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Systematics of Cltbadium L. (, )

Arriagada, Jorge Eugenio, Ph.D.

The Ohio State University, 1994

UMI 300 N. ZeebRd. Ann Arbor, MI 48106

SYSTEMATICS OF L. (ASTERACEAE, HELIANTHEAE)

DISSERTATION

Presented in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy in the Graduate School of The Ohio State University

By

Jorge E . Arriagada ******

The Ohio State University 1994

Dissertation Committee Approved by

Ralph E. J. Boerner Daniel J. Crawford Tod F. Stuessy Advisor^ Department of Plkfrt Biology Dedicated to my wife Violeta and my daughters Paula and Andrea

ii ACKNOWLEDGMENTS

Numerous people must be acknowledged for their assistance in helping me complete this dissertation. First and foremost I would like to thank my advisor, Professor Tod

F. Stuessy, for his invaluable help and guidance, constant encouragement and continuous support throught all phases of my graduate study. Appreciation is also owed to Professor Daniel Crawford for his suggestions on the final version of this thesis and for his interest in and support of all aspects of my professional development, and to Professor Ralph Boerner, chairman of the Department of Biology, for his advice and critical review of the manuscript.

Many institutions have been instrumental in helping make my dissertation successful. Financial support for field trips was provided by the National Science Foundation (INT-

9017 695), The Garden Club of America, The Tinker Foundation, The Organization of American States, The Department of Plant

Biology, OSU, and The Ohio State University Janice C. Beatley Herbarium Award.

Numerous friends and colleges have provided help and assistance during my field trips. Special thanks go to Peter Dobbeler for sharing with me his expertise on Costa Rican vegetation. I thank Santiago Diaz-Piedrahita, an outstanding Colombian synantherologist, who shared with me his botanical and historical expertise on Colombian Compositae. I also thank Jaime Aguirre, Primo Chavarria, Jorge Gomez Laurito, Hazel Hernandez, Luis J. Poveda, Jorge Rojas, Pablo Sanchez Vindas, Fausto Sarmiento, Cristina Velez-Nauer, and other people too numerous to mention. Special thanks go to Dr. Nicholas Hind, from the Royal Botanic Gardens, Kew, for his important comments and suggestions on my research, to Jose L. Panero, from MSC, for sharing with me important information concerning Andean Heliantheae, and to Javier Francisco-

Ortega, for his valuable comments on nomenclatural problems.

Discussions with other persons of the plant systematics program at The Ohio State University have also contributed to this study, including Karla Gengler, Paul 0. Lewis, Tao Sang, Jun Wen, and James Zech. Special thanks go to Dr. John Furlow (OS) for loan assistance. I also thank Dr. H. W. Lack for important information regarding types at B, and Hugo

Valdebenito and Fausto Sarmiento for obtaining valuable information about types at QPSL. I also thank the curators of the following herbaria from which specimens were borrowed for this study, and who provided, in some cases, space and valuable information during my visits: A, B, BM, CAS, COL, CR, DS, ECON, F, FSU, G, GH, K, LL, LP, MEDEL, MO, NY, P, UC,

US, WIS. Finally, I express sincere and special thanks to my wife, Violeta, for supporting me unconditionally during all the years spent as a graduate student.

iv VITA

November 26, 1949...... Santiago, .

1977 ...... Licenciado en Biologxa, Universidad de Concepcion, Concepcion,Chile.

1977-1988...... Instructor, Dept.of Botany, Universidad de Concepcion, Chile.

1987 ...... M.S., Universidad de Concepcion, Concepcion, Chile.

198 8 ...... Research Assistant, The Ohio State University Herbarium.

1989-1991...... Teaching Associate, The Ohio State University, , Ohio.

1991-199 2 ...... Lecturer, The Ohio State University, Columbus, Ohio.

1992-199 4 ...... Lecturer and Laboratory Instructor, The Ohio State University - Newark Campus.

v PUBLICATIONS

Stuessy, T.F. and J. Arriagada. 1989. Systematics of Desmanthodium (Compositae, Heliantheae). Amer. J. Bot. 76(6): 273. [Abstract].

Stuessy, T.F. and J. Arriagada. 1989. Systematics of Desmanthodium (Compositae, Heliantheae) . Ohio Acad.. Sci. 89(2): 7. [Abstract].

Arriagada, J.E. 1990. Chromosome studies in Clibadium (Compositae, Heliantheae). Ohio Acad. Sci. 90(2): 5. [Abstract].

Arriagada, J.E. Preliminary studies and infrageneric classification in the Clibadium (Compositae, Heliantheae). Ohio Acad. Sci. 90(2): 5. [Abstract].

Arriagada, J. and T.F. Stuessy. 1990. A new species and subgenus of Desmanthodium (Compositae, Heliantheae) from southern . Brittonia 42: 283-285.

Dxaz-Piedrahita, S. and J. Arriagada. 1992. Una nueva especie de Clibadium (Compositae, Heliantheae) para . Rev. Acad. Colomb. Cienc. 18: 301-304.

Stuessy, T.F. and J. Arriagada. 1993. Chromosome counts in Clibadium (Compositae, Heliantheae) from . Brittonia 45: 17 2-176.

Arriagada, J. 1993. Notas sobre las relaciones genericas de Clibadium (Compositae, Heliantheae). Rev. Acad. Colomb. Cienc. 18: 465-468.

vi Arriagada, J. 1994. Revision of the genus Clibadium (Compositae, Heliantheae). Ohio Acad. Sci. 94(2):2l. [Abstract] .

Arriagada, J. 1994. Systematics of Clibadium (Compositae, Heliantheae). Amer. J. Bot. 81: 140. [Abstract].

FIELD OF STUDY

Major field: Plant Biology Studies in Plant Systematics with Professor Tod F. Stuessy TABLE OF CONTENTS

DEDICATION...... ii ACKNOWLEDGEMENTS...... iii VITA...... v LIST OF TABLES...... X LIST OF FIGURES...... xi INTRODUCTION...... 1

TAXONOMIC HISTORY...... 5

GENERIC AFFINITIES ...... 12 CONCEPTS OF SPECIES AND VARIETIES...... 28 CHROMOSOME NUMBERS...... 31 HYBRIDIZATION...... 34 HABITAT AND DISTRIBUTION ...... 36 PLANT- INTERACTIONS...... 38 ETHNOBOTANY...... 42

PHYLOGENETIC ANALYSIS...... 48 MODES OF SPECIATION...... 58 MORPHOLOGY AND TAXONOMIC CRITERIA...... 64 ...... 7 6 EXCLUDED AND DOUBTFUL NAMES...... 235 APPENDICES

A. Numerical list of species...... 237 B. Index to numbered collections examined...... 239

viii C. Notas sobre las relaciones genericas de

Clibadium L. (Compositae,Heliantheae) ...... 260 D. Chromosome counts in Clibadium (Compositae, Heliantheae) from Latin America...... 265 E. Scatter diagram of representative collections of Clibadium grande. C. grandifolium, and C. pacificum...... 271

LIST OF REFERENCES...... 273

ix LIST OF TABLES

TABLE PAGE

1 . Characters and character states used in cladistic analysis of Clibadium and related genera. Characters 3, 4, 8, 12, 14, 15, 18, and 22 are ordered, but the rest of the multistate characters are unordered...... 19

2 . Character state matrix for phylogenetic reconstruction of Clibadium and related genera. See Table 1 for description of characters and states...... 22

3. Chromosome numbers of species of Clibadium.. 32 All u = 16.

4. Characters and character states used in cladistic analysis of species of Clibadium. Characters 4, 5, 7, 13, and 15 are ordered; the rest of multistate characters are unordered...... 50

Character state matrix for phylogenetic reconstruction of species of Clibadium. See Table 4 for description of characters and states...... 51

x LIST OF FIGURES

FIGURE PAGE

1. Distribution of Clibadium and related genera. Squares show disjunct distribution of Stachyceohalum...... 13

2. Strict consensus cladogram of Clibadium and related taxa...... 23

3. One of eight equally most parsimonious cladograms of Clibadium and related taxa generated by PAUP from 22 morphological characters. Indexes are: 0.787 (Consistency), 0.213 (Homoplasy), 0.730 (Retention). The solid, gray, and open bars represent synapomorphies, reversals, and parallelisms, respectively...... 24

4. Elevational ranges of species of Clibadium (based on herbarium data and personal observations) ...... 37

5. Strict consensus cladogram of taxa of Clibadium...... 54

6. One of twenty-five equally most parsimonious cladograms of taxa of Clibadium generated by

xi PAUP from 15 morphological characters. Indexes are 0.682 (Consistency), 0.320 (Homoplasy), 0.631 (Retention). The solid, gray, and open bars represent synapomorphies, reversals, and parallelims, respectively 55

7 . Flowering ranges of species of Clibadium (based on herbarium data and personal observations)...... 61

8. Three types of capitulescences in Clibadium. A, cymose, C. egaersii (from Stuessy, 1975, f. 50); B, thyrsoid, C. surinamense (from Stuessy, 1975, f . 49); C, paniculiform, C. glabrescens (from Aristeguieta, 1964, f. 62)...... 67

9. Distribution of Clibadium divaricatum. C. eggersii. and C . 1eptophyllum in Central, and northwest America...... 88

10. Distribution of Clibadium erosum and C. terebinthinaceum in the Caribbean...... 102

11. Distribution of Clibadium anceps. C . alomeratum. and C. trianae in Central and northwest ...... 112

12. Distribution of Clibadium microcephalum and C. sessile in and Panama, respectively...... 126

13. Distribution of Clibadium frontinoense and C. zaruchii in Ecuador...... 133

xii 14. Distribution of Clibadium rhytidophyllum and C. sprucei in Ecuador...... 138

15. Distribution of Clibadium cordatum and C. grandifolium in Colombia, and in Colombia, , Nicaragua, and Panama, respectively...... 146

16. Distribution of Clibadium glabrescens. C. manabiense. and C. oentaneuron in Colombia, Ecuador, and ...... 155

17. Distribution of Clibadium laxum and C. sodiroi in Ecuador...... 167

18. Distribution of Clibadium peruvianum and C. surinamense in , , and , and in the Caribbean, Central, and northern South America, respectively...... 177

19. Distribution of Clibadium arboreum and C. armanii in , Guatemala, Honduras, and Mexico, and in Brasil and , respectively...... 197

20. Distribution of Clibadium leiocarpum and C. sylvestre in the Caribbean, and northern South America...... 210

21. Distribution of Clibadium acuminatum and C. micranthum in Colombia, Panama, and Venezuela, and Bolivia and Peru, respectively...... 229

xiii 22. Scatter diagram of representative collections of Clibadium grande (black circles), C. arandifolium (hollow circles), and C. pacificum (hollow squares) comparing features previously believed to be diagnostic for separation of taxa. Numbers show number of ray florets in individuals heads (C. grande is constant with 6). Types of the three taxa indicated by "t." measurements were made on largest leaf; ray floret number on average from three heads...... 272

xiv SYSTEMATICS OF CLIBADIUM L.

(ASTERACEAE, HELIANTHEAE)

INTRODUCTION

The genus Clibadium L. is important systematically because of its broad geographical distribution throughout much of the Neotropics and for its many locally endemic species. The genus extends from southern Mexico throughout Central America, the Caribbean and Lesser Antilles, and northern South America, from sea level to 3,400 m, and with high concentrations of species in Colombia, Costa Rica, and

Ecuador. A few species [e.g., C. eggersii Hieron., C. surinamense L., and C. svlvestre (Aubl.) Baill.] have broad distributions from Honduras to Ecuador, but many are restricted to small areas (e.g., C. divaricatum S. F. Blake, prov. Tarapoto, Peru; C . rhytidophvllum Diels, Volcan Tungurahua, Ecuador; C. zaruchii H. Robinson, western slope of Cordillera Occidental, Colombia).

The affinities of Clibadium to other genera have never been examined closely. More recently the genus has been placed close to Desmanthodium Benth., Mart.,

Riencourtia Cass., and Stachycephalum Sch. Bip, by Stuessy (1977), and to Lantanopsis ex Griseb. and

1 by Robinson (1981). Clibadium is here defined by a unique combination of characteristics, most of which are individually present in Desmanthodium. Ichthyothere. Riencourtia. and Stachycephalum. but none of which has the same suite of diagnostic character states. Clibadium includes shrubs and small trees with usually loosely aggregated heads, herbaceous phyllaries arranged in 1-5 series, receptacle usually paleaceous throughout, ray corollas 2-4-lobed, disc corollas 4-5-lobed, purple to black anthers, and chromosome numbers all n = 16.

In addition to interesting patterns of distribution and complex generic relationships, Clibadium is also important in the Neotropical flora for several other reasons. First, the included species are shrubby, and in some cases woody, which makes them conspicuous and encountered often in many regions; floristic workers, therefore, frequently must deal with taxa of this group. Second, the drupe-like fruits are also used as food by birds (Feinsinger et. al. 2182) which indicates a significant role in wildlife resources and ecology. Third, one species of Clibadium (C. sylvestre) is used as a - poison by Indian tribes throughout Latin America, thus having an ethnobotanical role with medical potentials for the active compounds.

Despite the importance of Clibadium. basic systematic relationships of taxa within the genus have for a long time remained poorly understood to the extent that routine identification has been impossible. The only taxonomic treatment for the genus was done by Schulz (1912), which included 13 species and two varieties from the more than 30

species described by authors at that time. He divided the genus into two sections: Euclibadium DC. (characterized by having a naked receptacle and villous sterile ovaries) and Trixidium DC. (with paleaceous receptacle and pilose sterile ovaries). The studies of Schultz were based on morphological characters obtained from the study of limited herbarium material mostly deposited at Berlin-Dahlem. Although helpful, this work has not been satisfactory for revealing the basic relationships of the taxa involved. In addition, the valuable contributions by Blake (1917), Cuatrecasas (1954), Aristeguieta (1964), Stuessy (197 5), and Robinson (197 9 a,b) likewise have not provided a comprehensive treatment of the genus. Moreover, many new taxonomic names have been added based on characters that are extremely variable. Also, none of the previous contributions have ever assessed evolutionary relationships.

The present study is based on more than five years of work, including 3 months of field investigation and examination of more than 6,000 herbarium specimens. Also included is a cladistic study for helping resolve taxonomic limits and for developing hypotheses of evolutionary trends within the genus. Twenty-nine species are here recognized from more than 80 previously described. The two sections recognized by previous authors (Candolle, 1836; Schulz, 1912) are here considered as subgenera (as subg. Eggersii and subg. Clibadium) containing two and four sections, respectively. The major portion of this study is a taxonomic revision of the genus, including generic and specific synonymy, descriptions, distributions maps, representative specimens, and an artificial key to taxa. TAXONOMIC HISTORY

The genus Clibadium with the single species C. surinamense was published by Linnaeus in his Mantissa Plantarum (1771). Candolle (1836) described seven new species and one variety of Clibadium and divided these and other existing taxa into three sections based on pubescence and texture of achenes, number of series of female florets, and presence or absence of pales subtending disc florets. He placed Clibadium surinamense L. in Section Euclibadium because of drupe-like achenes, a single series of ray florets, and receptacles without pales. The single species, C. erosum. from the Caribbean, was placed in Section Trixidium based on dry ray achenes with villous apices, two series of ray florets, and epaleaceous receptacles. Candolle grouped the remaining species mostly from the Caribbean and northern South America, in Section Oswalda (Cass.) DC. because of dry and pubescent achenes, one series of ray florets, and epaleaceous receptacles. Oswalda had been named as a genus by (1817), based on Baillieria aspera Aubl.

Grisebach (1864), as a result of his study of the Flora of British West Indian Islands, described three new species of Clibadium (C. alexandri, C. badieri. and C. fraaiferum)

5 and subdivided the Caribbean taxa into two groups: Euclibadium DC. with only one series of female florets, and an unnamed group (clearly Candolle's Section Trixidium) containing species with two series of female florets. A few years later Hemsley (1881) listed fifteen herbaceous or half- shrubby species indigenous to America including C. acuminatum Benth. from Cocos Islands, C. asperum DC. from southern Mexico, Guatemala and Nicaragua, and C. leiocarpum Steez from Costa Rica.

A number of new species were added to Clibadium in the early 1900s. Based on material collected by A. Sodiro in subtropical and subandean regions of Ecuador, Hieronymus

(1900) described C. sodiroi and C. subsessilifolium. The number of species in the genus increased to 18 due to Greenman's (1903) descriptions of C . anceps. C . glomeratum. and C. pittieri based on material collected in Costa Rica by A. Tonduz.

Schulz (1912), in the only existing revision of the genus, recognized 19 species and two varieties. He accepted 15 previously described species and proposed four new ones: Clibadium armanii from Brasil, C. lehmannianum from Colombia,

C. micranthum from Peru, and C. remotiflorum from Brasil and Bolivia. Schulz also recognized two sections based on the presence of pales subtending disc florets and pubescence of sterile ovaries: Euclibadium DC. (including Candolle's section Oswalda) with 15 species, characterized by absence of pales subtending disc florets and villous sterile ovaries; and Trixidium with four species: C. eacrersii Hieron.; C. erosum (SW.) DC., C. fraaiferum. and C. terebinthjnaceum. characterized by paleaceous receptacles and sterile ovaries villous only at the apex.

Blake was a major contributor to the systematics of Clibadium. He (1917) agreed with Schulz's subdivision of the genus into two sections, and he added five new species: C. divaricatum and C. strigillosum from Peru; C. heterotrichum from Bolivia; C. polygynum from Nicaragua; and C. sprucei endemic to Volcan Tungurahua in Ecuador. He also transferred Desmanthodium trianae Hieron. to Clibadium. A few years later (1924), Blake described nine new taxa including three species from Costa Rica: C . grande. C. grandifolium. and C. schulzii from material collected by A. Tonduz. Clibadium parviceps also was described from material collected by A. Fendler in Venezuela, and Blake believed that the new taxon was related closely to C . acuminatum Benth., endemic to the Cocos islands. Blake (1926) added two new species, this time from Ecuador: C . laxum and C. microcephalum. The genus increased even further when Blake

(1937) described C. oligandrum based on material collected in Volcan , northern Guatemala, by A. F. Skutch in 1934. At the same time, he described C. qlabrescens from Colombia and C. leiocarpum var. strigosum from Costa Rica. One year later, Blake (1938) added another species to the list: C. mexiae from Ecuador. The last of the almost two dozen taxa described by Blake (1941) was C. sessile from the province of Chirigui, Panama.

Cuatrecasas (1954) added nine new species and one subspecies in Clibadium for the Flora of Colombia with interesting comments on possible relationships between and among closely related species. All the type specimens and additional material were collected by Cuatrecasas throughout the different departments along the Eastern and Western Cordillera of Colombia.

Additional comments were made on Clibadium by floristic workers in different regions of latin America. The distributional ranges of some species of Clibadium were reported as broader when Matuda (1950), in his Flora of Chiapas. listed C. arboreum. C. oligandrum and C. villosum growing near Volcan Tacana in southern Mexico. Clibadium heterotrichum, originally restricted to Bolivia, was reported by Cabrera (1962) as growing in Argentina. Aristeguieta (1964) mentioned three species in Venezuela together with one variety described by Steyermark, and he proposed a new species for that country: C. pediculatum. Nash (197 6), in the Flora of Guatemala, mentioned that more than 25 species of Clibadium have been recognized, from which two were cited in Guatemala (C. arboreum. C. eggersii) and a third one from

Chiapas, Mexico (C. leiocarpum). In recent years Robinson made several important specific additions to Clibadium. He (1979a) described a new species for Peru: C . varaasianum. based on fragments lacking the erect hispid pubescence typical of C . surinamense. More recently, however, Robinson (1992, p. 149) reconsidered C. varaasianum as "an unusually hirsute variant of the widespread C . surinamense." Robinson (197 9b) also described C. harlinaii and C. manabiense from Ecuador, based on material obtained by G. Harling. In 1988, Robinson described

C. rimachii based on a specimen collected by M. Rimachi (197 8) from Peru, but he later (1992) relegated it to synonymy of C. divaricatum Blake. Clibadium zarucchii from the region of Frontino, Colombia, was described by Robinson (1988) as a new species based on the most massive congestion of heads of any member of the genus. Diaz-Piedrahita and Arriagada (1992) described C. frontinoense from the same general region of C. zaruchii. a clearly related taxon also with massive head aggregations. The most recent publication by Robinson (1992) added four new species in Clibadium: C. funkiae from Antioquia, Colombia; and C. alatum, C. napoense. and C. zakii from Ecuador. This brought to 60 the total known species in the genus.

Since the synopsis of Clibadium by Schulz (1912), there has been no comprehensive revision of the genus. A large number of names have accumulated, resulting in more than 100 specific and infraspecific epithets within the genus. In the present treatment Clibadium is recognized to contain 29 species in two subgenera: subg. Paleata with two sections, and 5 species; and subg. Clibadium with four sections and 24 species. The subgeneric and sectional limits are supported by phylogenetic analyses (see details below), which also provide rationale for sequence of species. The classification presented in this treatment is as follows:

Clibadium L.

Subgenus Paleata Arriagada

Sect. Eggersia Arriagada 1. C. eggersii Hieron. 2. C. leotophvllum Cuatrec. 3. C. divaricatum S . F . Blake

Sect. Trixidium DC.

4. C. terebinthinaceum (Sw.) DC. 5. C. erosum (Sw.) DC.

Subgenus Clibadium

Sect. Glomerata Arriagada 6. C . anceps Greenm. 7. C. glomeratum Greenm. 8. C. trianae S. F. Blake 9. C. sessile S. F. Blake 10. C. microcephalum S. F. Blake 11. <2 frontinoense Diaz & Arriagada 12. C. zaruchii H. Robinson

13. C . rhytidoohyllum Diels 14. C. sprucei S. F. Blake

Sect. Grandifolia Arriagada

15. C. grandifolium S. F. Blake 16. C . cordatum Cuatrec.

17. C. manabiense H . Robinson 18. C. glabrescens S. F. Blake 19. C . pentaneuron S. F. Blake

Sect. Clibadium

20. C . laxum S. F. Blake

21. C. sodiroi Hieron. 22. C. surinamense L.

23. C. oeruvianum Poepp. 24. C . arboreum Donn. Sm.

25. C. armani (Balbis) Sch. Bip.

Sect. Qswalda (Cass.) DC. 26. C. leiocarpum Steetz

27. C. svlvestre (Aubl.) Baill. 28. C . acuminatum Benth. 29. C. micranthum 0. E. Schulz GENERIC AFFINITIES

Clibadium appears to have had a long evolutionary history. Based on the large number of species and on its overall broad range, it has diversified to a greater extent than any of the other closely related genera (Fig. 1). Clibadium has been suggested (Stuessy, 1977) as being related to Desmanthodium Benth., a small genus of eight species in Mexico and Central America, Ichthyothere Mart., with about seventeen species in Brasil, Colombia, Paraguay and Venezuela, and Stachycephalum Sch. Bip., consisting of two disjunct species in Argentina and Mexico. The genus was placed in an informal "Group 3" in subtribe Milleriinae (Stuessy, 1977). Robinson (1981), however, in a new subtribal classification of the Heliantheae, referred Clibadium to a new subtribe, Clibadiinae, Ichthyothere to the Melampodiinae, and Desmanthodium and Stachycephalum as the only members of the new subtribe, Desmanthodiinae. In the present study Clibadium is retained in subtribe Milleriinae with Desmanthodium. Ichthyothere. Riencourtia. and Stachycephalum as close relatives. The following discussion elaborates on this perspective.

The relationships among Clibadium. Desmanthodium. Ichthyothere. and Stachycephalum have been based on scattered

12 Fig. 1. Distribution of Clibadium and related genera. Squares show disjunct distribution of Stachycephalum. information from morphology, anatomy, chemistry and cytology. As for morphology, the adjudged close relationship between Clibadium and Ichthyothere is based primarily on

tubular ray corollas and relatively moderate aggregation of heads (Blake 1917; Stuessy, 1973). Stachycephalum. although

overall more distant morphologically from Clibadium. has radiate corollas and shows a looser aggregation of heads similar to those present in some species of Section Clibadium. Desmanthodium has a peculiar sac-like completely enclosing the ovary; this unique feature represents a definite morphological gap with the other three genera. But the general morphology of Desmanthodium. and particularly arrangement and aggregation of heads, shows close resemblance with some species of Section Glomerata of Clibadium.

Comparative anatomical studies of fruits of Clibadium. Desmanthodium and Ichthyothere show a strong similarity in the arrangement of the epidermis, hypodermis, phytomelanin and fiber layers (Stuessy and Liu, 1984). The similar pattern of the external phytomelanin surface in Clibadium and Ichthyothere argues for their close relationship in contrast to the smooth surface in Desmanthodium. The structure and development of the pericarp is similar and suggests evolutionary ties among Clibadium. Desmanthodium. Ichthyothere (Stuessy & Liu, 1983) as well as, somewhat surprisingly, with Galinsoga (Saenz, 1981) of subtribe

Galinsoginae. Based on general pericarp morphology and 15 features of phytomelanin, it is possible to suggest a relationship between Desmanthodium and Galinsoga. Both genera show a discontinuous phytomelanin layer associated with fiber bundles and a hypodermis with 1-2 layers of parenchyma cells. A relationship has been suggested previously between Desmanthodium and Galinsoga (Robinson,

1981) based on color and arrangement of resin ducts in the pericarp.

Flavonoid data indicate that Clibadium has a more simple biochemical profile than Desmanthodium in that the latter accumulates more mono-, di-, and tri-methyl derivatives (Bohm and Stuessy, 1981). Desmanthodium relates in flavonoid features to Clibadium somewhat more distantly than the latter does to Ichthyothere. The isolation of a fish poison compound (= tetrahydropyrane ichthyothereol) from some species of Clibadium as well as from Ichthyothere also suggests a close tie between the two genera (Czerson et al., 1979).

Chromosome counts give support to the concept of a closer relationship of Clibadium with Ichthyothere rather than with Desmanthodium. Clibadium is known as n = 16

(Turner and King, 1964; Coleman, 1968, 1982; Powell and Cuatrecasas, 197 0; Stuessy & Arriagada, 1993) and Desmanthodium as n = 18 (Keil and Stuessy, 1977; Robinson et al,, 1981) and n = 17 (Ralston et al. 1989). These numbers could be viewed as aneuploid increases from a base of x = 16 in Clibadium. Ichthyothere is Q = 16 and ca. 33 (Coleman, 197 0; Turner et al., 1979), also on an apparent base of x = 16 (it could, of course, have an ancestral base of x = 8).

Recent studies on Heliantheae based on chloroplast DNA restriction site analyses (Kim et al., 1989) indicate a tentative close relationship between Desmanthodium and Galinsoga. Although the generic sample is small and the results preliminary, the general difficulties of locating a satisfactory outgroup to Clibadium and its close relatives suggest that this hypothesis deserves attention. Because of the morphological conection between Desmanthodium and Clibadium. one could also assume that a reasonably close molecular relationship might exist between Clibadium and Galinsoga. In the most recent revision of Galinsoga (Canne, 1977), the genus was related with Cymophora Robin., Cass., Tricarpha Longpre and L., and not at all with

Clibadium and relatives. Nonetheless, in view of the preliminary molecular data, an examination of their possible ties to Clibadium seems in order.

Galinsoga R. & P. contains 14 species separated in three sections and distributed mainly in subtropical regions of Mexico, Central and South America (Canne, 1977). Section Galinsoga. and particularly one of the four lines of divergence within the section (including G. subdiscoidea Cronquist, G. mandonii Sch. Bip., and G. parviflora Cav.), might possibly have a with Clibadium. The species in this group all have very small ray corollas. Individuals within these taxa occasionally have reduced ligules or even lacking altogether. This particular line of divergence in Galinsoga apparently originated in Mexico (Canne, 1977), but G. mandonii is confined to the Andes of western South America. Chromosome numbers support a possible link between the two genera. All species of Clibadium surveyed are n = 16 (Stuessy & Arriagada, 1993). This number is also frequent among species of Galinsoga where n = 8, 16, 24 and 32 have been reported (Canne, 1983). Unsupportive of this relationship, however, is the habit in which all members of Clibadium are shrubs, and those of Galinsoga are herbs.

In my opinion, morphological features suggest that Galinsoga could be viewed as a somewhat distant ancestor for the entire subtribe Milleriinae. The ray florets arranged in 1-3 series could relate Galinsoga with section Eggersia of subgenus Paleata. but the presence of ligules in the former is a distinct difference. Another differentiation is hermaphroditic and fertile disc florets in Galinsoga and disc florets sterile (by abortion of ovaries; hence, functionally male) in all species of Clibadium.

Another taxon possibly considered as a close relative of Clibadium and other members of informal group 3 of Milleriinae (Stuessy 1977), might be Calea L. subgenus Calea

(subtribe Galinsoginae). Species in this subgenus resemble those of Clibadium in having numerous small heads arranged in close thyrses or modified umbels, and often with a reduced pappus. Most of the species of subgenus Calea are shrubs

(some are perennial herbs) and are distributed in Mexico and tropical South America, with some overlapping with Clibadium. The involucre with many series of phyllaries in Calea contrasts with the single series observed in all but one section (Sect. Glomerata) of Subgenus Clibadium. Chromosome numbers reported for Calea are n = 9 and 16. These cytological data give some additional support to the idea of affinities between Calea and Clibadium (also n = 16; Stuessy & Arriagada, 1993) .

In view of the previous discussion, and to provide a more definitive assessment of intergeneric relationships with Clibadium. a morphological cladistic analysis has been undertaken. Cladistic analysis is an approach that offers an explicit, repeatable and relatively objective methodology for reconstructing the branching patterns of phylogeny (Funk & Stuessy, 197 8; Duncan, 1980; Duncan & Stuessy, 1984; Stuessy, 1990; Quicke, 1993). One of the advantages of these methods is that the assumptions and steps involved in constructing a cladogram are clearly stated.

Materials and methods. Twenty-two characters and character states (Table l) were chosen, examined, and coded based on herbarium specimens. All characters were equally weighted in the cladistic analysis. Character states were polarized via outgroup analyses (Watrous & Wheeler, 1981; 19

Table l. Characters and character states used in the cladistic analysis of Clibadium and related genera.

Characters 3, 4, 8, 12, 14, 15, 18, and 22 are ordered, but the rest of the multistate characters are unordered.

DURATION.

1, annual (1); perennial (0). HABIT.

2, herbs (1); shrubs or trees (0). INVOLUCRAL .

2, Series: 4 (0); 3 (1); 2 (2); 1 (3). 4, Number: 5 or more (0); 4 (1); 3 or fewer (2). 2, Condition: herbaceous (0); membraneous (1). PALES.

£, presence (1); absence (0). 7, enclosure of achenes: yes (1); no (0). RAY FLORETS.

2, Number: 10 or more (0); 7 (1); 5(2); 1(3). 2, Color: yellow (0) ; white (1) . 10. Symmetry: bilateral (0); regular (1). 11. Ligule: present (0); absent (1). 12. Ligule condition: entire (0); 2-lobed (1); 3-lobed (2).

13. Pappus: present (0); absent (1). 14. Corolla (aligulate) lobes: 0 (0); 3 (1); 4 (2); 5 (3). 15. Achene shape: pyramidal (0); obovoid (1); fusiform (2). 16. Achene vesture: pubescent (0); glabrous (1). 20

Table l. Continued.

17, Achene surface: striated (0); smooth (1). DISC FLORETS. IB., Number: 75 (0); 45 (1); 25 (2); 15 (3); 1-8 (4) 19. Sterility: fertile (0); sterile (1). 20. Color: yellow (0); white (1). 21, Corolla lobes: 5 (0); 4 (1).

22, Anther bases: sagittate (0); subsagittate (l);obtuse (2); auriculate (3). Maddison et al., 1984; Nixon & Carpenter, 1993). Galinsoga and Calea were chosen as the most appropriate outgroup, as discussed previously. The data matrix (Table 2) was input with MacClade 3.0 (Maddison and Maddison, 1992) and the cladistic analysis was conducted using Swofford's (1993) PAUP version 3.1 (upgraded 3.1.1.). The most parsimonious cladograms were searched using branch-and-bound algorithm (Hendy & Penny, 1982). Character state optimization, ACCTRAN, was used in tree description. All the analyses were undertaken using the following options SWAP+ALT followed by USETREE for all trees in memory and SWAP=GLOBAL. Tree lengths and consistencies were recorded and consensus trees were compiled using the CONTREE program (Swofford, 1985).

A consensus tree, being less resolved than any of the cladograms from which it is calculated, has less explanatory power (Farris, 1983) than any one of them. Strict consensus trees are often much collapsed and uniformative (Bremer, 1990). As a result, any one of the competing cladograms will have more information and may be a better choice as a phylogenetic hypothesis (Mickevich & Farris, 1981; , 1985; Carpenter, 1988). Because of this, one of the most parsimonious trees has been chosen here in addition to the strict consensus cladogram, to illustrate hypotheses of generic relationships.

Results and Discussion. The results of the phylogenetic analysis are shown in Figs. 2 and 3. They Table 2. Character state matrix for phylogenetic reconstruction of Clibadium and related genera. See Table 1 for description of characters and states.

CHARACTERS TAXA l 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22

Calea 0 0 0 0 0 0 0 1 0 0 0 1 0 0 . 0 0 0 2 0 0 0 0 Clibadium 0 0 2 2 0 0 0 0 1 1 1 0 1 2 1 0 1 3 1 1 1 1 De smanthod ium 0 0 3 2 1 0 1 3 0 1 1 0 1 1 2 1 0 4 1 0 0 2 Galinsoga 1 1 1 0 0 0 0 2 0 03000000000 0 0 Ichthyothere 0 0 3 2 0 0 0 3 1 1 1 0 1 3 1 1 0 1 1 1 0 1 Riencourtia 1 1 3 1 0 1 0 3 1 1 1 0 1 1 1 1 1 4 1 1 1 3 Stachycephalum 0 1 3 2 1 0 1 3 1 0 0 2 1 1 1 1 1 4 1 1 0 3 23

Calea

Galinsoga

Clibadium

Riencourtia

Desmanthodium

Stachycephalum

Ichthyothere

Fig. 2. Strict consensus cladogram of Clibadium and related taxa. 24

Ancestor

Calea

12(1)a m 18(2)- - Galinsoga

i s(i y 4(2) Clibadium 3(2)1

14(3) 17(0) 2 0 ( 1)' Ichthyothere

3(3)

Riencourtia 2 ( 1 ). 18(4) 22(3)i

(0) (0) (2) Desmanthodium 9(0), 2(0), '5(2),

f Stachycephalum (0)(0)(2)

Fig. 3. One of eight equally most parsimonious cladograms of Clibadium and related taxa generated by PAUP from 22 morphological characters. Indexes are: 0.787.(Consistency), 0.213 (Homoplasy), 0.730 (Retention). The solid, gray, and open bars represent synapomorphies, reversals, and parallelisms, respectively. correspond to the strict consensus cladogram and to one ofeight equally parsimonious cladograms (consistency index of 0.787, i.e., 21 % homoplasy). The rationale for selecting this one particular tree was based on mayority-rule (rather than strict) consensus. Clibadium and its relatives Desmanthodium. ichthyothere. Riencourtia. and Stachycephalum share several character states (i.e., absence of pappus and ligule, regular symmetry in ray florets, black anthers, abortive ovaries and small heads tending toward tight aggregations). The ligulate ray florets, fertile disc florets, pappus, and anthers shortly saggitate at base observed in Calea and Galinsoga serve as the starting point for directionality observed in Clibadium and close relatives. Perfect disc florets in Calea and Galinsoga represent the primitive condition compared with abortive ovaries (= functionally staminate) in disc florets in all species and close relatives. Also the presence of pappus in Calea and Galinsoga represents the ancestral condition apparently lost in all members of subtribe Milleriinae, informal group 3 (Stuessy, 1977). The fully paleaceous condition of Galinsoga has its counterpart only in Subgenus Paleata of Clibadium (pales in the other sections are reduced and not always present in all species). The same paleaceous condition is found in all species of Ichthyothere. but it is absent in Desmanthodium (which has been placed close to Galinsoga based on chloroplast DNA studies, Kim et al., 1989), Riencourtia. and Stachycephalum. The sagittate condition of the base of anthers in Calea and Galinsoga indicates directionality of 26 this character to subsagittate in Clibadium and Ichthyothere. obtuse in Desmanthodium. and auriculate in Riencourtia and Stachycephalum. respectively.

Clibadium appears in a more derived position with respect to ichthyothere in the consensus analysis (Fig. 2), but in a more primitive position in the one most parsimonious tree (Fig. 3). Ray and disc corollas with five lobes, large number of disc florets (30-60), and striated achenes are the primitive character states represented in Ichthyothere. Clibadium has ray corollas with three or four lobes (occasionally two lobes), Desmanthodium and Riencourtia have three lobes, and Stachycephalum has two. The large number of disc florets observed in Ichthyothere (30-60) is also observed in Calea (50) and in Galinsoga (30-150). There is a clear decrease of disc floret number in Clibadium (3-22),

Riencourtia (6-8), Desmanthodium (4-6), and Stachycephalum (1-4). Chromosome numbers can also be placed on the cladogram to determine their relationship with morphological data. Clibadium. Ichthyothere. and Riencourtia (Robinson et al., 1981) are n = 16, whereas Desmanthodium has been reported as n = 17 and 18 (Stuessy & Keil, 1977; Robinson et al., 1981; Ralston et al., 1989). No chromosome number is known for Stachycephalum.

Recently, cladistic analysis has been presented of interrelationships among genera of Heliantheae (Karis, 1993). Based on 141 characters, a strict consensus cladogram resulting from 144 equally parsimonious trees was obtained (with a very low consistency index of 0.17). These results shows Clibadium in a more derived position with respect to ichthyothere. which agrees with the strict consensus tree presented here. According to Karis (1993), Desmanthodium occupies a more primitive position close to Galinsoga. which is contrary to the more derived position near Stachycephalum suggested here. Karis (1994), following the cladogram presented by Karis (1993), rearranged many of the genera into subtribes within Heliantheae. But, several genera were left unassigned to subtribes, and among these are included Clibadium. Desmanthodium. Ichthyothere. Riencourtia. and

Stachycephalum. CONCEPTS OF SPECIES AND VARIETIES

Species are the basic units of plant classification. Despite this obvious importance, much debate has ensued and almost no agreement exists regarding precise definitions of species (e.g., Stuessy, 1990; King, 1993; Rieseberg & Brouillet, 1994). Major concepts include autapomorphic, biological, biosystematic, cladistic (phylogenetic sensu Nixon & Wheeler, 1990), cohesion, ecological, evolutionary, genetic, morphological, paleontological, phenetic, recognition, taxonomic, and unitary. In plant taxonomy, the biological and morphological concepts are the two most widely used both conceptually and in practice (Stuessy, 1990).

The biological species concept is defined as "groups of interbreeding natural populations that are reproductively isolated from other such groups" (Mayr, 1969:314; Mayr & Ashlock, 1991:26). This concept emphasizes the importance of gene flow and reproductive isolation of species. The morphological species concept can be defined as "the smallest groups that are consistently and persistently distinct, and distinguishable by ordinary means" (Cronquist (1988:71). The morphological species concept is used in this study. It is assumed that the morphological discontinuities observed within Clibadium represent genetic variations that help

28 29 maintain the natural integrity of each of the recognized units. This assumption implies that the morphological concept here adopted does not exclude the biological concept and, as stated by Grant (1981), the morphological differences used to distinguish species are by-products of partial or complete reproductive isolation.

In this treatment all 29 species of Clibadium are recognized consistently by at least several morphological characters. All individuals showing continuous variation are grouped within a species and separated based on discontinuities from other such units. Some species of Clibadium are quite distinct morphologically, such as C. armanii. C. erosum. C. divaricatum. C. eggersii. C. frontinoense. C. leptoohvllum. C. surinamense. and _C,t zaruchii. Other species, however, are more polymorphic and show a great deal of variation, which makes circumscription and identification more difficult, e.g., C. grandifolium. C. parviceos. C. oentaneuron, C, peruvi&nym, C- .sy ly .e.S_tx.e , and C. terebinthinaceum. Another polymorphic species, Clibadium surinamense. shows great variation among populations, but it is still relatively distinct once the limits of this variation are understood. The lengthy synonymy given for this species, in particular, is evidence of its polymorphic nature.

For maximally predictive classification it is useful to formally recognize patterns of morphological variation within species, if such exist. These patterns, called subspecies or varieties, can be defined as groups of conspecific populations that are morphologically distinguishable and geographically coherent (i.e., at least partially allopatric; Stuessy, 1990). In this treatment infraspecific categories have not been used mainly because there are no discrete patterns of morphological variation among populations that correlate with distinct geographical distributions. However, a number of infraspecific taxa (including formae) have been recognized by previous workers (e.g., Schulz, 1912; Blake, 1937; Steyermark, 1953; Cuatrecasas, 1954), but they are judged here to be unsubstantiated. Some infraspecific taxa recognized previously (e.g., C. surinamense var. macrophyllum) did relate to geographical patterns, but these no longer exist due to larger numbers of specimens that now bridge earlier distinctions. CHROMOSOME NUMBERS

Chromosome numbers have been counted for 19 species of Clibadium (Table 3). The remaining taxa have been not counted, primarily because they are restricted endemics or are known only from the type collection. A chromosome number of n = 16 for all species has been reported (Stuessy & Arriagada, 1993). The base number for Clibadium. therefore, would most probably be x = 16. This level could be tetraploid based on ancestral x = 8, but no evidence is available to support this hypothesis. Absence of polyploidy and uniformity of chromosome number (n = 16) suggests speciation within Clibadium has occurred entirely at the diploid level. The same chromosome number of n = 16 has also been reported for two close generic relatives of Clibadium. Riencourtia Cass. (R. tenuifolia Gardn.) is known as n = 16 (Robinson et al., 1981) and Ichthyothere Mart. n. terminalis (Sprengel) Blake] as n = 16 and ca. 33 (Coleman, 1970; Turner et al. 197 9) . The n = ca. 33 is also probably based on x = 16. Desmanthodium Benth., the other close relative, is known chromosomally in D. fruticosum as n, = 18 (Keil & Stuessy, 1977; Robinson et al., 1981) and n = 17 (Ralston et al.,

1989) .

31 32

Table 3. Chromosome numbers of species of Clibadium

TAXON REFERENCE

£. anceos Greenm. Solbrig et al. (1972);

Robinson et al. (1981). £. arboreum Donn. Sm. Solbrig et al. (1972); Stuessy & Arriagada (1993; as C . erosum) . £. armanii (Balbis) Sch. Bip. Coleman (1968); Robinson et al. (1981); Coleman (1982; as

C. rotundifolium). £. cordatum Cuatrec. Stuessy & Arriagada (1993). £. eggersii Hieron. Stuessy & Arriagada (1993). £. erosum (Sw.) DC. Powell & King (1969). £. glabrescens S. F. Blake Stuessy & Arriagada (1993). £. glomeratum Greenm. Robinson et al. (1981) . £. grandifolium Powell & Cuatrecasas (197 0; as C . terebinthinaceum

subsp. colombiense). £. laxum S. F. Blake Stuessy & Arriagada (1993); Strother & Panero (1994; as C. alatum). £. leiocarpum Steetz Robinson et al. (1981); Stuessy & Arriagada (1993). 33

Table 3. Continued.

TAXON REFERENCE

£. microcephalum S. F. Blake Robinson et al. (1981);

Stuessy & Arriagada (1993).

£. pentaneuron S. F. Blake Stuessy & Arriagada (1993). £. rhytidophyllum Diels Stuessy & Arriagada (1993).

Q. sodiroi Hieron. Stuessy & Arriagada (1993; as C. mexiae). £. sprinamense L. Turner & King (1964); Powell & King (1969; as C. villosum); Powell & Cuatrecasas (197 0); Solbrig et al. (197 2); Robinson et al. (1981); Stuessy & Arriagada (1993).

£. sylvestrg, (Aubl.) Baill. Stuessy & Arriagada (1993; as C. asperum). £. trianae (Hieron.) S. F. Blake Stuessy & Arriagada (1993). HYBRIDIZATION

Interspecific hybridization in Clibadium is rare, due perhaps in part to normal allopatric distributions of closely related species. One case of apparent parapatric overlap, however, has been observed between two species restricted to a few provinces in Ecuador. Clibadium sodiroi Hieron. is found in provinces of Napo-Pastaza and Tungurahua, and C. microcephalum Blake, occurs in Canar, Napo-Pastaza, and

Santiago Zamora. However, in and around Puyo (Prov. Napo- Pastaza) , a slight overlap occurs in the distribution of these two species.

In the Napo-Pastaza area, morphological intermediates and meiotic chromosomal irregularities (fragments) suggest the occurrence of hybridization between C. microphyllum and

C. sodiroi by secondary contact between the two taxa (Stuessy & Arriagada, 1993). The zone of contact may have been accentuated by roads and other human activities in the region. Because the few individuals suspected as hybrids between these two species show intermediate morphology, pollen viability of the two parents and putative hybrids was examined. Acetocarmine and lactophenol- cotton blue stains were used, and positive pollen grain coloration by each was assumed to indicate pollen viability (Heuser & Morrison,

34 1964; Stuessy, 1990). A minimum of 100 pollen grains per specimen was examined. Pollen samples were taken from herbarium material and from flower buds preserved in modified Carnoy's solution and kept refrigerated. Pollen stainability in Clibadium microcephalum (King 6567; Stuessy & Jansen 4982;

Stuessv & Nesom 5823) and C. sodiroi (S t u o s s y-&_J,ans_gn._4.9_7_£; Stuessy & Nesom 5817) was 60-80 percent. The pollen stainbility in the suspected hybrids (Stuessy et al. 4960. 4969: Stuessy & Nesom 5814) was 0-5 percent, which does suggest hybridization. The very low percentage of pollen stainbility in the putative hybrids may result, in part, from the distant taxonomic position of the putative parents. Clibadium microcephalum belongs to Section Glomerata and its closest relative is C. sessile from Panama and Colombia. Clibadium sodiroi. on the other hand, belongs to Section Clibadium and is closest to C. laxum. also from Ecuador. HABITAT AND DISTRIBUTION

Species of Clibadium grow in different tropical habitats from southern Mexico (C. arboreum) to northern Peru and Argentina (C. peruvianum) and from sea level (C. eggersii) to

3,400 m (C. sprucei) (Fig. 4). Clibadium is found mainly in habitats that have been altered by human activities; most of the species are found along roadsides in cut-over secondary growth forests. A few species, however, still remain on the edges of tropical cloud forests or along small river creeks with original vegetation. Modification of habitats has resulted in vegetational alterations that have allowed some species of Clibadium to become secondarily parapatric, such as appears with C. microcephalum and C. sodiroi. discussed earlier. Clibadium eggersii (Stuessv et al. 4974) and C. surinamense (Stuessy et al. 4977) also have been found growing together at one locality in Napo-Pastaza, Ecuador. No hybrids have been detected here, however, perhaps because the species are from different subgenera (Subgenus Paleata in the former and Subgenus Clibadium in the latter).

Closely related species of Clibadium have allopatric distributions. This tendency fits with Stuessy & Sundberg"s hypothesis (1992) that closely related taxa in Heliantheae with the same chromosome number will tend to have allopatric distributional ranges (See Modes of Speciation).

36 37

< 300 600 900 1200 1500 1800 2100 2400 2700 3000 3300 3600 Subg. Paleata Sect. Eggersii eggersii divaricatum leptophyllum Sect. Trixidium erosum wwfiMt terebinthinaceum Subg. Clibadium Sect. Glomerata anceps I" -v :.S3 glomeratum trianae sessile immwrnmmmmMiil microcephalum ^v.vVitmsw^v;-y •: -t:"; ■••• •■ ;t:< ;v ; >;•••,•;■;•;■;• i frontinoense zaruchii rhytidophyllum E sprucei Sect. Grandifolia grandifolium------f a g k a a a ------cordatum feaaiaml manabiense glabrescens esss^ e s s s i pentaneuron Sect. Clibadium laxum b w » » m < « w a f » » i iM(iiatmai sodiroi surinamense l-;;ia s s a ;avB;iv;aif—S ia a s a a peruvianum arboreum Lmaiiaama^MmaW iMiiaaaa armanii taa-awmmgamwHa^ad Sect. Oswalda leiocarpum sylvestre E5HH33E23 acuminatum micranthum l-m-aK-K«^«»;-ffla-aaaa-K-K3

Fig. 4. Elevational ranges of species of Clibadium (based on herbarium data and personal observations) . PLANT-INSECT INTERACTIONS

Almost all species of Clibadium occur in populations of small size, ranging from one to few individuals. Occasionally large populations are found, however, such as in C. laxum (Stuessy & Arriagada 12352) and C. sylvestre (Arriagada 165). These populations are usually separated from each another by 500 m to 5 or more kilometers (pers. observ.). Because of the small populational sizes in species of Clibadium and substantial spatial separation, interactions between and their pollinators are likely to be extremely important for maintaining genetic cohesion.

Based on the considerable distance separating reproductive individuals in Clibadium. it initially seemed plausible that self-pollination and self-compatibility were likely reproductive mechanisms in the genus. It is known, however, that a high percentage of trees and shrubs in tropical regions are self-incompatible (Bawa, 1974), and that most tropical woody plants are insect-pollinated (Bawa, 1974; Janzen, 1984). Also, many pollinators are not species- specific, and they are able to migrate and pollinate over long distances. Large bees and sphinx have been

38 39 reported as large long-distance fliers, and they can move pollen over hundreds to thousands of meters (Janzen, 1984).

Personal field observations in Colombia, Costa Rica, and Ecuador on species of Clibadium indicate that by the time the anthers of the disc florets are mature and pollen dehisces, most of the carpellate florets of the same head have already been fertilized and fruits have been formed. This condition has been detected (pers. observ.) in C. eggersii. C. glomeratum. c. grandifolium. C. laxum. C. leiocarpum. and C. sylvestre. and it would favor cross-fertilization.

Two main groups of pollinators visiting species of Clibadium may be recognized based on their feeding preferences. The first is that take nectar from florets. An interesting case is Urania sp., a diurnal observed (L. Poveda, pers. comm.) visiting flowers of C. leiocarpum (Arriagada 308). This species migrates from Mexico to Bolivia at an average speed of 20 km/h for a 12-h day (Janzen, 1984). Another case is Anartia fatima F. (P. Dobbeler, pers. comm.), a butterfly abundant in disturbed areas from Mexico to Panama, that has been observed visiting Clibadium glomeratum (Arriagada 153), C. surinamense (Arriagada 151) and C. sylvestre (Arriagada 165). The second group is insects that use pollen as a food resource, which is a more highly specialized feeding behaviour (Janzen, 1984) . Among these type of pollinators visiting species of Clibadium (pers. observ.) are: Lasioglossum sp. (A. Solis, pers. comm.) a small greenish black bee common near forest edges and a common visitor to Clibadium surinamense (Arriagada 130) and Polymnia sp. (Arriagada 131); Bombus sp. (A. Solis, pers. comm.), a common bumblebee at high altitudes (1800-3000 m ) , visits several Compositae (Cirsium. Senecio) including C. leiocarpum (Arriagada 136). Another frequent visitor of species of Clibadium is the introduced Apis mellifera L. A common visitor of some low elevation (0-600 m) Costa Rican species of Clibadium is Simulium sp., Diptera (E. Chacon, pers. comm.). Although individuals of this species have been observed with pollen grains adhering to hairs on body and legs (pers. observ.), no information is available regarding precise feeding habits.

There are other insects interacting with species of Clibadium but their role as pollinators is unknown. Occasionally, species of Culex and Aedes have been seen visiting C. surinamense (P. Doebbeler, pers. comm.). Also Pseudomyrmex sp. (E. Chacon, pers. comm.) is found on C. surinamense and C. sylvestre (pers. observ.). An interesting relationship exists between flea beetles (Chrysomelidae) and Clibadium. These extremely small (2 mm long) beetles have been found (pers. observ.) in large quantities living inside the heads of two species of Clibadium. One species of subfamily Alticinae (C. Triplehorn, pers. comm.) has been found in capitulescences of C. microcephalum (Stuessy & Arriagada 12365) from Ecuador, and one species of subfamily 41

Eumolpinae was found on C . surinamense (Arriagada 304) from Costa Rica {D. Furth, pers. comm).

Another intriguing relationship has been described between beetles and ants in Clibadium microcephalum from Ecuador (Stuessy & Nesom, 1982). Scolytodes elongatus

Schedl. produce cavities in the stems for nesting. Two species of ants belonging to the genus Pachycondyla and one belonging to Pseudomyrmex have been reported living in the nesting places originally occupied by beetles. There is no evidence, however, of a mutualistic relationship between these species and C. microcephalum. One species of Scolytodes (A. Chacon, pers. comm.) visits C. glomeratum. C. leiocarpum. C. surinamense. and C. sylvestre. from Costa

Rica, but no nesting cavities have been observed on these species (pers. observ.). ETHNOBOTANY

Some species of Clibadium have been reported to be used as fish-poisons and as medicines by Indian tribes throughout Latin America (Prance, 197 2; Duke, 197 0, 197 5, 197 8). Local people and natives often refer to fish-poison plants as "barbasco", which is a generic term in Spanish-speaking countries of South America for ichthyotoxic plants.

Additional local names may also be used.

One of the first reports indicating the use of Clibadium species as fish-poisons comes from Ernst (1881) who reported

C. sylvestre (as .C..,_agp_erum) and C. surinamense as the species broadly used from Panama and the Guyanas to the Upper Orinoco region. Clibadium sylvestre (as C. asperum) has also been mentioned by Pulle (1938) as a common fish-poison distributed in the West Indies, and French Guyana, Brasil and Venezuela used among Tusmani and Temi Indians in the Orinoco region. Clibadium sylvestre is known as "ware koenamie" (true barbasco) and C. surinamense as "oeman koenami" (false barbasco) (Pulle, 1938). Others species of Clibadium also have been mentioned (Ernst, 1881; Hoehne, 1939) as being used as barbascos and medicines: C. eggersii (as C. polvgynum) from Guatemala; Clibadium rotundifolium from northern and southern ; and C . peruvianum (as C.

42 43 heterotrichum) and C. surinamense (as C. strigillosum) from Peru.

Of all the species cited as fish-poison plants, only C. sylvestre appears to possess such activity. No indication of use as barbasco has been found or mentioned on any herbarium label for any species other than C. sylvestre. Obviously, with the exception of C. eggersii. the other species mentioned in the literature as barbasco are sufficiently similar in morphology that misidentification is likely.

The active compounds in Clibadium sylvestre are the polyacetylenic alcohols ichthyothereol (Gorinsky et al., 197 3), also isolated from Ichthyothere terminalis. and cunaniol (Quilliam & Stables, 1968). Recently, ichthyothereol and its acetates have been also isolated from

C. erosum and £L±. surinamense (Czerson et al., 1979; Christensen & Lam, 1991). Re-examination of voucher specimens indicates that C. erosum (Mexico, Stuessy & Gardner 4574) corresponds to C. arboreum. and that C. surinamense (Costa Rica, Stuessy & Gardner 44 61: and Ecuador, King & Garvey 6871) is correct as cited. In spite of chemical evidence, no field report has mentioned the use of these two species as a fish-poison. Perhaps ichthyothereol is produced in minute amounts in these taxa, or it may be locally restricted to chemical races that are not recognized and used by natives. In any event, the isolated compounds strengthen taxonomic relationships of these species with 44

Ichthyothere. also known to contain similar acetylenic compounds (tetrahydropyrans; Gorinsky et al., 1973; see also Generic Affinities).

Clibadium sylvestre is one of three broadly distributed species in the genus (with C. eggersii and C. surinamense), and the only one reported as a fish-poison. Clibadium sylvestre is distributed in Brasil, Colombia, Costa Rica, Ecuador, Lesser Antilles, Panama, Peru, Suriname, and

Venezuela. There is a clear relationship between the broad distribution of C. sylvestre and its use by native peoples. Because of the large area covered by this taxon, there is considerable morphological variation in its vegetative structures. This variation does not correlate with any geographical distribution as well as with reported local domestication. It is probable that the current broad distribution of C. sylvestre could have been increased by migration (Vickers & Plowman, 1984) and domestication by native Indians [e.g., probable introduction in Venezuela by the Culinas Indians who migrated from Brasil (Rutter & Barozo-Ferreyra s.n., OS); the Choco Indians probably migrated from Brasil to Panama (Duke, 1970)].

The following citations obtained from herbarium labels illustrate the use, users, and some of the common names for the broadly distributed fish-poison plant Clibadium sylvestre. 45

Brasil. The Uaica Indians use the fruiting heads "udishihimok", the ("udihenuk"), and whole plant ("udishihi") as a fish-poison for small fish (Prance et al.

10529. ECON). The plant ("canabed" or "maku") is also cultivated by the Maku Indians (Prance et al.15555. ECON). In Portuguese the species is "cunambi" or "cunanbi"(= barbasco).

Colombia. Used by natives to stop bleeding, to cure wounds, and as barbasco, to kill fish (Garcia & Echavarria

232. MO; McDaniel 117 99. FSU). The Huitotos Indians cultivate this plant ("diare") in the Cara-Parana region (Ranghel-Galindo 4Q. COL). It is also cultivated by the Choco Indians together with manioc and platanos and locally named "huaca" and "catalina" (Gentry & Fallen 17253. COL). It is considered toxic to cattle (Romero Castaneda 6144. NY). The barbasco is commonly prepared using the tops of plants crushed with soil and mixed with water ( 5058. US); macerated and pounded leaves are put into a basket, dipped into standing water, and which stupify the fish (Zarucchi 1304. ECON, GH).

Ecuador. The Secoya Indians from Ecuador use C. sylvestre mixed with mashed-up fruits of Bactris aasipaes H.B.K. (Palmae) and shaped into balls that later are thrown into streams as bait to poison fish. Natives identify C. sylvestre as "to'teo" or "to'te eo" (Secoya name for pounded poison). Clibadium sylvestre is sometimes mixed with finely

chopped meat and thrown to the water where after a few minutes the effect is observed. Most accounts describe the fish as stunned and sometimes dying almost immediately, rising to the surface (Kamen-Kaye, 1977). This species is commonly cultivated and used by the Lowland Quichua Indians ("Panga ambi"; Mexia 7256. US; Prance et al. 10529. US) to stupify small fish (Maries & Palacios 39. F) and as pain killer (Gentry 9811. CAS). The Waorani Indians use it as fish-poison (Davis & Yost 924. ECON). Clibadium sylvestre is also cultivated by the Kofan Indians (Martin 61. ECON). It is broadly cultivated in house gardens by the Colorado Indians (Cazalet & Pennington 5169. NY), the Secoya, and Siona Indians (Vickers 210. F) and the product sold. Several kilograms of leaves are mashed by the Shuar Indians in water and the water agitated until foam appears (Limback 120. CAS).

Panama. There are only a few reports of C. sylvestre being used as fish-poison in Pamana (Allen 857. F, MO, NY,

US; Allen 4582. GH, MO). This lesser use may be because natives use different fishing techniques and/or plants in this region. One of the most common plants used is Piper auritum H.B.K., which is found throughout lowland Panama and used to attract fish without poisoning (, 1979). Use of Clibadium sylvestre to intoxicate fish has been banned by the

Panamanian government, apparently because it kills small as well as large fish (Stuessy, 197 6). It is still used illegally, however (Hartman & Duke 3866. OS). Peru. Clibadium sylvestre is commonly named "huaca" or "common barbasco", but also known locally as "wasu", "apach" ( s .n .. ECON), and "Chichka" (Woytkowski 35187. CAS). Crushed leaves (Killip & Smith 26834. F; Seibert 2022. US; Woytkowski 35185. F) or the whole plant (Mexia 6506a. NY) are macerated and used as fish-poisons. Clibadium sylvestre is

cultivated by the Culinas Indians, who may have brought the plant from their former home in Brasil (Rutter & Barozo- Ferreyra s.n., OS).

Suriname. Clibadium sylvestre has been cited by Archer (1934) as used mixed with Piper spp., and it is known as "Warakabakoro" or "tona" by the natives. According to Vickers & Plowman (1984), this species is broadly cultivated in house gardens in Amazonia, Panama and northern South America.

Venezuela. Plants ("conambi" or "Kuma") are cultivated and used by Arekunas Indians to kill fish (Cardona Puiq 2848.

NY). The cultivated plants are crushed and mixed with soil for use as a fish stupefactant ( & Clark 7 674. MO) . PHYLOGENETIC ANALYSIS

A phylogenetic analysis of Clibadium. based on morphological data, was undertaken to provide insights into ancestor-descendent relationships and the recognition of natural infrageneric groupings. To reconstruct the phylogeny of Clibadium the following assumptions were involved: (1) the genus should contain all known descendants from a common ancestor (i.e., be holophyletic); (2) characters must be chosen and divided into discrete states; (3) polarity of these characters must be hypothesized (Crisci & Stuessy, 1980; Watrous & Wheeler, 1981); and (4) a parsimonious branching diagram must be constructed based on shared derived character states (i.e., synapomorphies, Hennig 1966). One of the many and most used criteria to determine polarity of character states is outgroup comparison (Nixon & Carpenter, 1993). This method assumes that a character state shared by an outgroup and the taxon under investigation is primitive, and any changes from that state are derived. An alternative to this is the use of functional out-groups (Watrous & Wheeler, 1981; Stuessy, 1990; Quicke, 1993). Using some reliable criterion for dividing the study genus into one or more infrageneric groups and deciding upon which is more primitive, it is possible to use this one as a functional outgroup for the others. The main advantages of all these

48 49 methods is that the data, assumptions of character state polarity, and choice of branching algorithms are clearly presented and open to evaluation and criticism (Stuessy, 1990) .

Materials and Methods. Fifteen morphological characters encoding 38 character states were used in the phylogenetic analysis (Tables 4 and 5) The following explanations are provided regarding the selection and use of some characters. Some taxa (e.g., C. grandifolium, c. oeruvianum, surinamense. C. sylvestre) show great morphological variation, particularly in vegetative structures, causing difficulty in delimiting some characters and states (e.g., leaf morphology, pubescence). Vegetative characters, therefore, were not used in the analysis.

Character states in species of Clibadium were polarized using Ichthyothere as outgroup (see Generic Affinities). An example is number of lobes of ray corollas (# 7). Most species in Clibadium have ray corollas with three lobes, somewith four, and few with only two. Ichthyothere has ray corollas with five lobes (occasionally four). I assume that five lobes represents the primitive condition and reduction in number of lobes is derived. Characters 6-15 had states polarized by this method. Because Ichthyothere does not provide certainty about ancestral character states of phyllaries, subgenus Paleata has been used as functional ingroup to provide evolutionary directionality in characters 50

Table 4. Characters and character states used in the cladistic analysis of species of Clibadium. Characters 4, 5, 7, 13, and 15 are ordered; the rest of multistate characters are unordered.

PHYLLARIES.

1, number of series: 2 (0), 1 (1) ;

2, shape: ovate (0), orbicular (1) t 2, vestiture: strigose (0), pilose (l), glabrous (2) 4, number: 6 (0), 4(1), 2(2); £, venation: 9 (0),7 (1), 5 (2). RAY FLORETS.

£, number: 16-40 (0), 3-12 (1) ; 7, corolla lobes: 4 (0), 3 (l), 2 (2) ,* 2, pales: present (0), absent (1);

£, number of series: 4-7 (0), 1-2 (l) ; 10. venation: 7-9 (0), 3-5 (1). RAY ACHENE. 11, texture: hirsute (0), pilose (1), glabrous (2). DISC FLORETS. 12., number: 14-24 (0), 3-7 (1); 13. corolla lobes: 5 (0), 4 (1), 3 (2);

14. pales: present (0), absent (1); 15. pale venation: 3 (0), 2 (1), 1 (2), 0 (3). 51

Table 5. Character state matrix for phylogenetic reconstruction of species of Clibadium. See Table 4 for description of characters and states.

Taxa/characters l 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4 1 5 acuminatum 1 1 0 1 1 1 1 0 1 0 1 1 0 0 0 anceps 0 1 0 0 0 1 0 1 1 0 2 1 0 1 3 arbor eum 1 0 0 1 1 1 1 0 1 1 0 1 0 1 0 armanii 1 0 0 1 1 1 1 0 1 1 0 1 0 1 0 cordatum 1 1 0 2 2 1 1 0 1 1 1 1 0 0 1 divaricatum 1 0 0 0 1 0 0 0 0 1 1 1 1 0 0 eggersii 1 0 0 2 1 0 0 0 0 1 1 1 1 0 0 erosum 1 0 0 1 1 0 1 0 0 1 1 1 1 0 0 frontinoense 1 1 0 1 2 1 0 0 1 1 1 0 0 2 glabrescens 1 1 0 1 2 1 1 0 1 1 1 1 0 0 1 glomeratum 1 2 0 0 1 0 1 1 1 0 1 3 grandifolium 1 1 2 2 2 1 .1 0 1 1 1 1 0 1 1 laxum 1 0 0 1 1 1 1 0 1 1 1 1 1 0 0 leiocarpum 1 0 2 1 2 1 1 0 1 2 1 0 0 leptophyllum 1 0 0 2 1 0 0 1 0 1 1 0 0 manabiense 1 1 0 2 2 1 1 0 1 1 1 1 0 0 1 micranthum 1 1 1 1 1 1 1 0 1 2 0 2 0 0 microcephalum 1 1 2 1 1 1 0 0 1 1 1 0 0 1 3 pentaneuron 1 1 2 1 2 1 1 0 1 1 1 0 0 0 1 peruvianum 1 0 0 1 1 1 1 0 1 1 2 0 0 1 0 rhytidophyllum 1 1 0 1 2 1 2 0 1 1 2 0 0 0 2 sessile 1 1 2 1 0 1 2 0 1 2 0 0 1 3 sodiroi 1 0 0 1 1 1 1 0 1 1 1 0 1 1 0 sprucei 1 1 0 1 2 1 2 0 1 1 1 0 0 0 2 surinamense 1 0 0 1 2 1 1 0 1 1 1 0 1 1 0 sy lv estre 1 0 0 1 2 1 1 0 1 1 0 1 0 0 terebinthinaceum 1 0 2 1 1 1 0 1 1 0 0 0 0 trianae 0 1 2 0 0 1 0 1 1 1 0 0 1 3 zaruchii 1 1 0 1 2 1 0 0 1 1 2 0 0 0 2 52

1-5 (pubescence, shape, and venation). This subgenus was selected because it is closest to Calea and Galinsoga (here considered distant ancestors of Clibadium: see Generic Affinities) especially in paleaceous receptacles and large numbers of ray florets in 5-7 series.

The states of some morphological structures in Clibadium have undergone more than single evolutionary step changes. These multistate characters are common in systematic studies, and their value has been affirmed many times (e.g., Jansen,

1985). There is some disagreement, however, on whether these characters should be treated as ordered or not. According to Hauser & Presch (1991), ordered characters can produce more, equal, or less equally parsimonious trees and can increase, decrease or have no effect on tree resolution. Ordered characters can contribute to greater resolution of the cladogram (Slowinski, 1993), and useful information may be lost when ordering is discarded (Page et al., 1992). Some quantitative multistate characters (4, 5, 7, 13, and 15) in Clibadium have been ordered. As an example, the number of ray corolla lobes (# 7) among species of the genus are 4, 3, and 2. Because the outgroup shows a 5-lobed condition and because most of the species of Clibadium are 4-lobed, a few 3-lobed, and only one species has a 2-lobed corolla, it is likely that the evolutionary trend within this character is a linear reduction in number of lobes. 53

Most parsimonious and consensus cladograms were produced for species of Clibadium. For summarization of multiple solutions from parsimonious analyses, consensus techniques are useful tools (, 197 2; Nelson, 1979; Anderberg & Tehler, 1990), but they are often greatly collapsed and uniformative (Farris, 1983; Bremer, 1990). Because of the lower explanatory power of consensus cladograms (Mickevich & Farris, 1981; Miyamoto, 1985; Carpenter, 1988), a competing parsimonious (non-consensus) cladogram of Clibadium has also been chosen for presentation and discussion as an alternative phylogenetic hypothesis.

Results and Discussion. The results of the phylogenetic analysis of species of Clibadium are shown in Figs. 5 and 6.

They correspond to the strict consensus cladogram (Fig. 5) and to one (Fig. 6) from among 25 equally parsimonious trees (consistency index of 0.682, i.e., ca. 32% homoplasy). The particular tree selected show the following species-pairs having the highest frequency of occurrence among the 25 equally most parsimonious trees: C. eggersii with C. leptophyllum (68%), C. frontinoense with C. zaruchii (76%), C . rhytidophyllum with C. sprucei (68%), and C . surinamense with C . peruvianum (60%), in each case two very closely related species pairs, which in my opinion, are evolutionary sister species and must be closely allied in the cladogram.

In both cladograms (strict and non-consensus) the basic clades remain the same and are supported by the same eggersii leptophyllum divaricatum terebinthinaceum erosum anceps glomeratum trianae sessile microcephalum frontinoense zaruchii rhytidophyllum sprucei grandifolium cordatum manabiense glabrescens pentaneuron laxum sodiroi surinamense peruvianum arboreum armanii leiocarpum sylvestre acuminatum micranthum

Fig. 5. Strict consensus cladogram of taxa of Clibadium. 55

eggersii E ^ g e rsia leptophyllum divaricatum j?S4) *= terebinthinaceum 3 03 ( * ) C l. 4 erosum Trixidium anceps glomeratum trianae a sessile 4>2 microcephalum E o frontinoense 3 zaruchii rhytidophyllum sprucei grandifolium cordatum 5§ manabiense ■oo glabrescens 2 o (f> S pentaneuron 3 3 laxum S :3 “I sodiroi cn U surinamense -a peruvianum 03 .f i arboreum 33 armanii _ V leiocarpum sylvestre £a acuminatum micranthum

Fig. 6. One of twenty-five equally most parsimonious cladograms of taxa of Clibadium generated by PAUP from 15 morphological characters. Consistency index 0.682 (32% homoplasy). The solid, gray and open bars represent synapomorphies, reversals, and parallelims, respectively. 56

synapomorphies. Cladograms indicate that two main evolutionary lines exist within Clibadium. They are here

recognized taxonomically as subgenera.

Subgenus Paleata is defined by heads with a large number of ray florets (15-40) all subtended by pales and arranged into 4-7 series, and ray achene exocarp and base of phyllaries and pales fleshy (producing the unusual fleshy head). Two sections are here recognized: (1) Eggersia. characterized by ray corollas with four lobes, and ray pales with five veins; and (2) Trixidium. supported by ray corollas with three lobes, and ray pales with three veins.

The remainder of the species in subgenus Clibadium are grouped into four lineages herein also recognized as sections. This subgenus is characterized by heads with 3-10 ray florets arranged in 1-2 series, with or without disc pales, usually with fewer ray florets than disc florets, and with mature heads without drupaceous achenes and fleshy phyllaries and pales.

The topology of the cladogram (Fig. 6) of Clibadium warrants additional comment regarding evolutionary relationships between subgenera and among sections. Within Subgenus Paleata. Section Trixidium appears equally derived with respect to Section Eggersia. Section Clibadium appears derived with respect to Section Oswalda. where there is a marked decrease in ray floret number (3-9) ; ray corollas with three lobes; ray pales with five veins; disc corollas with four or five lobes; and disc florets no subtended by pales (except C . laxum which has 3-veined disc pales). Section Grandifolia is more derived than Section Clibadium. and the former has phyllaries with only five veins, ray corollas with three lobes, ray pales with four veins; and disc pales with two veins. Section Glomerata appears as most derived within subg. Clibadium. There is an increase in number of phyllaries (6); ray corollas with four lobes (although two lobes are present in a few species, e.g., C . rhytidoohyllum. C. sessile, and C. sprucei); ray pale veins reduced (3) or increased (7-9); and disc pales (when present) with only one vein. MODES OF SPECIATION

Revisions or monographs on any given taxon should include hypotheses regarding patterns of speciation (Stuessy, 1993) . These hypotheses are important because they serve as a stimulus to evolutionary biologists and other workers

(Sundberg & Stuessy, 1990). Different modes of speciation in plants have been proposed and extensively discussed (e.g., White, 1978; Templeton, 1981; Grant, 1991; King, 1993; Levin,

1993; Rieseberg & Brouillet, 1994).

The available data on reproductive biology and isolating mechanisms of species of Clibadium are scarce and incomplete. To hypothesize about possible modes of speciation in the genus, correlations with distributional patterns, habitat, altitude, phenology, reproductive strategies, and geographical barriers must be taken into account. The absence of polyploidy in the genus (see Chromosome Numbers) suggests that speciation has occurred only at the diploid level. Also, distributional patterns of closely related species are allopatric. These patterns of distribution and same chromosome level for all species of Clibadium agree with Sundberg & Stuessy's hypothesis (1990) that, all other factors being equal, closely related species in Heliantheae

58 59 with the same chromosome number tend to have allopatric distributional ranges.

To infer modes of speciation, one must examine sister species or very closely related species ("species pairs") among extant taxa. The discussion will focus on the following species pairs in Clibadium that belong to the same species group and occupy similar ecological zones: C . anceps and C. glomeratum; c . arboreum and C. armanif; C. eggersii and C . 1eptophy1lum; C. frontinoense and C. zaruchii: C. rhytidophy1lum and C. sprucei: C. surinamense and C, peruvianum; and C. svlvestre and C. leiocarpum.

Two modes of speciation seem likely to have occurred in Clibadium: geographical and quantum. Geographical speciation includes the gradual and continuous process of evolutionary divergence from differentiation of geographical races to the development of isolating mechanisms leading to the formation of geographical species through geographical isolation (Grant, 1991). Lately, this mode has been questioned by Levin (1993). He claimed that, considering that speciation involves many stochastic processes and the evidence of reduced gene flow among populations, it is more likely that species evolve in local populations (metapopulations) rather than by the gradual transformation from races to subspecies, as suggested by geographical speciation. Quantum speciation, on the other hand, is the budding off of a daughter species through a geographically isolated or semi-isolated peripheral population of an out-crossing species. The variation fixing forces include both selection and drift (Grant, 1991). Other terms are used instead of quantum such as speciation by catastrophic selection (Lewis, 1962); speciation involving population flush-crash cycles (Carson, 1971), speciation via founder effect and genetic transilience (Templeton, 1980); speciation involving a genetic revolution or peripatric speciation (Mayr, 1982) ; and speciation in local isolates (Levin, 1993)

Geographical speciation. Spatial isolation is probably the primary isolating mechanism that serves as a stimulus for speciation in Clibadium. The evidence is that most of the taxa of Clibadium are isolated geographically, especially closely related species, such as those mentioned earlier. Although, according to Levin (1993), this condition does not necessarily mean that they were originally geographically isolated. All these species pairs exist allopatrically, and they seem to have evolved gradually as evidenced by largely quantitative differences between them, in the few cases of geographical overlap, the taxa are isolated by habitat differences or phenology (Fig. 7).

Details of these closely related taxa are instructive. Clibadium anceps is distributed in Costa Rica and Panama in wet secondary forest between 1000 and 2200 m, whereas C. glomeratum is restricted to low elevation (sea level to 500 JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOT DEC Subg. Paleata Sect. Eggersii eggersu divaricatum leptophyllum Sect. Trixidium erosum E terebinthinaceum E Subg. Clibadium Sect. Glomerata anceps glomeratum trianae sessile microcephalum frontinoense itxtxmmiiiiMi zaruchii rhytidophyllum sprucei Sect. Grandifolia grandifolium cordatum rnanabiense t...... glabrescens pentaneuron Sect. Clibadium laxum sodiroi surinamense ...... •'...... peruvianum arboreum armanii Sect. Oswalda leiocarpum igsifflaasm ii sylvestre t—...... — acuminatum t.V.V.V.V.:...1...':. micranthum

Fig. 7. Flowering range of species of Clibadium (based on herbarium data and personal observations). Order of taxa is phylogenetic, based on cladistic analysis. m) in drier areas (i.e., open roadsides) only in and provinces in Costa Rica. Clibadiuia arboreum is distributed only in southern Mexico between 1000 to 2500 m and flowers from April to June, whereas C. armanii is distributed in gallery forests on sandy soil or at edges of wet savannas in the Brasilian provinces of Bahia, Brasilia,

Distrito Federal, Goias, Mato Grosso do Sul, Minas Gerais, Parana, Rio de Janeiro, and Sao Paulo, between 500-1600 m and flowers from November to March. Clibadium frontinoense and

C. zarucchii. both present in the region of Frontino, Colombia but in different phytogeographic zones, occur at different elevations and on different slopes in the Cordillera Occidental of Colombia. Clibadium rhytidophyllum and C . sprucei are found only in Ecuador with the former in the province of Pichincha, and the latter endemic to slopes of Volcan Tungurahua. Clibadium peruvianum is restricted to secondary lowland wet forests in Argentina, Bolivia, and Peru, whereas C. surinamense has its northern distribution reaching only to central Ecuador. Clibadium svlvestre has a broad distribution growing between 200 to 600 m, whereas C. leiocarpum is restricted to Costa Rica, Nicaragua, and Panama growing between 1400 to 3000 m.

Quantum speciation. The following species pairs are here hypothesized to have evolved by quantum speciation: C. eggersii and C. divaricatum. and c. eggersii and C. 1eptophy1lum. In each case, one of the species (C. eggersii. presumably the older) has a broader distribution with the derivative taxon being more restricted and peripheral. These species pairs have allopatric distributions. Although the most parsimonious cladogram (Fig. 6) shows C. divaricatum as being more distantly related to C. eggersii than to C. leptophvllum. and the consensus cladogram (Fig. 5) shows a trichotomy suggesting that the three species could have derived from a common ancestor, I believe, based mainly on distribution and phenology, that C. eggersii is likely the oldest taxon, and that C. divaricatum and C . 1eotoohvllum have speciated from it by means of similar mechanisms. Clibadium eggersii is found in Guatemala, Honduras,

Nicaragua, Costa Rica, Panama, Colombia, Ecuador, and Peru (in the departments of Amazonas, Loreto, and San Martin), and C. divaricatum occurs only in the Department of Huanuco,

Peru. Clibadium eggersii flowers throughout the entire year, whereas C. divaricatum flowers only from April to August.

The mountains surrounding the area of Tarapoto may have served as a barrier between the ancestral population and the peripheral segregate, restricting gene flow between them and therefore abetting speciation. At the same time, C. eggersii probably also gave rise to C . 1eptophyllum. an endemic species known only from the Anchicaya and Choco region in Colombia, growing along the Hoya of Rio San Juan. Clibadium eggersii flowers from January to December, whereas C. leptophvllum flowers from March through June. MORPHOLOGY AND TAXONOMIC CRITERIA

The classification presented here is based mainly on comparative morphology. In Clibadium some characters show considerable variation that cause difficulty in delimitation of species. Also, many structures appear to have undergone parallel changes and reversals during evolution of the group (refer to cladistic analysis presented earlier). At a practical level, patterns of variation in features need to be clearly presented to facilitate use of keys and descriptions. For these reasons, the understanding of morphology is fundamental for revealing systematic and evolutionary relationships within the genus.

Definitions of qualitative character states follow Radford (1986). The term achene is used over cypsela (favored by Spjut, 1994) because it represents traditional usage of the term (e.g., Blake, 1924; Cronquist, 1955). All measurements of quantitative characters were made on herbarium material. Because shrinkage of some parts occurs with dessication, reproductive structures were rehydratated with Aerosol OT (10% aqueous) before measuring.

Habit. All species of Clibadium exhibit a shrubby or weakly arborescent habit. Most of the species show a highly

64 65 branched stem (typically represented by C. surinamense). Some species have arching branched stems but with one main stem at the base (e.g., C. anceps, C. glabrescens. C. laxum. C. trianae), others possess a more or less erect stem (10 cm diam in C . cordatum and C . rhytidophylum), and still others are arching vines (e.g., C. pentaneuron).

Leaves. Shape, size, margin, and base of blades are sometimes useful in delimiting species of Clibadium. Blades of most taxa are ovate except in C. leptophyllum which is easily recognizable by its narrowly lanceolate leaves.

Clibadium rhvtidophvllum and £L_gQnLCJSi are likewise distinctive in their regular lanceolate leaves. The largest leaves (40 x 30 cm) are found in C. cordatum. C. frontinoense. C. grandifolium. and C. zaruchii (all in Section Grandifolia). and the smallest leaves (3 x 1.5 cm) in C. armanii. Most of the species have blades with a typical serrate margin, but C . erosum is distinctly erose. Some species (e.g., C. armanii. C . cordatum) have leaves with cordate or truncate bases in contrast to the typical attenuate condition in most other species of the genus. Most species of Clibadium have unwinged petioles, but partially or entirely winged petioles are found in C . laxum (in mature individuals, e.g., Harling & 11525; Jaramillo 6713). Clibadium sessile and C. leptophyllum have no or very short petioles, a feature that obviously helps in delimiting these taxa. 66

Capitulescence. The perspectives on capitulescence structure presented here are based in part on Cronquist's (1977) concepts of organization, arrangement of heads and

branches, peduncles, etc., for the Compositae. For simplicity of application of terminology, heads are treated as if they represent single flowers. Most members of the family have a basic cymose capitulescence (Cronquist, 1977), although solitary heads are also known. The basic cymose

capitulescence occurs in two different forms: monochasial and dichasial (Wyatt, 1982; Weberling, 1989). Monochasial capitulescences have a terminal head that opens first, with the other heads opening in a centrifugal order down the axis. Dichasial capitulescences also mature centrifugally, but they have only three heads arising from the same node in which the central head opens first followed by the two lateral ones shortly thereafter.

All species of Clibadium have complex monochasial capitulescences which, depending on the species, are arranged secondarily into cymes or thyrses. These secondary arrangements are sometimes organized further into compound cymose and thyrsoid aggregations (Fig. 8). These categories of capitulescences have also been recognized in other genera of Heliantheae (Beaman, 1990; Strother, 1991). In all cases the capitulescences are suboppositely branched with subtending bracts decreasing in size near the heads, with heads sessile or on short peduncles and secondarily aggregated into large clusters with up to 600 small, loosely A

Fig. 8. Three main types of capitulescences in Clibadium. A, cymose, C. eggersii. from Stuessy (197 5, £. 50) ; b , thyrsoid, ..surinamensje, from Stuessy (197 5 £. 49); C, paniculiform, C. alabrescens. from Aristeguieta (1964, £. 62). arranged (e.g., C. grandifolium) or tightly packed (e.g., C. zaruchii) heads. Aggregation of heads is illustrated in one extreme by C. eggersii with heads loosely aggregated (Fig. 8A), moderately aggregated as in C. surinamense (Fig. 8B), or densely aggregated as in C. glabrescens (Fig. 8C). Capitulescences can be very tighly congested ((e.g., C. frontinoense and C. zaruchii). Congested capitulescences also occur in all species of the related genus Desmanthodium. in particular D. congestum Arriagada & Stuessy. The more loose aggregation typical of C. eggersii is also found in many species of Ichthyothere. the other close relative of Clibadium.

Characterization of the total capitulescence of a taxon is complicated in Clibadium because there are many levels of organization. Estimation of the total number of heads is also difficult, particularly because the total number depends on the age of the branch. Many species flower for several months (Fig. 7), and the capitulescences continue to increase in size and complexity. This is probably one of the reasons why different types of capitulescences have been reported for the same species. For example, Clibadium svlvestre when young (e.g., Nee 7139) has a thyrsoidal capitulescence, whereas when older (e.g., Webster 167 82) it may be paniculiform. In Clibadium armanii. similarly, thyrsoidal capitulescences are found on young specimens (e.g., Harley & Castro 11295) whereas paniculiform ones occur on older ones (e.g., Irwin et al. 23772) . An even broader range of 69 variation occurs in C . surinamense. Here cymose capitulescences are found in young individuals (e.g., Knapp 1267: Maxon & Valentine 7 033) and panicles are observed in older ones (e.g., Dwyer 7197: Maxon & Harvey 6505: Stern et al. 720).

Occasionally an intermediate or slightly different capitulescence arrangement may be observed within a particular species. For example, some individuals of C. anceos (e.g., Dwver & Correa 8844) and of C, glabrescens. (e.g., Killip & Cuatrecasas 38862) deviate from the typically capitate panicle to a nearly umbellate one.

The capitulescence types observed among species of

Clibadium do not correlate with the proposed subgeneric and sectional classification of the genus. Similar morphological variation in capitulescences has been observed in other genera unrelated to Clibadium (e.g., Hieracium L., Beaman, 1990; Pappobulus S. F. Blake, Panero 1992; Simsia Pers., Spooner, 1990). Because of the considerable variation in capitulescences of some species of Clibadium. and due to the obvious cycles of capitulescences evolution within many of the different taxonomic sections, these characters have not been used in the cladistic analysis. However, C. frontinoense, C. laxum. C. trianae. and C. zaruchii have very distinctive capitulescences and these features can be diagnostic for specific recognition. Heads. Heads in Clibadium are heterogamous with white, tubular corollas, with carpellate and fertile peripheral florets (ray), and hermaphroditic and female-sterile (i.e., functionally male) central disc florets. This is simple monoecy, one of many breeding systems known to occur in the family (Lowry & Stuessy, 1982). The number of ray and disc florets is very important in establishing groups of species and for purposes of subsequent identification.

The number of carpellate florets ranges from 3-5 (C. ginc.eps, C. glomeratum, C. microcephalum. C. sessile, and C. trianae) to 30-40 (C. divaricatum. C. eggersii. and C. leptophyllum). The number of disc florets is 3 (C. microcephalum, C. zaruchii), 6 (C. anceps. C . rhytidophy1lum) or 24 (C. leptophyllum).

The number of heads per capitulescence varies in different taxa as follows: 24-28 heads in C. eggersii (Tyson

3-4.22) , 60-116 in C. trianae (Stuessy & Funk 5623) . 70-170 in C. surinamense (Stuessv & Funk 5510), 7 5 heads in c. sessile (Allen 497 0), 86-119 heads in C . anceps (Dwyer & Correa 8844: Stern et al. 1019), 94-106 in C. oentaneuron (Stuessy & Funk 5595), 98-100 heads in C. leiocarpum (Allen 1341), 100-130 in

C. glabrescens (Stuessy & Funk 5606), 219 heads in C. microcephalum (Stuessy & Arriagada 12362), and up to 600 in C. grandifolium (Croat 13372; Lewis et al. 2082). An unusual number of 245 heads in a cluster was counted in a young individual in C . trianae (Stuessv & Arriaaada 12337), 71 but the basic head and capitulescence organization was the same as observed in other specimens.

Involucres. The number of phyllaries, their arrangement into series, and their shape, texture, and venation are all taxonomically important in Clibadium. The phyllaries are herbaceous and spreading and usually reflex during fruiting (clearly seen in C. manabiense). The number and length of phyllaries are usually constant, and they are often pubescent over the upper one-third with a ciliate margin. Phyllaries in most species are organized in a single series. A few taxa (C. anceps. C. glomeratum. and C. trianae) , however, have two series. This exceptional arrangement may be due to a larger number of phyllaries (6) perhaps providing substantive protection as compensation for lack of pales. The most common number of phyllaries in the genus is three, with C. eggersii and C. leptophvllum with only two (this low number perhaps relating to a larger number of ray florets subtended by pales). Also uncommon is the 4-phyllaried condition in C. frontinoense and C. zaruchii. The shape of phyllaries ranges from ovate to orbicular with the upper one-third of the margin ciliate, and with apex obtuse (C. erosum. C. grandifolium. C. oentaneuron. C, sylvestre, terebinthinaceum) , acute (C. anceps. C. eggersii. C, glomeratum. C . laxum) or mucronate (C. armanii. C. cordatum. C. manabiense). Pubescence on the phyllaries is strigose in most species, but a few are glabrous (C. glomeratum. C. grandifolium, C. leiocarpum. C. microcephalum. 72 pentaneuron. C. sessile. C. terebinthinaceum. and C. trianae).

Pales. Two different morphological viewpoints exist with regard to bracts subtending ray florets in Clibadium. Stuessy, in a broad survey of Heliantheae (1977), has regarded them as inner phyllaries and hence part of the involucre. Funk, in her monograph of Montanoa (1982), considered them as pales (i.e., outer row of receptacular bracts; Gleason 1952, Carter, 1964, and Cabrera, 1978, also holds this viewpoint) because of their similarity of development with bracts substending disc florets. Following Briquet (1917), I use the term "pale" for those bracts subtending ray and disc florets, and the term phyllary for those forming the involucre. In Clibadium. the female florets are subtended by broad pales often resembling phyllaries but they differ in having fewer veins and less pubescence. In species with more than one series of ray florets, there is a gradual change (reduction in size and pubescence) in pale morphology from outermost to innermost ray florets. Disc florets in some species of Clibadium are also subtended by receptacular bracts (pales). Pales subtending the innermost ray florets are similar to those of the disc. Such a transition can be seen clearly in Clibadium eggersii. with 5-7 series of ray florets.

Taxonomically important variations in Clibadium occur in pales of ray and disc florets. Ray pales with 7 veins characterize all members of Section Oswalda and some members of Section Glomerata (C. anceps. C. glomeratum. C. trianae, C. sessile). Section Clibadium and Subgenus Paleata have pales with 5 veins, and Section Grandifolia is 4-veined. Disc floret pales are more uniform than those of ray florets. All members of subgenus Paleata. as well as all taxa in section Oswalda in subgenus Clibadium. are 3-veined. Taxa of section Grandifolia are 2-veined. All species but one (C. laxum) in Section Clibadium have epaleaceous receptacles.

Clibadium laxum has disc pales with three veins. Clibadium frontinoense. C . rhytidophy1lum. C. sprucei. and C. zaruchii of section Glomerata are the only species in the genus with only one conspicuous vein.

Ray corollas. The number of lobes of corollas is taxonomically valuable. Ten species are characterized by having 4-lobed corollas. This condition is found among some members of subgenus Paleata as well as among most members of Section Glomerata in subgenus Clibadium. Sixteen species are characterized by having 3-lobed corollas, including all species in sections Clibadium. Grandifolia. and Oswalda. In addition, two species from the Caribbean (C. erosiim and C. terebinthinaceum) belonging to subgenus Paleata share this condition. Three species in Section Glomerata (C. rhytidophy1lum. C. sessile, and C. sprucei) have 2-lobed corollas. 74

Disc corollas. The number of lobes of disc corollas also has taxonomic importance. All species in Section Eggersii of Subgenus Paleata have 4-5 lobes; Clibadium erosum and C. terebinthinaceum. both of Section Trixidium of the same subgenus, have disc corollas with 4 and 5 lobes, respectively. All species in sections Glomerata and Grandifolia are 5-lobed. The remainder of species in sections Clibadium and Oswalda are 5-lobed or 4-lobed. One species, Clibadium micranthum, has an unusual 3-lobed condition. There is no correlation between number of lobes in disc and ray corollas.

Achenes. All species have achenes with similar morphology and size proportional to florets and heads. Some species of Clibadium. however, possess unusual drupaceous achenes with orangish or greenish juice. This has been observed in the field in mature fruits of C. eggersii and C. laxum which release yellow- and orange juice, respectively, when squeezed. The drupe-like fruits have an inflated hypodermis with large cells and cell layers in which juice accumulates (Stuessy & Liu, 1984). This type of fruit is rare in Compositae, known only in a few other genera, e.g., Chrysanthemoides Fabr. (Calenduleae), and Neck, ex Cass. (Heliantheae).

Clibadium eggersii. C. erosum. C. divaricatum. C. leptophvllum. and C. terebinthinaceum of Subgenus Paleata possess an unusual head superficially resembling an aggregated fruit such as in Rubus ulmifolius Schott (Rosaceae). This conspicuous structure is the result of aggregation of 16-40 drupaceous achenes all of which develop fleshy exocarps at maturity. In addition, phyllaries and ray pales also become fleshy at their bases resulting in an inflated structure (even easily observable in dry herbarium specimens). This modified head is used as food by birds (e.g., C. eggersii in Costa Rica; Feinsinger et al. 2182). which could serve as dispersal agents.

Vestiture. Vestiture terminology in this treatment follows Lawrence (1951). All species of Clibadium have some type of pubescence on leaves (mainly undersurface and along veins of leaves), stems, phyllaries, and achenes. There are two types of trichomes: (1) uniseriate; and (2) biseriate (Zwillingshaare, Hess, 1938). Uniseriate trichomes are most common. Their length and density change with position on different structures in different species (e.g., C. arboreum. C. leiocarpum. C. surinamense. C. trianae). Biseriate trichomes are restricted to achenes of some species (e.g., C. armanii. C. leptophyllum. C. sprucei). Variation in ray achene pubescence does not correlate subgenerically or sectionally, but it is taxonomically helpful in characterizing individual species. Some taxa are glabrous (C. anceos. C. leiocarpum. C. micranthum, C. peruvianum, ,C.,_. rhvtidophvllum. C. sessile. C. zaruchii), and some are hirsute (Clibadium arboreum. C. armanii. and C. leptophyllum), or pilose (rest of genus). TAXONOMY

Clibadium L., Mant. Pi. 161. 17 71.--Type: C. surinamense L. Baillieria Aubl., Pi. Gui. 2. 804. t. 317. 1775.--Type: Baillieria aspera Aubl. [= C. surinamense L .]. Sw., Prod. Veg. Ind. Occ. 815. 1788. non P. Browne, Hist. Jamaic. 312. 1756.--Type: Trixis aspera Sw. [= Clibadium surinamense L .].

Oswalda Cass., in Diet. Sc. Nat. 59. 322. 1829.--Type:

Oswalda baillierioides Cass. [= C. surinamense L.]. Orsinia Bertol. ex DC., Prod. 5. 104. 1836.--Type: Orsinia eupatoria DC. [= C. armanii1. Trichapium Gilli. Feddes Repert. 94:312. 1983.--Type:

Trichapium strigosum Gilli [= C. manabiense H. Robinson].

Shrubs or small trees with a conspicuous single main stem to profusely highly branched at soil level; stem rounded or many-angled, striate, glabrous or strigose. Leaves opposite, with blades narrowly lanceolate to broadly ovate, subcoriaceous to coriaceous, 5-50 cm long, 2.5-40 cm wide, base attenuate, rounded or cordate; at apex acute, obtuse, mucronate; margin serrate, serrulate, finely irregularly serrate, dentate-serrate, crenate, erose; surfaces pilose,

76 strigose or glabrous, with veins sometimes prominent on abaxial surface. Capitulescences of disciform whitish cymose, thyrsoid or paniculiform arrangement of heads organized in open clusters or dense terminal or axillary glomerules, oppositely branched with subtending bracts decreasing in size near heads. Heads few to 600, heterogamous, few to 600, with marginal carpellate and fertile florets and central hermaphroditic and sterile florets; sessile or on short peduncles; involucre cupulate or funnelform (occasionally subglobose), of 2-6 imbricate, coriaceous phyllaries; phyllaries ovate or orbicular, 5-12- veined; at apex obtuse, acute or mucronate, with upper one- third of margin mostly ciliate; pales subtending carpellate florets often resembling phyllaries, ovate or orbicular, 3-9- veined; pales subtending hermaphroditic florets (sometimes absent) membranaceous, lanceolate, 1-3-veined, with upper one-third of margin ciliate. Ray florets 3-40, usually uniseriate or less often multiseriate, fertile; corollas tubular, white or yellow, 2-4 lobed; styles bifid, with strong marginal stigmatic lines; pappus absent, sometimes of tufts of short hairs (C. anceps) or two short awns (C. sylvestre); achenes obovoid, compressed, biconvex, brown- black. Disc florets 3-24, with tubular white corollas, 4-5- lobed; anthers purplish or black, sagittate at base; style undivided; ovaries sterile, linear, pilose at apex; with a 5- lobed nectary at base; pappus absent. Chromosome number, n = 16 (Stuessy & Arriagada, 1993) . 78

The subgeneric and specific taxa recognized in this study are delimited based on morphological characters complemented by geographical and ecological data. No varieties or subspecies are designated, because the observed morphological variation within species does not correlate with geography (i.e., it is scattered randomly among populations).

In this taxonomic treatment of Clibadium. herbarium acronyms follow Index Herbariorum (Holmgren et al., 1991), author abbreviations are from Authors of Plant Names (Brummitt & Powell, 1992), book title abreviations are based on TL-II (Stafleu & Cowan, 1976-88), and journal titles follow B-P-H/S (Lawrence et al., 1991), and the method of citing specimens on Interdocumentation Center (IDC) microfiches follows Hepper (1968) .

Key to species of Clibadium

1. Ray florets multiseriate; number of ray florets always greater than disc florets; all florets subtended by pales; mature heads with drupaceous achenes; phyllaries and pales fleshy (at base) at maturity; capitulescence thyrsoid. Clibadium subg. Eggersii

2. Capitulescence with fewer than 25 heads; each with more than 30 ray florets; ray corollas 4-lobed; ray pales 5-veined; Central and South America. 79

3. Heads 4-6 mm tall, with 2 phyllaries; ray florets 30-40; disc florets 8-24.

4. Leaves ovate; disc florets 8-9.... 1. C. eggersii

4. Leaves narrowly lanceolate; disc florets 24; ...... 2. C. leptophyllum

3. Heads 2-3 mm high, with 5-6 phyllaries; ray florets 24-39; disc florets 6-10 3. C. divaricatum

2. Capitulescence with 50-100 heads; each with fewer than 20 ray florets; ray corollas 3-lobed; ray pales 3-veined; Caribbean and Lesser Antilles.

5. Leaves with margin crenate-serrate, at apex acute; disc florets 7; disc corollas 5-lobed; and ...... 4. C . terebinthinaceum

5. Leaves with margin erose, at apex short-acuminate; disc florets 8; disc corolla 4-lobed; Puerto Rico and Lesser .Antilles ...... 5. C. erosum

1. Ray florets uniseriate; number of ray florets same or fewer than disc florets; ray florets subtended by pales only, or heads epaleaceous; mature heads with dry achenes; phyllaries and pales dry (never fleshy) at maturity; capitulescence cymose or paniculiform. Clibadium subg. Clibadium

6. Heads arranged in condensed capitulescences.

7. Heads epaleaceous. 80

8. Capitulescences of flat-topped clusters with 3-6 subsessile heads...... 6. C. anceps

8. Capitulescences of round-topped clusters with 10-30 sessile heads.

9. Leaves pilose on both sides, with margin crenate-serrulate; heads 10-20, 2-3 mm tall; Costa Rica and Panama 7. C. glomeratum

9. Leaves strigose on both sides, with margin dentate-serrate; heads 20-30, 5 mm tall; Colombia...... 8. C. trianae

7. Heads paleaceous throughout or with pales substending ray florets only.

10. Only ray florets subtended by pales.

11. Leaves sessile; ray florets 5; disc florets 6-8; ray corollas 2-lobed; phyllaries 7-12- veined...... 9. C. sessile

11. Leaves petiolate; ray florets 3; disc florets 3-4; ray corollas 4-lobed; phyllaries 4-6-veined 10. C. microcephalum

10. Ray and disc florets subtended by pales.

12. Leaves 20-35 cm long, 15-25 cm wide; heads with 4 phyllaries; Colombia.

13. Heads less than 6 mm tall, in condensed clusters 1.5-2 cm diameter ...... 11. C. frontinoense 81

13. Heads 8 mm tall, in condensed clusters 3-4 cm diameter...... 12. C. zaruchii

12. Leaves 7-14 cm long, 2.5-4 cm wide; heads with 3 phyllaries; Ecuador.

14. Leaves ovate to ovate-lanceolate; at apex acute; heads 4 mm tall...... 13. C. rhvtidophvllum

14. Leaves lanceolate; at apex narrowly acuminate; heads 6 mm tall...... 14. C. sprucei

6. Heads arranged in loose cymose or thyrsoid capitulescences.

15. Leaves 30-50 cm long, 10-40 cm wide; capitulescences with 200 or more florets.

16. Leaf bases attenuate, decurrent 1/3 of petiole; 400 or more heads per capitulescence; disc florets epalaceous not subtended by pales...... 15. C. grandif olium

16. Leaf base cordate with petiole naked; less than 300 heads per capitulescence; disc florets subtended by pales....16. C . cordatum

15. Leaves less than 20 cm long, 10 cm wide; capitulescences with less than 200 florets.

17. Ray florets only subtended by pales.

18. Disc corollas 4-lobed. 82

19. Leaves at base shortly cuneate, with margin crenate-toothed; heads 100 or more per capitulescence; disc florets 6 or fewer ...... 21. C. sodiroi

19. Leaves at base obtuse, with margin serrate to serrate-dentate; heads fewer than 100 per capitulescence; disc florets 10-14 22. C. surinamense

18. Disc corollas 5-lobed.

20. Leaves with veins yellowish underneath, at apex acute, at base attenuate, with margin finely irregularly serrate; disc florets 20 or more; phyllaries 3, 7 -veined...... 23. C. peruvianum

20. Leaves with veins light-green underneath, at apex acuminate, at base rounded, with margin serrate or mucronate-dentate; disc florets fewer than 10; phyllaries 3-4, 5 -veined.

21. Leaves membranaceous, 10-20 cm long, pilose, with margin serrate; Mexico, Nicaragua ...... 24. C . arboreum

21. Leaves subcoriceous, rigid, 5-7 cm long, hispidulous, with margin mucronate- dentate; Brasil, Paraguay....25. C. armanii

17. Ray and disc florets subtended by pales (sometimes 1-2 central disc florets not subtended by pales).

22. Disc corollas 3- or 4-lobed. 83

23. Disc corollas 3-lobed...... 29. C. micranthuxn

23. Disc corollas 4-lobed.

24. Capitulescence paniculifozm; stem many­ angled; leaves 14-30 cm long, with blade decurrent on petiole; phyllaries 5-7-veined ...... 20. C. laxum

24. Capitulescence thyrsoid; stem terete; leaves less than 14 cm long, with petiole naked; phyllaries 8-11-veined.

25. Leaves and stems hirsute; phyllaries dark brown-green, glabrous...... 26. C. leiocarpum

25. Leaves and stems strigillose; phyllaries light green, strigose...... 27. C. sylvestre

22. Disc corollas 5-lobed.

26. Leaves coriaceous, with veins conspicuous on abaxial surface.

27. Phyllaries at apex acuminate, strigose; ray florets 9-13; disc florets 15...... 17. C. manabiense

27. Phyllaries at apex obtuse, glabrous; ray florets 5-6; disc florets 5-8...... 19. C. oentaneuron

26. Leaves thin-pappery, smooth on abaxial surface. 84

28. Leaf margin serrulate, with teeth entire; phyllaries 5-veined; ray pales 4 -veined; disc pales 2-veined...... 18. C. glabrescens

28. Leaf margin serrate to pauci-serrate, with teeth mucronate; phyllaries 7-veined; ray pales 7 -veined; disc pales 3-veined...... 28. C. acuminatum

I. Clibadium L. subgenus Paleata Arriagada, subg. nov. --Typus: Clibadium eggersii Hieron.

Frutices capitulescentiis thyrsoideis, capitulis minus quam 100; feminei pluriseriati, floribus femineis plus quam floribus hermaphroditis disci; achenia feminea drupacea; phyllaria et paleae carnosae.

Shrubs with thyrsoid capitulescences with up to 100 heads; ray florets multiseriate; with number of ray florets always greater than disc florets; disc florets subtended by pales; mature achenes drupaceous; phyllaries and pales fleshy (at base) at maturity. Species 1-5.

IA. Clibadium L. section Eggersia Arriagada, sect. nov. --Typus: Clibadium eggersii Hieron.

Capitulescentiae minus quam 25 capitulis; flores feminei plus quam 30, corollae tetralobae; paleae quinquenervis. Capitulescences with fewer than 25 heads; ray florets more than 30, with corollas 4-lobed; ray pales 5-veined.

Species 1-3.

1. Clibadium eggersii Hieron., Bot. Jahrb. Syst. 28:

598. 1901.--Type: Ecuador. Manabx: Rio Rapallo near El Recreo, 6 Sep 1893, Eagers 153 09 (holotype: B, presumed destroyed, fragment at US!, photos: F! GH! MO! OS!; isotypes: K! photos: GH! OS!, P photo: OS!).

Clibadium pittieri Greenm., Proc. Amer. Acad. Arts 39: 98. 1903.--Type: Costa Rica. Limon: La Florida, Jul 1897, Pittier s.n [= 11290 CR accession number] (lectotype here designated: GH!; isolectotypes: CR!

US!) .

Clibadium pittieri Greenm. f. phrixium Greenm., Proc.

Amer. Acad. Arts 40: 38. 1904.--Type: Costa Rica. "Buissons a Luis", Nov 1897, Tonduz 1147 9 (lectotype

here designated, GH!; isolectotype: CR! US!).

Clibadium polygynum S. F. Blake, Contr. Gray Herb. 52:

32. 1917.--Type: Nicaragua. 1867, Seemann 88 (lectotype here designated, BM!, photos: GH! OS!; isolectotype: BM! photo: OS!, fragments at GH! US!).

Clibadium oropincruum s. F. Blake, Contr. U. S. Natl.

Herb. 22: 597. 1924. Wulffia sodiroi Hieron., Bot. 86

Jahrb. Syst. 29: 34. 1901.--Type: Ecuador, Santo Domingo de los Colorados, 500-1600 m, ca. 1894 [Hieronymus, 1901], Sodiro 21/3 (holotype: B, presumably

destroyed, fragment at US!).

Clibadium chocoense Cuatrec., Rev. Acad. Colomb. Ci.

Exact. 9: 236. 1954.--Type: Colombia. Choco: Rio San Juan, "cercanxas de Palestina", 31 May 1946, Cuatrecasas 21514 (holotype, F!, photo: US!).

Shrub 2-3 m tall; stems 2-3 cm in diameter, often

strigose with hairs 0.1-0.5 mm long. Leaves with blades ovate to ovate-lanceolate, 5-16 cm long, 4-10 cm wide, tripliveined, at apex acuminate, at base attenuate, with margin serrate, with both surfaces weakly or densely strigose with hairs 0.5-1 mm long; petioles 2-5 cm long, to 1 mm diameter, strigose with hairs 0.1-0.5 mm long. Capitulescence thyrsoid, with 10-25 heads; peduncles 1 mm long. Heads radiate, 5 mm high, 4 mm diameter; involucre cupulate, 4 mm diameter; phyllaries 2, ovate, 3-3.5 mm long, 2-2.5 mm wide, 5-7-veined, strigose, with hairs 0.1 mm long, the upper one-third ciliate, at apex acute. Ray florets 28- 40, subtended by ovate pales, 5-veined, 2-3 mm long, 1.5-2 mm wide, strigillose toward apex with hairs 0.1 mm long, with margin ciliate on upper one-third; corollas white, 1 mm long, with tube 0.6 mm long, 0.4 mm diameter, 4-lobed, with lobes deltate, shortly ciliate at margin; styles 1.5 mm long, with branches 0.5 mm long. Disc florets (4-) 8-9, subtended by 87 lanceolate pales, 2 mm long, 0.5 mm wide, 3-veined; corollas 1.5 mm long, (4)-5-lobed, with deltate lobes 0.5 mm long scarcely pubescent at apex; with throat 1 mm long, 0.5 mm in diameter; anthers black, 1.5 mm long; style 2.5 mm long, with branches 1.5 mm long; ovaries sterile, 1.5 mm long, 0.6 mm diameter, pilose toward apex with hairs 0.5 mm long. Achenes 2 mm long, 1.6 mm diameter, hirsute on upper one-third, with hairs 0.5 mm long. Chromosome number, n = 16.

Phenology. Flowering during the entire year throughout its distributional range. Distribution and Ecology (Fig. 9). Belize, Costa Rica,

Colombia, Ecuador, Guatemala, Honduras, Lesser Antilles, Nicaragua, Panama and Peru; modified wet habitats with secondary vegetation, along roadside; 0-1000 m.

Representative Specimens. Belize. District of Stann

Creek: along road and stream at Dry Creek, Croat 24522 (MO); near District of Cayo border, Dwyer 1119 8 (MO); on hillside, Hummingbird Highway, Gentle 8251 (F, MO, TX); 22 Mile Stann Creek River, Schirm 97 2 (F, GH, MO, NY). Colombia.

Antioquia: valley of Rio Anori between Dos Bocas and Anori, Denslow 2642 (WIS); carretera al mar cerca de Villa Arteaga, Gutierrez & Barkley 17C122 (COL); Zaragoza, Providencia, along Rio Anori, Soeiarto & Villa 2715 (COL). Caldas: Santa Cecilia, Cordillera Occidental, Vertiente Occidental, von Sneider 5168 (F). Choco: La Concepcion, 15 km E of Quibdo, Archer 1945 (BM, US); between Rio Nercua and Quebrada

Ambrosio near Camp Curiche, Duke 1147 0 (FSU); along Rio 88

Fig. 9. Distribution of Clibadium divaricatum ( ▼ ), C. eggersii ( • ), and C. leptophvllum (v ) in Central America and northwest South America. Mutata, Forero & Gentry 7 25 (COL); along Rio Mecana, Juncosa 17 33 (F); SE of Bahia de Solano, Q. Jellita, White & Warner 14 (MO). Meta: Sierra de la Macarena, Vereda El Tablazo, Chaparro et al. 81 (COL, F ) . Narino: Tumaco, La Guayacana, Romero-Castaneda 2974 (COL). Santander: Barranca Bermeja, between Sogamos and Carare Rivers, Haught 2047 (COL).

Valle: near Queremal, Alston 7901 (US); Cordillera Occidental, vertiente occidental, Hoya del rio Anchicaya, Cuatrecasas 137 09 (F) . Costa Rica. : about 8 km S of Colonia Blanca NE toward Volcan , Almada & Uakai 4023 (OS); about 2-4 km N of Bijagua on road to , Almada & Uakai 4 033 (OS); Cordillera de Tilaran, near Rio San Lorencito, NW of San Ramon, Arriagada 412 (OS); ca 15 mi S of

La Marina near Rio , Stuessy & Gardner 4465 (OS). Guanacaste: El Silencio, near Tilaran, Standley & Valerio 44639 (US). : Finca La Selva, confluence of Rio Puerto Viejo and Rio Sarapiqui, Denslow 79-104 (WIS); Tirimbina, 32215 (MO). Limon: Confluent du Puerto Viejo et du Sarapiqui, 7 399 (GH); Parque Nacional Braulio Carrillo, Quebrada Gonzalez, Chavarria 143 (CR); ca 10 mi S of Punta Cahuita, Croat 43195 (MO); Hacienda Tapezco- Hacienda La Suerte, Davidson 6717 (MO); Talamanca, catarata Rio San Box, Gomez-Laurito 8740 (CR); Port Lemon, Kuntze 1985 (NY, US); Guapiles, , Leon 660 (CR); La Selva, 3 km SE of Puerto Viejo, Opler 1602 (CR); Barra del Colorado, N side, Stevens 24140 (OS); Cerro Coronel, E of Rio , Stevens 23951 (OS); E of Laguna Dante, Stevens 24606 (CR); Finca Montecristo on the Rio Reventazon below Cairo, Standiey & Valerio 48507 (US). San Jose: San Ramon, Cataratitas, Poveda & Castro 4135 (CR) ; Basin of El General, Skutch 5217 (CR); Parque Nacional Braulio Carrillo, Estacion La Montura, Todzia 17 21 (F). Puntarenas: Peninsula, Lent 435 (MO); Rincon de Osa, 5 km W of Rincon, Liesner 2098 (OS); Golfo Dulce and Rio Terraba, Skutch 5271 (US), Ecuador. Pichincha: 20 km N of Santo Domingo de Los Colorados, Cazalet & Pennington 5188 (NY); Centinela, Canton Santo Domingo, 12 km E of Patricia Pilar, Dodson et al. 7 235 (MO); Reserva

Forestal Endesa, Rio Silanche, Jaramillo 67 00 (MO). Napo: 44 km E of El Chaco, Gentry 12409 (CAS, MO); Canton Mera,

Hacienda La Mascota, Mexia 7 024 (CAS); near Archidona, Mexia 7272 (CAS). Puyo, Prescott 4 05 (ECON). Pastaza: Colonia 24 de Mayo, to road Puyo, Puerto Napo, Lugo 2481 (MO); vicinity of Puyo, E foot-hills of the Andes, Skutch 44 86 (F, GH, MO, US). Guatemala. Alta Verapaz: Cubilquity, Turckheim 414 8 (F). Honduras. Atlantida: vicinity of La Ceiba, Danto River, Yuncker et al. 8699 (F, US). Comayagua: margins of Lake Yojoa, Williams & Molina 14621 (GH). Copan: Hacienda El Limon to El Paraiso, Blake 7 354 (US). Olancho: orillas del riachuelo Aguaquire, 3 0 km NE of Culmi, Nelson & Vargas 27 28 (MO). San Pedro Sula: Cuyamel, Carleton 458 (US). Nicaragua. Comarca del Cabo: Little Francia, Nelson 4830 (CAS, F, MO, NY). Jinoteca: Bocaycito, Neill 97 (MO). Matagalpa: NW Cerro Musun, trocha de Palan, Araquistain & Moreno 2440 (F); Macizos de Penas Blancas, SE side, Stevens et al. 2107 8a (OS); Rio San Juan: near Cano Chontaleno 20 km NE of El Castillo, Neill & Vincelli 3578 (OS). Zelaya: Cano Montecristo, 1 km S campamento La Grupera, Moreno 15098 (MO) ; Rio Punta Gorda, Atlanta, al SE de "La Richard", Moreno & Sandino 13072 (MO); 3 km W of Kuikuinita, S of Siuna, Neill 4207 (F); M t . Liveco near Siuna, Madregava, Nelson 5059 (F, GH, MO, NY); 0.8 km S of Serranias de Yolaina on road to Colonia Manantiales, Pipoly 1999 (F); ca. 13 km above Kukuria on road to San Jeronimo, Pipoly 37 59 (F, OS); Cano between Cerro La Pimienta and El Hormiguero, Pipolv 6005 (F) . Panama. Bocas del Toro: road from Fortuna Lake to Chiriqui

Grande, Hampshire & Whitefoord 492 (MO); 10-15 mi S from mouth of Changuinola River, Lewis et al. 97 5 (GH, MO, US) ;

Old Bank Island, vicinity of Chiriqui Lagoon, Wedel 2161 (MO) . Colon: vicinity of Guasimo on Rio Miguel de la Borda, Croat 10005 (MO); Santa Rita Ridge, end of road from Transisthmian Highway, et al. 47 85 (MO) ; along banks of Rio Guanche, Sullivan 181 (MO). Darien: Cerro Pirre, Duke 6088 (MO); tops of ridges separating Rio Jaque Valley from Pacific Ocean, Knapp & Mallet 3101 (MO); Camp Summit, along Sea level Canal Route 17, Oliver et al. 3667 (MO).

Panama: Campo Tres, 5 km NE of Altos de Pacora, Bussey 825 (MO); forests along headwaters of Rio Corso, Duke 11918 (MO, OS); Cerro Jefe, Dwyer et al. 9496 (MO); Serrania de Maje, between Rio Ipeti Grande and Charco Rico, Churchill & Nevers

4429 (MO); tributary of Rio Chagres, 5 mi SW of Cerro , Nowicke & Oliver 3434 (MO); El Llano-Carti road, 20.9 km from Interamerican Hwy, Mori & Kallunki 5590 (MO) . Veraguas: 5 mi NW of Santa Fe, Croat 23190 (MO); Guabal (Rio Dos Bocas), about 16 km NW of Santa Fe, Dressier 4798 (MO); 92

16 km NW of Santa Fe, on road to Calovebora, Mori & Kallunki 6164 (MO). San Bias: Nusagandi, McDonacrh et al. 165 (MO). Peru. Amazonas: Bagua, Mesones-Muro Highway below Montenegro 5.5 km, E of Olmos on or above the Rio Maranon, Hutchinson & Wright 37 21 (NY); Mendoza, Woytkowski 8310 (MO). Huanuco:

Pachitea, Codo de Pozuzo, Foster 9315 (OS). Loreto: above Pongo de Manseriche, Creek Carapisa, Mexia 6255 (F, GH, MO, NY, TEX, US). San Martin: Mariscal Caceres, Tocache Nuevo, Schunke 77 86 (MO, OS).

Clibadium eggersii is particularly distinctive at maturity by its globose heads containing 28-40 ray florets arranged in 5-7 series and placed in the axils of ovate pales. Because the phyllaries and pales (at base) become fleshy at maturity, the entire head takes on an unusual aspect suggestive of aggregate fruits of Rubus or species of Boraginaceae. This drupe-like fruit is rare in Compositae and known only in a few genera (e.g., Chrysanthemoides Fabr., Calenduleae; and Wulffia Neck, ex Cass., Heliantheae).

Clibadium eggersii is the oldest name for a group of previously described taxa that were based on variations in type and density of pubescence and ratio of ray to disc florets. For example, Clibadium chocoense was described by Cuatrecasas (1954, p. 236) as a new species closely related to C . polygynum but distinguished by the "less dense and adpressed pubescence and by the low number of disc florets". Clibadium pittieri was described by Greenman (1897) as having two different types of pubescence. In some individuals the vesture on stems and peduncles is hispidulous (e.g., Knapp & Mallet 3101. Weddel 2161. Wilbur & Teeri 13423), and in others it is strigose (Duke & Elias 1377 8. Gentry 12409. Neill & Vincelli 3578), but two collections are intermediate in these two types (Hampshire & Whiteford 492. Lewis et al.. 97 5). Forma phrixium was described by Greenman (1904) within C. pittieri based on the hispidulous condition. In my opinion, different types and densities of pubescence do not correlate with any other morphological features or particular distribution or habitat, and therefore do not warrant formal recognition. All the species and forms here considered as synonyms appear to be nothing more than morphological variation within the broad distributional range of C. eggersii. All of them share the same structure and organization of heads and florets.

Clibadium eggersii is very closely related to C. divaricatum and C. leptophvllum. They share a large number of ray florets (24-40) organized in 5-7 series, and 4-lobed ray and 4-5-lobed disc corollas. But C. eggersii differs from C . 1eptophyllum in having an ovate leaf (in contrast to the unusual narrowly lanceolate leaf in C. leptophyllum) and almost one-third the number of disc florets. Clibadium eggersii differs from C. divaricatum in having heads twice as large and with 2 phyllaries in contrast to 5-6 observed in £L_ divaricatum. This species honors botanist Henrik F. A. von Eggers (1844-1903), author of The Flora of Saint Croix and the

Virgin Islands (1879; Stafleu & Cowan, 1976).

2. Clibadium leptophyllum Cuatrec., Rev. Acad. Colomb. Cienc. Exact. 9: 237. 1954.--Type: Colombia. Choco: Rio San Juan, cercanias de Palestina, 14 Mar 1944, Cuatrecasas 16941 (holotype; F!, photo: F!; isotypes: COL! F ! U S !).

Shrub densely branched, stem elongated at nodes, densely strigose with straight hairs of 0.3-0.4 mm long. Leaves sessile with blades linear to linear-lanceolate, 10-16 cm long, 2-3 cm wide, tripliveined, at apex acuminate, at base narrowly attenuate, amplexicaule, with margin crenate, remotely serrate and distantly revolute, with both surface sparcely strigose with hairs up to 0.2 mm long. Capitulescence thyrsoid, with 10-25 heads; peduncles 2 mm long. Heads radiate, 3 mm high, 2.5 mm diameter; involucre subglobose, 2.5 mm diameter; phyllaries 2, ovate, 2.5 mm long, 2 mm wide, 5-7 veined, sparcely strigose, with hairs 0.5 mm long, upper one-third ciliate, at apex acute. Ray florets 30, subtended by ovate-lanceolate pales, 5-veined, 2,5 mm long, 1.5 mm wide, scarcely strigose at apex, with hairs 0.1 mm long; corollas white, glabra, 2.5 mm long, with tube 1.5 mm long, 0.4 mm in diameter, 4-lobed, with lobes deltate, 0.6 mm long, scarcely pubescent with hairs 0.1 mm long; styles l mm long, with branches 0.1-0.2 mm long. Disc florets 24, subtended by narrowly-lanceolate pales, 2.2 mm long, 0.5 mm wide, 3-veined, sparcely strigose with upper one third ciliate with hairs 0.1 mm long; corollas 1.5 mm long, with throat 1 mm long, 0.5 in diameter; 4-5 lobed, with deltate lobes, 0.3 mm long, scarsely strigose at apex, with hairs 0.1 mm long; anthers black l mm long; style 1-1.2 mm long, with branches 0.5 mm long; ovaries sterile, 1 mm long, pilose at apex with hairs 0.1 mm long. Achenes pyroide, 3 mm long, 2 mm wide, compressed, base attenuate-stipitate, hirsute on upper one-third, with hairs 0.1-0.2 mm long.

Chromosome number unknown. Phenology. Flowering from March through June. Distribution and Ecology (Fig. 9). Colombia. This species is known only from the Choco region; tropical rain forest, in secondary vegetation along roadside; 0-500 m.

Additional Specimens Examined. Colombia. Choco: Serrania de Baudo, along road between Las Animas and Pato on Rio Pato, ca 1 km Above Pato, Croat 56097 (MO); Anchicaya, Koie 4880 (US); Carretera Panamericana entre el Rio y el Rio Pato, Forero et al. 5527 (COL, MO); Hoya del Rio San Juan, arriba Palestina, Forero et al. 4163 (COL, MO).

Clibadium leptophyllum is easily distinguished by its linear-lanceolate and slightly amplexicaule leaves and small heads resembling drupe-like fruits. The leaf shape, unique among members of the genus, resembles unrelated species such 96

as Trixis pedunculosa Rich. Clibadium leptophyllum, endemic to the Choco region in Colombia, is closely related to C. eggersii with a broad distribution from Belize to northern Peru and C. divaricatum restricted to Northern Peru.

Clibadium leptophyllum differs from its close relatives in Central and South America by having narrow leaves and heads with 24 disc florets in contrast to 6-10 disc florets per head and ovate leaves in C. divaricatum and C. eggersii. Clibadium leptophyllum ia also related to C . erosum and C. terebinthinaceum in the shared multiseriate condition of ray florets, receptacle paleaceous throughout and the thyrsoid capitulescence. Clibadium leptophyllum differs from the relatives, however, in having a capitulescence with fewer than 25 heads, more than 30 ray florets, ray corollas with 4- lobes and ray pales 5-veined, all of which contrast in the Caribbean species with a capitulescence with more than 50 heads, fewer than 20 ray florets, with ray corollas 3-lobed, and ray pales 3 -veined.

3. Clibadium divaricatum S.F. Blake, Contr. Gray Herb. 52: 7. 1917.--Type: Peru. San Martin: Tarapoto, 1855-56, Spruce 4522 (lectotype: BM!, fragment at GH!, photo: OS!; isolectotypes: F! GH! K[2]! photo: OS!, P photo: OS!) . 97

Clibadium rimachii H. Robinson, Phytologia 65: 50. 1988.--Type: Peru. San Martin: Tarapoto, Carretera

de Tarapoto-Yurimaguas, km 12, 23 Aug 197 8, Rimachi 3877 (holotype: US!).

Shrub 2.5 m tall, stems striate, strigose with hairs 0.5 mm long. Leaves with blades ovate-lanceolate; 7-11 cm long, 2.5-4 cm wide, at apex acuminate, at base attenuate, with margin serrate, tripleveined, both surfaces weakly strigose, with hairs 0.5-1 mm long; petiole 1-2 cm long, 1 mm diameter, strigose, with hairs 0.5 mm long. Capitulescence thyrsoid, with up to 25 heads; peduncle 1 mm long. Heads 3-4 mm high, 3-4 mm diameter; involucre cupulate, 3.5 mm diameter; phyllaries 5-6, ovate, 3 mm long, 2.5 mm wide, 5-7 veined, at apex acute, strigose, with hairs 0.1 mm long, upper one-third ciliate. Ray florets 24-39, subtended by ovate pales; pales 3 mm long, 2.5 mm wide, 5-veined, arranged into 5-6 series; at apex acute, strigillose toward apex, upper one-third of margin ciliate; corollas white, 1.5-1.2 mm long, 1 mm wide, irregularly 4-lobed, with lobes deltate, with margin shortly and scarcely ciliate. Disc florets 6-10; corollas 1.5-1.2 mm long, 4-5 lobed, with deltate lobes, 0.4 mm long, with margin scarcely ciliate; throat 1 mm long, 0.5 mm diameter; subtended by lanceolate to filiform pales, 3-veined; at apex ciliate; anthers black, 1-1.2 mm long; ovaries sterile, 1.2 mm long, 0.3 mm diameter, hirsute on upper one-third. Achenes 2 mm long, 1.5 mm diameter, hirsute on upper one- third. Chromosome number unknown. 98

Phenology: Flowering from April through August. Distribution (Fig. 9). Peru, endemic to the Department of Tarapoto; wet montane forest on sandy rocky soil; 800-900 m. Additional Specimens Examined. Peru. Huanuco: Chinchavito, cerca a Cayumba, entre Huanuco y Tingo Maria, Ferreira 6805 (F); between Tingo Maria and Las Cuevas de las Lechuzas.MacRae 118 (F); Tingo Maria, Soukup 2260 (F).

Clibadium divaricatum is considered to belong to subgenus Paleata because of multiseriate ray florets (5-6), large number of ray florets (24-39, usually 4-6 times the number of disc florets), and with globose and paleaceous heads with fleshy and juicy phyllaries. It differs from C. eggersii and C. leptophyllum. also of the same subgenus, in having small heads (2-4 mm tall) surrounded by 5-6 phyllaries (three times the number observed in the other two species), and also in having gray light-green leaves (in contrast to the dark green leaves of its relatives).

The single collection, Rimachii 3877. was initially described as C. rimachii by Robinson (1988), based primarily on short pedicels (similar to those in C. pediculatum Aristeguieta). Critical examination of the type of C. rimachii. however, reveals that it is a minor variation of C. divaricatum (both type specimens are from exactly the same locality in the Department of Tarapoto, Peru). The capitulescence arrangement, head organization, and number 99 and structure of ray and disc florets are the same, except for slightly larger heads (4-5 mm) in C. rimachii in contrast to the smaller heads (2-3 mm) in C. divaricatum.

Blake (1917) listed three type specimens in the protologue of C. divaricatum (Spruce 4522) at BM, GH, and K.

Two more additional type specimens exist, one at F and one at P. The specimen at BM is here recognized as the lectotypetype. It is the only sheet labelled by Blake's handwriting as "type specimen". Also, it contains more complete information on collection date and habitat, and more importantly, it is a superior specimen showing all diagnostic characters.

IB. Clibadium L. section Trixidium DC., Prodr. 506. 1836. --Type: Clibadium erosum (Sw.) DC.

Capitulescence with 50-100 heads; ray florets fewer than

20; ray corollas 3-lobed; ray pales 3-veined; Caribbean and

Lesser Antilles. Species 4-5.

4. Clibadium terebinthinaceum (Sw.) DC. Prodr. 5: 5 06. 1836. Trixis terebinthinacea Sw., Prodr. 115.

1788.--Type: Jamaica (holotype: G-DC, IDC microfiche 28.925:11.1,2!). 100

Clibadium alexandri Griseb., Flora Brit. W. Ind. 30:368. 1864.--Type: Jamaica. S. Anns at Moncague, Gay's Hill, 29 Jul 1850, Prior s.n. (lectotype here designated: K!, photos: GH! OS!; isolectotype: K!,

photo: OS!).

Shrub up to 3 m tall, stems woody toward base, striate, strigose with hairs 1 mm long. Leaves with blades broadly ovate, 10-20 cm long, 4-14 cm wide, tripliveined, at apex acute to acuminate, at base obtuse, attenuate, with margin finely serrate to crenate-serrate, with upper surface glabrate to strigose, with lower surface strigose with hairs 0.5 mm long; petioles 3-5 cm long, striate, strigose with hairs 0.5 mm long. Capitulescence paniculiform with less than 100 heads loosely clustered in terminal open branches, peduncles 0.5 mm long. Heads radiate; 5 mm high, 4 mm diameter; involucre cupulate, 4 mm diameter; phyllaries 3, ovate to ovate-suborbicular, 4 mm long, 3 mm wide, light- green, 5-7-veined, abaxially glabrate to strigose, with hairs 0.5 mm long, margin upper one-third ciliate. Ray florets 12-

16, subtended by ovate pales, 3.5-4 mm long, 3 mm wide, 3-5 veined, strigillose toward apex; corollas white, 2.5 mm long, 0.5 mm diameter, 3-lobed, with lobes deltate, 0.5 mm long; styles 2.5 mm long, with branches 1.5 mm long. Disc florets 5-7, subtended by lanceolate pales, 3-veined; corolla white, 3 mm long, with throat 1 mm diameter, 5-lobed, with lobes 0.4 mm long; styles undivided, 2 mm long, with branches 0.5 mm long. Achenes obovoid, black, 3 mm long, 2 mm wide, villous 101 on the upper one-third. Chromosome number unknown. Phenology. Flowering from December to April. Distribution (Fig. 10). Endemic to Cuba, Jamaica and Tobago; ridge-type vegetation in secondary forests; 200-1500 m.

Representative Specimens. Cuba: Oriente: Sierra Maestra, above Firmeza, Ekman 168 (A, NY); Southern Oriente and Pico Turquino, Leon 10880 (NY). Sevilla: near Santiago,

Taylor 318 (NY). Jamaica. Clarendon: James Hill Savanna, Proctor 10341 (NY, US). Saint Andrew: Second-breakfast Spring, above Mount , Proctor 8313 (NY, US); Hardwar Gap, Yuncker 17 624 (F). Saint Catherine: Resource, Mountain

Diablo region, Hespenheide 1119 (DS, F, GH, MO, US). Portland: Vicinity of Mill Bank, Maxon & Killip 197 (F, GH, NY, US); Trail from Morces Gap to Vinegar Hill, Maxon & Killip 1323 (F, GH, NY, US); Blue Mountains, Stony River to

Nanny Town, Morley & Whitefoord 480 (A, MO); Nanny Town site, Morley & Whitefoord 49 3 (A, MO); near Clydesdale, Redher s.n. (GH). Arntully, Orcutt 3113 (CAS, DS, US); along trail up to Joe Hill, John Crow Mountains, above Moore Town, Yuncker 18827 (F). Saint Thomas: trail from Bath to Corn Puss Gap, Anderson & Sternberg 3301 (GH, US); vicinity of Cinchona, Chesternale to Silver Hill, Britton 357 (NY) ; Cockpit Country, Rocky wooded hillside, Britton 554 (NY); Mansfield, Rocks Devil's River, Britton 357 9 (NY); John Crow Mountains, Big Level, Britton 397 6 (NY); Union Hill, near Moneague, Britton & Hollick 2810 (NY); above Bowden Pen, along foot 102

'S ___

Fig. 1 0 . Distribution of Clibadium erosum ( ▼ ) and C. terebinthinaceum (®) in the Caribbean. 103 path to Bath via Cuna Cuna Pass, Crosbv & Anderson 1045 (F, LL, NY, UC); Tyres, near Troy, Harris 3183 (NY); Cinchona, Blue Mountains, Harris & Lawrence 15157 (UC); Hitchcock s.n. (MO); above Farm Hill Works, Orcutt 3324 (MO, US), Proctor 6864 (US). Tobago. Roxborough-Parlatuvier road, main ridge,

Cowan 1418 (GH, NY, US).

Clibadium terebinthinaceum is closely related to C. erosum. also from the Caribbean. Both species have ovate to broadly-ovate, 5-7-veined phyllaries, with obtuse apices, 3- lobed ray corollas, 3 -5-veined ray pales, and 3-veined disc pales. Clibadium terebinthinaceum differs by having glabrous phyllaries, 5-7 disc florets (half the number in C. erosum), 5-lobed disc corollas, and villous achenes.

The name C. terebinthinaceum has been erroneously applied to specimens of C. arandifolium. ranging from Costa

Rica to Colombia. This latter species is characterized by not only having large leaves, but also by having twice the number of disc florets than ray florets. Schulz (1912) suggested that due to similarity of head arrangement, C. terebinthinaceum should be considered a variety of C. pittieri (now C. eggersii) which is known only from mainland

Central and South America. These taxa, however, are quite distinct. 104

5. Clibadium erosum (Sw.) DC., Prodr. 5: 506. 1836.-- Type: Trixis erosa Sw., Prodr. Veg. Ind. Occ. 115.

17 88.--Type: Dominica. St. Kitts, 1826, Sieber 617 (holotype: BM!; isotype: G-DC, IDC microfiche 28.925:11.3,4!).

Clibadium fragiferum Griseb., Fl. Brit. W. Indian Is. 368. 1861.--Type: St. Vincent, 1817 [Urban, 1903]

Guilding s.n. (holotype: K!, photo: OS!).

Shrub or small tree to 5 m tall; stems woody at base, branches and twigs densely pubescent, with hairs 1 mm long.

Leaves with blades broadly ovate to ovate-oblong, membraneous, 8-25 long. 8-12 cm wide, tripliveined, at base attenuate, at apex shortly acuminate, with margin erose, coarsely and finely incised and serrate, larger teeth to 8 mm long, deltate, acute, densely strigose in both surfaces, with hairs 1 mm long; petioles 3-6 cm long. Capitulescence thyrsoid or paniculiform, 8-15 cm long. Heads subglobose, 5 mm high, nearly sessile or shortly pedicellated; involucre cupulate, 4 mm diameter; phyllaries (3)-4, ovate, 4 mm long, 3 mm wide, 5-7 veined, strigose with hairs 0.1 mm long, at apex obtuse, upper one-third ciliate. Ray flowers 12-24, subtended by ovate pales, 4 mm long, 3 mm wide, arranged into 3-5 series, 3-5 veined, abaxially strigose, upper one- third ciliate; corolla white or purple, 2.5 mm long, glabrous, 3-lobed, with lobes deltate, 0.5 mm long, shortly ciliate at margin; style 4 mm long, with branches 1 mm long. Disc florets 8-(14), subtended by ovate-lanceolate pales, 3 mm long, 3-veined, margin upper one-third ciliate; corolla tubulate, 3 mm long, 4-lobed, with lobes 0.5 mm long, at apex hispidulous; anthers black, 1.5 mm long; ovaries sterile, 2.5 mm long, at apex pilose, style undivided, 3 mm long. Achenes obovate, 2.5 mm long, 1.5 mm diameter, rounded at base, upper one-third hirsute, with hairs 0.5 mm long. Chromosome number, n = 16.

Phenology. Flowering from December through June. Distribution (Fig. 10). Lesser Antilles (Dominica, Guadeloupe, Martinique, Monserrat, Nevis, St. Kitts, St. Lucia, St. Vincent), Puerto Rico; secondary rain forest; 700-1500 m.

Representative Specimens. Dominica: Lisdara Estate, Proctor Cooper III 17 0 (NY); Saint George, S slope of Morne Macaque on road to Fresh Water Lake, Ernst 1088 (A; GH; US); summit of Morne trois Pitons, Ernst 2035 (US); upper slope Morne Trois Pitons, Hodge 7 36 (GH; NY); rainforests on the precipitous slopes of Morne Colla Anglais, Sylvania, Hodge

1037 (GH); mountains along Castle , vicinity of N base of Trois Pitons, Hodge 1201 (GH); forests upper slopes Morne Trois Pitons, Hodge 1396 (GH); woodlands about South Chiltern Estate, Hodge 1462: slopes Morne Anglais, Hodge 2241 (GH); Lisdara, Hodge 2380 (GH); forest border between Laudat and Freshwater Lake, Hodge & Hodge 187 0 (GH); near Grande Bay, Eggers 619 [see Urban (1902), for clarification of collector and date](UC; CAS); Freshwater Lake, Whitefoord 5143 (A; US); Bellevue, King 6305 (US); Glasham, King 6396 (US); Morne Diablotin, elfin woodland at the peak, Webster 13352 (US); vicinity of Fresh Water Lake, near Ladaut, Smith 10245 (NY; UC); along the old trans-island road at the Fresh Water Lake, SE flank of Morne Micotrin, Chambers 27 38 (GH; US); Morns Trois Pitons on NW slope, Chambers 27 56 (US); Deux Branches, trans-island highway cross to the Pagua River, Chambers 277 0 (US). Saint Joseph Parish, Layou Valley Road, Nicholson 4098 (A; US). Saint Peter, Morne Diablotins, at summit, dwarf Elfin Woodland, Wasshausen & Ayensu 416 (US). Guadeloupe: Basse Terre on road to Soufriere above Sainte Claude, Hespenheide & Wiseman 533 (GH); Basse terre, Pres de l'Stang as de Pique, Sastre & Clairou 2626 (MO); from Bains Jaunes on trail to Soufriere, Howard 11824 (A) ; forests above Bains

Jaunes, Howard 19387 (A; US); forests de Bains Jeunes, Stehle 86 (NY). Martinique: Saint Denis, Bois de Fonos, Pere Puss 316 (NY). Monserrat: crest of Chance's Mt, Howard 11911 (A); Chance's Mt, site of Pan Am crash, Howard & Howard 17 533 (A) ; Blakers, in the foothills, Shafer 485 (NY) . Nevis: Iron Gate, below Dodans Water Resource, Proctor 19 605 (A) ; Neavis Peak summit and adjacent ridge, Smith 10530 (A; NY, UC). Saint Kitt's: Auckley Estate, Britton & Cowell 17 9 (NY); summit area of Chance's Mountain Soufriere Hills, Proctor 19116 (A) . Santa Lucia: Quilesse, Beard 1157 (F, UC) . Saint Vincent: The Sanfriere, Beard 1360 (NY); Mount , Morton 594 6 (F). Puerto Rico: Mts. Oro Negro, Dona Juana, Alain 9422 (NY); Monte Cerrote, near Adjuntas, Britton & Brown 5420 (F, MO, NY, US); Catalina-Yunque Trail, 107

Luquillo Mountains, Britton & Bruner 7 586 (NY); Sierra de Naguabo, Barrio de Maizales, Britton & Shafer 2132 (NY, US); Monte Torrecilla, Brown et al.. 5530 (US); along the Guayama, Cayey Road, Britton et al. 6454 (NY); Luquillo National Forest, El Yunque, Mt. Britton area, Gentry & Zardini 50363 (MO); Caribbean National Forest, 7 mi N of Rio Blanco along Hwy 191, Hansen et al. 9422 (MO); Pico del Toro, Luquillo Forest, Hill 89 (NY); Cordillera Central, Bosque Toro Negro, near km 21 HM 3, Judd 525 (A, GH); Luquillo Experimental

Forest, Knight s.n. (WIS); near junction of routes 179 and 184, King & Garvey 7 012 (US); 2-4 mi SW of El Verde

Experimental Station, Little 13064 (A, F, US); El Yunque, Luquillo Mountains, Little 13554 (US); Summit cerro de Punta, Little 16320 (NY, US); El Yunque, El Hicaco, Otero 586 (F,

MO); Caribbean National Park, Sierra de Luquillo, Pico Este, Sauleda et al. 87 84 (F); Luquillo Insular Forest, El Yunque, Sargent B98 (US); El Yunque Resort, Schubert & Winters 419 (A, GH); Rio Piedras, Sierra de Naguabo, Stevenson 1558 (NY); Reserva Forestal de Toro Negro, about 2 mi N of cerro Maravilla, Stimson 1533 (GH); base of Cerro Punta, Stimson 1571 (CAS, GH, MO, NY); M t . Britton, Torborgh 346 (A); km 21 on Palmer to Florida, Wagner 9 8 (A, GH); at the summit of Mt. Britton, Wagner 116 (A); km 19 on route 191 in Luquillo Mts., Wagner 97 0 (MO, WIS); Luquillo Mountains, Wilson 150 (US).

Clibadium erosum is easily recognized by its peculiar erose leaf margin, which is unique within the genus. It is closely related to C. terebinthinaceum of Section Trixidium. 108

which is also restricted to the Caribbean and Lesser Antilles. They are separated from section Eggersia by having a capitulescence with more than 50 heads, less than 20 ray florets in each head, 3-4 phyllaries, 3-lobed ray corollas, and 3-5 veined ray pales, which contrast with fewer than 25 heads per capitulescence, heads with more than 20 ray florets, 2 or 5-6 phyllaries, 4-lobed ray corollas, and 5- veined ray pales in section Eggersia. Clibadium erosum differs from C. terebinthinaceum by the former having ovate and strigose phyllaries, twice the number of disc florets, 5- lobed disc corollas, and pilose achenes.

Clibadium fragiferum was described by Grisebach (1861) to include populations restricted to St. Vincent and Guadeloupe, and it was placed by Schulz (1912) within Section Trixidium together with C. eggersii. C. erosum, and C. terebinthinaceum. Schulz separated C. fragiferum from its close relative, C. eggersii. mainly by the presence of 6 phyllaries (probably the "6-folium" involucre described included the outermost series of pales as phyllaries). Although the original description is incomplete, the holotype and additional collections clearly indicate that C. fragiferum is one extreme of morphological variation for the most southern distribution of C. erosum in the Lesser

Antilles. 109

II. Clibadium L. subgenus Clibadium Euclibadium DC.. Prodr. 5: 505. 1836. --Type: Clibadium surinamense L.

Shrubs with racemose, cymose or thyrsoid capitulescences with more than 100 heads; heads mostly epaleaceous (only ray florets with pales); ray florets uniseriate; number of ray florets same or fewer than disc florets; mature achenes dry; phyllaries and pales never fleshy at maturity.

IIA. Clibadium L. section Glomerata Arriagada, sect. nov.--Typus: Clibadium glomeratum Greenm.

Folia late ovata; capitula in glomerulis densissimis; phyllaria 4-12 nervia; flores feminei 3-10, corollis 2- vel 4-lobatis, paleis 7-12-nervibus; flores hermaphroditi disci 4-10, corollis quinque lobatis, paleis uninervis.

Leaves broadly ovate; heads arranged in condensed capitulescences, with pales subtending ray florets only; phyllaries 4-12-veined; ray florets 3-10, with corollas 2- or 4-lobed, and pales 7-12-veined; disc florets 4-10, with corollas 5-lobed, and pales 1-veined. Species 6-14.

6. Clibadium anceps Greenm., Proc. Amer. Acad. Arts 39: 97. 1903.--Type: Costa Rica. "Forets de La Palma", 1459 m, 8 Sep 1898, Tonduz s.n. [= 12537 CR accession 110

number] (holotype: CR!, photo: US!, fragment at GH! : isotype: K!, photos: OS! US!).

Clibadium pilonicum Stuessy, Ann. Missouri Bot. Gard.

62: 1073-1074. 1975.--Type: Panama. C o d e mountains N of de Anton, 28 May 1967, Lewis. MacBryde. Oliver & Ridgway 1745 (holotype: MO!; isotype: UC!).

Arching or scandent shrub, 3-6 m tall; stems 2-5 cm diameter, running horizontal and rooting at the nodes, glabrate below to densely strigose above, with hairs 2 mm

long. Leaves with blades ovate to narrowly ovate, 7-15 cm long, 3-9 cm wide, at apex acuminate, at base cuneate, with margin serrulate remotely denticulate more or less revolute;

conspicuously 3 to 5-veined from near base, both surfaces glabrate to sparsely strigose with hairs up to 0.3 mm long; petioles less than 3 cm long, 1-1.5 mm diameter, strigose with hairs 0.2 mm long. Capitulescence at twig tips of 50- 120 heads, thyrsoid, with clusters of 3-6 sessile heads terminating all axes. Heads radiate, 3-4 mm high, 3-4 mm diameter; involucre funnelform, 3-4 mm diameter; phyllaries 6, ovate, 4 mm long, 2.5-3 mm wide, arranged in two series, abaxially strigose with hairs 0.3 mm long, the upper one- third of margin ciliate, at apex acute, 7-12 veined. Ray florets 3-5, subtended by ovate pales, 3 mm long, 2.5 mm wide, 7 -veined, at apex acute, with margin ciliate on upper- one-third; corollas white, 2 to 2.5 mm long, 1 mm diameter, 4-lobed, lobes 0.5 mm long, deltate, shortly ciliate at Ill margin; styles 3 mm long, with branches 1.5 mm long. Disc florets 4-10, epaleaceous, corollas 3 mm long, 5-lobed, lobes deltoid, 0.4 mm long; anthers black, 2 mm long; styles 3 mm long; ovary sterile, 2 mm long, 0.2 mm diameter, villous toward apex, with hairs 0.5 mm long. Achenes 2 mm long, 1.3 mm diameter, glabrous. Chromosome number, n = 16. Phenology. Flowering from May through August. Distribution (Fig. 11). Costa Rica and Panama; tropical rain forest, mainly in cut-over areas in wet secondary forest along roadside; 700-2200 m.

Representative Specimens. Costa Rica. Alajuela: along road to Finca Los Ensayos off Highway 15 ca 7.5 mi. N of , Croat 4347 0 (MO), along road between Pto. Viejo and

San Jose, 3 m N of Cariblanco, Croat 68217 (UC); Reserve, Atlantic slope, Rio Penas Blancas valley, Haber 7 37

(UC); near La Laguna 6-8 km S of Villa Quesada, Molina et al. 17508 (GH); along road from San Ramon northward through Balsa, Stevens 14136 (F, MO); 11 mi S of Quesada, Webster et al. 12198 (MO); 19 km N. of San Ramon, Wilbur & Stone 1067 8 (OS, MO). Cartago: 3-7 km beyond bridge in Tapanti, Almeda et al. 2987 (CAS); Tapanti Hydroelectric Reserve along Rio Grande de Orosi, Croat 36089 (UC); El Muneco, S of Navarro, Standley 33652 (GH, US); 10 km SW of Navarro, Wilbur & Stone 1057 5 (CAS, CR, GH, MO, OS); along a stretch of 9.3 km begining 3 km from bridge at Tapanti, Wilbur et al. 22349 (CAS); along Rio Grande de Orosi 11 km SSE of bridge at Tapanti, Wilbur et al. 22460 (CAS). San Jose: Cordillera Fig. 1 1 . Distribution of Clibadium anceos ( C. qlomeratum ( ▼ ) , and C... trianae ( A ) in Central America and northwest South America. Central, 7.5 km from San Jeronimo (N. of San Jose) Arriagada

408 (OS), between Bajo La Hondura and Alto La Palma, et a l . 4003 (CR); Alto de La Palma de San Ramon, Brenes 3817. 6335. 20586 (F), Rio Santa Clara Valley below La Palma NE San Jeronimo, Burger et al. 94 06 (F, OS); Basamento de Fila Lleskila, Gomez et al. 23246 (UC); 13 km NW of San

Isidro del General, Jimenez 2224 (CR, F); vicinity of El General, Skutch 2657 (GH, MO, NY, US), Vara Blanca de

Sarapiqui, Skutch 3260 (GH, MO, NY, US); Heredia: Rio Vueltas, E slope of Volcan near the Continental Divide, Burger & Liesner 6435 (F); Rio Vueltas on eastern slope Volcan Barva, Burger & Gentry 9048 (F); along route 113, 2 km SW of Alto Gallito, King & Guevara 6805 (MO); road between

San Rafael and Rios Las Vueltas, Stevens 13986 (UC). Puntarenas: 2.5 km NO from Estipulas, Arriagada 135b (OS); 6 km S of de Java Finca Las Cruces, San Vito de Java, Burch 4656 (MO); Monteverde, Veracruz River Valley S of Reserve, Haber et al. 53 62 (MO); Monteverde Cloud Forest Reserve, Feinsinger et al. 812093 (MO); Monteverde, Jimenez 1269 (CR); Manuel Antonio Parque Nacional, Punta , Khan et a l . 450 (BM); Monteverde, Cloud Forest Reserve, Koptur 177 (UC); 6 km S of San Vito de Java around Wilson's finca, Raven 21977 (F), 17.5 mi N. of Villa Neily on road to San Vito, Stuessy & Gardner 4517. 4518 (OS); 3 m W of San Vito, Sullivan 37 5 (FSU). Panama. Bocas del Toro: Carretera del Oleoducto, IRHE Fortuna Hydroelectric Project, Knapp & Vodicka 5633 (MO, UC). Chiriqui: Cerro Horqueta, Blum & Dwyer 2662 (FSU, MO), E side of Cerro Pando, Darcy & Darcv 114

6611 (MO); Cerro Colorado, road to Bocas del Toro, Folsom et al. 4753 (UC); Fortuna Dam site, top of the mountain, Folsom et al. 537 8 (UC); Palo Alto, E of Boquete, Stern et al. 1019 (GH, MO, US); Fortuna Dam area, along oleoducto road, Todzia et al. 2549 (CAS, MO); vicinity of Gualaca 11 mi from Planes de Hornito, Antonio 5041 (UC); E of road flowing into Rio Hornito near Quebrada Moro, Churchill 5485 (UC); vicinity of Fortuna Dam in Valley of Rio Chirigui, Croat 66523 (UC), Croat 66554 (UC); Fortuna Dam N of reservoir, D'Arcy & Todzia 15970 (UC, MO) . Code: Sawmill 7 km NE of El Cope, Antonio 1146 (UC); slopes Cerro Pilon near El Valle, Duke 12116 (MO, OS); Cerro Pilon, Dwyer & Lallathin 8669 (MO); Cerro Gaital Caracoral, Dwver & Correa 8844 (MO); 5.4 km along La Mesa from El Valle main road, Folsom 3648 (MO); Summit at Alto Calvario, Folsom & Robinson 2422 (UC); along road from La Pindeda to El Cope by way of Piedras Gordas, Hammel 2599 (UC); N side of Cerro Gaital near start of trail, Hartman 3963 (OS); Cerro Brewster on San Bias border, McPherson 7 529 (MO); above El Potroso sawmill at Continental Divide, Sytsma

1803 (UC).

Clibadium anceps is distinguished by lateral branches of the capitulescence forming more or less right angles to the main axis, fleshy phyllaries at maturity with a greenish juice, and the scandent condition. It is related to C. glomeratum and C. trianae forming a more or less homogeneous group within section Glomerata. These taxa are characterized by completely lacking disc pales. Clibadium anceos differs 115 from its close relatives, C . glomeratum and C . trianae. by fewer aggregated heads (see comments under the above- mentioned species), ovate and strigose phyllaries, 3-5 ray florets, 4-10 disc florets, and glabrate achenes.

Clibadium pilonicum. described by Stuessy (197 5) from Cerro Pilon, in C o d e Province, Panama, differs from typical C. anceps (also collected in the same area; Duke 12116. Dwyer & Lallathin 8669) in having heads more tightly aggregated and narrowly ovate leaves in contrast to broadly ovate leaves more typical for the species. Stuessy (197 5) also separated these two taxa by leaf venation (secondary midrib from the very base versus 5-30 mm above, respectively). After studying available material, I conclude that these diagnostic characters intergrade among specimens according to age and leaf position (e.g., Antonio 5041; Croat 66554; D 'Arcy & Todzia 1597 0: D 'Arcy & D 'Arcy 6611; Folsom & Robinson 2422; Folsom et al. 47 53. 537 8: Hammel 2599: Todzia et al. 2549). Therefore, both taxa can not be separated by macromorphology and are considered here to belong to the same taxon.

7. Clibadium glomeratum Greenm., Proc. Amer. Acad. Arts 39: 98. 1903.--Type: Costa Rica. Cartago: "Forets de Luis", 650 m, Nov 1897, Tonduz 7 330 [= 11508 CR accession number] (lectotype here designated: GH!; isolectotypes: F! GH! K! US [5]!). Shrub 1.5-4 m tall, 8 cm diameter at base, branches densely pilose to tomentose, with hairs 1-2 mm long. Leaves with blades broadly ovate, 10-20 cm long, 5-15 wide, at apex acute, at base obtuse, subtruncate, subcordate or cordate; margin irregularly crenate-serrulate to serrulate-dentate; scabrous pubescent above, densely strigose beneath mainly on veins, hairs 1-2 mm long; petioles 5-8 cm long, 1-1.5 mm diameter, strigose with hairs 0.5-1 mm long. Capitulescence thyrsoid, branches scabrous to tomentose with hairs 1-2 mm long; glomerules 0.5-1.0 cm diameter with 10-20 tightly sessile heads. Heads 3-4 mm high, 2-3 mm diameter; phyllaries 6, orbicular, 3 mm long, 2.5 mm wide, arranged in two series, nearly glabrous below, ciliate on the upper one- third, 9-12-nerved. Ray florets 5, subtended by ovate pales, 3 mm long, 2 mm wide, glabrous, 7 -veined, margin ciliate at upper on-third; corollas 1 mm long, tube 0.6 mm long, 0.5 mm diameter, 4-lobed, with deltate lobes, scarcely pubescent at apex; styles 1 mm long, branches 0.5 mm long. Disc florets 5, epalaceous, corollas l mm long, 5-lobed, lobes deltate, 0.5 mm long, throat 0.6 mm long, 0.3 mm diameter; anthers black, 1 mm long; style undivided, 1 mm long; ovary sterile, villose toward apex with hairs 0.5 mm long. Achenes 2 mm long, 1.5 mm diameter, villous on upper one-third, with hairs 0.5 mm long. Chromosome number, n = 16.

Phenology. Flowering mid-June though late September; fruits maturing from October through early April. Distribution (Fig. 11). Only known from Costa Rica; second growth tropical rain forest, open roadside; from sea 117 level to 500 m (occasionally individuals found up to 1400 m, e.g., Holm & litis 121: Lent 1127).

Representative Specimens. Costa Rica. Cartago: about 7 km NE Santa Cruz on road beyond Agua, Almeda & Nakai 4171

(CAS, OS); 10 km S of Tapanti along E slope above the Rio Grande de Orosi, Burger & Stolze 57 35 (MO); small valley near Santa Cruz, S slope of Volcan , Holm & litis 121 (US); El Muneco, King 67 66 (MO, US); hillside overlooking Rio

Tambor, 3 km E of Cachi, Lent 860 (NY); along roadside between La Jicotea and de Turrialba, Poveda & Saenz 1508 (CR); 5.7 mi S of Orosi on road to Tapanti, Stuessy & Gardner 4492 (OS). Heredia: Yerba Buena NE of , Standley & Valerio 497 62 (US). Limon: above unnamed tributary of Rio Siquirres, Dodge et al. 5603 (GH). Puntarenas: vicinity Delta of the Rio Esquinas, Golfo Dulce, Allen 5637 (CAS); Panamerican Hwy 2, km 37, from 011a Cinco River to Puerto Jimenez, Arriagada 152 (OS); Ricon de Osa near village, toward Panamerican Hwy 2, Arriagada 153 (OS); road from to Panamerican Hwy 2, 1cm 5.9 after La Purruja, Arriagada 154 (OS); slopes above Golfito along the trail to the television tower, Burger & Matta 4744 (NY); 5 km W of Rincon de Osa, Osa Peninsula, Burger & Gentry 9010 (CR); dried stream bed paralleling airport at Rincon, Duke 16117 (OS); Pacific road about 7.5 mi from Rincon near Rio

Riyitos, Peninsula de Osa, Kennedy 1910 (MO, UC); Rincon de Osa, Liesner 1713 (OS); Corcovado National Park, Liesner 2815

(MO); along the camino al Pacifico, W of Rincon de Osa, Osa 118

Peninsula, Raven 20863 (F); 18.5 mi N Villa Neily on gravel road to San Vito, Stuessy & Gardner 4519 (OS); Rincon de Osa, Utley & Utley 105 (OS); Finca Las Cruces, S of San Vito, Webster 21911 (CR); near airstrip Rincon, Osa Peninsula, Weston 4550 (CAS); telecommunication Hill above the town of Golfito, Wilbur et al. 227 85 (CAS). San Jose: Bosquecillos entre las Nubes y Cacajalde Coronado, Gomez-Laurito 5515 (CR); along route 216, ca 2-3 km N of Nubes, King 677 8 (MO, UC, US). Panama. Colon: along Rio Escandalosa, 19 mi E of Transisthmian Hwy on road to Salamanca, Huft & Knapp 1621 (UC) . Clibadium glomeratiim is characterized by having terminal thyrsoid capitulescences with 10 to 20 sessile heads organized in dense glomerules 0.5 - 1 cm diameter. Stems, pedicels and leaves of most of the specimens are densely soft-tomentose (well illustrated in Liesner 2815). The density of pubescence, however, varies to moderate (e.g.,

Allen 5637) or even to sparingly and shortly pubescent (Burger & Liesner 6791). This latter collection also possesses narrowly lanceolate-ovate leaves instead of the more typical broadly ovate ones.

The close relatives of Clibadium glomeratum are C. anceps and C. trianae. These species are placed in Section Glomerata. which includes species with heads arranged into dense glomerules, and with 5-8 ray florets, ray corollas 4- lobed, and disc corollas 5-lobed. Clibadium glomeratum differs from its close relatives in the arrangement and 119 organization of its heads, it has an open and more rounded capitulescence in contrast to a more flat-topped capitate capitulescence in C . anceps. The distribution of these three species also differs. Clibadium glomeratum is restricted to Costa Rica (0-100 m), Clibadium anceps is found in Costa Rica and Chiriqui province in Panama (1200-2100 m ) , and C. trianae occurs in Colombia y Ecuador (1000-2300 m).

Greenman (1903), in the protologue of C. glomeratum. mentions two type specimens (GH and CR) which are indeed labelled with his handwriting and with "n. sp." Additional uncited syntypes without Greenman's annotations also exist at F, K, and US. I choose the specimen at GH as lectotype because it is specifically cited in the protologue, labelled as "n. sp." in Greenman's hand, and also because it is a more complete specimen.

8. Clibadium trianae (Hieron.) S. F. Blake, Contrib. Gray Herb. 52: 6. 1917. Desmanthodium trianae Hieron., Bot.

Jahrb. Syst. 19: 52. 1894.--Type: Colombia. Bogota, Tenasuia, 1853, Triana 1317 (holotype: B, presumed destroyed, photos: F! GH! OS! TEX!; isotypes: BM!, fragment at GH! photos: GH! US!, K!, p photo: OS!, US!).

Clibadium subsessilifolium Hieron.. Bot. Jahrb. Syst. 29: 32. 1900--Type: Ecuador, "Crescit in selvis

subtropicis et subandinis," ca. 1894 [Hieronymus, 1901], 120

Sodiro 22/1 (holotype: B, presumed destroyed, fragment at US!, photos: F! GH! MO! NY! OS!).

Clibadium sychnocephalum S. F. Blake, Contrib. U.S.

Natl. Herb. 603. 1924.--Type: Colombia. Cauca: near Rio Flautas, 26 Jan 1906, Pittier 1211 (holotype: U S !; isotype: F!, photo: US!).

Clibadium congestum Cuatrec., Rev. Acad. Colomb. Cienc. Exact. 9: 237. 1954.--Type: Colombia. Antioquia: 10 km E. de Sonson, subparamo, 18 Mar 1949, Scolnik. Barva. Lopez & Barkley 224 (holotype: F!, photo: US!).

Arching branched shrub, 3-4 m tall, stem 3 cm diameter, branches subangulate forming a right angle, tomentose with hairs 1-2 mm long. Leaves with blades ovate-lanceolate to widely ovate, 10-18 cm long, 8-13 cm wide, at apex acuminate, at base atenuate exceptionally rounded and truncate; both surfaces strigose with hairs 1 mm long, at margin irregularly dentate-serrate to serrate, teeth ending in a shortly mucro; petioles 2-6 cm long, strigose with hairs 1 mm long. Capitulescence thyrsoid, branches scabrous to lightly strigose with hairs 1 mm long. Heads radiate, organized in globose glomerules, 1.0 to 1.5 cm diameter with 20-30 tightly heads. Heads 5 mm high, 4 mm diameter; phyllaries 6, ovate, 4.5 mm long, 3.5 mm wide, nearly glabrous below, margin ciliate on upper one-third, 9-12 veined, acute at apex. Ray florets 5, subtended by ovate pales, 4 mm long, 3 mm wide, 7- 121 veined, glabrous, ciliate at upper one-third; corolla white, 3.5 mm long, tube 3 mm long, 2 mm diameter, 4-lobed, lobes deltate, shortly ciliate at margin; styles 3 mm long, branches 1.5 mm long. Disc florets 5-6, epalaceous, corolla 3.5 mm long, throat 2.5 mm long, 1 mm diameter, 5-lobed, lobes deltate, with margin scarcely ciliate; anthers black, 2.5 mm long, style 2.5 mm long, undivided; ovaries sterile, 2 mm long, 1.5 mm diameter, pilose toward apex, with hairs 1-2 mm long. Achenes obovate 2.5 mm long, 1.8-2 mm diameter, villose on upper one-third, with hairs 1.5 mm long. Chromosome number, n = 16. Phenology. Flowering from May to September. Distribution (Fig. 11). Colombia, Ecuador and

Venezuela, in secondary growth moist tropical forest, mostly in cut areas on roadside (sometimes on rocky slopes); 1000- 2200 m.

Representative Specimens. Colombia. Antioquia: Municipio de Frontino km 18.5 of road Nutibara-Murri, Zarucchi et a l . 5566 (MO). Cundinamarca: W of Bogota on road to El Colegio, Barclay et al. 3299 (COL); Cordillera Oriental, Salto de , Cuatrecasas 61 (COL; F); Cuatrecasas 8182 (COL; F; US); Hauuht 6488 (US); Killip 33982 (BM); Philipson et a l . 2407 (F); Eastern Cordillera, ca 18 km ENE of La Mesa, Kina et al. 5824 (US); near km 8 on road to El Salto from El Colegio, Stuessy et al. 5528 (WIS). Putumayo: vertiente oriental de la Cordillera entre El Silencio y La Cabana, Cuatrecasas 11521 (COL). Tolima: 122

Cordillera Central, Hoya Rio Combeima, Canon del Combeima, El Silencio, Cuatrecasas et al. 27 639 (US). Ecuador. Pichincha: 30 km W of San Juan on gravel road to Chiriboga, Stuessv & Arriagada 12337 (OS); 1 km E of Chiriboga on road to Quito, Stuessy & Arriagada 12347 (OS); 9 km E of Tandapi on road to Aloag, Stuessy & Arriagada 12353 (OS), 25 km E of

Tandapi on route to Aloag, Stuessy & Arriagada 12355 (OS). Venezuela. Tachira: ad ripam dextram rivi Tachira ad limina reipublicae Colombiae, Charoin & Jacquemoud 1327 4 (F); W forested slopes along Quebrada Agua Azul, over slate-shale substrate, S of El Reposo, 14 km SE of Delicias, Stevermark &

Liesner 118392. 118648 (MO)

Clibadium trianae has sessile heads densely aggregated in ternately arranged glomerules. Also it has achenes with fleshy exocarp releasing an orange juice when squeezed. Its general morphology, particularly the densely aggregated small heads, relate it to C. anceps, C. glomeratum. and C. sessile (also pointed out by Blake, 1917), all of which are now grouped in Section Glomerata.

Clibadium congesturn, described by Cuatrecasas (1954) and suggested to be a close relative of C. trianae is clearly a morphological variation of the latter. Within the same population some individuals show tomentose stems, petioles and leaves, which were key features for the delimitation of C. congestum (e.g., Stuessv & Funk 5680). Clibadium trianae 123 has mature leaves with a serrate margin with a tiny mucro at the apex of each tooth. Also common are branches forming a right angle to the stem. The heads (usually 5-7 per cluster) are arranged in different degrees of condensation.

Clibadium subsessilifolium, a clear synonym of trianae. was described originally by Hieronymus as a distinct species mainly on the basis of large numbers of phyllaries (10). Dissection of one head from the type fragment indicates that the number of phyllaries is 5-5 (which coincides with the number commonly found in C. trianae). Clibadium trianae was originally referred by Hieronymus to Desmanthodium: Blake (1917) corrected the misplacement.

An herbarium name, authored by Schulz-Bipontinus, and commemorating H. Karsten (1817-1908) exists (Karsten s.n.). The material, collected in Colombia, Paramo de Bocona, consists of one simple leaf, and a capitulescence with a condensed cluster of heads. The name was apparently never published.

Clibadium trianae commemorates Jose Jeronimo Triana (1834-1890), Columbian botanist who worked on the Prodromus

Florae Novo Granatensis (1862-67; Stafleu & Cowan, 1986; Diaz-Piedrahita, 1991) . 124

9. Clibadium sessile S. F. Blake, Ann. Missouri Bot. Gard. 28: 475. 1941.--Type: Panama. Chirigui: near Bajo Chorro, 20-22 Jul 1940, Woodson & Schery 658 (holotype: US!; isotypes: GH! MO!).

Clibadium subauriculatum Stuessy, Ann. Missouri Bot. Gard. 62: 1074. 197 5.--Type: Panama. Chiriqui, Bocas del Toro: Robalo Trail, N slopes of Cerro Horgueta, 5-7 Aug 1947, Allen 497 0 (holotype: GH!; isotypes: BM! MO!).

Shrub 4 m tall; stems sometimes prostrate, glabrous to strigose with hairs 1 mm long. Leaves with blades narrowly ovate; 8-20 cm long, 5-11 cm wide, broadest near the middle; at apex acuminate, at base cuneate, with margin serrate, upper surface glabrous, lower surface strigillose, with hairs 0.5-1 mm long; petioles 2-6 cm long, 2 mm diameter, winged, wings decreasing toward nodes to become subauriculate at base. Capitulescence paniculiform, capitate, with 7 0 to 100 sessile heads. Heads radiate, 3.5-4.5 mm high; involucre cupulate, 3-4 mm diameter; phyllaries 3, ovate to obovate, 3-

4 mm long, 2.5-3 mm wide, 7-12-veined. abaxially glabrous, at apex obtuse, with margin upper one-third ciliate. Ray florets 5, subtended by ovate to oblanceolate pales, 7-9 veined, strigose abaxially with hairs 0.5 mm long and shortly ciliate at apex; corollas white, 2 mm long, 0.5 mm diameter, 2-lobed, lobes irregularly deltate, 0.3 mm long, at margin shortly ciliate; styles 2 mm long, branches 1 mm long. Disc florets 6-8; corollas white, 3 mm long, throat 2 mm long, 1 125 nun diameter, 5-lobed, lobes deltate, with margin scarcely pubescent, 0.5 mm long; anthers black, 1,6 mm long; styles 3 mm long, ovaries sterile, pilose toward apex with hairs 1 mm long. Achenes 2 mm long, 1.5 mm diameter, glabrous. Chromosome number unknown. Phenology. Flowering from July to August. Distribution (Fig. 12). Panama; rare, only in and around Cerro Horqueta; lower montane rain forest on open hillsides; 1400-1900 m.

Specimens Examined. Panama. Chiriqui: Cerro Horqueta along high trail to summit, Cochrane et al. 6287 (WIS); Cerro Horqueta Cordillera de Talamanca above Boquete, Cochrane et al. 6289 (MO, WIS); Cerro Colorado 30 km above bridge over Rio San Felix, Croat 37102 (UC); Cerro Colorado 34.1 km beyond bridge over Rio San Felix, Croat 37 311 (MO); Bajo

Chorro, D'Arcy 10911 (MO); road from Alto Quiel to Cerro Punta, near Bajo Chorro, Hartman 3916 (OS).

Clibadium sessile is an endemic species restricted to Chiriqui province in Panama. It is specifically similar to C . glomeratum from Costa Rica and Panama in having tightly clustered heads, but the former differs by having winged petioles with a subauriculate base that makes C. sessile easily distinguishable. Winged petioles are also observed in specimens of Clibadium laxum from Ecuador (e.g., Stuessy et al. 4942. Werling & Leth-Nissen 84), a quite unrelated species. 126

Fig. 12. Distribution of Clibadium microcephalum

( a ) and C. sessile (® ) in Ecuador and Panama, respectively. 127

Clibadium sessile has its closest relationship with C. microcephalum from Ecuador. Both species are included in Section Glomerata and have involucres with three glabrous phyllaries and disc florets without pales, but they differ by phyllaries 7-12-veined in the former and 5-7-veined in the latter. They also differ in head structure with C. sessile with 5 ray and 6-8 disc florets and C. microcephalum with 3 ray and 3-4 disc florets. A unique characteristic of this species is the presence of tubular ray corollas with only two lobes in contrast to the common 4-lobed condition observed in

C. microcephalum.

10. Clibadium microcephalum S. F. Blake, Wash. Acad. Sci. 16: 418. 1926.--Type: Ecuador. Tungurahua: Valley of , between Banos and Cashurco, 25 Sep 1923, Hitchcock 2187 3 (holotype: US!; isotypes: GH! NY! ) .

Shrub 2-4 m tall; stem subangulate, strigillose, with hairs 0.5 mm long. Leaves with blades ovate, 16-24 cm long, 10-14 cm wide, at apex acuminate, at base cuneate to cuneate- rounded, narrowly decurrent on the upper one-third of the petiole, with margin shortly serrate to serrulate, sparcely strigose both sides with hairs 1 mm long; petiole 3-6 cm long, strigillose with hairs 1 mm long. Capitulescence paniculiform, ternate at apex of stem and formed by two aggregated groups of 3-4 heads each one. Heads radiate, 5 mm 128 high, 2-3 mm diameter; involucre oblong cylindric, 2-3 mm diameter; phyllaries 3, suborbicular-ovate, 2.5-3 mm long, 2- 2.5 mm wide, 5-7 veined, at apex obtuse, with ciliate upper one-third, glabrous dorsally. Ray florets 3, subtended by, ovate pales, 2.5 mm long, 2 mm wide, 5 -veined, strigillose toward apex, with hairs 1 mm long; corollas white, 2 mm long, tube 1.2-1.4 mm long, 0.5 diameter, 4-lobed, lobes deltate, shortly ciliate at margin. Disc florets 3, epaleate; corolla white, 1.5 -1.8 mm long, 0.5 mm diameter, shortly 5-lobed, lobes deltate, 0.2 mm long, scarcely pilose at apex, with hairs 0.5 mm long; anthers black, 1 mm long; styles 2 mm long, undivided, ovaries sterile, pilose toward apex with hairs 0.5 mm long. Achenes obovoid, 2 mm long, 1.5 mm diameter, pilose on upper one-third, with hairs 1 mm long.

Chromosome number, n = 16. Phenology. Flowering from June to August. Distribution (Fig. 12). Ecuador; tropical humid forests, mainly in cut-over areas; 900-2800 m.

Representative Specimens. Ecuador. Canar: Guayaquil, Cuenca road, ca. 10 km E of Cochencai, Gentry et al. 30826 (MO); about 40 km E of the bridge at Guayaquil, King & Almeda 7735 (CAS, MO). Napo-Pastaza: road Baeza-Napo, 20 km S of Baeza, Cosanga, 3 km w of the village, Balslev & Madsen 10341 (F); Mera. rastrojo.Harling et al. 10124 (F, WIS); Parayaca, ca 10 km E of Canelos, Lugo 4554 (MO); 2 km of Baeza on gravel road to Tena, Stuessy & Arriagada 12363 (OS); 6 km S of Cosanga on gravel road to Tena, Stuessv & Arriagada 12365 (OS); 7 km N of Tena to Archidona along gravel road, Stuessy & Arriagada 12369 (OS); 2 km N of Archidona on road to Baeza, Stuessy & Arriagada 1237 0 (OS); 0.5 km W of Mera on road to Banos, Stuessy & Jansen 4965 (OS), 5 km E of Mera on road to Puyo, Stuessv & Jansen 4968 (OS); 34.6 km N of Puyo on road to Tena, Stuessy & Jansen 4980 (OS); 62 km N of Puyo on road to Tena, Stuessy & Jansen 4982 (OS). Tungurahua: entre Huambalo y Cotalo, Solis 97 23 (F); along the road to Puyo, ca 2 km E of Rio Negro, King 6560 (MO, US); along the

■road to Tena, ca 7 kms N of Puyo, King 6567 (MO; US); 11.7 km E of Rio Veroade on road to Puyo, Stuessy & Jansen 4960 (OS) ; 5.3 km E of Rio Verde on road to Puyo, Stuessy & Jansen 4963 (OS); 3.3 km E Rio Topo on road to Puyo along Rio Pastaza, Stuessv & Nesom 5814 (OS); 4.8 km E of Rio Topo on road to

Puyo, Stuessy & Nesom 5816 (OS); 10.4 km W of Mera on road to Banos, Stuessy & Nesom 5822 (OS); 11.9 km W of Mera on road to Banos, Stuessv & Nesom 5823 (OS).

Clibadium microcephalum is distinguishable within the genus by its subangular stem, acuminate leaf apices, and cuneate leaf bases narrowly decurrent on the upper one-third of the petioles. Clibadium microcephalum is closely related to C. sessile. Both species are similar in having 3 glabrate phyllaries with obtuse apices, glabrate ray pales, and disc florets without pales, and with 5-lobed corollas. Clibadium microcephalum differs by having orbicular 5-7-veined phyllaries (vs. ovate and 7-12-veined in C. sessile). 3 ray florets with corollas 4-lobed (vs. 5 ray florets with 130 corollas irregularly 2-lobed), ray pales 5-veined (vs. 7-9- veined) 3-4 disc florets (vs. 6-8), and pilose achenes (vs. glabrate in C. sessile). Clibadium microcephalum is distributed only in Canar, Napo-Pastaza, and Tungurahua provinces, Ecuador, whereas C. sessile is restricted to

Chiriqui province, Panama.

Clibadium microcephalum has been suggested as being related to C. grandifolium from Costa Rica (Blake, 1926) because of the large leaves (24 cm long, 14 cm wide) and congested capitulescences. But the detailed structure of the head renders C . microcephalum distinctive. Clibadium grandifolium has large heads with 6-8 ray and 8-12 disc florets, in contrast to 3 ray and 3-4 disc florets in C. microcephalum.

Clibadium microcephalum has also been suggested as closely related to C. sodiroi with which it may hybridize (based on intermediate morphology and chromosomal abnormalities; see Hybridization). Although similar in morphology, the heads of C. microcephalum are arranged in small glomerules, whereas those of C. sodiroi are more loosely arranged. The morphological differences (see species descriptions) result in placement of these two species in different taxonomic sections. 131

11. Clibadium frontinoense Diaz & Arriagada, Rev. Acad. Colomb. Cienc. 18: 301. 1992.--Type: Colombia.

Antioquia: Frontino, Corregimiento de Nutibara, cuenca del Rio Cuevas, 9 Jul 1986, Sanchez. Orrego. Silva. Martinez. Restrepo & Acevedo 213 (holotype: MEDEL!;

isotype: COL!)

Shrub 1.5 m tall, branches shortly strigose with hairs 0.5 mm long. Leaves with blades ovate, membraneous, 28-28.5 cm long, 20-21.5 cm wide, at apex acuminate, at base cordate slightly truncate and decurrens; with margin twice serrate; upper surface sparcely strigose, main vein conspicuous and shortly pubescens; lower surface light green, scarcely strigose with hairs 0.5-1 mm long, main vein thick, secondary veins 6-7 each side, minor veins reticulated; petioles 8-10 cm long, shortly strigose, with hairs 0.2 mm long. Capitulescence capituliform, with tightly condensed clusters of heads, 1.5-2.5 cm diameter, with opposite, ovate leaves at base, blades 10.5 cm long, 3.8 cm wide, branches up to 10 cm long, densely pilose, with hairs 1 mm long. Heads radiate, subglobose, 6 mm high, 3-4 mm diameter, sessile or shortly pedicellate; bracts narrowly ovate or linear, 10-35 mm long decreasing toward top; phyllaries 4, ovate, 6.5 mm long, 3.5 mm wide, at apex acute, abaxially strigose with hairs 1 mm long, with margin upper one-third ciliate, 4-5 veined. Ray florets 8, subtended by ovate pales, 5.5-6 mm long, 3.4-3.6 mm wide, 3 -veined, with margin upper one-third ciliate; corollas white, glabrous, 3.6 mm long, 1.6 mm diameter, 4- 132

lobed, lobes deltate, 1 mm long, at margin shortly ciliate; styles 3 mm long, branches 1.6 mm long. Disc florets 5, subtended by lanceolate pales, 4.5 mm long, 1.8-2.0 mm wide, 1-veined, at apex obtuse, with margin upper one-third ciliate; corolla white, 5 mm long, 5-lobed, lobes 1.5 mm long, apex hirsute; ovaries sterile, pilose toward apex, hairs 1 mm long. Achene not seen. Chromosome number unknown. Phenology. Flowering probably in June and July. Distribution. (Fig. 13). Colombia, known only from the type locality in the Frontino region, east slope of the Cordillera Occidental; 1300 m.

Clibadium frontinoense is characterized by highly congested capitulescences similar to the close relative, C. zarucchii. also from the Frontino region in the western slope of the Occidental Cordillera. The tightly congested clusters of heads in C. frontinoense. however, are 1.5-2 cm diameter, whereas those in C. zaruchii are 3-4 cm diameter. The heads in C. frontinoense are smaller than those of C. zaruchii (6 mm vs. 8 mm long, respectively). Clibadium frontinoense also has double the number of ray florets of C. zaruchii (8 ray florets vs. 4, respectively). Further, the leaves in C. frontinoense are slightly decurrent from a cordate base in contrast to attenuate leaves in C. zaruchii. 133

Fig. 13. Distribution of Clibadium frontinoense (•) and C. zaruchii ( a ) in Colombia. 134

12. Clibadlum zaruchii H. Robinson, Phytologia 65: 51. 1988.--Type: Colombia. Antioguia: Frontino, km 14 of road Nutibara-Murri, 23 Sep 1987, Zarucchi. Brant & Castano 5674 (holotype: US!; isotypes: COL! MO!).

Shrub 3 m tall; stem many-angled, often flattened near new branches, densely strigose, with hairs 1 mm long; internodes 6-9 cm long. Leaves with blades ovate, 10-20 (35) cm long, 6-15 (25) cm wide, membraneous, at apex acuminate, at base atenuate (new leaves), truncate-rounded, and shortly decurrent on the petiole, with margin serrulate to irregularly serrate, both surfaces strigose, with hairs 1 mm long; petioles 2-5 cm long, sulcate, narrowly winged, strigillose with hairs 1 mm long, marginate by the decurrent leaf blades. Capitulescence paniculiform, with clusters of heads tightly condensed (ca. 100 heads, 3-4 cm diameter) at the end of trichotomously divided terminal branches. Heads sessile or shortly pedicellated, 8 mm high, 3-4 mm diameter; phyllaries 4, ovate, ovate-lanceolate, 7-8 mm long, 3-4 mm wide, at apex acute, 4-5 veined, abaxially strigose with hairs 1 mm long, with margin ciliate on upper one-third. Ray florets 4, subtended by ovate-lanceolate pales, 6-7 mm long, 3-4 mm wide, 3-veined; corolla white, glabrous, narrowly tubulose, 4.5 mm long, 2 mm diameter, 4-lobed, with lobes 1 mm long shortly pilose at margin, with hairs 0.5 mm long; styles 3.5 mm long, branches 2 mm long. Disc florets 4 (-5), subtended by lanceolate pales, 5-5.5 mm long, 2 mm wide, 1- veined, apex slightly ciliate; corollas light-purple, 135 tubulose, 5 mm long. 2.4 mm diameter, 5-lobed, with lobes l mm long, at apex slightly hirsute, with hairs 0.2 mm long; styles undivided, 2.5 mm long; ovaries sterile pilose, with hairs 1 mm long. Achenes obovoid, 2.5 mm long, 1.8-2.1 mm diameter, glabrous. Chromosome number unknown. Phenology. Flowering from June to September. Distribution (Fig. 13). Known only from Colombia, Municipio de Frontino; wet montane vegetation, disturbed areas along roadside, west slope of Occidental Cordillera; 1850-1870 m.

Additional Specimen Examined. Colombia. Antioquia: Municipio de Frontino, Corregimiento La Blanquita, Alto de

Cuevas, 14.5 km W of Nutibara on road to La Blanquita,

Callejas et al. 6842 (MO).

Clibadium zaruchii is easily recognized by its condensed capitulescence (ca. 100 heads) and large (3-4 cm diameter) tightly congested clusters of heads at the tops of trichotomous branches. These congested clusters of heads are the largest of any species in the genus. Clibadium zaruchii has a close relationship with C . frontinoense. Both species are restricted to the Frontino region in Antioquia, but they have allopatric distributions. Clibadium zaruchii grows in wet montane vegetation on the western slope of Cordillera Occidental, whereas C. frontinoense grows in a drier zone on the eastern slope of the same Cordillera. These two slopes differ in the amount of light and precipitation (Diaz- 136

Piedrahita, pers. comm.). In addition to geographical distribution and ecology, there are morphological differences between these closely related species (see discussion under the former species).

13. Clibadium rhytidophyllum Diels, Notizbl. Bot. Gart.

Berlin 14: 341. 1939.--Type: Ecuador. Pichincha: "zwischen Nono und Mindo bei Guarumas," 17 Oct 1937, Sydow 204 (type material not located; presumed destroyed)--Neotype (here designated): Ecuador. Pichincha: 21 km N of Nono, 2-3 km N of Guarumas, 21

Jul 1977, Stuessv. Padilla & Jansen 4863 (OS!).

Clibadium harlinaii H. Robinson, Phytologia 44: 282. 197 9.--Type: Ecuador. Carchi: road Tulcan - Maldonado, ca. 13 km SE of Maldonado, 1 Mar 1974, Harling & Anderson 12363 (holotype: GB; isotype: US!, photos: CAS!

F! GH! MO [2] ! ) .

Shrub to arching shrub 3 m tall; stems and branches strigose-pilose, older branches tomentose with hairs 1 mm long. Leaves with blades lanceolate to lanceolate-ovate, subcoriacea, 7-13 cm long, 2-5 cm wide, densely strigose with hairs 0.6-1 mm long, at apex briefly acuminate, at base obtuse, with margin serrate to serrate-serrulate, veins prominent; petioles 1-2 cm long, strigose with hairs 1 mm long. Capitulescence thyrsoid, with 3-terminal glomerules. Heads radiate, ovoid-subglobose, shortly pedicellate, 4-4.5 mm high, 3.3-4 mm diameter; phyllaries 3, ovate, 4.5 mm long, 2.5-3 mm wide, with margin shortly ciliate at the upper one- third, 4-5-veined. Ray florets 5-10, subtended by ovate pales, 4 mm long, 3 mm wide, with margin at upper one-third ciliate, 3 -veined; corolla white, tubulate, 2 mm long, 0.6 mm diameter, 2-lobed, lobes irregularly deltate, 0.5 mm long, scarcely pilose toward apex. Disc florets 5-10, subtended by ovate to ovate-rhomboid, 3 mm long, 1.5-2 mm wide, at apex acuminate, with margin shortly ciliate at the upper one- third, 1-veined; corolla white, 3 mm long, 0.8 mm diameter, 5-lobed, lobes 0.6 mm long, scarcely strigose toward apex; anthers black, 1-1.2 mm long, glabrous; styles 1.5 mm long, undivided, ovaries sterile, 2 mm long, 1 mm diameter, upper half pilose, with hairs 0.5 mm long. Achenes obovoid with a short apical disc, 2 mm long, 1.5 mm diameter, glabrous. Chromosome number, n = 16. Phenology. Flowering from June to December. Distribution (Fig. 14). Ecuador: Carchi and Pichincha provinces; high mountain humid forest; 2000-2600 m.

Additional Specimens Examined. Ecuador. Pichincha: 4 km S of Guarumos on road along Tandayapa River, Stuessy et al. 4871 (OS), 28.7 km W of Aloag on road to Sto. Domingo de Los Colorados, Stuessy et al. 4896 (OS); along road between Nono and Nanegal, 89.8 km before Tandayapa, Croat 49326 (MO).

Clibadium rhytidophyllum is closely related to C. sprucei. also from Ecuador, but the latter is endemic to 138

Fig. 14. Distribution of Clibadium_rbvtidooftvlXuxn

( •) and C. sprucei (a ) in Ecuador. 139

Volcan Tungurahua. Clibadium rhyt idophyllum is distinguished by having ovate leaves, heads arranged in terminal thyrses with three levels of aggregation, phyllaries greenish-white, and glabrous achenes. Its achenes are also characterized by the presence of a short ring-shaped apical projection similar to that seen in C. glabrescens from Venezuela. That is, however, regarded as an evolutionary parallelism.

Clibadium harlingii was described by Robinson (197 9) as a distinct species from C . rhytidophyllum because of a more glomerate capitulescence, pinnate venation of leaves and glabrous ray achenes. Study of available material indicates an overlapping of characters observed in both taxa. Also, the ring-shaped apical projection, both in ray and disc achenes, occurs in some florets of C. harlingii. (e.g., Croat 49326). Because there seems no clear distinction between the two species, therefore, they are here treated as synonyms.

After having sent letters to B, C, and G inquiring about type material, and having received negative responses, I reluctantly conclude that the type is lost or destroyed. Therefore, I have herein selected a neotype collected from the type locality (= topotype). 14 0

14. Clibadium sprucei S. F. Blake, Contr. Gray Herb. 52: 5. 1917.--Type: Ecuador. Tungurahua: Jan 1859, Spruce 5826 (holotype GH!; isotypes: BM! fragment at GH!

photos: GH! OS!, K! photo: OS!, NY!, P photo: OS!).

Shrub 2-3 m tall; stems subangulate, branches densely strigose with hairs 1 mm long. Leaves with blades ovate- lanceolate to lanceolate, 7-14 cm long, 2-4 cm wide, at apex largely acuminate, at base truncate, attenuate, 3-nerved, with margin crenate-serrate, above dark-green, scabrous, below pilose with hairs 1-1.5 mm long; petioles 0.5-1 cm long, densely strigose-hispidulous with hairs 1.5 mm long. Capitulescence thyrsoid, with dense axillary or terminal cluster of heads, on peduncles 3-6 cm long, with 2 bracts at base, glomerules, 1-1.5 cm diameter (including up to 20 aggregated heads). Heads 6 mm high, 4 mm diameter; phyllaries 3, oblongo-ovate, 5 mm long, 4.5 mm wide, at apex acute, densely strigose with hairs 1 mm long, with upper one- third ciliate, 4-5 nerved. Ray florets 5-7, subtended by ovate pales, 4.5 mm long, 3.5 mm wide, 3-veined, strigose with hairs 1 mm long; corolla white, tubulose, 4 mm long, 1.5 mm diameter, 2-lobed, lobes irregularly deltate, scarcely pilose toward apex, hairs 0.5 mm long. Disc florets (7)-10, subtended by lanceolate pales, 4 mm long, 2 mm wide, 1- veined, slightly ciliate toward apex margin; corolla white, tubulose, 3 mm long, 1 mm diameter, 5-lobed, lobes regularly deltate, 0.5 mm long, scarcely ciliate toward apex; anthers black; style undivided, 2.5 mm long; ovaries sterile, 2 mm 141 long, 1 nun diameter, pilose toward with hairs 0.8 mm long. Achenes brown-black, 2.5 mm long, 1.8-2 mm wide, apex pilose with hairs 1 mm long. Chromosome number unknown. Phenology. Flowering from January through August. Distribution (Fig. 14). Ecuador, known only in the vicinity of the Tungurahua Volcano and from the slopes of the Paramos de Matanga; tropical evergreen wet forest with secondary vegetation and cultivated areas; 1800- 3100 m.

Additional Specimens Examined. Ecuador. Chimborazo: Chontapamba between Puela and Banos, Lugo 7 63 (F, MO, WIS) . Morona-Santiago: E slopes of Paramos de Matanaga, ca. 30-40 km S of Sigsig along road towards Gualaquiza, Lutevn & Cotton 11197 (F). Tungurahua: Chaupi, Dodson & Thien 1849 (MO, WIS); Eastern Cordillera, NE slope of Volcan Tungurahua, Rimbach 7 00 (F); San Antonio E slope of Tungurahua Volcano, Tate 577 (US); from Banos toward Chaupi and Volcan Tungurahua, Stuessy & Nesom 5811 (OS); up path from Banos to Volcan Tungurahua, Stuessy & Arriagada 12400 (OS).

Clibadium sprucei has its closest relationship with C . rhytidophyllum. both species being endemic to Ecuador but restricted to different regions. Clibadium sprucei is confined close to the Tungurahua Volcano, whereas C. rhytidophyllum only has been reported from the provinces of Carchi (border with Colombia) and Pichincha (250 km N of

Tungurahua Province). Both species occur in similar habitat 142

(2000-3000 m) and have similar general morphology, with head structure and organization in dense clusters (hence both included in Section Glomerata). Clibadium sprucei is easily

recognized, however by its more lanceolate leaves in contrast to the more ovate leaves of C . rhytidoohvllum. The heads of

C. sprucei are arranged on axillary and terminal branches whereas in C . rhvtidophyllum they are only in terminal thyrsoid clusters. Both species have achenes with fleshy exocarps, but only C . rhytidoohyl1um produces an orange sap

(e.g., Stuessy et al. 4863). Clibadium sprucei lacks the ring-shaped apical projection seen in specimens of C. rhytidophy1lum. In addition, C. sprucei has pilose achenes which contrasts with the glabrous condition in C. rhyt idophyllum.

This species honors British botanist and plant collector Richard Spruce (1817-1893), who collected in South America between 1849 and 1864 (Stafleu & Cowan, 1985).

IIB. Clibadium L. section Grandifolia Arriagada, Sect. nov. Typus: Clibadium grandifolium Blake.

Folia ovata vel lanceolato-ovata; capitulescentiae cymarum laxarum vel thyrsiformum; phyllaria guinquenervia; flores feminei 3-9, corollis trilobatis, paleis tetranervibus; flores hermaphroditi disci 4-10, corollis quinquelobatis, paleis binervibus. 143

Leaves ovate to ovate-lanceolate; heads arranged in loose thyrsoid or paniculiform capitulescences, paleaceous; phyllaries 5-veined; ray florets 3-9, with corollas 3-lobed, and pales 4 -veined; disc florets 4-10, with corollas 5-lobed, and pales 2-veined. Species 15-19.

15. Clibadium grandifolium S. F. Blake, Contr. U.S. Natl. Herb. 22: 599. 1924.--Type: Costa Rica. Limon: Llanuras

de Santa Clara, along Rio Pacuare, Apr 1896, Smith 6614 (holotype: US!; isotype: K!, photo: OS!).

Clibadium grande S. F. Blake, Contr. U.S. Natl. Herb. 22: 601. 1924.--Type: Costa Rica. Limon: La Florida, 18 Jun 1897, Pittier 11280 (holotype: GH!, photos: GH! US!; isotype: US!).

Clibadium pacificum Cuatrec., Rev. Colomb. Cienc. Exact.

9:238. 1954.--Type: Colombia. Valle: Rio Calima entre La Herradura de Ordonez y Pena de Campotriste, 3 Mar 1944, Cuatrecasas 16683 (holotype: F!, photo: US!; isotypes: COL! F! photo: US!, US!).

Clibadium terebinthinaceum (Sw.) DC. subsp. colombiense

Cuatrec., Rev. Colomb. Cienc. Exact. 9: 240. 1954.-- Type: Colombia. Valle: Hoya del Rio Digua, lado derecho, La Elsa, 9 Nov 1943, Cuatrecasas 15298 (holotype: F!,

photo: US!; isotypes: COL! US!). Shrub to small tree 2-6 m tall; stems 3 cm diameter toward base, striate, strigose with hairs 1 mm long. Leaves with blades broadly ovate; 15-40 cm long, 10-30 cm wide, at apex acute to acuminate, at base obtuse to shortly attenuate, sometimes decurrens on upper third petiole; with margin serrate, upper surface glabrate to strigose, lower surface strigose with hairs 0.6-0.8 mm long; petioles 4-19 cm long, 3-4 mm diameter, glabrate to strigillose with hairs 0.5 mm long. Capitulescence paniculiform, with 100 and up to 600 clustered heads in terminal open branches, peduncles 0.5 mm long. Heads globose to subglobose, 4-6 mm high, 3-5 mm diameter; involucre cupulate; phyllaries 3, light green, ovate to suborbicular-ovate, 3-5 mm long, 3-4 mm wide, 5- veined, abaxially glabrate to strigillose with hairs 1 mm long, with margin upper one-third ciliate. Ray florets 6-8

(-10), subtended by ovate pales, 3-4 mm long, 2-2.6 mm wide, strigillose toward apex with hairs 0.5 mm long, with margin upper one-third ciliate, 4 -veined; corollas white, 3 mm long, 0.5 mm diameter, 3-lobed, lobes deltate, 0.4 mm long, scarcely pubescent at apex; styles 2.5 mm long, branches 1.2 mm long. Disc florets 8-12 (-14), epaleate (exceptionally l-

2 disc florets subtended by narrowly lanceolate pales); corollas white, 3-4 mm long, throat 2.5 mm long, 1 mm diameter, 5-lobed, lobes 0.5 mm long, scarcely pubescent at apex; anthers black, 2 mm long; styles 3 mm long, ovaries sterile, 1.5 mm long, willose at apex. Achenes 2.5 mm long, 1.7 mm diameter, villose on upper one-third, hairs 0.5 mm long. Chromosome number, n = 16. 145

Phenology. Flowering from March through September.

Distribution (Fig. 15). Colombia, Costa Rica, Nicaragua and Panama; tropical moist forest, secondary vegetation, disturbed forests mainly along roadside; 0-2000 m.

Representative Specimens. Colombia. Antioquia: alrededores de Villa Arteaga, Araque & Barkley 18C740 (F, US); Municipio Frontino, Corregimiento La Blanquita, 14.5 km W Nutibara, Calleias et al. 6610 (MO); Quebrada Congo, 6 km E of Guapa, 53 km S of Turbo, Haught 467 8 (NY, US). Municipio de Mutata, 2.5 km N of Mutata on road to Chigorodo, Zarucchi et al. 5036 (MO). Boyaca: Mt. Chapon, NW of Bogota, Lawrence

135 (F, GH, NY, MO, US). Cauca: Cordillera Occidental, Carpinterias, bosque entre los cerros Munchique y Altamira, Perez Arbelaez & Cuatrecasas 6142 (COL, F, US). Choco: Rio

Truando, between La Nueva and La Esperanza, Duke 9886 (FSU); Rio El Valle between El Valle and Choco Indian Village, Gentry & Fallen 17249 (MO, OS). Municipio de Novita, ladera N del Cerro Torra, camino al Alto del Oso, Forero et al. 3312 (MO); Carretera Panamericana, entre el Rio San Pablo y el Rio Pato, Forero et al. 5540 (MO); Carretera Itsmina-Quibdo, entre El Dos y Las Animas, 21 Apr 197 9, Forero et al. 547 6 (MO); 15 km desde Pizarro a La Porquera, Confluencia de la Quebrada Carpio, Fuchs et al. 217 28 (COL). Municipio San Jose del Palmar, Hoya del Rio Torito, Finca "Los Guaduales", Forero et al. 6853 (MO); vertiente Oriental vereda de Rio

Negro, Ramos et al. 1599 (MO); Bahia Solano, near Cuidad Mutis, Killip & Garcia 3357 5 (COL, US). Municipio de Pizarro, 146

Fig. 15. Distribution of Clibadium cordatum ( ▲ ) and C. arandifolium (®) in Colombia, and in Colombia, Costa Rica, Nicaragua, and Panama, respectively. entre Boca de Pepe y Pie de Pepe, Espina 1991 (MO). Tolima: at bridge over Saldana to house of Manuel Puertas, Core 1599 (F, US); El Fresno, basque abierto, Cuatrecasas 9383 (COL, F ) . Valle: Cordillera occidental, Hoya del Rio Digua, lado izquierdo, Piedra de Moler, Cuatrecasas 15025 (F); Quebrada del Rio San Juan, arriba de Queremal, Las Colonias, Cuatrecasas 23932 (F). Cordillera Central; Hoya del Rio

Guadalajara, Cuatrecasas et al. 27 601 (NY, US); Cordoba, wooded cliffs of Rio Dagua, Killip 5268 (US); Sabaletas, km 29 of highway from Buenaventura to Cali, Killip & Cuatrecasas 28862 (CAS, F, MO, NY, US, WIS). Costa Rica. Cartago: beyond Pavones, 14 km from IICA Turrialba, Carlson 3405 (F, NY). Heredia: Finca La Selva OTS Field Station, Hammel 924 0 (MO); near Cariblanco N Volcan Poas, Williams 20263 (F) . Limon: SE of Siquirres near Rio Pacuare, Arriagada 406 (OS); Vatsi, W from Bribri, Arriagada 413 (OS); near Rio Zent, Jan 1900, Pittier 16068 (GH); cerca Catarata La Paz camino a Sarapiqui, Poveda & Castro 4138 (CR, MO); Rio Pacuare, Llanuras de Santa Clara, Smith 6614 (GH, US). Puntarenas: Santa Cecilia, Stevens 7 53 (US). San Jose: Parque Nacional Braulio Carrillo, Del Bajo de La Hondura, 13 km a Gu&piles, Gomez-Laurito 6419 (CR); near Rio Sucio, Estacion Carrillo, Sanchez et al. 510 (MO). Nicaragua. Zelaya: ca. 6.3 km S of bridge at Colonia Yolania, Stevens 4 816 (OS). Panama. Bocas del Toro: Santa Catalina along riverbank, Blackwell et al. 2749 (MO); SE y NE del campamento Changuinola 1 del IRHE, Correa & Taylor 3349 (MO); Rio Changuinola near Changuinola, Dwyer 512 8 (MO); 2 km W of Continental Divide, trail to elfin forest, Hampshire & Whitefoord 97 6 (BM); Chiriquicito to 5 mi S along Rio Guarumo, Lewis et al. 2082 (MO, UC); vicinity of Fortuna Dam along trail on continental divide, McPherson 1107 3 (MO). Chiriqui: Fortuna Dam area on Kaolin Hill, D'Arcy et a l . 15904 (MO, UC); N of San Felix at Chriqui-Bocas del Toro border, Mori & Kallunki 5897 (MO); Cerro Colorado 31.6 km from Rio San Felix bridge, Sullivan 32 8 (MO). Code: El Valle de Anton, vicinity Finca Tomas Arias, Allen 3621 (MO); 7 km HE of El Cope, Antonio 1136 (MO); 2.5 mi above El

Valle on road to La Mesa, Croat 1337 2 (OS); road from Penonome to Coclecito 9 km N of El Llano Grande, D'Arcy & Hammel 12294 (MO); Caribbean side of divide at El Cope,

Hamilton & Davidse 27 29 (UC); around Limon N of Alto Calvario, Folsom 5813 (MO); 44 km N of Penonome on road to Coclesito, Hammel 1691 (UC); along road from La Pindeda to El Cope by way of Piedras Gordas, Hammel 2621 (UC); N of Cerro Gaital vicinity of El Valle. Hammel 3891 (MO); 3 km beyond El Valle de Anton, Hartman 3936 (OS); Luteyn & Kennedy 1638 (MO) ; foot of Cerro Pilon, above El Valle de Anton, Porter et a l . 4651 (MO, UC); N of La Mesa on Finca Gabriela, Stuessy & Gardner 4534 (OS). Colon: along Rio Escandalosa on road to Salamanca, Huft & Knapp 1621 (MO). Darien: NO trails to Cerro Pirre along banks of Rio Perrecenega, D'Arcy & McPherson 16230 (MO); Cocalita near Colombian border, Dwyer

5111 (US); vicinity of mine at Cana, Croat 37 677 (MO); lower slopes of Alturas de Nique along Rio Coasi, Hartman 12254 (MO). Panama: hill S of Guacuco 8 km E of Ipeti, Hamilton & D'Arcy 1391 (MO). Veraguas: roadside between 149

Santa Fe and San Jose, D'Arcy 10322 (MO); 3-5 mi N of Santa Fe, Gentry 3034 (OS); NW of Santa Fe 11 km from Escuela

Agricola de Piedra, Mori et al. 3860 (MO, WIS).

Clibadium grandifolium is easily recognizable by its large leaves, even up to 4 0 cm long (e.g., Lawrence 135: Killip & Garcia 33575: Zarucchi 5036) with attenuate bases and by paniculiform capitulescences with 100-600 heads (the largest number counted for a single capitulescence in any

species) clustered together in terminal branches. The heads may change from 6 mm long (Stuessy & Gardner 4533). 4-5 mm long (Hammel 3891) to (rarely) 3 mm long (Williams 20263). but the basic head structure remains the same. Also the leaves, in juvenile specimens, are small (10-15 cm long, 8-10 cm wide) which makes C. grandifolium superficially resemble C. asperum. The strigose condition of the involucral bracts of the latter is the distinguishing feature. Clibadium grandifolium is closely related to C. cordatum. both species being included in Section Grandifolia. Clibadium grandifolium differs in having glabrate phyllaries with obtuse apices, pilose achenes, and disc florets without pales. Clibadium grande was described by Blake (1924) as a distinct species, related to C . grandifolium. based on one specimen from Costa Rica, that consisted "only of an inflorescence and a detached leaf, probably from the lower part of the stem...(p. 601)." In my opinion, the differences emphasized by him are not sufficient for formal recognition 150 at any level. In addition, both species have overlaping distributions in Limon Province. The overlap of diagnostic characters of these species, as shown in a scatter diagram (Fig. 22), supports the decision to treat them as conspecific. Even though C. grandifolium was published simultaneously with C . grande in the same paper, I have selected the former for use because it communicates better the main chracteristics of the species (and also of the Section).

16. Clibadium cordatum Cuatrec., Rev. Acad. Colomb. Cienc.

Exact. 9: 237. 1954.--Type: Colombia. Valle: Cordillera Occidental, Hoya del Rio Anchicaya, bosques entre y Miramar, 15 Apr 1943, Cuatrecasas 1437 8 (holotype:

F!, photo: US!; isotype: US!).

Shrub 2-4 m tall; stem subangulate, 4 cm diameter, branches strigose-pilose with hairs 1 mm long. Leaves with blades ovate, rounded-ovate, membraneous, 20-50 cm long, 20- 40 wide, densely strigose with hairs 0.6-1 mm long, at apex briefly acuminate, at base deeply cordate (mainly in old leaves); with margin serrate to crenate-serrate, veins prominents, 7-9 veins; largely petiolated, 16-22 cm long, 1.5-2.5 mm diameter, striate, strigose with hairs 1 mm long.

Capitulescence cymose or paniculiform, broadly branched. Heads ovoide, 4-5 mm high, 3 mm diameter, shortly pedicellated or sessile, clustered in aggregations at 151 terminal branches; phyllaries 3, obovate to obovate-ovate, 4-5 mm long, 3-4 mm wide, at apex mucronate-acuminate, strigose with hairs 1 mm long, with margin ciliate at upper half, 5-veined. Ray florets 7, paleate, subtended by ovate pales, 4.5 mm long, 3-3.5 mm wide, strigose with hairs 0.3- 0.4 mm long, with margin shortly ciliate at upper half, 4- veined; corolla white, 2 mm long, tube 1.4 mm long, 0.5 mm diameter, 3-lobed, lobes irregularly deltate, 0.5 mm long, shortly ciliate at margin; styles 2.5 mm long, branches 1 mm long. Disc florets 8 (-9), subtended by lanceolate pales, 3- 4 mm long, 0.5-1 mm wide, 2-veined, with margin shortly ciliate at apex; corolla white, 3 mm long, 0.5 mm diameter, 5-lobed, lobes deltate, 0.5 mm long, scarcely pubescent at apex; anthers black, 1-1.5 mm long; style 2 mm long undivided, ovaries sterile, pilose toward apex, with hairs 1 mm long. Achenes 2 mm long, 1.6 diameter, pilose on the upper one-third, with hairs 1-1.5 mm long. Chromosome number, n = 16.

Phenology. Flowering from March to December. Distribution (Fig. 15). Colombia, mostly restricted to Choco and Valle; growing in cut-over areas with secondary vegetation; 300-600 m.

Representative Specimens. Colombia. Choco: headwaters of Rio Tutunendo, E of Quibdo, Archer 2187 (NY); Lunar de Jurubida, afluente izquierdo, Quebrada Isla, Barbosa 6646 (MO); carretera Anserma-nuevo-San Jose del Palmar, Alto del Galapago, Forero et al. 2257 (COL); carretera San Jose del 152

Palmar-Novita, alrededores de Curundu, Forero et al. 2347 (COL), Forero et al 3517 (MO); carretera Tutunendo-El , alto Rio Atrato, Forero et al. 5871 (COL, OS); Municipio de San Jose del Palmar, Hoya de la quebrada La Cristalina, Forero et al. 7 594 (MO); 12 km E of Quibdo, ca. 3 km W of Tutunendo, Gentry & Renteria 23881 (COL); along bank of Rio

Virudo above mangroves, Jeffrey et al. 100 (COL). Valle:

Cordillera Occidental, vertiente occidental, Hoya del Rio Anchicaya, Quebrada del Retiro, Cuatrecasas 13696 (COL, F, US, paratypes). Municipio Dagua, Corregimiento El Danubio, Alto Anchicaya, Devia 624 (MO).

Clibadium cordatum is distinguishable within the genus by its large leaves (20-50 cm long, 15-40 cm wide, the largest leaves known for the entire genus) with cordate bases (young leaves, however, are somewhat attenuate, e.g., Forero et al. 5871). This species is sometimes mistaken for C. grandifolium from Costa Rica and Panama, which also has large leaves but with a more typical attenuate base. In addition, C . cordatum has condensed clusters of heads (1 cm diameter) arranged in a paniculiform capitulescence. Both species are included in Section Grandifolia because of similarities of head arrangement (see sectional description). Clibadium cordatum differs from C. grandifolium in having strigose phyllaries with mucronate apices (glabrous and obtuse in the latter), and disc florets subtended by lanceolate, 2-veined pales (no disc pales in C. grandifolium). 153

The holotype consists of two parts (F accession numbers: 1362889. 1362890). The second sheet has been labeled by the author as "holotypus (part)". This sheet contains a single leaf that represents the size and cordate condition that really characterizes the species. Also the isotype consists of two parts (US accession numbers: 2220748. 2220749) showing the main features of the taxon.

17. Clibadium manabiense H. Robinson, Phytologia 44: 280. 1979.--Type: Ecuador. Manabi: Road to Santo Domingo-Chone, Flavio Alfaro, 11 May 1968, Harling.

Storm & Storm 9410 (holotype: GB, photos: CAS! F! GH! MO!; isotype: US!).

Trichapium strigosum Gilli, Feddes Repert. 94: 303. 1983.--Type: Ecuador. Waldfragment bei Sto. Domingo de los Colorados, 19 Jun 197 5, Gilli 113 (holotype: W, photos: CAS! F! MO [2]!).

Shrub 2.5 m tall; stem striate, strigose, with adpressed hairs l mm long. Leaves with blades ovate to ovate- lanceolate, coriaceous, 8-15 cm long, 3-8 cm wide, at apex acuminate, at base cuneate to rounded cuneate, with margin crenate-serrate, upper surface strigose with hairs 0.5 mm long, lower surface scarcely pilose with hairs 1 mm long, 3 main veins, primary and secondary veins prominent underside; petioles 0.5-2.5 cm long, striate, strigose with hairs 1 mm long. Capitulescence thyrsoid, at terminal branches, branches strigose with hairs 1 mm long. Heads subglobose, 4- 5 mm high, 4-5 diameter; phyllaries 2-3, suborbicular, 4-5 mm long, 3.5-4 mm wide, abaxially strigose with hairs 0.5-0.8 mm long, at apex markedly acuminate-mucronate, with margin

shortly ciliate on upper half, 5-veined. Ray florets 9-13, subtended by ovate pales, 4-4.5 mm long, 3.5-4 mm wide, 5- veined, strigose with hairs 0.5 mm long, with margin ciliate on upper one-third; corollas white, tubulate, 2.5 mm long, 1 mm diameter, 3-lobed, lobes 0.5 mm long, at apex scarcely pubescent; styles 1.5 mm long, branches 1 mm long. Disc florets 15, subtended by lanceolate pales, 3-3.5 mm long, 1.5-2 mm wide, with margin ciliate on upper one-third, 2- veined; corollas white, 3 mm long, throat 1 mm diameter, 5- lobed, lobes 0.5 mm long, at apex scarcely pubescent; anthers dark brown-purple, 1.5 mm long; styles undivided, 2 mm long, ovaries sterile, l mm long, 0.6 mm diameter, at apex pilose with hairs 1 mm long. Achenes obovoid, 2.5 mm long, 2 mm diameter, at apex pilose with hairs 1 mm long. Phenology. Flowering from July to September. Distribution (Fig. 16). Only known from the provinces of Manabi and Pichincha in Ecuador; secondary vegetation; 100-2000 m.

Additional Specimens Examined. Ecuador. Pichincha: carretera Quito-San Juan-Chiriboga-Empalme, km 69 a 3 km de la carretera a Santo Domingo de Los Colorados, Zack 1194 (F, MO); Waldfragment bei Sto. Domingo de Los Colorados, Gilli 155

A ®

Fig. 16. Distribution of Clibadium glabrescens ( ▼ ) , c. manabiense ( @ ) , and C. penfcageuron (A ) in Colombia, Ecuador, and Venezuela. 156

113 (W; photo: CAS, F, MO); Reserva Ecologica "Rio Guajalito", km 59 de la carretera antigua Quito-Santo Domingo de Los Colorados, 3.5 km NE de la carretera, Jaramillo & Zak 7797 (MO); Carretera Quito-San Juan-Chiriboga-Empalme-Santo Domingo de Los Colorados, km 59, a 15 km al NO de la carretera, Zak 1318 (F, MO).

Clibadium manabiense is characterized by a large number (13) of ray florets [the largest within subgenus Clibadium. together with C. micranthum (10-14), which is more distantly related], a large number (15) of disc florets, and acuminate tips of phyllaries.

Clibadium manabiense is closest to C . pentaneuron and both share many character states that emphasize their similarity (e.g., phyllaries 5-veined, ray corollas 3-lobed, ray pales 4 -veined, disc corollas 5-veined, disc pales 2- veined, and pilose achenes). Clibadium manabiense can be distinguished most easily however, by the 2-3 strigose phyllaries which contrast with 4 glabrate phyllaries (occasionally 3) in C . pentaneuron.

18. Clibadium glabrescens S. F. Blake, J. Wash. Acad. Sci. 27: 381. 1937.--Type: Colombia. Putumayo: mountains between Mocoa and Sibundoy, 19 May 1935, Archer 3415 (holotype: US!, photo: US!; isotypes: COL

[2]! LL!). 157

Clibadium napoense H. Robinson, Phytologia 73: 152. 1992.--Type: Ecuador. Napo: Canton Napo, Zatzayacu, 28 Mar 1935, Mexia 7110A (holotype: US!; isotypes: LL!

U C ! U S !).

Spreading shrub 2.5 m; stem with branches slender, striate, glabrous to strigose with hairs 1 mm long, internodes 3-12 cm long. Leaves with blades ovate, thin- papery, 8-18 cm long, 4-9 cm wide, at apex caudate-acuminate, at base cuneate, w i t h margin serrate to serrulate; above dark green, roughly strigose with hairs 1 mm long; beneath light green, sparcely strigose with hairs 0.6 mm long; petioles slender, slightly striate, 1-2 cm long, strigillose with hairs 0.6-1 mm long, ocasionally glabrous. Capitulescence paniculiform on terminal branches. Heads sessile or shortly pedicellate, subglobose, 3.5-4.5 mm high, 3-3.5 mm diameter; phyllaries 4, ovate-suborbicular, 3.5-4 mm high, 3-3.5 mm wide, 5-veined, at apex obtuse, upper half strigose with hairs 0.6 mm long, with margin ciliate on upper one-third. Ray florets 5-6, subtended by ovate pales, 3-3.5 mm long, 3-3.5 mm wide, upper half strigose, with hairs 0.5 mm long, with margin ciliate on upper one-third; corollas white, tubular, 2 mm long, 0.8 mm diameter, 3-lobed, lobes 0.4 mm long, at apex scarcely to densely strigillose; styles 1.5 mm long, branches 0.5 mm long. Disc florets 8-11, subtended by ovate-lanceolate pales, 3-3.5 mm long, 1.5-2 mm wide, at apex scarcely ciliate, 2-veined; corollas, white, 3 158 mm long, 0.5 mm diameter, 5-lobed, lobes deltate, 0.4 mm long, at apex scarcely pubescent; anthers black 1.5 mm long; ovaries sterile, 2 mm long, 1.5 mm diameter, pilose toward apex. Achenes obovoid, at base rounded, pilose at upper one- third, with hairs 1.1-5 mm long. Chromosome number, n = 16. Phenology. Flowering from March to September. Distribution (Fig. 16). Colombia, Ecuador and Venezuela, eastern slope of Andes; low humid tropical forest, along wooded stream banks; 500-1000 m.

Specimens Examined. Ecuador. Napo-Pastaza: near

Canelos, Mexia 6915a (US); 17 km E of Veracruz, Stuessy & Jansen 4971. 4977 (OS); 43 km N of Puyo on road to Tena, Stuessy & Jansen 4981 (OS); Tena: vacant lot near gas station on S side of river, Stuessy & Jansen 4984 (OS), N edge of Archidona on road to Baeza, Stuessy & Jansen 4985 (OS). Los Rios: between Quevedo and Naranjal, Mexia 6668 (NY, US); Canton Vinces, near Quevedo, Mexia 6595 (US). Venezuela. Merida: Distrito Campo Elias, Bosque de San Eusebio, Ruiz Teran 1139 (MO).

Clibadium glabrescens is a taxon characterized by its strigose leaves and capitulescences of open clusters of heads arranged in panicles at the ends of terminal branches. Clibadium glabrescens has C. pentaneuron as its closest relative. Both species (included in Section Grandifolia) share 5 -veined phyllaries with obtuse apices, 5-6 ray florets with 3-lobed corollas, 4 -veined pales subtending ray florets, 159 disc florets with 5-lobed corollas, 2-veined disc pales, and pilose achenes. Clibadium glabrescens differs from C. pentaneuron by having 4 strigose phyllaries and 8-11 disc florets vs 3-4 glabrate phyllaries and 5-8 disc florets in C. pentaneuron.

All the characters of C. glabrescens coincide with those used to characterize the new species, C. napoense H. Robinson, from Napo, Ecuador (the same eastern slope of the Cordillera). Clibadium napoense has the same floral structure and vegetative characteristics as C. glabrescens. The only character that differs is the occasional presence of 3 phyllaries (although some heads have 4 phyllaries, e.g., Mexia 7110: Stuessy & Jansen 4981), contrasting with the more usual 4-phyllary condition in C. glabrescens.

19. Clibadium pentaneuron S. F. Blake, Contr. U.S. Natl.

Herb. 22: 598. 1924.--Type: Colombia, 1906, Lehmann 1256 (holotype: GH!, fragment and photo: US!; isotypes: F [2]! photo: US!, K! photo: OS!, NY!).

Clibadium oileorubrum Cuatrec., Rev. Acad. Colomb. Cienc. Exact. 9: 238. 1954.--Type: Colombia. Huila: Comisaria Caqueta, filo divisorio de la Cordillera Oriental, 21 Mar 1940, Cuatrecasas 8422 (holotype: F!, photo: US!). 160

Clibadium sarmentosum Cuatrec., Rev. Acad. Colomb.

Cienc. Exact. 9: 239. 1954.--Type: Colombia. Valle: Hoya del Rio Cali, lado derecho del Rio Pichinde, cuchilla de Los Carpatos, 24 Jul 1946, Cuatrecasas 21660 (holotype: F!, photo: US!; isotypes: COL!, P photo: OS!).

Clibadium scandens Cuatrec., Rev. Acad. Colomb. Cienc.. Exact. 9: 239. 1954.--Type: Colombia. Putumayo: Valle de Sibundoy, La Cabana, 2 Jan 1941, Cuatrecasas 11596 (holotype: F!, photo US!; isotype: COL!).

Clibadium funkiae H. Robinson, Phytologia 73: 149. 1992.--Type: Colombia. Antioquia: 3 Ion SE of Santa Elena on road from Medellin to Santa Elena and Rio Negro, 15 km IJ of Rio Negro, 30 Jul 1979, Stuessv & Funk 57 09 (holotype: US!; isotype: OS [2]!).

Clibadium zakii H. Robinson, Phytologia 73:153.

1992.--Type: Ecuador. Bolivar: carretera Chillanes- Bucay, en la Hacienda Tiquibuso del Sr. Gonzalo Gomez, 10 Sep 1987, Zak & Jaramillo 2881 (holotype: US!;

isotypes: F! MO!)

Arching or scandent shrub, up to 2 m tall; stem scabrous to strigose with appressed hairs of 1 mm long. Leaves with blades ovate to ovate-lanceolate, thick, coriaceous, 7-16 cm long, 3-7 cm wide, 5 -nerved, primary and secondary veins prominent underside, at apex acuminate, at base cuneate to rounded cuneate, with margin serrate, teeth less than 1 mm high; dark green and sparcely strigose both sides, with hairs 0.5-1 mm long; petioles 10-15 mm long scabrous to strigose with hairs 1 mm long. Capitulescence paniculiform, always trichotomously divided at ending of branches and stems. Heads sessile, or on 2 mm long pedicels, subglobose, 4-5 mm high, 3-4 mm wide; phyllaries 3-4, suborbicular, 4-5 mm high, 3-4 mm wide, 5 -veined, at apex obtuse, glabrous, slightly hirsute and ciliate on upper one-third. Ray florets 5-6, white, subtended by ovate pales, 4 mm long, 3 mm wide, 4 -veined, slightly hirsute on upper one third; corolla 2 to 2.5 mm long, throat l to 1.5 mm diameter, 3-lobed, lobes irregularly deltate, 0.5 mm long, at apex scarcely pubescent; styles 1,5 mm long, branches 1 mm long. Disc florets 5-8, subtended by (exceptionally, epaleaceous central disc floret), lanceolate pales, 3.5 mm long, 1.5 mm wide, at apex slightly ciliate, 2- veined; corolla white, 2.5 mm long, tube 1.5 mm long, 0.5 mm diameter, 5-lobed, lobes 0.5 mm long, deltate, at apex slightly pilose; anthers black, 1 mm long; styles 2 mm long undivided; ovaries sterile, 1.5 mm long, 1 mm diameter, at apex pilose, with hairs 0.5 mm long. Achenes 2 mm long, 1.5 mm wide, pilose on upper half with hairs 0.5 mm long. Chromosome number, n = 16. Phenology. Flowering June to January, in fruit August to January. Distribution (Fig. 16). Colombia. Ecuador; cloud forest to subparamo; (100-), 1400-2800 m. 162

Representative Specimens. Colombia. Antioquia: Municipio de Frontino, on road to Murri, 15 km W of Nutibara, Brant & Martinez 137 8 (BM, K, MO); 13 km from Nutibara; forest edge east of road, McPherson 13396 (MO); km 17 of road Nutibara-La Blanquita, region de Murri, Zarucchi et al. 7 097 (MO); Santa Elena, Archer 1150 (US); along road from Medellin to Rio Negro, Killip et al. 39842 (US); bajo la cumbre de Santa Elena, camino entre Medellin y Rio Negro, Pelaez et al. 510 (US); San Antonio de Pereira, Hermano Daniel 4205 (US); near Las Palmas, camino entre Medellin y El Retiro, Medina 249 (GH); Diaz 47 (COL); San Puro, Hermano Tomas 17 5 (F).

Boyaca: 9 km N of Arcabuco on road to Bambita, ca. 5 km S of Boyaca-Santander state line, Stuessy & Funk 5595 (OS). Caqueta: 30 km SE of Guadelupe on road to Florencia, Gentry et al. 9036A (COL, MO). Caldas: Salento, Rio Santa Rita, Killip & Hazen 897 6 (GH). Choco: Carretera Anserma-Nuevo-San Jose del Palmar, entre el Alto del Galapago y San Jose del Palmar, Forero et al. 227 9 (MO, OS). Cundinamarca: Eastern

Cordillera, ca. 20 km NE of Fusagasuga, King & Guevara 57 54 (F, NY, US) ; ca. 18 km NE of Fusagasugci, King & Guevara 57 64 (F, NY, US); Montes abajo del Tequendama, en la base del Cerro del Ermitano, Uribe 4098 (COL); Montes SE de Santandercito, cerca al Cerro El Ermitano, Uribe 4917 (COL, US). Narino: Municipio de Barbacoas, 1 km de Junin, carretera Tumaco-Pasto, Benavides 4688 (MO). Putumayo; Valle de Sibundoy, 2 km SW of Sibundoy, Bristol 519 (GH, US) .

Santander-Boyaca: cerca al limite, en la carretera de Gambita a Arcabuco, Uribe 6310 (COL, US). Valle: Cordillera 163

Occidental, vertiente occidental, Hoya del Rio Anchicaya, entre Sabaletas y la Quebrada del Tatabro, Cuatrecasas 22034

(F); Bajo Calima, 15 km N of Buenaventura, at end of "Gasolina" road, Faber-Langendoen & Hurtado 1856 (MO); Carton de Colombia concession, Gentry et al. 4 0293 (MO); Concesion Pulpapel/Buenaventura, Monsalve 624 (MO), Monsalve 1601 (MO).

Clibadium pentaneuron is restricted mostly to high elevations (1400-2800 m) were it shows thick coriaceous leaves and heads 4-5 mm tall arranged in panicles of open clusters (6-10 cm wide) at the ends of trichotomously divided branches. Four specimens (Faber-Langendoen & Hurtado 1856: Gentry 40293; Monsalve 624. 1601) collected below 100 m, however, have ovate-lanceolate to lanceolate leaves, thin blades, heads with 3 ray and 6 disc florets all subtended by pales, and strigose phyllaries and pales. Also each head has a long pedicel (0.4 mm long) contrasting with the short one

(0-2 mm long) observed in typical high altitude populations. Probably one of the reasons for considerable morphological variation observed in these lower elevation specimens is the modified habitat where these plants are growing. The lack of nutrients (P, Ca, K, Mg, Cu, B, Mn, Zn) and high concentration of Al (according to labels of the four cited specimens) characterizes poor soils of the region. This degraded soil condition may be the result of the activity of the pulp industry (Carton de Colombia) located in that area. 164

Clibadium pentaneuron has a close relationship with C. glabrescens and both species (included in section Grandifolium) have heads with orbicular 5-veined phyllaries with obtuse apices, 5-6 ray florets with 3-lobed corollas, subtended by 4-veined orbicular pales, disc florets subtended by 2-veined pales, corollas 5-lobed, and pilose achenes. Clibadium pentaneuron differs from C. glabrescens in its glabrate phyllaries and 5-8 disc florets vs. strigose and 8-

11 in C. glabrescens.

Clibadium scandens was described by Cuatrecasas (1954) based only on the type material, in which the branches are scandently arranged and with heads organized in more condensed clusters. Cuatrecasas (1954) also described two new taxa, C. pileorubrum and C. sarmentosum. which were related to C . pentaneuron. Reexamination of these types clearly suggests that C. pileorubrum differs from C. pentaneuron only in having smaller leaves (3-4 cm long, 2-3 cm wide; vs. 7-16 cm long, 4-7 cm wide, respectively) and more acuminate pales (acute in C. pentaneron). Clibadium sarmentosum is a densely pilose variation of the more normally scabrous to shortly strigose pubescence in C. pentaneuron. Therefore, and because there is no geographic integrity to any of these minor morphological variations, they are treated as synonyms. IIC. Clibadium L. section Clibadium Clibadium L., Mant. Pi. 161. 1771.--Type: Clibadium

surinamense L.

Leaves ovate to ovate-lanceolate; heads arranged in cymose, thyrsoid or paniculiform capitulescences; with only ray florets subtended by pales; phyllaries 5-7-veined; ray florets 3-6, with ray corollas 3-lobed, and pales 5 -veined; disc florets 5-14, with corollas 4- or 5-lobed. Species 20-

25.

20. Clibadium laxum S. F. Blake, Wash. Acad. Sci. 16: 418 1926.--Type: Ecuador. Guayas: Teresita, 3 km W of Bucay 5-7 Jul 1923, Hitchcock 20430 (holotype: US!; isotypes: GH! NY!).

Clibadium alatum H. Robinson, Phytologia 73: 151-152.

1992. Type: Ecuador. Carchi: environs of Maldonado, 3 Jun 197 8, Madison. Plowman. Kennedy & Besse 4940

(holotype: US!; isotype: F!).

Arching branched shrub up to 4 m tall, one main stem at base in juvenile, 2.5-3.5 cm diameter, branched at base in older plants; stem many-angled, often flattened near new branches, densely short strigose with hairs 1 mm long. Leaves with blades 14-23 (-30) cm long, 8-15 (-23) cm wide, at apex acuminate, at base truncate-rounded, and shortly decurrent on the petiole, with margin serrate to irregularly serrate, both surfaces strigillose, with hairs l mm long; petioles sulcate above, marginate by the decurrent blades, 3- 6 cm long, strigillose with hairs 1 mm long. Capitulescence paniculiform, with many loosely cluster of heads, usually ternately divided on peduncles up to 6 cm long. Heads 5-6 mm high, 3-4 mm diameter; phyllaries 3-4, obovate to suborbicular, 4 mm long 2.5-3.5 mm wide, 5-7 veined, green below, whitish above, at apex acute, strigose with hairs 1 mm long, with margin ciliate to the upper one-third. Ray florets 9, subtended by ovate pales, 3.5-4 mm long, 2.5-3 mm wide, strigose with hairs 0.8 mm long, at apex acute, 5- nerved; corolla white, glabrous, 2 mm long, 0.8 mm diameter, 3-lobed, lobes deltate, 0.5 mm long, at apex scarcely pubescent; styles 1.5 mm long, branches 1 mm long. Disc florets 8-14, subtended by lanceolate (occasionally, 1-2 epaleaceous central florets), pales, 3-3.5 mm long, 1.5 mm wide, at apex scarcely ciliate, 3-veined; corolla white, 1-8- 2 mm long, l mm diameter, 4-lobed, lobes deltate, 0.5 mm long, at apex scarcely pubescent; anthers black, 1 mm long; ovaries sterile, 2 mm long, 1.5 mm diameter, pilose toward apex with hairs 1 mm long. Achenes 2.5 mm long, 1.5-2 mm diameter, pilose toward upper one-third, with hairs l mm long. Chromosome number, n = 16. Phenology. Flowering from May through October.

Distribution (Fig. 17). Ecuador, broadly distributed in most of the central provinces; tropical wet forest, common along streams; 600-1800 m. Fig. 17. Distribution of Clibadium laxum ( • ) and C. sodiroi ( t ) in Ecuador. Representative Specimens. Ecuador. Azuay: 18 km E of Jesus Maria border with Canar, Gentry et al. 28484 (MO). Bolivar: Balsapampa, Harling et al. 9680 (F) . Carchi: Maldonado-Tulean road, km 2-6, Werling & Leth-Nielsen 84 (F). Chimborazo: foothills of the western cordillera near the village Bucay, Camp 3826 (MO, NY); N edge of Bucay on route to Teresita, Stuessy et al. 4942 (OS). Cotopaxi: foothills above Valencia near cane mill, Mathias et al. 5191 (F). El Oro: Monos, 14 km WNW of , Fosbera et al. 22969 (US); 13 km W of Pinas on road to Santa Rosa, King & Almeda 7 97 2 (MO); 36.6 km ESE of La Avanzada on gravel to Pinas, Stuessy & Nesom 5867

(OS). Esmeraldas: Ventanas, km 319 on Quito - San Lorenzo, Jativa et al. 804 (UC, NY); near La Playa Grande, Cayapa River, Jdtiva et al. 2102 (NY); Parroquia de Concepcion, bank of Rio Santiago above Playa Rica, Mexia 8467 (F; GH; K, MO; NY; US); down Rio Palavi, 1 km from camp, Lorentzen & Gelpi 3792 (MO). Guayas: Foothills of the western cordillera near the village of Bucay, Camp 3826 (GH; MO; NY; US). Leon: Canton Pijili, Hacienda Salento, Mexia 6696 (LL; US). Los Rios: Rio Palenque Biological Station, km 56 route Quevedo- Santo Domingo, Dodson et al. 6385 (OS). Pichincha:

Cooperativa Santa Marta 2 along Rio Verde, 2 km SE of Santo Domingo de Los Colorados, Dodson et al. 7 589 (MO); Palmitopamba, ca 10 km NW of Nanegal, Harling et al. 11525 (MO); Reserva Forestal Endesa, Rio Silanche, km 113 route Quito - Pto. Quito, Jaramillo 6713 (MO); Jaramillo 7 559 (MO);

9 km N of Sta Rosa on dirt road to Mindo, Stuessy et al. 4877 (OS); 4 km W of Santo Domingo de Los Colorados, Stuessy et 169

al. 4902 (OS); 13.5 km NW of Nono on road to Mino, Stuessy &

Arriagada 12318 (OS); l km W of Santo Domingo de Los Colorados on road to El Carmen, Stuessy & Arriagada 12352 (OS). Zamora: 4 km NE of Cumbaratza on road to Zumbi,

Stuessy & Nesom 5901 (OS); 0.5 km NE of Zamora on road to Timbera, along banks of Rio Zamora, Stuessy & Nesom 5897 (OS) .

Clibadium laxum is characterized by having an arching branched stem but with one main stem at base, large leaves in old individuals (30 cm long, 20 cm wide; e.g, Camp 3826. Stuessy et al. 4942. Werling & Leth-Nissen 84), capitulescences with open clusters and loosely arranged heads, and drupe-like achenes that produce a deep orange juice when squeezed. Clibadium laxum is closely related to C. sodiroi (which has an eastern distribution in low humid tropical forests between 100 and 1000 m ) . Both species are placed in Section Clibadium. have 3-4, ovate, stigose phyllaries with acute apices, 3-lobed ray corollas, 4-lobed disc corollas, 5-veined disc pales, and achenes with the upper one-third pilose. Clibadium laxum differs from C. sodiroi by having three times the number of ray florets (9 vs. 3), and disc florets subtended by pales.

Clibadium alatum was described by Robinson (1992) based on material with winged petioles from ,

Ecuador. After studying the types and available specimens, it is clear that this represents only one extreme of the morphological variation within C . laxum. the basic 17 0

characters overlap completely. The large leaves with distinct wings on the petioles that were described as unique in C. alatum. are also found in many specimens of C . laxum (e.g., Camp 3826: Jativa & Epling 804: Stuessy et al. 4942; Werling et al. 84). The apex of the ray achenes in C. alatum is similar to that observed in C. laxum (e.g., Dodson et al.

7 589: Harling et al. 9680: Mexia 8467). The protologue of C. laxum gives the involucre as having 1-2 phyllaries, but reexamination of the type indicates 3-4, the typical condition in C. alatum.

21. Clibadium sodiroi Hieron., Bot. Jahrb. Syst. 29:37. 1900.--Type: Ecuador, Pichincha: near Canzacoto, "crescit locis asperis regionis subtropicae et subandinae", ca. 1894 [Hieronymus, 1901], Sodiro 22/2 (holotype: B, presumed destroyed, fragment at US!,

Photos: F ! GH! M O ! OS!).

Clibadium mexiae S. F. Blake, Journal Wash. Acad. Sci.

28: 489. 1938.--Type: Ecuador. Napo-Pastaza: near Puyo, dense forest, 350 m, 20 Feb 1935, Mexia 6949 (holotype: US!; isotype: LL!).

Clibadium pallidum Diels, Notzbl. Bot. Gart. Berlin, 14: 340. 1939.--Type: Ecuador. Napo-Pastaza: bei Puyo, Feb 1938, Sydow 807 (holotype: not located). 171

Shrubs and trees up to 6 m tall; stem up to 5 cm diameter, densely tomentose, with gray hairs 2 mm long. Leaves below capitulescence, rugose, with blades broadly ovate, 30 cm long, 20-25 cm wide, at apex acuminate, at base broadly rounded or slightly cordate, some slightly decurrent on the petiole, 3-5 pliveined, with margin crenate-dentate.

Leaves from branches, ovate, 6-15 cm long, 5-10 cm wide, at apex falcate-acuminate, at base subtruncate or shortly cuneate; densely pilose with straigh hairs 1-1.5 mm long; with margin crenate-toothed. Capitulescence paniculiform, convex or flattish at twigs tip, many headed (up to 219 heads), branches densely and griseously tomentose-pilose with spreading hairs, hairs 1.5-2 mm long. Heads, small, obovoid- subglobose, 3-4 mm high, 2-3 mm diameter; phyllaries 3, whitish, ovate, 3-3.5 mm high, 3 mm wide, 5-7 veined, strigose with hairs 1 mm long, at apex acute, shortly ciliate on upper one-third margin. Ray florets 3 (-5), subtended by suborbicular pales, 3-3.5 mm long, 3 mm wide, 5 veined, at apex obtuse, shortly ciliate at upper one-third margin; corolla white, tubulose, 2 mm long, 1 mm diameter, subequally 3-lobed, lobes irregularly deltate, 0.5 mm long, at apex scarcely pubescent; styles 1.5 mm long, branches 0.6-1 mm long. Disc florets 6, epaleaceous; corolla white, 2 mm long, l mm diameter, 4 lobed, lobes 0.5 mm long, at apex shortly pilose; anthers black, 1 mm long; ovaries sterile, 2 mm long, 1-1.5 mm diameter, pilose toward apex with hairs 1 mm long. 17 2

Achenes broadly obovate, rounded at base, 2.5 mm long, 2 mm diameter, densely short pilose at upper one-third, with hairs 0.5-0.8 mm long. Chromosome number, n, = 16. Phenology. Flowering from June to September. Distribution (Fig. 17). Only in Ecuador; tropical low humid forest, on steep slopes and roadsides; 300-1000 m.

Representative Specimens. Ecuador. Napo-Pastaza: road N from El Chaco, Quito-Lago Agrio Road, 20 km E of Baeza, Gentry & Miller 54996 (MO); cabeceras del Rio Bobonaza, ca 12 km E of Puyo, Lugo 4605 (MO); Near Puyo, Mexia 694 6 (UC); Carretera Hoi1in-Loreto, 3 km E Rio Pucuno, camino al caserio El Pacto, Neill et al. 8525 (MO); forest at Puyo, Prescott 404 (DS, NY); 5 km E of Mera on road to Puyo (3.6 km W of

Shell-Mera), Stuessv & Jansen 4967 (OS), 4968 (OS); 2.7 km E of Puyo on road to Veracruz, Stuessy & Jansen 4969 (OS); 17 km N of Puyo on road to Tena, Stuessy & Jansen 497 8 (OS);

34.6 km N of Puyo on road to Tena, Stuessy & Jansen 4980 (OS); Village of Topo, near Rio Topo, ca 2 km toward Hacienda La Mascota, Stuessv & Jansen 4987 (OS); w edge of Puyo along roadside, Stuessy & Nesom 5817 (OS); 3.6 lcm W of Puyo on road to Banos, Stuessy & Nesom 5819 (OS); eastern foot-hills of the Andes, Skutch 4402 (A, F, GH, MO, NY). Santiago-Zamora: near Mendez, Camp E-884 (NY). Tungurahua: Rio Verde, Har1ing et al. 10153 (F) .

Average mature individuals of Clibadium sodiroi have very soft pubescence on leaves and stems. Mature fruits also 17 3 produce a small quantity of orange juice (stems and heads also sometimes produce small amounts of orange sap; pers. field observ.). The large leaves (20 cm long, 16 cm wide) below the capitulescence are often cordate or subcordate at the base, which is a useful diagnostic field character. Also helpful is the large number of small (3-4 mm tall) heads per capitulescence: 219 heads in young individuals (e.g., Stuessv & Arriagada 12363); 819 heads were counted in one mature and highly branched individual (Stuessy & Arriagada 12362).

Clibadium sodiroi is closely related to C. laxum. but it differs in having a small number (3) of ray florets and 6 disc florets without pales, contrasting with 9 ray florets in C . laxum arranged in two series, and 8-14 disc florets subtended by pales.

This species commemorates Luigi Sodiro (1836-1909), clergyman, professor of botany and plant collector in Ecuador

(Stafleu & Cowan, 1985).

22. Clibadium surinamense L., M a n t . PI. 294. 1771. --Type: Suriname, exact locality and date unknown, Allamand s.n. (neotype: K, Hostmann 647. designated by Hind in Jarvis, Barrie, Allan & Reveal, 1993).

Trixis aspera Sw., Prodr. Veg. Ind. Occ. 815. 1788. 17 4

Clibadium asperum (Aubl.) DC., Prodr. 5: 506. 1836. Baillieria aspera Aubl., Hist. Pi. Guianae 2: 804,

£. 317. 1775.--Type: French Guyana, "habitat Caiennae & Guianae locis inclutis," 1762-64 [Urban, 1903], Aublet s.n. (holotype: BM!, photos: GH! OS!; isotype: G-DC, IDC 28.925:1.5,6!).

Oswalda baillerioides Cass., in Diet. Sc. Nat. 59. 322. 1829.

Clibadium caracasanum DC.. Prodr. 5: 506. 1836.--Type:

Venezuela. Distrito Federal: Caracas, 1830, Vargas 292 (holotype: G-DC, IDC 28.925:1.3,4!; photo: GH!).

Clibadium trinitatis DC., Prodr. 5:505. 1836.--Type: Trinidad, 1825, Sieber 71 (holotype: G-DC, IDC 28.925: 1.2!; isotypes: K! photo: OS!, P photo: OS!).

Clibadium villosum Benth., Plant. Hartweg. 2. 205. 1845.--Type: Colombia. Bogota: near the village of Tena, Feb-Apr 1843, Hartweg 1139 (lectotype here designated: K!, fragment at US!, photos: GH! OS! US [2]!; isolectotypes: BM! photo: OS!, K! photo: OS!, G photos: F ! MO! OS!).

Clibadium lehmannianum O.E. Schulz, Bot. Jahrb. Syst.

46:620. 1912.--Type: Colombia. Playa, Rio Timbiqui, 17 5

1910, Lehmann 9 056 (holotype: B destroyed, photos:

F! GH! MO! NY! US!; isotype: K! photo: OS!.

Clibadium lanceolatum Rusby, Descr. S. Amer. Pi. 150.

1920.--Type: Colombia. Magdalena: Santa Marta Mts., 1898-99, Smith s.n. (holotype: NY!, photos: US [3]!).

Clibadium surinamense var. macrophyllum Steyerm.,

Fieldiana, Bot. 28: 629. 1953.--Type: Venezuela. Sucre: Cerro Turumuquire, north-facing slopes between La Trinidad and Quebrada El Boqueron, SW of Cocollar, 3 May 1945, Steyermark 62437 (holotype: F!; isotype: US!).

Shrub, 1.5-4 (5) m tall; stems 1-2 cm diameter at base, hispidulous to scabrous with hairs 0.1 mm long. Leaves with blades lanceolate to broadly ovate, rough chartaceous, 5-12 (-20) cm long, 2-6 (-12) cm wide, at apex acute to acuminate, at base obtuse (sometimes attenuate or slightly decurrent to petiole), with margin serrate, crenate-serrulate or serrate- dentate, upper surface weakly to moderate hispidulous with hairs 0.5 mm long (occasionally with close erect hispid pubescence), dark green, lower surface strongly hispidulous, pale gray green; petioles 1.5-5 cm long, hirsute with hairs 0.2-0.5 mm long. Capitulescence thyrsoid or paniculifor, with 10-180 heads. Heads sessile, obovoid, 3-5 mm high, 3-4 mm diameter, yellowish; phyllaries 3-4, coriaceous, concave, broadly ovate, 4 mm long, 3-5 mm wide, at apex acute, strigose with hairs 0.5-0.8 mm long, with margin upper one- 17 6 third ciliate, 5 -veined. Ray florets 3-5, subtended by ovate pales, 4 mm long, 3-4 mm wide, 5-veined, at apex acute, strigose with hairs 0.5 mm long, with margin upper one-third ciliate; corollas 2 mm long, 0.5 mm diameter, 3-lobed, lobes irregularly deltate, 0.4 mm long, scarcely pubescent at apex; styles 3 mm long, branches 1.5 mm long. Disc florets 10-14, epaleaceous; corollas tubular, 3 mm long, throat 2 mm long,

0.5 mm diameter, 4-lobed, lobes irregularly deltate, 0.5 mm long, scarcely pubescent at apex; anthers purple to black, 1.6-1.8 mm long; styles 4 mm long, branches 2.5 mm long; ovaries sterile, 2 mm long, 0.5 mm diameter, villous at apex, with hairs 0.5 mm long. Achenes 2.5 mm long, 2 mm diameter, villose on upper half. Chromosome number, n = 16. Phenology. Flowering through the entire year.

Distribution (Fig. 18). Brasil, Colombia, Costa Rica, Cuba, Ecuador, El Salvador, Guatemala, Honduras, Jamaica, Lesser Antilles (Grenada, Martinique, St. Vincent), Nicaragua, Panama, Suriname, and Venezuela (introduced to Borneo, Java, Mauritius, and Sumatra); 0-1000 (-2000) m.

Specimens Examined: Brasil. Amazonas: Rio Branco, Campus Universitario, Coelho et al. 1739 (F). Sao Paulo, Rio Solimoes, Barrio de Odorio, Lleras et al. P17 37 6 (F); Rio Negro near San Luis, Maia et al. 144 (MO); Municipio de Tefe: banks of Lago Tefe, Plowman et al. 12080 (F); Rio Mucajax, Posto Mucajai, Prance et al. 11151 (F); San Gabriel da

Cachoeira, Spruce 2136 (F); Rio Branco: below Serra Sabang, Maauire & Maguire 40336 (NY), Rio Cotinga, between Vista 17 7

— 20

— 10

— 10

20

— 30

— 40

Fig. 18. Distribution of Clibadium peruvianum {y ) and CL surinamense ( @ ) in Argentina, Bolivia, and Peru, and in the Caribbean, Central, and northern South America, respectively. 17 8

481 (US); 8 km de Sabanalarga, Callejas et al. 2285 (F, MO) ; between Salgar and Main Medellin-Quibdo Hwy, Croat 69937 (MO); Heliconia, Hermano Daniel 3986 (F); valley of Rio Anori between Dos Bocas & Anori, Denslow 4 07 (WIS); 26 km S of Zaragoza, Planta Providencia, Denslow 227 6 (MO, WIS); Angostura, Fosberg 21619 (US). Cauca Valley, 10 km N of Valdivia, Gentry & Fallen 17 942 (MO); Angelopolis, Gutierrez

& Barklay 17 0657 (TEX, MO, US); entre Medellin y El Retiro, Mejia 86 (GH); entre El Retiro y La Ceja, Uribe 4340 (COL),

Bajo Magdalena, Casabe, Laguna del Miedo, 5873 (COL, MO). Medellin, along Medellin-Santa Fe de Antioquia,Alto de Boqueron, Zarucchi & Brant 5286 (MO). San Jeronimo, km 10 of road San Jeronimo-Medellin, Zarucchi et al. 5443 (MO). Bolivar: Boca Verde on Rio Sinu, Pennell 4219 (NY). Caldas: near La Dorada, Haught 2124 (US); Cordillera Central,

Salento, Rio Santa Rita, Killip & Hazen 897 6 (US); Rio Quindio, above , Pennell et al. 87 04 (GH). Santa Cecilia, Pueblo Rico, Sneidern 5167 (F, US). Caqueta: Rio

Orteguaza, Quebrada de Miramar, Cuatrecasas & Soderstrom 27121 (US); 6 km SE Florencia, Davidse et al. 5649 (COL). Cauca: Chisquio, Finca Los Derrumbos, Asplund 10544 (US) ; Timbio, Cuatrecasas & Perez-Arbelaez 6112 (F); Popay&n, carretera al Tambo, Garcia-Barriga et al. 1277 2 (COL); Cauca valley, 10 km N of Valdivia, Gentry & Fallen 17 942 (COL); Mosquera, El Tambo, Idrobo & Fernandez .69 (COL); Sneidern 1540 (US); Rio Sucio, Pennell & Killip 7217 (GH, US); El Saladito above Cali, road to Buenaventura, Pittier 749 (F). Choco: Cerro del Torrea, Silverstone-Sopkin 1254 (MO); Lloro, 50 km S of Quibdo, Archer 2058 (US). Cundinamarca: Sueva y Gacheta, Cuatrecasas & Jaramillo 11994-A (COL, F); Estacion Santana, Dugand & Jaramillo 3854 (US); Caparrapi, Hacienda "Saldana", Garcia-Barriga 7716 (COL); Cerinza, Rio Blanco

Valley, Grant 97 59 (US); 4 km NE of Fusagasuga, King & Guevara 57 91 (F, US), 3 km SE of Chipaque, 5814 (F), ca. 13 km ENE of La Mesa, 5852 (F, US); 28 km from Bogota via Salto de Tequendama-El Colegio road, Luteyn et al. 4833 (F, MO); Cano Negro, afluente del Rio Negrito, Murillo et al. 560 (COL, F, NY); 69 km NW of Bogota, Olsen & Escobar 596 (WIS); Pacho, colinas del Rio Patasia, Uribe 5715 (COL); Quetame, vereda de Monterredondo, Uribe 6957 (COL). Huila: Hacienda Pensilvanica, Little 8114 (COL, US); San Agustin, Parque Arqueologico, Romero Castaneda 6689 (COL). Magdalena: cercanias de Bonda, Diaz-Piedrahita 181 (COL). Meta: carretera de Bella Vista a Pinalito, Echeverry & Jaramillo 2322 (COL); Villavicencio, along Quebrada Canabrava, Killip 34462 (COL); along the Coho Rosa Blanca, Kirkbride 393 (COL). Narino: Gorgonilla Island, Killip & Garcia 33061 (COL). Santander: region de Ocana: entre La Maria y

Jurisdicciones, Cuatrecasas & Rodriguez 27933 (F); La Cabuya, Cuatrecasas et al. 12086 (COL, F, GH); Salida de Durania, camino de la Donjuana, Garganta 853 (F); frontera Colombo- Venezolana, Hoya del Rio de Oro, Garcia-Barriga & Lozano 18325 (COL); Magdalena Valley, vicinity of Barranca Bermeja, Haught 1914 (COL), Haught 2047 (NY); vicinity of Chinacota,

Killip & Smith 207 83 (US); about 1 km S of Floridablanca, Langenheim 3264 (COL); Campo Capote and vicinity, Nee & Mori 180

37 80 (MO); Puerto Araujo, Renteria et al. 1831 (COL, MO); Rio de Oro, Cano del Norte, Solange 9 (COL); La Mosquitera, Woronow & Juzepczuk 47 88 (US). Valle: Calima Valley, NW of

Restrepo, Bridgeman 55 (US); between El Saladito and Felidia, Croat & Watt 7 0571 (MO); Bajo Calima along road from Buenaventura to Bahia Malaga, Croat & Watt 7 0305 (MO); Rio

Naya, Puerto Merizalde, Cuatrecasas 14004 (US); Rio Cajambre, Cuatrecasas 17534 (F) ; Pichinde, El Abismo, Cuatrecasas 18653 (F, US), entre Las y La Carbonera, 22478 (F), Rio San

Juan abajo de Queremal, 23808 (F), Monte La Guarida, La Carbonera, 22163 (COL); Buga, El Vinculo, Jardin Botanico,

Cuatrecasas et al. 27 504 (NY, US); Bolivar, via Primavera al Alto de los viejos, Devia 977 (MO); Cordillera Occidental; entre El Valle y Choco, Dugand & Jaramillo 3014 (US); carretera Ansermanuevo, 26 km E de San Jose del Palmar, Forero et al. 2824 (MO); Zarzal, between Caserio La Paila and Zarzal, Gentry et al. 54125 (MO); Cartago, Hacienda La Silvia, Jaramillo 4030 (NY); Rio Digua Valley, Chorrera La Elsa, Killip 34815 (COL); San Antonio, W of Cali, Killip & Garcia 33867 (GH, US); Hacienda Valparaiso, Zarzal-

Bugalagrande, Perez-Arbelaez & Cuatrecasas 6441 (COL, US). Vaupes: Bajo Vaupes, Gutierrez & Schultes 944 (COL, GH); Rio Kuduyari middle and lower course, Schultes & Cabrera 17 861

(US). Costa Rica. Alajuela: Rio Catarata, NW of San Ramon, Barringer 3211 (CR, F); slopes above Rio , Burger & Stolze 5144 (US); Canton de Upala along Rio Chimurria Grayum et al.; 9041 (F); l km S Rio Penas Blancas on road 142 between Florencia and Fortuna, Hammel et al.14039 (CR); , S side of dam, Funk et al.10654 (CR); 5-10 mi W of on route CA-1, Stuessy & Gardner 4451 (OS); ca. 9 mi N of Zarcero on route 15, Stuessy & Gardner 4460 (OS); ca. 1 mi S of La Marina, Stuessy & Gardner 4463 (OS); 1 mi Quesada side of junction to Muelle and Santa Clara, Weston 4407 (CR, OS). Cartago: S slope Irazu Volcano, near , Arriagada 142 (OS); San Gerardo, Reserva Biologica, Arriagada 143 (OS); 5 km SE of Rio General from San Isidro del General to , Arriagada 149 (OS); Valley of Rio Revetazon, 3 km SE of Turrialba, Holm & litis 7 (NY); Linda Vista Ltda. Nursery, near Cartago, Raven 22048 (F. DS, NY); 5.6 mi SSE of Paraiso on road to Cachi, Stuessy & Gardner 4489 (OS); E slopes of Tuis and N of Platanillo, Wilbur & Stone 10661 (CR, F, OS). Guanacaste: Guayabo on road to

Aguas Claras, Almeda et al. 3145 (OS); Parque Rincon de la Vieja, Hacienda Santa Maria, Herrera 939 (F). Heredia: road to Puerto Viejo, km 54 from Barva, Arriagada 163 (OS); from Puerto Viejo to Horquetas, km 10, Arriagada 164 (OS); Finca La Selva, OTS field station, Gravum 27 00 (MO), McDowell 715 (F), Chacon 832 (F); 4 mi N of San Isidro, S slope of Volcan

Poas, Turner & Turner 15030 (CR). Limon: from Guapiles to San Jose, 14 km before Guapiles, Arriagada 310 (OS); Parque Nacional Tortuguero, Estacion Agua Fria, Robles 17 22 (F); near Carmen Station on Indiana Branch, Standley & Valerio 48375 (GH); W of Limon, road to Station Reta, Utley 6059 (F).

Puntarenas: 5 km before , Montes del Aguacate, Arriagada 131 (OS); Villa Neily to San Vito, 15 km N of Villa Neily, Arriagada 150 (OS); between Villa Neily and Paso 182

Canoas Arriagada 151 (OS); km 10, after locality La Mona, between Golfito and , Arriagada 155 (OS); km 160 on route 2, between Volcan and San Isidro, Arriagada 159 (OS).

San Jose: La Palma, road to Dominical, Almeda et al. 3304 (OS); San Pedro, Campus of University of Costa Rica, Gentry 948 (CR); vicinity of El General, Skutch 27 33 (GH). Ecuador. Bolivar: Balsapampa, Harling et al. 967 0 (F, MO).

Canar: ca. 77 km ESE of Guayaquil, King & Garvey 6871 (CAS, MO, US). El Oro: 4 8.5 km ENE of La Avanzada to Pinas, Stuessy & Nesom 587 3 (OS). Esmeraldas: Quininde to Esmeraldas, Holm-Nielsen et al. 7 07 0 (F). Guayas: vicinity of Naranjito, Camp 3601 (OS); Milagro, Hitchcock 20261 (GH, NY); Naranjal, junction with road to Guayaquil, King & Garvey 7005 (CAS, MO, US); l km NW of Milagro, Stuessy et al. 4935 (OS); 18.1 km N Naranjal, banana plantations, Stuessy & Nesom 5856 (OS). imbabura: Lita, Acosta Solis 12285 (F). Loja: 24 km S of Catamayo, road to Cariamanga, King & Almeda 7 951

(CAS, MO). Los Rios: Hacienda Clementina on Rio Pita, Asplund 5389 (F); Canton Vinces, km 7 0 Quevedo-Palenque, Dodson & Gentry 9817 (CAS, MO); Estacion Biologica Pedro Franco Davila, Dodson & Gentry 127 03 (MO); near Quevedo, Hacienda San Jose, Mexia 6595 (CAS, F, US); trail from Quevedo to El Naranjal, Mexia 6668 (CAS, NY); Rio Cristal, near Montalvo, Jativa & Epling 103 (CAS, NY); 1 km E of Montalvo near Pisagua, Stuessy & Jansen 4928 (OS). Manabi:

13 km E of Alajuela, road to Quevedo, Stuessy et al. 4919 (OS). Napo: Puerto Francisco de Orellana, town at Rio Napo, Balslev & Madsen 10652 (F, MO); Rio Napo between Coca and 183

Hacienda , Harlina & 11876 (MO); Santa Rosa, Lugo IS2 (F, WIS), 224 (A, CAS), Ahuano, 228 (F), Puerto Napo, 1027 (CAS, WIS), Tierra Colorada, ca. 3 km N of Coca, 2926 (F); San Jose de Payamino, 40 km W of Coca, Irvine

363 (F); Zaruma et al. 66 (F). Pastaza: cabeceras at Rio Bobonaza, ca. 12 km E Puyo, Lugo 1343 (F); Chiquita, Lugo

1371 (A, CAS, F, WIS); Puerto Ubilla, Lucto 17 03 (F), Puerto Sarayacu, 3890 (MO), Sarayaquillo, 3986 (F), vicinity of El Porvenir, ca. 5 Ion of Puyopungu, 4941 (A, CAS, F, WIS). Tungurahua: La , Valley of Rio Pastaza, Asplund 7 614 (A, F); bridge over Rio Blanco, Stuessy & Jansen 4954 (OS).

Zamora-Chinchipe: near Zamora, Knight 711 (WIS). French Guyana: vicinity of Cayenne, Broadway 37 6 (GH, US), Broadway 634 (US); vicinity of Orstom Research Station, Croat 53802

(MO); Montagne des Chevaux, Gentry & Morawetz 63181 (MO); Crique Jacques, Wachenheim s.n . (US). Honduras. El Paraiso: entre El Junquillo y Teupasenti, Molina 11863 (F, NY); Sierra El Chile, entre El Junquillo y El Robledal, Molina 14147 (F, NY, US); Cordillera Dipilto, highway to Las Manos, Molina 30573 (F, MO), near border of Nicaragua, 3 km N of Las Manos, Williams et al. 42259 (F, US). De Gracias a Dios: caserio de Rus-Rus, Nelson & Romero 407 8 (MO). Morazan: Cordillera de Misico o Volcan de Guaimaca, Molina 3148 (F, GH, US). Olancho: poblacion de Culmi, Nelson & Romero 4653 (MO). Java. Nirmala Estate, Kalimun area, Balgooy & Wiriadinata 2903 (A); Batavia, ad ripas fluminis Tjiliwong, Schiffner 2819 (K); Gunnong Salak, Sinclair 10052 (K) . Mauritius: Perrier Nature reserve near Mare aux Vacoas, Lorence 1852 (K). Nicaragua. Boaco: Cerro Mombachito, Aranda et al. 32 (OS); ca. 5 km NW de Camoapa, Grijalva 2859 (F); El Tablon, 2 km N of Camoapa, Moreno 10613 (CAS, F). Chontales: road from Juigalpa to Santo Tomas, Neill 7394 (GH); E of El Naranjal, between Juigalpa and La Libertad, Stevens & Krukoff 22445 (F); ca. 4.5 km S of Highway 7. road to Granada Williams 20029 (WIS). Jinotega: Nueva Guinea, Stevens 4928 (OS); N slope of Volcan Yali, Stevens 15140 (F, OS); along road from Hwy 3 through La Fundadora, Stevens et al. 15401 (F). Madriz: ca. 5 km SW of San Juan de Rio Coco, Stevens et al. 17702 (F, OS). Matagalpa: carretera al Tumma, 10 km NE ciudad de Matagalpa, Guzman et al. 7 01 (CAS, F); road between

La Danta and La Luna, Stevens 9625 (OS). Nueva Segovia: 3.7 mi S of Los Mancos on route 15, Jansen & Harriman 547 (OS); 12.5 km NW de Matagalpa, carretera al Tuma, Moreno 10313 (F) ; Rio San Juan: 2 km NW of Sabalos, Moreno 23298 (OS). Zelaya: ca. 5 km S de Waslala, Grijalva & Moreno 1180 (F); on road to Los Pintos Miller & Sandino 1221 (CAS, F); Mt. Liveco near Siuna, Madregava, Nelson 5059 (CAS); N road between Puerto Cabezas and Rio Wawa, Stevens & Krukoff 10659 (F); along Rio Sucio, ca. 0.5 km E first suspension bridge,

Stevens 12330 (OS); SW of Bluefields, Stevens & Krukoff 197 99 (F). Panama. Canal Zone: Barro Colorado Island, Wheeler Trail, Hood 977 (F); 20 mi S of Colon, Kina 5257 (US); Pipeline Road, 1 km N of Rio Agua Salud, Schmalzel et al. 1023 (MO); behind Frijoles Train Station, Stimson 5274 (COL); Chiva Chiva Trail near Miraflores Lake, Tyson 1411 (MO); Ft. Sherman to Gatun Locks, Tyson 6286 (FSU); Pipeline Road, 10 mi N Gamboa, Tvson 6312 (FSU); Fort Kobe road, Woodson et al. 1408 (GH, NY). Chiriqui: 32 mi W of Santiago, Croat 10715 (MO); El Cope sawmill, Hammel 407 2 (MO); 15 km ESE of Concepcion on highway 1, Hartman 3926 (OS); El Llano de Hicaco, Schmalzel 1989 (MO); Rio San Cristobal, 2 mi W David, Tvson 919 (FSU); vicinity of Boquete, Woodson & Schery 7 31 (GH, MO, US) . Code: Llano Bonito N of Las Margaritas, Seibert 519 (NY). Colon: vicinity Sardinella, Blum & Tvson 487 (FSU); along Rio Boqueron, E of Salamanca, Knapp et al.

5825 (MO); 1 mi N of Col6n-Panama line, Lazor & Tyson 2983 (MO). Darien: vicinity Cana gold mine, Croat 37 982 (MO). Herrera: 5 mi S of Ocu, Wilbur et al. 12084 (CR, MO). Los Santos: 3 km S of Macaracas, Hartman 3942 (OS). Panama: La Campana, Cerro Campana, Ebinaer 3 62 (US); 4.5 mi N junction road to Chepo and Cerro Azul, Stuessy & Gardner 4540 (OS); road to Cerro Campana, Siallivan 443 (MO) ; road from Cerro Azul to Cerro Jefe, Tyson 6317 (M)); road from Chepo to El

Llano, Tyson & Smith 4122 (FSU, MO); forests near Arraijan, Woodson et al. 1359 (NY). Veraguas: 18 km W of las Minas, Hammel 4353 (MO); 8 km NE of Sona, Hartman 3932 (OS); Laguna La Yeguada, Luteyn 1447 (MO); vicinity of San Jose on

Santiago-Santa Fe road, Nee 817 6 (MO). V e n e z u e l a . Amazonas: Atabapo, N Rio Orinoco and W Rio El Ejercito, Guanchez 2088 (MO). Anzoategui: Mendocero, orilla del Rio Chive, Pittier 15061 (US). Aragua: above Guayas on road to

Los Teques, Steyermark 56890 (F, US); Parque Nacional, Williams 12241 (F, US). Avila: savana de Monte, Vogl 7 08 (F, GH). Bolivar: La Gran Sabana, junction of road Santa Elena-Cabanayen, Davidse et al. 4748 (MO); 5 km S of San

Ignacio de Yuruani, Liesner 24447 (MO); La Paragua, Killip 37592 (US); Igarape below Serra Sabag, Maguire & Maauire

40336 (MO); Mount Roraima, Glycon Swamp and vicinity, Steyermark 58625 (F, GH) ; Distrito Roscio: Cerro Ceita, Hato Divina Pastora, Steyermark & Liesner 12747 0 (MO); E Rio

Parguaza, 125 km N of Alcabala of Puerto Ayacucho, Steyermark et al. 131743 (MO); Cerro Bolivar, between Tunnel E-5 and Morrison-Kundsen Camp, Wurdack 35740 (GH). Distrito

Federal: La Florida, near Caracas, Alston 5210 (US); Alston 5324 (NY, US); Bailev & Bailev 252 (US); Pittier 7893 (F, GH, MO); Pittier 9664 (GH, NY); road near Turmerito, Pittier 13596 (LL). Merida: arriba Santo Domingo, Aristeguieta 3268 (NY); carretera Sta. Barbara del Zulia-Encontrado, Bernardi

361 (NY); along Rio Albarregas, Monte Serpa, Steyermark 5592 8 (F, NY); between San Isidro Alto and Santa Cruz de , Steyermark 56567 (F); Los Chorros, Eggers 3057 (GH); near Colonia Tovar, Fendler 640 (GH, MO). Miranda: Quebrada de Las Comadres, near Las Mostazas, Allart 230 (NY, US); Km 109 Caracas-Oriente road, 13 km E of Caucagua, Berry 917 (MO); Agua Fria, Gines 4481 (US). Monagas: El Paramo, NE Las Delicias, Steyermark 62029 (F, MO); Tabay, Gehriger 361 (F, MO); Agua Caliente, Gehriger 494 (F, US). Sucre: Peninsula de Paria, Steyermark & Liesner 120954 (MO). Sumatra. Toba: Toetoepan, Boeea 5901 (A); vicinity of Tomoean Dolok, Asahan, Boeea 9 055 (A). Suriname: 13 km NW of Parimaribo, Archer

2684 (LL); Scotelweg, 2, Archer 2647 (F, GH, US), Archer 27 25 (US); Archer 2716 (LL); Wredenburg Weg, Archer 2857 (US); 187

Patamacca Kreek, Borsboom 12293 (F) ; Paramaribo, near Agric. Experim. Station, Koster 49 (GH); Potaro-Siparuni Region, Kvist et al. 230 (F); near Kabelstation, Laniouw 12 056 (MO); vicinity Agricultural Experiment Station at Paramaribo, Maauire & Stahel 227 07 (A) : Kaieteur National Park, Pipoly & Gharbarran 9962 (MO); forest of the Station, Groningen,

Samuels 345 (GH).

Clibadium surinamense is distinguished by ovate to ovate-lanceolate leaves with a well marked reticulate and strongly hipidulous venation of the abaxial surface. Clibadium surinamense is the most common species of the genus, distributed broadly from Nicaragua to Ecuador. Because of its distribution, this species shows some variation in its vegetative structures, but the head structure and organization and inflorescence arrangement remain constant throughout. The size, shape, and pubescence of the leaves are subject to considerable fluctuation along its distributional range. To illustrate this variation, variety macrophy 1 lum was described by Steyermark (1953) to include Venezuelan specimens with very large leaves (e.g., 17 cm long, 10 cm wide, Steyermark 5656720 cm long, 11 cm wide, Bernardi 361). However, this condition is scattered among specimens along the Andean Cordillera and without any particular distributional pattern. Clibadium villosum. described by Bentham (1845) from Colombian material, is clearly an excessively tomentose variation of C. surinamense. Lanceolate leaves also occur (C. lanceolatum Rusby) as well 188 as densely strigose leaves (C. lehmannianum O.E. Schulz). This variation is scattered among individuals mainly at higher elevations.

Clibadium surinamense is closely related to C. peruvianum. but the former differs by having 5-veined phyllaries, 10-14 disc florets, 4-lobed disc corollas, and villose achenes. Clibadium surinamense is distributed from Nicaragua to Ecuador, whereas C . peruvianum is restricted to Argentina, Bolivia, and Peru.

Since Schulz (1912) in his monograph of the genus

Clibadium synonymized C. sylvestre with C . surinamense. confusion has existed with the application of the name surinamense and asperum. Personal observations of the type specimens of Baillieria aspera and Baillieria sylvestre at BM confirms (as suggested previously by Blake, 1917) that the names written on the sheets have been transposed, it is clear from the protologues that the labelled Baillieria sylvestris sheet corresponds to the type of B. aspera. which clearly falls biologically within C. surinamense.

23. Clibadium peruvianum Poepp. ex DC., Nov. Gen. Sp. Plant. 3: 1. 1845.--Type: Peru, Amazonas, 1832, Poeppig 35 (holotype: BM!, photos: GH! OS!; isotypes: BM! P,

photo: OS!; G-DC, IDC microfiche 28.925:1.1!). 189

Clibadium remotiflorum 0. E. Schulz, Bot. Jahrb. Syst.

46: 621. 1912.--Type: Bolivia. Cochabamba, 1891, Bang 1203 (lectotype here designated: K!).

Clibadium heterotrichum S. F. Blake, Contr. Gray Herb.

52: 3. 1917.--Type: Bolivia. La Paz: Polo-Polo, near Coroico, Oct-Nov 1912, Buchtien s.n. (holotype: BM! photos: GH! OS!; isotypes: K! MO!, photo: OS!).

Clibadium osilogynum S. F. Blake, J. Wash. Acad. Sci.

21:329-330. 1931.--Type: Peru. Cuzco: Quispicanchi, Marcapata Valley, near Chilechile, 21 Feb 1929, Weberbauer 7 864 (holotype: US!; isotypes: F! GH!).

Clibadium vargasianum H. Robinson, Wrightia 6: 46. 1979.

Type: Peru. Madre de Dios: Manu, Carbon Salvacion, 24 Nov 1965, Vargas 1692 8 (holotype: US!, photos: CAS! F ! G H !).

Shrub 2-3 m tall; stem striate, purplish with green spotting, branches densely strigose with hairs 1 mm long. Leaves with blades ovate-lanceolate to ovate, 8-10 cm long, 3-7 cm wide; at apex acute, at base attenuate (rounded- cuneate), with margin finely irregular serrate, 3-veined, veins yellowish, strigose-pubescent on both surfaces with hairs 0.5-1 mm long; petioles 1-3 cm long, strigose with hairs 1 mm long. Capitulescence paniculiform, terminal, with heads shortly pedicellated. Head subglobose, 5 mm high, 4 mm diameter; phyllaries 3, broadly ovate, 4.5-5 mm long, 3.5-4 mm wide, at apex acute, 7 -veined, with margin upper one-third ciliate, strigose-pubescent on lower side with hairs 0.5 mm long, light-green with fine green dark lines. Ray florets 5, subtended by ovate pales, 4-4.5 mm long, 3.5-4 mm wide, 5 veined, at apex acute, strigose with hairs 0.5 mm, with margin upper half ciliate; corolla white, tubulosa, 3 mm long, l mm diameter, 3-lobed, lobes deltate, 0.5 mm long, at apex scarcely ciliate; styles 2 mm long, branches 1 mm long. Disc florets 19-20, epaleaceous; corolla tubulate, white, 3 mm long, 1 mm diameter, 5-lobed, lobes 0.5 mm long, scarcely ciliate at apex; anthers black, 1.5 mm long; style 1 mm long, undivided, ovaries sterile, 1.5-2 mm long, 1-1.5 mm diameter, glabrous to scarcely pubescent at apex with hairs 0.2-0.5 mm long. Achenes subglobose, black, 2 mm long, 1.5 mm diameter, glabrous, slightly papillose on upper one-third. Chromosome number unknown. Phenology. Flowering throught the entire year.

Distribution (Fig. 18). Argentina, Bolivia and Peru; growing in secondary lowland tropical wet forests, open and grazed selva, riverbanks, and seasonally inundated areas; 100

-600 m (-1200 m) .

Representative Specimens. Argentina. Jujuy: Ledesma, Rio Agua Negra, Logname & Cuezzo 7 534c (LP); Logname & Cuezzo

7 399 (LP). Tucuman: Famailla, Quebrada de Lules, Venturi

1180 (GH; LP); San Rafael, camino al Potrero de las Tablas, Venturi 1180a (LP); Bolivia. Reis Bolivia, Rusby 2145 (NY). 191

Arce: 7.4 km E of Emborozu on road to Bermejo, Solomon 10025

(MO). Murillo: La Paz, Valley of the Rio Zongo, Solomon 7 515 (MO); 45. 5 km below the dam at lago Zongo, Solomon 12945

(MO). Pando: Nicolas Suarez,cerca de Cobija en las riberas del rio Acre, Fdez. Casas & Susanna 8048 (MO). Beni, km 28 carretera de Yucumo a Rurrenabaque, Rio Hondo, Smith & Quintana 12855 (MO). Chapare: Cochabamba, Locotal, Steinbach 9018 (GH; MO); 115 km NE of Cochabamba, near Chimore, Eyerdam 24802 (F); 143 km NE of Cochabamba, north bank of Rio , Eyerdam 24 694 (F) Yungas, Bang 361 (GH, MO; NY); Rio Juntas, Kuntze s.n. (NY; MO); Yapacani, Kuntze s.n. (NY; US). km 9 road Cochabamba-Todos Santos, Piers 564 (LP). Peru. Amazonas: Bagua, ca. 5 km S of La Peca, Barbour 4299 (F), 43 km NE of Chiriaco to Puente Valenzuela, 4386 (F); ridge above Cikan Ece Creek, Berlin 1633 (MO). 13 km SW of Chiriaco on road to Bagua Chica, Gentry et al. 23125 (F, MO). Bagua, along Rio Santiago, 3-5 km above mouth, Wurdack 2224 (F, US).

Ayacucho: Ocarrapa, Killip & Smith 22450 (LP), Aina, between Huanta and Rio Apurimac, 22810 (F), Rio Apurimac Valley, near Kimpitiriki, 22860 (US), 22916 (F, NY, US), 23034 (F, GH, NY,

US). Cuzco: Paucartambo, Atalaya, Vargas 13983 (US); Callanga, Woytkowski 37 6 (US). Huanuco: Pachitea, Honoria, Bosque Nacional de Iparia, Schunke 167 2 (F, OS, US), Rio

Sungarayacu, NW de Puerto Inca, 2959 (F; OS). Junin: ca. 18- 24 km N of San Ramon, Dillon & Turner 1454 (F, MO, OS); La Merced, Killip & Smith 24049 (F, NY, US), Huacapistana, 24180

(F, NY), Colonia Perene, 25068 (F, NY), Puerto Yessup, 26305 (NY), Puerto Bermudez, 26474 (NY). Loreto: Maynas, camino a Santa Maria, Ayala 501 (MO), Rio Itaya, 3104 (F); Iquitos, W edge of the city, Croat 17465 (MO), 5 km Sw of Iquitos, 18142 (MO, OS), along Rio Amazonas S of Iquitos, 19316 (CAS, F, GH, LL, MO, NY), Rio Napo near Varadera de Indiana, 20238 (MO);

Rio Ampiyacu, environs Pucuraquillo, Davis et al. 7 89 (US) ; Maynas, Rio Itaya arriba de San Juan de Muniches, Diaz et a l .: 650 (F; MO, OS); Punchana, near Iquitos, Ferreyra & Corner 3358 (US); Tahuampa near Rio Amazonas between Punchana and Santa Clara de Nanay, Gentry et al. 21604 (F), Rio Gueppi, tributary of Rio Putumayo, 21861 (F), Quebrada Yamamono, Rio Amazonas above mouth of Rio Napo, 27992 (F; MO, OS), Tourist camp on Rio Amazonas between Indiana and mouth of Rio Napo, 29189 (F, MO), Mishana, Rio Nanay, halfway between Iquitos and Santa Maria de Nanay, 28811 (F), Alto Amazonas, Andoas, Rio Pastaza near Ecuador border, 29668 (F,

MO), Rio Itaya, trail behind Puerto Muniches, 31334 (F, MO); Iquitos, Killip & Smith 27 061 (F, NY), 27263 (NY, US); Mishuyacu, near Iquitos, King 87 5 (NY; US); Florida, Rio

Putumayo, at mouth of Rio Zubineta, Klug 2348 (A, F, GH, MO, NY, US); Hendrix 149 (F); McDaniel & Rimachi 17 089 (MO).

Puerto Almendras, MacRae 35 (F); Moore et al.; 142 (F); Rio Ampiyacu, Pebas and vicinity, Plowman et al. 6487 (ECON, F); Iquitos, Maynas, Mishana, Rio Nanay Ramirez 59 (MO); Quistococha, 9 may 1877, Revilla & Froehner 244 6 (MO); Carretera de Zungaro Cocha, cerca de la Quebrada de Shushuna, Rimachi 5992 (MO); carretera de Pena Negra, km 1.5 de Quisco Cocha, Rimachi 7400 (MO); Hacienda San Ramon, 6 km S of Yurimaguas, Straw 2441 (US); Rio Ampiyacu environs 193

Pucuraquillo, Turner et al. 7 89 (F); Lower Rio Nanay, Williams 328 (F, LL), 4££ (LL), 1929 (F), 1998 (US), Alto Rio Itaya, 3344 (F), Lower Rio Huallaga, 4304 (F, GH, US), Alto Rio Huallaga, 6684 (F). Madre de Dios: Tambopata, Puerto Maldonado, Barbour 5386 (F); Rio Alto Madre de Dios between Shintuya and Boca Manu, Gentry et al. 27 259 (F, MO), Pakitsa, entrance to Manu Park, 27359 (OS), road between Shintuya and Salvacion, 27389 (MO); near the confluence of Rio Tambopata and Rio La Torre, 39 km SW of Puerto Maldonado, Smith et a l . 5ia (US), 1698 (US); Tambopata, Rio Piedras, Vargas 18619 (US), Aeropuerto Maldonado, 18788 (US). Pasco: Oxapampa, 2-4 km N of Mallampampa, Smith & Canne 5830 (F). San Martin: San Martin, near Tarapoto on trail to Juan Guerra, Belshaw 3316 (GH, LL, MO, NY); Juanjui, Shapaja, Johns & Ramirez 239 (OS);

Mariscal Caceres, Tocache Nuevo, between Puerto Pizana and Quebrada Salas, Plowman & Schunke 11647 (F). Alto Rio Huallaga, Williams 5655 (F).

Clibadium peruvianum is easily recognized by its leaves with yellowish veins on the abaxial surface, the striate and green spotted stem, the dark green striated phyllaries, and the open panicle. This is an interesting species in which the heads and the capitulescence appears as in C. surinamense but the leaves are like those of C, sylvestfd- Clibadium peruvianum is closely related to C. surinamense. both placed in Section Clibadium. but the former differs by its strigose 7 -veined phyllaries, 5 ray and 20 disc florets, 5-lobed disc corollas, and glabrous achenes vs. 5-veined phyllaries, 3-5 ray and 10-14 disc florets, 4-lobed disc corollas, and villous achenes in C. surinamense. Clibadium surinamense has a broad distribution from Nicaragua to Ecuador, whereas C . peruvianum has a southern distribution restricted to northern Argentina, Bolivia, and Peru.

Clibadium psilogynum was described by Blake (1931), based only on type material collected from the Cuzco region in Peru. These three type specimens are one extreme of morphological variation observed in C . peruvianum involving larger leaves (15 cm), irregularly serrate margins, fewer disc florets, and sometimes 1 or 2 disc florets with a small pale.

Clibadium heterotrichum was described as distinct from C . peruvianum by Blake (1937), based on the harsh spreading pubescence of the stems and larger heads (6 mm) of the former. Examination of more ample material, however, indicates that this pubescence and floral variation grades into C . peruvianum and vitiates specific recognition.

24. Clibadium arboreum Donn. Sm., Bot. Gaz 14: 26. 1889.--Type: Guatemala. Alta Verapaz: Pansamala, Jun 1886, von Turckheim 929 (holotype: US!; isotype: GH!

K! NY [2]! US!; photos: OS! US!). 195

Clibadium donnell-smithii J. M. Coult., Bot. Gaz. 16: 98. 1891.--Type: Guatemala. Guatemala, Feb 1890, Smith 2347 (holotype: US!; isotype: K!, photos: OS!

US! ) .

Clibadium pueblanum S. F. Blake, Contr. U.S. Natl. Herb.

22: 601. 1924.--Type: Mexico. Puebla: Huauchinango, Pahuatlan, 4 Oct 1914, Salazar s.n. (= 1038790 US accession number] (holotype: US!; isotype: GH!).

Clibadium oligandrum S. F. Blake, Brittonia 2: 342.

1937.--Type: Guatemala. Quezaltenango: Volcan Zunil, 3 Aug 1934, Skutch 927 (holotype: A!; isotype: BM!, photos: F ! GH! N Y ! OS! US!).

Shrubs or small trees up to 6 m tall; stems densely branched, with branches densely pilose, with hairs 1 mm long. Leaves with blades ovate to broadly ovate, 10-20 cm long, 8- 18 cm wide, at apex acuminate or long acuminate, at base more or less rounded and then cuneate, triplinerved far above the base, with margin serrate; upper surface scabrous, lower surface densely short pilose with hairs 0.5 mm long. Capitulescence paniculiform, with small or large cluster of heads leafly at base. Heads subglobose, sessile or shortly pedicellated, 5 mm high, 3.5 mm diameter; phyllaries 3 (-4), broadly ovate, striate, 5 mm long, 4-5 mm wide, 5-7 veined, at apex acute or slightly obtuse to obtusely pointed, strigose with hairs 1 mm long, with margin upper one-third 196 ciliate. Ray florets 3-6, subtended by broadly ovate pales, 4.5 mm long, 4 mm wide, 5 -veined, strigose with hairs 0.5 mm long, at apex obtuse, scarcely ciliate; corollas white, 2.5 mm long, l mm diameter, 4 lobed, lobes 0.2 mm long, at apex scarcely pubescent; styles 2 mm long, branches 1.5 mm long, tips recurved. Disc florets 4-11, epaleaceous; corollas white, 2-3 mm long, tube 1 mm long, expanded into a campanulate throat, 0.5 mm diameter, 5-lobed, lobes deltate, 1 mm long, at apex scarcely hirsute; ovaries sterile, 1.5 mm long, 0.5 mm diameter, hirsute at upper one-third with hairs 0.5 mm long. Achenes blackish, obovoid, 2 mm long, 1.5 mm diameter, hirsute at apex with hairs 0.5 mm long. Chromosome number, n = 16. Phenology. Flowering from April through June (October- November).

Distribution (Fig. 19). Belize, Guatemala, Honduras and Mexico; toward Pacific slopes, humid dense forest in stream- valleys associated with Carpinus. Magnolia. Podocarpus. and Ouercus. damp or wet secondary forest along roadside; sea level to 2,500 m.

Representative Specimens. B e l i z e . Belize: 1.5 mi S of mile 22 on Western Highway, Croat 23863 (OS). Cayo: along

Hummingbird at mile 28, Dwyer 11415 (OS); Rio Frio near Augustine, McDaniel 14482 (F). Hummingbird Highway, E of Over-the-Top Camp, Whitefoord 2421 (F). Cortez: Montana

Santa Ana along Rio , Molina 3 601 (GH). O l a n c h o : Izabal, 6-8 km S of Modesto Mendez, Harmon 2552 (GH). 197

'20

30

Fig. 19. Distribution of Clibadium arboreum ( • ) and C. armanii (*) in Belize, Guatemala, Honduras, and Mexico, and in Brasil and Paraguay, respectively. Toledo: Columbia Forest Reserve, ca. 1-2 mi N of entrance, Croat 24155 (OS); vicinity of San Jose Myan Indian Village, 6.7 mi N of Columbia Forest Station, Croat 24409 (OS). Guatemala. Alta Verapaz: ca. 6 km NE of Panzos, Croat 4157 9 (CAS, MO); city limits of Coban, King & Renner 7106 (CAS, MO); ca. 5 km NE of San Pedro Carcha, King & Renner 7122 (CAS, MO); along Rio Frio between Tactic and Santa Cruz, Molina & Molina 12260 (F); between Limon and Chisec, Steyermark 45106 (F); Cubilquitz, Turckheim 7 903 (F, GH, MO, NY, US). Chimaltenango: Quisache, Standley 62042 (US); Escuintla: Finca Monterrey, S slope of Volcan de Fuego, Standley 64559 (F, GH). Guatemala: Izabal: Puerto Mendez, on Toquela road, Contreras 9951 (CAS), Donnell-Smith 2347 (GH); road in pineapple plantations from Highway ca. 14 mi S of Puerto Barrios.Croat 4177 6 (CAS); 6-8 km S of Modesto Mendez, Harmon 2553 (MO); along Rio Bonita, Steyermark 4167 0 (F); between Puerto Mendez and Cienaga, Tun Ortiz 2468 (F, US).

Peten: La Libertad and vicinity, Aguilar 322 (MO, NY) ; Dolores, in pineland, 2.5 km E, Contreras 287 3 (F), San Pedro, on Cadenas Road, 7026 (CAS, F) ; 32 km S of Poptun, Harmon 282 6 (NY, US); La Cumbre, on Pusila road, 2 km,

Lundell & Contreras 20044 (F); Ceibal, Molina 15860 (NY, US); , on shore of Lago Petdn-Itza, N of Flores, Ratcliff 10 (CAS); 20 km N of Santa Ana, Tim Ortiz 1359 (F, MO, NY); Santa Elena, en camino a La Libertad, 14 km SW, Tun

Ortiz 1996 (CAS, F ) . Quezaltenango: S of San Martin Chile Verde on road to Colomba, Standley 85097 (F); between Finca Pirineos and Patzulin, Standley 867 86 (F). Quiche: Finca Chaita, Zona Reyna, Skutch 17 97 (GH); Retalhuleu: ca. 27 km W of Mazatenango, King & Renner 7 020 (CAS, MO); San Marcos: Finca El Porvenir along Rio Cabus above Potrero Matasan, Steyermark 37 622 (F); 8-10 km W of San Marcos, slopes of Tajumulco Volcano, Williams et al. 26903 (F, NY, US). Solola: Volcan Atitlan, Steyermark 47 342 (F). Suchitepequez:

Volcan Santa Clara, between Finca El Naranjo and upper slopes, Steyermark 46626 (F, GH). Honduras. Cortes: along Rio Santa Ana, Molina 3601 (F, US); Aldea La Pita, 5 km SO of Puerto Cortes, Nelson et al. 3109 (MO); Agua Azul, Williams & Molina 1447 4 (F, GH, US); Santa Barbara: 24 km E of Santa

Rita de Copan, Harmon & Dryer 4 031 (MO, NY). Mexico. Chiapas: Municipio of Jitotol, 7 mi N of Jitotol, Breedlove 15423 (F, LL, NY); Municipio of Solosuchiapa, 3-5 km above Solosuchiapa along road to Tapilula, Breedlove 26410 (MO);

Municipio de Palenque 6-12 km S of Palenque on road to Ocosingo, Breedlove 26504 (MO); Municipio of Motozintla de Mendoza, near Niquivil, Breedlove 42714 (MO); El Chinchon,

Burnham & Spicer 114 (MO); 23 mi SE of Palenque from Palenque to Chancala, Croat 40283 (MO); Monte Tacana, Matuda 2422 (F, NY); road 195 between Solosuchiapa and Jitotol, Maxwell 217 (MO); 4 mi N of Jitotol on road to Pueblo Nuevo Solistahuacan, Raven & Breedlove 19974 (F, LL); Municipio of San Pedro Chenalho near the Colonia Choro, Shilom Ton 2496 (NY); Municipio of Ocozocoautla de Espinosa SW of the Presa de Malpaso, Shilom Ton 3311 (F, NY). Jalisco: 11-12 mi S of

Talpa de Allende, McVaugh 21414 (LL, NY). Mexico: Ejido Lazaro Cardenas, 100 m del puente del ferrocarril, Cowan 2427 (MO). Oaxaca: 4.5 mi S of Valle Nacional, Croat 39712 (MO); S of Valle Nacional and about 109 road miles NNE of Oaxaca.Cronguist 9631 (GH, NY); Tuxtepec, Chiltepec, Martinez Calderon 128 (MO); about 29.7 mi W of Tuxtepec along road to Ixcatlan, Stevens 1390a (MO). Tehuantepec, 27.8 km al NW de La Chiviza, carretera a Santiago Lachiguiri, Torres & Martinez 5917 (F). Puebla: 4 km NE of Villa Juarez Marcks &

Marcks 826 (WIS); Barranca, Teoxolo near Honey Station, Pringle 10802 (LL); below Teziutlan near Ocostoc, Sharp 45808

(MO). Veracruz: Estacion Biologica de los Tuxtlas, Calzada 500 (F, MO). Yecuatla, 3 km adelante de Santa Rita, Sierra de Chiconquiaco, Calzada 4716 (F). San Andres Tuxtla, Cima del

Vigia, Estacion de Biologia Tropical Los Tuxtlas, Cedillo 288 (F); road from Coyame to Tabanca, Cruz 286 (LL); NW of

Santiago Tuxtla, Dressier & 50 (GH, MO, NY); Cascada de Texolo, Fay & Calzada 7 89 (F) ; Highway 131 between Huatusco and Coscomatepec, Gibson & Gibson 2512 (NY); 1 km from Los Mangos on road to Juan Diaz Covarrubias, Gomez-Pompa 4517 (GH); Las Choapas, Gomez-Pompa & Nevling 1546 (GH) ;

Coscomatepec, 5 km NE of Coscomatepec, 1 Ion W of turnoff to La Candelaria, Hansen & Nee 7 586 (F);. Jalapa, MacDaniels 860 (F); San Andres Tuxtla, MatInez-Calderon 17 25 (F, MO); Xico, gorge at Puente Acabaloya, ca. l km SE of Xico Viejo Nee & Taylor 26315 (F); Atzalan, between Puente de Rieles and La Calavera, 9.5 km NE of Altotonga on road to Tlapacoyan, Nee & Didds 24914 (F); Soteapan, along road 13 km E of Tebanca, Nee & Hansen 187 92 (F); Barranca of Teoxolo near Jalapa, Pringle 10802 (F, GH, LL); roadsides at Zacuapan, 201

Purpus 37 05 (F, GH); Cerro de Cuahutlapan E of Orizaba, Rosas 836 (F) ; near field station of UNAM between Sontecomapan and Montepio, Stevens 1104 (MO).

Clibadium arboreum is characterized by its leaves, petioles, and larger foliar veins being densely hirsute with soft, yellowish hairs. It can also be distinguished by its broadly ovate leaves with a finely serrate margin, rounded base, and acuminate apex. The leafly paniculiform capitulescence is mostly wider than high and most of the young branches irregularly exceed the central (terminal) branch.

Clibadium arboreum in Belize, Guatemala, and Mexico is the most northern species of the genus. Clibadium arboreum is closely related to C. surinamense and C . peruvianum forming a relatively homogeneous group (Sect. Clibadium) characterized by 3-4 strigose ovate phyllaries with acute apices, 3-4 ray florets subtended by pales, 4-20 disc florets without pales, and 5-lobed disc corollas (except 4-lobed in C . surinamense).

25. Clibadium armanii (Balb.) Sch. Bip., Linnaea 30:180. 1859. Eupatorium armani Balb., Hort. Taurin. Stirp. 1. 27. £.£. 1810.--Type: Brasil, locality and date unknown, Armani s.n. (holotype: TO?). A letter, fax, and telephone inquiry to TO yielded no information regarding 202

type material. The plate in the protologue, however, is diagnostic for this taxon.

Orsinia euoatoria DC., Prodr. 5. 104. 1836. Clibadium rotundifolium DC., Prodr. 5:505. 1836. Type: Brasil. Bahia: Dec 1832, Blanchet 632 ((lectotype here chosen: NY!). Former syntypes (= lectoparatypes fide Hansen & Seberg, 1984): Blanchet 360 (F!, photo: OS!), Blanchet s.n. (P, photo: OS!), Blanchet 722 (G-DC, IDC 28.924:111.5!), Gaudichaud 466 (P, photo: OS!), Salzman s.n. (P, photo: OS!), Salzman. 1830 (G-DC, IDC

28.924:111.6!) .

Shrub 3-4 m tall;.stems with branches terete, scabridly hispid, with hairs 1 mm long. Leaves with blades broadly ovate, edges recurved. 4-7 cm long, 3-5 cm wide, subcoriaceous, at apex obtuse and mucronate at tip, at base rounded or shortly attenuate, with margin mucronate dentate, pinni-nerved, 5-7-nerved at the base, pale green on both surfaces, scabridly hispid with hairs 1 mm long; petioles 3- 10 mm long, scabridly hispid with hairs 1 mm long.

Capitulescence thyrsoid. Heads ovoid, sessile, 5-6 mm high, 4 mm diameter; phyllaries 3, coriaceous, concave, broadly ovate, 4.5 mm long, 3.5-4 mm wide; 5-veined; whitish green, at apex shortly acuminate to mucronate; strigose with hairs 1 mm long. Ray florets 4, subtended by broadly ovate pales, 4 mm long, 3.5 mm wide; 5-veined; at apex shortly acuminate; strigose with hairs 0.5 mm long; corolla white, tubular, 3.5 203 mm long, 1.5 mm diameter, 3-lobed, lobes irregularly deltate,

0.3 mm long, at apex scarcely ciliate. Disc florets 5-8, epaleaceous;, corollas white, tubular, 4.5 mm long, 2 mm diameter, 5-lobed, lobes deltate, 0.8-1 mm long, at apex scarcely ciliate; ovaries sterile, 2 mm long, 1-1.5 mm diameter, hirsute with hairs 1 mm long. Achene broadly obovoid, 3 mm long, 2-2.5 mm diameter, at apex hirsute with hairs 1 mm long. Chromosome number, n = 16. Phenology. Flowering from November through March.

Distribution (Fig. 19). Brasil and Paraguay; gallery forest, disturbed areas, on sandy soil or at edges of wet savanna; 500-1600 m.

Representative Specimens. Brasil. Bahia: Municipio de Ilheus, Rodovia Uruguca, Belem & Aguiar 1289 (NY); Municipio de Santa Cruz de Cabralia, Brito & da Vinha 12 (US); Chapadao Occidental de Bahia, 12 km N of Correntina, Harlev et al. 2187 4 (US); Municipio de Itabuna, King & Mori 8000 (MO, US);

Municipio de Ilheus, 4 km near Distrito Industrial, King & Silva 8008 (MO, US); Municipio de Ilheus, 2 km S of Olivenga, Mori et al. 13643 (US), 20 km S of Olivenga, Mori et al.

13719 (US); between Itabuna and Itajuba, Pereira & Pabst 8443 (F, LP); Santa Cruz de Cabralia Reserva Biologica, Santos 1938 (US); 2 km before da Vila do Rio do Engenho, Capoeira Alta, Silva & Sobral 2168 (US). Brasilia: Taguatinga Norte, Chagas & Silva 252 (CAS); area do Cristo Redentor, da Silva &

Alvarenga 936 (US); Bacia do Rio Sao Bartolomeu, Heringer et al. 2784 (US); Reserva Ecologica do IBGE, Heringer et al. 7560 (MO); Chapada da Cotagem, ca. 20 km NE of Brasilia, Irwin et al. 9642 (GH); E of Lake Paranoa, Irwin et al. 11209 (NY); Sobradinho, D.F., Irwin et al. 11381 (GH, NY, US); Cachoeira Piripiripau ca. 15 km S of Planaltina, Irwin et al.

26443 (NY, UC, US); Sobradinho, beira d'agua, Pereira 9064 (LP); 32 km N bridge at Asa Norte on road to Alto Paraiso,

King & Almeda 8205 (CAS, US); Pereira & Pabst 4950 (F). Distrito Federal: Cachoeira Piripiripau, ca. 15 km S of Planaltina, Irwin et al. 26443 (F). Goias: Chapada dos Veadeiros ca. 29 km N of Alto Paraiso, Anderson 67 39 (F, MO, NY, US); Serra do Caiapo ca. 12 km S of Caiaponia, Anderson 9603 (F, MO, NY, US); 5 km W of Cristalina, Hatschbach 43850

(US); Municipio de Jatai, Hatschbach & Kummrow 34993 (UC) ; ca. 15 km W of Veadeiros, Irwin et al. 12674 (US, NY); Serra dos Pirineus 20 km N of Corumba de Goias, Irwin et al. 187 43 (DS); 58 km NE of Catalao, Irwin et al. 25432 (F, NY); Chapada dos Veadeiros, 21 km N of Alto Paraiso, King & Almeda 8286 (MO, US); 5 km NE to 5 km NW of Cristallina, Kina & Bishop 8966 (US). Mato Grosso do Sul: Campo Grande, Cabrera & Zardini 29986 (OS); about 12 km SW of base camp, Harley et al. 11249 (NY, MO, UC); 7 km S of Xavantina on Aragargas road, Harley & Souza 11081 (NY, MO, UC); Municipio Rio Brilhante Fazenda Bela Vista, Hatschbach 26091 (UC). Curitiba, Rio Pandai, Hatschbach 4617 4 (F). Serra do Roncador, 86 km N of Xavantina, Irwin et al. 16352 (NY, UC, US); Contraforte Central ca. 58 km NE of Catalao, Irwin et al. 25432 (GH, US); Campo Grande, University Park, Kraoovickas & Cristobal 34 351 (MO); ca. 1 km from km 259 205

Xavantina-Cachimbo road, Philcox et al. 3292 (NY). Minas

Gerais: entre Queimado and Rio Douro, Glaziou 9484 (NY); Brejo ca. 35 km W of Montes Claros road to Agua Boa, Irwin et al. 23772 (F, GH, MO, NY, US); Morro das Pedras ca. 40 km NE of Patrocinio, Irwin et al. 25681 (MO, NY); ca. 3 km S of Ouro Preto, Irwin et al. 29507 (F, NY, US); Serra do Cipo at

Chapeu de Sol, King & Bishop 8437 (US); Pirapitinga, Macedo

1771 (MO); Municipio de Belo Horizonte, Brejo, Magalhaes 3183 (US); Passo Quatro, Itanbamtu, Nunes 116 (US); Widgren 166 (GH); Municipio of Betim, Fazenda do Cabui, Williams 5119 (GH), Fazenda do Cabui, near Contagem, 5122 (LP), Municipio de Belo Horizonte, Bento Pires, 5198 (GH). Parana: Ibarare Dusen 16424 (F, NY) ; Municipio de Ibaiti, Guai, Hatschbach 25957 (UC); Municipio de Senges, Fazenda Morungava Rio Funil, Smith et al. 14 842 (NY, US). Rio de Janeiro: Corva Grande, Itaguai, Duarte 4464 (US); Fazenda Santa Cruz, Glaziou 94 89 (US); Itaguai, Fazenda Patioba, Monteiro 3100 (US); Valley of the Rio Campo Bello, Mt. Itatiaya, Smith 1601 (F, GH, US).

Sao Paulo: Parque do Estado, Coleman 265 (US); 6 km N of city of Paraguagu Paulista, Eiten & Eiten 5880 (MO, US); Municipio de Itapetininga, Estacao Experimental de Itapatininga, Instituto Forestal, Giannotti 4567 (F); Horto Botanico Cantareira, Puttemans 209 (LP); between Caraguatatuba and Ubatuba, Smith & McWilliams 15415 (US); Jaragua, Usteri 16711 (NY); Piracicaba, Viegas 3803 (CAS). Paraguay. Alto Parana: Fiebrig 5815 (GH, US). Amambay: Sierra de Amambay, Roias 10053 (A, F); Parque Nacional Cerro Cora, Zardini et al. 4049 (US); Zardini et al. 4185 (US). San Pedro: Sierra de 206

Maracayu, ca. Igatimi, Delessert 5625 (F); Colonia Guayaibi, 20 km N of San Estanislao, Krapovickas et al. 14274 (UC).

Clibadium armanii has characteristic pale-green scabrid rigid leaves, whitish-green phyllaries, and stems and petioles tinged slightly red. Leaves are shortly petiolate (3-10 mm long), broadly obovate, at base more or less attenuate, rounded or sometimes cordate, and regularly serrate with minute mucros. The midrib and secondary veins are conspicuously yellowishly marked. This species is closely related to C . arboreum and C. surinamense. Clibadium armanii differs from its close relatives by having leaves subcoriaceous (mucronate at apex), 5-veined shortly acuminate phyllaries, and 5-8 disc florets with corollas hispidulous at the apex of the lobes.

Although C. armanii is clearly distinguishable by the characters mentioned above, the taxon shows variation along its distributional range. Some specimens have attenuate leaf bases (e.g., Eiten & Eiten 5880). whereas others show a more rounded to cordate base (e.g., Krapovickas & Cristobal 34351; hence the specific epithet rotundifolium). Some specimens show very small leaves (2 cm long, 1.5 cm wide) with a typical deltoid shape (Kina & Almada 8205); others, lanceolate leaves (Irwin et al. 12394); still others with large (7 cm long, 6 cm wide) basal leaves with attenuate bases and small leaves along the peduncles (Harley et al. 11249). There is also variation with regard to the typical 207 mucronate dentate margin: it is irregularly serrate to slightly erose in Eiten & Eiten 5880.

IID. Clibadium L. section Oswalda (Cass.) DC., Prodr. 5:505. 1836.

Oswalda Cass., in Diet. Sci. Nat. 59. 322. 1829. --Type: Clibadium sylvestre (Aubl.) Baill.

Leaves ovate-lanceolate; heads arranged in thyrsoid capitulescences; heads paleaceous; phyllaries 7-11-veined; ray florets 4-14, with corollas 3-lobed, and pales 7 -veined; disc florets 5-22, ray corollas 3, 4, or 5-lobed, with pales 3-veined. Species 26-29.

26. Clibadium leiocarpum Steetz, in Seem. Bot. Voy. Herald 152. 1853.--Type: Panama. Veraguas: volcano of Chiriqui, 7 000 ft, Feb 1849, Seemann 1591 (holotype: K!

photos: F! GH! OS! US!; isotype: BM! photos: F! OS! US! ) .

Clibadium schultzii S. F. Blake, Contr. U. S. Nat. Herb. 22:602. 1924.--Type: Costa Rica. San Jose: thickets at Copey, 1800 m, Mar 1898, Tonduz 11915 (holotype: U S !, isotypes: G H ! US!).

Clibadium leiocarpum Steetz var. strigosum S. F. Blake, Jour. Wash. Acad. Sci. 27: 382. 1937.--Type: Costa 208

Rica. San Jose: Cerro de Piedra Blanca, above Escazu, 31 Jan 1924, Standlev 32593 (holotype: US!).

Shrub 2-5 m tall; stems usually densely branched, striate-angulate, 4-10 cm at base, strigose to densely pilose tomentose with hairs to 1 mm long. Leaves with blades lanceolate to ovate, 5-20 cm long, 2-10 cm wide, at apex acuminate, at base narrowly atenuate, with margin serrate; both surfaces hirsute-tomentose to strigose with hairs to 1 mm long; petioles 2-6 cm long, 1-3 mm diameter, tomentose to tomentose-lanate with hairs 1 mm long. Capitulescence paniculiform, with up to 100 heads; peduncles 0.5-1.0 mm long. Heads radiate, 5 mm high, 3-3.5 mm diameter; phyllaries 3, suborbicular to broadly ovate, 3.5-4.5 mm long,

3-4 mm wide, 8-9 veined, abaxially glabrous, at apex acute, with margin ciliate. Ray florets 7-9, subtended by ovate pales, 3.5-4 mm long, 3-3.5 mm wide, glabrous, scarcely ciliate at apex, 7 -veined; corollas white, 1.5 mm long, 0.5 mm diameter, glabrous, 3-lobed, lobes 0.4 mm long; styles 2 mm long, branches 0.5-0.8 mm long. Disc florets 10-15, subtended by lanceolate pales (ocassionally 1-2 central disc florets epaleaceous), 2.5 mm long, 1-1.5 mm wide, 3-veined; corollas white, 2.5-3 mm long, throat 2 mm long, 1 mm diameter, lobes 0.5 mm long, at apex strigillose with hairs 0.3 mm long; anthers black, 1.5 mm long; styles undivided; ovaries sterile, 2 mm long, 1 mm diameter, glabrous. Achenes 1.5-1.7 mm long, 1 mm diameter, glabrous or sparcely pilose 209 at apex. Chromosome number, n = 16.

Phenology. Flowering from December to September. Distribution (Fig. 20). Costa Rica, Nicaragua and Panama; secondary wet forest, common along roads and disturbed slopes; 1400-2800 m.

Representative Specimens. Costa Rica. Alajuela: between and , few km W Atenas toward Cerro del Aguacate, Arriagada 405 (OS); Cordillera Central S slope VolCcin Poas near Poacito, Arriagada 407 (OS) ; W slope of Volcan Barva on route to Concordia, N of , Arriagada 409 (OS); Cordillera Central, 7 mi N of Carrizal, Croat 35500 (UC); Monteverde Reserve Atlantic slope, Rio Penas Blancas, Haber 835 (UC); Cerro Azahar, San Ramon, Gomez-Laurito 937 8 (CR, F); ca 9 mi N of Zarcero on route 15,

Stuessy & Gardner 4462 (OS); Volcan Poas Park , just S of main entrance, Stuessy & Gardner 447 9 (OS); Volcan Poas along upper rim, Turner & Turner 15042 (CR); slopes Volcan Poas from rim of crater 2 mi down road to , Wilbur & Teeri 13669 (CAS); near River, Cordillera Central, 10 km N of Zarcero, Williams et al. 28923 (F). Cartago: San Francisco 10 km after Paraiso on route to Turrialba, Arriagada 146 (OS); Molina, 3.5 km NE Tres Rios, between San Jose and Cartago, Arriagada 400 (OS); Tapanti Watershed Preserve ca. 20 miles SW of Paraiso, Croat 47 008 (UC); Carretera 2 E of San Rafael, Parque Pasador El Cristo Rey, Khan et al. 587 (CR); S slopes Volcan Irazu, ca. 4 km N Pocayas, King 6831 (MO, UC, US); 7.6 mi S of Cartago on route 210

Fig. 20. Distribution of Clibadium leiocarpum (T ) and C. sylyestre ( • ) in the Caribbean, Central America, and northern South America. 2, Stuessv & Gardner 4494 (OS); Tapanti Hydroelectric project, 4-10 km beyond entrance to project, Utlev & Utley 5158 (F). Hillside 2/3 of the way from Cot to , Weston 4479 (OS, UC); near La Sierra 25 km S of Cartago, Cordillera de Talamanca, Williams et al. 28141 (US). Heredia: from Pueblo Barva to San Miguel on , km 23, Arriagada 162

(OS); S slope Volcan Barva, 1 km N. of Porrosanti, toward W from Rio Ciruelas, Arriagada 411 (OS); Barva de Heredia on route to Volcan Barva, Arriagada 308 (OS); Vara Blanca on route to Volcan Poas along coffee plantation, Arriagada 309 (OS); Rio Vueltas, E slope of Volcan Barba near the Continental Divide, Barringer & Christenson 3365 (F); Alto del Roble a la vera Rio Patria, Gomez-Laurito 8599 (CR); along route 113, ca 2 km SW of Alto Gallito, King 6805 (US); between San Rafael and Rio Las Vueltas N slope of Cerro Chompipe, Stevens 13986 (F); up to 1 Ion from church and school, Vara Blanca, Weston & Weston 3420 (UC); near road 0.5-2 km above Hacienda Montecristo, San Rafael-Monte de la

Cruz, Weston et al. 5624 (UC); wooded slopes Volcan Barva near village of Sacramento, Wilbur & Teeri 137 01 (CAS, CR, F,

GH, OS). Puntarenas: Cotonsito along road to Sitio Cordillera de Talamanca, Davidse 24607 (CR); Monteverde, Monteverde Cloud Forest Reserve, roadside of road paralleling the Sendero Nuboso, Feisinaer et al. 82-218-2 (BM); 6 km S of San Vito de Java at Finca Las Cruces, Gillis & Gillis 10162 (GH, OS); Monteverde, Haber 1189 (UC); near entrance to

Monteverde, Turner & Turner 15024A (CR, F ) . San Jose: W slope Volcan Irazu, Finca Alto Pizote, Arriagada 415 (OS); Cerro de la Muerte, Carretera interamericana, between Hortensia and Division, 23 km N of San Isidro, Arriagada 417 (OS); along , ca 16 km W of Santa Ana, King 6746 (MO, UC, US); between Aserri and Tabarca, Standley 34088 (US); along Rio Blanco NE of El Copey, Standley 41894 (US); near , Weston 3001 (UC); 105 km point Pan-American Highway, below Cerro de la Muerte, Weston 4714 (UC);

Cordillera de Talamanca 25 km N of San Isidro del General along Pan American Highway, Williams et al. 28560 (F). Nicaragua. Granada: volcan , Plan de Las Flores, Aracruistain & Moreno 655 (F) , 1157 (OS) ; Griialva et al. 2109

(OS); Griialva 2509 (OS); Miller et a l . 1422 (F); Moreno & Henrich 9165 (OS); Neill 2920 (OS). Panama. Chiriqui: vicinity of El Valle, lower Rio Anton, Allen 118 (MO); vicinity Boquete, lumber road into the hills E of Rio Caldera, Allen 4653 (M); 8 km W of Cerro Punta, vicinity of Las Nubes, Almeda & Nakai 3515 (CAS); Burica Peninsula, San Bartolo Limite, 20 km W of Puerto Armuelles, Busey 534 (MO,

UC); aprox 2 km en el camino Cerro de la Muerte proximo a campamento de la Universidad Estatal de Florida, Correa s.n. (MO); SW del Campamento Fortuna, desde Finca Pitti hasta el filo del Cerro Punta, Correa et al. 27 81 (F). Croat 13518 (MO); in road to Cerro Colorado and Escopeta, above Rio San Felix bridge, Croat 334 04 (MO); roadside above Cerro Punta, D'Arcy & D'Arcy 6537 (MO, UC); Poblado de Fortuna above Gualaca, D'Arcy et al. 15860 (UC); Hill E of Audubon Cabin, S of Cerro Punta, Hamilton & Krager 3857 (UC); Alto Los Guerra road W of Bambito, Hamilton & Stockwell 3656 (UC); Nueva Suiza road to Cerro Punta, Hamilton et al. 715 (UC); road from Alto Quiel to cerro Punta, ca 2-3 air mi due W of Cerro Horqueta, Hartman 3917 (OS); 17 km E of Rio Sereno on road from California, Hartman 3920 (OS); slopes of ridges to the S of Cerro Pelota, Knapp 1532 (F, MO); 4.1 mi from Boquete on road to David, Kirkbride .94 (F) ; Vicinity of Boquete, from

Boquete to 3 mi N, Lewis et al. 596 (MO, UC); Pastures around El Boquete, Pittier 3136 (US); Volcan Baru, Schmalzel et al. 1571 (UC); Valley of the upper Rio Chiriqui Viejo, vicinity of Monte, Seibert 245 (NY); slopes of Volcan Baru near town of Cerro Punta, Stern & Chambers 80 (MO); E of Vicinity of Boquete.Stern et al. 1066 (UC); 3.7 km E of bridge NE of cerro Punta on road to Bajo Grande, Stevens 18186 (UC); halfway between Cerro Punta and Bambito, Wilbur et al. 10884 (CAS, F); about 2 km W of La Garita, 3 km WNW of Cerro Punta, Wilbur et al. 15271 (F); Vicinity of Casita Alta, Volcan de Chiriqui, Woodson et al. 908 (NY).

Clibadium leiocarpum is distinguished from other species of the genus by its typically glabrate phyllaries (rarely strigillose or ciliate at apex), black and glabrous achenes, and densely strigose peduncles and axes of capitulescences. The density of pubescence varies among populations, to the extent that Clibadium schulzii was described as a distinctive taxon due to more copious pubescence. Also, variety strigosum was based on specimens (Standley 23593) having petioles and blades strigose and achenes (sometimes) sparsely villose at apex. In my opinion, this taxon does not warrant 214 formal recognition because the above-mentioned pubescence type is broadly distributed among populations without any geographical pattern (e.g., Costa Rica, San Isidro, Arriagada 160; San Jose, Arriagada 415: Cartago, King 6831: Alajuela, Stuessv & Gardner 4456: Heredia, Weston et al. 5624).

Clibadium leiocaroum has C. sylvestre as its closest relative, but the latter differs by strigose phyllaries, with obtuse apices, which are white-green when dried in contrast to brown-green in C. leiocarpum. a typically thyrsoid capitulescence (in contrast to paniculiform in C. sylvestre) and also by the presence of an unusual rudimentary pappus of two short awns. Moreover, Clibadium leiocarpum has a broader phenology (see Fig. 7) than C. sylvestre. and it is adapted to higher elevations (1400-2800 m in the former and 0-600 m in the latter). Clibadium leiocarpum also lacks acetylenic compounds isolated from C. sylvestre which are responsible for fish-poisoning (see Ethnobotany section).

27. Clibadium sylvestre (Aubl.) Baill., Hist. PI. 8: 307. 1886. Baillieria sylvestris Aubl., Hist. Pi. Guianae 2: 807. 1775.--Type: Guyana, 1762-64 [Urban, 1903], Aublet

s.n. (holotype: BM! , photos: GH! OS!).

Clibadium havanense DC., Prodr. 5: 506. 1836.--Type: Cuba. La Havana, 1826, de la Ossa s.n. (holotype: G-DC, IDC microfiche 28.925:1.8!; photos: F! GH!). 215

Clibadium vargasii DC., Prodr. 5: 506. 1836.--Type: Venezuela. Caracas, 1829, Vargas 13 (holotype: G-DC, IDC microfiche 28.925:1.7!; photo: A!).

Clibadium barbasco (H.B.K.) DC., Prodr. 5: 507. 1836. Bailleria barbasco H.B.K., Nov. gen. sp. (qt) 4: 289. 1818.--Type: Venezuela. Misiones del Orinoco, "in umbrosis Javitae," 30 May-7 May 1800 [Sandwith, 1968], Humboldt & s.n. (holotype: P, photos: F!

GH! OS!; isotype: P, photo:OS!).

Clibadium badieri Griseb., Flor. Brit. W. Ind. 368. 1864.--Type: Dominica, 1832 [Hedge & Lamond, 1970],

Imray 145 (holotype: K!, photos: A! OS!).

Clibadium strigillosum S. F. Blake, Contr.Gray Herb. 52: 4. 1917.--Type: Peru. Huaca: Rio Huallaja, 1835,

Mathews 13 60 (lectotype here designated: BM!, fragment at GH! photos: GH! OS!; isolectotypes: K[2]! photo: OS!, P, photos: F! GH! MO! OS!).

Clibadium latifolium Rusby, Descr, S. Amer. Pi. 150. 1920.--Type: Colombia. Magdalena: "in open places in alluvial forest on banks of river Buritaca, 2 miles from the sea," 28 Sep 1898, Smith 2014 (holotype:

NY!, fragment at US!, photos: US [3]!; isotypes: F! photo: US!, K ! photo: OS!, P photo: OS!, US!). 216

Clibadium appressipilum S. F. Blake, Contr. U.S. Natl. Herb. 22: 600. 1924.--Type: Panama. Darien: Boca de

Cupe, 13 Apr 1908, Williams 698 (holotype: US!; isotype: NY! photo: OS!).

S. F. Blake, Contr. U.S. Natl. Herb. 22: 600. 1924.--Type: Panama. Canal Zone: alluvial bottom near Bohio, 12 Feb 1912, Maxton 47 67 (holotype:

U S !) .

Shrubs 2-4 m tall; stems strigillose, with hairs 0.1 mm long. Leaves with blades ovate to ovate-lanceolate; 8-20 cm long, 4-12 cm wide, at apex acuminate to caudate, at b ase attenuate at base, with margin serrate, pauci-serrate, dentate, crenate-serrate; both surfaces strigillose (usually more abundant along main veins) with hairs 1 mm long; petioles 1-5 cm long, to 2 mm diameter, strigillose with hairs 1 mm long. Capitulescence thyrsoid, with about 100 heads at end of leafy twigs; peduncles 1-2 mm long. Heads radiate, 4-5 mm high, 3-4 mm diameter; involucre cupulate; phyllaries 3(-4), ovate to obovate, 3-4 mm long, 2-3 mm wide, abaxially strigose, with hairs 0.5 mm long, 7-11-veined. Ray florets 5-9, subtended by ovate pales, 3-4 mm long, 2-3 mm wide, 5-7-veined, abaxially shortly strigose, with margin upper one-third shortly ciliate; corollas 1.5-2 mm long, 0.3- 4 mm diameter, 3-lobed, 0.4 mm long, styles 2-2.5 mm long, branches l mm long; reduced pappus of two short awns. Disc 217

florets 9-22, subtended by lanceolate pales, 2 mm long, 1 mm wide, 3-veined; corollas 2.5 mm long, throat 2 mm long, lobes 0.5 mm long, pubescent abaxially, tube 0.3 mm diameter;

anthers 1.4 mm long, styles 4 mm long, ovaries sterile, 1.2 mm long, villous (frequently only the upper one-third). Achenes 2 mm long, 1.5 mm diameter, puberulent at apex (upper one-third). Chromosome number, n = 16. Phenology. Flowering from May to September, occasionally as late as January. Distribution (Fig. 20). Broadly distributed: Brasil, British Guiana, Colombia, Costa Rica, Ecuador, Lesser

Antilles, Panama, Peru, Surinam and Venezuela; tropical rain forest, disturbed forest with secondary vegetation, along roadside, also in disturbed soils, commonly cultivated; 200- 600 (-1600) m.

Representative Specimens. Brasil. Amazonas: Rio Branco, Boa Vista, Ducken 1323 (F, GH, US); Basin of Rio Jurua, upper Rio Jurupary, Krukoff 5212 (A, MO, US); Carana, Vaupes, Rio Negro, Lemos Froes 212 89 (US); Yanomanmo Indian village, Prance & Ramos 2 3617 (MO, US); Roraima, Indian trail from Surucucu to Uaica, Prance et al.: 10529 (ECON, F, US); Posto Mucajai, Rio Mucajai, Prance et al.: 11151 (GH, US);

Basin of Rio Negro, Rio Uneiuxi, Maku Indian village, Prance et al. 15555 (ECON, F, US); Rio Negro, between Uaupes (Sao Gabriel) and Sao Felipe, Schultes & Pires 8965 (ECON, US); Acre, Rodovia Rio Branco-Porto Velho, Ubiratan et al. 171 (US). Maranhao: Maracassume River region, Froes 17 38 (A, MO, 218

NY). Para: Belem, near Instituto Agronomico do Norte, Archer

8249 (US); Rivers Moju and Acara, Austin & Cavalcante 4097 (MO); Castanhal, on Belem-Braganza railway, Barbosa da Silva 33 (US); Tapana, near Para, Killip & Smith 30232 (NY, US) ;

Instituto Agronomico do Norte, Silva 193 (US). British Guyana. Tumatumari, Indian Clearing, Gleason 286 (US); Rockstone, in and about the village, Gleason 619 (GH). Northwest District: Mabaruma Compound, Archer 2245 (US); Red Hill on the Barima River, Archer 2517 (US). Pomeroon

District: Kabakaburi, De La Cruz 3289 (F, GH, MO), Tabla, 1228 (GH, US) , 3229 (US) , Waini River, 3679 (F, GH, MO, US) . Issorora, Aruka River, Hitchcock 17 543 (GH). Yarikita Police Station, Hitchcock 17 658 (GH, US), Sand Hill, Bootooba, Demerara River, Persaud 45 (F); Basin of Essequibo River,

Smith 2182 (GH). Colombia. Amazonas: Leticia, ca. 1.5 hr walk E, estimated 4 mi Muroodoo, McDaniel et al. 2366 (US), 11799 (F, FSU, MO); vicinity of Leticia, Rio Amazonas, Schultes & Lopez 104 00 (US). Bolivar: Municipio de San

Jacinto, Cerro Maco, Zarucchi & Cuadros 3988 (US). Caqueta: Rio Caqueta, La Pedrera, Schultes 5 87 9 (ECON). Choco: Lloro, 50 km S of Quibdo, Archer 2049 (US); Rio Atrato, Samurindo, Cuatrecasas & Llano 24 084 (US). Pizarro, carretera Pie de Pepe-Barrecul, Espina 193 0 (MO, US). Istmina, alto Rio San Juan, Espina et al. 1144 (MO). Acandi, Untuiia, Reserva Indigena Cuna de Arquia, Forero 361 (MO). Unguia, Forero et al. 1984 (COL). Rio Sucio, Parque Nacional Los Catios, Forero et al. 17 67 (COL, MO); Hoya del Rio San Juan, Rio Bicordo, Forero et al. 4 622 (MO, OS); Andagoya, Forero et al. 5139 (OS). Quibdo, Puente de Cabi, Cordoba et al. 021 (MO); San Francisco Icho, Rio Icho, Garcia & Echavarrla 232 (MO); trail from Unguia along Rio Tigre, Gentry & Aguirre 15203 (COL); vicinity of Ijito, Choco Indian village, Gentry & Fallen 17253 (COL, MO, US); Rio San Juan between Dipurdu and San Miguel, Gentry & Fallen 17 695 (COL, MO, US); along Rio Pavarando, Gentry & Leon 20253 (MO, OS); Rio Yuto between Lloro and La Vuelta, Gentry & Renteria 24341 (MO, OS); a 9 km de Leticia, por la carretera Leticia-Tarapaca, Gutierrez et al. s.n. (US); Andagoya, Killip 35123 (COL, US); Rio El Salto, 9 km W of Andayoga, Lellinger & de la Sota 448 (COL). Rio Sucio, Truando, Romero-Castaneda 6144 (COL, NY); Tilupo, Romero-Castaneda 6254 (COL). Parque Nacional Natural Ensenada de Utria, Bahia Solano, Espina et al. 3668 (MO). Magdalena:

Aracataca, hacienda de "Macaraquilla", Dugand & Garcia- Barriga 2485 (US). Fundacion, 3 leguas N Santa Rosa, Romero- Castaneda 11155 (F, MO, NY). Narino: El Palmar, Sneidern 4522 (A, GH, LL, MO, NY, US). Putumayo: Puerto Limon, Plowman 2153 (A. ECON, US). Vaupes: Riberas del Rio Inirida, Fernandez 1934 (COL, US); Rio Kuduyari, Garcia-Barriga et al. 15802 (ECON, COL); Alto Vaupes, alrededores de Miraflores,

Gutierrez & Schultes 7 53 (GH); Rio Piraparana, Schultes & Cabrera 17153 (COL, GH, US); Mitu and vicinity, Rio Parana- Pichuna, Zarucchi 1304 (ECON, GH). Costa Rica. Cartago: 6 mi S of Cartago on route 2, Stuessy & Gardner 4493 (OS).

Puntarenas: Esquinas forest preserve, Allen 527 2 (F); Herradura, 2 km N Rio Cana Blanca, Arriagada 132 (OS); between Estipulas and Villanueva, Arriagada 133 (OS). Naranjito, San Jose, Arriagada 134 (OS); 3.5 km from NW of Estipulas to Naranjo, Arriagada 135C (OS). Quepos, Parque National Manuel Antonio, Arriagada 165 (OS). Las Cruces Botanical Gardens, Croat 4447 5 (MO). Monteverde, road before Reserve, Koptur 177 (MO). Cerro Amigos, Judziewicz 4581 (MO); 7 0 mi SE of San Isidro del General, Stuessy & Gardner 4507 (OS); 1.9 mi N of Villa Neily on gravel road to San Vito,

Stuessv Sc Gardner 4509 (OS) . Ecuador. Los Rios: Pichilingue, Acosta Solis 10771 (F). Canton Vinces, Hacienda El Naranjal, Mexia 667 8 (CAS, LL, US). Morona Santiago: basin of Rio Morona, Limbach 120 (CAS, MO). Napo: confluence of Quiwado and Tiwaeno Rivers, Davis & Yost 924 (ECON, F); Rio Pucino, bridge Aguarico Gentry 9811 (CAS, MO, OS); San Jose de Payamino, 40 km W of Coca, Irvine 104 (F); Santa Rosa at Rio Napo, Lugo 2052 (MO); Hacienda Aguinda, Rio Arajuno,

Maries & Palacios 39 (F); Rio Aguarico, village of Dureno, Martin 61 (ECON). Canton Napo, near Archidona, Mexia 7 256 (CAS, US). Rio Aguarico, Vickers 210 (F). Pastaza: basin of Rio Pastaza, 44 (NY); Cabeceras, Chiquita, at Rio Bobonaza, Lugo 1360 (MO); Ollaaard et al. 35322 (MO); Village of Rio Chico (8 km from Puyo), Shemluck 292 (ECON, F). Pichincha: 20 km W of Santo Domingo de Los Colorados, Cazalet & Pennington 5169 (NY); Gentry 9594 (MO). Zamora-Chinchipe:

Near Pangui, Hart 1567 (A, OS). Dominica: Botanic Garden at Roseau, Hodge 901 (GH); Carib Trail from Salybia to Hatton Garden, Hodge 3195 (GH) , gj^hli_& Stehle 6.40.8. (US) . Grenada: Saint George, Black Forest, Broadway s.n. (F). Saint Vincent: Mount Brisbane, Morton 594 6 (GH). Panama. 221

Canal Zone: Barro Colorado Island, w side of Orchid Island, Croat 122 83 (NY); Madden Dam near Rio Chagres, Seibert 559 (GH, MO, US). Chiriqui: N of Gualaca on Rio Chiriqui, Antonio 2 881 (CAS); 10-11 mi W of Puerto Armuelles Croat

22017 (MO); roadside above Boquete past Finca "Princesa Janca", D'Arcy & D'Arcy 6333 (CAS); gravel bar in Rio

Chiriqui Viejo, D'Arcy & D'Arcy 6590 (CAS) . Code: vicinity of El Valle, N rim, Allen 224 (GH); on way to Nombre de Dios, above road 18 km past Sardinilla, Croat 2609 0 (MO). Colon: 1 mi E from Puerto Pilon along dirt road to Maria Chiquita, Correa & Haines 234 (F); Croat 11364 (MO); Portobello Road, ca. 1 mi W of Portobello, Gentry 17 27 (F); ridges and drainages NW of Mina Boqueron # 1, Knapp & Sytsma 2401 (MO); ca. 2 km NW of Salamanca at bridge over Quebrada Bonita, Nee 6963 (MO); along Rio Guanche, 0.5-1 km upstream from Puerto

Pilon-Portobelo road, Nee 7139 (CAS); along Rio Culebra, above Santa Isabel, Pittier 4157 (US); Rio Guanche, Systma 1606 (CAS). Darien: vicinity of Yape, Allen 857 (NY); Rio Pierre, Croat & Porter 15520 (MO); E of Pinogana on Rio Tuira, Hartman & Duke 387 0 (OS); Rio Morti, ca. 6 mi upstream from Morti Abajo, Duke 10171 (MO); Cuipo forests near Santa Fe, Duke 12282 (MO); along Rio Chico from Nazareth to upstream, Hahn 195 (MO); Choco village, Piji Vasal, Folsom

4564A (CAS); Rio Chucunaque, ca. 6 mi NNW of Yaviza, Hartman 3876 (OS); ca. 2 mi W of Pinogana on Rio Tuira, Hartman & Duke 3871 (OS); Rio Jacque Valley, Knapp & Mallet 3207 (MO); Penas Bay near hotel, Tyson 5525 (MO); near Yaviza, Schmalzel & Al verson 1207 (CAS) ; Rio Paya along the shore, Stern et al. 236 {CAS, GH, MO); vicinity of El Real along trail to Rio

Pirre, Stern et al. 3 00 (GH). Panama: Rio Canina, near Jenine, Duke 3828 (CAS); Forests along headwaters of Rio Corso Duke 11927 (MO); San Jose Island, along road between Bodega Bay and Rio Mata Puerco, Duke 12516 (MO); beginning of El Llano-Carti-Tupile road, Dwyer & Nee 11988 (CAS, MO); Goofy Lake toward Cerro Azul, Dwyer & Stimson 8066 (CAS) . Isla del Rey, Correa 115 (A); Isla Viveros, on S side, Hartman 39 64 (OS); Isla Pacheca, Hartman 3937 (OS); Isla San

Jose, Jonhston 1069 (US); Isla Salaga, Knapp 327 3 (MO); unnamed tributary of Rio Chagres, 5 mi SW Cerro Brewster, Lewis et al. 33 82 (MO); junction of Rios Pacora and Corso, Oliver 2385 (CAS); banks of Mamoni River above Chepo, Pittier 4730 (GH, US). Veraguas: Isla de Coiba, Antonio 2224 (CAS); Mountains of S Azuero Peninsula, Hammel 5453 (MO) . San Bias: Ustopo, D'Arcy 9494 (MO); Ailigandi, in forest near waterfall, D'Arcy & Hammel 12255 (MO); near stream above

Puerto Obaldia, Gentry 1541 (MO). Peru. Amazonas: Quebrada Chichijam entsa, Rio Canepa, Ancuash 461 (MO); Confluence of Icikiti and Canepa, Berlin 625 (MO); 2 km up Quebrada Huampami, Berlin 984 (MO) ; al lado de Huampami, Monte, Kayap

1460 (OS, MO); Requena, Rio Ucayali, Peters 121 (MO). Huanuco: Tingo Maria-Aguaytia road, Gentry et al. 36138 (MO). Pachitea, Honoria, Bosque Nacional de Iparia, Schunke 1320 (COL, OS, US) ; Cucharas, on shores of Rio Hualaga, Woytkowski 5018 (MO). Junin: Huacapistana, on Rio Chanchamayo between Tarma and San Ramon, Fosberg 56191 (MO, US). Loreto: Quebrada Shanuce above Yurimaguas, Croat 18069 (MO); Rio Itaya, ca. 10 km above Iquitos, Croat 19138 (MO); Puerto Leguia, Rio Pachitea, Killip & Smith 26834 (F, GH, LP, NY) ; Maynas, km 8 Carretera Quistococha-Varillal, McDaniels et al. 27433 (US); vicinity of Aguaytia, Mathias & Tavlor 5056 (F). Iquitos, Muena-Cano, Mexia 6506a (CAS, GH, LL, NY); Above Pongo de Manseriche, Creek Carapisa, Mexiae 6254 (CAS,

F, MO, NY, US); Rio Yaguasyacu, affluent of Rio Ampiyacu, Brillo Nuevo and vicinity, Plowman et al. 6852 (ECON, MO); Purus River at the village of San Bernardo, Rutter & Barozo-

Ferrevra s.n. (OS); Ucayali, Pucapanga, Saaasteaui & Aldave 5733 (US); Lower Rio Huallaga, Puerto Arturo, Yurimaguas, Williams 4967 (F, LL); Garnitanacocha, Rio Mazan, Schunke 310

(CAS, F, GH, NY, US). San Martin: Lamas, N of San Antonio, BelShaw 3577 (CAS, F, GH, LL, MO, NY, US, WIS). Huascayacu, Brown s.n. (ECON). Mariscal Caceres, Juanjui, Ferreira 4515 (US); Bulzon de Utchya, Johns & Ramirez 202 (OS). Mariscal Caceres, Tocache Nuevo, Quebrada Luis Salas, Schunke 6629 (MO). San Martin, Tarapoto, Woytkowski 35185 (F, MO), 35187 (CAS). San Roque, Williams 7363 (F, US). Suriname. Sandrij I, Archer 27 67 (US); Carolina and vicinity, Archer 2893 (US), Archer 2919 (LL, US); Konashen-area, Essequibo River, Saparimo, Jansen-Jacobs et al. 1819 (MO); Paramaribo, Agricultural Experimental Station, Samuels s.n. (MO).

Venezuela. Anzoategui: Mendocero, orillas Rio Chive, Pittier 15059 (US). Bolivar: Cerro Auyantepui, Cardona Puig

264 (US); Confluencia Carun, Rio Paragua, Cardona Puig 2848 (NY). Carabobo: La Cumaca near San Diego, Alston 59 05 (BM, NY, US); Upper Guaremales, road from Puerto Cabello to San 224

Felipe, Pittier 8964 (GH, NY, US), Merida: entre Santo Domingo-Merida y La Mucuy, Aristeguieta 4932 (F). Monagas: near Capirito, Tamayo 3520 (MO). Tachira: along Quebrada Agua Azul, S of El Reposo, Steyermark & Liesner 118741 (MO). Territorio Federal Amazonas: San Carlos de Rio Negro, Clark & Clark 7 67 4 (MO); Atabapo, near Culebra on Rio Cunucunuma,

Liesner 17 877 (MO); Rio Cunucunuma at base of trail to Huachamacari, Liesner 25925 (MO).

Clibadium sylvestre is one of three broadly distributed species in the genus (C. egcrersii and C. surinamense are the others). Because of the large area covered by this species, ranging from Nicaragua to Peru, there is considerable morphological variation among populations throughout the range. Variation is observed mainly in vegetative structures. Leaves vary from irregularly dentate (Lewis et al. 3382) to almost crenate-serrate (Allen 4582); from large (20 cm; e.g..Croat 22557). medium (15 cm; e.g., Nee 6963) to small (8 cm; e.g., Harlow 2) . Pubescence also displays considerable variation, such as in the formerly recognized Clibadium appressipilum. with many slender stems and short appressed hairs (and also a toothed margin, e.g., Allen 857. Dwyer 2508. Johnston 1069). These characters do not correlate well with geography, however, and hence vitiate varietal recognition.

Clibadium sylvestre has its close relative in C. leiocarpum but differs by the strigose white-green (when 225 dried) phyllaries with obtuse apices in contrast to brown- green phyllaries with acute apices in C. leiocarpum. a tipically paniculiform capitulescence (vs. thyrsoid), villose achenes (vs. glabrous), and also by a rudimentary pappus of two short awns (absent in C. leiocarpum. but also found in C. anceps . an unrelated species of section Glomerata).

Clibadium sylvestre is adapted to lower elevations from 0- 600 m, whereas C. leiocarpum occurs 1400-2800 m. Clibadium sylvestre seems to be the only species producing acetylenic compounds which are responsible for the observed fish- poisoning activity (see Ethnobotany section).

Blake (1924) described Clibadium appressipilum to include only Panamanian elements, but this taxon falls clearly within the variation observed in C. sylvestre. The type of C. appressipilum has phyllaries 9-11-veined, whereas . C. sylvestre usually has phyllaries 7-11-veined; the pales subtending ray florets in C. appressipilum are 5-7-veined but in C. sylvestre they are always 7-veined. Despite these minor differences (that largely overlap), C. appressipilum clearly belongs within C. sylvestre. Both taxa occur in the same geographical area. In addition to similar morphology and overlapping distributions, the documented use of C. appressipilum as a fish poison (cf. Allen 857. NY), also supports its inclusion within C. sylvestre.

Blake (1924, p. 601), when describing C. appressipilum. indicated that the nearest ally of this species was C. 226

caudatum (also suggested by Robinson, 197 9) "in which the leaves are coarsely serrate, the branches of the inflorescence densely hispidulous with spreading hairs, and the phyllaries 12- to 16-nerved." However, Blake in his description of C . caudatum indicated close relationships with C . arboreum and C. surinamense. It is not clear on what character states these suggestions were based.

Earlier workers (Schulz, 1912; Blake, 1917; Stuessy, 197 5) mentioned a possible confusion in the application of the name C . asperum with C. sylvestre. also described by

Aublet in the same publication (both names as species of Baillieria). Schulz (1912) synonymized C. sylvestre with C. surinamense. Personal observations of the type specimens of

Baillieria aspera and Baillieria sylvestre at BM, and comparisons with protologues, confirmed (as suggested previously by Blake, 1924) that the names written on the sheets have been transposed. Therefore, the sheet labelled as Baillieria aspera corresponds to the type of Baillieria sylvestris. and vice versa.

28. Clibadium acuminatum Benth., Bot. Voy. Sulph. 114. 1844.--Type: Costa Rica. , 3-7 Apr 1838, Barclay 217 9 (lectotype here designated: MO!;

isolectotype: BM!, K[2]! photo: OS! fragment at US!). 227

Clibadium parviceps S. F. Blake, Contr. U. S. Natl.

Herb. 22:598. 1924.--Type: Venezuela. Aragua: Colonia Tovar, 1856-1857, Fendler 1967 (holotype: GH!, fragment at US!, photos: F! US!; isotype: K!, photo: O S !).

Clibadium sneidernii Cuatrec., Rev. Acad. Colomb. Cienc. Exact. 9: 240. 1954.--Type: Colombia. Narino: El Palmar (Cordillera Occidental, vertiente occidental), 25 Oct 1944, von Sneidern 4522 (holotype: F!, photo: US!)

Clibadium pediculatum Aristeg., Fl. Venez. (Comp.) 10:

393. 1964.--Type: Venezuela. Merida: La Carbonera, 24 Sep 1956, Aristecruieta 247 9 (holotype: VEN!; isotypes:

N Y ! U S !).

Shrub up to 5 m tall, woody at base; stems with branches slender, striate, strigose to densely strigose with hairs 1 mm long. Leaves with blades ovate, to ovate-lanceolate; membraneous; 6-20 cm long, 3-10 cm wide; 3 to 5-veined at base, at apex acuminate to largely acuminate, at base acute or atenuate, with margin serrate, pauci-serrate, serrulate or crenate-serrate, teeth usually mucronate; above dark green, slightly strigose with hairs 0.5 mm long, beneath pale green, scarcely pilose or shortly pilose mostly on the main veins with hairs 1 mm long; petioles slender, 1.5 to 5 cm long, strigose with hairs 1 mm long. Capitulescence thyrsoid, trichotomously branched. Heads subglobose, 4-5 mm high, 3-4 mm diameter, sessile or shortly pedicellate; phyllaries 3, suborbicular-orbicular, 3-4 mm long, 2.5 to 3.5 mm wide, 7- veined, at apex acute, abaxially greenish, strigose with hairs 0.5 mm long, with margin shortly ciliate at upper one- third. Ray florets 4-6, subtended by ovate pales, 3.5 mm long, 2.5 mm wide, 7 -veined, at apex acute, strigose with hairs 0.5 mm long, with margin scarcely ciliate at upper one- third; corolla white, 3 mm long, 1 mm diameter, glabrous, 3- lobed, lobes irregularly deltate, 0.4 mm long, scarcely pilose at apex; styles 1.5 mm long, branches 1 mm long. Disc florets 5-10, subtended by lanceolate pales (occasionally few central florets without pales), 3 mm long, 1-1.5 mm wide, glabrous, margin upper one-third shortly ciliate, 3-veined; corolla white, tubulate, 3 mm long, 1 mm diameter, 5-lobed, lobed deltate, 0.5 mm long scarcely pilose at apex; anthers black, 1 mm long; ovaries sterile 1.5 mm long, 0.8 mm diameter, scarcely pilose at upper apex. Achenes obovoid, rounded at base, 2 mm long, 1.7 mm diameter, pilose at upper one-third, with hairs 1 mm long. Chromosome number unknown. Phenology. Flowering from March through December.

Distribution (Fig. 21). Colombia, Costa Rica (Cocos Island), and Venezuela; mostly on edges of secondary tropical wet forest; 300-1300 m.

Representative Specimens. Venezuela. Aragua: Altos del Choronx, Badillo 1945 (F); cerca de La Cumbre de Choronx, Williams 10478 (F [2], US); Peninsula Paraguana, Cerro Santa 229

w

Fig. 21. Distribution of Clibadium acuminatum (® ) and C. micranthum (v ) in Colombia, Panama, and Venezuela, and Bolivia and Peru, respectively. Ana, Breteler 4286 (MO); along road between Colonia Tovar-

Caracas, Croat 544 69 (MO). Miranda: Cerros del Bachiller, S of Santa Cruz, Steyermark & Davidse 1167 30 (MO). Distrito Federal: Entre Aguas Megras-El Junquito, Badillo 2069 (F);

Parque El Avila, vertiente N, Benitez 2517 (F); Cordillera de la Costa, El Junquito, Steyermark 56956 (US); Fila de Agua Negra, Whetzel et al. 216 (US); El Junquito, Killip & Rohl 37165 (US). Merida: Tovar, alrededores de Zea, Badillo 6457 (F). Miranda: Parque Nacional Guatopo, Nee 17 658 (F) . Trujillo: alrededores de la via Flor de Patria-Bocono, Perez- Levis 7 9 (F).

Clibadium acuminatum is most closely related to C. micranthum. Both species are included (together with C.

leiocarpum and C. sylvestre) in section Oswalda. Clibadium acuminatum and C. micranthum have strigose ovate leaves, thyrsoid capitulescences, orbicular 7-veined phyllaries with acute apices, ray florets subtended by 7 -veined pales, and disc florets subtended by 3-veined pales. Clibadium acuminatum differs from its close relative in having strigose phyllaries (pilose in C. micranthum). 4-6 ray florets (vs. 10-14), 5-10 disc florets (vs. 10-11) with 5-lobed corollas (vs. 3-lobed), and pilose achenes (vs. glabrous).

Clibadium acuminatum was considered by Blake as closely related to C. parviceps based on general morphology, except for minor differences in number and shape of phyllaries, pales, and by more acuminate leaf apices. Blake (1924) 231 suggested this relationship based on the examination of one specimen of C. acuminatum (Steward 326) from the type locality. He noticed similarity in general morphology but the specimen had only 2 narrow phyllaries in contrast to 3 more ovate phyllaries observed in C. parviceps. After having studied the same material (Steward 326) and the type specimens of C . acuminatum and C. parviceps. I have observed some heads of C . acuminatum with 2 or 3 ovate phyllaries (2 phyllaries are less frequently found, however). The characters of both species overlap to such extent that synonymy for C. parviceps seems appropriate.

Four syntypes of C . acuminatum collected by Barclay have been located. The two specimens at K and the one at BM are not well preserved and therefore they do not comunicate well the diagnostic characters of the species. However, the one at MO (ex BM) here chosen as lectotype, is the best preserved and has all diagnostic characters.

29. Clibadium micranthum 0. E. Schulz, Bot. Jahrb. Syst. 46: 625. 1912.--Type: Peru, 17 80 [Dahlgren, 1940], Ruiz 3 (holotype: B destroyed; photos: F! GH! MO!).

Branched shrub, 2 m tall; stems with branches striate, pilose with hairs 0.5-1 mm long. Leaves with blades ovate to ovate-lanceolate, membraneous, 8-15 cm long, 4-8 cm wide, at apex largely acuminate, at base atenuate, with margin shortly serrate to serrulate; upper surface glabrous to shortly strigose with hairs 0.5 mm long, lower surface glabrous; petioles 1-2.5 cm long, strigose with hairs 1 mm long. Capitulescence thyrsoid, trichotomously divided, branches slightly striate, strigose. Heads subglobose, 3 mm long, 2 mm diameter, sessile or shortly pedicellated; phyllaries 3, ovate to suborbicular, 3 mm long, 2-2.5 mm wide, at apex acute, abaxially short pilose with hairs 0.3 mm long, with margin upper one-third shortly ciliate, 7 -veined. Ray florets 10-14, subtended by ovate to oblong-ovate pales, 3 mm long, 2 mm wide, 7 -veined, shortly pilose, with hairs 0.4 mm long, with margin upper one-third shortly ciliate; corolla white, glabra, 1.5 mm long, 0.5 mm diameter, 3-lobed, lobes 0.3 mm long, glabrous; styles 1.5 mm long, branches 0.5 mm long. Disc florets 10-11, subtended by lanceolate pales, 2.5 mm long, 1-1.5 mm wide, 3-veined; corolla tubulate, white, 1.5 mm long, 0.5 mm diameter, 3-lobed, lobes irregularly deltate, 0,4 mm long, glabrous; anthers black, 1 mm long; ovaries sterile 1.5 mm long, 0.5 mm diameter, glabrous. Achene obovoid, 1.5-2 mm long, 1-1.4 mm diameter, rounded at base, glabrous (exceptionally scarcely pilose at apex). Chromosome number unknown. Phenology. Flowering from April through December.

Distribution (Fig. 21). Bolivia and Peru; secondary vegetation, disturbed areas; 700-17 00 m.

Representative Specimen. Bolivia. La Paz: Yungas: Polo- Polo bei Coroico, Buchtien 230 (F, NY); Maripi, Rushy 2144 (GH, NY, US); near Yungas, Rusby 2146 (GH, NY, US). Peru. Amazonas: trail E from La Peca into Serrania de Bagua, 5-10 trail km from La Peca, Gentry et al. 22846 (F). Valle del Rio Santiago, Quebrada Caterpiza, 2-3 km atras de la comunidad de Caterpiza, Tungui 335 (F), 496 (F). Cuzco: Quispicanchis, Inambari, Vargas 1547 2 (US); La Convencion, along edges of Rio Mapitunuari and Rio Klause, below and on descent to Camp

I, Dudley 10091 (F); from Luisiana to Rio Apurimac & Campa Indian's, Dudley 11411 (F); near Rio Yanamayo, below Pillahuata, Pennell 14035 (F, US). Huanuco: Leoncio Prado, Hermilio Valdizan, cerca de La Divisora, Rimachi 5006 (CAS), Schunke 9350 (F), 11260 (CAS). Junin: ca. 51 km NE of Tarma on road to San Ramon, Dillon & Turner 1428 (F, OS); 26.3 km NE of central plaza in Acobamba (NE of Tarma) on carretera 20 B to Oxapampa, Jones & Davidson 9144 (MO); Schunke Hacienda, above San Ramon, Killin & Smith 24822 (F, GH, LP, NY, US); La Merced, Killip & Smith 23550 (F, NY) ; Pichis Trail, Yapas, Killip & Smith 25494 (NY); Chanchamayo, Fondo Romero,

Pampatigre, above Santa Ana (SE of La Merced), Stein & Todzia 2340 (MO, OS). Loreto: Iquitos, Williams 8205 (LP); Puerto Leguia, Killip & Smith 26834 (LP). Madre de Dios: Iberia, vicinity Rio Tahuamanu, Seibert 2022 (LP); Nishuiya, Velarde-

Nunez 24 66 (LP).

Clibadium micranthum is characterized by the presence of small heads (2.5-3 mm long) arranged in loosely cymose- thyrsoid capitulescences. It is closely related to C. acuminatum, also of section Oswalda. Clibadium micranthum 234

differs from C . acuminatum because of pilose phyllaries (strigose in C. acuminatum). 10-14 ray florets (vs. 4-6), 10- 11 disc florets with corollas 3-lobed (vs. 5-10 and 5-lobed), and glabrous achenes (vs. pilose).

Clibadium micranthum appears superficially similar to C. microcephalum from Ecuador, a distant species in different

section Glomerata). They differ clearly in head structure and organization. Clibadium microcephalum has glabrous phyllaries with obtuse apices, 3 ray florets with 4-lobed corollas, 3-4 epaleaceous disc florets with 5-lobed corollas, and pilose achenes. DOUBTFUL AMD EXCLUDED NAMES

Clibadium agreste Mart, ex Baker, FI. Bras. 6: 157. 187 3.--

Type: Brasil. Sao Paulo, Ponte Alta, "dans la Serra do urbano ou das Divisoes," Dec 1827, Sello 4920 (holotype: B, presumably destroyed, photo: GH!). = Ichthyothere agreste Mart, ex Baker, FI. Bras. 4: 157. 187 3.

Clibadium angustifolium Mart, ex Baker, FI. Bras. 6: 154.

1873.--Type: Brasil. Sao Paulo, Ponte Alta, "pres de la cascade," Pohl 472 {holotype: K, photo: GH!; isotype: K, photo: OS!). = Ichthyothere linearis Mart, ex Baker, FI. Bras. 4: 154. 1873.

Clibadium? commelinoides DC., Prodr. 5:506. 1836.--Type: Brasil. Maypuri, 1836, Schomburgk 247 (holotype: K, photo: OS!; isotype: BM photo: OS!, K photo: O S !). = Ichthyothere cunabi Mart, ex Baker, Fl. Bras. 4: 154. 1873.

Clibadium foetidum Alemand ex Linn., Mant. Pi. 294. 1771. nomen nudum.

Clibadium hispidulum (Wright ex Griseb.) Gomez de la Maza & Molina, Anal. Soc Espan. Hist. Nat. 19: 27 6. 1890.

235 236

Lantanopsis hispidula Wright ex Griseb., Mem. Am. Acad.

N. S. 8, 513. 1863.--Type: Cuba. Oriente, "prope villam Monte Verde dictam," Jan-Jul 1859, Wright 1315 (holotype: K photo: OS!; isotypes: BM photo: OS!, P photo: OS!). = Lantanopsis hispidula Wright ex Griseb.

Clibadium neriifolium (H.B.K.) DC., Prodr 5: 507. 1836. Baillieria neriifolia H.B.K., Nov. Gen. sp. [ed. fol.]

4: 289. 1818.--Type: Venezuela. Federal District: Silla de Caracas, 21 Nov -7 Feb 1799-1800 [Sandwith, 1968], Humboldt & Bonpland (holotype: P). = neriifolia (H.B.K.) Schultz Bip., Wedd. Chlor. And. 1:67. 1856.

Clibadium pedunculosum (Rich.) DC., Prodr. 5: 507. 1836-- Type: French Guyana, 17 92, Leblond 339 (holotype: G, photos: F! MO!). = Trixis pedunculosa Rich., Actes Soc.

Hist. Nat. Paris 1. 112. 17 92.

Clibadium schomburgkii Schultz Bip., Schomb. Faun, et Flor.

Guian. 940. 1848. nomen nudum. APPENDIX A

NUMERICAL LIST OF SPECIES

237 238

NUMERICAL LIST OF SPECIES

1. £. eggersii 2. £. leptophyllum 3. £. divaricatum 4. £. terebinthinaceum 5. £. erosum 6. £. anceps 7. £. glomeratum 8. £. trianap 9. £. sessile 10. £. microcephalum 11. £. frontinoense 12. £. zaruchii 13. £. rhvtidophyllum 14. £. sprucei 15. £. grandifolium 16. £. cordatum 17. £. manabiense 18. £. glabrescens 19. £. oentaneuron 20. £. laxum 21. £. S.Odiroi 22. £. surinamen.se 23. £. peruvianum 24. £. arboreum 25. £. armanii 26. £. leiocarpum 27. £. sy l .vest re. 28. £. acuminatum 29. £. micranthum APPENDIX B

INDEX TO NUMBERED COLLECTIONS EXAMINED

239 240

INDEX TO NUMBERED COLLECTIONS EXAMINED

The numbers in parentheses refer to the corresponding species in the text and in the Numerical List of Species presented above. Collections are cited under the name of the principal collector.

Acosta Solis 9723(10); 10771(27); 12285(4);

Aguilar 322 (24) ; Alain 9422(5); Allart 230(4); Allen 118(26); 224 (27); 857(27); 3621(15); 4653(26); 5272 (24); 5637(7);

Almeda 2987 (6); 3145(4); 3304(4); 3515 (26); 4023 (3); 4033(3); 4171(7); Alston 5210 (4); 5324 (4); 5905(27); 7901(3);

Ancuash 461(27); Anderson 3301 (4) ; 6739(25); 9603(25); Antonio 1136(15); 1146 (6);2224 (27) ; 2881(27); 5041(6); Aranda 32(4); Araque 18C740(15);

Araquistain 655 (26); 1157(26); 2440(3); Archer 107 (22); 481(22); 1150(19); 1945(3); 2049(27); 2058 (22); 2187(16); 2245(27); 2517 (27); 2647 (22); 2684(22); 2716 (22); 2725(22);

2767 (27); 2857 (22); 2893 (27); 2919(27); 241

8249 (27) Aristeguieta 3268(22); 4932(27); Arriagada 131(22); 132(27); 133(27); 134(27); 135b(6); 135c (27) ; 142 (22); 143 (22); 146 (26); 149(22); 150(22); 151(22); 152(7); 153(7); 154(7); 155 (22); 159(22); 160(26); 162 (26); 163 (22);

164 (22); 165(27); 308(26); 309(26); 310(22);

400 (26); 405(26); 406(15); 407(26); 408(6); 409(26); 411(26); 412(1); 413(18); 415(26);

417 (26) ; Asplund 5389 (22) ; 7614(22); 10544 (22); Austin 4097 (27) ;

Ayala 501(23); 3104(23); Badillo 1945 (28) ; 2069 (28); 6457 (28); Bailey 252 (22);

Balgooy 2903 (22); Bang 361(23); Barbosa 33(27); 6646(16); Barbour 4299 (23) ; 4386 (23); 5386(23); Barclay 3299 (8); Barringer 3211(22); 3365(26); 4003(6); Balslev 10341 (10); 10652 (22); Beard 1157(5); 1360(5); Belem 1289(25); Belshaw 3316 (23) ; 3577(27);

Benavides 4688(19); Berlin 625 (27); 984 (27); 1633 (23); Bernardi 361(22); Benitez 2517 (28);

Berry 917 (22) ; Biolley 7 399(1); Blackwell 2749(15); Blake 7354 (1); Blum 487 (22); 2662 (6) ; Boeea 5901(22);

Borsboom 12293(22); Brant 137 8(19); Brenes 3817(6); 6335(6); 20586 (6); Breedlove 15423(24); 26410(24); 26504(24); 42714(24) Breteler 4286(28);

Bridgeman 55(22); Bristol 519(19); Brito 12(25); Britton 179(5); 357(4); 554 (4); 2132(5); 2810(4); 3579(4); 3976(4); 5420(5); 6454 (5); 7586(5);

Broadway (s.n.)(24); 376(22); 634(22); Brown s.n. (27); 5530(5); Buchtien 230(29); Burger 47 44 (7) ; 5144 (22); 5735 (7); 6435 (6); 9010(7); 9048(6); 9406(6); Burch 4656(6); 114(24); Bussey 534 (26) ; 825(1); Cabrera 29986(25); Calzada 500(24); 4716(24); Callejas 2285 (4); 6610(15); 6842 (12); 243

Camp 884(21); 3601(22); 3826(20);

Carlson 3405(15); Cardona Puig 264 (27);2848 (27); Carleton 458(1); Cazalet 5169(27); 5188(1); Cedillo 288(24); Chacon 832 (22) ;

Chagas 252(25); Chambers 2738(5); 2756(5); 2770 (5);

Chaparro 81(1); Charpin 13274 (8) ; Chavarria 143 (1); Churchill 4429(1); 5485(6); Clark 7 674 (27) ;

Cochrane 6287 (9) ; 6289 (9) ; Coelho 1739(22) Coleman 265(25); Contreras 2873(24); 7026(24); 9951(24);

Cordoba 21 (27): Correa 115 (27) ; 234 (27); 3349(15); 2781(26); S.n. (26) ; Cowan 1418(4); 2427 (24); Croat 10005(1); 10715(22); 11364(27); 12283(27) 13372(15); 13518(20); 15520(27); 17465(23) 18069(27); 18142(23); 19138(27); 19316(23) 20238(23); 22017(27); 23190(1); 23863(24); 24155(24); 24409(24); 24522(1); 26090(27);

33404(26); 35500(26); 36089(6); 37102(9); 37311(9); 37677 (15); 37982 (22); 38398(22); 244

39712(24); 40283 (24); 41579(24); 41776(24); 43195(1); 43470 (6); 44475(27); 47008(26); 49326(13); 53802 (22); 54469 (28); 56097 (2); 66523 (6); 66554(6); 68217 (6); 69937 (22); 70305 (22) ; 70571 (22) ;

Cronquist 9631(24); Crosby 1045(4);

Cruz 286 (24) ; Cuatrecasas 61(8); 6112(22); 8182(8); 11521(8); 11994-A(22); 12086(22); 13696(16); 14004(22); 15025 (15); 17534(22); 18653(22); 22034(19); 22478(22); 22163(22); 23808(22); 23932(15); 24084(27); 27121(22); 27504(22); 27601(15); 27639 (8) ; 27933 (22) ; D'Arcy 6537 (26) ; 6611(6); 6333 (27); 6590(27); 9494 (27); 10322(15); 10911 (9) ; 12255(27); 12294(15); 15860(26); 15904(15); 15970(6);

16230 (15) ; Davidse 4748 (22) ; 5649(22); 24607 (26); Davidson 6717(1); Davis 789 (23); 924(27); De la Cruz 1228 (27); 3229(27); 3289 (27); 3459 (22); 3679 (27); Delessert 5625(25); Denslow 79-104 (1); 407(22); 2276(22); 2642 (1); Devia 624 (16); 977 (22); Dxaz-Piedrahita47 (19); 181(4); 650(23); Diers 564(23); Dillon 1428(29); 1454(23); Dodge 5603(7); Dodson 1849(14); 6385(20); 7235(1); 7589(20); 9817(22); 12703(22); Donell-Smith 2347 (24) ; Dressier 50(24); 4798(1); Duarte 44 64 (25); Ducken 1323 (27) ;

Dudley 10091 (29); 11411(29); Dugand 2485 (27) ; 3014(22); 3854(22); Duke 3828 (27); 6088 (3); 9886(15); 10171(27); 11470(3); 11918(3); 11927(27); 12116(6); 12282(27); 12516(27); 16117(7);

Dusen 16424(25); Dwyer 5111(15);5128(15) ; 8066 (27); 8669 (28); 8844(6); 9496(1); 11198(1); 11415(24); 11988 (27) ; Ebinger 362 (22); Eggers 619(5); 3057 (22); Echeverry 2322(4); Eiten 5880(25); Ekman 168(4);

Ernst 1088(5); 2035(5); Espina 1144(27); 1930(27); 1991(15); 3668(27) Eyerdam 24694 (23); 24802 (23); Faber-Langendoen 1856(19); Fay 789 (24) ; Feisinger 812093(6); 822182(26); Fendler 640(22); Fernadez 1934 (27) ;

Fernandez-Casas 804 8 (23) ; Ferreira 3358(23); 4515(27); 6805(3); Fiebrig 5815(25); Folsom 2422 (6) ; 3648(6); 4564A(27); 4753 (6); 5378(6);

5813(15); Forero 361 (27) ; 725 (1); 1767(27); 1984(27); 2257 (16); 2279 (19); 2347 (16); 2824 (4) ;3312(15) ; 3517(16); 4163(2); 4622(27); 5139(27);

5476(15); 5527 (2); 5871(16); 6853 (15); 7594(16); 5540(15); Fosberg 21619(22); 22969(20); 56191(27);

Foster 9315(1); Froes 17 38(27); Fuchs 21728(15); Funk 10654(22); Garcia-Barriga 7716(22); 12772(22); 15802(27); 18325(22)

Garcia 232(27);

Garganta 853(22) / Gentle 8251(1); Gentry 94 8 (22); 1541(27); 1737(27); 3034 (15);

9036A(19); 9594 (27); 9811(27); 12409 (1); 15203 (27) ; 17249(15); 17253 (27); 17695 (27) 17942 (22) ; 20253 (27); 21604 (23); 21861(23) 22846(29); 23125(23); 23381(16); 24341 (27)

27259(23); 27359 (23); 27389(23); 27992 (23) 28484 (20) ; 28811(23); 29189(23); 29668(23) 30826(10); 31334(23); 36138(27); 40293 (19) 50363(5); 54125(22); 54996(21); 63181(22); Gehriger 361(22);

Giannotti 4567(25); Gibson 2512(24); Gill 44 (27); Gilli 113(26); Gillis 10162(20); Gines 4481(22); Glaziou 9484(25); 9489 (25); Gleason 286 (27) ; 619(27); Grant 97 59 (22); Grayum 27 00 (22); 9041(22);

Grijalva 1180(22); 2109(26); 2509(26); 2859(22); Gomez 23246(6)

Gomez-LauritO 5515 (7) ; 6419(15); 8599(26); 8740(1) 9378(26); Gomez-Pompa 1546(24); 4517(24); Guanchez 2088(22);

Gutierrez 753(27); 944 (22); 170122(1); 170657 (22); (S.n.)(27);

Guzman 7 01 (22); Haber 737(6); 835 (26); 1189 (26); 5362(6); Hahn 195(27);

Hamilton 715(26); 1391(15); 2729(15); 3656(26); 3857 (26);

Hammel 1691(15); 2599(6); 2621(15); 3891(15); 4072(22); 4353(22); 5453(27); 9240(15); 14039 (22) ; 248

Hampshire 492 (1); 97 6(15); Hansen 7586(24); 9422(5); Harley 11081 (25); 11249 (25); 21874(25); Harling 9670 (22); 9680 (20); 10124(10); 10153(21);

11525(20); 11876(22); Harmon 2552 (24) ; 2553(24); 2826(24); 4031(24); Harris 3183(4); 15157(4);

Hart 1567(27); Hartman 387 0 (27); 3871 (27); 3876(27); 3916(9); 3917 (26); 3920(26); 3926(22); 3932 (22); 3936(15); 3937(27); 3942(22); 3963 (6);

3964 (27); 12254(15);

Hatschbach 25957 (25); 26091(25); 34993(25); 43850(25); 46174 (25) ; Haught 1914 (22) ; 2047 (22) ; 2124 (22) ;

4678(15); 6488(8); Heringer 2784 (25); 7560 (25) ; Hermano-Daniel 3986(22); 4205(19); Hermano Tomas 175(19); Hendrix 149(23); Herrera 939(22); Hespenheide 533(5); 1119(4); Hitchcock 16875 (4); 17021(22); 17543(27); 17658 (27);

20261 (22); s.n. (4); Hill 89(1); Hodge 736 (5); 901(27); 1037 (5); 1201(5); 1396 (5); 1462(5); 1870(5); 2241(5); 2380(5); 3195 (27) ; Holm 7 (22) ; 121 (7); Hood 977 (22) ; Howard 11824 (5) ; 11911(5); 17533 (5); 19387 (5); Huft 1621(15);

Hutchinson 3721(1); Idrobo 69(22); Irvine 104(27); 363(22); Irwin 393(22); 9642(25); 11209(25); 11381(25); 12674 (25); 16352(25); 18743 (25); 23772 (25); 25432(25); 25681(25); 26443(25); 29507(25); Jansen 547 (22) ;

Jansen-Jacobs 1819(27); Jaramillo 4030 (22) ; 6700(3); 7559(20); 7797 (17) Jativa 103 (22); 804 (20); 2102(20);

Jeffrey 100(16); Jimenez 1269(6); 2224(6); Johnston 1069(27); Jones 9144(29); Johns 202(27); 239(23); Judd 525(5); Judziewicz 4581(27);

Juncosa 1733(3); Kayap 14 60(27);

Kennedy 1910(7); Khan 450 (6); 587 (26); Killip 5268(15); 8976(22); 20783(22); 22450(23);

22810(23); 22860(23); 23034(23); 23550(29); 24049 (23); 24180(23); 24822 (29) ;25068 (23) ; 25494 (29);26305(23); 26474 (23); 26834(29); 27061(23); 27263(23); 31232(27); 33061(22) 33575 (15) ;33867 (22);33982 (8) ; 34462 (22); 34815(22); 35123 (24); 37165 (28); 37592(22) 38862(15); 39842(19); King 875(20); 5257 (22); 5754 (19); 5764(19); 5791(22); 5814 (22); 5824(22); 5852 (22); 6305(5); 6396(5); 6567 (10); 6560(17); 6746 (20); 6766 (7); 6778(7); 6805 (26); 6831(26); 6871(22); 7005(22); 7106(24); 7012(5); 7122(24); 7020(24); 7972 (20); 7735(17); 7951(22); 8000(25); 8008(25); 8205(25); 8286(25); 8437 (25); 8966(25); Kirkbride 94 (26) ; 393 (22); Klug 2348(23); Knapp 1532 (26); 2401(27); 3101(1); 3207(27); 3273 (27); 5633(6); 5825(22);

Knight 711(4); s.n.(5); Koie 4880(2); Koptur 177 (27)

Koster 49 (22);

Krapovickas 14274(25); 34351(25); Krukoff 5212(27);

Kuntze s.n.(23); Kvist 230(22); Lagenheim 3264(22); Lanjouw 12056(22); Lawrence 135(15); Lazor 2983(22); Lellinger 448(27); Lemos-Froes 21289(27);

Lent 435 (3); 860(7); Leon 660(3); 10880(4); Lleras P17 376 (22);

Lewis 596 (26); 975(1); 2082 (15); 3382(27); Liesner 1713 (7) ; 2098 (1); 2815(7); 17877 (27); 24447 (22) ; 25925 (27) ; Limbach 120 (27) ; Little 8114(22); 13064(5); 13554(5); 16320(5);

Logname 7399(23); 7534c(23); Lorentzen 3792(20); Lugo 152 (22); 224(22); 228(22); 763 (14); 1027 (22); 1343(22); 1360(27); 1703(22); 2052(27); 2481(1); 2926(22); 3890(22); 3986(22); 4554(10); 4605(21); 4941(22); Lundell 20044(24);

Luteyn 11197(14); 1447(22); 1638(15); 4833(4); Macedo 1771(25); MacDaniels 860(24); 27433(27);

McPherson 7529 (6); 11073(15); 13396 (19) ; McDowell 715(22); MacRae 35(23); 118(3); Magalhaes 3183(25); Maguire 22707 (22); 22956(22); 40227 (22); 40336 (22) Maias 144 (22); Marcks 826(24); Maries 33 (27); Martin 61(27); Martinez 128(24); 1725(24);

Mathias 5056(27); 5191(20); Matuda 2422 (24) ; Maxon 197 (4) ; 1323 (4) ; Maxwell 217 (24) ; McDaniel 2366(27); 11799(27); 14482(24); 17089(23); McDonagh 165(1); McVaugh 21414(24); Medina249(19) ; Mexia 6254(27); 6255(1); 6506(27); 6595(18);

6668(18); 6678(27); 6696(20); 6915a(18); 6946 (21); 7024 (1); 7256 (27); 7272 (1); 8467 (20)

Mejia 86 (22); Miller 1221(22); 1422(26); Molina 314 8 (22) ; 3601(24); 11863 (22); 12260(24); 14147(22); 15860(24); 17508(6); 30573(22);

Monsalve 624(19); 1601(19); Monteiro 3100(25); Moreno 9165(26); 10313(22); 10613(22); 13072(1); 15098(1); 23298(22); Mori 3860(15); 5590(1); 5897(15); 6164(1);

13643(25); 13719(25); Morley 480 (4); 493 (4);

Moore 142(23); Morton 5946 (27) ; Murata 4451(22); 253

Murillo 560 (22) ; Nelson 2728 (1) ; 3109 (24); 4078 (22); 4653 (22); 4830 (1) ; 5059 (22) ; Nee 3780 (22); 6963 (24); 7139(27); 8176(4); 17658(28); 18792 (24); 24914(24); 26315(24); Neill 97(1); 2920 (26); 3578 (1); 4207(1); 7394(22); 8525 (21);

Nicolson 4098(5); Nowicke 3434(1); Nunes 116(25); Ollgaard 35322(27); Oliver 2385(27); 3667(1); Olsen 596 (22) ;

Opler 1602(1); Orcutt 3113(4); 3324(4); Otero 586(5);

Pelaez 510(19); Pennell 4219 (22); 7217 (22); 8704(22); 14035(29);

Pere Duss 316(5); Pereira 4950(25); 8443(25); 9064(25); Perez Arbelaez 6142(15); 6441(22);

Perez Levis 79(28); Persaud 45 (27) ; 341(22); Peters 121(27); Philcox 3292(25); Philipson 2407(8); Pittier 749 (22) ; 3136(26); 4157 (27); 4730(27); 7893 (22); 8964(27); 9664(4); 13596 (22); 15059(27); 15061(22); 16068(15); Pipoly 1999(1); 3759(1); 6005(1); 9962(4); Plowman 2153(27); 6487(23); 6852(27); 11647(23); 12080(22) Poveda 1508(7); 4135(1); 4138(15); Prance 10529(27); 11151(4); 15555(27); 23617(27);

Pringle 10802 (24); Proctor 17 0(5); 6864(4); 8313(4); 10341(4); 19116(5);

19605(5); 32215(5); Prescott 404(21); 405(1); Porter 4651(15); 4785(1); Purpus 37 05 (24); Puttemans 209(25); Ramirez 59(23);

Ramos 1599(15); Ratcliff 10(24); Raven 19974 (24); 20863 (7); 21977(6); 22048(22); Redher s.n.(4); Renteria 1831(22); Revilla 2446(23); Rimachi 5006(29); 5992(23); 7400(23); Rimbach 7 00(14);

Robles 17 22 (22); Rodriguez 169(22); Rojas 10053(25);

Romero-Castaneda 2974(3); 6144(27); 6254(27); 6689(22); 11155 (27) ; Rosas 836(24);

Ruiz Teran 1139(18); Rusby 2144(29); 2145(23); 2146(29); 255

Rutter (s.n.)(27); Sagastegui 57 33(27); Samuels 345(22); s.n. (27); Sanchez 510(15); Santos 1938(25); Sastre 2626(5); Schipp 972 (1) ; Schultes 5879(27); 8965(27); 10400(27); 17153(27); 17861 (4) ; Schunke 310(27); 1320(27); 1672(23); 2959(23); 6629(27); 7786(3); 9350(29); 11260(29);

Seibert 245(26); 519(22); 559(27); 2022(29); Shafer 485(5); Sharp 45808(24); Shemluck 292(27); ShiIon Ton 2496(24); 3311(24); Silva 193(27); 936(25); 2168(25); Silverstone 1254(22); Skutch 1797(24); 2657(6); 2733(22); 3260(6); 4402 (21); 4486(1); 5217(1); 5271(1); Smith 538(23); 1601(25); 1698(23); 2182(27);

5830(23); 6614(15); 10245(5); 10530(5); 12855(23); 14842(25); 15415(25); Sneidern 1540(22); 4522(27); 5167(22); Spruce 2136(22); Soejarto 2715(1); Solange 9(22); Solomon 7 515(23); 10025(23); 12945(23); 256

Soukup 2260 (3) ; Standley 33652 (6); 34088(26); 41894(26); 44639 (1); 48375(22); 48507(1); 49762 (7); 62042(24); 64559 (24); 85097(24); 86786(24); Stein 2340(29); Steiribach 9018(23); Stern 80(26); 236(27); 300(27); 1019(6); 1066(26); Stevens 7 53(18); 1104(24); 1390a(24); 4816(15); 4928(22); 9625(22); 10659(22); 12330(22); 13986(26); 13986(6); 14136(6); 15140(22); 15401(22); 17702(22); 18186(20);

19799(22); 21078a(l); 22445(22); 23951(1); 24140(1); 24606(1); Steyermark 37622(24); 41670(24); 45106(24); 46626(24);

47342(24); 55928(24); 56567 (25); 56956(28);

56890(22); 58625(4); 62029(25); 116730(28); 118392(28); 118648(28); 118741(27); 120954 (25); 127470(22); 131743 (4); stehle 86(5); 6408(27); Straw 2441(23); Stuessy4462(26); 4465(1); 4451(22); 4460(22); 4463(22); 4479(26); 4489(22); 4493(27); 4494(26); 4507(27); 4509(27); 4517(6); 4518(6); 4519(7); 4534(15); 4540(22); 4863 (13); 4871(13); 4877 (20); 4896(13); 4902(20); 4919(22); 4928(22); 4935(22);

4942(20); 4954(22); 4960(10); 4963(10); 4965(10); 4968(10); 4969(21); 4971(18); 4977 (18); 4978(21); 4980(10); 4981(18); 4982(10); 4984 (18); 4985(18); 4987(21); 5528(8); 5595(19); 5709(19); 5814(10); 5816(10); 5817(21); 5822 (10); 5823 (10); 5856 (22); 5867(20); 5873 (22); 12318 (20); 12352 (20); 5901(20); 5897(20); 12337 (8); 12347(8); 12353(8); 12355(8); 12363(10);

12365(10); 12369(10); 12370(10); 12400(14); Sullivan 181(1); 328(15); 375(6); 433(22); 4492(7) Systma 1606(27); 1803(6); Sauleda 87 84(5); Sargent B9 8(5); Schmalzel 1023 (4); 1207 (27); 1571(26); 1989 (22); Schubert 419(5); Stevenson 1558(5);

Stimson 1533(5); 1571(5); 5274(22); Tamayo 3520(27); Tate 577(14); Taylor 318 (4) ; Terborgh 346(5);

Todzia 1721(1); 2549(6); Tonduz 1147 9(1); Torres 5917(24); Tun Ortiz 1359(24); 1996(24); 2468(24); Tunqui 335 (29);496 (29); Turckheim 4148 (1); 7 903 (24) ; Turner 7 89 (23) ; 15024A(26); 15030(22); 15042 (26); Tyson 919(22); 1411(22); 5525(27); 6286(22); 258

6312 (22); 6317 (22); 4122 (22);

Ubiratan 171(27); Uribe 4098(19); 4340(22); 4917(19); 5715(22); 5873(22); 6310(19); 6957 (22); Usteri 16711(25); Utley 105(7); 5158(26); 6059(22); Vargas 13983(23); 15472(29); 18619(23); 18788(23); Velarde-Nunez 2466(29); Venturi 1180(23); Vickers 210(27); Viegas 3803 (25); Vogl 7 08(22); von Sneider 5168(1); Wachenheim s.n.(4);

Wagner 98(5); 116(5); 970(5); Wasshausen 416(5); Webster 12198 (6) ; 13352(5); 21911(7); Wedel 2161(1); Werling 84(20); Weston 4550 (7) ; 4479 (26); 3420(26); 5624 (26);

3001(20); 4714 (26); 4407 (22); Whetzel 216(28); Widgren 166(22); Wilbur 10575(6); 10661(22); 10678(6); 10884(26); 12084 (22); 13669(26); 13701(26); 15271(26); 22349 (6); 22460 (6) ;22785 (7 ) ;

Williams 328(23); 468(23); 1929(23); 1998(23); 3344(23); 4304 (23); 4967(26); 5119(4); 259

5122(4); 5198(4); 5655(23); 6684(23);

7363(27); 8205(29); 10478(28); 12241(25); 14474(24); 14621(1);(25); 20263(15); 26903(24); 28141(26); 28560(26); 28923(26); 42259 (25) ; Wilson 150(1); White 14(1); Whitefoord 2421(24); 5143(5); * Woytkowski 376(23); 5018(27); 8310(1); 35185(27); Woronow 4788(22); Woodson 731(22); 908(26); 1359(22); 1408(22); Wurdack 2224(23); 35740(22); Yuncker 8699(1); 17624(22); 18827(4); Zack 1194(17); 1318(17);

Zardini 4049(25); 4185 (25); Zarucchi 3988(27); 1304(27); 5036(15); 7097(19); 5286(22); 5443(22); 5566(8);

Zaruma 66 (22). APPENDIX C

NOTAS SOBRE LAS RELACIONES GENERICAS DE CLIBADIUM L. (COMPOSITAE, HELIANTHEAE)

260 261

NOTAS SOBRE LAS RELACIONES GENERICAS DE Clihadium L. (COMPOSITAE, HELIANTHEAE)

por Jorge E. Arriagada* Resumen

Arriagada, J.E.: Notas sobre lu relaciones genericas deCtibadium L. (Compositae, Helian- theae). Rev. Acad. Colomb. Cienc. 18 (71): 465-468. 1993. ISSN 0370-3908.

La inform acidn recolectada para la rcvisidn del gtaeroClibadium (Compoiitae, Heliantheae), actualmente en deiairollo, ha permitido eitablecer relaciones preliminarei con loi gineros afineiDetmanthodium Bench., Ichthyothere Mart, y Stachycephalum Sch. Bip. con lot que parece formar un grupo natural. Del miimo modo aunque de mane- ra m il diitante, la informaci6n dliponible insinua reladonei de aflnidadei con loi gine- ros Galinsoga R. & P. y Calea L.

Palabraa clave: Compoiitae, Heliantheae, Clibadium, Detmanthodium, Ichthyothe­ re, Stachycephalum.

Abstract

In the coune of a reviiion ofClibadium L. (Compoiitae, Heliantheae) I obtained information that luggeit that thii genus forms a natural group with the related genera Detmanthodium Benth., Ichtyothefe Mart, andStachycephalum Sch. Bip. The genera Ga­ linsoga R. & P. andCalea L. appear to be more distantly related. <

El genero Clibadium L. (Compositae, Helian­ La primera insinuacion de afinidades genericas theae) comprende alrededor de 35 especies arbusti- fue sugerida por Bentham (1873) cuando incluyo a vas distribuidas desde el sur de Mexico hasta el nor- Clibadium conjuntamente conDeamanthodium te de Peru con concentraciones principales en Benth. y Riencourtia Cass, en la subtribu Milleri- Costa Rica, Panama, Colombia y Ecuador (Diaz & inae. La inclusion deClibadium en esa subtribu y Arriagada, 1992). por lo tan to su afinidad generica no fue explicada por el autor. RecientementeClibadium ha sido rela- cionado (Stuessy, 1977) conDeamanthodium * Depart am ento de Botanica, UnWenidad de Concep- Benth, un pequeno genero con ocho especies distri­ ci6n, Caailla 2407, Concepcion, Chile. Direcci6n buidas en Mexico y Centroamerica (Arriagada & actual: Dept, of Botany, The Ohio State University, Stuessy, 1991), con Ichthyothere Mart., con alre­ Columbus, Ohio 43210. U.S.A. dedor de 25 especies (Robinson et al., 1981) distri- 262

REV. ACAD. COLOMB. CIENC. VOL. XVIII. NUMERO 7I.MARZO DE I99S

buidai en Centro y Sudamlrica, y conStachyce­ Los datos basados en flavonoides indican que phalum Sch. Bip, con solo dot especies disyunta- Clibadium e Ichthyothere tienen un perfil bioqui- mente distribuidas en Argentina y Mexico. En una mico mas primitivo que Deamanthodium debido a revision de la clasificacion subtribal de las Helian­ que este ultimo acumula mas mono, di y tri-metil theae, Stuessy (1977) ubico a Clibadium en un derivados (Bohm & Stuessy, 1981). Deamantho­ informal “Grupo 3 ” dentro de la subtribu Mille- dium se relaciona con Clibadium de manera mas rinae conjuntamente conDeamanthodium, Ich­distante que conIchthyothere debido a las dife- thyothere y Stachycephalum. Robinson (1981), sin rencias en los derivados de flavonoides (Bohm & embargo, dividio la tribu Heliantheae intensiva- Stuessy, 1982). Sin embargo, el aislamiento de mente y coloco a Clibadium en la nueva subtribu ichthyothereol y de sus acetatos en algunas especies Clibadiinae, aIchthyothere en la subtribu Melam- de Clibadium, enrelacion con los mismos compues- podinae y a Deamanthodium y Stachycephalum tos obtenidos deIchtyothere, sugiere una cercania como unicoa miembros de la nueva subtribu' Dea- entre estos dos generos e indica una mayor distan- manthodiinae. En el presente estudioClibadium es cia con Deamanthodium el cual carece de estos retenido en la subtribu Milleriinae junto conDee- compuestos (Czerson et al., 1979). manthodium, Ichthyothere y Stachycephalum como generos afineo. La siguiente discusion se desa- El recuento cromosomico apoya las afini­ rrolla en torno a esta perspective. dades entreDeamanthodium, Clibadium e Ichthyo­ there, pero la mayor relacion de afinidad se presen­ Las relaciones existentes entre estos generoa ta entre estos dos ultimos mas que conDeaman­ se basan en escasa y disperse information obtenida. thodium. Clibadium es conocido como n « 16 a partir de estudios morfologicos, anatomicos, quf- (Turner & King, 1964; Coleman, 1968, 1982; micos y citologicos. Las relaciones morfologicas Powell & Cuatrecasas, 1970; Stuessy & Arriagada, entre Clibadium e Icnthyothere se basan en las submitido) e Ichthyothere como n - 16 (Turner et caractensticas de las corolas tubulosas de las flores al, 1979) y ca. 33 (Coleman, 1970), este ultimo marginales, en las anteras pequenas y angostas con puede ser considerado como un tetraploide infra- aplndices agudos (Blake, 1917) y en la relativa generico de x - 16 y podria, por supuesto, ser tam­ agregacion de cabezuelas. Strachycephalum tam ­ bien un octaploide de un numero base ancestral de bien presents agregacion de cabezuelas similar a la x - 8. Deamanthodium es conocido como n - 18 observada en algunas especies deClibadium, pero (Keil & Stuessy, 1977) y n - 17 (Ralston et al., el habito y la morfologia general son similares a las 1989). Estos numeros cromosomicos problamente observadas en Deamanthodium, razon por la cual sean el resultado de un aumento aneuploide del las afinidades entreDeamanthodium y Stachyce­ numero basico x - 16 encontrado enClibadium. phalum no han sido cuestionadas (Robinson, 1981). Deamanthodium tiene una peculiar bractea que Estudios recientes de la tribu Heliantheae recubre totalmente al ovario, la cual lo distingue de basados en los sitios de restriction del ADN cloro- los otros tres generos; sin embargo, el ordenamien- plastidial (Kim et al., 1989) sugieren una relacion to y agregacion de sus capitulos, lo asemejan con entre Deamanthodium y Galineoga. Aunque la algunas especies de la section "Gloraerata” de muestra generica analizada es pequena y los resul- Clibadium (Arriagada, en prep.). tados son aun preliminares, esta relacion coincide con la sugerida por Robinson (1981), basada en el Estudios comparativos de la anatomia de los color y arreglo de los conductos resiniferos. Debi­ frutos de Clibadium, Deamanthodium e Ichtyo- do a la cercana relacion morfologica y posicidn there indican una gran similitud en la disposition taxonomies deDeamanthodium y Clibadium, es de las capas de la epidermis, hipodermis, fitomela- posible asumir la misma relacion entreClibadium nina y fibras. (Stuessy & Liu, 1984). La similitud y Galineoga. El posible vinculo entre estos. dos g£- en la estructura celular de la superficie externa de neros se apoya en el numero cromosomico. Todas la capa de fitomelanina enClibadium e Ichthyo­ las especies de Clibadium que han sido examinadas there sugiere una relation mas proxima que con hasta la fecha tienen un numero cromosomico de Deamanthodium. Clibadium e Ichthyothere com- n “ 16. Este numero es tambien frecuente entre parten una superficie tuberculada mientras que especies de Galineoga, donde tambien han sido Deamanthodium presenta una capa de fitomelanina registrados n « 8, 24 y 32 (Canne, 1983; Robinson totalmente lisa. La similitud en la estructura del et al., 1981). Lo que parece debilitar esta relacion pericarpo hace muy dificil determinar las exactas es la condicion del habito. Mientras todos los relaciones entre estos tres generos. Debido a la miembros de Clibadium son tipicamente arbusti- carencia de estudios evolutivos detallados sobre vos, la condicion herbacea prevalece entre las espe­ estos generos (sin embargo, estudios sobreCliba­ cies de Galineoga (Canne, 1977). Sin embargo, no dium y Deamanthodium estan actualmente en desa- existe en la fecha unanimidad con respecto a la rrollo), resulta muy dificil determinar la direction condicion herbacea como caracter primitivo o deri- evolutiva de estos caracteres. Solamente con base vado. Posiblemente el prototipo ancestral para toda en la estructura mas simple del pericarpo se puede la tribu Heliantheae sea herbaceo (Stuessy, 1977). especular que Clibadium e Ichthyothere parecen ser mas primitives al compararlos con Deaman­ Tambien es importanle considerar otros as- thodium. pectos morfologicos que relacionan a Clibadium 263

ARRJAGADA.J.C.: RELACIONES GENERICAS PE CLIBADIUM (COMPOSITAE. HELIANTHEAE).

con Galintoga y que al miamo tiempo, permiten Las observaciones aqui presented as revelan ubicar a eate ultimo como un lejano ancestro para afinidades entreClibadium, Ichthyothere, Detman­ toda la subtribu Milleriinae. Las floras marginalea thodium y Strachycephalum, al tiempo que sugie­ ordenadaz en 1-3 series relacionanGalintoga a con ren relaciones con los generosGalintoga y Calea. la seccion Eggertia (Arriagada en prep.) deCliba­ S61o cuando los demaa gdneroa de las Heliantheae dium, pero asi mismo la presencia de ligulaa en aean e studied os, se podra determinar cuan cercanas Galintoga puede ser interpretada como una condi­ eon estas afinidades. cion primitive. Las flores del disco hermafroditao y fertiles en Galineoga, son funcionalmento mas- culinas por reduccion (ovarios esteriles) en todas las especies de Clibadium. La base sagitada de las BibUograffa anteras en Galineoga parece representor una carac- terfstica primitivo. La condlcidn subaagitada de las Arriagada, J.E. Si T.F. Stuessy. 1990. A new specks and anteras observada enClibadium sugiere la diraccio- subgenus of Daimanthodhtm (Compositae, Hellan- nalidad de este caracter, el que cambiaa integro y these) from Southern Mexico. Brittonia 42: 283-285. obtuso en Ichthyothere y Detmanthodium, respec- tivamente. Bentham, C. 1873. Notes on the Classification, History, and Geographical Distribution of Compositae. J . La presencia de pappus en todas las especies Linn. Soc. Lon. Bot. 13:431-433. de Galineoga represents una condicion ancestral perdida en todos los miembros del informal "Gru- Blahe, S.F. 1917. Notes on the systematic position ofCliba- po 3" de la subtribu Milleriinae. La condiddn dium, with descriptions of some new species. Conn. paleacea de Galintoga tiene su contraparte solo en Gray Herb. 52: 1-8. la seccidn Eggersia deClibadium (las paleas en las otras secciones son reducidas y comunmente pre­ Bohra, BA. & T.F. Stnessy. 1981. Flavonol derivatives of senter en las floras marginalea). La misma condi- the genus Clibadium (Compositae). Phytochemistry d6n paleacea se encuentra en todas las especies de 20: 1053-1053. Ichthyothere y, esta ausente en el g£neroDetman­ thodium el cual ha sido colocado cerca deGalin­ ------. Sc ------. 1981. Flavonol derivatives a t Dei- toga con base en estudios del ADN cloroplastidlal mmthodium (Compositae). Phytochemistry 20: (Kim et al., 1989). 1573-1575.

Otro genero que presents afinidades y que ------. & ------. 1983. Flavonol derivatives of Ith- merece ser considerado como posible ancestro de th y o th m terminalis. Phytochemistry 21: 2761-2763. Clibadium y consecuentemente de los demos miem­ bros del informal “Grupo 3" de la subtribu Mille­ Canne, J.M. 1977. A revision of the genusGalimoga (Com­ riinae, es el genero Calea L. (subtribu Galinsogi- positae Heliantheae). Rhodora 79: 319-389. nae), subgeneroEucalea Benth. & Hook. Las espe­ cies en este subgenero se asemejan a las especies de . . 1983. Cytologies! and morphological observa­ Clibadium por tener cabezuelas pequeiiss y sums- tions in Galimoga and related genera (Asteraceae). rosas organized as en co rim boa apretados. La mayo- Rhodora 85: 355-366. rfa de estas especies son arbustos y algunas pocas son hierbas perennes, y se distribuyen en Mexico y Coleman, J.R . 1968. Chromosome numbers in some Brazi­ en la zona tropical de Suramerica. (Canne, 1983; lian Compositae. Rhodora 70: 228-240. Robinson & Greenman, 1896; Wussow et al., 1985), coincidiendo en gran parte con la distribu- ..... — . 1970. Additional chromosome numbers in Bra­ cion deClibadium. zilian Compoiitae. Rhodora 72: 94-99.

El involucro en Calea con varias series de fila- — . 1982. Chromosome numbers of Angiosperms rias contrasts con la unica serie observada en cast collected in the state of Sao Paulo. Rev. Brasil. Genet. todas las especies deClibadium, con la excepci6n 3: 353-349. de la seccionEggersia (Arriagada, en prep.). Las flo­ ras marginalea liguladas y las floras fertiles del disco Czcrsen, H., F. Bohlmann, T.F. Stuessy & N.H. Fischer. sugieren direccionalldad de estos caracteres hacia 1979. Sesquiterpenoid and scetylenic constituents Clibadium, y tambien hacia Detmanthodium e of seven Clibadium species. Phytochemistry 18: 257- Ichthyothere. Ademas, la presencia de papus o es- 260. camas y las anteras cortamente sagitadas en la base, son algunas de las condiciones que permiten consi- Dfaz-Picdrahlta, S. Sc J.E. Arriagada. 1992. Una nueva espe- derar este genero como posible ancestro deCliba­ cie de Clibadium (Compositae) para Colombia. Rev. dium. La idea de proximidad generica se ve refor- Acad. Colomb. Cienc. 18: 301-303. zada ademas por el numero cromosomico, que en Calea ha sido registrado como n «* 16, 18, 19 y 24 Kcil, D J. tc T.F. Stuessy. 1977. Chromosome counts in lo cual indica cierta afinidad conClibadium cuyo Compositae from Mexico and the . numero cromosomico es n " 16. Amer.J. Bot. 64: 795-798. 264

REV. ACAD. COLOMB. CIENC. VOL. VD1. NUMERO 71, MARZO DE 1995

Kim, K. J., R. Jansen, BX. Turner, T. Mabry Be G. Netom. Stnessy, T.F. 1977. Heliantheae - systematic review. 1989. Phylogenetic Implication of chloroplaat DNA In: Hcywood, V.H.,JJI. Harborne L BX. Turner rcitrlctlon ilte variation In tbe tribe Heliantheae (eds.) The Biology and Chemistry of the Compositae. (Aateraceae). Amer.J. Bot. 76: 252. Academic Press. London, pp: 621-671.

Powell, A.M. 3c J . Cuatrecasas. 1970. Chromosome numbers ————— . Sc H. -Y. Liu. 1985. Anatomy of the pericarp of In Compositae: Colombian and Vcnesuclan species. Clibadium, Detmanthodium andIchthyothere (Com- Ann. Missouri Bot. Card. 55: 574-579. positae, Heliantbcae) and systematica implications. Rhodora 85: 213-227. Roblaaon, BX. Be JM. Grccnmaa. 1896. Revision of the Turner, BX. Be RBi. EC lag. 1964. Chromosome numbers In Mexican and Central American species of the genua the Compositae. VIH. Mexican and Central American Calea. Proc. Amcr. Acad. Arts Sci. 32:20-50, species. Southwest. Nat. 27-39.

Robinson, H. 1981. A revision o f the tribal and sub tribal ------, J . Bacon, L. Urbatsch Be B. Simpson. 1979. limits of the Heliantheae (Asteraceae). Smithsonian Chromosome numbers In South American Com­ Contr. Bot. 51. positae. Amer.J. Bot. 66: 173-178. ------., AJd. Powell, RX9. KingIs J J . Weedln. 1981. Wesson, J R ., L E . Urbatsch, Be G A , Sullivan. 1985. Calea Chromosome numbers In Compositae, XU: Helian­ (Asteraceae) In Mexico, Central America, and Jamai­ theae. Smithsonian Contr. Bot. 52. ca. Syst. Bot. 10:241-267. APPENDIX D

CHROMOSOME COUNTS IN CLIBADIUM L. (COMPOSITAE, HELIANTHEAE) FROM LATIN AMERICA

265 266

Chromosome counts in Clibadium (Compositae, Heliantheae) from Latin America

T o d F. St u e ssy a n d Jo r g e E. A r r ia g a d a

Stuessy, Tod F. & Jorge E. Arriagada (Department of Plant Biology, The Ohio State University, Columbus, OH 43210, U.S.A.). Chromosome counts inCli­ badium (Compositae, Heliantheae) from Latin America. Brittonia 45: 172-176. 1993.—New meiotic chromosome counts are reported from 86 populations from Mexico to Ecuador of 13 species ofClibadium (Compositae, Heliantheae). These, plus previous reports, yield a total of 120 counts from 20 of the approximately 39 species recognized in the genus (Arriagada, in prep.). All counts aren = 16 with a few fragments or B-chromosomes observed sporadically in some popula­ tions. One population, containing what appear morphologically to be interspecific hybrids between C.mexiae and C.microcephalum, showed one tetravalent and fragments. Species in both historical sections Clibadium( andTrixidium, created by Candolle and maintained by Schulz) and from the five sections to be proposed in a new classification of the genus (Arriagada, in prep.) have now been counted. The uniformity of chromosome numbers withinClibadium is correlated with observed allopatric distribution of closely related taxa. Key words: Compositae, Heliantheae,Clibadium, Latin America, chromosomes, allopatric speciation.

Clibadium L. is a genus of approximately 1981; Turner & King, 1964). These studies, 39 species found throughout Latin America based primarily on meiotic bud material, (Arriagada, in prep.). Its center of distri­ have shown numbers of n ~ 16, ca. 16, 16 bution is in Central America and there are + frags., 16-17, ca. 17, 17-20, and 24. Be­ particular concentrations in Costa Rica, Co ­ cause chromosome numbers are useful in lombia, and Ecuador. Taxonomically there understanding the reproductive behavior of has never been a comprehensive m o n o ­ populations and hence the taxonomic limits graph, and the genus is in need of systematic of species (Stuessy, 1990), and because two attention. The earlier studies of Candolle chromosomal levels have been reported in (1836), Schulz (1912), and Blake (1917) the genus previously, it seemed worthwhile helped to provide an initial' understanding to carry out additional surveys within Cli­ of the group, but comprehensive investi­ badium. gations are now underway in the laboratory Because most of the species of Clibadium of the authors to provide syntheses of re­ have been shown to have the same chro­ lationships. In the course of expeditions to mosome number (n = 16), the opportunity collect material for morphological and also exists to examine chromosome number chemical work, material for cytological and distribution in the context of isolating studies also was obtained. mechanisms in the Heliantheae, as suggest­ There have been several chromosome ed by Sundberg and Stuessy (1990). The ba­ counts reported previously for taxa of Cli­ sic idea is that closely related species with badium (Coleman, 1968, 1982; Powell & the same chromosome number tend to have Cuatrecasas, 1970; Powell & King, 1969a, allopatric distributional ranges. 1969b; Solbriget al., 1972; Robinson et al.. The purposes of this paper, therefore, are

Brittonia. 45(2). 1993. pp. 172-176. ISSUED: 24 May 1993 © 1993, by The New York Botanical Carden. Bronx. NY 10458-5126 267

1993| STUESSY & ARRIAGADA: CLIBADIUM

T a b l e I P r e v io u s c h r o m o s o m e c o u n t s in Clibadium. M o s t n - 16 w it h m in o r v a r ia t io n s a s n o t e d

Number o f populations Tftjion examined Referenced) C anceps Grccnm. 2 Robinson et al. (1981); Solbrig et al. (1972). C. arboreum Donn. Sm. 1 Solbrig et al. (1972). C. armanii (Balbis) Sch. Bip. 4* Coleman (1968); Robinson et al. (1981). C. erosum DC. 4 Powell and King (1969a). C. glomeratum Greenm. 2 Robinson et al. (1981). C. leiocarpum Steetz 2* Robinson et al. (1981). C. microcephalum Blake t Robinson et al. (1981). C. eggersii Hieron. 1 Solbrig et al. (1972; as C. polygynum Blake). C. rotundifoiium DC. t Coleman (1982). C. surinamense L. 15* Turner and King (1964); Powell and King (1969b; as C. viltosum Benth.); Powell and Cuatrecasas (1970); Solbrig et al. (1972); Robinson et al. (1981). C. terebinthinaceum (Sw.) DC. subsp. I Powell and Cuatrecasas (1970). colontbiense Cuatr. * See text for discussion of the original report ofh “> 24 by Coleman (1968). k One n ~ ca. 17 count was also reported. * Four counts have been reported as it ■ ca. 16 (Solbrig et al.. 1972), n ” 16-17 (Robinson et al., 1981),n ** 16 + frag. (Robinson et al., 1981), andn — 17-20 (Robinson et al., 1981).

to: (1) survey additional taxa of Clibadium Although most of the counts clearly show for meiotic chromosome numbers; (2) relate 16 bivalents in meiosis, a few variations the variation in chromosome number to the were encountered. The geographically wide- developing taxonomy of the genus (Arria- ranging C. surinamense ordinarily has n -* gada, in prep.); and (3) correlate cytological 16 throughout its distribution, but 16 II + data with known patterns of geographic dis­ 1 - 4 fragments have been recorded as well tribution to determine the role of chromo­ as 16 II + 5 B ’s CSF 5743). Five B-chro- some number and geography as isolating mosomes also were encountered in one pop­ mechanisms in the group. ulation of C. trianae (SF 5623), but this is the only collection (out of six) with such Materials and Methods variations and is not characteristic of the species. Four fragments were seen in one Young dowering buds were collected in population of C. mexiae (SPJ4969) as well the field and preserved in modified Ca m o y ’s as in one of C. microcephalum (SPJ 4960). fixative (4 chloroform, 3 absolute ethanol, The significance of fragments is uncertain, 1 glacial acetic acid). In the laboratory, the but based on morphology it does seem likely buds were transferred to 70 % ethanol and that hybridization between the two species stored under refrigeration. Conventional is occurring in and around Puyo, Ecuador. acetocarmine squash techniques’were used These fragments ma y relate to meiotic ab­ and meiotic figures were documented by normalities in hybrid individuals. In this camera lucida. regard, SPJ 4978 is likely to be a hybrid that has n =■ 14 II + 1 IV + l fragment and Results is found in the same geographic area. Eighty-six meiotic counts are reported for Discussion 13 species of Clibadium (Appendix) of which eight are documented for the first time: C. Although Clibadium has never been co m ­ asperum, C. cordatum. C. glabrescens, C.prehensively revised taxonomically, a sur­ laxum. C. mexiae, C. pentaneuron, C. rhy-prisingly large number of chromosome tidophyllum, and C. trianae. All counts are counts have been reported previously in the n - 16. genus (Table I). Eleven species have been 268

BRITTONIA

counted from 34 populations and most re- Schulz (1912) recognized two sections in ports have been n = 16 with a few variations Clibadium. Section Clibadium (Eucliba- similar to those reported here. dium DC.) contained the majority of the The original report for C. armanii was n species, i.e., all those characterized by a sin­ = 24 (Coleman, 1968: 232), with the state­ gle outer series of carpellate florets and with­ ment, “The haploid complement of C. ar- out pales subtending the disc florets. Species mani [sic] revealed a dimorphism in regard of section Trixidium DC., on the other hand, to size with 16 larger and 8 decidedly small­ had several series of carpellate florets and er chromosomes. ... The possibility that the pales subtending the disc florets. Despite 8 smaller chromosomes are B chromosomes 120 counts from 20 species of the genus, can not as yet be excluded.” In a subsequent most counts are from species of section Cli­ study of new material of C. rotundifolium badium except for two from C. eggersii of (a name believed to be a synonym of C. section Trixidium. The five sections pro­ armanii), Coleman (1982: 539) reported posed in the new classification of Clibadium clearly n = 16 with the comment, “The ab­ (Arriagada, in prep.) have now also been sence of such smaller chromosomes in the sampled, and all taxa examined to date have material studied in the present investigation n = 16. It seems unlikely that further sam­ supports the suggested supernumerary na­ pling will reveal other numbers in the genus. ture of those chromosomes.” The conclu­ Because of the uniformity of chromo­ sion at that time, therefore, was that all spe­ some number within Clibadium, an oppor­ cies of the genus are known as n = 16, tunity exists to test the hypothesis of Sund- including C. armanii. berg and Stuessy (1990), who stated that all Based on the incorrect n — 24 count in other factors being equal, closely related taxa C. armanii, showing 16 larger bivalents and with the same chromosome number will 8 smaller or even probably “B chromo­ tend to have allopatric distributional rang­ somes,” Coleman (1968) suggested that Cli­ es. Certainly the general tendency in closely badium is based on* = 8, an opinion echoed related species of Clibadium is for allopatric by Robinson et al. (1981). These latter distributions, for example: (1) C. eggersii workers speculated that the n = 24 count Hieron. and C. divaricatum Blake, both in could have arisen by hybridization between section Trixidium, with the former in Gua­ n = 16 and n — 32 cytotypes, but the latter temala, Honduras, Nicaragua, Costa Rica, number has not yet been reported in the Panama, Colombia, Ecuador, and Peru (in genus. The later report (Coleman, J982) of the departments of Amazonas, Loreto, and new material of the same species as n = 16, San Martin), and the latter with a range re­ however, causes reinterpretation of the base stricted to Dept. Huanuco, Peru; (2) C. rhy- number hypothesis. tidophyllum Diels and C. sprucei Blake, both Desmanthodium Benth. and Ichthyothere found only in Ecuador with the former in Mart, are considered here the two genera the province of Pichincha, and the latter most closely related to Clibadium (Stuessy, endemic to the slopes of Volcan Tungura- 1977). These are known chromosomally as hua (Prov. of Tungurahua); (3) C. remoti- n = 18 in the former (Keil & Stuessy, 1977; jlorum O. E. Schulz and C. surinamense L., one recent report by Ralston et al., 1989, is the latter being the species in the genus with n — 17 for D. fruticosum) and n = 16 and the broadest distribution ranging from ca. 33 in the latter (Coleman, 1970; Turner Honduras, through Central America and the et al., 1979). If these counts are correct, then Caribbean to Venezuela, Brazil, Colombia, the n = ca. 33 report for one species of Ich­ and Ecuador (mainly in the northern prov­ thyothere may be viewed as an infrageneric inces of Carchi and Pichincha), whereas its tetraploid from an x - 16 base. The base close relative C. remotijlorum shows a more number for Clibadium, therefore, would be southern distribution in Argentina, Bolivia, most probably x = 16. This level could Ecuador (mainly in the southern provinces be tetraploid based on ancestral x- = 8, but of Caiiar, Chimborazo, Cotopaxi, and Tun­ no evidence is available to support that gurahua), and Peru. Other closely related hypothesis. species pairs, for which chromosome counts 19931 STUESSY A ARRIAGADA: CLIBADIUM

are not yet available, include: (1) C. gran- Candolle. A. P. de. 1836. Prodromus systematis na- difolium Blake and C. pacificum Cuatr., with turalis regni vegetabilis 5: 505-506. Treutte! and Wurtz. Paris. the former in Costa Rica and Panama, and Coleman, J. R. • 968. Chromosome numbers in some the latter restricted to the provinces of An- Brazilian Compositae. Rhodora 70: 228-240. tioquia and Boyaca, in Colombia and (2) C. . 1970. Additional chromosome numbers in frontinoense Diaz & Arriagada and C. za- Brazilian Compositae. Rhodora 72: 94-99. —— . 1982. Chromosome numbers of angiosperms rucchii H. Robins., both distributed in the collected in the state of S3o Paulo. Revista BrasiL region of Frontino, Colombia, but in dif­ Genet. 3: 533-549. ferent phytogeographic zones, including dif­ K e l l ,D . J . & T.F. Stuessy. 1977. Chromosome counts ferent elevations and slopes in the Cordill­ in Compositae from Mexico and the United States. era Occidental of Colombia. Amer. J. Bol 64: 795—798. Powell, A. M. & J. Cuatrecasas. 1970. Chromosome O n e case of apparent parapatric overlap­ numbers in the Compositae: Colombian and Ven­ ping has been observed between two spe­ ezuelan species. Ann. Missouri Bot. Gard. 57:374- cies: C. mexiae Blake and C. microcepha­ ' 379. lum Blake. Morphological intermediates and &R.M . King. 1969a. Chromosome numbers in the Compositae: West Indian species. Sida 3: meiotic chromosomal irregularities (dis­ 319-320. cussed earlier) suggest the occurrence of hy­ ■ '■■■■ & ------. 1969b. Chromosome numbers in bridization between the two taxa. Both spe­ the Compositae: Colombian species. Amer. J. Bot. cies are restricted to a few provinces in 56: 116-121. Ecuador and are largely separated geograph­ Ralston, B., G. Nesom & B. L. Turner. 1989. Doc­ umented plant chromosome numbers 1989. 1. ically. Clibadium mexiae is found in the Chromosome numbers in Mexican Asteraceae with provinces of Napo-Pastaza and Tungura- special reference to the tribe Tageteae. Sida 13:359- hua, and C. microcephalum occurs in San- 368. tiago-Zamora, Canar, and Napo-Pastaza. In Robinson, K , A. M. Powell, R. M. King& J. F. Wee- dln. 1981. Chromosome numbers in Compositae, and around Puyo (Prov. Napo-Pastaza), XII: Heliantheae. Smithsonian Contr. Bot. 52: 1- however, where a slight overlap occurs in 28. the distribution of these two species, hy­ Schulz, O. E. 1912. BeitrSge zur Kenntnis der Gat- bridization has been suspected. The zone of tung Clibadium. Bot. Jahrb. Syst. 46: 613-628. contact ma y have been accentuated by roads Solbrig, O. T., D. W. Kyhos, M. Powell & P. H. Raven. 1972. Chromosome numbers in Compositae VIII: and other hum a n activities in the region. Heliantheae. Amer. J. Bot. 59: 869-878. Stnessy, T.F. 1977. Heliantheae—systematic review. Acknowledgments Pages 621-671. In: V. H. Heywood et al., editors. The biology and chemistry of the Compositae. Ac­ Appreciation is expressed to: NSF- for ademic Press, London. support to the senior author under Grants . 1990. Plant taxonomy: The systematic eval­ DEB-75-20819 and INT-76-84454; Na­ uation of comparative data. Columbia Univ. Press, tional Geographic Society for support of field New York. Sandberg, S. D. & T. F. Stuessy. 1990. Isolating collections in Panama; and V. A. Funk, R. mechanisms and implications for modes of sped- C. Hartman, R. Jansen, G. Nesom, and I. ation in Heliantheae (Compositae). PI. Syst. Evol. Padilla for help with field studies. [suppl. 4], 77-97. Turner, B. L~, J. Bacon, L. Urbatsch& B. Simpson. Literature Cited 1979. Chromosome numbers in South American Compositae. Amer. J. Bot. 66:173-178. Blake, S. F. 1917. Notes on the systematic position & R. M. King. 1964. Chromosome numbers of Clibadium with descriptions of some new spe­ in the Compositae. VIII. Mexican and Central cies. Contr. Gray Herb. 52: 1-8. American species. Southw. Naturalist 9: 27-29. 27 0

______BRITTONIA [VOL. 45

Appendix

T axa a n d vo u ch ers f o r n ew ch r o m o so m e c o u n t s in C libadium All n — 16 with variations as noted. Vouchers de­ SF 5705; above Medellin on road to Rio Negro posited at OS. H ” Hartman; LJ - La Duke& Jansen; and Santa Elena,SF 5707; 6 km NW of Santa SF “ Stuessy & Funk; SPJ ■ Stuessy, Padilla & Jansen. Elena, SF 5708. Boyaca: 20 km N of Arcabuco, • - first count for the taxon. SF 5591; 4 km NE of Moniquira. SF 5599; 7 km S of Chinavita, SF 5641; 7 km S of Tibara, SF •C asperum (Aubl.) DC. 5640; 9 km WNW of Guateque, S F 5641. Caldas: ECUADOR. Pichincha: 4 km W of Santo Do­ 31 km W ofTolima-Caldas state line,SF 5697; 2 mingo de Los Colorado*,SPJ 4902; ca. 24 km W km N of Neira, SF 5698; 19 km N of Aranzazu, of Santo Domingo on road to ElCarmen , SPJ SF 5700; 38 km N of Aranzazu, SF 5703; 18 km 4905. PANAMA. Darien: Rio Chucunaque, ca. 6 S of Supia, SF 5713; 4 km SW of Rio Surio, SF mi NNW of Yaviza, H 3876. 5715 (+1 frag.). Cauca: 70 km S of Popayan, SF °C. cordatum Cuatr. ■ 5744b; 17 km NE of Popayan, SF 5747; 49 .km COLOMBIA. Valle: 50 km N of Rio Digua, SF SW of Popayan, SF 5746 (+2 frags.). Cundlna- 5725; 2 km W ofQueremal, SF 5737; 6 km W of marca: 24 km NE of Fusagasuga on road to Bo­ Queremal on gravel road to Buenaventura,SF gota, SF 5510; 15 km SW ofSubia, SF5511; 18 5742. km NE of Fusagasuga, SF 5517b (+1 frag.); 10 C. eggersii Hieron. km S of Arbelaez, SF 5519; 2 km E of jet paved ECUADOR. Manabi: 42.8 km SW of El Carmen, rd from Rafael Reyes to La Mesa and dirt road to SPJ 4911. Napo-Pastaza: 34.2 km E of Veracruz, El Colegio, SF 5524; 4km from Pacho, SF 5571; SPJ 4974, 4975 (ca. 16). 2 km SE of Caquiza, SF 5643; 5 km SE of Que- C. erosum DC. tame, SF 5645; 24 km SE of Quetame, SF 5646; MEXICO. Veracruz: 5.7 mi N of road to Xochiapa 18 km NW of Villeta, SF 5666. Meta: 8 km E of on Hwy 185, U 404 (ca. 16). Villavicencio, SF 5649. Narifio: 3 km S of Cha- *C. glabrescens S. F. Blake chaqui, SF 5756. Tolima: 17 km W of Mariquita, COLOMBIA. Santander: 15 km NE o f Bucara- SF 5670. Santander: 38 km NE ofSanta Ana,SF manga on road to Pamplona,SF 5606. 5601; 1 km S of Los Curos, SF 5602; overlooking ®C laxum S. F. Blake city of Bucaramanga, SF 5603; 15 km NE of Bu- ECUADOR. Chimborazo: N edge o f Bucay, SPJ caramanga, S F 5604. Valle: 17 km W of Buga, SF 4942. 5719; 42 km W of Buga, SF 5723; 13 km S of C. leiocarpum Steetz Buga, SF 5727; 2 km W of La Treinta, SF 5732; PANAMA. Chlriqui: road from Alto Quiel to Cer- 4 km W of Queremal, SF 5743 (+5 B’s). EC­ ro Punta, 2-3 mi W of Cerro Horqueta, H 3917; UADOR. Gnayas: 7 km SE of road to Guayaquil 17 km E of Rio Sereno on road from California, and Milagro on road to Bucay,SPJ 4935 (+3 H 3920 (ca. 16). frags.), 4936. Manabi: 13 km E of Alajuela, SPJ *C. mexiae S. F. Blake 4919. Napo-Pastaza: 2 km E of Puyo, SPJ 4969; ECUADOR. Napo-Pastaza: 2.7 km E of Puyo on 17 km S of Veracruz, SPJ 4971 (ca. 16); 34.2 km road to Veracruz,SPJ 4969 (+4 frags.); Village of E of Veracruz, SPJ 4977; 43 km N of Puyo, SPJ Topo near Rio Topo, SPJ 4987. 4982; Tena, SPJ 4984; N edge of Archidona,SPJ C. microcephalum S. F. Blake 4985. Tungurahua: 0.5 km of bridge over Rio ECUADOR. Napo-Pastaza: 0.5 km W of Mere on Blanco.SPJ4954. PANAMA. Chlriqui: 2 km NW road to Banos,SPJ 4965; 17 km N of Puyo on of Potrerillo Arriba. H 3919; 12 km E of Rio Sere­ road to Tena.SPJ 4978 (14 I I .+ 1 IV + 1 frag.). no, H 3921; 19 km S of El Gato del Volcan, H Tungttrahua: 11.7 km E of Rio Verde on road to 3924; 3 km SE ofSanta Marta, H 3925; 15 km Puyo, SPJ 4960 (+4 frag.). ESE of Concepcion,H 3926 (ca. 16); Los Santos, •C. pentaneuron S. F. Blake 21 km NE ofTonosi, H 3943. Veraguas: ca. 4 km COLOMBIA. Anfioquia: 3 km SE o f Santa Elena NW of Santiago,H 3895; 5 km NNW of San Fran­ and Rio Napo,SF 5709. Boyaca: 9 km N of Ar- cisco, H 3933. cabuco on road to Bambita,SF 5595. °C. trianae (Hieron.) S. F. Blake •C. rhytidophyllum Diels COLOMBIA. Cundinamarca: 8 km SW on road ECUADOR. Pichincha: 21 km N o f Nono, SPJ to El Salto, SF5S28. Nariflo: ca. 4 km E of Encano, 4863; 28.7 km W of Aloag on road to Santo Do­ SF5770. Putumayo: 1 km W ofSantiago, SF5775. mingo de Los Colorado*,SPJ 4896b. Santander: 24 km SE of Pamplona,SF 5623 (+5 C. surinamense L. B’s); 26 km SW of Berlin, SF 5627. Tolima: 25 COLOMBIA. Antioquia: 4 km S ofSanta Barbara, km W of Palma, SF 5680. APPENDIX E SCATTER DIAGRAM OF REPRESENTATIVE COLLECTIONS OF CLIBADIUM GRANDE. C. GRANDIFOLIUM. AND C. PACIFICUM

271 27 2

40 e © • ® u 8 A *® % 30 ®

H5 © 11 ►J 8 ® t[S(s)[Q]

20 ®

x. x 10 20 30 40 Leaf length (cm)

Fig. 22. Scatter diagram of representative collections of Clibadium grande (black circles), C. grandifolium (hollow circles), and C. pacificum (hollow squares) comparing features previously believed to be diagnostic for separation of taxa. Numbers show number of ray florets in individual heads (C. grande is constant with 6). Types of the three taxa indicated by "t." Leaf measurements were made on largest leaf; ray floret number based on average from three heads. LIST OF REFERENCES

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