A REVISION OF THE GENUS CYATHEA

A REVISION OF THE GENUS CYATHEAAuthor(s): Rolla TryonSource: Contributions from the Gray Herbarium of Harvard University, No. 206 (1976), pp.19-98Published by: Harvard University HerbariaStable URL: http://www.jstor.org/stable/41764719 .

Accessed: 11/09/2014 20:44

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .http://www.jstor.org/page/info/about/policies/terms.jsp

.JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range ofcontent in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new formsof scholarship. For more information about JSTOR, please contact [email protected].

.

Harvard University Herbaria is collaborating with JSTOR to digitize, preserve and extend access toContributions from the Gray Herbarium of Harvard University.

http://www.jstor.org

This content downloaded from 129.115.103.99 on Thu, 11 Sep 2014 20:44:57 PMAll use subject to JSTOR Terms and Conditions

http://www.jstor.org/action/showPublisher?publisherCode=huh

http://www.jstor.org/stable/41764719?origin=JSTOR-pdf

http://www.jstor.org/page/info/about/policies/terms.jsp


A REVISION OF THE GENUS CYATHEA

Rolla Tryon

Cyathea is a genus of forty neotropical species primarily found in the mountains of the Greater Antilles, Central America, and tropical South America. It is the dominant element in the tree fern flora of cloud forests and is the most frequent genus of Cyatheaceae occurring at higher altitudes. Two species grow at 4200 m in the Andes, the highest altitude re- corded for the family.

The genus is here defined in a restricted sense, following my earlier study of the generic classification of the Cyatheaceae (Tryon, 1970). The primitive species of Cyathea , with a complex leaf architecture and hemitelioid indusium, are related to Trichipteris and to Cnemidaria. Recent intensive studies, including species revisions of all three genera, have served to strengthen the earlier generic proposals. Trichipteris (Barring- ton, 1974), Cyathea , and Cnemidaria (Stolze, 1974) have emerged more clearly as separate evolutionary lines with truly intermediate species few or lacking.

Trichipteris is technically separated from Cyathea by the absence of an indusium, while Cyathea has a well-developed or at least small (hemitelioid) indusium. Some species of Trichipteris have lamina scales regularly associated with the sorus and these scales may be mistaken for indusia. They are, however, thin and clearly cellular, narrowed basally and attached at one point, attenuate at the apex, and often erect. The smallest indusia in Cyathea are somewhat thickened, with the cells ob- scured, broadly attached at the base, rounded at the apex, and appressed to the leaf tissue. The closest relation between Cyathea and Trichipteris is seen in Cyathea multiflora and Trichipteris nigra and (or) T. Schiedeana.

Stolze ( 1974) has clarified the definition of Cnemidaria and Cyathea by removing from Cnemidaria a few species that I had earlier placed there which lack large-porate spores. These and some other related species are now seen to represent an independent line of lamina reduction in Cyathea. The closest relationship with Cnemidaria is evidently represented by Cyathea species with a more complex lamina.

KEY TO SCALY GENERA OF CYATHEACEAE a. Petiole scales structurally conform, the cells of the body similar in orientation, shape, and (usually) in size and color, or a single row of cells at the edge may be differentiated as cilia or denticulations Sphaeropteris.

a. Petiole scales structurally marginate, with a narrow multicellular marginal band to a broad margin of cells different in orientation, size, and (usually) in shape and color from those of the central portion b. b. Petiole scales with a dark, opaque apical seta; spores 16 in a sporangium, rarely 64 c. c. Petiole lacking spines, or with corticinate spines, each bearing (when young) a scale at the apex, croziers lacking spines Alsophila.

c. Petiole with squaminate spines, many large and black, with a slender apex, these

19



20 ROLLA TRYON

also present on the croziers, petiole scales borne on the petiole surface. . Nephelea. b. Petiole scales lacking an apical seta, the apex rounded to filamentous; spores 64 in a sporangium, rarely 16 or 32 d. d. Indusium absent Trichipteris. d. Indusium present e. e. Spore exine lacking apertures or with variously distributed small pits or pores; costae and costules with adaxial trichomes; veins free, or in two species sometimes areolate, in two species the lamina l-pinnate-pinnatifid and the petiole either alate or densely long-pubescent Cyathea.

e. Spore exine with three large equatorial pores, often also with smaller pits or pores; costae and costules lacking trichomes on the adaxial side, or rarely present; basal veins usually forming areolate, or if free then connivent to the sinus, or in one species running to the margin above the sinus and the lamina l-pinnate- pinnatifid, the petiole nearly glabrous and non-alate Cnemidaria.

Ecology and Geography

Cyathea is predominently a genus of the cloud forest, although it is also

well-represented in wet montane forests. The cloud forest environment, usually developed on steep mountain slopes, provides nearly constant high humidity and considerable light since the tree fern crowns occupy part of the relatively open canopy. Several species, such as C. pallescens , C.

suprastrigosa, and C. caracasana also grow at higher altitudes in Chusquea thickets and in subparamo scrub. Other species grow in lowland rain forest below 500 m, but only five of these, including C. conformis, C. decorata and C. parva are wholly lowland species. The lowland forest habitat is different from the cloud forest in providing a significant dry season or dense shade, as well as higher temperatures. Some species occasionally grow in other, specialized habitats, for example, C. parvula in wet savannahs, C. Harrissii in wet pinelands, C. platylepis among sandstone rocks, and C. Delgadii in gallery forest. Among the 15 species that grow at altitudes of 500 m or less, 11 of them have hemitelioid indusia. The significance of this correlation is not clear since these 11 species are morphologically quite diverse and might be expected to have considerable ecological differences.

Cyathea arborea is a pioneer species, invading landslides where it

frequently establishes handsome colonies. It also invades artificial barren sites, such as road cuts and road fills. Since the many cloud forest species usually grow on steep mountain slopes, they probably also grow in land- slide areas. However, little is known about their successional position. Some may be pioneers, as C. arborea, while others may enter the new community later.

The distribution of the genus ( Map 1) correlates closely with the mountains of tropical America. Cyathea is notably absent from the Yucatan Peninsula, from the Llanos region of Colombia and Venezuela, the Orinoco Basin, most of the Amazon Basin, and from the Bolivian-Paraguayan lowlands. Table 1 and Map 2 provide information on the occurrence of species, varieties and endemics in countries and regions. The center of di- versity is in Venezuela and Colombia, south to Peru. In this region there



REVISION OF CYATHEA 21

are 31 taxa and 19 endemics. Colombia is the country with the most diverse Cyathea flora, with 22 taxa and 6 endemics. Ecuador seems to be under- represented. It is usually rich in species of cloud forest groups; for

example, in Hymenophijllum section Sphaerocionium (Morton, 1947), there are 19 species in Colombia, 16 in Ecuador and 22 in Peru. It is expected that additional collecting of tree ferns in Eucador will increase the number to 15 or more.

In southeastern Brazil there is a single species, Cyathea Delgadii, and only three species occur in Mexico and Guatemala. The paucity of species in these regions is peculiar for there are considerable areas of wet montane

Maps 1-2. 1, Distribution of Cyathea (occurrence on isolated islands on Map 2); 2. Numbers of species and varieties, and number of endemics in countries and regions (see Table 1), the left or single bar is the number of taxa ( 1-22 ) , the right bar is the number of endemics ( 1-6 ) .



22 ROLLA TRYON

forest and also smaller ones of cloud forest; previous adverse climates may account for the limited number. The situation is probably similar to that in west tropical Africa, where a history of vegetational and climatic change evidently occurred and where there are only three species of Cyatheaceae (Alston, 1959).

There are seven species having wide ranges of 3000 miles (4800 km) or more in extent. They are frequent species with rather broad altitudinal ranges. Successful migration of species is perhaps most dependent on their broad ecological tolerance, as may be seen in those which are the source for populations occurring on isolated islands. These insular species are related to the three most widely distributed continental species: the Cocos Island endemics, Cyathea Alphonsiana and C. notabilis to C. multiflora ; the Galapagos endemic, C. Weatherbyana to C. andina; and C. Delgadii of Ilha Trinidade is widespread in South America.

Table 1. geographic occurrence of taxa and endemics of cyathea Mexico to Nicaragua: 3 taxa, 1 endemic. 1. multiflora, 20b. divergens var. Tuerck-

heimii, 29. fulva. Costa Rica and Panama: 10 taxa, 3 endemics. 1. multiflora, 9. petiolata, 10. impar,

12. conformis, 20a. divergens var. divergens, 29. fulva, 32. suprastrigosa, 33. Delgadii, 36e. caracasana var. Maxonii, 38. gracilis. Cocos Island: 2 endemic species. 4. Alphonsiana, 5. notabilis.

Greater Antilles: 8 taxa, 4 endemic. 2. andina, 6. parvula, 17. arborea, 30. Harrisii, 31. furfuracea, 35. dissoluta, 36c. caracasana var. caracasana, 38. gracilis. Lesser Antilles , Trinidad and Margarita: 2 species, 1 endemic. 17. arborea, 34. tenera.

French Guiana , Surinam and British Guiana: 3 taxa, no endemics. 2. andina, 20a. divergens var. divergens, 33. Delgadii.

Venezuela: 14 taxa, 2 endemic. 2. andina, 7. platylepis, 11. Steyermarkii, 15. speciosa, 20a. divergens var. divergens, 22. simplex, 29. fulva, 33. Delgadii, 36. caracasana vars. a, b, c, d. 37. Lechleri, 39. ebenina.

Colombia: 22 taxa, 6 endemic. 1. multiflora, 2. andina, 7. platylepis, 9. petiolata, 12. conformis, 13. decorata, 14. parva, 15. speciosa, 16. Haughtii, 18. Alstonii, 19. peladensis, 20a. divergens var. divergens, 21. pallescens, 25. straminea, 29. fulva, 33. Delgadii, 36. caracasana vars. a, b, c, d, 38. gracilis, 39. ebenina.

Ecuador: 9 taxa, 1 endemic. 1. multiflora, 2. andina, 20a. divergens var. divergens, 21. pallescens, 23. corallifera, 36. caracasana vars. a, b, c, d.

Galapagos Islands: 1 endemic species. 3. Weatherbyana. Peru: 13 taxa, 5 endemic. 1. multiflora, 2. andina, 8. Vilhelmii, 20a. divergens var.

divergens, 21. pallescens, 26. Ruiziana, 27. microphylla, 28. multisegmenta, 33. Del- gadii, 36a. caracasana var. boliviensis, 37. Lechleri, 39. ebenina, 40. Dudleyi.

Bolivia: 7 taxa, 1 endemic. 1. multiflora, 2. andina, 21. pallescens, 24. boliviana, 33. Delgadii, 36a. caracasana var. boliviensis, 37. Lechleri.

Argentina , Paraguay and Brazil: 3 species, no endemics. 1. multiflora, 2. andina, 33. Delgadii.

Ilha Trinidade: 1 species, not endemic. 33. Delgadii.

Speciation

In the scaly Cyatheaceae, speciation is based mainly on eco- geographic isolation and on evolutionary migration (Tryon, 1971, 1972). There are




examples of speciation by geographic isolation in the island endemics: Cyathea Alphonsiana , C. notahilis and C. Weatherbyana. Others involve geographic isolation, possibly preceded by evolution during migration: C. diver gens var. Tuerckheimii in Guatemala and Mexico, from var. diver gens in Costa Rica; C. caracasana var. Maxonii of Costa Rica from the C. caracasana complex in Colombia; and probably C. tenera of the Lesser Antilles, and C. furfuracea of the Greater Antilles, from C. Delgadii of South America. The relatively uniform collections of C. multiflora from

Nicaragua to British Honduras possibly indicate a migration from the more diverse populations of the species in Costa Rica.

There are 13 local endemics in Colombia south to Bolivia where the

continuity of habitats suggests that geographic isolation is not a factor in their endemism. Try on and Gastony (1975) have proposed the hypothesis that such local endemics have become ecologically specialized to the extent that other suitable environments are either absent or exist only as small areas distant to the present range.

Most species of Cyathea are morphologically and eco-geographically distinct, thus their classification is relatively clear. This is especially apparent in the species having hemitelioid and cyatheoid indusia. These distinctive species probably represent relatively early speciation, which would be consistent with their primitive position in the genus. Among the more advanced species, with sphaeropteroid indusia, there are two groups in which speciation appears to be more recent. There are 12 taxa within these groups (species 29-34, 35-36) that are not clearly differentiated due to intergrading variation. Pleistocene climatic changes may have been instrumental in generating migrations and disjunctions leading to the evolution of these taxa.

Amphiploid speciation does not seem to be a recent phenomenon in

Cyathea , although several putative hybrids (see below) suggest that it

might be present. The four species that have been determined cytologi- cally all have a chromosome number of n-6 9: Cyathea arborea , C. furfuracea , C. Harrisii , and C. parvula (Walker, 1966). Considerably more

cytological sampling is needed before the status of polyploidy in Cyathea can be known. The relatively high number of n=6 9 also suggests polyploidy, but this probably occurred very early in the evolution of the family since all of the scaly genera of Cyatheaceae share this number.

Hybrids

Two hybrids have been recently recognized by Conant (1975) in the

Cyatheaceae on the basis of an analysis of several characters and field observations on the hybrids and parental species in the same site. He also discusses five other putative hybrids in Alsophila and Nephelea. These

proposals provide a new perspective on some of the described species in the family. In addition to the hybrid, proposed by Conant, involving



24 ROLLA TRYON

Cyathea arborea and Cnemidaria hórrida (Hemitelia Wilsonii), I am recognizing nine others, with varying degrees of confidence. On the basis of the evidence, an interpretation of them as hybrids seems more realistic than treating them as variants of a species or recognizing them as endemics. Putative hybrids present special problems that require careful field study. Hybrids have not been suggested involving parental species that are similar in their indusium and laminar structure. However, such hybrids may also exist, perhaps with some frequency, but they cannot be identified from herbarium materials at hand. Discriminating field study and collecting will be required before the frequency of hybridization in the genus is known.

Morphology

The petiole always bears large scales at the apex of spines, where they are usually caducous, or they may be on smaller emergences. In addition to these large petiole scales , there is usually also a covering of small to minute scales on the petiole surface, especially on the abaxial side. The

large scales, which have characters of systematic importance, are on the abaxial side of the petiole. Other large scales are borne on the adaxial side. The basal scales, especially, are longer and narrower than those on the abaxial side. They tend to be similar in many species, and often lack the differentiated margin characteristic of the genus.

The body of the petiole scale includes the whole structure except for

peripheral processes such as cilia, teeth or setae. The edge of the scale is the outer row of cells which bears the peripheral processes. The margin of the scale is the cellularly differentiated part of the body on each side of the rather uniformly elongate-celled central portion . Scales that are uni-

formly colored are concolorous (Fig. 1). They are usually whitish. Light brown scales may be nearly concolorous (Fig. 2), the margins being only slightly lighter than the central portion. Scales that have the margins definitely lighter in color than all or part of the central portion are bicolorous. If the central portion is dark and the margins are light, the scale is concordantly bicolorous (Fig. 3), while if the light color of the margin ex- tends into the central portion, the scale is discordantly bicolorous (Fig. 4).

The large petiole scales are best examined on a crozier because all details of the cellular construction are present at this young state. The base of a recently expanded leaf will also usually have scales in good condition. However, the scales on older leaves are more or less eroded, especially the more delicate margins, or they may have been lost.

The small scaly indûment on the petiole surface is called scurf , and when it is composed of definite, although small, scales these are referred to as

large scales of the scurf (Figs. 5, 6). The minute scales of the scurf are referred to as squamellae (Figs. 7, 8). Either kind may be crested (Fig. 6) and the squamellae may be strongly dissected and trichomoid (Fig. 8).




The petiole scurf is best examined in recently expanded leaves. It is usually not fully developed in croziers, and it often becomes abraded in older leaves.

The pinnae and pinnules vary from sessile to very long-stalked. In some species the length of the stalk relative to the width of the basal segment is a useful character. If the stalk is shorter than the width of the basal segment of the pinnule, the pinnule is referred to as short-stalked. It is long- stalked when the stalk is definitely longer than the width of the basal segment. Short- and long-stalked pinnae are defined in a similar manner.

The term pubescence or pubescent is applied in reference to the under surface (abaxial side) of segments or axes, such as the costa and pinna- rachis, when there is a sufficient number of trichomes to form a uniformly distributed indûment. This may be dense to somewhat sparse. If the trichomes are scattered, reference is made to their individual characteristics.

The terminology applied to the scales on the under surface of the pinnae is used in a manner similar to that for the petiole scales.

The lower epidermis in Cyathea is basically of a uniform type (Figs. 10, 12, 14, 16). The stomates are nearly surrounded by a subsidiary cell with

strongly undulating walls. They conform to the polocytic pattern of Van Cotthem (1970). The upper epidermis (Figs. 9, 11, 13, 15) usually shows greater differences in the size of cells and in the undulations of the walls.

The position of the sorus is sometimes an important character, especially in species with hemitelioid indusia. The following terms are applied: the costal position is very near the costa; subcostal is slightly removed; medial is about half the distance between the costa and the segment margin; supramedial is somewhat beyond this; and submarginal sori are close to the margin.

The indusium is considered to be hemitelioid (Fig. 17) if it partially surrounds the base of the receptacle. Small hemitelioid indusia are referred to as minute. This term seems preferable to the more descriptive one, "scale-like," previously proposed (Try on, 1970). That term may cause some confusion with the true laminar scales in Trichipteris which may be mistaken for indusia. Cyatheoid indusia completely surround the base of the

receptacle, but do not completely envelop the sorus. Four types of cyatheoid indusia are recognized: discoid, if nearly flat; meniscoid, if lens-

shaped with somewhat raised edges (a shallow saucer); cyathiform, if a cup (Fig. 18), usually enveloping about half of the sorus; and urceolate, if taller, open, and somewhat contracted at the apex. Sphaeropteroid indusia are globose, completely envelop the sorus (Fig. 19) and often have an apical umbo. In some species of Cyathea the sphaeropteroid indusium is very fragile and ephemeral, for example, in C. straminea , C. pallescens, and C. caracasana var. caracasana. In them the mature indusium breaks into fragments that fall away, leaving only an irregular basal portion of the indusium, if any, attached to the receptacle. Relatively young sori are re-

quired to observe the presence of the indusium in these species.



26 ROLLA TRYON

Figs. 1-16. 1-8. Petiole scales and scurf of Cyathea. 1-4, Petiole scale types, enlarged and diagrammatic (see text): 1, Concolorous (whitish); 2, Nearly concolorous (light brown); 3, Con- cordantly bicolorous; 4, Discordantly bicolorous. 5-8, Petiole scurf types, enlarged; 5, Large scales of scurf, C. divergens var. divergem, Venezuela, Steyermark et al. 103580 (gh); 6, Crested large scales of scurf, C. pallescens, Ecuador, Sparre 16946 (gh); 7, Squamellae, source as in 5; 8, Trichomoid squamellae, C. boliviana, Bolivia, Steinbach 9512 (gh). 9-16. Upper and lower lamina epidermis of Cyathea, X 90. 9-10, Cyathea andina, Peru, Killip à- Smith 23995 (gh): 9, upper; 10, lower; 11-12, C. speciosa, Venezuela, Steyermark 61970 (us): 11, upper; 12, lower; 13-14, C. divergens var. divergens, Colombia, Killip 11392 (gh): 13, upper; 14, lower; 15-16, C. Delgadii, Peru, Spruce 4729 (gh): 15, upper; 16, lower.




Figs. 17-22. 17-19. Photographs of indusium types of Cyathea, x 20. ±7, Hemitelioid indusium, C. multi flora, Honduras, Yuncker 4745 (f); 18, Cyathiform indusium, C. arborea , Grenada, Proctor 17270 (a); 19, Sphaeropteroid indusium, C. Lechleri, Venezuela, Steyermark 75444 (gh). 20. Large paraphyses in sori of C. notabilis, Cocos Island, Holdridge 5161 (gh), X 20. 21-22. Scan- ning electron micrographs, spores of Cyathea furfuracea, Cuba, Eggers 5211 (gh): 21, X 1200; 22, X 5000.



28 ROLLA TRYON

Paraphyses are borne on the elevated receptacle among the sporangia. They are usually moderately long and slender, but in Cyathea notabilis (Fig. 20) they are much enlarged.

The spores of Cyathea have not been treated here in detail because of the current research of Gerald J. Gas tony on the palynology of the Cy- atheaceae (Gastony, 1974; Gastony & Tryon, 1976). The scanning electron micrographs included here provide some information on the exine and perine layers in Cyathea . The exine may be verrucate, as in C. furfuracea (Figs. 21, 22), or minutely pitted as in C. arborea (Fig. 23). Spores of the latter are usually without perine, as in Fig. 23, but in a few collections some spores were partially covered by a matrix, including large spherical bodies (Fig. 24) that appear to represent perine. In other species the

perine consists of a fairly dense network of slender rods superimposed on a relatively smooth surface as in C. caracasana var. Maxonii (Fig. 25, 26), or on a coarsely tuberculate surface as in C. diver gens var. diver gens (Fig. 27), and C. Delgadii (Figs. 28, 29). The surface of the slender rods, when examined at high magnification, is seen to consist of fused pelleted material (Figs. 26, 29).

Illustrations

Drawings of the leafy parts: pinnae, pinnules and segments, have been taken, unless otherwise indicated, from central pinnae, from their central pinnules, and from the central portion of a central pinnule. The drawings of enlarged portions of fertile pinnules have the venation and position of the receptacle shown on some of the segments, and the indusia and indûment (if any) are shown on others.

Acknowledgements The support of the National Science Foundation through Grants GB4184 to Rolla

M. Tryon and GB31170 to Rolla M. Tryon and Alice F. Tryon is gratefully acknowl- edged.

I am indebted to Dr. Alice F. Tryon for aid in many aspects of the research, and for the photographs of indusia and SEM micrographs of the spores. Dr. Gerald J. Gastony has provided help in a number of technical matters. The drawings have been prepared by Sarah Landry, Lydia Wunsch and Mary Robbins.

The officers and staff of the following herbaria have been generous in the loan of specimens for study and have extended courtesies during visits to their institutions: Botanische Garten und Museum, Berlin-Dahlem; British Museum (Natural History); Botanical Museum and Herbarium, Copenhagen; Field Museum of Natural History; Royal Botanic Gardens, Kew; Missouri Botanical Garden; New York Botanical Garden; Muséum National d'Histoire Naturelle, Paris; Naturhistoriska Riksmuseet, Stockholm; and United States National Herbarium, Smithsonian Institution.




Figs. 23-29. Scanning electron micrographs, spores of Cyathea. 23-24, C. arborea: 23, Dominican Republic, Gastony et al. 702 (gh), X 1200; 24, Puerto Rico, Conant ir Conant 51 (gh), X 10000. 25-26, C. caracasana var. Maxonii, Costa Rica, Wilbur ir Stone 10623 (gh): 25, X 1200; 26, X 60000. 27. C. diver gens var. diver gens, Costa Rica, White ir Lucansky 1968-30 (gh), X 1200. 28- 29. C. Delgadii , Brazil, Tryon ir Tryon 6658 (gh): 28, X 1200; 29, X 35000.



30 ROLLA TRYON

SYSTEMATIC TREATMENT

Cyathea J. E. Smith

Cyathea Sm. Mém. Acad. Turin 5: 416. 1793. type: Cyathea arborea (L.) Sm. (Polypodium arhoreum L. ). Hemitelia R. Br. Prod. Fl. Nov. Holl. 158. 1810. type: Cyathea multiflora Sm. (no

combination by R. Br. ) . Disphenia Presi, Tent. Pterid. 55. 1836, nom. superfl. type: the same as that of

Cyathea. Cormophyllum Newm. Phytol. 5: 237. 1856, nom . superfl. type: the same as that of

Cyathea.

Cyathea is an American genus of 40 species with its principal development in mountain forests of the Andes from Venezuela and Colombia to Bolivia. Some species of Trichipteris have a scale associated with the sorus that may be mistaken for an indusium, and some species of Cyathea have a thin, fragile, and ephemeral indusium. Problems of identification of the two genera are discussed in the introduction. The generic characters are also given there, in the key to the American scaly genera. Ten indusiate

putative hybrids of Cnemidaria, Cyathea and Trichipteris species are treated in a section at the end of the Systematic Treatment.

Key to Species Groups of Cyathea a. Indusium hemitelíoid, variously developed, usually not completely surrounding the base of the receptacle (rarely surrounding it and then asymmetrical), or discoid to deeply cyathiform b. b. Indusium hemitelioid ( rarely surrounding the receptacle as an asymmetrical disk ) , when large often splitting at maturity into 2-3 segments c. c. Pinnae 1 -pinnate or more complex d. d. Pinnules pinnatifid to 1-pinnate, the ultimate segments predominantly less than half -joined, or completely separated, lamina with a gradually to abruptly reduced apex Group of Cyathea multiflora , species 1-8.

d. Pinnules entire to lobed, or rarely a few segments less than half -joined, rarely deeply lobed and then the lamina with a strictly conform apical pinna Group of Cyathea petiolata, species 9-12.

c. Pinnae entire to pinnatifid Group of Cyathea speciosa , species 13-16. b. Indusium cyatheoid ( discoid to deeply cyathiform ) not splitting at maturity Group of Cyathea arborea, species 17-19.

a. Indusium sphaeropteroid, completely enclosing the immature sorus and usually with an apical umbo (rarely very delicate, especially in C. caracasana, C. palles cens, C. straminea and C. tenera, evanescent and persisting if at all only as an irregular basal disk or portion), very rarely less developed and open on the distal side e. e. Petiole scales whitish to light brownish, concolorous or nearly so, or discordantly bicolorous with the whitish to brownish color of the differentiated margins extending into part of the otherwise dark colored, elongate-celled central portion of the body; petiole scurf whitish to light brownish, or rarely brown at the base of the

rtiole Lamina

wholly bipinnate-pinnatifid, or tripinnate only at the base of the pin- f*

Lamina wholly bipinnate-pinnatifid, or tripinnate only at the base of the pinnae g- g. Flattish scales absent on the pinnules beneath, or if present, then lacking peripheral processes of strongly contrasting color to the body Group of Cyathea pallescens, species 20-22.

g. Some flattish scales on the pinnules beneath with dark, opaque peripheral processes, or apical processes, of strongly contrasting color to the body, or with brown cilia the same color as the central portion of the bicolorous body Group of Cyathea straminea, species 23-26,




f. Lamina tripinnate to quadripinnate nearly throughout Group of Cyathea microphylla, species 27-28.

e. Petiole scales brown and nearly concolorous, or slightly to definitely concordantly bicolorous with light brown to brown margins; petiole scurf brown or absent. . . . h. h. Petiole scales light brown to brown, concolorous to slightly bicolorous Group of Cyathea fulva, species 29-34.

h. Petiole scales (usually all to rarely several) with a reddish-brown to usually dark reddish-brown or atropurpureous central body, or with dark areas or streaks, bicolorous with definitely lighter colored (sometimes narrow) margins i. i. Petiole usually aculeate, to abundantly tuberculate, with abundant (sometimes caducous) scaly scurf; petiole tan to brown or sometimes darker Group of Cyathea caracasana, species 35-36.

i. Petiole lacking spines, nearly or quite smooth or with some scattered tubercules, scaly scurf absent or rarely sparsely present; petiole dark reddish-brown to usually atropurpureous, at least basally Group of Cyathea Lechleri , species 37-40.

Group of Cyathea multiflora KEY TO SPECIES 1-8

a. Paraphyses long, much enlarged; Cocos Island 5. C. notabilis. a. Paraphyses short, to long and slender b. b. Indusium partly to completely arching over the sorus, or (rarely) concealed by the sorus and curving erect c. c. Ultimate segments with simple fertile veins and then usually entire, or with forked fertile veins and coarsely dentate; Central America, Colombia to Bolivia, northern Brazil 1. C. multiflora. c. Ultimate segments with forked fertile veins, entire, finely dentate, crenate to crenately lobed to deeply lobed d. d. Sori supramedial to submarginai e. e. Basal pinnules of the lower pinnae long-stalked or nearly so; Greater Antilles, French Guiana to Colombia and adjacent Brazil, south to Bolivia. . . 2. C. andina.

e. Basal pinnules of the pinnae sessile or nearly so; Galapagos Islands 3. C. Weatherbyana.

d. Sori costal to medial f. f. Sori subcostal to medial, pinnules up to ca 1.5-2 cm broad and 6-8 cm long; Venezuela and Colombia 7. C. platylepls.

f. Sori strictly costal, pinnules ca 3-4 cm broad and 10-14 cm long; Peru 8. C. Vilhelmii.

b. Indusium appressed or subappressed to the segment surface, nearly or quite concealed by the sorus g- g. Petiole smooth or slightly muricate; Cocos Island 4. C. Alphonsiana. g. Petiole aculeate, especially toward the base and usually abundantly so; Greater Antilles 6. C. parvula.

The first five species of this group, including two continental species and three insular endemics, are clearly related. Cyathea multiflora and C. andina are not wholly distinctive species; however, it is in the areas where

they are allopatric that variations occur which approach intermediate

specimens. In the range of sympatry, from Colombia to Bolivia, there is little or no evidence of intergradation.

Cyathea Weatherbyana of the Galapagos Islands is most closely related to C. andina. Cyathea Alphonsiana and C. notabilis of Cocos Island are both closest to C. multiflora. It is remarkable to have two endemics on an island, both derived from the same source species. The most divergent of these, C. notabilis, has unusual enlarged paraphyses (Fig. 44). It perhaps is the older, while the less distinctive C. Alphonsiana may be a younger



32 ROLLA TRYON

endemic. It is also possible that each was derived from a different variation of C. multiflora. The other three species, C. parvula, C. platylepis and C. Vilhelmii are all distinctive but are probably related to C. multiflora and C. andina. Among this group of primitive species, Cyathea parvula appears to be closest to C. arborea and other derived species.

1. Cyathea multiflora Sm. Figs. 30-34. Map 3.

Cyathea multiflora Sm. Mém. Acad. Turin 5: 416. 1793. Holotype: Amer. Merid., Shakespeare , Herb. Banks, bm! photo gh,ny,us, fragm. us! The locality was erroneously published as Jamaica, see Maxon, Bull. Torrey Bot. Cl. 38: 545-550, t. 35. 1911.

Hemitelia multiflora (Sm.) Spreng. Syst. Veg. 4: 126. 1827. Not by R. Br. Fl. Nov. Holl. 158. 1810, as often cited.

Alsophila multiflora ( Sm. ) Presi, Tent. Pterid. 61. 1836. Amphicosmia multiflora (Sm. ) Gardner, Lond. Jour. Bot. 1: 441. 1842. Hemitelia nigricans Presi, Epim. Bot. 31. 1849. Holotype: "Guatemala," ad ripas

fluvii S. Juan, (Nicaragua), Friedrichsthal, W. See Maxon, Bull. Torrey Bot. Cl. 38: 545-550. 1911.

Amphicosmia nigricans (Presi) Moore, Ind. Fil. 61. 1857. Hemitelia obscura Mett. Ann. Sci. Nat. V, 2: 264. 1864. Holotype: Prov. de

Barbacoas, via de Tuquerres, (Colombia), 1600 m, Triana. Isotype: p! fragm. ex col, gh! fragm. ex k,ny!

Hemitelia denticulata Hook. Syn. Fil. 31. 1865. Holotype: "Elizabeth Island, Pacific, near Pitcairn/' Cuming 1360, k! This is clearly Cyathea multiflora and probably from Panama. Holttum, Blumea 12: 274. 1965 excludes it from Pacific species of Cy- atheaceae.

Hemitelia Lindigii Baker, Syn. Fil. 454. 1874. Holotype: Alto del Trigo, Andes of Bogotá, (Colombia), Lindig 310, k. Isotype: p! photo gh.

Hemitelia Hartii Baker, Tour. Bot. 24 (n.s. 15): 243. 1886. Holotype: Chiriqui Lagoon, Panama, Hart 43, k! photo gh. Isotypes: ny! p! photo gh,us!

Alsophila decussata Christ, Pittier, Prim. Fl. Costaric. 3(1) ( Filices 2nd Mém. ) : 41. 1901. Holotype: Vallé de Diquis, Costa Rica, Pittier 12027. Isotypes: ny! p! us!

Alsophila leucolepis var. puhescens Christ, Pittier, Prim. Fl. Costaric. 3(1) (Filices 2nd Mém.): 42. 1901. Syntypes: Cañas Gordas, Costa Rica, Pittier 10981, us!, 10989 (not seen), 10992, p! us! Alsophila acutidens Christ, Bull. Herb. Boiss. II, 6: 186. 1906, based on C. leucolepis

var. puhescens Christ. Hemitelia squarrosa Rosenst. Fedde, Rep. Spec. Nov. 22: 2. 1925. Holotype: Finca

Gebr. Hundriesser, Costa Rica, Brade ò- Brade 405, Herb. S. Birger, s-pa! Isotypes: b! us!

Cyathea acutidens (Christ) Domin, Pterid. 262. 1929. Cyathea austroamèricana Domin, Pterid. 263. 1929, nom. nov. for Hemitelia nigri-

cans Presi, not Cyathea nigricans Mett. Cyathea columbiana Domin, Pterid. 263. 1929, nom. nov. for Hemitelia obscura

Mett., not Cyathea obscura (Mett.) Copel. Cyathea Hartii (Baker) Domin, Pterid. 264. 1929. Cyathea Lindigii (Baker) Domin, Pterid. 264. 1929. Cyathea squarrosa (Rosenst.) Domin, Acta Bot. Bohem. 9: 161. 1930. Hemitelia squamulosa Losch, Mitteil. Bot. Staatssaml. München 1 : 20. 1950, ex char.

Holotype: Orosi, Costa Rica, Kupper 798, m, photo bm! Cyathea leucolepismata Alston, Jour. Wash. Acad. Sci. 48: 231. 1958. Holotype:

near San Diego de Colorado, between Umbria and Puerto Asis, Putumayo, Colombia, Ewan 16748, bm. Isotype: us!




Cyathea multiflora is a variable species, especially in its pinnule in- dûment, petiole scales, segment margins, and paraphyses. In British Honduras and Guatemala to Nicaragua the pinnules have few trichomes beneath and mostly few, brown scales. To the south in Costa Rica to Bolivia, the pinnules are often pubescent beneath with long whitish trichomes and often have whitish scales. The petiole scales are usually relatively small, brown and nearly concolorous, while in Ecuador and Colombia especially, they may be large ( Sparre 18482, GH, Fig. 30) or large and bicolorous ( Cuatrecasas 13918 , F), or they may be mixed with whitish nearly concolorous scales. The segments vary from entire to coarsely toothed. The specimens that have a long pubescence on the pinnules beneath often have long, slender, whitish paraphyses in the sorus, while those with only a few trichomes beneath usually have short and brownish paraphyses.

The variation in these several characters shows some correlations with

geography, and with each other. However, they are weak and do not

provide a basis for the recognition of other taxa. The more uniform ex- treme, especially in Central America north of Costa Rica, is interpreted as

resulting from a geographic differentiation from the more variable southern populations.

British Honduras and Guatemala to Panama, Colombia south to Bolivia, northern Brazil. In dense forest on steep mountain slopes, along creek banks and in ravines, also in open forest or cloud forest, from near sea level, mostly below 500 m, to 1500 m in Central America and up to 2300 m in Colombia, mostly 500-2000 m in South America. Stem often ca 1 m tall, to 5 m, leaves to 2.5 m long.

selected specimens. British Honduras (Belize). Tomash River, Schipp S923 (gh); Camp 34, Schipp S774 (gh). Guatemala. Alta Verapaz: near Chirriacté on Petén highway, Standley 91674 (us). Izabal: Livingston, D earn 483 ( f,gh,mo,ny ) ; Rio Frio, Cerro San Gil, Steyermark 39965 (f,gh,us). Honduras. Atlântida: above Lancetilla, Y uncker 4745 (f,mo); near Tela, Standley 53132 , 53945 (f,gh). Nica- ragua. Cabo Gracias á Dios: Laimos Creek, southwest of Waspan, Bunting ir Licht 396 (gh). Zelaya ( "Bluefìelds" ) : Río Escondido, Bahia de Bluefìelds, Molina 1964 , 2013 (f); Río Rama, Río Escondido, Cerro San Isidro, Proctor , Jones ir Facey 26954 , 27102 (ny). Costa Rica. Alajuela: Quebrada Marin, 7 km east of Ciudad Quesada, Burger ir Stolze 4941 (gh,ny), 4982 (gh). Heredia: Finca La Selva, Río Puerto Viejo, Burger ir Stolze 5828, 5876 (gh); Cariblanco, Nisman 112 (gh). Limon: near Río Madre de Dios, Scamman 6998 (gh). San José: vicinity of El General, Skutch 2161 ( f, gh, mo, ny ) ; along Río Hermosa, Williams , Molina , Williams ir Gibson 28434 (f). Cartago: Río Grande del Orosi, Tryon ir Tryon 7023 (gh), Burger ir Stolze 6094 (gh); vicinity of Pejivalle, Standley ir Valerio 47189 (gh). Puntarenas: San Vito, Me Alpin 2257 (gh); Canton Golfito, Nisman 139 (gh); Osa, near Rincón, Nisman 88 (gh). Panama. Bocas del Toro: vicinity of Chiriqui Lagoon, Wedel 2858 (mo). Herrera: vicinity of Las Minas, Stern , Eyde ir Ayensu 1755 (gh). Colon: Porto Bello, Maxon 5776 (gh,mo,ny). Panamá: La Campana, Welch 19613 (mo); Cerro Campana, Madison 779 (gh). Canal Zone: Quebrada Salamanca, Steyermark ir Allen 17141 (mo). Colombia. Bolivar: Cascade Chorron, south of Antizales, Pennell 4408 (f,gh,mo,ny,us). Antioquia: 8 km west of Valdivia, Madison 805 (gh); Porcesito, valle del Rio Medellin, Hodge 6872, 6873 (gh,us). El Valle: La Cumbre,



34 ROLLA TRYON

Killip 5701 ( GH, us ) ; Monte La Guarida, Cuatrecasas 22161 (f,us); El Cairo, Cuatrecasas 13918 (f,gh,us). Cauca: Río Ortega, north of Tambo, Pennell ù- KiUip 8058 (gh,ny,us); Cauca valley, Río Sucio, Pennell ir Killip 7230 (us). Nariño: Río Guabo, Ewan 16809 (gh,us). Caquetá: Quebrada del Río Hacha, Cuatrecasas 8779 (f,us). Putumayo: San Diego del Colorado, Ewan 16784 (us). Ecuador. Pichincha: Mindo, Sydow 299 (us); Toáchi, Sparve 18482 (gh). Napo-Pastaza: near Archidona, Mexia 7318 (us). Los Ríos: Cerro Mombre, Río Pita, Asplund 5557 (ny,us). Chimborazo: Chimborazo, Spruce 5741 (ny,us). Cañar: east of El Triunfo, Guayaquil- Cuenca road, Madison 913 (gh). Santiago-Zamora: Bombaiza, south of Gualaquiza, Sparre 19099 (gh). Peru. Amazonas: Laguna Pomacocha, Soukup 5260b (gh). Loreto: Gamitanacocha, Río Mazán, Schunke 117 ( gh,mo,ny,usm ) ; San Antonio, Río Itaya, Killip ò- Smith 29379 (f,ny,us). Junin: near La Merced, Killip ir Smith 23930 (f,us), 23979 (f,ny,us). Cuzco: near San Lorenzo, Prov. Quispicanchis, Vargas 16069 (gh); Atalaya, Prov. Paucartambo, Vargas 16274 (gh). Bolivia. La Paz: Mapiri, Williams 1272 (gh,ny,us). Brazil. Upper Amazonas, Traill 1382 (gh,ny,us).

2. Cyathea andina ( Karst. ) Domin Figs. 35-37. Map 4.

Cyathea andina ( Karst. ) Domin, Pterid. 263. 1929. Hemitelia andina Karst. Linnaea 28: 452. 1856. Holotype: Santa Martha, (Colom-

bia), 2500 m, Karsten. Fragment of type collection: Sierra Nevada de Santa Martha, (San Miguel), 6000 ft. Karsten , Herb. Mett. b! photo gh. Chosen over Hemitelia servitensis Karst, because figured in Karst. Fl. Columb. 2: t. 197, f. I.

Hemitelia servitensis Karst. Linnaea 28: 451. 1856. Holotype: Prov. Cundinamarca Novo-Granatae 2000 m, Karsten. Isotypes: Servita, Bogotá, (Colombia), Karsten , Herb. Mett. b! p! photo gh.

Hemitelia Boryana Mett. ex Kuhn, Linnaea 36: 161. 1869. Lectotype: French Guiana, Leprieur 265a, p! det. Mett. Isolectotype: Leprieur 265 , us! Isolectoparatype: French Guiana, Poiteau, k! photo gh. The "a" was evidently added to the number, perhaps by Mettenius to distinguish the specimen from others under the same number. The sheet at us, however, is also this species.

Hemitelia escuquensis Karst. Fl. Columb. 2: 181, t. 196. 1869. Holotype: Sinus Maracaibensis prope Escuque, 100 m, Karsten. Isotype: Hemitelia Escuquensis spec, nov. Escuque, Venezuela, Karsten 28, Herb. Mett. b!

Hemitelia Joadii Baker, Ann. Bot. 5: 187. 1891. Holotype: Santa Marta, (Colombia), Dec. 1863, Joad, k!

Hemitelia Leprieurii Jenm. W. Ind. Guiana Ferns 47. 1898. Holotype: Cayenne, French Guiana, Leprieur, Herb. Jenman, ny, photo gh.

Cyathea Boryana ( Kuhn ) Domin, Pterid. 263. 1929. Cyathea escuquensis ( Karst. ) Domin, Pterid. 264. 1929. Cyathea Joadii (Baker) Domin, Pterid. 264. 1929. Cyathea circumdentata Kramer, Acta Bot. Neerland. 3: 491. 1954, nom. nov. for

Hemitelia Leprieurii Jenm., not Kze. 1844, not Cyathea Leprieurii ( Kze. ) Domin.

The indusium of Cyathea andina is usually larger than that of C. multiflora and often splits into two persistent parts at maturity. The sori are borne at the fork of a vein that is simple basally and forks toward, or sometimes very close to, the margin of the segments. The few collections from French Guiana, British Guiana and adjacent Brazil are not typical of the species. They are interpreted as geographically peripheral variations somewhat comparable to those of Cyathea multiflora in Nicaragua and Guatemala.

Hispaniola and Puerto Rico, French and British Guiana to Colombia, south to Bolivia, northern Brazil. In montane forest, occasionally in open




places in forests, 250-900 m in the Greater Antilles, and 50-2500 m, usually 1000-1500 m in South America. Stem to 8 m tall, leaves to 4 m long.

selected specimens. Haiti. Vicinity of Plaisance, Leonard 9308 (gh,ny,us). Dominican Republic. Distr. Sabaneta, Monte Cristi, Valeur 566 (gh); La Cumbre, Santo Domingo, Ekman H11499 (ny,us). Puerto Rico. Maricao Forest, Little 21704 (gh); Maricao, Hioram 808 (us); near Adjuntas, Stevens 5950 (ny); Adjuntas, Sintenis 4102 (mo). French Guiana. Near Saul Village, near Crique Cochow, Bier- horst FG87 , FG110 (gh). British Guiana (Guyana). Basin of Shodikar Creek, Essequibo tributary, A. C. Smith 2873 ( f,gh,mo,ny,us ) . Brazil. Roraima: Serra Tepequem, Prance , Forero , Pena ir Ramos 4470 (ny,us). Venezuela. Bolivar: Auyan- tepui, Schnee 1487 (ven). Zulia: Sierra de Peri já, between Río Negro and Río Apon, Vareschi 3183 (ven). Colombia. Magdalena: Santa Marta, H. H. Smith 1017 , 2641 ( F, gh, mo, ny, us ) . Santander: vicinity of Barranca Bermaja, Magdalena valley, Haught 1358 (gh,us). Cundinamarca: Fusagasuga, André 2375 (ny). Meta: Cordillera La Macarena, Idrobo ó- Schultes 748 (a,gh,us). Putumayo: Mocoa, Cuatrecasas 11325 (us). Ecuador. Chugiriraja, André 1134 (ny). Peru. San Martin: La Divisoria, Ferreyra 1694 (us). Junín: east of Quimiri Bridge, near La Merced, Killip ò- Smith 23995 (gh,ny,us); Prov. Tarma, Esposto 659 (usm); Chanchamayo valley, Schunke 53 (f). Puno: Prov. Carasaya, Vargas 17536 (gh). Bolivia. La Paz: Tajlini, northeast of La Paz, Quesada 3 ( us ) .

3. Cyathea Weatherbyana (Morton) Morton Figs. 38-39. Map 5.

Cyathea Weatherbyana (Morton) Morton, Amer. Fern Jour. 59: 65. 1969. Hemitelia Weatherbyana Morton, Leafl. West. Bot. 8: 188. 1957. Holotype: Inde-

fatigable Island, Galapagos Islands, Howell 9227 , us! Isotype: cas. Paratypes (all Galapagos Islands): Villamil, Albemarle Island, Stewart 894 , cas, us; James Bay, James Island, Stewart 896 , 897 , cas, us; Wreck Bay, Cahtham Island, Stewart 895, cas, us. Isoparatypes : Stewart 894 , gh! mo!; 895 , 896 , 897, f! gh! mo! ny!

The whitish, strongly dissected scales on the pinna-rachises and costae are a distinctive character of Cyathea Weatherbyana. The indusium is

moderately large and membranous. This endemic species is clearly related to the continental C. andina.

Galapagos Islands, in the fern-sedge zone and the Miconia zone on mountain sides at 450-800 m. Stem to 6 m tall.

additional specimens. Galapagos Islands. Santa Cruz (Indefatigable): Mt. Crocker, Wiggins 18561 (gh,us); Colinvaux 357 (gh); above Fortuna, Harling 5084 (gh).

4. Cyathea Alphonsiana Gómez Figs. 40-41. Map 6.

Cyathea Alphonsiana Gómez, Amer. Fern. Jour. 61: 166, f. 1, 2. 1971. Holotype: Twin Mountains, upper Wafer Valley, Cocos Island, Costa Rica, Gómez 3394, cr. Isotypes: gh! us!

Cyathea Alphonsiana is most closely related to C. multiflora, from which it differs especially in its minute, appressed indusium and usually entire segments with forked veins. The specimens cited below were also indicated as paratypes by Gómez, although not seen by him.

Cocos Island, in dense forest, 450 m. Stem to 4 m tall. additional specimens. Cocos Island. Holdridge 5160 (gh); W. L. Schmitt 129,

130,131 (us).



36 BOLLA TRYON

Figs. 30-39. 30-34, Cyathea multiflora: 30, Petiole scales, Ecuador, Sparre 18482 (gh), X Vz' 31, Pinnules, Costa Rica, Scamman 6998 (gh), X 32, Pinnules, source as in 30, X ft; 33, Portion of fertile pinnule, source as in 30, X 2; 34, Portion of fertile pinnule, source as in 31, X 2. 35-37, C. andina: 35, Pinnules, Colombia, Mutis 3291 (us), X 36, Pinnules, British Guiana, A. C. Smith 2873 (gh), X 37, Portion of fertile pinnule, Puerto Rico, Little 21704 (gh), X 38-39, C. Weatherbyana, Galapagos Islands: 38, Pinnules, Stewart 897 (mo), X 39, Portion of fertile pinnule, Wiggins 18561 (gh), X 2.




5. Cyathea notabilís Domin Figs. 42-44. Map 7.

Cyathea notabilis Domin, Acta Bot. Bohem. 9: 141. 1930, nom. nov. for Álsophila notabilis Maxon, not Saporta, Mém. Soc. Géol. France II, 8: 329. 1868.

Álsophila notabilis Maxon, Contrib. U.S. Nat. Herb. 24: 39, t. 12. 1922. Holotype: Wafer Bay, Cocos Island, Costa Rica, Pittier 12355, us! Paratype: same locality, Pittier 16228, us! Isoparatype: gh!

Cyathea notabilis is a unique species with its much enlarged paraphvses, many of them forming a conspicuous tuft at the apex of the receptacle. There are ten or fewer sporangia in a sorus, in the materials I have seen, while other Cyathea species with smaller paraphyses have more sporangia. Gastony (1974) reports five species of Cyathea with 44 up to 106 sporangia per sorus. The apparent reduction of reproductive capacity in C. notabilis may be interpreted as the development of precinctiveness (Carlquist, 1966) in this insular endemic. Among the ferns, it is perhaps the only example of this phenomenon.

Cyathea notabilis also differs from C. Alphonsiana of Cocos Island in its predominently simple fertile veins and in its minute indusia.

Cocos Island, in forested ravines, 50 m. Stem to 2 m tall.

additional specimens. Cocos Island. Jiménez 3146 (gh); Holdridge 5161 (gh); Schmitt 128 ( gh,us ) .

6. Cyathea parvula (Jenm.) Domin Figs. 45-47. Map 8.

Cyathea parvula (Jenm.) Domin, Pterid. 264. 1929. Álsophila parvula Jenm. Jour. Bot. 17 (n.s.8): 258. 1879. Holotype: Jamaica, Jen-

man 97, K, photo gh,ny,us, fragm. us! One sheet (of four) in the Jenman Herbarium, ny! lacks data but is very similar to the holotype and may be considered authentic.

Hemitelia microsepala Jenm. Jour. Bot. 24 (n.s.15): 266. 1886. Holotype: Bull Head, Clarendon Parish, Jamaica. Sherring ?. Three sheets without data in the Jenman Herbarium, ny! are probably authentic.

Hemitelia parvula (Jenm.) Baker, Ann. Bot. 5: 188. 1891. Álsophila aquilina Christ, Engl. Bot. Jahrb. 24: 83. 1897, "1898." Holotype: Monte-

verde, Cuba, Eggers 5117, fragm. ex Christ, us! Isotypes: f! p! photo gh, fragm. ex k, ny!

Álsophila gracilis Underw. & Maxon, Bull. Torrey Bot. Cl. 29. 577. 1902. Holotype: Baracoa, Prov. Santiago, Cuba, Pollard, Palmer ö- Palmer 255, us! Isotypes: f! gh! mo! Paratypes: Monteverde, Cuba, Wright 951, ny! us. Isoparatypes : gh! mo! ny!

Álsophila aquilina var. Maxonii Rosenst. Fedde, Rep. Spec. Nov. 6: 179. 1908. Holo- type: valley of Río Bayamita, Sierra Maestra, Cuba, Maxon 3910 . Isotypes: gh! ny! us!

Hemitelia miniuscula Maxon, Jour. Wash. Acad. Sci. 18: 316. 1928. Holotype: Anse-à-Toleur, Morne Colombeau, Massif de Nord, Haiti, Ekman H 4365, c. Isotype: us!

Cyathea aquilina (Christ) Domin, Pterid. 262. 1929. Cyathea gracilescens Domin, Pterid. 262. 1929, nom. nov. for Álsophila gracilis

Underw. & Maxon, not Cyathea gracilis Griseb. Cyathea parvula var. microsepala (Jenm.) Domin, Acta Bot. Bohem. 9: 145. 1930. The petiole scales of Cyathea parvula are often concolorous and whitish;

they vary to bicolorous with a darker center. The pinnules often have light to dark brown búllate scales and (or) flattish scales beneath. There is



38 ROLLA TRYON

considerable variation in the leaf texture between plants growing in the open sun and those growing in the forest. Although the indusium is usually small, it is especially well- developed in Conant ¿r Hodgdon 646 , Puerto Rico (GH), in which it may extend about halfway around the receptacle and arch over the lower portion of the sorus. These variations, among others, are found scattered through the Greater Antilles and are not correlated with each other nor with their geography.

A notable feature of Cyathea parvula, at least in the Maricao Mountains of western Puerto Rico, is that the stems may produce positively geo- trophic branches, some of which extend to the ground and take root. A similar phenomenon has been reported in Alsophila Manniana of Africa

by Halle (1966, as Cyathea Manniana). Cuba, Jamaica, Hispaniola and Puerto Rico. In rain forest, shaded

ravines, and mountain valleys, or more often in open woods, along streams, at the edge of woods, in thickets or savannahs, and along roadsides and in

secondary growth, from sea level to 2000 m. Stem to 9 m tall, frequently ca 1-2 m, leaves to 2 m long, usually less.

selected specimens. Cuba. Oriente: northern spur of Sierra Maestra, west of Rio Yao, Morton & Acuna 3487 (gh,mo,ny); Sierra Nipe, near Woodfred, Shafer 3059 (gh,ny); Sierra de Moa, Howard 5859 , 5963 (gh,mo,ny); Wright 889 (gh,mo,ny). Jamaica. Mount Diabolo, Portland, Maxon &■ Killip 441 (f,gh,ny); Tweedside, St. Andrew, Proctor 4460 (mo); Corn Puss Gap, St. Thomas, Proctor 5513 (gh); Mason River Field Station, Clarendon, Riba 209 (gh). Haiti. Vicinity of Port de Paix, Leonard & Leonard 12316 (mo); vicinity of Plaisance, Leonard 9350 (us); vicinity of Marmelade, Leonard 8227 (f,gh,mo); Bayeux, Morne Brigand, Ekman H2873 (ny). Dominican Republic. Vicinity of San Francisco de Macoris, Duarte (Pacificador), Abbott 2067 ( gh,ny ) ; Pilón de Azúcar, vicinity of Laguna, Samaná Peninsula, Abbott 294 , 445 (gh); Distr. San José de las Matas, Santiago, Valeur 317 (f,mo,ny); Seibo, from Miches to Higuey, Gastony , Jones ò- N orris 656, 659, 660 (gh,ny). Puerto Rico. Río Maricao, Britton , Stevens & Hess 2482 (f,mo,ny); Maricao State Forest, Tryon ir Tryon 7083 (gh); Espino, Sintenis 5960 (a,f,mo,ny); near Mayaguez, Heller 4595 ( f,gh,mo,ny ) .

7. Cyathea platylepis ( Hook. ) Domin Figs. 48-51. Map 9.

Cyathea platylepis ( Hook. ) Domin. Pterid. 264. 1929. Hemitelia platylepis Hook. Second Cent. Ferns t. 100. 1861. Holotype: near San

Carlos, Rio Negro, "Brazil," (Venezuela), Spruce 3127 (not 3027 as published), k! photo gh. Isotypes: gh! p! fragm. ex Christ, ny!

Hemitelia minima Morton, Fieldiana: Bot. 28: 9. 1951. Holotype: Ptari-tepuí, Boli- var, Venezuela, Steyermark 59481 , f!

Cyathea platylepis is a distinctive species of the Roraima sandstone and derived soils of the Guayana region of Venezuela and Colombia. The fertile pinnae are characteristically dimorphic. The large indusium is

usually half-globose or nearly so; or it sometimes is about % of a deep cup. Venezuela and Colombia. In rocky woods, bamboo thickets, mossy

forest, moist scrubby woods and boggy savannahs, 125-2200 m. Stem

usually short, ca 1 m tall, to 3.5 m, leaves usually 1-1.5 m long, or less.




Figs. 40-51. 40-41, Cyathea Alphonsiana, Cocos Island, Schmitt 131 (us): 40, Pinnules, X Vz; 41, Portion of fertile pinnule, X 2. 42-44, C. notabilis, Cocos Island, Pittier 16228 (gh): 42, Pinnules, X xk; 43, Portion of fertile pinnule, X 2%; 44, Sorus with few sporangia and many enlarged paraphyses, X 15. 45-47, C. parvula: 45, Pinnules, Cuba, Maxon 3910 (us), X 46, Portion of fertile pinnule, Jamaica, Proctor 5060 (mo), X 2; 47, Portion of fertile pinnule, Haiti, Leonard & Leonard 14535 (us), X 2. 48-51, C. platylepis: 48, Sterile pinna, Venezuela, Maguire 33083 (us), X 49, Fertile pinnules, source as in 48, X 50, Portion of fertile pinnule, source as in 48, X 2; 51, Portion of fertile pinnule, Colombia, Schultes à- Cabrera 15079 (gh), X 2.



40 ROLLA TRYON

Maps 3-9. 3, Cyathea multiflora; 4, C. andina; 5, C. Weatherbyana; 6, C. Alphonsiana; 7, C. notabilis; 8, C. parvula; 9. C. platylepls.




selected specimens. Venezuela. Bolivar: Cerro Venamo, Steyermark ir Nilsson 122 (us,ven); Chimatá Massif, Steyermark 74905 , 75451 (ny,us,ven); Auyan-tepui, Steyermark 93922 (gh,us,ven); Amazonas: Cerro de la Neblina, Maguire, Wuraack ir Bunting 37261 (gh,ny,us); Maroa, Río Guainía, Lt. Williams 14328 (f); Cerro Yapacana, Steyermark ir Bunting 103084 (gh,ny); Cerro Autana, Steyermark 105183 (ven). Colombia. Vaupés: Río Vaupés, Mitú, Schuttes , Raffauf ir S o ej art o 24221 (gh); Río Apaporis, Cachivera de Jirijirimo, Schuttes ir Cabrera 12469 (gh,us); Cerro Isibukuri, Schuttes ir Cabrera 15079 (gh). Amazonas: Río Caquetá, vicinity of La Pedrera, Schuttes 5857 ( gh ) .

8. Cyathea Vilhelmii Domin Figs. 52-53. Map 10.

Cyathea Vilhelmii Domin, Pterid. 264. 1929, nom . nov. for Hemitelia Lechleri Mett., not Cyathea Lechleri Mett.

Hemitelia Lechleri Mett. Fil. Lechl. 2: 28. 1859. Lectotype: Tatanara, (Puno), Peru, Lechler 2654 , Herb. Mett. b! Isolectotype: Fragm. ex Rosenst. us! Lectoparatype: Tatanara, Peru, Lechler 2650 , Herb. Mett. b!

Hemitelia Lechleriana Diels, Nat. Pflanz. 1(4): 131. 1899, nom. superfl. Syntypes: Lechler 2650, 2654, Herb. Mett. b!

The correct name of this species depends on the nomenclatural status of Hemitelia Lechleriana Diels, which, if legitimate, would provide the earliest epithet under Cyathea. Diels' name is not obviously superfluous since Mettenius did not cite Lechler collection numbers, and since the specimens in Mettenius' herbarium may have been duplicates, with the original sheets at Leipzig. However, since it is now impossible to resolve the status of these specimens with respect to Leipzig, I believe it is better to consider them as the type collections, which they may have been, and therefore Hemitelia Lechleriana becomes superfluous.

The costal sori and tripinnate-pinnatifid lamina are distinctive characters of Cyathea Vilhelmii. It is probably closely related to the previous species, Cyathea platy lepls.

Group of Cyathea petiolata key to species 9-12

a. Lamina with a gradually to abruptly reduced apex, sori submarginal; Panama, Colombia 9. C. petiolata.

a. Lamina with a conform apical pinna b. b. Stalk of apical pinna not articulate, sori about medial c. c. Pinnules subcordate at base, those of the upper pinnae and the conform apical pinna regularly lobed; Panama 10. C. impar.

c. Pinnules mostly cuneate to subcuneate at base, those of the upper pinnae and the conform apical pinnae entire; Venezuela 11. C. Steyermarkii.

b. Apical pinna articulate, sori submarginal; Panama, Colombia. . . 12. C. conformis. The species of this group are all quite distinctive and, with the exception

of Cyathea petiolata , are known from only one or a few collections. They form a natural group with reduced lamina architecture that appears to be the modern representative of a line leading toward Cnemidaria. Cyathea petiolata and C. conformis , especially, show affinities to Cnemidaria in their pinnules which sometimes have areolate venation. However, Cnemi- daria amabilis , the most primitive species of the genus (Stolze, 1974) has



42 ROLLA TRYON

pinnatifid pinnae and large-porate spores and there is no species of Cyathea that is really close to it. Considerable evolution has probably occurred since the origin of Cnemidaria from hemitelioid indusiate cyatheas.

9. Cyathea petiolata ( Hook. ) Tryon, comb. nov. Figs. 54-58. Map 11.

Hemitelia petiolata Hook. Sp. Fil. 1: 31, t. 16. 1844. Holotype: Isthmus of Panama, Dr. Sinclair, k ( t. 16 ) .

Hemistegia marginalis Presi, Gefässb. Stipes der Farm 47. 1847 (preprint from Abh. Böhem. Ges. Wiss. V, 5: 355. 1848), nom. superfl.

Cyathea panamensis Domin, Pterid. 264. 1929, nom. superfl. An intended nom. nov. for Hemitelia petiolata Hook., not Cyathea petiolata J. Sm. Hook. Jour. Bot. 3: 419. 1841, but the latter is a nom. nud.

Hemitelia Woronovii Maxon & Morton, Amer. Fern Jour. 36: 91. 1946. Holotype: Peñas Blancas, Antioquia, Colombia, Woronow b- Juzepczuk 4549 , le. Isotype: us. Paratypes: Peñas Blancas, Antioquia Colombia, Woronow ò- Juzepczuk 4584 , le, us; Norosi-Tiquisio Trail, Lands of Loba, Bolivar, Colombia, Curran 136 , us!; Barranca Bermeja, Magdalena Valley, between Sogamoso and Colorado Rivers, Santander, Colombia, Haught 1337, us. Isoparatypes: Haught 1337, f! gh! ny!

Cnemidaria petiolata ( Hook. ) Copel. Gen. Fü. 97. 1947. Cyathea Woronovii (Maxon & Morton) Stolze, Fieldiana: Bot. 37: 81. 1974.

Cyathea petiolata is variable in its leaf architecture. The material described as Hemitelia Woronovii is especially different with its large ultimate segments, but in the absence of other characters, this and other variations do not seem to have taxonomie significance.

Two collections from Peru, Dudley 10616 (gh) and 13214 (us), are sterile but probably represent a new species allied to Cyathea petiolata.

Panama and Colombia. In wet forest, along streams and on hillsides, 30-700 m. Stem to 5 m tall, leaves to 1.5 m long.

selected specimens. Panama. Panamá: near Cerro Azu, road to Cerro Jefe, Blum , Godfrey ò- Tyson 1700, 1701 (flas). Canal Zone: Frijoles, Killip 2802 (gh,mo); Caño Quebrada, Pittier 6820 (gh,us); near Gatun, Maxon 4645 (gh); Chagres, F endler 417 (gh,mo), 421 (mo). San Bias: Hills of Siridi, near Puerto Obaldia, Pittier 4412 (gh). Colombia. Chocó: Bahi Solano, near Ciudad Mutis, Killip ò- Garcia 33624 (ny). Santander: 25 km east of Puerto Wilches, Magdalena Valley, Elias 4 (us). Nariño: Gorgona Island, KiUip ir Garcia 33173 (gh,ny), Barclay 908 (gh).

10. Cyathea impar Tryon, spec. nov. Figs. 59-60. Map 12.

Folium 1 m longum; petiolus squamis structura marginata discordanter bicoloribus vel concoloribus pallide brunneis; lamina obovata bipinnato-lobata, pinna apicali con- formi 1-pinnato-lobata non articulata; pinnae ad 20 cm longae apice sensim acuminato; pinnulae pagina inferiore breviter pubescenti; sori mediales in venis simplicibus; indusium hemitelioideum semicyathiforme vel fere cyatheoideum. Holotype: northeast slope of Cerro Jefe, on road to Buenos Aires, Prov. Panamá, Panamá, 2600 ft, 27 January 1966, Tyson, Dwyer è- Blum 3264, mo.

A distinctive species from Cerro Jefe which is considered by Lewis (1971) to be a center of local endemism.




Maps 10-21. 10, Cyathea Vilhelmii; 11, C. petiolata; 12, C. impar; 13, C. Steyermarkii; 14, C. conformis; 15, C. decorata; 16, C. parva; 17, C. speciosa; 18, C. Haughtii; 19, C. arborea, the line between Puerto Rico and Tortola indicates the geographic break in indusium variation (see text); 20, C. Alstonii; 21, C. peladensis.



44 ROLLA TRYON

Figs. 52-63. 52-53, Cyathea Vilhelmii, Peru, Lechler 2654 (b): 52, Pinnules, X Vi ; 53, Portion of fertile pinnule, X IV2. 54-58, C. petiolata: 54, Pinnules, Panama, Maxon 5769 (gh), X V2; 55, Pinnules, Colombia, Killip ir Garcia 33173 (gh), X %; 56, Pinnules, Colombia, H aught 1337 (gh), X V2 ; 57, Portion of fertile pinnule with areolate venation, source as in 54, X 1 V2; 58, Portion of fertile pinnule with free venation, source as in 56, X 1%. 59-60, C. impar, Panama, Tyson et al. 3264 (mo): 59, Pinnules, X V2; 60, Fertile pinnule, X 2. 61-63, C. Steyermarkii, Venezuela, Steyermark 105194 (gh): 61, Apex of lamina, X 62, Pinnules, X x/a' 63, Portion of fertile pinnule, X 1^.




Figs. 64-71. 64-66, Cyathea conformis, Panama, Stern ò- Chambers 188 (gh): 64, Apex of pinna, X 65, Base of apical segment of pinna, enlarged; 66, Portion of fertile pinnule, X 2. 67-68, C. decorata, Colombia, Killip 5257 (gh): 67, Pinnae, X 68, Portion of fertile pinna, X 2. 69-71, C. parva , Colombia, KiUip 5254 (ny): 69, Pinna, X 70, Portion of fertile pinna, X 1%; Portion of alate petiole, diagrammatic, enlarged.



46 ROLLA TRYON

11. Cyathea Steyermarkii Tryon Figs. 61-63. Map 13.

Cyathea Steyermarkii Tryon, Rhodora 74: 449. 1972. Holotype: Cumbre del Cerro Autana, Amazonas, Venezuela, Steyermark 105194, gh!

This species is known only from the small summit of Cerro Autana, which is a single massive projection rising some 1000 m above the surrounding land (Steyermark, 1974).

Venezuela. In woods, ca 1250 m. Stem 1 m tall, leaves ca 1.5 m long.

12. Cyathea conformis (Tryon) Stolze Figs. 64-66. Map 14.

Cyathea conformis (Tryon) Stolze, Fieldiana: Bot. 37: 80. 1974. Hemitelia conformis Tryon, Rhodora 62: 1, f. 1. 1960. Holotype: Piñas Bay, Pro v.

Darien, Panamá, Stern ù- Chambers 188, gh! Isotypes: a! mo! us! Cnemidaria conformis (Tryon) Tryon, Contrib. Gray Herb. 200: 51. 1970. The articulate apical pinna of the lamina is unique in the genus, and

the articulate apical pinnule of the pinnae is (Fig. 65) unique in the family. This species evidently illustrates the evolution of reduced lamina architecture by abortion of the apical portion of the lamina and of the pinnae. However, the process may be based on a different type of leaf modification in other species.

Panama and Colombia.

additional specimens. Seeman 990 (bm). Panamá. Seeman (k). Colombia. Valle or Cauca: Bay of Chocó, 1848, Seeman (bm).

Group of the Cyathea speciosa KEY TO SPECIES 13-16

a. Pinnae deeply lobed to pinnatifid b. b. Petiole not alate, densely and usually persistently scaly; rachis abundantly long pubescent 13. C. decorata.

b. Petiole prominently green alate, sparingly scaly; rachis lacking long trichomes. . . 14. C. parva.

a. Pinnae entire to shallowly lobed c. c. Larger pinnae ca 20 cm or more long, stalked pinnae acute to caudate, rarely some subobtuse 15. C. speciosa.

c. Larger pinnae ca 2-4 cm long, all pinnae obtuse 16. C. Haughtii. A group of distinctive and probably not closely related species. They all

represent a continuation of the trend in reduction of lamina architecture evident in the previous group. The most reduced is Cyathea Haughtii with a small 1-pinnate lamina. Although these species of Cyathea have their lamina architecture comparable to that of Cnemidaria , they are evidently convergent in this character. A closer relationship with Cnemidaria is seen in the previous Cyathea petiolata group.




13. Cyathea decorata (Maxon) Tryon, comb. nov. Figs. 67-68. Map 15.

Hemitelia decorata Maxon, Jour. Arn. Arb. 27: 439, t. 1. 1946. Holotype: Rio Yuru- mangui, El Valle, Colombia, Cuatrecasas 15737 , us. Isotype: f! Paratypes (all us): El Valle: Córdoba, Dagua Valley, Killip 5257 ; Costa del Pacifico, Río Cajambre, Cuatrecasas 17429 ; Agua Clara, Buenaventura to Cali, Killip ò- Cuatrecasas 38914. Chocó: Río Condoto, between Quebrada Guarapo and Mandinga, Killip 35192 ; Corcovada Region, upper San Juan, Killip 35334 . Isoparatypes: Killip 5257 , gh! ny!, Cuatrecasas 17429, f!, Killip 35192 , gh!

The segments of Cyathea decorata are long pubescent on both surfaces and the petiole may be densely and persistently scaly throughout.

Colombia, in wet forest, 5-200 m. Stem to 1 m tall, leaves to ca 1.25 m long.

additional specimen. Colombia. Valle: 22 km. from Buenaventura on road to Rio Calima, Barrington 504 ( gh ) .

14. Cyathea parva (Maxon) Tryon, comb. nov. Figs. 69-71. Map 16.

Alsophila parva Maxon, Jour. Wash. Acad. Sci. 34: 48, f. 1. 1944. Holotype: near Córdoba, Dagua Valley, El Valle, Colombia, Killip 5254 , us! Isotypes: gh! ny!

The petiole and rachis are broadly, membraneously green alate (Fig. 71), a character unique in the genus. Other species may have the rachis green alate only toward the apex of the lamina. The indusium is small and appressed.

Colombia. In forest, ca 90 m. Stem a few cm tall, leaves to 75 cm long.

15. Cyathea speciosa Willd. Figs. 72-74. Map 17.

Cyathea speciosa Willd. Sp. Pi. 5: 490. 1810. Holotype: Caripe (Venezuela), Humboldt, Herb. Willd. 20179, b! photo gh. Isotype: Fragm. ex Herb. Kaulf, Herb. Luerssen, p!, photo gh.

Hemitelia speciosa (Willd.) Kaulf. Enum. Fil. 252. 1824. Hemitelia integrifolia Kl. Linnaea 18: 539. 1844. Syntypes: Caracas (Venezuela),

Otto 671, Moritz 107. Lectotype: Otto 671, b! Hemitelia Lindenii Hook. Icones Pl. t. 706. 1848. Holotype: Caracas (Venezuela),

Linden 663, k! Cyathea integrifolia ( Kl. ) Domin, Pterid. 264. 1929. Cnemidaria integrifolia (Kl.) Tryon, Contrib. Gray Herb. 200: 52. 1970. Cnemidaria Lindenii (Hook.) Tryon, Contrib. Gray Herb. 200: 52. 1970.

The shape of the apex of the pinnae close to the apex of the lamina varies from usually obtuse to rarely attenuate. The lamina apex is sometimes gradually reduced, or it may be abruptly reduced; rarely it is a nearly conform apical pinna.

There is a correlated pattern of variation in the position of the sori and the lobing of the margin of the pinnae. When the margin is quite entire, the sori are submarginai or nearly so (Fig. 74), but when the margin is lobed, the sori are nearly medial, especially those on the basal veins



48 ROLLA TRYON

(Fig. 73). The extremes have often been recognized as distinct species but they clearly represent variations within a single species. Hooker ( Species Filicum t. XIIIB ) illustrated the variation with entire pinnae as Hemitelia speciosa. Klotsch pointed out that this was not the species of Willdenow, and named it Hemitelia integrifolia. Hooker did not respond to this suggestion because he later described the lobed variation as a new

species, Hemitelia Lindenii, a second name for Hemitelia speciosa sens, str. This confusion is only of historical interest since it now involves a

single species. Venezuela and Colombia. In dense wet forest, cloud forest, in ravines

and gorges, 1000-1700 m. Stem to 2 m tall, leaves to 2 m long. selected specimens. Venezuela. Monagas: southwest of Caripe, Steyermark 61970

(mo, NY, us, ven). Distrito Federal: Cerro Naiguatá, Steyermark 92132 (gh,ven); cerca de Agua Negra, Vittier 13740, 13741 (ven). Aragua: Parque Nacional Henry Pittier, Steyermark 89750 (gh,ny,ven); Parque Nacional, Pittier 14997 (ven); Colonia Tovar, F endler 46 (gh,mo). Colombia. Magdalena: Sierra Nevada de Santa Marta, Martin 3395 (mo,us); Santa Marta, H. H. Smith 1124 (gh,mo).

16. Cyathea Haughtii (Maxon) Tryon, comb. nov. Figs. 75-76. Map 18.

Alsophila Haughtii Maxon, Jour. Wash. Acad. Sci. 34: 46. 1944. Holotype: Cerro Armas, Santander, Colombia, Haught 1957 , us! Isotype: us!

Cyathea Haughtii , with a very small stem and leaves 20-30 cm (to 40

cm) long, is one of the smallest species in the family. Colombia. Sandstone cliffs, ca 1400 m.

Group of Cyathea arborea KEY TO SPECIES 17-19

a. Lamina bipinnate-pinnatifid, to partly tripinnate; indusium with entire margin; West Indies 17. C. arborea.

a. Lamina pinnate-pinnatifid; Colombia b. b. Ultimate segments not pubescent above or beneath, a few long trichomes may be present and almost confined to the costules; indusium with the margin irregular, sparingly short dentate-ciliate 18. C. Alstonii.

b. Ultimate segments pubescent above and beneath; indusium with the margin long- ciliate with trichomes similar to those on the adjacent leaf surface. 19. C. peladensis. The complete indusium of Cyathea arborea with an entire margin is a

unique character in the genus. The few collections of C. Alstonii and of C. peladensis have several differences which may prove to be species characters. The petiole scales of C. Alstonii are bicolorous, the lamina is herbaceous in texture, there are very few brownish búllate scales with obtuse to acuminate apex on the costules beneath, and the indusium is

cyathiform to deeply cyathiform. The petiole scales of C. peladensis are concolorous, the lamina is rigidly coriaceous, there are many light brown to brown búllate scales with a long-attenuate apex on the costules beneath, and the indusium is meniscoid to cyathiform (Fig. 84).

These three species are distinctive but evidently form a natural group, probably derived from a species such as Cyathea parvula.




Figs. 72-80. 72-74, Cyaťhea speciosa: 72, Pinna, Colombia, H. H. Smith 1124 (mo), X Ví' 73, Portion of fertile pinna with nearly lobed margin, Venezuela, Steyermark 61970 (ny), X 1%; 74, Portion of fertile pinna with nearly entire margin, Venezuela, F endler 46 (gh), X 1%. 75-76, C. Haughtii , Colombia, Haught 1957 (us): 75, Leaf, X Vz; 76, Fertile pinnae, X 1%. 77-78, C. arborea , Cuba, Maxon 3909 (gh)? 77, Pinnules, X %; 78, Portion of fertile pinnule, X 2. 79-80, C. Alstonii , Colombia, Idrobo à- Schuttes 1130 (us): 79, Pinna, X 80, Portion of fertile pinna, X 1%.



50 ROLLA TRYON

17. Cyathea arborea ( L. ) Sm. Figs. 77-78. Map 19.

Cyathea arborea (L. ) Sm. Mém. Acad. Turin 5: 417. 1793. Polypodium arboreum L. Sp. PL 2: 1092. 1753. Holotype: Morne de la Calebasse,

Martinique. Filix arborescens, pinnulis dentatis Plumier, Fil. Amer. (Traité Foug. ) 1, t. 1; Descr. Pl. Amer. 1, t. 1, 2. Plum. Fil. Amer. t. 2, which Linnaeus did not cite is the same as Descr. Pi. t. 2, except that the sorus and indusium are shown.

Cyathea serra Willd. Sp. PL 5: 491. 1810. Holotype: "Caracas," Bredemeyer , Herb. Willd. 20169, b! photo gh, fragm. us! A variation with narrow and lobed fertile pinnules. The species is not known from Venezuela. The specimen may have been obtained from a cultivated plant, or more likely from Puerto Rico where Bredemeyer visited before going to Venezuela.

Cyathea guadalupensis Spreng. Acad. Caes. Leopold. Nova Acta 10: 233. 1821, ex char. In Guadalupa, Bertier, Perrin.

Hemitelia serra (Willd.) Desv. Mém. Soc. Linn. Paris 6: 321. 1827. Disphenia arborea ( L. ) Presi, Tent. Pterid. 56. 1836. Cyathea arborea var. nigrescens Hook. Sp. Fil. 1: 17. 1844. Type: The same as that

of Polypodium arboreum L. Hemitelia arborea (L. ) Fée, Mém. Fam. Foug. 5 (Gen. Fil.): 350. 1852. Although

there is reference neither to Linnaeus, nor to Polypodium arboreum, the basionym can be readily traced through the reference to Presi.

Cormophyllum arboreum (L.) Newm. Phytol. 5: 238. 1854. Cyathea nigrescens (Hook.) J. Sm. Ferns Brit. For. 242. 1866. Cyathea barbata Kuhn, Linnaea 36: 164. 1869. Holotype: Guadaloupe, V Herminier

3, b! fragm. ny! Isotype: p! photo gh, fragm. ex k, ny!

The petiole scales are usually whitish and concolorous, or they may have tinges of very light brown centrally or basally. They vary to rarely also tinged with brown along the edges ( Hodge 7, Dominica, gh) or brown at the base ( Proctor 21785, Martinique, gh). Sterile leaves usually have a white scale at the base of the costule beneath.

The indusium is always complete and surrounds the receptacle. It varies in its development from discoid or meniscoid with an only slightly, if at all, elevated margin, to deeply cyathiform. A geographic analysis of this variation (Table 2) indicates that the moderately well-developed indusia (deeply meniscoid) are rather equally distributed throughout the range, while the better developed indusia form 77 % of the sample from the Greater Antilles, and the least developed ones form 51% of the sample from the Lesser Antilles. The variation is not clinal, but exhibits a sharp break between the two regions (Table 1, Map 19).

Juvenile plants of Cyathea arborea have, among other characters, a narrower lamina than adult leaves. A specimen from Dominica ( Lellinger 642, gh) has some plants with leaves 9 cm long, the lamina narrowly ovate-acuminate and pinnate-pinnatifid. Another from Puerto Rico ( Jones 10995, gh) has a stem about 1 cm long and erect, with roots to 25 cm long, the larger leaves are 30 cm long with the lamina 13 cm long, pinnate- pinnatisect and broadly ovate-acuminate. Specimens of leaves with the lamina about 30 cm long essentially have the adult characters of an ovate, bipinnate-pinnatifid lamina.




Greater Antilles and Lesser Antilles south to Grenada. Cyathea arborea typically grows in montane forests, in humid ravines, along water courses and on mountain slopes, from sea level to 1200 m, usually from 500-800 m. It frequently persists in cutover land, along forest borders and in forest clearings. It is successful as a pioneer species, often becoming established in disturbed habitats such as landslides, road cuts and spills and on aban- doned land. Stem to 10 m tall; leaves to 4 m long.

selected specimens. Cuba. Pinar del Rio: Rangel, upper Taco-Taco River, León 12721 (ny). Las Villas (Santa Clara): Loma Ventana, Trinidad Mountains, Howard 5221 (gh,mo,ny); Mina Carlotta, Sierra de San Juan, Senn 386 (f,gh,mo). Oriente: Rio Bayamito, Sierra Maestra, Maxon 3906 (gh,ny), 3909 (gh,ny,us); Rio Navas, Shafer 4379 (gh,mo,ny); Firmeza to Gran Piedra, Shafer 8957 (gh,mo,ny); Sierra de Nipe, Ekman 1805, 3368 (ny); Baracoa, Pollard , et al 45 ( f,gh,mo,ny ) . Jamaica. Valley of Rio Grande, 7 miles s. of Port Antonio, Gastony 38 (gh); Bloxburgh, Port Royal mountains, Maxon 8757 (gh,ny); between House Hill and Cuna Cuna Gap, Maxon 8886 (gh,ny); above Bowden Pen, to Bath via Cuna Cuna Pass, Crosby ir Anderson 1044 (f,gh,ny); Crooks Bottom, n. of Ipswich, Maxon ir Killip 1449 (f,gh, ny); Quashiba Mountain, 1 mile w. of Georges Plain, Wilson ir Webster 518 (a,us); Hermitage Dam, St. Andrew Parish, Proctor 3914 (mo); near Cambridge, Chrysler 1850 (mo). Haiti. Vicinity of Plaisance, Leonard 9365 (gh,ny); vicinity of St. Louis du Nord, Leonard ir Leonard 14256 (gh,ny); vicinity of Port de Paix, Leonard ir Leonard 15707 (mo); Bayeux, Morne Brigand, Eckman H 2954 (ny); Petit Borgne, Nash 470 (ny). Dominican Republic. Sonador, Prov. La Vega, Valeur 396 (f,mo); near Barahona, Türckheim 2716 (f,gh,mo,ny); Laguna, Samaná Peninsula, chiefly on the Pilón de Azúcar, Abbott 410 (gh,ny); Río Cataline, Las Cañitas, Abbott 2710 (gh), 2712 (ny); Liali, Abbott 2612 (gh) between Miches and El Seibo, Gastony , Jones ir N orris 702 (gh,ny). Puerto Rico. Sierra de Luquillo, Sintenis 1375 (a,mo,ny); Adjuntas, Sintenis 4242 (a,gh); El Verde Experiment Station, Little 13071 (a,f,ny); vicinity of Barran- quitas, Britton et al. 6622 (f,ny); Fajardo and vicinity, Rio Aribe, Britton ir Shafer 1693 (f,gh,mo,ny); 14 miles ne. of Mayaguez, Heller 4467 (f,gh,mo,ny); between Caguas and Cayey, Underwood ir Griggs 287 (ny); Sierra de Naguabo, Rio Icaco and adjacent hills, Shafer 3518 (f,gh,mo,ny). St. Thomas. Hopedale, Crown, Britton ir Marble 1422 (f,ny). Tortola. Sage Mountain, Fishlock 381 (ny), D'Arcy 48D (gh). Saba. Boldingh 2170 (gh). St. Kitts. Britton ir Cowell 165 , 286 , 331 (ny); Box 377 (f). Nevis. Proctor 19481 (a). Montserrat. Shafer 175 , 343 (f,ny); Howard 11905 (gh); Proctor 19169 (a).

Table 2. geographic variation in the development of the indusium of Cyathea arborea. (Based on three mature indusia from each of 140 collections, see Map 19.)

Deeply Deeply Cyathiform Cyathiform Meniscoid Meniscoid Discoid

Greater Antilles 88 collections 15 53 18 2 0 % of collections 17 60 20 3 0

Puerto Rico 25 collections 3 14 6 2 0 % of collections 12 56 24 8 0

Lesser Antilles 52 collections 0 10 15 20 7 % of collections 0 19 29 38 13

Tortola to Nevis 11 collections 0 2 2 6 1 % of collections 0 18 18 54 9



52 ROLLA TRYON

Guadeloupe. Matouba, Scamman 8142 (gh); Bois de Bains, Duss 4322 in 1896 (ny); Morne Papaye, Duss 4322 in 1892 (ny); south of La Citerne, Proctor 20135 (a). Dominica. Hodge 6 (gh,mo), 7 (gh); Hodge ù Hodge 2444, 2880 (gh); Lloyd 333, 653 (ny); Lellinger 354 , 642 (gh); Wilbur 8055 , 8288 (ny). Martinique. Duss 1604 (f,gh,mo,ny); Sieber, Syn. Fil. 194 ( MO,NY-fragm. ) ; Stehlé 3309 (f), 6081 (mo); Bailey ò- Bailey 279 (gh,ny). St. Lucia. Howard 11353 (ny); Proctor 17829 (gh). St. Vincent. Eggers 6610 (f,gh), 6921 (f), 6996a (f,gh); Morton 5193 (gh). Grenada. Eggers 6083 (f,gh,us); Broadway (f,gh); Sherring (ny).

18. Cyathea Alstonii Tryon, spec. nov. Figs. 79-80. Map 20.

Folia ad 75 cm longa; petiolus ca 25 cm longus squamis structura marginata discordanter valde bicoloribus vel fere concoloribus; lamina 1-pinnato-pinnatifìda; pinnae ad 12 cm longae sessiles, pagina inferiore glabra vel squamis paucis bullatis brunneis; sori plerumque subcostales ad furcam venarum; indusium cyatheoideum vel profunde cyatheoideum margine irregulari breviter dentato-ciliato. Holotypus: Pico Renjifo, Central Mountains, Sierra de la Macarena, Intend. Meta, Colombia, Philipson, Idrobo ò - Jaramillo 2157 (bm, photo gh). Paratypus: Macizo Renjifo, Cordillera La Maca- rena, Meta, Colombia. Idrobo ir Schultes 1130 (us).

An endemic of the Macarena, growing on sandstone rocks at 1300-1900 m.

19. Cyathea peladensis ( Hieron. ) Domin Figs. 81-84. Map 21.

Cyathea peladensis (Hieron.) Domin, Pterid. 263. 1929. Alsophila peladensis Hieron. Hedwigia 45: 233, t. 13. 1900. Holotype: Cerro Pelado

(upper Magdalena Valley), Colombia, Stubel 1259, b!, photo gh. All leaves of the holotype of Alsophila peladensis at Berlin are sterile, in

spite of the fact that Hieronymus described sori as ". . . soris in bifurca- tione venarum medio inter marginem et costam sitis usque 8-jugis." The "fertile" pinna illustrated in t. 13, f. 3a shows numerous small búllate scales, rather than sori. More recent collections of this species are fertile and are very similar in other characters to the sterile Stübel collection.

Colombia, 2300-2400 m. Stem to 3 m tall, leaves to 1 m long. additional specimens. Colombia. Huila-Cauca boundary: Head of Río Villa-lobos,

southwest of Pitalito, Cordillara Central, Fosberg 19950 (us). Huila: Comisaría del Caquetá, Cuatrecasas 8507 (f). Putumayo: Cerro Portachuelo, Sibundoy to Pepino, Soejarto 1569 ( gh ) .

Group of Cyathea pallescens KEY TO SPECIES 20-22

a. Fertile veins regularly forked, rarely a few simple b. b. Large scales of the petiole scurf flattish, elongate to sometimes ovate; Mexico to Panama, British Guiana to Colombia and south to Peru 20. C. diver gens.

b. Large scales of the petiole scurf mostly ( or many of them ) crested, expanded into two or more planes above the base and usually dissected, at least apically, or rarely larger scales absent and only squamellae present; Colombia to Bolivia 21. C. pallescens.

a. Fertile veins all simple; Venezuela 22. C. simplex.




The single collection of Cijathea simplex is unique among species with large ultimate segments in its simple fertile veins. Cyathea diver gens and C. pallescens are both highly variable species. Within the range of C. pallescens , the two can nearly always be separated, in addition to the petiole scurf characters mentioned in the key, by the long-stalked pinnules of C. diver gens and the sessile to short-stalked pinnules of C. pallescens. Only rarely does C. pallescens have some of the pinnules long-stalked.

The three species are probably a natural group. A species similar to Cyathea pallescens was probably ancestral to the four local endemics of the next group, and perhaps to C. fulva of the following group.

Map 22. Geography of variation in pinnule characters of Cyathea divergens: Left series, top to bottom: Hernandez ò- Sharp X472 (us), Johnson 966 (gh), Stone ir Stone 2700 (gh), Johnson 967 (gh), Pennell 9311 (ny), Cuatrecasas 13919 (gh), Ewan 16031 (gh), Harling et al. 10178 (gh). Right series, top to bottom: Tonduz 11789 (ny), Ulloa 102 (gh), Wilbur et al. 11077 (ny), Steyermark et al. 105688 (gh), Steyermark & Wurdack 717 (ven), Steyermark 93918 (us), Poeppig (p).



54 ROLLA TRYON

20. Cyathea divergens Kze. Map 22.

Cyathea divergens is a highly variable species, for example, the pinnules may be glabrous beneath or there may be indûment of trichomes, flattish scales, or búllate scales. The size of the pinnae and pinnules varies, and also their shape and the length of their stalks. The variation is highest in Colombia and Venezuela. A selection of the variation in pinnule characters is presented in Map 22, where a weak correlation with geography may be observed.

It is not possible to recognize two species, but the Mexican and Guate- malan element, var. Tuerckheimii, is sufficiently distinctive to merit infraspecific rank. The geography reinforces the characters of the variety and there is a relatively strong break in the variation between Mexico- Guatemala and Costa Rica-Panama. Table 3 provides the principal differences between var. divergens and var. Tuerckheimii .

Mexico and Guatemala, Costa Rica and Panama, British Guiana to Colombia and south to Peru.

20a. Cyathea divergens var. divergens Figs. 85-87. Map 23.

Cyathea divergens Kze. Linnaea 9: 100. 1834. Holotype: Pampayaco, Peru, Jul. 1829, Poeppig (Diar. J 1 63), lz, destroyed. Authentic specimens: Poeppig 219 (Diar. 1152), b! p! photo gh, fragm. ex b,ny!, ex k,ny!

Cyathea equestris Kze. Linnaea 9: 100. 1834. Holotype: Cerro de Cristobal, Pampa- yaco, Peru, Jul. 1829. Poeppig , lz, destroyed (Kze. Frarnkr. 1: t. 76), fragm. k! det. Kze., photo gh. In synonomy of Cyathea divergens by Mett. Ann. Sci. Nat. V, 2: 265. 1864.

Cyathea globularis Presi, Epim. Bot. 30. 1849, ex char. Holotype: Nova Granada, Linden.

Cyathea equestris var. boconensis Karst. Linnaea 28: 456, 1856, ex char, and Karst. Fl. Columb. 2: t. 185, f. 7. Holotype: Páramo de Bocono, Sierra de Merida, (Vene- zuela), Karsten.

Cyathea calva Karst. Fl. Columb. 2: 175, t. 192. 1869. Holotype: Sierra de Merida, Escuque, 1000 m, Venezuela, Karsten. Type collection: Escuque, 1000 m, Karsten 127 , b! fragm. ex b,ny!, ex Rosenst. us!

Cyathea petiolulata Karst. Fl. Columb. 2: 163, t. 185. 1869. Holotype: Merida, Venezuela, 2000-2500 m, Karsten. Authentic specimen: Merida, Karsten 135, b!

Cyathea petiolulata var. boconensis (Karst.) Karst. Fl. Columb. 2: 164. 1869, based on C. equestris var. boconensis , not Cyathea boconensis Karst.

Cyathea firma Kuhn, Linnaea 36: 163. 1869. Holotype: Merida, Venezuela, Funck ir Schlim 1228. Fragm. us!

Alsophila subaspera Christ, Pittier, Prim. Fl. Costaric. 3(1) ( Filices 2nd Mém. ) : 43. 1901. Syntypes: forêts du Copey, Costa Rica, Tonduz 11787 y a! gh! ny! p! us!, 11802 p! fragm. ex b,ny!, ex br,ny!, 12183 , ny!

Cyathea petiolulata var. pastoensis Hieron. Engl. Bot. Jahrb. 34: 437. 1904, ex char. Holotype: prope Altaquer et San Pablo, Cordillera de Pasto, Colombia, Leh- mann 81.

Cyathea pelliculosa Christ, Bull. Herb. Boiss. II, 4: 946. 1904. Syntypes: Costa Rica, Wercklé 45 , p!, Wercklé 50.

Cyathea divergens var. minor Rosenst. Fedde, Rep. Spec. Nov. 22: 2. 1925. Lecto- type: La Palma, Costa Rica, Brade ir Brade 108. Isolectotype: ny! (Brade ir Brade 853 is excluded as a type, it is Cyathea gracilis.)

Cyathea subaspera (Christ) Domin, Pterid. 263. 1929. Cyathea calva var. firma (Kuhn) Domin, Acta Bot. Bohem. 9: 102. 1930.




Cyathea divergens var. hirta Losch, Mittl. Bot. Staatssaml. München 1: 20. 1950, ex char. Holotype: Chirripó Grande, Costa Rica, Kupper 1265 , m.

A collection from the Chimantá Massif in the Guayana region of Vene- zuela, Steyermark ir Wurdack 717, has very coriaceous, narrow leaves, the pinnae ca 35-40 cm long, rather few long-stalked pinnules and an acute lamina apex. This may represent a local Tepui endemic but the evidence is insufficient to recognize it as a separate variety. Another collection from Auyan-tepuí, Guayana, ( Steyermark 93918) is rather intermediate between the above collection and typical var. divergens in Venezuela. A specimen from Colombia (Killip 11392) has a more or less cyatheoid indusium which is very thin, breaking into segments at maturity.

Differences from the superficially similar Cyathea corallifera are mentioned under that species.

Costa Rica and Panama, British Guiana west to Colombia and south to Peru. In humid mountain forests, often cloud forests and often persisting in cutover land. In Central America at elevations of 900-2600 m, usually at 1400-1800 m; in South America from 1300-2500 m, usually at 1600-2200 m. Stem to 15 m tall, the leaves to 6 m long.

selected specimens. Costa Rica. Heredia: Isla Bonita, Vallé du Sarapiqui, Pittier 14160 (mo,ny); Cinchona, upper Sarapiqui valley, Scamman 7862 (gh). Guanacaste: northern (Salazar) slopes of Volcán Orosi, White ir Lucansky 1968-74 (gh). San Jose: vicinity of El General, Skutch 2964 (gh,mo,ny); 25 km. n. of San Isidro del General, toward Cerro de la Muerte, Gastony ó- Gastony 755 , 759 (gh). Cartago: El Cedrai near Naranjo river, Nisman 124 (gh); Pacayas, foot of Volcán Turrialba, Scamman 7005 (gh). Puntarenas: Cantón Golfito, Nisman 138 , 140 (gh); Monte- verde, Nisman 156 (gh). Panama. Chiriquí: Casita Alta, Volcán de Chiriqui, Woodson , Allen ò- Seibert 922 (gh,mo,ny); Cerro Punta, Alien 3529 (mo); above El Boquete, Maxon 5691 (gh,ny); Río Piedra Candela, road to Las Mellisas, Mc Alpin 2192 (gh); Cerro Horqueta northwest of Boquete, Burch 548 (gh,ny). British Guiana (Guyana). Mount Roraima, Tate 428 (ny), 479 (ny). Venezuela. Bolivar: Ptari- tepui, Steyermark 59588 , 59811 (gh,mo,ny,us); Cerro Venamo, Steyermark , et al. 92701 (gh,ven); Auyan-tepuí, Steyermark 93918 (gh,ny,ven); Meseta de Jauá, Steyermark 98097 (gh); Sarvén-tepuí, Wurdack 34115 (gh,ny,us); Churi-tepui (Muru-tepui), Wurdack 34285 (gh,ny,us); Chimantá Massif, headwaters of Rio Tirica, Steyermark 6- Wurdack 717 ( gh,ny,us,ven ) . Amazonas: Sierra Parima, Steyer- mark 105973 (gh). Lara: 10 km. ne. de El Blanquito, Dist. Jiménez, Steyermark et al. 103580 (gh,ny). Merida: Loma de San Jacinto, Bernardi 1868 (ny). Colombia. Magdalena: Cincinati region, Espina ò- Giacometto A134 (g); Sierra Nevada de Santa Marta, H. H. Smith 2225 ( f,gh,no,ny,us ) . Bolivar: Antizales, Pennell 4455 ( f,gh,mo,ny ) . Norte de Santander: Loso, north of Toledo, Killip ù Smith 20407 (gh,ny). Antioquia: San Antonio del Prado, Daniel 3017 (gh). Caldas: above Salento, Pennell 9311 (gh,ny) south of Salento, Hazen 9693 (gh,ny,us). Cundin- amarca: Fusagasuga, Lindig 282 (gh); Bogotá, Lindig 309 (ny); 20 km. south of Bogotá, Barrington 464 (gh). Valle (Valle del Cauca) : La Cumbre, Killip 5684 (gh), 11392 (gh,ny,us); Monte Frio, Yanaconas, Killip ò- Garcia 33711 (f,gh,ny); El Cairo, entre Darién y Mediacanoa, Cuatrecasas 13919 (f,gh). Cauca: km. 19, Cali to Buenaventura, Barrington 500 (gh). Huila: 30 km. east-southeast of Baraya, Little 8846 (us); Hacienda Balsillitas, Meta to El Cedrai, Little 8040 , 8044 (gh,us); Rio Sauza, sw. of Alajandria, Little 8535 (gh,us). Nariño: 2 km. above San Juan, Ewan 16031 (us). Ecuador. Tungurahua: Rio Verde, Harling et al. 10178 (gb,gh). Peru. Huánuco: Cuslii, Macbride 4819 (f); Pampayacu, Kanehira 120 (gh). Cuzco: Sahuayacu, Chaupiorcco, Bües 840 (us).



56 ROLLA TRYON

Table 3. distinguishing characters of Cyathea diver gens var. diver gens and var. Tuerckheimii.

var. divergens var. Tuerckheimii Pinna stalk Usually long-stalked (the stalk Usually ¿hort-stalked (the stalk

longer than the width of the shorter than the width of the basal pinnule ) , especially the basal pinnule ) . basal pinnae and those toward the base.

Pinnule stalk Usually long-stalked (the stalk Usually short-stalked (the stalk longer than the width of the shorter than the width of the basal segment of the pinnule), basal segment), or all except often twice as long, especially a few basal pinnules so, or pin- on basal pinnules. nules sessile.

Pinnule shape Often long-triangular, or taper- Usually tapering to the apex ing to the apex from just above from about the middle. the base.

Apex of lamina Long acuminate. Acute to acuminate. Orientation of Drooping from beyond the mid- No data. the lamina die. Distribution Costa Rica and Panama, Brit- Mexico and Guatemala,

ish Guiana west to Colombia and south to Peru.

20b. Cyathea divergens var. Tuerckheimii (Maxon) Tryon, comb. nov. Figs. 88-90. Map 24.

Cyathea Tuerckheimii Maxon, Contrib. U.S. Nat. Herb. 13: 4. 1909. Holotype: near Cobán, Alta Verapaz, Guatemala, Feb. and Nov. 1907, Türckheim II 1645, us! Isotypes: f! gh! mo! ny! Paratypes: between Cobán and Tactic, Guatemala, Dec. 1907, Türckheim II 2031, us!; near Cobán, Guatemala, April, 1887, Türckheim (J. D. Smith no. 1238), us! Isoparatypes: Türckheim II 2031, gh! mo! ny!, Smith 1238, gh! ny!

Cyathea Sartorii Salomon, Nomencl. Gefässkrypt. 144. 1883, nom. nud. Mirador, Mexico, Sartorius, fragm. ex b,ny! bears this name.

The more abrupt, rather than elongate, lamina apex of Cyathea divergens var. Tuerckheimii suggests that the leaf probably does not have the pronounced drooping habit of var. divergens . However, field observations are needed to confirm this character. S kutch 1162, Guatemala, is a variation with a cyatheoid indusium.

Veracruz and southern Mexico to Guatemala. Predominantly in cloud forests, especially those of Quercus and Liquidambar at elevations of 750- 3100 m, usually at 1500-2500 m. Stem to 10 m tall, the leaves to 3 m long.

selected specimens. México. Veracruz : Cerro de San Cristobal, H. Xolocotzi ò- Sharp X-1163, 1164 (us); Cordoba, Farlow (gh); Orizaba, Pringle 6088 (gh); Mirador, Purpus 15312a (f). Chiapas: Laguna Ocotal Grande, Municip. de Ocosingo, Dressier 1688 (gh,ny,us); Sierra de Soconusco, H. Xolocotzi ir Sharp X-472 (us). Guatemala. Huehuetenango: San Juan Atitlán, Skut eh 1162 (gh); Cerro Negro, 2 miles e. of Las Palmas, Steyermark 51677 (f,gh,us); between San Mateo Ixtatán and Nuca, Steyermark 49787 (f,gh,us). San Marcos: southeast portion of Volcán Tacaná, Steyermark 36394 (f,us). El Quiche: Nebaj, Skutch 1698 (gh). Chimaltenango: Chichavac, Skutch 743 (us). Alta Verapaz: Samac, lohnson 960, 961 (f,gh,ny);




Chihob, Johnson 966 (gh,mo). Baja Verapaz: 10 km. north of Pantin, Hellwig ir Whitaker 1433 (gh). Zacapa: Sierra de las Minas, Steyermark 29801 (f), 30057 (f,us). Jalapa: Volcán Jumay, Steyermark 32466 (f,us).

21. Cyathea pallescens ( Sod. ) Domin Figs. 91-92. Map 25.

Cyathea pallescens (Sod.) Domin, Pterid. 263. 1929. Chosen over Cyathea cystolepis Sod. because of the more certain typification.

Alsophila pallescens Sod. Ree. Crypt. Vase. Prov. Quit. 20. 1883. Holotype: bosques de Nanegal, Ecuador, Sodiro. Type collection: p! photo gh.

Cyathea cystolepis Sod. Ree. Crypt. Vase. Prov. Quit. 15. 1883. Syntypes: Crecen en la pendiente occidental del Atacazo y del Corzón, 1600, 2000 m, Ecuador, Sodiro. Authentic specimen: in sylv. apud Niebly, Ecuador, 1/1883, Sodiro , p! fragm. ex Rosenst. us!

Hemitelia cystolepis (Sod.) Baker, Ann. Bot. 5: 187. 1891. Cyathea Borjae Sod. Crypt. Vase. Quit. 504. 1893, ex char. Holotype: en la par-

roguia de Santo Domingo, Ecuador, Sodiro. Hemitelia suhcaesia Sod. Crypt. Vase. Quit. 522. 1893. Holotype: bosques subandi-

nos de la cordillera occidental hasta 2,800 m, Ecuador, Sodiro. Specimens seen: Niebly, Sodiro , ny!; Canzacoto, 5/1882, Sodiro , p! photo gh; Ecuador, 4/1874, Sodiro , us!

Cyathea asperata Sod. Sert. Fl. Ecuad. 2: 9. 1908. Holotype: Pichincha, Ecuador, 3/1906, Sodiro. Type collection: ny! us!

Cyathea asperata var. brevipes Sod. Sert. Fl. Ecuad. 2: 10. 1908, ex char. No specimen cited.

Cyathea brachypoda Sod. Sert. Fl. Ecuad. 2 : 8. 1908. Holotype: Sylv. subandin. vulc. Atacazo, Ecuador, Sodiro. Specimens seen: Atacazo, 7/1906, Sodiro , gh! mo!; Atacazo, 7/1907, Sodiro , ny! us!

Cyathea muricatula Sod. Sert. Fl. Ecuad. 2: 10. 1908. Holotype: Corazón, Ecuador, 12/1907, Sodiro. Type collection: mo! p! us!

Cyathea nitens Sod. Sert. Fl. Ecuad. 2: 3. 1908. Holotype: Silv. sub-andin. occid. vulc. Corazón, Ecuador, Sodiro. Type collection: Corazón, 12/1907, Sodiro , mo! p! us!

Cyathea ochroleuca Sod. Sert. Fl. Ecuad. 2: 11. 1908. Holotype: Silvis subandin. occid. vulc. Atacazo, Ecuador, 7/1907, Sodiro. Type collection: Atacazo, Julio 8/1907, Sodiro , p! photo gh; Atacazo, 7/1907, Sodiro , ny! us!

Cyathea subinermis Sod. Sert. Fl. Ecuad. 2: 10. 1908. Holotype: Silvis subandinis vulcani Atacazo, Ecuador, 7/1907, Sodiro. Specimens seen: Pichincha et Atacazo, 1906-1907, Sodiro , gh! mo! p! photo gh.

Cyathea tungurahuae Sod. Sert. Fl. Ecuad. 2: 12. 1908. Holotype: Silvis subandin. vulc. Tungurahuae, Ecuador, Sodiro . Type collection: Tungurahua, 8/1901, Sodiro, p! photo gh,us!

Cyathea subcaesia ( Sod. ) Domin, Pterid. 264. 1929.

The accurate typification of many of Sodiros names is difficult because (a) he did not use collection numbers; (b) he often did not cite collection data fully, or he combined two or more collections in one citation; and (c) there is no catalogue of the Sodiro herbarium that remains in Ecuador. I believe that the most complete set of Sodiro's ferns is at Paris, but because of doubt I often hesitate to designate them as holotypes or lecto- types. These do not include, of course, the new species described by Baker from some 200 specimens sent earlier by Sodiro to Kew.

Cyathea pallescens is a highly variable species in several of its characters. The petiole scales are usually not long persistent, although they are in some specimens from Colombia ( Madison 861). The petiole scales



58 ROLLA TRYON

Figs. 81-90. 81-84, Cyathea peladensis, Colombia, Soejarto 1569 (gh): 81, Apical portion of stem, X Va', 82, Pinnae, X V2; 83, Portion of fertile pinna, X 2; 84, Receptacle and indusium, enlarged. 85-90, C. diver gens: 85-87, var. divergem: 85, Apical portion of stem, Venezuela, Steyer- mark ir Wurdack 717 (ny), X V2; 86, Pinnules, Costa Rica, Scamman 5884 (gh), X xk; 87, Portion of fertile pinnule, source as in 86, X 2. 88-90, var. Tuerckheimii: 88, Pinnules with short stalks, Guatemala, Skutch 1162 (gh), X V2; 89, Pinnule with longer stalk, Guatemala, Johnson 961 (gh), X V2 ; 90, Portion of fertile pinnule, source as in 88, X 2.




Fios. 91-99. 91-92, Cyathea pallescens, Ecuador, Mexia 7600: 91, Pinnules, (gh), X xk' 92, Portion of fertile pinnule, (us), X 2. 93-94, C. simplex, Venezuela, Maguire 35194 (gh): 93, Pin- nules, X 94, Portion of fertile pinnule, X 2%. 95-96, C. corallifera , Ecuador, 1874, Sodiro (mo): 95, Pinnules, X 96, Portion of fertile pinnule, X 1%. 97-99, C. boliviana, Bolivia, Steinbach 9512 (gh): 97, Pinnules, X Vz' 98, Portion of fertile pinnule, X 2%; 99, Scale from costa, enlarged.



60 ROLLA TRYON

are usually strongly bicolorous, but in some collections they may be whitish, tinged with brown or with brown streaks as in Dudley 11149 , from Peru, or they may be more irregularly tinged with brown. The petiole scurf may consist only of the characteristic crested squamellae, or it may be of appressed and strongly flattened, more or less entire to crested, squamellae at the base of the petiole. The pinnules are sometimes finely pubescent above. They may be essentially glabrous beneath or they may have one or more of the following types of indûment, which may be sparse to abundant: narrow, elongate, tìattish, whitish scales, whitish to brown búllate scales, with a short or long tip, large trichomes, and

strongly dissected small, whitish scales. Spruce 5367 from Ecuador, has at least some of the indusia urceolate, or partially open on one side.

Colombia to Bolivia. Usually in cloud forests, but sometimes above in

paramillo woods or thickets, and sometimes below in montane forest, at elevations of 1200-3500 m, most often at 2500-3000 m. Stem to 10 m tall, leaves to 4.5 m long.

selected specimens. Colombia. Cundinamarca: below Alto de Cuchuco, 7 km southwest of Sibate, Tryon ò- Try on 6113 (gh,ny); Manzanos, Cordillera de Bogotá, Lindig 289 (gh,us). Tolima: 30 km west of Cajamarca, Madison 861 (gh). Cauca: region de Moscopán, Cuatrecasas 23486 (f). Huila: Commis, del Caqueta, Cuatrecasas 8418 , 8438 (f); 20 km southeast of Garzón, Little 9391 (us). Nariño: 3 km north of Victoria, Rio Chingual drainage, Ewan 16182 (gh,us). Ecuador. Carachi: Canton Tulcan, near Pun, Mexia 7600 (f,gh,mo,us). Pichincha: vicinity of Huigra, Rose ir Rose 22508 (gh,ny,us), 22605 (gh,ny); between Nono and Tandayápi (Tandayapa), Sparre 16753 (gh), Lockwood 800 (gh). Peru. Cuzco: Cordillera Vilcabamba, Pro v. Convencion, Dudley 10864, 10950B (gh), 11149 (gh,us); Camino Panticolla, Prov. Convencion, Vargas 19857 (gh). Bolivia. Cochabamba: Sailapata, Cardenas 3150 (gh); between Pojo and Comarapa, on the Cochabamba-Santa Cruz road, Ugent ir U gent 5111 (gh,us).

22. Cyathea simplex Tryon, spec. nov. Figs. 93-94. Map 26.

Folia ca 1.5 m longa; petiolus ca 50 cm longus squamis structura marginata discordanter valde bicoloribus marginibus ciliatis; lamina bipinnato-pinnatifida; pinnulae pagina inferiore fere glabra; sori inframediales in venis simplicibus; indusium sphaeropteroideum. Holotypus: Cerro Yutaje, Serranía Yutaje, Rio Manapiare, Terr. Amazonas, Venezuela, Maguire ò- Maguire 35194, ny. Isotype: gh,us.

A very distinctive species, with simple fertile veins in relative large ultimate segments.

Venezuela, 1250 m. Group of Cyathea straminea

KEY TO SPECIES 23-26 a. Flattish scales on the costa, and sometimes on the costules, beneath ca 1-2 mm long, and some with dark, opaque peripheral or apical processes (Fig. 99) b. b. Squamellae of the petiole scurf nearly entire to slightly dissected c. c. Larger pinnules 3-5 cm broad, short-stalked, whitish scales and triohomes on the segments beneath; Ecuador 23. C. corallifera.

c. Larger pinnules 1-2 cm broad, sessile to subsessile; brownish scales and few trichomes on the segments beneath; Colombia 25. C. straminea.

b. Squamellae of the petiole scurf strongly dissected into trichome-like arms, larger pinnules 1-2 cm broad, sessile to subsessile, Bolivia 24. C. boliviana.




a. Flattish scales on the costa, and sometimes on the costules, beneath up to 3.5 mm long, with the marginal cilia the same color as the brown central portion of the body; Peru 26. C. Ruiziana.

A group of four apparently quite distinctive species, although further collections of them will certainly demonstrate variation in some of the characters. The characters of the petiole scurf are evidently important; they are presented in Table 4 to provide additional information to that given in the key. The species represent a radiation from an ancestral type similar to Cyathea pallescens.

23. Cyathea corallìfera Sod. Figs. 95-96. Map 27.

Cyathea corallifera Sod. Ree. Crypt. Vase. Prov. Quit. 11. 1883. Syntypes: Crece en los declives del Corazón y del Atacazo ( Miligallí-S. Florencio), 1500-2000 m, Ecua- dor, Sodiro. Authentic specimens: Corazón, 1882, Sodiro, p! photo gh; Atacazo, prop. Canzacoto, 5/1882, Sodiro , ny!

Cyathea aspidioides Sod. Ree. Crypt. Vase. Prov. Quit. 14. 1883, not (Bl. ) Moritz, Syst. Verz. Java 108. 1845. Syntypes: Crece en los bosques del Corazón y del Atacazo, 1800-2000 m, Ecuador, Sodiro. Authentic specimens: Corazón, prop. Miligally, 10/ 1896, Sodiro, ny!; Corazón, 10/1891, Sodiro, ny!; 1874, Sodiro, mo!

Cyathea corallifera var. alsophilacea Sod. Ree. Crypt. Vase. Prov. Quit. 12. 1883, ex char. No specimen cited.

Cyathea corallifera var. firma Sod. Ree. Crypt. Vase. Prov. Quit. 12. 1883. No specimen cited. Specimen seen: Ecuador, Sodiro, fragm. ex k,ny!

Cyathea corallifera var. ortholoba Sod. Ree. Crypt. Vase. Prov. Quit. 12. 1883. No specimen cited. Authentic specimen: Corazón, 1800 m, 5/1882, Ecuador, Sodiro, ny!

Cyathea aspidiiformis Domin, Acta Bot. Bohem. 9: 94. 1930, nom. nov. for Cyathea aspidioides Sod., not ( Bl. ) Moritz.

Cyathea corallifera is similar to some forms of C. diver gens in its large pinnae, pinnules and ultimate segments. It differs, in addition to the characters of pinnule indûment provided in the key, in having the large pinnules sessile to short-stalked. Large pinnules, when sometimes found in C. diver gens, are always long-stalked.

Ecuador, montane forests, 1400-2000 m. Stem to 5 m tall, leaves to 5 m

long- additional specimens. Ecuador. 1897, Sodiro (a); Corazón, Sodiro (ny); valley of

"Tallatanga" (Pallatanga), Rimbach 67 (us).

24. Cyathea boliviana Tryon, spec. nov. Figs. 97-99. Map 28.

Petíolus squamis structura marginata discordanter valde bicoloribus, furfure squamulis bicoloribus valde dissectis et squamis subplanis albidis elongatis setis peripheralibus fuscatis opacis; lamina bipinnato-pinnatisecta; pinnae ad 50 cm longae sessiles vel subsessiles; pinnulae longe acuminatae, pagina inferiore squamis paucis vel multis subplanis albidis vel brunneis setis peripheralibus fuscatis opacis; sori inframediales ad furcam venarum; indusium sphaeropteroideum. Holotype: Incachaca, Prov. Chapare, Dept. Cochabamba, Bolivia, Steinbach 9512, gh. Isotypes: F, gh, mo, ny, us. Paratype: Camarapa, Dept. Santa Cruz, Bolivia, Steinbach 8572, gh.

Cyathea boliviana grows in forests at 2000-2200 m. Stem up to 15 m tall.



62 ROLLA TRYON

25. Cyathea straminea Karst. FIGS. 100-101. MAP 29.

Cyathea straminea Karst. Linnaea 28: 457. 1856. Holotype: Cresit cum C. Quin- diuensi in decli vitate montes glacialis vulcaniei Tolima, Colombia, Karsten , (Fl. Columb. 2: t. 182, t. 183, f. III.) Isotype: Quindiu, Karsten , fragm. b!

The nearly concolorous, straw-colored petiole scales are the most distinctive character of Cyathea straminea .

Colombia, montane forest, ca 2000-2500 m.

additional specimens. Colombia. Caldas: above Salento, Pennell 9695 (gh,ny,us). Cauca: Km 55 on road from Torotó to Inza, Barrington 481 , 482 (gh). Putumayo: Laguna de la Cocha, Páramo de Santa Lucia, Cuatrecasas 11856 (f,us). Nariño: 5 km southwest of La Graja Botana, Pasto, Barrington 505 (gh).

26. Cyathea Ruiziana Kl. FIGS. 102-104. MAP 30.

Cyathea Ruiziana Kl. Linnaea 20: 439. 1847. Holotype: In Peruviae Andium nemoribus, Ruiz 72, b! The label also bears the name Panatahuas, formerly a province west of Huanúco. Isotype: us!

The holotype is a single pinna. Further characters are present in the Macbride collection cited below and especially notable among them are the very long large scales of the scurf (see Table 4), which densely in- vest the lower portion of the petiole.

Huanúco, central Peru. In montane forest at about 2000 m, stem to 9 m tall.

additional specimens. Peru. Huanúco: Huacachi, near Muña, Macbride 4135 (f,us).

Table 4. petiole scurf characters of species 23-26.

Large scales Squamellae 23. Cyathea corallifera Mostly entire or very nearly Slightly to moderately dis-

so, mostly bicolorous. sected, whitish to very light brown, mostly concolorous.

24. Cyathea boliviana Elongate, mostly whitish, Strongly dissected into tri- with dark, opaque peripheral chome-like arms, bicolorous. processes.

25. Cyathea straminea Mostly entire or nearly so, Slightly to strongly dissected, some bicolorous. mostly concolorous.

26. Cyathea Ruiziana Borne toward the base of the Elongate, nearly entire, or petiole: elongate, 3-4 mm somewhat dissected toward long, whitish, mostly con- the apex, mostly concolor- colorous. ous toward the base of the

petiole, mostly bicolorous beyond.




Maps 23-31. 23-24, Cyathea divergens: 23, var. divergens; 24, var. Tuerckheimii; 25, C. pallescens; 26, C. simplex; 27, C. corallifera; 28, C. boliviana; 29, C. straminea; 30, C. Ruiziana; 31, C. microphylla.



64 ROLLA TRYON

Figs. 100-108. 100-101, Cyathea straminea, Colombia, Barrington 482 (gh): 100, Pinnules, X 101, Portion of fertile pinnule, X 2%. 102-104, C. Ruiziana, Peru, Macbride 4135 (f): 102, Basal %rds of pinna toward apex of lamina, X %; 103, Pinnules, X Vz; 104, Portion of fertile pinnule, X 2. 105-106, C. microphyUa, Peru, Vargas 11858 (gh): 105, Pinnae, X 106. Sterile pinnules, X 2. 107-108, C. multisegmenta, Peru, Dudley 11326 (gh): 107, Pinnules, X 108, Portion of fertile pinnule, X 4%.




Group of Cyathea microphylla KEY TO SPECIES 27-28

a. Tertiary segments entire to basally 2-lobed; many small, dark brown, búllate scales on the pinnule-rachis beneath 27. C. microphylla.

a. Tertiary segments usually with 5 or more segments or lobes, some with fewer; light brown, búllate scales and trichomes on the pinnule-rachis beneath 28. C. multisegmenta. These two Peruvian species share unusual characters: the highly com-

plex lamina, with small ultimate segments, and the unusually broad petiole scales. From the available data, the two species differ widely in their leaf size. Cyathea microphylla has small leaves up to about 75 cm long, while C. multisegmenta has large leaves to at least 4 m long.

27. Cyathea microphylla Mett. FIGS. 105-106. MAP 31.

Cyathea microphylla Mett. Fil. Lechl. 1: 23, t. 3, f. 1-6. 1856. Holotype: St. Gavan, (San Gavan, Puno), Peru, Lechler 2160 , lz, destroyed.

Diacalpe microphylla (Mett.) Moore, Ind. Fil. c. 1857.

There is obvious confusion in the Lechler collection (or collections) of this species. Mettenius cites only Lechler 2160 from St. Gavan but he also cites the same number and locality under Cyathea Schanschin. There is a sheet of Lechler 2160 at Paris, correctly identified as Cyathea Schanschin (=C. Delgadii). Hooker (Second Century of Ferns, t. 99) also cites Lechler 2160 but from Tatanara (also in Puno, Peru). In the Mettenius Herbarium at Berlin, there is Cyathea microphylla ( Lechler 2569 ) from Tatanara, and the same collection at Paris. I have not been able to deter- mine if the different combinations of number and locality represent three collections, or if there was confusion in the labeling of a single collection. In any event, Cyathea microphylla is a very distinctive species and the original illustrations and the specimen at Berlin authentically represent the name.

Southern Peru, in thickets and forests at elevations of 2100-2800 m. Stem to 1.25 m tall, with a diameter of 1.5-2.5 cm.

additional specimens. Peru. Cuzco: Prov. Convencion, 73°22'W, 12°37'S, Dudley 10943 (gh). Puno: Valle Grande, Sandia, Vargas 11858 (gh); Tatanara, Lechler 2569 ( b,p, photo gh, fragms. f,us ) .

28. Cyathea multisegmenta Tryon, spec. nov. FIGS. 107-108. MAP 32.

Petiolus 1-2 m longus squamis structura marginata fere concoloribus pallide brunneis vel albidis, furfure squamulis albidis dissectis; lamina 1.25-3 m longa plerumque quadripinnata; pinnae sessiles vel subsessiles; pinnulae pagina inferiore squamis subplanis pallide brunneis et squamis bullatis albidis vel pallide brunneis necnon tri- chomatibus; sori subcostales in venis simplicibus vel ad furcam venarum; indusium sphaeropteroideum. Holotype: 73°37'W, 12°38'S, Prov. Convención, Dept. Cuzco, Peru, Dudley 11326 , na. Isotypes: gh,us,na.



66 ROLLA TRYON

The single collection was made in very dense cloud forest at about 1800 m, the stem was 2.5 m tall, and the leaves to 4 (or more ) m long.

Group of Cyathea fulva KEY TO SPECIES 29-34

a. Pinnules with few or no scales beneath and abundantly to moderately long- pubescent, or in Brazil sometimes with few long trichomes or with several scales; petiole scales usually light brown, the scurf of early caducous squamellae, or sometimes with larger scales also present; Costa Rica and Panama, British Guiana to Colombia south to Bolivia, Brazil, Ilha Trindade 33. C. Delgadii.

a. Pinnules with few or no scales beneath and with few or no long trichomes, or with several to many scales and then sometimes short-pubescent to moderately long- pubescent; petiole scales often brown to dark brown, the scurf of usually persistent squamellae and larger scales b. b. Squamellae of the petiole scurf strongly flattened and appressed, or erect and many crested (in South America rarely also ciliate-dissected ) ; continental North and South America c. c. Upper surface of the ultimate segments glabrous or with a few large trichomes, especially toward the apex; base of the petiole lacking reduced pinnae; Mexico to Panama, Venezuela and Colombia 29. C. fulva.

c. Upper surface of the ultimate segments pubescent, usually some of the trichomes long, slender, whitish and tortuous; base of the petiole usually with subaphlebioid pinnae; Costa Rica 32. C. suprastrigosa.

b. Squamellae of the petiole scurf erect, entire, ciliate or branched, rarely some crested; Antilles, Trinidad and Margarita d. d. Segments pubescent above, pinnules usually long-triangular, short-stalked, especially on the central pinnae; Jamaica and Dominican Republic 30. C. harrisii.

d. Segments with a few long trichomes above or none, pinnules with parallel sides, tapering to an acute to acuminate apex, sessile to subsessile or rarely a few short- stalked e. e. Indûment of rachis of (often caducous) scaly scurf beneath, rarely trichomes may also be present; costa with large, flat, usually brownish scales beneath; cos tules with usually brownish búllate scales beneath; Greater Antilles 31. C. furfuracea.

e. Indûment of rachis of trichomes beneath, these ( or their bases ) persistent, very rarely also with some scaly scurf; costa lacking large, flat scales beneath, or if present these usually wholly or partly whitish; costules with whitish búllate scales beneath; Lesser Antilles, Trinidad, Margarita 34. C. tenera.

This is a natural group of not wholly distinctive species. Their relation among the previous species is to Cyathea pallescens. Cyathea fulva is rather close to the next group, especially to C. caracasana var. Maxonii .

29. Cyathea fulva ( Mart. & Gal. ) Fée FIGS. 109-113. MAP 33.

Cyathea fulva (Mart. & Gal.) Fée, Mém. Fam. Foug. 9: 34. 1857. Alsophila fulva Mart. & Gal. Mém. Acad. Brüx. 15: 78, t. 23. 1842. Holotype: Talea,

Cordillère Orientale d'Oaxaca, México, Galeotti 6346 , br, photo gh, fragm. ex br, us! Isotype: p! photo gh.

Cyathea Schlechtendalii Kze. Bot. Zeit. 1845: 288, ex char. Herb. Schlechtendal. A specimen in Herb. Kuhn, b! bears this name.

Cyathea aurea Kl. Allg. Gartenzeit. 24: 105. 1856. Karsten. In Fl. Columb. 2: 149, t. 178, Karsten cites this species from Caracas, between Petaquire and Colonia Tovar, Venezuela. Fragments: Herb. Schlechtendal, Karsten 7, b! and Karsten 7 ex b,ny! are authentic.




Maps 32-37. 32, Cyathea multisegmenta; 33, C. fulva; 34, C. Harrisii; 35, C. furfuracea; 36, C. suprastrigosa; 37, C. Delgadii.



68 ROLLA TRYON

Cyathea aurea var. squamosa Karst. Linnaea 28: 459. 1857. Holotype: vicinity of Caracas, between Petaquire and Colonia Tovar, Venezuela, Karsten. Isotype: b!

Cyathea flaccida Mett. Ann. Sci. Nat. V, 2: 265. 1864, nom. nud. Nova Granada, Prov. Carabobo, Funck <b Schlim 413 was cited; l, photo ny,p! fragm. us!

Cyathea furfuracea Christ, Bull. Herb. Boiss. II, 4: 950. 1904, not Baker, 1874. Holotype: Costa Rica, 1903, Wercklé , Herb. Christ, p! fragm. ex Christ, us! Isotype: ny!

Cyathea onusta Christ, Bull. Herb. Boiss. II, 4: 950. 1904. Holotype: Costa Rica, Wercklé 41 , Herb. Christ, p!

Cyathea papyracea Christ, Bull. Herb. Boiss. II, 4: 946. 1904. Holotype: Costa Rica, Wercklé 52, Herb. Christ, p! (2ème feuille). Gastony, Contrib. Gray Herb. 203: 147. 1973, indicates that this sheet should represent the name; the 1ère feuille, lacking the number, is Nephelea mexicana.

Cyathea conspersa Christ, Bull. Herb. Boiss. II, 5: 260. 1905, nom. nov. for Cyathea furfuracea Christ, not Baker.

Cyathea Underwoodii Christ, Bull. Herb. Boiss. II, 6: 183. 1906. Syntypes: Navarro, Costa Rica, 1903, Wercklé , Herb. Christ, p! ny! fragm. ex Christ, ny!; 1901-1905, Wercklé , ny! us!; 1905, Wercklé , Herb. Christ, p!

Cyathea delicatula Maxon, Contrib. U.S. Nat. Herb. 13: 4. 1909. Holotype: between Tactic and Cobán, Alta Verapaz, Guatemala, Türckheim II 1629 , us! Isotype: gh!

Cyathea mollis Rosenst. Fedde, Rep. Spec. Nov. 22: 2. 1925, not Copel. 1917. Lectotype: La Palma, Costa Rica, Brade b- Brade 631 , Herb. S. Birger, s-pa! Isolecto- type: b! Lectoparatype: La Palma, Costa Rica, Brade ir Brade 630 , Herb. S. Birger, s-pa! Isolectoparatypes : b! ny! A sheet with the label indicating "630 and 631," us!

Cyathea molliuscula Domin, Acta Bot. Bohem. 9: 138. 1930, nom. nov. for Cyathea mollis Rosenst. not Copel.

Cyathea fulva usually has whitish to very light brown búllate scales on the under surface of the costae and cos tules. It often grows at lower elevations in Central America than other species with a sphaeropteroid indusium, and it is the only species among them that occurs in Nicaragua and Honduras.

In Costa Rica Cyathea fulva is sometimes close to specimens of C. caracasana var. Maxonii that have only some of the petiole scales partly reddish-brown to atropurpureous, the others being brown as in C. fulva. These two taxa often grow at different elevations, for example, about 25% of the 24 localities for C. fulva are below 2000 m, while over 50% of 18 localities for var. Maxonii are above that altitude.

Southern Mexico to Panama, Colombia and Venezuela. In Mexico and Central America in Liquidambar forest, Podocarpus forest and oak forest, humid forest to cloud forest, persisting in cleared land, at 800-2600 m, in Costa Rica usually below 2000 m. In Venezuela and Colombia in dense woods, cloud forests, on steep slopes and sandstone escarpments, 1300- 4200 m, the highest in Colombia in subparamo forest. Stem up to 10 m tall and 25 cm in diameter, leaves to 3 m long.

selected specimens. México. Hidalgo: near Zacualtipan, Martinez 1 (gh), 26 (mo,us). Puebla: between Zacapoaxtla and Xalacapa, Sharp 45960 (gh). Veracruz: Cuantlancillo (Orizaba), Copeland 15 (gh); 12 km south of Misantla, Lockwood 706 (gh). Guererro: Teotepec, Dist. Galeana, Hinton 14274 (gh,ny). Oaxaca: 35 km from Huatla, on road to Teotitlan, Madison 618 (gh). Chiapas: Colonia Ach'lun, Municip. Tenejapa, Ton 892 (ny). Guatemala. San Marcos: between San Rafael Pie de la Cuesta and Palo Verde, Williams , Molina ir Williams 25670 (f). Quezaltenango: El Pocito, south of San Martin Chile Verde, on road to Colomba, Standley 85050 ( f ) .




Alta Verapaz: Sacomum, Johnson 964 (f,gh,us). Zacapa: between El Picacho and Cerro de Monos, Steyermark 42791 (f,gh,us). Honduras. Morazán: Montaña del Tigre, Molina 10255 (f); Mt. Uyuca, Williams ir Molina 11093 (f). Nicaragua. Matagalpa: road to La Fuñadora, north of Sta. María de Ostuma, Williams, Molina ir Williams 24972 (f). Costa Rica. Alajuela: La Paña de Zarcero, Canton Alfaro Ruiz, A. Smith 1538 (ny); 19 km north of San Ramón, Wilbur ir Stone 10109 (gh,us). Heredia: Cinchona, upper Sarapaqui valley, Scamman 7584 (gh); Cariblanco, Nisman 113 , 160 (gh). San José: La Palma, Maxon 417 (ny); above La Hondura, Gastony ir Gastony 774 (gh). Cartago: 7 km south of Tapanti, Tryon ir Tryon 6988, 6989 , 7123 , 7124 (gh); 10 km south of Tapanti, Burger ir Stolze 5667 (gh), 5683 (gh,ny). Puntarenas: Monteverde, Palmer 76 (ny); San Vito, Mc Alpin 2259 (gh). Panamá. Chiriqui: vicinity El Boquete, Maxon 5046 (gh,ny); vicinity of Cerro Punta, Allen 1522 ( f,gh,mo,ny ) . Venezuela. Sucre: Cerro Turumuquire, Steyermark 62467 (gh, mo, ny, us, ven ) . Distrito Federal: El Junquito, Pittier 13946 ( f,ny,us,ven ) . Ar agua: Colonia Tovar, Fendler 52 (f,gh,mo). Carabobo: east of Los Tanques, south of Borburata, Steyermark ir Steyermark 95396 (ny,ven). Falcon: Sierra de San Luis, Steyermark 98914 (gh). Merida: Carbonerro, White ir Lucansky 1969-237 (ny,us); Cerro El Toro, Bernardi 1876 (ny). Bolivar: Amurí-tepuí, Chimantá Massif, Steyer- mark ir Wurdack 1328 ( gh,ny,us,ven ) . Amazonas: Cerro Parú, Cowan ir Wurdack 31390 ( gh, ny, us). Colombia. Antioquia: 10 km east of Sonsón, Scolnik, López ir Barkley 19An214 (mo). Santander: Pamplona to Toledo, Killip ir Smith 19962 (gh,ny,us). Cundinamarca: Tena, Laguna de Pedro Palo, Fernandez ir Mora 1451 (us). Cauca: El Tambo, Munchique, Agredo 431 (f,us). Tolima: above Anaime, on road south of Cajamarca, Barrington 474 (gh). Huila: inter La Resina et Andaluzia, Juzepczuk 6595 ( us ) .

30. Cyathea Harrisii Maxon FIGS. 114-115. MAP 34.

Cyathea Harrisii Maxon, No. Amer. Fl. 16: 81. 1909. Holotype: Blue Mountain Peak, Jamaica, Underwood 2502 , ny. Isotype: Maxon 1371 , "= Underwood 2502 ," us!

Cyathea Harrisii is characterized, among the Jamaican species of the

genus, by its usually long-triangular pinnules that are pubescent on the

upper surface of the segments. Cyathea suprastrigosa of Costa Rica is its closest relative, rather than species of the Greater Antilles.

Jamaica and Hispaniola. Blue Mountain Peak, Jamaica, in rain forest, 2100-2200 m; and Sierra de Ocoa, Dominican Republic, in mossy forest and wet pinelands, persisting after clearing, 2300-2500 m. Stem to 3.5 m tall, leaves to 2 m long.

additional specimens. Jamaica. Blue Mountain Peak, St. Thomas, Proctor 4378 (mo), Faull 12582 (mo), Chrysler 2049 (gh,mo), Maxon 9866 (gh,mo,ny), 9889 (gh,ny). Dominican Republic. 13 km from Valle Nuevo on road to San José de Ocoa, La Vega, Gastony, Jones ir N orris 720, 730 (gh,ny); Sierra de Ocoa, Azua, Ekman H 11678 (f,gh,ny).

31. Cyathea furfuracea Baker FIGS. 116-118. MAP 35.

Cyathea furfuracea Baker, Syn. Fil. 450. 1874. Lectotype: Portland Gap, Jamaica, July, 1843, Purdie, k! fragm. ny! Isolectotype : bm! photo gh. Lectoparatype : Jamaica, Bancroft, k! fragm. ny!

Cyathea monstrabila Jenm. Jour. Bot. 19 (n.s. 10): 275. 1881. Holotype: Portland Gap, Jamaica, Nock, Herb. Jenm., ny! fragm. us! (A monstrous form).



70 ROLLA TRYON

Cyathea asperula Maxon, Contrib. U.S. Nat. Herb. 17: 179. 1913. Holotype: near Constanza, Santo Domingo, Türckheim 3056 , us! Isotypes: f! gh! mo!

Cyathea Brittoniana Maxon, Jour. Wash. Acad. Sci. 14: 139. 1924. Holotype: Mt. Alegrillo, Puerto Rico, Britton , Stevens ir Hess 2620 , us! Isotype: f! Paratypes: Monte de la Prenda, Oriente, Cuba, Eggers 5211 , us; Pinal de Santa Ana, Oriente, Cuba, Eggers 5031 , us; Paradis, Barahona, Dominican Republic, Abbott 1590 , us; Maricao, Puerto Rico, Hioram 809 , us. Isoparatypes : Eggers 5031 , f! fragm. ex k,ny!; Eggers 5211 , gh! fragm. ex k,ny! (Trinidad: Pendler 80, and Britton , Hazen ir Mendelson 1351 are excluded as paratypes; they are Cyathea tenera).

There is considerable variation in the indûment on the under surface of the pinnules in Cyathea furfur acea. While this shows some correlation with geography, it is not sufficiently strong to suggest the recognition of varieties. Specimens from Cuba often have numerous long trichomes beneath and some have few scales. Jamaica specimens usually have several to numerous trichomes and many scales, while in those of Hispaniola there is usually a moderate investment of both trichomes and scales. The few collections from Puerto Rico include most of this range of variation.

Most of the collections cited by Maxon as Cyathea Brittoniana are the relatively pubescent and slightly scaly variation of C. furfuracea. They could be interpreted as intermediates with Cyathea tenera, which also has variations approaching C. furfuracea. However, considering the gap between the range of the two taxa, from Puerto Rico to Montserrate, it seems best to interpret this variation as local infraspecific variation within two closely related species.

Maxon s Cyathea Brittoniana is a source of confusion because he included in it Trinidad collections of Cyathea tenera , while including in C. tenera Greater Antillean collections which are C. furfuracea.

Cuba, Jamaica, Hispaniola and Puerto Rico. On steep slopes of montane forest, in wet ravines, Podocarpus cloud forest and elfin woodland, rarely 300 to 900-2200 m. Stem to 8 m tall, and 25 cm in diameter, leaves to 3 m long.

selected specimens. Cuba. Santa Clara: Buenos Aires, Trinidad Hills, Jack 7273 (f,gh,ny). Oriente: La Bayamesa, Sierra Maestra, Ekman 7070 (ny); Gran Piedra, Maxon 4070 (f,gh,ny); Pico Turquino, Leon 11154 (ny), Ekman 5430 (ny). Jamaica. Mt. Horeb trail, St. Andrew, Crosby , Hespenheide ir Anderson 308 (f,gh, mo, ny ) ; Morce's Gap, St. Andrew, Maxon ir Killip 660 (f,gh,ny); Sugar Loaf Peak, Portland, Proctor 4360 (mo); Hardwar Gap, Portland, Try on ir Try on 6982 , 6983 (gh); Corn Puss Gap, St. Thomas, Proctor 3996 (gh,mo,us); Blue Mountain Peak, St. Thomas, Maxon ir Killip 1146 (f,gh,ny). Haiti. Vicinity of Furcy, Leonard 4670 , 5355 ( gh,ny ) ; Morne Haut Piton, vicinity of Bassin Bleu, Leonard ir Leonard 15165 ( gh,mo,ny ) ; Morne Tranchant, Massif de la Selle, Ekman H3217 (ny). Dominican Republic. Vicinity of Paradis, Barahona, Abbott 1664 (gh); La Ciénega, La Vega, Gastony ir N orris 173 , 197 , 213 (gh); La Cumbre, Santo Domingo, Ekman H 11496 (ny); Monción, Lagunas de Cenobi, Monte Cristi, Ekman H12865 (f,gh); Sierra de Neiba, near Haitian border, Gastony , Jones ir N orris 431 (gh,ny), 469 (gh). Puerto Rico. Monte del Estado, Maricao Forest, Little 21702 (gh); Cerro de La Punta, km 18.5, road 143, Conant 596 , 603, 685 (gh).




Figs. 109-118. 109-113, Cyathea fulva: 109, Apical portion of stem, Venezuela, Steyermark & Steyermark 95396 (ven), X %} 110, Pinnules, source as in 109, X 111, Pinnules, Mexico, Martinez 1 (gh), X 112, Portion of fertile pinnule, source as in 109, X 1%; 113, Portion of fertile pinnule, source as in 111, X 2. 114-115, C. Harrisii, Jamaica, Maxon 9866 (gh): 114, Pinnules, X Vz' 115, Portion of fertile pinnule, X 2. 116-118, C. furfuracea : 116, Pinnules, Domini- can Republic, Ekman H12865 (gh), X 117, Pinnules, Haiti, Leonard ir Leonard 15165 (gh), X 118, Portion of fertile pinnule, Jamaica, Maxon & Killip 660 (gh), X 2.



72 ROLLA TRYON

32. Cyathea suprastrigosa (Christ) Maxon FIGS. 119-120. MAP 36.

Cyathea suprastrigosa (Christ) Maxon, No. Amer. Fl. 16: 83. 1909. Hemitelia suprastrigosa Christ, Pittier, Prim. Fl. Costaric. 3 (1) (Filices, 2nd Mém.):

44. 1901. Holotype: forêts de TAchiote, Volcán de Poas, Costa Rica, 2000 m, Tonduz 10701 , fragm. ex Christ, ny! Isotypes: Herb. Bonap. p! photo gh,us! fragm. ex us,ny!

Cyathea conspicua Christ, Bull. Herb. Boiss. II, 6: 178. 1906. Holotype: Volcán Turrialba, Costa Rica, 1905, Wercklé, p! Isotype: us!

Cyathea suprastrigosa is usually characterized, in addition to the characters mentioned in the key, by a strongly outward and downward bent petiole base and by auriculate basal inferior segments, especially on the larger pinnules. The distinctive delicate trichomes on the upper surface of the segments may be lacking in older leaves.

Costa Rica. Montane forest, cloud forest and subparamo woodland, on

slopes or in poorly drained sites, 2000 to usually 2400-2800 to 3000 m. Stem to 5 m tall and 10 cm in diameter, leaves to 1 m long.

selected specimens. Costa Rica. Alajuela: Volcán Poás, Nisman 100 , 102 (gh), Jiminez 1022 (gh,ny). San José-Cartago: near El Trinidad, Burger ir Stolze 5238 , 5964 (gh); 3 km from El Empalme, toward Cerro de la Muerte, Gastony ir Gastony 747 , 748 (gh). Cartago: vicinity of Millsville, Pan-American Highway, Holm ir litis 533 , 601 (a,f,mo), 605 (a,mo); 10 miles southeast of El Empalme, Wilbur ir Stone 9972 (gh).

33. Cyathea Delgadii Sternb. FIGS. 121-123. MAP 37.

Cyathea Delgadii Sternb. Flor, der Vorwelt 1: 47, t. B. 1820 (also Flor. Monde Prim. 4: 51, t. B. 1826). Holotype: Gancho do Generale Delgado in via ad Caldas Novas, Goyaz, Brazil, Pohl , prc, fragm. ex prc,gh! A fragm. ex l,us! is probably from the same collection.

Cyathea vestita Mart. Denkschr. Bot. Ges. Regensberg 2: 146. 1822. (Icon. Pl. Crypt. Bras. 75, t. 52). Holotype: uncertain without an examination of the original materials. Minas Geraes, Pohl 663 , w, photo and fragm. bm! may be authentic.

Cyathea oligocarpa Kze. Linnaea 9: 101. 1834. Holotype: Pampayaco, Peru, Poeppig (Diar .1101 ), LZ, destroyed. Isotypes: Poeppig 218 , b! mo! p!

Cyathea hirtula Mart. Icon. Pi. Crypt. Bras. 76, t. 53. 1834. Holotype: Uncertain without an examination of the original materials, Almada, Serra do Mar, Bahia, Brazil, Max. Neuwied, br photo and fragm. bm! is authentic.

Cyathea schanschin Mart. Icon. Pi. Crypt. Bras. 77, t. 54. 1834. Holotype: Uncertain without an examination of the original materials, São Paulo, Brazil, Martius , bm! photo gh, and Curibiba, São Paulo, Brazil, Martius , m, photo bm are probably authentic.

Cyathea denticulata Goldm. Nova Acta Acad. Caesar. Leopold. -Carol. 19, Suppl. 1: 466. 1843, ex char.

Cyathea abruptecaudata Fée, Crypt. Vase. Brésil 1: 183, t. 62, f. 2. 1869. Holotype: Rio de Janeiro, Brazil, Glaziou 2284 , Herb. Fée-Herb. Cosson, p! photo gh, fragm. ex Fée, ny! Isotype: us!

Cyathea Feei Fée, Crypt. Vase. Brésil 1: 179, t. 66, f. 2. 1869. Lectotype: Rio de Janeiro, Brazil, Glaziou 2286 , Herb. Fée-Herb. Cosson, p! photo gh, fragm. ex Fée, ny! Isolectoparatype : Brazil, Claussen 114, p!

Cyathea sphaerocarpa Fée, Crypt. Vase. Brésil. 1: 180, t. 53, f. 2. 1869. Holotype: Rio de Janeiro, Brazil, Glaziou 2283 , Herb. Fée-Herb. Cosson p! photo gh.

Cyathea pilosa Baker, Syn. Fil. 19. 1874. Holotype: Andes of east Peru, (Tarapoto),




Figs. 119-128. 119-120, Cyathea suprastrigosa , Costa Rica, Gastony ir Gastony 747 (gh) : 119, Pinnules, X 120, Portion of fertile pinnule, X 2. 121-123, C. Delgadii: 121, Pinnules, Brazil, Dusén 11781 (gh), X 122, Pinnules, Bolivia, Williams 1334 (gh), X Vzi 123, Portion of fertile pinnule, Costa Rica, Nisman 122 (gh), X 2. 124-126, C. tenera: 124, Pinnules, Guadeloupe, Proctor 20134 (a), X Vz; 125, Portion of fertile pinnule, source as in 124, X 2; 126, Pubescent rachis, Grenada, Proctor 16995 (a), enlarged. 127-128, C. dissoluta , Jamaica, Proctor 5819 (mo): 127, Pinnules, X %; 128, Portion of fertile pinnule, X 2.



74 ROLLA TRYON

Spruce 4729. k. Isotypes: gh! p! photo gh, fragm. (probably ex p) f! fragm. ex Rosenst. us! fragm. ex bm,us!

Cyathea Copelandii Kuhn & Luerrsen, Abhandl. Naturw. Ver. Bremen 7: 278. 1882. Holotype: Trinidad (Ilha da Trindade, Brazil), Ralph Copeland, fragm. Herb. Luerssen, p! photo gh.

Cyathea hypotricha Christ, Bull. Herb. Boiss. II, 4: 947. 1904. Holotype: Costa Rica, Wercklé , Herb. Christ, p! fragm. ny!

Cyathea schanschin var. oligocarpa (Kze. ) Hieron. Hedwigia 45: 230. 1906. Cyathea bahiensis Rosenst. Fedde, Rep. Spec. Nov. 20: 90. 1924. Holotype: São

Gómala, gebiet des Rio Femmeas, Bahia, Brazil, Luetzelburg 534 , m, photo bm. Cyathea micromera Rosenst. Fedde, Rept. Spec. Nov. 20: 90. 1924. Holotype:

Apertada-hora, Goyaz, Brazil, Luetzelburg 12841, m, photo bm. Cyathea trindadensis Brade, Archiv. Instit. Biol. Veg. Rio Janeiro 3: 1, t. 1, f. 1.

1936. Holotype: Ilha de Trindade, Brazil, Campos Porto 579, rb. Isotype: f!

Brazilian specimens have considerable variation in the lamina indûment but there is relatively little variation in other parts of the range. In Brazil the sterile leaves, especially, may have few trichomes on the under surface and there are also sometimes some scales beneath. This latter variation, for example, in Try on & Try on 6659 and in Sehnem 4213 , is a very close match to the material from Ihla Trindade, previously considered to be an endemic species.

While the characters employed in the key are all variable, they form a combination that serves to distinguish the species. In addition, the pinna- rachis is usually pubescent, a character otherwise usually seen only in Cyathea tenera.

Costa Rica and Panama, around the Amazon Basin in British Guiana to Colombia, south to Bolivia, to southeast Brazil and northeastward to Ceará, Brazil. In primary forest, cloud forest, persisting in partially cleared forest, (in central Brazil) in gallery forest, along streams, usually 500-1200 m, sometimes lower to 100 m, or in Costa Rica and Colombia to 2000 m, and in Peru to 2700 m. Stem to 10 m tall, and 15 cm in diameter, leaves to 3 m long.

selected specimens. Costa Rica. Alajuela: Carrizal toward Cariblanco, Gastony è- Gastony 790, 794 (gh); La Marina, Nisman 78 (gh); above Los Angeles, north of San Ramón, Stone 2752 (gh,us). Heredia: Cariblanco, Nisman 110, 115 (gh). Limón: La Trocha, Guápiles, Nisman 133, 135 (gh). Cartago: east of Tuis, on road to San Joaquin, White ò- Lucansky 1968-182, -184, -191 (gh). Puntarenas: Rincon, Osa Peninsula, Lent 459 (gh); between San Vito de Java and Villa Neilly, Nisman 144 (gh). Panamá. Codé: crater of El Valle de Antón, Wilbur, Weaver ò- Correa 11121 (gh,ny). Panamá: Cerro Campana, McDaniel 6888 (mo). British Guiana (Guyana). Makubere Savanna, Kanuku Mountains, For. Dept. Brit. Guian. WB401 (ny); Roraima Range, McConnell ö Queich 605 (ny); Mount Roraima, Im Thum 92 (us). Venezuela. Sucre: Cerro Espejo, Peninsula de Paria, Steyermark ò- Rabe 96085 (gh). Miranda: Quebrada Humboldt, Guayabitos, Vareschi 5819 (ven). Bolivar: Meseta de Jaua, Río Kanarakuni, Steyermark 98224 (gh). Colombia. Santander: Mesa de los Santos, Killip ir Smith 15338 (gh,ny,us). Cundinamarca: Fosca, Río Sáname, Garda-Barriga 17581 (ny). Vichada: Northwest of San Jose de Ocune, Haught 2813 (gh). Meta: north end of Cordillera de Macarena, Smith ir Idrobo 1539 ( gh, mo, ny, us). Nariño: Junin to Altaquer, Municip. Altaquer, Soejarto 1451 (gh). Peru. San Martin: Rioja, northwest of San Martin, Soukup 5223 (gh). Huánuco: Tingo Maria, Tryon à- Tryon 5219 (f,gh). Cuzco: Cordillera Vilcabamba, Dudley 10058, 10268, 11258 (gh). Puno: Lechler 2160 (p). Bolivia. Santa Cruz: Buena




Vista, Prov. Sara, Steinbach 5310 ( f,gh,mo,ny ) . La Paz: Polo-Polo bei Coroico, X, XI, 1912, Buchtien ( f,gh,mo,ny ) ; Apolo, Williams 1333 (us). Argentina. Corrientes: Estancia El Plata, Dept. Ituzaingó, Meyer 6278 (a,us). Paraguay. Tobatis, Hassler 4004 ( GH, NY ) ; Sierra de Amaambay, Hassler 10442 (mo,ny,us); Carapegua, Rojas 3010 (gh,p). Brazil. Roraima: Serra Tepequem, Prance , Forero , Pena ir Ramos 4499 (ny). Ceará: Serra do Araripe, Duarte 1346 (ny). Mato Grosso: Serra do Roncador, 86 km north of Xavantina, Irwin , Grear, Souza ir Santos 16305 (gh,ny). Goias: Riberon Grande, Municip. Jatai, Macedo 2161 , 2173 (us). Distrito Federal: Parque Municipal do Gama, 25 km south of Brasilia, Irwin , Souza ir Santos 10138 (ny). Minas Gerais: Itacolumi, Sehnem 4213 (a), Macedo 2825 (us); Viçosa, Mexia 4582 (f,gh,mo,ny). Rio de Janeiro: Monte Serrât, Mt. Itatiaya, Smith 1593 (f,gh,ny); Riberão Campo Belo, Mt. Itatiaya, Try on ir Try on 6658 , 6659 , 6661 , 6671 (gh). Guanabara: Foresta de Tijuca, Martins 262 (gh). São Paulo: Morro das Pedras, Iguapé, Brade 8220 (gh,ny); Monte Alegre do Sul, Kuhlman 1792 (gh); 27 km northwest of Moji-Mirim, Eiten , Eiten ir Sota 2127 (gh,ny,us). Paraná: Porto de Cima, Dusén 623a (gh,mo,us), 11781 (f,gh,mo); Valhinas, Dusén 10760 (f,gh,mo, ny). Santa Catarina: Blumenau, Haerchen (Ros. Fil. Austrobras. 50a) (f,mo). Ilha Trinidade (South Trinidad). Pico Desejado, Mello Filho 961 (Segadas-Viana 2058) (us); "Discovery," 13 Sept. 1901 (us).

34. Cyathea tenera ( Hook. ) Moore FIGS. 124-126. MAP 38.

Cyathea tenera (Hook.) Moore, Ind. Fil. 274. 1861. Alsophila tenera Hook. Sp. Fil. 1: 49. 1844. Holotype: St. Vincent, Caley , k! fragm.

ex k,ny! Isotype: bm! photo gh, fragm. ex bm,us!

Cyathea tenera is close to both C. furfuracea and to C. Delgadii. It is maintained as a species especially on the basis of its geography and the lack of a clinal relationship of intermediates. A few collections of C. tenera have large, brown, flattish scales beneath and some have búllate scales on the costules beneath.

Cyathea tenera appears to represent the initial element in a migration accompanied by evolution from C. Delgadii in South America. The migration evidently continued to the Greater Antilles with the further speciation which resulted in C. furfuracea .

Lesser Antilles, Montserrat to Grenada, Trinidad and Isla Margarita. In rain forest and montane woods and in thickets, 400-1000 m. Stem to 9 m

tall, leaves to 3 m long, usually less. selected specimens. Montserrat. Howard 11906 (gh). Guadeloupe. 1840, L'Her-

minier 62 (gh); Proctor 20134 (a). Dominica. Hodge ir Hodge 1973 (gh,ny); Lloyd 170 , 391 (ny). Martinique. Sieber Exsicc. 374 (mo,ny). St. Lucia. Proctor 17781 ( gh ) . St. Vincent. Eggers 6859 ( f,gh ) , 6868 ( f ) ; Morton 5492 ( ny ) , 6213 ( gh ) . Grenada. Howard 10638 (gh); Nov. 1904, Broadway (f,gh). Trinidad. F endler 80 (gh,no,ny); Broadway 5514 (f,gh,mo), 7408 (f,mo), 9968 (gh,ny). Venezuela. Isla Margarita: Johnston 143 ( f,gh,ny,us ) .

Group of Cyathea caracasana KEY TO SPECIES 35-36

a. Jamaica: pinnules with a few to several long trichomes and a few brown búllate scales beneath, sometimes also a few brown flattish scales on the costa, indusium firm, when over-mature the segments persistent 35. C. dissoluta.

a. South America, Costa Rica: pinnules usually with different indûment beneath, or none; Cuba, and Hispaniola: (var. caracasana) ultimate segments short, whitish pubescent beneath, and indusium delicate, evanescent 36. C. caracasana.



76 ROLLA TRYON

This group shows its closest relationship with Cyathea fulva. The complex and variable C. caracasana is the most common species of the Andes. Cyathea dissoluta is a rare and questionable endemic of Jamaica. While technically close to C. caracasana , it does not seem to fall within the variation of that species.

35. Cyathea dissoluta Jenm. FIGS. 127-128. MAP 39.

Cyathea dissoluta Jenm. Jour. Bot. 19 (n.s. 10) : 52. 1881. Holotype: woods near the Cinchona Plantation, Jamaica, 5000 ft., Jenman 1, k! Isotypes: Herb. Jenm. ny! fragm. and photo ex ny,us!

Cyathea dissoluta is a rare, variable species of Jamaica. It is quite possibly a hybrid, or a series of hybrids, but field study is required to provide a basis for a better evaluation of its status.

Jamaica. Montane rain forest, ca. 1200 m.

additional specimens. Jamaica. Jenman (ny); Campbell 7731 (f,ny); Hart (p); Morce's Gap, St. Andrew, Sherring 6 (us); near Cinchona, St. Andrew, Harris 7731 (us); above Murdock's Gap, Portland, Proctor 5819 (mo); Arntully Gap, St. Thomas, Sabonabière (gh).

36. Cyathea caracasana ( Kl. ) Domin

Cyathea caracasana is a complex and highly variable species. The principal variation is in the squamellae of the petiole scurf, which may be crested, strongly dissected or entire, in the length of the pinnule stalk, the shape of the pinnules, the abundance and kinds of trichomes and scales on the segments, costae and pinna-rachises beneath, and in the texture and corresponding persistence of the segments of the indusium. Considerable study has not resulted in the morphological-geographical correlations and group distinctions to be expected of either species or geographic varieties. However, groups can be formed that may have evolutionary validity and I have chosen to recognize them in a formal classification.

Greater Antilles, Costa Rica, Venezuela to Colombia and south to Bolivia.

KEY TO VARIETIES OF CYATHEA CARACASANA. a. Ultimate segments glabrous, or with 1 to few large trichomes or rarely pubescent above b. b. Pinnules sessile to short-stalked or rarely long-stalked and the pinnule broadly long-triangular c. c. Ultimate segments lacking trichomes beneath, or long-pubescent, or rarely short- pubescent; indusium firm to thin, when over-mature, the segments persistent, at least in part d. d. Pinnules usually tapering to the apex from beyond the middle, to very narrowly long-triangular, sessile to short-stalked; indûment beneath usually rather abundant, of various kinds of trichomes and scales; Venezuela and Colombia to Bolivia 36a. var. boliviensis.

d. Pinnules, especially toward the base of the central and basal pinnae, broadly long-triangular or nearly so, short-stalked to rarely long-stalked; indûment beneath sparse or absent; Venezuela, Colombia and Ecuador. . . 36b. var. meridensis.

c. Ultimate segments short-whitish pubescent beneath; indusium delicate, more or




less evanescent and often persisting, if at all, as an irregular basal disk or parts (segments dull beneath); Cuba, Jamaica, Hispaniola, Venezuela, Colombia and Ecuador 36c. var. caracasana.

b. Pinnules, especially basal ones of the central and basal pinnae, long-stalked, indû- ment beneath sparse or absent; very narrowly long-triangular to tapering to the apex from beyond the middle; Colombia and adjacent Venezuela (Zulia), Ecuador 36d. var. chimb or azensis.

a. Ultimate segments short-pubescent above, at least at the margin, sometimes sparingly so, usually short-pubescent beneath; Costa Rica 36e. var. Maxonii.

36a. Cyathea caracasana var. boliviensis ( Rosenst. ) Tryon, comb. nov. FIGS. 129-130. MAP 40.

Cyathea mexicana var. boliviensis Rosenst. Fedde, Rep. Spec. Nov. 25:56. 1928. Holotype: Hacienda Simaco supra Tipuani, Bolivia, Buchtien 5140. Isotypes: f! gh! ny! us/

Cyathea grenadensis Trevis. Atti Instit. Venat. II, 2: 164. 1851, nom. nud. Linden 1022 is cited.

Cyathea Lindeniana Presi, Epim. Bot. 30. 1852. Holotype: Nova Granada, Prov. Maraquita, (Colombia), Linden 1022. Isotypes: fragm. Herb. Mett. b! fragm. ex bm, us!, BR photo GH,K photo GH.

Cyathea boconensis Karst. Linnaea 28: 458. 1856. Holotype: none cited, but probably: Monte glacialis Meridensis, 1000 m, cited in Fl. Columb. 2: 171, t. 190. Isotype: Páramo de Bocono, Serrania de Mérida (Venezuela), Karsten , fragm. Herb. Mett. b!

Cyathea frondosa Karst. Fl. Columb. 1: 149, t. 74. 1860. Holotype: Andes of Bogotá, (Colombia), 2700 m, Karsten. Isotype: Karsten 196, b! (Lindig 196 is indicated by another label on the same b sheet, p! ) .

Cyathea Mettenii Karst. Fl. Columb. 1: 113, t. 56. 1860. Holotype: Andes of Bogotá, (Colombia), 2700 m, Karsten. Isotype: Bogotá, 9000 ft., Karsten , b!

Cyathea squamipes Karst. Fl. Columb. 1: 199, t. 99. 1861. Holotype: Mérida, (Venezuela), 1000-1500 m, Karsten , w photo and fragm. bm! Cyathea patens Karst. Fl. Columb. 2: 173, t. 191. 1869; not Hort. 1851, nom. nud.

Holotype: Guadelupe, Bogotá, (Colombia), 2900 m, Karsten. Isotype: Colombia, Karsten , b! is similar to t. 191, f. 1.

Cyathea lepidopoda C. Chr. Ind. Fil. 193. 1905, nom. superfl. An intended nom. nov. for "Cyathea squamipes Sod." not Karst, but Sodiro uses C. squamipes Karst, with reference to the Fl. Columb.

Cyathea Herzogii Rosenst. Meded. Rijks. Herb. Leiden 19: 7. 1913. Holotype: Yungas de San Mateo, Bolivia, Herzog 1990 , l photo gh. Isotypes: s! us!

Cyathea catacampta Alston, Jour. Wash. Acad. Sci. 48: 231. 1958. Holotype: between Rio Miraflores and Rio San Martin, Volcán de Cumbal region, Nariño, Colombia, Ewan 16153, bm. Isotype: us!

This is the most common and widely distributed of the varieties. In its variation it tends toward other sympatric varieties, especially var. meridensis and var. chimbor azensis. In Bolivia there is a variation with the segments slightly pubescent above (Cardenas 3059) or strongly so, ( Herzog 1990) that is similar in this character to var. Maxonii of Costa Rica.

Venezuela and Colombia, south to Bolivia. Montane forest, cloud forest, subparamo forest and Chusquea thickets, persisting after clearing, 1000 to

usually 2200-3000 to 4200 m. Stem to 15 m tall, leaves to 3 m long. selected specimens. Venezuela. Trujillo: cerca La Puerta, Vareschi 7492 (ven).

Mérida, (Venezuela), 2000 m, Karsten. Chuchilla, 20 km west of Mérida on road to La Azulita, Tryon ò- Tryon 5773 (gh). Tachira: south of San Vincente de la Revancha, Steyermark, Dunsterville ir Dunst er- ville 100748 (gh); debajo de Páramo de La Negra, Steyermark ir Rabe 96956 (gh).



78 ROLLA TRYON

Colombia. Magdalena: Sierra de Perijá, 23 km east of Codazzi, Grant 10948 (gh,us); Sierra Nevada de Santa Marta, Smith 1020 ( f,gh,mo,ny,us ) . Norte de Santander: Pamplona to Toledo, Killip ò Smith 19835 , 19857 , 19861 (gh,ny,us). Antioquia: Santa Elena, Archer 1251 (us); 8 km west of Valdivia, Madison 804 (gh). Santander: Mesa de los Santos, Killip ir Smith 15088 (gh). Cundinamarca: Páramo San Miguel, Gutierrez ò- Jaramillo 279 (gh,ny); Fusugasuga, Cuatrecasas 8035 (f,us), 8037 (f,gh,us). Valle: Piedra de Moler, Río Dagua, Cuatrecasas 15095 (f,gh,us); Los

Maps 38-45. 38, Cyathea tenera; 39, C. dissoluta; 40-44, C. caracasana; 40, var. boliviensis; 41, var meridensis; 42, var. caracasana; 43, var. chimborazensis; 44, var. Maxonii; 45, C. Lechleri.




Farallones, Alto del Buey, Cuatrecasas 17886 , 18043 (f,us). Tolima: above Anaime, on road south of Cajamarca, Barrington 477 (gh). Cauca: Cerro Munchique, Try on ir Tryon 6009 (gh,ny). Huila: Río Villalobos, vicinity of Río Suazita, Schultes ir Villa- real 5185 (gh, mo, NY, us). Putumayo: Valle de Sibundoy, Cuatrecasas 11675 (f,us). Nariño: Volcán de Chiles, Ewan 16012 (gh,us). Ecuador. Carchi: Rio Chingual drainage, Ewan 16307 (gh,us). Pichincha: confluence Río Tandápi and Río Pilatón, Aloag to Santo Domingo, Sparre 14049 (gh). Loja: 45 km north of Loja on road to Cuenca, Madison 918 (gh). Zamora-Chinchipe: km 18, road to Loja-Zamora, Sparre 16310, 16312 , 16313 , 16488 (gh). Peru. Amazonas: entre Ingenio y Pomacocha, López , Sagástegui ir Collante 4312 (gh). Huánuco: Carpish, Tryon ir Tryon 5326 (f,gh,ny). Loreto: Divisoria, Prov. Cornei Portillo, Ferreyra 1074 (gh), 1696 (us, usm). Junin: Oxapampa, Soukup 2335 (f,gh). Cuzco: Machu Picchu, Iltis , Iltis i Ugent 1025 (gh,us). Puno: San Juan del Oro, valle del Alto Tambopata, Ferreyra 16684 (gh). Bolivia. La Paz: Yungas, Bang 562 (gh,mo,ny); Sailapata, Prov. Ayo- paya, Cardenas 3059 (gh,us). Cochabamba: Prov. Chapare, Steinbach 9046 (f,gh, mo, US ) , 9425 ( F,GH,MO,NY,US ) , 9449 (f,GH,MO,ny).

36b. Cyathea caracasana var. meridensis ( Karst. ) Tryon, comb. nov. FIGS. 131-132. MAP 41.

Cyathea meridensis Karst. Fl. Columb. 2: 161, t. 184. 1869, ex char. Holotype: Mérida, (Venezuela), 2000 m, Karsten.

Alsophíla petiolulata Mett. Ann. Sci. Nat. V, 2: 263. 1864, nom. nud. The epithet is credited to Karsten (-Cyathea petiolulata Karst.?), although Lindig 285, cited, gh! is Cyathea caracasana var. meridensis.

Cyathea parvifolia Sod. Sert. Fl. Ecuad. 2: 7. 1908. Holotype: Silv. subandin. occ. Pichincha, Sodiro. Isotypes: 3/1901, Sodiro, a! p! photo gh,us!

This variety is rather uniform in its pinnule shape and indûment, al-

though it does grade into var. boliviensis in these characters. Some specimens approach other varieties, for example, Barrington 455 has a few of the long-stalked pinnules characteristic of var. chimborazensis , and Cuatrecasas 6710 , Killip & Smith 17821 are pubescent on the upper surface of the segments as in var. Maxonii.

Venezuela, Colombia and Ecuador. In wet forests and woodlands, cloud forest and thickets, 1800 to usually 2400-2800 to 3400 m. Stem 8 m tall.

selected specimens. Venezuela. Trujillo: between Boconó and Guaramacal, Steyermark 104864 (gh). Mérida: Laguna Coromoto, Vareschi ir Vareschi 5822 (ven). Tachira: 20 km south of San Vincente de la Revancha, Steyermark, Dunster- ville ir Dunsterville 100967 (gh). Bolivar: Ilu-tepuí, Gran Sabana, Maguire 33530 (ny,us). Colombia. Santander: Páramo Rico, Kiüip ir Smith 17821 (gh,ny,us). Cundinamarca: San Miguel, Cuatrecasas 6710 (f,gh,ny); Bogotá to Fusugasuga, Barrington 455 (gh). Valle: La Cumbre, Killip 11404 (gh,ny,us). Tolima: above Volcancitos, old Quindio Trail, Killip ir Hazen 9496 (gh,ny). Nariño: La Granja Botana, Pasto, Barrington 506 (gh). Ecuador. Loja: 5 miles north of San Lucas, Wiggins 10999 (ny,us).

36c. Cyathea caracasana var. caracasana FIGS. 133-134. MAP 42.

Cyathea caracasana ( Kl. ) Domin, Pterid. 262. 1929. Alsophila caracasana Kl. Linnaea 18: 541. 1844. Holotype: Caracas, (Venezuela),

Moritz 117. Isotypes: gh! p! (Domin, Pterid. Dominica, t. 10, f. 8 and t. 18, f. 1, figures Moritz 117). Moritz 394 (gh) is nearly identical and Domin cites a sheet bearing both numbers.



80 BOLLA TRYON

Figs. 129-138. Cyathea caracasana. 129-130, var. boliviensis: 129» Pinnules, Colombia, Ewan 16012 (gh), X Vz; 130, Portion of fertile pinnules, Bolivia, Herzog 1990 (us), X 2. 131-132, var. meridensis, Colombia, Killip & Hazen 9496 (gh): 131, Pinnules, X Vz' 132, Portion of fertile pinnule, X 2. 133-134, var. caracasana: 133, Pinnules, Jamaica, Proctor 22547 (gh), X 134, Portion of fertile pinnule, Venezuela, Fendler 56 (mo), X 2. 135, var. chimborazensis: Pinnules, Colombia, Cuatrecasas 18186 (us), X Vz> 136-138, var. Maxonii: Pinnules, Costa Rica, Nisman 63 (gh), X %; 137, Portion of fertile pinnule, Costa Rica, Scamman 5885 (gh), X 2; 138, Petiole scale, Costa Rica, Nisman 108 (gh), X 4.




Hemitelia crenata Sod. Ree. Crypt. Vase. Prov. Quit. 18. 1883. Syntypes: Ecuador, Volcán Atacazo: Canzacoto, Boloňa, Sodiro. Lectotype: Hacienda Boloňa, 1550 m, 1882, Sodiro , p! photo gh.

Cyathea fulva Sod. Ree. Crypt. Vase. Prov. Quit. 13. 1883, not (Mart. & Gal.) Fée, 1857. Lectotype: Tambo-loma, Prov. Riobamba, Ecuador, 10/1882, Sodiro , p! photo gh. Isolectotypes : fragm. ex b,ny!, fragm. ex k,ny! The petiole of this collection agrees with the description. An excluded element, under the same label, is Cyathea pallescens (ny).

Hemitelia Sherringii Jenm. Jour. Bot. 24 (n.s. 15): 266. 1886. Holotype: Rose Hill, Jamaica, 11/1886, Sherring , k! Isotypes: us! and a series of sheets ny! mostly without data. Baker, Ann. Bot. 5: 8. 1891 says "only one plant seen."

Cyathea ocanensis Baker, Ann. Bot. 5: 184. 1891. Holotype: Ocaña, Colombia, Kalbreyer 608 , k. Dept. Santander, Nuga, Kdhreyer 608 , b! is evidently an isotype.

Cyathea crenata (Sod.) Christ, Farnkr. 323. 1897. Cyathea Sodiroi C. Chr. Ind. Fil. 195. 1905, nom. nov. for Cyathea fulva Sod., not

( Mart, and Gal. ) Fée. Cyathea producta Maxon, Jour. Wash. Acad. Sci. 12: 438. 1922. Holotype: Palma

Mocha Peak, Sierra Maestra, Oriente, Cuba, Léon 11181 , us! Isotype: ny! Alsophila caracasana var. petiolularis Domin, Mem. Roy. Czech Soc. Sci. n.s. 2

( Pterid. Dominica ) : 95. 1929, ex char. Syntypes : Venezuela, Caracas, Büschel ; Colonia Tovar, Tittier 10011 ; Caracas, Büschel.

Cyathea suhindusiata Domin, Mem. Roy. Czech Soc. Sei. n.s. 2 (Pterid. Dominica): 67, t. 8, f. 4-6, t. 9, f. 2. 1929. Holotype: Caracas, (Venezuela), Büschel. Venezuela, 1855, Büschel , bm! photo gh, is probably an isotype.

Cyathea Sherringii (Jenm.) Domin, Pterid. 264. 1929. Cyathea caracasana var. petiolularis (Domin) Domin, Acta Bot. Bohem. 9: 103.

1930. Trichipteris caracasana (Kl.) Tryon, Contrib. Gray Herb. 200: 45. 1970.

The disjunct distribution of var. caracasana has probably been a major reason for its extensive synonomy. However, the characters of indûment and indusium, mentioned in the key, are quite uniform throughout its

range. The continental-Greater Antilles distribution is similar to that of

Cyathea andina and C. gracilis. Cuba, Jamaica and Hispaniola, Venezuela, Colombia and Ecuador.

Montane rain forest, 1200-2000 m. Stem to 8 m tall.

selected specimens. Cuba. Oriente: between Río Yara and Rio Palmamocha, Sierra Maestra, Ekman 14362 (us). Jamaica. Port Royal Mountains, Jenman (ny); Hardwar Gap, Portland, Froctor 16503 (mo), 22547 (gh). Dominican Republic. Above Gros Figuier, Sierra de los Comisarios, Prov. Barahona, Ekman 6769 (us). Venezuela. Distrito Federal: near Avila, Delgado 3 (gh,us). Aragua: Colonia Tovar, Fendler 56 (gh,mo). Mérida: southwest of Canaguá, Steyermark 56432 ( f,gh,mo,ny ) . Colombia. Cundinamarca: Tequedama, Holton 69 (ny). Meta: Macizo Renjifo, Cordillera La Macarena, Idrobo ir Schuhes 1105 (us). Ecuador. Chimborazo: Chimborazo, Sodiro (ny).

36d. Cyathea caracasana var. chimborazensis ( Hook. ) Tryon, comb. nov. FIG. 135. MAP 43.

Alsophila chimborazensis Hook. Syn. Fil. 37. 1866. Holotype: Chimborazo, Ecuador, 3000-4000 ft., Spruce 5743 , k. Isotype: p! fragm. ex. Rosenst. us!

Cyathea chimborazensis (Hook.) Hieron. Hedwigia45: 230. 1906.

The characteristic long-stalked pinnules of var. chimborazensis rarely occur in var. boliviensis also, to the south of their area of sympatry. The



82 BOLLA TRYON

pinnule indûment is variable: for example, Little 8906 has rather numerous trichomes beneath, Cuatrecasas 8674 has búllate and flat scales beneath, and Bristol 384 is slightly pubescent on the upper surface of the segments.

Colombia and adjacent Venezuela, Ecuador. Wet montane forest, persisting in clearings, 1000 to usually 1600-2300 to 3000 m. Stem to 8 m tall, leaves to 3 m long.

selected specimens. Venezuela. Zulia: Sierra de Perija, southwest of Pishikakao and Iria, Steyermark, Dunsterville ir Dunstervitte 105693 (gh). Colombia. Chocó: 5°55' N, 76° IO' W, Madison 821 (gh). Norte de Santander: Vicinity of Toledo, KiUip Ò- Smith 20100 (gh,ny,us). Santander: east of Las Vegas, Killip ò- Smith 15861 (ny,us). Valle: La Cumbre, Killip 11363, 11364 (gh,ny,us); Piedra de Moler, Río Dagua, Cuatrecasas 15191 (f,us). Cauca: km 19 from Cali to Buenaventura, Barring- ton 498, 502 (gh). Huila: between Gabinete and Andalucía, Cuatrecasas 8674 (f,us); near Resina, Guadalupe to Florencia, Little 9083 (gh,us); 25 km eastsoutheast of Baraya, Little 8906 (gh,us). Putumayo: Valle de Sibundoy, Bristol 384 (gh,us). Nariño: above San Juan, Ewan 16031 (gh).

Figs. 139-146. 139-140. Cyathea Lechleri, Peru, Spruce 4723: 139, Pinnules, (mo), X 140, Portion of fertile pinnule, (gh), X 2. 141-142, C. gracilis , Jamaica, Maxon <Lr Killip 944: 141, Pinnules, (gh), X Vz; 142, Portion of fertile pinnule, (f), X 2. 143-144, C. ebenina, Venezuela, Fendler 50 (gh): 143, Pinnules, X %; 144, Portion of fertile pinnule, X 2. 145-146, C. Dudleyi, Peru, Dudley 10867B (gh): 145, Pinnules, X 146, Portion of fertile pinnule, X 2%.




36e. Cyathea caracasana var. Maxonii (Maxon) Try on, comb. nov. FIGS. 136-138. MAP 44.

Cyathea Maxonii Maxon, No. Amer. Fl. 16: 82. 1909. Holotype: 5 miles south of Cartago, Costa Rica, Maxon 524 , ny. Isotype: us!

Cyathea membranulosa Christ, Bull. Herb. Boiss. II, 7: 271. 1907. Syntypes: San Pascon, Costa Rica, 1500 m, Wercklé (Herb. Instit. Nat. Costaric. 17024), Herb. Christ, p! ; La Palma, Costa Rica, 1500 m, Wercklé (Herb. Instit. Nat. Costaric. 17082), Herb. Christ, p!

The pinnules of var. Maxonii are often long-triangular, although they sometimes taper to the apex from about the middle; they are usually short-stalked, but rarely sessile. The petiole scale often has dark streaks, or areas (Fig. 138), rather than a dark central portion. In some collections the scales are mostly dark brown and rather few of them have the typical dark central portion. While var. Maxonii is not as well-defined as the other varieties, it is the only one that is completely allopatìe.

Costa Rica. Montane forest and cloud forest, 915-2800 m, usually above 2000 m. Stem to 9 m tall and 30 cm in diameter, leaves to 4 m long.

selected specimens. Costa Rica. Alajuela: 31 km from Carrizal toward Cariblanco, Gastony ir Gastony 789 (gh); 19 km north of San Ramón, White ir Lucansky 1968-26 (gh). Heredia: Volcán Barba, near Porrosati, Burger ir Stolze 6007 (gh,ny); Volcán Barba, Scamman ir Holdridge 7868 (f,gh,ny). San José: 22 km north of San Isidro del General toward Cerro de la Muerte, Gastony ir Gastony 760 (gh). Cartago: 7 km south of Tapanti, Try on ir Tryon 7022 , 7025 (gh); La Chonta, km 55 Pan-American Highway, Nisman 105 , 107 , 108, 177 (gh). Puntarenas: San Vito, Mc Alpin 2263 (gh).

Group of Cyathea Lechleri KEY TO SPECIES 37-40

a. Lamina bipinnate-pinnatifid throughout or tripinnate only at the base of central to basal pinnae; larger ultimate segments usually ca 10 mm long or more b. b. Pinnules short-stalked, or when long-stalked the basal segments not decurrent on the pinnule stalk which is only somewhat lighter, concolorous, or darker beneath than the pinna-rachis c. c. Pinnules with nearly parallel sides, tapering to an acute to acuminate apex, sessile to short-stalked throughout the lamina; Venezuela, Peru, Bolivia. 37. C. Lechleri.

c. Pinnules broadly long-triangular and long-stalked, especially toward the base of central to basal pinnae, usually long-acuminate; Jamaica, Costa Rica, Colombia. . . 38. C. gracilis.

b. Pinnules long-stalked and the basal segments decurrent onto the pinnule stalk which is of strongly contrasting lighter color beneath to the dark pinna-rachis; Venezuela, Colombia, Peru 39. C. ebenina.

a. Lamina tripinnate nearly throughout; larger ultimate segments small, ca. 5 mm long; Peru 40. C. Dudleyi.

A very natural group of distinctive, mostly Andean species. Cyathea Lechleri relates the group to either C. caracasana, or perhaps to C. platy -

lepls. All of the other species are clearly specialized in the pinnule and lamina complexity characters mentioned in the key.

37. Cyathea Lechleri Mett. FIGS. 139-140. MAP 45.

Cyathea Lechleri Mett. Fil. Lechl. 2: 32. 1859. Holotype: St. Gavan, (San Gavan,



84 ROLLA TRYON

Puno), Peru, Lechler, lz, destroyed, fragm. probably ex lz: Lechler 2309 , Herb. Mett. b!

Cyathea castanea Baker, Syn. Fil. 451. 1874. Holotype: Andes of Peru, (Tarapoto), Spruce 4723 , k. Isotypes : gh! mo! p! us!

Cyathea purpurea Morton, Fieldiana: Bot. 28: 7. 1951. Holotype: between Agüita and summit of Cerro Duida, Amazonas, Venezuela, Steyermark 58395 , f. Isotype: us!

The Venezuelan materials of Cyathea Lechleri are somewhat variable. The petiole and rachis may be pubescent or with only scattered trichomes, and the petiole color varies from atropurpureous to reddish-brown.

Venezuela, Peru and Bolivia. Montane forest, dense cloud forest and bamboo thickets, 1600-2200 m in Venezuela, 800-1500 m in Peru and Bolivia. Stem to 9 m tall, leaves to 2.5 m, or 6 m in one collection.

selected specimens. Venezuela. Aragua: to Los Carayaca from Colonia Tovar, White ir White 1970-31 (ny). Bolivar: Auyan-tepui, Vareschi 4782 (us,ven); Churi- tepuí (Muru-tepui), Wurdack 34257 (gh,ny,us); Chimantá-tepuí (Torono-tepuí ), Chimantá Massif, Steyermark 75444 ( f,gh,ny,ven ) . Amazonas: Cerro de la Neblina, Maguire , Wurdack ir Maguire 42369 , 42370 (us); Cerro Sipapo (Paráque), Maguire ò - Politi 28557 (gh,ny,us); Serrania Parú, Cowan ir Wurdack 31365 (gh,ny,us). Peru. Huánuco: Cerros dei Sira, Rio Lulla-Pichis watershed, Dudley 13007 , 13213 , 13262 (gh). Bolivia. La Paz: San Carlos, Mapiri region, Buchtien 291 (gh,ny); San José, Mapiri, Cardenas 1009 (gh).

38. Cyathea gracilis Griseb. FIGS. 141-142. MAP 46.

Cyathea gracilis Griseb. Fl. Brit. W. Indies 704. 1864. Holotype: Fox Gap, Jamaica, Purdie , k!

Cyathea gracilis, in Jamaica, Costa Rica, and Colombia, expresses the

relatively strong biogeographic affinity between those countries. In addition to the characters mentioned in the key, the species has very long, gradually acuminate pinnules (especially the larger ones) and a very firm, dull indusium.

Jamaica, Costa Rica and Colombia. Montane forest and cloud forest, 900-1600 m in Jamaica, ca 1500 m in Costa Rica, 1700 m in Colombia. Stem to 2 m ( probably more ) tall, leaves to 3 m long.

selected specimens. Jamaica. Morce's Gap, St. Andrew, Underwood 1536 (ny); Mt. Horeb Trail, St. Andrew, Crosby , Hespenheide ir Anderson 316 (f,gh,ny); below New Haven Gap, St. Andrew, Maxon ir Killip 944 (f,gh,ny); east slope of John Crow Mountains, 2 % miles southwest of Ecclesdown, Proctor 5711 (mo); Crown Peak, John Crow Mountains, Proctor 4583 (mo). Costa Rica. San José: La Palma, Brade ir Brade , 853 (us); above La Hondura, Gastony ò- Gastony 772 , 773 (gh); Las Nubes, Scamman 7004 (gh). Colombia. Antioquia: 8 km west of Valdivia, Madison 803 (gh).

39. Cyathea ebenina Karst. FIGS. 143-144. MAP 47.

Cyathea ebenina Karst. Linnaea 28 : 461. 1856. Holotype: between Caracas and Puerto Cabello, Venezuela, Karsten. Isotype: Caracas, Karsten , b!

A very distinctive species with long-stalked pinnules with the basal segments decurrent onto the upper part of the pinnule stalk.




Venezuela, Colombia and Peru. Montane forest, 2100-2500 m. Stem to 3 m tall, leaves to 1.5 m long.

selected specimens. Venezuela. Aragua: Colonia Tovar, F endler 50 (gh), Moritz 393 (f,gh,p). Colombia. Valle: La Cumbre, Killip 11386 (gh,ny,us); Gibralter, north of Las Brisas, Cuatrecasas 22525 (f,us). Huila: 25 km eastsoutheast of Baraya, Little 8905 (gh,us). Peru. Amazonas: 3 km north of Lake Pomacocha, Prov. Bongara, Hutchison ir Wright 6814 (gh,ny).

40. Cyathea Dudleyi Try on, spec. nov. Figs. 145-146. Map 48.

Folia ad 1.25 m longa; petiolus atrovinosus vel atropurpureus fere laevis squamis structura marginata caducis fere concoloribus fuscis; lamina tripinnata; pinnae ad 30 cm longae; pinnulae pagina inferiore squamis bullatis brunneis; segmenta ultima ad 5 mm longa; sori subcostales in venis simplicibus; indusium sphaeropteroideum. Holo- typus: 77°33/ W, 12°37' S, Prov. Convencion, Dept. Cuzco, Peru, Dudley 10867B, gh. Paratypes: same locality, Dudley 10738 , 10867, gh.

Except in its lamina complexity, Cyathea Dudleyi appears close to C. Lechleri. It is very similar to C. microphylla in its lamina architecture, but that species has very different petiole scales. The species is named for Dr. T. R. Dudley, who has added greatly to our knowledge of Peruvian ferns through his extensive collections, especially those from the Cordillera Vilcabamba.

Cordillera Vilcabamba, Peru. Very wet, dense cloud forest, 2600-2700 m. Stem 0.6-1.2 m tall, leaves to ca 1 m long.

Putative Hybrids

The following hybrids are proposed with varying degrees of confidence. The first two, 41 and 42, are certainly hybrids. Conant (1975) has presented a detailed discussion of 41, Cnemidaria hórrida X Cyathea arborea , including observations of its occurrence with both parents. Cnemidaria hórrida X Cyathea parvula is rather similar to 41 and Cnemidaria hórrida is certainly one parent. The Cyathea species involved needs to be con-

Maps 46-48. 46, Cyathea gracilis; 47, C. ebenina; 48, C. Dudleyi.



86 ROLLA TRYON

firmed by field study. The other, less certain, hybrids have each been assessed individually. In some, the lamina architecture is suggestive of a hybrid, especially those involving a species of Cnemidaria. Others are rarely collected and are intermediate between common species. In all cases, the interpretation as a hybrid seems more probable than that of a variation of a species or as an endemic species.

41. Cnemidaria hórrida X Cyathea arborea FIG. 147.

Hemitelia Wilsonii Hook. Syn. Fil. 30. 1865. Holotype: Mansfield near Bath, Jamaica, 1000 ft., Wilson 731 , k! Isotype (probable) : gh!

Cyathea Wilsonii (Hook.) Domin, Pterid. 264. 1929.

This hybrid will key out to species 1-8, 9-12 or usually 17-19. It may be readily distinguished from the species of Cyathea in the Greater Antilles, C. andina , C. parvula and C. arborea , by its predominantly ad- nate pinnules. The spores are abortive. The relatively well- developed indusium varies from about half cyathiform to fully cyathiform. Conant (1975) has presented a full discussion of the hybrid status of Hemitelia Wilsonii.

additional specimens. Jamaica. Claverty Cottage, Herb. Jenman (ny); vicinity of Castleton Gardens, St. Mary, Proctor 18419 (a). Haiti. Morne Chapelet, Massif du Nord, Ekman H4835 (ny,s); Poste Marie Congo, Massif du Nord, Ekman H4846 (s,us). Dominican Republic. Laguna, Península de Samaná, Ekman H15014 (gh,ny); Sanchez, Península de Samaná, Ekman H14752 (ny); Liali, Abbott 2660 (gh,ny,us); Miches to Higuey, Seibo, Gastony, Jones ò- N orris 654 (gh,ny). Puerto Rico. Sabana Road, Luquillo National Forest, Municip. Río Grande, 5 March, 1970, Kepler (El Verde Field Station, Puerto Rico Nuclear Center), Conant 626 , 627 (gh); El Verde, Municip. Rio Grande, Conant 546 (gh); Maricao, Hess 371 (fragm. ex us,ny), Hioram 804 (fragm. ex us,ny); Quebradilles, Hioram 182 (fragm. ex us,ny); El Yunque Road, Luquillo National Forest, Municip. Rio Grande, 5 March, 1970, Kepler (El Verde Field Station, Puerto Rico Nuclear Center).

42. Cnemidaria hórrida X Cyathea parvula FIG. 148.

Alsophila sessilifolia Jenm. Jour. Bot. 20 (n.s. 11): 325. 1882. Lectotype: Mans- field, near Bath, Jamaica, Wilson 520 , k! Isotype: bm! photo gh,mo! (an exact match of the lectotype), fragm. ex bm,us!, photo us. Lectoparatypes : Wilson 513 , bm, Wilson Al, bm. Isolectoparatype: Wilson 513 , ny!

Hemitelia sessilifolia (Jenm.) Jenm. West Ind. Guiana Ferns 44. 1898. Cyathea sessilifolia (Jenm.) Domin, Pterid. 263. 1929.

This hybrid will key to species 1-8, or to species 9-12. The only Greater Antillean species in these groups are Cyathea andina and C. parvula. The hybrid differs from both of them in its large, entire, acuminate pinnule- segments near the apex of the pinnae. The spores are evidently abortive. Thé leaf architecture indicates one parent to be Cnemidaria hórrida ; the other is probably Cyathea parvula , which also occurs in Cuba, Jamaica, and Puerto Rico. This hybrid may be distinguished from the previous one, Cnemidaria hórrida X Cyathea arborea , by its mostly sessile or very short-




Figs. 147-154. Hybrids. 147, Cnemidaria hórrida X Cyathea arborea, pinnules, Dominican Re- public, Ekman HI 5014 (gh), X 148, Cnemidaria hórrida X Cyathea parvula, pinnules, Puerto Rico, Conant 612 (gh), X 149, Cnemidaria grandifolia var. obtusa X Trichipteris aspera , Por- tion of lower pinna, Grenada, Sherring (k), X %; 150-151, Cnemidaria spectabilis var. spectabilis X Cyathea tenera: 150, Portion of pinna, Trinidad, Broadway ò- Hombersley (us), X 151, Portion of fertile pinna, Trinidad, Broadway 5719 (mo), X 1%; 152, Cyathea arborea X Tri- chipteris aspera , pinnules, Jamaica, Proctor 4000 (mo), X %; 153, Cyathea arborea X Trichipteris sp., pinnules, Cuba, Wright 950 (gh), X 154, Cyathea divergens var. divergens X Trichipteris stipularis, pinnules, Costa Rica, White & Lucansky 1972-067 (gh), X



88 ROLLA TRYON

stalked pinnules, the brown rather than white scales on the abaxial surface of the pinnules, and its small, rather than well-developed, indusium. Except for the small indusium, the Cyathea parent could as well be C. furfuracea .

additional specimens. Cuba. Rancho Mundito, Pinar del Rio, Jan. 18, 1953, Acuña (us). Puerto Rico. Road 143, Municip. Rio Grande, Conant 612 (gh); El Toro Negro, Jan. 14, 1966, Woodbury (El Verde Field Station, Puerto Rico Nuclear Center).

43. Cnemidaria grandifolia var. obtusa X Trîchipteris aspera FIG. 149.

Alsophila Elliottii Baker, Ann. Bot. 6: 96. 1892. Lectotype: St. Catherine's Peak, Pyrenees, Grenada, Sherring , k! Lectoparatype: Antoine, Bellevue, Grenada, Elliott , k.

Hemitelia Elliottii (Baker) Maxon, Contrib. U.S. Nat. Herb. 17: 415. 1914. Cyathea Elliottii (Baker) Domin, Acta Bot. Bohem. 9: 113. 1930.

This hybrid will key to species 9-12 of Central America and northern South America. The leaf architecture suggests a hybrid with a species of Cnemidaria, and the small indusium an exindusiate species as the other parent. The proposed parents are both relatively common in Grenada. Trichipteris aspera is an Antillean species, including Cyathea muricata.

additional specimen. Grenada. Herb. Jenman (ny).

44. Cnemidaria spectabilis var. spectabilis X Cyathea tenera FIGS. 150-151.

Hemitelia Hombersleyi Maxon, Jour. Wash. Acad. Sci. 25: 528, f. 1. 1935. Holotype: Aripo Road, via Arima, Trinidad, June 12, 1925, Broadway ò- Hombersley , us! Para- types (all Trinidad); Blanchisseuse Road, Broadway 6118 , us; Las Lapas Road, Broadway 5913 , us, Hombersley 208 us! Isoparatypes: Broadway 5913, mo! Hom- bersley 208, gh! ny!

Cyathea Hombersleyi (Maxon) Stolze, Fieldiana: Bot. 37: 81. 1974.

This hybrid will key to species 9-12, 13-16, or often to 17-19, none of them growing in Trinidad. The relatively simple lamina architecture suggests a species of Cnemidaria, while the relatively well-developed indusium indicates another parent with a large indusium. Cyathea tenera is the only species in Trinidad with a large ( sphaeropteroid ) indusium. None of the material seen is suitable to assess the degree of development of the spores.

Apparently known from three plants: (a) Broadway 6118; (b) Broad- way 5913 and Hombersley 208 ; (c) June 12, 1925, Broadway ir Hombers- ley, Broadway 5719, and 1925, Hombersley .

additional specimens. Trinidad. Aripo Road, via Arima, 1925, Hombersley (gh, ny), and also Broadway 5719 (mo), are from the same plant as the type.

45. Cyathea arborea X Trichipteris armata

This hybrid has generally been identified as Hemitelia calolepis (see 47),




but the type of that name is from Cuba, where Trichipteris armata does not grow. It will key to species 1-8 and of these, only Cyathea parvula, with a small indusium, is in Jamaica. Most of the sporangia appear to be poorly developed and well-developed spores have not been observed. The pinna architecture and whitish petiole scales suggest Cyathea arborea as one parent, and the long trichomes on the pinna-rachis and pinnules beneath as well as the toothed segments suggest Trichipteris armata as the other. The large hemitelioid indusium is consistent with this parentage.

specimens seen. Jamaica. Wilson 681 (ny); St. Thomas: between House Hill and Cuna Cuna Gap, Maxon 8968 (ny,us); Corn Puss Gap, Proctor 4001 (mo,us); Moody's Gap, Underwood 1550 (ny,us); Bath, 1895, Gilbert (gh,mo,ny).

46. Cyathea arborea X Trichipteris aspera FIG. 152.

Hemitelia Lewisii Morton & Proctor, Amer. Fern Jour. 41: 49. 1951. Holotype: Corn Puss Gap, St. Thomas, Jamaica, Proctor 4000 , us. Isotypes: ij,mo! ph.

Cyathea Lewisii (Morton & Proctor) Proctor, Bull. Inst. Jam. Sei. Ser. 5: 21. 1953.

The collections will key to species 1-8. Among these only Cyathea parvula , with an aculeate petiole and usually smaller indusium is in

Jamaica. This hybrid has a tuberculate petiole and the indusium is attached about halfway around the receptacle. The material is too mature to allow assessment of the spore development. Eight other tree ferns were listed by Morton and Proctor as occurring in the same locality as Hemi- telia Lewisii. Among these, the brownish scales on the costules and the

moderately developed indusium suggest the indicated parentage.

47. Cyathea arborea X Trichipteris sp. FIG. 153.

Hemitelia calolepis Hook. Syn. Fil. 29. 1865. Holotype: Monte Verde, Cuba, Wright 950 , K. Isotypes: f! gh! mo! us!

Cyathea calolepis (Hook.) Domin, Pterid. 263. 1929.

This hybrid will key to species 1-8 and among them only Cyathea parvula occurs in Cuba. The hybrid may be distinguished by its tuberculate petiole and the pinna architecture that is similar to C. arborea.

Cyathea parvula has an aculeate petiole and different pinna architecture. The sporangia are often not well-developed and the spores appear to be abortive. The whitish scales of the petiole and whitish búllate scales on the under surface of the lamina, as well as the pinna architecture suggest C. arborea as one parent. The relatively small indusium, attached from about /4 to /2 around the receptacle suggests an exindusiate species as the other. However, there is no information on which Trichipteris species were growing in the same place as the hybrid.

additional specimens. Cuba. W right 891 ( GH,NY ) .



90 ROLLA TRYON

48. Cyathea divergens var. divergens X Trichipteris stipularis FIG. 154.

Cyathea Holdridgeana Nisman & Gomez, Amer. Fern Jour. 61: 168, f. 3, 4. 1971. Holotype: La Chonta, km 55 Interamerican Highway, Prov. Cartago, Costa Rica, Nisman 104 , cr. Isotypes: f,gh! Paratypes: (all La Chonta, Herb. Gómez); Gómez 2542 , 2560, 2577; Goldgewicht ir Gómez 2675; km 54 Interamerican Highway, J. A. Saenz.

This hybrid will key to species 1-8, and among these only Cyathea multiflora grows in Costa Rica. It has sessile to short-stalked basal pinnules on the pinnae, rather than long-stalked ones as in the hybrid. These long- stalked pinnules suggest C. divergens var. divergens as one parent, and the small indusium, minute to about around the receptacle, indicates an exindusiate species as the other. Trichipteris stipularis is a common species in the Cordillera de Talamanca and may be the other parent. The spores are unusual in lacking perine and many of them have the outer layer of the exine peculiarly irregular.

Cyathea Holdridgeana is possibly a local endemic species of Costa Rica. Luis D. Gómez (pers. comm.) informs me that neither of the proposed parents were growing in the two localities where he has seen it. However, its characters are suggestive of a hybrid and I prefer to treat it as one until further field studies indicate that it is an endemic species.

additional specimens. Costa Rica. 16 km south of El Empalme, White , Nortis ir S oeder 1972-067 (gh).

49. Cyathea divergens var. Tuerckheimii X Cyathea fulva

The long-stalked pinnules, especially the basal ones, suggest Cyathea divergens var. Tuerckheimii as one parent, and the brown petiole scales suggest Cyathea fulva as the other. The petiole scurf is consistent with the parentage; it is brown as in C. fulva , although not crested, and there are some whitish small scales, as in C. divergens. The indusium is sphaeropteroid, but the sori are too young to assess the development of the sporangia or spores.

specimens seen. México. 20 km antes Huatusco, carretera Puente Nacional a Huatusco, Veracruz, N exiling ir Gomez-Pompa 2443 (gh).

50. Cyathea tenera X Trichipteris sp.

Hemitelia trinitensis Jenm. West Ind. Guiana Ferns 46. 1898. Trinidad. A specimen in TRiN, fragm. bm!, photo bm,gh, is probably the holotype.

Cyathea trinitensis (Jenm.) Domin, Pterid. 264. 1929.

This hybrid will key to species 1-8, or to 17-19, none of which occur in Trinidad. The indusium varies from hemitelioid to discoid, which is suggestive of a hybrid. More adequate material is required.




NAMES OF UNCERTAIN IDENTITY AND EXCLUDED NAMES

The following list includes (a) those names that apply to species of Cyathea that I have been unable to place more precisely, ( b ) those names that cannot be accurately identified to genus without an examination of the original materials, and (c) a few names to be referred to other genera. No attempt has been made to include the much larger number of names placed in Cyathea by Domin and others. These are dealt with under other revisions, Nephelea (Gastony, 1973), Cnemidaria (Stolze, 1974), Tri- chipteris (Barrington, 1974), Alsophila (Conant, MS) and Sphaeropteris (Windisch, MS).

Cyathea aphlebioides Christ, Bull. Herb. Boiss. II, 6: 179. 1906. There are three sheets under this name in Herb. Christ, p! One, with two petiole bases and the lower portion of a rachis with basal pinnae has the abortive ( subaphlebioid ) pinnae em- phasized by Christ in the name. The epithet aphlebioides was substituted for the original "decrescens." Lectotype: Navarro, 1400 m, Wercklé , C. Rica, 1905. Another sheet: Navarro, Costa Rica, Wercklé, 1905, is Cyathea Delgadii and the third: C. Rica, Navarro, 1400 m, Wercklé, 1905, is Cyathea fulva (fragm. ex Christ, us).

Cyathea arborea var. pusilla Bosco, Nuov. Giorn. Bot. Ital. n.s. 45: 141. 1938. Páramos de Portrerillos, 3200 m, and Plan de Sapote, 1800-2100 m, Ecuador, Crespl.

Cyathea azuayensis Sod. Crypt. Vase. Quit. 644. 1893. Prov. Azuay, Ecuador, 3000 m, Rimbach 23. Sodiro indicated a close comparison with Cyathea incarta, which suggests that this is a species of Nephelea and perhaps N. incarta.

Cyathea caduca Christ, Bull. Herb. Boiss. II, 7: 271. 1907. San José, Costa Rica, Jardin de Camiol Wercklé, 1906. The spiny trunk indicates that this name refers to a species of Nephelea.

Cyathea denudans Kze. Linnaea 18: 349. 1844. Mexico, Leibold, Herb. Roemer, lz, destroyed.

Cyathea Dyeri Sod. Crypt. Vase. Quit. 515. 1893. Chimborazo, cerca Saltuco, Ecuador, 300-600 m, Sodiro. A portion of a rachis with three pinnae: Saltuco, Prov. Bolivar, 9/1872, Sodiro, p! is a species of Cyathea.

Cyathea Eggersii Hieron. Engl. Bot. Jahrb. 34: 438. 1904. El Recreo, Río Puntilla, Prov. Manabi, Ecuador, Eggers 15320. Isotypes: f! gh! A species of Cyathea but not identifiable from the pinna material seen.

Cyathea fulva var. minor Sod. Ree. Crypt. Vase. Quit. 14. 1883. Holotype: Chim- borazo cerca de S. Pablo de Atenas, Prov. de Riobamba, Ecuador, Sodiro.

Cyathea furfuracea Sod. Sert. Fl. Ecuad. 2: 7. 1908, not Baker, 1874. Pichincha, Ecuador, Sodiro. Cyathea pichinchae C. Chr. Ind. Fil. Suppl. 21. 1913, nom. nov. for Cyathea furfuracea Sod., not Baker.

Cyathea hexagona Fée, Mém. Fam. Foug. 8: 111. 1857. Huatusco, Mexico, Schaffner 237. A sheet at k! photo gh, is a mixture of a Cyathea species and Nephelea mexicana. Gastony (Contrib. Gray Herb. 203: 147. 1973), excluded the Nephelea material as inconsistent with the description. The Cyathea is either C. fulva or C. diver gens var. Tuerckheimii.

Cyathea hirtula var. multisorosa Karst. Linnaea 28: 462. 1856. Montalban and Cuirgua, Venezuela, Karsten.

Cyathea Jurgensenii Fourn. Mex. PI. 1: 135. 1872. Jurgensen 874, p! photo gh is a large pinnae which could be either Cyathea fulva or C. diver gens var. Tuerckheimii; fragm. ex ny,us!

Cyathea Mettenii var. caucana Hieron. Engl. Bot. Jahrb. 34 : 437. 1904. Supra Paletará, Cauca, Colombia, Lehmann 3482.

Cyathea patens Houlst. & Moore, Gard. Mag. Bot. 3: 330. 1851, nom. nud. Hort, from Jamaica, the meager description is not sufficient to effect valid publication.

Cyathea pilosa Sampaio, Bol. Mus. Nac. Rio 1: 13. 1923, nom. nud. Cyathea puberula Sod. Ree. Crypt. Vase. Prov. Quit. 16. 1883. Rio Peripe, cerca San

Miguel de los Colorados, Ecuador, Sodiro. A small specimen: Peripe, 8/1875, Sodiro, p! is a species of Cyathea.

Cyathea pygmaea Hort. Gard. Chron. III, 15: 663. 1894, nom. nud. Cyathea subindusiata var. chontiUa Domin, Mem. Roy. Czech Soc. Sci. n.s. 2

(Pterid. Dominica): 68. 1929. Andes of Ecuador, 11,000 ft., Spruce.



92 ROLLA TRYON

Cyathea Schottland Kze. Linnaea 23: 246. 1850, nom. nud.9 Mexico. Cyathea Trejoi Christ, Bull. Herb. Boiss. II, 5: 733. 1905. San Pablo, Chiapas,

Mexico, 1500 m, 1903, Munch. The spiny trunk, a fragment ex Christ, ny! and the provenence refer this name to Nephelea mexicana.

Cyathea venosa (Kuhn) Domin, Pterid. 264. 1929. Hemitelia venosa Kuhn, Linnaea 36. 161. 1869. Caracas, Appun, fragm. b! photo gh, fragm. ex k,ny! The fragmentary material is not adequate for determination of the genus. Stolze (Fieldiana: Bot. 37: 81. 1974) came to the same conclusion.

Hemitelia Lindenii Hort. Gard. Chron. III, 15: 663. 1894, nom. nud.

LITERATURE CITED

Alston, A. H. G. 1959. The ferns and fern-allies of West Tropical Africa. Fl. West Trop. Africa, Ed. 2, Suppl. pp. 1-89.

Barrington, D. S. 1974. A revision of Trichipteiis ( Cyatheaceae ) . Ph.D. Thesis, Harvard University.

Carlquist, S. 1966. The biota of long-distance dispersal, III. Loss of dispersibility in the Hawaiian flora. Brittonia 18: 310-335.

Conant, D. S. 1975. Hybrids in American Cyatheaceae. Rhodora 77: 441-455. Gastony, G. J. 1973. A revision of the fern genus Nephelea. Contrib. Gray Herb.

203: 81-148. . 1974. Spore morphology in the Cyatheaceae, I. The perine and spor-

angial capacity: general considerations. Amer. Jour. Bot. 61: 672-680. and Ř. Ťryon. 1976. Spore morphology in the Cyatheaceae, II. The

genera Lophosoria, Metaxya , Sphaeropteris , Alsophila and Nephelea. Amer. Jour. Bot. ( in press ) .

Hallé, F. 1966. Etude de la ramification du tronc chez quelques fougères arbores- centes. Adansonia, n.s. 6: 405-424.

Lewis, W. H. 1971. High floristic endemism in low cloud forests of Panama. Biotropica 3: 78-80.

Morton, e. V. 1947. The American species of Hymenophyllum section Sphaerocion- ium. Contrib. U.S. Nat. Herb. 29: 139-201.

Steyermark, J. A. 1974. The summit vegetation of Cerro Autana. Biotropica 6: 7-13. Stolze, R. G. 1974. A taxonomie revision of the genus Cnemidaria (Cyatheaceae).

Fieldiana: Bot. 37: 1-98. Tryon, R. 1970. The classification of the Cyatheaceae. Contrib. Gray Herb. 200: 3-53. , 1971. The process of evolutionary migration in species or Selaginella.

Brittonia 23: 89-100. . 1972. Endemic areas and geographic speciation in tropical American

ferns. Biotropica 4: 121-131. and G. J. Gastony. 1975. The biogeography of endemism in the Cyathea-

ceae. Brit. Fern Gaz. 11: 73-79. Van Cotthem, W. 1970. Comparative morphological study of the stornata in the

Filicopsida. Bull. Tard. Bot. Nat. Bele. 40: 81-239. Walker, Ť. G. 1966. Á cytotaxonomic survey of the Pteridophytes of Jamaica. Trans.

Roy. Soc. Edinb. 66: 169-237. Windisch, P. G. 1976. The systematics of the group of Sphaeropteris hirsuta (Cyathea-

ceae). Mem. New York Bot. Gard, (in press). TAXA, PUTATIVE HYBRIDS AND DISTRIBUTION OF CYATHEA

1. Cyathea multiflora Sm.: British Honduras, Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Ecuador, Peru, Bolivia, Brazil.

2. Cyathea andina (Karst.) Domin: Hispaniola, Puerto Rico, French Guiana, British Guiana, Venezuela, Colombia, Ecuador, Peru, Bolivia, Brazil.

3. Cyathea Weatherbyana (Morton) Morton: Galapagos Islands. 4. Cyathea Alphonsiana Gómez: Cocos Island. 5. Cyathea notabilis Domin: Cocos Island. 6. Cyathea parvula (Jenm.) Domin: Cuba, Jamaica, Hispaniola, Puerto Rico.




7. Cyathea platylepis (Hook.) Domin: Venezuela, Colombia. 8. Cyathea Vilhelmii Domin: Peru. 9. Cyathea petiolata (Hook.) Try on: Panama, Colombia. 10. Cyathea impar Tryon: Panama. 11. Cyathea Steyermarkii Tryon: Venezuela. 12. Cyathea conformis (Tryon) Stolze: Panama, Colombia. 13. Cyathea decorata (Maxon) Tryon: Colombia. 14. Cyathea parva (Maxon) Tryon: Colombia. 15. Cyathea speciosa Willd.: Venezuela, Colombia. 16. Cyathea Haughtii (Maxon) Tryon: Colombia. 17. Cyathea arborea (L.) Sm.: Cuba, Jamaica, Hispaniola, Puerto Rico, Lesser An-

tilles south to Grenada. 18. Cyathea Alstonii Tryon: Colombia. 19. Cyathea peladensis (Hieron.) Domin: Colombia. 20. Cyathea divergens Kze.

20a. var. divergens: Costa Rica, Panama, British Guiana, Venezuela, Colombia, Ecuador, Peru.

20b. var. Tuerckheimii (Maxon) Tryon: Mexico, Guatemala. 21. Cyathea pallescens (Sod.) Domin: Colombia, Ecuador, Peru, Bolivia. 22. Cyathea simplex Tryon: Venezuela. 23. Cyathea corallifera Sod. : Ecuador. 24. Cyathea boliviana Tryon: Bolivia. 25. Cyathea straminea Karst. : Colombia. 26. Cyathea Ruiziana Kl. : Peru. 27. Cyathea microphylla Mett. : Peru. 28. Cyathea multisegmenta Tryon: Peru. 29. Cyathea fulva (Mart. & Gal.) Fée: Mexico, Guatemala, Honduras, Nicaragua,

Costa Rica, Panama, Venezuela, Colombia. 30. Cyathea Harrisii Maxon: Jamaica, Hispaniola. 31. Cyathea furfuracea Baker: Cuba, Jamaica, Hispaniola, Puerto Rico. 32. Cyathea suprastrigosa (Christ) Maxon: Costa Rica. 33. Cyathea Delgadii Sternb.: Costa Rica, Panama, British Guiana, Venezuela, Colom-

bia, Peru, Bolivia, Argentina, Paraguay, Brazil, Ilha Trindade. 34. Cyathea tenera (Hook.) Moore: Lesser Antilles, Trinidad, Margarita. 35. Cyathea dissoluta Jenm.: Jamaica. 36. Cyathea caracasana ( Kl. ) Domin.

36a. var. boliviensis (Rosenst. ) Tryon: Venezuela, Colombia, Ecuador, Peru, Bolivia.

36b. var. meridensis (Karst.) Tryon: Venezuela, Colombia, Ecuador. 36c. var. caracasana: Cuba, Jamaica, Hispaniola, Venezuela, Colombia, Ecuador. 36d. var. chimborazensis (Hook.) Tryon: Venezuela, Colombia, Ecuador. 36e. var. Maxonii (Maxon) Tryon: Costa Rica.

37. Cyathea Lechleri Mett.: Venezuela, Peru, Bolivia. 38. Cyathea gracilis Griseb.: Costa Rica, Jamaica, Colombia. 39. Cyathea ebenina Karst.: Venezuela, Colombia, Peru. 40. Cyathea Dudleyi Tryon: Peru. 41. Cnemidaria hórrida X Cyathea arborea: Jamaica, Hispaniola, Puerto Rico. 42. Cnemidaria hórrida X Cyathea parvula: Cuba, Jamaica, Puerto Rico. 43. Cnemidaria grandifolia var. obtusa X Trichipteris aspera: Grenada. 44. Cnemidaria spectabilis var. spectabilis X Cyathea tenera: Trinidad. 45. Cyathea arborea X Trichipteris armata: Jamaica. 46. Cyathea arborea X Trichipteris aspera: Jamaica. 47. Cyathea arborea X Trichipteris sp.: Cuba. 48. Cyathea divergens var. divergens X Trichipteris stipularis: Costa Rica. 49. Cyathea divergens var. Tuerckheimii X Cyathea fulva: Mexico. 50. Cyathea tenera X Trichipteris sp.: Trinidad.



94 BOLLA TRYON

Index to Exsiccatae Abbott 294(6); 410(17); 445, 747(6); 1590,

1664(31); 2067(6); 2612(17); 2660(41); 2710, 2712(17).

Acuña 6834(31); 12347(6). Agredo 431(29). Allen 1522(29); 3529(20a). André 1134, 2375(2). Archer 125l(36a). Aristeguieta 856(15). Asplund 5557 ( 1 ) . Augusto 854(6); 1401(17). Bailey, L. H. 741(17). Bailey & Bailey 279(17). Bang 562, 2318(36a). Barclay 908(9). Barrington 453, 454(36a); 455(36b); 456

(36a); 464(21); 474(29); 477(36a); 481, 482(25); 493(36a); 498(36d); 500(20a); 502(36d); 504(13); 505(25); 506(36b); 521(36a).

Beard 625(17). Bernardi 1868(20a); 1879(29). Bierhorst FG87, FG110(2). Blum et al. 1700, 1701(9). Box 377(17). Brade 8220(33). Brade & Brade 108(20a); 287(29); 288

(36e); 405, 496(1); 629(20a); 630, 631 (29); 853(38).

Bridarolli 4341(33). Bristol 384(36d). Britton & Britton 7210(17). Britton & Cowell 165, 286, 331(17); 4217

(6). Britton & Marble 691(6); 1422(17). Britton & Shafer 1693(17). Britton & Wilson 5328(17). Britton et al. 1351, 2285(34); 2409(17);

2482(6); 2620(31); 6292, 6622, 8173, 8404(17).

Broadway 1904, 5514(34); 5719, 5913(44); 7408, 9968(34).

Buchtien 291(37); 5139(33); 5140(36a). Bunting & Licht 396, 1246(1). Burch 548(20a). Burchell A703, 1896(33). Burger & Liesner 6793(1). Burger & Matta 4237(1). Burger & Ramirez 3964(29). Burger & Stolze 4941, 4982(1); 5238(32);

5667, 5683(29); 5828, 5876(1); 5963 (36e); 5964(32); 5968, 6007(36e); 6094 (1).

Cabrera 246(36a). Campbell 7731(35). Campos Porto 579(33). Carabia 3544(6).

Cardenas 1009(37); 3059(36a); 3150(21). Cardona, F. 2075(7). Carriker 7(20a). Chambers 2664(17). Chase 6211a(6). Chrysler 1850(17); 2049(30); 4586(31). Chrysler & Roever 4837(9). Clément 1656(17); 1663(6). Clute 89(31); 181(38); 289(17). Colinvaux 357(3). Colwell 585(17). Conant 596, 603(31); 612(42); 626(41);

685(31). Conant & Hodgdon 646(6). Conzatti & Gómez 2361(29). Copeland 15(29). Cornman 695(9). Cowan & Wurdack 30229, 31365(37);

31390(29). Crosby & Anderson 1044(17). Crosby et d. 308(31); 316(38). Cuatrecasas 5492(36a); 6710(36b); 8035,

8037(36a); 8418, 8438(21); 8507(19); 8674(36d); 11325(2); 11465(36d); 11- 675(36a); 11856(25); 13918(1); 13919 (20a); 15000, 15095(36a); 15191(36d); 15593(20a); 15737, 17429(13); 17886, 18043(36a); 18186, 18245(36d); 20994 (36a); 21611, 21649(36d); 21925, 21934 (36a); 22161(1); 22525(39); 23486(21); 23803(20a); 23811(36a).

Cuatrecasas & Jaramillo 12015(36a). Cuatrecasas et al. 12436, 12584(36a);

12728(36a). Cuming 1360(1). Curran 136(9). Daniel 2983(29); 3017(20a); 4175(29). D'Arcy 48D(17). Deam 483(1). Delgado 3(36c); 60(15). Duarte 1346(33). Dudley 10058, 101 15B, 10268, 10286B

(33); 10462, 10588(36a); 10738, 10861 (40); 10864(21); 10867B(40); 10943 (27); 10950B(21); 11020(36a); 11149 (21); 11258(33); 11275(36a); 11326(28); 11327(36a); 13007(37); 13055(36a); 13213, 13262(37); 13276, 13358(36a).

Dusén 623a, 3219, 10382, 10760, 11781, 16980(33).

Duss 1604, 1609, 4322(17). Eggers 5031(31); 5117(6); 5211(31); 6083;

6610(17); 6859, 6868(34); 6921, 6996a (17); 14899, 15320(21).

Eiten et al. 2127(33). Ekman 1638, 1805(17); H2873(6); H2954

(17); 3097(6); H3217(31); 3368(17);




H4365(6); H4835, H4846(41); 5430; H6393(31); H6769(36c); 7070, H11496 (31); H11499(2); H11678(30); H12865 (31); H14362(36c); H14752(41).

Elias 4(9). Englesing 290(1). Espina & Giacometto A134(20a). Espinosa 918(21). Esposto 659(2). Ewan 16012(36a); 16031(20a,36d); 16-

153(36a); 16182(21); 16307(36a); 16784, 16809(1).

Faull 12582(30). Fawcett 8419(6). Fendler 46(15); 50(39); 52(29); 56(36c);

80(34); 417, 421(9). Fernandez & Mora 1451(29). Ferreyra 1074(36a); 1694(2); 1696, 16684

(36a). Figueiras 8382(36d). Fisher 125(37). Fishlock 381(17). Flack 914(33). Foldats 4948(7). Fosberg 19950(19). Fredholm 3236(17). Fuertes 1515, 1320b(31). Funck 663(15). Funck & Schlim 413(29). Galeotti 6346(29). Garcia 50(15). Garcia-Barriga 17581(33). Gardner 1907(33). Gastony 10, 38(17). Gastony & Gastony 747, 748(32); 755

(20a); 757, 758(29); 759(20a); 760(36e); 761(32); 762(29); 772, 773(38); 774 (29); 779, 780(1); 789(36e); 790, 794 (33).

Gastony et al. 173, 197(31); 208(6); 213, 252, 253(31); 272(6); 324, 431, 469(31); 654(41); 656, 659, 660(6); 702(17); 720, 730(30).

Glaziou 2283, 2284, 2286, 15776(33). Göll et al. 401(17). Gómez 2370(32); 3349(4). Grant 10298(36a); 10323(21); 10324(36a);

10345, 10354(36a); 10743(2); 10948 (36a).

Grant & Fosberg 9308(21). Grubb et al. 1016(36a); 1334(36a). Gutierrez & Jaramillo 279(36a). Hamilton 243(6); 3247(29). Harling 5084(3). Harling et al. 10178(20a). Harris 10661(6). Hart 43(1). Hassler 4004(33), 10442(33). Hatch & Wilson 220(1).

Haught 1337(9); 1358(2); 1957(16); 2813 (33).

Hazen 9693(20a); 11834(1). Heller 4467(17); 4595(6). Heller & Heller 285(17); 705(1). Hellwig & Whitaker 1433(20a). Henri-Stanislas 1619(29). Herzog 1990(36a). Hespenheide 957, 1327(6); 1466(31). Hess 342(2); 371(41). Hinton 14274(29). Hioram 182, 804(41); 809(31); 1298(17);

7311(6). Hodge 6, 7(17); 6872, 6873(1). Hodge & Hodge 1973(34); 2444, 2880(17). Holdridge 1966(31); 2150(6); 5160(4);

5161(5). Holm & Iltis 533, 601, 605(32). Holton 69(36c). Hombersley 208(44). Howard 5221, 5224(17); 5859, 5963(6);

6465a, 6565(17); 10638(34); 11353, 11905(17); 11906(34).

Howard & Nevling 15776(17). Howard & Proctor 15033(6). Howell 9227(3). Husnot 398(17). Hutchison & Idrobo 3144(1). Hutchison & Wright 6814(39). Idrobo & Schultes 748(2); 1105(36c); 1130

(18). Iltis et al. 1025(36a). Im Thum 92(33). Irwin & Soderstrom 6282(33). Irwin et al. 10138, 16305(33). Jack 7273(31); 7424(17). Jenman 97(6). Jimenez 599(29); 872(1); 1022(32); 3146

(5). Johnson, H. 960, 961(20b); 962, 964(29);

966, 967(20b). Johnston, J. R. 143(34). Jones, G.N. 10995(17). Jones & Norris 1026(31); 1078(6); 1188

(17). Juzepczuk 6595(29). Kalbreyer 608(36c). Kanehira 120(20a). Karsten 7(29); 28(2); 127, 135(20a); 196

(36a). Kenoyer 5(9). Killip 2802(9); 5096(20a); 5254(14); 5257

(13); 5684(20a); 5701(1); 11364(36d); 11386(39); 11392(20a); 11404(36b); 12142(9); 35192(13).

Killip & Garcia 33173, 33624(9); 33711 (20a).

Killip & Hazen 9496(36b). Killip & Lasser 37747(15).



96 ROLLA TRYON

Killip & Smith 15088, 15338(33); 15861 (36d); 15886(36b), 17821(36b); 19277 (2); 19835, 19857, 19861(36a); 19962 (29); 20100(36d); 20407(20a); 23979, 23980(1); 23995, 24014, 24572(2); 29379(1).

Kuhlman 1792(33). Lavraste 1899(6). Lechler 2160(33); 2309(37); 2569(27);

2654(8). Lehmann BT890(20a). Lellinger 354, 642(17). Lemos 18911(33). Lent 459(33). León 11154, 11176(31); 11181(36c);

11900, 12296(17), 12721(17). León & Clément 4001, 8097(17). León et al. 9865(6); 9926, 10070(17);

10309(6); 10535(31); 13919(17). Leonard 4670, 5355(31); 8227(6); 8377

(31); 9308(2); 9350(6); 9365(17). Leonard & Leonard 12316(6); 14256(17);

14535(6); 14595, 15165(31); 15707(17). Leprieur 265, 265a(2). Levis & Maurel 3834(6). Levy 468(1). L'Herminier 3(17); 62(34). Linden 1022(36a). Lindig 115(36b); 196, 242(36a); 248(36b);

282(20a); 284(36a); 285(36b); 286, 287, 288(36a); 289(21); 306(36a); 308 (29); 309(20a); 310(1); 355(2).

Little 7332(36d); 8040, 8044(20a); 8120, 8497(36a); 8535(20a); 8643, 8847(21); 8905(39); 8906, 9083(36d); 9391(21); 13071, 13424(17); 21702(31); 21704(2); 21708(6).

Lloyd 170(34); 333(17); 391(34); 653(17). Lockwood 706(29); 800(21). Lopez et al. 4312(36a). Luetzelburg 534, 12841(33). Macbride 4135(26); 4819(20a); 4869(33). Macedo 2161, 2173, 2825(33). Madison 618(29); 779(1); 802(36a); 803

(38); 804(36a); 805(1); 821(36d); 861 (21); 888(36d); 892(36a); 913(1); 933, 1076(36a).

Maguire 33083(7); 33530(36b). Maguire & Maguire 35194(22). Maguire & Politi 28557(37). Maguire et al. 37261(7); 42369, 42370

(37); 42472(7). Martin 3395(15). Martinez 1, 26(29). Mathews 1828(33). Maxon 225(29); 318(36e); 351(20a); 417

(29); 524(36e); 569(29); 137l(30); 1913 (6); 2449(17); 2643(31), 2698(31); 2863, 3906, 3909(17); 3910(6); 3918(17); 4077, 4132(6); 4645(9); 5046(29); 5199

(20a); 5505, 5508(29); 5555, 5691(20a); 5769(9); 5776(1); 6876(9); 7070(31); 8757, 8886(17); 8968(45); 9216, 9260, 9410, 9499, 9516, 9586, 9694(31); 9866, 9889(30); 10143(31).

Maxon & Harvey 8090(38). Maxon & Killip 441(6); 483(17); 660(31);

944(38); 1146(31); 1449(17). McAlpin 2192(20a); 2222, 2230, 2257(1);

2259(29); 2263(36e); 2360(1). McConnell & Queich 605(33). McDaniel 6888(33). Mello Filho 961(33). Meyer, T. 6278A(33). Mexia 4582(33); 7138(1); 7600(21). Mickel 2265(29); 3228(36e). Mille 171(21). Miller 407(20b). Molina 1964, 2013(1); 10255(29). Molina et al. 17154, 17164, 17240, 17440,

17968(1); 17990, 17994(33). Moore, H. E. 3484(29). Moore et al. 9829(7). Morton 5193(17); 5492, 6213(34); 9273,

9397(31); 9446(6). Morton & Acuña 3047, 3487(6). Morton & Alain 8971(6). Moritz 177(36c); 393(39); 394(36c). Münch 24(20b); 76(29). Mutis 3159, 3173, 3291, 3294(1). Nash 470(17). Nash & Taylor 1744(31). Nevling & Gomez-Pompa 2443(49). Nicolson 2177(34). Nisman 9(1); 10(32); ll(36e); 29, 34(33);

36, 37(1); 43(20a); 44, 45(1); 61(29); 78(33); 88(1); 99(38); 100, 102, 103 (32); 104(48); 105, 107, 108(36e); 110 (33); 112(1); 113(29); 115, 119, 121, 122(33); 124(20a); 133, 135(33); 138 (20a); 139(1); 140(20a); 144(33); 145, 148(1); 150(29); 156(20a); 160, 161, 162(29); 168(20a); 169(29); 170, 171 (32); 172, 175(1); 177(36e).

Orcutt 4648(17); 5158(31); 5786(17). Otto 671(15). Palmer, C. W. 76(29). Pennell 4408(1); 4455(20a); 7118, 7503

(36a); 9311(20a); 9695(25). Pennell & Killip 5701, 7230, 8058(1). Phelps & Hitchcock 484(37). Philipson 2395(33). Philipson et al. 2157(18). Pittier 218(15); 4412(9); 6016(29); 6820

(9); 7367, 8070(15); 9359(29); 10011 (36c); 10981, 10992(1); 11853(15); 12027(1); 12355(5); 13728(29); 13740, 13741(15); 13946(29); 14160(20a); 14- 997(15); 16228(5).

Poeppig 218(33); 219, 1152(20a).



BEVISION OF CYATHEA 97

Pohl 663(33). Pollard et al. 45(17); 255(6). Prance et al. 4470(2); 4499(33). Pringle 6088(20b). Proctor 3914(17); 3996(31); 4000(46);

4001(45); 4360(31); 4378(30); 4460(6); 4583(38); 4607, 4889(17); 5060, 5513 (6); 5711(38); 5819(35); 11453(31); 11- 544(6); 16503(36c); 16995(34); 17270 (17); 17781(34); 17829, 19169, 19481 (17); 20134(34); 20135, 21785(17); 22547(36c).

Proctor et al. 26954, 27102(1). Purpus 438, 3809, I5312a(20b). Quesada 3(2). Riba 219(31). Riba et al. 260(29). Rimbach 67(23). Rohr 307(33). Rojas 3010(33). Rose & Rose 22508, 22605(21). Rose & Russell 20598, 20643(33). Rose et al. 4391(17). Ruiz 72(26). Ruiz, A. & A. Smith 209(6); 1538(29). Sampaio 4817, 4818(33). Sargent 412(6). Sauvalle 31(17). Scamman 5880(1); 5884(20a); 5885(36e);

6105(1); 6517(17); 6998(1); 7004(38); 7005(20a); 7006, 7007, 7584, 7860(29); 7862(20a); 8116(17); 8142(17).

Scamman & Holdridge 6997, 7436(1); 7861(29); 2864A(32); 7864®, 7867, 7868, 7869(36e); 7870(32); 7880(1).

Schipp S774, S923(l). Schmalz 200(33). Schmitt 128(5); 129, 130, 131(4). Schnee 1487(2). Schnell 1045(1). Schomburgk 1124(33). Sohultes 5857(7). Schultes & Cabrera 12469, 13407, 15079,

16330(7). Schultes & Lopez 9497(7). Schultes & Villarreal 5185(36a). Schultes et al. 24221(7); 24423(7). Schunke 53(2); 117(1). Scolnik et al. 19An214(29). Seemann 990(12); 1120(1). Segadas-Viana 2058(33). Sehnem 4213(33). Seifriz 68(2). Senn 386(17). Sermolli 182(15). Shafer 175, 343(17); 1713, 3059(6); 3518

(17); 4162, 4217(6); 4379, 7998(17); 8066, 8333(6); 8957(17).

Sharp 45960(29). Sherring 51(34).

Sieber 194(17); 374(34). Sintenis 409, 1375, 1788, 2415(17); 4102

(2); 4242(17); 5960(6); 5961(17); 6076 (6); 6086(17); 6156(2).

Skutch 743, 1162, 1698(20b); 2161(1); 2964(20a).

Smith, A. C. 2873(2). Smith, C. E. et al. 4140(29). Smith, G. W. & H. H. Smith 1349(17). Smith, H. H. 1017(2); 1020(36a); 1124

(15); 2225(20a); 2227, 2641(2). Smith, H. H. & G. W. Smith 292(34). Smith, J. D. 1567(1). Smith, L. B. 1593(33). Smith, S. G. & Idrobo 1539(33). Sneidern 2189(36a); 5365(20a). Soejarto 1451(33); 1562, 1568(36a); 1569

(19); 2050(36a). Sota, de La 5053(36e); 5263(33). Soukup 1052(2); 2335, 5017(36a); 5223

(33); 5260b(l). Sparre 14049, 16310, 16312, 16313; 16-

488(36a); 16753, 16946(21); 18482(1); 18738(36a); 19099(1).

Spruce 3127(7); 4723(37); 4729(33); 53- 67(21); 5741(1); 5743(36d).

Standley 27496(9); 53132, 53945(1); 85- 050(29); 90427(20b); 91674(1).

Standley & Torres 51288(20a). Standley & Valerio 47189(1). Stehlé 3309, 6081(17). Steinbach 5310(33); 8572(24); 9046, 9425,

9449(36a); 9512(24). Stern & Chambers 188(12). Stern et al. 1135(20a); 1755(1). Stevens 5950(2). Stewart 894, 895, 896, 897(3). Steyermark 29801, 30057, 32464, 32466,

36394(20b); 39965(1); 42791, 43498 (29); 48784, 49787, 51677(20b); 56- 026(36a); 56432(36c); 58395(37); 59- 588, 59811(20a); 61970(15); 62467, 74842(29); 74905(7); 74989, 75444(37); 75451(7); 75472(29); 89750, 92132(15); 93787(7); 93918(20a); 93922(7); 94017 (37); 97872(7); 98019(37); 98097(20a); 98224(33); 98914(29); 104864(36b); 105183(7); 105194(11); 105973(20a).

Steyermark & Allen 17141(1). Steyermark & Bunting 103084(7). Steyermark & Nilsson 122(7). Steyermark & Rabe 96085(33); 96956(36a). Steyermark & Steyermark 95396(29). Steyermark & Wurdack 717(20a); 934(7);

1328(29). Steyermark et al. 92504(7); 92701(20a);

100748(36a); 100967(36b); 103580 (20a); 105693(36d); 109074(7); 109268, 109395, 109396, 109556(20a).

Stimson 1272(17); 1273(6).



98 ROLLA TRYON

Stolm 209(6). Stone 2447(29); 2752(33). Stone & Stone 2700(20a); 2719(1). Stork 1040, 1785, 2330(20a). Stübel 1259(19). Sydow 299(1). Tate 112(15); 428(20a); 668, 865(37). Taylor, N. 225(17); 510(6). Taylor, Wm. R. 1374(1). Tessmann 3025(1). Ton 892(29). Tonduz 10103(17); 10701(32); 11787,

11789, 11802, 12183(20a). Torres 53(20a). Tovar 4201(36a). Traill 1382(1). Tryon & Tryon 5219(33); 5326, 5773,

6009(36a); 6113(21); 6658, 6659, 6661, 6671(33); 6962(17); 6913(33); 6982, 6983(31); 6988, 6989(29); 7022(36e); 7023(1); 7025(36e); 7083(6); 7123 (29); 7124(29).

Tschudi 147(15). Türckheim 1238(20b); II 1629(29); II

1645, II 2031(20b); 2716(17); 3056 (31).

Tyson & Blum 1647(9). Tyson et al. 3264 (10). Ugent & Ugent 5111(21). Ulloa 5(33); 102, 103(20a). Underwood 49(17); 11433, 1527, 1530,

1535(31); 1536(38); 1550(45); 1623, 2102(6); 2383(31); 2502(30); 2633, 2634(31); 2858(17); 2889(6); 3143, 3150, 3210, 3216, 3219(31); 3398, 3401(6).

Underwood & Earle 609(17); 714(6); 715(17); 730(6); 731(17); 1173, 1184, 1367(6).

Underwood & Griggs 287(17). Valeur 317(6); 396(17); 566(2). Vareschi 3183(2); 4782(37); 5819(33);

6458, 6462(29); 7492(36a). Vareschi & Foldats 4869(37). Vareschi & Pannier 621(29). Vareschi & Vareschi 2822(36b). Vargas 11754(1); 11858(27); 16069(1);

16206, 16274, 16544(1); 17536(2); 19- 857(21).

Wacket 224(33). Watt 184(38); 196(31). Weber, C. 6235(31). Webster, R. 1850(17). Wedel 2858(1). Welch 19613(1). Wercklé 41(29); 45(20a); 52(29). Wetmore Se Woodworth 113, 114(9). White, R. 1972067(48). White & Lucansky 196823(33); -27, -28,

-29, -30, -31, -32, -55, -74(20a); -77, -78, -79, -80(33); -96(20a); -101, -102, -103(29); -110(1); -114(29); -124, -138, -156, -158(20a); -163, -164(33); -165; -166; -167(20a); -181 (1); -182, -184, -191(33); 1969237 (29); -243(36c); 1970140(36c).

White & White 197031(37). White, et al. 1972067(48). Wiggins lO999(30b); 18561(3). Wilbur 8055, 8288(17). Wilbur & Stone 9972(32); 10109(29);

10623(36e). Wilbur et al. 11077(20a); 11121(33). Williams, LI. 10723(15); 12250(29);

14328(7). Williams, R. S. 1272(1); 1333, 1334(33). Williams, L. O. & Molina 11093(29). Williams, L. O. et al. 24260(1); 24972,

25670, 27674, 27683(29); 28434, 28658(1).

Wilson, K. A. 524(31). Wilson & Murray 560(6); 638(31). Wilson & Webster 482(31); 518(17). Wilson, N. 332(17); 513, 520(42); 681

(45); 731(41). Wood & Atchison 7472(17). Woodson et al. 922(20a). Wright, C. 889(6); 891, 950(47); 951(6). Wurdack 1653(36a); 34115(20a); 34257

(37); 34285(20a). Wydler 444(17). Xolocotzi & Sharp X472, X1163(20b). York 194(31). Yuncker 4745(1); 17532(31).



Documents

A REVISION OF THE GENUS CYATHEA