ILLINOIS LIBRARY

,M URBANA-CHAMPAIGN

BIOLOGY

Botany

NEW SERIES, NO. 34

PTERIDOPHYTA OF PERU

Part VI

22. Marsileaceae-28. Isoetaceae

Rolla M. Tryon

Robert G. Stolze

With the collaboration of:

R. James Hickey

Benjamin 011gaard

December 30, 1994

Publication 1461

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

Information for Contributors to Fieldiana

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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

ACKNOWLEDGMENTS 1

22. MARSILEACEAE 2

Marsilea 2

23. SALVINIACEAE 5

Salvinia 6

Azolla 8

24. PSILOTACEAE 11

Psilotum 11

25. EQUISETACEAE 12

Equisetum 12

26. LYCOPODIACEAE 16

Huperzia 19

Lycopodium 52

Lycopodiella 58

27. SELAGINELLACEAE 66

Selaginella 66

28. ISOETACEAE 88

Isoetes 89

ADDENDUM 97

1 . Species to Be Added to the Pterido-

phyte Flora, Parts I-V 97

2. Consideration of Pteridophyte Diversity

in Respect to Ecology and Geography ... 98

DEPARTMENTS OF PERU 103

COMPREHENSIVE INDEX TO NAMES . . 1 04

1. Marsilea: M. ancylopoda; M. vestita .... 3

2. Salvinia: S. minima; S. auriculata 7

3. Azolla: A. filiculoides\ A. microphylla .... 9

4. Psilotum: P. nudum . .13

14

18

53

5. Equisetum: E. giganteum; E. bogotense

6. Huperzia: H. eversa; H. binervia; H.

linifolia var. tenuifolia; H. cuneifolia .

1. Lycopodium: L. clavatum; L. thyoides

8. Lycopodiella: L. caroliniana var. meridi-

onale; L. cernua 59

9. Selaginella: S. peruviana; S. haema-

todes; S. exaltata 67

10. Isoetes: I. andicola; I. lechleri; I. di-

spora 90

List of Tables

1 . Data on the diversity of pteridophyte

species in Peru

100

List of Maps

1 . The most species-rich departments in

Peru 99

2. The main vegetational zones in Peru and

their ferns . .101

111

Back cover: Selaginella haematodes

PTERIDOPHYTA OF PERU

Part VI

22. Marsileaceae-28. Isoetaceae

Introduction

This sixth and final part of the "Pteridophyta

of Peru" contains the aquatic fern families Mar-

sileaceae and Salviniaceae as well as the "fern al-

lies" Psilotaceae, Equisetaceae, Lycopodiaceae,

Selaginellaceae, and Isoetaceae. A brief section on

the diversity and ecology are included, written by

the senior author. The comprehensive index con-

tains not only the names in this final part but those

of the other five parts as well. The general style,

typography, form of citations, and so forth follow

the previously published parts. These matters are

adequately dealt with in Part I (Fieldiana: Botany,

n.s., No. 20, 1 989), and it is not necessary to repeat

them here.

Treatment of the Lycopodiaceae has been con-

tributed by Benjamin 011gaard and that of Isoeta-

ceae by R. James Hickey. Both are recognized spe-

cialists in these two families. The other genera are

a joint effort of Rolla M. Tryon and Robert G.

Stolze, each critically reviewing the treatments

prepared by the other.

Type collections from Peru are mentioned in

the nomenclature but are not repeated in the spec-

imen citations. They are, however, included in the

Peru range and ecology. The nomenclature of the

genera and species is not intended to be complete.

It includes all names based on Peru material and

other names that are considered useful to mention.

Abbreviations of periodicals generally follow the

system of Botanico-Periodicum-Huntianum

(1968), while those of books and authors generally

follow the system of Taxonomic Literature (TL-

2, 1976 et seq.). The acronyms for herbaria follow

Index Herbariorum and are also provided below.

Acknowledgments

Benjamin 011gaard produced a recent Index to

the names of Lycopodiaceae, as well as taxonomic

treatments for the family in "The ferns and fern

allies of Guatemala" and in the "Flora of Ecua-

dor." R. James Hickey contributed Isoetaceae for

"The ferns and fern allies of Guatemala" and is

currently working on a monograph of the family.

Consequently, both have a keen understanding of

the problems in these pteridophyte families, and

for their outstanding efforts in the production of

this Flora the authors wish to express their deep

appreciation.

We would like to extend special thanks to Blanca

Leon (USM) for her invaluable assistance in pre-

paring loans and arranging for their packing and

shipment from this important Peruvian herbari-

um, as well as from the Universidad Nacional de

Trujillo (HUT). Rolla M. Tryon appreciates the fa-

cilities provided by the Department of Biology,

University of South Florida, Tampa, and the aid

of Alice F. Tryon in the preparation of the treat-

ment of Selaginella.

The illustrations were contributed by Field Mu-

seum scientific illustrator Zorica Dabich, who cre-

ated the original drawings and adapted the rest

from those previously used in the Fieldiana: Bot-

any publication "The ferns and fern allies of Gua-

temala." Her art work, which now has appeared

in all six parts of this Flora, is an invaluable com-

plement to the descriptions. We are extremely

grateful to her. Thanks also go to Bent Johnsen,

free-lance artist of Copenhagen, Denmark, for his

drawing ofHuperzia. We also appreciate the valu-

able comments presented by reviewers of the

manuscript.

We are grateful to the officers of the following

institutions for granting loans of their material or

allowing us to examine specimens in their her-

baria: Herbarium Jutlandicum, Aarhus Univer-

sitet, Denmark (AAU); Field Museum of Natural

History, Chicago (F); Harvard University, Cam-

bridge, Massachusetts— most Gray Herbarium

(GH), some Arnold Arboretum (A); Herbarium

Truxillense, Universidad Nacional de Trujillo,

Trujillo, Peru (HUT); Missouri Botanical Garden,

St. Louis (MO); Miami University, Oxford, Ohio

(MU); Museum National d'Histoire Naturelle, Par-

is (P); University of California, Berkeley (uc);

United States National Herbarium, Smithsonian

Institution, Washington, D.C. (us); and Herbario

FIELDIANA: BOTANY, N.S., NO. 34, DECEMBER 30, 1994, PP. 1-123

San Marcos, Universidad Nacional Mayor de San

Marcos, Lima, Peru (USM).

Materials, mostly types, have also been studied

in the following herbaria (acronyms follow Index

Herbariorum, ed. 8): AWH, B, BKL, BM, BONN, BR,

C, CGE, CPUN, CR, DUKE, G, GB, GL, HB, HEID, K,

KRA, L, LG, LIL, LINN, LP, MA, MICH, MOL, MSC, NO,

NY, PR, Q, QCA, RB, S, SAPF, U, UPS, W, WIS, and Z.

This project has been supported in part by grant

#BSR-85-16358 from the National Science Foun-

dation, Systematic Biology Program. The work

would not have been possible without this assis-

tance. However, any opinions and conclusions ex-

pressed are those of the authors and do not nec-

essarily reflect the views of the Foundation.

Family 22. MARSILEACEAE

Marsileaceae Mirb., Hist. nat. veg. (Lam. & Mirb.)

5: 126. 1802. TYPE: Marsilea L.

Stem short- to long-creeping, slender, often

branched, hardly indurated, bearing trichomes.

Leaves ca. 1-40 cm long, with 4, 2, or no leaflets

at the apex of the petiole, circinate in the bud. Sori

borne within sporocarps, indusiate, with indehis-

cent, short- to long-stalked, not annulate, mega-

sporangia and microsporangia; heterosporous,

spores without chlorophyll.

The Marsileaceae are a family of three genera,

all in tropical America. The genera (Marsilea, Reg-

nellidiwn, and Pilularid) are clearly distinct and

not very closely allied.

There are numerous detailed studies on the

Marsileaceae centering on the form and devel-

opment of the elaborate reproductive structures

and simplified leaves. These plants have also been

useful in physiological and experimental work on

factors influencing leaf form. However, some of

the basic problems concerning the nature of the

leaflets and the sporocarp are not resolved, and

the phyletic relationships of the three genera are

not wholly certain.

Pilularia L. is not known from Peru. However,

Pilularia americana A. Br. has been collected near-

by in Bolivia (as P. mandonii) and it probably also

grows in Peru. It is a small plant with no leaflets

on the petiole; the filiform leaves (petioles) are 1-

1 0 cm long. A single subglobose sporocarp is borne

at the base of a leaf. Unless sporocarps are present,

the plant can easily be taken for a species of Cy-

peraceae, which grows in wet habitats that are sim-

ilar to those of Marsilea.

I. Marsilea

Marsilea L., Sp. pi. 1099. 1753; Gen. pi. ed. 5,

485. 1754. TYPE: Marsilea quadrifolia L.

Figure 1.

Plants palustral or aquatic. Stem usually long-

creeping, commonly bearing long roots at the nodes

of the stem or along the internodes. Leaves with

the petiole terminated by 2 adjacent pairs of nar-

rowly cuneate to broadly flabellate leaflets that are

glabrous or pubescent. Veins more or less anas-

tomosing, usually connected at the margin. Sori

borne within 1 to several stalked, indurated spo-

rocarps attached to the petiole or at its base, en-

closed by a diaphanous indusium, with microspo-

rangia and megasporangia. Megaspores somewhat

ellipsoidal with an apical papillalike laesura, the

surface papillate. Microspores spheroidal, trilete,

the surface slightly rugulose.

Marsilea is a nearly worldwide genus of perhaps

50 species, with about 1 2 in America. The tropical

American species are poorly known for there are

relatively few collections and these often lack spo-

rocarps. The species of Marsilea are most diverse

and common in regions that support vernal pools

or shallow pools that dry out, at least along the

borders. Mexico is one center of diversity, Aus-

tralia and Africa are others.

The following treatment is based directly on that

of Johnson (1986). When necessary, specimens are

indicated as cited by Johnson.

The sporocarp is borne on a stalk that is con-

nected to the petiole. If this stalk is attached by

its end to the sporocarp, there is neither raphe nor

inferior tooth. If the stalk is attached along its side

(the raphe), there may be an inferior tooth (the

end of the stalk). The superior tooth is an extension

of the sporocarp near the inferior tooth (fig. Id).

FIG. 1 . Marsilea ancylopoda: a, habit; b, leaflet; c, stem and roots, with sporocarp. Marsilea vestita: d, sporocarps.

(a, b from Alston 6354, Venezuela, F, c from Alston 5874, Venezuela, F, d from Sandberg, United States (Idaho), F.)

FIELDIANA: BOTANY

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

The sporocarps may be very long-lived (to 100

years; Johnson, 1986) and they presumably are

dispersed by waterfowl. Species also may be dis-

persed as a result of discarded aquaria.

JOHNSON, D. M. 1986. Systematics of the New

World species ofMarsilea (Marsileaceae). Syst.

bot. monogr. 11: 1-87.

References

GUPTA, K. M. 1962. Marsilea, Bot. monogr. 2,

pp. 1-113, Council Sci. India Res., New Delhi.

Key to Species of Marsilea

a. Roots borne only at the nodes of the stem (rarely a few to 1 cm distant from a node); raphe present

b

b. Superior tooth acute 3. M. vestita

b. Superior tooth blunt or absent c

c. Sporocarps borne below the ground or stem level; leaflets essentially glabrous

2. M. ancylopoda

c. Sporocarps borne above the ground or stem level; leaflets pubescent 4. M. mollis

a. Roots borne at the nodes of the stem and along the internodes d

d. Sporocarps 1-4, borne on the basal 'A of the petiole 1. M. deflexa

d. Sporocarps 5-20, borne on the basal % of the petiole M. crotophora (see Comments)

1. Marsilea deflexa A. Br., Monatsber. konigl.

preuss. akad. wiss. Berlin. 1863: 421. 1864.

TYPE: Piauhy, Brazil, Gardner 2760 (holo-

type, G; isotypes, BM, G, K, P, all cited by John-

son).

Plants forming dense colonies. Roots at nodes

and internodes. Sporocarps 1-4, on basal '/4 of the

petiole, 3.5-6 mm long with conspicuous lateral

ridges.

In a pool, ca. 59 m; a single collection from San

Martin.

Mexico and Central America; Venezuela and

Colombia, south to Brazil; Paraguay and Peru.

San Martin: Morales, Tarapoto, Vie 6866 (G, K, P, all

cited by Johnson).

2. Marsilea ancylopoda A. Br., Monatsber. ko-

nigl. preuss. akad. wiss. Berlin. 1863: 434.

1864. TYPE: Guayaquil, Ecuador, Jameson

304 (holotype, G; isotypes, BM, G, K, all cited

by Johnson). Originally as James 304, cor-

rected later (A. Br., 1871, op. cit. 1870: 717).

Figure la-c.

Roots borne only at the nodes (rarely a few 2-

3 cm distant). Raphe present. Superior tooth ab-

sent or slightly raised. Sporocarps 2.5-6 mm long,

borne below the ground or stem level, 1 at the base

of a petiole. Leaflets of non-floating leaves gla-

brous or with a few trichomes.

Northern Peru, Tumbes to La Libertad, near sea

level.

Mexico; Florida and Greater Antilles; Venezue-

la, Colombia to Peru; Brazil; Argentina; Uruguay.

Tumbes: Laguna de Salitoral Grande, Coronado 235

(GH, uc). Lambayeque: San Nicolas, entre Lambayeque

et Chiclayo, Santos Quiroz 2401 (F, GH). Prov. Lam-

bayeque, Yencala, Santos Quiroz 71 (HUT). Laguna Boro,

Chiclayo, Leon 600 (USM). La Libertad: Prov. Trujillo,

Huaman, Angulo & Lopez 348 (GH).

3. Marsilea vestita Hooker & Grev., Icon. fil. 2:

t. 159. 1830. LECTOTYPE (designated by

Johnson, Syst. bot. monogr. 11: 62. 1986):

Columbia River, Scouler 338 (K; isolecto-

types, GH, NY). Figure Id.

FIELDIANA: BOTANY

Marsilea uncinata A. Br., Amer. J. Sci. ser. 2, 3: 55.

1 847. TYPE: Arkansas, Englemann 33 (holotype,

MO; isotypes, K, M, MO, all cited by Johnson).

Marsilea mucronata A. Br., Amer. J. Sci. ser. 2, 3: 55.

1847. TYPE: Near Devil's Lake, North Dakota,

Geyer 71 (holotype, MO; isotypes, K, NY, all cited

by Johnson).

Plants in diffuse or dense colonies. Roots only

at the nodes (rarely a few to 10 mm distant). Raphe

present. Superior tooth acute, often hooked at apex.

Sporocarp 3.0-7.6 mm long with lateral ridges.

Leaflets of non-floating leaves covered by over-

lapping trichomes on both surfaces.

All of the above pertains to ssp. vestita; ssp.

tenuifolia (A. Br.) Johnson is endemic to central

Texas. It differs from ssp. vestita in having very

narrow leaflets with sparse trichomes or none

abaxially.

Known in Peru only from the Department of

Lambayeque.

Southern British Columbia east to western Min-

nesota; south to Louisiana, Florida; Mexico; Peru.

The Peru and Florida localities may be introduc-

tions.

Lambayeque: Chiclayo, Leon (MICH, det. Johnson).

Prov. Lambayeque, Salazar HI (USM).

4. Marsilea mollis Rob. & Fern., Proc. Amer.

Acad. 30: 123. 1895. TYPE: San Diego, Chi-

huahua, Mexico, Hartman 604 (holotype, GH;

isotypes, F, GH, MSC, NY, uc, all cited by John-

son).

Roots borne only at the nodes (occasionally to

10 mm distant). Raphe present. Superior tooth

absent or to 0.2 mm long. Sporocarps at or near

petiole base, 1.7-6.7 mm long, on recurved pe-

duncles, borne above the ground level. Leaflets of

non-floating leaves adaxially glabrous to sparingly

pilose, abaxially densely pilose.

2000-3900 m, Cajamarca south to Puno.

Arizona and Texas south to northern South

America, south to Argentina, Chile and Brazil.

The following sterile specimens are considered

to represent this species. All have been seen by the

present authors except the Hill collections from

Puno. All have been placed here by Johnson except

the Nunez et al. collection from Cuzco.

Cajamarca: Cajamarca, Muller & Gutte 27322 (USM).

Junin: Huancayo, Kunkel 422 (GH). Huancavelica: Prov.

Angaraes, 4 km W of Huanta, Stork & Morton 10808 (F,

uc). Cuzco: Prov. Espinar, Yauri, Nunez et al. 7810 (F,

MO), Vargas 13524 (GH). Puno: Towards Juliaca, Hill

562 (K), 563 (K).

Comments

Marsilea crotophora D. M. Johnson, Syst. bot.

monogr. 11: 46. 1986. TYPE: Mato Grosso,

Brazil, Hatschbach & Scherer 30470 (holo-

type, us; isotypes, c, LP, M, MICH, NY, uc, all

cited by Johnson).

This species may be found in the upper Amazon

or its tributaries. It may be distinguished from

Peruvian species by the roots being borne on the

internodes as well as the nodes, the lack of a raphe

and superior tooth, and the 5-20 sporocarps borne

on the basal % of the petiole, the lowest one borne

well above the petiole base.

Family 23. SALVINIACEAE

Salviniaceae Reichenb., Bot. damen kunst. freunde

pflanzenw. 255. 1828. TYPE: Salvinia Se-

guier.

Azollaceae Wettst. Handb. syst. bot. 2: 77. 1903.

TYPE: Azolla Lam.

Plants floating in water or stranded in wet mud.

Stem elongate, small, and slender, often branched,

not indurated, usually bearing trichomes. Floating

leaves ca. 0.5-2.5 cm long, not circinate in the

bud. Sori borne on a lobe of a leaf or on a branched

leaf, indusiate, with the sporangia stalked, not an-

nulate, the sori either megasporangiate or micro-

sporangiate; heterosporous, spores without chlo-

rophyll.

The structure bearing the sporangia is consid-

ered to be a sorus, surrounded by an indusium,

and accordingly the special term sporocarp is not

used.

The living Salviniaceae are represented by two

genera, Salvinia and Azolla, floating aquatics of

wide distribution, especially in the tropics. Al-

though Azolla is sometimes included in a separate

family, it has many basic similarities with Sal-

vinia.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

Key to Genera of Salviniaceae

a. Leaves in whorls of 3, 2 floating, ca. 5-25 mm long, with anastomosing veins and not lobed, 1

submerged, highly branched I. Salvinia

a. Leaves alternate, ca. 0.5-1.5 mm long, without veins, bilobed II. Azolla

I. Salvinia

Salvinia Seguier, Fl. veron. 3: 52. 1754. TYPE:

Salvinia natans (L.) All. (Marsilea natans L.)

Figure 2.

Plants floating aquatics. Stem short or elongate,

slender, bearing trichomes and no roots. Leaves

borne in whorls of 3, dimorphic, 2 of them green,

floating, entire, oblong to suborbicular, ca. 0.5-2.5

cm long, often pubescent, usually papillate on the

upper surface, one of them submerged, pendent in

the water, highly branched, bearing many tri-

chomes. Veins anastomosing. Sporangia borne in

stalked sori on the submerged leaf, enclosed by

the indusium, with either megasporangia or mi-

crosporangia. Spores trilete, enclosed within a vac-

uolate tapetal formation, the megaspore surface

perforate, the microspore surface somewhat ru-

gulose.

Salvinia is a widely distributed genus of about

ten species, with seven of them in America.

Shaparenko (1956) recognized four sections, and

these were adopted by de la Sota (1962); they are

not recognized here because the genus is a small

one.

The descriptions of the species are freely adapt-

ed from Stolze(1983).

References

SHAPARENKO, K. K. 1956. History of the Sal-

vinias (in Russian). Act. Inst. Bot. Komarov.

Ser. VIII Paleobotania 2: 1-44.

SOTA, E. R. DE LA. 1962-1964. Contribution al

conocimenti de las "Salviniaceae" neotropi-

cales. I-V. Darwiniana 12: 465-520, 612-623;

13: 529-536.

STOLZE, R. G. 1983. Ferns and fern allies of Gua-

temala, III. Fieldiana: Bot., n.s. 12: 1 1-13.

Key to Species of Salvinia

a. Trichomes at the apex of the papillae, on the upper surface of the floating leaves, separate (not joined)

at their tip 1 . S. minima

a. Trichomes at the apex of the papillae, on the upper surface of the floating leaves, joined at their tip

. 2. S. auriculata

1. Salvinia minima Baker, J. Bot. 24: 98. 1886.

TYPE: Santa Catarina, Brazil, Muller 479

(holotype, K; isotype, BM). Figure 2a-c.

Stele of the stem more or less circular in cross-

section. Floating leaves oblong-elliptic, the apex

obtuse (rarely retuse), the larger ones 0.6-1.3 cm

long, their upper surface papillate, each papilla

with apical trichomes, these separate (not joined)

at the apex.

130-160 m, Loreto and Madre de Dios.

Southeastern United States; Mexico and Central

America; South America, south to Argentina and

Uruguay.

This species has often been called S. rotundifolia

but that name = S. auriculata.

FIG. 2. Salvinia minima: a, habit; b, portion of leaf showing venation; c, papillae with free, spreading trichomes.

Salvinia auriculata: d, habit; e, papillae with joined trichomes. (Adapted from Stolze, Ferns and fern allies of

Guatemala, 1983.)

FIELDIANA: BOTANY

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

Loreto: Iquitos, McDaniel 11776 (MO). Near Iquitos,

Ellenberg 8301 (F). Rio Itaya, Iquitos, Revilla 467 (F,

MO, us, USM). Lupuna Cocha, Iquitos, Tryon & Tryon

5188 (F, GH, us, USM). Madre de Dios: Prov. Tambopata,

Tambopata Nature Reserve, 30 air km SSW of Puerto

Maldonado, Barbour 5373 (MO). Prov. Manu, Parque

Nacional Manu, Foster 12206 (F).

2. Salvinia auriculata Aublet, Hist. pi. Guiane

Franc. 2: 969. 1775. TYPE: French Guiana,

Terr. Caux, Aublet (p?). Figure 2d-e.

Salvinia rotundifolia Willd., Sp. pi. ed. 4, 5: 5 1 7. 1 8 1 0.

TYPE: Brazil, Hoffmannsegg(see Morton, Contr.

U.S. Natl. Herb. 38: 75. 1967).

Stele of the stem U-shaped in cross-section.

Floating leaves oblong-elliptic and nearly circular,

obtuse or retuse at the apex, larger floating leaves

1 .5-2.5 cm long, their upper surface papillate, each

papilla with 4 apical trichomes, these joined at the

apex.

At ca. 100 m, Loreto and Madre de Dios.

Cuba; Mexico south to Argentina.

Loreto: Rio Amazonas, SE of Iquitos, Croat 19296

(MO). Prov. Maynas, mouth of Rio Nanay, Gentry el al.

21715 (MO, uc). Yanamono, Gentry et al. 42574 (F, MO).

Madre de Dios: Prov. Tambopata, Tambopata Nature

Reserve, 30 air km SSW of Puerto Maldonado, Barbour

4888 (MO).

II. Azolla Lam., Encycl. 1: 343. 1783. TYPE:

Azolla filiculoides Lam. Figure 3.

Floating aquatics. Stem short, elongate, slender,

sometimes bearing trichomes, and usually short

roots. Leaves ca. 0.5-1.5 mm long, bilobed, the

upper lobe usually minutely to strongly papillate,

without veins. Sporangia borne in short-stalked

sori enclosed by an indusium, with either 1 mega-

sporangium or several microsporangia. Spores tri-

lete, the megaspores irregularly marked, with a

perforate surface, the microspores smooth, em-

bedded in massulae.

The megaspore has accessory structures usually

referred to as a columella bearing floats. These are

partially enclosed by a band of sporoderm, the

collar, which is between the megaspore proper and

the floats. In the living species the floats are either

three or nine in number. The microsporangia con-

tain a few to several massulae, within which the

microspores are embedded. Projecting structures

called glochidia are often on the outer surface of

the massulae.

Azolla is a widely distributed genus of six spe-

cies, with four of them (sect. Azolla) in America

and in Peru. It is notable in having the most com-

plex reproductive structures of any plant and in

having a colony of Anabaena azollae within the

lobes of each leaf. This blue-green alga has the

ability to fix atmospheric nitrogen, and accord-

ingly Azolla is the center of wide interest as a green-

manure (Lumpkin & Plucknett, 1980), especially

for the rice paddies of Southeast Asia.

Azolla is a very distinctive genus, but the species

are poorly known due to the fact that sterile ma-

terial is difficult to identify. Fertile material is un-

common. Accordingly, the species in Peru and their

characters are only provisionally known. In spite

of evidence to the contrary, the treatment of Sven-

son (1944) is largely followed. The key is adapted

from his key to species of the Americas and the

megaspore characters are taken from Perkins et al.

(1985). Species names may be assigned to one of

the four species adopted, although this does not

mean that the material is accurately determined.

The following Peru collections have not been iden-

tified to species: Cajamarca: Dillon 2919. La Li-

bertad: Killip & Smith 21511; D. Smith et al. 2225.

Cuzco: Herrera 2616, 2618; Cook & Gilbert 237,

241, 1078; Cook 1963; Maldonado 17. Puno: Bar-

clay 9273.

Species of Azolla, as they are known, are not

restricted to particular environments as differ-

ences in range and habitat statements might imply.

Accordingly, the range, habitat, and altitude are

given for the genus as a whole in Peru, rather than

for each species.

In wet mud of riverbanks or stream banks, where

stranded by higher water, usually floating in stand-

ing water of lakeshores, ponds, or small streams,

also in ditches, in stagnant or fast-flowing water,

less often in swamps or wet parts of cultivated

land, near sea level to 4100 m, Cajamarca, Ama-

zonas, and Loreto, south to Puno and Moquegua.

FIG. 3. Azolla filiculoides: a, habit; b, megasporangium; c, sporocarp indusia with microsporangia; d, massula

with nonseptate trichomes. Azolla microphylla: e, megasporangium; f, massula with septate trichomes. (Adapted from

Stolze, Ferns and fern allies of Guatemala, 1983.)

FIELDIANA: BOTANY

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

References

LUMPKIN, T. A., AND D. L. PLUCKNETT. 1980.

Azolla, botany, physiology, and use as a green-

manure. Econ. Bot. 34: 1 1 1-153.

PERKINS, S. K., G. A. PETERS, T. A. LUMPKIN, AND

H. E. CALVERT. 1985. Scanning electron mi-

croscopy of perine architecture as a taxonomic

tool in the genus Azolla Lamarck. Scanning

Electron Microscopy 4: 1719-1734.

SVENSON, H. K. 1944. The New World species

of Azolla. Amer. Fern J. 34: 69-84.

Key to Species of Azolla

a. Plants dichotomously branched, 0.5-1.5 cm long; leaves less than 1 mm long; megaspore collar with

or without filaments b

b. Plants small, 0.5-1.0 cm in diameter; leaves small (0.5 mm long), nearly orbicular and nearly

smooth; glochidia not septate; megaspore collar with dense filaments, the megaspore surface more

or less smooth, obscured by dense filaments 1 . A. caroliniana

b. Plants often larger, 1.0-1.5 cm in diameter; leaves with the upper lobe 0.7 mm long, the lower

lobe much larger, more or less obovate; glochidia septate; megaspore collar lacking filaments,

megaspore surface with large and small pits, but few filaments 2. A. mexicana

a. Plants pinnately branched, 1-6 cm long; leaves 1 mm long; megaspore collar lacking filaments . .c

c. Plants often 2-6 cm long; leaves oblong to ovate; glochidia not septate (or with 1-2 septa distally);

megaspore surface with raised hexagonal markings 3. A. filiculoides

c. Plants 1-2 cm long; leaves nearly orbicular; glochidia septate, some septa located below the middle;

megaspore surface rugulate-verrucate 4. A. microphylla

1 . Azolla caroliniana in the sense of Svenson and

other authors, not Willd., as usually thought.

Plants dichotomously branched, 0.5-1.0 cm in

diameter. Leaves divaricate (not closely imbri-

cate), nearly orbicular, nearly smooth, small, 0.5

mm long. Glochidia not septate. Megaspore collar

with dense filaments on its surface (magnification

x200 or more), rugulate-verrucate beneath fila-

ments.

Eastern North America; Mexico and the Carib-

bean; introduced in Europe and South America.

The name is used in the sense of Svenson and

most authors (not Willd.) until a definitive treat-

ment is published.

Jermy (Somerfeltia 6: ix-x. 1987) has pointed

out that according to the Ph.D. thesis of D. Dun-

ham (Portsmouth Polytechnic) the type of Azolla

caroliniana is A. filiculoides. This leaves Azolla of

eastern North America presumably without a

name.

Amazonas: Rio Santiago, S of La Poza, Berlin 3711

(MO, uc). La Libertad: Prov. Bolivar, Laguna de las Ichus,

Lopez & Sagdstegui 3238 (GH, MO). San Martin: Prov.

Mariscal Caceres, 2-4 km W of Tocache Nuevo, Plow-

man 11437 (F). Loreto: Prov. Maynas, Rio Itaya, Revilla

632 (F, MO, uc.). Prov. Maynas, Yanuyaca, Gentry et al.

32119 (GH). Ayacucho: 44 km N of Tambo, Luteyn &

Lebron-Luteyn 6369 (uc, us). Cuzco: Prov. Cuzco, Say-

lla, Fernandez et al. 455 (USM). Puno: Santa Rosa, Staf-

ford 495 (F). Ichu, Soukup 354 (F).

2. Azolla mexicana Presl, Abh. Bohm. Ges. Wiss.

Ser. V, 3: 150. 1845. TYPE: Mexico, Schiede

& Deppe (holotype, PR).

Plants dichotomously branched, often 1.0-1.5

cm in diameter. Leaves divaricate (not closely im-

bricate) with the upper lobe ca. 0.7 mm long, the

lower one much larger. Glochidia septate. Mega-

spore collar glabrous; megaspore surface (magni-

fication x 200 or more) coarsely rugose, also with

pits, and with many dense filaments.

North America; Mexico; Central America;

northern half of South America.

Tumbes: Casa Fernandez, Haught 183 (us). Loreto:

Prov. Maynas, Iquitos, Tryon & Tryon 5213 (GH, us).

Ancash: Lago Santa Cruz Chico, Huascaran National

Park, D. Smith et al. 9261 (F, HUT, MO). Junin: Huancayo,

Kunkel 421-lh (GH). Acopalca, Kunkel 421 (GH). Cerca

Carhuamayo, Ferreyra 5242 (USM). Cuzco: Pampa de

Anta, Iltis & Ugent 785 (GH, uc, us, USM). 3.5 km NW

of Saylla, Iltis et al. 1208 (GH, us). Prov. Paucartambo,

Paucartambo, Plowman & Davis 4920 (F). Madre de

Dios: Prov. Manu, Parque Nacional Manu, Foster 9872

10

FIELDIANA: BOTANY

(MO). Puno: Pucara, Hunnewell 15874 (GH). Lake Titi-

caca, Shepard 1511 (GH, us). Moquegua: Ilo, Stafford

926 (F).

3. Azolla filiculoides Lam., Encycl. 1: 343. 1783.

TYPE: "Magellan Region," Herb. Lamarck

(p). Figure 3a-d.

Plants pinnately branched, often 2-6 cm long.

Leaves closely appressed, imbricate, oblong to

ovate, ca. 1 mm long. Glochidia not septate (rarely

1-2 "septa" toward the apex). Megaspore collar

more or less glabrous; megaspore surface (mag-

nification x 200 or more) prominently rugose with

hexagonal markings, with few filaments.

Southern South America north to western North

America and to Alaska; introduced in Europe, Asia,

and Australia. Azolla filiculoides var. rubra (R. Br.)

Strasb., treated as a species A. rubra R. Br. by

Perkins et al. (1985), is not considered to occur in

Peru.

Cajamarca: Prov. Contumaza, Rio San Benito, San

Benito, Sagdstegui et al. 12509 (HUT, MO). Amazonas:

Prov. Bongara, Laguna Pomacocha, Wurdack888 (F, GH,

uc, us). La Libertad: Prov. Trujillo, Tschudi, Shimo-

kowa 7294 (HUT). Playa de Chan Chan, Angulo 18 (us).

Loreto: Paraiso, above Iquitos, Rio Amazonas, Fosberg

29076 (us). Huanuco: Chasqui, Macbride 3306 (F, us);

3307 (F, us). Lima: Prov. Lima, Laguna de Villa, Corona-

do 5 (GH, MO, uc, us); Tryon & Tryon 5458 (GH); Cerrate

2766 (USM). Huancavelica: Prov. Tayacaja, Pampas, Stork

& Morton 10243 (F, uc, us). Cuzco: Cook & Gilbert 197

(us). Prov. Paucartambo, Huilabamba, Balls B7632 (F,

uc, us). Arequipa: Prov. Arequipa, Rio Yarabamba,

Ponce 32 (USM).

Family 24. PSILOTACEAE

Psilotaceae Kanitz, Novenyrends. Attek. 43. 1887.

TYPE: Psilotum Sw.

Stem flaccid to somewhat indurated, lacking in-

dument and roots. Leaves (or pinnae) small to

minute, borne alternately, with a single vein or

none. Two or 3 sporangia joined in a sessile, thick-

walled synangium. Homosporous. Spores without

chlorophyll.

The Psilotaceae are a family of two genera, Tme-

sipteris Bernh. of western Malesia, Australia, and

the Pacific, and Psilotum, which is pantropical in

distribution.

After an extensive series of studies, D. W. Bier-

horst concluded that the Psilotaceae were primi-

tive elements in the Filicopsida. This interpreta-

tion of the position of the family provoked further

interest in its phylogeny and resulted in a sym-

posium on the taxonomic and morphological re-

lationships of the Psilotaceae (White et al., 1977).

The Psilotaceae have a number of characters

that occur in the Filicopsida, such as the details

of early ontogeny of the leaf, the subterranean,

cylindrical gametophytes, multiflagellate sperm,

and several aspects of the rhizoids and gametangia.

These and the evidence from the structure and wall

formation of Psilotum spores are indicative of re-

lationships with the ferns. However, Cooper- Driv-

er (1977) has shown that the chemical evidence

does not support this relationship.

References

4. Azolla microphylla Kaulf, Enum. fil. 273. 1 824.

TYPE: "California," Chamisso (LE?). Figure

3e-f.

Plants pinnately branched, small, 1-2 cm long,

many leaves orbicular, 1 mm long, most leaves

closely imbricate. Glochidia septate. Megaspore

collar glabrous; megaspore surface slightly fila-

mentous, slightly pitted, rugulate (high magnifi-

cation) to verrucate.

Western and northern South America; southern

North America; West Indies.

Loreto: Yurimaguas, Killip & Smith 27707 (us); Prov.

Requena, Rio Ucayali, Encarnacion E-1101 (us). Aya-

cucho: Puquio, Gentry et al. 23296 (F, MO, us). Cuzco:

Prov. Anta, Pampas de Anta, Tryon & Tryon 5364 (GH,

us, USM). Yauri, Nunez et al. 7813 (F, MO).

COOPER-DRIVER, G. 1977. Chemical evidence

for separating Psilotaceae from the Filicales.

Science 198: 1260-1261.

WHITE, R. A., ET AL. 1977. Taxonomic and mor-

phological relationships of the Psilotaceae. Brit-

tonia 29: 1-68.

I. Psilotum

Psilotum Sw., J. Bot. (Schrader) 1800 (2): 8, 109.

1 80 1 , nom. nov. for Hoffmannia Willd. (not

Sw.) and with the same type.

Hoffmannia aphylla Willd. = Psilotum nudum (L.)

Beauv. Figure 4.

Plants terrestrial, rupestral or usually epiphytic.

Stem compactly branched in the substrate, the ae-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

11

rial stems green, erect to pendent, with several

dichotomous divisions, ca. 20 cm to 1 m long,

without roots. Leaves minute, alternate, leafy stem

glabrous. Synangia large, borne singly in the axil

of forked fertile leaves. Spores elongate-ellipsoid-

al, monolete, the laesurae '/2 to % the spore length,

the surface coarsely rugose to shallowly and com-

pactly verrucate.

Psilotum is a highly distinctive genus of wide

distribution in the tropics and with some exten-

sions to subtemperate regions. Notable characters

of the genus are the dichotomously branched stem,

the bifurcate sporophylls, and the large synan-

gium. Psilotum has a simple morphology, proba-

bly through reduction from more complex ances-

tors.

1. Psilotum nudum (L.) Beauv., Prodr. Fam. Ae-

theog. 112. 1805. Figure 4.

Lycopodium nudum L., Sp. pi. 1100. 1753. TYPE:

LINN 1257.1.

Psilotum triquetrum Sw., J. Bot. (Schrader) 1800 (2):

109. 1801, nom. super/I, for Lycopodium nudum

L. and with the same type.

Plants pendent or usually erect. Branches 3-an-

gled, to 1 .2 mm wide, leaves distant, 2-3 mm long,

triangular-subulate.

Epiphytic, especially on palms and tree ferns,

200-600 m, San Martin south to Madre de Dios.

American and Old World tropics and subtrop-

ics.

P. complanatum Sw. may be a distinct species.

It has flattened branches 2 mm or more wide and

is especially distinctive in the Pacific islands.

San Martin: Dist. Chazuta, Rimachi 7945 (MO). Dist.

Tocache Nuevo, Schunke 4084 (F, us). Gramalote ad

Saposoa, Woytkowski 7300 (GH, MO, us). Loreto: Lower

Rio Huallaga, Killip & Smith 29244 (Coll. W. J. Dennis).

Padre Isla, Iquitos, Encarnacion 26314 (MO, us). Prov.

Requena, Rio Ucayali, Ayala et al. 3753 (MO). Huanuco:

Tingo Maria, Moran 3687 (USM). Madre de Dios: Prov.

Tambopata, Cuzco Amazonico Lodge, 15 km NE of

Puerto Maldonado, Nunez 12214 (MO); Gentry et al.

68958 (MO). Prov. Manu, Parque Nacional Manu, Foster

11336(F).

Family 25. EQUISETACEAE

Equisetaceae A. P. DC., Fl. Franc. (Lam. & DC.)

ed. 3, 2: 580. 1805. Type: Equisetum L.

Stem jointed, more or less indurated, the aerial

usually green, the subterranean with numerous wiry

roots. Leaves each with a single vein, borne in

whorls, joined in a sheath. Sporangia thin-walled,

borne on stalked, peltate sporangiophores that form

a strobilus. Homosporous. Spores with chloro-

phyll and bearing elaters.

I. Equisetum

Equisetum L., Sp. pi. 1061. 1753; Gen. pi. ed. 5,

484. 1754. TYPE: Equisetum fluviatile L. Fig-

ure 5.

Terrestrial, palustral or aquatic. Stem subter-

ranean, short- to long-creeping, freely branched,

bearing erect, aerial stems that are jointed, lon-

gitudinally ridged, and usually hollow, ca. 10 cm

to 8 m long, often with whorls of branches. Leaves

small, in whorls at nodes, the lower portion lat-

erally fused in sheaths, the upper portion in more

or less prolonged teeth. Sporangia large, several

borne on each stalked, peltate, apically flattened

sporangiophore, in a condensed terminal strobi-

lus. Spores spheroidal with circular aperture, and

4 paddle-shaped elaters, the surface with small

granulate and large spherical deposits.

The Equisetum strobilus is composed of spo-

rangiophores that are sometimes called sporo-

phylls. A detailed study of these structures by Page

(1972) shows they are partly modified from a leaf

and partly from a cauline appendage; thus the term

sporangiophore is more appropriate. The small

leaves that surround each node are laterally fused

below and form coarse teeth above.

References

HAUKE, R. L. 1961-1962. A resume of the tax-

onomic reorganization of Equisetum subgenus

FIG. 4. Psilotum nudum: a, habit; b, portion of fertile branch with sporangium. (Adapted from Stolze, Ferns and

fern allies of Guatemala, 1983.)

12

FIELDIANA: BOTANY

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

13

5cm

14

FIELDIANA: BOTANY

Hippochaete, I-IV. Amer. Fem J. 51: 131-137; Equisetum subgenus Equisetum. Nova Hedwig-

52: 29-35, 57-63, 122-130. ia 30: 385^55.

HAUKE, R. L. 1963. A taxonomic monograph of PAGE, C. N. 1972. An interpretation of the mor-

the genus Equisetum subgenus Hippochaete. phology and evolution of the cone and shoot of

Beih. Nova Hedwigia 8: 1-123. Equisetum. J. Linn. Soc. Bot. 65: 359-397.

HAUKE, R. L. 1973. A taxonomic monograph of

Key to Species of Equisetum

a. Main aerial stem large, usually 10 mm or more wide (pressed), rarely less (to 5 mm), usually 2-5 m

tall, with a hollow center; strobilus sessile or nearly so 1 . E. giganteum

a. Main aerial stem small, to 2 mm wide (pressed), usually 0.5m tall or less, with a solid center; strobilus

stalked 2. E. bogotense

1 . Equisetum giganteum L., Syst. nat. ed. 10: 1318.

1759. TYPE: Plumier, PI. Amer. (ed. Bur-

man) t. 125, f. 2 (1757 as to the plate). Figure

5a-b.

Equisetum myriochaetum Schlect. & Cham., Linnaea

5: 623. 1 830. TYPE: Misantla, Veracruz, Mexico,

Schiede and Deppe 833 (holotype, B?).

Equisetum schaffneri Milde, Verb. Zool.-Bot. Ges.

Wien 1 1: 345. 1861. TYPE: Orizaba, Mexico, W.

Schaffner 315 (holotype, B?).

Stem usually 2-4 m tall, often bearing an apical

strobilus, usually 1 5-25 mm wide (10 mm or more,

rarely 5 mm or less), with a central hollow. En-

dodermis surrounding each of the vascular bun-

dles. Sheaths of the main stem usually light brown,

with often deciduous, rather indurated teeth.

Branches in regular whorls, with 8-10 ridges.

Strobilus sessile or nearly so.

In wet, usually open areas, in sloughs, on the

banks of streams, rivers, and irrigation ditches, in

boggy areas, and in streambeds, 1900-3500 m,

Piura, Cajamarca and Amazonas, south to Are-

quipa and Puno.

Greater Antilles; Mexico and Central America;

Venezuela and Colombia, south to Chile, Argen-

tina and Brazil.

Equisetum myriochaetum may be a species, as

Hauke and others have it, or it may be an extreme

variation of E. giganteum, as treated here. Most

likely it should be recognized as a subspecies, as

is E. ramosissimum ssp. debile, but we do not wish

to make a new combination at this time.

Equisetum giganteum has the turbercles on the

ridges of the stem square or flattened in profile

while the turbercles of E. myriochaetum appear

rather similar to the teeth of a saw. The former

species has two to several lines of stomates on each

side of a groove of the stem, while the latter species

has a single line of stomates. Characters of the

strobilus (apiculate or not) and of the sheath color

(different from the stem or not) are not species-

specific.

Equisetum schaffneri has a variable mixture of

the characters of E. giganteum and of E. schaffneri.

It has a wider distribution than a sterile hybrid

should have, and it occurs in regions in which only

one of the putative parents is known (Venezuela

and in Peru, for example). Accordingly, it may be

a natural intermediate of the extremes of E. gi-

ganteum. The abortive spores present in some col-

lections (not all have strobili) are also present in

some other species: in E. bogotense and E. var-

iegatum (Hauke, 1963, 1978) where hybridization

is not concerned.

Hauke (Ph.D. dissertation, University of Mich-

igan) cites the following collections of E. myri-

ochaetum from Peru: Ayacucho, Killip & Smith

22757 (us); Cuzco, Cook & Gilbert 1372 (us). Also

the following collections of E. x schaffneri from

Peru: Lima, Rose 18762 (NY, us); Ucayali?, Herre-

ra 1392 (us); Cuzco: Cook & Gilbert 1091 (us).

FIG. 5. Equisetum giganteum: a, habit; b, apex of branch with strobilus. Equisetum bogotense: c, habit; d, apex

of branch with strobilus. (a, b from Lowell 535, Ecuador, F; c, d from Cuatrecasas 1 1807, Colombia, F.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

15

Piura: Hills of Chiarnique, Barbour 2160 (MO). Lam-

bayeque: Olmos to Jaen, Correll & Smith P792 (GH).

Cajamarca: Prov. Cutervo, Llatun, Mostacero et al. 1740

(HUT). Prov. Celendin, alrededores de Celendin, Mos-

tacero et al. 835 (MO, uc). Amazonas: Prov. Bagua, 3.9

km NE of Chiriaco, Barbour 4360 (F, MO). Utcabamba,

Pennell 15586 (GH, us). La Libertad: Prov. Trujillo, valle

de Rio Chicama, Miiller & Krebs 12137 (USM). San Mar-

tin: Tarapoto, Martin & Plowman 1843 (F, GH). Prov.

Mariscal Caceres, 45 km NE of Tingo Maria, Tryon &

Tryon 5270 (GH). Ancash: Prov. Santa, alrededores de

Jimbe, Mostacero et al. 1096 (F). Huanuco: Prov. Huanu-

co 6 km S of Huanuco, Stork & Morton 9376 (F, GH, MO,

uc). Lima: Villa Lagoon (Prov. Lima, Chorrillos), Coro-

nado 6 (GH, uc, us). Pasco: Oxapampa, Soukup 2677

(us). Junin: 3 km N of La Merced, Tryon & Tryon 5442

(GH, uc). Ucayali: Prov. Coronel Portillo, Arboretum

von Humboldt, Diaz et al. 687 (F, MO, us). Huancavelica:

Prov. Tayacaja, Huari, Sounders 1148 (F, GH, us). Aya-

cucho: Ayna, Killip & Smith 22757 (F). Apurimac: Prov.

Abancay, Trancapata, Vargas 9617 (F, uc, us). Cuzco:

Aguas Calientes, Solomon 3180 (F, MO); 39 km E of

Urohuasi on Abancay-Cuzco road, Gentry et al. 23399

(MO, uc). Madre de Dios: Prov. Vlanu. Parque Nacional

del Manu, Foster 9796 (MO); 9856 (MO). Arequipa: Yura,

Solomon 2912 (F, MO, USM). Puno: Puno, Angulo 1760

(HUT).

2. Fquisetum bogotense HBK., Nov. gen. sp. pi.

1: 42. 1815. TYPE: "Prope Santa Fe de Bo-

gota et Alto del Roble, Columbia," Humboldt

& Bonpland (holotype, P?). Figure 5c-d.

Steins usually 20-50 cm tall, commonly bearing

an apical strobilus, 1-2 mm wide, solid in the

center. Endodermis single, surrounding the ring of

vascular bundles. Sheaths with short, brown, pa-

pery teeth. Branches irregular, with 4 ridges.

Strobilus stalked.

In soil or among rocks, in wet open places, in-

cluding banks of creeks, rivers, and irrigation

ditches in moist to wet seeps and clay road banks,

100-4200 m, most often above 2000 m, Piura

south to Puno and Tacna.

Costa Rica and Panama; Galapagos Islands; the

Andes of Venezuela and Colombia, south to Bo-

livia; to southern Argentina and Chile.

Piura: Prov. Huancabamba, Cerro Blanco, Stork 114 15

(GH, uc, us). Cajamarca: Prov. Contumaza, Guzmango

to Santiago, Sagdstegui & Lopez 10584 (F, MO). San Pa-

blo, Hutchison & Wright 5060 (F, MO, uc, us). Amazonas:

Hills WNW of Pomacocha, Wurdack 9445 (GH, uc, us,

USM). La Libertad: Above Yamobamba, Conrad 2728

(MO). San Martin: NW corner of Rio Abiseo National

Park, Young & Leon 4534 (HUT, USM). Ancash: Prov.

Recuay, Lopez et al. 7596 (MO, us). Huanuco: Prov.

Huanuco, Cerro Carpish, Luteyn & Lebron-Luteyn 5483

(F, uc, us). Lima: Prov. Huarochiri, Sullivan et al. 1103

(F, MO); Prov. Canta, Dist. Huaros, Pennell 14715 (F,

GH, us). Pasco: Prov. Oxapampa, Los Chacos, near Oxa-

pampa, Smith & Pretel 1514(f, MO, USM). Junin: Above

Conception, Correll & Smith P736 (GH). Huancavelica:

Prov. Huancavelica, Yauli, Stork & Morton 10863 (F,

uc). Ayacucho: Santiago, about 67 km from Nasca on

road to Puquio, Correll & Smith PI 51 (GH). Cuzco: Prov.

Urubamba, Pomatales, Nunez 7327 (F, MO). Arequipa:

Chacra Pachacutac, Balls B5868 (F, uc, us). Puno: San

Gabon (San Gaban) to Ollachea, Dillon et al. 1239 (MO).

Tacna: Prov. Tarata, Chuvire, Metcalf 30407 (MO, uc,

us).

Family 26. LYCOPODIACEAE

Contributed by Benjamin 011gaard

Lycopodiaceae Mirbel, in Lam. & Mirbel, Hist,

nat. veg. 4: 293. 1802. ("Lycopodia"). TYPE:

Lycopodium L.

Plants terrestrial or epiphytic, erect to pendu-

lous herbs or climbers. Stems dichotomously

branching, sometimes also with lateral branching,

protostelic, with the xylem arranged radially or in

parallel bands, or forming an incomplete cylinder

(Phylloglossuni). Leaves simple, with 1 simple vein,

arranged in low alternating spirals or irregular al-

ternating whorls, or seemingly decussate, homo-

phyllous or heterophyllous, isophyllous or aniso-

phyllous. Sporophylls like the foliage leaves, or

modified, in some groups specialized and aggre-

gated into distinct strobili. Sporangia solitary, ax-

illary or on the upper side of the sporophyll base,

homosporous, unilocular, reniform to subglobu-

lar, short-stalked, dehiscing by a transverse slit,

which divides each sporangium into 2 nearly equal

or strongly unequal valves. Spores without chlo-

rophyll, tetrahedral, with a trilete scar. Gameto-

phytes monoecious, tuberous, subterranean, and

mycorrhizal and without chlorophyll or surface-

living and green.

The family occurs in almost all humid regions

of the world, both warm and cold. It is here treated

as consisting of four genera including Phylloglos-

sum (Australia, New Zealand). Up to 14 genera

have been recognized by other authors (Holub,

1975, 1983, 1985, 1 99 1 ; Wagner & Beitel, 1992),

the genera of these authors generally correspond-

ing to the infrageneric taxa of 011gaard ( 1 987, 1 989,

1992); for a review of the classification, an index

to the family, and an overview of the Neotropical

species, see these publications. Most of the Pe-

ruvian species are illustrated in 011gaard (1988).

16

FIELDIANA: BOTANY

The total number of species in the family probably

exceeds 350, and approximately 185 of these are

tropical American. The Andes have a particularly

high diversity, and many species are conspicuous

elements in the montane and alpine vegetation of

these mountains.

This treatment includes 62 species for Peru, but

the real number is likely to be higher. Although

largely all available material has been consulted

for this treatment, it is evident that much collect-

ing needs to be done before a complete presen-

tation can be made. Many species are represented

by a single or few collections, and intensive efforts

in limited areas such as those of Blanca Leon and

Kenneth Young in the Rio Abiseo National Park,

of Abundio Sagastegui in the north of Peru, and

of the late David N. Smith in several areas have

turned up several species not formerly known from

Peru and widely disjunct from other known oc-

currences, as well as species new to science.

References

HOLUB, J. 1975. Diphasiastrum, a new genus in

Lycopodiaceae. Preslia 47: 97-1 10.

HOLUB, J. 1983. Validation of generic names in

Lycopodiaceae: With a description of a new ge-

nus Pseudolycopodiella. Folia Geobot. Phyto-

tax. 18: 439-442.

HOLUB, J. 1985. Transfers of Lycopodium spe-

cies to Huperzia: With a note on generic clas-

sification in Huperziaceae. Folia Geobot. Phy-

totax. 20: 67-80.

HOLUB, J. 1991. Some taxonomic changes with-

in Lycopodiales. Folia Geobot. Phytotax. 26:

81-94.

0LLGAARD, B. 1987. A revised classification of

the Lycopodiaceae sens. lat. Opera Bot. 92: 1 53-

178.

0LLGAARD, B. 1988. Lycopodiaceae, in G. Har-

ling and L. Andersson, Flora of Ecuador 33: 1-

155.

0LLGAARD, B. 1989. Index of the Lycopodi-

aceae. Biol. Skr. Dan. Vid. Selsk. 34: 1-135.

0LLGAARD, B. 1992. Neotropical Lycopodi-

aceae—An overview. Ann. Missouri Bot. Gard.

79:687-717.

ROTHMALER, W. 1 944. Pteridophytcn-Studien I,

Feddes Repert. Spec. Nov. Regni Veg. 54: 55-

82.

TRYON, A. F., AND B. LUGARDON. 1991. Spores

of the Pteridophyta. Springer- Verlag, New York.

TRYON, R. M., AND A. F. TRYON. 1982. Lyco-

podiaceae, in Ferns and allied plants, with spe-

cial reference to tropical America. Springer- Ver-

lag, New York.

WAGNER, W. H., AND J. M. BEITEL. 1992. Mod-

ern North American Lycopodiaceae and their

generic classification. Ann. Missouri Bot. Gard.

79: 676-686.

Key to Genera of Lycopodiaceae

a. Stem dichotomies equal (isotomous) throughout, plants without elongate, indeterminate main stems,

but sometimes with equally thick, somewhat heteroblastic branches; roots usually forming 1 basal

tuft; sporophylls and vegetative leaves alike, or the sporophylls, if smaller, persisting, not subpeltate,

not ephemeral; spores foveolate or fossulate I. Huperzia

a. Stem dichotomies unequal (anisotomous) throughout, the branches differentiated into elongate, in-

determinate, rhizomatous, or creeping, or trailing, main stems, and usually determinate aerial branch-

let systems; sporophylls strongly modified, ephemeral, unlike vegetative leaves, usually subpeltate,

aggregated in compact, terminal strobili; spores reticulate or rugate b

b. Strobili erect, sessile or pedunculate, borne on branchlet systems that arise in a dorsolateral position

on the main stem; side walls of sporangium epidermis cells sinuate, lignified throughout; spores

reticulate II. Lycopodium

b. Strobili pendulous and sessile, or strobili erect and terminating simple erect branches that arise

dorsally on the creeping stem, side walls of sporangium epidermis cells straight, not lignified,

except for nodular or semiannular thickenings; spores rugate III. Lycopodiella

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

17

FIG. 6. Huperzia eversa: a, habit; b, portion of fertile division with sporangia. Huperzia binervia: c, habit; d, portion

of fertile division with sporangia. Huperzia linifolia var. tenuifolia: e, habit; f, leaf from basal division; g, portion of fertile

division with sporangia. Huperzia cuneifolia: h, habit; i, expanded leaves from basal division; j, fertile shoot tip. (a from

Holm-Nielsen et al. 3873, Ecuador, AAU; b from Holmgren 674, Ecuador, s; c, d from Matthews, Peru, K; e, f, g from

Holm-Nielsen et al. 4422, Ecuador, AAU; h, i, j from Hutchison & Wright 6943, Peru, p.)

18

FIELDIANA: BOTANY

I. Huperzia

Huperzia Bernh., J. Dot. (Schrader) 1800 (2): 126.

1801. LECTOTYPE (designated by Roth-

maler, Feddes Repert. Spec. Nov. Regni Veg.

54: 59. 1944): Huperzia selago (L.) Martius

& Schrank (Lycopodium selago L.). Figure 6.

Plananthus Mirbel in Lamarck & Mirbel, Hist. nat.

veg. 3: 476. 1802. TYPE: Plananthus selago (L.)

Beauv. (Lycopodium selago L.).

Lycopodium subgen. Selago Baker, Handb. Fern-Al-

lies 8. 1887. TYPE: Lycopodium selago L.

Lycopodium subgen. Urostachya Pritzel, Nat. Pflan-

zenfam. 1 (4): 592. 1901. TYPE: Lycopodium se-

lago L.

Urostachys (Pritzel) Herter, Beih. Bot. Centralbl. 39:

249. 1922. TYPE: Urostachys selago (L.) Herter

(Lycopodium selago L.).

Phlegmariurus Holub, Preslia 36: 21. 1964. TYPE:

Phlegmarius phlegmaria (L.) Sen & Sen (Lyco-

podium phlegmaria L.).

Plants epiphytic or terrestrial, pendent, re-

curved, erect or ascending, isotomously branched

throughout (except in connection with bulbil for-

mation in the group of H. selago, and sprouting

from the base of old plants), the branches all sim-

ilar (homoblastic), or in some terrestrial species

differentiated (heteroblastic) into prostrate (some-

times subterranean) versus erect, aerial branches.

Roots arising from the stem stele, descending

through the cortex to the stem base, here emerging

as 1 basal tuft, or in heteroblastic species some-

times emerging directly along the underside of

prostrate shoots. Shoots homophyllous or hetero-

phyllous. Sporophylls and vegetative leaves alike,

not peltate, without mucilage cavities, persisting

and green after sporangium dehiscence, the grad-

ual or abrupt constriction of distal divisions of

heterophyllous species associated or not with pres-

ence of sporangia. Sporangia axillary, reniform,

isovalvate, with a short slender stalk; side walls of

sporangium epidermis cells sinuate, thickened and

lignified together with the inner walls. Spores fo-

veolate or fossulate. Gametophytes (not known for

any Peruvian species) usually subterranean (in ter-

restrial plants), mycorrhizal, cylindrical with ra-

dial or bilateral symmetry, with pluricellular, uni-

seriate trichomes among the gametangia.

The genus is virtually cosmopolitan, occurring

in tropical, temperate, arctic, and alpine environ-

ments. Species diversity is highest throughout the

tropics in evergreen montane forests, and in the

wet Andean grass and shrublands in South Amer-

ica.

There are approximately 300 species world-

wide, 49 in Peru. Seven species are endemic to

Peru, four are shared only with Bolivia and more

southerly regions, 26 occur also in Ecuador or far-

ther north in the Andes and the Neotropics, while

12 species occur both to the north and south of

Peru.

Species delimitation is problematic almost

throughout the genus due to the simple morphol-

ogy of the group and to plasticity of the characters

(011gaard, 1992). Morphogenesis seems to be

somewhat unstable in most species and may be

modified by external factors. Most characters are

plastic within a species (e.g., stem thickness, num-

ber of leaf orthostichies, leaf crowding, leaf direc-

tion, color, degree of heterophyllous differentia-

tion). Hybridization seems to occur rather

frequently, contributing to the blurring of species

limits, and the putative hybrids often have nor-

mally developed spores. As a consequence, species

recognition is often based on some experience and

comparison with identified material, rather than

a set of definite characters.

Measurements and terminology. The descrip-

tions and key characters are based on dried ma-

terial, unless otherwise indicated. Very often mea-

surements and solid shape of organs in life deviate

strongly from the dried condition.

Branching is isotomous throughout the genus,

except in connection with sprouting (lateral

branching) from the base of old or injured indi-

viduals. Isotomy results in the formation of equal-

ly thick branches each with an equal amount of

vascular tissue. However, many species of high

Andean grasslands develop heteroblastic branch

pairs; i.e., the isotomous branches differentiate into

distinct aspects and functions due to different leaf

development, as exemplified by the sporangiate,

erect, aerial shoots and the sterile, prostrate or

even subterranean, basal, rejuvenating shoots of

Huperzia crassa and H. hypogaea. Homoblastic

branches have the same aspect and function and

similar leaf development.

Phyllotaxis is poorly understood in the Lyco-

podiaceae. It seems to be irregularly organized

morphogenetically and quite variable within one

species, sometimes even within the same individ-

ual. Stevenson (Bot. J. Linn. Soc. 72: 8 1-1 00. 1 976)

interpreted the phyllotaxis of Huperzia lucidula

(Michx.) Trevisan (temperate North America) as

consisting of low alternating spirals, and found a

definite relation between the number of orthosti-

chies and the number of protoxylem lobes in the

stele. In many species this relation is different or

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

19

possibly absent, and transitions from spiral to ir-

regularly whorled phyllotaxis is common, al-

though whorled arrangement is the more com-

mon. The number of longitudinal leaf ranks is

readily observed in some cases but often must be

estimated, or calculated, from the number of leaves

in each leaf whorl, or from the number of leaves

contained in one rotation of the leaf spiral. As the

leaves of a whorl or a rotation of the spiral alter-

nate with the adjacent whorls or spirals above and

below, the number of leaves in two whorls or ro-

tations makes up the number of longitudinal leaf

ranks of a shoot. For the sake of brevity, I have

described leaf arrangement in terms of leaf whorls,

although this is not everywhere completely accu-

rate.

Reference

ROLLERI, C. 1981. Sinopsis de las especies de

Lycopodium L. (Lycopodiaceae-Pteridophyta)

de la seccion Crassistachys Herter. Revista Mus.

La Plata (n.s.) Bot. 13: 61-113.

Key to Species of Huperzia

a. Plants erect, or ascending to erect, terrestrial or epiphytic and shoot apices erect; leaf margins smooth,

fimbriate, denticulate or erose, or leaves papillate b

b. Leaf margins denticulate (at least of some leaves, sometimes minutely and remotely denticulate)

by pointed teeth, ciliolate, fimbriate, or leaves papillate c

c. Leaves densely papillate abaxially, at least at the apex 12. H. weberbaueri

c. Leaves not papillate d

d. Leaves of upper, sporangiate divisions loosely to closely appressed, abaxially strongly convex

throughout; leaf margins fimbriate by irregularly shaped and directed pale and soft teeth;

plants bright red or tinged with red e

e. Erect shoots predominantly quadrangular 25. H. tetragona

e. Erect shoots hexagonal or terete f

f. Leaves of upper, sporangiate branches triangular-lanceolate to triangular-ovate, with

smooth abaxial epidermis, arranged in alternating whorls of 3-4 24. H. attenuata

f. Leaves of upper, sporangiate branches strongly uneven abaxially by protruding, blister-

like epidermal cells, arranged in alternating whorls of 5 23. H. sagasteguiana

d. Leaves of upper, sporangiate divisions patent to recurved (sometimes appressed and green

in H. affinis), abaxially flat or convex; leaf margins ciliate or denticulate; plants green . . . g

g. Leaves borne in alternating whorls of 4-5(-6), denticulate or long-ciliate, narrowly tri-

angular-lanceolate, linear-lanceolate or nearly subulate, (2.5-)3-6 mm long, 0.5-1.5 mm

wide h

h. Sporangiate leaves usually with long-ciliate margins, 1-1.5 mm wide at the somewhat

clasping base, 4-6 mm long, spreading to appressed 10. H. affinis

h. Sporangiate leaves denticulate, to 1 mm wide at the base, 2.5-5 mm long, not clasping,

spreading to strongly reflexed 9. H. eversa

g. Leaves borne in alternating whorls of (6-)7-l 1, denticulate, subulate to linear-subulate,

4-10 mm long i

i. Leaves borne in alternating whorls of (6-)7-8, ascending to spreading or sharply

reflexed, straight to strongly recurved, usually evenly tapering, with very sparsely to

densely and sharply denticulate to short-ciliolate margins throughout 7. H. reflexa

i. Leaves borne in alternating whorls of 8-11, usually sharply bent upward from a

perpendicular junction to the stem and then gently to strongly claw-like recurved,

narrowed shortly above the base, with rather densely denticulate-ciliolate margins at

base, usually sparsely denticulate or smooth at apex 8. H. acifolia

b. Leaf margins entirely smooth, or uneven, rugose to erose-rugose, without pointed teeth or cilia,

not fimbriate or papillate, or papillate only on leaf base margins j

j. Epiphytes with spreading leaves; leaves filiform, up to 0.5 mm wide, (6-) 10-1 7 mm long, leaf

bases often red . . . 36. H. wilsonii

20

FIELDIANA: BOTANY

j. Terrestrial plants with spreading, reflexed or appressed leaves; leaves linear-filiform to nearly

orbicular, not red k

k. Leaves wide-spreading to reflexed or patent-ascending, the widest ones linear-filiform to

lanceolate, without a basal swelling (air sac) 1

1. Broadest leaves of sporangiate divisions linear-filiform to subulate, 1.5 mm wide or less

m

m. Leaves softly herbaceous, usually straight or slightly recurved, 0.5-1.3 mm wide,

usually flat abaxially n

n. Leaves linear to linear-subulate, 0.8-1.3 mm wide at base, slightly convex to

canaliculate adaxially 1 . H. hippuridea

n. Leaves linear-filiform, 0.5-0.8 mm wide at the base, canaliculate abaxially ....

2. H. lechleri

m. Leaves coriaceous, usually sigmoid, 1.2-1.5 mm wide, usually bisulcate abaxially .

4. H. binervia

1. Leaves lanceolate, the narrowest ones 1.5 mm wide or more o

o. Leaves reflexed to patent-ascending, thickly coriaceous, adaxially evenly convex, or

with slightly prominent vein, margins flat or involute, smooth 5. H. weddellii

o. Leaves papery to thinly subcoriaceous, folded slightly down along the vein, margins

revolute, minutely rugose 6. H. brongniartii

k. Leaves, at least of sporangiate divisions, appressed, the narrowest ones linear-lanceolate to

broadly triangular-cordate and appressed, with or without a basal swelling, or patent to

reflexed and oblong, ovate or wider p

p. Shoots strongly heteroblastic, with deeply subterranean, elongate, horizontal shoots bear-

ing stiffly erect aerial branches 22. H. hypogaea

p. Shoots homoblastic to heteroblastic, above the ground or shallowly subterranean, not

deeply subterranean q

q. Sporangiate divisions with shortest leaves 6 mm or longer, linear to widely lanceolate

or widely triangular-ovate r

r. Aerial shoots somewhat club-shaped, compactly caespitose, narrowed and some-

what etiolated at the base and here with pale and irregularly appressed leaves;

shoots strongly heteroblastic, the basal, prostrate shoots short and with much

reduced leaves 21. H. saururus

r. Aerial shoots usually equally thick throughout or tapering, loosely to densely caes-

pitose, with all leaves nearly alike; shoots homoblastic or heteroblastic; prostrate

shoots (if any) with leaves the same size or larger than those of aerial shoots . . s

s. Shoots heteroblastic, with prostrate basal, rejuvenating, often rooting shoots

bearing erect, fingerlike aerial shoots t

t. Aerial shoots often short, unbranched, appearing narrower than creeping

shoots; leaves of creeping shoots larger than in aerial shoots

20. H. andina

t. Aerial shoots well developed, simple or branched, not appearing narrower

than creeping shoots; leaves of creeping and aerial shoots of nearly equal size

u

u. Leaves dull, pruinose, usually red-tinged, abaxially rugose from protrud-

ing, blisterlike epidermis cells (rarely smooth) v

v. Leaves 5-9 x 1-2 mm, with a prominent basal air cavity causing a

perpendicular appearance of the junction to the stem in sporangiate

leaves 18. H. crassa

v. Leaves 9-12 x 2-2.5 mm, without a prominent basal air cavity, ap-

pressed from the base 19. H. nesselii

u. Leaves dull to somewhat glossy, green, abaxially smooth w

w. Leaves of aerial shoots lanceolate to widely lanceolate, slightly long-

acuminate, (1.8-)2-2.5(-3) mm wide, strongly curved upward at apex

. . 17. H. darwiniana

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI. 21

w. Leaves of aerial shoots lanceolate, evenly tapering, 1.5-2 mm wide,

straight to slightly curved upward at apex 16. H. macbridei

s. Shoots homoblastic, all essentially alike x

x. Leaves of basal divisions linear to linear-subulate, up to 1 .5 mm wide, usually

bisulcate abaxially, usually somewhat sigmoid 4. H. binervia

x. Leaves of basal divisions linear-lanceolate or wider, 1.5—4 mm wide, not

bisulcate abaxially, sigmoid or not y

y. Plants rather fragile and slender; stems 2-3.5 mm thick excluding leaves;

leaves 5-10 x 1.5-2 mm, distal divisions ca. 5-10 mm in diameter in-

cluding leaves 15. H. capellae

y. Plants large, robust; stems 3-8 mm thick excluding leaves; leaves 5-12

x 2-4 mm, distal divisions ca. (7-) 1 0-25 mm in diameter including leaves

z

z. Leaves of aerial shoots evenly tapering to short-acute, straight or curved

apex aa

aa. Leaves triangular-lanceolate to widely triangular-ovate, straight to

slightly curved upward; leaf base margins in sporangiate leaves

usually slightly revolute, sometimes subauriculate

11. H. kuesteri

aa. Leaves lanceolate to widely lanceolate, usually strongly curved

upward (rarely patent-ascending and somewhat sigmoid); leaf bas-

es not revolute or auriculate 14. H. polylepidetorum

z. Leaves of aerial shoots slightly long-acuminate, strongly curved upward

at apex 1 7. H. darwiniana

q. Sporangiate divisions with longest leaves up to 6 mm long, lanceolate or wider .... bb

bb. Leaves (longest ones) up to 4 mm long, ovate, elliptic, or cordate cc

cc. Broadest leaves ovate or elliptic, (1.8-)2-2.6(-3.5) mm long, (1. 2-) 1.5-2(2. 4)

mm wide, sporangia 1-1.5 mm wide 26. H. sellifolia

cc. Broadest leaves widely ovate to widely suborbicular-cordate or triangular-

cordate, 2-4 mm long, 2— 4(-5) mm wide, sporangia 1.5-2.5 mm wide ....

28. H. brevifolia

bb. Leaves, even shorter ones, 4 mm or longer, or if shorter, then lanceolate . . . dd

dd. Stem, excluding leaves, 1-1.5 mm thick, sporangia 2-2.5 mm wide, subgl-

obose 27. H. engleri

dd. Stem, excluding leaves, usually more than 2-4 mm thick; sporangia 1.5-2

mm wide, flattened ee

ee. Leaves triangular-ovate to widely ovate, usually red-tinged, leaf bases

with a strongly prominent air cavity 29. H. hohenackeri

ee. Leaves lanceolate, green, leaf bases without or with a slightly prominent

air cavity 13. H. colanensis

a. Plants pendulous, or initially erect with nodding to pendulous shoot apices, epiphytic or terrestrial; leaf

margins entire ff

ff. Leaves more or less sharply dimorphic: basal divisions with long, expanded leaves, apical divisions

constricted, with appressed, reduced, decussate or subdecussate leaves; or the entire plant covered by

appressed, short, broad, decussate leaves gg

gg. Plants robust, stem excluding leaves at least 2 mm thick at the base, without red color-

ation hh

hh. Constricted divisions sharply quadrangular, 3-8 mm thick including leaves; sporangiate leaves

3-8 mm long, sharply carinate, often with a conduplicate apex 41. H. molongensis

hh. Constricted divisions bluntly quadrangular to subterete, 2-4 mm thick; sporangiate leaves 3

mm long or less, rounded to bluntly carinate throughout abaxially, not conduplicate apically

ii

ii. Plants entirely pendulous, expanded leaves of proximal divisions in usually densely crowd-

ed, alternating whorls of 3, the whorls 2.5-5 mm apart; sporangiate leaves 2-3 mm long

22 FIELDIANA: BOTANY

42. H. campiana

ii. Plants erect to scandent, with recurved shoot tips and constricted divisions, expanded

leaves of proximal divisions in distant, alternating whorls of 4, the whorls 6-9 mm apart;

sporangiate leaves 1.2-1.6 mm long 9. H. pruinosa

gg. Plants slender, stem excluding leaves usually 1 mm thick or less (rarely to 1 .7 mm) at the base;

stem base with or without red coloration jj

jj. Expanded leaves of basal divisions very uniform in size, shape, and position throughout, closely

situated, with almost continuously overlapping leaf margins (pressed specimens), borne in

alternating whorls of 3 at the base, broadly lanceolate to ovate, acute; stems not red

44. H. ericifolia

jj . Expanded leaves uniform or variable in shape and position, close to distant and not continuously

overlapping, decussate, subdecussate or in alternating whorls of 3 at the base, linear-subulate

to ovate or oblanceolate; stems with or without red coloration kk

kk. Expanded leaves decussate or subdecussate, lanceolate to ovate, the broadest ones 2-3.5

mm wide 11

11. Constricted terminal divisions usually not sharply distinct from expanded divisions,

with irregularly sized and directed leaves; expanded divisions usually well developed

and more extensive than constricted divisions 45. H. myrsinites

11. Constricted terminal divisions sharply distinct from expanded divisions, with reg-

ularly sized and uniformly directed leaves throughout; expanded leaves restricted

to a short zone at the base (rarely lacking), and sometimes inserted in short zones

of constricted divisions 43. H. heteroclita

kk. Expanded leaves decussate or in whorls of 3, linear-subulate to lanceolate or oblan-

ceolate, the broadest ones 2 mm wide or less, or if wider then less than 6.5 mm long

and oblanceolate mm

mm. Expanded leaves decussate or in whorls of 3, linear-subulate to lanceolate, the

broadest ones 1-2 mm broad, the longest ones 8-15 mm long nn

nn. Expanded leaves linear, or linear-subulate to linear-lanceolate, 0.5-1 mm

broad 47. H. subulata

nn. Expanded leaves linear-lanceolate to lanceolate, 1.3-2 mm

broad 46. H. phylicifolia

mm. Expanded leaves decussate, lanceolate to oblanceolate, the broadest ones 1.5-3

mm broad, the longest ones less than 6.5 mm long 48. H. cuneifolia

ff. Leaves uniform throughout, long and expanded, or gradually smaller and more appressed, but

not predominantly decussate or subdecussate toward the shoot apices oo

oo. Plants very delicate, the longest leaves less than 6(-8) mm long, acicular, less than 0.5 mm

broad; stems less than 1 mm thick at base excluding leaves pp

pp. Non-sporangiate leaves of narrow terminal divisions closely appressed to the stem, with

cuneate to rounded base; leaves of basal divisions usually strongly curved upward and

inward from a patent base, or sigmoid 39. H. curvifolia

pp. Non-sporangiate leaves of terminal divisions with a patent, rounded to auriculate or

subhastate leaf base; leaves of basal divisions spreading to ascending, often unilaterally

curved 40. H. tenuis

oo. Plants slender to robust, longest leaves of basal divisions more than 8 mm long; leaves filiform

or linear to lanceolate; stems 0.5-5 mm thick at base, excluding leaves qq

qq. Leaves of basal divisions alternate or paired, or in occasional whorls of 3, more than 13

mm long, 1—4 mm broad, with a narrowed, twisted, often petiolelike and approximately

perpendicular lamina base, leaves of terminal divisions alternate to whorled; stem base

usually 1 mm or less thick excluding leaves 35. H. linifolia

qq. Leaves of basal divisions predominantly or entirely whorled, without petiolelike lamina

bases, 8-23 mm long; stem base 0.5-5 mm or more thick rr

rr. Leaves closely appressed throughout, linear-subulate to linear-lanceolate, apically con-

vex to conduplicate abaxially ss

ss. Leaves borne in whorls of 7-8, linear-subulate, falcate- appressed

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

23

33. H. funiformis

ss. Leaves borne in whorls of 4-5, linear to linear-lanceolate, straight to slightly

recurved at apex 34. H. buesii

IT. Leaves, at least in basal divisions, ascending to patent, filiform to lanceolate, flat,

concave, or convex tt

tt. Leaves filiform to linear, or linear-subulate from an auriculate leaf base, up to 1

mm broad; the leaves, if linear and 1 mm broad, borne in whorls of 5 or more

uu

uu. Leaf bases of non-sporangiate leaves auriculate and often somewhat overlap-

ping neighboring leaf bases of the same whorl; lamina usually less than 0.7

mm broad above the auriculate base 38. H. sarmentosa

uu. Leaf bases of non-sporangiate leaves without auricles; lamina 0.2-1 mm broad

vv

w. Leaves filiform, 0.3-0.5 mm broad just above the base ww

ww. Leaves not twisted at base, perpendicularly spreading to slightly

upward curved, or ascending, (6-) 10- 17 mm long, with green or

bright red bases; stems usually 1.5-2 mm thick . . . 36. H. wilsonii

ww. Leaves twisted at base, obliquely falcate-ascending, 6- 1 0(- 1 2) mm long,

with green leaf bases; stems usually 1-1.5 mm thick

37. H. polycarpos

vv. Leaves linear, 0.7-1 mm broad, (10-)14-19 mm long ... 3. H. arcuata

tt. Leaves linear- lanceolate to lanceolate, the broadest ones more than 1.5 mm broad;

or the non-sporangiate leaves of basal divisions linear, 1 mm broad or more, and

borne in whorls of 3-4 xx

xx. Leaf margins minutely uneven by individually protruding epidermis cells,

especially near the apex; leaves lanceolate, 2—4 mm broad

32. H. rosenstockiana

xx. Leaf margins smooth; leaves linear to lanceolate, 1-3.5 mm broad yy

yy. Stem base 2.5-5 mm thick excluding leaves; leaves, at least of basal di-

visions, brightly shining, firmly coriaceous throughout, not twisted at base,

2.5-3.5 mm broad; sporangia 1.7-3 mm broad .... 30. H. hartwegiana

yy. Stem base (1-) 1.5-2 mm thick; leaves dull or slightly shining, firmly her-

baceous to subcoriaceous, twisted or straight at base, 2-2.5 mm broad;

sporangia 1-2.2 mm broad 31. H. taxifolia

1. Huperzia hippuridea (Christ) Holub, Folia

Geobot. Phytotax. 20: 73. 1985.

Lycopodium hippurideum Christ in Pittier, Primit. Fl.

Costar. 3 (1): 56. 1901. TYPE: Costa Rica, El

Paramo, 3000 m, massif de Buena Vista, 1897,

Pittier 10619 (holotype, P!; isotype, us!).

Urostachys hippurideus (Christ) Nessel, Barlappge-

wachse 88. 1939.

Urostachys poseidonis Herter, Revista Sudamer. Bot.

10: 122-123. 195 3. TYPE: Ecuador, Prov.Chim-

borazo, Penipe, 3400 m, Rose in Mi lie 35 (ho-

lotype, us!; isotypes, GH!, NY!).

Lycopodium poseidonis (Herter) Morton, Amer. Fern

J. 54: 42. 1964.

Shoots homophyllous, equally thick throughout,

10—35 mm in diameter including leaves. Stems

excluding leaves 2.5-4 mm thick at the base, ta-

pering to ca. 2-3 mm upward. Leaves borne in

irregular alternating whorls of 5-8, spreading to

reflexed, sometimes sharply reflexed and ap-

pressed to the stem, linear to linear-subulate, even-

ly tapering from the base or the middle, (10-)1 1-

19 mm long, 0.8-1.3 mm wide, not, or rarely,

twisted at base, adaxially with a slightly prominent

vein, with smooth, usually slightly revolute mar-

gins, and indistinctly to prominently and widely

decurrent bases. Sporangia 1.5-2 mm wide.

Plants ascending to stiffly erect from a decum-

bent base, up to 60 cm tall, sparsely branched.

Upper montane forest, especially near the forest

limit, usually on the forest floor, in semishade, alt.

24

FIELDIANA: BOTANY

2500-3500 m, Cajamarca, San Martin, Pasco,

Cuzco.

Central America; Andes from Venezuela to Bo-

livia.

Huperzia hippuridea belongs to a group of close-

ly related taxa of high montane forests throughout

tropical America, including also H. arcuata B. 011g.

(Colombia, Ecuador), H. lechleri, H. montana

(Underw. & Lloyd) Holub (Greater Antilles), H.

nuda (Nessel) B. 011g. & Windisch (Brazil).

Hutchison & Bismarck 6455 and Matthews 963

(presumably from Chachapoyas) have thicker

stems and more crowded and coriaceous leaves

than usual for the species, thus approaching Hu-

perzia \veddellii (Herter) Holub and H. loxensis

B. 011g. Matthews 963 is a mixed collection, sheets

of the same number at BM pertaining to H. mac-

bridei.

Cajamarca: Prov. Hualgayoc, Hda. Taulis, 4.6 km be-

yond Palmito junction on the road to La Playa, 2740 m,

Hutchison & Bismarck 6455 (uc, USM). San Martin: Prov.

Mariscal Caceres, NW corner of Rio Abiseo National

Park, Chochos, 3500 m, Young & Leon 4660 (AAU).

Prov. Mariscal Caceres, Puerta del Monte, 3100-3300

m, Young & Leon 4436 (AAU), 3450 m, Leon & Young

1305 (USM). Pasco: Prov. Oxapampa, Santa Barbara,

3200-3300 m, D. Smith 8176 (AAU, USM). Cuzco: Prov.

Paucartambo, Cerro Macho Cruz, Parque Nacional

Manu, 3400 m, Leon 2303 (USM). Department unknown:

(possibly from Chachapoyas) Matthews 963 (G).

2. Huperzia lechleri (Hieron.) Holub, Folia Geo-

bot. Phytotax. 20: 74. 1985.

Lycopodium lechleri Hieron., Bot. Jahrb. Syst. 34: 57 1 .

1905. LECTOTYPE (designated by Rolleri, Re-

vista Mus. La Plata (n.s.) Bot. 13 (78): 82. 1981):

Peru (Dept. Puno), Tabina, Lechler, ed. Hohe-

nacker 20/2 (B!; isotypes, G!, K!, P!, UPS!).

Vrostachys lechleri (Hieron.) Nessel, Barlappgewachse

85. 1939.

Vrostachys lechleri Hieron. var. lehmannii Nessel, Re-

vista Sudamer. Bot. 6: 61. 1940. SYNTYPES:

BONN, Herb. Nessel 148\, Ecuador, Spruce, Palla-

tanga 1858, marked 7; Ecuador: unknown collec-

tor "Aus dem Herbar Bonaparte, Paris," anno-

tated "Ur. lehmannii Boge.").

Urostachys lehmannii (Nessel) Herter, Index Lyco-

podiorum 67. 1949.

In most features resembling Huperzia hippuri-

dea closely, but differing consistently in the nar-

rower, linear to nearly filiform leaves 10-23 mm

long and 0.5-0.8 mm wide at the base, adaxially

usually canaliculate, with a slightly prominent vein

at the base, and the margins not revolute, the spo-

rangia 1-1.8 mm wide.

Upper montane forest, especially near the forest

limit, usually on the forest floor, in semishade, alt.

1 800-3600 m, Junin, Huancavelica, Cuzco, Puno.

Peru and Bolivia.

Closely related to Huperzia hippuridea (see the

preceding species for discussion).

Junin: Prov. Tarma, Carpapata, Soukup 3477 (F, GH).

Huacapistana, 1800-2400 m, Killip & Smith 24503 (F,

GH, us). Huancavelica: Prov. Tayacaja, Marcavalle, be-

tween Huachocolpa and Tintay, 2600 m, Tovar 4085

(USM), Tovar 4089 (USM). Cuzco: Valle Occobamba [Oco-

bamba], 1900 m, Bites 856 (us). Prov. Paucartambo,

Valle de Pilcopata, near Pillahuata, 2500 m, Foster &

Wachter 7501 (AAU). Pillahuata, 2850 m, Fitzpatrick &

Willard (F). San Ignacio, Huadquina, Biies 1410 (us).

Puno: Ollachea to San Gaban, 1000-2000 m, Dillon et

al. 1156 (AAU, F, MO). Prov. Carabaya, road San Gaban

(Lanlacuni Bajo) to Macusani, near Ollachea, 2600 m,

Maas et al. 6111 (AAU, USM). Department unknown:

Lechler 2023a (K). Vargas 16767 (GH).

3. Huperzia arcuata B. 011g. in Harling and An-

dersson, Fl. Ecuador 33: 15, t. ID. 1988.

TYPE: Ecuador, Prov. Carchi, Road El Angel

to Tulcan, 3500 m, Holm-Nielsen et al. 5341

(holotype, AAU!; isotypes, GB!, QCA!, us!).

Plants ascending to erect from a decumbent base,

with nodding shoot apices, up to 30 cm tall (ter-

restrial), or recurved to pendulous and up to 90

cm long (epiphytes), sparsely branched. Shoots

homophyllous, equally thick throughout, 25-30

mm in diameter, or tapering to ca. 12 mm in-

cluding leaves. Stem excluding leaves 2-3 mm thick

at base, tapering to ca. 1.5 mm upward. Leaves

borne in irregular alternating whorls of 6—7 near

the base, upward of 5-6, spreading to falcately

ascending, linear to subulate from a slightly wid-

ened lamina base, ( 1 0-) 14-19 mm long, (0.5-)0.7-

1 mm wide above the widened base, gradually

tapering in the distal half or so, adaxially flat to

slightly concave, or with slightly revolute margins,

with abaxially slightly prominent vein and smooth

margins, with slightly widened lamina base, twist-

ing the lamina to a vertical position. Vegetative

leaves of basal divisions often spreading to re Hexed

and not twisted. Sporangia 1 .3-2 mm in diameter.

Upper montane forest, near timberline, on the

forest floor, in semishade, or epiphytic, alt. 2700-

3500 m, Amazonas, San Martin.

Southern Colombia to Peru.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

25

Closely related to Huperzia hippuridea, from

which it differs by its obliquely falcate-ascending,

twisted leaves and the nodding to pendulous shoot

tips. Huperzia arcuata also resembles H. dicho-

toma (Jacq.) Trevisan (tropical America) but is

larger in all parts and restricted to timberline forest

habitats. The Peruvian collections generally have

short leaves, 8-15 mm long.

Cajamarca: Prov. Cutervo, Llama to Huambos, 2700

m, Lopez et al. 6591 (AAU, OH, MO). Cutervo, Raimondi

3126 (B, USM). Trail Socota to Tambillo, 3300 m, Stork

& Morton 10174 (F, K, uc). Amazonas: Prov. Chacha-

poyas, Cerros Calla Calla, W side, 45 km above Balsas,

3100 m, Hutchison & Wright 5784B (uc). Cerros Calla

Calla, E side, 1 9 km above Leimebamba, 3 1 00 m, Hutch-

ison & Wright 5559 (F, GH, M, MO, NY, p, uc). Cano Santa

Lucia just E of Chachapoyas, 2000-2400 m, Wurdack

600 (F, GH, K, uc).

Amazonas: Chachapoyas, Matthews (E). San Martin:

Prov. Mariscal Caceres, Rio Abiseo National Park, Puer-

ta del Monte, 3450 m, Young 1627 (AAU, USM). Prov.

Mariscal Caceres, near La Playa Camp, 2700 m, Young

& Leon 4954 (AAU).

4. Huperzia binervia (Herter) B. 011g., Opera Bot.

92: 169. 1987.

Lycopodium binervium Herter, Bot. Jahrb. Syst. 43,

Beibl. 98: 48. 1909. TYPE: Peru, Prov. Chacha-

poyas, Matthews (holotype, P!; isotypes, BM!, GL!,

K!).

Urostachys binervius (Herter) Nessel, Barlappge-

wachse 110. 1939.

Plants ascending to erect from a decumbent base,

up to 50 cm tall. Shoots homophyllous, equally

thick throughout, 1 5-30 mm in diameter includ-

ing leaves or with gradually shorter leaves and

tapering to 5-8 mm. Stems excluding leaves 3-6

mm thick at the base, sometimes tapering to 2-

2.5 mm, ridged by decurrent leaf bases and mar-

gins. Leaves borne in very close alternating whorls

of 6-9, spreading or somewhat reflexed to ascend-

ing or appressed upward, slightly recurved or sig-

moid, linear to linear-subulate, 7-16 mm long,

1.2-1.5 mm wide, upwards sometimes reduced to

3-7 mm long, not twisted at base, thick, coria-

ceous, adaxially convex and shallowly rounded or

with a slightly to sharply prominent veinal ridge,

abaxially convex and shallowly rounded, when

dried usually with a prominent vein and irregular

longitudinal ridges or wrinkles, or sometimes the

vein sunken and hence the adjacent leaf tissue ap-

pearing as 2 longitudinal veins, with smooth mar-

gins, green. Stomates often forming 2 irregular,

slightly sunken longitudinal bands along the vein.

Sporangia 1.5-2 mm wide.

Open shrubland and grassland (jalca), seepage

areas, 2000-3100 m, Cajamarca, Amazonas.

Endemic.

Related to Huperzia loxensis B. 011g. (southern

Ecuador) and H. hippuridea.

5. Huperzia weddellii (Herter) Holub, Folia Geo-

bot. Phytotax. 20: 78. 1985.

Lycopodium weddellii Herter, Bot. Jahrb. 43: Beibl.

98: 45. 1909. TYPE: Peru, Dept. Puno, Carabaya,

1847, Weddell 4684 (holotype, P!; isotype frag.,

BONN, Herb. Nessel 1701 in part).

Urostachys weddellii (Herter) Nessel, Barlappge-

wachse91. 1939.

Plants erect, or erect from a decumbent base,

very robust, up to 30 cm tall, or to 50(-70) cm

long, sparsely branched. Shoots homophyllous, al-

most equally thick throughout, (14-) 16-2 3 mm in

diameter including leaves. Stems excluding leaves

3-6 mm thick at the base, tapering to 3—4 mm in

diameter. Leaves almost uniform throughout,

borne in alternating whorls of 6-7, ascending to

spreading or reflexed, straight, curved upward or

recurved, lanceolate, acute to slightly acuminate

or short-acute, 6-ll(-12) mm long, 1.5-2.5 mm

wide, thick and coriaceous, often glaucous, rarely

with red tips, adaxially convex with a slightly

prominent vein, usually irregularly concave abax-

ially (dried), with vein abaxially obscure to dis-

tinctly and widely prominent, sometimes in as-

cending leaves abaxially convex and rounded with

obscure vein, with usually smooth to minutely un-

evenly rugose, narrowly sclerified, translucent

margins, or rarely papillate near the base. Leaf

bases prominently decurrent, with a small, some-

times indistinct swelling (air sac), especially in

sporangiate leaves. Sporangia 1.5-2.2 mm in di-

ameter.

Terrestrial in exposed habitats in timberline for-

est and lower shrub paramo, 2600-3900 m, Ama-

zonas, Puno.

Ecuador to Bolivia.

Amazonas: Prov. Bagua, Cordillera Colan, NE of La

Peca, ca. 3 1 70 m, Barbour 3446 (MO).

6. Huperzia brongniartii (Spring) Trev., Atti Soc.

Ital. Sci. Nat. 17:248. 1874.

26

FIELDIANA: BOTANY

Lycopodium brongniartii Spring, Bull. Acad. roy. Sci.

Bruxelles 8 (2): 515. 1841. TYPE: Bolivia, Yun-

gas, D'Orbigny 227 (holotype, P!; isotypes, BONN,

Herb. Nessel 169\ in part, BR!).

Lycopodium taxifolium Sw. var. brongniartii (Spring)

Baker, Handb. Fern Allies 16. 1887.

Urostachys brongniartii (Spring) Nessel, Arch. Bot.

Est. S. Paulo 1: 388. 1927.

Plants erect from an ascending base, up to 50

cm tall, sparsely branched. Shoots homophyllous,

almost equally thick throughout, 20-30 mm in

diameter including leaves. Stems excluding leaves

2.5—4 mm thick near the base, sometimes tapering

to 1.5-2 mm upward, somewhat ridged by decur-

rent leaf bases. Leaves uniform throughout, borne

in alternating, rather distant whorls of 4-5, wide-

spreading to somewhat reflexed, usually straight,

not twisted at the base, lanceolate, with long-acute

apex, papery to subcoriaceous and opaque, 9-1 2(-

15) mm long, 2-3 mm wide, almost flat, with

prominent vein adaxially, or folded slightly down

along the vein, with slightly revolute, minutely

rugose margins. Sporangia ca. 2.5 mm wide.

Terrestrial in humid montane forest, ca. 2900-

3300 m, Cuzco.

Colombia to Bolivia.

Related to Huperzia rosenstockiana (Herter)

Holub with which it shares the minutely rugose

leaf margins. The specimens cited correspond to

the Bolivian representatives of the species.

Cuzco: Prov. Urubamba, Machu Picchu, hillside above

Rio Mandor, 2920 m, Peyton 1350 (AAU). Alturas de

Rio Calzada, Huadquina, 3300 m, Sues 756 (us). Prov.

Paucartambo, Parque Nacional Manu, Road Acjanaco

to Pilcopata, bridge between Pillahuata and La Esperan-

za, 2750 m, Leon 2228 (USM); Prov. Paucartambo, Par-

que Nacional Manu, below Acjanaco, 2950 m, Leon &

Cano 2129 (USM).

7. Huperzia reflexa (Lam.) Trev., Atti Soc. Ital.

Sci. Nat. 17: 248. 1874.

Lycopodium reflexum Lam., Encycl. 3: 653. 1789.

TYPE: Martinique, Comm. Joseph Martin (ho-

lotype, P, Herb. Lam. 4421).

Plananthus reflexus (Lam.) Beauv., Prodr. Aeth. 100.

1805.

Urostachys reflexus (Lam.) Herter, Beih. Bot. Cen-

tralbl. 39: 249. 1922.

Plants erect or ascending from a decumbent base,

soft, usually loosely caespitose, 10-30(-40) cm tall.

Shoots homophyllous, almost equally thick

throughout, 7-1 5 mm in diameter including leaves.

Stems excluding leaves 1 .5-3(-4) mm thick at base,

sometimes tapering to 1-1.5 mm in diameter,

ridged by decurrent leaves or almost smooth.

Leaves borne in alternating irregular whorls of

(6-)7-8(-9), ascending to spreading or sharply re-

flexed, straight to strongly recurved, linear-subu-

late, widest just above the base, 4-8 mm long, 0.5-

1(-1.2) mm wide, softly herbaceous to subcoria-

ceous, adaxially convex, or concave near the base,

abaxially flat, or slightly concave to convex, with

an obscure to somewhat prominent vein, with flat

to revolute, very sparsely to densely denticulate

to short-ciliolate margins. Leaf bases often some-

what decurrent. Sporangia 1-1.5 mm in diameter.

A frequent pioneer on permanently moist, dis-

turbed ground, mainly in the montane forest zone,

also in peat bogs, or rarely as a low epiphyte.

Tropical America.

Closely related to Huperzia eversa, H. acifolia,

and H. affinis.

Huperzia reflexa is variable with respect to leaf

and stem size, direction and crowding of leaves,

and leaf margin characters. Part of the variation

undoubtedly reflects variable growth conditions,

but very often several distinct forms can be found

growing intermixed in the same habitat, indicating

that genetic differences exist. The variation pat-

terns are complex and in need of a detailed study.

A variety with smaller dimensions stands some-

what apart and is recognized taxonomically. The

remaining material, referred to the type variety, is

polymorphic.

Key to Varieties

a. Shoots including leaves 9-15(-20) mm in diameter, leaves (5-)6-8 mm long, 0.7-1 (-1.2) mm wide,

with regularly denticulate to short-ciliolate margins, sporangia ca. 1.5 mm wide . . 7a. var. reflexa

a. Shoots including leaves 7-10 mm in diameter, leaves 4-5(-6) mm long, 0.5-0.7 mm wide, with

sparsely and remotely denticulate margins, sporangia ca. 1 mm in diameter 7b. var. minor

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

27

7a. Huperzia reflexa var. reflexa

Lycopodium bifidum Willd., Sp. pi. ed. 4, 5: 53. 1810.

TYPE: Venezuela, Cuchilla de Guajana Guajana,

Humboldt & Bonpland 474 (holotype, B, Herb.

Willd. 194211; isotype, p, Herb. Humb.!).

Lycopodium reversum Presl, Reliq. haenk. 1:82. 1825.

TYPE: Ecuador, Guayaquil, Haenke (holotype,

PRC!).

Lycopodium reflexum Lam. var. majus Spring, Mem.

Acad. roy. Belg. 15 [Mon. Lye. 1]: 26. 1842.

Huperzia reflexa (Lam.) Trev. var. bifida (Willd.) Trev.,

Atti Soc. Ital. Sci. Nat. 17: 248. 1874.

Lycopodium reflexum Lam. var. densifolium Baker,

Handb. Fern-Allies 11. 1887. SYNTYPES: Hart-

weg 1480 (K!); Moritz 2266 (= 226bl K!); Brazil,

Glaziou 15797 (K!).

Lycopodium densifolium (Baker) Underw. & Lloyd,

Bull. Torrey Bot. Club 33: 106. 1906.

Lycopodium mexiae Copel., Univ. California Publ.

Bot. 19: 294, t. 47. 1941. TYPE: Peru, Dept.

Huanuco, Churubamba, trail Cotirarda to Mer-

cedes, 1875 m, Mexia 8193a (holotype, uc!; iso-

types, GH!, K!, MICH).

Urostachys stellae-polaris Herter, Revista Sudamer.

Bot. 10: 121. 1953. TYPE: Colombia, Cundina-

marca, Guayabetal to Monte Redondo, SE of

Quetame, 1 300-1 500 m, Pennell 1801 (holotype,

us!).

Lycopodium stellae-polaris (Herter) Morton, Amer.

Fern J. 54: 72. 1964.

Huperzia bifida (Willd.) Holub, Folia Geobot. Phy-

totax. 20: 71. 1985.

Huperzia mexiae (Copel.) Rolleri and Deferrari, No-

tas Mus. La Plata, Bot. 21 (100): 156. 1988.

cucho: Prov. La Mar, eastern Massif of the Cord. Central

opposing the Cord. Vilcabamba between Tambo, San

Miguel, Ayna, and Hda. Luisiana, 1 570 m, Dudley 11826

(GH). Ayna between Huanta and Rio Apurimac, 750-

1000 m, Killip & Smith 23198 (NY). Cuzco: Prov. Con-

vencion, Hda. Guayanay, 1800 m, Vargas 13248 (GH).

Quispicanchi, Mandor, Marcapata, 1000 m, Vargas 5220

(us).

7b. Huperzia reflexa var. minor (Spring) B. 011g.

in Marling and Andersson, Fl. Ecuador 33:

26. 1988.

Lycopodium reflexum Lam. var. minus Spring, Mem.

Acad. roy. Belg. 15 [Mon. Lye. 1]: 26. 1842. SYN-

TYPES: Brasil, Pr. Rio de Janeiro, Gaudichaud

(P!); Langsdorff (M, Herb. Mart.l); Brazil, Serra

dos Orgaos, fr. Majo, Guillemin (P!); In sylvis

prov. Paraensis, Martins (M!); Brazil, in prov.

Minarum, Claussen (P!).

Lycopodium brutum Herter, Bot. Jahrb. 43: Beibl. 98:

47. 1909. TYPE: Trinidad, Hooker ded. 1845

(holotype, P!).

Humid montane forest, road banks, alt. 1000-

2400 m, Huanuco, Cuzco, Puno. Brazil; Trinidad;

Ecuador to Bolivia; probably more widespread.

This variety seems identical to Huperzia par-

vifolium (Nessel) Rolleri and Deferrari from SE

Brazil. It often grows intermixed with individuals

of var. reflexa.

Landslides, road banks, and other open or dis-

turbed habitats in montane forest, alt. 900-3400

m, Cajamarca, Amazonas, San Martin, Huanuco,

Pasco, Junin, Ayacucho, Cuzco.

Throughout humid mountainous regions of

tropical America.

Foster & Smith 9095 (USM) is very robust and

tall, with thicker stems and slightly wider leaves

than usual for this variety.

Huanuco: Prov. Huanuco, Road Huanuco to Tingo

Maria, N of Carpish Pass, 2350-2450 m, Plowman &

Rury 11146 (F, GH). Cuzco: Prov. Urubamba, Machu

Picchu, Soukup 167 (F). Near Machu Picchu, 2000 m,

Tryon & Tryon 5409 in part (GH, us, USM). Puno: Prov.

Sandia, below Cuyocuyo, 2900 m, Ferreyra 16624 (USM).

8. Huperzia acifolia (Rolleri) Rolleri and Defer-

rari, Notas Mus. La Plata, Bot. 21 (100): 155.

1988.

Cajamarca: Prov. Cutervo, El Pajonal, San Andres,

2200 m, Llatas & Suarez 2828 in part (F). Prov. Cutervo,

La Pucarilla, between Socota and San Andres, 2500 m,

Sanchez Vega el al. 5923 (AAU, F). Amazonas: Prov.

Chachapoyas, roads to Molinopampa, 2700 m, Sanchez

Vega et al. 2218 (AAU). San Martin: Prov. San Martin,

Dist. Tarapoto, road Tarapoto to Yurimaguas, km 9-

1 3, ca. 700 m, Rimachi 4102 (F, GH, NY). Huanuco: Prov.

Huanuco, Chinchao to Puente Duran, 1900 m, Ochoa

14597 (F, us). Carpish Pass, 2700 m, Hodge 6294 (GH,

us). Pasco: Manto at Yaupi, Woytkowski 6534 (MO, us).

Prov. Oxapampa, Ulcumanu SW of Oxapampa, road to

Maria Teresa and Llaupi, 2150-2450 m, Foster et al.

7688 (AAU). Junin: Tarma, Chanchamayo, above La

Merced, ca. 2000 m, Weberbauer 2006 (G, MOL). Aya-

Lycopodium acifolium Rolleri, Revista Mus. La Plata

(n.s.) 14: 2, /. 1-2. 1985. TYPE: Peru, Dept. Aya-

cucho, Cearrapa, between Huanta and Rio Apu-

rimac, 1 500 m, Killip & Smith 22368 (holotype,

us!).

Plants erect, or erect from a decumbent base,

1 5-30 cm tall, or up to 60 cm long. Shoots homo-

phyllous, almost equally thick throughout, 10-20

mm in diameter including leaves. Stems excluding

leaves 2-5(-6) mm thick at the base, sometimes

tapering to 1-3 mm thick. Leaves uniform

throughout, borne in alternating, usually densely

28

FIELDIANA: BOTANY

crowded whorls of 8-1 1, usually sharply bent up-

ward from a perpendicular junction to the stem

and then gently to strongly clawlike recurved, lin-

ear-subulate, widest just above the pale and soft-

herbaceous base, 6-10 mm long, 0.7-1(-1.3) mm

wide at the base, narrowed shortly above the base,

subcoriaceous at apex, adaxially convex distally,

abaxially flat to convex with obscure to somewhat

prominent vein (sometimes sunken when dried),

with rather densely denticulate-ciliolate margins

at base, usually sparsely denticulate or smooth at

apex. Sporangia 1-1.5 mm wide.

Moist banks in montane forest at mid-altitudes,

alt. 1350-2040 m, San Martin, Huanuco, Pasco,

Junin, Ayacucho, Cuzco.

Venezuela to Bolivia.

Closely related to Huperzia rejlexa and H. un-

guiculata B. 011g. (Colombia, Ecuador). Some ma-

terial earlier tentatively referred to H. unguiculata

from Ecuador and Peru belongs here, and thus H.

unguiculata is yet unknown from Peru.

San Martin: Prov. Rioja, Pedro Ruiz— Moyobamba

road, km 390-394, Venceremos, 1910-2040 m, D. Smith

4522 (AAU, USM). Prov. Mariscal Caceres, 60 km NE of

Tingo Maria, La Divisoria pass through Cerro Azul, 1 500

m, Tryon & Tryon 5269 (GH, us). Huanuco: Prov. Leon-

cio Prado, Dist. Daniel Alomia Robles, Road Tingo Maria

to Pucallpa, La Divisoria, 1600 m, J. Schunke V. 3082

(F, G, GH, NY, us); Ferreyra 1690 (USM). Pasco: Prov.

Oxapampa, Rio Boqueria, ca. 26 km from Oxapampa

via Rio Yamaquizu, 1870 m, D. Smith et al. 1772 (AAU,

USM). Oxapampa [as Junin], 1600 m, Soukup 2675 (F,

GH, us). Junin: Prov. Tarma, road to La Mina Pichita,

9 km from the Tarma-La Merced road, ca. 1 3 km from

San Ramon, 1 600 m, Skog et al. 5055 (us). Chancha-

mayo Valley, above La Merced at Cumbre Yacunay,

2000 m, Hutchison 11 94 A (F, M, NY, uc, us, USM). Cuzco:

Prov. Paucartambo, Kosnipata, San Pedro, 1 350 m, Var-

gas 10219 (MO, uc).

9. Huperzia eversa (Poiret) B. 011g. in Harling and

Andersson, Fl. Ecuador 33: 28. 1988.

Lycopodium reflexum Willd., Sp. pi. ed. 4, 5: 52. 1810,

not Lam. 1789. TYPE: Ecuador: Tungurahua,

(Humboldt) Nee D. D. (holotype, B, Herb. Willd.

19419\).

Lycopodium eversum Poiret, in Lam., Encycl. 3: 556.

1814 [1813].

Lycopodium reflexum Lam. var. polycarpum Sodiro,

Recens. crypt, vase. Quit. 90. 1883. TYPE: Ec-

uador, valle de Nanegal, Sodiro (not located).

Lycopodium polycarpum (Sodiro) Underw. & Lloyd,

Bull. Torrey Bot. Club 33: 105. 1906, not L. poly-

carpos Kunze 1835.

Lycopodium ecuadoricum Herter, Bot. Jahrb. 43: Beibl.

98: 48. 1909. LECTOTYPE (designated by B.

011g. in Harling and Andersson, Fl. Ecuador 33:

28. 1988): Ecuador, Andium nemoribus humidis,

Jameson 74 (P!).

Huperzia ecuadorica (Herter) Holub, Folia Geobot.

Phytotax. 20: 72. 1985.

Plants erect or erect from a decumbent base,

soft, often large, much-branched and caespitose,

up to 30(-50) cm tall. Shoots homophyllous,

equally thick throughout, (2.5-)3-6(-10) mm in

diameter including leaves. Stems excluding leaves

1.5-2.5 mm thick at the base, sometimes tapering

to 1-2 mm, prominently ridged by decurrent leaf

bases (dried). Leaves uniform throughout, borne

in irregular, alternating, subdistant to densely

crowded whorls of (4-)5(-6), wide-spreading to

sharply reflexed, usually strongly recurved, linear-

lanceolate, widest in the basal half, (2.5-)3-5 mm

long, 0.5-1 mm wide, softly herbaceous to sub-

coriaceous, adaxially convex with obscure vein,

abaxially irregularly concave (dried), with obscure

to somewhat prominent vein, with slightly revo-

lute, denticulate-ciliolate margins. Leaf base with

prominently decurrent vein and margins. Sporan-

gia 1-1.5 mm in diameter.

Terrestrial, as a pioneer on landslides, road

banks, and other open, moist habitats in upper

montaine forest, alt. 2400-3400 m, Piura, Ama-

zonas, San Martin, Huanuco, Pasco, Ayacucho,

Cuzco.

Costa Rica; Andes from Venezuela to Bolivia.

Huperzia eversa resembles H. rejlexa, but has

shorter and relatively wider leaves, which are usu-

ally strongly recurved, so that that shoots appear

much more slender. It also differs by usually form-

ing very densely branched individuals.

Piura: Prov. Huancabamba, Loma Redonda (Sapa-

lache to Chinguela), 2400 m, Sagdstegui et al. 10174

(AAU, MO). Amazonas: Prov. Chachapoyas, middle east-

ern Calla-Calla slopes, 3000-3200 m, Wurdack 1770 (F,

GH, NY, uc, us, USM). San Martin: Prov. Mariscal Ca-

ceres, Rio Abiseo National park, NW corner, Chochos,

3400 m, Young 3693a (AAU). Huanuco: Playapampa,

2700 m, Macbride 4486 (F). Cushi, trail to Tambo de

Vaca, Bryan 622 (F, us). Pasco: Prov. Oxapampa, Dist.

Oxapampa, Rio San Alberto, Abra Esperanza, 2400-

2700 m, Foster et al. 10300 (AAU). Ayacucho: Prov. La

Mar, eastern Massif of the Cord. Central opposing the

Cord. Vilcabamba between Tambo, San Miguel, Ayna,

and Hda. Luisiana, 2920 m, Dudley 11972 (F, GH). Cuz-

co: Prov. Paucartambo, Parque Nacional Manu, Camino

Eriksson, Leon & Huapalla 2387 (AAU). Michehuanunca,

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

29

Huadquina, 2700 m, Biies 745 (us). Department un-

known: Matthews 1082 (E).

10. Huperzia affinis Trev., Atti Soc. Ital. Sci. Nat.

17: 248. 1874.

Lycopodium affineGrev. & Hooker, Bot. Misc, 2: 364.

1831, not Bory 1804. LECTOTYPE (designated

by Nessel, Barlappgewachse 97. 1939): Ecuador,

Pichincha, 1824, Herb. Greville, Jameson (K!;

possible isotypes, E!, NY!, us!).

Lycopodium blepharodes Maxon, Contr. U.S. Natl.

Herb. 17: 423. 1914.

Huperzia blepharodes (Maxon) Holub, Folia Geobot.

Phytotax. 20: 71. 1985.

Plants erect from a decumbent base, at least up

to 25 cm tall. Shoots homophyllous, equally thick

throughout, 5-10 mm in diameter including leaves.

Stems excluding leaves 1.5-3 mm thick, upward

sometimes slightly tapering. Leaves borne in rath-

er close alternating whorls of 5, almost covering

the stem, patent to loosely appressed, narrowly

triangular-lanceolate to almost subulate, with

slightly widened base, (3.5-)4-6 mm long, 1-1.5

mm wide, abaxially convex and evenly rounded

or with prominent veinal ridge, at least at base,

somewhat clasping the sporangia, with long, flac-

cid marginal cilia and small teeth. Sporangia 1.5-

2 mm in diameter.

Terrestrial on banks in upper montane forest

and grassland, alt. 2700-3100 m, Amazonas,

Huanuco.

Colombia to Peru.

The description above applies to the Peruvian

specimens. These are generally smaller, are more

compact, and have more appressed and abaxially

more convex leaves than specimens from Colom-

bia and Ecuador. Huperzia affinis is variable with

respect to crowding and direction of the leaves. It

is distinguished from H. reflexa var. reflexa, H.

eversa, and H. pearcei (Baker) Holub (Bolivia) by

its wider and more convex leaf bases and the slen-

der cilia on the leaf margins.

Amazonas: Prov. Chachapoyas, Cerros Calla Calla, W

side, 45 km above Balsas, 3100 m, Hutchison & Wright

5830 (F, GH, NY, uc, USM). Huanuco: Mito, 2700 m,

Bryan 389a (F). Prov. Huanuco, Punta de Panao, As-

plund 13712 (s). Panao, 2700 m, Macbride 3602 (F).

1 1 . Huperzia kuesteri (Nessel) B. 011g., Opera Bot.

92: 169. 1987.

Urostachys kuesteri Nessel, Repert. Spec. Nov. Regni

Veg. 35: 182, t. 172. 1934. LECTOTYPE (des-

ignated by 011gaard in Harling and Andersson,

Fl. Ecuador 33: 37. 1988): Ecuador, Loja to Za-

mora, 3500 m, 1875, "/. M. J. Mission" Quito

415 (BONN, Herb. Nessel 23 /!).

Plants ascending to erect, massively robust and

large caespitose plants, with almost completely

homoblastic divisions, up to 40 cm tall. Shoots

homophyllous, almost equally thick throughout,

or slightly tapering upward and the leaves grad-

ually shorter and wider, 12-25 mm in diameter

including leaves near the base, sometimes tapering

to (7-) 10-1 5 mm. Stems excluding leaves 3-8 mm

thick at the base, tapering to 3-5 mm upward,

almost completely concealed by leaves. Leaves

uniform throughout or upward slightly reduced,

densely crowded, borne in usually regular alter-

nating whorls of 4-5, ascending to closely imbri-

cate, rarely patent to recurved in basal divisions,

straight to slightly upwardly curved, narrowly tri-

angular-lanceolate near the base to widely trian-

gular-ovate in terminal divisions, widest just above

the leaf base, 8-12 mm long and 1.5—4 mm wide

in basal divisions, upward (4-)5-10 mm long, 2.5-

4 mm wide, evenly tapering, with acute, upward-

curved apex, thick, adaxially flat or slightly con-

cave with flush or slightly prominent vein, abax-

ially rounded with slightly to strongly prominent,

narrow vein, with a short, flattened basal swelling

(air sac), shining or pruinose, green or rarely with

red-tinged margins, with smooth to uneven or ru-

gulate, opaquely to transparently sclerified mar-

gins. Leaf base margins usually slightly revolute,

sometimes subauriculate in apical divisions. Spo-

rangia 2-2.5 mm wide.

Low, wet mossy and grassy paramos, alt. 2750-

3400 m, Piura, Lambayeque. Southern Ecuador

and northernmost Peru.

This species is related to Huperzia macbridei

but can be distinguished from this by the homo-

blastic, evenly spreading branching habit, wider

leaves, and greater size.

Plura: Purchased in the market of Huancabamba,

Friedberg7631b(GH). Lambayeque: Prov. Ferranafe, Dist.

Incahuasi, Laguna Tembladera to Cerro Negro, 3300 m,

Sagdstegui et al. 12818 (AAU), 12848 (F in part).

12. Huperzia weberbaueri (Nessel) Holub, Folia

Geobot. Phytotax. 20: 78. 1985.

Urostachys weberbaueri Nessel, Revista Sudamer. Bot.

6: 162, /. 10, f. 41. 1940. TYPE: Peru, Dept.

Amazonas, Ostlich Chachapoyas, Tambo Ven-

30

FIELDIANA: BOTANY

tillas, 2400-2600 m, Rosen 672 (holotype, BONN,

Herb. Nessel 1831).

Lycopodium papillatum Rolled, Amer. Fern J. 65: 3.

1975. TYPE: Peru, Dept. Amazonas, Prov. Cha-

chapoyas, Cerro Malcabal (Cerro Tumbe), 3-6

km SW of Molinopampa, 2900 m, Wurdack 1456

(holotype, us!; isotypes, GH!, K!, uc!, USM!).

Huperzia papillata (Rolleri) Holub, Folia Geobot.

Phytotax. 20: 75. 1985.

Plants erect from an ascending base, large, caes-

pitose, homoblastic, up to 30(-40) cm tall. Shoots

homophyllous, 10-15(-20) mm in diameter in-

cluding leaves, sometimes tapering to 7 mm in

diameter, strikingly whitish papillate in life. Steins

excluding leaves 2—4 mm thick. Leaves uniform

throughout, or slightly smaller upward, borne in

more or less regular, alternating whorls of 5, patent

to ascending, upward curved, lanceolate, 7-10(-

15) mm long, 2-2.5 mm wide, not or slightly de-

current, without a prominent basal swelling (air

sac), abaxially convex, with an obscure veinal ridge,

adaxially flat to concave (dried), densely long-pap-

illate on margins and abaxially, smooth adaxially.

Sporangia ca. 1.5 mm wide.

Wet grassy and boggy paramos with some shel-

ter, and lower puna 3000-3600 m, Lambayeque,

Amazonas, San Martin, Cuzco.

Southernmost Ecuador and Peru.

The densely papillate, bright whitish leaves in

the living plants are unique in the genus. The whit-

ish color is often lost after drying, especially heat

drying.

Lambayeque: Prov. Ferranafe, Dist. Incahuasi, La-

guna Tembladera, 3 1 50 m, Sagdstegui et al. 12790 (AAU).

Amazonas: Prov. Chachapoyas, Leimebamba to Calla

Calla, 3100 m, Sanchez V. 5 JO (AAU); Boeke 1809 (AAU).

NE of Tambo de Ventilla, Cerro de Fraijaco (Huaui to

Huni), 3450 m, Pennell 15883 (GH). Cerros Calla Calla,

3100 m, Hutchison & Wright 5564 (F, GH, MO, NY, uc,

us). San Martin: Prov. Mariscal Caceres, NW corner of

Rio Abiseo National Park, 3400 m, Young & Leon 4768

(AAU). Prov. Mariscal Caceres, Chochos Valley, 3300 m,

Young & Leon 4877 (AAU). Huallaga, valley of Apison-

cho, 30 km above Jucusbamba, 3600 m, Hamilton &

Holligan 1216, 1217 (us). Cuzco: Prov. Paucartambo,

Cord, de Tres Cruces, 3600 m, Vargas 12195 (GH). Prov.

Paucartambo, Pavayoc, 2100 m, Woytkowski 555 (MOL,

USM).

13. Huperzia colanensis B. 011g., sp. nov.

Planta erecta vel ascendens; surculi homoblastici us-

que ad 30 cm alti, homophylli 6-10(-12) mm diametro

foliis inclusis, omnino crassitie aequali vel parum an-

gustati. Caulis 2-4 mm crassus foliis exclusis. Folia om-

nino uniformes vel sursum parum minores, 8-10-faria,

patentes usque ad ascendentes vel perpendiculares usque

ad parum reflexa, recta vel incurva, lanceolata, aequate

angustata vel acuminata, (3-)4-6(-7) mm longa, 1.5-2

mm lata, parum decurrentes, abaxialiter convexa, ad

basin vix vel parum ventricosa, plerumque angulo me-

diali subacuto instructa, adaxialiter plana usque ad con-

cava (siccata), subcoriacea, hypostomatica, plerumque

polita, marginibus laevibus, scleroideis et translucidis.

Sporangia ca. 1.5-2 mm lata.

Plants erect from an ascending base, loosely

caespitose, homoblastic, up to 30 cm tall. Shoots

homophyllous, 6-10(-12) mm in diameter includ-

ing leaves, equally thick throughout or slightly ta-

pering. Stems, excluding leaves, 2-4 mm thick.

Leaves uniform throughout, or slightly smaller up-

ward, borne in more or less regular, alternating

whorls of 4-5, patent to ascending or perpendic-

ular to slightly reflexed in basal divisions, straight

to upward curved, lanceolate, widest ca. '/4 of the

leaf length above the base, evenly tapering in the

distal % or slightly acuminate, straight to slightly

upward curved, (3-)4-6(-7) mm long, 1.5-2 mm

wide, somewhat decurrent, without or with a

slightly prominent basal swelling (air sac), abaxi-

ally convex, rounded or usually with a rather sharp

medial ridge, adaxially flat to concave (dried),

somewhat coriaceous, usually glossy on both sides,

the margins smooth, sclerified and somewhat

translucent. Sporangia ca. 1.5-2 mm wide.

TYPE— Peru, Dept. Amazonas, Prov. Bagua,

Cord. Colan NE of La Peca, 10,800 ft, 8 Sep 1978,

Barbour 3384 (holotype, MO!; isotypes, AAU!, USM!).

Andean grassland, open patches in shrub para-

mo, alt. 3140-3600 m, Amazonas, San Martin.

Endemic.

The specimen cited from San Martin deviates

from the other material by a softer texture and less

glossy leaves with less sclerified margins and is

included in the species with some doubt.

Huperzia colanensis appears to be intermediate

in position between the Huperzia brevifolia and

H. saururus groups of 011gaard (1987, 1989). It

does not appear to be intimately related to other

Peruvian species.

Amazonas: Prov. Bagua, Cord. Colan NE of La Peca,

3 1 70 m, Barbour 3446 (MO). Prov. Bagua, Cord. Colan

NE of La Peca, ridge W of peaks, 3 140 m, Barbour 3197

(MO), 3198 (AAU). San Martin: Dist. Huallaga, Valley of

Rio Apisoncho, 3600 m, Hamilton & Holligan 1218

(us).

14. Huperzia polylepidetorum B. 011g. in Harling

and Andersson, Fl. Ecuador 33: 42,/ 6B. 1 988.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

31

TYPE: Ecuador, Prov. Azuay, Laegaard

55117 (holotype, AAU!; isotype, QCA!).

Plants ascending to erect, large, robust, caes-

pitose, nearly homoblastic, up to 35 cm tall, or

more than 50 cm long. Shoots homophyllous,

equally thick throughout, or slightly tapering up-

ward and the leaves gradually shorter, 10-25 mm

in diameter including leaves. Stems excluding

leaves 5-7 mm thick at the base, sometimes ta-

pering to 2 mm thick, usually not completely con-

cealed by leaves. Leaves uniform throughout, or

gradually shorter upward, well spaced to densely

crowded, borne in more or less regular, alternating

whorls of 5-6, patent-ascending to arcuate-ap-

pressed, strongly curved upward and inward, lan-

ceolate to widely lanceolate, acute or short-acute,

6-12 mm long, (2-)2.5-3(-3.5) mm wide, some-

what fleshy, abaxially convex or rarely flattened,

with an obscure to prominent adaxial and abaxial,

long decurrent median ridge, with an indistinct air

sac, adaxially concave or flat, with smooth, slightly

involute margins, green throughout or with red-

tinged tips. Sporangia 2.5-3.5 mm wide.

In open Polylepis forest among mossy rocks,

3900-4300 m, Ancash.

Ecuador and Peru.

The Peruvian collection was made in shaded

Polylepis woods, and the specimen differs slightly

from Ecuadorean plants by the more patent-as-

cending leaves, which tend to be abaxially flat to

slightly concave in the basal divisions, while the

distal, more exposed divisions have abaxially con-

vex leaves. It thus seems close to Huperzia wed-

dellii.

Ancash: Prov. Huari, Huascaran National Park, Que-

brada de Yuraccocha, a lateral valley of Quebrada Ruri-

chinchay, 3900-4300 m, D. Smith et al. 12716 (AAU, F).

1 5. Huperzia capellae (Herter) Holub, Folia Geo-

bot. Phytotax. 20: 71. 1985.

Urostachys capellae Herter, Revista Sudamer. Hot. 10:

114-115. 1953. LECTOTYPE (designated by

Roller!, Revista Mus. La Plata, n.s., Bot. 13 (71):

78. 1981): Ecuador, prov. Napo [as Imbabura],

E of Volcan de Cayambe, along trail between Rio

Boqueron and Rio Arturo, 1 1,000 ft, Drew E 3 14

(us!; isotype, MSC!).

Lycopodium capellae (Herter) Morton, Amer. Fem J.

54: 72. 1964.

Plants ascending to erect, sparsely to densely

branched, without prostrate-ascending, rejuvenat-

ing shoots, up to ca. 35 cm tall. Shoots homo-

blastic, homophyllous to gradually slightly hetero-

phyllous, 12-20 mm in diameter at the base

including leaves, usually tapering to 5-10 mm.

Stems excluding leaves 2-3 mm thick at the base,

often brown to reddish brown, terete to promi-

nently ridged by decurrent leaf bases. Leaves of

basal divisions (and shaded shoots) borne in ir-

regular, rather distant, alternating whorls of 5-6,

usually patent to ascending, not covering the stem,

linear-lanceolate, 7-10 mm long, 1.5-2 mm wide,

adaxially flat, shining, abaxially flat to slightly con-

vex, dull to shining green, usually with long-de-

current leaf bases. Leaves of distal divisions grad-

ually shorter, closer, and more appressed, borne

in alternating whorls of 4-5(-6), ascending to ar-

cuate-appressed, straight to strongly upward or

unilaterally curved or twisted, lanceolate to broad-

ly lanceolate, often slightly cuspidate above the

middle, (5-)6-8 mm long, 1.5-2 mm wide, adax-

ially concave (dried) or flat to shallowly rounded

in life, abaxially convex, rounded and irregularly

wrinkled (dried), with slightly prominent, short to

long, narrowly decurrent basal swelling (air sac),

with smooth to densely irregularly rugulate, slight-

ly to distinctly, translucently sclerified margins,

shining to dull green. Sporangia ca. 2 mm wide.

Grass paramos, usually growing in partial shade

among grasses, 3600 m, Cajamarca, Cuzco.

Andes from Venezuela to Peru.

Known from two collections in Peru, matching

the Ecuadorean plants perfectly.

Cajamarca: Prov. Chota, Laguna Yahuarcocha, above

Incahuasi, 3600 m, Sagdstegui et al. 12908 (F). Cuzco:

Prov. Paucartambo, Parque Nacional Manu, vicinity of

El Mirador, 3600 m, Leon 2270 (USM).

16. Huperzia macbridei (Herter) B. 011g., Opera

Bot. 92: 169. 1987.

Urostachys macbridei Herter, Revista Sudamer. Bot.

10: 115. 1953. TYPE: Peru, Dept. Huanuco, 3-

6 mi NW of Mito, 1 1,000 ft, Macbride & Feath-

erstone 1922 (holotype, us!; isotypes, B!, F!, MA!,

s!).

Lycopodium macbridei (Herter) Morton, Amer. Fem

J. 54: 72. 1964.

Plants erect from a decumbent base, loosely to

densely caespitose, with basal, prostrate-ascend-

ing, rejuvenating shoots at the periphery, up to 32

cm tall. Shoots homophyllous or with leaves grad-

ually slightly reduced upward, equally thick

throughout or slightly tapering upward, 1 5-20 mm

32

FIELDIANA: BOTANY

in diameter including leaves near the base, often

tapering to 6-9 mm in diameter. Stems excluding

leaves 2—4 mm thick at the base, upward tapering

to ca. 2 mm, almost completely concealed by

leaves. Leaves uniform throughout, or upward

slightly reduced, borne in irregular, alternating

whorls of 4-5(-6), densely crowded, spreading to

ascending in basal divisions, upward often grad-

ually becoming closely imbricate, straight or slightly

upward curved, narrowly lanceolate to lanceolate,

widest near the base, evenly tapering, 10-15 mm

long and 1.5-2.5 mm wide in basal divisions, up-

ward 6-11 mm long, 1.5-2 mm wide, without or

with a slightly pronounced basal swelling (air sac),

adaxially concave with flush or slightly prominent

vein, abaxially rounded, with slightly prominent

and often slightly darker and long decurrent vein,

with smooth to slightly uneven, somewhat invo-

lute, indistinctly sclerified margins, green to yel-

lowish green. Sporangia ca. 2.5 mm wide.

Terrestrial in low shrub paramos, at the edge of

woods, 2500-3600 m, Amazonas, La Libertad,

Huanuco, Ayacucho, Cuzco.

Southern Ecuador and Peru.

Individuals of shaded or sheltered habitats (e.g.,

Macbride 3487) tend to retain juvenile leaf mor-

phology, being completely homophyllous with rel-

atively wide and long, patent-ascending rather than

appressed and gradually reduced leaves and often

with a more loosely caespitose growth habit. Mat-

thews 963 is a mixed collection, sheets of the same

number at G pertaining to H. hippuridea.

Amazonas: Chachapoyas (presumably), Matthews 963

(BM). La Libertad: Prov. Bolivar, Nevado del Cajamar-

L] u ilia, puna, Ferreyra 1350 (USM). San Martin: Prov.

Mariscal Caceres, Rio Abiseo National Park, Puerta del

Monte, 3350 m, Leon & Young 1500 (AAU, USM). Prov.

Mariscal Caceres, Rio Abiseo National Park, Valle de

Chochos, 3450 m, Leon 1870 (AAU, USM). Huanuco: Mito,

2700 m, Bryan 381 (F). Cani, 7 mi NE of Mito, 2500

m, Macbride 3487 (F, us). Ayacucho: Prov. La Mar, east-

ern Massif of the Cord. Central opposing the Cord. Vil-

cabamba between Tambo, San Miguel, Ayna and Hda.

Luisiana, 3200-3500 m, Dudley 11989 (F, GH). Cuzco:

Paucartambo, Tres Cruces, 3600 m, Bikes 2214 (MO).

Prov. Paucartambo, Parque Nacional Manu, vicinity of

El Mirador, 3600 m, Leon 2269 (USM).

17. Huperzia darwiniana (Nessel) B. 011g., comb,

nov.

Urostachys darwinianus Nessel [Barlappgewachse 80,

/. 3,f. 20. 1939, nom. nud.], Revista Sudamer.

Bot. 6: 161. 1940. TYPE: "Columbien: Andinum

montibus nemoribus, Ruiz" (holotype, BONN,

Herb. Nessel 1331).

Plants erect, or erect from a prostrate to as-

cending base, sometimes loosely caespitose, with

basal, prostrate-ascending, rejuvenating shoots at

the periphery, and erect, fingerlike shoots in the

center, at least up to 35 cm tall. Erect shoots homo-

phyllous or almost so, equally thick throughout,

10-15 mm in diameter including leaves, or some-

times tapering to ca. 8 mm upward. Stems ex-

cluding leaves 3-5 mm thick at the base, some-

times tapering to 2 mm, usually completely

concealed by leaves. Leaves uniform throughout,

or slightly reduced upward, borne in alternating

whorls of 4-7, in distal fully sporangiate divisions

usually 4—5, crowded, loosely to closely imbricate,

slightly upward curved and usually with a strongly

upward curved apex, lanceolate to widely lanceo-

late, slightly long-acuminate from a wide and

somewhat rounded base, 8-12 mm long, (1.8-)2-

2.5(-3) mm wide, upward sometimes reduced to

7 mm long, with an inconspicuous, narrow, long-

decurrent basal swelling (air sac), adaxially con-

cave to slightly convex at the base, abaxially con-

vex and rounded, with a prominent, often darker

colored (dried) ridge along the vein, smooth and

often shining, with smooth, slightly sclerified mar-

gins, green, not or scarcely pruinose. Sporangia 2-

3 mm wide.

Boggy paramo and jalca, humid rocks, 2700-

3600 m, Huanuco, Pasco, Cuzco.

Peru and Bolivia.

The information on the type label is in Nessel's

handwriting, and the geographic information is as

dubious as on numerous other Nessel labels. Spec-

imens in Geneva, "Peruvia, Herb. Pavon" (G!),

are likely to be isotypes. See Comments under Hu-

perzia polyclada.

Huanuco: Cushi, trail to Tambo de Vaca, Bryan 677

(F, us). Playapampa, ca. 2700 m, Macbride 4475 (F, us).

Pasco: Prov. Oxapampa, trail to summit of Cord. Yana-

chaga via Rio San Daniel, 3 1 50-3300 m, D. Smith 7710

(AAU, USM). Cuzco: Cerro Puncuyoc, 4500 m, Biles 563

(us). Prov. Urubamba, near Wenner Gren ruins, 3400-

3600 m, Metcalf 30745 in part (uc). Trail Puyupata to

Sayaccmarca, 3600 m, Vargas 2910 (us). Department

unknown: Perou (1839-1840), Gay (p). Peru, Pearce (us

1431555).

18. Huperzia crassa (Willd.) Rothm., Feddes Re-

pert. Spec. Nov. Regni Veg. 54: 60. 1944.

Lycopodium crassum Willd., Sp. pi. ed. 4, 5: 50. 1810.

TYPE: Ecuador, Antisana, Humboldt & Bon-

pland 226 3 (holotype, B,Herb. mild. 194171; iso-

types, BM!, P!).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

33

Urostachys crassus (Willd.) Nessel, Barlappgewachse

75. 1939.

Urostachys pilgerianus Nessel, Revista Sudamer. Bot.

6: 161, t. 9,f. 32. 1940. TYPE: "Peru, Cordillere,

Rautenstock et Mann, in 1 90 1 " (holotype, BONN,

Herb. Nessel 124\).

Huperzia pilgeriana (Nessel) Holub, Folia Geobot.

Phytotax. 20: 75. 1985.

Plants erect from a prostrate to ascending base,

slightly to strongly heteroblastic, loosely to densely

caespitose with prostrate-ascending or shallowly

subterranean basal shoots, these bearing often nu-

merous erect, fingerlike shoots in the center, at

least up to 40 cm tall. Erect shoots homophyllous

or almost so, equally thick throughout, or slightly

tapering upward, 5-10 mm in diameter including

leaves. Stems excluding leaves 2-3 mm thick, usu-

ally completely concealed by leaves. Leaves uni-

form throughout, or slightly reduced upward, borne

in irregular, alternating whorls (or low spirals) of

5-7, densely crowded, usually closely imbricate,

straight (or slightly upward curved in shaded di-

visions), linear-lanceolate to lanceolate, 5-9 mm

long, (1-) 1.2-2 mm wide, upward sometimes re-

duced to 4.5 mm long, with a prominent, short to

long decurrent basal swelling (air sac) causing a

perpendicular appearance of the very leaf base in

sporangiate leaves, adaxially concave to slightly

convex, with a raised veinal ridge near the base,

abaxially convex and rounded or with a prominent

veinal ridge, slightly to strongly rugose by pro-

truding, blisterlike epidermal cells, with smooth

to rugose margins, green to brick red or dark red,

usually strongly pruinose. Sporangia 1-2 mm wide.

Terrestrial in high paramo and superparamo,

puna, 3600-4850 m, La Libertad, Ancash, Junin,

Cuzco, Puno.

As here delimited, Huperzia crassa is a widely

distributed polymorphic species, occurring from

Mexico to Panama, Hispaniola, and in the Andes

from Venezuela to Bolivia.

The Peruvian material represents a disjunct

population, widely separated from the Ecuadorian

populations, and it may be distinct at least at the

variety level. Therefore, the synonyms included

for the type variety in Ollgaard (1 988) are not cited

here.

The holotype of Urostachys pilgerianus most

likely is a mislabeled duplicate of Weberbauer 6626

(F, GH, MOL, us) from Dept. Junin, Nevado Runa-

tullu, SE of Jauja, on rocks, 4400-4500 m, as this

collection matches it exactly. Mislabeled (or re-

labeled?) specimens abound in Nessel's herbari-

um. The type and the Weberbauer collection both

have creeping basal shoots and are rather densely

tufted. The leaves are twisted unilaterally, but oth-

erwise this corresponds to most of the Peruvian

material.

La Libertad: Prov. Santiago de Chuco, Jalca de Ques-

nada (Quiruvilca to Huamachuco), 4000 m, Sagdstegui

& Fabris 7574 (MO, NY). Prov. Santiago de Chuco, Dist.

Calchicadan, Escalerilla-Conzuzo road near Tamboras,

3960 m, Saunders 882 (F, GH). Ancash: Prov. Carhuaz,

Huascaran National Park, Quebrada Ishinca, 4380-4500

m, D. Smith 9480 (AUU). Prov. Carhuaz, Huascaran Na-

tional Park, Quebrada Honda, 4300-4750 m, D. Smith

et al. 11643 (F). Cord. Blanca, above Vicos, toward Le-

jiacocha, 3600 m, Hutchison & Wright 4323 (F, GH, NY,

uc). Prov. Huaylas, Huascaran National Park, Quebrada

Alpamayo, 4750 m, D. Smith et al. 9715 (AUU, F). Junin:

Mount La Juntay, near Huancayo, 4700 m, Killip &

Smith 22114 (F, NY, us). Cuzco: Cerro Salamanca, Valle

Lucumayo, 2200 m, Biies 569 (us). Prov. Urubamba,

Abra de Malaga, 4300 m, Chavez 2822 (MO); 4 1 50-4230

m, Molau & Ohman 1639 (GB). Puno: "Cord, jugis pr.

Tabina," Lechler 2043 (E). Cord, near Agapata and Sa-

chapata, Lechler 2028 (BR, G, s).

1 9. Huperzia nesselii (Nessel) Roller! & Deferrari,

NotasMus.LaPlata,Bot.21 (100): 156. 1988.

Urostachys nesselii Nessel, Revista Sudamer. Bot. 6:

161. 1940 [Barlappgewachse 7 8, / 12. 1939, nom.

inval.]. TYPE: "Ost Peru, Cuzco, 1868, Rein-

hardt" (holotype, BONN, Herb. Nessel 132, in part!).

Plants erect from a prostrate to ascending base,

slightly to strongly heteroblastic, loosely to densely

caespitose with prostrate-ascending basal shoots,

these bearing numerous erect, fingerlike shoots in

the center, up to 20 cm tall. Erect shoots homo-

phyllous or almost so, equally thick throughout,

10-15 mm in diameter including leaves. Stems

excluding leaves 2-6 mm thick, usually completely

concealed by leaves. Leaves uniform throughout,

borne in irregular, alternating whorls (or low spi-

rals) of 5-6, densely crowded, closely imbricate,

straight to irregularly curved upward or twisted at

the apex, linear-lanceolate to lanceolate or nar-

rowly elliptic, 9-12 mm long, 2-2.5 mm wide,

without a prominent basal swelling, adaxially con-

cave to slightly convex, with a raised veinal ridge

near the base, abaxially flat especially at the base

to rounded especially at the apex, with a broadly

prominent medial ridge especially at the base,

slightly rugose because of protruding, blisterlike

epidermal cells, with smooth to slightly uneven

margins, pruinose. Sporangia 2-3 mm wide.

34

FIELDIANA: BOTANY

High Andine grassland near the timber line,

3000-3500 m, Junin.

Endemic.

The material referred to this species including

the type seems to represent a single gathering, re-

ferable to Weberbauer. The information on the

type label is in Nessel's handwriting, and the geo-

graphic information is dubious as are numerous

other Nessel labels.

The species seems most closely related to Hu-

perzia saururus with respect to size and growth

habit; however, it differs by the open growth of

basal portions of the plant, and the ridged leaf

bases, and smooth margins. From Peruvian H.

crassa it differs by the large, rather flat leaves with-

out a prominent basal air sac.

Junin: Tarma, mountains W of Huacapistana, 3400-

3500 m, Weberbauer 2222 (G, MOL). "Lagasca ex herb.

Swartz, ex Tarma Provincia in Peruvia, n. 2222" (BONN,

Herb. Nessel 132 in part).

20. Huperzia andina (Rosenst.) Holub, Folia

Geobot. Phytotax. 20: 70. 1985.

Lycopodium andinum Rosenst., Repert. Spec. Nov.

Regni Veg. 5: 239. 1908, not Herter 1909. TYPE:

Bolivia, La Paz, Murusata, 5000 m, Buchtien 173

(holotype, s!).

Urostachys andinus (Rosenst.) Nessel, Barlappge-

wachse 76. 1939.

Plants with prostrate, rooting shoots from which

1 to several, usually short, erect, fingerlike shoots

arise, sometimes loosely caespitose, up to 10(-15)

cm tall. Prostrate shoots densely covered with 8-

15 mm long, unilaterally upward curved, linear,

red-tinged leaves. Erect shoots homophyllous or

almost so, equally thick throughout or slightly ta-

pering upward, ca. 6-10 mm in diameter including

leaves. Steins of erect shoots excluding leaves 2-

3 mm thick, usually completely concealed by

leaves. Leaves of erect shoots uniform throughout,

or somewhat reduced upward, borne in close, ir-

regular, alternating whorls of 4-5, usually closely

imbricate, straight to unilaterally or irregularly

twisted and often somewhat deformed, linear-lan-

ceolate to lanceolate or triangular-lanceolate,

evenly to abruptly tapering from the slightly wid-

ened base or slightly acuminate, 5-8(-l 1) mm long,

l-2(-2.5) mm wide, upward sometimes reduced

to 4-6 mm long, with a prominent basal swelling

(air sac), adaxially concave to canaliculate apical-

ly, abaxially convex and rounded, smooth without

protruding, blisterlike epidermal cells, with smooth

to irregularly rugose, often strongly sclerified mar-

gins, green to reddish-tinged. Sporangia 2-2.5 mm

wide.

In boggy grassland, open turfy puna, wet ground

at lakes, alt. (3300-)4000-4700(-5000) m, Caja-

marca, La Libertad, Ancash, Junin, Huancavelica,

Cuzco, Puno.

Peru and Bolivia.

Some collections, especially from lower alti-

tudes, are referred to this species with doubt. Leon

& Young 1566, 1621 (AAU), from San Martin, Rio

Abiseo National Park, alt. 3650 m, and Leon &

Young 1105 (USM), from La Libertad, Prov. Pataz,

Rio Abiseo National Park, Cueva de Manachaqui,

3600-3800 m, are smaller, more slender, and en-

tirely green but share the growth habit with rela-

tively large leaves on the creeping shoots and rath-

er slender erect shoots.

Cajamarca: Prov. Contumaza, Jalca El Chuno (Pozo

Chuno), 4500 m, Sagdstegui et al. 9380 (AAU, F). La

Libertad: Prov. Santiago de Chuco, Laguna La Victoria

(road to Conzuzo), 4000 m, Sagdstegui et al. 6180 (GH,

MO). Huillilas, N of Cachicadan, 4000 m, Stork & Norton

10003 (F, uc). Ancash: Prov. Yungay, Dist. Yungay, Lake

Llanganuco, ca. 3500 m, Sounders 503 (F). Junin: Prov.

Huancayo, Dist. Huancayo, Huaytapallana, Huancayo-

Pariahuanca road, ca. 4480 m. Sounders 1151 (GH). Dist.

Huancayo, ca. 28 km E of Huancayo, 4500 m, Tyron &

Tryon 5469 in part (F, GH, us). Huancavelica: Prov. Ta-

yacaja, Millpu, puna de Tocas, between Colcabamba and

Paucarbamba, 4000 m, Tovar 1945 (USM). Marcapata,

3600 m, Stafford 1007 (F, K). Puno: Carabaya, Aricoma

Lake, 4400 m, Stafford 1118 (K).

21. Huperzia saururus (Lam.) Trev., Atti Soc. Ital.

Sci. Nat. 17: 249. 1874.

Lycopodium saururus Lam., Encycl. 3: 653. 1789.

TYPE: He de Bourbon (Reunion), Commerson

(holotype, P, Herb. Law.!).

Lycopodium elongatum Sw., Syn. Fil. 175. 1806.

TYPE: Peru, Prov. Junin, Tarma, collector un-

known (holotype, s!).

Urostachys saururus (Lam.) Herter, Repert. Spec. Nov.

Regni Veg. 19: 162. 1923.

Urostachys elongatus (Sw.) Herter, Index Lye. 60. 1 949.

Lycopodium sanctae-barbarae Rolleri, Darwiniana 16:

129,/ 1 D-E, t. 1, 3B. 1970. TYPE: Argentina,

Prov. Jujuy, de la Sola 2883 (holotype, LP).

Huperzia sanctae-barbarae (Rolleri) Rolleri and De-

ferrari, Notas. Mus. La Plata, Bot. 21 (100): 156.

1988.

Plants erect, very compactly caespitose with

basal, prostrate, rejuvenating shoots from which

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

35

stiffly erect, sparsely branched, fingerlike shoots

arise, at least up to 50 cm tall. Erect shoots homo-

phyllous or almost so, usually narrow and some-

what etiolated at the base and gradually becoming

wider and green upward, 5-10 mm in diameter

including leaves at the base, upward usually 8-15

mm. Steins excluding leaves (2-)3-5 mm thick,

usually completely concealed by leaves except at

the etiolated base. Leaves uniform throughout or

gradually shorter upward in old shoots, often

smaller, narrower, pale and irregularly appressed

at the etiolated base of shoots, borne in irregular,

alternating whorls (or low spirals) of 6-8, densely

crowded, closely appressed, the sporophylls ap-

pressed from a nearly perpendicular leaf base,

straight or somewhat secund, linear to linear-lan-

ceolate, the uppermost sometimes lanceolate to

widely lanceolate, 10-12(-15) mm long, l-2(-3)

mm wide, upward sometimes reduced to 5-8 mm

long, sporophylls with a prominent basal swelling

(air sac), adaxially concave to slightly convex,

abaxially convex and rounded throughout or at

least at the apex, smooth, with minutely rugose,

thickened, sclerified margins, especially in the bas-

al leaves, green and shining to reddish-tinged or

rusty-colored and sometimes strongly pruinose.

Sporangia 2-2.5 mm wide.

Moist shrubland and grassland (jalca), often

among rocks, alt. 3300—4500 m, Cajamarca, La

Libertad, Ancash, Pasco, Junin, Huancavelica,

Cuzco.

Peru to Argentina and Chile; Kerguelen; Tristan

da Cunha; alpine regions of tropical and southern

Africa; Madagascar; Reunion; Mauritius.

Contrary to the view of Rolleri (Amer. Fern J.

67: 109-120. 1977), the present author finds the

Neotropical distribution of Huperzia saururus re-

stricted to Peru and more southern regions in

America. The Central American and north Andine

material referred to H. saururus by Rolleri belongs

to various different species.

Some plants of this species are distinctly prui-

nose and reddish-tinged, similar to Huperzia cras-

sa, but can usually be distinguished by the leaf

shape, the etiolated character of the base of the

erect shoots, and the lack of blisterlike, protruding

epidermis cells on the abaxial leaf surface.

Cajamarca: Contumaza, Jalca El Chuno, 4500 m, Sa-

gdstegui et al. 9559 (AAU, F, NY). Pozo Kuan, 3790 m,

Sagdstegui et al. 13089 (AAU, F). Pozo Kuan, Laguna el

Toro, 4100 m, Sagdstegui 9452 (F). La Libertad: Prov.

Santiago de Chuco, Pampas de la Julia, 3600 m, Sagds-

tegui et al. 11123 (AAU, GH). Jalca de Quiruvilca, 4200

m, Lopez 1506 (us). Ancash: Prov. Huari, Huascaran

National Park, Quebrada Rima Rima, a lateral valley of

Quebrada Carhuazcancha, 4200-4440 m, D. Smith et

al. 1231 1 (AAU, F). Prov. Yungay, Huascaran National

Park, Quebrada Ranicuray, 4000-4300 m, D. Smith et

al. 9142 (AAU, F). Pasco: Pasco, Huayllay, ex herb. Cruck-

shanks in 1830 (GH). Junin: San Jose, ca. 3960 m, Mac-

bride & Featherstone 1 1 1 1 (F, G, us). Huancavelica: Prov.

Tayacaja, above Hda. Tocas, between Colcabamba and

Paucarbamba, 3200 m, Tovar 1967 (USM). Cuzco: Neva-

do Sallcantay, 3900-4200 m, Biies 744 (us). Prov. Uru-

bamba, Dist Chinchero, Cuper, Hatun Wayk'o quebra-

da, 3300 m, Sallo ex Franquemont 281 (AAU, F). Prov.

Paucartambo, Pfuyucalla?, Huilcacunca, 4000 m, Var-

gas 9840 in part (MO). Madre de Dios: Pinasniocj, Pan-

tiacolla, Pass, 3600 m, Cook & Gilbert 1867 (us). De-

partment unknown: Peru, Rusby 354A (us).

22. Huperzia hypogaea B. 011g. in Harling and

Andersson, Fl. Ecuador 33: 58. 1988. TYPE:

Ecuador, Prov. Carchi, 0llgaard & Balslev

8517 (holotype, AAU!; isotype, QCA!).

Plants with short or up to 36 cm long, horizon-

tal, usually subterranean, isotomously branching

shoots, from which stiffly erect, aerial shoot sys-

tems arise. Subterranean divisions with pale ap-

pressed leaves, 2-4 mm thick including leaves.

Aerial shoots up to 40 cm tall including erect sub-

terranean divisions, stiffly erect, homophyllous,

equally thick throughout, or slightly tapering, 4-

6(-8) mm thick including leaves. Stems of aerial

shoots 1.5-3 mm thick excluding leaves, partly to

completely concealed by leaves, brownish to bright

red, with smooth to somewhat rugose epidermis.

Leaves borne in alternating, irregular whorls of 4-

5, arcuate-appressed to closely appressed, abaxi-

ally convex and rounded or with a prominent, long

decurrent veinal ridge, with or without a slightly

prominent basal swelling (air sac), adaxially con-

cave to slightly convex, linear-lanceolate to lan-

ceolate, 4-6(-7) mm long, l-1.8(-2) mm wide,

upward often reduced to 3.5-4 mm long, red-tinged

to entirely red, with smooth to unevenly rugose

and sclerified margins. Sporangia 1-1.3(-1.5) mm

wide.

Wet and boggy paramo and jalca, alt. 3000-

4300 m, Lambayeque.

Colombia to northern Peru.

Usually a very easily recognizable species of

marshes and wet depressions of the lower paramos

with a soft substrate where the subterranean run-

ner-shoots are well developed. Where the species

grows on more solid substrates, e.g., open soil in

landslides and road cuts, supraterranean creeping

shoots with well-developed, patent-ascending

36

FIELDIANA: BOTANY

leaves replace the runner shoots, and make the

distinction from slender forms of Huperzia crassa

more subtle. In such cases the difference of leaf

curvature, leaf size, and the number of leaf or-

thostichies are helpful characters.

Lambayeque: Prov. Ferranafe, Dist. Incahuasi, La-

guna Tembladera to Cerro Negro, 3300 m, Sagdstegui

et al. 12847 (AAU, F, GH).

23. Huperzia sagasteguiana B. 011g., sp. nov.

Species cum habitu Huperziae hypogaeae et H. atten-

uatae. A H. hypogaea differt surculis horizontalibus epi-

gaeis vel non profunde hypogaeis, foliis brevioribus,

marginibus denticulato-fimbriatis, epidermide abaxiali

rugulosa cellulis protrusis pustuliformibus. A H. atten-

uata differt foliis 1 0-fariis, epidermide abaxiali foliorum

rugulosa, basibus foliorum minus ventricosis.

Plants with up to 20 cm long, horizontal, epi-

terranean and creeping, or shallowly subterranean,

isotomously branching shoots, rooting along the

underside, bearing stiffly erect, up to 20 cm tall

aerial shoot systems with short to long intervals

between branches. Creeping shoots with loosely

appressed to upward curved, secund leaves, 3-5

mm thick including leaves. Aerial shoots stiffly

erect, homophyllous, terete, equally thick through-

out, 4-6 mm thick including leaves. Stems of aerial

shoots 1.5-3 mm thick excluding leaves, partly to

completely concealed by leaves, brownish to bright

red, with rugosely striate epidermis. Leaves borne

in alternating, irregular whorls of 5, arcuate-ap-

pressed to closely appressed, abaxially strongly

convex and rounded to apically indistinctly cari-

nate, with a prominent, short to long-decurrent

veinal ridge, with a somewhat prominent basal

swelling (air sac), adaxially concave (dried), lan-

ceolate to triangularly ovate-lanceolate, 3-5 mm

long, 1.5-2 mm wide, red-tinged to entirely red,

abaxially wrinkled (dried) and strongly uneven due

to the protruding blisterlike epidermal cells, with

numerous irregularly shaped and directed, soft and

pale marginal processes. Sporangia 1-1.5 mm wide.

TYPE— Peru, Dept. La Libertad, Prov. Pataz,

Paso de Alaska, carretera a Tayabamba, en ladera

abierta de Gramineas, Jalca, 3900 m, Lopez &

Sagdstegui 8177 (holotype, MO; isotypes, AAU!, GH!,

NY!).

Terrestrial in jalca vegetation, 3900-3950 m, La

Libertad.

Endemic.

The growth habit resembles Huperzia hypogaea,

but the trailing stems are usually on or just below

the surface of the ground, and the leaves are short-

er, strongly pruinose, with many blisterlike cells

on the abaxial leaf epidermis, and have densely

denticulate-fimbriate margins. Huperzia attenu-

ata is rather similar but is less distinctly hetero-

blastic, has a smooth abaxial leaf epidermis, and

has a more sharply prominent basal abaxial air

sac on the upper leaves, and the leaves are borne

in whorls of three to four.

La Libertad: Prov. Pataz, between Retama and La Paz,

3950 m, Lopez & Sagdstegui 3590 (GH).

24. Huperzia attenuata (Spring) Trev., Atti Soc.

Ital. Sci. Nat. 17: 249. 1874.

Lycopodium attenuatum Spring, Mem. Acad. roy. Belg.

24 [Mon. Lye. 2]: 8. 1849. LECTOTYPE (des-

ignated by Lellinger, Proc. Biol. Soc. Wash. 89:

717. 1977): Ecuador (Prov. Pichincha), in declivi-

tate montis Pichincha, Hartweg 1470 (us!; isolec-

totypes, BM!, GL!, K!, NY!, P!).

Urostachys attenuatus (Spring) Nessel, Barlappge-

wachse 101. 1939.

Plants ascending to erect, rather small, caespi-

tose with erect shoots in the center, and prostrate-

ascending, rejuvenating basal shoots in the pe-

riphery, up to 20 cm tall, or up to 30 cm long.

Shoots homophyllous to slightly gradually hetero-

phyllous, terete to hexagonal, equally thick

throughout, 4-7 mm in diameter including leaves.

Steins excluding leaves 1 .5-2 mm thick at the base,

sometimes tapering to 1-1.5 mm in diameter up-

ward, completely concealed by leaves, tinged with

red or brick-red. Leaves borne in alternating whorls

of 3—4, forming 6 or 8 usually regular ranks, ap-

pressed to the stem throughout, in basal divisions

lanceolate and convex abaxially, upward gradually

changing to narrowly triangular-ovate with acute

or slightly acuminate apex, abaxially strongly con-

vex to subcarinate near the apex, with a prominent

basal swelling (air sac), (4-)5-6 mm long, in basal

divisions 1-1.5 mm wide, distally 1.5-2.2 mm

wide, with fimbriate margins, tinged with red. Spo-

rangia 1.5-2 mm wide.

Exposed cushion vegetation in paramo, pioneer

vegetation on landslides, road banks, etc., in the

paramo zone, alt. 3400-3500 m, Cajamarca, La

Libertad, San Martin.

Costa Rica; Ecuador and Peru to Bolivia.

An easily recognizable species, characterized by

fimbriate leaf margins. Huperzia attenuata is a

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

37

close relative of H. tetragona (see the following for

discussion).

C'ajamarca: Prov. Cajamarca, N of Canal Cumbe Mayo,

3400-3500 m, Sanchez. V. & Molau 3746 (AAU). La

Libertad: Prov. Huamachuco, Pallar-Huaguil, road to

Tayabamba, 3400 m, Lopez & Sagdstegui 8148 (AAU, F,

MO). San Martin: Prov. Mariscal Caceres, Rio Abiseo

National Park, NW sector, Valle de Chochos, 3450 m,

Leon 1872 (AAU, USM). Prov. Mariscal Caceres, Rio Abi-

seo National Park, Pampa de Cuy, 3450 m, Leon &

Young 1401A (USM).

25. Huperzia tetragona (Hooker & Grev.) Trev.,

Atti Soc. Ital. Sci. Nat. 17: 248. 1874.

Lycopodium tetragonum Hooker & Grev., Icon. Fil.

1: t. 109. 1829. TYPE: Ecuador (Prov. Pichin-

cha), Prope verticem montis Pichincha, Jameson

(E!, K!).

Lycopodium catharticum Hooker, Ann. Nat. Hist. 1 :

430-^3 1 , /. 14. 1 838. SYNTYPES: Ecuador (Prov.

Azuay), "Asuay of the Equator," W. Turner (K!);

Ecuador (Prov. Canar), from the mountains of

Pillzhum, Jameson (K!).

Lycopodium myrsinites Lam. var. minus Spring, Mem.

Acad. roy. Belg. 15 [Mon. Lye. 1]: 29. 1842, based

on Lycopodium catharticum Hooker and with the

same type.

Lycopodium tetragonum Hooker & Grev. var. patu-

lum Spring, Mem. Acad. roy. Belg. 24 [Mon. Lye.

2]: 12. 1849, based on Lycopodium catharticum

Hooker and with the same type.

Urostachys tetragonus (Hooker & Grev.) Nessel, Bar-

lappgewachse 135. 1939.

Urostachys catharticus (Hooker) Nessel, Barlappge-

wachse 135. 1939.

Plants ascending to erect, rather small, caespi-

tose with erect shoots in the center, and usually

prostrate to ascending rejuvenating basal shoots

in the periphery, up to 20(-30) cm tall. Shoots

homophyllous, or gradually slightly heterophyl-

lous, quadrangular and equally thick throughout,

2.5-3.5(-4) mm in diameter including leaves.

Steins excluding leaves 1-2 mm thick. Leaves dec-

ussate throughout, imbricate, concealing the stem,

in basal divisions widely lanceolate to triangular-

ovate, abaxially rounded and evenly decurrent,

upward triangular, carinate at least in the upper

half, somewhat decurrent or abaxially with a

prominent basal swelling (air sac), 3.5-5(-7) mm

long, 2-4 mm wide at the base, evenly tapering

into an acute or slightly acuminate apex, orange

to bright red-tinged at the margins throughout,

with fimbriate margins. An occasional branch may

be hexagonal or terete, with leaves borne in alter-

nating whorls of 3. Sporangia 1.3-2.5 mm in di-

ameter.

Exposed cushion vegetation, pioneer vegetation

on landslides, road banks, etc., in the lower part

of paramos and jalca, 2750-3600 m, Cajamarca,

La Libertad, San Martin, Huanuco, Cuzco.

Columbia to Bolivia.

The red or reddish, quadrangular, erect shoots

make this species an easily recognizable one. In-

complete material of some epiphytic species with

quadrangular constricted shoots have been fre-

quently misidentified as this species. They are

readily separated because of the fimbriate leaf

margins in Huperzia tetragona.

Huperzia tetragona is closely related to H. at-

tenuata. The occasional aberrant hexagonal or te-

rete shoots in some specimens of//, tetragona are

virtually indistinguishable from slender shoots of

H. attenuata. The two species are also ecologically

rather similar and are often found to grow inter-

mixed. There are numerous examples of species

in which the number of leaves in each whorl is

variable in the same population. The difference of

terete and quadrangular shoots in Huperzia atten-

uata and H. tetragona may be due to such simple

and taxonomically overrated variation within

populations of one species.

Cajamarca: Prov. Chota, Laguna Yahuarcocha, above

Incahuasi, 3600 m, Sagdstegui et al. 12901 (AAU). Prov.

Cajamarca, N of canal Cumbe Mayo, 3400-3500 m,

Sanchez Vega & Molau 3747 (AAU). La Libertad: Pataz,

Puerta del Monte, ruta de Huaylillas, 3200 m, Lopez &

Sagdstegui 3450 (GH). San Martin: Prov. Mariscal Ca-

ceres, Rio Abiseo National Park, Chochos valley, 3425

m, Young 3507 (AAU); 3300 m, Young & Leon 4878

(AAU). Huanuco: 3-6 mi NW of Mito, ca. 3350 m, Mac-

bride & Featherstone 1923 (F, s, us). Playapampa, ca.

2750 m, Macbride 4890 (F, o, us). Cuzco: Paucartambo,

Huillcacunca to Huaisampilla, 3800 m, Vargas 9859

(MO, uc).

26. Huperzia sellifolia B. 011g. in Harling and An-

dersson, Fl. Ecuador 33: 68, / 12C. 1988.

TYPE: Ecuador, Prov. Carchi, B. 0llg. & Bal-

slev 8432 (holotype, AAU!).

Plants erect or ascending, usually forming large,

caespitose, homoblastic plants, often more than

25 cm tall, with up to 45 cm long shoots. Shoots

homophyllous, almost equally thick throughout,

(5-)6-8(-10) mm in diameter including leaves.

Stems excluding leaves (2.5-)3—4 mm thick, al-

most completely concealed by leaf bases, except

in basal divisions. Leaves almost uniform

throughout, usually densely crowded throughout,

borne in rather regular alternating whorls of 4-5,

loosely imbricate or ascending to patent or sharply

38

FIELDIANA: BOTANY

reflexed and appressed to the stem, usually dis-

tinctly sigmoid, widely lanceolate to ovate or el-

liptic, acute or obtuse, (1.8-)2-2.6(-3.5) mm long,

(1.2-)1.5-2(-2.4) mm wide, coriaceous, usually

brightly red-tinged, abaxially convex to concave

with prominent vein in basal part of lamina,

rounded in the apex, with prominent, short to long-

decurrent basal swelling (air sac), with usually

slightly involute, narrowly sclerified, darker,

smooth to slightly uneven margins, adaxially with

slightly to distinctly prominent vein. Sporangia 1-

1.5 mm in diameter.

Terrestrial in exposed wet paramos, alt. 3 1 50-

3650 m, Piura, Lambayeque, Amazonas, San

Martin.

Andes of southern Colombia to northern Peru.

Piura: Purchased in the market of Sullana, Friedberg

3933c (GH). Lambayeque: Prov. Ferranafe, Dist. Inca-

huasi, Laguna Tembladera, 3150 m, Sagdstegui et al.

12799 (AAU, F). Amazonas: Prov. Bagua, Cord. Colan

NE of La Peca, ca. 3300 m, Barbour 3382 (AAU, MO,

USM). San Martin: Prov. Mariscal Caceres, Rio Abiseo

National Park, Paredones, 3650 m, Leon & Young 1564

(AAU).

27. Huperzia engleri (Herter) B. 011g., Opera Bot.

92: 169. 1987.

Lycopodium engleri Herter, Bot. Jahrb. Syst. 43, Beibl.

98: 45. 1909. TYPE: Peru, Dept. Huanuco, Prov.

Huamalies, mountains SW of Monzon, 3200-

3300 m, Weberbauer 3359 (holotype, B!; isotype,

BONN, Herb. Nessel 17 3\, MOL!).

Urostachys engleri (Herter) Nessel, Barlappgewachse

94. 1939.

Plants erect or ascending, up to 1 0 cm tall. Shoots

homophyllous, equally thick throughout, 8-10 mm

in diameter including leaves. Stems excluding

leaves 1-1.5 mm thick, densely papillate. Leaves

well spaced at base, crowded upward, borne in

alternating whorls of 4, patent to somewhat re-

flexed, widely lanceolate to oblong-lanceolate, acute

to obtuse, 4-5.5 mm long, 1.5-2 mm wide, slightly

convex abaxially with slightly prominent vein, with

evenly decurrent base, without a basal swelling,

adaxially with a shallow groove along the vein,

with minutely rugose margins. Sporangia 2-2.5

mm wide, nearly globose.

Habitat information according to protologue:

Bog at grass steppe interrupting the shrub for-

mation.

Endemic, Huanuco.

Possibly related to Huperzia rufescens (Hooker)

Trev. (Ecuador) and H. brevifolia but apparently

adapted to more sheltered habitats. The sporangia

are disproportionately large, thick, and volumi-

nous and do not open as is otherwise usual in dried

specimens. This, plus the rarity of the species, sug-

gests hybridity, but there is no other evidence of

hybridity, because the spores seem normally de-

veloped. The collection by Jelski from the Nessel

Herbarium needs to be confirmed; it was not found

in Krakow.

Department unknown: "Ost-Cordilere, Jelskr [pos-

sibly Dept. Cajamarca, Prov. Cutervo] (BONN, Herb. Nes-

sel 1731).

28. Huperzia brevifolia (Grev. & Hooker) Holub,

Folia Geobot. Phytotax. 20: 71. 1985.

Lycopodium brevifolium Grev. & Hooker, Hooker Bot.

Misc. 3: 104. 1832. TYPE: Peru, in Herb. Lam-

bert, Ruiz and Pavon (BM?; possible isotypes, FI,

Herb. Webb; AAU photo ex FI!, o!).

Urostachys rufescens (Hooker) Nessel var. brevifolius

(Grev. & Hooker) Nessel, Barlappgewachse 103.

1939.

Urostachys brevifolius (Grev. & Hooker) Herter, Index

Lye. 54. 1949.

Plants erect or ascending, stiff and robust, form-

ing small to large, caespitose, homoblastic plants,

often more than 25 cm tall. Shoots homophyllous,

almost equally thick throughout, (5-)7-12 mm in

diameter including leaves. Stems excluding leaves

(3-)4-7 mm thick at the base, slightly tapering

upward, usually concealed by leaf bases. Leaves

almost uniform throughout, densely crowded, or

sometimes more spaced in basal divisions, borne

in alternating whorls of 4— 5, ascending or perpen-

dicular to sharply reflexed, with straight to upward

curved apex, widely ovate or triangular-ovate to

widely suborbicular-cordate or triangular-cordate,

acute to almost obtuse or mucronulate, 2-4 mm

long, 2-4(-5) mm wide, stiffly coriaceous, green-

or red-tinged, abaxially concave, with sharply

prominent vein, with sharply prominent, some-

what flattened, short-decurrent basal swelling (air

sac), with slightly revolute, strongly sclerified,

darker and somewhat translucent, slightly erose

margins. Sporangia 1.5-2.5 mm wide.

Terrestrial in wet paramos, jalca, alt. 2700-4 1 20

m, Piura, Lambayeque, Cajamarca, Amazonas,

Huanuco, Pasco, Cuzco.

Costa Rica; Colombia to Peru.

The total distribution range is uncertain, as there

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

39

is considerable regionally characteristic variation

in the material referred to Huperzia brevifolia.

Populations from Colombia, southern Ecuador and

Peru are slightly different and usually recognizable;

perhaps they are worth taxonomic recognition.

Huperzia brevifolia differs from H. rufescens and

H. sellifolia by its larger size, thick stems, and stiff

and almost prickly leaves and by the frequent lack

of red coloration.

Dudley 11109 deviates by its slender shoots with

very short leaves.

Piura: Purchased in the market of Sullana, Friedberg

3933e (GH). Lambayeque: Prov. Ferranafe, Dist. Inca-

huasi, Laguna Tembladera, 3150 m, Sagdstegui 12792

(F, GH), 12848 (GH in part). Cajamarca: From Chiclayo

market, Leon (AAU, USM). Amazonas: Prov. Chachapo-

yas, Leimebamba-Chilchos trail, Boeke 2130 (AAU). Prov.

Bagua, Cord. Colan NE of La Peca, 3300 m, Barbour

3381 (AAU, MO). Huanuco: Playapampa, 2700 m, Mac-

bride 4476 (F, us). Cushi, trail to Tambo de Vaca, Bryan

678 (F, us). Pasco: Prov. Oxapampa, trail to summit of

Cord. Yanachaga via Rio San Daniel, 3350-3620 m, D.

Smith 7742 (AAU, USM). Junin: Prov. Tarma, mountains

W of Huacapistana, 3400-3500 m, Weberbauer 2225

(MOL). Cuzco: Prov. Paucartambo, Paso del Gallo, 2900

m, Vargas 4279 in part (uc, us). Urubamba, Machu

Picchu, Runcuraccay-Sayamarca pass, 4120 m, Peyton

288 (AAU). Prov. La Convention, summit ridge of Vil-

cabamba, ca. km 28 along trail from Hda. Luisiana and

Rio Apurimac, 3330-3410 m, Dudley 11109 (GH, us).

29. Huperzia hohenackeri (Herter) Holub, Folia

Geobot. Phytotax. 20: 73. 1985.

Lycopodium hohenackeri Herter, Bot. Jahrb. 43: Beibl.

98: 46. 1 909. TYPE: Peru, Dept. Puno, in summis

Cordiller. jugis pr. Tabina, Lechler 2043 (B!, BR!,

K, NY!, p, s!).

Urostachys hohenackeri (Herter) Nessel, Barlappge-

wachse 105. 1939.

Plants erect or ascending, forming small to large,

caespitose, homoblastic plants, up to 40 cm tall.

Shoots homophyllous or with gradually smaller

leaves upward, equally thick throughout, 10-13(-

1 5) mm in diameter including leaves, or tapering

to (5-) 7-10 mm in diameter upward. Stems ex-

cluding leaves 5-6 mm thick at the base, upward

tapering to (2-)3— 4 mm thick, partly to completely

concealed by leaves. Leaves borne in more or less

regular, alternating low spirals or whorls of 5-6,

often well spaced in basal divisions, densely

crowded upward, those of basal non-sporangiate

divisions ascending to spreading, widely triangu-

lar-lanceolate or ovate-lanceolate, 4.5-5.5 mm

long, 2.5-3 mm wide, with short to long, evenly

decurrent leaf bases. Leaves of upper, sporangiate

divisions patent to loosely imbricate, triangular-

ovate to ovate or widely ovate, acute, 3-5 mm

long, 2-3.5 mm wide, softly to firmly coriaceous,

green- to red-tinged, adaxially concave, or slightly

convex and with a prominent veinal ridge at the

base, abaxially usually slightly concave at the base,

above the base convex and rounded or with a

slightly to strongly prominent veinal ridge, with a

strongly prominent, narrow basal swelling (air sac),

with upward curved apex, with usually strongly

sclerified, irregularly rugose to minutely erose

margins. Sporangia 1.7-2 mm wide.

Terrestrial on banks, steep, mossy slopes in wet

paramos, jalca, alt. 3400-4000 m, Cuzco, Puno.

Colombia to Peru.

Huperzia hohenackeri resembles most the large

individuals of H. brevifolia, with which it shares

the general aspect of size and growth habit. It dif-

fers in the longer, more acute and usually ap-

pressed leaves. Metcalf 30745 deviates by having

recurved leaves throughout.

Cuzco: Prov. Urubamba, trail Piajupata to Sayacmar-

ca, 3600 m, Vargas 2911 (us). Near Wenner Gren ruins,

3400-3600 m, Metcalf 307 45 (uc in part). Prov. La Con-

vencion, Alturas de Rio Calzada, Huadquina, ca. 3660

m, Bties 755 (us). Puno: Tabina, in summis Cordilleras,

julio 1854, Hohenacker (BONN, Herb. Nessel 199). Cor-

dillera near Tabina, Hohenacker 4204 (BONN, Herb. Nes-

sel 199). Cord, near Ayapata and Sachapata, Lechler

2028 (E). Department unknown: Cordillera, 1902, Lan-

doni (BONN, Herb. Nessel 199).

30. Huperzia hartwegiana (Spring) Trev., Atti.

Soc. Ital. Sci. Nat. 17: 248. 1874.

Lycopodium hartwegianum Spring, Mem. Acad. roy.

Belg. 24 [Mon. Lye. 2]: 14. 1849. LECTOTYPE

(designated by Ollgaard in Harling and Anders-

son, Fl. Ecuador 33: 76. 1988): Colombia, Prov.

Popoyan, Hartweg 1466 (K! annotated by Spring;

isolectotypes, BM!, G!, LG!, NY!).

Lycopodium caracasicum Herter, Hedwigia 49: 88, /.

3a. 1909. TYPE: Venezuela, Caracas, Jan. 1856,

Gollmer (holotype, B!).

Urostachys hartwegianus (Spring) Nessel, Barlappge-

wachse 90. 1939.

Urostachys caracasicus (Herter) Nessel, Barlappge-

wachse 90. 1939.

Huperzia caracasica (Herter) Holub, Folia Geobot.

Phytotax 20: 71. 1985.

Plants robust, recurved or pendulous, at least

up to 30 cm long. Shoots homophyllous to grad-

ually heterophyllous, 15-30 mm in diameter in-

cluding leaves in basal divisions, sometimes ta-

pering to 10-15 mm in diameter. Stems excluding

40

FIELDIANA: BOTANY

leaves 2.5-3.5(-5) mm thick at the base, tapering

to 1.5-2 mm. Leaves of basal divisions borne in

often regular alternating whorls of 4-5, falcately

patent-ascending to imbricate, radially directed to

somewhat secund, almost completely covering the

stem and sporangia, lanceolate, coriaceous, usu-

ally shining, 14-23 mm long, 2.5-3.5 mm wide,

widest below the middle to near the base, abaxially

convex at the base, usually slightly concave toward

the tip, with smooth, flat to somewhat involute

margins, the vein slightly prominent at the base

and apically obscure. Leaves of terminal divisions

conform, or usually smaller, 7-10(-13) mm long,

1 .5-3 mm wide, often somewhat clasping with the

base widened, softer and less shining. Sporangia

1.7-3 mm in diameter.

Rupestral in montane forest, alt. 2400-3100 m,

Ancash, Cuzco.

Southern Mexico; Guatemala; Andes from Ven-

ezuela to Peru.

Huperzia hartwegiana is widely distributed and

variable. The species is often epiphytic outside

Peru, but in Peru all records are from rocky sub-

strates. The variation seems partly correlated with

habitat type. The species frequently occurs in sites

that are too dry for the other Huperzia species. In

such exposed rupestral sites, the plants initially

grow erect, then recurved, and may develop rather

short and rigid shoots only. Such forms were rec-

ognized as Lycopodium caracasicum by Herter and

others. In moist, shaded, epiphytic situations long,

lax, pendulous forms are developed. Huperzia

hartwegiana is related to H. taxifolia.

Ancash: Prov. Bolognesi, Quero E of Huasta, Cerrate

2471 (USM). Cuzco: Prov. Cuzco, Cadena de Pactatayan,

Vargas 17044 (GH). Prov. Urubamba, Ollantaytambo,

3050 m, Plowman & Davis 4741 (GH). Cusuchaca, 2400

m, Vargas 22971 (GH). Valle de Urubamba, Herrera

3423 (c). Valle de Urubamba, ca. 1 0 km from the village,

Leon 446 (USM). Quebrada Las Penas, 3100 m, Chavez

3493 (MO).

31. Huperzia taxifolia (Sw.) Trev., Atti Soc. Ital.

Sci. Nat. 17: 248. 1874.

Lycopodium taxifolium Sw., Prodr. 138.1788. TYPE:

A remounted specimen, marked as type, without

original annotation, in the Regnellian type her-

barium in s!, is possibly the holotype.

Lycopodium passerinoides Kunth, in HBK., Nov. gen.

sp. 1: 41. 1816. TYPE: Peru, prope Olleras et

Aipate, alt. 747 hexp., Humboldt (holotype, P,

Herb. Humb.l; isotypes, B, Herb. Willd. 194101,

BONN, Herb. Nessel 160\).

Lycopodium nitens Schlecht. & Cham., Linnaea 5:

623. 1830. TYPE: Mexico, in arboris vetustis

prope Jalapam, Schiede & Deppe (holotype, B!;

isotype, BM!).

Huperzia passerinoides (Kunth) Trev., Atti Soc. Ital.

Sci. Nat. 17: 248. 1874.

Huperzia passerinoides (Kunth) Trev. var. nitens

(Schlecht. & Cham.) Trev., Atti Soc. Ital. Sci. Nat.

17: 248. 1874.

Lycopodium schwendeneri Herter, Bot. Jahrb. 43: Beibl.

98: 50. 1909. TYPE: Not designated. Mexico in-

dicated as locus classicus by Herter 1923.

Urostachys taxifolius (Sw.) Herter, Repert. Spec. Nov.

Regni Veg. 19: 162. 1923.

Phlegmariurus taxifolius (Sw.) Love & Love, Taxon

26: 324. 1977.

Plants lax and pendent, or sometimes recurved

from an erect and somewhat rigid stem base, up

to 50(-70) cm long. Shoots usually gradually ta-

pering from ca. 20-30 mm in diameter including

leaves at the base, to 4-10 mm in distal divisions

of fully developed plants, sometimes not, or only

slightly tapering, rarely abruptly constricted. Stems

excluding leaves 1.5-2 mm thick at the base, ta-

pering to 1-1.5 mm. Leaves usually reduced and

modified upward, borne in irregular alternating

whorls of 3-5. Leaves of basal divisions spreading

to ascending or somewhat appressed, often twisted

at the base, narrowly lanceolate, widest in the low-

er half, firmly herbaceous to subcoriaceous, 14-

23 mm long, 2-3 mm wide, almost flat or some-

what concave adaxially, with flat or slightly rev-

olute, smooth margins, with evident to somewhat

prominent vein abaxially. Leaves of middle and

terminal divisions usually gradually shorter, nar-

rower and more appressed, abaxially more convex,

with involute margins. Leaves of fully sporangiate

divisions often distinctly 6-( 1 0-) ranked, with wid-

ened, clasping base, partly covering the sporangia,

often abruptly contracted into a short to long, nar-

row, involute apex, 3-8 mm long, l-1.5(-2) mm

wide. Sporangia 1-1.5 mm wide.

Usually epiphytic in lower to mid-altitude wet

montane forest, alt. 1300-1400 m, Cajamarca,

Amazonas.

Central America; West Indies; northern tropical

South America.

Huperzia taxifolia is a variable species, the de-

limitation of which is problematic, especially out-

side Peru. The Peruvian specimens cited below

are unusual in having the leaves arranged in al-

ternating whorls of 5. Hutchison & Bismarck 6379,

unusual in the nearly conform sporophylls and

uniformly patent-ascending leaves throughout, is

apparently juvenile.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

41

Cajamarca: Prov. Hualgayoc, Hda. Taulis, between

the Casa Hacienda and Palmito, 2000 m, Hutchison &

Bismarck 6379 (uc, USM). Amazonas: Prov. Bongara,

vicinity of Campamiento Ingenio 1-3 km up road to

Pomacocha (and Rioja) from camp along the Rio In-

genio, 1300-1400 m, Hutchison & Wright 3825 (F, GH,

uc).

32. Huperzia rosenstockiana (Herter) Holub, Fo-

lia Geobot. Phytotax. 20: 76. 1985.

Lycopodium rosenstockianum Herter, Hedwigia 49:

90. 1909. TYPE: Ecuador, Ost-Abhang des Tun-

gurahua, 3000 m, Rimbach 130 (holotype, Herb.

Rosenstock, not located; isotypes, s!, uc!, us!).

Urostachys rosenstockianus (Herter) Nessel, Bar-

lappgewachse 174. 1939.

Plants lax, pendulous, rarely recurved from an

ascending base, at least up to 1 50 cm long. Shoots

homophyllous, or with gradually slightly smaller

leaves upward, 20-25(-30) mm in diameter in-

cluding the leaves in basal divisions, sometimes

tapering to 10-14 mm. Stems excluding leaves

(1 .5-)2-3(-5) mm thick at the base, tapering to 1-

2 mm. Leaves of basal divisions borne in alter-

nating irregular whorls of 3-5, irregularly spread-

ing to ascending, usually somewhat twisted at the

base, with oblique to vertical lamina, lanceolate

to narrowly oblong, widest at or below the middle,

with widely cuneate to somewhat rounded base,

with short to long acute apex, 10-15 mm long, 2-

4 mm wide, subcoriaceous and opaque to herba-

ceous and translucent, almost flat, with prominent

vein adaxially, or sometimes folded down along

the vein, with slightly revolute, minutely rugose

margins with individually protruding epidermis

cells, especially at the apex. Leaves of middle and

terminal divisions conform, or gradually smaller

and more ascending to loosely appressed, more

densely crowded. Leaves of densely sporangiate

divisions usually widest just above the base, lan-

ceolate to narrowly triangular, with a rounded base,

6-9 mm long, 1.5-2.5 mm wide, partly covering

the sporangia. Sporangia 1.5-2 mm wide.

Usually epiphytic, rarely terrestrial, in cloud for-

est and elfin forest above 3000 m, San Martin.

Colombia to northern Peru.

Huperzia rosenstockiana resembles H. taxifolia,

but details of the leaf margin, color, and ecology

are quite distinct. Closely related to H. brong-

niartii with which it shares the rugulate leaf mar-

gins. Huperzia socratis (Herter) Rolleri & Defer-

rari (type from Colombia) seems to be a synonym

of the present species.

San Martin: Prov. Mariscal Caceres, Rio Abiseo Na-

tional Park, Puerta del Monte, 3400 m, Leon & Young

1303 (AAU, USM). Prov. Mariscal Caceres, Rio Abiseo

National Park, Laguna de Chochos, 3300 m, Young &

Leon 4855 (AAU).

33. Huperzia funiformis (Spring) Trev., Atti Soc.

Ital. Sci. Nat. 17: 248. 1874.

Lycopodium funiforme Spring, Bull. Acad. roy. Sci.

Bruxelles 8 (2): 516. 1841. LECTOTYPE (des-

ignated by Spring, Mon. Lye. 2: 49. 1949): Gua-

deloupe, L'Herminier (P!; isolectotypes, BR!, GH!,

NY!, uc!).

Urostachys funiformis (Spring) Herter in Urban, Symb.

Antill. 9: 387. 1925.

Plants robust, flaccidly pendulous, ropelike, at

least up to 250 cm long. Shoots homophyllous,

almost equally thick throughout, 5-10(-15) mm

in diameter including leaves, or sometimes taper-

ing from a thicker base with patent-ascending

leaves. Stems excluding leaves 1 .5-3(-5) mm thick

at the base, ridged by decurrent leaf bases. Leaves

almost uniform throughout, or slightly shorter up-

ward, borne in crowded alternating whorls of 7-

8, densely covering the stem, usually closely fal-

cate-appressed throughout, not twisted, linear-su-

bulate, widest just above the base, 6-10(-12) mm

long, 1-1.5 mm wide, evenly tapering into a long

pungent apex, firmly herbaceous to coriaceous, dull

to shining, abaxially strongly convex, often api-

cally conduplicate, with smooth margins. Sporan-

gia 1-1.5 mm wide.

Lower montane rain forest, up to alt. ca. 1 300

m, Huanuco.

West Indies; southern Mexico to Panama; Guy-

ana; Venezuela to Peru.

Huperzia funiformis is a very distinct species,

not easily confused with other species due to its

smooth ropelike shoots.

Huanuco: SW slope of the Rio Llullapichis watershed,

on the ascent of Cerros del Sira, 1290 m, Wolfe 12330

(GH).

34. Huperzia buesii (Herter) B. 011g., comb. nov.

Urostachys buesii Herter, Revista Sudamer. Bot. 10:

126. 1954 [as 1953]. TYPE: Peru, Tranca de Ya-

racuaya, 1800 m, Bues 742 (holotype, us!).

Lycopodium buesii (Herter) Morton, Amer. Fern J.

54: 72. 1964.

Plants robust, pendent, up to 50 cm long. Shoots

homophyllous, almost equally thick throughout,

42

FIELDIANA: BOTANY

ca. 10-20 mm in diameter including leaves, or

tapering to 10 mm. Steins excluding leaves 2-3

mm thick, prominently ridged by decurrent leaf

bases. Leaves almost uniform throughout, or

slightly shorter distally, borne in close alternating

whorls of 4-5, densely covering the stem, usually

closely appressed throughout, slightly recurved at

the apex, not twisted, linear to linear-lanceolate,

widest just above the slightly rounded base, 12-

19 mm long, 1-2 mm wide, evenly tapering into

a long apex, firmly herbaceous to subcoriaceous,

shining, abaxially convex, somewhat involute, with

smooth margins. Leaves of sporangiate divisions

sometimes reduced to 6-8 mm. Sporangia 1.5-2

mm wide.

A rare endemic. Epiphytic in forest, 1800 m,

Cuzco.

Perhaps related to Huperzia funiformis and H.

aristei (Nessel) Rolleri and Deferrari (?Colombia)

with which it shares the high number of leaves in

each whorl and the general leaf shape. However,

the leaves are longer and more diverging. The sim-

ilarity to the juvenile plants of H. sotae (Rolleri)

Rolleri and Deferrari (NW Argentina to Bolivia,

e.g., Venturi 2955, isotype in us!) is close.

Cuzco: Prov. La Convention, Choquellohuanca, 1 800

m, Marin 2247 (F).

35. Huperzia linifolia (L.) Trev., Atti Soc. Ital.

Sci. Nat. 17: 248. 1874.

Lycopodium linifolium L., Sp. pi. 1 100. 1753. TYPE:

Plumier, Traite foug. Amer. /. 166, f. C, from an

unspecified locality, either Martinique or His-

paniola (see Proctor, Fl. Lesser Antilles 2: 26.

1977).

Urostachys linifolius (L.). Herter, Repert. Spec. Nov.

Regni Veg. 19: 154. 1923.

Plants pendulous, usually with flaccidly hanging

divisions, the terminal divisions often aggregated

in fasciculate clusters, up to 60(-80) cm long. Shoots

homophyllous or gradually heterophyllous, equal-

ly thick throughout, 20-30(-45) mm in diameter

including leaves, or gradually tapering to (7-) 1 0-

15 mm in diameter in terminal, densely sporan-

giate divisions. Stems excluding leaves 0.5-1 mm

thick at the base, slightly tapering upward, almost

straight to somewhat flexuous, pale greenish in life.

Leaves of basal divisions spirally arranged, single,

or in occasional pairs or whorls of 3, not predom-

inantly whorled, forming about 6 indistinct lon-

gitudinal ranks, subdistant, soft-herbaceous, wide-

spreading to ascending, straight to slightly falcate,

usually with the lamina vertical due to a twist of

the lamina base, linear-lanceolate to lanceolate,

widest in the basal '/3 or '/4, distinctly narrowed

into a petiolelike, twisted, usually perpendicular

to deflexed lamina base, (10-) 13-24 mm long, 1-

5 mm wide, flat, or with slightly revolute, smooth

margins. Leaves of middle and terminal divisions

spirally arranged, paired or borne in irregular to

regular, alternating whorls of 3, conform, or usu-

ally narrower, (5-) 10- 15 (-20) mm long, (1-)1.5-

2(-3) mm wide, often with a long narrow apex

from a subauriculate, non-twisted lamina base.

Sporangia 1-1.5 mm wide.

Southern Mexico to Panama; West Indies;

northern South America.

Huperzia linifolia is a widespread and poly-

morphic species. Its variation is especially com-

plex in the northern Andes. The two varieties keyed

out below are fairly well characterized morpho-

logically and are geographically well defined.

Key to Varieties

a. Shortest sporophylls of densely sporangiate divisions with distinctly widened, usually not twisted

lamina base, usually ascending to loosely appressed, opaquely green; stem usually opaque-stramin-

eous, usually above 1000 m elevation 35a. var. tenuifolia

a. Shortest sporophylls of densely sporangiate divisions usually conform to leaves of basal divisions,

or shorter, usually with a twisted, narrow to slightly widened lamina base, perpendicular to spreading-

ascending, pale, transparently green or brownish green; stem, at least in terminal divisions, usually

transparently stramineous to reddish brown, the vascular tissue usually visible through the cortex,

usually below 600 m elevation 35b. var. jenmanii

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

43

35a. Huperzia linifolia var. tenuifolia (Nessel) B.

011g. in Harling and Andersson, Fl. Ecuador

33: 90. 1988.

Urostachys linifolius (L.) Herter var. tenuifolius Nes-

sel, Revista Sudamer. Bot. 6: 164. 1940 [Bar-

lappgewachse 162, Abb. 39, / 14. 1939, nom.

nud.]. LECTOTYPE (designated by 011gaard, in

Harling and Andersson, Fl. Ecuador 33: 90. 1 988):

Peru, Prov. Huanuco, Pampayacu, Poeppig

["Moritz 210 und 1155"] (BONN, Herb. Nessel

38 1\; isotype, G!).

Wet forests of the eastern Andine slopes, alt.

100-2700 m, Cajamarca, Amazonas, San Martin,

Huanuco, Pasco, Junin, Cuzco.

Colombia?, Ecuador and Peru.

Cajamarca: Tabillo, Jelski 1022 (KRA). Huancabamba,

Andre Kl 763 (F, GH, NY, us). Tambillo, Jelski 1022 (KRA).

Amazonas: Prov. Bongara, 5 km N of end of Lake Po-

macocha, on road to Rioja, 2000 m, Hutchison & Wright

6791 (F, GH, NY, uc). Prov. Chachapoyas, Cerros Calla

Calla, 10 km above Leimebamba, 2700 m, Hutchison &

Bennett 4743 (F, GH, M, MO, NY, uc, us). San Martin:

Tarapoto, monte Campana, Spruce 4627 (BR). Prov. Rio-

ja, Pedro Ruiz-Moyobamba road, km 390, Venceremos,

1750 m, D. Smith 4372 (AAU). Prov. Rioja, Pedro Ruiz-

Moyobamba road, km 29 1 , 1 850 m, Gentry et al. 45502

(AAU). Pasco: Prov. Oxapampa, Quebrada San Alberto,

on the borders of Parque Nacional Yanachaga-Chemi-

llen, Leon et al. 944 (F, USM). Junin: La Merced, Chan-

chamayo, Soukup 1007 (F); 1000 m, Esposto 11978 (USM).

Cuzco: Prov. La Convention, Potrero, Sapan Sachayoc,

2300 m, Vargas 13643 (GH). Valley of Santa Ana, 1000-

1 500 m, Herrera 2628 (us). Department unknown: Peru,

Ruiz & Pavon (E, G).

35b. Huperzia linifolia var. jenmanii (Underw. &

Lloyd) B. 011g. & Windisch, Bradea 5: 13.

1987.

Lycopodium jenmanii Underw. & Lloyd, Bull. Torrey

Bot. Club 33: 1 12. 1906. TYPE: Guyana, Monica

River, Jenman (holotype, NY!; isotypes, BONN,

Herb. Nessel 388 in part, E!).

Urostachys jenmanii (Underw. & Lloyd) Nessel, Bar-

lappgewachse 158. 1939.

Huperzia jenmanii (Underw. & Lloyd) Holub, Folia

Geobot. Phytotax. 20: 74. 1985.

Epiphytic in lowland rain forest up to ca. 600

m, San Martin.

Throughout the Amazonian basin and the

Guianas.

San Martin: Prov. Mariscal Caceres, Palo Blanco, W

of Puente, Tocache Nuevo, J. Schunke V. 5738 (F, NY,

s, us).

36. Huperzia wilsonii (Underw. & Lloyd) B. 011g.,

Opera Bot. 92: 170. 1987.

Lycopodium wilsonii Underw. & Lloyd, Bull. Torrey

Bot. Club 33: 111. 1906. TYPE: Puerto Rico,

Luquillo Mountains, Wilson 271 (holotype, NY!).

Lycopodium trichodendron Herter, Bot. Jahrb. 43:

Beibl. 98: 49. 1909. SYNTYPES: Guadeloupe,

L'Herminier (P!); Bory 103 (P!).

Lycopodium andinum Herter, Bot. Jahrb. 43, Beibl.

98: 49. 1909, not Rosenst. LECTOTYPE (des-

ignated by 011gaard in Harling & Andersson, Fl.

Ecuador 33: 97. 1988): Ecuador, Eraser, in Herb.

Drake (P!; isolectotype, G!).

Lycopodium lindavianum Herter, Hedwigia 49: 90.

1909, nom. nov. for L. andinum Herter, not Ro-

senst., and with the same type.

Lycopodium stamineum Maxon, Smithsonian Misc.

Coll. 56 (29): 2, pi. 2. 1912. TYPE: Panama, Chi-

riqui, above El Boquete, ca. 1 750 m, Maxon 5636

(holotype, us!).

Urostachys wilsonii (Underw. & Lloyd) Herter, Re-

pert. Spec. Nov. Regni Veg. 19: 163. 1923.

Lycopodium arcanum Maxon in Yuncker, Field. Mus.

Publ. Bot. 17: 310, t. 3. 1938. TYPE: Honduras,

Comayagua, above El Achote, above plains of

Siguatepeque, 1800 m, Yuncker et al. 6149 (ho-

lotype, us!).

Huperzia lindaviana (Herter) Holub, Folia Geobot.

Phytotax. 20: 74. 1985.

Plants erect, arcuate-spreading to pendulous, up

to 20(-30) cm long. Shoots homophyllous, 15-

2 5 (-30) mm in diameter including leaves, equally

thick throughout, or in some slender individuals

gradually tapering to 1—1.5 cm. Stems excluding

leaves (l-)1.5-2(-3) mm thick at the base, often

tapering to (0.7-)1 mm, prominently ridged by

decurrent leaf bases, pale green to stramineous,

often with bright red spots on leaf bases. Leaves

usually uniform throughout, borne in alternating,

irregular whorls of 6-7, in basal divisions, upward

often in whorls of 4-5, perpendicularly spreading

to ascending, straight to upward curved, usually

not twisted at the base, linear to filiform, (6-) 10-

1 7 mm long, 0.3-0.5 mm wide at the base, quickly

narrowed to ca. 0.2 mm due to involution, grad-

ually tapering, adaxially canaliculate to involute,

often with a prominent vein abaxially near the

base. Leaves of terminal divisions in old plants

sometimes gradually reduced to 6(-4) mm long.

Decurrent leaf bases usually not wider than the

lamina base, often bright red. Sporangia 1-1.5 mm

wide.

Epiphytic in lower montane forests, up to alt.

2750 m, San Martin.

West Indies; southern Mexico to Panama; An-

dean South America south to Peru.

44

FIELDIANA: BOTANY

Huperzia wilsonii is related to H. dichotoma and

H. mandiocana (Raddi) Trev. (Brazil) and to H.

polycarpos. With these it shares the bottlebrush-

like growth habit, and with H. mandiocana the

bright red coloration of the leaf bases in some

individuals. The absence or presence of red color

seems uncorrelated with other characters. The in-

tensity and size of the coloration are variable.

Pendent individuals of H. wilsonii are rather

similar to H. polycarpos, but the latter has shorter,

more flattened leaves and uniformly falcate leaves

due to a basal twist, and this species apparently is

always pendulous. The direction of leaves is cor-

related with growth habit. In erect-growing plants

the leaves are wide-spreading to almost reflexed,

while in pendent plants the leaves are somewhat

ascending.

San Martin: Prov. Rioja, Pedro Ruiz-Moyobamba

road, km 390, Venceremos, 1750 m, D. Smith 437 2 A

(MO). Prov. Mariscal Caceres, Rio Abiseo National Park,

trail La Playa camp-Papayas, 2650-2750 m, Young &

Leon 4968 (AAU). Department unknown: Ex herb. Pavon

(o, P). Peru, M. Matthews (G, K). Peru, Matthews 1082

(o).

37. Huperzia polycarpos (Kunze) B. 011g., Opera

Bot. 92: 169. 1987.

Lycopodium polycarpos Kunze, Linnaea 9: 5. 1834.

TYPE: Peru, ad Cassapi [Casapi?, Huanuco],

Poeppig (w!).

Urostachys polycarpos (Kunze) Herter, Index Lyco-

podiorum 76. 1949.

Urostachys cuatrecasasii Herter, Revista Sudamer. Bot.

10: 123. 1953. TYPE: Colombia, Valle, Cord.

Occidental, Vertiente Occidental, hoya del rio Di-

gua, Piedra de Moler, 900-1 180 m, Cuatrecasas

15143A (holotype, us!).

Lycopodium cuatrecasasii (Herter) Morton, Amer. Fem

J. 54: 72. 1964.

Plants pendulous, with flaccidly hanging divi-

sions, up to 30 cm long. Shoots homophyllous,

almost equally thick throughout, 10-15 mm in

diameter including the leaves. Stems excluding

leaves 1-1.5 mm thick near the base, tapering to

0.5-1 mm, somewhat ridged by decurrent leaf bas-

es. Leaves almost uniform throughout, densely

crowded, borne in alternating, irregular whorls of

6—7 in basal divisions, upward of (3-)4 whorls,

wide-spreading to ascending, often subsecund,

usually with vertical lamina due to a twist at the

base, linear to filiform, 6-10(-12) mm long, 0.4-

0.5 mm wide, evenly tapering, usually oblique-

falcately curved, adaxially flat or shallowly cana-

liculate, often with an abaxially prominent vein

near the leaf base, with green decurrent leaf bases.

Sporangia ca. 1 mm wide.

Pendulous epiphyte in montane forest, ca. 2000

m, Huanuco?, Junin.

Costa Rica to Colombia; Peru.

Huperzia polycarpos is close to H. wilsonii, which

see for discussion. From H. sarmentosa it is dis-

tinguished by the lack of leaf base auricles. From

H. mollicoma (Spring) Holub (tropical America),

it differs in the more patent, twisted leaves, with

much less prominent veins.

Junin: Chanchamayo Valley, 2000 m, C. Schunke 529

(F).

38. Huperzia sarmentosa (Spring) Trev., Atti Soc.

Ital. Sci. Nat. 17: 248. 1874.

Lycopodium sarmentosum Spring, Mem. Acad. roy.

Belg. 24 [Mon. Lye. 2]: 13. 1849. LECTOTYPE

(designated by 011gaard in Harling and Anders-

son, Fl. Ecuador 33: 100. 1988): Ecuador, Qui-

tonian Andes, Jameson 41 (K!).

Urostachys sarmentosus (Spring) Herter, Repert. Spec.

Nov. Regni Veg. 19: 165. 1923.

Plants slender, pendulous, with flaccidly hang-

ing divisions, up to 100(-200) cm long. Shoots

homophyllous or almost so, equally thick through-

out, 15-25 mm in diameter including leaves in

basal divisions, or sometimes tapering to ca. 8 mm

in diameter. Stems excluding leaves 1-1.5 mm

thick at the base, often tapering to 0.4-0.8 mm,

slightly to prominently ridged by decurrent leaf

bases. Leaves borne in alternating irregular whorls

of 4-7, spreading to ascending, or rarely loosely

appressed in terminal divisions, usually twisted at

the lamina base, straight to slightly upward curved,

linear to linear-subulate, with distinctly widened,

auriculate, usually strongly revolute, often over-

lapping lamina bases, 7-15 mm long, 0.4-0.7 mm

wide just above the auriculate base, usually ca. 0.3

mm wide in the middle, usually somewhat revo-

lute, with obscure to prominent vein, with smooth

margins, or the auricles irregularly dentate. Leaves

of terminal divisions often densely crowded, con-

form, or gradually reduced to 5-7 mm long. De-

current leaf bases narrower than the lamina base.

Sporangia 1-1.3 mm in diameter.

Epiphytic or rarely hanging from banks, in up-

per montane forest, alt. 2100-3450 m, Amazonas,

San Martin.

Ecuador and northern Peru.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

45

Huperzia sarmentosa is quite variable with re-

spect to leaf length. It resembles H. polycarpos,

and pendulous forms of//, wilsonii but differs from

these by its auriculate lamina base.

Amazonas: Chachapoyas, Leimebamba-Lajasbamba

trail, Boeke 2059 (MO, us, USM). San Martin: Prov. Maris-

cal Caceres, Rio Abiseo National Park, NW sector, Valle

de Chochos, 3450 m, Leon 2161 (AAU, USM).

39. Huperzia curvifolia (Kunze) Holub, Folia

Geobot. Phytotax. 20: 72. 1985.

Lycopodium curvifolium Kunze, Linnaea 9: 5. 1835.

TYPE: Peru (Dept. Huanuco), sylvar. densarum

arboribus prope Pampayaco (Pampayacu), 1829,

Poeppig(LZ, holotype, destroyed; isotypes, AWH!,

B!, BR!, E!, oi, K!, M!, NY!, wi).

Lycopodium tenue Willd. var. tenuissimum Spring,

Mem, Acad. roy. Belg. 24 [Mon. Lye. 2]: 2 1 . 1 849,

based on L. curvifolium Kunze and with the same

type.

Urostachys curvifolius (Kunze) Nessel, Barlappge-

wachse 129. 1939.

Plants delicate to extremely delicate, flaccidly

pendulous, up to 60 cm long. Shoots gradually

(rarely abruptly) heterophyllous, 2.5-6 mm in di-

ameter including the leaves in basal divisions, ta-

pering to 0.3-1.5(-2) mm in diameter including

leaves in terminal divisions. Stems excluding leaves

0.4-0.7 mm thick at the base, tapering to 0.2-0.4

mm upward, usually not concealed by leaves.

Leaves of basal divisions borne in alternating whorls

of 4-5, patently sigmoid to strongly falcately up-

ward curved, acicular-nliform, ong and widely de-

current, 2-4(-5) mm long, 0.2-0.3 mm wide at the

base, extremely delicate, abaxially convex, flat or

canaliculate adaxially. Vegetative leaves of ter-

minal constricted divisions alternate, or borne in

alternating irregular whorls of 3, closely appressed,

lanceolate to linear-lanceolate, somewhat clasping

with the base, 1.2-3.5 mm long, 0.2-0.6 mm wide

at the base, usually strongly involute, often sharply

carinate at the base, short to long decurrent. Spo-

rophylls usually few and scattered, often unilat-

eral, conform, or shorter and wider, lanceolate to

widely ovate, or subcordate and short to long acu-

minate, usually widely spreading, 1-3 mm long,

0.3-0.8 mm wide, sometimes scarcely exceeding

the sporangia, abaxially with a prominent vein, or

carinate. Sporangia 0.6-0.8 mm wide.

Epiphytic in lower, wet montane forest, alt. 400-

1750 m, San Martin, Ucayali, Huanuco.

Costa Rica to Peru.

Huperzia curvifolia is variable in size and com-

pactness, seemingly not only as a response to en-

vironmental factors. It is easy to recognize by the

strongly upward curved leaves of basal divisions

and the appressed non-sporangiate leaves of the

terminal divisions. The smallest forms with hair-

like terminal shoots represent the most fragile and

slender extreme known in the family. Only these

are represented in Peru. They correspond to the

type collection. Other, stronger forms to the north

in the Andes and in Central America approach

slender forms of H. acerosa and H. filiformis.

San Martin: Boqueron pass, 92 km from Tingo Maria

on road to Pucallpa, 400 m, Allard 22098 (us). Prov.

Rioja, Pedro Ruiz-Moyobamba road, km 390, Vencere-

mos, 1750 m, D. Smith 437 2A (USM). Ucayali (as Lo-

reto): Prov. Coronel Portillo, Boqueron Padre Abad, 400

m, J. Schunke V. 3046 (F, GH, us), 480 m, Hutchison et

al. 6054 (F, GH, uc, us, USM).

40. Huperzia tenuis (Willd.) Trev., Atti Soc. Ital.

Sci. Nat. 17: 248. 1874.

Lycopodium tenue Willd., Sp. pi. ed. 4, 5: 55. 1810.

TYPE: Ecuador, Valley of Vinajacu, near Loja,

Humboldt & Bonpland 3363 (holotype, B, Herb.

mild. 194231; isotypes, BM!, LG!, P!).

Urostachys tenuis (Willd.) Nessel, Arch. Bot. Est. S.

Paulo 1: 401. 1927.

Plants delicate to extremely delicate, flaccidly

penduolous, at least up to 75 cm long. Shoots grad-

ually heterophyllous, 3-8 mm in diameter includ-

ing the leaves in the basal divisions, tapering to

1-3 mm. Stems excluding leaves 0.5-0.7 mm thick

at the base, tapering to ca. 0.3 mm upward. Leaves

of basal divisions borne in irregular alternating

whorls of 3-5, or spirally arranged, forming 6-10

indistinct longitudinal ranks, usually densely

crowded, spreading to ascending, straight to some-

what curved, often secund, acicular, tapering from

a subauriculate base, 3-5(-7) mm long, 0.3-0.5

mm wide, decurrent. Vegetative leaves and spo-

rophylls of constricted terminal divisions almost

conform, borne in alternating irregular whorls of

3, or irregularly alternate, patently diverging from

the stem, with upward curved tips, widely ovate

to subhastate, 1-3 mm long, 0.5-1.4 mm wide,

clasping at base, tapering to abruptly narrowed

into a short to long acuminate apex, bluntly to

sharply carinate, usually unilaterally, discontinu-

ously, but often densely sporangiate. Sporangia

0.8-1.2 mm wide.

46

FIELDIANA: BOTANY

Usually epiphytic, in high montane forests, alt.

1350-3300 m, Amazonas, San Martin, Cuzco.

Costa Rica; Andes from Venezuela to Peru.

Huperzia tenuis exhibits the same type of size

variation as mentioned for H. curvifolia. Some

forms are as extremely fragile and slender but dif-

fer by the spreading leaves in terminal divisions.

The typical forms of the species are also ecologi-

cally distinct, being restricted to the uppermost

montane forests, while the extremely thin forms

(LI. Williams 7348 and Spruce 4621) appear to

occur only at lower altitudes.

Amazonas: Prov. Chachapoyas, Cerros Calla Calla, 1 8

km above Leimebamba on road to Balsas, 3000-3100

m, Hutchison & Wright 6932 (F, GH, MO, NY, uc, us,

USM); Wurdack 1762 (F, G, GH, K, NY, s, uc, USM). San

Martin: Tarapoto, Monte Campana, Spruce 4621 (BM,

BR, P). Prov. Mariscal Caceres, NW corner of Rio Abiseo

National Park, Puerta del Monte, 3100-3300 m, Young

& Leon 4458 (AAU). Prov. Mariscal Caceres, Chochos

valley, 3100 m, Young 2667 (AAU, USM). San Roque,

1 350-1 500 m, LI. Williams 7348 (F, us). Cuzco: Camino

a Huarcan, caqui Acobamba [Ocobamba], Valle Neapi-

llo, 2400 m, Biies 1376 (NY). Department unknown:

Poeppigin 1829 (MO). Lehmann 5021 (F, GH, us).

41. Huperzia molongensis (Herter) Holub, Folia

Geobot. Phytotax. 20: 75. 1985.

Lycopodium molongenseHerter, Bot. Jahrb. 43: Beibl.

98: 51. 1909. TYPE: "Molong," Pearce (holo-

type, K!) [erroneously as "Ostaustralische Pro-

vinz: Gebirgswalder Neu-Siid- Wales" in the pro-

tologue].

Urostachys molongensis (Herter) Nessel, Barlappge-

wachse240. 1939.

Plants robust to very robust, pendulous, at least

up to 1 50 cm long. Shoots heterophyllous, usually

not sharply differentiated, 20-35 mm in diameter

including the expanded leaves in basal divisions,

distally gradually to abruptly constricted to 3-8

mm in diameter including the shorter, imbricate

leaves in the terminal divisions. Stems excluding

leaves 2-3.5 mm thick at the base, tapering to 1-

1.5 mm. Expanded leaves of basal divisions borne

in alternating whorls of 3, patent to ascending, or

slightly recurved, lanceolate to lanceolate-ovate,

acute, 10-18 mm long, 3-6(-7) mm wide, flat, or

with slightly revolute margins, coriaceous, often

shining, usually twisted at the base. Leaves of con-

stricted divisions decussate or subdecussate, im-

bricate, discontinuously sporangiate, ovate to sub-

cordate, acute, sharply carinate and clasping with

the base, usually conduplicate in the apex, usually

shining, (2.5-)3-10 mm long, 3-6 mm wide. Spo-

rangia 1.5-2.5 mm in diameter.

Usually epiphytic, in montane, wet cloud forest

on the eastern Andean slopes, 3100-3550 m,

Amazonas, San Martin.

Andes of Venezuela, south to northern Peru.

This species is easily distinguished because of

its large size and the sharply quadrangular ter-

minal divisions. The expanded leaves are some-

times developed only along a very short portion

of the stem base and are sometimes lacking in

herbarium specimens.

Amazonas: Prov. Chachapoyas, Leimebamba, road to

Balsas, 3240 m, Luteyn 11370 (NY). Cerros Calla Calla,

3450-3550 m, Wurdack 1707 (F, GH, NY, us); 3360 m,

Hutchison & Wright 6998 (c, E, F, G, GH, K, M, MO, NY,

s, uc, us). San Martin: Prov. Mariscal Caceres, NW

corner of Rio Abiseo National Park, trail to Mirador,

3 100-3300 m, Young& Leon 4457 (AAU). Prov. Mariscal

Caceres, Rio Abiseo National Park, Puerta de Monte,

3400 m, Leon & Young 1308 (AAU). Prov. Mariscal Ca-

ceres, Rio Abiseo National Park, Chochos Valley, 3100

m, Young 2620 (AAU).

42. Huperzia campiana B. 011g., in Harling and

Andersson, Flora of Ecuador 33: 109. 1988.

TYPE: Ecuador, Prov. Loja, E of Nudo de

Cajanuma, 0llgaard et al. 57861 (holotype,

AAU!; isotype, QCA!).

Plants rather robust, pendulous, at least up to

1 m long. Shoots heterophyllous, the basal divi-

sions ca. (15-)20-30 mm in diameter including

the expanded leaves, the distal quadrangular di-

visions abruptly constricted to 2-4 mm thick in-

cluding the imbricate, reduced leaves. Stems ex-

cluding leaves 2-4 mm thick at the base, tapering

to 0.5-1.5 mm, pale greenish or brownish. Ex-

panded leaves of basal divisions usually uniform

throughout, borne in alternating whorls of 3,

spreading to perpendicular, lanceolate to widely

lanceolate-ovate, usually widest below the middle,

(7.5-)10-15 mm long, 2.5-4(-4.5) mm wide, often

somewhat shorter near the contraction of the shoot,

twisted at the base, with flat to revolute margins,

herbaceous to subcoriaceous, dull to somewhat

shining. Leaves of terminal constricted divisions

decussate, or rarely in alternating whorls of 3 near

the base of constricted divisions, usually sporan-

giate throughout, appressed and clasping with their

bases, bluntly to sharply carinate, widely ovate to

triangular-ovate, short-acuminate or mucronu-

late, 2-3 mm long, 1.5-2 mm wide, the sporo-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

47

phylls equalling or slightly exceeding the sporan-

gia. Sporangia 1.2-2 mm in diameter.

Epiphytic in upper cloud forest and elfin forest,

alt. 2400-3500 m, San Martin, Huanuco.

Ecuador; Peru.

Bryan 779 deviates from typical Huperzia cam-

piana by the relatively wide expanded leaves and

slightly thicker constricted divisions.

San Martin: Prov. Mariscal Caceres, Rio Abiseo Na-

tional Park, NW sector, Valle de Chochos, 3450 m, Leon

2179 (AAU). Huanuco: Cushi, trail to Tambo de Vaca,

Bryan 7 19 A (F, G, us).

stricted divisions and is tentatively referred to this

species.

Amazonas: Leimebamba, 2100 m, Woytkowski 7749

(F, MO, NY, USM).

44. Huperzia ericifolia (Presl) Holub, Folia Geo-

bot. Phytotax. 20: 72. 1985.

Lycopodium ericaefolium Presl, Reliq. haenk. 1: 77.

1825. TYPE: "Peruvia, verosimiliter Luzon,"

Haenke (holotype, PRC!; isotype, BR!).

Urostachys phlegmaria (L.) Herter var. ericaefolius

(Presl) Nessel, Barlappgewachse 217. 1939.

43. Huperzia heteroclita (Poiret) Holub, Folia

Geobot. Phytotax. 20: 73. 1985.

Lycopodium heteroclitum Poiret, in Lam., Encycl. 3:

554. 1814 [1813]. LECTOTYPE (designated by

011gaard in Harling and Andersson, Fl. Ecuador

33: 1 12. 1988): America calidiore, Herbier de A.

N. Desvaux (P!).

Plants pendulous, lax, at least up to 80 cm (pos-

sibly to 400 cm) long. Shoots heterophyllous, with

expanded, perpendicular leaves, often only in the

first 5-20 cm of the basal divisions, upward

abruptly constricted, sharply to bluntly quadran-

gular, 2-3 mm thick including the small, imbricate

leaves, the constricted divisions often interrupted

by short zones with expanded leaves. Stems ex-

cluding leaves 1-1.7 mm thick at the base, tapering

to 0.7-1 mm, often bright red. Expanded leaves

decussate, or subdecussate, perpendicular to the

stem, lanceolate to widely lanceolate, widest in the

middle, acute, 6-9 mm long, 2-3.5 mm wide, soft-

herbaceous, usually twisted at the base. Leaves of

constricted divisions decussate, closely imbricate,

usually discontinuously sporangiate, ovate-cor-

date, acute, bluntly to sharply carinate to condu-

plicate in the apex, 1.5-4 mm long, 2-2.5 mm

wide, dull, sometimes with shallowly erose, or in-

distinctly dentate margins. Sporangia ca. 1.5 mm

wide.

Epiphytic in upper wet montane forest, ca. 2 100

m, Amazonas.

Southern Ecuador and northern Peru.

The species is usually recognizable by its long,

sharply quadrangular constricted shoots with ex-

tensive non-sporangiate zones and sharply cari-

nate, apically conduplicate leaves. However, the

specimen cited below deviates from the typical in

the terete rather than sharply quadrangular con-

Plants slender, pendulous, at least up to 6 cm

long. Shoots heterophyllous, in the basal divisions

ca. 8-14 mm in diameter including the expanded

leaves, then abruptly constricted to 1.5-2 mm in-

cluding the imbricate, reduced leaves of the qua-

drangular, terminal divisions. Stems excluding

leaves ca. 1 mm thick, tapering to ca. 0.5 mm,

pale greenish. Expanded leaves usually uniform in

position, shape and size throughout, borne in al-

ternating whorls of 3, or decussate upward, almost

continuously overlapping throughout (pressed

specimens), widely lanceolate to oblong-lanceo-

late, acute to obtuse and mucronate, 7-9 mm long,

2-3 mm wide, with slightly revolute margins, the

lamina usually twisted at the base to a vertical

position. Leaves of terminal, constricted divisions

decussate, discontinuously sporangiate, appressed

and clasping with their bases, carinate to condu-

plicate in the apex, lanceolate to ovate- or trian-

gular-ovate, short to long acuminate, 2-4 mm long,

1.5-2 mm wide, the sporangiate leaves 1.5-2.5x

longer than the sporangia. Sporangia 1-1.5 mm

wide.

Wet lower montane forest, alt. 625-2040 m, San

Martin, Huanuco.

Ecuador to Bolivia.

Huperzia ericifolia is closely related to H. di-

chaeoides (Maxon) Holub (Central America to Ec-

uador) and H. aqualupiana (West Indies, N Ven-

ezuela to NW Colombia). Its distinctness from the

latter is doubtful, although H. ericifolia can usually

be distinguished by its narrower, more acute ex-

panded leaves in whorls of three, while H. aqualu-

piana usually has ovate, subacute to obtuse, dec-

ussate expanded leaves.

Problems of typification were discussed by 011-

gaard in Harling and Andersson, Fl. Ecuador 33:

113. 1988.

48

FIELDIANA: BOTANY

San Martin: Prov. Rioja, Pedro Ruiz-Moyobamba

road, km 390, Venceremos, 2040 m, Smith & Vdsquez

4589 (AAV, USM). Jingo Maria, 625-1 100 m, Allard20935

(us). Huanuco: SW slope of Rio Llullapichis watershed,

ascent of Cerros del Sira, 1290 m, Dudley 13029 (GH,

us).

45. Huperzia myrsinites (Lam.) Trev., Atti Soc.

Ital. Sci. Nat. 17: 249. 1874.

Lycopodium myrsinites Lam., Encycl. Meth. Bot. 3:

654. 1789. TYPE: Dominican Republic, "S. Do-

mingue," Comm. Joseph Martin, (holotype, P,

Herb. Lam.l).

Urostachys myrsinites (Lam.) Herter, Repert. Spec.

Nov. Regni Veg. 19: 166. 1923.

Lycopodium skutchii Maxon, Proc. Biol. Soc. Wash.

46: 159 (1933). TYPE: Guatemala, Chimaltenan-

go, Chichavac, alt. 2400-2700 m, Skutch 243 (ho-

lotype, us!).

Plants pendulous, slender, at least up to 65 cm

long. Shoots heterophyllous, usually not all sharp-

ly differentiated, the basal divisions 10-18 mm in

diameter including the expanded leaves, distal di-

visions gradually or abruptly constricted to 1.5-3

mm thick including the reduced, imbricate leaves.

Stem excluding leaves ca. 1 mm thick at the base,

tapering to ca. 0.5 mm, pale greenish or reddish.

Expanded leaves of basal divisions decussate or

subdecussate, often irregularly shaped, subdistant

to densely crowded and somewhat overlapping,

ascending to perpendicular, ovate-lanceolate or

lanceolate, acute, usually the widest ones with a

rounded base, 6-11 mm long, 1.5-3 mm wide,

usually flat, straight to somewhat recurved. Leaves

of terminal constricted divisions highly variable,

often with complete reduction series, and with re-

current series to expanded shape, decussate or sub-

decussate, continuously or discontinuously spo-

rangiate. Transitional leaves with widely ovate

base, and short to long acuminate apex, appressed

and clasping, with the wide base abaxially rounded

to bluntly carinate, with straight to recurved apex,

2.5-5 mm long, 1.5-2 mm wide. Shortest leaves

with base conform, but with straight to falcate

apex, bluntly to sharply carinate, scarcely exceed-

ing the sporangia, ca. 2 mm long. Sporangia 1-1.3

mm in diameter.

Epiphytic in montane forest, ca. 2500 m, Ca-

jamarca.

Hispaniola; Central America; Trinidad; Guy-

ana; Venezuela to Peru.

For illustration of the species, see A. R. Smith

(Pteridophytes, in Breedlove: Flora of Chiapas,

part 2: 347,f.82a-b. 1981).

Huperzia myrsinites is closely related to H. phy-

licifolia. Species of the group of Huperzia quad-

rifariata (Bory) Rothm. (SE Brazil) were errone-

ously placed under H. myrsinites by several earlier

authors.

Cajamarca: Prov. Hualgayoc, Hda. Taulis, 2500 m,

Hutchison & Bismarck 6391 (F, NY, us). Prov. Contu-

maza, Bosque de Cachil, 2450 m, Sagdstegui 14873 (AAV).

46. Huperzia phylicifolia (Poiret) Holub, Folia

Geobot. Phytotax. 20: 75. 1985.

Lycopodium phylicifolium Poiret, in Lam., Encycl. 3:

546. 1814 [1813]. LECTOTYPE (designated by

011gaard in Harling and Andersson, Fl. Ecuador

33: 117. 1988): "Habitat in Chili," Herbier de A.

N. Desvaux, Donnee par Mme Vve Lavallee en

1896(p!).

Urostachys phylicifolius (Poiret) Nessel, Barlappge-

wachse 246. 1939.

Lycopodium congestifolium Spring, Mem. Acad. roy.

Belg. 15 [Mon. Lye. 1]: 70. 1842. TYPE: Peruvia,

Dombey, H. M. P., Herb. Deless. (P!).

Lycopodium nubigenum Herzog, Meded. Rijks-Herb.

27: 2. 1915. TYPE: Bolivia, Comarapa, Herzog

1967 (holotype, L!; isotypes, B!, z!).

Urostachys nubigenus (Herzog) Nessel, Barlappge-

wachse 246. 1939.

Plants slender, pendulous, up to 150 cm long.

Shoots heterophyllous, in the basal divisions ca.

10-20(-25) mm in diameter including the ex-

panded leaves, then abruptly (rarely gradually)

constricted to (l-)1.5-2(-2.5) mm in diameter in-

cluding the imbricate, reduced leaves. Stems ex-

cluding leaves 0.7-1.2 mm thick at the base, up-

ward tapering to ca. 0.5 mm, greenish to bright

red. Expanded leaves of basal divisions borne in

alternating whorls of 3, or decussate, subdecussate,

or alternate, usually widely spaced in alternate-

leaved stem portions, perpendicular to the stem

to falcately ascending, lanceolate to linear-lanceo-

late, widest at or below the middle, (4-)6-10(-13)

mm long, (1-)1 .5-2 mm wide, softly to firmly her-

baceous, with flat to slightly revolute margins, the

lamina twisted to a vertical position. Expanded

leaves near the constriction often sporangiate.

Leaves of constricted terminal divisions decussate,

or subdecussate, continuously or discontinuously

sporangiate, appressed and clasping with their bas-

es, abaxially rounded to carinate, widely lanceo-

late to widely ovate or subcordate, acute to mu-

cronate or cuspidate, rarely with an elongate flat

apex, 1.7-2(-4) mm long (in gradually hetero-

phyllous shoots sometimes longer), 1-1.5 mm wide,

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

49

equalling to more than twice as long as the spo-

rangia. Sporangia 1.2-1.5 mm wide.

Epiphytic or rarely rupestral in upper montane

and elfin forest, 2300-3700 m, Cajamarca, Ama-

zonas, Huanuco, Huancavelica, Apurimac, Cuzco,

Puno.

Due to uncertain delimitation of the species the

total distribution of this species is unsettled. In an

inclusive sense it is wide-ranging: from Costa Rica,

along the Andes south to northern Argentina, and

SE Brazil.

Huperzia phylicifolia occurs in a wide range of

epiphytic and rupestral habitats with highly vari-

able exposure and humidity. Its growth habit (i.e.,

size, leaf size, texture and crowding, and stem col-

or), responds strongly to the environmental con-

ditions. It is closely related to H. myrsinites, H.

subulata, H. biformis (Hooker) Holub (Brazil), and

H. erythrocaulon (Fee) Holub (Brasil). Forms of

the species corresponding to the type of Lycopo-

dium nubigenum, with expanded leaves slightly

longer and widest just above the base, are preva-

lent in the southern part of the range (Peru, Bo-

livia, Argentina).

Cajamarca: Prov. Cutervo, Jelski 1013 (p). Amazonas:

Chachapoyas, Matthews 1081 (G, K, p). Huanuco: Cani,

NE of Mito, 2800 m, Macbride 3546 (F, us). Cushi, trail

to Tambo de Vaca, Bryan 719B (F, GH, us). Huancaveli-

ca: Prov. Tayacaja, Marcavalle, between Huachocolpa

and Tintay, 2700 m, Tovar 4751 (GH, USM). Apurimac:

Prov. Abancay, Ampay, 3400-3700 m, Vargas 8388 (MO).

Cuzco: Prov. La Convencion, Dist. Vilcabamba, Yu-

paanqui, 2700 m, Davis et al. 1210 (AAV, F, GH). Prov.

Paucartambo, Paso de Aguila, Pillawata, 2450 m, Vargas

22995 (GH). Puno: Prov. Sandia, 2300 m, Weberbauer

633 (G, MOL). Prov. Carabaya, Ayapata-Kahuallayoc,

2600-3600 m, Vargas 10741 (GH). Tabina, Lechler2023

(us).

47. Huperzia subulata (Poiret) Holub, Folia Geo-

bot. Phytotax. 20: 77. 1985.

Lycopodium subulatum Poiret, in Lam., Encycl. 3:

544. 1814 [1813]. LECTOTYPE (designated by

Ollgaard in Harling and Andersson, Fl. Ecuador

33: 1 19. 1988): HerbierdeA. N. Desvaux, Donnee

par Mme Vve Lavallee en 1896 (P!).

Urostachys subulatus (Poiret) Nessel, Arch. Hot. Est.

S. Paulo 1: 420. 1927.

Urostachys ewanii Herter, Revista Sudamer. Bot. 10:

126-127. 1953. TYPE: Colombia, drainage of

upper Rio Pascual, above Cordoba, 2840 m, Ewan

16526 (holotype, us!; isotype, NO!).

Lycopodium ewanii (Herter) Morton, Amer. Fern J.

54: 72. 1964.

Plants slender, pendulous, up to 2 m long. Shoots

heterophyllous, ca. 15-25 mm in diameter in-

cluding the expanded leaves in the basal divisions,

abruptly constricted to 1.5-2 mm in diameter in-

cluding the reduced and appressed leaves of the

terminal divisions. Stems excluding leaves ca. 1

mm thick at the base, upward tapering to less than

0.5 mm, pale greenish to bright red. Expanded

leaves of basal divisions borne in alternating, often

irregular whorls of 3, or often upward subdecus-

sate, spreading to perpendicular to the stem, nar-

rowly lanceolate to linear-subulate, straight to

obliquely falcate-ascending, (6-) 10- 15 mm long,

0.5-1 mm wide, twisted at the base, flat, soft-her-

baceous. Leaves of terminal constricted divisions

subdecussate, or in alternating whorls of 3 just

above the constriction of the shoots, sporangiate

almost throughout, closely imbricate, widely ovate

with obtuse to acute apex, abaxially rounded to

carinate, 1.3-2 mm long, 1.3-1.5 mm wide,

equalling or slightly exceeding the sporangia. Spo-

rangia ca. 1 mm wide.

Epiphytic, in cloud forest and elfin forest, alt.

1 500-3400 m, Amazonas, San Martin, Huanuco,

Cuzco.

Costa Rica; Colombia to Peru.

Huperzia subulata is closely related to H. phy-

licifolia but seems to tolerate a narrower range of

growth conditions, being restricted to the most

humid forests of the eastern slopes of the Andes.

Amazonas: Prov. Bagua, Serrania de Bagua, ca. 12-18

trail km E of La Peca, 1 800-1 950 m, Gentry et al. 22904

(AAV, MO, USM). San Martin: Prov. Mariscal Caceres,

Rio Abiseo National Park, Puerta del Monte, 3400 m,

Leon & Young 1340 (AAU). Huanuco: Pampayacu, Hda.

at mouth of Rio Chinchao, Bryan 725 (F, G, us). Cuzco:

Prov. La Convencion, Cord. Vilcabamba, Hda. Lu-

isiana-Rio Apurimac, 3250 m, Dudley 11 105 (F, GH, us,

USM). Prov. La Convencion, Chontapampa, Valle San

Miguel, 1500 m, Bties 2987 (GH).

48. Huperzia cuneifolia (Hieron.) Holub, Folia

Geobot. Phytotax. 20: 72. 1985.

Lycopodium cuneifolium Hieron., Bot. Jahrb. Syst. 34:

572. 1905. LECTOTYPE (designated by Nessel,

Barlappgewachse 252. 1939): Costa Rica, Volcan

Barba, Hoffman 50 (B!).

Urostachys cuneifolius (Hieron.) Nessel, Barlappge-

wachse 252. 1939.

Plants pendulous, delicate, up to 60 cm long.

Shoots heterophyllous, in the basal divisions (5-)7-

1 3 mm in diameter including the expanded leaves,

50

FIELDIANA: BOTANY

then abruptly constricted to 1-2 mm in diameter

including the reduced, imbricate leaves in the ter-

minal, constricted, quadrangular divisions. Stems

excluding leaves 1 mm thick, or less, at the base,

tapering to ca. 0.5 mm, greenish to bright red.

Expanded leaves of basal divisions decussate,

spreading to perpendicular or slightly reflexed,

spathulate to lanceolate or lanceolate-obovate,

usually widest above the middle, obtuse or mu-

cronulate, 2-6.5 mm long, 1.5-3 mm wide, flat,

the lamina twisted to a vertical position, softly to

firmly herbaceous. Leaves of terminal constricted

divisions decussate, often sporangiate in rather

short zones of the divisions, appressed and clasp-

ing with their bases, widely ovate to subcordate,

1.5-2 mm long, 1-1.5 mm wide, carinate, with

falcate to reflexed mucronulate tips, the sporo-

phylls equalling or slightly exceeding the sporan-

gia. Sporangia ca. 1 mm wide.

Epiphytic or occasionally on banks in upper

montane forest and elfin forest, ca. 3000-3100 m,

Amazonas and San Martin.

Costa Rica; Andean Venezuela; Colombia and

Peru.

Amazonas: Prov. Chachapoyas, Cerros Calla Calla, 19

km above Leimebamba, 3100 m, Hutchison & Wright

6943 (F, GH, K, M, P, us); Wurdack 1744 (F, GH, NY, us,

USM). San Martin: Prov. Mariscal Caceres, NW sector

of Rio Abiseo National Park, near Mirador, 3000-3100

m, Leon 27/9(AAu).

49. Huperzia pruinosa (Herter) Holub, Folia Geo-

bot. Phytotax. 20: 76. 1985.

Lycopodium pruinosum Herter, Bot. Jahrb. Syst. 43,

Beibl. 98: 52. 1909. TYPE: Peru, Dept. Ama-

zonas, Prov. Chachapoyas, Tambo Ventillas,

2400-2600 m, Weberbauer (err. as Ule in pro-

tologue) 4410 (holotype, B!; isotypes, BONN, Herb.

Nessel 62 l\,o\).

Lycopodium durissimum Herter, Bot. Jahrb. Syst. 43,

Beibl. 98: 52. 1909. TYPE: Peru, "Voyage a

FEquateur et au Perou, 1876-77, Guayaba mars

1877" (err. "Colombia, Guayabal" in proto-

logue), Vidal-Seneze ["Senege"] (holotype, P!; iso-

type, BONN, Herb. Nessel 6221).

Urostachys pruinosus (Herter) Nessel, Arch. Bot. Sao

Paulo 1:420. 1927.

Urostachys durissimus (Herter) Nessel, Barlappge-

wachse 240. 1939.

Huperzia durissima (Herter) Holub, Folia Geobot.

Phytotax. 20: 72. 1985.

Plants erect and distally recurved, unbranched

or sparsely branched at base, distally with densely

tassel like ramification, at least up to 70 cm long.

Shoots heterophyllous, the basal divisions with

expanded leaves ca. 10-18 mm in diameter in-

cluding leaves, distally abruptly constricted to ca.

2-2.5 mm including the reduced, imbricate leaves.

Stems excluding leaves up to 5 mm thick at the

base, upward tapering to ca. 0.5-1.5 cm, rigid,

dark brown to purplish brown. Expanded leaves

of basal divisions borne in alternating, irregular,

distant whorls of 4, the whorls 6-9 mm apart,

ascending to spreading or sharply reflexed, lan-

ceolate, straight or recurved, (6-)8-10 mm long,

2.5-3 mm wide, acute, flat, coriaceous, with

smooth, revolute margins, often adaxially con-

cave. Leaves of recurved, constricted divisions sub-

decussate, densely crowded, sporangiate almost

throughout, imbricate, widely ovate with obtuse

to acute apex, abaxially rounded to carinate, 1.2-

1.6 mm long and wide, equalling or slightly ex-

ceeding the sporangia. Sporangia 1-1.3 mm wide.

Indicated as terrestrial scandent shrub, in ceja

scrub (protologue) and as an epiphyte in high Swie-

tenia trees (Nessel, in Hoehne, Fl. Bras. 11: 88.

1955).

Endemic, Amazonas and San Martin.

The two taxonomically synonymous basionyms

were published simultaneously. The epithet prui-

nosa is maintained due to the quality of its type

material and greater certainty of its origin.

San Martin: Mount Organero, ceja, 1900 m, Melin 96

(s). "Tarapoto, 1908, G. Huebner" (BONN, Herb Nessel

6211).

Comments

Huperzia polyclada (Sodiro) Rolleri and Deferrari,

Notas Mus. La Plata, Bot. 21(1 00): 156.1988.

Lycopodium polycladum Sodiro, Crypt, vase. Quit.

561.1893. TYPE: Ecuador, Mojanda, Sodiro (not

located).

The type locality in Ecuador contains several

taxa of the Huperzia crassa group. Due to the ab-

sence of authoritative material and the relatively

scarce information of the protologue it is not pos-

sible to certify to which of these taxa the name

applies. However, none of the taxa correspond to

the Peruvian material that was placed in this spe-

cies by Rolleri (1980). This material is here re-

ferred to H. darwiniana.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

51

II. Lycopodium

Lycopodium L., Sp. pi. 1100. 1753. TYPE: Ly-

copodium clavatum L. Figure 7.

Lepidotis Mirbel, in Lam. & Mirbel, Hist. nat. veg. 3:

477. 1802. TYPE: Lycopodium clavatum L.

Lycopodium subgen. Lepidotis Baker, Handb. Fern-

Allies 8. 1887. TYPE: Lycopodium clavatum L.

Lycopodium subgen. Rhopalostachya Pritzel, Nat.

Pflanzenfam. 1 (4): 601. 1901. TYPE: Lycopo-

dium clavatum L.

Diphasiastrum Holub, Preslia 47: 104. 1975. TYPE:

Lycopodium complanatum L.

Diphasium Rothm., Feddes Repert. Spec. Nov. Regni

Veg. 54: 64. 1944. TYPE: Diphasium jussiaei

(Poiret) Rothm. (= Lycopodium jussiaei Poiret).

Austrolycopodium Holub, Folia Geobot. Phytotax. 26:

87, 9 1 . 1 99 1 . TYPE: Lepidotis magellanica Beauv.

(= Lycopodium magellanicum (Beauv.) Sw.).

Plants anisotomously branched, with elongate,

indeterminate, subterranean, creeping, or scan-

dent, plectostelic main stems (rhizomes), which,

in a dorsolateral position, give rise to usually de-

terminate, ascending to erect, dendroid or spread-

ing, repeatedly inclinate-bilaterally branched,

branchlet systems. Roots emerging directly along

the underside of main stems, with plectostelic main

roots. Branchlet leaves uniform or anisophyllous.

Strobili erect, simple or forked, sessile or borne

on simple or forked peduncles. Sporoph ylls peltate

without a basal mucilage cavity, or subpeltate with

a thin basal decurrent wing and with a basal mu-

cilage cavity. Sporangia attached to the sporophyll

base, reniform, with a short thick stalk, isovalvate

or slightly anisovalvate, their epidermis cells with

thin, lignified, sinuate side walls, without partial

thickenings. Spores reticulate. Gametophytes ob-

conic to convoluted disc-shaped, subterranean,

mycoparasitic, lacking pluricellular uniseriate tri-

chomes among the gametangia.

The generic description and synonymy above

includes only the Neotropical representatives. For

a fuller general treatment of the genus, see 011gaard

(1987).

The genus Lycopodium as here circumscribed

has perhaps 40 species mainly in temperate regions

of both the Northern and Southern Hemispheres,

with eight species in the Neotropics and five in

Peru, all of these, except L. vestitum, with wide

distributions. It includes several of the genera of

Holub (1975, 1983, 1985, 1991), also advocated

by Wagner and Beitel (1992). These genera cor-

respond to the sections in 011gaard (1987, 1989).

Four of the nine sections are represented in the

Neotropics. These sections are very distinct groups

of species. None of them are connected by inter-

mediate species or by intersectional hybrids.

Reference

WILCE, J. H. 1965. Section Complanata of the

genus Lycopodium. Beih. Nova Hedw. 19: 1-IX,

1-233, t. 1-40.

Key to Species of Lycopodium

a. Branchlets radially symmetrical, isophyllous b

b. Branchlet leaves with a colorless hair tip, or a colorless membranous apex; sporophylls subpeltate,

with a basiscopic, membranous wing on the stalk (sect. Lycopodium) c

c. Branchlet leaves hair-tipped, or with a short membranous apex; branchlets green; strobili sessile

or borne on short or long, simple or branched peduncles 1 . L. clavatum

c. Apical half or more of branchlet leaves colorless, membranous; branchlets grayish to silvery

white; strobili sessile or borne on short, simple, indistinct peduncles 2. L. vestitum

b. Branchlet leaves green throughout, without membranous or hairlike tips; sporophylls peltate, with

a slender, terete, wingless stalk (sect. Magellanica) 3. L. magellanicum

a. Branchlets dorsiventral, flattened, anisophyllous, with dimorphic or trimorphic leaves d

d. Branchlet leaves decussate, arranged in 4 ranks, with widened lateral leaves, and 1 rank of dorsal

and 1 rank of ventral, median narrow leaves (sect. Complanata) 5. L. thyoides

d. Branchlet leaves alternate, with 2 ranks of of wide, dorsolateral leaves, and ca. 3 indistinct ranks

of narrow, ventral, apically membranous leaves (sect. Diphasium) 4. L. jussiaei

FIG. 7. Lycopodium clavatum: a, aerial shoot, habit. Lycopodium thyoides: b, aerial shoot, habit; c, tip of branchlet,

adaxial side; d, tip of branchlet, abaxial side. (Adapted from Stolze, Ferns and fern allies of Guatemala, 1983.)

52

FIELDIANA: BOTANY

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

53

1. Lycopodium clavatum L., Sp. pi. 1101. 1753.

LECTOTYPE: Herb. Burser XX: 49 (UPS, des-

ignated by Jonsell & Jarvis, Regnum Veg. 1 27:

63. 1993).

Lepidotis clavata (L.) Beauv., Prod. Aetheogamie 108.

1805.

Plants creeping, trailing, or hanging over banks.

Main stem usually above ground with ascending

to stiffly erect, up to at least 50 cm tall, repeatedly

unequally branched aerial shoots, with strongly

diverging to almost parallel branchlets. Ultimate

branchlets terete. Leaves borne in low alternating

spirals or whorls of 6-8(-10), patent to ascending

or imbricate, linear-acicular, 6-8(-10) mm long

and 0.5-0.8 mm wide, terminating in a long col-

orless trichome or short membranous apex, with

smooth to sparsely denticulate margins. Strobili

sessile or pedunculate. Peduncles, if present, erect,

up to 30 cm long, simple, or branched and bearing

up to 6 pedicellate strobili. Strobili 1.5-6(-8) cm

long, ca. 6 mm in diameter including sporophylls,

sometimes forked. Sporophylls borne in alternat-

ing whorls of 5-6, subpeltate, with triangular-ovate

to rhombic-ovate, acuminate exterior face, with

usually widely scarious, dentate to erose-laciniate

margins. Sporangia 1.3-1.6 mm wide, their epi-

dermal cells with strongly sinuate side walls with

pocket-like in- and evaginations. Spores reticulate

on all faces.

Almost cosmopolitan, in humid temperate and

boreal regions of the Northern Hemisphere and

on tropical mountains in the Old and New World.

The species is highly variable and adaptive to ex-

ternal factors. The discovery of tetraploids in Ja-

pan indicates that genetic variation is also in-

volved. There is as yet no indication of polyploidy

in the Neotropical representatives of the species.

Lycopodium clavatum varies almost continu-

ously from amply branched plants with diverging

branches and spreading, soft leaves, and long-

branched peduncles, growing in moist, warm, shel-

tered habitats, to small, compact, parallel-branched

plants with more imbricate and firm leaves, and

lacking, or short, simple or once forked peduncles,

belonging to cold, exposed habitats. The latter

forms are here recognized as ssp. contiguum. Cor-

responding monostachyous forms are found in the

Arctic and high mountains of New Guinea. The

two subspecies recognized here are often consid-

ered to be distinct species. However, there are many

intermediate forms that cannot be placed in one

or the other taxon with certainty. These inter-

mediates form normal spores and have normal

meiosis. Also there are rare examples of single

individuals showing characters of one subspecies

in the growth of one year and of the other the

following year. As the two forms are usually rec-

ognizable and are ecologically rather distinct, I

treat them as subspecies.

Key to Subspecies of Lycopodium clavatum

a. Strobili rarely forked, borne on elongate, simple or branched peduncles; aerial shoots amply branched,

with more or less diverging branchlets; sporophylls short to long acuminate .... 1 a. ssp. clavatum

a. Strobili sessile, or borne on short, simple, often indistinct peduncles, frequently forked; aerial shoots

sparsely branched, usually stiffly erect, with parallel branchlets; sporophylls usually with a long, wide,

scarious apex Ib. ssp. contiguum

la. Lycopodium clavatum ssp. clavatum

Lycopodium aristatum Willd., in L., Sp. pi. ed. 4, 5:

17. 1810. TYPE: Venezuela, Silla de Caracas,

Humboldt & Bonpland (holotype, B, Herb. Willd.

79557!).

Lycopodium piliferum Raddi, PI. Bras. nov. gen. 1 :

79, t. 3. 1825. Based on L. aristatum Willd. and

with the same type.

Lycopodium trichophyllum Desv., Mem. Soc. Linn.

Paris 6: 184. 1827 [as trychophyllum]. LECTO-

TYPE (designated by 011gaard in Harling and An-

dersson, Fl. Ecuador 33: 125. 1988): "Habitat in

Brasilia, no. 41 in herb. deA. N. Desvaux, Donnee

par Mme Vve Lavallee en 1895" (P!).

Lycopodium clavatum L. var. aristatum (Willd.) Spring,

Flora 21 (1): 173. 1838.

Lycopodium eriostachys Fee, Crypt. Vase. Bresil 1:

224. 1869. TYPE: Brasilia fluminensi, Serra dos

Orgaos, Glaziou 1788 (BR!, c!, P!, RB!).

Clearings exposed habitats in montane forest,

700-3200 m, Piura, Cajamarca, Amazonas, La

Libertad, San Martin, Loreto, Huanuco, Pasco,

Junin, Ucayali, Ayacucho, Cuzco, Puno.

54

FIELDIANA: BOTANY

Humid temperate and boreal regions of the

Northern Hemisphere, montane in the tropics.

Absent from Australia.

For more complete synonymy, see 011gaard

(1988).

Cajamarca: Prov. Cutervo, Cutervo-Socota, 2320 m,

Lopez & Sagdstegui 5331 (F, MO). Prov. Hualgayoc,

Culquirrumi, 2448 m, Sanchez Vega 2099 (AAU). Ama-

zonas: Prov. Chachapoyas, Pipus, between Chachapoyas

and Molinopampa, 2100 m, Sanchez Vega et al. 2180

(AAU). La Libertad: Prov. Bolivar, above Longotea, 2800

m, Sagdstegui 14189(r). San Martin: Prov. Rioja, Pedro

Ruiz-Moyobamba road, km 390-394, Venceremos,

1910-2040 m, D. Smith 4524 (AAU). Zepelacio, near

Moyobamba, 1 1 00-1 600 m, King 3360 (MO, s, us). Tara-

poto, Spruce 4732 (BR, E, G, p, us). Huanuco: Prov. Leon-

cio Prado, Dist. Rupa Rupa, near Tingo Maria, 700 m,

King 337 (F). Prov. Huanuco, Dist. Churubamba, Santo

Toribio, 1500 m, Mexia 8144 (F, MO, s, uc, us). Pasco:

Prov. Oxapampa, Cord. San Gutardo, 2600 m, Leon 512

(AAU). Cord. Yanachaga, 1 2 km E of Oxapampa-Villa

Rica road, 2100-2200 m, Gentry & Smith 35922 (AAU).

Junin: Prov. Tarma, Chanchamayo Valley, above La

Merced at Cumbre Yacunay, 2000 m, Hutchison 1 194

(F, M, s, uc, us). Huacapistana, Killip & Smith 24255

(us). Ucayali: Prov. Coronel Portillo, La Divisoria, ca.

20 km NNE of Tingo Maria on road to Pucallpa, 1 600

m, Dillon 2641 (F). Ayacucho: Prov. La Mar, eastern

Massif of Cord. Central opposing the Cord. Vilcabamba,

between Tambo, San Miguel, Ayna and Hda. Luisiana,

1 300 m, Dudley 1 1 953 (F). Cearapa, Killip & Smith 22459

(us). Cuzco: Valle del Urubamba, Machu Picchu, 2400

m, Herrera 3197 (c, F). Tres Cruces, Gentry et al. 23458

(us). Puno: Below Limbani, 3100-3200 m, Brandbyge

523 (AAU). At Lake Titicaca near Huancano, 2000 m,

Charpin AC- 12704(0).

Ib. Lycopodium clavatum ssp. contiguum

(Klotzsch) B. 011g. in Harling & Andersson,

Flora of Ecuador 33: 126. 1988.

Lycopodium contiguum Klotzch, Linnaea 18: 519.

1844. LECTOTYPE (designated by 011gaard in

Harling & Andersson, Flora of Ecuador 33: 126.

1988): Ecuador, Paramo de Tiopullo [between

Cotopaxi and Illiniza], Hartweg 1474 (B!; isolec-

totypes, BM!, GL!, K.!, NY!, P!).

Lycopodium vestitum Poiret, var. herbaceum Spring,

Mem. Acad. roy. Belg. 24 [Mon. Lye. 2]: 45. 1 849.

TYPE: Colombia, Sierra Nevada, Moritz (holo-

type, B!; isotype, LG!).

Lycopodium clavatum L. var. pseudo-contiguum

Christ, Primitiae Fl. Costar. 3 (1): 55. 1901. TYPE:

Costa Rica, Cerro de las Vueltas, 3000 m, versant

W du massif de Buena Vista, Pittier 10467 (ho-

lotype?, BR!; isotype?, P!).

Lycopodium herbaceum (Spring) Hieron., Bot. Jahrb.

Syst. 34: 575. 1905.

Clearings and exposed habitats in uppermost

montane forest and in grass paramo, frequent in

disturbed paramo, 2600-3800 m, Piura, Caja-

marca, Amazonas, San Martin, Ancash, Pasco,

Huancavelica, Apurimac, Cuzco.

Costa Rica; Andes from Venezuela to Peru.

For more complete synonymy, see 011gaard

(1988).

Piura: Prov. Huancabamba, Cuello del Indio, 2800 m,

Lopez et al. 8883 (AAU). Cajamarca: Prov. San Miguel,

near El Tingo (Agua Blanca), 3100 m, Mostacero et al.

1130 (AAU). Cutervo, Jelski 1016 (us). Amazonas: Prov.

Chachapoyas, Saullamur-Calla Calla, 3250 m, Sanchez

Vega et al. 2136 (AAU). San Martin: Prov. Mariscal Ca-

ceres, NW corner of Rio Abiseo National Park, 3500 m,

Young & Leon 4712 (AAU). Ancash: Prov. Carhuaz,

Huascaran National Park, Quebrada Ishinca, 3900 m,

D. Smith et al. 9588 A (AAU). Prov. Yungay, Huascaran

National Park, Quebrada Ranincuray, 3850 m, D. Smith

et al. 9067 (USM). Pasco: Prov. Oxapampa, Dist. Huan-

cabamba; Sta. Barbara, above Lanturachi, 3300-3500

m, Foster et al. 10398 (AAU). Huancavelica: Prov. Tay-

acaja, Montepungo, 5 km E of Surcubamba, 300 m,

Stork & Horton 10386 (F). Prov. Tayacaja, vicinity of

Huachocolpa, 3000-3100 m, Tovar 3950 (USM). Apu-

rimac: Prov. Abancay, Cerro Turronmocco, NW of Nev-

ado Ampay, 3500 m, Brandbyge 422 (AAU). Cuzco: Prov.

Urubamba, Antakillqa, 3300 m, Franquemont 285 (F).

2. Lycopodium vestitum Poiret, in Lam., Encycl.

3: 546. 1814. LECTOTYPE (designated by

011gaard in Harling & Andersson, Flora of

Ecuador 33: 128. 1988): Ecuador (Prov. Loja),

prope Loxam, Humboldt (p-Humboldt!; iso-

types, B, Herb. Willd. 194281, BR!, LG!, us!).

Lycopodium scariosum Hooker, Icon. PI. 1 : /. 89. 1 836,

not Forst. TYPE: Peru, Casapi, Matthews 1765

(holotype, K!; isotype, IVY!).

Lycopodium albidum Baker, J. Bot. 25: 37. 1887.

TYPE: Ecuador, Prov. Loja, Mataba 1883, Hu-

beck (holotype, K!).

Rhizome creeping, scrambling or hanging over

banks, usually above ground, 3-5 mm in diameter

including leaves. Aerial shoot systems stiffly erect,

sparsely branched, with almost parallel branches,

silvery whitish. Branchlet leaves appressed, linear-

lanceolate, 5-9 mm long, 0.6-0.8 mm wide,

herbaceous in less than half of their length, with

colorless, translucent, membranous, slightly

widened, coarsely erose-laciniate apex. Strobili

sessile or borne on a short, simple, indistinct pe-

duncle. Sporophylls borne in alternating whorls of

5-6, subpeltate, 5-7 mm long, 1.5-2 mm wide,

narrowly triangular-ovate, with a long membra-

nous apex, with widely membranous, erose-den-

tate to laciniate margins. Sporangia 1.2-2 mm

wide, their epidermis cells with strongly sinuate

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

55

side walls with pocketlike evaginations. Spores re-

ticulate on all faces.

Clearings, road banks, and exposed sites in up-

per montane forest, and in the lower paramos,

2400-3500 m, Piura, Cajamarca, Amazonas, San

Martin.

Southern Ecuador and northern Peru.

Lycopodium vestitwn is closely related to L. cla-

vatum, especially to ssp. contiguum. In spite of the

striking appearance of this species, it is only doubt-

fully distinct from L. clavatum. Individuals with

intermediate characters between L. vestitwn and

both subspecies are frequent. These have normal

meiosis and normal spores.

Piura: Prov. Huancabamba, Cord. Chinguela (Sapa-

lache-El Carmen), 2900 m, Sagdstegui et al. 10223 (AAU).

Above Huancabamba, 3400-3500 m, Weberbauer 6145

(F, GH, us). Cajamarca: Prov. Cutervo, Fortaleza de Santa

Cruz, San Andres, 2400 m, Llatas & Suarez 2866 (F).

Cutervo, Jelski 990 (p), 994, 1007 (KRA). Amazonas: Prov.

Bagua, Cord. Colan E of La Peca, 3200 m, Barbour 3261

(AAU). Prov. Chachapoyas, Cerros Calla Calla, E side,

19 km above Leimebamba on road to Balsas, 3100 m,

Hutchison & Wright 5545 (F, GH, MO, P, uc, us, USM).

San Martin: Rio Abiseo National Park, Mariscal Ca-

ceres, NW corner of the Park, 3300 m, Young 27 54 (AAU,

USM). Rio Abiseo National park, Mariscal Caceres, La-

guna del Eco, Chochos, 3450 m, Young & Leon 4901

(USM).

3. Lycopodium magellanicum (Beauv.) Sw., Syn.

Fil. 180. 1806.

Lepidotis magellanica Beauv., Prod. Aetheogamie 102.

1805. TYPE: Fretum magellanicum, unknown

collector (holotype presumably in G, Herb. De-

lessen n.v.\ isotype, P!, Herb. Palisot de Beauvois,

comm. M. de Jussieu).

Lycopodium spurium Willd., Sp. pi. ed. 4, 5: 28. 1810.

TYPE: Ecuador (Prov. Tungurahua), Quito, in

vulcano Tungurahua, Humboldt (holotype, B,

Herb. Willd. 193641).

Lycopodium pichinchense Hooker, Icon. pi. 1: /. 55.

1837. LECTOTYPE (designated by B. 011gaard,

Biol. Skr. Dan. Vid. Selsk. 34: 62. 1989): Ecuador,

Pichincha, 10,000 ft, Col. Hall (38?) (K!).

Austrolycopodium magellanicum (Beauv.) Holub, Fo-

lia Geobot. Phytotax. 26: 91. 1991.

Rhizome subterranean or occasionally above

ground, 1-2 mm thick excluding leaves. Aerial

shoots ca. 1 mm thick excluding leaves at origin,

up to 25 cm long excluding peduncles and strobili,

repeatedly unequally branched, with spreading to

ascending branchlets. Branchlets 4-7 mm in di-

ameter including leaves. Branchlet leaves acicular,

flattened, smooth, with a long, pointed, non-pilif-

erous apex, 3-5 mm long, 0.6-0.8 mm wide.

Strobili sessile, or terminating somewhat indis-

tinct, simple or up to twice-forked peduncles, 3-

4 mm thick, 2-10(-l 5) cm long, often forked. Spo-

rophylls borne in alternating whorls of 4, peltate,

with a slender, terete, wingless stalk, with widely

ovate, short to long acuminate exterior face at-

tached to the stalk near the center, with narrowly

scarious, almost smooth to shallowly erose-den-

ticulate margins. Sporangia ca. 2 mm wide; spor-

anguium epidermis with evenly sinuate side walls.

Spores with unornamented proximal faces, with

an irregular, fine-meshed reticulum on distal faces.

Creeping or hanging over banks in grass paramo,

usually in relatively dry paramos, but also often

in mossy bogs, 2750-4000 m, Lambayeque, Ca-

jamarca, La Libertad, San Martin, Ancash, Huan-

uco, Cuzco.

Hispaniola; Costa Rica; Andes from Venezuela

to Chile and Argentina.

The subterranean rhizomes seem to enable this

species to survive in vegetation that is burned fre-

quently.

Lycopodium magellanicum is possibly identical

to certain forms included in L. fastigiatum R. Br.

of New Zealand and Australia.

Lambayeque: Ferranafe, Laguna Tembladera, Inca-

huasi, 31 50 m, Sagdstegui et al. 12791 in part (F). Prov.

Ferranafe, ca. 7 km NW of Incahuasi, near Cerro Puna-

machay, 3300-3550 m, Dillon & Skillman 4125 (F). Ca-

jamarca: Cajabamba-Luchubamba, 3800 m, Sagdstegui

et al. 11202 (AAU). San Miguel, Millan (El Tingo-Taulis),

3000 m, Sagdstegui 9526 (NY). Cutervo, Jelski 993, 1012,

1017 (KRA). La Libertad: Bolivar, Laguna de los Ichus,

3600 m, Lopez & Sagdstegui 3239 (GH). San Martin:

Prov. Mariscal Caceres, Parque Nacional Rio Abiseo,

Paredones, 3650 m, Leon & Young 1606 (USM). Prov.

Mariscal Caceres, Parque Nacional Rio Abiseo, Cho-

chos, 3400 m, Young & Leon 4896B (USM). Ancash:

Prov. Carhuaz, Huascaran National Park, Quebrada Ish-

inca, 3900 m, D. Smith et al. 9588B (AAU). Prov. Huari,

Huascaran National Park, Quebrada Pachachaca, 3700-

3860 m, D. Smith et al. 12547 (USM). Huanuco: Yanano,

2000 m, Macbride 4928 (F). Cuzco: Paucartambo, 3700

m, Vargas 13938 (GH). Paucartambo, Parque Nacional

Manu, Cerro Macho Cruz, 3200 m, Leon & Aguilar 2345

(USM).

4. Lycopodium jussiaei Poiret, in Lam., Encycl.

3: 543. 1814. TYPE: Perou (holotype, P, Herb.

Jussieu 6581).

Lycopodium jussiaei Poiret var. microphyllum Poiret,

in Lam., Encycl. 3: 543. 1814. TYPE: Amerique

meridionale, Herb. Bonpland (holotype, P!).

Lycopodium haenkei Presl, Reliq. haenk. 1: 78. 1825.

56

FIELDIANA: BOTANY

TYPE: Peru (Dept. Huanuco), In montosis Pe-

ruviae ad Huanocco, Haenke (holotype, PRC!;

possible isotype, K!).

Lycopodium heterophyllum Sprengel, Syst. Veg. ed.

16, 4: 13. 1827. TYPE: Peruvia, Humboldt (ho-

lotype, B, Herb. Willd. 194251).

Lycopodium scariosum Forst. var.jussieui(Poirei) Ba-

ker, Handbook Fern Allies 29. 1887.

Diphasium jussiaei (Poiret) [Presl ex] Rothm., Feddes

Repert. Spec. Nov. Regni Veg. 54: 65. 1944 [as

jussieui].

Rhizomes creeping, scrambling to scandent, rig-

id, 2-4 mm thick, usually above ground, with leaves

radially arranged, uniform, 3-5 mm long, ca. 1

mm wide, linear-lanceolate, with widely membra-

nous, irregularly obtuse apex. Aerial shoot systems

5-75 cm tall, branched almost from the base, in

large individuals with a main axis almost conform

to the rhizome, upward gradually changing to an-

isophyllous, bearing alternating, fan-shaped

branchlet systems. Branchlets dorsiventral, ani-

sophyllous, flattened, 4-6(-8) mm wide including

leaves, with 2 dorsolateral ranks of wide leaves

and 2-3 indistinct ventral ranks of narrow leaves.

Dorsolateral leaves flattened in the plane of the

branchlet system, or inclinate to it, obliquely el-

liptic, with the acroscopic margin 2-3 mm long,

1—1.5 mm wide, forward and ventrally curved,

mucronate to short hair-tipped, short to long de-

current. Ventral leaves appressed, lanceolate-su-

bulate, with membranous apices. Peduncles up to

50 cm long (rarely absent), simple or up to twice-

forked, bearing 1-3 strobili, terete. Strobili (l-)3-

10 cm long, 4—6 mm in diameter including spo-

rophylls. Sporophylls borne in alternating whorls

of 4, subpeltate, with ovate, more or less acumi-

nate exterior face, 4-6 mm long, ca. 2 mm wide,

with narrowly membranous, shallowly erose-den-

ticulate margins. Sporangia 1.5-2 mm wide, the

side walls of epidermis cells evenly sinuate and

finely curled. Spores reticulate, with large, regular

meshes on distal faces, and unornamented prox-

imal faces.

Usually a vigorous scrambling to scandent plant,

common in clearings and on road banks and open

places in montane forest, 1700-3700 m, Piura,

Cajamarca, Amazonas, San Martin, Huanuco,

Pasco, Huancavelica, Ayacucho, Cuzco, Puno.

Jamaica; Dominican Republic; Costa Rica;

Venezuela; Andes south to Bolivia; Brazil (Ita-

tiaia).

A variable species, apparently due to environ-

mental factors. Lycopodium jussiaei is closely re-

lated to L. scariosum (Indonesia to New Zealand)

and to plants from Chile and Juan Fernandez rec-

ognized as L. gayanum Remy. The plants referred

to Lycopodium jussiaei are generally larger and

coarser than those species. A modern revision of

the group is needed. For more complete synony-

my, see 011gaard (1988).

Piura: Huancabamba, Loma Redonda (Sapalache-

Chinguela), 2400 m, Sagdstegui et al. 10203 (AAU). Ca-

jamarca: Prov. Cutervo, 10 km NW of Socota, 3200 m,

Stork & Horton 10143 (F, G, MO). Cutervo, Jelski 1025

(KRA). Amazonas: Prov. Bagua, Cord. Colan, E of La

Peca, 3200 m, Barbour 3262 (AAU, MO, USM). Prov. Cha-

chapoyas, Puma-urcu SE of Chachapoyas, 3100-3200

m, Wurdack 1158 (F, s, uc). San Martin: Prov. Mariscal

Caceres, NW corner of Rio Abiseo National Park, 3300

m, Young & Leon 4882 (AAU). Huanuco: Prov. Huanuco,

Carpish Pass, 3300 m, Hodge 6291 (AAU, F). Churubam-

ba, Pampa Hermosa, 1400 m, Mexia 8146 (F, o, MO, s).

Pasco: Cord. Yanachaga, 1 2 km E of main Oxapampa-

Villa Rica Road, 2100-2200 m, Gentry & Smith 35924

(AAU, MO). Oxapampa, 1970 m, Smith & Pretel 1645

(AAU). Huancavelica: Prov. Tayacaja, Montepungo, 5 km

E of Surcubamba, 3000 m, Stork & Horton 10384 (F).

Ayacucho: Prov. La Mar, eastern Massif of Cord. Central

opposing the Cord. Vilcabamba between Tambo San

Miguel, Ayna and Hda. Luisiana, Dudley 1 1 973 (F). Cuz-

co: Prov. Paucartambo, Valle de Pilcopata, Patria-Pil-

lahuata, 2000 m, Foster & Wachter 7475 (AAU). Puno:

Below Limbani, 3100-3200 m, Brandbyge 558 (AAU).

5. Lycopodium thyoides Willd., Sp. pi. ed. 4, 5:

18.1810. TYPE: Venezuela, Silla de Caracas,

Humboldt (holotype, B, Herb. Willd. 193521).

Lycopodium complanatum L. var. tropicum Spring,

in Mart., Flora Bras. 1 (2): 1 16. 1840.

Lycopodium complanatum L. var. thyoides (Willd.)

Christ, in Schwacke, PI. Nov. Mineiras 2: 42.

1900 [as thujoides].

Lycopodium complanatum L. var. validum Weath-

erby, Proc. Amer. Acad. Arts Sci. 45: 414. 1910,

based on L. thyoides Willd. and with the same

type.

Diphasiastrum thyoides (Willd.) Holub, Preslia 47: 108.

1975.

Plants with creeping, trailing to scandent rhi-

zomes, usually above ground, or hanging over

banks. Rhizomes terete, 1.2-2.5 mm in diameter

excluding leaves. Rhizome leaves relatively dis-

tant, borne in irregular spirals, or subverticillate,

subulate, appressed to ascending. Aerial shoots as-

cending to erect, 10-50 cm tall, with vegetative

portions up to ca. 30 cm tall. Main upright axis

terete to somewhat flattened, bearing lateral, flat-

tened, fan-shaped branchlet systems. Ultimate

branchlets flattened, dorsiventral, anisophyllous,

1.5-3 mm wide including leaves, with trimorphic,

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

57

decussate leaves. Upper, median branchlet leaves

with pointed, subulate to acicular, appressed, 1-2

mm long, free blades, and a conspicuous, ca. 0.4-

0.6 mm wide, prominently decurrent base. Lateral

branchlet leaves bilaterally compressed, long-de-

current, 2.5-7 mm long including bases, the free

blades 1-3 mm long, appressed to spreading, acu-

minate to long-pointed, the leaf bases 0.6-1 .5 mm

wide, with almost parallel to distinctly diverging

margins, often curved down. Ventral leaves in-

conspicuous, acicular, without decurrent base, 1-

2 mm long. Peduncles 10-25 cm long, terete, bear-

ing 4-9 pedicellate strobili. Strobili 1 .5-5 cm long,

2-4 mm in diameter including sporophylls, often

with protracted sterile tips. Sporophylls usually

borne in alternating whorls of 3, subpeltate, widely

deltoid-ovate, long-cuspidate, ca. 2-3 mm long,

1.5-2 mm wide, with almost entire, widely mem-

branous margins. Sporangia 1.5-2 mm wide, with

side walls of epidermal cells evenly sinuate. Spores

densely reticulate on all faces.

Clearings, road banks, open habitats, and sec-

ondary scrub in upper montane forest, and in the

lowest paramos, alt. 2000-3400 m, Piura, Lam-

bayeque, Cajamarca, Amazonas, La Libertad, San

Martin, Ancash, Huanuco, Pasco, Junin, Huanca-

velica, Ayacucho, Cuzco, Puno.

Throughout moist mountainous regions of trop-

ical America, south to northern Argentina.

The present application of the name Lycopo-

dium thyoides corresponds to the "L. thyoides-

complex" of Wilce (1965). I have not attempted

to treat the infraspecific variation. The Peruvian

material referred to this species is highly variable

and may belong to more than one species, but also

external factors apparently greatly affect the growth

habit of the individuals. Ferreyra 15111 (USM),

from Dept. Cajamarca, Prov. Celendin, Celedin,

ca. 3100 m, has unusually wide branchlets with

very wide lateral leaves.

For more complete synonymy, see 011gaard

(1988).

Cajamarca: Prov. Chota, Chota Tacabamba road, km

14, 2800 m, Smith & Vdsquez 3549 (AAU, F). Prov. San

Miguel, El Tingo, 3100 m, Mostacero et al. 1123 (AAU,

F). Amazonas: Prov. Chachapoyas, Leimebamba-Calla

Calla road, km 12-15, 2960-3100 m, Smith & Vdsquez

4992 (AAU). Donile-Cohechan, Soukup 4160 (F, us). La

Libertad: Prov. Pataz, Cerro Colpar, above Yalen, 3300-

3700 m, Young 3046 (AAU, USM). Prov. Bolivar, above

Longotea, 2800 m, Sagdstegui 14195 (F). San Martin:

Rio Abiseo National Park, NW corner, 2800 m, Young

1480 (AAU); Rio Abiseo National Park, Laguna del Eco,

Chochos, 3450 m, Young & Leon 4902 (USM). Ancash:

Prov. Yungay, Huascaran National Park, Quebrada

Ranincuray, 3650-3900 m, D. Smith et al. 10336 (AAU,

USM). Huanuco: Prov. Huanuco, Huanuco-La Union road

km 32, 2940-3100 m, Huapalla 2201 (AAU). Carpish

Pass, 2850 m, Asplund 12748 (s). Pasco: Above Oxa-

pampa (Cerro Corporation), ca. 2300 m, collector un-

known (F). Junin: Huacapistana, 1 800-2400 m, Killip &

Smith 24255 (F). Huancavelica: Prov. Tayacaja, Mar-

cavalle, between Huachocolpa and Tintay, 2800 m, 70-

var 4765 (USM). Ayacucho: Ccarrapa, between Huanta

and Rio Apurimac, 2500 m, Killip & Smith 22293 (F,

us). Cuzco: Prov. Calca, below Kachin, 2700-3800 m,

Sallo ex Franquemont 244 (AAU, F). Prov. Paucartambo,

Marcachea, Achirani, 2500-3000 m, Vargas 11133 (F,

MO, uc). Paucartambo, Woyt kowski 6744 (MO, us). Puno:

Below Limbani, 3100-3200 m, Brandbyge 546 (AAU).

Prov. Sandia, between Sandia and Cuyo-Cuyo, 3100-

3300 m, Ferreyra 16813 (USM). Tabina, Lechler 2034 (E,

G, s).

III. Lycopodiella

Lycopodiella Holub, Preslia 36: 22. 1964. TYPE:

Lycopodiella inundata (L.) Holub. Figure 8.

Lycopodium subgen. Cernuistachys Herter, Bot. Jahrb.

43: Beibl. 98: 29. 1909. TYPE: Lycopodium cern-

uum L.

Palhinhaea Vascon. & Franco, Bol. Soc. Brot. II, 41:

24 (1 967). TYPE: Palhinhaea cernua (L.) Vascon.

& Franco (= Lycopodium cernuum L.).

Lycopodium sect. Caroliniana Bruce, Amer. Fern J.

66: 136. 1976. TYPE: Lycopodium carolinianumL.

Pseudolycopodiella Holub, Folia Geobot. Phytotax.

18:441. 1983.

Lepidotis auct., not Mirbel 1802.

Lycopodium subgen. Lepidotis auct., not Sprengel,

Anleit. ed. 2, 2, 1: 108. 1817, nor Baker 1887.

Piura: Prov. Huancabamba, Dist. Sondor, Cerro La

Viuda, 2170 m, Sagdstegui et al. 8197 (AAU, MO). Piura:

Loma Redonda, 2400 m, Sagdstegui 10175 (AAU). Lam-

bayeque: Prov. Ferranafe, Incahuasi (Sinchigual-Laguna

Tembladera, 3000 m, Sagdstegui et al. 12870 (AAU, F).

Plants with prostrate, rooting, indeterminate,

isophyllous or anisophyllous, horizontally branch-

ing shoots, and dorsally arising, erect, simple

strobiliferous branches (sect. Lycopodiella and sect.

Caroliniana), or with trailing to arching or looping

FIG. 8. Lycopodiella caroliniana var. meridionale: a, habit; b, portion of stem and branch. Lycopodiella cernua:

c, habit; d, ultimate branchlet, showing sterile leaves and strobilus; e, apex of strobilus. (Adapted from Stolze, Ferns

and fern allies of Guatemala, 1983.)

58

FIELDIANA: BOTANY

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

59

indeterminate runner shoots that root with usually

long intervals, and branch occasionally in the hor-

izontal plane giving off horizontal branchlet sys-

tems, and usually bearing 1 vertical, dorsally aris-

ing, dendroid branch system on each loop (sect.

Campylostachys). Dendroid branch system with a

series of subdecussate, spreading to hanging, fla-

bellate branchlet systems, which in turn may ter-

minate in sessile, nodding to pendulous strobili.

Sporophylls subpeltate, with a median basiscopic

wing, or with coalescent basal membranes that

almost enclose the sporangia (sect. Campylosta-

chys). Sporangia on the sporophyll base, or axil-

lary (sect. Lycopodielld), strongly anisovalvate, or

isovalvate (sect. Caroliniand), the epidermis cells

with thin, straight, non-lignified side walls, but

with lignified, nodular, or semiannular thicken-

ings. Spores rugate. Gametophytes green, tuberous

and lobed on the upper side, surface-living, with

mycorrhiza in the base.

Almost all moist temperate and tropical regions

of the world. Perhaps 40 species, the majority of

these in the Americas and probably nine in Peru,

all fairly widely distributed.

The generic description above covers only the

Andean representatives of the genus. Although

growth habit and morphological details are quite

diverse, features of branching, sporangium anat-

omy, spores, and gametophytes indicate that Ly-

copodiella is a natural entity.

In sect. Campylostachys, most species have erect,

dendroid branch systems arising from the dorsal

side of horizontal, looping main axes (e.g., Lyco-

podiella cernud), but in some species the corre-

sponding dendroid branch system is scandent,

hanging, or creeping. In these cases the dendroid

branch system can be distinguished from the hor-

izontal branch system by the orientation of lateral

branchlet systems on the main axes. In the hori-

zontal system they are distichous, all arranged in

the horizontal plane. In the dendroid branch sys-

tem the lateral branchlets are polystichous, ar-

ranged in an irregularly subdecussate manner.

Zimmer, in a review in Bot. Jahrb. Syst. 112:

414, 1991, claimed that Lepidotis is the correct

name for this genus as lectotypified by Rothmaler

(1944). However, Rothmaler's lectotypification

does not refer to the protologue of Lepidotis and

is superceded by the choice of Lycopodium cla-

vatum (Pichi Sermolli, Webbia 26: 145-149. 1971).

Key to Species of Lycopodiella

a. Strobili erect, terminating simple, erect branches that arise dorsally on the creeping stem b

b. Creeping shoots isophyllous or nearly so; leaves of the erect branch crowded, borne in alternating

whorls of 6 or more c

c. Sporangia subglobular, strongly anisovalvate, ca. 1 mm wide; Sporophylls and vegetative leaves

with the same color and texture; leaves of erect branch with flattened, appressed base (sect.

Caroliniand) 1. L. alopecuroides

c. Sporangia reniform, isovalvate, 1-1.5 mm wide; Sporophylls and vegetative leaves with different

color and texture; leaves of erect, branch strongly upward curved from a subterete, diverging

base (sect. Caroliniand) 3. L. contexta

b. Creeping shoots strongly anisophyllous, with wide and long ventrolateral leaves and narrow dorsal

leaves; leaves of the erect branch borne in distant, alternating whorls of 4 or 5 (sect.

Caroliniand) 2. L. caroliniana var. meridionalis

a. Strobili nodding to pendulous, terminating spreading to pendulous branchlets that are borne on

dendroid, erect or scandent, amply branched shoot systems (sect. Campylostachys) d

d. Branchlet leaves closely imbricate, flattened throughout, lanceolate, with fimbriate margins and

densely hairy leaf bases 9. L. riofrioi

d. Branchlet leaves spreading to ascending, subterete, angular or apically flattened, acicular to linear-

lanceolate, glabrous or hairy, not fimbriate e

e. Leaves of the main axes closely appressed, less than 4 mm long; stems usually densely hairy,

dendroid shoot system flexible, usually bending to hanging from an erect base

6. L. descendens

e. Leaves of main axes patent to ascending or reflexed or, if loosely appressed, more than 4 mm

long; stems glabrous or hairy f

f. Dendroid branch system usually indeterminate, up to 4 m long, scandent to creeping, or all

60

FIELDIANA: BOTANY

axes scandent to creeping; main axes robust; leaves of main axes coriaceous, apically flat-

tened, usually strongly upward curved and hook-shaped, with hairy leaf bases

7. L. glaucescens

Dendroid branch system determinate, vertical, not scandent or climbing g

g. Lateral branchlet systems of dendroid branch system recurved to long-pendulous; strobili

3-5 mm in diameter; sporophylls usually more than 3 mm long, with irregularly dentate

margins 8. L. pendulina

g. Lateral branchlet systems of dendroid branch system erect or spreading to horizontal and

distally nodding; strobili 2.5-3 mm in diameter, sporophylls rarely more than 2 mm long,

with coarsely dentate to erose-laciniate margins h

h. Branchlet systems divaricate to somewhat aggregate, patent to horizontal, usually with

gently recurved tips; strobilus-bearing branchlets softly nodding at tip 4. L. cernua

h. Branchlet systems stiffly ascending or suberect, often densely aggregated, pointed up-

ward, not recurved; strobilus-bearing branchlets usually sharply reflexed at tip, or only

the strobilus reflexed 5. L. camporum

1 . Lycopodiella alopecuroides (L.) ( 'ran li 1 1 . Amer.

Fern J. 71: 97. 1981.

Lycopodium alopecuroides L., Sp. pi. 1102. 1753.

TYPE: Eastern temperate North America, Vir-

ginia, Kalm (LINN 1257.7).

Lycopodium matthewsii Hooker, Icon. pi. 1: /. 26.

1836. TYPE: Peru, Bagasan, [Bagazan, Dept.

Amazonas] Matthews 1778 (holotype, K.!; isotype,

BM!).

Lepidotis alopecuroides (L.) Rothmaler, Feddes Re-

pert. Spec. Nov. Regni Veg. 54: 66. 1944.

Lycopodiella matthewsii (Hooker) Holub, Folia Geo-

bot. Phytotax. 18: 441. 1983.

Horizontal shoots creeping and firmly rooted

throughout, up to 25 cm long, sparsely and un-

equally branched, densely covered on all sides by

almost uniform, somewhat upwardly secund, or

spreading to perpendicular leaves, 7-12 mm wide

including leaves, bearing stiffly erect, dorsally aris-

ing, simple, strobiliferous branches. Leaves linear-

lanceolate to subulate, flat, with smooth to den-

ticulate-ciliate margins, soft, light green, 5-8 mm

long, 0.5-1 mm wide. Erect branches ca. 5-8 mm

in diameter including leaves, up to 30 cm tall in-

cluding the strobilus, with radially arranged leaves

borne in alternating whorls of 6 or more, like leaves

of horizontal shoots, or slightly narrower, usually

more ascending. Strobili up to 12 cm long, 10-18

mm in diameter including sporophylls. Sporo-

phylls arranged as peduncle leaves, narrowly lan-

ceolate to lanceolate from a subpeltate base, not

coalescent at base, with few to many spreading to

hooked teeth on margins, 6-10 mm long, 1-1.7

mm wide at base. Sporangia axillary, subglobular,

ca. 1 mm wide, strongly anisovalvate, almost com-

pletely concealed by sporophyll bases, the epider-

mal cells with straight, unlignified side walls with

semiannular lignified thickenings. Spores rugate,

with a distinct equatorial rim.

On open, wet, peaty or sandy soil, in the jalca

or montane forest zone, 2400-3450 m, Cajamarca,

Amazonas, San Martin, Huanuco, Pasco.

Eastern temperate North America; Cuba;

throughout continental tropical America.

As here delimited, in the wide sense, Lycopo-

diella alopecuroides is a polymorphic species, con-

sisting of several elements. In the narrow sense,

the species is restricted to temperate North Amer-

ica and Cuba, while other elements are present in

South America. High altitude forms in the Andes

are relatively compact, wide-leaved, and robust

and perhaps deserve recognition as a species of

which the correct name is Lycopodiella matthews-

ii; the material cited below belongs to this form.

Cajamarca: Prov. Cutervo, El Suro, 2400 m, Sanchez

Vega et al. 6019 (AAU, F). Cumbemayo 10 km WSW of

Cajamarca, 3550 m, Molau & Sanchez Vega 840 (GB).

Amazonas: Prov. Chachapoyas, 1-5 km W of Molino-

pampa, 2400 m, Wurdack 1381 (F, GH, uc, us). Lei-

mebamba-Calla Calla, km 12-15, 2960-3100 m, Smith

& Vdsquez 5000 (AAU). Chachapoyas, 2400-2600 m,

Weberbauer 4397 (G). Prov. Bagua, Cord. Colan E of La

Peca, 3500 m, Barbour 3197, 3200 A (AAU, MO). San

Martin: Prov. Mariscal Caceres, Rio Abiseo National

Park, NW corner of the park, 3425m, Young 3475 (AAU);

3450 m, Leon 1854 (AAU). Huanuco: Tingo Maria, Mor-

row 11131 (GH). Pasco: Prov. Oxapampa, San Gotardo,

36 km W of Oxapampa, 2850 m, D. Smith 2753 (AAU).

2. Lycopodiella carol in iana (L.) Pichi-Sermolli var.

meridionalis (Underw. & Lloyd) B. 011g. &

Windisch, Bradea 5: 27. 1987.

Lycopodium meridionale Underw. & Lloyd, Bull. Torr.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

61

Bot. Club 33: 121. 1906. TYPE: Puerto Rico, dry

savannahs, Luquillo Mountains, Wilson 94 (ho-

lotype, NY!).

Lycopodium carolinianum L. var. meridionalis Nes-

sel. Arch. Bot. Est. S. Paulo 1: 431. 1927.

Pseudolycopodiella meridionalis (Underw. & Lloyd)

Holub, Folia Geobot. Phytotax. 18: 442. 1983.

Horizontal steins creeping and firmly rooted

throughout, to ca. 30 cm long, sparsely and un-

equally branched, anisophyllous, with long and

wide lateral leaves, and usually shorter and nar-

rower dorsal leaves, 7-1 2(-l 5) mm wide including

leaves, bearing 1 to few, dorsally arising, stiffly

erect, simple, strobilus-bearing branches. Lateral

leaves 3-5(-7) mm long, (l-)1.5-2.5(-3) mm wide

at the base, triangular-ovate to lanceolate, oblique-

ly spreading to falcately recurved, tapering into an

acute to long pointed apex. Dorsal leaves arranged

in (l-)2-4 longitudinal ranks (often on the same

individual), widely lanceolate to subulate, diverg-

ing to appressed, straight to upward curved,

(1.5-)3-4(-5) mm long, l-1.3(-2) mm wide at the

base. Erect branches up to 30 cm tall including

the strobilus, 1-1.5 mm thick excluding leaves,

bearing small, 3-5 mm long, acicular leaves in

remote, alternating spirals or irregular whorls of

4-5. Strobili up to at least 13 cm long, 3-5 mm

thick with appressed sporophylls. Sporophylls

borne in alternating whorls of 4 or 5, subpeltate,

rhombic or ovate-acuminate or ovate-cuspidate to

triangular-lanceolate, 3.5-6 mm long, (l-)1.5-2(-

2.5) mm wide, with entire to erose-denticulate,

minutely fimbriate-denticulate margins. Sporan-

gia reniform, isovalvate, borne on the sporophyll

stalk, ca. 1.5-2 mm wide. Spores rugate, with a

distinct equatorial rim.

Terrestrial or sometimes epilithic, usually on

wet ground, on road banks, slopes, ledges, grass-

lands in the jalca zone, on seepages, 1350-2400

m, Amazonas, San Martin.

Widely distributed in tropical America, rare in

the Andes.

The distinctness of the present taxon from the

type variety (temperate North America) has been

doubted (e.g., Proctor, 1977, pp. 32-33), because

the distinguishing characters usually indicated for

them are widely overlapping, and seem to be based

entirely on features related to the size of the plants.

Amazonas: Chachapoyas, 1-5 km W of Molinopampa,

2400 m, Wurdack 1382 (F, GH, NY, us, USM). San Martin:

San Roque, 1350-1500 m, LI. Williams 7763 (F, us).

3. Lycopodiella contexta (Mart.) Holub, Folia

Geobot. Phytotax. 18: 441. 1983.

Lycopodium contextum Mart., Icon. Crypt. Brasil. 38,

t. 20, f. 1. 1834. TYPE: In campis graminosis

apricis in summo monte Arara-Coara, fluvii Ja-

pura cataractae imminente, alt. ca. 1200 p., Mar-

tins (holotype, M!).

Lycopodium alopecuroides L. var. contextum (Mart.)

Baker, Handb. Fern Allies 19. 1887.

Lycopodium sprucei Baker, Handb. Fern Allies 24.

1887. TYPE: Venezuela, San Carlos del Rio Ne-

gro, Spruce 3151 (holotype, K!; isotype, BONN,

frag. Herb. Nessel 200\ in part).

Lycopodium alopecuroides L. subsp. contextum (Mart.)

Hassler, Trab. Inst. Bot. Farm. Buenos Aires 45:

92. 1928.

Lepidotis contexta (Mart.) Rothm., Feddes Repert.

Spec. Nov. Regni Veg. 54: 66. 1944.

Pseudolycopodiella contexta (Mart.) Holub, Folia

Geobot. Phytotax. 26: 93. 1991.

Horizontal shoots creeping and firmly rooted

throughout, up to at least 30 cm long, sparsely and

unequally branched, isophyllous, with somewhat

upwardly secund, ascending, acicular leaves, 3-7

mm in diameter including leaves, bearing stiffly

erect, dorsally arising, simple, or sometimes 1-2 x

forked, up to at least 40 cm tall, vegetative or

strobiliferous aerial shoots. Erect branches 4-6 mm

in diameter including leaves, densely foliose, iso-

phyllous. Leaves of erect branches borne in alter-

nating irregular whorls of 6-7, subterete at the base

(angular when dry), sometimes apically flattened,

strongly upward curved from a strongly diverging

to almost perpendicular base, 4-5(-7) mm long,

ca. 0.5 mm wide, with conspicuously acroscopical-

ly adnate and long decurrent leaf bases, with

smooth margins. Strobili up to 8 cm long, 3-4 mm

in diameter with appressed sporophylls. Sporo-

phylls borne in alternating whorls of 4-5, subpel-

tate, widely triangular-ovate to ovate-lanceolate

at base, with a long, narrow apex, 4.5-7 mm long,

1-1.5 mm wide, with subentire to slightly erose-

dentate margins at the base. Sporangia borne on

the sporophyll base, reniform, isovalvate, 1-1.5

mm wide.

Terrestrial, humid places on white sand, grass-

lands, and river margins, ca. 1 20 m, Loreto.

Circum-Amazonian, periphery of the Amazo-

nian lowland of Brazil, Peru, Colombia, and Ven-

ezuela.

Loreto: Manfinfa, on the upper Rio Nanay, Ll. Wil-

liams 1 106 (F, us); Vicinity of Iquitos, ca. 120 m, Revilla

4340 (F).

62

FIELDIANA: BOTANY

4. Lycopodiella cernua (L.) Pic.-Ser., Webbia 23:

165. 1968.

Lycopodium cernuum L., Sp. pi. 1103. 1753. TYPE:

LINN 1 257. 1 3, see Proctor, Ferns of Jamaica, Lon-

don 29. 1985.

Lepidotis cernua (L.) Beauv., Prodr. Aetheogamie 108.

1805.

Lycopodium cernuum var. capillaceum Spring, Mem.

Acad. roy. Belg. 15 [Mon. Lye. 1]: 80. 1842. TYPE:

Venezuela, Edo. Monagas, Guanaguana, Hum-

boldt 473 (holotype, B, Herb. Willd. 194291).

Lycopodium capillaceum (Spring) Hieron., Bot. Jahrb.

Syst. 34: 573. 1905.

Palhinhaea cernua (L.) Vase. & Franco, Bol. Soc. Brot.

41:25. 1967.

Plants with arching-looping runner shoots, with

up to 1 m tall, erect, dendroid branch systems,

these with several, subdecussate to alternate, high-

ly compound, spreading to horizontal, 5-15(-20)

cm long lateral branchlet systems. Ultimate

branchlets nodding, 3— 4(-6) mm in diameter in-

cluding leaves. Branchlet leaves usually borne in

densely crowded, alternating whorls or low spirals

of 3-5, usually 3-4 mm long, ca. 0.3 mm diameter,

acicular, terete to angular (dried), with often con-

spicuously acroscopically adnate and decurrent leaf

bases, gradually changing from patent-reflexed and

distant on main axes, to patent, upward curved

and densely crowded in ultimate branchlets, oc-

casionally with sparse, lax trichomes or minute

spinules; leaf bases often with longer, irregularly

crisped or branched trichomes, these rarely also

on stern surfaces. Strobili usually numerous, ses-

sile, terminating ultimate branchlets, 4-10(-20)

mm long, 2.5-3 mm in diameter. Sporophylls usu-

ally borne in alternating whorls of 5, ovate-deltoid,

short to long cuspidate, ca. 2 mm long, ca. 1 mm

wide, with membranous, coarsely erose-laciniate

margins. Sporangia globose, 0.5-0.8 mm in di-

ameter, strongly anisovalvate. Spores rugate,

without a distinct equatorial rim.

A common pioneer species on road cuts and

moist disturbed soil, along rivers, in forest clear-

ings, etc., from sea level up to 3000 m, Cajamarca,

Amazonas, San Martin, Loreto, Huanuco, Pasco,

Junin, Ayacucho, Cuzco, Puno.

Pantropic.

Where Lycopodiella cernua grows intermixed

with L. descendens, intermediates may occur.

Llanos & Chimouy 61 (USM), from Dept. Ca-

jamarca: Prov. Jaen, San Patricio, Santa Rita,

Chontali, 1 600 m, with the growth habit of L.

cernua, deviates from typical forms of the latter

by the densely crowded and strongly reflexed leaves

on the main erect axis, by the somewhat flattened

branchlet leaves, and by the densely hairy leaf bas-

es and stem surfaces. It closely resembles forms

from similar altitudes in Ecuador (Prov. Zamora-

Chinchipe) and Bolivia and may represent a dis-

tinct element, but more field observations and ma-

terial are desirable for an assessment of its status.

Cajamarca: Prov. Cutervo, La Pucarilla-San Andres,

2400 m, Sanchez Vega et al. 5999 (AAU, F). San Ignacio,

San Jose de Lourdes, Villarica, 1650 m, Diaz 2053 (F).

Amazonas: Prov. Bagua, road Chiriaco-Puente Vene-

zuela, ca. 200-300 m, Barbour 4326 (AAU, MO). Prov.

Chachapoyas, ca. 1800 m, Raimondi 350 (USM). San

Martin: Prov. Mariscal Caceres, Dist. Tocache Nuevo,

J. Schunke V. 3713 (F, us). La Divsioria, 59 km from

Tingo Maria on road to Pucallpa, Allard 22175 (us).

Loreto: Prov. Maynas, Dist. Iquitos, near mouth of Rio

Nanay, Rimachi 449 (F, MO, USM). Balsapuerto, 220 m,

Klug 2970 (F, MO, s, uc, us). Huanuco: Prov. Huanuco,

Dist. Churubamba, Hda. Mercedes, 1875 m, Mexia 8193

(F, GH, s, us). Vilcabamba, 2000 m, Macbride 5014 (F,

GH, MO, us). Pasco: Prov. Oxapampa, Palcazu valley,

300 m, D. Smith 3927 (AAU). Pichis Trail, Eneiias, 1700

m, Killip & Smith 25688 (F, us). Junin: Chanchamayo

Valley, 1200 m, C. Schunke 188 (F). La Merced, Killip

& Smith 23755 (us). Ayacucho: Prov. La Mar, eastern

Massif of the Cord. Central opposing the Cord. Vilca-

bamba between Tambo, San Miguel, Ayna and Hda.

Luisiana, 1570 m, Dudley 11889 (GH, us). Cuzco: Pil-

copata-Atalaya, 700 m, Vargas 13309 (GH). Puno: Olla-

chea to San Gaban, 1000-2000 m, Dillon et al. 1166 (F,

MO).

5. Lycopodiella camporum B. 011g. & P.

G.Windisch, Bradea 5: 24, t. 3. 1987. TYPE:

Brasil, Est. Minas Gerais, Mun. Santana do

Riacho, Serra do Cipo, Prado et al. 69 (ho-

lotype, HB!; isotypes, AAU!, RB!, SP!, SPF!).

Palhinhaea camporum (B. 011g. and Windisch) Holub,

Folia Geobot. Phytotax. 26: 93. 1991.

Plants with arching-looping runner shoots, with

up to 1 m tall, stiffly erect, dendroid branch sys-

tems, these with several, subdecussate to alternate,

often densely aggregated, stiffly ascending, 5-10

(-20) cm long lateral branchlet systems. Ultimate

branchlets 2.5-7 mm in diameter including leaves,

stiffly ascending to erect, only the strobiliferous

ones rather sharply recurved at the tip. Branchlet

leaves usually borne in densely crowded, alter-

nating low spirals or oblique whorls of 5-7, 2.5-

4 mm long, ca. 0.3-0.5 mm in diameter, acicular,

terete or angular (dried), sometimes flattened in

the lower half, with often slightly acroscopically

adnate and long decurrent leaf bases, patently ar-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

63

cuate-ascending to arcuate-appressed, entirely

smooth to densely hairy on leaf bases, rarely with

soft trichomes on leaf margins. Strobili usually

numerous, sessile, terminating tips of ultimate

branchlets with a sharp bend, to 2.5 cm long, 2-

3 mm in diameter. Sporophylls borne in alternat-

ing whorls of 5-7, widely ovate, short to long cus-

pidate, 1.5-2 mm long, 0.6-0.9 mm wide, with

scarious, coarsely erose-laciniate margins. Spo-

rangia globose, ca. 0.6 mm in diameter, strongly

anisovalvate.

Swamps, moist campo-grasslands, on peaty or

sandy soil, often in the light-open vegetation ad-

jacent to gallery-vegetation at rivers, subject to

flooding during the rainy season; alt. 1 20-1 800 m,

San Martin, Junin, Loreto.

Amazonian Colombia, Peru and Bolivia; sa-

vanna region of Venezuela and Guyana, campos

vegetation in Brazil.

Lycopodiella camporum is a widespread species

which has been confused with L. pendulina and

synonyms thereof. The distinctive branching habit

and characteristic ecology make L. camporum a

rather easily recognizable and ecologically well-

defined taxon, although detail characters of mor-

phology do not seem strong.

San Martin: Tarapoto, LI. Williams 5960 (F). Junin:

Prov. Satipo, Gran Pajonal, 1000-1 200 m, Pena & Oven-

tini 915 (USM). Loreto: Prov. Maynas, Quisto Cocha,

Iquitos, 120 m, Sagdstegui & Aldave 5821 (GH, MO).

6. Lycopodiella descendens B. 011g., in Harling

and Andersson, Fl. Ecuador 33: 143. 1988.

TYPE: Ecuador, Prov. Pastaza, Mera, B. 0llg.

& Balslev 9076 (holotype, AAU!; isotypes, F!,

QCA!).

Palhinhaea descendens (B. 011g.) Holub, Folia Geo-

bot. Phytotax. 26: 93. 1991.

Runner shoots arching-looping, usually densely

hairy, with relatively distant, appressed, subulate

leaves, with initially erect, then recurved and de-

scending, amply branched, up to at least 75 cm

long dendroid branch systems, these with several

subdecussate to alternate, highly compound,

spreading to hanging lateral branchlet systems, with

usually densely hairy main axis with relatively dis-

tant, appressed, subulate leaves. Ultimate branch-

lets 3-4 mm in diameter including leaves. Branch-

let leaves borne in alternating, densely crowded

whorls or low spirals of 5-6, 2—4 mm long, ca. 0.3

mm thick, subterete, acicular to quadrangular, with

long-decurrent and acroscopically adnate, hairy leaf

bases, ascending and upward curved, glabrous, or

with few lax trichomes above the base. Strobili

usually numerous, sessile, terminating ultimate

hanging branchlets, 5-20 mm long, ca. 3 mm in

diameter. Sporophylls usually borne in alternating

whorls of 5, ovate-deltoid, cuspidate, 1.5-2 mm

long, ca. 1 mm wide, with membranous, coarsely

erose-laciniate margins. Sporangia globose, ca. 0.5

mm in diameter, strongly anisovalvate. Spores ru-

gate, without a distinct equatorial rim.

Road banks, moist disturbed soil in lower, wet

montane forest, alt. 1000-1 500 m, Amazonas, San

Martin, Loreto, Huanuco.

Lower eastern slopes of the Andes in Peru and

Ecuador.

Lycopodiella descendens is closely related to L.

cernua, and is sometimes seen growing intermixed

with it. In such situations intermediate individuals

may be found. In addition to the key characters

mentioned, L. descendens very often differs from

L. cernua by its longer Strobili.

Amazonas: Prov. Bagua, Mesones-Muro highway, 286

km E of Olmos, 8 km E of Montenegro, 650 m, Hutchi-

son & Wright 3820 (F, uc, us). San Martin: Prov. Lamas,

Dist. Lamas, Rio Curiyacu, affluent of Rio Cumbasa, ca.

450 m, Belshaw 3585 (GH, MO, NY, us, us). Prov. Maris-

cal Caceres, Tocache Neuvo-Juanjui road, km 88, 850

m, D. Smith 2152 (AAU). Prov. Moyobamba, Jepelacio,

Cerro Shallcahurco, 1320 m, Fernandez & Clemants 27

(USM). Loreto: Pumayacu, between Balsapuerto and

Moyobamba, 600-1200 m, Klug 3232 (F, G, GH, K, MO,

NY, s, us). Huanuco: Prov. Huanuco, Dist. Churubamba,

Hda. Mercedes, 1640 m, Mexia 8195 (F, GH, K, s, uc,

us). SW slope of the Rio Llullapichis watershed, on the

ascent of Cerros del Sira, ca. 1000 m, Dudley 13125 (GH).

7. Lycopodiella glaucescens (Presl) B. 011g., Ope-

ra Bot. 92: 176. 1987.

Lycopodium glaucescens Presl, Reliq. haenk. 1: 81.

1825. TYPE: Peru (Huanuco), In montanis ad

Huanocco, Haenke (holotype, PRC!).

Palhinhaea glaucescens (Presl) Holub, Folia Geobot.

Phytotax. 26: 93. 1991.

Runner shoots long, robust, arching-looping to

scandent, with initially erect, amply branched, up

to several meters long, bending to long-scandent

dendroid branch systems, these with several

spreading or nodding to long pendulous, usually

alternate, up to at least 40 cm long lateral branchlet

systems. Leaves of main axes patent to reflexed,

usually strongly upward curved to hook-shaped,

3-5 mm long, up to 1 mm wide, with terete to

64

FIELDIANA: BOTANY

quadrangular, hairy leaf bases, apically flattened,

usually coriaceous. Ultimate branchlets 3-5 mm

in diameter including leaves, the sterile ones often

tapering to less than 2 mm in diameter. Leaves of

ultimate branchlets borne in densely crowded, al-

ternating whorls or low spirals of 4-6, acicular to

lanceolate, usually with flattened apex, 2.5-4 mm

long, 0.3-0.8(-1.0) mm wide, usually strongly up-

ward curved or hook-shaped from a spreading or

perpendicular to reflexed base, soft herbaceous to

rigidly coriaceous, hairy only on leaf base. Strobili

variable, 5-30 mm long, 3-4.5(-5) mm in diam-

eter. Sporophylls borne in alternating whorls of 5,

lanceolate-ovate, apically acute-acuminate, 2-3

mm long, 0.8-1.2 mm wide, with shallowly den-

tate-ciliate margins, of herbaceous, pale green tex-

ture throughout. Sporangia globose, strongly an-

isovalvate, 0.7 mm in diameter. Spores rugate,

without a distinct equatorial rim.

Clearings and open places in wet upper montane

forest, alt. 1600-3700 m, Cajamarca, San Martin,

Huanuco, Pasco, Cuzco.

Costa Rica; Colombia to Peru; possibly farther

north and south.

A remarkable species with often very long scan-

dent dorsally arising shoots. Under favorable con-

ditions, these may reach a length of several meters.

Lycopodiella glaucescens is highly variable with

respect to strobilus size, size, coarseness, and di-

rection of the branchlet systems. Plants of exposed

habitats are usually relatively short and compact

and with rigid, coriaceous leaves, approaching the

growth habit of L. pendulina, while individuals

from protected habitats are longer, more lax, and

soft and with long pendulous branchlets.

Cajamarca: Prov. Cutervo, La Pucarilla (San Andres-

Socota), 2420 m, Lopez & Sagdstegui 5459 (GH). San

Martin: Huallaga. Valley of Rio Apisoncho, 30 km above

Jucusbamba, Hamilton & Holligan 1248 (K). Prov.

Mariscal Caceres, Rio Abiseo National Park, hill past

Las Palmas, 2650-2750 m, Young 4000 (USM). Huanuco:

Prov. Huanuco, Carpish, 2850 m, Asplund 13076 (s).

Playapampa, 3000 m, Macbride 4506 (F, G, us). Pasco:

Prov. Oxapampa, Cord. San Gutardo, 2800 m, Leon

523a (AAU, USM). Cuzco: Prov. La Convencion, Cord.

Vilcabamba, 2825 m, Dudley 10782 (USM).

8. Lycopodiella pendulina (Hooker) B. 011g., Ope-

ra Dot. 92: 176. 1987.

Lycopodium pendulinum Hooker, Icon. pi. 1: /. 90.

1837. TYPE: Peru, Casapi, Matthews 1776 (ho-

lotype, K.!).

Lycopodium cernuum L. var. pendulinum (Hooker)

Baker, Handb. Fern Allies 23. 1887.

Palhinhaea pendulina (Hooker) Holub, Folia Geobot.

Phytotax. 26: 93. 1991.

Runner shoots robust, shallowly arching, with

up to ca. 40 cm tall, erect dendroid branch systems,

these with several subdecussate to upward alter-

nate, sparsely subequally branched, up to 1 2(-20)

cm long, usually long pendulous lateral branchlet

systems. Leaves of main axes loosely appressed,

upward curved, 5-7 mm long, up to 1 mm wide.

Ultimate branchlets (3-)5-6(-9) mm in diameter

including leaves, rarely tapering to 2 mm in di-

ameter. Branchlet leaves borne in densely crowded

alternating whorls or low spirals of 5-7, acicular,

terete to quadrangular, or sometimes apically flat-

tened, (3-)4-6 mm long, up to 1 mm wide, patent-

ascending, upward curved, softly to firmly her-

baceous, glabrous, or with few marginal cilia. Leaf

bases and stem surfaces usually glabrous, rarely

with short crisped trichomes. Strobili 10-20 mm

long, 5-6 mm in diameter. Sporophylls borne in

alternating whorls of 5-6, lanceolate-ovate, short

acuminate, (2.5-)3-3.5 mm long, ca. 1 .5 mm wide,

with irregularly dentate, narrowly membranous

margins, pale greenish, of herbaceous texture

throughout. Sporangia globose, strongly aniso-

valvate, ca. 1 mm in diameter. Spores rugate,

without a distinct equatorial rim.

Wet grassland, open spaces or clearings in the

upper montane forest and, 2200-3600 m, Piura,

Amazonas, Huanuco, Cuzco.

Costa Rica; Andes from Venezuela to Bolivia,

southeastern Brazil.

Lycopodiella pendulina is most closely related

to L. glaucescens, and distinguished from the latter

by its stiffly erect main shoots with thick, sparsely

ramified, weeping branchlet systems and large

Strobili.

Piura: Prov. Huancabamba, Loma Redonda (Sapa-

lache-Chinguela), 2400 m, Sagdstegui et al. 10217 (AAU).

Amazonas: Prov. Chachapoyas, Cerros Calla Calla, E

side, 1 8 km above Leimebamba on road to Balsas, 3 100

m, Hutchison & Wright 5663 (BR, c, E, F, G, GH, M, MO,

NY, s, uc, USM). Prov. Bagua, Cord, de Colan E of La

Peca, 3400 m, Barbour 3199 (AAU, MO). Huanuco: Prov.

Huanuco, Carpish, 2800 m, Asplund 13105 (s). Chushi,

trail to Tambo de Vaca, Bryan 681 (F, us). Cuzco: Prov.

La Convencion, Cord. Vilcabamba, 2825 m, Dudley

10707, 10782 (GH, MO).

9. Lycopodiella riofrioi (Sodiro) B. 011g., Opera

Bot. 92: 176. 1987.

Lycopodium riofrioi Sodiro, Crypt, vase. Quit. 582.

1893. TYPE: Ecuador (Prov. Pichincha), In silv.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

65

occid. m. Pichincha ad Gualea, 9/888, Sodiro (ho-

lotype, si 2 1230!).

Lycopodium pensum Lell., Proc. Biol. Soc. Wash. 89:

7 1 7, t. 2. 1 977. TYPE: Costa Rica, Prov. Heredia,

6 mi from San Rafael de Heredia on slopes of

Volcan Barba [Cerro Chompipe], McAlpin 216

(holotype, DUKE; isotype, GH!).

Palhinhaea riofrioi (Sodiro) Holub, Folia Geobot.

Phytotax., Praha 26: 93. 1991.

Growth habit as in Lycopodiella pendulina, up

to 2 m tall. Branchlet leaves imbricate, densely

crowded, borne in alternating whorls of 4-6, lan-

ceolate to ovate-lanceolate, acuminate, 2.5-4 mm

long, ca. 1 mm wide, with densely ciliate-fimbriate

margins, with a very short, patent, terete-suban-

gular base. Stems and leaf bases densely hairy.

Strobili 1-2.5 cm long, 3-5 mm thick including

sporophylls. Sporophylls borne in alternating

whorls of 5, ca. 2.5 mm long, ca. 1 mm wide.

Sporangia 0.7-1 mm in diameter.

Wet montane forest, in clearings and on road

banks, 1700-2600 m. To be expected in north-

ernmost Peru.

Costa Rica; Panama; Venezuela; Colombia to

southern Ecuador.

A very distinctive species with its smooth-ap-

pearing branchlets covered by closely imbricate,

scalelike leaves.

Family 27. SELAGINELLACEAE

Selaginellaceae Milde, Hoher. Sporenpfl. Deutschl.

Schweiz 136. 1865, as Selaginelleae. TYPE:

Selaginella Beauv.

Stem indurated or not, branched, bearing rather

few, often long roots usually at a branch of the

stem. Leaves simple, ca. 0.5-10 mm long, with 1

(very rarely 2) vein(s). Sporangia short-stalked,

single, near the axil of a leaf. Heterosporous, spores

without chlorophyll.

The Selaginellaceae are a distinctive family in-

cluding the single genus Selaginella, only distantly

related to others such as the Lycopodiaceae and

Isoetaceae. Heterospory, and the presence of ves-

sels in some species of subgenus Selaginella, in-

dicate the family is specialized.

I. Selaginella

Selaginella Beauv., Magasin Encycl. 5: 478. 1804;

and Prod. Fam. Aetheog. 101. 1805, nom.

conserv. TYPE: Selaginella spinosa Beauv.,

nom. nov. for Lycopodium selaginoides L. =

Selaginella selaginoides (L.) Link. Figure 9.

Bryodesma Sojak. Preslia 64: 154. 1992. Type: Bry-

odesma rupestris (L.) Sojak (Lycopodium rupestre

L.) = Selaginella rupestris (L.) Spring.

Terrestrial, rupestral, or rarely epiphytic. Stem

slender, branched, sometimes dichotomously,

prostrate-creeping or with ascending branches, or

pendent-epiphytic, or erect from a usually stolon-

iferous base. Leaves ca. 1.0-10 mm long, with

usually 1 vein, borne in a close spiral or usually

alternate in 4 ranks. Sporangia large, borne in or

near the axil of a well-differentiated sporophyll, in

a quadrangular or (in S. deflexa and S1. selagi-

noides) a cylindrical strobilus. Megasporangia

commonly basal in the strobilus, usually with 4

megaspores, larger than the microsporangia and

different in size and color. Microsporangia usually

borne above the megasporangia and with many

microspores. Megaspores tetrahedral-globose, tri-

lete, the laesurae % to usually equaling the radius,

often with a more or less prominent equatorial

ridge, rugose-reticulate, rugose, papillate, tuber-

culate, granulate, rarely nearly smooth on the

proximal face. Microspores tetrahedral-globose,

trilete, often compressed or the proximal face more

or less depressed, the laesurae '/2 to equaling the

radius, usually finely to coarsely echinate, rugose,

papillate, perforate-cristate, or granulate.

Selaginella is a large genus of perhaps 700 spe-

cies, with about 40 of them in Peru, or expected

there. The descriptions are drawn from Peru ma-

terial and are intended to be parallel within related

groups of species. The genus has not been ame-

nable to segregation although five subgenera are

FIG. 9. Selaginella peruviana: a, habit; b, tip of branch with aerial root. Selaginella haematodes: c, habit; d,

portion of branch, abaxial side, showing axillary and lateral leaves. Selaginella exaltata: e, habit; f, portion of secondary

branch, adaxial side, showing median leaves, (a, b from Solomon & Nee 17895, Bolivia, F; c, d from Taylor 5049,

Brazil, F; e, f from Bohs 2215, Ecuador, F.)

66

FIELDIANA: BOTANY

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

67

currently recognized (Jermy, 1986), and Sojak has

recently recognized a second genus, Bryodesma,

for the S. rupestris group.

The genus is probably more diverse in Peru than

presently known. Species are difficult to identify,

and many may appear similar to the field botanist

unacquainted with special specific characters. At

the same time, a better understanding of the vari-

ation within species and the morphology of ju-

venile plants will lead to an increase in synonyms.

The species have been placed in a sequence that,

presumably, extends from primitive to advanced.

Species 1-2 are homophyllous, while the remain-

der of the species are heterophyllous.

Species 3-10 are prostrate and not articulate

(both primitive characters), while species 11-12

are prostrate and articulate. These species relate

to the former group by their small leaves and often

intricate stems. Species 13-15 have erect, not ar-

ticulate stems that are usually short and pinnately

branched. Species 16-25 are also erect and not

articulate but usually have a main stem with ap-

pressed leaves toward the base and flabellate

branches. Species 1 7 may be unbranched or with

irregular branches. Species 26-30 are prostrate and

articulate, as are species 11-12. However, these

have large leaves and mostly separate stems. Spe-

cies 31-34 are articulate and erect (both derived

characters). Species 35 is a climbing species and

36 is a highly derived, xeric species.

The following treatment is based to a large ex-

tent upon the published work of A. H. G. Alston

(1934), especially the (posthumous) publication

with A. C. Jermy and J. M. Rankin (1981). The

Koller and Scheckler (1986) paper is among the

many that could be cited, but it is selected because

of the great potential of their work to the classi-

fication of the genus.

References

ALSTON, A. H. G. 1934. Notes on Selaginella,

VI. J. Dot. 72: 223-226.

ALSTON, A. H. G., A. C. JERMY, AND J. M. RANKIN.

1981. The genus Selaginella in tropical South

America. Bull. Brit. Mus. (Nat. Hist.) Bot. 9:

233-330.

BAKER, J. G. 1887. Handbook of the fern-allies.

George Bell & Sons, London.

JERMY, A. C. 1986. Subgeneric names in Selag-

inella. Brit. Fern Gaz. 13: 117, 118.

KOLLER, A. L., AND S. E. SCHECKLER. 1986. Vari-

ations in microsporangia and microspore dis-

persal in Selaginella. Amer. J. Bot. 773: 1274-

1288.

SOJAK, J. 1992. Generische problematik der Se-

laginellaceae. Preslia 64: 151-158.

TRYON, R. M. 1 955. Selaginella rupestris and its

allies. Ann. Mo. Bot. Gard. 42: 1-99.

VALDESPFNO, I. A. 1993. Notes on Neotropical

Selaginella (Selaginellaceae), including new spe-

cies from Panama. Brittonia 45: 315-327.

Key to Species of Selaginella

a. Leaves of branches and stems more or less uniform, spirally arranged; main stem prostrate, creeping

b

b. Leafy stem radially symmetrical; leaves all alike; base of the lower leaves abruptly adnate, distinct

from the stem 1 . S. sellowii

b. Leafy stem somewhat dorsiventral; upper leaves differ from the lower leaves on the same portion

of the stem; base of the lower leaves strongly decurrent on the stem 2. S. peruviana

a. Leaves of the branches and often of the main stem dimorphic, in 4 ranks with 2 smaller median

ranks on the upper side of the leafy stem and 2 larger lateral ranks; main stem erect, assurgent at the

apex, to prostrate c

c. Main stem very short, erect, bearing leafy branches that form a rosette; branches strongly involute

when dry 36. S. convolute

c. Main stem long, erect, assurgent at the apex, to prostrate; branches straight or nearly so when dry

d

d. Main stem erect, uniformly red or reddish, especially below the basal branch, not articulate

e

e. Acroscopic edge of lateral leaves ciliate or ciliolate, at least near the base; erect stem and

branched portion ca. 10-25 cm, mostly 10-15 cm long 21. S. erythropus

68

FIELDIANA: BOTANY

e. Acroscopic edge of lateral leaves minutely denticulate to essentially entire; erect stem and

branched portion mostly 30-65 cm long 22. S. haematodes

d. Main stem erect to prostrate, brownish, greenish, or stramineous, rarely red and then articulate

(with a break, a dark ring, a swollen area, or usually a constriction) f

f. Main stem climbing or scrambling to 8 m long, articulate 35. S. exaltata

f. Main stem erect to prostrate, not exceeding 1 m long, articulate or not g

g. Strobilus with sporophylls only on the upper side of the leafy stem 10. S. ramosissima

g. Strobilus tetragonous, sporophylls spirally arranged on the stem h

h. Branches pubescent, especially on the branch axils, trichomes sometimes sparse . . .

30. S. articulata

h. Branches glabrous i

i. Leaves tightly clasping the stems when dry 9. S. microphylla

i. Leaves spreading, ascending or appressed, not clasping the stems when dry . . . . j

j . Plants with an erect main stem and often a flabelliform arrangement of branches;

basal part of the main stem usually with appressed leaves, these dimorphic or

not; rhizophores usually confined to the basal portion of the main stem, or rarely

above the first branch k

k. Ultimate leafy branches 5-20 mm broad 1

1. Axis of primary branches fractiflex, branches sometimes flagelliform; main

stem articulate 3 1 . S. parkeri

1. Axis of primary branches straight or nearly so, branches not flagelliform;

main stem not articulate m

m. Median leaves more or less acute to acuminate, not or hardly aristate

n

n. Median leaves inequilateral, the outer side larger, the costa acentric

24. S. praestans

n. Median leaves equilateral, the sides nearly equal in size, the costa

central 23. S. quadrifaria

m. Median leaves aristate o

o. Primary branches usually 1 -pinnate, sometimes 2-pinnate with

branches nearly parallel from the first branch to the apex of the

main stem 16. S. bombycina

o. The main stem and branches irregularly or dichotomously divided

P

p. Branches usually few, erect, of irregular lengths, not forming a

flabelliform arrangement 17. S. chrysoleuca

p. Branches forming a broad, flabelliform arrangement

1 8. S. speciosa

k. Ultimate leafy branches 5 mm broad or less q

q. Axis of primary branches fractiflex r

r. Median leaves oblong, with a basal auricle that is entire or

nearly so 32. S. geniculata

r. Median leaves ovate-lanceolate, with a ciliolate basal auricle

34. S. stellata

q. Axis of primary branches straight or nearly so s

s. Main stem articulate 33. S. asperula

s. Main stem not articulate t

t. Base of the main stem slender, 0.25-0.50 mm broad; leaves

short, the longest 2 mm long . . 13. S. novae-hollandiae

t. Base of the main stem stouter, 0.75-3.5 mm broad; longest

leaves 2-5 mm long u

u. Median leaves aristate 25. S. haenkeana

u. Median leaves acute to acuminate, rarely a few aristate

. v

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

69

v. Leaves of the main stem distal to the second branch,

strongly ascending to spreading (patent); stem and

leaves near the axil of the first branch usually 4-5

mm broad 19. S. lechleri

v. Leaves of the main stem distal to the second branch,

appressed; stem and leaves near the axil of the first

branch 1-3 mm broad 20. S. anceps

j. Plants with a prostrate or assurgent, sometimes intricate, or rarely erect stem;

basal part of the main stem with dimorphic leaves; lateral leaves spreading;

rhizophores usually borne throughout the main stem, or at least above the first

branch w

w. Main stem articulate x

x. Leafy branches slender, 2-4 mm broad, often intricate y

y. Lateral leaves with ciliolate margins at the base, without a prolonged

auricle 12. S. lingulata

y. Lateral leaves with entire margins, with 1 or 2 prolonged, ciliolate

auricles 11. S. diffusa

x. Leafy branches 4-20 mm broad, not intricate z

z. Leafy branches 8-20 mm, often 10 mm broad aa

aa. Base of the lateral leaves without prolonged auricles

28. S. trisulcata

aa. Base of the lateral leaves with 1 or 2 prolonged auricles, the inner

sometimes an acute lobe bb

bb. Leafy main stem 12 mm or less broad, glabrous; branches

forming an open, dichotomous arrangement

29. S. poeppigiana

bb. Leafy main stem 1 5-20 mm broad, often pubescent beneath,

especially on the axils, trichomes sometimes sparse; branches

at the apex forming a broad, compact, flabellate arrangement

30. S. articulata

z. Leafy branches 4-8 mm, usually 5 mm broad cc

cc. Apical portion of the main stem, and often of the primary branches,

forming a somewhat rhomboid arrangement, the apex acute to

acuminate 26. S. kunzeana

cc. Apical portion of the main stem, and often of the primary branches,

forming a broadly ovate to obdeltate arrangement, the apex obtuse

to truncate 27. S. silvestris

w. Main stems not articulate dd

dd. Median leaves with a short, acute apex often extended in a cusp

5. S. truncata

dd. Median leaves aristate or subaristate ee

ee. Edges of lateral leaves entire, denticulate, or ciliolate ff

ff. Leafy branches, when present, 5-20 mm, mostly ca. 1 5 mm broad;

main stem with no, or rarely few primary branches, these strongly

ascending, mostly dichotomous 17. S. chrysoleuca

ff. Leafy branches, 3-12 mm, mostly ca. 5 mm broad; main stem

with many, spreading branches gg

gg. Stems erect; rhizophores confined to the base of the main

stem or not beyond the first branch hh

hh. Ultimate branches short, the apex acute

1 5. S. flagellata

hh. Ultimate branches longer, with the apex truncate to

somewhat rounded 14. S. xiphophylla

gg. Stems prostrate, creeping, sometimes intricate, forming a mat;

70 FIELDIANA: BOTANY

rhizophores extending to the stem apex, or above the first

branch to lh the length of the stem ii

ii. Stems usually intricate, forming a mat; penultimate leafy

branches 2-4 mm broad 6. S. t a ra pot ens is

ii. Stems elongate, discrete; penultimate leafy branches 4-10

mm broad jj

jj. Median leaves orbicular, the base truncate; lateral leaves

oblong with an obtuse apex 8. S. product a

jj. Median leaves ovate to oblong, the base with an ex-

tended, round lobe; lateral leaves elongate-ovate, with

an acute apex 7. S. seemannii

ee. Edges of lateral leaves ciliate, the cilia most abundant from the base

to the mid-part of the leaf kk

kk. Median leaves without a prolonged auricle .... 3. S. revoluta

kk. Median leaves with a prolonged auricle 11

1 1 . Main stem short, erect; rhizophores at base of the stem or

mostly below the first branch ... 13. S. novae-hollandiae

1 1 . Main stems creeping, often intricate and forming a mat;

rhizophores extending above the first branches

. . 4. S. brevifolia

1 . Selaginella sellowii Hieron., Hedwigia 39: 306.

1900. LECTOTYPE (by R. Tryon, 1955):

Brazil, Paria de San Diego Sellow, in 1821

(holotype, B; frag., NY).

Selaginella rupestris f. amazonica Milde, Fil. Eur. At-

lant. 263. 1867. TYPE: Peru, Cajamarca, (Rio)

Maranon, Bonpland (holotype, B; isotype, BM).

Selaginella amazonica (Milde) Hieron., Hedwigia 39:

310. 1900, not S. amazonica Spring, 1840.

Selaginella mildei Hieron., in Engler & Prantl, Nat.

Pflanzenfam. 1 (4): 671. 1902, nom. nov. for S.

amazonica (Milde) Hieron.

Main stem prostrate, creeping, assurgent at the

tip, not articulate, pinnately branched throughout,

green to brownish, glabrous. Primary branches

mostly simple, 1 -pinnate, or often 2-pinnate, the

ultimate branches as broad as the main stem or

nearly so, 1-1.5 mm broad including the leaves.

Rhizophores throughout the stems, mostly at the

axils of the branches. Leaves similar (homophyl-

lous), spirally arranged, subulate, long-triangular

to broadly Ungulate to lanceolate, the edges whit-

ish, denticulate, the apex setate, the seta opaque,

milky-white, the base abruptly adnate, ciliate or

ciliolate, rarely lacking cilia, abruptly distinct from

the stem in form and color, glabrous.

On exposed or wooded, rocky bluffs, or among

stones, from ca. 300 to 2900 m, Lambayeque, Ca-

jamarca, Ancash, and Cuzco.

Central Mexico, Cuba, Venezuela, and Colom-

bia south to Argentina, and eastern Brazil.

The species belongs to the Sartorii series of the

Selaginella rupestris group and appears most closely

allied to S. sartorii Hieron. Geographically it is

wide-ranging from Mexico and Cuba through South

America. It is readily distinguished from the nine

others of series S. sartorii and other Peruvian spe-

cies by the glabrous leaf bases, especially by the

opaque, milky-white setae on the leaves. This and

S. peruviana have elongate stems that differ from

the radially symmetrical plants of S. convoluta.

Lambayeque: Hacienda Valor, Ellenberg 3613 (GH).

Cajamarca: between Llaconora and Namora, Correll &

Smith P893 (GH, MO). Between Jaen and Quemado, Fe-

rreyra & Sanchez 19668 (USM). Ancash: Mancos, Yun-

gay, Tryon & Tryon 6551 (GH). Cuzco: Biies 608 (us);

Herrera 3009a (us).

2. Selaginella peruviana (Milde) Hieron., Hed-

wigia 39: 307. 1900. Figure 9a-b.

Selaginella rupestris f. peruviana Milde, Fil. Eur. At-

lant. 263. 1891. TYPE: Peru, Huanuco, Ruiz 98

(holotype, B; isotype, NY).

Selaginella peruviana var. dombeyana Hieron., Hed-

wigia 39: 308. 1900. PARATYPE: Peru, Dombey

14 (B).

Selaginella sheldonii Maxon, Proc. Biol. Soc. Wash.

31: 171. 1918. TYPE: United States, Quanah

Mountains, Oklahoma, Sheldon 233 (holotype,

us).

Main stem prostrate, creeping, not articulate,

pinnately branched throughout, glabrous, green to

brownish. Leafy branches forming compact to dif-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

71

fuse mats with discrete, simple or 1 -pinnate, some-

times 2-pinnate stems. Rhizophores borne

throughout the stems at the axils of branches.

Leaves generally similar (homophyllous), spirally

arranged, setate, the seta usually more or less lu-

tescent, or often not well differentiated in color,

the base adnate, glabrous, sometimes denticulate,

the edges usually ciliate, or sometimes somewhat

pubescent, leaves on the underside of the stem

differ from those on the upper side, on the same

part of the stem, those on the upper side linear,

acuminate to long-triangular, the lower leaves su-

bulate, acuminate, broadest at or near the base, to

linear-lanceolate, gradually distinct from the stem.

Mats on bluffs, dry rocky slopes, ledges or crev-

ices of cliffs, exposed or in light shade, igneous,

calcareous, or sandstone rock, 1750—4400 m, Ca-

jamarca south to Puno.

Southwestern United States south to Puebla,

Mexico; Peru to Argentina.

The species is recognized as the least specialized

and basic species within the series Eremophilae,

in the treatment of the Selaginella rupestris group

(Tryon, 1955). Geographically it is the widest

ranging species of that series. On the basis of the

slender leaves bearing long setae, it is most closely

related to S. arizonica Maxon, a species of north-

west Mexico and the states of Texas and Arizona.

However, it is distinguished from this as well as

Peruvian selaginellas by the lower leaves that are

strongly decurrent on the stem. Among the Pe-

ruvian selaginellas, it is most closely allied with

S. sellowii, another species with homophyllous

leaves.

Cajamarca: Between Cajamarca and San Juan, Mutter

& Gutte 8869 (USM). Ancash: Near Mancos, Tryon &

Tryon 6550 (GH). Huanuco: Prov. Huanuco, Plowman

& Rury 11103 (F, GH, USM), Prov. Huanuco, Stork &

Morton 9388 (F, uc). Lima: Prov. Canta, Punte San Jose,

Lopez & Riccio 10080 (GH). Prov. Huarochiri, Leon 583

(USM). Junin: Above Conception, Correll & Smith P748

(GH, MO). Prov. Tarma, lit is et al. 108 (GH, wis). Apu-

rimac: Prov. Andahuaylas, Stork & Morton 10719 (F,

uc). Cuzco: Near Pisac, McDaniel & Gorski 11402 (GH,

MO). Urubamba valley, Leon 443 (F, USM). Prov. Uru-

bamba, Chicon, Vargas 11058 (F, uc). Puno: Near Puno,

Solomon 2895 (F, MO).

3. Selaginella revoluta Baker, J. Bot. 21:141.1883.

TYPE: Venezuela, Amazonas, near Mau-

pures, Spruce 3621 (holotype, K; isotypes, CGE,

us).

Selaginella demissa Christ, Bull. Herb. Boissier 2, 1:

75. 1883. TYPE: Peru, Cerro Canchahuaya, Hu-

ber 1421 (holotype, p).

Main stem short, creeping, often forming com-

pact mats, not articulate, stramineous, glabrous,

the basal part with spreading leaves, and often

smaller, appressed, ascending leaves. Primary

branches short, distant, usually 1 -2-pinnate, ex-

tending to the base of the main stem, 3-5 mm

broad including the leaves, the terminal branches

short. Rhizophores extending from the base of the

main stem to the apex of the branches. Lateral

leaves above the first branch not imbricate,

spreading, elongate-ovate, broadest at the base,

0.5-0.2 mm long, ca. 3 x longer than broad, acute,

with a round, ciliolate, scarcely prolonged basal

lobe, without conspicuous whitish borders, the

edges with long cilia, these denser at the base.

Median leaves suborbicular, broadest at the cen-

ter, aristate, the arista '/2 as long as the lamina,

cuspidate, the auricles not prolonged, often with

narrow, whitish borders, the edges denticulate.

In wet, primary forests, 100-1100 m, Loreto

and Huanuco.

Costa Rica and Panama; in South America from

the Guianas south to Peru and Brazil.

The compact, creeping habit of the plants with

short, lateral branches bearing discrete, wide-

spread leaves along the main stem form a dis-

tinctive growth pattern. The species was allied with

Selaginella product a by Alston et al. (1981), but

distinguished by the strongly tapering, ciliate, more

acute lateral leaves, sometimes pubescent on the

upper surface near the margin. However, that spe-

cies also has broader, more delicate leafy stems.

Loreto: Prov. Maynas, Mishuyacu, Klug 1220 (F, us).

Prov. Maynas, below Iquitos, Moran 3651 (MO). Gami-

tanacocha, Schunke 322 (F, GH, us, USM). Huanuco: Tin-

go Maria, Allard 20485 (MO), 20846 (us).

4. Selaginella brevifolia Baker, J. Bot. 21:83.1883.

TYPE: Brazil, Janarate Cochoeira, Rio Ne-

gro, Spruce 2547 (holotype, K; isotypes, BM,

CGE).

Main stem prostrate, creeping, it and its branch-

es forming a mat of stems, sometimes elongate to

15 cm, not articulate, stramineous, glabrous, the

basal part with spreading leaves and often smaller,

appressed, ascending leaves. Primary branches 2-

3 -pinnate, alternate, distant, extending to the base

of the main stem, 2-3 mm broad including the

leaves. Rhizophores above the first branch, up to

72

FIELDIANA: BOTANY

'/3 the length of the stem. Lateral leaves ovate,

acute, ca. 1-2 mm long, without prolonged auri-

cles, ciliate especially on the acroscopic margin,

the cilia usually long, discrete. Median leaves ovate-

elliptic, acuminate to aristate, the arista '/4 the lam-

ina length, the base with a broad, long, ciliolate

auricle, without conspicuous whitish borders, the

edges denticulate.

On soil and among rocks, along roadsides, 400-

2400 m, Cajamarca, San Martin, and Huanuco.

Venezuela, Colombia, Peru, and western Ama-

zonas, Brazil.

Although the plants are small and mosslike in

habit, growing over soil or among rocks, they are

well collected and known to be wide-ranging in

South America and in Peru.

The species is considered close to the Greater

Antillean Selaginella cordifolia but with longer

aristae on the median leaves (Alston et al., 1981).

It also resembles S. novae- hollandiae, but the

strongly prostrate habit with rhizophores above

the median part of the main stem contrast with

the erect habit and rhizophores, mainly at the base

of the stem in S. novae- hollandiae.

Cajamarca: Prov. Contumaza, Guzmango, Sagdstegui

3932 (GH). Prov. Cajamarca, El Molino, Sagdstegui 7995

(F). Contumaza, Lleden, Sagdstegui et al. 10887 (F). San

Martin: Boqueron Pass, between Tingo Maria and Pu-

calpa, Allard22124 (us). Huanuco: Cushi, trail to Tambo

de Vaca, Bryan 718 (F, us).

5. Selaginella truncata A. Braun, Index Sem, Hort.

Bot. Berol. Appendix 1857: 15. SYNTYPES:

Colombia, Cundinamarca, Bogota, Andes of

New Grenada, Karsten (B, BM); Susumuco,

Triana 696 (BM), 238 (NY).

Selaginella weberbaurei Knox, Trans. Bot. Soc. Ed-

inburgh 35:282. 1950. Only spores are described.

"TYPE": based on a specimen collected by We-

berbauer, named by Hieronymus, B?.

Main stem prostrate, assurgent, not articulate,

stramineous, glabrous, the basal part with spread-

ing leaves and often smaller appressed, ascending

leaves. Primary branches 1-2-pinnate, pinnately

branched, extending to the base of the main stem,

the central branches distant on the main stem,

strongly ascending, forming an open, dichotomous

division of the branches, the ultimate branches as

broad as the main stem, 4-6 mm broad including

the leaves. Rhizophores slender, threadlike,

throughout the main stem. Lateral leaves uniform

in length, 2-3 mm long, oblong to broader near

the base, the apex obtuse, the base truncate, with

narrow, whitish borders and delicate, long cilia at

the edges, especially dense at the base. Median

leaves ovate, the apex acute, sometimes with a

short cusp, the base truncate, with whitish borders,

the edges regularly ciliate or ciliolate, more strong-

ly so at the base.

Epiphytic in dense jungle, or on rotted logs, in

dense forests, 365-700 m, San Martin and Cuzco.

Venezuela and Colombia south to Bolivia.

The compact, deep green leaves, imbricate on

the stems, give a distinctive aspect to the plants.

Ecological information on collections indicates

the plants are epiphytic on trees 15 ft above the

ground. This is corroborated by the dense rhizo-

phores on the branches and the stems that are free

from soil.

Selaginella applanata A. Braun, Ann. Sci. Nat.

Bot. 5, 3: 274. 1865, may be a synonym; uc has

a fragment of an isotype, Peru, Puno, San Gavan,

(Gaban) Lechleri 2405.

San Martin: Prov. Lamas, San Antonio, Belshaw 3569

(H, GH, uc, us). Guayrapurima, Tarapoto, Spruce 4024

(GH, us). Pasco: Prov. Oxapampa, Cordillera de San Ma-

tias, D. Smith 2000 (F). Cuzco: Prov. Quispicanchis, entre

Inombaris y Quincemil, Vargas 1 1690 (GH); entre Quin-

cemil y San Lorenzo, Vargas 16481 (GH).

6. Selaginella tarapotensis Baker, J. Bot. 21: 98.

1883. TYPE: Peru, Mt. Guayrapurima, near

Tarapoto, Spruce 4625 (holotype, K; isotypes,

BM, CGE, us).

Main stem prostrate or assurgent at the tip, often

forming a mat of stems, not articulate, stramin-

eous, glabrous, the basal part with spreading leaves,

and often smaller, appressed, ascending leaves.

Primary branches 3-4-pinnate, alternate, distant,

extending to the base of the main stem, 2-4 mm

broad including the leaves, the ultimate branches

short. Rhizophores filamentous, dense at the base,

extending nearly to the stem apex. Lateral leaves

oblong-lanceolate, but not imbricate on the main

stem, 0.5-2.0 mm long, approximately more than

3 x longer than broad, obtuse to acute, the auricles

not prolonged, without conspicuous whitish bor-

ders, the edges short-denticulate. Median leaves

rhomboidal to ovate, aristate, the arista slender,

usually whitish, '/2 as long to nearly equal the length

of the lamina, the base not or hardly prolonged,

often with narrow, whitish borders that extend

into the whitish arista, the edges denticulate.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

73

Rarely epiphytic, in deep cloud forests, and on

shaded banks bordering roads, 500-1 200 m, Ama-

zonas and San Martin south to Cuzco.

Costa Rica south to Bolivia.

The truncate leaf bases on both lateral and me-

dian leaves, along with the rhomboid shape and

prominent aristata on the median leaves, distin-

guish the species. This is one of the slender-

stemmed, mat-forming species most closely re-

sembling Selaginella revoluta.

Selaginella calosticha Spring, Nouv. Mem. Acad.

roy Sci. Belg. 24: 206. 1849. TYPE: Venezuela,

Caracas, Funck & Schlim 3321 (K), may be an

earlier name for this species. Two Allard collec-

tions from Peru— 20821 and 21206— are cited as

S. calosticha in Alston et al. (1981).

Amazonas: Prov. Bagua, Montenegro, Hutchison &

Wright 3822 (F, GH, uc, us). Monte Campana, near Tara-

poto, Spruce cf4625 (us). San Martin: Tocache Nuevo-

Juanjui, Smith et al. 2132A (uc). Huanuco: Prov. Tingo

Maria, Cueva de las Pavas, Aldave & Fernandez 5578

(GH). Pasco: Oxapampa, Cordillera San Matias, Leon et

al. 321 (F, MO, USM). Junin: Satipo, Pichanaki, Leon 222

(F, USM). Cuzco: Prov. Paucartambo, Quebrada Quita

Calzon, Leon et al. 2936 (USM). Achirani, Vargas 11147

(F).

7. Selaginella seemannii Baker, J. Bot. 21: 244.

1883. TYPE: Colombia, Choco, Cacaqual Is-

land, Seemann 1006 (holotype, K; isotype, BM).

Main stem assurgent at the tip, ca. 5-25 cm long,

not articulate, stramineous, glabrous, the basal part

with spreading leaves, and often smaller, ap-

pressed, ascending leaves. Primary branches 2-3-

pinnate, arising a short distance above the base of

the main stem, sometimes gradually reduced,

forming a flabelliform arrangement, the ultimate

leafy branches short, unequal, 4-6 mm broad in-

cluding the leaves. Rhizophores mostly near the

base to the mid-portion of the main stem. Lateral

leaves elongate-ovate, broadest at the base,

spreading above the first branch, approximate,

imbricate toward the apex, 2-3 mm long, mem-

branous, acute, the base truncate or 1 side enlarged

and rounded, with narrow, whitish, denticulate

edges, or sparsely denticulate along the base. Me-

dian leaves elongate-ovate, the apex acuminate to

subaristate, the arista less than '/4 the lamina length,

the base truncate or with an enlarged, rounded

outer side, with narrow, whitish, denticulate edges,

dentate at the base.

On open floor of dense forest, in deep ravines,

or dense, seasonally flooded forest, 130-625 m,

Loreto and Huanuco.

Costa Rica, Colombia, Suriname, Ecuador, and

Peru.

The flabelliform arrangement of the branches,

with rhizophores concentrated at the base of the

stem, distinguishes this from other slender-

stemmed species such as S. producta and S. re-

voluta that clearly are prostrate, creeping.

Although the species is reported to be wide-

ranging from Costa Rica to Peru, we have seen

collections only from Loreto, and Huanuco in Peru.

Loreto: Prov. Maynas, Calentura, Killip & Smith 29141

(us). Prov. Maynas, Iquitos, Tryon & Tryon 5195 (F, GH,

us, USM). Prov. Maynas, Yanamono Tourist Camp, van

der Werffet al. 9959 (uc). Rio Itaya, LI. Williams 238

(F, us). Huanuco: Tingo Maria, La Cueva de las Pavas,

Allard 20514 (us).

8. Selaginella producta Baker, J. Bot. 21: 243.

1883. LECTOTYPE (chosen by Alston et al.,

1981): Brazil, Amazonas, between Barcellos

and San Gabriel, Spruce 2043 (holotype, BM;

isotype, CGE).

Main stem prostrate or assurgent, 1 0-20 cm long,

not articulate, several stems usually arising at the

base of the plants, stramineous to somewhat green-

ish, glabrous, the basal part with spreading leaves

and often smaller, appressed, ascending leaves.

Primary branches 2-3 -pinnate, with distant sec-

ondary divisions spreading, sometimes forming a

broad, flabelliform arrangement, or with elongate,

alternatively pinnate branches as long as the main

stem or nearly so, 5-10 mm broad including the

leaves, the ultimate leafy branches short, unequal

in length. Rhizophores mostly at the stem base, or

extending to the central part of the main stem and

branches. Lateral leaves oblong to ovate, spread-

ing, approximate to somewhat imbricate above

the first branch, 2-5 mm, usually 4 mm long,

broader at the base, obtuse to subacute, the base

truncate, or with an enlarged, round basal lobe,

without conspicuous whitish borders, the edges

sparsely denticulate with the teeth somewhat dens-

er at the base. Median leaves suborbicular, broad-

est at the center, the apex acuminate to subaristate

and cuspidate, the arista usually V* or less the lam-

ina length, the base truncate, with or without nar-

row, whitish margins, the edges evenly denticulate.

Terrestrial, or a trunk epiphyte, on rotted logs,

or commonly on white sand of the forest floor,

100-860 m, Loreto, Huanuco, and Pasco.

74

FIELDIANA: BOTANY

Tobago and Trinidad, French Guiana west to

Colombia and south to Peru and Brazil.

The species is compared to Selaginella revoluta

by Alston et al. (1981). However, it differs in

broader lateral leaves that have a truncate base

and lack cilia. The imbricate leaves covering most

of the branches resemble those of S. truncata but

are clearly broader and may have a whitish sur-

face.

Loreto: Prov. Maynas, Iquitos, Killip & Smith 27322

(GH, us). Iquitos, Klug 198 (F, us). Iquitos, Revilla 4285

(F, MO). Iquitos, McDaniel et al. 22107 (F). Mishana,

Solomon 3595 (MO). Prov. Maynas, Alpahuayo, van der

Werff et al. 10262 (F, MO). Huanuco: Rio Llullapichis,

Cerros del Sira, Wolfe 12264 (GH). Pasco: Prov. Oxa-

pampa, Palcazu, Foster 9532 (MO, USM).

9. Selaginella microphylla (HBK.) Spring, Bull.

Acad. roy Sci. Bruxelles 10: 234. 1843.

Lycopodium microphyllum HBK., Nov. gen. sp. 1: 37.

1816. TYPE: Colombia, Cauca, Qulcace, Bon-

pland (holotype, p?, photo, GH; isotype, BM).

Main stem short, creeping, usually it and its

branches forming a compact mat, not articulate,

stramineous or greenish, glabrous. Primary

branches often intricate, 1-2-pinnate, short, dis-

tant, extending to the base of the main stem, slen-

der, ca. 0.5 mm broad, including the leaves. Rhi-

zophores abundant at the base, extending nearly

to the apex of the main stem and branches. Lateral

leaves extending nearly to the base of the stems,

minute, inserted obliquely, 0.25-0.50 mm long,

ovate or ovate-elliptical, tightly clasping the stem

and enveloping the median leaves when dry, im-

bricate, less so on the ultimate branches, the apex

acute with a more or less extended cusp, the bor-

ders whitish, ciliolate to fimbriate along the edges,

especially at the base. Median leaves ovate, about

half as long as the lateral leaves, subacute, not

auriculate, the edges with regular, dense cilia, the

border broad, whitish.

Terrestrial in open forest, on soil or among damp

rocks, in deep shade, 800-2700 m, Cajamarca, La

Libertad, Huancavelica, Apurimac, and Cuzco.

Costa Rica and Panama, Venezuela along the

Andes south to Argentina, east to southern Brazil,

Paraguay and Uruguay.

The abundant cilia along the borders of the

leaves, and especially the lateral leaves that strong-

ly clasp the dry stems, readily characterize the spe-

cies. The slender, intricate stems have a growth

form that resembles some species in the Selagi-

nella rupestris group.

The species is known from a broad geographic

range through South America and has been widely

collected in Peru. Specimens from southern Co-

lombia collected by Bonpland on his travels in

South America with Humboldt represent the type.

Cajamarca: Prov. Contumaza, El Tunel, Sagdstegui et

al. 12565 (MO). La Libertad: Prov. Pataz, Vista Florida,

Leon & Young 1086 (USM). Huancavelica: Prov. Tuya-

caja, Surcubamba, Tovar 3696 (GH). Apurimac: Prov.

Abancay, Vargas 16584 (GH). Cuzco: Prov. Convention,

Sahuayaco, Biles 834 (us). Urubamba Valley, San Mi-

guel, Cook & Gilbert 1782 (us).

10. Selaginella ramosissima Baker, J. Bot. 23: 295.

TYPE: Peru, San Martin, near Tarapoto,

Spruce 4088 (holotype, K; isotypes, BM, GH,

NY).

Main stem erect, or assurgent at the tip, short,

less than 10 cm long, not articulate, greenish, gla-

brous, the basal part with spreading leaves and

often smaller, appressed, ascending leaves. Leafy

branches 2-pinnate, ascending, longest in the cen-

tral part of the stem, the secondary branches

straight, ascending, forming a short, deltate ar-

rangement of the stems, ca. 3 mm broad. Rhizo-

phores mostly at the base extending to the first

branch. Lateral leaves slender, elongate, broadest

at the center, closely placed to somewhat imbricate

especially on the secondary branches, oblong, 0.5-

1 .0 mm long, membranous, acute, the base trun-

cate, not prolonged, with narrow, whitish borders,

the edges denticulate especially at the base. Me-

dian leaves slender, elongate-ovate, or somewhat

broader at the center, membranaceous, acuminate

to aristate, the arista '/4 or more the length of the

lamina, the base more or less equal, without a

prolonged auricle, the borders narrow, whitish, the

edges denticulate. Strobilus dorsiventral with 2

ranks of sporophylls on the upper surface of the

leafy stem.

Along roadside or bank of stream below the

forest, 200-2700 m, Cajamarca, San Martin, Lima,

and Cuzco.

Ecuador and Peru.

The strobilus is dorsiventral with two ranks of

sporophylls on the upper part of the leafy stem.

This readily distinguishes the species from all oth-

ers in Peru. The thinner, membranaceous texture

of the leaves also is a characteristic of the Peruvian

plants.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

75

Cajamarca: Prov. Hualgayoc, Soukup 3872 (us). Lima:

Prov. Cantua, Acleto 758 (USM). Cuzco: Biies in Junio,

1930(F).

1 1 . Selaginella difTusa (Presl) Spring, Bull. Acad.

roy. Sci. Bruxelles 10: 143. 1843.

Lycopodium diffusum Presl, Reliq. haenk. 78. 1825.

TYPE: Panama, Haenke (holotype, PR; isotype,

B).

Selaginella eggersii Sodiro, Crypt, vase. Quit. 605.

1893. TYPE: Ecuador, Valle Takatanga, Sodiro

(holotype, not found; isotype, us).

Selaginella atirrensis Hieron., in Engler & Prantl, Nat.

Pflanzenfam. 1 (4), 71 1. 1 90 1 . LECTOTYPE: (by

Alston, 1955), Costa Rica, Cartago, near Atirro,

Donn. Smith 5103 (holotype, us; isotype, NY).

Main stem elongate, prostrate, assurgent at the

tips, sometimes compact and matlike, articulate,

stramineous, glabrous, the basal part with spread-

ing leaves, and often smaller, appressed, ascending

leaves. Primary branches distant, ascending, un-

equally forked, 2-3-pinnate, extending to or nearly

to the base of the main stem, 2-4 mm broad in-

cluding the leaves, the ultimate branches short.

Rhizophores hairlike, extending nearly to the apex

of the branches. Lateral leaves spaced or approx-

imate, not imbricate, 1-2 mm long, elliptical-lan-

ceolate, acute, short, with a prolonged auricle, of-

ten densely ciliolate, sometimes with whitish

borders, the edges sparsely ciliolate to nearly en-

tire. Median leaves elongate-ovate, broadest near

the middle, the apex long-acuminate, the base with

a prolonged, Ungulate auricle that is usually cilio-

late, with or without a whitish border, the edges

finely denticulate.

In moist places in deep forest, on steep banks,

on logs or among rocks, sometimes forming large

mats, 1100-2100 m. Huanuco, and Pasco south

to Cuzco.

Costa Rica; Trinidad; South America east to

Suriname, and in the Andes south to Bolivia.

Plants appear to have lateral leaves of more than

one form. Most abundant are slender, elliptical

leaves with entire margins. Other, larger leaves

are broader with the base abundantly ciliolate, and

the cilia sometimes extending along the margins

to the mid-region of the leaf. The ciliolate base

resembles that of the median leaves.

Selaginella atirrensis Hieron. is included here

although it has been recognized as a distinct spe-

cies on the basis of its acute median leaves. It is

not easily distinguished on leaf shape and possibly

intergrades with S. diffusa.

Huanuco: Tingo Maria, Croat 21245 (MO, uc).

Churubamba, Mexia 8256 (F, GH, MO, uc, us). Tingo

Maria, Tryon & Tryon 5227 (F, GH, us, USM). Junin: near

La Merced, Killip & Smith 23949 (F, GH). Chanchamayo

valley, Schunke 197 (F). Tarma, Pan de Azucar, Velarde

5490 (GH). Pasco: Prov. Oxapampa, between Oxapampa

and Paucartambo, D. Smith 1466 (MO). Ucayali: Bo-

queron, Ferreyra 8128 (USM). Cuzco: Prov. La Conven-

cion, Rio Mapitunuari, Dudley 11352 (GH, MO). Kos-

nipata, Sta. Isabel, Vargas 23011 (GH).

12. Selaginella lingulata Spring, Nouv. Mem.

Acad. roy. Sci. Belg. 24: 224. 1849. TYPE:

Ecuador, Pichincha, Jameson (holotype, K;

isotype, BM).

Selaginella intacta Baker, J. Bot. 21: 335. 1883. TYPE:

Ecuador, San Nicolas, Sodiro (holotype, K).

Main stem prostrate, intricate, usually compact,

forming a dense mat, articulate, stramineous, gla-

brous, the basal part with spreading leaves and

often smaller, appressed, ascending leaves. Leafy

branches 1-3-pinnate, usually pinnately branched,

distant, extending to the base of the main stem,

the ultimate branches short, 2-4 mm broad. Rhi-

zophores extending nearly to the stem apex. Lat-

eral leaves closely placed to distant, 1-3 mm long,

spreading on the upper part of the stems and pen-

ultimate branches, oblong, acute, the basal lobes

more or less equal, not prolonged, with long, dense

cilia, without whitish borders, the edges entire.

Median leaves elongate-ovate, the apex usually

long-acuminate, the base with 2 equally long au-

ricles, ciliate at the base, without conspicuously

whitish borders, the edges denticulate above.

On soil or among rocks, in shade, sometimes

forming dense mats on hillsides or on steep banks,

sometimes in deep forests, 1000-1300 m, Pasco

and Junin.

Colombia south to Bolivia.

The long marginal cilia at the base of the leaves

form a small fringe especially on the lateral leaves.

The median leaves are clearly more slender than

the lateral ones and have a long-acuminate apex

in the material examined. However, the apex of

the median leaves was considered as acute in the

treatment by Alston et al. (1981).

The axillary leaves are distinctive, somewhat

larger than the median and lateral leaves, and more

or less ovate, acute, usually with one prolonged

auricle and few long cilia.

Pasco: Pichis trail, San Nicolas (as Junin), Killip &

Smith 26056 (F, GH, us). Junin: Above San Ramon,

76

FIELDIANA: BOTANY

Schunke A251 (us), Chanchamayo Valley, Schunke 230

(us), 755 (us). Prov. Tarma, San Ramon, Tryon & Tryon

5448 (F, GH, us).

13. Selaginella novae-hollandiae (Sw.) Spring,

Bull. Acad. roy. Sci. Bruxelles 10: 234. 1843.

Lycopodium novae-hollandiae Sw., Syn. fil. 184, 410.

1806. TYPE: "Nova Hollandiae," probably an

error for Nova Granada, not located.

Selaginella pearcei Baker, J. Bot. 22: 246. 1884. TYPE:

Peru, Huanuco, Cordilleras of Pozuzo, Pearce 249

(holotype, K).

Selaginella chionoloma Crabbe & Jermy, Amer. Fem

J. 63: 137. 1973. TYPE: Peru, Cuzco, Cuquipata,

Herrera 1636 (holotype, us; isotype, BM).

Main stem short, erect or assurgent, usually less

than 20 cm long, not articulate, stramineous, gla-

brous, the basal part with spreading leaves and

often smaller, appressed, ascending leaves, or the

basal part with only appressed leaves. Primary

branches 2-3-pinnate, often forming a complex

system of branches broadest at the base, the pen-

ultimate branches straight or nearly so, 1-3 mm

broad including the leaves. Rhizophores at base of

the main stem, mostly below the first branch. Lat-

eral leaves 0.5-2.0 mm long, elliptical to some-

what ovate, acute, not auriculate, the base cilio-

late, sometimes with narrow whitish borders, the

edges ciliolate, often densely so. Median leaves

ovate, usually more or less elongate, the apex acu-

minate to aristate, the arista '/4 the lamina length,

the base with a prolonged, Ungulate, ciliolate au-

ricle, or a somewhat enlarged, round, ciliate base,

the borders narrow, whitish, the edges ciliolate or

denticulate.

On damp, shaded ledges or crevices of rock fac-

es, or moist banks along streams, wooded hillsides,

open woods, dense forests, sometimes carpeting

the forest floor, rarely epiphytic, 400-3600 m,

Lambayeque, south to Puno.

Nicaragua south to Bolivia and Argentina.

This differs from other slender-stemmed species

in lacking articulations on the stems and having

generally smaller leaves. It is one of the most com-

mon species of tropical America and is wide-rang-

ing in Peru.

Lambayeque: Along Olmos-Jaen road, Correll & Smith

P826 (GH, MO, us). Cajamarca: El Molino, Sagdstegui et

al. 7995 (F, MO, uc). Amazonas: Prov. Bagua, Chacha-

poyas, Hutchison & Wright 5782 (F, GH, MO, uc, us, USM).

La Libertad: Prov. Otuzco, road to Paranday, Lopez

1044 (us). San Martin: Rio Sion, Schunke 3494 (F, us).

Loreto: Balsapuerto, Killip & Smith 28488 (F, GH). Hua-

nuco: Tingo Maria (as San Martin), Allard 21130 (MO,

uc). La cueva de las Pavas, Schunke 3257 (F, GH, us).

Lima: Matucana, Bryan 48 (F, MO). Junin: Prov. Tarma,

San Ramon, H. Iltis & C. Iltis 18 (GH, wis). E of Quimiri

Bridge, Killip & Smith 23945 (F, GH). Ucayali: Boqueron

Padre Abad (as Loreto), Schunke 3065 (F, GH, us). Huan-

cavelica: Surcubamba, Tovar 3682 (GH). Ayachucho:

Ccarrapa, Killip & Smith 22331 (F, GH, us). Apurimac:

Prov. Abancay, Vargas 11083 (GH). Cuzco: Chincheros,

King et al. 293 (USM). Machu Picchu, Leon 457 (USM).

Puno: Prov. Carabaya, Ollachea, Vargas 6914 (us).

14. Selaginella xiphophylla Baker, J. Bot. 22: 296.

1884. TYPE: Peru, San Martin, Mt. Guay-

rapurima, near Tarapoto, Spruce 3990 (ho-

lotype, K; isotypes, BM, CGE, GH, NY).

Main stem erect, ascending, 10-14 cm long, not

articulate, stramineous to greenish, glabrous, the

basal part with appressed, ascending leaves. Pri-

mary branches 1-2-pinnate, alternate, distant, ex-

tending to, or nearly to the base of the main stem,

the lower branches usually longest, the penulti-

mate leafy branches 6-12 mm broad. Rhizophores

confined to the lower Vs-'/z of the main stem, rarely

above. Lateral leaves ascending to somewhat ap-

pressed, those above the first branch approximate,

2-6 mm long, oblong-lanceolate, the apex acute,

the base truncate, with denticulate edges especially

at the base. Median leaves ovate-lanceolate, grad-

ually attenuate, aristate, the arista as long as the

lamina, the base truncate, the outer side ciliolate,

with narrow, whitish denticulate margins, the teeth

extending onto the arista.

Panama, Colombia, and Peru.

We have seen only a single collection, the type

material from San Martin listed above. The spe-

cies is also reported from Panama and Colombia

by Valdespino (1993). The small size, erect habit

of the plants, the rhizophores borne in the basal

l/3-l/2 of the main stem, and median leaves with

an arista as long as the lamina are characteristics

by which the species can be recognized.

15. Selaginella flagellata Spring, Bull. Acad. roy.

Sci. Bruxelles 10: 228. 1843. TYPE: French

Guiana, Rio Inini, "source of the Rio Oya-

pok," Leprieur (holotype, LG; isotype, P).

Selaginella regularis Baker, J. Bot. 22: 277. 1884.

TYPE: Peru, San Martin, near Tarapoto, Spruce

3977 (holotype, K; isotypes, BM, GCE).

Main stem erect, short, 5-20 cm long, the base

2-3 cm below the leafy branches, not articulate,

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

77

stramineous, glabrous, the basal part with spread-

ing leaves, and often smaller, appressed, ascending

leaves. Primary branches 2-3-pinnate, sessile or

subsessile, distant, spreading, the central ones

longest, the lateral branches ascending in a broad,

frondlike arrangement, the penultimate leafy

branches straight, 3-5 mm broad. Rhizophores

basal, or mostly below the lowest branches. Lat-

eral leaves 1-3 mm long, elliptic, or broadest at

the base, attenuate to acute, the base truncate, the

borders not conspicuously whitish, the edges den-

ticulate, Median leaves ovate to somewhat elon-

gate, aristate, the arista l/4 as long as the lamina,

the base not auriculate, the borders narrow, whit-

ish or not, the edges denticulate, especially at the

base.

Terrestrial, or climbing in cloud forest, among

moist rocks and on cliffs, 200-2000 m, San Mar-

tin, Loreto, and Cuzco.

Mexico south to French Guiana and Brazil, and

in the Andes to Bolivia.

The short, ultimate branches clearly differ from

other slender, erect-stemmed species in Peru. There

is a general resemblance to Selaginella xiphophylla

in the erect habit and rhizophores confined to the

basal part of the main stem.

San Martin: Juan Jui, Klug 4230 (F, GH, us). Loreto:

Pongo de Manseriche, Mexia 6333A (uc). Cuzco: Prov.

Convention, Echarate, Sues 787 (us).

16. Selaginella bombycina Spring, Nouv. Mem.

Acad. roy. Sci. Belg. 24: 191. 1849. TYPE.

Peru, San Martin, Matthews (Mathews) 1781

(holotype, K).

Main stem erect, 25-30 cm long, 1 -pinnate, not

articulate, stramineous, glabrous, the basal part

with appressed, ascending leaves. Primary branch-

es alternate, mostly 1 -pinnate, extending in nearly

parallel alignment from the first branch to the apex

of the stem, 10-16 mm broad including the leaves,

the axes straight or nearly so. Rhizophores largely

at the base of the main stem, rarely with a few

among the main branches. Lateral leaves imbri-

cate on the lateral branches, oblong, slender 5-10

mm long, 4 or 5 x longer than broad, obtuse, the

base with a slightly enlarged, round lobe, usually

densely ciliate, without conspicuous whitish bor-

ders. Median leaves suborbicular, asymmetric, the

outer side larger, aristate, the arista '/2 to equal the

length of the lamina, with a somewhat longer basal

auricle, the base ciliolate, without conspicuous

whitish borders, cilia often dense along the outer

edges.

On hillsides and deep ravines of wet forests,

700-1 100 m, San Martin and Huanuco. In Peru

nearly restricted to the area around Tingo Maria.

Costa Rica south to Peru.

The plants usually have long primary branches

that are 1 -pinnate. This feature and the ciliate lat-

eral leaves distinguished the species.

San Martin: Prov. Mariscal Caceres, 4 km de Puerto

Pizana, Schunke 4893 (F, us). Huanuco: Tingo Maria,

Tryon & Tryon 5255 (GH, us). Prov. Leoncio Prado,

Tingo Maria, Plowman & Kennedy 5732 (F, GH, USM).

Prov. Leoncio Prado, Rupa Rupa, Schunke 10172 (F,

MO).

17. Selaginella chrysoleuca Spring, Bull. Acad.

roy. Sci. Bruxelles 10: 226. 1843. TYPE: Bo-

livia, D'Orbigny, (holotype, P).

Selaginella sprucei Hooker, Sec. cent, ferns, t. 83. 1861.

TYPE: Peru, San Martin, Mt. Campana, near

Tarapoto, Spruce 4623 (holotype, K; isotypes, BM,

CGE, us).

Main stem below the first branch short, usually

less than 5 cm long, erect, not articulate, greenish,

glabrous, the basal part with spreading leaves and

often smaller, appressed, ascending leaves, or the

basal part only with appressed, ascending leaves.

Leafy branches dichotomously divided 1 or 2 x ,

often with 1 longer branch, sometimes with tuft-

like branches arising from a slender creeping run-

ner, ultimate leafy branches 5-20 mm, mostly 1 5

mm broad, axes of primary branches straight. Rhi-

zophores mostly at the base, sometimes extending

to l/3 the length of the stem, but not among the

broad, leafy branches. Lateral leaves spreading

above the first branch, imbricate, 5-10 mm long,

oblong to linear, 3-5 x longer than broad, the apex

acute to obtuse, asymmetrical, with the upper (ac-

roscopic) edge somewhat larger, round, and finely

denticulate, with narrow whitish borders, the edg-

es finely denticulate. Median leaves elongate-ovate

to suborbicular, asymmetrical, acuminate to aris-

tate, the arista sometimes Vz the lamina length, the

base truncate with a slightly enlarged, round lobe,

the borders narrow, whitish, the edges finely den-

ticulate.

Among mosses on rocks, in damp soil, on shad-

ed slopes of ravines, on moist riverbanks, and in

dense, dark cloud forests, 400-2000 m, Amazo-

nas, south to Cuzco.

78

FIELDIANA: BOTANY

Panama, Venezuela to Bolivia.

Similarities in the general aspect of the broad,

leafy branches, in the asymmetrical shape of the

median leaves with a long arista and dentate mar-

gins, suggest an alliance with Selaginella speciosa.

Amazonas: Prov. Bagua, La Peca, Barbour 2839 (MO).

Prov. Bagua, near Montenegro, Hutchison & Wright 3821

(F, GH, uc, us). Above Montenegro, Wurdack 1879 (F,

GH, uc, us). San Martin: Road from Tarapoto to Yu-

rimaguas, Kennedy 3544 (F, us). Between Tarapoto and

Yurimaguas, Knapp & Mallet 8474 (F, MO). Tarapoto-

Yurimaguas road, McDaniel 13805 (GH, MO). Near Tara-

poto, Spruce 4628 (GH). Ucayali (as Loreto): Between

Tingo Maria and Pucallpa, Ferreyra 1008 (USM). Huanu-

co: Cerros del Sira, Rio Llullapichis Watershed, Dudley

13360 (GH). Cuzco: Prov. La Convention, Dudley 10355

(GH, MO).

18. Selaginella speciosa A. Braun, Ann. Sci. Nat.

Bot. 5, 3: 274. 1865. TYPE: Colombia, Bo-

gota, Triana (holotype, B?; isotype, BM).

Selaginella huberi Christ, Bull. Herb. Boissier 2, 1:

73. 1901. TYPE: Peru, "entre Ucayali et Hualla-

ga," Huber 1547 (holotype, P).

Main stem erect, 30-70 cm long, not articulate,

green to brownish, glabrous, the basal part with

appressed, ascending leaves. Primary branches di-

chotomous, long, 1 -pinnate, with 2 or more long

divisions, the lower ones divided at short intervals

forming a frondlike, somewhat flabelliform ar-

rangement, 10-20 mm broad including the leaves,

the axes straight. Rhizophores at the base of the

main stem. Lateral leaves closely placed to im-

bricate, linear-oblong, to broader above the base,

8-10 mm long, 5-6 x longer than broad, asym-

metrical, the acroscopic side larger, the apex ob-

tuse, the base truncate, the borders sometimes

whitish at the base, the edges finely denticulate.

Median leaves orbicular to somewhat ovate,

asymmetric with the outer side larger, acuminate

to aristate, the arista often nearly '/2 as long as the

lamina, the auricles not prolonged, borders nar-

row, whitish, the edges finely denticulate.

In deep, shady forests, in rain forests, in moist

ravines, rarely epiphytic, 100-600 m, Amazonas

and Loreto.

Colombia, Ecuador, and Peru.

This and Selaginella chrysoleuca have excep-

tionally broad leafy branches and asymmetrical

median leaves that are long aristate. Both species

occur in dense forest in the Andes, but S1. speciosa

occurs mainly in the north of Peru while S. chry-

soleuca extends south to Cuzco.

Amazonas: Huampami, Kayap 1311 (uc). Loreto: Be-

tween Yurimaguas and Balsapuerto, Killip & Smith 28340

(F, us). Balsapuerto, Klug 2921 (F, GH, MO, us). Prov.

Maynas, Iquitos, McDaniel & Rimachi 17473 (GH). Si-

erra del Pongo, Mexia 6283 (F, GH, MO, uc, us). Gamita-

nacocha, Rio Mazan, Schunke 109 (F, GH, uc, us).

1 9. Selaginella lechleri Hieron., in Engler & Prantl,

Nat. Pflanzenfam. 1 (4): 683. 1901. LEC-

TOTYPE (chosen by Alston et al., 1981):

Peru, Puno, near San Gavan (San Gaban),

Lechler 2159 (holotype, B?; isotype, BM).

Main stem erect, 1 2-40 cm long, not articulate,

stramineous, glabrous, the basal part with ap-

pressed, ascending leaves. Primary branches

3-pinnate, forming an extended, flabelliform ar-

rangement, distant, the lower stalked, those above

sessile or nearly so, penultimate leafy branches

straight or nearly so, ultimate leafy branches slen-

der, 3-5 mm broad. Rhizophores confined to the

base of the main stem. Lateral leaves of the main

stem, above the first branch, ascending, coarse,

mostly strongly imbricate, 2-5 mm long, oblong,

acute, the auricles not prolonged, without con-

spicuous whitish borders, the edges denticulate to

short ciliate, the cilia denser at the base. Median

leaves elongate, ovate, acuminate, the base some-

times unequal, with a slightly enlarged auricle, not

or sparsely ciliolate, without conspicuous whitish

borders, the edges slightly denticulate or entire.

Terrestrial on clay or sandy soil, along trails or

on stream banks in primary forest or rain forests,

1 30-1 200 m, Amazonas and Loreto south to Puno.

Colombia and Peru.

This taxon seems to intergrade with Selaginella

anceps but is maintained here in its traditional

sense. The broad, erect stems with appressed leaves,

the primary branches forming a broad, flabelli-

form arrangement, and small size of the leaves on

the ultimate branches are similar to those of S.

anceps.

Amazonas: E of Cinkan, Berlin 254 (F, MO, uc). Loreto:

Prov. Maynas, Puca Urquillo, Plowman et al. 6628 (F,

us). La Victoria, LI. Williams 2919 (F, us). Pasco: Prov.

Oxapampa, between Puerto Bermudez and San Matias,

Leon et al. 333 (F, USM). Puerto Bermudez (as Junin),

Killip & Smith 26467 (F, GH). Cuzco: between Mistiana

and Keros, Scolnik 868 (uc). Madre de Dios: Alto Madre

de Dios, Rauh & Hirsch PI 570 (GH). Puno: Prov. Caraba-

ya, Qllachea a Quillabamba, Vargas 17539 (GH).

20. Selaginella anceps (Presl) Presl, Abh. Bonn.

Ges. Wiss. 5, 3: 581. (Bot. Bemerk., 151).

1844.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

79

Lycopodium anceps Presl, Reliq. haenk. 1: 80. 1825.

TYPE: "Luzon," but probably from Peru, Haenke

(holotype, PR, photo, GH).

Lycopodium gracile Poiret, in Lam., Encycl. suppl. 3:

551. 1814. TYPE: Peru (Holotype, Herb. Des-

vaux P, photo OH).

Selaginella gracilis (Poiret) Hieron., Hedwigia 58: 293.

1917, not Moore, 1886.

Main stem erect, 20-25 cm or longer, not artic-

ulate, 2-3.5 mm broad at the base, stramineous,

glabrous, the basal part with appressed, ascending

leaves. Primary branches 3-4-pinnate, forming an

elongate, flabelliform arrangement, distant, stalked,

the penultimate branches straight or nearly so, the

ultimate leafy branches slender, 1-2 mm broad.

Rhizophores confined to the base of the main stem.

Lateral leaves above the first branch, appressed,

2-3 mm long, elongate-ovate, broadest at the base,

acute, with a somewhat prolonged, ciliate auricle,

without conspicuous whitish borders, long ciliate

at the base of the leaf. Median leaves ovate-elon-

gate, acuminate with a somewhat prolonged au-

ricle or an enlarged, ciliolate basal lobe, the bor-

ders not conspicuously whitish, the edges usually

prominently ciliolate.

Disturbed forests, primary and secondary rain

forests, steep hillsides in wet forests, 1 50-1 500 m,

San Martin and Loreto to Cuzco and Madre de

Dios.

Costa Rica south to Venezuela and to Bolivia.

On the basis of similarities in the small, imbri-

cate leaves densely covering the ultimate branches

Selaginella anceps appears to be allied to 5. lech-

leri. However, the usually denticulate base of the

median leaves in S. lechleri is distinct from the

usually prominently ciliolate one in S. anceps.

San Martin: Between Tocache Nuevo and Juanjui,

Croat 58038 (F, MO). Loreto: Pongo Manseriche, Mexia

6371 (GH, MO, uc, us). Huanuco: Pampayacu, Kanehira

118 (GH, us). Pasco: Prov. Oxapampa, Palcazu, Foster

& d'Achille 10177 (USM). Junin: Rio Perene, Killip &

Smith 25104 (F, GH, us). Perene, Seibert 2193 (F, MO,

us). Ayacucho: Estrella, between Huanta and Rio Apu-

rimac, Killip & Smith 22671 (F, GH, us). Cuzco: Prov.

Convencion, Dudley 11381 (GH, MO). Madre de Dios:

Prov. Manu, Atalaya, Foster & Wachter 7456 (MO).

2 1 . Selaginella erythropus (Mart.) Spring, in Mar-

tius, Fl. bras. 1 (2): 125. 1840.

Lycopodium erythropus Martius, Icon. pi. crypt. 39.

1 834. TYPE: Brazil, Piaui, Martius (holotype, BR).

Plants 10-25 cm, mostly 10-15 cm long. Main

stem erect and simple below the lowest, well-de-

veloped branch, not articulate, reddish, glabrous,

the basal part with appressed, ascending leaves.

Primary branches often irregular, mostly 2-pinnate,

ultimate branches not exceeding 5 mm broad, in-

cluding the leaves. Rhizophores usually confined

to the base of the main stem, reddish. Lateral

leaves strongly imbricate, reddish, especially the

borders, the color extending into the marginal cil-

ia, elongate-oblong, 0.25-0.5 mm long, the acro-

scopic side somewhat larger, acuminate (not aris-

tate), with a rounded, denticulate basal lobe, the

upper edges ciliate or ciliolate. Median leaves

elongate-ovate, strongly imbricate, somewhat

asymmetrical, acute to aristate, the base more or

less truncate, the borders whitish, the edges den-

ticulate, somewhat denser at the base.

On moist riverbanks, steep, wet road banks,

thickets in dense forests, sometimes forming dark

green mounds on the floor of lowland rain forests,

250-1800 m, Amazonas south to Cuzco.

Costa Rica; Colombia to Bolivia; Brazil.

Among the species with erect main stems, Se-

laginella erythropus is distinguished by the reddish

color of the main stem that often extends into the

branches and rhizophores, and by the small size

of the plants. It also differs from S. haematodes

by the ciliate or ciliolate lateral leaves.

Amazonas: Prov. Bagua, between Aramango and

Montenegro, Lopez et al. 4212 (GH, MO). San Martin:

Juan Jui, Klug 4267 (GH, MO, us). Tarapoto, Spruce 3989

(GH, us). Junin: Colonia Perene, Killip & Smith 24918

(F, GH, us). La Merced, Macbride 5549 (F, us). Ayacucho:

Estrella, between Huanta and Rio Apurimac, Killip &

Smith 22664 (F, us). Cuzco: Prov. La Convencion, Dud-

ley 11438 (GH).

22. Selaginella haematodes (Kunze) Spring, in

Martius, Fl. bras. 1 (2): 126. 1840. Figure

9c-d.

Lycopodium haematodes Kunze, Linnaea 9: 9. 1834.

TYPE: Peru, San Martin, Mission Tocache, Poep-

pig, in June 1830 (isotype, K).

Selaginella filicina Spring, Nouv. Mem. Acad. roy.

Sci. Belg. 24: 189, 1849. SYNTYPE: Peru, Mat-

thews (Mathews) 40 36 (K). Other collections from

Venezuela are also cited.

Plants 30-65 cm long. Main stem erect, not

articulate, bright red below the branches, glabrous,

the basal part with appressed, ascending leaves.

Primary branches usually red, ultimate branches

3-8 mm broad including the leaves, the basal part

80

FIELDIANA: BOTANY

with appressed, ascending leaves. Rhizophores near

the base of the main stem, deep red. Lateral leaves

often reddish, oblong to elongate-ovate, broader

at the base, 0.25-2.0 mm long, acute, the base

truncate, without conspicuous whitish borders, the

upper (acroscopic) edge finely denticulate, the teeth

longer toward the apex, or nearly entire. Median

leaves slender, elongate-ovate, acuminate to aris-

tate, the arista '/4 or less the lamina length, the base

truncate, with narrow, whitish borders, the edges

denticulate especially near the apex.

Frequent along riverbanks, on steep, moist hill-

sides, or scattered on floor of dense forests, pri-

mary forests, rocky hillsides, or along trails, 135-

900 m, San Martin and Loreto, south to Madre

de Dios.

Panama, and Venezuela south to Bolivia.

The large, deep green, branches borne on long,

simple, red-colored stems, readily distinguish the

species. The vivid red color extends into the base

of the main branches and also to the rhizophores

that are confined to the basal part of the main

stem.

San Martin: Tocache Nueve, Schunke 7077 (F, us).

Prov. Mariscal Caceres, Uchiza, Schunke 3202 (F, GH,

us). Tarapoto, Spruce 4036 (GH, us). Loreto: San An-

tonio, Rio Samira, Ayala & Arevalo 4227 (F, MO). Puerto

Arturo, Killip & Smith 27747 (F, GH, us). Cueva de las

Pavas, Fukushima 6521 (F, GH, MO). Huanaco: Prov.

Huanuco, Mexia 8302 (GH, MO, uc). Pasco: Pichis Trail,

Killip & Smith 26214 (F). Junin: La Merced, Soukup

2369 (F, GH). Madre de Dios: Cocha Cashu, lake of Rio

Manu, Foster et al. 3417 (F).

23. Selaginella quadrifaria Alston et al., Bull. Brit.

Mus. (Nat. Hist.) Bot. 9: 261. 1981. TYPE:

Peru, Prov. Loreto, Sierra del Pongo, Mexia

6282 (holotype, BM; isotypes, GH, uc, us).

Main stem erect, 20-70 cm long, not articulate,

stramineous, glabrous, the basal part with ap-

pressed, ascending leaves. Primary branches 3-4-

pinnate, distant, ascending, usually dichotomously

divided, forming a broad, flabellate arrangement of

ultimate branches, 8-20 mm broad including the

leaves, the axes straight. Rhizophores confined to

the base of the main stem. Lateral leaves coarse,

rigid, imbricate, 4-8 mm long, oblong or broader

near the base, equilateral, with a central costa, the

apex acute, the base truncate, with an enlarged,

rounded, ciliolate lobe, without conspicuous whit-

ish borders, the edges denticulate. Median leaves

ovate, equilateral, with a central costa, acute, the

base truncate, the edges sometimes narrow, whit-

ish, finely denticulate.

In dense forest or rain forest, on slopes, on well-

drained soils, 450-860 m, Amazonas to Pasco.

Colombia and Peru.

Among related species with an erect stem and

appressed leaves Selaginella quadrifaria is gener-

ally distinguished by the median leaves that are

equilateral and not or hardly aristate. Differences

from S. praestans are noted under that species.

Amazonas: Prov. Bagua, near Montenegro, Hutchison

& Wright 382 IB (uc). Prov. Bagua, near Campamento,

Wurdack 1869 (GH, us). Huanuco: Cerro del Sira, Rio

Llullapichis, Wolfl2263B (F, us). Pasco: (as Junin) Puer-

to Yessup, Killip & Smith 26372 (F, us). Prov. Oxapam-

pa, valle del Palcazu, Iscozacin, Foster 4556 (F). Cabeza

de Mono, near Iscozacin, D. Smith 3752 (uc).

24. Selaginella praestans Alston et al., Bull. Brit.

Mus. (Nat. Hist.) Bot. 9: 260. 1981.

Selaginella sprucei A. Braun, Ann. Sci. Nat. Bot. 5,

3: 277. 1865, not Hooker 1861. TYPE: Peru, San

Martin, near Tarapoto, Guayrapurina, Spruce

4788 (not 4708 as originally cited) (holotype, G;

isotypes, BM, NY).

Main stem erect, 20-70 cm long, not articulate,

green to brownish, glabrous, the basal part with

appressed, ascending leaves. Primary branches 1-

2-pinnate, distant, ascending, sparingly pinnate,

the basal ones often longer, the axes straight, ul-

timate branches 10-15 mm broad including the

leaves. Rhizophores confined to the base of the

main stem. Lateral leaves above the first branches,

ascending, imbricate, those below coarse, elon-

gate-ovate to oblong, broadest at the base, 7-8 mm

long, 3 or 4 x longer than broad, inequilateral, the

acroscopic side larger, the costa acentric, acute to

obtuse, the base truncate, or with a slightly en-

larged, ciliolate auricle, without conspicuous whit-

ish borders, the edges delicately ciliolate, some-

times denticulate, Median leaves suborbicular to

somewhat ovate, asymmetric, inequilateral, the

outer side larger, acute, the base truncate, without

conspicuous whitish borders, the edges ciliolate or

entire.

In dense shade of deep woods or rain forests,

and along stream sides, 630-1500 m, San Martin,

Huanuco, and Pasco.

Colombia to Peru.

The species is similar to Selaginella quadrifaria

in the robust size of the plants and coarse leaves

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

81

appressed to the main stem. It can be distinguished

from that species by the inequilateral shape and

acentric costa of the lateral and median leaves in

contrast to the equilateral leaves and centric costa

in 5. quadrifaria.

San Martin: Valley of Huallaga, between Tocache

Nuevo and Juanjui, Croat 58057 (MO). Huanuco: Prov.

Huanuco, Cotirarda, Mexia 8196 (BM, GH, MO, uc, us).

Pasco: Prov. Oxapampa, between Iscozacin and Villa

America, D. Smith 2840 (F).

25. Selaginella haenkeana Spring. Bull. Acad. roy.

Sci. Bruxelles 10: 225. 1843. TYPE: probably

Peru, "cordilleris chilensibus," Hdenke (ho-

lotype, PR; photo, GH; isotypes, B, us).

Selaginella dimorpha Klotzsch, Linnaea 18: 523. 1844.

TYPE: "Auf den Anden von Chile," Hdenke (ho-

lotype, B; isotypes, PR, us). Probably the same

Peru collection cited above.

Main stem erect, up to 40 cm or more long,

usually unbranched Vs or more above the base of

the main stem, the base stout, 1-3 mm broad, not

articulate, greenish to stramineous, glabrous, the

basal part with appressed, ascending leaves. Pri-

mary branches 1 -pinnate, sessile or subsessile, dis-

tant, spreading, often arching upward, the central

ones usually longest, slender, the axes straight, ul-

timate branches not more than 5 mm broad in-

cluding the leaves. Rhizophores basal, or below

the lowest branch. Lateral leaves 2-3 mm long,

oblong, acute, without prolonged auricules, with

long, discrete cilia especially dense at the leaf base.

Median leaves ovate to somewhat elongate, aris-

tate, the arista slender, '/4 or less the lamina length,

the base not auriculate, the borders narrow, whit-

ish or not, the edges long-ciliate, especially long

and dense at the base.

Terrestrial, among rocks, or climbing, in cloud

forest and dense forest, 1 700-2000 m, Amazonas,

and Pasco.

Colombia, south to Bolivia.

Well-developed plants often have an unusual

branching system with a long, slender central stem

and widely spaced, 1 -pinnate lateral branches

bearing short, ultimate divisions.

Amazonas: Prov. Bagua, La Peca, Barbour 2820 (F,

MO, uc), 2967 (F, MO). Pasco: Pichis Trail, Dos de Mayo

(as Junin), Killip & Smith 25843 (F, GH).

26. Selaginella kunzeana A. Braun, Ann. Sci. Nat.

Bot. 5, 3: 296. 1865. LECTOTYPE (chosen

by Alston et al., 198 1): Peru, Huanuco, Pam-

payacu, Poeppig 195 (holotype, B; isotype, BM,

P).

Main stem prostrate long-creeping to assurgent,

articulate, stramineous, glabrous, the basal part

with spreading leaves and often smaller, ap-

pressed, ascending leaves. Primary branches 2-3-

pinnate, ascending, mostly widely distant, these as

well as the apex of the main stem forming a rhom-

boid arrangement, with an acute to acuminate apex,

ultimate branches 5-8 mm broad including the

leaves. Rhizophores extending above the lowest

branches, mostly in the lower half of the stem.

Lateral leaves above the first branch distant, wide-

spreading, imbricate at the stem apex, 2-3 mm

long, oblong, elongate with somewhat parallel sides,

acute, the base with a prolonged auricle forming

a round or slender lobe, with narrow, whitish, fine-

ly denticulate edges, the teeth especially dense at

the base. Median leaves slender, elliptic, the apex

long-acuminate to aristate, the arista to '/4 the lam-

ina length, the base prolonged with one, some-

times two, large, auricles, with narrow, whitish,

finely denticulate edges.

Trailing on steep banks, in dense woods, often

in disturbed areas, 500-1950 m, Huanuco, south

to Cuzco.

Mexico to Panama; Venezuela, Andes of Co-

lombia to Peru.

A widely distributed species occurring from

Mexico to Peru, with a broad range in Peru, often

in disturbed places. The rhomboid arrangement

of the terminal, leafy branches distinguishes this

from other articulate, prostrate species.

Huanuco: Prov. Huanuco (as San Martin), Tingo Ma-

ria, Allard 20361 (us); Asplund 12140 (us). Hacienda

Mercedes, Mexia 8197 (F, GH, MO, uc, USM). Pasco: south

of Pozuzo, Teppner 79/269 (us). Ucayali: Divisoria, Ce-

rro Azul, Tryon & Tryon 5272 (F, GH, us, USM). Aya-

cucho: Prov. La Mar, southwest of Hacienda Luisiana,

Dudley 11721 (GH). Estrella, between Huanta and Rio

Apurimac, Killip & Smith 22663 (GH, us). Cuzco: Prov.

Quispicanchis, Marcapata, Vargas 3178 (MO, us). Kos-

nipata, Velarde 7987 (GH).

27. Selaginella silvestris Asplund, Ark. f. Bot. 20 A

(7): 30. 1926. TYPE: Bolivia, Sur Yungas, El

Chaco, Asplund 1140 (holotype, UPS; frag., BM;

photo, GH).

Main stem prostrate, long-creeping to decum-

bent, articulate, stramineous, glabrous, the basal

part with spreading leaves, and often smaller, ap-

82

FIELDIANA: BOTANY

pressed, ascending leaves. Primary branches 2-3-

pinnate, often dichotomous, forming a broadly

ovate to obdeltate arrangement, the ultimate leafy

branches short, with a round to truncate apex, 4-

8, usually 5 mm broad including the leaves. Rhi-

zophores mostly at the base of the main stem,

extending above the first branch. Lateral leaves

distant, spreading above the first branch, 3-4 mm

long, oblong, somewhat broader near the base,

asymmetrical, the apex obtuse to acute, the auri-

cles not prolonged, without conspicuous whitish

borders, the edges denticulate, especially at the

base. Median leaves elongate-ovate, usually acu-

minate, or if aristate the arista less than 'A the

lamina length, the base unequal with 1 prolonged,

curved auricle that may be up to % the length of

the lamina, the borders not conspicuously whitish,

the edges prominently denticulate.

In humid forests, tropical rain forests, wooded

ravines or thickets, also among rocks, rarely on

rock walls or epiphytic, 650-3000 m, Cajamarca,

and Amazonas, south to Ayacucho and Cuzco.

Mexico south to Panama; Venezuela and Co-

lombia south to Bolivia.

The long-creeping stems with abundant rhizo-

phores and prolonged auricles on the median leaves

suggest an alliance with Selaginella trisulcata.

Cajamarca: Cueva San Andres, Velarde 6984 (GH).

Prov. Cutervo, Guta de las Huacharos, San Andres, Lopez

& Sagdstegui 5404 (GH). Amazonas: Prov. Chachapoyas,

Quebrada Molino, Wurdack 636 (F, GH, uc, us); Prov.

Bagua, 25 km E of La Peca, Barbour 2962 (F, MO). San

Martin: Prov. Mariscal Caceresa, Rio Abiseo Nacional

Park, Young & Leon 4957 (HUT, USM). Huanuco: Prov.

Huanuco, Carpish, Asplund 12842 (us). Pasco: Oxapam-

pa, Leon et al. 913 (F). Junin: Yucapata, Yaupi, Woyt-

kowski 6627 (MO, us). Huancavelica: Prov. Tayacaja,

Marcavalle, Tovar4754 (GH). Ayacucho: Ccarrapa, Killip

& Smith 22359 (us). Cuzco: Machu Picchu, Peyton &

Peyton 1320 (GH, MO); Nunez 7516 (MO, uc).

28. Selaginella trisulcata Asplund, Ark. f. Bot.

20A (7): 34. 1926. TYPE: Bolivia, La Paz, El

Chaco, Asplund 1482 (holotype, UPS; photo,

GH).

Selaginella poeppigiana var. peruviana A. Braun, Ann.

Sci. Nat. Bot. 5, 3: 295. 1865. TYPE: Peru, Puno,

near Tabina, Lechler 2015 (holotype, B?).

Main stem prostrate, elongate, creeping, only

rarely assurgent at the apex, articulate, pale green

or sometimes light brown, glabrous, the basal part

with spreading leaves, and often smaller, ap-

pressed, ascending leaves. Primary branches 3-4-

pinnate, distant, extending to the base of the main

stem, more or less equally divided, in a flabelliform

arrangement, the terminal branches nearly the same

length, 8-12 mm broad including the leaves. Rhi-

zophores long, coarse, stramineous, throughout the

stems. Lateral leaves above the first branch spread-

ing, distant, imbricate only on the ultimate leafy

branches, 6-8 mm long, oblong-lanceolate, the sides

more or less parallel, symmetrical, acute, the auricles

not prolonged, without whitish borders, the edges

denticulate, sometimes sparingly. Median leaves

elongate-ovate, long-acuminate to subaristate, the

arista less than V* the lamina, the base with a pro-

longed, Ungulate auricle, the borders narrow, whitish

or not, denticulate along the edges.

In moist ravines, in secondary forest, primary

rain forests, in soil among rocks, 730-2700 m,

Amazonas, south to Puno.

Ecuador to Bolivia.

The general aspect of the leaves and system of

branches is similar to those of Selaginella poep-

pigiana and S. articulata, but the lateral leaves in

those species have prolonged auricles that are not

found in 5". trisulcata. The stem also has two vas-

cular bundles.

Amazonas: Prov. Bongara, N of Pedro Ruiz, Smith &

Vasquez 4896 (MO, uc, USM). Huanuco: Carpish Pass,

Allard 20996 (us). Pasco: Prov. Oxapampa, Smith &

Canne 5824 (MO). Prov. Oxapampa, Canyon de Huan-

cabamba, Leon 670 (F, GH). Junin: Carpapata, above

Huacapistana, Killip & Smith 24467 (us). Yaupe, Woyt-

kowski 6331 (MO, us). Ayacucho: Between Huanta and

Rio Apurimac, Killip & Smith 22751 (F, us). Cuzco: Ca.

5 km N of Aguas Calientes, Solomon 3162 (MO). Portero,

8 km W of Quillabamba, Tryon & Tryon 5394 (F, GH,

us).

29. Selaginella poeppigiana (Spring) Splitg.,

Tijdschr. Naturl. Gesch. Physiol. 7: 443. 1840.

Selaginella sulcata subsp. poeppigiana Spring, Flora

(Jena), 21: 185. 18 38. TYPE: Ecuador, Pichincha,

Jameson (holotype, K).

Main stem prostrate, long-creeping, ca. 20-40

cm long, articulate, stramineous, sometimes light

brown at the apex, glabrous, the basal part with

spreading leaves, and often appressed, ascending

leaves. Leafy branches 3-4-pinnate, dichotomous,

in a subflabellate arrangement, extending to the

base of the main stem, the ultimate branches sim-

ilar in length or nearly so, 8-10 mm broad. Rhi-

zophores at the base of the main stem, extending

to the upper branches nearly to the apex. Lateral

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

83

leaves above the first branch distant, spreading to

patent, 4-6 mm, usually 5 mm long, oblong, sym-

metrical or nearly so, acute, the base narrowed

with 1 or 2, prolonged, acute or Ungulate, ciliate

auricles, without conspicuous whitish borders, the

edges finely denticulate. Median leaves elongate-

ovate, long-acuminate or subaristate, the arista less

than 'A the lamina length, the base with a pro-

longed, Ungulate, often ciliate auricle, without con-

spicuous borders, the edges sparsely denticulate.

Creeping, prostrate, forming dense mats on

shaded floor of wet forest, trailing along banks and

roadsides, or in rock crevices, 900-2800 m, Ama-

zonas south to Puno.

Ecuador and Peru.

The long, creeping stems, with abundant rhi-

zophores throughout and in a subflabelliform ar-

rangement, suggest alliances with Selaginella tri-

sulcata and S. articulata. However, the leafy

branches are not as broad, and usually more open

and dichotomous than in those species.

Amazonas: Prov. Bagua, La Peca, Harbour 2493 (F,

MO, uc), 281 9 (F, MO). La Peca, Serrania de Bagua, Gentry

et al. 23065 (F, MO, uc). San Martin: Prov. Moyobamba,

Ferreyra 18518 (USM). Tarapoto, Spruce 4626 (GH, us).

Loreto: Upper Rio Macusari, below Ecuadorian border,

McDaniel 10984 (GH, MO). Huanuco: Pampayacu, Ka-

nehira 182 (GH, us). Pasco: Prov. Oxapampa, Rio San

Alberto, Leon 645 (F, USM). Junin: Above San Ramon,

Killip & Smith 24657 (F, GH); Chanchamayo Valley,

Schunke 192 (F), 194 (F), 231 (F). Cuzco: Prov. Paucar-

tambo, Suecia, Woytkowski 175 (USM). Puno: Churu-

mayo, Soukup 434 (GH).

30. Selaginella articulata (Kunze) Spring, Flora

(Jena), 21: 182. 1838.

LycopodiumarticulatumKunze,Linnaea9: 10, 1834.

TYPE: Peru, Tocache, Poeppig, in 1830 (not lo-

cated).

Main stem prostrate, long-creeping, articulate,

stramineous or light brown near the apex, pubes-

cent especially at the apex and branch axils, or the

main stem glabrescent. Leafy branches usually

3-pinnate, the apex forming a broad, compact, fla-

belliform arrangement, the ultimate branches 10-

20 mm broad. Rhizophores long, coarse, numer-

ous, extending 3/4 the length of the stem, stramin-

eous or golden-colored. Lateral leaves closely

placed to somewhat imbricate at the apex, those

on the leafy branches oblong, spreading, approx-

imate, 8-10 mm long, oblong, somewhat broader

near the base, nearly symmetrical, obtuse to acute,

with a prolonged, round auricle, the edges narrow,

whitish, denticulate to entire. Median leaves short,

ovate, the apex attenuate, the base with 2 pro-

longed, round auricles, the edges narrow, whitish,

finely denticulate.

Plants subrepent, on floor of rain forest, along

streams and roadsides, 350-2000 m, Amazonas

and Loreto south to Pasco and Ucayali.

Panama; Suriname and French Guiana; Peru,

Bolivia, Brazil, Paraguay, and Argentina.

The large, flabellately aligned branches with im-

bricate leaves on the terminal portion of the stems

compose form-flattened sprays that distinguish this

from other Peruvian species. The prolonged basal

auricles on the outer side of the median leaves are

also exceptional.

Amazonas: Prov. Bagua, Montenegro, Hutchison &

Wright 3749 (F, GH, MO, uc, us). Prov. Bagua, near Cam-

pamento, Wurdack 1892 (F, GH, uc, USM). San Martin:

Prov. Mariscal Caceres, Tocache Nuevo, Schunke 6916

(F, MO, uc). Tarapoto, Spruce 4627 (GH, us). Loreto:

Prov. Maynas, Poeppig, in 1831 (MO). Pasco: Prov. Oxa-

pampa, Smith & Salick 8366 (MO). Ucayali: Padre Abad,

Schunke 3074 (F, GH).

3 1 . Selaginella parkeri (Hooker & Grev.) Spring,

Bull. Acad. roy. Sci. Bruxelles 10: 146. 1843.

Lycopodium parkeri Hooker & Grev., Bot. Misc. 2:

388. 1831. TYPE: Guyana, Demerara, Parker

(holotype, K).

Selaginella pedata Klotzsch, Linnaea 18: 521. 1844.

TYPE: Guyana, Schomburgk 118 (holotype, B?;

isotype, BM).

Selaginella fragilis A. Braun, Ann. Sci. Nat. Bot. 5, 3:

305. 1865. TYPE: Brazil, Amazonas, Rio Vaupes,

Spruce 2533 (holotype, B?; isotype, BM, CGE).

Selaginella brachylepis Christ, Bull Herb. Boissier 2,

1: 74. 1901. TYPE: Peru, "entre Ucayali et Hua-

llaga," Huber (holotype, P).

Main stem erect, 20-25 cm long, articulate, stra-

mineous, or rarely red, glabrous, the basal part

with appressed, ascending leaves. Primary branch-

es 3-4-pinnate, sessile, the axis fractiflex, the

branches forming compact, frondlike, flabelliform

sprays, ultimate branches 5-12 mm broad includ-

ing the leaves. Rhizophores at or near the base of

the main stem, well below the first branch. Lateral

leaves above the first branch, coarse, spreading to

somewhat ascending, usually approximate, 4-6

mm long, oblong, broader just above the base,

acute, the base usually with a prolonged, dentic-

ulate auricle, sometimes 2, 1 an acute lobe, with

narrow whitish borders or not, the edges dentic-

84

F1ELDIANA: BOTANY

ulate, the teeth somewhat longer and denser at the

base. Median leaves lanceolate, the apex long-acu-

minate, the base with a prolonged, peltate, Un-

gulate auricle, or sometimes 2, without conspic-

uous borders, the edges finely denticulate.

Primary forests, sometimes in wet or swampy

forests, on sand or clay soils, usually in deep shade,

140-650 m, Amazonas and Loreto, south to Pas-

co.

Colombia and Peru; the Guianas, Amazonian

Venezuela; Brazil.

The plants have stems with one vascular bundle,

and there are short auricles on the axillary leaves

similar to those on the median leaves. The long

main stems bearing flagelliform branches that ap-

pear somewhat frondlike, and the fractiflex axes

of the primary branches clearly suggest alliances

between this and S. geniculata. Some specimens

with unusually attenuated, flagelliform branches

have been recognized as a distinct species, Selag-

inella fragilis.

Amazonas: Prov. Bagua, Montenegro, Hutchison &

Wright 3821 A (uc). San Martin: Alto Rio Huallaga, Juan

Jui, King 3831 (F, GH, both mixed with S. anceps). Lo-

reto: Prov. Requena, van der Werffet al. 9966 (MO, uc),

10108 (MO, uc). H nan urn: Prov. Pachitea, Bosque Na-

tional de Iparia, Schunke 2258 (F, GH, us). Pasco: Prov.

Oxapampa, Quebrada Castilla, Leon & Young 1036 (F,

GH). Prov. Oxapampa, Iscozacin, D. Smith 3735 (MO,

uc).

32. Selaginella geniculata (Presl) Spring, Bull.

Acad. roy. Sci. Bruxelles 10: 230. 1843.

Lycopodium geniculatum Presl, Reliq. haenk. 1 : 80.

1 825. TYPE: "Luzon," but doubtless Peru or Ec-

uador, Haenke (holotype, PR; photo, BM, GH).

Selaginella ferruminata Spring, Bull. Acad. roy. Sci.

Bruxelles 10: 231. 1843, TYPE: Peru, Pangoa,

Matthews (Mathews) 1083 (holotype, K.; isotypes,

GH, us).

Selaginella elongata Klotzsch, Linnaea 18: 522. 1844.

TYPE: Peru, Cuchero, Ruiz 94 (holotype, B).

Selaginella nodosa Presl, Abh. Bohm Ges. Wiss. 5 (2):

580 (Bot. Bermerk. 50). 1844. TYPE: Peru, Cas-

sapi, Poeppig (not located).

Selaginella tomentosa Spring, Nouv. Mem. Acad. roy.

Sci. Belg. 24: 231. 1849. TYPE: Colombia, Cau-

ca, Gorgona Isl., Hinds (holotype, K.).

Main stem erect, 10-50 cm or longer, articulate,

stramineous, glabrous, or sometimes pubescent,

the basal part with appressed, ascending leaves.

Primary branches 3-4-pinnate, sessile above the

lowest branches, forming flat, imbricate sprays in

an extended flabelliform, frondlike arrangement,

axis of the primary branches fractiflex, the ulti-

mate leafy branches 2-5 mm broad. Rhizophores

borne at the base of the main stem, or below the

first branch. Lateral leaves of the main stem, above

the first branches, coarse, ascending or spreading,

usually approximate, those below distinct, peltate,

2-5 mm long, elongate-ovate, broadest above the

base, acute, the base with a short, acute lobe, with-

out conspicuous whitish borders, the edges den-

ticulate, especially at the base. Median leaves im-

bricate, oblong, acute to acuminate with a long,

Ungulate auricle, without conspicuous whitish bor-

ders, the edges entire or nearly so.

On white sand or clay soil in high forest or rain

forest, usually terrestrial but sometimes on tree

trunks, or on rocks, 120-1500 m, Amazonas and

Loreto south to Cuzco and Madre de Dios.

Colombia, Ecuador and Peru.

The axillary leaves are truncate; there are two

vascular bundles at the base of the main stem, and

there are peltate leaves on the main stem, below

the first branch. These characters distinguish the

species from other erect, articulate species of Peru.

An alliance with Selaginella parkeri is indicated

by the long main stem, the alignment of branches,

and the fractiflex axes of the primary branches;

however, the leafy branches are not as broad as in

S. parkeri.

Amazonas: Rio Cenepa, vicinity of Huampami, Berlin

633 (F, MO, uc); Ancuash 1042 (MO, uc). Prov. Bagua,

above Cascadas de Mayasi, Wurdack 1894 (F, GH, uc,

us, USM). San Martin: Prov. Mariscal Caceres, Cam-

panilla, Schunke 4118 (F, us). Loreto: Between Yuri-

maguas and Balsapuerto, Killip & Smith 28250 (F, GH,

us). Prov. Maynas, between Rio Momon and Rio Mo-

monicillo, McDaniel 16121 (F, GH, MO). Huanuco: Prov.

Huanuco, Vargas 5288 (MO). Pasco: Prov. Oxapampa,

between Puerto Bermudez and Hito San Matias, Leon

et al. 327 (F, USF, USM). Junin: Satipo, Ridoutt 11397

(USM). Ucayali: Boqueron del Padre Abad, Skog et al.

5128 (us). Cuzco: Marcapata, Quispicanchis, Vargas 3781

(MO, us). Madre de Dios: Prov. Manu, Vargas 16643

(GH).

33. Selaginella asperula Spring, in Martius, Fl.

bras. 1 (2): 127. 1840. SYNTYPES: "ad flu-

vium Amazonum," Martius (M), "et ad flu-

men S. Francisco," Martius (M).

Main stem slender, erect, sometimes arching,

and rooting at the apex, usually 1 -pinnate, 1 5-30

cm long, articulate, stramineous, glabrous, the basal

part with appressed, ascending leaves. Primary

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

85

branches distant, often forming tufts of slender,

dichotomous branches along the main stem,

sometimes rebranching into uniformly slender ul-

timate leafy branches 1-3 mm broad, the axes

straight or nearly so. Rhizophores mostly at or near

the base of the main stem. Lateral leaves above

the first branch ascending or wide-spreading, 0.5-

3 mm long, elongate-ovate, somewhat broader just

above the base, acute, with an auricle sometimes

forming a small, acute tooth, the edges narrow,

whitish, ciliolate, these longer at the base. Median

leaves ovate, asymmetrical, acute, with a pro-

longed, ciliate auricle, the edges narrow, whitish,

denticulate, the teeth longer and denser at the base.

Steep slopes and road banks, open areas, thick-

ets and moist areas in dense forests, 680-1 300 m,

Amazonas and Loreto to Junin.

Amazonian Venezuela and Brazil, south to Bo-

livia.

The diffuse branches that form a uniformly slen-

der, dichotomous, leafy branch system, with small,

strongly imbricate leaves, are distinctive among

the Peruvian species. The species has a single stele

at the base of the erect stem, a character that sep-

arates it from Selaginella geniculata.

Amazonas: Prov. Bagua, 43 km NE of Chiriaco, Bar-

hour 4458 (MO, USM). San Martin: San Rogue, LI. Wil-

liams 7415 (F, us). Prov. Rioja, Woytkowski 6036 (GH,

MO, us). Loreto: Rio Huallaga. above Lagunas, Croat

18097 (F, GH, MO). Huanuco: Tingo Maria (as San Mar-

tin) Allard 20823 (MO). Pasco: Prov. Oxapampa, Parque

Nacional Yanachaga-Chemillen, Leon et al. 989 (F). Ju-

nin: San Ramon, Killip & Smith 24785 (F, GH, us). Prov.

Chanchamayo, La Merced, Macbride 5504 (F, us); La

Merced, D. Smith 4063 (MO, uc).

34. Selaginella stellata Spring, in Martius, Fl. bras.

1 (2): 129. 1840. LECTOTYPE (designated

here): Brazil, Prov. Para. Obidos, Martius

(holotype, M).

Selaginella conduplicata Spring, in Martius, Fl. bras.

1 (2): 129. 1840. TYPE: Brasil, Prov. Para, Mar-

tius (holotype, M).

Selaginella calcarata A. Braun, Ann. Sci. Nat. 5, 3:

305. 1865. TYPE: ?, several syntypes are cited.

Main stem erect to sometimes creeping, 10-30

cm long, articulate, stramineous, glabrous, the basal

part with appressed, ascending leaves. Primary

branches 3-4-pinnate, usually with an extended,

flabelliform, frondlike arrangement, branches dis-

tant, the lowest often longest, or in creeping stems

the branches of similar length, axes of the branches

fractiflex, ultimate leafy branches slender, 2-5 mm

broad. Rhizophores abundant on the lower part of

the stem nearly to the first branch, or in creeping

stems among the upper branches. Lateral leaves

above the first branches distant, ascending, those

below distant, peltate, 1—4 mm long, lanceolate,

slightly broader above the base, acute, the base

with a prolonged, membranaceous auricle on the

acroscopic side and usually a sharp tooth on the

basiscopic side, with narrow, whitish, denticulate

edges. Median leaves very small, usually '/2 the

length of the lateral leaves, ovate-lanceolate, acu-

minate or subaristate, with a prolonged, Ungulate,

ciliate basal auricle, with narrow, whitish, dentic-

ulate edges.

In dense stands on clay or often sandy soil in

lowland woods or rain forests and secondary for-

ests. Common in humid, disturbed areas, in chac-

ras, especially old ones, ca. 100-250 m, Loreto

and Junin.

Guianas, Amazonian Brazil west to Peru.

We follow Alston (Repert. Spec. Nov. Regni

Veg. 40: 309. 1936) in accepting the name Selag-

inella stellata over S. conduplicata, both published

by Spring in the same work.

The species is readily distinguished by the un-

usually prolonged, ciliate auricles on axillary, me-

dian, and lateral leaves. The bright stramineous

color of the fractiflex axes contrasting with the

deep green leaves resembles what is seen in Se-

laginella geniculata, and there are peltate leaves

on the lower part of the main stem and branches

similar to that species.

Loreto: Iquitos, Croat 18306 (MO, us), 20057 (GH, MO,

uc). Iquitos, road to San Juan, Mexia 6496 (F, GH, MO,UC).

Prov. Maynas, Picuruyacu, Revilla 123 (F, MO, uc). Ga-

mitanacocha, Schunke 144 (F, GH, uc, us). Iquitos, Tryon

& Tryon 5166 (F, GH, us). Junin: Chanchamayo Valley,

La Merced, Soukup 1117 (F), 7775 (F). La Merced, Mac-

bride 5503 (F).

35. Selaginella exaltata (Kunze) Spring, Bull.

Acad. roy. Sci. Bruxelles 10: 234. 1843. Fig-

ure 9e-f.

Lycopodium exaltatum Kunze, Linnaea 9: 8. 1834.

TYPE: Peru, San Martin: "ab Uchiza ad Toache,"

Poeppig, July 1830, Diar. 1953 (not located).

Selaginella strobilifera Christ, Bull. Herb. Bossier 2,

1: 72. 1901. TYPE: Peru, "entre Rios Huallaga

et Ucayali," Huber 1515 (holotype, P; isotype,

MG).

Main stem erect, scandent up to 8 m long, ar-

ticulate, 2-6 mm broad including the leaves, stra-

86

FIELDIANA: BOTANY

mineous, often pubescent. Primary branches widely

spreading, 2-3-pinnate, often pubescent. Rhizo-

phores few, borne in axils of branches. Lateral

leaves distant, clasping on the main stem when

dry, approximate, or imbricate on the ultimate

branches, obtuse, elongate, 1-3 mm long, 3 or 4 x

longer than broad, acute, or sometimes entire,

somewhat acuminate, the base truncate, some-

times with conspicuous whitish borders. Median

leaves acentric, the outer side larger, ovate, acu-

minate to aristate, the arista '/4 the lamina length,

glabrous, the base entire.

In lowland forests, rain forests, secondary for-

ests, and disturbed areas, scandent or scrambling

over shrubs, often forming dense thickets, 30-760

m, Amazonas south to Madre de Dios.

Costa Rica south to Bolivia and western Brazil.

The scandent habit of these plants, often form-

ing dense thickets, is exceptional among species

of Selaginella. The stems are noted to be up to 8

m long, but in most collections they are indicated

as 3 m. The habit of these plants, with dense

branches bearing closely placed leaves, forms a

compact cover over other vegetation.

Amazonas: Prov. Bagua, Montenegro, Sagdstegui et

al. 7158 (F, GH, HUT, MO, uc). Prov. Bagua, above Cas-

cadas de Mayasi, Wurdack 1800 (F, GH, uc, us). San

Martin: between Tocache Nuevo and Juanjui, Croat

58035 (F, MO). Lamas, Naranjal, Knapp & Mallet 6958

(F, MO, uc). Loreto: Bosque Nacional von Humboldt,

Gentry & Revilla 20418 (F, MO). Prov. Maynas, Plowman

et al. 6502 (F, GH, us). Nanay, Woytkowski 5126 (GH,

MO, us). Huanuco: Casa de Koepke, Rio Llullapichis,

Dudley 12484 (GH). Bosque Nacional de Iparia, Schunke

1299 (F, GH, MO, us). Pasco: Prov. Oxapampa, Cabeza

de Mono, D. Smith 3768 (MO, uc). Puerto Bermudez (as

Junin), Killip & Smith 26498 (us). Madre de Dios: Prov.

Tambopata, Tambopata Nature Reserve, Barbour 4881

(F, MO). Manu National Park, Gentry et al. 27209 (F, MO).

36. Selaginella convoluta (Arnott) Spring, in Mar-

tius, Fl. bras. 1 (2): 131. 1840.

Lycopodium convolutum Arnott, Mem. Wernerian Nat.

Hist. Soc. 5: 199. 1824 TYPE: Brasil, Rio de

Janeiro, Jameson (holotype, E).

Main stem erect, very short, not articulate, green

to brownish, glabrous. Primary branches forming

an open rosette curling inward when dry, 1-2-

pinnate, with the ultimate branches, including the

leaves, 3-5 mm broad. Rhizophores few at the base

of the main stem. Lateral leaves rigid, strongly

imbricate, 1-2 mm long, ovate-lanceolate or

somewhat elliptic, acuminate to acute, strongly

imbricate, the base with a prolonged, densely cil-

iate auricle, the acroscopic side with a broad, whit-

ish border, the edges irregularly denticulate. Me-

dian leaves broadly ovate to somewhat elliptical,

obliquely oriented, acute or short aristate, the base

not prolonged, the borders not conspicuously

whitish, the edges regularly dentate.

On dry desert slopes, 1250 m, Cajamarca.

Guatemala; Greater Antilles; Guyana; Peru;

Brazil and Paraguay.

The habit of these plants forming a rosette of

stems, with lateral branches curling inward, readi-

ly characterizes the species. The leaves are di-

morphic; both lateral and median leaves are

strongly imbricate and have whitish margins and

dentate edges.

Selaginella convoluta is widely distributed in

Central and South America, but only a single col-

lection has been seen from Peru. The inrolled,

brownish, leafy stems may be easily overlooked

on the dry, desert slopes where the plants occur.

Cajamarca: Prov. Celendin, Balsas-Celendin road, Rio

Maranon valley, D. Smith 6179 (MO).

Comments

The following species are accepted by Alston et

al. (1981) as growing in Peru. Adequate material

of these has not been seen and a better evaluation

of them is desired than is now possible.

a. Selaginella acanthostachys Baker, J. Bot. 21:

99. 1 883. TYPE: Peru, San Martin, Spruce 4328

(holotype, K).

b. Selaginella- asplundii Crabbe & Jermy, Fern

Gaz. 11: 257. 1976. TYPE: Peru, Huanuco,

Asplund 12822 (holotype, s; isotype, BM).

c. Selaginella muscosa Spring, in Martius, Fl. bras.

1 (2): 120. 1840. Type: Brazil, Luschnatt (ho-

lotype, M).

d. Selaginella suavis (Spring) Spring, Bull. Acad.

roy. Sci. Bruxelles 10: 229. 1843. Selaginella

sulcata ssp. suavis Spring. Flora (Jena) 21: 185.

1838. TYPE: Brazil, Sellow (holotype, K; iso-

type, B?).

e. Selaginella wolfii Sodiro, Crypt, vase. Quit. 620.

1893. TYPE: Ecuador, "woods of the western

region, 1800 m." Sodiro (holotype?, p).

The following species was credited to Peru on

the basis of the type, which is now considered to

have been collected in Panama.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. VI.

87

f. Selaginella horizontalis (Presl) Spring, Bull.

Acad. roy. Sci. Bruxelles 10: 226. 1843. Lyco-

podium horizontale Presl, Rel. haenk. 1: 78.

1825. TYPE: Panama, Haenke (holotype, PR),

not from Peru as originally thought.

Family 28. ISOETACEAE

Isoetaceae Reichenb., Hot. Damen Kunst. Freunde

Pflanzenw. 309. 1828. TYPE: Isoetes L.

Stem erect, subterranean, not indurated, bearing

numerous, fleshy to wiry, dichotomously branched

roots. Leaves numerous, spirally arranged, ligu-

late, acicular with a single vein and 4 rows of air

chambers, each transversely partitioned by mul-

ticellular septa. Sporangia foliar, large, single, ses-

sile at or near the adaxial leaf base. Megaspores

tetrahedral-globose and trilete, without chloro-

phyll, germinating endosporically to form a small,

short-lived megagametophyte. Microspores ellip-

soidal and monolete, germinating endosporically

to form a small megagametophyte producing 4

multiflagellate spermatozoa.

An ancient genus with representatives known

from the lower Cretaceous; represented today by

a single, distinctive, and cosmopolitan genus ex-

hibiting distant affinities with Selaginella and Ly-

copodium.

Species recognition is greatly hindered by tre-

mendous phenotypic plasticity and active speci-

ation, both through hybridization and polyploidy

as well as through diploid divergence (Taylor &

Hickey, 1992). Species identificatiop is also ham-

pered by these phenomena but is made even more

difficult by a suite of specialized characters not

found elsewhere in the plant kingdom. Unique

terminology, therefore, is an historic addendum

to Isoetes systematics. Most of these novel terms

were first introduced by Alexander Braun in 1 864.

The use of them, although inconvenient at first,

adds clarity and uniformity of understanding.

Within the body of this treatment I employ them

in the following contexts: corm— the "stem" of the

plant, that part to which the leaves and roots are

attached; fossa— the groove (or one of several) on

the side of the corm (lateral fossa), or extending

all the way around the corm (circumbasal fossa)

and from which roots are derived; synchronous

roots acropetal roots— in Isoetes, roots are initi-

ated with a distinctive rhizotaxy (Paolillo, 1963)

with root initiation simultaneous (synchronous) or

not (acropetal); subula— the distal, non-laminate

portion of the leaf; ala— the lateral, proximal lam-

inate portion of the leaf, not including the ex-

panded mid vein or sporangial regions; scales —

leaf primordia that have been arrested early in

ontogeny and have become sclerified and black-

ened; these initially surround the corm apex during

dormancy and, as new leaves are initiated, are

displaced outward, and hence may be found sur-

rounding the fully developed leaves on a corm;

ligule— a multicellular, planar, and glandular tis-

sue growing out of a small pit (foveola) in the

proximal portions of the adaxial leaf surface, just

distal to the sporangium; the ligule is composed

of a thickened central region (the cushion) and

peripheral, delicate margins; labium— a nonglan-

dular, planar tissue that is apparently a modifi-

cation of the proximal rim of the foveola; this

tissue can be quite extensive and may be errone-

ously identified as the ligule (it has also been called

"pseudoligule"); velum— a tissue of leaf derivation

that completely or partially covers the sporangi-

um, the latter in most species somewhat impressed

within the leaf tissue.

References

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HICKEY, R. J. 1984. Chromosome numbers of

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HICKEY, R. J. 1986. Isoetes megaspore surface

morphology: Nomenclature, variation, and sys-

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MARSDEN, C. R. 1976. Morphological variation

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PAOLILLO, D. J. 1963. The developmental anat-

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PFEIFFER, N. 1922. A monograph of the Isoe-

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TAYLOR, W. C., AND R. J. HICKEY. 1992. Hab-

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262.

88

FIELDIANA: BOTANY

I. Isoetes

Isoetes L., Sp. pi. 2: 1 100. 1753. TYPE: /. lacus-

tris L.

Stylites Amstutz, Ann. Missouri Hot. Gard. 44: 121.

1957. TYPE: S. andicola Amstutz = /. andicola

(Amstutz) Gomez. Figure 10.

Aquatic, amphibious or terrestrial perennials.