1838. TYPE: Brazil, Sellow (holotype, K; iso- Main stem erect, very short, not articulate, green type, B?). to brownish, glabrous. Primary branches forming e. Selaginella wolfii Sodiro, Crypt, vase. Quit. 620. an open rosette curling inward when dry, 1-2- 1893. TYPE: Ecuador, "woods of the western

pinnate, with the ultimate branches, including the region, 1800 m." Sodiro (holotype?, p). leaves, 3-5 mm broad. Rhizophores few at the base of the main stem. Lateral leaves rigid, strongly The following species was credited to Peru on imbricate, 1-2 mm long, ovate-lanceolate or the basis of the type, which is now considered to somewhat elliptic, acuminate to acute, strongly have been collected in Panama.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 87 f. Selaginella horizontalis (Presl) Spring, Bull. with root initiation simultaneous (synchronous) or Acad. roy. Sci. Bruxelles 10: 226. 1843. Lyco- not (acropetal); subula the distal, non-laminate podium horizontale Presl, Rel. haenk. 1: 78. portion of the leaf; ala the lateral, proximal lam- 1825. TYPE: Panama, Haenke (holotype, PR), inate portion of the leaf, not including the ex- not from Peru as originally thought. panded midvein 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 Family 28. ISOETACEAE dormancy and, as new leaves are initiated, are displaced outward, and hence may be found sur- Isoetaceae Reichenb., Hot. Damen Kunst. Freunde rounding the fully developed leaves on a corm; Pflanzenw. 309. 1828. TYPE: Isoetes L. ligule a multicellular, planar, and glandular tissue growing out of a small pit (foveola) in the Stem erect, subterranean, not indurated, bearing proximal portions of the adaxial leaf surface, just numerous, fleshy to wiry, dichotomously branched distal to the sporangium; the ligule is composed roots. Leaves numerous, spirally arranged, ligu- of a thickened central region (the cushion) and late, acicular with a single vein and 4 rows of air peripheral, delicate margins; labium a nonglan- chambers, each transversely partitioned by mul- dular, planar tissue that is apparently a modifi- ticellular septa. Sporangia foliar, large, single, ses- cation of the proximal rim of the foveola; this sile at or near the adaxial leaf base. Megaspores tissue can be quite extensive and may be errone- tetrahedral-globose and trilete, without chloro- ously identified as the ligule (it has also been called phyll, germinating endosporically to form a small, "pseudoligule"); velum a tissue ofleaf derivation short-lived megagametophyte. Microspores ellip- that completely or partially covers the sporangi- soidal and monolete, germinating endosporically um, the latter in most species somewhat impressed to form a small megagametophyte producing 4 within the leaf tissue. multiflagellate spermatozoa.

An ancient genus with representatives known References from the lower Cretaceous; represented today by a single, distinctive, and cosmopolitan genus ex- BRAUN, A. 1864. Les especes d'lle Sardaigne. hibiting distant affinities with Selaginella and Ly- Ann. Sci. Nat. 5: 306-377. copodium. FUCHS, H. P. 1982. Zur heutigen Kenntnis von Species recognition is greatly hindered by tre- Vorkommen und Verbreitung der siidameri- mendous phenotypic plasticity and active speci- kanischen Isoetes-Arten. Proc. Kon. Ned. Akad. ation, both through hybridization and polyploidy Wetensch. CSS: 205-260. as well as through diploid divergence (Taylor & HICKEY, R. J. 1984. Chromosome numbers of Hickey, 1992). Species identificatiop is also ham- Neotropical Isoetes. Amer. Fern J. 74: 9-13. pered by these phenomena but is made even more HICKEY, R. J. 1986. Isoetes megaspore surface difficult by a suite of specialized characters not morphology: Nomenclature, variation, and sys- found elsewhere in the plant kingdom. Unique tematic importance. Amer. Fern J. 76: 1-16. terminology, therefore, is an historic addendum MARSDEN, C. R. 1976. Morphological variation to Isoetes systematics. Most of these novel terms and taxonomy of Isoetes muelleri A. Br., J. Ade- were first introduced by Alexander Braun in 1 864. laide Bot. Gard. 1: 37-54.

The use of them, although inconvenient at first, PAOLILLO, D. J. 1963. The developmental anat- adds clarity and uniformity of understanding. omy of Isoetes. Illinois Biol. Monograph 31:1- Within the body of this treatment I employ them 130. in the following contexts: corm the "stem" of the PFEIFFER, N. 1922. A monograph of the Isoe- plant, that part to which the leaves and roots are taceae. Ann. Missouri Bot. Gard. 9: 79-232. attached; fossa the groove (or one of several) on TAYLOR, W. C., AND R. J. HICKEY. 1992. Hab- the side of the corm (lateral fossa), or extending itat, evolution, and speciation in Isoetes. Ann. all the way around the corm (circumbasal fossa) Missouri Bot. Gard. 79: 613-622. and from which roots are derived; synchronous WEBER, U. 1922. Zur anatomic und systematik roots acropetal roots in Isoetes, roots are initi- der gattung Isoetes L., Nova Hedwigia 63: 2 1 9- ated with a distinctive rhizotaxy (Paolillo, 1963) 262.

88 FIELDIANA: BOTANY I. Isoetes to completely covering the ligule, generally thick and fleshy. Sporangium elliptic to ovate, embed- Isoetes L., Sp. pi. 2: 1 100. 1753. TYPE: /. lacus- ded in the leaf or partially emergent, exposed or tris L. completely covered by the velum. Megaspores laevigate, reticulate, echinate, tuberculate or cristate,

Stylites Amstutz, Ann. Missouri Hot. Gard. 44: 121. 250-1 ,200 Mm in diameter. Microspores laevigate, 1957. S. andicola Amstutz = /. andicola = TYPE: tuberculate, or echinate, 25-50 MHI long. 2n 22- 10. (Amstutz) Gomez. Figure 132.

Aquatic, amphibious or terrestrial perennials. The cosmopolitan genus Isoetes has been vari- Stem to or elon- globose, vertically horizontally ously divided into sections based on megaspore gate, buried in substrate. Leaves terete, trigonal surface morphology. Recent studies, however, (broadly 3-angled) or triquetrous (sharply suggest that spore morphology is extremely vari- 3-angled). Ligule delicate, to ca. 8 mm, glandular, able and highly subject to convergence. auriculate. Labium variable in expression, minute

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 pigmentation basally; corm globose to slightly elongate horizontally; megaspores laevigate to minutely tuberculate or cristate 9. I. boliviensis

1. I soctes panamensis Maxon & Morton, Ann. In ephemeral bodies ofwater such as rain pools,

Missouri Hot. Gard. 26: 272-273. 1939. riverbanks, and marshes, from sea level to 1 500 TYPE: Panama, Prov. Panama, vicinity of m, Tumbes. Bejuco, Woodson et al. 1685 (holotype, us!; Western Guatemala to Panama; coastal Ecua- isotypes, GH!, NY!). dor and Peru; Brazil and Paraguay. Rarely collected.

Isoetes pacifica Svenson, Amer. Fern J. 34: 123. 1944. Throughout its range, this species exhibits con- TYPE: Prov. east of Ecuador, Guayas, Chanduy, siderable variation in the shape, size and distri- Svenson 1 1022 (holotype, BKL; isotypes, GH!, NY!). bution of the bacula on megaspores. Microspores Isoetes savanarum Gomez, Phytologia 49: 339. 1891. also in surface but TYPE: Costa Rica, Prov. Guanacaste, Gomez vary morphology, no parallel 17088, as 7055 (holotype, CR!). trends are evident. The single tetraploid count for this species is from Costa Rican material; it would Corm globose, 7-34 mm in diameter, 2-3-lobed, not be surprising to find distinctive reproductively dichotomous roots arising synchronously within isolated elements within /. panamensis, with the the circumbasal fossa(e). Leaves to 100, erect, 160- Peruvian material presumably assigned to /. pa- 540(-760) mm long, 6-16 mm wide at the base, cifica Svenson. 0.5-1. 5(-2) mm wide at mid-length; alae colorless Isoetes panamensis is the only Peruvian species and hyaline proximally, stramineous to bright green with either triquetrous subulae or baculate mega- or bluish green and chartaceous distally, 2-4 mm spores and as such is a distinctive element in the wide at the sporangium, 35-185(-230) mm long flora. Gomez's description of the ligule in /. sa- ([17-]22-30[-40]% of the total leaf length), apices vannarum refers to the condition of the labium. attenuate; subula triquetrous (sharply 3-angled), stramineous to bright green to bluish green, the Tumbes: South of Zarumilla, Ellenberg 1380 (GH, u). apex attenuate to subacute, with 3 groups of fibrous bundles forming distinct longitudinal ridges, 2. Isoetes novo-granadensis Fuchs, Caldasia 8: continuing to the apex; stomates present; scale 314-315. 1969. TYPE: Colombia, Comisaria leaves present. Sporangium obovate, tan, unspot- del Putumayo, 3250 m, Cuatrecasas 11770 ted, 6-12 mm long, 3.5-7 mm wide, basal. Velum (us!). rudimentary, 0.2-0.5 mm wide along the lateral edges of the sporangium, absent along the upper Isoetes dichotoma Mora-Osejo & Hagemann, Mutisia edge of the sporangium. Ligule delicate, ephem- 43: 5. 1977. TYPE: Colombia, Dept. Narino, a deltate eral, occasionally represented by whitish, Volcan Galeras, 3900 m, Hagemann & Leist 1773 to triangular fragment, 3-4 mm long, 2.5-4.5 mm (COL, HEID!). wide. Labium depressed-ovate, entire, laciniate or shallowly scalloped, 800-2,800 Mm high, 2,500- Corm slightly elongate horizontally to globose, 4,500 Mm wide. Megaspores white, not shiny, 380- up to 15 mm tall and 30 mm in diameter, 2-lobed, = 580 (x 496) Mm in diameter, baculate, the bacula roots arising synchronously within a continuous occasionally laterally confluent; equatorial and circumbasal fossa. Leaves to about 80, stiffly erect, proximal ridges narrowly triangular to deltoid in up to 200 mm long, often with reflexed tips, 6-10 cross-section, equatorial ridges often undulate. mm wide at the base, 2-7 mm wide at mid-length, = Microspores ash-gray, 2740 (x 34) Mm long, dark green to brown proximally, bright to dark = 20-33 (x 26) Mm wide, sparsely echinate, to green distally; alae dark green to nearly black, = papillate, to laevigate. 2n 44. membranaceous, 1.54 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 embedded, basal, hyaline, unspotted, 1-3 mm long, leaf length), parallel-sided, their apices attenuate; 1-2 mm wide. Velum incomplete, extending (10-) subula trigonal (broadly 3-angled in cross-section, 50-75% down the sporangium. Ligule quickly erect to slightly reflexed, heavily cutinized, the apex ephemeral, not seen. Labium not evident. Mega- = sharply acute; fibrous bundles absent; stomates spores reddish brown en masse, 30-36.7 (x 33.2) = absent; scale leaves absent. Sporangium elliptic to Mm long, 25-28.4 (x 26.4) nm wide, shortly ovate, diaphanous to light brown, concolorous, 4- echinate. 9 mm long, 3.5-5 mm wide, basal. Velum rudimentary, represented by a narrow ridge of tissue TYPE Peru, Dept. Ayacucho, Prov. Ayacucho, along the lateral portions of the sporangium, ab- Laguna Yaurihuiri, about 205 km from Nazca on sent along the upper edge of the sporangium. Lig- the road to Abancay, rocky and stony slopes, dom- ule sagittate, brown, 7-9.5 mm high, 3.5-5 mm inant in site with seeping water, 4300 m, Brand- wide, delicate; the margins especially ephemeral, byge 321 (holotype, AAU!). the cushion occasionally persistent as a widely High-altitude seeps, 4300 m, Ayacucho. ovate remnant, ca. 2 mm high x 2.5 mm wide. Known only from the type specimen. Labium obsolete. Megaspores white to light gray, = in not lustrous, 880-1,120 (;c 1,015) m di- 4. Isoetes andicola (Amstutz) Gomez, Brenesia 1 8: ameter, reticulate; equatorial and proximal ridges 4. 1980. Figure lOa-c. straight, distinct, as high or slightly higher than broad, generally obscured by the dense muri. Mi- Stylites andicola Amstutz, Ann. Missouri Bot. Gard. = crospores dark brown, 37-52 (x 47) /urn long, 44: 121. 1957. TYPE: Peru, Dept. Lima, Prov. = = Huarochiri, above in 4750 30-43 (x 37) nm wide, smooth. 2n 132. Casapalca, alpine bogs, m, Amstutz 2000 (holotype, MO). Stylites gemmifera Rauh, in Rauh & Falk, Sitzungs- Cuzco. Wet boggy soil, 3500-3600 m, ber. Heidelberg Acad. Wiss. Math. Naturwiss. Kl. Colombia, Ecuador and Peru. 1959: 11. 1959. TYPE: Peru, Dept. Lima, Prov. above von Known from Peru only by this single collection. Huarochiri, Casapalca, Appen (holotype, HEID). The species is common in moist paramos through- Isoetes andicola var. gemmifera (Rauh) Gomez, Bre- out Ecuador and Colombia where it either grows nesia 18: 4. 1980. as a submerged aquatic or, more commonly, as a terrestrial species deeply embedded in surround- Corm vertically elongate, 20-67 mm tall, 11-15 ing vegetation. The Peruvian material is typically mm in diameter, 1 -lobed (2-3-lobed in juvenile terrestrial, with only the upper half to upper third plants), roots arising acropetally along a single (2- of the leaves protruding above the substrate. 3) lateral fossa(e) (in juveniles arising synchronously within a continuous circumbasal fossa). Cuzco: Prov. Paucartambo, camino peatonal de El Leaves 200 or more, stiffly erect in terrestrial in- Mirador a Cerro Macho Cruz, 3500-3600 m, Leon 2246 dividuals, weakly recurved in submerged individ- (F). uals, 30-50(-75) mm long, 3.5-6 mm wide at the base, 4-7 mm wide at mid-length, dark brown to 3. Isoetes parvula Hickey, sp. nov. black proximally, bright to dark green distally; alae colorless and hyaline to dark brown or black, char- Ab aliis speciebus altitudis celsis statute parvulo, squamis scleroticis praesentibus, et megasporis subfari- taceous proximally, membranaceous distally, 1- nosis differt. 2.8 mm wide at the sporangium, 20-37 mm long (55-85% of the leaf length), each apex obtuse; su- Corm globose, 3-5 mm in diameter, bilobed; bula trigonal, highly cutinized, strongly reflexed in dichotomous roots arising synchronously within terrestrial plants, weakly reflexed to arcuate in sub- the circumbasal fossa. Leaves 8-15, stiffly reflexed, merged plants, the apex attenuate to acute; fibrous 20-30 mm long, 3-4 mm wide at the base, 0.3- bundles absent; stomates absent, scale leaves ab- 0.5 mm wide at mid-length; alae hyaline, 0.5-1 sent. Sporangium elliptic to obovate, superficial to mm wide at the sporangium, 0.5-1 mm long (25- emergent, chocolate-brown, concolorous, 2.5-5.5 50% of the total leaf length), each apex attenuate; mm long, 1.7-3 mm wide, positioned 3.5-9 mm subula half-terete (?), bright green, not highly cu- above the leaf base. Velum rudimentary, repre- tinized, straight to slightly reflexed, the apex at- sented by a narrow ridge of tissue along the lateral tenuate; fibrous bundles not evident; stomates not portions of the sporangium, absent along the upper seen; scales present. Sporangia circular to elliptic, edge of the sporangium. Ligule triangular, weakly

92 FIELDIANA: BOTANY cordate, off-white, 3-4.5 mm high, 2-2.5 mm wide; the total leaf length), each apex attenuate; subula the margin delicate and ephemeral, the cushion trigonal, highly cutinized, straight to slightly re- persistent, deltate, 2-3 mm high, 1 .8-2.3 mm wide. flexed, the apex blunt, subacute; fibrous bundles Labium deltate to depressed-ovate, entire, dark not evident; stomates absent; scales absent. Spo- brown, 280-320 urn high, 180-240 Mm wide. rangia ovate to obovate, essentially superficial, Megaspores mottled, gray on brown, or uniformly basal, light brown, concolorous or occasionally with = brown, not lustrous, 460-720 (x 559) pm in scattered dark spots, 2.5-4.5 mm long, 2-3.5 mm diameter, laevigate to obscurely pustulate, the pus- wide. Velum completely absent. Ligule ephemeral, tules occasionally merging laterally to form short occasionally represented by a dark auriculate frag- meandriform rugae; equatorial and proximal ridg- ment, 2.5 mm long by 2 mm wide. Labium light es straight, distinct, as high or slightly higher than to dark brown, narrowly oblong, often cryptic, 1 40- = broad. Microspores dark brown, 35-43.8 (x 39) 1 80 Mm long, 40-60 fim wide, the apex asymmet- = Mm long, 28.8-37.5 (x 32) pm wide, scabrate. ric, erose. Megaspores gray, not lustrous, laevigate = 2n 44. Vegetative reproduction by cortical gem- to minutely tuberculate, dimorphic, showing Types mae. I and II spores (sensu Marsden 1976), 750-1,125 = (x 920) Mm wide, the proximal and distal ridges In wet, boggy soil or inundated plains of puna laevigate, straight, distinct. Microspores gray en vegetation from 4100 to 4900 m, Lima, Pasco = masse, 37.5-42.5 (x 40.3) urn long, 26.3-32.5 (numerous collections), Junin, Cuzco, Puno. = (x 29.9) wide, laevigate to slightly papillate. Peru and Bolivia. Mm The inclusion of Stylites as a generic synonym of Isoetes is well supported by the early develop- TYPE Peru, Dept. Lambayeque, Prov. Ferre- ment of the sporophyte and the internal anatomy fiafe, Dist. Incahuasi, Jalca, Laguna Tembladera, of the plants (Hickey, unpubl.) as well as the con- Cerro Negro, 3300 m, Skillman et al. 12850 (ho- firmation of the chromosome number as 2n = 44. lotype, MU!; isotypes, F!, HUT!). Earlier reports of 2n = 48-52 are incorrect. High-altitude paramo (aquatic?), 3300 m, Lam- bayeque. Lima: Prov. Huarochiri, Laguna Caprichosa, 4800 m, Known only from the type specimen. near Mina Rauh 186 (M). Lago Aguascocha, Caprichosa, Isoetes dispora is, despite being known only from 4780 m, Hutchison & Tovar 4244 (c, E, F, o, L, MICH, the type collection, a very distinctive species. The MU, s, uc, us, wis). Pasco: Prov. Pasco, Cerro de Pasco, combination of dimorphic, laevigate megaspores, ca. 4300 m, Asplund 11830 (s). Junin: Prov. Jauja, Pa- or less chayo, 3800 m, Ameghino (MU). Prov. Huancayo, La- nearly superficial sporangia, more trigonal guna Condoray, 4700 m, Marshall, 6 Sept. 1961 (BM). subulae, no velum, and a strap-shaped labium dif- Hunter Prov. Junin, 13-14 km N of Junin, Karrfalt & ferentiate it from all other known collections of 22 (USM). Prov. Tarma, Lago Junin, east side near road, Isoetes. The presence of dimorphic megaspores is 13 km north of Junin, 4100 m, Hutchison et al. 5890 (F, taken as an indication of hybridity, often NY, uc, us). Cuzco: In high Andes above Cuzco, toward normally Puno, Gomez AS2189 (CR). Prov. Chumbivilcas, Laguna involving unbalanced genomes. The materials of Huarmicocha, 4600 m, Curlier 241 (USM). Puno: Prov. Skillman et al., however, have certain anomalies Carabaya, de Lacka Macusani, 4360 m, Vargas pampa suggesting that alternative possibilities should be 7128 (us). entertained. Notable among these is the absence of any other collected taxa from the Lambayeque 5. Isoetes dispora Hickey, sp. nov. Figure lOf. region. Hybrids are typically found in lower numbers than the sum of their parental taxa and it Differt ab aliis speciebus Isoetes, megasporis dimor- seems unlikely that three individual Fl hybrids phis necnon laevibus, sporangiis paene superficiaribus, would be the sole collections from a mixed subulis plus minusve triquetris et labiis ligulatis. population of quillworts. Furthermore, there is no ev- Corm globose to slightly elongate vertically, ca. idence of vegetative reproduction in these speci- 20 mm wide, ca. 30 mm high, 2-lobed, dichoto- mens, suggesting that all three have had separate mous roots arising synchronously from a complete origins from spore. In addition, there is a previ- circumbasal fossa. Leaves ca. 30, stiffly erect, dark ously undescribed regularity to the production of brown to dark green proximally, dark green dis- the two spore types in this material. Careful distally, to 65 mm long, 6-8 mm wide at the base, section of individual sporangia showed that each 3.5-6 mm wide at mid-length; alae dark green, sporangium contained 20 spores, ofwhich 1 were chartaceous to sub-membranaceous, 2-3 mm wide large and fully formed and contained cytoplasm, at the sporangium, 10-20 mm long (15-34% of whereas the other 10 were flattened in the equa-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 93 = tonal plane and contained no cytoplasm. Because gray en masse, 28-35 (x 31.7) nm long, 21-28 = of space limitations within the sporangia, the spores (x 24.9) Mm wide, with acute to truncate echinae. derived from each megaspore mother cell retained Vegetative reproduction by cortical budding. their tetrad orientation, and so it was possible to observe directly the end products of all meiotic TYPE Peru, Dept. Cajamarca, Prov. Celendin, events ( 1 total) in two separate sporangia. In all desvio a Guagal siguiendo la ruta a Celendin, 3700 cases tetrads consisted of two large globose spores m, Sanchez Vega et al. 2021 (holotype, MU!; iso- and two smaller flattened ones. Taken together, types, ASU!, F!, CPUN). these data are inconsistent with previous data on High-altitude aquatic endemic, 3700 m, so far Isoetes hybrids, leading us to believe that these known only from Cajamarca. plants are either sexual or apogamous. This species is dedicated to Dr. Walter Hewit- Apogamy in the genus is relatively rare, being son, a student of the fern genus Osmunda and the confined to a few species in Australia and India. man who first introduced me to the excitement of In virtually all known cases ofapogamy in Isoetes, both botany and Isoetes. microspores are either rare or lacking and the A delicate and distinctive species, with the gen- megaspores are irregular in shape, often forming eral appearance of a rigid Isoetes boliviensis, but

Type HI spores or having single contact faces, i.e., with stiffer leaves and no velum. The large mega- diplospores. Such is not the case in Isoetes dispora. spores suggest polyploidy. Additional collections The presence ofnormal microspores (less than 10% should be sought to determine the chromosome abortion) and the extreme regularity of megaspore number of this species and to encompass its vari- production suggest meiotic consistency, albeit ir- ation. regular. The presence of two types of megaspores in this species may, therefore, reflect differential 7. Isoetes saracochensis Hickey, sp. nov. resource allocation among meiotic products in a fashion analogous to that seen during megaspore Species foliis latis, subulus valde trigonis, apicibus acutis velis fere com- development in Selaginella and angiosperms and corneisque, sporangiis basalibus, pletis et megasporis laevibus ad ieniter rugulatis a con- during oogenesis in mammals. generibus diversa.

6. Isoetes hewitsonii Hickey, sp. nov. Corm globose, 2-lobed, 17-23 mm in diameter, with dichotomous roots arising from a continuous Species nova L. dispora proxima, cujus megasporis circumbasal fossa. Leaves 13-14, stiffly erect, 55- laevibus, velo carenti et subulis infirme triquetris habet. 135 6-10 wide 5-7 Differt megasporis uniformibus et foliis angustatis ha- mm long, mm basally, mm bens apicibus attenuatisque minus corneis. wide at mid-length; alae chartaceous, light to dark brown, extending 1.2-3 mm wide at the sporan- Corm globose, slightly elongate laterally, ca. 1 5 gium, 35-75 mm long (46-73% of the leaf length), mm wide, 2-lobed, with fleshy dichotomous roots apices acute; subula highly cutinized and strongly arising from a continuous circumbasal fossa. Leaves trigonal, with 3 distinct, rounded ridges continuing light green, 10-20, erect, 50-160 mm long, 3.5-6 to the apex; leaf apices acute, sharp, corneous; mm wide at the base, 1.5-1.6 mm wide at mid- fibrous bundles absent; stomata absent; scale leaves length; alae light green to hyaline, chartaceous, absent. Sporangium elliptic to ovate, concolorous, 0.9-1.5 mm wide at the sporangium, 23-27 mm 2.5-6 mm long, 2.5-3.5 mm wide, basal. Velum long (36-46% of the total leaf length), each apex virtually complete, extending (%)% or more down attenuate; subula weakly triquetrous, highly cutin- from the upper edge of the sporangium. Ligule ized, straight, the apex attenuate, scales absent. delicate, transparent, broadly to narrowly ovate Sporangia elliptic, embedded, although only with a weakly cordate base, 1 .9-2.7 mm high, 1 .7- slightly so for the megasporangia, light tan, con- 2 mm wide, ephemeral, frequently degraded in colorous, 2.8-4 mm long, 1.5-2 mm wide, basal. mature leaves. Labium light to dark brown, a Velum completely absent. Ligule widely ovate, au- transverse ridge along the lower border ofthe fove- riculate, ephemeral, ca. 1.1 mm high by 0.6 mm ola, entire, 100-150 Mm high, 500-800 Mm wide. = wide, the margins entire. Labium tan to pale green, Megaspores white to gray, 400-620 (x 515) Mm depressed-ovate, ca. 12 Mm high by 54 (j.m wide. in diameter, laevigate or with indistinct to distinct, = Megaspores white, laevigate, 650-750 (x 715) short, low meandriform rugae; equatorial and Mm in diameter; proximal ridges straight, distinct, proximal ridges straight, distinct, with equatorial the equatorial ridge distinct or not. Microspores ridges as broad as high and proximal ridges fre-

94 FIELDIANA: BOTANY quently twice as broad as high. Microspores dark 20-55 mm long (46-75% of the total leaf length), = gray to dark brown, 30-36 (x 33.5) t*m long, each apex attenuate; subula terete, dark green, the = 22-28 (x 24.3) ^m wide, with clavate or echinate apex attenuate to subacute, corneous; fibrous bun- projections. dles absent; stomates absent; scale leaves absent. Sporangium obovate to elliptic, hyaline, unspot- TYPE Peru, Dept. Puno, Laguna Saracocha, ted, 4-19 mm long, 3.5-4 mm wide, basal. Velum

14,000 ft, Tutin 1424 (holotype, BM; isotype, BM). complete to nearly complete, extending 75-100% Submerged or emergent in high-altitude lakes, down the sporangium. Ligule ephemeral, ovate to 3900-4400 m, Cuzco, Puno. very widely ovate, with an auriculate base, 1.2- Endemic. 1.8 mm high, 0.8-2 mm wide, the more persistent This species is fairly well described despite the cushion ovate to widely ovate, ca. 0.8-1 mm high, few collections available. It is distinctive but ob- 0.4-0.6 mm wide. Labium not evident. Mega- viously belongs in an alliance close to 7. lechleri. spores white, laevigate, typically shiny, showing = The extensive velum and the distinctive labium variable abortion, 280-440 (x 340) ^m wide, are shared characteristics that are quite unusual in the equatorial ridges indistinct in larger spores, the the genus for this part of the world. The /. lechleri proximal ridges sharp, distinct. Microspores gray = complex (below), however, is distinctive unto it- in mass, 33.8-38.8 (x 36.2) m long, 26.3-30 = self, having a number of unique characters (hor- (x 28.1) Mm wide, with acute to truncate pro- = izontally elongate corm, turgid leaf condition) that jections. 2n 44. set it apart from 7. saracochensis.

In high-altitude ponds and streams, typically Cuzco: near Sicuna, 3950 m, Tutin 1406 Chectuyoc, submerged, 33004750 m, Cajamarca, San Mar- (BM). Prov. Cafias, Laguna Langui y Layo, Chavez 2313, tin, Ancash, Junin, Cuzco, Puno. 2321 (MO). Puno: Prov. Carabaya, Lake Chungara, 4400 Ayacucho, m, de Macedo & Enriquez (GH). Prov. Lampa, Laguna Peru and Bolivia. Lagunilla, ca. 4300 m, Tutin 1407a, 1420 (BM). The high-altitude ponds, lagunas, and streams of south-central Bolivia and central Peru are re- 8. Isoetes lechleri Mett., Fil. Lechl. 2: 36. 1859. plete with laevigate-megaspored plants that have TYPE: Peru, "in laguna cacuminis, Cordler. the same overall form but that differ from one pr. Agapata," Lechler 1937 (holotype, B!; iso- population to another in the extent of pigmenta-

types, G! [2 sheets], UPS!, frag. s!). Figure lOd-e. tion, the size and rigidity ofthe leaf, and the amount of lateral growth in the corms. Quite a few of these Isoetes soda A. Br., Verb. Bot. Ver. Brandenb. 4: 332. variants have been formally recognized as discrete 1862. TYPE: in cacuminis Peru, laguna (mixta species and even more are now published (Fuchs- cum pi an tula repente ignota), Lechler 1937b (ho- Eckert, 1 982) as nom. nud. Nearly all populations lotype, B!). show some of meiotic as ev- Isoetes glacialis Asplund, Ark. Bot. 20A: 34-35. 1926. degree abnormality TYPE: Bolivia, Dept. La Paz, Prov. Murillo, idenced by irregularly shaped or sized spores. To Jainvags stationem La Cumbre, 4700 m, Asplund date, all of these populations have been found to 4041 (holotype, UPS!; isotypes, G!, M!, s!). = be tetraploids, with 2n 44. The publication Isoetes laevis Weber, Hedwigia 63: 252. 1922. TYPE: = (Hickey, 1 984) of2n 44 for 7. "ticlioensis" Fuchs Peru, Dept. Ancash, Cordillera regia uber Caraz, I. 7. are re- auf dem Grunde eines Tumpels, vollig unterge- ined., glacialis, and herzogii (in part) taucht, 4400 m, Weberbauer 3111 (holotype, B!). ferable to this assemblage. The distinctive color- Isoetes 63: 246. 1922. peruviana Weber, Hedwigia ation, corm shape, velum structure, and ligule form TYPE: Peru, Dept. Junin, Prov. Tarma, Beige strongly suggest a common origin for these plants. westl. von Huacapistana, 3500 m, Weberbauer All of these plants have had a com- 2228 (holotype, B!; isotype, B!; frag., UPS!; photos, undoubtedly s and UPS of B). mon hybrid origin, and it would appear that rapid speciation via reciprocal gene silencing is occur- Corm (globose) laterally elongate, distinctly bi- ring. Each population, or perhaps even subsets lobed [more so in damaged specimens], 1 5-44 mm within a population, is differentially silencing the wide, 2-5 mm high; dichotomous roots arising tetraploid genome down to the diploid level. As synchronously within the circumbasal fossa. Leaves this phenomenon proceeds, genetic isolation slow- to 20-40, stiffly turgid and erect, 88-160(-240) ly builds up and functional spores become rarer mm long, 8-19 mm wide at the base, 2-3 mm and rarer. We are left, then, with a mosaic of vari- wide at mid-length; alae dark green to dark brown, able morphologies, established by drift or perhaps nearly black, 2-3.5 mm wide at the sporangium, selection, which are partially isolated from each

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. other and perpetuated by vegetative reproduction Chanco & Montoya 188 (MU). Pasco (as Junin): Huaron, 4200 11828 G (corm budding). An examination of only a few lake, m, Asplund (c, [2 sheets], NY, s, UPS, us.). Junin: Prov. Yauli, Ticlio, small lake, 4750 m, As- populations gives a false sense of discreteness, and plund 11669 (s). Ayacucho: Prov. Huanta, Mt. Razu- after a broad can one observe the com- only survey huillca, 4000-4100 m, Weberbauer 7500 (F, NY, us). plete intergradation in forms. All of these popu- Cuzco: Prov. Canas, Laguna de Langui y Layo, 3900 m, Chavez 2323 Prov. lations are, therefore, best included within a single (GH). Paucartambo, Leon & Young 2833 (F, USM). Puno: Cordillera Real, Laguna Rinconada, species concept, /. lechleri. 4680 m, Thomasson (s). Sagdstegui et al. 13098 and 14347, while of the general form of 7. boliviensis, have much longer, 9. Isoetes boliviensis Weber, Nova Hedwigia 63: narrower, more slenderly tapering leaves, with 247. 1922. TYPE: Bolivia, circa La Paz, via some pigmentation along the bases. Provisionally, ad Coroico, Lancha, 5000 m, Mandon 1532 it has been identified as 7. lechleri off. Likewise, (holotype, G!; isotypes, BM!, G!). Leon & Young 2833 is provisionally assigned to 7. lechleri. This collection has the darkened leaf Corm globose to slightly elongate laterally, 5- bases and alae, laevigate megaspores, and exten- 1 6 mm in diameter, 2-lobed; dichotomous roots sive velum coverage typical of 7. lechleri. It differs arising synchronously within the circumbasal fos- in having a large (up to 5 mm long), strongly au- sa. Leaves to 40, delicate, flexuous, erect to laxly riculate ligule more typical of 7. andina ofEcuador, spreading, 70-1 20(-240) mm long, 6-1 6 mm wide Colombia, and Venezuela and in having emargin- at the base, 1-1.5 mm wide at mid-length; alae ate ala apices reminiscent of those seen in 7. kar- hyaline to light brown, 1-2 mm wide at the spo- stenii (s.l.) of Colombia and Venezuela. rangium, 20-30 mm long (1 2-33% of the total leaf The species is best distinguished by its brownish length), each apex attenuate (rarely truncate); su- (almost caramel) coloration and the distinctly te- bula terete, bright green, the apex attenuate; fi- rete leaves that are turgid to the point of being brous bundles absent; stomates absent; scale leaves brittle. Herbarium material is best identified by absent. Sporangium obovate to elliptic, hyaline, coloration, the presence of a horizontally elongate unspotted, 5-8 mm long, 3-5 mm wide, basal. corm (in most populations), the laevigate mega- Velum incomplete to complete, extending 6-100% spores, the more or less complete velum, and the down the sporangium. Ligule ephemeral, widely widely ovate ligule which has a tendency toward ovate, with an auriculate base, ca. 2 mm high, 1.7 an auriculate base. mm wide, the more persistent cushion narrowly Isoetes lechleri is a plant of high-altitude (over depressed-ovate, often bilobed, ca. 0.2 mm high, 3500 m) lakes, streams, and pools. It grows in both 0.6 mm wide. Labium not evident. Megaspores shallow and deep water, with an individual plant's white to light gray, laevigate to minutely tuber- form being strongly influenced by water depth. culate, shiny, 380-460 (x = 433) /nm wide, the Shallow plants have short, stiffly rigid leaves. Plants equatorial and proximal ridges sharp, distinct. Mi- of deeper water "etiolate": the leaves are a bit = crospores gray in mass, 26.3-40 (x 33) um long, more lax and considerably longer. Vegetative re- 20-30 (x = 25.3) /mi wide, with acute to truncate production, by corm budding, is very common projections. 2n = 22. and may be a primary means of reproduction. The plants often form dense mats in quiet water; in In shallow water, edges of lakes, ponds, and ver- streams and shallows the plantlets are often broken nal pools, 4100-5000 m, Cajamarca, San Martin, off and transported by animals or wave/current Ancash, Lima, Pasco, Junin, Ayacucho, Cuzco, action. In Bolivia one between two road, cutting Puno. lakes, was covered washed-out completely by plants Peru and Bolivia. and corms. A very distinctive plant due to its small and relatively delicate appearance; several collectors Cajamarca: Pozo Kuan, 3790 m, Sagdstegui et al. have even suggested that it is an annual. This is 13098 & 14011 (F, HUT, MU). Prov. Contumaza, Pozo the only Peruvian species with lax, flexuous leaves. Kuan, 3900 m, Sagdstegui et al. 14347 (F, HUT, MU). San Martin: Prov. Mariscal Caceres, forest on the edge It has a partial to complete velum normally cov- of Laguna de Chochos, NW corner of Rio Abiseo Na- ering about 75% of the sporangium, corms that tional Park, 3300 m, Young & Leon 4856 (MU). Ancash: are slightly elongate horizontally, and megaspores Prov. Huari, Huascaran National Park, 4440-4490 m, that are laevigate to minutely tuberculate. It is D. Smith et al. 12355 (HUT). Prov. Yungay, Laguna de in of Llanganuco, 4100-4200 m, Enderlin (USM). Lima: Prov. found shallow (= ephemeral?) portions ponds Cajatambo, Raura, perimetro de Ada, 4800-4900 m, and lakes. The microspores are quite distinctive

96 FIELDIANA: BOTANY in appearance, having acute or club-shaped pro- nearly complete velum, and has megaspores that jections which themselves are spiny. Microspores vary from tuberculate to rugulate to nearly lae- of some /. lechleri collections are similar. vigate. Isoetes herzogii is a high-altitude, nearly = The only counts for this species, 2n 22, are emergent plant of streams and shallow pools. from Bolivian material.

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). Addendum Lima: Prov. Huarochiri, Ticlio Pass, between Lima and Oroya, 4840 m, Hutchison et al. 6081 (c, E, F, G, GH, 1. to Be Added to the HEID, L, LIL, M, MICH, MO, NY, s, uc, wis). Junin: 5 km Species Pteridophyte above Hacienda Cochas, 34 km W of la Oroya-Huan- Flora, Parts I-V cayo Route 3 on road to Pachacayo, 4425 m, Keeley & Keeley 11087, 11088 (MU, occ). The following species have come to the atten- tion of the authors as growing in Peru, after their treatment had been Comments generic published.

Trichomanes arbuscula Desv., Mem. Soc. Linn. Isoetes triquetra A. Braun, Verh. Bot. Vereins Prov. Paris 6: 326. 1827. TYPE: "Crescit in Guia- Brandenburg 3 (4): 332-333. 1862. TYPE: na," P, Herb. Desvaux. Peru, Sachapata (am ostlichen Abhang der Cordillera von Peru), in pascuis humidis, Trichomanes coriaceumKunze, Linnaea9: 105. 1834. Lechler 3337 (holotype, B!; photos, s and UPS TYPE: "Prope Collares, Brasiliae," Poeppig in ofe). 1832 (Diar. 2981), not located.

Calamaria triquetra (A. Braun) Kuntze, Rev. gen. pi. This species has an alate rachis and petiole and 2:828. 1891-1893. is sometimes subdimorphic. Illustrations are Isoetes lechleri var. triquetra (A. Braun) Gomez, Bre- Hooker & Icones t. 204. 1831 nesia 18: 5. 1980. Greville, Filicum, (as T. bancroftii) and Vareschi, Flora de Vene-

zuela, Vol. 1 (Helechos), t. 47. 1969. The type material for this species is very frag- mentary. There are no megaspores and the micro- Loreto: Prov. Maynas, Puerto Almendras, van der Werff spores are of a very common form (echinate). The et al. 9845 (MO), 9795 (MO), det. K. U. Kramer. most distinctive features of the specimen are the trigonal subula, the moderate labium, and the large Trichomanes crispumL., Sp.pl. 1097. 1753. TYPE: auriculate ligule. In these regards, the type of L. "Martinica," Plumier, Traite foug. Amer., t. triquetra is nearly identical to /. andina of Ecua- 86. 1705. dor, Colombia, and Venezuela but differs from all other known Peruvian collections. I have argued This is a critical and polymorphic species (see (Taxon 35: 243-246. 1986) for the conspecificity Windisch, Bradea 6: 99. 1992 for discussion). of these taxa. The major argument against the common identity is distribution (some 1 ,400 km Huanuco: Prov. Pachitea, region of Pucallpa, Wall- separate the type locality of I. triquetra from the nofer 11-261088 (z), det. K. U. Kramer. nearest "good" locality of /. andina).

Pterozonium paraphysatum (A. C. Sm.) Lell., Mem. Isoetes 63: 250. 1922. herzogii Weber, Hedwigia New York Bot. Card. 17: 13. 1967. TYPE: Bolivia, Tunari, 4300 m, Herzog2083 (holotype, M; isotypes, L!, M!, us!; frag., UPS!). Syngramma paraphysata A. C. Sm., Bull. Torrey Bot. Club 58: 301. 1931. TYPE: Venezuela, Amazo- Although not recorded from Peru, this species nas, Cerro Duida, Tate 441 (NY; frag., us). is geographically proximate in Bolivia and should

be expected in Peru. The species has stiffly erect, This species is distinguished by the scales (vs. narrow, subulate leaves with attenuate apices. It trichomes) on the stem, the keeled lamina, and lacks the pigmentation typical of /. lechleri, has a relatively distant veins.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 97 Huanuco: Prov. Pachitea, region of Pucallpa, low, elfin partments that have the most endemics are those 1750 110-13988 det. K. U. forest, m, Wallnofer (z), with the most species. This suggests that disrup- Kramer. tions of ranges during the Pleistocene may have occurred throughout the Andes of Peru. However, Lindsaea phassa Kramer, Bot. Helvetica 101: 207. there is need for more detailed knowledge of the 1991. TYPE: Peru, Loreto, Rio Ampiyaco, Pleistocene in the Andean region, such as that for vicinity of Pucaurquillo, Davis et al. 849 (ho- Europe published by Lang (1992). lotype, uc). Paratypes: Loreto: Prov. Maynas, There are few studies oflocal diversity, although Yanamono Explorama tourist camp, van der the ecological work of Young and Leon (1989, Werffet al. 9869 (MO, z), 7994 (MO, z). 1991) on pteridophytes of Peru supplies critical new data. Their analysis ofa low elevation forested Kramer provides a discussion of the relation- area in central Peru, including 61 species within

1 2 that and ship of this new species and an illustration (fig. , hectares, suggests edaphic topographic p. 228) of it. gradients affect the diversity of species. Their work on a high elevation forest, on the east slope of the 2. Consideration of Pteridophyte Diversity in Peruvian Andes, including 1 09 species in about 5 Respect to Ecology and Geography square km, shows greatest pteridophyte diversity in the montane rain forest. They suggest that the This treatment of the pteridophytes of Peru in- high and nearly constant humidity is a factor help- cludes 28 families, 118 genera, and 1,060 native ing to account for the species richness. or adventive species. The account includes all fam- The Andes are the major feature of Peru that ilies of the Pteridophyta except for four that occur has molded the vegetation and distribution pat- only in the Old World and the Hymenophyllopsi- terns ofthe biota. The principle types of vegetation daceae ofnorthern South America. It includes most and species diversity are in large part based upon of the genera in the Americas and about one-third altitudinal zones affecting temperature and pre- ofthe Neotropical species. This indicates that Peru cipitation. The species of Peruvian pteridophytes ranks high in pteridophyte diversity. It is not sur- are treated here in four main zones, based upon prising that among the 24 departments of Peru, their ecology. They include Montana Ferns, Ceja those with the greatest number of species are also Ferns, Sierra Ferns and Loma Ferns. The ecolog- those that are ecologically diverse. The six most ical zones are shown on Map 2, along with a di-

1 table 1 are in speciose departments (map ; ) ecolog- agrammatic section showing the zones profile. ically strongly diverse, and they include relatively Montana Ferns occur in the forest region that large areas of montane rain forest and/or cloud covers the largest part of Peru. On the east flank forest. All of these have been relatively accessible of the Andes, this extends up to 1 800 m. This to collectors since the late 1 8th century, and with region receives 1500-3500 mm of rain annually, the exception of Pasco they are all large depart- usually between October and April. It is difficult ments. About 90% of the species and 86% of the to recognize floristic elements among several hun- endemics are in one or more of the six most spe- dred species in the Montana, but they may be ciose departments. characterized by their large size or by their epi- It is doubtful whether any additional families phytic habit. The large ones have creeping stems, will be added to the pteridophyte flora except by often forming large colonies with continued growth segregation of those already present. A few genera throughout the year. The leaves may be more than may be added, such as Microlepia in Ecuador and a meter long and are often modified for vegetative Bolivia, Pilularia, known to occur in Bolivia, and reproduction. There may be a relatively large Maxonia, known in Ecuador. Additional species number in a given locality, but many may be rare most likely will be found in the northern depart- species. The large-leaved species include Nephro- ments ofAmazonas and Cajamarca and the south- lepis biserrata (leaves to 3.6 m), Lygodium volubile ern departments of Madre de Dios and Puno. The (to 12 m), Hemidictyum marginatum (to 3 m), number also may be increased by addition to large Adiantum pectinatum (to 2.5 m), and the tree ferns, genera such as Selaginella and Thelypteris. mostly ca. 3 m. Some of the large-leaved species The 1 30 species endemic to Peru are included that form conspicuous colonies are Dennstaedtia in 1 Table and some of them are also on Map 1 . cicutaria, Gleichenia bifida, Dicranopteris pectin- The map shows that there appears to be no clear ata, and Pteridium aquilinum var. arachnoideum. geographic center of endemism. Rather, the de- 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 species in Peru.

Department 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 barren, or with very sparse vegetation. In the north support snow fields and glaciers. The temperature of Peru, these desert conditions are less pro- is relatively cool and the air is dry. The vegetation nounced. At the northernmost tip of Peru there is largely consists of grasses, sedges, semi-desert a small forested area. The otherwise barren coastal shrubs and cacti. Locally there are small woods, zone is relieved, at intervals, by green, irrigated especially those dominaed by Polylepis. The puna, valleys, and by the naturally verdant Lomas. The the land above the limits of agriculture, extends ferns of this region are part of a unique vegetation from 4000 m up to the limit of vascular plants, at that occurs along the coast ofPeru and Chile, north about 5200 m. Here cold prevails and low cushion to about 8 degrees south latitude. This vegetation plants are the principal vegetational feature. develops in response to local physiographic con- The Sierra Ferns are seasonally dormant but the ditions. In winter there is more or less constant dormancy may be intermittent due to infrequent fog and "guara" (a dense mist) on certain hills and rains during the dry season. The available mois- upper parts of valleys. The summer months are ture is largely in the soil or rock crevices. Many continuously dry. A rather lush vegetation may species occur where there is local seepage or where develop where unusual moist conditions occur. conditions are suitable for condensation of at- The long, continuously dry periods evidently re- mospheric moisture. The annual rainfall, between strict the flora. Loma Ferns include a selection of 500 and 1 1 00 mm, is by no means deficient but the Sierra species including Polypodium pycno- the dry air, winds, and strong insolation at the carpum, Adiantum subvolubile, and A. digitatum high altitudes combine to diminish the moisture. and, less commonly, Ophioglossum reticulatum, One of two main kinds of Sierra Ferns is the Anogramma leptophylla, Woodsia montevidensis, mesic type, typical of the locally moist habitats. and Cheilanthes peruviana. It is of interest to note Leaves of these plants die during the dry season that while about 80% of the Loma seed plants are unless they are in an unusually favorable place. endemic, there are no endemic ferns. This may be Species characteristic of this habitat include due to slower evolution ofthe ferns but more likely Adiantum poiretii, A. digitatum, Cystopteris fra- reflects their superior means of dispersal. The gilis, Asplenium peruvianum, and Woodsia mon- spores undoubtedly are blown from the Sierra to tevidensis. Asplenium peruvianum and the closely the Lomas with sufficient frequency to prevent related A. gilliesii are exceptional in this group in endemism. 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 References plantlet develops at what appears to be the tip of the rachis. LANG, G. 1992. Some aspects of European late- Xeric species are the second characteristic type and post-glacial flora history. Acta Bot. Fennica of Sierra Ferns. These plants retain their leaves 144: 1-17. but they become curled during the dry season, YOUNG, K. R., AND B. LEON. 1 989. Pteridophyte reviving during the brief moist periods. Species diversity in the central Peruvian Amazon: Im- characteristic of this group include Cheilanthes in- portance of edaphic specialization. Brittonia 41: carum, C. pruinata, C. bonariensis, C. scariosa, 388-395. Polypodium pycnocarpum, Pellaea ternifolia, and YOUNG, K. R., AND B. LEON. 1991. Diversity, Notholaena nivea. ecology and distribution of high-elevation pte- Loma Ferns occur on the Pacific side of the ridophytes within Rio Abiseo National Park, Andes, usually near the coast. This region becomes north-central Peru. Fern Gaz. 14: 25-29. progressively drier toward the ocean and finally

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 Bolivia 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 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 hackelianum 4-6 1 maxonii 5-112 hartwegii 4-135 reptans 5-153 Acrostichum 2-81 hayesii 4-136 rufum 2-47 acuminatum 4-154 haynaldii 4-136 scariosum 2-32 adenolepis 4- 1 66 heterophyllum 5-63 schomburgkii 4- 1 44 albescens 4-121 hickenii 4-136 serratifolium 4-101 alcicorne 5-181 horridulum 4-137 serratum 4-101

alienum 4- 1 00 huacsaro 4-137 serrulatum 5-83 apodum 4-157 /zysm';t4-138 setigerum 4- 1 60 aureonitens 2-48 ilvense 4-94 sinuatum 2-34 aureum 2-83 insigne 4-6 1 sorbifolium 4-105 auricomum 4-125 japurense 4-107 sphenophyllum 4-169 bakeri 4-125 juglandifolium 4-6 1 squamipes 4-161 barbatum 4-125 laminarioides 4-139 stenopyllum 4-161 bonariense 2-28 lanuginosum 2-32 suberectum 4-61 calaguala 4-137 latifolium 4-140 tambillense 4- 1 62 calomelanos 2-18 lechlerianum 4-141 tartareum 2-19 calophyllum 4-166 leprosum 4- 1 42 tectum 4-162 castaneum 4- 1 27 leptophyllum 4- 1 46 tenuiculum 4-163 caudatum 4-64 lindenii 4-142 tereticaulon 2-40 cervinum 4-85 /inrf/gii'4-103 thalictroides 2-50 chrysoconium 2-20 lingua 4-142 thelypteris 3-6 chrysolepis 4-6 1 litanum 4-143 trifoliatum 2-21 chrysophyllum 2-16 luridum 4- 1 44 unitum 4-146 ciliatum 4-128 mathewsii 4-144 villosum 4-156 citrifolium 2-92 minutum 4-146 Actinostachys 1-33 cladotrichum 4-125 moorei 4-167 digitata 1-33 curvans 4- 1 66 muscosum 4-147 pennula 1-36 cuspidatum 4- 1 29 nicotianifolium 4-102 Adiantopsis 2-34 danaeifolium 2-83 nigrescens 4-148 chlorophylla 2-36 decoration 4-129 nivosum 4-148 paupercula 2-34 denticulatum 4-131 nodosum 1-15 radiata 2-36 dichotomum 1-33 oligarchicum 4-102 ternata 2-36 digitat um 1-33 orbignyanum 4-151 Adiantum 2-52 discolor 4- 1 30 osmundaceum 4-62 alarconianum 2-67 dissimile 4- 1 66 pachyphyllum 4-151 amabile 2-56 diversifolium 4- 1 30 paleaceum 4-152 anceps 2-68 ebeneum 2- 1 8 pandurifolium 4-105 capillus-veneris 2-58 elegans 1-34 papillosum 4-152 cayennense 2-64 elongatum 4- 1 62 patinii 4-153 ceciliae 2-63 engelii 4-131 peltatum 4- 1 69 chilense 2-57 erinaceum 4-132 petiolosum 4-154 concinnum 2-58 erythrolepis 4- 1 32 phlebodes 4-107 crenatum 2-57 eximium 4- \ 33 piloselloides 4- 1 54 cuneatum 2-56 fendleri 3-68 plumbicaule 4-60 decorum 2-56 flabellatum 4-169 plumosum 4-155 deflectens 2-62 flaccidum 4-133 poeppigianum 4-156 delicatulum 2-62 fractiseriale 4-60 polybotryoides 4-6 1 digitatum 2-61 furfuraceum 4- 1 64 polypodioides 5-137 dolabriforme 2-62 glabellum 4- 1 34 preslianum 4-128 filiforme 2-62 graminioides 5-72 propinquum 4-156 flagellum 2-62 guamanianum 4-135 quitense 4-157 fructuosum 2-64

104 FIELDIANA: BOTANY fuliginosum 2-64 Allantodia Ananthacorus 2-89 guianense 2-120 asplenioides 4-75 angustifolius 2-89 henslovianum 2-59 Alsophila 1-116 Aneimia 1-24 hirtum 2-64 armigera 1-123 Anemia 1-24 humile 2-66 aterrima 1-1 14 adiantifolia 1-24 imbricatum 2-60 austral is 1-1 16 buniifolia 1-24 incisum 2-67 blechnoides 1-111 c limit a 1-25 kalbreyeri 2-63 capensis 1-118 ferruginea 1-27 kaulfussii 2-66 caracasana 1-133 flexuosa 1-27 killipii 2-66 conjugata 1-128 haenkei 1-29 laetum 2-59 contracta 1-109 hirsuta 1-28 lancea 2-121 cuspidata 1-120 hispida 1-30 latifolium 2-66 dombeyi 1-123 humilis 1-29 lobatum 2-60 elongata 1-115 myriophylla 1-28 lucidum 2-69 engelii 1-116 oblongifolia 1-29 lunulatum Burm. 2-62 erinacea 1-118 pastinacaria 1-28 lunulatum Houtt. 4-40 floribunda 1-123 phyllitidis 1-29 macrocladum 2-64 frigida 1-126 repens 1-30 macrophyllum 2-70 incana 1-120 tomentosa 1-27 mathewsianum 2-67 infesta 1-123 villosa 1-25 mexiae 2-64 kalbreyeri 1-124 Anemirhiza 1-24 microsorium 2-62 ft/ftp// 1-125 adiantifolia 1-24 moorei 2-56 lasiosora 1-125 ^ner/a 2-92 obliquum 2-57 latevagans 1-124 Anetium 2-92 orbignyanum 2-60 fcc/z/en 1-128 citrifolium 2-92 palmatum 2-6 1 macrosora 1-115 Anisogonium 4-67 patens 2-6 1 melanopus 1-124 fraxinifolium 4-67 pauperculum 2-34 microdonta 1-127 pinnatifidum 4-89 pectinatum 2-62 H/gra 1-124 Anogramma 2-22 pedatum 2-61 nigripes 1-124 chaerophylla 2-23 peruvianum 2-68 pallescens 1-131 leptophylla 2-23 petiolatum 2-66 paucifolia 1-118 Antrophyum 2-84 phillipense 2-62 peruviana 1-123 brasilianum 2-87 phyllitidis 2-70 phegopteroides 1-127 cajenense 2-87 pilosum 2-63 pilosissima 1-125 ensiforme 2-85 platyphyllum 2-68 podophylla 1-124 guayanense 2-87 poeppigianum 2-69 poeppigii 1-115 lanceolatum 2-87 poiretii 2-57 procera 1-123 lineatum 2-86 pulverulentum 2-65 pruinata 1-109 plantagineum 2-84 raddianum 2-56 pterorachis 1-123 Arachniodes 4-35 radiatum 2-36 pubescens 1-126 aspidioides 4-35 rhizophyllum 2-62 pycnocarpa 1-123 denticulata 4-37 ruizianum 2-62 quadripinnata 1-109 ochropteroides 4-38 scalare 2-69 rostrata 1-111 Argyrochosma 2-37 serratodentaum 2-65 sprucei 1-115 nivea 2-38 sessilifolium 2-59 swartziana 1-122 stuebeliana 2-38 speciosum 2-61 tarapotensis 1-125 Aspidium 4-2 1 steerei 2-62 tryonorum 1-128 abruptum 3-46 strictum 2-121 w/e/ 1-128 articulation 4-96 subaristatum 2-62 Amauropelta 3-9 atrorubens 3-24 subvolubile 2-59 breutelii 3-9 biolleyi 3-62 sulphureum 2-58 cheilanthoides 3-34 biserratum 5-5 1 terminatum 2-65 concinna 3-29 braunianum 4-24 tetraphyllum 2-64

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 105 gongylodes 3-40 claussenii 5-39 nidus 5-1 1 guianense 4-48 concinnum 5-4 nigricans 5-46 heracleifolium 4-27 congestum 5-41 ocanniense 4-85 hispidulum 3-41 conquisitum 5-18 oligophyllum 5-30 incanum 3-52 crassifolium 4-76 otites 5-48 karstenii 4-9 cristatum 5-24 partitum 5-21 kunzei 4-29 cuneatum 5-44 parvulum 5-15 macrophyllum 4-29 cuspidatum 5-26 pearcei 5-13 macrourum 3-44 davisii 5-30 perkinsii 5-18 martinicensis 4-24 delicatulum 5-25 peruvianum 5-34 meniscioides 4-48 delitescens 5-71 plantagineum 4-86 microchlaena 4-9 dentatum 5-35 plantaginifolium 4-86 navarrense 3-15 denticulosum 4-80 poloense 5-35 nemophilum 4-10 denudatum 5-48 praemorsum 5-46 rtoMe 4-38 desvauxii 4-77 procerum 4-77 nodosum 4-96 dimidiatum 5-46 projectum 5-33 opulentum 3-43 discolor 5-48 pseudoangustum 5-12 orbiculatum 4-53 discrepans 5-36 pteropus 5-36 paleaceum 4-36 divaricatum 5-23 pulchellum 5-35 parallelogramma 4-36 drepanophyllum 5-42 pumilum 5-14 patulum 4-36 eggersii 4-87 purdieanum 5-47 pectinatum 5-53 erosum 5-46 purpurascens 5-32 pendulum 4-98 escaleroense 5-14 quitense 5-35 pilosulum 3-19 escragnollei 5-30 raddianum 5-39 platyphyllum 4-52 expansum 4-75 radicans 5-19 poeppigii 4-29 extensum 5-15 repandulum 5-28 protensum 4-19 falcinellum 5-28 repens 5-21 pusillum 3-17 ferulaceum 4-90 resiliens 5-15 pycnolepis 4-54 flabellulatum 5-21 rhizophyllum 5-20 rostratum 1-111 flavescens 4-85 rhomboideum 5-22 rotundatum 4-42 flavidum 5-30 roemerianum 4-85 rutaceum 5-17 flexuosum 4-77 rosenstockianum 5-37 scalare 3-15 foeniculaceum 5-27 ruizianum 5-43 semihastatum 3-57 formosum 5-14 rusbyanum 5-30 sprengelii 3-31 /ragife 5-34 rutaceum 5-17 stipulare 3-44 fragrans 5-27 salicifolium 5-43 trianae 4-47 fuscopubescens 4-72 sandwichianum 4-72 truncatulum 4-40 gillesianum 5-34 scolopendrium 5-4 uliginosum 3-3 gilliesii 5-33 serra 5-45 Asplenium 5-2 haenkeanum 5-24 serratum 5-11 abrotanoides 5-27 hallii 5-18 sessilifolium 5-37 abscissum 5-41 haplophyllum 5-13 shepherdii 4-80 achilleifolium 5-32 harpeodes 5-39 spruceanum 5-18 aethiopicum 5-46 hastatum 5-44 squamosum 5-26 alatum 5-40 integerrimum 5-28 striatum 4-79 alienum 4-72 jamesonii 5-26 stuebelianum 5-12 amazonicum 5-13 juglandifolium 5-28 tabinense 5-38 angustum 5-12 kapplerianum 5-29 te*> 5-22 arboreum 4-80 kunzei 4-87 ternatum 5-22 auriculatum 5-44 laetum 5-31 theciferum 5-48 auritum 5-42 fcc/z/m 4-85 tomentosum 2-46 balliviani 5-21 lindbergii 4-78 trapezoides 5-48 bangii 5-38 lividum 5-47 tricholepis 5-30 barbaense 5-37 longicaudatum 5-19 trichomanes-dentatum 5-34 blechnoides 5-60 lorentzii 5-33 trilobatum 5-22 brasiliense 5-40 macrophyllum 4-77 trilobum 5-48 callipteris 4-83 macrurum 5-19 triphyllum 5-22 canelense 5-30 marginatum 4-90 tucumanense 5-26 caracasanum 4-79 marinum 5-2 tuerckheimii 5-29 castaneum 5-17 mathewsii 5-28 tungurahuae 4-76 caucense 4-85 maxonii 5-18 uniseriale 5-2 1 celtidifolium 4-83 melanopus 5-3 1 vargasii 5-17 centripetale 4-87 monanthemum 5-16 vastum 4-73 cicutarium 5-24 monanthes 5-16 venulosum 4-76 cirrhatum 5-20 myriophyllum 5-25 virens Desv. 5-3 1 cladolepton 5-24 neogranatense 5-29 virens Presl 5-30

106 FIELDIANA: BOTANY vomeriforme 5-28 maxonii 5-61 Campyloneurum 5-158 wagneri 5-17 meridense 5-64 abruptum 5-172 weberbaueri 5-49 nigrosquamatum 5-61 aglaolepis 5-167 Athyrium 4-88 nudum 5-56 amphostenon 5-165 achilleifolium 5-32 obtusifolium 5-67 angustifolium 5-168 ambiguum 4-7 1 occidentale 5-58 angustipaleatum 5-169 "bradearum" 4-88 orientale 5-58 aphanophlebium 5-161 celtidifolium 4-83 ornifolium 5-62 asplundii 5-167 dombei 4-90 pectinatum 5-58 brevifolium 5-170 dombeyi 4-90 penna-marina 5-62 caespitosum 5-162 expansum 4-75 peruvianum 5-62 chlorolepis 5-168 ferulaceum 4-90 polypodioides Raddi 5-60 coarctatum 5-164 filix-femina 4-88 polypodioides (Sw.) Kuhn 5-64 decurrens 5-132 flexuosum 4-77 pteropus 5-63 densifolium 5-166 fumaroides 4-92 rubicundum 5-65 fasciale 5-163 haenkeanum 5-24 scandens 5-70 praestans 4-87 schomburgkii 5-66 fuscosquamatum 5-163 Austrolycopodium 6-52 serrulatum 5-60 heterolepis 5-168 magellanicum 6-56 sprucei 5-66 inflatum 5-164 Azolla 6-8 squamulosum 5-65 irregulare 5-166 caroliniana 6-10 stenophyllum Presl 5-65 jamesonii 5-172 filiculoides 6-11 stenophyllum (Klotzsch) Mett. lapathifolium 5-162

var. rw&ra 6- 1 1 5-65 /atara 5-170 mexicana 6-10 stipitellatum 5-65 lorentzii 5-167 microphylla 6- 1 1 magniftcum 5-132 rubra 6- 1 1 tr(angulare 5-59 nitidissimum 5-171 trilobum 5-68 nodosum 5-172 unilateral 5-60 occultum 5-161 Balantium volubile 5-70 ophiocaulon 5-162 karstenianum 1-105 Blotiella2-lll pascoense 5-171 Blechnum 5-56 glabra2-lll phyllitidis 5-169 acutum 5-64 lindeniana 2-113 repens 5-162 andinum 5-61 Bolbitis 4-98 serpentinum 5-163 angustifolium 5-64 aliena4-100 solutum 5-172 arborescens 5-62 bradeorum 4-103 sphenodes 5-164 asplenioides 5-60 crenata 4-101 taeniosum 5-169 auratum 5-67 guianensis 4-109 trichiatum 5-161 auriculatum 5-68 fci7/(p4-102 vulpinum 5-167 binervatum 5-63 Iindigii4-103 Ceradenia 5-72 blechnoides 5-60 macrophylla 3-68 capillaris 5-90 brasiliense 5-6 1 nicotianifolia 4-102 curvata 5-72 buchtienii 5-67 oligarchica 4-102 dendroxa 5-88 caudatum 5-58 pandurifolia 4-105 discolor 5-86 chilense 5-62 portoricensis 4-101 farinosa 5-89 ciliatum 5-68 serrata 4-101 herrerae 5-88 cognatum 5-68 serratifolia 4-101 longipinnata 5-85 columbiense 5-67 stuebelii 4-103 meridensis 5-85 confluens 5-59 Botrychium 1-6 mirabilis 5-91 cordatum 5-62 cicutarium 1-8 pearcei 5-87 delicatum 5-68 lun an a 1-6 pilipes 5-90 divergens 5-63 mirifica 1-58 praeclara 5-89 ensiforme 5-64 schaffner 1-6 terrestris 5-87 fragile 5-64 underwoodianum 1-6 Ceratopteris 2-50 fraxineum 5-59 virginianum 1-8 pteridoides 2-50 glandulosum 5-58 virginicum 1-8 richardii 2-50 gracile 5-59 Bryodisma 6-66 thalictroides 2-50 heterophyllum 5-63 Byrsopteris 4-35 Ceropteris kunthianum 5-64 aristata 4-35 adiantoides 2- 1 9 1'herminieri 5-61 Ceterach lanceola 5-60 aspidioides 3-13 lechleri 5-68 Caenopteris Cheilanthes 2-23 lehmannii 5-61 achilleifolia 5-32 andina 2-27 longifolium 5-59 myriophylla 5-25 arequipensis 2-33 loxense 5-65 Calamaria bonariensis 2-28 magellanicum 5-68 triquetra 6-97 borsigniana 2-37 malacothrix 5-68 Camptosorus 5-2 buchtienii 2-34

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 107 cantangenste 2-32 blumei 5-72 Culcita 1-103 chlorophylla 2-36 Ctenitis 4-5 coniifolia 1-103 concolor 2-3 1 ampla 4-10 macrocarpa 1-103 crenata 4-94 andicola 4- 1 9 Cuspidaria 5-145 elegans 2-31 biserialis 4-15 furcata 5-145 farinosa 2-30 catocarpa 4-10 Cyathea 1-129 fasciculata 2-27 distans 4-5 andina 1-130 fractifera 2-25 hirsuto-setosa 4-16 arborea 1-129 fraseri 2-28 honesta 4-14 aterrima 1-1 14 glandulosa 2-106 karsteniana 4-17 bipinnatifida 1-126 glauca 2-30 microchlaena 4-9 caracasana 1-133 hostilis 2-\\Q mollicoma 4-17 castanea 1-135 hypoleuca 2-34 nemophila 4-10 cuspidata 1-120 incarum 2-33 nigrovenia 4-9 delgadii 1-132 intramarginalis 2-23 protensa 4-2 1 divergens 1-131 lonchophylla 2-33 pulverulenta 4- 1 7 dudleyi 1-135 macleanii 2-27 refulgens 4-7 ebernina 1-135 marginata 2-30 sloanei 4-10 elongata 1-117 mathewsii 2-27 subincisa 4-10 equestris 1-131 micropteris 2-23 submarginalis 4-8 erinacea 1-118 mollis 2-29 Ctenopteris 5-72 frigida 1-126 moritziana 2-25 amylacea 5-87 fulva 1-135 myriophylla 2-3 1 anfractuosa 5-99 incana 1-120 notholaenoides 2-25 apiculata 5-94 kalbreyeh 1-124 obducta 2-29 asplenifolia 5-104 lasiosora 1-125 obtusata 2-\\\ athyrioides 5-113 latevagans 1-124 orbignyana 2-29 capillaris 5-90 lechleri 1-135 ornatissima 2-32 congesta 5-89 macrosora 1-115 parallelogramma 2-110 contacta 5-107 meridensis 1-135 peruviana 2-32 crispata 5-72 mexicana 1-133 pilosa 2-27 cultrata 5-108 microdonta 1-127 pilosa x pruinata 2-27 discolor 5-86 microphylla 1-132 poeppigiana 2-30 dolorensis 5-104 multiflora 1-130 pruinata 2-27 ecuadorensis 5-110 multisegmenta 1-132 radiata 2-36 farinosa 5-89 n/#ra 1-125 rigida 2-34 ./zrma 5-100 nigripes 1-124 rufopunctata 2-29 gracilis 5-96 oligocarpa 1-132 saundersii 2-25 herrerae 5-88 oreites 1-115 scariosa 2-32 heteromorpha 5-105 pallescens 1-131 sinuata 2-34 lanigera 5-106 panamensis 1-131 squamosa 2-34 leucosticta 5-111 petiolata 1-131 tripinnata 2-32 longipinnata 5-85 phegopteroides 1-127 Cheiroglossa 1-8 longiuscula 5-111 pilosa 1-132 palmata 1-9 major 5-96 pilosissima 1-125 Christella 3-39 melanosticta 5-98 poeppigii 1-115 dentata 3-42 meridensis 5-85 polystichoides 1-118 hispidula 3-41 moniliformis 5-98 primaeva 1-125 parasitica 3-39 obovata 5-95 procera 1-123 Christensenia 1-13 peruviana 5-102 pubens 1-127 Chrysodium phlegmaria 5-97 pubescens 1-127 lindigii 4-103 pilosissima 5-101 pungens 1-123 serratum 4-101 pseudocapillaris 5-94 quindiuensis 1-116 Cnemidaria 1-136 pseudonutans 5-103 rufescens 1-1 14 alatissima 1-138 pteropus 5-95 ruiziana 1-132 cocleana 1-139 r/ge5 5-101 schanschin 1-132 horrida 1-138 rigescens 5-102 sprucei 1-115 nervosa 1-139 semihirsuta 5-112 subtropica 1-128 speciosa 1-139 sericeo-lanata 5-106 wte/ 1-128 uleana 1-138 ste//a 5-106 vilhelmii 1-130 Cochlidium 5-72 subflabelliformis 5-107 willdenowiana 1-123 graminioides 5-72 subimpressa 5-94 Cyclodium 4-47 pumilum 5-82 subsessilis 5-95 guianense 4-48 serrulatum 5-82 taxifolia 5-113 meniscioides 4-48 Coptophyllum 1-24 tunguraguae 5-95 trianae 4-47 buniifolium 1-24 venulosa 5-72 Cyclopeltis 4-29 Cryptosorus 5-72 yungensis 5-113 semicordata 4-29

108 FIELDIANA: BOTANY Cyclosorus 3-39 coniifolia 1-103 bogotense 4-70 dentatus 3-42 dissecta 2-100 bombonasae 4-8 1 gongylodes 3-40 erosa 2-100 bonapartii 4-75 Cyrtomium 4-38 gigantea 1-105 bradeorum 4-88 dubium 4-38 karsteniana 1-105 brasiliense 4-79 falcatum 4-40 montevidensis 4-94 buchtienii 4-72 nobile 4-38 obtusifolia 2-100 callipteris 4-83 Cyrtophlebium 5-158 pearcei 1-99 caracasanum 4-79 phyllitidis 5-169 polypodioides 2-95 celtidifolium 4-83 repens 5-162 rubiginosa 2-98 centripetale 4-88 Cystodium 1-103 sellowiana 1-105 chimborazense 4-87 Cystopteris 4-92 spruceana 1-105 costale 4-77 fragilis 4-92 stuebelii 1-105 crassifolium 4-76 translucens 4-92 Dicranoglossum 5-49 cristatum 4-80 desvauxii 5-147 cuneifolium 4-82 furcatum 5-145 delitescens 5-32 Danaea 1-15 panamense 5-145 denticulosum 4-80 cuspidata 1-19 polypodioides 5-148 diplazioides 4-71 elliptica 1-18 subnudum 5-147 divergens 4-70 grandifolia 1-17 Dicranopteris 1-46 drepanolobium 4-8 1 humilis 1-20 o^nw 1-42 eggersii 4-88 jamaicensis 1-19 bancroftii 1-39 expansum 4-75 longifolia 1-17 ftCfrfa 1-41 ferulaceum 4-90 moritziana 1-19 brittonii 1-44 flavescens 4-85 nodosa 1-17 dichotoma 1-46 flexuosum 4-76 oblanceolata 1-18 flexuosa 1-47 fraxinifolium 4-67 stenophylla 1-19 linearis 1-47 fuscopubescens 4-72 trichomanoides 1-20 longipinnata 1-45 fuscum 4-87 wendlandii 1-20 nervosa 1-47 gracilescens 4-72 Davallia pectinata 1-49 grande 4-78 arborescens 2-100 pennigera 1-39 grandifolium 4-82 concinna Presl 2-100 peruviana 1-45 hians 4-70 concinna Schrader 5-4 pruinosa 1-42 induratum 4-78 glauca 2-99 remota 1-44 lechleri 4-85 inaequalis 2-103 rubiginosa 1-46 legalloi 4-84 multiflora 5-52 schomburgkiana 1-47 lehmannii 4-72 thecifera 5-48 seminuda 1-47 lindbergii 4-78 Davalliopsis simplex 1-39 lonchophyllum 4-80 elegans 1-84 ve/ata 1-42 macrodictyon 4-87 Dennstaedtia Bernh. 2-95 yungensis 1-44 macrophyllum 4-77 Dennstaedtia Moore 2-95 Didymochlaena 4-40 melanopus 5-3 1 Dennstaedtia lunulata 4-40 melanosorum 4-75

arborescens 2- 1 00 sinuosa 4-40 moritzianum 4-73 bipinnata 2-99 truncatula 4-40 obscurum 4-85 cicutaria 2-98 Didymoglossum 1-76 obtusum 4-79 concinna 2-100 angustifrons 1-86 oxylobum 4-77 dissecta2-100 hymenoides 1-86 pactile 4-87 distenta 2-95 krausii 1-87 paucijugum 4-84 erosa 2-100 membranaceum 1-88 pedatum 4-74 flaccida 2-95 muscoides 1-86 pinnatifidum 4-86 glauca 2-99 reptans 1-87 plantagineum 4-86 globulifera 2-99 sphenoides 1-86 plantaginifolium 4-86 mathewsii 2-100 Diphasiastrum 6-52 praestans 4-87 obtusifolia 2-100 thyoides 6-57 preslianum 4-76 pearcei 1-99 Diphasium 6-52 remotum 4-74 punctilobula 2-95 jussiaei 6-56 riedelianum 4-85 rubiginosa 2-98 Diplazium 4-65 roemerianum 4-85 sprucei 2-101 aberrans 4-87 rostratum 4-75 wercklei 2-101 alienum 4-72 sandwichianum 4-72 Deparia altissimum 4-70 shepherdii 4-80 mathewsii 2-100 ambiguum 4-7 1 striatum 4-79 Dicksonia 1-105 angelipolitanum 4-79 stuebelianum 4-80 arborescens 1-105 appolinaris 4-77 subnudum 4-79 berteriana 1-105 arboreum 4-80 subobtusum 4-78 bipinnata 2-99 asplenioides 4-75 tabalosense 4-79 cicutaria 2-98 bicolor 4-74 tarapotense 4-73

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 109 tungurahuae 4-76 euchlora 3-25 poiteana 3-65 unilobum 4-81 extensa 3-43 prasina 4-45 vastum 4-73 falcata 3-75 protensa 4-2 1 venulosum 4-75 fibrillosa 4-13 ptarmiciformis 3-17 verapax 4-85 filix-mas 4-35 pteroidea 3-25 werckleanum 4-80 /wrva 3-19 pubescens 4-35 Diplopterygium 1-37 gigantea 3-68 pulverulenta 4- 1 7 bancroftii 1-39 glandulosa 3-49 ps/7/a 3-17 Doryopteris 2-43 glandulosolanosa 3-20 pyramidata 3-58 concolor 2-3 1 heterophlebia 4-45 quadrangularis 3-41 crenulans 2-44 hirsuto-setosa 4- 1 6 refulgens 4-7 lomariacea 2-44 honesta 4-14 resinosofoetida 3-34 lorentzii 2-44 incana 3-52 retrorsa 3-27 palmata 2-44 jamesonii 3-56 rimbachii 3-19 pedata 2-44 jurgensenii 3-75 rivulorum 3-32 Drynaria juruensis 3-61 n&//.s 3-26 acuminata 5-156 karsteniana 4- 1 7 rw/a 3-20 Dryopteris 4-35 karstenii 4-9 ruiziana 3-35 ampla 4-10 killipii 4-33 saffordii 4-37 anceps 3-68 laevigata 3-24 salzmannii 3-73 ancyriothrix 3-63 /ec/z/m' 3-69 scalaris 3-16 andicola 4-19 leprieurii 3-5 1 sellensis 3-3 1 andreana 3-76 leptosora 4-14 semihastata 3-57 angustifolia 3-74 leucothrix 3-32 serrata 3-71 arcana 3-76 limaensis 3-20 simplicifrons 3-68 aristata 4-35 lindigii 3-30 sprengelii 3-31 aspidioides 3-13 lingulata 3-75 stuebelii 3-22 assurgens 3-30 linkiana 3-14 subandina 3-20 asterothrix 3-62 lomatosora 3-15 subincisa 4- 1 6 atrorubens 3-24 longicaudata 4-44 submarginalis 4-8 balbisii 3-3 1 lugubriformis 3-58 supina 3-26 bangii 3-42 macbridei 3-19 supralineata 3-52 biformata 3-60 macrophylla 3-68 tarapotensis 4- 1 5 biserialis 4- 1 5 macrostegia 4-38 tetragona 3-64 boqueronensis 3-26 macrotis 3-57 thelypteris 3-6 brachyodus 3-49 mapiriensis 3-49 thomsonii 3-22 brachypus 3-18 megalodus 3-64 tremula 3-9 brausei 3-28 mercurii 3-31 m'sto 3-6 1 canadasi 3-2 1 microchlaena 4-9 uliginosa 3-3 catocarpa 4-10 microsora 4-14 valdepilosa 3-50

caucaensis 3-28 mi// 3-28 vasta 4- 1 7 cheilanthoides 3-34 mollicoma 4-11 wallichiana 4-36 christii 4-45 mollis 3-42 warmingii 3-56 chrysodioides 3-69 multiformis 3-35 yungensis 4-14 clypeata 3-65 nemophila 4-10 coarctata 3-32 nigrovenia 4-9 columbiana 3- 1 5 w'teHs 3-35 Elaphoglossum 4-111 comosa 3-49 ochropteroides 4-38 concinna 3-29 oligocarpa 3-15 albescens 4-121 consobrina 3-73 oligophylla 3-45 alipes 4- 1 22 contermina 3-32 opaca 4-45 amazonicum 4-122 corazonensis 3-27 pachyrhachis 3-3 1 amphioxys 4-123 ctenitis 4-5 paleacea 4-36 amplum 4-123 decussata 3-49 paludosa 4-48 angustius 4- 1 24 deflexa 3-30 parallelogramma 4-36 apodum 4-123 deltoidea 3-46 patens 3-44 atropunctatum 4- 1 24 dmsa 3-34 patula 4-36 atrosquamatum 4- 1 24 densiloba 3-52 paucinervata 4-45 auricomum 4-125 dentata 3-42 pavoniana 3-23 bakeri 4-125 denticulata 4-37 pellucido-punctata 4-44 bangii 4-167 desvauxii 3-71 permollis 3-70 barbatum 4-125 diplazioides 3-14 peruviana 3-23 blepharoglottis 4- 1 26 dumetorum 3-27 phacelothrix 3-22 calaguala 4-137 eggersii 3-59 pilosohispida 3-27 cam pt oil-pis 4- 1 26 engelii 3-26 pilosula 3-19 cardenasii 4-126 ensiformis 3-75 platyloba 4-15 castaneum 4-127

110 FIELDIANA: BOTANY caudatum 4-154 minutum 4-146 williamsiorum 4-165 chloodes4-127 molle4-152 williamsii 4-123

ciliatum 4- 1 28 moorei 4-167 zebrinum 4-165 concinnum 4-128 moyeri 4-147 Enterosora 5-72 conforme 4-111 muscosum 4-147 campbellii 5-72 craspedotum 4-128 nastukiae 4- 1 48 parietina 5-84 crinipes 4-160 nidiformis4-148 trichosora 5-84 cuspidatum 4-129 nigriscens 4-148 trifurcata 5-83 decoratum 4- 1 29 nigrocostatum 4- 1 40 Equisetaceae 6- 1 2 denticulatum 4-131 nivosum 4-148 Equisetum 6-12 dichroum4-129 obovatum 4-149 bogotense 6-16

discolor 4- 1 30 obtusum 4- 1 49 fluviatile 6- 1 2

diversifrons 4- 1 30 oculatum 4-150 giganteum 6- 1 5 eatonianum 4-127 odontolepis 4- 1 50 myriochaetum 6- 1 5 elegantipes 4-131 orbignyanum 4- 1 50 ramosissimum elongatum 4-162 ornatum 4-149 ssp. debile 6- 1 5 engelii 4-131 oxyglossum 4-151 schaffneri 6-15 x ensiforme 4-131 odphyllum 4- 1 50 schaffneri 6- 1 5 erinaceum 4-132 pachyphyllum 4-151 variegatum 6- 1 5 erythrolepis 4- 1 32 pachyrrhizum 4-152 Eriosorus 2-3 eximium 4-133 paleaceum 4-152 accrescens 2-6 flaccidum 4-133 palorense 4-153 aureonitens 2-6 fortipes 4-133 papillosum 4-153 cheilanthoides 2-6 glabellum 4-134 pascoense 4-153 elongatus 2-8 glossophyllum 4-134 patinii 4-153 flabellatus 2-8 glutinosum 4-154 pattersoniae 4-166 flexuosus 2-7 gracillimum 4- 1 34 peltatum 4-169 lechleri 2-8 guamanianum 4-135 petiolosum 4-154 orbignyanus 2-7 hartwegii 4-135 pichinchae 4-158 rufescens 2-4 hayesii 4- 1 36 piloselloides 4-154 ruizianus 2-20 haynaldii 4-136 pilosius 4-155 scandens 2-7 hickenii 4-136 plicatum 4-152 stuebelii 2-6 hieracioides 4-137 plumosum 4-155 wurdackii 2-7 "hikenii" 4-137 poeppigianum 4-15 Eschatogramme 5-145 horridulum 4-137 potomogeton 4-166 desvauxii 5-147 huacsaro 4-137 preslianum 4-128 furcata 5-147 hystrix 4-138 propinquum 4-156 polypodioides 5-148 jucundum 4-138 pseudohirtum 4-158 subnuda 5-147 killipii 4-13S pumilio 4-156 Eupodium 1-15 laminarioides 4-139 punae 4-157 kaulfussii 1-15 Ianatum4-139 quitense 4-157 Iasioglottis4-139 raywaense 4-157 latevagans 4- 1 40 rimbachii 4-158 Feea latifolium 4-140 rosenstockii 4-158 diversifrons 1-91 latum 4-123 rubellum 4-159 heterophylla 1-94 lawyerae 4-141 ruficomus 4-159 humboldtii 1-94 laxisquama 4-141 rufum4-159 trollii 1-91

lechlerianum 4-141 russelliae 4- 1 60 leprosum 4-142 schomburgkii 4-144 leptophyllum 4-146 setigerum 4-160 Glaphyropteris 3-48 lindbergii 4- 1 26 siliquoides 4-136 decussata 3-49

lindenii 4-142 simulans 4- 1 60 Gleichenia 1-37 lingua 4-142 spatulatum 4-155 affinis 1-42 linguaeforme 4- 1 34 squamipes 4-161 bancroftii 1-39 litanum 4-143 stenophyllum 4-161 bifida 1-41 lloense 4- 1 32 sty riacum 4-161 boliviensis 1-43 longipes 4- 1 44 subciliatum 4-143 buchtienii 1-42 longius 4-143 tambillense 4-162 costaricensis 1-43 luridum 4-144 tectum 4-162 flexuosa 1-47 macilentum 4- 1 44 tenue 4-163 glauca 1-37 mathewsii 4- 1 44 tenuiculum 4-163 hypoleuca 1-43 megalurum 4-145 to men tell um 4-163 lechleri 1-44 megarhizon 4-146 velongum 4- 1 64 leucocarpa 1-44 meladenium 4-145 vittarioides 4- 1 64 longipinnata 1-45 melancholicum 4-145 vulcanicum 4-164 mathewsii 1-41 metallicum 4-146 wardiae 4-165 mellifera 1-4

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. Ill nervosa 1-47 discolor 5-86 variabilis 5-115 nitidula 1-43 dudleyi 5-92 venulosa 5-72 pectinata 1-49 elongata 5-143 werfii 5-92 pennigera 1-39 erecta 5-98 xiphopteroides 5-101 peruviana 1-45 farinosa 5-88 youngii 5-97 polypodioides 1-37 firma 5-100 Gymnogramma pruinosa 1-42 flabelliformis 5-102 aureonitens 2-7 remota 1-44 flexuosa 2-7 diplazioides 3-14 revoluta 1-42 gracilis 5-96 elongata 2-8 rigida 1-47 graminioides 5-72 ferruginea 2-20 rubiginosa 1-46 herrerae 5-88 flabellata 2-8 simplex 1-39 heteromorpha 5-105 flexuosa 2-7 subandina 1-43 immixta 5-115 goudotii2-\2 tomentosa 1-41 jamesonii 5-82 jamesonii 2-20 truncata 1-37 jamesonioides 5-93 lechleri 2-8 tuberculata 1-43 lanceolata 5-143 mathewsii 2-4 velata 1-42 lanigera 5-105 mohriaeformis 2-4 yungensis 1-44 laxa 5-107 ochracea 2- 1 9 Glyphotaenium 5-72 lehmanniana 5-104 orbignyana 2-1 trifurcatum 5-83 leucosticta 5-111 pearcei 2-2 1 Goniophlebium 5-71 limbata 5-8 1 peruviana 2- 1 9 incanum 5-125 linkiana 3-14 polypodioides 3-14 pectinatum 5-121 longipinnata 5-85 pumila 2-84 semipinnatum 5-140 major 5-96 reniformis 2- 1 5 subauriculatwn 5-72 marginella 5-72 rufescens 2-4 Goniopteris 3-52 mathewsii 5-91 stuebelii 2-6 biolleyi 3-62 melanosticta 5-97 Gymnopteris 2-46 crenata 3-52 meridensis 5-85 aliena 4-100 eggersii 3-59 mirabilis 5-91 nicotianifolia 4-102 juruensis 3-6 1 moniliformis 5-98 pandurifolia 4-105 pennata 3-64 myosuroides 5-82 rufa 2-47 poiteana 3-65 myriophylla 5-114 tomentosa 2-46 pyramidata 3-58 nigrolimbata 5-81 tetragona 3-64 obliquata 5-72 m'sto 3-6 1 paramicola 5-8 1 Hecistopteris 2-84 Grammitis 5-72 parietina 5-84 pumila 2-84 albidula 5-87 peruviana 5-102 Hemidictyum 4-90 alsopteris 5-110 phlegmaria 5-97 marginatum 4-90 amylacea 5-87 pichinchae 5-110 Hemionitis 2-46 andicola 5-103 pichinchensis 5-110 brasiliana 2-87 and/mz 5-109 pilipes 5-90 cajenensis 2-87 anfractuosa 5-99 pilosissima 5-100 lanceolata 2-87 apiculata 5-94 praeclara 5-89 lineata 2-86 aromatica 5-100 pseudocapillaris 5-94 palmata 2-48 asplenifolia 5-104 pseudonutans 5-103 pinnata 2-48 assurgens 5-99 pumila 5-82 plantaginea 2-84 athyrioides 5-113 recondita 5-95 rufa 2-47 basalis 5-109 revoluta 5-143 tomentosa 2-46 bipinnata 5-92 rigens 5-101 Hemiphlebium bishopii 5-86 rigescens 5-102 kapplerianum 1-88 blepharidea 5-112 ruiziana 2-7 Hemitelia 1-129 blepharolepis 5-108 sectifrons 5-72 andina 1-130 bryophila 5-81 semihirsuta 5-111 horrida 1-138 Z>wes/7 5-112 senilis5-107 lechleri 1-131 campbellii 5-72 sericeo-lanata 5-106 multiflora 1-130 capillaris 5-90 serrulata 5-82 nervosa 1-139 cheilanthoides 2-6 sprucei 5-84 petiolata 1-131 chrysleri 5-104 squamulosa 5-143 rufescens 1-1 14 congesta 5-89 subflabelliformis 5-106 speciosa 1-139 crispata 5-72 subsessilis 5-95 subincisa 1-139 cultrata 5-108 taxifolia 5-113 uleana 1-138 curvata 5-72 terrestris 5-87 Hicriopteris daguensis 5-109 trichosora 5-83 bancrofti 1-39 david-smithii 5-109 trifurcata 5-83 Histiopteris 2-115 dendroxa 5-88 truncicola 5-109 incisa 2-115 dependens 5-104 tunguraguae 5-94 vespertilionis 2-115

112 FIELDIANA: BOTANY Hoffmannia 6- 1 1 pilgeriana 6-34 mexiae 1-59 aphylla 6- 1 1 polycarpos 6-37 microcarpum 1-68 Holodictyum 5-2 polyclada 6-5 1 mirincum 1-58

Homoeotes 1-76 polylepidetorum 6-3 1 molle 1-65 heterophylla 1-94 pruinosa 6-5 1 nuil tialat urn 1-73 Huperzia 6-19 quadrifariata 6-49 multiflorum 1-63 acerosa 6-46 reflexa 6-27 myriocarpum 1-62 acifolia 6-28 var. bifida 6-28 nigrescens 1-63 affinis 6-30 var. minor 6-28 nigricans 1-63 andina 6-35 var. reflexa 6-28 pedicellatum 1-57 aqualupiana 6-48 rosenstockiana 6-42 peltatum 1-57 arcuata 6-25 sagasteguiana 6-37 peruvianum 1-75 aristei 6-43 sanctae-barbarae 6-35 platylobum 1-68 attenuata 6-37 sarmentosa 6-45 plumieri 1-72 bifida 6-28 saururus 6-35 plumosum 1-72 blepharodes 6-30 selago 6-19 poeppigianum 1-76 binervia 6-26 sellifolia 6-38 polyanthos 1-59 brevifolia 6-39 socratis 6-42 polycarpum 1-64 brongniartii 6-27 sotae 6-43 procerum 1-65 buesii 6-42 subulata 6-50 pulchellum Hooker 1-70 campiana 6-47 taxifolia 6-41 pulchellum Mett. 1-65 capellae 6-32 tenuis 6-46 pyramidatum 1-74 caracasica 6-40 tetragona 6-38 reniforme 1-64 colanensis 6-3 1 unguiculata 6-29 rimbachii 1-64 crassa 6-33 weberbaueri 6-30 ruizianum 1-69 cuneifolia 6-50 weddellii 6-26 rupestre 1-82 curvifolia 6-46 wilsonii 6-44 simplex 1-69 darwiniana 6-33 Hydroglossum 1-33 speciosum 1-70 dichaeoides 6-48 oligostachyum 1-33 spectabile 1-70 dichotoma 6-26 Hymenodium sprucei 1-76 durissima 6-51 kunzeanum 4-151 superbum 1-72 ecuadorica 6-29 Hymenophyllum 1-50 tarapotense 1-75 engleri 6-39 adiantoides 1-66 tenerrimum 1-66 ericifolia 6-48 amabile 1-70 tomentosum 1-73 eversa 6-29 andinum 1-62 tortuosum 1-57 filiformis 6-46 apiculatum 1-59 trapezoidale 1-69 funiformis 6-42 axillare 1-62 trianae 1-61 hartwegiana 6-40 beyrichianum 1-68 trichomanoides 1-61 heteroclita 6-48 calodictyon 1-56 trichophyllum 1-65 hippuridea 6-24 ciliatum 1-67 trifidum 1-76 hohenackeri 6-40 contortum 1-64 tunbridgense 1-50 hypogaea 6-36 crispatulum 1-68 undulatum 1-63 kuesteri 6-30 crispum 1-67 valvatum 1-68 jenmanii 6-44 cristatum 1-55 verecundum 1-75 lechleri 6-25 dendritis 1-59 Hymenostachys lindaviana 6-44 dependens 1-72 diversifrons 1-91 linifolia 6-43 dicranotrichum 1-50 Hypoderris var. jenmanii 6-44 ectocarpon 1-57 stuebelii 4-103 var. tenuifolia 6-44 elegans 1-66 Hypolepis 2-106 loxensis 6-26 elegantulum 1-70 bogotensis 2-1 10 lucidula 6-19 endiviifolium 1-63 flexuosa 2-110 macbridei 6-32 fendlerianum 1-64 hostilis 2-110 mandiocana 6-45 ferax 1-61 nigrescens 2-109 mexiae 6-28 fragile 1-69 obtusata 2-110 mollicoma 6-45 fucoides 1-55 parallelogramma 2-110 molongensis 6-47 fusagasugense 1-73 pteroides 2-111 montana 6-25 fusugasugense 1-73 stuebelii 2-109 myrsinites 6-49 hirsutum 1-66 tenuifolia 2- 106 nesselii 6-34 interruptwn 1-72 nuda 6-25 karstenianum 1-71 papillata 6-3 1 lamella turn 1-58 Isoetaceae 6-88 parvifolium 6-28 latipes 1-59 Isoetes 6-89 passerinoides 6-4 1 lindenii 1-71 andicola 6-92 var. nitens 6-4 1 lineare 1-66 var. gemmifera 6-92 pearcei 6-30 lobatoalatum 1-74 andina 6-97 phylicifolia 6-49 mathewsii 1-60 boliviensis 6-96

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 113 dichotoma 6-9 1 Lepicystis 5-125 loxensis 5-65 dispora 6-93 incana 5-125 meridensis 5-64 glacialis 6-95 Lepidotus auct. 6-58 M<&Z 5-56 herzogii 6-97 Lepidotus Mirbel 6-52 obtusifolia 5-67 hewitsonii 6-94 alopecuroides 6-6 1 ornifolia 5-62 karstenii 6-96 cernua 6-63 pteropus 5-63 lacustris 6-89 clavata 6-54 schomburgkii 5-66 laevis 6-95 contexta 6-62 serrulosa 5-62 lechleri 6-95 magellanica 6-56 squamulosa 5-65 var. triquetra 6-97 Lepisorus 5-140 stenophylla 5-65 novo-granadensis 6-91 Leptochilus stipitellata 5-65 pacifica 6-9 1 alienus 4-100 volubilis 5-70 panamensis 6-9 1 bradeorum 4-103 Lomaridium parvula 6-92 crenatus 4-101 semicordatum 1-101 peruviana 6-95 guianensis 4-109 Lomariopsis 4-105 saracochensis 6-94 killipii 4-102 erythrodes 4-107 savanarum 6-9 1 lindigi 4-103 fendleri4-106 soda 6-95 nicotianifolius 4-102 japurensis 4-106 "ticlioensis" 6-95 oligarchicus 4- 1 03 latipinna 4-107 triquetra 6-97 pandurifolium 4-105 nigropaleata 4-108 serratifolius 4-101 sorbifolia 4-105 serratus 4-101 Jamesonia 2-8 stuebelii 4-103 Lonchitis 2-113 alstonii 2-12 Leptocionium 1-50 blepharum 2-14 dicranotrichum 1-50 hirsuta 2-113 boliviensis 2-12 fucoides 1-56 lindeniana 2-113 cinnamomea 2-14 pedicellatum 1-57 ;?tea 2-80 glutinosa 2-13 Leptopteris 1-20 tenuifolia 2-106 goudotii 2-12 Lindsaea 2-115 Lophidium 1-33 imbricata 2- 1 3 arcuata 2-117 elegans 1-34 paleacea 4-54 divaricata 2-118 flabellum 1-34 peruviana 2- 1 1 guianensis 2-120 latifolium 1-33 pulchra 2-11 hemiglossa 2-122 poeppigianum 1-36 rotundifolia 2-10 lancea 2-121 Lophosoria 1-107 scalaris 2- 1 2 latifrons 2-122 pruinata 1-109 scammanae 2-1 1 portoricensis 2-120 quadripinnata 1-107 phassa 6-98 Loxoma 1-98 schomburgkii 2-122 Loxoscaphe 5-4 Lacostea spruceana 2-118 concinna 5-4 tanaica 1-89 sprucei 2-118 thecifera 5-48 Lastrea stricta2-121 Loxsoma 1-98 cheilanthoides 3-34 taeniata 2-118 Loxsomopsis 1-99 nitens 3-35 tarapotensis 2-118 costaricensis 1-99 poeppigiana 3-57 trapeziformis 2-115 lehmannii 1-99 poiteana 3-65 ulei2-122 notabilis 1-99 pubescens 4-3 1 Litobrochia pearcei 1-99 recedens 4-3 1 horizontalis 2-77 Lycopodiaceae 6-16 ntt/w 3-26 Lomagramma 4-109 Lycopodiella 6-58 scabriuscula 3-44 guianensis 4-109 sect. Campylostachys 6-60 Lastreopsis 4-3 1 Lomaria 5-56 sect. Caroliniana 6-58 amplissima 4-33 acwta 5-64 sect. Lycopodiella 6-60 effusa 4-33 andina 5-61 alopecuroides 6-6 1 exculta 4-35 angustifolia 5-64 camporum 6-63 killipii 4-33 arborescens 5-62 caroliniana

recedens 4-3 1 aurata 5-67 var. meridionalis 6-6 1

tenera 4-3 1 caudata 5-66 cernua 6-63 Lecanium chilensis 5-62 contexta 6-62 membranaceum 1-88 cordata 5-62 descendens 6-64 Lellingeria 5-72 cuspidata 5-64 glaucescens 6-64 apiculata 5-94 divergens 5-63 inundata 6-58 major 5-96 ensiformis 5-64 matthewsii 6-6 1 myosuroides 5-82 euphlebia 1-101 pendulina 6-65 phlegmaria 5-97 fragile 5-64 riofrioi 6-65 pseudocapillaris 5-94 fraxinea 5-59 Lycopodium 6-52 subsessilis 5-95 heterophylla 5-63 sect. Caroliniana 6-60 tunguraguae 5-95 linariaefolia 5-68 sect. Complanata 6-52

114 FIELDIANA: BOTANY sect. Diphasium 6-52 gayanum 6-57 spurium 6-56 sect. Lycopodium 6-52 geniculatum 6-85 stamineum 6-44 sect. Magellanica 6-52 glaucescens 6-64 stellae-polaris 6-28 subg. Cernuistachys 6-58 gracile 6-80 subulatum 6-50 subg. Lepidotis auct. 6-58 haematodes 6-80 taxifolium 6-41 subg. Lepidotis Baker 6-52 haenkei 6-56 var. brongniartii 6-27 subg. Rhopalostachya 6-52 hartwegianum 6-40 tenu? 6-46 subg. Selago 6-19 herbaceum 6-55 var. tenuissimum 6-46 subg. Urostachya 6- 1 9 heteroclitum 6-48 tetragonum 6-38 acifolium 6-28 heterophyllum 6-57 var. patulum 6-38 affine 6-30 hohenackeri 6-40 thujoides 6-57 albidum 6-55 hypogaea 6-19 thyoides 6-57 alopecuroides 6-6 1 hippurideum 6-24 trichodendron 6-44 var. contextum 6-62 jenmanii 6-44 trichophyllum 6-54 ssp. contextum 6-62 jussiaei 6-56 trychopyllum 6-54 anceps 6-80 var. microphyllum 6-56 unguiculata 6-29 andinum 6-35 lechleri 6-25 vestitum 6-55 arcanum 6-44 linifolium 6-43 weddellii 6-26 aristatum 6-54 lucidula 6-19 wilsonii 6-44 articulatum 6-84 macbridei 6-32 Lygodium 1-30 attenuatum 6-37 magellanicum 6-56 digitatum 1-32 bifidum 6-28 mathewsii 6-6 1 mexicanum 1-33 binervium 6-26 meridionale 6-6 1 micans 1-32 blepharodes 6-30 mexiae 6-28 oligostachyum 1-33 brevifolium 6-39 microphyllum 6-75 polymorphyum 1-33 brongniartii 6-27 molongense 6-47 radiatum 1-32 brutum 6-28 myrsinites 6-49 scandens 1-30 Z>Mes 6-42 var. minus 6-38 venustum 1-30 capellae 6-32 -4 1 volubile 1-32 capillaceum 6-63 nubigenum 6-49 caracasicum 6-40 nova-hollandiae 6-77 carolinianum 6-58 nudum 6-12 Macrothelypteris 3-3 var. meridionalis 6-62 papillatum 6-31 torresiana 3-3 catharticum 6-38 parkeri 6-84 Marattia 1-13 cernuum 6-64 parvifolium 6-28 a/ara 1-15 var. capillaceum 6-64 passerinoides 6-4 1 kaulfussii 1-15 var. pendulinum 6-65 pendulinum 6-65 laevis 1-15 clavatum 6-54 pensum 6-68 Marginaria 5-125 ssp. clavatum 6-54 phlegmaria 6-19 angustifolia 5-169 ssp. contiguum 6-55 phylicifolium 6-49 polypodioides 5-125 var. aristatum 6-54 pichinchense 6-56 Marginariopsis 5-7 1 var. pseudocontiguum 6-55 piliferum 6-54 Marsilea 6-2 var. thyoides 6-57 polycarpos 6-45 ancylopoda 6-4 var. validum 6-57 polycarpum 6-29 crotophora 6-5 congestifolium 6-49 polycladum 6-5 1 deflexa 6-4 contextum 6-62 poseidonis 6-24 mollis 6-5 contiguum 6-55 pruinosum 6-5 1 natans 6-6 convolutum 6-87 reflexum Lam. 6-27 mucronata 6-5 crassum 6-33 var. densifolium 6-28 uncinata 6-5 cuatrecasasii 6-45 var. majus 6-28 vestita 6-4 cuneifolium 6-50 var. minus 6-28 ssp. tenuifolia 6-5 curvifolium 6-46 var. polycarpum 6-29 ssp. vestita 6-5 densifolium 6-28 reflexum Willd. 6-29 Marsileaceae 6-2 diffusum 6-76 reversum 6-28 Mecodium 1-50 durissimum 6-5 1 riofrioi 6-65 apiculatum 1-59 ecuadoricum 6-29 rosenstockianum 6-42 contortum 1-64 elongatum 6-35 sanctae-barbarae 6-2 1 dendritis 1-59 engleri 6-39 sarmentosum 6-45 endiviifolium 1-63 ericaefolium 6-48 saururus 6-35 fendlerianum 1-64 eriostachys 6-54 scariosum 6-55 /mzx 1-61 erythropus 6-80 vsa.jussieui 6-57 mexiae 1-59 eversum 6-29 schwendeneri 6-41 microcarpum 1-68 ewanii 6-50 selaginoides 6-66 multiflorum 1-63 exaltatum 6-86 selago 6- 1 9 myriocarpum 1-62 fastigiatum 6-56 skutchii 6-49 polyanthos 1-59 funiforme 6-42 sprucei 6-62 trichomanoides 1-61

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 115 undulatum 1-64 percussa 5-144 tarapotense 4-8 5-157 trapezoides 4-53 Megalastrum 4- 1 1 persicariifolia 3-50 andicola 4- 1 8 piloselloides 5-152 valdepilosum biseriale 4-15 recreense 5-157 villosum 4- 1 8 hirsutosetosum 4-16 reptans 5-163 Nephrolepis 5-49 honestum 4-13 rosmarinifolia 5-154 biserrata 5-51 cordifolia 5-52 leptosorum 4-14 squamulosa 5-156 microsorum 4-14 tecta5-154 hirsutula 5-54 multiflora 5-52 mollicomum 4- 1 7 thurnii 5-156 occidentalis 5-53 pansamalense 4- 1 8 ulei 5-156 5-53 platylobum 4-15 vacciniifolia 5-152 pectinata 5-52 pulverulentum 4-17 Microlepia 2-95 pendula rivularis 5-52 spectabile 4- 1 6 flaccida 2-95 subincisum 4-16 inaequalis 2-103 Neuromanes vastum 4-17 polypodioides 2-95 pinnatum 1-92 villosum 4-1 1 speluncae 2-95 Neurophyllum hostmannianum 1-93 yungense 4-14 Micropolypodium 5-72 Melpomene 5-96 pseudotrichomanoides 5-72 pinnatum 1-92 Meniscium 3-66 Microstaphyla Niphidium 5-173 andreanum 3-76 bangii 4-167 albopunctatissimum 5-174 americanum 5-173 angustifolium 3-74 moorei 4-167 arborescens 3-70 Mildella 2-23 anocarpos 5-177 cristatum 4-80 intramarginalis 2-23 carinatum 5-176 crassifolium 5-174 falcatum 3-75 Mohria 1-23 5-173 giganteum 3-68 longifolium macbridei 5-176 guyanense 3-68 vittaria 5-177 jurgensenii 3-75 Nephelea 1-118 macrophyllum 3-68 cuspidata 1-120 Notholaena 2-37 opacum 4-45 erinacea 1-118 arequipensis 2-33 salzmannii 3-73 incana 1-120 awm* 2-28 serratum 3-71 polystichoides 1-118 bonariensis 2-28 Meringium Nephrodium brackenridgei 2-32 buchtienii 2-34 fucoides 1-56 antioquoianum 4-26 Mertensia 1-46 brachypus 3-18 Candida 2-37 2-32 bancroftii 1-39 canadasii 3-21 cantangensis 2-40 Z>(/zrfa 1-41 "carazanense" 3-27 chrysophylla dichotoma 1-46 caucaense 3-28 /ra?ri 2-28 2-33 flexuosa 1-47 conspersum 3-42 lonchophylla 2-37 laevigata 1-37 corazonense 3-27 marantae mo//w 2-29 longipinnata 1-45 crassipes 3-3 1 mathewsii 1-41 deflexum 3-30 nivea 2-38 nervosa 1-47 eggersii 3-59 obducta 2-29 2-32 pectinata 1-49 firmifolium 4-47 peruviana sinuata 2-34 pennigera 1-39 funestum 4-21 pruinosa 1-42 gardnerianum 3-52 stuebeliana 2-33 remota 1-44 jamesonii 3-56 sulphurea 2-37 revoluta 1-42 kunzeanum 3-46 tectaria 2-34 tenera 2-40 rigufa 1-47 lagerheimii 4-8 simplex 1-37 fecA/m 3-69 tomentosa 2-34 tomentosa 1-41 leprieurii 3-51 trichomanoides 2-37 ve/afa 1-42 lizarzaburui 4-25 Metaxya 1-109 longipilosum 3-14 rostrata 1-111 macradenium 3-21 Odontomanes Microgramma 5-148 macrotis 3-57 hostmannianum 1-93 acatallela 5-152 microsorum 4-14 Oleandra 4-96 acuminata 5-157 nemorale 3-62 articulata 4-96 baldwinii 5-155 nigrovenium 4-9 distenta 4-96 ochropteroides 4-38 hirta 4-98 chrysolepis 5-151 pilosohispidum 3-27 lehmannii 4-97 ciVuzfcz 5-153 polyphyllum 4-53 micans 4-98 neriiformis 4-96 fuscopunctata 5-144 quadrangulare 3-4 1 geminata 5-157 resinosofoetidum 3-34 nodosa 4-96 latevagans 5-151 retrorsum 3-27 pilosa 4-98 Iindbergii5-157 schizotis 3-44 Olfersia 4-55 4-64 lycopodioides 5- 1 54 sodiroi 4-26 caudata megalophylla 5-157 supinum 3-26 cervina 4-57

116 FIELDIANA: BOTANY ciliata 4-128 eurybasis 5-121 euphlebia 1-101 corcovadensis 4-57 filicula 5-119 latifolia 1-101 Onoclea funicula 5-116 semicordata 1-101 polypodioides 1-37 hygrometrica 5-124 Plananthus 6-19 Ophioglossum 1-8 pectinata 5-121 reflexus 6-27 coriaceum 1-12 plumula 5-118 selago 6- 1 9 crotalophoroides 1-12 ptilodon 5-123 Platycerium 5-181 ellipticum 1-12 venturii 5-122 alcorne 5-181 lusitanicum 1-12 Pellaea 2-40 andinum 5-181 nudicaule 1-11 atropurpurea 2-40 Plecosorus 4-49 opacum 1-12 cordifolia 2-4 1 mexicanus 4-49 palmatum 1-9 crenulans 2-44 peruvianus 2-111 pendulum 1-8 dealbata 2-38 speciosissimus 4-49 peruvianum 1-9 lorentzii 2-44 Pleopeltis5-140 petiolatum 1-9 /vea 2-38 angusta 5-140 reticulatum 1-9 ovata 2-4 astrolepis 5-143 scandens 1-30 peruviana 2-41 fuscopunctata 5-144 scariosum 1-11 sagittata 2-41 lanceolata 5-143 tenerum 1-11 tenera 2-40 macrocarpa 5-142 vulgatum 1-8 ternifolia 2-41 percussa 5-144 ypanemense 1-11 weddeliana 2-4 1 pinnatifida 5-135 Osmunda 1-21 wrightiana 2-41 revoluta 5-143 adiantifolia 1-24 Peltapteris4-167 squamulosa 5-143 cervina 4-57 moorei 4-167 Pleuridium cicutaria 1-8 peltata4-169 albopunctatissimum 5-174

cinnamomea 1-21 peruviana 4- 1 70 Pleurosorus 5-2 flexuosa 1-27 Peltochlaena 4-47 Poecilopteris hirsuta 1-28 Phanerophlebia 4-38 crenata 4-101 humilis 1-29 nobilis 4-38 Polybotrya 4-57 lunaria 1-6 Phegopteris aequatoriana 4-64 oblongifolia 1-29 cochleata 4-55 alfredii 4-65 palustris 1-23 dictyophylla 4-38 altescandens 4-6 1 dwZ>/a 4-38 andina 4-6 1 phyllitidis 1-29 laevigata 3-24 appressa 4-64 polypodioides 5-64 lechleri 4-45 caudata 4-64 regalis 1-23 membranacea 3-69 cervina 4-57 spectabilis 1-23 mo/fo 3-70 crassirhizoma 4-60 virginiana 1-8 pycnolepis 4-55 decorata 4-62 refulgens 4-7 fractiserialis 4-60 Phlebodium 5-125 fulvastrigosa 4-65 Paesia2-106 aureum 5-134 glandulosa 4-63 amazonica 2-106 decumanum 5-135 hickeyi 4-65

anfractuosa 2-106 Phlegmariurus 6- 1 9 juglandifolia 4-6 1 glandulosa 2-106 taxifolius 6-4 1 kalbreyeri 4-6 1 viscosa 2-106 Phyllitis 5-4 lechleriana 4-63 Palhinhaea 6-58 scolopendrium 5-4 lomarioides 4-65 camporum 6-63 Phylloglossum 6-16 macbridei 4-60 cernua 6-63 Pilularia 6-2 nutans 4-65 descendens 6-64 americana 6-2 osmundacea 4-62 glaucescens 6-64 mandonii 6-2 plumbicaulis 4-60 pendulina 6-65 Pityrogramma 2-16 polybotryoides 4-6 1

riofrioi 6-66 austroamericana 2- 1 8 pubens 4-62 Parablechnum chrysoconia 2-20 puberulenta 4-63 ciliatum 5-68 chrysophylla 2-18 serratifolia 4-57 Paraceterach ebenea 2-18 sorbifolia 4-57 marantae 2-37 ferruginea 2-20 subelliptica 4-63 Parkeria ochracea 2- 1 9 suberecta 4-6 1 pteridoides 2-50 pearcei 2-21 Polypodium 5-125 Pecluma 5-116 perelegans 2- 1 9 abitaguae 5-86 absidata 5-119 peruviana 2-19 abruptum 3-58 boliviano 5-120 presliana 2-20 absidatum 5-119

camptophyllaria 5-122 tartarea 2- 1 9 acrodontium 5-101 choquetangensis 5-116 trifoliata 2-21 acrosorum 5-177 curvans 5-119 Plagiogyria 1-101 adiantiforme 4-3 1 dispersa 5-125 costaricensis 1-101 adnatum 5-133 divaricata 5-120 denticulata 1-101 aglaolepis 5-168

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 117 albopunctatissimum 5-174 dentatum 3-41 5-129 alternifolium 5-108 denticulatum 4-37 5-170 americanum 5-173 dependens 5-104 5-107 amphostenon 5-166 dichotomum 1-46 leptophyllum 2-23 amplum 4-10 discolor 5-86 leucatomos 5-134 andinum 5-109 dispersum 5-125 x leucosporum 5-139 anfractuosum 5-99 disectum 2-100 leucosticton Fee 5-111 angustifolium 5-168 dissimile 5-129 leucosticton Klotzsch 5-136 angustipaleatum 5-169 divaricatum 5-120 levigatum 5-134 anocarpos 5-177 dolorense 5-104 limbatum 5-81 apiculatum 5-94 5-83 lomariiforme 5-125 oppression 5-132 lonchitis 4-49 arboreum 1-129 ecostatum 5-85 longicaudatum 4-44 aristatum 4-35 ecuadorense 5-1 10 longifolium 5-132 aromaticum 5-100 effusum 4-33 longisetosum 5-114 articulatum 5-133 euchlorum 3-25 longiusculum 5-111 asplenifolium 5-104 eurybasis 5-121 longum 5-108 asplundii 5-167 exaltatum 5-49 loretense 5-156 astrolepis 5-143 falcatum 4-40 loriceum 5-129 athyrioides 5-113 farinosum 5-89 lycopodioides 5-154 aureum 5-134 fasciale 5-163 macrocarpum Presl 5-135 azuyense 5-94 fendleri 5-132 macrocarpum Willd. 5-143 balaoense 5-138 fibrillosum 4-13 macrophyllum 4-44 fez/Mrii 3-31 fUicula5-H9 marginellum 5-72 biauriculatum 5-131 filix-femina 4-88 mathewsii 5-91 fiiix-mas 4-35 medullare 1-112 binervatum 5-63 firmum 5-100 megalodus 3-64 biseriale 4- 1 5 flabelliforme 5-102 megalolepis 5-137 blepharideum 5-112 flavopunctatum 4-42 megalophyllum 5-157 blepharolepis 5-108 fragile 4-92 melanostictum 5-97 bolivianum 5-120 fraseri 5-136 meridense 5-85 bombycinum 5-138 fraxinifolium 5-132 microdontum 1-127 brachyodus 3-49 fulvescens 2-111 mo//e 3-41 brevifolium 5-170 furfuraceum 5-136 mollendense 5-135 bryophilum 5-81 fuscopunctatum 5-144 moniliforme 5-98 bryopodum 5-135 giganteum 5-133 monosorum 5-138 buchtienii 5-136 gilliesii 5-130 montevidense 4-5 1 l>ues/7 5-112 glaucophyllum 5-134 monticola 5-99 caceresii 5-133 glaucum 1-109 muricatum 4-55 caespitosum 5-162 globuliferum 2-99 murorum 5-139 camptophyllarium 5-123 gracile 5-96 capillare 5-90 gracillimum 5-108 myosuroides 5-82 chacapoyense 5-131 guianense 4-109 myriophyllum 5-114 chartaceum 5-129 haynaldii 4-26 nigrolimbatum 5-81 chnoodes 5-129 herzogii 5-100 /j/tefu 3-35 chrysolepis 5-151 heteromorphum 5-105 nitidissimum 5-172 ciliatum 5-153 heterophlebium 4-45 nodosum 5-172 circinatum 5-119 hirsutulum 5-54 obliquatum 5-72 coardatum 5-164 honestum 4-13 occultum 5-161 cochleatum 4-54 horridum 1-138 oligocarpum 3-15 concinnum 3-29 x huancayanum 5-139 oligophlebium 4-45 cordatum 5-131 hygrometricum 5-124 ophiocaulon 5-162 cordifolium 5-52 incanum 5-137 opposition 3-32 crassifolium 5-174 jamesonii 5-82 parietinum 5-84 crenatodentatum 4-54 jamesonioides 5-93 patens 3-44 crenatum 3-65 karstenianum 4- 1 7 pavonianum 3-23 crispatum 5-167 pearcei 5-87 crossii 3-23 kunzeanum 5-131 pectinatum 5-121 crystalloneuron 5-139 lachniferum 5-123 pedicellata 1-15 cultratum 5-108 /aftfwm 5-130 pendulum 5-95 curvans 5-1 19 laevigatum 5-134 penna-marina 5-62 curvatum 5-72 lanceolatum 5-142 pennatum 3-64 dasypleuron 5-131 lanigerum 5-106 percussum 5-144 decumanum 5-135 lapathifolium 5-162 persicariifolium 5-157 decurrens 5-132 lasiopus 5-130 peruvianum 5-102 decussatum 3-49 latevagans 5-151 phlegmaria 5-97

118 FIELDIANA: BOTANY phyllitidis 5-169 subauriculatwn 5-71 trapezoides 4-54 pichinchae 5-1 10 subflabelliforme 5-106 VW//H 4-52 pichinchense 5-110 subincisum 4- 1 6 yungense 4-52 pilipes 5-90 submarginale 4-8 Polytaeniwn 2-84 piloselloides 5-152 subscabrum 5-1 10 brasilianum 2-87 pilosissimum 5-100 subsessile 5-95 cajenense 2-87 plantagineum 4-27 subvestitwn 5-135 guayanense 2-87 platylobum 4- 1 5 surucuchense 5-131 lanceolatum 2-86 plumula 5-118 taeniosum 5-168 Pronephrium 3-66 polypodioides 5-137 tarapotense 4- 1 5 Pseudolycopodiella 6-58 pozuzoense 5-90 taxifolium 5-113 contexta 6-62 prasinum 4-45 tectum 5-154 meridionalis 6-62 preslianum 5-132 tenuiculum 5-96 Psilogramme 2-8 procerum 1-123 tetragonum 3-64 Psilotaceae 6- 1 1 pruinatum 1-109 thomsonii 3-22 Psilotum 6-11 pseudoaureum 5-134 thurnii 5-156 complanatum 6-12 pseudocapillare 5-94 thyssanolepis 5-137 nudum 6-12 pseudonutans 5-103 tottum 3-40 Pteridanetium 2-92 pteroideum 3-25 trichiatum 5-161 citrifolium 2-92 pteropus 5-95 trichosorum 5-84 Pteridium 2-105 ptilodon 5-124 trifoliatum 4-21 aquilinum 2-105

pubescens Hooker & Grev. 5- 1 30 trifurcatum 5-83 arachnoideum 2-105 pubescens L. 4-35 triseriale 5-132 caudatum 2-105 pulverulentum 4-17 m'ste 3-60 Pteris 2-70 punctatum 4-45 truncicola 5-109 altissima 2-76 pungens 1-123 tunguraguae 5-95 amazonica 2-106 pycnocarpum 5-135 tweedianum 5-135 amp/a 2-78 pycnolepis 4-55 M/e/5-156 angustifolia 2-90 quadripinnatum 1-109 vacciniifolium 5-152 aquilina 2-105 ratibori 5-135 variabile 5-115 arachnoidea 2-105

remotum 5-136 vastum 4- 1 7 atropurpurea 2-40 repens 5-162 venturii 5-122 awrea 2-28 reticulatum 3-66 venulosum 5-72 bakeri 2-80 richardii 5-133 villosum 4-11 biaurita 2-77 rigens 5-101 vittaria 5-177 concolor 2-3 1 rigescens 5-102 vulgare 5-125 consanguinea 2-77 rigidum 4-53 vulpinum 5-167 coriacea 2-74 rivulare 5-52 xantholepis 5-135 cretica 2-81 rivulorum 3-32 xiphopteroides 5-101 decomposita 2-80 rosmarinifolium 5-154 yungense 5-113 decurrens 2-77 rostratwn 1-111 Polystichopsis 4-35 deflexa 2-74 nfe 3-26 ochropteroides 4-38 edentula 2-74 rufum 3-20 Polystichum 4-49 farinosa 2-30 ruiz-lealii 5-135 amplissimum 4-33 fraseri 2-78 ruizianum 3-35 boboense 4-54 furcata 5-145 rusbyi 5-135 bonapartii 4-38 grandifolia 2-79 salicifolium 3-74 cochleatum 4-54 haenkeana 2-78 saxatile 4-54 dubium 4-38 horizontalis 2-77 sectifrons 5-72 gelidum 4-54 imbricata 2-13 semicordatum 4-29 haenkeanum 4-54 incisa 2-115 semihirsutum 5-111 lehmannii 4-51 interrupta 3-40 semipinnatifidum 5-140 lonchitis 4-49 intramarginalis 2-23 .wrote 5- 107 mexiae 4-5>l faV/i/Hi 2-78

sericeo-lanatum 5-106 montevidense 4-5 1 kunzeana 2-76 serpentinum 5-163 moritzianum 4-55 lechleri 2-78 serrulatum 5-83 muricatum 4-55 //'nea/a 2-90 sessilifolium 5-131 nudicaule 4-52 livida 2-80 sloanei 4-10 orbiculare 4-53 lonchitoides 2-113 solutum 5-172 orbiculatum 4-53 longifolia 2-70 sororium 5-129 paleaceum 4-54 /wa'fito 2-69 speluncae 2-95 platyphyllum 4-52 muricata 2-74 sphenodes 5-164 polyphyllum 4-54 /v2 2-38 spixianum 5-86 pycnolepis 4-54 notholaenoides 2-25 sprucei 5-84 sodiroi 4-54 orbiculata 2- 1 3 squamulosum 5-156 speciosissimum 4-49 ovate 2-43 subandinum 5-130 torresianum 3-3 palmata 2-44

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 119 6-74 pedata 2-80 Salviniaceae 6-5 producta 6-81 peruviana 2-41 Schaffheria 5-2 quadrifaria ramosissima 6-75 petiolulata 2-79 Schizaea 1-33 podophylla 2-80 dichotoma 1-33 regularis 6-77 revoluta 6-72 polita 2-74 digitata 1-33 propinqua 2-75 elegans 1-34 rupestris 6-66 pungens 2-75 nstulosa 1-34 f. amazonica 6-71 f. 6-7 1 quadriaurita 2-75 flabellum 1-34 peruviana reticulata 2-76 incurvata 1-36 sartorii 6-71 reticulatovenosa 2-76 pennula 1-36 seemannii 6-74 6-66 rigida 2-34 poeppigiana 1-36 selaginoides sellowii 6-71 ruffa 2-47 pusilla 1-34 sheldonii 6-7 1 sagittata 2-41 Selaginella 6-66 silvestris 6-82 speciosa 2-76 acanthostachys 6-87 6-79 splendens 2-79 amazonica (Milde) Hieron. 6-7 1 speciosa 6-66 sulphured 2-37 amazonica Spring 6-7 1 spinosa A. Br. 6-8 1 ternifolia 2-41 anceps 6-79 sprucei Hooker 6-78 transparens 2-76 applanata 6-73 sprucei trichomanoides 2-37 arizonica 6-72 stellata 6-86 6-86 tripartita 2-8 1 articulata 6-84 strobolifera sulcata vespertilionis 2-115 asperula 6-85 verttftz 2-78 atirrensis 6-76 ssp. poeppigiana 6-83 suavis 6-87 vittata 2-8 1 bombycina 6-78 ssp. tomentosa 6-85 Pteropsis brachylepis 6-84 trisulcata vittarioides 2-9 1 brevifolia 6-72 6-83 Pterozonium 2-14 calcarata 6-86 truncata 6-73 brevifrons 2-15 calosticha 6-74 wolfii 6-87 paraphysatum 6-96 chionoloma 6-77 weberbaueri 6-73 6-77 reniforme 2-15 chrysoleuca 6-78 xiphophylla Ptilophyllum conduplicata 6-86 Selaginellaceae 6-66 bicorne 1-91 convolute 6-87 Selaginelleae 6-66 hostmannianum 1-93 cordifolia 6-73 Selenodesmium lambertianum 1-95 demissa 6-72 rigidum 1-85 martiusii 1-95 diffusa 6-76 Sitobolium 2-95 pellucens 1-96 dimorpha 6-82 punctilobulum 2-95 eggersii 6-76 Solanopteris 5-179 elongata 6-85 bifrons5-181 Ragatelus erythropus 6-80 bismarckii 5-180 crinitus 1-94 exaltata 6-86 brunei 5-180 Regnellidium 6-2 ferruminata 6-85 tuberosa 5-180 Soromanes 4-57 Rhipidopteris 4-167 filicina 6-80 4-57 flabellata 4-\69 flagellata 6-77 serratifolia 1-50 peltata 4-169 fragilis 6-84 Sphaerocionium adiantoides 1-66 rusbyi 4-167 geniculata 6-85 ciliatum 1-67 sphenophylla 4-169 gracilis Moore 6-80 1-67 Rumohra 4-3 1 gracilis (Poiret) Hieron. 6-80 crispum adiantiformis 4-3 1 haematodes 6-80 elegans 1-66 1-70 aspidioides 4-31 haenkeana 6-82 elegantulum berteriana 4-3 1 horizontalis 6-87 fragile 1-69 huberi 6-79 hirsutum 1-67 intacta 6-76 interruptum 1-72 Saccoloma 2-101 kunzeana 6-82 karstenianum 1-71 1-71 elegans 2-103 lechleri 6-79 lindenii lobatoalatum 1-74 inaequale 2-103 lingulata 7-76 wercklei 2-101 microphylla 6-75 microcarpum 1-68 /no//e 1-65 Saffordia 2-48 mildei 6-7 1 /rtrfwta 2-50 nodosa 6-85 multialatum 1-73 Salpichlaena 5-68 nova-hollandiae 6-77 nigricans 1-63 hookeriana 5-70 parkeri 6-84 plumieri 1-72 1-72 lomarioidea 5-70 pearcei 6-77 plumosum volubilis 5-70 pedata 6-84 pyramidatum 1-74 ruizianum 1-69 Salvinia 6-6 peruviana 6-7 1 auriculata 6-8 var. dombeyana 6-71 simplex 1-69 minima 6-6 poeppigiana 6-83 spectabile 1-71 natans 6-6 var. peruviana 6-83 tenerrimum 1-66 rotundifolia 6-8 praestans 6-8 1 tomentosum 1-73

120 FIELDIANA: BOTANY trichophyllum 1-65 Tectaria 4-2 1 dudleyi 3-33 valvatum 1-68 andina 4-25 dumetorum 3-27 Sphaeropteris 1-112 antioquiana 4-26 eggersii 3-59 u talma 11 pa 1-115 brauniana 4-24 enigmatica 3-16 aterrima 1-1 14 decurrens 4-29 ensiformis 3-75 bradei 1-115 draconoptera 4-26 erythrothrix 3-59 elongata 1-115 fraxinea 5-51 euchlora 3-25 hirsuta 1-114 haynaldii 4-26 extensa 3-43 horrida 1-1 12 heracleifolia 4-27 exuta 3-37 macrosora 1-1 14 incisa 4-24 falcata 3-75 medullaris 1-112 kunzei 4-29 frigida 3-18 quindiuensis 1-116 lizarzaburui 4-25 funckii 3-18 rufescens 1-114 martinicensis 4-24 furfuracea 3-34 Steiropteris 3-46 plantaginea 4-27 furva 3-19 gardneriana 3-52 poeppigii 4-29 gardneriana 3-52 incana 3-52 sodiroi 4-26 gigantca 3-68 valdepilosa 3-50 transiens 4-25 glandulosa 3-50 Stenochlaena trifoliata 4-27 glandulosolanosa 3-20 angusta 4-106 vivipara 4-25 gongylodes 3-40 fendleri 4-106 Thelypteris 3-5 grandis 3-46 japurensis 4-107 abrupta 3-58 guyanensis 3-68 vesfita 4-106 aequatorialis 3-45 hispidula 3-41 Sticherus 1-37 ancyriothrix 3-63 hutchisonii 3-29 o#z/ii5 1-42 andicola 3-16 interrupta 3-40 6//K/WS 1-41 andreana 3-76 invisa 3-46 buchtienii 1-42 angustifolia 3-74 jamesonii 3-56 laevigatus 1-37 arborea 3-24 juruensis 3-61 lechleri 1-44 arborescens 3-70 killipii 3-60 longipinnatus 1-45 arcana 3-76 laevigata 3-24 mathewsii 1-41 arenosa 3-33 lancea 3-74 nitidulus 1-43 argentina 3-21 leoniae 3-18 penniger 1-39 arrecta 3-36 leprieurii 3-5 1 pruinosus 1-42 aspidioides 3-13 leucothrix 3-32 revolutus 1-42 assurgens 3-30 Hmaensis 3-20 rubiginosus 1-46 atrorubens 3-24 limbata 3-9 simplex 1-39 balbisii 3-31 lindigii 3-30 tuberculatus 1-43 biformata 3-60 lingulata 3-75 velatus 1-42 biolleyi 3-62 linkiana 3-14 yungensis 1-44 brachyodus 3-50 lomatosora 3- 1 5 Stigmatopteris 4-42 brachypus 3-18 longifolia 3-70 alloeoptera 4-45 brausei 3-28 loretensis 3-30 ecuadorensis 4-45 canadasii 3-21 lugubriformis 3-58 guianense 4-48 caucaensis 3-28 macbridei 3-19 heterophlebia 4-45 cheilanthoides 3-34 macrophylla 3-68 ichtiosma 4-44 chrysodioides 3-69 macrotis 3-57 lechleri 4-45 clivalis 3-45 mapiriensis 3-49 longicaudata 4-44 clypeata 3-65 maxoniana 3-71 meniscioides 4-48 coarctata 3-32 megalodus 3-64 opaca 4-45 comosa 3-49 membranacea 3-69 paludosa 4-48 comptula 3-23 mercurii 3-3 1 pellucidopunctata 4-44 concinna 3-29 micula 3-33 prasina 4-45 confluens 3-6 millei 3-28 rotundata 4-42 consobrina 3-73 multiformis 3-35 Struthiopteris conspersa 3-42 navarrensis 3-15 maxonii 5-61 contermina 3-32 nemoralis 3-62 Sr>>///es 6-89 corazonensis 3-27 nitens 3-35 andicola 6-92 ctenitoides 3-37 oligocarpa 3-15 gemmifera 6-92 curta 3-60 oligophlebia 3-3 Syngramma decussata 3-48 opposita 3-32 brevifrons 2- 1 5 deflexa 3-30 opulenta 3-43 paraphysata 6-97 deltoidea 3-46 pachyrhachis 3-3 1 demissa 3-21 palustris 3-6 densa 3-34 parasitica 3-39 Taenitis dentata 3-41 patens 3-43 desvauxii 5-147 depilata 3-45 pavoniana 3-23 furcata 5-148 diplazioides 3-14 pennata 3-64

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 121 pennellii 3-50 angustatum 1-83 spruceanum 1-91 peripae 3-60 angustifrons 1-86 sprucei 1-86 peruviana 3-23 ankersii 1-90 subsessile 1-90 phacelothrix 3-22 applerianum 1-88 tanaicum 1-89 pilosohispida 3-27 arbuscula 6-97 tenerum 1-83 pilosula 3-19 bancroftii 6-97 trollii 1-91 pinnatifida 3-56 bicorne 1-91 tuerckheimii 1-90 poiteana 3-65 botryoides 1-92 undulatum 1-97 proboscidea 3-36 brachyblastos 1-81 vandenboschii 1-97

ptarmiciformis 3- 1 7 capillaceum 1-84 Trichopteris 1-122 pteroidea 3-25 cellulosum 1-85 Triplophyllum 4-19 pusilla 3-17 ciliatum 1-67 acutilobum 4-2 1 quadrangularis 3-4 1 collariatum 1-81 dicksonioides 4-2 1 resinosofoetida 3-34 coriaceum 6-97 funestum 4-2 1 reticulata 3-66 crinitum 1-94 protensum 4-19 retrorsa 3-27 crispum 6-97 Trismeria 2-16 rudis 3-25 cristatum 1-97 aurea 2-16

rufa 3-20 delicatum 1-98 microphylla 2-2 1 ruiziana 3-35 diaphanum 1-83 trifoliata 2-21 salicifolia 3-74 diversifrons 1-91 Trogonospora 3-2 salzmannii 3-73 ekmanii 1-88 scalaris 3-15 datum 1-96 schunkei 3-63 elegans Rich. 1 -84 Urostachys 6-19 semihastata 3-57 elegans Rudge 1 -9 1 andinus 6-35 serrata 3-71 flaccida 2-95 attenuatus 6-37 sodiroi 3-62 fragile 1-69 bifida 6-28 sprengelii 3-3 1 fucoides 1-56 brevifolius 6-39 subandina 3-20 haenkeanum 1-98 capellae 6-32 supina 3-26 heterophyllum 1-94 caracasicus 6-40 tetragona 3-64 hirsutum 1-67 catharticus 6-38 thomsonii 3-22 hookeri 1-88 cuatrecasasii 6-45 torresiana 3-3 hostmannianum 1-93 binervius 6-26 totta 3-40 humboldtii 1-94 brogniarti 6-27 tristis 3-60 hymenoides 1-86 buesii 6-42 tryonorum 3-62 hymenophylloides 1-83 crassus 6-34 urbanii 3-64 krausii 1-87 cuneifolius 6-50 valdepilosa 3-50 kunzeanum 1-81 curvifolius 6-46 Tmesipteris 6-1 1 lambertianum 1-95 darwinianus 6-33 Todea 1-20 leptophyllum 1-83 durissimus 6-5 1 Trachypteris 2-48 lucens 1-95 elongatus 6-35 aureonitens 2-48 martiusii 1-94 engleri 6-39 induta 2-50 membranaceum 1-88 ewanii 6-50 pinnata 2-48 muscoides 1-86 funiformis 6-42 Trichipteris 1-120 opacum 1-84 hartwegianus 6-40 conjugata 1-128 pedicellatum 1-90 hippurideus 6-24 corcovadensis 1-120 pellucens 1-96 hohenackeri 6-40 dombeyi 1-123 pellucidum 1-96 jenmanii 6-44 excelsa 1-120 peltatum 1-57 kuesteri 6-30 flava 1-125 pennatum 1-92 lechleri 6-25 frigida 1-126 pilosum 1-95 var. lehmannii 6-25 infesta 1-123 pinnatum 1-92 lehmannii 6-25 kalbreyeri 1-124 plumosum 1-96 linifolius 6-43 lasiosora 1-125 plumula 1-95 var. tenuifolius 6-44 latevagans 1-124 poeppigii 1-89 macbridei 6-32 lechleri 1-127 polyanthos 1-59 mexiae 6-28 microdonta 1-127 polypodioides 1-89 molongensis 6-47 nigra 1-124 prieurii 1-84 myrsinites 6-49 nigripes 1-124 punctatum 1-86 nesselii 6-34 phegopteroides 1-127 pyxidiferum 1-82 nubigenus 6-49 pilosissima 1-125 radicans 1-81 phlegmaria procera 1-123 reptans 1-87 var. ericaefolius 6-48 pubescens 1-126 rigidum 1-85 phylicifolius 6-49 serpens 1-126 rupestre 1-82 pilgerianus 6-34 tryonorum 1-128 scandens 1-76 polycarpos 6-45 Trichomanes 1-76 sellowianum 1-97 poseidonis 6-24 accedens 1-98 sphenoides 1-86 pruinosus 6-5 1

122 FIELDIANA: BOTANY reflexus 6-27 pyxidifera 1-82 erenata 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 >uH/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

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Pnhliratinn 1403

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[RIDOPHYTA OF PERI

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PTERIDOPH^

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