revision of the genus clibadium (asteraceae, heliantheae)

Revision of the Genus Clibadium (Asteraceae, Heliantheae)Author(s): Jorge E. ArriagadaReviewed work(s):Source: Brittonia, Vol. 55, No. 3 (Jul. - Aug., 2003), pp. 245-301Published by: Springer on behalf of the New York Botanical Garden PressStable URL: http://www.jstor.org/stable/3218446 .Accessed: 05/10/2012 04:33

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Revision of the genus Clibadium (Asteraceae, Heliantheae)

JORGE E. ARRIAGADA

Arriagada, J. E. (Department of Biological Sciences, St. Cloud State University, St. Cloud, MN 56301-4498, U.S.A.; e-mail: [email protected]). Revi- sion of the genus Clibadium (Asteraceae, Heliantheae). Brittonia 55: 245-301. 2003. Clibadium L. (Asteraceae, Heliantheae) is a genus of 29 species distributed throughout Latin America, from Mexico to Peru, and in the West Indies, with high numbers of species in Costa Rica, Colombia, and Ecuador. Clibadium includes shrubs and small trees; usually with loosely aggregated capitula; herbaceous phyllaries arranged in 1-5 series; receptacles usually paleaceous throughout; corollas of pistillate florets 2-4-lobed; corollas of the staminate florets 4-5- lobed; purple to black anthers; and chromosome numbers all n = 16. Two sections of species previously recognized are here considered as subgenera (subg. Paleata and subg. Clibadium) containing two and four sections, respectively. Clibadium subg. Paleata contains five species distributed in sects. Eggersia (3 spp.) and Trixidium (2 spp.), and subg. Clibadium has 24 species distributed among sects. Clibadium (6 spp.), Glomerata (9 spp.), Grandifolia (5 spp.), and Oswalda (4 spp.).

Key words: Asteraceae, Heliantheae, taxonomy, Clibadium, Paleata, Eggersia, Trixidium, Glomerata, Grandifolia, Oswalda, Mexico, Central America, West In- dies, South America.

Arriagada, J. E. (Department of Biological Sciences, St. Cloud State University, St. Cloud, MN 56301-4498, U.S.A.; e-mail: [email protected]). Revi- sion of the genus Clibadium (Asteraceae, Heliantheae). Brittonia 55: 245-301. 2003. Clibadium L. (Asteraceae, Heliantheae) es un genero distribuido a trav6s de Am6rica Latina, desde M6jico a Peru, y en West Indies, con un gran numero de especies en Costa Rica, Colombia, y Ecuador. Clibadium incluye arbustos y arboles pequefios; generalmente con capitulos laxamente agregados; filarias her- baceas, ordenadas en 1-5 series; receptaculos generalmente palaceos; corolas de las florets pistiladas con 2-4 16bulos; corolas de las florets estaminadas con 4-5 16bulos; anteras purpuras a negras; y numeros cromos6micos todos n = 16. Dos grupos de especies previamente reconocidas son aqui considerados como subg6- neros (subg. Paleata y subg. Clibadium) que tienen dos y cuatro secciones, res- pectivamente. Clibadium subg. Paleata contiene cinco especies distribuidas en sects. Eggersia (3 spp.) y Trixidium (2 spp.), y subg. Clibadium incluye 24 especies distribuidas entre las sects. Clibadium (6 spp.) Glomerata (9 spp.), Gran- difolia (5 spp.), y Oswalda (4 spp.).

Clibadium L. has a broad geographical distribution throughout much of the Neo- tropics and it includes several locally endemic species. The genus extends from southern Mexico throughout Central Amer-

ica, the Caribbean, and tropical South America, and has high numbers of species in Costa Rica, Colombia, and Ecuador. A few species (e.g., C. eggersii Hieron., C. surinamense L., and C. sylvestre (Aubl.)

Brittonia, 55(3), 2003, pp. 245-301. ISSUED: 17 July 2003 ?) 2003, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.

246 BRITTONIA [VOL. 55

Baill.), have broad distributions; others are restricted only to small areas (e.g., C. divaricatum S. F Blake, from Prov. Tarapoto, Peru; C. rhytidophyllum Diels, from Volcain Tungurahua, Ecuador; and C. zarucchii H. Rob., from the western slope of Cordillera Occidental, Colombia). The affinities of Clibadium to other genera have not been examined closely. Stuessy (1977) placed it close to Desmanthodium Benth., Ichthyo- there Mart., Riencourtia Cass., and Stachy- cephalum Sch. Bip.; more recently, Robin- son (1981) placed it close to Lantanopsis C. Wright ex Griseb. and Riencourtia. The genus Clibadium is here defined by a unique combination of characteristics, most of which are also 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; usually with loosely aggregated capitula; herbaceous phyllaries arranged in 1- 5 series; receptacles usually paleaceous throughout; corollas of pistillate florets 2- 4-lobed; corollas of the staminate florets 4- 5-lobed; purple to black anthers; and a chromosome number of n = 16.

In addition to its interesting pattern of distribution and complex generic relationships, Clibadium is important in the neotro- pical flora for three other reasons. First, the species are shrubby, and in some cases small trees, which makes them conspicuous and frequently encountered in many regions; floristic workers, therefore, frequently must deal with plants of this genus. Sec- ond, the drupe-like fruits are used as food by birds (Feinsinger et al. 2182) and might indicate a significant role in relation to wildlife resources and ecology. Third, one species of Clibadium, C. sylvestre, might be a potential source of chemical compounds of medicinal interest, based on its use as a fish-poison by Indian tribes throughout Lat- in America (Arriagada, 1995a; Gorinsky et al., 1973).

The basic systematic relationships of the taxa of Clibadium have been poorly under- stood, to the extent that routine identification has been impossible. Schulz (1912) completed the first taxonomic treatment which included 13 species and two varie-

ties. He divided the genus into two sections: Euclibadium DC. (naked receptacles and villous sterile ovaries of staminate florets) and Trixidium DC. (paleaceous receptacles and pilose sterile ovaries of staminate florets). The studies of Schulz were based on morphological characters obtained from limited herbarium material, mostly depos- ited at Berlin-Dahlem (B). Although helpful, Schulz's work has not been satisfactory for revealing the basic relationships of the taxa involved. The valuable contributions by Blake (1917), Cuatrecasas (1954), Aris- teguieta (1964), Stuessy (1975), and Rob- inson (1979a, 1979b) likewise have not provided a comprehensive treatment of the genus. Moreover, many new names have been added for taxa based on characters that are extremely variable.

Twenty-nine species of Clibadium are here recognized. The two sections previously recognized by de Candolle (1836) and Schulz (1912), are here considered as subgenera Eggersia and Clibadium containing two and four sections, respectively.

Taxonomic History

Clibadium, with a single species, C. surinamense, was published by Linnaeus (1753). De Candolle (1836) described seven new species and one variety and placed those into three sections based on pubescence and texture of cypselae, numbers of series of pistillate florets, and presence or absence of paleae subtending staminate florets. De Candolle (1836) placed Clibadium surinamense L. in sect. Clibadium (as Eu- clibadium) based on its drupe-like cypselae, single series of ray florets, and receptacles without paleae. A single species from the Caribbean, C. erosum, was placed in sect. Trixidium based on its dry pistillate cypselae with villous apices, two series of pistillate florets, and epaleaceous receptacles. De Candolle grouped eight species, mostly from the Caribbean and northern South America, in sect. Oswalda (Cass.) DC., based upon its dry and pubescent cypselae, one series of pistillate florets, and epaleaceous receptacles. Grisebach (1864), as a result of his study of the flora of British West Indian Islands, recognized two more

2003] ARRIAGADA: ASTERACEAE, HELIANTHEAE 247

new species of Clibadium, C. alexandri and C. fragiferum, and placed the Caribbean taxa into two groups: Clibadium (as Eucli- badium), with only one series of pistillate florets, and an unnamed group (correspond- ing to de Candolle's sect. Trixidium), containing species with two series of pistillate florets. A few years later, Hemsley (1881) listed fifteen herbaceous or half-shrubby species of Clibadium indigenous to the Neotropics including C. acuminatum Benth. from Cocos Islands; C. asperum DC. from southern Mexico, Guatemala, and Nicara- gua; and C. leiocarpum Steetz from Costa Rica. A number of new species were added to Clibadium in the early I900s. Based on material collected by Sodiro in subtropical and subandean regions of Ecuador, Hie- ronymus (1901) described C. sodiroi Hie- ron. and C. subsessilifolium Hieron. The number of species in the genus increased to 18 with Greenman's (1903) descriptions of C. anceps Greenm., C. glomeratum Greenm., and C. pittieri Greenm. based on material collected in Costa Rica by Tonduz.

Schulz (1912), in the first revision of the genus, recognized 19 species and two va- rieties. He accepted the 15 previously described species and proposed four new ones: Clibadium armanii 0. E. Schulz from Brazil, C. lehmannianum 0. E. Schulz from Colombia, C. micranthum 0. E. Schulz from Peru, and C. remotiflorum 0. E. Schulz from Brazil and Bolivia. Schulz also recognized two sections based on the presence of paleae subtending staminate florets and pubescence of sterile ovaries: sect. Cli- badium (as Euclibadium) (including de Candolle's sect. Oswalda) with 15 species, characterized by absence of paleae subtending staminate florets and sterile ovaries villous throughout; and sect. Trixidium with four species-C. eggersii Hieron., C. erosum (Sw.) DC., C. fragiferum, and C. terebinthinaceum-characterized by receptacles paleaceous throughout and sterile ovaries villous only at the apex.

Blake (1917) was a major contributor to the systematics of Clibadium. He agreed with Schulz's subdivision of the genus into two sections, and added five new species: C. divaricatum S. F Blake and C. strigillosum S. F Blake, from Peru; C. heterotri-

chum S. F Blake, from Bolivia; C. polygynum S. F Blake from Nicaragua; and C. sprucei S. F Blake, endemic to Volcdn Tun- gurahua in Ecuador. He also transferred Desmanthodium trianae Hieron. to Cliba- dium. A few years later, Blake (1924) described nine additional new taxa including three species from Costa Rica: C. grande S. F Blake, C. grandifolium S. F Blake, and C. schulzii S. F Blake, all from material collected by A. Tonduz. Blake also described Clibadium parviceps S. F Blake from material collected by A. Fendler in Venezuela, and believed that the new species was related closely to C. acuminatum Benth., endemic to the Cocos Islands. Blake (1926) added two new species, this time from Ecuador: C. laxum S. F Blake and C. microcephalum S. F Blake. The genus increased even further when Blake (1937) described C. oligandrum S. F Blake based on material collected by Skutch on Volcdn Zu- nfl, Guatemala. At the same time, he described C. glabrescens S. F Blake from Co- lombia and C. leiocarpum var. strigosum S. F Blake from Costa Rica. One year later, Blake (1938) added another species to the list: C. mexiae S. F Blake from Ecuador. The last of the almost two dozen species of Clibadium described by Blake (1941) was C. sessile S. F Blake, from Prov. Chiriqui, 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 of the specimens were collected by Cuatreca- sas along the eastern and western cordille- ras of Colombia.

Additional comments on Clibadium have been made by floristic workers in different regions of Latin America. Broader distributional ranges of some species of Cliba- dium were reported by Matuda (1950) in his flora of Chiapas; he listed C. arboreum, C. oligandrum, and C. villosum as growing near Volcdn Tacand in southern Mexico. Clibadium heterotrichum, originally restricted to Bolivia, was reported by Cabrera (1962) for Argentina. Aristeguieta (1964) mentioned three species in Venezuela together with one variety described by Ste-


yermark, and he proposed a new species for that country: C. pediculatum Aristeg. Nash and Williams (1976) in the Flora of Gua- temala, cited two for Guatemala (C. arboreum and C. eggersii) and a third one from Chiapas, Mexico (C. leiocarpum).

In recent years Harold Robinson made several important additions of species to Clibadium. He described a new species for Peru: C. vargasianum H. Rob., based on leaf fragments lacking the erect hispid pubescence typical of C. surinamense (Rob- inson, 1979a). More recently, however, Robinson (1992, p. 149) reconsidered C. vargasianum as "an unusually hirsute variant of the widespread C. surinamense." Robinson (1979b) also described C. harlingii H. Rob. and C. manabiense H. Rob. from Ecuador, based on material obtained by G. Harling. In 1988, Robinson described C. rimachii H. Rob. based on one specimen collected by Rimachi from Peru, but he later (1992) relegated it to synonymy under C. divaricatum Blake (Robinson, 1992). Clibadium zarucchii H. Rob., from the region of Frontino, Colombia, was described by Robinson (1988) as a new species based on the most massive congestion of capitula of any member of the genus. Diaz-Piedra- hita and Arriagada (1992) described C. frontinoense S. Diaz & Arriagada from the same general region of C. zarucchii; it is a clearly related taxon also with massive aggregations of capitula. The most recent pub- lication on Clibadium by Robinson (1992) added four new species: C. funkiae H. Rob. from Antioquia, Colombia; and C. alatum H. Rob., C. napoense H. Rob., and C. zakii H. Rob. from Ecuador. This brought to 85 the number of species named in the genus.

Generic Affinities

Stuessy (1977) and Arriagada (1993) have suggested that Clibadium is related to: Desmanthodium, a genus of eight species in Mexico and Central America; Ichthyothere, with about 17 species in Brazil, Colombia, Paraguay, and Venezuela; and to Stachy- cephalum, consisting of two disjunct species in Argentina and Mexico. Clibadium was placed in an informal "Group 3" in the Milleriinae by Stuessy (1977). Robinson

(1981) referred Clibadium to a new subtribe, the Clibadiinae H. Rob.; Ichthyothere to Melampodiinae; and Desmanthodium and Stachycephalum to another new subtribe, the Desmanthodiinae H. Rob. Ac- cording to Karis (1993), the limits between these subtribes (including Galinsoginae) should be further studied because many of the genera will most likely have to be re- classified as to subtribe. In the present study Clibadium is retained in Milleriinae, with Desmanthodium, Ichthyothere, Riencourtia, and Stachycephalum as close relatives.

The relationships among Clibadium, Desmanthodium, Ichthyothere, and Stachy- cephalum have been based on incomplete information from morphology, anatomy, chemistry, and cytology. The proposed close relationship between Clibadium and Ichthyothere is based primarily on tubular corollas in the pistillate florets and relative- ly moderate aggregation of the capitula (Blake, 1917; Stuessy, 1973). Stachyceph- alum, although overall more distant morphologically from Clibadium, has radiate corollas in the pistillate florets and shows a looser aggregation of capitula, similar to those present in some species of sect. Cli- badium. The genus Desmanthodium has a peculiar sac-like bract completely enclosing each ovary, a unique feature that represents a definite morphological gap relative to the other three genera. The general morphology of Desmanthodium, and particularly the arrangement and aggregation of capitula, show close resemblance to some species of Clibadium sect. Glomerata.

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 & Liu, 1983). The similar pattern of the ex- ternal phytomelanin surface in Clibadium and Ichthyothere argues for their close relationship, in contrast to the smooth surface in Desmanthodium. The structure and de- velopment of the pericarp is very similar and suggests evolutionary ties among Cli- badium, Desmanthodium, Ichthyothere (Stuessy & Liu, 1983) as well as, somewhat surprisingly, Galinsoga Ruiz & Pav. (Saienz, 1981). Based on general pericarp


morphology and features of the 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 paren- chyma cells. A relationship between Des- manthodium and Galinsoga, based on color and arrangement of resin ducts in the pericarp, was suggested by Robinson (1981).

Chemical analyses indicate that Cliba- dium has a simpler biochemical profile than Desmanthodium in that the latter accumulates more mono-, di-, and trimethyl deriv- atives (Bohm & Stuessy, 1981). Desman- thodium is somewhat more distant from Clibadium than from Ichthyothere on the basis of flavonoids. The presence of a fish poison (=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 Cliba- dium to Ichthyothere than to Desmantho- dium. Clibadium has n = 16 (Turner & King, 1964; Coleman, 1968, 1982; Powell & Cuatrecasas, 1970; Stuessy & Arriagada, 1993); and Desmanthodium has n = 18 (Keil & Stuessy, 1977; Robinson et al., 1981) and n = 17 (Ralston et al., 1989). These numbers could be viewed as dysploid increases from a base of x 16 in Cliba- dium. Ichthyothere has n 16 and 33 (Coleman, 1970; Turner et al., 1979) with an apparent base of x = 16.

Recent studies on Heliantheae based on chloroplast DNA restriction site analyses (Kim et al., 1989) indicate a close relationship between Desmanthodium and Galin- soga. Although the sampling of species is small and the results are 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 similarities of Desmanthodium and Cliba- dium, one could also assume that a reason- ably close molecular relationship might exist between Clibadium and Galinsoga. In the most recent revision by Canne (1977), Galinsoga was considered to be similar to

Cymophora H. Rob., Sabazia Cass., Tri- carpha Longpre, and Tridax L., and not at all to Clibadium and its relatives. Nonethe- less, in view of the preliminary molecular data, an examination of their possible ties to Clibadium seems in order.

Galinsoga contains about 14 species separated into three sections and distributed mainly in subtropical regions of Mexico, Central America, and South America (Can- ne, 1977). Section Galinsoga and, particularly, one of the four lines of divergence within the section (including G. subdiscoi- dea Cronq., G. mandonii Sch. Bip., and G. parviflora Cav.) might possibly have a link with Clibadium. The species in that group all have very small ray corollas. This particular line of divergence in Galinsoga ap- parently originated in Mexico (Canne, 1977), even though G. mandonii is confined to the Andes of western South America. Chromosome numbers support a possible link between the two genera. The number, n = 16, that characterizes Clibadium, is also frequent among species of Galinsoga in which n = 8, 16, 24, and 32 have been reported (Canne, 1983). Growth habit is discordant; however, all members of Cli- badium are shrubs and those of Galinsoga are herbs.

In my opinion, morphological features suggest that Galinsoga could be viewed as a somewhat distant link for the entire Mil- leriinae. The ray florets arranged in 1-3 series could relate Galinsoga to Clibadium sect. Eggersia; the presence of ligules in the former is a distinct difference. Another difference is the presence of hermaphroditic and fertile disc florets in Galinsoga and only disc florets that are functionally staminate 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. subg. Calea (a member of Galinsoginae). Species in this subgenus resemble those of Cliba- dium in having numerous small capitula arranged in closed thyrses or modified um- bels, and, often, cypselas with a reduced pappus. Most of the species of subg. Calea are shrubs (some are perennial herbs) and are distributed in Mexico and tropical South


America. Involucres with many series of phyllaries in Calea contrasts with the single series observed in all but one section (sect. Glomerata) of subg. Clibadium. Chromo- some numbers reported for Calea are n = 9 and 16. The cytological data give some additional support to the idea of affinities between Calea and Clibadium (also n = 16; Stuessy & Arriagada, 1993).

Morphology and Taxonomic Criteria

The present classification of the species of Clibadium is based mainly on comparative morphology. In Clibadium, some characters show considerable variation (Arria- gada, 1995b). The variation causes difficul- ty in delimitation of species. Also, many structures appear to have undergone parallel changes and reversals during evolution of the genus. 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 cypsela (favored by Spjut, 1994) is used instead of achene despite its traditional usage (e.g., Blake, 1924). All measurements of quanti- tative characters were made on herbarium material. Because shrinkage of some parts occurs with desiccation, reproductive structures were rehydratated with Aerosol OT (10% aqueous) before measurement.

GROWTH HABIT

All species of Clibadium exhibit a shrubby or weakly arborescent habit. Most of the species show a highly branched stem (typically represented by C. surinamense). Some species have arching branched stems but with one stem at the base (e.g., C. anceps, C. glabrescens, C. laxum, C. trianae), others possess a more or less an erect stem (10 cm diam. in C. cordatum and C. rhyti- dophylum), and still others are arching or scandent shrubs (e.g., C. pentaneuron).

LEAVES

Shapes, sizes, margins, and bases of blades are sometimes useful in delimiting species. Blades of most taxa are ovate except in Clibadium leptophyllum, which is easily recognizable by its narrowly lanceolate leaves. Clibadium rhytidophyllum and C. sprucei are likewise distinctive in their consistent lanceolate leaves. The largest leaves, to 50 X 40 cm, are found in C. cordatum, C. frontinoense, C. grandifolium, and C. zarucchii, and the smallest, ca. 3 X 1.5 cm, are found in C. armanii. Most of the species have blades with serrate margins, but C. erosum is distinctly erose. Some species (e.g., C. armanii and C. cordatum) have leaves with cordate bases in contrast to the attenuate condition in most other species of the genus. Most species of Clibadium have unwinged petioles; partial- ly or entirely winged petioles are found in C. laxum (in mature individuals, e.g., Har- ling & Anderson 11525, MO; Jaramillo 6713, MO). Clibadium sessile and C. leptophyllum have no or very short petioles, a feature that helps in delimiting these taxa.

CAPITU LESCENCE

The perspectives on capitulescence structure presented here are based in part on Cronquist's (1977) concepts of organization, arrangement of capitula and branches, and peduncles for the Asteraceae. For sim- plicity of application of terminology, capitula are treated as if they represent single flowers. Most members of the family have a basic cymose capitulescence (Cronquist, 1977), although solitary capitula are also known. The basic cymose capitulescence occurs in two different forms: monochasial and dichasial (Wyatt, 1982; Weberling, 1989). Monochasial capitulescences have a terminal capitulum that opens first, with the other capitula opening in a centrifugal order down the axis. Dichasial capitulescences also mature centrifugally, but they have only three capitula arising from the same node in which the central capitulum opens first, followed by the two lateral ones shortly thereafter.

All species of Clibadium have complex monochasial capitulescences that, depend-


ing on the species, are arranged secondarily into cymes or thyrses. These secondary ar- rangements are sometimes organized further into compound cymose and thyrsoid aggregations. 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 capitula, with capitula sessile or on short peduncles, and secondarily aggregated into large clusters with up to 600 small, loosely arranged (e.g., C. grandifolium) or tightly packed (e.g., C. zarucchii) capitula. The range of aggregation of capitula is illustrated by C. eggersii with capitula loosely aggregated, C. surinamense with capitula moderately aggregated, and C. glabrescens with capitula densely aggregated, and with very densely aggregated capitula (e.g., C. frontinoense and C. zarucchii). Densely aggregated capitulescences also occur in all species Desmanthodium, in particular, in D. congestum Arriagada & Stuessy. The more loose aggregation typical of C. eggersii is also found in many species of Ichthyothere.

Characterization of the total capitulescences of a taxon in Clibadium is compli- cated because there are many levels of organization. Estimation of the total number of capitula is also difficult, particularly because the total number depends on the age of the branch. Many species flower for several months, and the capitulescences contin- ue 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, C. sylvestre has a thyrsoidal capitulescence when young (Nee 7139); it can have paniculiform capitulescences when older (Web- ster 16782). In C. armanii, thyrsoidal capitulescences are found on young specimens (Harley & Castro 11295) and paniculiform ones occur on older plants (Irwin et al. 23772). An even broader range of variation occurs in C. surinamense, in which cymose capitulescences are found in young individuals (Knapp 1267; Maxon & Val- entine 7033) and panicles are observed in older ones (Dwyer 7197, Maxon & Harvey 6505, Stern et al. 720). Occasionally, an in-

termediate or slightly different capitulescence arrangement can be observed in a particular species. For example, some individuals of C. anceps (Dwyer & Correa 8844) and of C. glabrescens (Killip & Cua- trecasas 38862) range from typically capitate to nearly umbellate panicles.

The capitulescence types observed among species of Clibadium do not correlate with the proposed subgeneric and sec- tional classification of the genus. Similar morphological variation in capitulescences has been observed in genera unrelated to Clibadium (Hieracium L., Beaman, 1990; Pappobulus S. F Blake, Panero, 1992; and Simsia Pers., Spooner, 1990). Because of the considerable variation in capitulescences of some species of Clibadium, and because of the obvious cycles of capitulescences evolution within the different taxonomic sections, characters of the capitulescence have not been used in infrageneric classification. However, C. frontinoense, C. laxum, C. trianae, and C. zarucchii have very distinctive capitulescences and these features can be diagnostic for specific recognition.

CAPITULA

Capitula in Clibadium are disciform and heterogamous: peripheral pistillate florets and functionally staminate central florets with white, tubular corollas. This is simple monoecy, one of many breeding systems known to occur in the family (Lowrey & Stuessy, 1982). The numbers of pistillate and staminate florets is very important in recognizing groups of species and for pur- poses of identification.

The number of pistillate florets ranges from 3-5 (e.g., Clibadium anceps, C. glomeratum, C. microcephalum, C. sessile, and C. trianae) to 30-40 (C. divaricatum, C. eggersii, and C. leptophyllum). The range in number of staminate florets is three (C. microcephalum) to 24 (C. leptophyllum).

The number of capitula per capitulescence varies in different taxa, as follows: 24-28 in Clibadium eggersii (Tyson 3422), 60-116 in C. trianae (Stuessy & Funk 5623), 70-170 in C. surinamense (Stuessy & Funk 5510), 75 in C. sessile (Allen


4970), 86-119 in C. anceps (Dwyer & Co- rrea 8844; Stern et al. 1019), 94-106 in C. pentaneuron (Stuessy & Funk 5595), 98- 100 in C. leiocarpum (Allen 1341), 100- 130 in C. glabrescens (Stuessy & Funk 5606), 219 in C. microcephalum (Stuessy & Arriagada 12362), and up to 600 in C. grandifolium (Croat 13372; Lewis et al. 2082). An unusual number of 245 capitula in a cluster was counted in a young individual in C. trianae (Stuessy & Arriagada 12337), but the basic capitulum and capitulescence organization was the same as observed in other specimens.

INVOLUCRES

The numbers of phyllaries, their arrangement into series, and their shapes, textures, and venation are all taxonomically useful in Clibadium. The phyllaries are herbaceous and spreading during anthesis and usually reflex during fruiting (clearly seen in C. manabiense). The numbers and lengths of phyllaries are usually constant within the species. Phyllaries are often pubescent over the upper one-third, with ciliate margins. Phyllaries in most species are organized in a single series; C. anceps, C. glomeratum, and C. trianae, however, have two series. The most common number of phyllaries in the genus is three; C. eggersii and C. leptophyllum have only two (this low number perhaps relating to a larger number of ray florets subtended by paleae). Clibadium frontinoense and C. zarucchii have 4 phyllaries per capitulum. The shapes of phyllaries range from ovate to orbicular with the upper one-third of the margin ciliate and with apex obtuse (C. erosum, C. grandifolium, C. pentaneuron, C. sylvestre, C. 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 tends toward glabrous in others (C. glomeratum, C. grandifolium, C. leiocarpum, C. microcephalum, C. pentaneuron, C. sessile, C. terebinthinaceum, and C. trianae).

PALEAE

The term "palea" is here used for those bracts subtending any floret, and the term

"phyllary" for those bracts forming the involucre (Briquet, 1917; Cabrera, 1978; Funk, 1982; Stuessy, 1977). In Clibadium, the pistillate florets are subtended by broad paleae, which often resemble phyllaries but differ in having fewer veins and less pubescence. In species with more than one series of pistillate florets, there is a gradual change, a reduction in size and pubescence of the paleae from the outermost to the innermost pistillate florets. Disc florets in some species are also subtended by paleae (paleae). Paleae subtending the innermost pistillate florets are similar to those of subtending the staminate florets. Such a tran- sition can be seen clearly in Clibadium eggersii, with 5-7 series of pistillate florets.

Taxonomically important variations in Clibadium occur in the paleae. Paleae with seven veins subtend pistillate florets in all members of sect. Oswalda and in some members of sect. Glomerata (C. anceps, C. glomeraitum, C. trianae). Section Cliba- dium and subg. Paleata have paleae with five veins; in sect. Grandifolia paleae are 4- veined. Staminate floret paleae are more uniform than those of pistillate florets. All members of subg. Paleata, as well as all taxa in subg. Clibadium sect. Oswalda, have 3-veined paleae. Taxa of sect. Gran- difolia (except C. grandifolium) have 2- veined paleae. All species but one (C. laxum) in sect. Clibadium have epaleaceous receptacles. In C. laxum, paleae of staminate florets have three veins. Clibadium frontinoense, C. rhytidophyllum, C. sprucei, and C. zarucchii of sect. Glomerata are the only species in the genus with only one conspicuous vein.

COROLLAS OF PISTILLATE FLORETS

Ten species have 4-lobed corollas. This condition is found among some members of subg. Paleata as well as among most members of subg. Clibadium sect. Glomerata and C. arborea in sect. Clibadium. Sixteen species have 3-lobed corollas, including all but one species in sects. Clibadium (except C. arboreum), Grandifolia, and Oswaldca. Two species from the Caribbean (C. erosum and C. terebinthinaceum, subg. Paleata) have 3-lobed corollas. Three species in sect.


Glomerata (C. rhytidophyllum, C. sessile, and C. sprucei) have 2-lobed corollas.

COROLLAS OF STAMINATE FLORETS

All the species in subg. Paleata sect. Eg- gersii have 4-5 lobes; C. erosum and C. terebinthinaceum, both of subg. Paleata sect. Trixidium, have disc corollas with 4 and 5 lobes, respectively. All species in sect. Glomerata and Grandifolia are 5- lobed. The remainder of species in sect. Cli- badium and Oswalda are 5-lobed or 4- lobed. One species, Clibadium micranthum, has an unusual, 3-lobed corolla. There is no correlation between number of lobes in corollas of staminate and pistillate florets.

CYPSELAE

All species have cypselae with similar morphology. Cypsela size is proportional to the size of the florets and capitula. Some species of Clibadium possess unusual drupaceous cypselae with orangish or greenish juice. This has been observed in the field in mature fruits of C. eggersii and C. laxum, which release yellow-green and orange juice, respectively, when the cypsela is squeezed. The drupe-like fruits have an inflated hypodermis with large cells and cell layers in which the juice accumulates (Stuessy & Liu, 1983). This type of fruit is rare in Asteraceae and is known only in two other genera: Chrysanthemoides Fabr. (Ca- lenduleae) and Wulffia Neck. ex Cass. (=Tillesia Meyer) of the Heliantheae.

Clibadium eggersii, C. erosum, C. divaricatum, C. leptophyllum, and C. terebinthinaceum of subg. Paleata possess unusual cypselae superficially resembling an aggregated fruit such as that found in Rubus ul- mifolius Schott (Rosaceae). This conspicuous structure is the result of aggregation of 16-40 drupaceous cypselae, all of which develop fleshy exocarps at maturity. In addition, phyllaries and paleae of pistillate florets also become fleshy at their bases, re- sulting in an inflated structure (even easily observable in dry herbarium specimens). This modified capitula is used as food by birds (C. eggersii in Costa Rica; Feinsinger et al. 2182), which serve as dispersal agents.

VESTITURE

Vestiture terminology follows Lawrence (1951). All species of Clibadium have some type of pubescence on the leaves (mainly on the abaxial surface and along veins of leaves), stems, phyllaries, and cypselae. There are two types of trichomes: (1) uniseriate; and (2) biseriate ("Zwillingshaare" of Hess, 1938). Uniseriate trichonmes are most common. Their length and density vary with position on different structures in different species (e.g., C. arboreum, C. leiocarpum, C. surinamense, C. trianae). Biseriate trichomes are restricted to the cypselae of some species (C. armanii, C. leptophyllum, C. sprucei). Even though the pubescence of the cypselae does not correlate subgenerically or sectionally, is taxonomically helpful in characterizing individual species. Some taxa have glabrous cypselae (C. anceps, C. leiocarpum, C. micranthum, C. peruvianum, C. rhytidophyllum, C. sessile, C. zarucchii), and somne have hirsute cypselae (Clibadium arboreum, C. armanii, and C. leptophyllum), and others have pilose cypselae (rest of genus).

Taxonomy

Clibadium L., Mant. P1. 161. 1771. TYPE: C. surinamense L.

Baillieria Aubl., Hist. P1. Guiane 2: 804. t. 317. 1775. TYPE: Baillieria aspera Aubl. [=C. surinamense L.].

Trixis Sw., Prodr. 115. 1788. non P Browne, Civ. Nat. Hist. Jamaica 312. 1756. TYPE: Trixis aspera Sw. [=Clibadium surinamense L.].

Oswalda Cass. in E Cuvier, Dict. Sci. Nat. 59: 322. 1829. TYPE: Oswalda baillierioides Cass. [= C. surinamense L.].

Orsinia Bertol. ex DC. Prodr. 5: 104. 1836. TYPE:

Orsinia eupatoria DC. [=C. armanii (Balb.) Sch.Bip.].

Trichapium Gilli, Feddes Repert. 94: 312. 1983. TYPE: Trich apupi umn trigosu.-m Gilli r= -a mna-,a biense H. Rob.].

Shrubs or small trees, with a conspicuous single stem or profusely branched at soil level; stems terete or angled, or striate, glabrous or strigose. Leaves opposite, sessile or petiolate; blades narrow-lanceolate to broad-ovate, subcoriaceous to coriaceous, 5-50 X 2.5-40 cm, bases attenuate, rounded, or cordate, apices acute, obtuse, or mu-


cronate, margins serrate, serrulate, finely irregularly serrate, dentate-serrate, crenate, or erose, surfaces pilose, strigose, or glabrous, with veins sometimes prominent on abaxial surfaces. Capitulescences cymose, thyrsoid, or paniculiform, capitula organized in open clusters or dense terminal or axillary glomerules, branching opposite, subtending bracts decreasing in size near capitula. Ca- pitula to 600(-819), heterogamous, with peripheral pistillate florets and central functionally staminate florets, sessile or on short peduncles; involucres cupulate or funnelform (occasionally subglobose), phyllaries 2-6, imbricate, subequal, coriaceous; phyllaries ovate or orbicular, 5-12-veined, apices obtuse, acute, or mucronate, upper one- third of margin mostly ciliate; paleae subtending pistillate florets often resembling phyllaries, ovate or orbicular, 3-9-veined;

paleae subtending staminate florets (sometimes absent) membranaceous, lanceolate, 1-3-veined, with upper one-third of margin ciliate. Pistillate florets 3-40, usually uniseriate, less often multiseriate, fertile; corollas white or yellow, tubular, 2-4 lobed; styles bifid, with strong marginal stigmatic lines; pappus usually absent, sometimes of tufts of short hairs (C. anceps) or two short awns (C. sylvestre). Staminate florets same number or fewer than pistillate florets, corollas tubular, white, with 5 triangular lobes; anthers black; style undivided. Cyp- selae obovoid, compressed, biconvex, brown-black. Staminate florets 3-24, corollas white, tubular, 4-5-lobed; anthers purplish or black, sagittate at base; styles undivided; ovaries sterile, linear, with a 5- lobed nectary at style base, pilose at apex, pappus absent. Chromosome number, x 16 (Stuessy & Arriagada, 1993).

Key to the Species of Clibadium

1. Pistillate florets multiseriate; numbers of pistillate florets always greater than number of staminate florets; all florets subtended by paleae; mature capitula with drupaceous cypselae; phyllaries and paleae fleshy (at base) at maturity; capitulescences thyrsoid -Clibadium subg. Paleata

2. Capitulescences with <25 capitula; each capitulum with >30 pistillate florets; pistillate corollas 4-lobed; pistillate paleae 5-veined; Central America and South America.

3. Capitula 4-6 mm high, phyllaries 2; pistillate florets 30-40; staminate florets 8-24. 4. Leaves ovate to ovate-lanceolate; staminate florets 8-9 -C. eggersii 4. Leaves narrowly lanceolate; staminate florets 24 -C. leptophyllum

3. Capitula 2-3 mm high, phyllaries 5-6; pistillate florets 24-39; staminate florets 6-10 --------- -------------------------------------------------------------------------------------------------------------------------------------------C. divaricatum

2. Capitulescence with 50-100 capitula; each capitulum usually <14 pistillate florets; pistillate corollas 3-lobed; paleae 3-veined; Caribbean.

5. Leaf margins finely serrate to crenate-serrate, apically acute; staminate florets 5-7; staminate corollas 5-lobed; Cuba and Jamaica -C. terebinthinaceum

5. Leaf margins erose, finely incised and serrate, apically short-acuminate; staminate florets (8-)14; corollas 4-lobed; Puerto Rico and Lesser Antilles -C. erosum

1. Pistillate florets uniseriate; number of pistillate florets same or fewer than number of staminate florets; only pistillate florets subtended by paleae, or capitula epaleaceous; mature capitula with dry cypselae; phyllaries and paleae dry (never fleshy) at maturity; capitulescences cymose or paniculiform

----------------------------------------------------------------------Clibadium subg. Clibadium 6. Capitula arranged in congested capitulescences.

7. Capitula epaleaceous. 8. Capitulescences flat-topped with clusters of 3-6 subsessile capitulae -C. anceps 8. Capitulescences round-topped with clusters of 10-30 sessile capitula.

9. Leaves pilose on both sides, margins crenate-serrulate; capitula 10-20, 2-3 mm high; Costa Rica and Panama -C. glomeratum

9. Leaves strigose on both sides, margins dentate-serrate; capitula 20-30, 5 mm high; Colombia -- ---------------------------------------------------------------------------------------------------------------------------------------------------------------C. trianae

7. Capitula paleaceous throughout or with paleae subtending pistillate florets only. 10. Only pistillate florets subtended by paleae.

11. Leaves sessile; pistillate florets 5; staminate florets 6-8; pistillate corollas 2-lobed; phyllaries 7-12-veined -C. sessile

11. Leaves petiolate; pistillate florets 3; staminate florets 3; pistillate corollas 4-lobed; phyllaries 4-6-veined -C. microcephalum

10. All florets subtended by paleae.


12. Leaves 10-35 x 6-25 cm; phyllaries 4. Colombia. 13. Capitula 6 mm high, in congested clusters 1.5-2 cm diam. - - C. frontinoense 13. Capitula 8 mm high, in congested clusters 3-4 cm diam. - --- ---- C. zarucchii

12. Leaves 7-14 X 2-5 cm; phyllaries 3. Ecuador. 14. Leaves ovate to ovate-lanceolate, apices briefly acuminate; capitula 4-4.5

mm high --- ---- -------------- --- ---- ------ C. rhytidophyllum 14. Leaves lanceolate; apices long-acuminate; capitula 6 mm high ------ C. sprucei

6. Capitula arranged in loose capitulescences. 15. Leaves 20-50 X 10-40 cm; capitulescences >200 capitula.

16. Leaf bases attenuate, decurrent on distal 1/3 of petioles; ?400 or more capitula per capitulescence; staminate florets not subtended by paleae ---- ---- C. grandifolium

16. Leaf base cordate, not decurrent on petioles; <300 capitula per capitulescence; staminate florets subtended by paleae. - --------- ------ ------ C. cordatum

15. Leaves less than 20 x 10 cm; capitulescences <400 capitula. 17. Paleae subtending only pistillate florets.

18. Staminate corollas 4-lobed. 19. Leaf bases broadly rounded or slightly cordate, margins crenate-toothed;

capitula - 100 or more per capitulescence; staminate florets 6 -- C. sodiroi 19. Leaf bases obtuse, margins serrate to crenate-serrate to serrate-dentate; ca-

pitula <100 per capitulescence; staminate florets 10-14 - C. surinamense 18. Staminate corollas 5-lobed.

20. Leaves with veins yellowish abaxially, apices acute, bases attenuate, margins finely irregularly serrate; staminate florets 19; phyllaries 3, 7-veined

-------------- ----------- --- -------------------------- ------- C. peruvianum 20. Leaves with veins light-green abaxially, apices acuminate, bases rounded,

margins serrate or mucronate-dentate; staminate florets <10; phyllaries 3-4, 5-veined.

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

21. Leaves subcoriaceous, rigid, 5-7 cm long, hispidulous, margins mucronate-dentate. Brazil, Paraguay -- ----------- C. armanii

17. Paleae subtending all florets (sometimes 1-2 central florets not subtended by paleae). 22. Staminate corollas 3- or 4-lobed.

23. Staminate corollas 3-lobed --- -------------- C. micranthum 23. Staminate corollas 4-lobed.

24. Capitulescences paniculiform; stems many-angled; leaves 14-30 cm long, blades decurrent on petioles; phyllaries 5-7-veined -- C. laxum

24. Capitulescences thyrsoid; stems terete; leaves <14 cm long, blades not decurrent on petiole; phyllaries 7-11-veined.

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

25. Leaves and stems strigillose; phyllaries light green, strigose --- -- C. sylvestre 22. Staminate corollas 5-lobed.

26. Leaves coriaceous, veins conspicuous on abaxial surfaces. 27. Phyllaries apically acuminate, strigose; pistillate florets 9-13; stami-

nate florets 15 ------------------ C. manabiense 27. Phyllaries apically obtuse, glabrous; pistillate florets 5-6; staminate

florets 5-8 ----- -------------------------- C. pentaneuron 26. Leaves thin-papery, veins not conspicuous on abaxial surface.

28. Leaf margins serrulate, teeth entire; phyllaries 5-veined; pistillate paleae 4-veined; staminate paleae 2-veined -------------------------------------- ----------------- ------------------------------------------------------------------------------- C. glabrescens

28. Leaf margins serrate to pauci-serrate, teeth mucronate; phyllaries 7- veined; pistillate paleae 7-veined; staminate paleae 3-veined ---------

--------------- ------------- ----- ----- ------ ------ ------ ------- ----- C. acuminatum

CLIBADIUM L. subg. PALEATA Arriagada, subg. nov.

TYPE: Clibadium eggersii Hieron.

Frutices capitulescentiis thyrsoideis, capitulis usque 100; flores pistillati pluriseriati, magis numerosis quam

flores staminati; achenia drupacea; phyllaria et paleae carnosae.

Shrubs with thyrsoid capitulescences, with ?100 capitula; pistillate florets multiseriate; number of pistillate florets always greater than number of staminate florets;


FIG. 1. Clibadium eggersii Hieron. (Lewis et al. 3434): A. Habit (X3/5); B. outer floret (x I I 4/5); C. Head (X5 9/10). [After Ann. Missouri Bot. Gard. 62: 1079. 1975]

staminate florets subtended by paleae; mature cypselae drupaceous; phyllaries and paleae fleshy (at base) at maturity.

CLIBADIUM L. sect. EGGERSIA Arriagada, sect. nov.

TYPE: Clibadium eggersii Hieron.

Capitulescentiae capitulis usque 25; flores pistillati plus 30, corollis quatrilobis; paleae quinquenerves.

Capitulescences with <25 capitula; pistillate florets more than 30, corollas 4- lobed; paleae of pistillate florets 5-veined.

CLIBADIUM EGGERSII Hieron., Bot. Jahrb. Syst. 28: 598. 1901. (Fig. 1). TYPE: EC- UADOR. Manabi: Rio Rapallo near El Recreo, 6 Sep 1893, Eggers 15309 (LEC- TOTYPE, here designated: K, photos: GH,

OS; ISOLECTOTYPES: P, photo: OS; US, photos: F, GH, MO, OS). Clibadium pittieri Greenm., Proc. Amer. Acad. Arts

39: 98. 1903. TYPE: COSTA RICA. Lim6n: La Florida, Jul 1897, Pittier 1129() (I.ECTOTYPE, here designated: GH; ISOLECTOTYPI'S: CR, US-2).

Clibadium pittieri Greenm. f. /Phrixium Greenm., Proc. Amer. Acad. Arts 40: 38. 1904. TYPE: COS- TA RICA: Buissons a Luis, Nov 1897, Tonduz 11479 (LECTOTYPE, here designated: GH; ISOLEC- TOTYPE: CR, US).

Clibadium polygynum S. F Blake, Contr. Gray Herb. 3(52): 6. 1917. TYPE: NICARAGUA. 1867, See- mann 88 (LECTOTYPE, here designated: BM, photos: GH, OS; ISOLECTOTYPE: BM, photo: OS).

Clibadium propinquum S. F Blake, Contr. U.S. Natl. Herb. 22: 597. 1924. Wulffia sodiroi Hieron., Bot. Jahrb. Syst. 29: 34. 1901. TYPE: ECUADOR: San- to Domingo de los Colorados, 500-1600 m, ca. 1894 (Hieronymus, 1901), Sodiro 21/3 (HOLO- TYPE: B, destroyed, fragment: US).

Clibadium chocoense Cuatrec., Rev. Acad. Coloimb. Ci. Exact. 9: 236. 1954. TYPE: COLOMBIA. Cho- c6: Rio San Juan, cercanias de Palestina, 31 May


1946, Cuatrecasas 21514 (HOLOTYPE: F, photo: US).

Shrubs, 2-3 m tall; stems branched, 2-3 cm diam., often strigose, hairs 0.1-0.5 mm long. Leaves petiolate; blades ovate to ovate-lanceolate, 5-16 X 4-10 cm, 3- veined, apices acuminate, bases attenuate, margins serrate, faces weakly to densely strigose, hairs 0.5-1 mm long; petioles 2-5 cm long, to 1 mm diam., strigose, hairs 0.1- 0.5 mm long. Capitulescences thyrsoid, 10- 25 capitula; peduncles 1 mm long. Capitula 5 mm high, 4 mm diam.; involucres cupulate; phyllaries 2, ovate, 3-3.5 X 2-2.5 mm, 5-7-veined, strigose, hairs 0.1 mm long, the distal one-third ciliate, apices acute, bases fleshy at maturity. Pistillate florets 30-40, in 5-7 series, subtended by ovate, 5-veined paleae, 2-3 X 1.5-2 mm, paleae strigillose toward apex, hairs 0.1 mm long, margins ciliate on distal one-third, bases fleshy at maturity; corollas white, 1 mm long, tubes 0.6 mm long, 0.4 mm diam., lobes 4, deltoid, shortly ciliate at margins; styles 1.5 mm long, branches 0.5 mm long. Staminate florets 8-9, subtended by lanceolate paleae, paleae 2 mm long, 0.5 mm wide, 3-veined; corollas 1.5 mm long, (4)-5-lobed, with deltoid lobes 0.5 mm long scarcely pubescent at apex; throats 1 mm long, 0.5 mm diam.; anthers 1.5 mm long, black; styles 2.5 mm long, branches 1.5 mm long; ovaries 1.5 mm long, 0.6 mm diam., pilose distally, hairs 0.5 mm long. Cypselae drupaceous, 2 X 1.6 mm, hirsute on distal one-third, hairs 0.5 mm long. Chromosome number, n = 16 (Stuessy & Arriagada, 1993).

Distribution, ecology, and phenology. Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Ecuador, and Peru; in modified wet habitats with secondary vegetation and along roadsides; 0- 1000 m; flowering throughout the year.

Selected specimens examined: GUATEMALA. Alta Verapaz: Cubilquity, 1913, Tiirckheim 4148 (F).

BELIZE. Cayo: 12 Apr 1972, Dwyer 11198 (MO). Stann Creek: along rd. and stream at Dry Creek, 14 Jun 1973, Croat 24522 (MO).

HONDURAS. Atlantida: vic. of La Ceiba, Danto River, Jun-Aug 1938, Yuncker et al. 8699 (F, US). Co- mayagua: margins of Lake Yojoa, 9 Aug 1948, Wil- liams & Molina 14621 (GH). Copan: Hacienda El Li- m6n to El Paraiso, 12-13 May 1919, Blake 7354 (US). Olancho: orillas del riachuelo Aguaquire, 30 km NE

of Culmi, 1-4 May 1975, Nelson & Vargas 2728 (MO). San Pedro Sula: Cuyamel, 1 Mar 1923, Carle- ton 458 (US).

NICARAGUA. Comarca del Cabo: Little Francia, 15 Mar 1971, Nelson 4830 (CAS, F, GH, MO, NY). Jinotega: Bocaycito, 5 May 1975, Neill 97 (MO). Ma- tagalpa: NW Cerro Musun, trocha de Palan, 14 May 1980, Araquistain & Moreno 2440 (F). Rio San Juan: near Canio Chontalefio 20 km NE of El Castillo, 18- 21 Apr 1978, Neill & Vincelli 3578 (OS). Zelaya: Canio Montecristo, 1 km S campamento La Grupera, 7 Feb 1982, Moreno 15098 (MO); Madregava, near Siu- na, 19 Mar 1971, Nelson 5059 (GH).

COSTA RICA. Alajuela: ca. 8 km S of Colonia Blanca NE toward Volcan Santa Maria, 28 Feb 1978, Almeda & Nakai 4023 (OS); Cordillera de Tilaran, near Rfo San Lorencito, NW of San Ram6n, 12 Sep 1991, Arriagada 412 (OS). Guanacaste: El Silencio, near Tilaran, 25 Jan 1926, Standley & Valerio 44639 (US). Heredia: Finca La Selva, confluence of Rio Puerto Viejo and Rio Sarapiquf, 20 Dec 1976, Denslow 79-104 (WIS); along "Starkey Road" 4.5 km SE of bridge at Puerto Viejo, 23 Jul 1979, Stevens 13310 (OS). Lim6n: Confluent du Puerto Viejo et du Sara- piqui, Jan 1893, Biolley 7399 (GH); 6 km SW of Gua- piles, 18 Jul 1964, Jimenez s.n. (UCR); Barra del Col- orado, N side, 26 Jan 1986, Stevens 24140 (OS). Pun- tarenas: Isla Violin, Sierpe, 10 May 1977, G6mez- Laurito & Bermudez 2660 (UCR); Rinc6n de Osa, 5 km W of Rinc6n, 13 Feb 1974, Liesner 2098 (OS). San Jose: San Ram6n, Cataratitas, 15 Aug 1986, Po- veda & Castro 4135 (CR).

PANAMA. Bocas del Toro: rd. from Fortuna Lake to Chiriquf Grande, 15 Mar 1985, Hampshire & Whi- tefoord 492 (MO). Col6n: vic. of Guasimo on Rio Mi- guel de la Borda, 23 Apr 1970, Croat 10005 (MO). Darien: vic. of Boca Quebrada Venado, Rio Turqueza, 29 Jun 1967, Bristan 1110 (OS); Cerro Pirre, Oct 1962, Duke 6088 (MO). Panama: Campo Tres, 5 km NE of Altos de Pacora, 9 Mar 1973, Bussey 825 (MO); forest along headwaters of Rio Torzo, 9 Jun 1967, Duke 11918 (MO, OS). San Blas: Nusagandi, 18 Jul 1986, McDonagh et al. 165 (MO). Veraguas: 5 mi NW of Santa Fe, 18-19 Mar 1973, Croat 23190 (MO); lower montane wet forest, 6-7 km W of Santa F6, 18 Feb 1974, Nee 9854 (OS).

COLOMBIA. Antioquia: valley of Rio Anorn between Dos Bocas and Anorf, 24 Jan 1975, Denslow 2642 (WIS). Caldas: Santa Cecilia, Cordillera Occi- dental, W slope, 21 Nov 1945, von Sneider 5168 (F). Choc6: La Concepci6n, 15 km E of Quibdo, 20 Apr- 23 May 1931, Archer 1945 (BM, US); Panamerican Hwy., Rio Pat6, 21 Apr 1979, Forero et al. 5418 (OS). Meta: Sierra de la Macarena, Vereda El Tablazo, 29 Mar 1973, Chaparro et al. 81 (COL, F). Nariiio: Tu- maco, La Guayacana, 29 Jul 1951, Romero-Castaneda 2974 (COL). Santander: Barranca Bermeja, between Sogamos and Carare rivers, 29 Oct 1936, Haught 2047 (COL). Valle: near Queremal, 4 Mar 1939, Alston 7901 (US).

ECUADOR. Manabi: 43.8 km SW of El Carmen, 23 Jul 1977, Stuessy et al. 4911 (OS). Napo: 44 km E of El Chaco, 4 Nov 1974, Gentry 12409 (CAS, MO). Pastaza: Colonia 24 de Mayo, on rd. to Puyo, Puerto


Napo, 23 Jun 1972, Lugo 2481 (K, MO). Pichincha: 20 km W of Santo Domingo de Los Colorados, 29 Oct 1961, Cazalet & Pennington 5188 (K, NY).

PERU. Amazonas: Bagua, Mesones-Muro High- way below Montenegro, 5.5 km E of Olmos on or above the Rio Marafion, 22 Jan 1964, Hutchinson & Wright 3721 (NY); Mendoza, 19 Aug 1963, Woy- kowski 8310 (MO). Huanuco: Pachitea, Codo de Po- zuzo, 19 Oct 1982, Foster 9315 (OS). Loreto: above Pongo de Manseriche, Creek Carapisa, 11 Dec 1931, Mexia 6255 (F, GH, K, MO, NY, TEX, US). San Mar- tin: Mariscal Caceres, Tocache Nuevo, 26 Jul 1974, Schunke 7786 (MO, OS).

Clibadium eggersii is particularly distinctive (at maturity) with its globose capitula containing 28-40 pistillate florets arranged in 5-7 series, each floret subtended by an ovate palea. Because the phyllaries and paleae (at base) become fleshy at maturity, the entire capitulum takes on an unusual aspect suggestive of aggregate fruits of Rubus.

Clibadium eggersii is the oldest name in a series of names based on specimens with variation in type and density of pubescence and the 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 and distinguished by the "less dense and adpressed pubescence and by the low number of disc florets." Clibadium pittieri was described by Greenman (I 897) 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 13778, Gentry 12409, Neill & Vincelli 3578); two collections are intermediate between these two types (Hampshire & Whitefoord 492, Lewis et al., 975). Cliba- dium pittieri f. phrixium (Greenman, 1904) was based on the hispidulous condition. The different types and densities of pubescence do not correlate with any other morphological features or particular geographic distributions or habitats, and therefore do not warrant formal recognition. All the species and subspecific taxa here considered as synonyms correspond to morphological variation within the broad geographic distributional range of C. eggersii. All of them share the same structure and organization of capitula and florets.

Clibadium eggersii is morphologically similar to C. divaricatum and C. leptophyllum. They share large numbers of pistillate florets (24-40) organized in 5-7 series, 4- lobed corollas in pistillate florets, and 4-5- lobed corollas in staminate florets. Cliba- dium eggersii differs from C. leptophyllum in having ovate leaves (vs. unusual narrowly lanceolate leaves in C. leptophyllum) and less than one-third the number of staminate florets. Clibadium eggersii differs from C. divaricatum in having capitula twice as large and with 2 phyllaries (vs. 5-6 characteristic of C. divaricatum).

CLIBADIUM LEPTOPHYLLUM Cuatrec., Revista Acad. Colomb. Ci. 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, US).

Shrubs, densely branched; stems densely strigose, hairs straight, 0.3-0.4 mm long. Leaves sessile; blades linear to linear-lanceolate, 10-16 X 2-3 cm, 3-veined, bases narrow-attenuate, amplexicaule, margins crenate, remotely serrate, and distantly revolute, apices acuminate, faces sparsely strigose, hairs to 0.2 mm long. Capitulescences thyrsoid, capitula 10-25; peduncles 2 mm long. Capitula disciform, 3 mm high, 2.5 mm diam.; involucres subglobose, 2.5 mm diam.; phyllaries 2, ovate, 2.5 X 2 mm, 5- 7-veined, scarcely strigose, hairs 0.5 mm long, distal one-third ciliate, apices acute, bases fleshy at maturity. Pistillate florets 30, multiseriate, subtended by ovate-lanceolate, 5-veined paleae, 2.5 X 1.5 mm, the paleae sparsely strigose at apex, hairs 0.1 mm long, bases fleshy at maturity; corollas white, glabrous, 2.5 mm long, tubes 1.5 mm long, 0.4 mm diam., lobes 4, deltoid, 0.6 mm long, scarcely pubescent, hairs 0.1 mm long; styles 1 mm long, branches 0.1-0.2 mm long. Staminate florets about 24, subtended by narrowly lanceolate paleae 2.2 X

0.5 mm, 3-veined, sparsely strigose with distal one-third ciliate with hairs 0.1 mm long; corollas 1.5 mm long, throats 1 mm long, 0.5 mm diam.; lobes 4-5, deltoid, 0.3 mm long, sparsely strigose at apex, hairs 0.1 mm long; anthers black 1 mm long;


styles 1-1.2 mm long, branches 0.5 mm long; ovaries, 1 mm long, pilose at apex, hairs 0.1 mm long. Cypselae drupaceous, 3 X 2 mm, compressed, bases attenuate-stip- itate, distal one-third, hirsute, hairs 0.1-0.2 mm long. Chromosome numbers unknown.

Distribution, ecology, and phenology. Colombia. This species is known only from the Choco and El Valle regions; tropical rain forest, in secondary vegetation along roadside; 0-500 m; flowering from March through June.

Selected specimens examined: COLOMBIA. Cho- c6: Serrania de Baud6, along rd. between Las Animas and Pato on Rio Pato, ca. 1 km above Pato, 18 Apr 1983, Croat 56097 (MO); Anchicaya, 14 Aug 1954, K0ie 4880 (US); Hoya del Rio San Juan, arriba Pales- tina, 28 Mar 1979, Forero et al. 4163 (COL, MO). Valle de; Cauca: new rd. Cali-Buenaventura, Km 85, El Boquer6n, 9 Oct 1974, Maas 2021 (GH).

Clibadium leptophyllum is easily distinguished by its linear-lanceolate and slightly amplexicaule leaves and small capitula resembling drupe-like fruits. Clibadium leptophyllum is known from the Choco region in Colombia, and is closely related to C. eggersii, which occurs from Belize to northern Peru, and to C. divaricatum, which is restricted to northern Peru.

Clibadium leptophyllum differs from its close relatives in Central America and South America by having narrow leaves and capitula with ca. 24 staminate florets in contrast to ovate leaves and 4-10 staminate florets per capitulum in C. divaricatum and C. eggersii. Clibadium leptophyllum is also similar to C. erosum and C. terebinthinaceum in the multiseriate condition of pistillate florets, receptacles paleaceous throughout, and thyrsoid capitulescences. Cliba- dium leptophyllum differs from those relatives, however, in having capitulescences with <25 capitula, capitulum with -30 pistillate florets, corollas of pistillate florets with 4-lobes, and paleae of pistillate florets 5-veined, all of which contrast in the Carib- bean species which have capitulescences with >50 capitula, ?20 pistillate florets, corollas of pistillate florets 3-lobed, and paleae of pistillate florets 3-7-veined.

CLIBADIUM DIVARICATUM S. F Blake, Contr. Gray Herb. 52: 7. 1917. TYPE: PERU.

San Martin: Tarapoto, 1855-56, Spruce 4522 (LECTOTYPE, here designated: BM; photo, OS; ISOLECTOTYPES: F; GH; K-2, photo: OS; P, photo: OS).

Clibadium rimachii H. Rob., Phytologia 65: 50. 1988. TYPE: PERU. San Martin: Tarapoto, Carre- tera de Tarapoto-Yurimaguas, Km 12, 23 Aug 1978, Rimachi 3877 (HOLOTYPE: US).

Shrubs, ca. 2.5 m tall; stems striate, strigose, hairs 0.5 mm long. Leaves petiolate; blades ovate-lanceolate, 7-11 X 2.5-4 cm, apices acuminate, bases attenuate, margins serrate, 3-veined, both faces weakly strigose, hairs 0.5-1 mm long; petioles 1-2 cm long, 1 mm diam., strigose, hairs 0.5 mm long. Capitulescences thyrsoid, to 25 capitula; peduncle 1 mm long. Capitula 2-3 mm high, 3-4 mm diam.; involucres cupulate, 3.5 mm diam.; phyllaries 5-6, ovate, 3 X 2.5 mm, 5-7 veined, apices acute, strigose, hairs 0.1 mm long, distal one-third ciliate, bases fleshy at maturity. Pistillate florets 24-39, subtended by ovate paleae; paleae 3 X 2.5 mm, 5-veined, arranged in 5-6 series, bases fleshy at maturity; apices acute, strigillose toward apex, distal one-third of margin ciliate; corollas white, 1.5-1.2 X 1 mm, irregularly 4-lobed, lobes deltoid, margins short and scarcely ciliate. Staminate fiorets 6-10; corollas 1.5-1.2 mm long, 4-5 lobed, lobes, 0.4 mm long, margins scarcely ciliate; throats 1 mm long, 0.5 mm diam.; subtended by lanceolate to filiform paleae, 3- veined; apices ciliate; anthers black, 1-1.2 mm long; ovaries, 1.2 mm long, 0.3 mm diam., hirsute on distal one-third. Cypselae drupaceous, 2 X 1.5 mm, hirsute on distal one-third. Chromosome numbers unknown.

Distribution, ecology, and phenology. Peru, known only from Dept. Huainuco; wet montane forest on sandy and rocky soil; 800-900 m; flowering from April to Au- gust.

Selected specimens examined: PERU. Huanuco: Chinchavito, cerca de Cayumba, entre Huanuco y Tin- go Maria, 9 Feb 1959, Ferreira 6805 (F); between Tingo Maria and Las Cuevas de las Lechuzas, MacRae 118 (F); Tingo Maria, Feb 1944, Soukup 2260 (F).

Clibadium divaricatum belongs to subg. Paleata because of 5-6 series of pistillate florets, large number of pistillate florets (24-39, usually 4-6 times the number of


staminate florets), and globose, paleaceous capitula with fleshy and juicy phyllaries. It differs from C. eggersii and C. leptophyllum, also of the same subgenus, in having small capitula (3-4 mm high) surrounded by 5-6 phyllaries to three times the number of phyllaries observed in the other two species, and also in having gray to light-green leaves (in contrast to the dark green leaves of its relatives).

Rimachi 3877 was initially described as C. rimachii by Robinson (1988), based primarily on short pedicels (similar to those in C. pediculatum Aristeg.). Critical examination of the type of C. rimachii reveals, however, that it is a minor variant of C. divaricatum (both type specimens are from the same locality). The arrangement of capitula in the capitulescences, organization of capitula, and numbers and structures of pistillate and staminate florets are the same, except for slightly larger capitula (4-5 mm) in C. rimachii (vs. the smaller capitula (3- 4 mm) in C. divaricatum).

Blake (1917) listed three type specimens in the protologue of C. divaricatum (Spruce 4522) at BM, GH, and K. Two specimens exist, one at F and one at P The sheet at BM is the only one labelled by Blake's handwriting as "type specimen." Also, it contains more complete information on collection date and habitat, and probably, most importantly, it is a superior specimen, showing all diagnostic characteristics.

CLIBADIUM L., sect. TRIXIDIuIM DC., Prodr. 506. 1836.

TYPE: Clibadium erosum (Sw.) DC. Capitulescences with 50-100 capitula;

pistillate florets <20, corollas 3-lobed; paleae of pistillate florets 3-veined.

CLIBADIUM TEREBINTHINACEUM (Sw.) DC., Prodr. 5: 506. 1836.

Trixis terebinthinacea Sw., Prodr. 115. 1788. TYPE: JAMAICA. No specific locality, Swartz s.n. (HO- LOTYPE: S, n.v.; ISOTYPE: G-DC, IDC microfiche 28.925:11.1,2).

Clibadium alexandri Griseb., Fl. Brit. W.I. 30: 368. 1861. TYPE: JAMAICA. St. Anns at Monague, Gay's Hill, 29 Jul 1850, Prior s.n. (LECTOTYPE, here designated: K, photos: GH, OS; ISOLEC-

TOTYPE: K, photo: OS).

Shrubs, to 3 m tall; stems woody toward base, striate, strigose, hairs, 1 mm long. Leaves petiolate; blades broad-ovate, 10-20 X 4-14 cm, 3-veined, apices acute, bases obtuse, attenuate, margins finely serrate to crenate-serrate, adaxial surface glabrate to strigose, abaxial surface strigose, hairs 0.5 mm long; petioles 3-5 cm long, striate, strigose, hairs 0.5 mm long. Capitulescences thyrsoid, <100 capitula loosely clustered in terminal open branches, peduncles 0.5 inni long. Capitula radiate; 5 mm high, 4 mm diam.; involucres cupulate, 4 mm diam.; phyllaries 3, ovate to ovate-suborbicular, 4 X 3 mm, light-green, 5-7-veined, abaxially glabrate to strigose, hairs 0.5 mm long, margins distal one-third ciliate, bases fleshy at maturity. Pistillate florets 12(-16), subtended by ovate paleae, 3.5-4 X 3 mm, 3- veined, strigillose toward apex, bases fleshy at maturity; corollas white, 2.5 mm long, 0.5 mm diam, 3-lobed, lobes deltoid, 0.5 mm long; styles 2.5 mm long, branches 1.5 mm long. Staminate florets 5-7, subtended by lanceolate paleae, 3-veined; corollas white, 3 mm long, throats 1 mm diam., 5- lobed, lobes 0.4 mm long; styles undivided, 2 mm long, branches 0.5 mm long. Cyp- selae drupaceous, obovoid, black, 3 X 2 mm, villous on the distal one-third. Chro- mosome numbers unknown.

Distribution, ecology, and phenology. Known only in Cuba, Jamaica, Haiti, Trin- idad, and Tobago; in ridge-type vegetation in secondary forests; 200-1500 m; flowering from December to April.

Selected specimens examined: CUBA: Oriente: Si- erra Maestra, above Firmeza, 9 Sep 1917, Ekman 8771 (A); southern Oriente and Pico Turquino, Jul 1922, Leon 10880 (NY). Sevilla: near Santiago, 10 Sep 1906, Taylor 318 (NY).

JAMAICA. Clarendon: James Hill Savanna, 23 Jun 1955, Proctor 10341 (NY, US). Saint Andrew: Sec- ond-breakfast Spring, above Mount Airy, 13 Dec 1953, Proctor 8313 (NY, US). Saint Catherine: Re- source, Mountain Diablo region, 12 Aug 1965, Hes- penheide 1119 (DS, F, GH, MO, US). Saint Thomas: trail from Bath to Corn Puss Gap, 29 Jul 1966, An- derson & Steinberg 3301 (GH, US); vic. of Cinchona, Chesterdale to Silver Hill, 2-10 Sep 1906, Britton 357 (NY); above Bowden Pen, along foot path to Bath via Cuna Cuna Pass, 4 Aug 1963, Crosby & Anderson 1045 (OS).

HAITI. Masif du Nord-St. Louis du Nord, 22 Apr 1925, Ekman 3854 (GH).

TRINIDAD. Aripo Rd., 14 May 1932, Broadwilay


8032 (GH); St. Andrew, abandoned U.S. Army Base, 20 Jun 1965, Crosby 21 (GH).

TOBAGO. Roxborough-Parlatuvier rd., main ridge, 4 Apr 1959, Cowan 1418 (GH, NY, US); Forest Res- ervoir, 20 Jan 1953, Hunnewell 20003 (GH).

Clibadium terebinthinaceum is morphologically similar to C. erosum, also from the Caribbean. Both species have ovate to broadly-ovate, 5-7-veined phyllaries with obtuse apices, corollas of pistillate florets 3- lobed, paleae of pistillate florets 3-5- veined, and paleae of staminate florets 3- veined. Clibadium terebinthinaceum differs by having phyllaries glabrous, 5-7 staminate florets (vs. 8(-14) in C. erosum), corollas of staminate florets 5-lobed (vs. 4- lobed), and cypselae villous (vs. hirsute).

The name C. terebinthinaceum has been erroneously applied to specimens of C. grandifolium, which ranges from Nicaragua to Colombia. The latter species is characterized not only by having large leaves, but also by having more staminate florets than pistillate florets. Schulz (1912) suggested that because of similarity of the capitulum arrangement, C. terebinthinaceum should be considered a variety of C. pittieri (now C. eggersii), which is known only from mainland Central America and South America. These taxa, however, are quite distinct.

CLIBADIUM EROSUM (Sw.) DC., Prodr. 5: 506. 1836. Trixis erosa Sw., Prodr. 115. 1788. TYPE: DOMINICA. St. Kitts, no specific locality, Masson s.n. (HOLOTYPE: S; ISOTYPE: n.v. G-DC, IDC microfiche 28.925:11.3,4). Clibadium fragiferum Griseb., Fl. Brit. W.I. 368.

1861. TYPE: ST. VINCENT, 1817, Guilding s.n. (HOLOTYPE: K, photo: OS).

Shrubs or small trees, to 5 m tall; stems woody at base, branches and twigs densely pubescent, hairs 1 mm long. Leaves petiolate; blades broad-ovate to ovate-oblong, membraneous, 8-25 X 8-12 cm, 3-veined, bases attenuate, apices shortly acuminate, margins erose, coarsely and finely incised and serrate, larger teeth to 8 mm long, deltoid, acute, densely strigose in both faces, hairs 1 mm long; petioles 3-6 cm long. Ca- pitulescences thyrsoid, 8-15 cm long. Ca- pitula subglobose, 5 mm high, nearly ses-

sile or shortly pedicellate; involucres cupulate, 4 mm diam.; phyllaries (3)-4, ovate, 4 X 3 mm, 5-7 veined, strigose, hairs 0.1 mm long, apices obtuse, distal one-third ciliate, bases fleshy at maturity. Pistillate florets 12(-24), subtended by ovate paleae, 4 X 3 mm wide, arranged in 3-5 series, 3- veined, abaxially strigose, distal one-third ciliate, bases fleshy at maturity; corollas white or purple, 2.5 mm long, glabrous, 3- lobed, lobes deltoid, 0.5 mm long, shortly ciliate at margins; styles 4 mm long, branches 1 mm long. Staminate florets 8- (14), subtended by ovate-lanceolate paleae, 3 mm long, 3-veined, margins distal one- third ciliate; corollas tubulate, 3 mm long, 4-lobed, lobes 0.5 mm long, apices hispidulous; anthers black, 1.5 mm long; ovaries, 2.5 mm long, apices pilose, styles undivided, 3 mm long. Cypselae, drupaceous, obovate, 2.5 X 1.5 mm, rounded at base, distal one-third hirsute, hairs 0.5 mm long. Chromosome number, n = 16 (Stuessy & Arriagada, 1993).

Distribution, ecology, and phenology. Puerto Rico, Monserrat, St. Kitts, Nevis, Dominica, Guadeloupe, Martinique, St. Lu- cia, and St. Vincent: secondary rain forest; 700-1500 m; flowering from December to June.

Selected specimens examined: PUERTO RICO. Mts. Oro Negro, Dofia Juana, 10 Jun 1962, Alain 9422 (NY); Catalina-Yunque Trail, Luquillo Mountains, 23-26 Feb 1923, Britton & Bruner 7586 (NY).

SAINT KITTS. Auckley Estate, 8 Sep-5 Oct 1901, Britton & Cowell 179 (NY).

NEVIS. Nevis Peak summit and adjacent ridge, 11 Apr 1956, Smith 10530 (A, NY, UC).

MONSERRAT. Chance's Mt, site of Pan Am crash, 18 Feb 1970, Howard & Howard 17533 (A); Blakers, in the foothills, 8 Feb 1907, Shafer 485 (NY).

GUADELOUPE. Basse Terre, pres de l'Etang de Pi- que, Sastre & Clairou 2626 (MO); Base FelTre on rd. to Soufri6re, 8-11 Jul 1969, Hespenheide 533 (GH).

DOMINICA. Lisdara Estate, 8 Mar 1933, Cooper 170 (NY); vic. of Fresh Water Lake, nr. Ladaut, 26 Mar 1956, Smith 10245 (K, NY, UC).

MARTINIQUE. Saint Denis, Bois de Fongs, 1885, Pere Duss 316 (NY).

SANTA LUCIA. Quilesse, Sep 1945, Beard 1157 (F, UC).

SAINT VINCENT The Soufriere, 26 Nov 1945, Beard 1360 (GH, NY); Richmond Peak, Morne Garou Mountains, 15 Apr 1947, Morton 4959 (GH).

Clibadium erosum is easily recognized by its peculiar, erose leaf margins, which


are unique within the genus. It is closely related to C. terebinthinaceum of sect. Tri- xidium, which is also restricted to the Ca- ribbean. They are separated from sect. Eg- gersia by having capitulescences with >50 capitula, ?24 pistillate florets in each capitulum, 3-4 phyllaries, corollas of pistillate florets 3-lobed, and paleae of pistillate florets 3-5 veined, (vs. ?25 capitula per capitulescence, capitula with >20 pistillate florets, 2 or 5-6 phyllaries, corollas of pistillate florets 4-lobed, and paleae of pistillate florets 5-veined in sect. Eggersia). Cli- badium erosum differs from C. terebinthinaceum by its ovate and strigose phyllaries, more staminate florets, corollas of staminate florets 4-5-lobed, and cypselae pilose.

Clibadium fragiferum was described by Grisebach (1864) to include populations restricted to St. Vincent and Guadeloupe and was placed by Schulz (1912) in sect. Tri- xidium, 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 that he described included the outermost series of paleae as phyllaries). Al- though the original description is incomplete, the holotype and other collections clearly indicate that C. fragiferum is an extreme of the morphological variation that is found in plants from the most southern part of the distribution of C. erosum in the Less- er Antilles.

CLIBADIUM L. subg. CLIBADIUM

CLIBADIUM L. sect. EUCLIBADIuM DC., Prodr. 5: 505. 1836.

TYPE: Clibadium surinamense L. Shrubs, with racemose, cymose, or thyr-

soid capitulescences with > 1 00 capitula; capitula frequently epaleaceous (only pistillate florets with paleae); phyllaries and paleae never fleshy at maturity; pistillate florets uniseriate; number of pistillate florets similar to number of staminate florets; mature cypselae dry.

CLIBADIUM L. sect. GLOMERATA Arriagada, sect. nov.

TYPE: Clibadium glomeratum Greenm.

Folia late ovata; capitula in glomerulis densissimis; phyllaria 4-12-nervia; flosculi feminei 3-10, corollis 2- vel 4-lobis, paleis 7-12-nervibus; flosculi staminati 4-10, corollis quinque lobis, paleis uninervibus.

Leaves commonly ovate; capitula arranged in condensed capitulescences; paleae subtending pistillate florets; phyllaries 4-12-veined; pistillate florets 3-10, corollas 2- or 4-lobed, paleae 7-12-veined; staminate florets 4-10, corollas 5-lobed, paleae 1-veined.

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. (HOLOTYPE: CR; photo: US; ISOTYPES: GH; K, photos: OS, US).

Clibadium pilonicum Stuessy, Ann. Missouri Bot. Gard. 62: 1073-1074. 1975. TYPE: PANAMA. Cocle: mountains N of El Valle de Ant6n, 28 May 1967, Lewis, MacBryde, Oliver, & Ridgway 1745 (HOLOTYPE: MO; ISOTYPE: UC).

Arching or scandent shrubs, 3-6 m tall; stems 2-5 cm diam., running horizontal and rooting at the nodes, glabrate proximally to densely strigose distally, hairs 2 mm long. Leaves petiolate; blades 7-15 X 3-9 cm, ovate to narrow-ovate, apices acuminate, bases cuneate, margins serrulate remotely denticulate more or less revolute; conspicuously 3-5-veined from near base, both faces glabrate to sparsely strigose, hairs up to 0.3 mm long; petioles <3 cm long, 1-1.5 mm diam., strigose, hairs 0.2 mm long. Ca- pitulescences thyrsoid, flat-topped, 50-100 capitula, with clusters of 3-6 sessile capitula terminating all axes. Capitula radiate, 3-4 mm high, 3-4 mm diam.; involucres funnelform, 3-4 mm diam.; phyllaries 6, 4 X 2.5-3 mm, ovate, arranged in two series, abaxially strigose, hairs 0.3 mm long, the distal one-third of margin ciliate, apices acute, 7-12 veined. Pistillate florets 3-5, subtended by ovate paleae, 3 X 2.5 mm, 7- veined, apices acute, margins ciliate on distal-one-third; corollas white, 2 to 2.5 mm long, 1 mm diam., 4-lobed, lobes 0.5 mm long, deltoid, shortly ciliate at margin; styles 3 mm long, branches 1.5 mm long. Staminate florets 4-10, epaleaceous, corol-

20031 ARRIAGADA: ASTERACEAE, HELIANTHEAE 263

las 3 mm long, 5-lobed, lobes deltoid, 0.4 mm long; anthers black, 2 mm long; styles 3 mm long; ovary 2 X 0.2 mm, villous toward apex, hairs 0.5 mm long. Cypselae 2 X 1.3 mm, glabrous. Chromosome number, n = 16 (Robinson et al., 1981).

Distribution, ecology, and phenology. Costa Rica and Panama; tropical rain forest, mainly in cutover areas in wet secondary forests along roadsides; 700-2200 m; flowering from May to August.

Selected specimens examined: COSTA RICA. Al- ajuela: along rd. between Puerto Viejo and San Jose, 3 m N of Cariblanco, 28 Sep 1987, Croat 68217 (UC); along rd. from San Ram6n northward through Balsa, 10 Sep 1979, Stevens 14136 (F, MO, OS). Cartago: 3-7 km beyond bridge in Tapanti, 9 Jul 1977, Almeda et al. 2987 (CAS); 10 km SW of Navarro, 29 Aug 1968, Wilbur & Stone 10575 (CAS, CR, GH, MO, OS). Heredia: Rio Vueltas, E slope of Volcdn Barva near the Continental Divide, 22-24 Nov 1969, Burger & Liesner 6435 (F). Puntarenas: 2.5 km NO from Estipulas, 17 Aug 1990, Arriagada 135b (OS); 17.5 mi N of Villa Neily on rd. to San Vito, 6 Nov 1976, Stuessy & Gardner 4517 (OS). San Jose: Cordillera Central, 7.5 km from San Jer6nimo, N. of San Jose, 10 Sep 1991, Arriagada 408 (OS).

PANAMA. Bocas del Toro: Carretera del Oleoduc- to, IRHE Fortuna Hydroelectric Project, 19 Jun 1982, Knapp & Vodicka 5633 (MO, UC). Chiriquf: Cerro Horqueta, 24 Jul 1966, Blum & Dwyer 2662 (FSU, MO). Cocle: Sawmill 7 km NE of El Cope, Antonio 1146 (UC); slopes Cerro Pil6n near El Valle, 10 Jun 1967, Duke 12116 (MO, OS).

Clibadium anceps is distinguished by lateral branches of the capitulescences forming more or less right angles to the main axis, fleshy phyllaries at maturity with a greenish juice, and the scandent habit. It is related to C. glomeratum and C. trianae, and the three form a more or less homo- geneous group within sect. Glomerata. The three taxa are characterized by completely lacking paleae subtending staminate florets. Clibadium anceps differs from its close relatives, C. glomeratum and C. trianae, by fewer aggregated capitula (see comments under the above mentioned species), ovate to suborbicular, strigose phyllaries, 3-5 pistillate florets, 4-10 staminate florets, and glabrate cypselae.

Clibadium pilonicum, described by Stuessy (1975) from Cerro Pil6n, in Prov. Cocl6, Panama, differs from the typical C. anceps (also collected in the same area; Duke 12116, Dwyer & Lallathin 8669) in

having capitula more tightly aggregated and narrowly ovate leaves, (vs. loosely aggregated capitula and ovate to narrowly ovate leaves). Stuessy (1975) also separated these two taxa by leaf venation (secondary midrib from the very base vs. 5-30 mm). After studying herbarium specimens, I have con- cluded that these expressions of diagnostic characters intergrade among specimens ac- cording to age and leaf position (see Anto- nio 5041; Croat 66554; D'Arcy & Todzia 15970; D'Arcy & D'Arcy 6611; Folsom & Robinson 2422; Folsom et al. 4753, 5378; Hammel 2599; Todzia et al. 2549). There- fore, the two taxa cannot be separated by macromorphology as suggested by Stuessy.

CLIBADIUM GLOMERATUM Greenm., Proc. Amer. Acad. Arts 39: 98. 1903. TYPE: COSTA RICA. Cartago: Forets de Luis, 650 m, Nov 1897, Tonduz 7330 (LECTO- TYPE, here designated: GH 11508; iso- LECTOTYPES: F, GH, K, US 5).

Shrubs, 1.5-4 m tall, 8 cm diam. at base, branches densely pilose to tomentose, hairs 1-2 mm long. Leaves petiolate; blades broadly ovate, 10-20 X 5-15 cm, apices acute, bases obtuse, subtruncate, subcordate or cordate; margins irregularly crenate-serrulate to serrulate-dentate; pilose on both surfaces, hairs 1-2 mm long; petioles 5-8 cm long, 1-1.5 mm diam., strigose, hairs 0.5-1 mm long. Capitulescences thyrsoid, branches scabrous to tomentose, hairs 1-2 mm long; glomerules 5-10 mm diam. with 10-20 tightly sessile capitula. Capitula 3- 4 mm high, 2-3 mm diam.; phyllaries 6, orbicular, 3 X 2.5 mm, arranged in 2 series, nearly glabrous below, ciliate on distal one- third, 9-12-nerved. Pistillateflorets 5, subtended by ovate paleae, 3 X 2 mm wide, glabrous, 7-veined, margin ciliate at distal one-third; corollas 1 mm long, tube 0.6 mm long, 0.5 mm diam., 4-lobed, lobes deltoid, scarcely pubescent at apex; styles 1 mm long, branches 0.5 mm long. Staminateflo- rets 5, epalaceous, corollas 1 mm long, 5- lobed, lobes deltoid, 0.5 mm long, throats 0.6 mm long, 0.3 mm diam.; anthers black, 1 mm long; styles undivided, 1 mm long; ovaries villose toward apex, hairs 0.5 mm long. Cypselae 2 X 1.5 mm wide, villous


on distal one-third, hairs 0.5 mm long. Chromosome number, n = 16 (Robinson et al., 1981).

Distribution, ecology, and phenology. Only known from Costa Rica and Panama; in second-growth tropical rain forests, open roadsides; from sea level to 500 m (occasionally found to 1400 m; see Holm & Iltis 121; Lent 1127); flowering from mid-June to late September, fruits maturing from Oc- tober to early April.

Selected specimens examined: COSTA RICA. Car- tago: ca. 7 km NE Santa Cruz, on rd. beyond Agua, 8 Mar 1978, Almeda & Nakai 4171 (CAS, OS). He- redia: Yerba Buena, NE of San Isidro, 22-28 Feb 1926, Standley & Valerio 49762 (US). Lim6n: above unnamed tributary of Rio Siquirres, 4 Dec 1929, Dodge et al. 5603 (GH). Puntarenas: Panamerican Hwy. 2, Km 37, from Olla Cinco River to Puerto Jimenez, 28 Aug 1990, Arriagada 152 (OS). San Jose: Bosquecillos entre Las Nubes y Cacajalde Coronado, 14 Jun 1980, G6mez-Laurito 5515 (CR).

PANAMA. Col6n: along Rio Escandalosa, 19 mi E of Transisthmian Hwy. on rd. to Salamanca, 28 Mar 1982, Huft & Knapp 1621 (UC); N side of Cerro Gai- tal, 10 Jul 1976, Hartman 3963 (OS).

Clibadium glomeratum is characterized by having thyrsoid capitulescences with 10 to 20 sessile capitula organized in dense glomerules 5-10 mm diam. Stems, pedicels, and leaves of most of the specimens are densely soft-tomentose (well illustrated in Liesner 2815). The density of pubescence varies, however, to moderate (Allen 5637) or even to sparingly and shortly pubescent (Burger & Liesner 6791). The 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. Cli- badium glomeratum differs from its close relatives in having open, more rounded capitulescences (vs. more flat-topped, capitate capitulescences in C. anceps). Clibadium glomeratum has been found only in Costa Rica and Panama (0-500, rarely to 1400 m), Clibadium anceps is found in Costa Rica and Chiriqui Province, Panama (700- 2200 m), and C. trianae occurs in Colom- bia, Venezuela, and Ecuador (1000-2200 m).

Greenman (1903), in the protologue of Clibadium glomeratum, mentioned two

specimens of Tonduz 7330/11508, (one at GH and the other at CR), which are indeed labelled with his handwriting and with "n. sp." Additional duplicates of Tonduz 7330/ 11508, without Greenman's annotations, exist at F, K, and US. I have chosen the specimen at GH as the lectotype because it is specifically cited in the protologue, labelled as "n. sp." in Greenman's hand, and also because it is the most complete specimen.

CLIBADIUM TRIANAE (Hieron.) S. E Blake, Contr. Gray Herb. 52: 6. 1917.

Desmanthodium trianae Hieron., Bot. Jahrb. Syst. 19: 52. 1894. TYPE: COLOMBIA. Bogota, Tena- suia, 1853, Triana 1317 (LECTOTYPE, here designated: BM; ISOLECTOTYPES: GH, photos: GH, US; K; P, photos: OS, US).

Clibadium subsessilifolium Hieron., Bot. Jahrb.Syst. 29: 32. 1900. TYPE: ECUADOR. Crescit in selvis subtropicis et subandinis, ca. 1894, Sodiro 22/1 (LECTOTYPE, here designated: US; photos: F, GH, MO, NY, OS).

Clibadium sychnocephalum S. F Blake, Contr. U.S. Nat]. Herb. 603. 1924. TYPE: COLOMBIA. Cau- ca: near Rio Flautas, 26 Jan 1906, Pittier 1211 (HO1(LOTYPE: US; ISOTYPE: F; photo: US).

Clibldilim congestum Cuatrec., Revista Acad. Co- lomb. Ci. Exact. 9: 237. 1954. TYPE: COLOM- BIA. Antioqufa: 10 km E. de Sons6n, subpdiramo, 18 Mar 1949, Scolnik, Barva, L6pez & Barkley 224 ( HOLOTYPE: F; photo: US).

Arching branched shrubs, 3-4 m tall; stems 3 cm diam., branches forming a right angle, tomentose, hairs 1-2 mm long. Leaves petiolate; blades ovate-lanceolate to widely ovate, 10-18 X 8-13 cm, apices acuminate, bases attenuate exceptionally rounded and truncate; both faces strigose, hairs 1 mm long, margins irregularly dentate-serrate to serrate, teeth ending in short mucro; petioles 2-6 cm long, strigose, hairs 1 mm long. Capitulescences thyrsoid, branches scabrous to lightly strigose, hairs 1 mm long, 5-7 capitula per cluster, radiate, organized in globose glomerules, 1.0 to 1.5 cm diam. with 20-30 tightly aggregated capitula. Capitula 5 mm high, 4 mm diam.; phyllaries 6, 4.5 X 3.5 mm, ovate, nearly glabrous abaxially, margins ciliate on distal one-third, 9-12 veined, apices acute. Pistil- late florets 5, subtended by ovate paleae, 4 X 3 mm, 7-veined, glabrous, ciliate at distal one-third; corollas 3.5 mm long, white,


tubes 3 mm long, 2 mm diam., 4-lobed, lobes deltoid, shortly ciliate at margins; styles 3 mm long, branches 1.5 mm long. Staminate florets 5-6, epalaceous, corollas 3.5 mm long, throats 2.5 mm long, 1 mm diam., 5-lobed, lobes deltoid, margins scarcely ciliate; anthers 2.5 mm long, black; styles 2.5 mm long, undivided; ovaries 2 mm long, 1.5 mm diam., pilose toward apex, hairs 1-2 mm long. Cypselae 2.5 X 1.8-2 mm, obovate, villous on distal one- third, hairs 1.5 mm long. Chromosome number, n = 16 (Stuessy & Arriagada, 1993).

Distribution, ecology, and phenology. Colombia, Venezuela, and Ecuador, in secondary growth moist tropical forests, mostly in cut areas on roadsides, sometimes on rocky slopes; 1000-2200 m; flowering from May to September.

Selected specimens examined: COLOMBIA. Anti- oquia: Mun. Frontino, Km 18.5 of rd. Nutibara-Murri, 21 Sep 1987, Zarucchi et al. 5566 (MO). Cauca: Cor- dillera Central, 1 1-13 Jun 1922, Pennell & Killip 6609 (GH). Cundinamarca: W of Bogota on rd. to El Co- legio, I I Apr 1972, Barclay et al. 3299 (COL); nr. Km 8 marker on rd. NE to El Salto from El Colegio, 13 Jul 1979, Stuessy & Funk 5528 (OS). Narinlo: Laguna de la Cocha, ca. 20 km E of Pasto, 21 Feb 1979, Mea- cham 2 (OS). Putumayo: vertiente oriental de la Cor- dillera entre El Silencio y La Cabana, 31 Dec 1940, Cuatrecasas 11521 (COL); I km W of Santiago, on gravel rd. to Pasto, 3 Aug 1979, Stuessy & Funk 5775 (OS). Santander: 26 km SW of Berlin on rd. to Bu- caramanga, Stuessy & Funk 5627, 19 Jul 1979 (OS). Tolima: Cordillera Central, Hoya Rio Combeima, Canon del Combeima, El Silencio, 3 Mar 1969, Cua- trecasas et al. 27639 (US).

VENEZUELA. Tachira: ad ripam dextram rivi Ta- chira ad limina Reipublicae Colombiae, Charpin & Jacquemoud 13274 (F, K); W forested slopes along Quebrada Agua Azul, over slate-shale substrate, S of El Reposo, 14 km SE of Delicias, Steyermark & Lies- ner 118392 (MO).

ECUADOR. Pichincha: 30 km W of San Juan on gravel rd. to Chiriboga, 15 Aug 1991, Stuessy & Arria- gada 12337 (OS).

Clibadium trianae has sessile capitula densely aggregated in ternately arranged glomerules. Its general morphology, particularly the densely aggregated small capitula, allies it with C. anceps, C. glomeratum, and C. sessile, as was also pointed out by Blake, 1917.

Clibadium congestum, described by Cua- trecasas (1954), and suggested to be a close

relative of C. trianae, is clearly a morphological variation of the latter. Within single populations, some, but not all, individuals show tomentose stems, petioles, and leaves, which were key features for the delimitation of C. congestum (e.g., Stuessy & Funk 5680). Clibadium trianae has mature leaves with serrate margins, and with a tiny mucro at the apex of each tooth. Branches forming a right angle to the stem are common. The capitula (usually 5-7 per cluster) are arranged in different degrees of congestion.

Clibadium subsessilifolium was originally applied by Hieronymus to what he considered a distinct species, mainly on the basis of large numbers of phyllaries (10). Dis- section of one capitulum from the type fragment indicates that the number of phyllaries is 5-6 (which coincides with the number commonly found in C. trianae). Clibadium trianae was originally referred by Hierony- mus to Desmanthodium; Blake (1917) cor- rected the misplacement.

Clibadium trianae commemorates Jose Jeronimo Triana (1834-1890), Columbian botanist who worked on the Prodromus Flo- rae Novo Granatensis, 1862-1867 (Stafleu & Cowan, 1986).

CLIBADIUM SESSILE S. F Blake, Ann. Mis- souri Bot. Gard. 28: 475. 1941. TYPE: PANAMA. Chiriqui: near Bajo Chorro, 20-22 Jul 1940, Woodson & Schery 658 (HOLOTYPE: US; ISOTYPES: GH, MO).

Clibadium subauriculatum Stuessy, Ann. Missouri Bot. Gard. 62: 1074. 1975. TYPE: PANAMA. Chi- riquf: Robalo Trail, N slopes of Cerro Horqueta, 5-7 Aug 1947, Allen 4970 (HOLOTYPE: GH; Iso- TYPES: BM, MO).

Shrubs, 4 m tall; stems sometimes pros- trate, glabrous to strigose, hairs 1 mm long. Leaves sessile; blades 8-20 X 5-11 cm, broadest near the middle; narrowly ovate; apices acuminate, bases cuneate, margins serrate, adaxially glabrous, abaxially strigillose, hairs 0.5-1 mm long; petioles 2-6 cm long, 2 mm diam., winged, wings decreasing toward nodes to become subauriculate at base. Capitulescences paniculiform, capitate with 70-100 sessile capitula. Capitula radiate, 3.5-4.5 mm high; involucres cupulate, 3-4 mm diam.; phyllaries 3, 3-4 X 2.5-3 mm, ovate to obovate, 7-


12-veined, abaxially glabrous, apices obtuse, margins ciliate on distal one-third. Pistillate fiorets 5, subtended by ovate to oblanceolate paleae, 7-9 veined, strigose abaxially, hairs 0.5 mm long and shortly ciliate at apex; corollas 2 mm long, 0.5 mm diam., white, 2-lobed, lobes irregularly deltoid, 0.3 mm long, margins shortly ciliate; styles 2 mm long, branches 1 mm long. Sta- minate fiorets 6-8; corollas white, 3 mm long, throats 2 mm long, 1 mm diam., 5- lobed, lobes deltoid, margin scarcely pubescent, 0.5 mm long; anthers black, 1.6 mm long; styles 3 mm long, ovaries pilose toward apex, hairs 1 mm long. Cypselae 2 X 1.5 mm, glabrous. Chromosome numbers unknown.

Distribution, ecology, and phenology. Panama; rare, only on and around Cerro Horqueta; in lower montane rain forest on open hillsides; 1400-1900 m; flowering from July to August.

Selected specimens examined: PANAMA. Chiri- qui: Cerro Horqueta along high trail to summit, 2 Jan 1975, Cochrane et al. 6287 (WIS); Bajo Chorro, 21 Mar 1977, DArcy 10911 (MO); rd. from Alto Quiel to Cerro Punta, near Bajo Chorro, 29 Jun 1976, Hart- man 3916 (OS).

Clibadium sessile is known only from Chiriqui Province, Panama. It is superficially similar to C. glomeratum from Costa Rica and Panama in having tightly clustered capitula; winged petioles with subauriculate bases make C. sessile easily distinguish- able. Winged petioles are also observed in specimens of Clibadium laxum from Ec- uador (Stuessy et al. 4942, Werling & Leth- Nissen 84), a quite unrelated species. Cli- badium sessile has its closest relationship with C. microcephalum from Ecuador. Both species have involucres with three glabrous phyllaries and staminate florets without paleae; they differ by phyllaries 7-12-veined in C. sessile (vs. 5-7-veined in C. microcephalum). They also differ in capitulum structure, C. sessile has 5 pistillate and 6- 8 staminate florets (vs. 3 pistillate and 3 staminate florets).

CLIBADIUM MICROCEPHALUM S. F Blake, J. Wash. Acad. Sci. 16: 419. 1926. TYPE: ECUADOR. Tungurahua: Valley of Pas- taza River, between Banios and Cashurco,

25 Sep 1923, Hitchcock 21873 (HOLO-

TYPE: US; ISOTYPES: GH, NY).

Shrubs, 2-4 m tall; stems strigillose, hairs 0.5 mm long. Leaves petiolate; blades 16-24 X 10-14 cm, ovate, apices acuminate, bases cuneate to cuneate-rounded, narrowly decurrent on the distal one-third of the petiole, margins shortly serrate to serrulate, sparcely strigose both faces, hairs 1 mm long; petioles 3-6 cm long, strigillose, hairs 1 mm long. Capitulescences paniculiform, ternate at apex of stem and formed by two aggregated groups of 3-4 capitula on each one. Capitula radiate, 5 mm high, 2-3 mm diam.; involucres oblong cylindric, 2-3 mm diam.; phyllaries 3, suborbicular- ovate, 2.5-3 X 2-2.5 mm, 4-6 veined, apices obtuse, ciliate on distal one-third, glabrous dorsally. Pistillate fiorets 3, subtended by ovate paleae, 2.5 X 2 mm, 5-veined, strigillose toward apex, hairs 1 mm long; corollas 2 mm long, white, tubes 1.2-1.4 mm long, 0.5 mm diam., 4-lobed, lobes deltoid, shortly ciliate at margins. Staminate florets 3, epaleate; corollas 1.5-1.8 mm long, 0.5 mm diam., white, shortly 5-lobed, lobes deltoid, 0.2 mm long, scarcely pilose at apex, hairs 0.5 mm long; anthers I mm long, black; styles 2 mm long, undivided, ovaries pilose toward apex, hairs 0.5 mm long. Cypselae 2 X 1.5 mm, obovoid, pilose on distal one-third, hairs 1 mm long. Chro- mosome number, n = 16 (Robinson, 1981; Stuessy & Arriagada, 1993).

Distribution, ecology, and phenology. Ecuador; tropical humid forests, mainly in cut-over areas; 900-2800 m; flowering from June to August.

Selected specimens examined: ECUADOR. Caniar: Guayaquil, Cuenca Rd., ca. 10 km E of Cochencai, Gentry, et al. 30826 (MO); ca. 40 km E of the bridge at Guayaquil, 21 Jan 1979, King & Almeda 7735 (CAS, MO). Napo-Pastaza: rd. Baeza-Napo, 20 km S of Baeza, Cosanga, 3 km W of village, 26 Oct 1976, Balslev & Madsen 10341 (F); 2 km of Baeza on gravel rd. to Tena, 17 Aug 1991, Stuessy & Arriagada 12363 (OS). Pichincha: 9 km N of Santa Rosa on dirt rd. to Mindo, 21 Jul 1977, Stuessy et al. 4877(OS). Tun- gurahua: entre Huambal6 y Cotal6, Acosta Solis 9723 (F); 5.3 km E of Rio Verde on rd. to Puyo, 30 Jul 1977, Stuessy & Jansen 4963 (OS).

Clibadium microcephalum is distinguished by its angled stems, acuminate leaf


apices, and cuneate leaf bases that are narrowly decurrent on the distal one-third of the petioles. Clibadium microcephalum is closely related to C. sessile. Both species are similar in having 3 glabrate, obtuse phyllaries, glabrate paleae of pistillate florets, staminate florets without paleae, and corollas of staminate florets 5-lobed. Cli- badium microcephalum differs by having orbicular, 5-7-veined phyllaries (vs. ovate, 7-12-veined in C. sessile), 3 pistillate florets with corollas 4-lobed (vs. 5 pistillate florets with corollas irregularly 2-lobed), paleae of pistillate florets 5-veined (vs. 7- 9-veined), 3 staminate florets (vs. 6-8), and pilose cypselae (vs. glabrate). Clibadium microcephalum is known only from provs. Caniar, Napo-Pastaza, Pichincha, and Tun- gurahua, Ecuador; C. sessile is only known from Prov. Chiriqui, Panama.

Blake (1926) considered Clibadium microcephalum to be related to C. grandijo- lium on the basis of the similar-sized large leaves and congested capitulescences. How- ever, the detailed structure of the capitula is quite different. Clibadium grandifolium has large capitula with 6-8 pistillate and 8-12 staminate florets vs. the 3 pistillate and 3 staminate florets in C. microcephalum.

Clibadium microcephalum has also been considered related to C. sodiroi with which it might hybridize, based on intermediate morphology and chromosomal abnormali- ties. Capitula of C. microcephalum and C. sodiroi are similar in morphology; those of C. microcephalum are arranged in small glomerules vs. capitula of C. sodiroi are more loosely arranged. It should be noted, however, the total morphological differences result in placement of these two species in different taxonomic sections in the genus.

CLIBADIUM FRONTINOENSE S.Diaz & Arria- gada, Revista Acad. Colomb. Ci. Exact. 18: 301. 1992. TYPE: COLOMBIA. Anti- oquia: Frontino, Corregimiento de Nuti- bara, cuenca del Rio Cuevas, 9 Jul 1986, Sanchez, Orrego, Silva, Martlfnez, Res- trepo, & Acevedo 213 (HOLOTYPE: ME- DEL; ISOTYPE: COL).

Shrubs, 1.5 m tall; stems branched, short-

ly strigose, hairs 0.5 mm long. Leaves petiolate; blades 28-28.5 X 20-21.5 cm, ovate, membraneous, apices acuminate, bases cordate, slightly truncate and decurrent; margins twice serrate; adaxial surface sparsely strigose, main veins conspicuous and shortly pubescent; abaxial surface light green, scarcely strigose, hairs 0.5-1 mm long, main veins thick, secondary veins 6- 7 each side, minor veins reticulated; petioles 8-10 cm long, shortly strigose, hairs 0.2 mm long. Capitulescences of tightly condensed clusters of capitula, 1.5-2.5 cm diam., with opposite, ovate leaves at base, blades 10.5 X 3.8 cm, branches to 10 cm long, densely pilose, hairs 1 mm long. Ca- pitula radiate, subglobose, 6 mm high, 3-4 mm diam., sessile or shortly pedicellate; bracts narrowly ovate or linear, 10-35 mm long decreasing toward top; phyllaries 4, ovate, 6.5 X 3.5 mm, apices acute, abaxially strigose, hairs 1 mm long, margins ciliate on distal one-third, 4-5 veined. Pistil- late fiorets 5, subtended by ovate paleae, 5.5-6 X 3.4-3.6 mm, 3-veined, margins with distal one-third ciliate; corollas 3.6 mm long, 1.6 mm diam., white, glabrous, 4-lobed, lobes deltoid, 1 mm long, margins shortly ciliate; styles 3 mm long, branches 1.6 mm long. Staminate fiorets 8, subtended by lanceolate paleae, 4.5 X 1.8-2.0 mm, 1- veined, apices obtuse, margins ciliate on distal one-third; corollas 5 mm long, white, 5-lobed, lobes 1.5 mm long, apices hirsute; ovaries pilose toward apex, hairs 1 mm long. Cypselae not seen. Chromosome number unknown.

Distribution, ecology, and phenology.- Colombia; known only from the type locality in the Frontino region, on the eastern slope of the Cordillera Occidental; 1300 m; flowering in June and July.

Clibadium frontinoense is characterized by highly congested capitulescences similar to those of its close relative, C. zarucchii, also from the Frontino region, but on the western slope of the Occidental Cordillera. The tightly congested clusters of capitula in C. frontinoense are 1.5-2.5 cm diam. (vs. 3-4 cm diam. in C. zarucchii). The capitula in C. frontinoense are smaller than those of C. zarucchii (6 mm vs. 8 mm high). Cli- badium frontinoense has 8 pistillate florets


(vs. 4). Further, the leaves in C. frontinoense are slightly decurrent from a cordate base vs. attenuate in C. zarucchii.

CLIBADIUM ZARUCCHII H. Rob., Phytologia 65: 51. 1988. TYPE: COLOMBIA. Anti- oquia: Frontino, Km 14 of rd. Nutibara- Murri, 23 Sep 1987, Zarucchi, Brant, & Castano 5674 (HOLOTYPE: US; ISOTYPES:

COL,, MO).

Shrubs, 3 m tall; stems angled, often flattened near new branches, densely strigose, hairs 1 mm long; internodes 6-9 cm long. Leaves petiolate; blades ovate, 10-20(-35) X 6-15(-25) cm, membraneous, apices acuminate, bases attenuate (new leaves), truncate-rounded, and shortly decurrent on the petiole, margins serrulate to irregularly serrate, both faces strigose, hairs 1 mm long; petioles 2-5 cm long, sulcate, narrowly winged, strigillose, hairs 1 mm long, marginate by the decurrent leaf blades. Ca- pitulescences paniculiform, with clusters of capitula tightly condensed (ca. 100 capitula/ cluster, 3-4 cm diam.) at the end of trichotomously divided terminal branches. Capitula sessile, or shortly pedicellated, 8 mm high, 3-4 mm diam.; phyllaries 4, 7-8 X 3-4 mm wide, ovate, ovate-lanceolate, apices acute, 4-5 veined, abaxially strigose, hairs 1 mm long, margins ciliate on distal one-third. Pistillate florets 4, subtended by ovate-lanceolate paleae, 6-7 X 3-4 mm, 3- veined; corollas 4.5 mm long, 2 mm diam., white, glabrous, narrowly tubulate, 4-lobed, lobes 1 mm long, margin shortly pilose, hairs 0.5 mm long; styles 3.5 mm long, branches 2 mm long. Staminate fiorets 4(- 5), subtended by lanceolate paleae, 5-5.5 X 2 mm, 1-veined, apex slightly ciliate; corollas light-purple, tubulate, 5 mm long. 2.4 mm diam., 5-lobed, lobes 1 mm long, apices slightly hirsute, hairs 0.2 mm long; styles undivided, 2.5 mm long; ovaries pilose, hairs 1 mm long. Cypselae 2.5 X 1.8- 2.1 mm, obovoid, glabrous. Chromosome numbers unknown.

Distribution, ecology, and phenology.- Known only from Mun. Frontino, Colom- bia; in wet montane vegetation, disturbed areas along roadsides, west slope of Occi-

dental Cordillera; 1850-1870 m; flowering from June to September.

Selected specimen examined: COLOMBIA. Antio- quia: Mun. Frontino, Corregimiento La Blanquita, Alto de Cuevas, 14.5 km W of Nutibara on rd. to La Blanquita, Callejas et al. 6842 (MO).

Clibadium zarucchii is easily recognized by its condensed capitulescences (ca. 100 capitula) and large (3-4 cm diam.) tightly congested clusters of capitula at the tops of trichotomously divided branches. The congested clusters of capitula are the largest of any species in the genus. Clibadium zarucchii is closely related to C. frontinoense. Both species are restricted to the Frontino region in Antioqufa, but they have allopat- ric distributions. Clibadium zarucchii grows in wet montane vegetation on the western slope of Cordillera Occidental; 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 pre- cipitation (Diaz-Piedrahita, pers. comm.). The morphological differences of these two species are discussed under C. frontinoense.

CLIBADI)1UM RHYTIDOPHYLLUM Diels, Notizbl. Bot. Gart. Berlin-Dahlem 14: 341. 1939. TYPE: ECUADOR. Pichincha: "zwischen Nono und Mindo bei Guarumas," 17 Oct 1937, Sydow 204 (HOLOTYPE: B, destroyed). NEOTYPE, here designated: Ec- uador. Pichincha: 21 km N of Nono, 2- 3 km N of Guarumas, 21 Jul 1977, Stuessy, Padilla, & Jansen 4863 (OS).

Clibacdium harlingii H. Rob., Phytologia 44: 282. 1979. TYPE: ECUADOR. Carchi: rd. Tulcan-Mal- donado, ca. 13 km SE of Maldonado, 1 Mar 1974, Harling & Anderson 12363 (HOLOTYPE: GB; Iso- TYPEI: US; photos: CAS, F, GH, MO 2).

Arching shrubs, 3 m tall; stem 10 cm diam.; stems and branches strigose-pilose, older branches tomentose, hairs 1 mm long. Leaves petiolate; blades ovate to ovate-lanceolate, subcoriaceous, 7-13 X 2-5 cm, densely strigose, hairs 0.6-1 mm long, apices briefly acuminate, bases obtuse, margins serrate to serrate-serrulate, veins prominent; petioles 1-2 cm long, strigose, hairs 1 mm long. Capitulescences thyrsoid, with 3 terminal glomerules. Capitula radiate, ovoid-subglobose, short-pedicellate, 4-4.5


mm high, 3.3-4 mm diam.; phyllaries 3, ovate, 4.5 X 2.5-3 mm, margin shortly ciliate at distal one-third, 4-5-veined. Pistil- late fiorets 5-10, subtended by ovate paleae, 4 X 3 mm, margins ciliate on distal one- third, 3-veined; corollas 2 mm long, 0.6 mm diam., white, tubulate, 2-lobed, lobes irregularly deltoid, 0.5 mm long, scarcely pilose toward apex. Staminate fiorets 5-10, subtended by ovate to ovate-rhomboid paleae, 3 X 1.5-2 mm, apices acuminate, margins shortly ciliate on distal one-third, 1- veined; corollas 3 mm long, 0.8 mm diam., white, 5-lobed, lobes 0.6 mm long, scarcely strigose toward apex; anthers 1-1.2 mm long, black, glabrous; styles 1.5 mm long, undivided, ovaries 2 mm long, 1 mm diam., distal half pilose, hairs 0.5 mm long. Cyp- selae 2 X 1.5 mm, with short ring-shaped apical projection, exocarp fleshy, glabrous. Chromosome number, n = 16 (Stuessy & Arriagada, 1993).

Distribution, ecology, and phenology. In Carchi and Pichincha Provs, Ecuador; in high montane humid forests; 2000-2600 m; flowering from June to December.

Selected specimens examined: ECUADOR. Pichin- cha: 13.5 km NW of Nono on dirt rd. to Mino, 14 Aug 1991, Stuessy & Arriagada 12311 (OS); 4 km S of Guarumos on rd. along Tandayapa River, 21 Jul 1977, Stuessy et al. 4871 (OS); along rd. between Nono and Nanegal, 89.8 km before Tandayapa, 16 Dec 1979, Croat 49326 (MO).

Clibadium rhytidophyllum is morphologically closely related to C. sprucei, which is known only from the vicinity of Volcain Tungurahua, Ecuador. Clibadium rhytidophyllum is distinguished by having lanceolate to lanceolate-ovate leaves, capitula arranged in terminal thyrses with three levels of aggregation, phyllaries greenish- white, and glabrous cypselae. Its cypselae are also characterized by the presence of a short ring-shaped apical projection similar to that seen in C. glabrescens from Vene- zuela.

Clibadium harlingii was described by Robinson (1979b) as distinct from C. rhytidophyllum because of more glomerate capitulescences, pinnate leaf venation, and glabrous cypselae. My study of specimens indicates overlap of characters in both taxa. Also, the ring-shaped apical projection on

ovaries of pistillate and staminate florets (typical of C. rhytidophyllum) also occurs in some florets of C. harlingii (e.g., Croat 49326). There seems to be no clear distinc- tion between the two species, therefore, they are here treated as synonyms.

CLIBADIUM SPRUCEI S. F Blake, Contr. Gray Herb. 3(52): 5. 1917. TYPE: ECUADOR. Tungurahua: Jan 1859, Spruce 5826 (HO- LOTYPE: GH; ISOTYPES: BM; GH, photos: GH, OS; K, photo: NY, OS; P n.v., photo: OS).

Shrubs, 2-3 m tall; stems subangulate, branches densely strigose, hairs 1 mm long. Leaves petiolate; blades 7-14 X 2-4 cm, lanceolate, apices long-acuminate, bases truncate, attenuate, 3-nerved, margins crenate-serrate, adaxially dark-green, scabrous, abaxially pilose, hairs 1-1.5 mm long; petioles 0.5-1 cm long, densely strigose-hispidulous, hairs 1.5 mm long. Capitulescen- ces thyrsoid, with dense axillary or terminal clusters of capitula, on peduncles 3-6 cm long, 2 bracts at base, glomerules 1-1.5 cm diam. (including ?20 aggregated capitula). Capitula 6 mm high, 4 mm diam.; phyllaries 3, oblongo-ovate, 5 X 4.5 mm, apices acute, densely strigose, hairs 1 mm long, distal one-third ciliate, 4-5 nerved. Pistil- late fiorets 5-7, subtended by ovate paleae, 4.5 X 3.5 mm wide, 3-veined, strigose, hairs 1 mm long; corollas 4 mm long, 1.5 mm diam., white, tubulate, 2-lobed, lobes irregularly deltoid, scarcely pilose toward apex, hairs 0.5 mm long. Staminate fiorets (7)-10, subtended by lanceolate paleae, 4 X 2 mm, 1-veined, slightly ciliate toward apex margin; corollas 3 mm long, 1 mm diam., white, tubulate, 5-lobed, lobes regularly deltoid, 0.5 mm long, scarcely ciliate toward apex; anthers black; styles undivided, 2.5 mm long; ovaries 2 mm long, 1 mm diam., pilose toward, hairs 0.8 mm long. Cypselae 2.5 X 1.8-2 mm, with fleshy exocarp, brown-black, apex pilose, hairs 1 mm long. Chromosome numbers unknown.

Distribution, ecology, and phenology. Ecuador, known only from the vicinity of Tungurahua Volcano and Chimborazo on the slopes of the Pairamos de Matanga; in tropical evergreen wet forest with second-


ary vegetation and cultivated areas; 1800- 3100 m; flowering from January to August.

Selected specimens examined: ECUADOR. Chim- borazo: Chontapamba between Puela and Bafios, 15 Mar 1969, Lugo 763 (F, MO, WIS). Morona-Santia- go: E slopes of Paramos de Matanaga, ca. 30-40 km of Sigsig, 22 Jan 1985, Luteyn & Cotton 11197 (F). Tungurahua: Chaupi, 1 Apr 1962, Dodson & Thien 1849 (MO, WIS); up path from Bafios to Volcan Tun- gurahua, 19 Aug 1991, Stuessy & Arriagada 12400 (OS).

Clibadium sprucei is morphologically close to C. rhytidophyllum; both are endemic to Ecuador but in different regions. Cli- badium sprucei is confined close to the Tungurahua Volcano; C. rhytidophyllum has been reported only from provs. Carchi (border with Colombia) and Pichincha (250 km N of Prov. Tungurahua). Both species occur in similar habitats (2000-3000 m) and have similar general morphology, particularly in structure of the capitulescence and in the congested capitulescences. Cli- badium sprucei is easily recognized by its lanceolate leaves (vs. the ovate leaves of C. rhytidophyllum). The capitula are arranged on axillary and terminal branches (vs. only in terminal thyrsoid clusters). Cypselae in both species have fleshy exocarps; only C. rhytidophyllum produces an orange sap (e.g., Stuessy et al. 4863). Clibadium sprucei lacks the ring-shaped apical projection seen in specimens of C. rhytidophyllum. In addition, C. sprucei has pilose cypselae (vs. glabrous).

This species was named to honor the British botanist and plant collector Richard Spruce (1817-1893), who collected in South America from 1849 to 1864 (Stafleu & Cowan, 1985).

CLIBADIUM L. sect. GRANDIFOLIA Arriaga- da, sect. nov.

TYPE: Clibadium grandifolium Blake.

Folia ovata vel lanceolato-ovata; capitulescentiae cymarum laxae vel thyrsiformes; phyllaria quinque- nervia; flores pistillati 5-9, corollis trilobis, paleis qua- drinervibus; flores staminati 5-11, corollis quinquelo- bis, paleis binervibus.

Leaves ovate to ovate-lanceolate; capitula arranged in loose thyrsoid or paniculiform capitulescences, paleaceous; phylla-

ries 5-veined; pistillate florets 5-9, with corollas 3-lobed, and paleae 4-veined; staminate florets 5-11, with corollas 5-lobed, and paleae 2-veined.

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; ISO- TYPE; K; photo: OS). Clibadium grande S. F Blake, Contr. U.S. Natl.

Herb. 22: 601. 1924. TYPE: COSTA RICA. Li- m6n: La Florida, 18 Jun 1897, Pittier 11280 (HO- LOTYPE: GH; photos: GH, US; ISOTYPE: US).

Clibadium pacificum Cuatrec., Revista Acad. Co- lomb. Ci. Exact. 9: 238. 1954. TYPE: COLOM- BIA. Valle: Rfo Calima entre La Herradura de Ord6fiez y Pefia de Campotriste, 3 Mar 1944, Cuatrecasas 16683 (HOLOTYPE: F; photo: US; iso- TYPES: COL; F, photos: US).

Clibadium terebinthinaceum (Sw.) DC. subsp. colombiense Cuatrec., Revista Acad. Colomb. Ci. Exact. 9: 240. 1954. TYPE: COLOMBIA. Valle: Hoya del Rio Digua, lado derecho, La Elsa, 9 Nov 1943, Cuatrecasas 15298 (HOLOTYPE: F; pho- to: US; ISOTYPES: COL, US).

Shrubs to small trees, 2-6 m tall; stems 3 cm diam. toward base, striate, strigose, hairs 1 mm long. Leaves petiolate; blades 20-40 X 10-30 cm, broadly ovate, apices acute to acuminate, bases obtuse to shortly attenuate, sometimes decurrens on distal third of petiole; margins serrate, adaxial surface glabrate to strigose, abaxial surface strigose, hairs 0.6-0.8 mm long; petioles 4- 19 cm long, 3-4 mm diam., glabrate to strigillose, hairs 0.5 mm long. Capitulescences paniculiform, with 400-600(-819) clustered capitula in terminal open branches, peduncles 0.5 mm long. Capitula globose to subglobose, (3) 4-6 mm high, 3-5 mm diam.; involucres cupulate; phyllaries 3, light green, ovate to suborbicular-ovate, 3- 5 X 3-4 mm, 5-veined, abaxially glabrate to strigillose, hairs 1 mm long, margins ciliate on distal one-third. Pistillate forets 6- 8(-10), subtended by ovate paleae, 3-4 X 2-2.6 mm, strigillose toward apex, hairs 0.5 mm long, margins ciliate on distal one- third, 4-veined; corollas 3 mm long, 0.5 mm diam., white, 3-lobed, lobes deltoid, 0.4 mm long, scarcely pubescent at apex; styles 2.5 mm long, branches 1.2 mm long. Staminate fiorets 8-12(-14), epaleate (ex-


ceptionally, 1-2 disc florets subtended by narrowly lanceolate paleae); corollas 3-4 mm long, throats 2.5 mm long, 1 mm diam., white, 5-lobed, lobes 0.5 mm long, scarcely pubescent at apex; anthers 2 mm long, black; styles 3 mm long, ovaries 1.5 mm long, villose at apex. Cypselae 2.5 X 1.7 mm, pilose on upper one-third, hairs 0.5 mm long. Chromosome number unknown.

Distribution, ecology, and phenology. Nicaragua, Costa Rica, Panama, and Co- lombia; in tropical moist forests, secondary vegetation, disturbed forests, mainly along roadsides; 0-2000 m; flowering from March to September.

Selected specimens examined: NICARAGUA. Ze- laya: ca. 6.3 km S of bridge at Colonia Yolania, 29- 31 Oct 1977, Stevens 4816 (OS).

COSTA RICA. Cartago: beyond Pavones, 14 km from IICA Turrialba, 26 Jan 1957, Carlson 3405 (F, NY). Heredia: Finca La Selva OTS Field Station, 19 Jul 1980, Hammel 9240 (MO). Lim6n: SE of Siquirres near Rio Pacuare, 10 Sep 1991, Arriagada 406 (OS). Puntarenas: Santa Cecilia, 8 Jul 1923, Stevens 753 (US). San Jose: Parque Nacional Braulio Carrillo, Del Bajo de La Hondura, 13 km a Guapiles, 23 Mar 1981, G:i,nez-Laurito 6419 (CR).

PANAMA. Bocas del Toro: Santa Catalina, along riverbank, 4 Dec 1967, Blackwell et al. 2749 (MO). Chiriqui: Fortuna Dam area on Kaolin Hill, 31 Jul 1984, DArcy et al. 15904 (MO, UC). Cocle: El Valle de Ant6n, vic. Finca Tomas Arias, 5 Aug 1946, Allen 3621 (MO); 2.5 mi above El Valle on rd. to La Mesa, 11 Feb 1971, Croat 13372 (OS). Col6n: along Rio Escandalosa on rd. to Salamanca, 28 Mar 1982, Huft & Knapp 1621 (MO). Darien: NO trails to Cerro Pirre, along banks of Rio Perrecenega, 31 Mar 1985, D'Arcy & McPherson 16230 (MO). Panama: hill S of Gua- cuco, 8 km E of Ipeti, 18 Sep 1982, Hamilton & D 'Arcy 1391 (MO). Veraguas: roadside between San- ta F6 and San Jos6, 6 Dec 1975, DArcy 10322 (MO); 3.5 mi N of Santa F6, 12 Dec 1971, Gentry 3034 (OS).

COLOMBIA. Antioquia: Mun. Frontino, Corregi- miento La Blanquita, 14.5 km W Nutibara, 11 Jun 1988, Callejas et al. 6610 (MO). Boyaca: Mt. Chap6n, NW of Bogota, 1932, Lawrence 135 (F, GH, K, MO, NY, US). Cauca: Cordillera Occidental, Carpinterfas, bosque entre los cerros Munchique y Altamira, Perez Arbelaez & Cuatrecasas 6142 (COL, F, US). Choc6: Rfo El Valle, between El Valle and Choc6 Indian Vil- lage, 6 Aug 1976, Gentry & Fallen 17249 (MO, OS); Rio Truando, between La Nueva and La Esperanza, Duke 9886 (FSU); Mun. San Jos6 del Palmar, Hoya del Rio Torito, Finca "Los Guaduales," Forero et al. 6853 (MO). Tolima: at bridge over Saldania to house of Manuel Puertas, Core 1599 (F, US). Valle: Cordil- lera Central, Hoya del Rio Guadalajara, 27 Feb 1969, Cuatrecasas et al. 27601 (NY, US). Vaupes: Rio Ku- duyari, middle and lower course, 16 Oct 1952, Schul- tes 17861 (GH).

Clibadium grandifolium is easily recognizable by its large leaves, to 40 cm long (Lawrence 135; Killip & Garcia 33575; Zarucchi 5036) with obtuse to shortly attenuate bases and by paniculiform capitulescences with 100-600 capitula (819 capitula, the largest number counted for a single capitulescence in any species) clustered together on terminal branches. The capitula ranges from 6 mm high (Stuessy & Gardner 4533) to 4-5 mm high (Hammel 3891) to (rarely) 3 mm high (Williams 20263). In juvenile specimens, the leaves are smaller (10-15 X 8-10 cm) and superficially resemble leaves of plants of C. sylvestre. Stri- gose phyllaries distinguish C. sylvestre. Cli- badium grandifolium is closely related to C. cordatum and differs in having glabrate, obtuse phyllaries, pilose cypselae, and staminate florets without paleae.

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 em- phasized by him are not sufficient for formal taxonomic recognition at any level. The overlap of diagnostic characters mentioned by Blake for these species supports the de- cision to treat them as conspecific.

CLIBADIUM CORDATUM Cuatrec., Revista Acad. Colomb. Ci. Exact. 9: 237. 1954. TYPE: COLOMBIA. Valle: Cordillera Occidental, Hoya del Rio Anchicaya2, bosques entre Pavas y Miramar, 15 Apr 1943, Cuatrecasas 14378 (HOLOTYPE:

F-1362889, F-1362890; photo: US; ISOTYPE: US 2220748, US-2220749).

Shrubs, 2-4 m tall; stems subangulate, 10 cm diam., branches strigose-pilose, hairs 1 mm long. Leaves largely petiolate; blades 20-50 X 20-40 cm, ovate, rounded-ovate, membraneous, densely strigose, hairs 0.6-1 mm long, apices briefly acuminate, bases deeply cordate (mainly in old leaves); margins serrate to crenate-serrate, veins 7-9, prominent; petioles 16-22 cm long, 1.5-2.5 mm diam., striate, strigose, hairs 1 mm long. Capitulescences cymose or paniculi-


form, broadly branched. Capitula ovoid, 4- 5 mm high, 3 mm diam., shortly pedicellate or sessile, clustered in aggregations at terminal branches; phyllaries 3, 4-5 X 3-4 mm, obovate to obovate-ovate, apices mucronate-acuminate, strigose, hairs 1 mm long, margins ciliate at upper half, 5- veined. Pistillate fiorets 7, paleate, subtended by ovate paleae, 4.5 X 3-3.5 mm, strigose hairs 0.3-0.4 mm long, margins shortly ciliate at distal half, 4-veined; corollas 2 mm long, white, tubes 1.4 mm long, 0.5 mm diam., 3-lobed, lobes irregularly deltoid, 0.5 mm long, shortly ciliate at margin; styles 2.5 mm long, branches 1 mm long. Staminate fiorets 8(-9), subtended by lanceolate paleae, 3-4 X 0.5-1 mm, 2-veined, margins shortly ciliate at apex; corollas 3 mm long, 0.5 mm diam., white, 5-lobed, lobes deltoid, 0.5 mm long, scarcely pubescent at apex; anthers 1-1.5 mm long, black; styles 2 mm long, undivided; ovaries pilose toward apex, hairs 1 mm long. Cypselae 2 X 1.6 mm, pilose on the distal one-third, hairs 1-1.5 mm long. Chromosome number, n = 16 (Stuessy & Arriagada, 1993).

Distribution, ecology, and phenology. Colombia, mostly restricted to Choco and Valle; growing in cut-over areas with secondary vegetation; 300-600 m; flowering from March to December.

Selected specimens examined: COLOMBIA. Cho- c6: headwaters of Rfo Tutunendo, E of Quibd6, 20- 21 May 1931, A rcher 2187 (NY); Mun. San Jose del Palmar, Hoya de la Quebrada La Cristalina, Forero et al. 7594 (MO); carretera San Jose del Palmar-Novita, alrededores de Curundu, 26 Feb 1977, Forero et al. 3517 (COL, MO). Valle: Cordillera Occidental, vertiente occidental, Hoya del Rio Anchicaya, Quebrada del Retiro, Cuatrecasas 13696 (COL, F, US, PARA-

TYPES); Mun. Dagua, Corregimiento El Danubio, Alto Anchicaya, Devia 624 (MO).

Clibadium cordatum is distinguished by its large leaves (20-50 X 15-40 cm, the largest leaves known for the entire genus) with cordate bases (young leaves are somewhat attenuate, e.g., Forero et al. 5871). This species is sometimes mistaken for C. grandifolium of Costa Rica and Panama, which has also large leaves, but with more attenuate bases. In addition, C. cordatum has condensed clusters of capitula (1 cm diam.) arranged in paniculiform capitulescences. Clibadium cordatum differs from C.

grandifolium in having strigose, mucronate phyllaries (vs. glabrous and obtuse) and staminate florets subtended by lanceolate, 2- veined paleae (vs. no paleae subtending staminate florets).

CLIBADIUM MANABIENSE H. Rob., Phytologia 44: 280. 1979. TYPE: ECUADOR. Ma- nabi: rd. to Santo Domingo-Chone, Fla- vio Alfaro, 11 May 1968, Harling, Stormii, & Storm 9410 (HOLOTYPE: GB; photos: CAS, F, GH, MO; ISOTYPE: US).

Trichapium strigosum Gilli, Feddes Repert. 94: 303. 1983. TYPE: ECUADOR. Waldfragment bei Santo Domingo de los Colorados, 19 Jun 1975, Gilli 113 (HOLOTYPE: W; photos: CAS, F, MO-2).

Shrubs, 2.5 m tall; stems striate, strigose, hairs 1 mm long, adpressed. Leaves petiolate; blades ovate to ovate-lanceolate, coriaceous, 8-15 X 3-8 cm, apices acuminate, bases cuneate to rounded cuneate, margins crenate-serrate, adaxially strigose, hairs 0.5 mm long, scarcely pilose abaxially, hairs 1 mm long, 3 main veins, primary and secondary veins prominent; petioles 0.5-2.5 cm long, striate, strigose, hairs 1 mm long. Cap itulescences thyrsoid, at terminal branches, branches strigose, hairs 1 mm long. Capitula subglobose, 4-5 X 4-5 mm diam.; phyllaries 2-3, 4-5 X 3.5-4 mm, suborbicular, abaxially strigose, hairs 0.5- 0.8 mm long, apices markedly acuminate- mucronate, margins shortly ciliate on distal half, 5-veined. Pistillate forets 9-13, subtended by ovate paleae, 4-4.5 X 3.5-4 mm, 5-veined, strigose, hairs 0.5 mm long, margins ciliate on distal one-third; corollas 2.5 mm long, 1 mm diam., white, tubulate, 3- lobed, lobes 0.5 mm long, apices scarcely pubescent; styles 1.5 mm long, branches 1 mm long. Staminate florets 15, subtended by lanceolate paleae, 3-3.5 X 1.5-2 mm, margin ciliate on distal one-third, 2-veined; corollas 3 mm long, white, throats 1 mm diam., 5-lobed, lobes 0.5 mm long, apices scarcely pubescent; anthers 1.5 mm long, dark brown-purple; styles undivided, 2 mm long, ovaries 1 mm long, 0.6 mm diam., apices pilose, hairs 1 mm long. Cypselae 2.5 X 2 mm, obovoid, apices pilose, hairs 1 mm long. Chromosome number unknown.

Distribution, ecology, and phenology.-


Only known from provs. Manabi and Pi- chincha in Ecuador; secondary vegetation; 100-2000 m: flowering from July to Sep- tember.

Selected specimens examined. ECUADOR: Pichin- cha: carretera Quito-San Juan-Chiriboga-Empalme, Km 69 a 3 km de la carretera a Santo Domingo de Los Colorados, 17 Sep 1986, Zack 1194 (F, MO); Re- serva Ecol6gica 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).

Clibadium manabiense is characterized 9-13 pistillate florets, 15 staminate florets, and acuminate phyllaries. Clibadium manabiense is morphologically similar to C. pentaneuron (e.g., phyllaries 5-veined, pistillate corollas 3-lobed, pistillate paleae 4- 5-veined, staminate corollas 5-lobed, staminate paleae 2-veined, and cypselae pilose). Clibadium manabiense can be distinguished easily by the 2-3 strigose phyllaries (vs. 3-4 glabrate phyllaries in C. pentaneuron).

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; pho- to: US; ISOTYPES: COL-2, LL).

Clibadium napoense H. Rob., Phytologia 73: 152. 1992. TYPE: ECUADOR. Napo: Cant6n Napo, Zatzayacu, 28 Mar 1935, Mexia 71/IA (HOLO- TYPE: US; ISOTYPES: LL, UC, US).

Spreading shrubs, 2.5 m tall; stems with branches slender, striate, glabrous to strigose, hairs 1 mm long, internodes 3-12 cm long. Leaves petiolate; blades 8-18 X 4-9 cm, ovate, thin-papery, apices caudate-acuminate, bases cuneate, margins serrate to serrulate; adaxially dark green, roughly strigose, hairs 1 mm long; abaxially light green, sparcely strigose, hairs 0.6 mm long; petioles slender, slightly striate, 1-2 cm long, strigillose (ocasionally glabrous), hairs 0.6-1 mm long. Capitulescences paniculiform, on terminal branches. Capitula sessile or short-pedicellate, subglobose, 3.5-4.5 mm high, 3-3.5 mm diam.; phyllaries 4, 3.5-4 X 3-3.5 mm, ovate-suborbicular, 5-veined, apices obtuse, distal half strigose, hairs 0.6 mm long, margins ciliate on distal one-third. Pistillate florets 5-6,

subtended by ovate paleae, 3-3.5 X 3-3.5 mm, distal half strigose, hairs 0.5 mm long, margins ciliate on distal one-third, 4- veined; corollas 2 mm long, 0.8 mm diam., white, tubular, 3-lobed, lobes 0.4 mm long, apices scarcely to densely strigillose; styles 1.5 mm long, branches 0.5 mm long. Sta- minate fiorets 8-11, subtended by ovate- lanceolate paleae, 3-3.5 X 1.5-2 mm, apices scarcely ciliate, 2-veined; corollas 3 mm long, 0.5 mm diam., white, 5-lobed, lobes deltoid, 0.4 mm long, apices scarcely pubescent; anthers 1.5 mm long, black; ovaries 2 mm long, 1.5 mm diam., pilose toward apex. Cypselae obovoid, with short ring-shaped apical projection, bases rounded, pilose at distal one-third, hairs 1.1-5 mm long. Chromosome number, n = 16 (Stuessy & Arriagada, 1993).

Distribution, ecology, and phenology. Colombia, Venezuela, and Ecuador on the eastern slope of the Andes; in low humid tropical forest, along wooded stream banks; 500-1000 m; flowering from March to Sep- tember.

Selected specimens examined: COLOMBIA. Nari- ino: El Palmar, 25 Oct 1944, von Sneidern 4522 (GH). Santander: 15 km NE of Bucaramanga on rd. to Pom- plona, 17 Jul 1979, Stuessy & Funk 5606 (OS).

VENEZUELA. Merida: Distrito Campo Elias, Bosque de San Eusebio, Rufz Terdn 1139 (MO).

ECUADOR. Los Rios: between Quevedo and Na- ranjal, 7-8 Nov 1934, Mexia 6668 (NY, US). Napo- Pastaza: near Canelos, Mexia 6915a (US). Tena: N edge of Archidona on rd. to Baeza, Aug 1977 Stuessy & Jansen 4985 (OS). Zamora: ca. 5 km NE of Zamora on rd. to Timbera, 6 Sep 1979, Stuessy & Nesom 5897 (OS).

Clibadium glabrescens is characterized by strigose leaves and capitulescences of open clusters of capitula arranged in panicles at the ends of terminal branches. Cli- badium glabrescens is closest to C. pentaneuron. Both species share 5-veined, obtuse phyllaries, 5-6 pistillate florets with 3- lobed corollas, 4-veined paleae subtending pistillate florets, staminate florets with 5- lobed corollas, 2-veined staminate paleae, and pilose cypselae. Clibadium glabrescens differs from C. pentaneuron by having 4 strigose phyllaries and 8-11 staminate florets (vs. 3-4 glabrate phyllaries and 5-8 staminate florets).

Most characteristics of Clibadium gla-


brescens coincide with those used to char- acterize C. napoense, from Napo, Ecuador, the same eastern slope of the Cordillera. The only difference is the occasional presence of 3 phyllaries (although some capitula have 4 phyllaries, e.g., Mexia 7110; Stuessy & Jansen 4981) vs. the common 4- phyllary condition in C. glabrescens.

CLIBADIUM PENTANEURON S. E Blake, Contr. U.S. Natl. Herb. 22: 598. 1924. TYPE: COLOMBIA, 1906, Lehmann 1256 (HO- LOTYPE: GH; ISOTYPES: F-2, photo: US; GH, photo: US; K, photo: OS; NYO). Clibadium pileorubrum Cuatrec., Revista Acad. Co-

lomb. Ci. Exact. 9: 238. 1954. TYPE: COLOM- BIA. Huila: Comisarfa Caqueta, filo divisorio de la Cordillera Oriental, 21 Mar 1940, Cuatrecasas 8422 (HOLOTYPE: F; photo: US).

Clibadium sarmentosum Cuatrec., Revista Acad. Colomb. Ci. Exact. 9: 239. 1954. TYPE: COLOM- BIA. 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-n.v., photo: OS).

Clibadium scandens Cuatrec., Revista Acad. Co- lomb. Ci. Exact. 9: 239. 1954. TYPE: COLOM- BIA. Putumayo: Valle de Sibundoy, La Cabafia, 2 Jan 1941, Cuatrecasas 11596 (HOLOTYPE: F; photo US; ISOTYPE: COL).

Clibadium funkiae H. Rob., Phytologia 73: 149. 1992. TYPE: COLOMBIA. Antioqufa: 3 km SE of Santa Elena on rd. from Medellin to Santa Elena and Rio Negro, 15 km NW of Rio Negro, 30 Jul 1979, Stuessy & Funk 5709 (HOLOTYPE: US; ISO- TYPE: OS-2).

Clibadium zakii H. Rob., Phytologia 73: 153. 1992. TYPE: ECUADOR. Bolivar: carretera Chillanes- Bucay, en la Hacienda Tiquibuso del Sr. Gonzalo G6mez, 10 Sep 1987, Zak & Jaramillo 2881 (HO-

LOTYPE: US; ISOTYPES: F, MO).

Arching or scandent shrubs, to 2 m tall; stem scabrous to strigose, hairs appressed, 1 mm long. Leaves petiolate; blades ovate to ovate-lanceolate, thick, coriaceous, 7-16 X 3-7 cm, 5-nerved, primary and secondary veins abaxially prominent, apices acuminate, bases cuneate to round-cuneate, margins serrate, teeth less than 1 mm high; both faces dark green and sparcely strigose, hairs 0.5-1 mm long; petioles 10-15 mm long scabrous to strigose, hairs 1 mm long. Capitulescences paniculiform, always trichotomously divided at ending of branches and stems. Capitula sessile, or on 2 mm long pedicels, subglobose, 4-5 X 3-4 mm; phyllaries 3-4, 4-5 X 3-4 mm, suborbic-

ular, 5-veined, apices obtuse, glabrous, slightly hirsute and ciliate on distal one- third. Pistillate fiorets 5-6, subtended by ovate paleae, 4 X 3 mm, white, 4-veined, slightly hirsute on distal one-third; corollas 2 to 2.5 mm long, throats 1 to 1.5 mm diam., 3-lobed, lobes irregularly deltoid, 0.5 mm long, apices scarcely pubescent; styles 1.5 mm long, branches 1 mm long. Stami- nate florets 5-8, subtended by lanceolate paleae (exceptionally, epaleaceous central disc floret), 3.5 X 1.5 mm, apices slightly ciliate, 2-veined; corollas 2.5 mm long, white, tubes 1.5 mm long, 0.5 mm diam., 5-lobed, lobes 0.5 mm long, deltoid, apices slightly pilose; anthers 1 mm long, black; styles 2 mm long, undivided; ovaries 1.5 mm long, 1 mm diam., apices pilose, hairs 0.5 mm long. Cypselae 2 X 1.5 mm, piloses on distal half, hairs 0.5 mm long. Chro- mosome number, n = 16 (Stuessy & Arria- gada, 1993).

Distribution, ecology, and phenology. Colombia, Ecuador; in cloud forests to sub- paramos; (100-)1400-2800 m; flowering June to January, fruiting from August to January.

Selected specimens examined: COLOMBIA. Anti- oquia: Mun. Frontino, on rd. to Murri, 15 km W of Nutibara, 17 Oct 1987, Brant & Martfnez 1378 (BM, K, MO); 3 km SE of Santa Elena on rd. to Medellin to Santa Clara and Rio Negro, 30 Jul 1979, Stuessy & Funk 5709 (OS). Boyaca: 9 km N of Arcabuco on rd. to Bambita, ca. 5 km S of Boyaca-Santander state line, 16 Jul 1979, Stuessy & Funk 5595 (OS). Caldas: Sa- lento, Rio Santa Rita, Killip & Hazen 8976 (GH). Ca- queta: 30 km SE of Guadelupe on rd. to Florencia, 29 Jul 1922, Gentry et al. 9036A (COL, MO). Choc6: Carretera Anserma-Nuevo-San Jos6 del Palmar, entre el Alto del Galapago y San Jos6 del Palmar, 29 Aug 1976, Forero et al. 2279 (MO, OS). Cundinamarca: Eastern Cordillera, ca. 20 km NE of Fusagasuga, King & Guevara 5754 (F, NY, US). Nariiio: Mun. Barba- coas, I km de Junin, carretera Tumaco-Pasto, Bena- vides 4688 (MO); El Palmar, Cordillera Occidental, vertiente occidental, 25 Oct 1944, von Sneidern 4522 (WIS). Putumayo: Valle de Sibundoy, 2 km SW of Sibundoy, 31 Jan 1963, Bristol 519 (GH, US). San- tander-Boyaca: cerca al limite, en la carretera de Gambita a Arcabuco, Uribe 6310 (COL, US). Valle: Cordillera Occidental, vertiente occidental, Hoya del Rio Anchicaya, entre Sabaletas y la Quebrada del Ta- tabro, Cuatrecasas 22034 (F); Bajo Calima, 15 km N of Buenaventura, 25 Jul 1988, Faber-Langendoen & Hurtado 1856 (MO); Cart6n de Colombia Concession, 15 Feb 1983, Gentry et al. 40293 (MO).

ECUADOR. Bolivar: 10 Sep 1987, Zak & Jara- millo 2881 (MO). Zamora: 13.4 km NE of Cumba-


ratza on gravel rd. to Zumbi, 6 Sep 1979, Stuessy & Nesom 5902 (OS).

Clibadium pentaneuron is found mostly at high elevations (1400-2800 m) where it has thick coriaceous leaves and capitula 4- 5 mm high arranged in panicles of open clusters (6-10 cm wide) at the ends of trichotomously divided branches. However, four specimens, Faber-Langendoen & Hur- tado 1856; Gentry 40293; and Monsalve 624, 1601, which were collected below 100 m, have ovate-lanceolate to lanceolate leaves with thin blades, capitula with 3 pistillate and 6 staminate florets all subtended by paleae, and strigose phyllaries and paleae.

Clibadium pentaneuron is closely related to C. glabrescens. Both have orbicular, obtuse, 5-veined phyllaries, 5-6 pistillate florets with 3-lobed corollas, subtended by 4- veined orbicular paleae, staminate florets subtended by 2-veined paleae, corollas 5- lobed, and cypselae pilose. Clibadium pentaneuron differs from C. glabrescens in its glabrate (vs. strigose) phyllaries and 5-8 (vs. 8-l 1) staminate florets.

Clibadium scandens (Cuatrecasas, 1954) was based only on the type material, in which capitula are in more condensed clusters. Cuatrecasas (1954) also described C. pileorubrum and C. sarmentosum. Reex- amination of the types of those names clearly suggests that C. pileorubrum differs from C. pentaneuron only in having smaller leaves (3-4 X 2-3 cm vs. 7-16 X 3-7 cm, respectively) and more acuminate paleae (ovate in C. pentaneron). Clibadium sarmentosum is a densely pilose variant of the more commonly scabrous to shortly strigose-pubescent C. pentaneuron. Also, there are no geographic distribution patterns to any of these minor morphological variations. They are treated as synonyms here.

CLIBADIUM L. sect. CLIBADIUM

Clibadium L. Mant. P1. 161. 1771.

TYPE: Clibadium surinamense L. Leaves ovate to ovate-lanceolate; capit-

ula arranged in cymose, thyrsoid, or paniculiform capitulescences; with only pistillate florets subtended by paleae; phyllaries

5-7-veined; pistillate florets 3-9, with pistillate corollas 3-4-lobed, and paleae 5- veined.

CLIBADIUM LAXUM S. F Blake, J. Wash. Acad. Sci. 16: 418. 1926. TYPE: ECUA- DOR. Guayas: Teresita, 3 km W of Bu- cay, 5-7 Jul 1923, Hitchcock 20430 (HO- LOTYPE: US; ISOTYPES: GH, NY).

Clibadium alatum H. Rob., Phytologia 73: 151-152. 1992. TYPE: ECUADOR. Carchi: environs of Maldonado, 3 Jun 1978, Madison, Plowman, Kennedy, & Besse 4940 (HOLOTYPE: US; ISOTYPE:

F).

Arching branched shrubs, to 4 m tall, one main stem at base in juvenile plants, 2.5- 3.5 cm diam., branched at base in older plants; stems many-angled, often flattened near new branches, densely short strigose, hairs 1 mm long. Leaves petiolate, winged, blades ovate, 14-23(-30) X 8-15(-23) cm, apices acuminate, bases truncate-rounded, shortly decurrent on the petiole, margins serrate to irregularly serrate, both faces strigillose, hairs 1 mm long; petioles sulcate above, marginate by the decurrent blades, 3-6 cm long, strigillose, hairs 1 mm long. Capitulescences paniculiform, with many loose clusters of capitula, usually ternately divided on peduncles to 6 cm long. Capit- ula 5-6 mm high, 3-4 mm diam.; phyllaries 3-4, 4 X 2.5-3.5 mm, obovate, 5-7 veined, adaxially whitish, abaxially green, apices acute, strigose, hairs 1 mm long, distal one-third margin ciliate. Pistillateflorets 9, subtended by ovate paleae, 3.5-4 X 2.5- 3 mm, strigose, hairs 0.8 mm long, apices acute, 5-nerved; corollas 2 mm long, 0.8 mm diam., white, glabrous, 3-lobed, lobes deltoid, 0.5 mm long, apices scarcely pubescent; styles 1.5 mm long, branches 1 mm long. Staminateflorets 8-14, subtended by lanceolate (occasionally, 1-2 epaleaceous central florets), paleae, 3-3.5 X 1.5 mm, apices scarcely ciliate, 3-veined; corollas 1.8-2 mm long, 1 mm diam., white, 4-lobed, lobes deltoid, 0.5 mm long, apices scarcely pubescent; anthers 1 mm long, black; ovaries 2 mm long, 1.5 mm diam., pilose toward apex, hairs 1 mm long. Cyp- selae 2.5 X 1.5-2 mm, pilose toward distal one-third, hairs 1 mm long. Chromosome


number, n = 16 (Stuessy & Arriagada, 1993).

Distribution, ecology, and phenology. Ecuador, widely distributed in most of the central provinces; in tropical wet forests, common along streams; 600-1800 m; flowering from May to October.

Selected specimens examined: ECUADOR. Azuay: 18 km E of Jesus Maria, border with Caniar, 12 Apr 1980, Gentry et al. 28484 (MO). Bolivar: Balsapampa,

Harling et al. 9680 (F). Carchi: Maldonado-Tulcan

rd., Km 2-6, Werling & Leth-Nielsen 84 (F); 62.9 km W of Tufino, ca. 2 km E of Maldonado, 20 Jul 1992, Panero & Clark 3039 (OS). Chimborazo: foothills of the western cordillera nr. village Bucay, 8-15 Jun 1945, Camp E-3826 (MO, NY); N edge of Bucay on rte. to Teresita, 26 Jul 1977, Stuessy et al. 4942 (OS). Cotopaxi: foothills above Valencia near cane mill, 20 Jul 1960, Mathias et al. 5191 (F). El Oro: Monos, 14 km WNW of Zaruma, 15 Feb 1945, Fosberg et al. 22969 (US); 40.6 km ENE of La Avanzada on gravel rd. to Las Pifias, 4 Sep 1979, Stuessy & Nesom 5870 (OS). Esmeraldas: Ventanas, Km 319 on Quito, San Lorenzo, I Aug 1967, Jdtiva et al. 804 (UC, NY). Guayas: foothills of the western cordillera near the village of Bucay, 8-15 Jun 1945, Camp E-3826 (GH, K, MO, NY, US). Le6n: Cant6n Pijili, Hacienda Sa- lento, 18 Nov 1934, Mexia 6696 (LL, US). Los Rios: Rio Palenque Biological Station, Km 56, rd. Quevedo- Santo Domingo, 2 Oct 1976, Dodson et al. 6385 (OS).

Pichincha: Palmitopanba, ca. 10 km NW of Nanegal, 23 Jan 1974, Harling et al. 11525 (MO); 13.5 km NW of Nono on rd. to Mino, 15 Aug 1991, Stuessy & Arriagada 12352 (OS). Zamora: 4 km NE of Cum- baratza on rd. to Zumbi, 6 Sep 1979, Stuessy & Nesom 5901 (OS).

Clibadium laxum has arching branched at base stems, large leaves in old individuals (30 X 23 cm wide; e.g, Camp E-3826, Stuessy et al. 4942, Werling & Leth-Nissen 84), capitulescences with open clusters and loosely arranged capitula, and drupe-like cypselae that produce a deep orange juice when squeezed. Clibadium laxum is closely related to C. sodiroi (which has a more easterly distribution in the low humid tropical forests, between 100 and 1000 m). Both species have 3-4, ovate, acute, strigose phyllaries, 3-lobed corollas in pistillate florets, 4-lobed corollas in staminate florets, and cypselae with the distal one-third pilose. Clibadium laxum differs from C. sodiroi by having 9 (vs. 3-5) pistillate florets and staminate florets subtended by paleae. Robinson (1992) described Clibadium alatum on material with winged petioles from Carchi Province, Ecuador. After studying

the types and other specimens, I find that this represents only one extreme of the morphological variation within C. laxum. Ex- pressions of the 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 cypselae in C. alatum is similar to that observed in C. laxum (e.g., Dodson et al. 7589; Harling et al. 9680; Mexia 8467). The protologue of C. laxum gives the involucre as having 1-2 phyllaries; my reexamination of the type of C. alatum indicates 3-4 phyllaries the typical condition in C. alatum.

CLIBADIUM SODIROI Hieron., Bot. Jahrb. Syst. 29:37. 1900. TYPE: ECUADOR. Pi- chincha: prope Canzacoto, crescit locis asperis regionis subtropicae et subandi- nae, ca. 1894, Sodiro 22/2 (LECTOTYPE, here designated: US; photos: F, GH, MO, OS).

Clibdlium mexiae S. F Blake, J. Wash. Acad. Sci. 28: 489. 1938. TYPE: ECUADOR. Napo-Pastaza: near Puyo, dense forest, 360 m, 20 Feb 935, Me- xiai 6949 (HOLOTYPE: US; ISOTYPE: LL).

Clibaidium pallidum Diels, Notzbl. Bot. Gart. Berlin- Dahlem 14: 340. 1939. TYPE: ECUADOR. Napo- Pastaza: bei Puyo, Feb 1938, Sydow 807 (HOLO-

TYPE: B, destroyed).

Shrubs and trees, to 6 m tall; stems to 5 cm diam., densely tomentose, hairs, gray, 2 mm long. Leaves petiolate; blades below capitulescences ovate, 30 X 20-25 cm, apices acuminate, bases broadly rounded or slightly cordate, some slightly decurrent on the petiole, 3-5 pliveined, margins crenate- dentate; blades from branches, ovate, 6-15 X 5-10 cm, apices falcate-acuminate, bases subtruncate or shortly cuneate; densely pilose, hairs 1-1.5 mm long; margins crenate- toothed. Capitulescences paniculiform, con- vex or flattish at twig tips, many capitula (to 219), branches densely and griseously tomentose-pilose with spreading hairs, hairs 1.5-2 mm long. Capitula small, obovoid- subglobose, 3-4 mm high, 2-3 mm diam.; phyllaries 3, 3-3.5 X 3 mm, whitish, ovate, 5-7 veined, strigose, hairs 1 mm long, apices acute, margin shortly ciliate on distal


one-third. Pistillate florets 3(-5), subtended by suborbicular paleae, 3-3.5 X 3 mm, 5- veined, apices obtuse, margin shortly ciliate at distal one-third; corollas 2 mm long, 1 mm diam., white, tubulate, subequally 3- lobed, lobes irregularly deltoid, 0.5 mm long, apices scarcely pubescent; styles 1.5 mm long, branches 0.6-1 mm long. Sta- minate florets 6, epaleaceous; corollas 2 mm long, 1 mm diam., white, 4 lobed, lobes 0.5 mm long, apices shortly pilose; anthers 1 mm long, black; ovaries 2 mm long, 1- 1.5 mm diam., pilose toward apex, hairs I mm long. Cypselae 2.5 X 2 mm, broadly obovate, rounded at base, densely short pilose at distal one-third, hairs 0.5-0.8 mm long. Chromosome number unknown.

Distribution, ecology, and phenology. Ecuador; in tropical low humid forests, on steep slopes, and roadsides; 300-1000 m; flowering from June to September.

Selected specimens examined: ECUADOR. Napo- Pastaza: N from El Chaco, Quito-Lago Agrio rd., 20 km E of Baeza, 22 Jul 1986, Gentry & Miller 54996 (MO); forest at Puyo, 16 Feb 1953, Prescott 404 (DS, NY); eastern foot-hills of the Andes, Aug 1939, Skutch 4402 (A, F, GH, K, MO, NY); W edge of Puyo along roadside, I Sep 1977, Stuessy & Nesom 5817 (OS); 2 km N of Archidona on rd. to Baeza, 18 Aug 1991, Stuessy & Arriagada 12370 (OS). Pichincha: Carre- tera Quito-San Juan-Chiriboga-Empalme-Sto. Do- mingo de Los Colorados, 25 Sep 1986, Zak 1318 (GH). Santiago-Zamora: near M6ndez, 5-6 Nov 1944, Camp E-884 (NY). Tungurahua: Rio Verde, no date, Harling et al. 10153 (F); along rd. to Tena, ca. 7 km N of Puyo, 22 Jan 1974, King et al. 6567 (MO, US).

Average mature individuals of Clibadium sodiroi have very soft pubescence on leaves and stems. Mature fruits also produce a small quantity of orange juice. Stems and capitula also sometimes produce small amounts of orange sap (pers. field observ.). The large leaves (20 X 16 cm) 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 capitula (3-4 mm high) per capitulescence: 219 capitula have been observed in young individuals (Stuessy & Arriagada 12363); 819 capitula were counted in one mature and highly branched individual (Stuessy & Arriagada 12362).

Clibadium sodiroi is closely related to C.

laxum; it differs in having 3-5 pistillate florets and 6 staminate florets without paleae vs. 9 pistillate florets in C. laxum arranged in two series, and 8-14 staminate florets subtended by paleae.

CLIBADIUM SURINAMENSE L., Mant. P1. 294. 1771. (Fig. 2). TYPE: SURINAME, exact locality unknown, 1843, Hostmann 647 (NEOTYPE: K, designated by Hind in Jar- vis et al., 1993; ISONEOTYPE: NY). Trixis aspera Sw., Prodr. I15. 1788. TYPE: FRENCH

GUYANA, habitat "Caiennae & Guianae locis inclutis," 1762-64, Aublet s.n. (HOLOTYPE: BM; photos: GH, OS; ISOTYPE: G-DC, IDC 28.925: 1.5,6).

Clibadium asperumn (Aubi.) DC., Prodr. 5: 506. 1836. Baillieria aspera Aubl., Hist. PI. Guianae 2: 804, t. 317. 1775. Oswalda baillerioides Cass. Dict. Sci. 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).

Clibadiun villo.sum Benth., P1. Hartw. 2: 205. 1845. TYPE: COLOMBIA. Bogota: near the village of Tena, Feb-Apr 1843, Hartweg 1139 (LECTOTYPE, here designated: K; ISOLECTOTYPES: BM, photo: OS; K, photo: OS; G, photos: F, MO, OS; US, photos: GH, OS, US-2).

Clibadium lehmannianum 0. E. Schulz, Bot. Jahrb. Syst. 46: 620. 1912. TYPE: COLOMBIA. Playa, Rio Timbiqui, 1910, Lehmann 9056 (LECTOTYPE:

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 L. var. macrophyllum Ste- yerm., Fieldiana, Bot. 28: 629. 1953. TYPE: VEN- EZUELA. Sucre: Cerro Turumuquire, north-fac- ing slopes between La Trinidad and Quebrada El Boquer6n, SW of Cocollar, 3 May 1945, Steyer- mark 62437 (HOLOTYPE: F; ISOTYPE: US).

Shrubs, 1.5-4 (5) m tall; stems 1-2 cm diam. at base, hispidulous to scabrous, hairs 0.1 mm long. Leaves petiolate; blades lanceolate to broadly ovate, rough chartaceous, 5-12(-20) X 2-6(-12) cm, apices acute to acuminate, bases obtuse (sometimes attenuate or slightly decurrent to petiole), margins serrate, crenate-serrulate or serrate- dentate, adaxial surface weakly to moderate hispidulous, hairs 0.5 mm long (occasionally with close erect hispid pubescence),


r,3

I /~~~~~~~~~

A V FIG. 2. Clibadium surinamense L. (Croat 10920): A. Habit (X 1/2). (Rosario 31): B. Outer floret (X10 3/5);

C. Head (X3 1/5). [After Ann. Missouri Bot. Gard 62: 1076. 1975]

dark green, abaxial surface strongly hispidulous, pale gray green; petioles 1.5-5 cm long, hirsute, hairs 0.2-0.5 mm long. Ca- pitulescences thyrsoid or paniculiform, with 10-80 capitula. Capitula sessile, obovoid,

3-5 mm high, 3-4 mm diam., yellowish; phyllaries 3-4, coriaceous, concave, broadly ovate, 4 X 3-5 mm, apices acute, strigose, hairs 0.5-0.8 mm long, margins distal one-third ciliate, 5-veined. Pistillateflorets


3-5, subtended by ovate paleae, 4 X 3-4 mm, 5-veined, apices acute, strigose, hairs 0.5 mm long, margins distal one-third ciliate; corollas 2 mm long, 0.5 mm diam., 3- lobed, lobes irregularly deltoid, 0.4 mm long, scarcely pubescent at apex; styles 3 mm long, branches 1.5 mm long. Staminate fiorets 10-14, epaleaceous; corollas tubular, 3 mm long, throats 2 mm long, 0.5 mm diam., 4-lobed, lobes irregularly deltoid, 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 2 mm long, 0.5 mm diam, villous at apex, hairs 0.5 mm long. Cypselae 2.5 X 2 mm, villose on upper half. Chromosome number, n = 16 (Stuessy & Arriagada, 1993).

Distribution, ecology, and phenology. Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Guy- ana, Suriname, French Guiana, Ecuador, Brazil, and introduced to Borneo, Java, Su- matra, and Mautitius; 0-1000(-2000) m; flowering throughout the year.

Selected specimens examined: GUATEMALA. Iza- bal: 22 Jul 1977, Croat 41776 (MO).

HONDURAS. El Paraiso: entre El Junquillo y Teu- pasenti, 26-27 Apr 1963, Molina 11863 (F, NY). Gra- cias a Dios: caserfo de Rus-Rus, 21 Jul 1977, Nelson & Romero 4078 (MO). Morazan: Cordillera de Mi- sico o Volcdn de Guaimaca, 15 Jun 1950, Molina 3148 (F, GH, US). Olancho: poblaci6n de Culmi, 17-22 Jul 1978, Nelson & Romero 4653 (MO).

NICARAGUA. Boaco: Cerro Mombachito, 29 Nov 1983, Aranda et al. 32 (OS). Chontales: rd. from Jui- galpa to Santo Tomas, 6 Jul 1976, Neill 7394 (GH, OS). Granada: forest on Mombacho Volcano, 5 Jan 1967, Williams 20029 (WIS). Jinotega: N slope of Volcdn Yali, 25 Oct 1979, Stevens 15140 (F, OS). Ma- driz: ca. 5 km SW of San Juan de Rio Coco, 29 Jun 1980, Stevens et al. 17702 (F, OS). Matagalpa: carretera al Tuma, 10 km NE ciudad de Matagalpa, 8 Sep 1980, Guzma'n et al. 701 (CAS, F). Nueva Segovia: 3.7 mi S of Los Mancos on Rt 15, 7 Jan 1978, Jansen & Harriman 547 (OS). Rio San Juan: 2 km NW of Sabalos, 23 Feb 1984, Moreno 23298 (OS). Zelaya: ca. 5 km S de Waslala, 16 Sep 1982, Grijalva & Mo- reno 1180 (F).

COSTA RICA. Alajuela: ca. 9 mi N from Zarcero on Rte. 15, 1 Nov 1976, Stuessy & Gardner 4461 (OS). Cartago: S slope Irazu Volcano, near Buenos Aires, 23 Aug 1990, Arriagada 142 (OS). Guanacas- te: Guayabo on rd. to Aguas Claras, 20 Jul 1977, Al- meda et al. 3145 (OS). Heredia: rd. to Puerto Viejo, Km 54 from Barva, 28 Aug 1990, Arriagada 163 (OS). Limon: from Guapiles to San Jos6, 14 km before Guapiles, 5 Sep 1991, Arriagada 310 (OS). Puntare-

nas: 5 km before Quepos, Montes del Aguacate, 17 Aug 1990, Arriagada 131 (OS). San Jose: La Palma, rd. to Dominical, 30 Jul 1977, Almeda et al. 3304 (OS).

PANAMA. Canal Zone: Barro Colorado Island, Wheeler Trail, Hood 977 (F); Pipeline Road, 1 km N of Rio Agua Salud, 26 Sep 1982, Schmalzel et al. 1023 (MO). Chiriqui: 15 km ESE of Concepci6n on Hwy. 1, 1 Jul 1977, Hartman 3926 (OS); El Llano de Hi- cacao, 3 Jul 1984, Schmalzel 1989 (MO). Cocle: Llano Bonito N of Las Margaritas, 26 Jul 1955, Seibert 519 (K, NY); 1.3 km NW of Ant6n, 3 Jul 1976, Hartman 3930 (OS). Col6n: vic. Sardinella, Blum & Tyson 487 (FSU). Darien: vic. Cana gold mine, 29 Jul 1976, Croat 37982 (MO). Herrera: 5 mi S of Oc6, 1 Jun 1970, Wilbur et al. 12084 (CR, MO). Los Santos: 3 km S of Macaracas, 11 Jul 1976, Hartman 3942 (OS). Panama: La Campana, Cerro Campana, 8 Jul 1960, Ebinger 362 (US); vic. of TV tower on Continental Divide, Cerro Pefion, 18 Dec 1973, Nee 8890 (OS). Veraguas: 18 km W of las Minas, 8 Aug 1978, Ham- mel 4353 (MO); 8 km NE of Sona, 4 Jul 1976, Hart- man 3932 (OS).

COLOMBIA. Antioquia: 8 km de Sabanalarga, 8 Apr 1986, Callejas et al. 2285 (F, MO). Bolivar: Boca Verde on Rfo Sinu, 13-14 Feb 1918, Pennell 4219 (NY). Boyaci: Sierra Nevada del Cocuy, near Bachira, 25 Aug 1957, Grubb et al. 715 (MSU). Caldas: near La Dorada, 29 Oct 1936, Haught 2124 (US). Caqueta: Rio Orteguaza, Quebrada de Miramar, Jan 1969, Cua- trecasas & Soderstrom 27121 (US); 6 km SE Floren- cia, 9 Jan 1974, Davidse et al. 5649 (OS). Cauca: Chisqufo, Finca Los Derrumbos, Asplund 10544 (US); 49 km SW of Popayan on rd. to Pasto, 2 Aug 1979, Stuessy & Funk 5748 (OS). Choc6: Cerro del Torra, 8 Aug 1982, Silverstone 1254 (MO); 4 km from Pacho on rd. to Zipaquira, Stuessy et al. 5571, 14 Jul 1979 (OS). Cundinamarca: Km 5 E of La Mesa on hwy. to Bogota-La Mesa, 27 Aug 1991, Arriagada 304 (OS); Sueva y Gacheta, Cuatrecasas & Jaramillo 11994-A (COL, F). Huila: Hacienda Pensilvanica, Lit- tle 8114 (COL, US). Magdalena: Alto Rio Buritaca. Alto de Mira, 14 Jul 1989, Madrinan & Barbosa 237 (GH). Meta: carretera de Bella Vista a Pifialito, Eche- verry & Jaramillo 2322 (COL). Narifio: Cordillera Oriental, regi6n del Sarare, La Cabuya, 12-14, 17, 24 Oct 1941, Cuatrecasas et al. 12086 (GH). Quindio: 24 km SW of Armenia on rd. to Caicedonia, 5 Aug 1979, Stuessy & Funk 5782 (OS). Santander: region de Ocania: entre La Maria y Jurisdicciones, 3 Mar 1969, Cuatrecasas & Rodriguez 27933 (F); 15 km NE of Bucaramanga on rd. to Pamplona, 17 Jul 1979, Stuessy & Funk 5604 (OS). Tolima: 40 km E of Ar- menia on rd. to Girardot, 5 Aug 1979, Stuessy & Funk 5783 (OS). Valle: 13 km S of Dagua on rd. from Lo- boguerrero to Cali, 1 Aug 1979, Stuessy & Funk 5727 (OS). Vaupes: Rio Kuduyarf, middle and lower course, 16 Oct 1961, Schultes & Cabrera 17861 (US).

VENEZUELA. Amazonas: Atabapo, N Rio Ori- noco and W Rio El Ej6rcito, Gua'nchez 2088 (MO). Anzoategui: Mendocero, orilla del Rio Chive, Pittier 15061 (US). Aragua: above Guayas, on rd. to Los Teques, Apr 1944, Steyermark 56890 (F, US). Bolivar: La Gran Sabana, jct of rd. Santa Elena-Cabanayen, 3


Dec 1973, Davidse et al. 4748 (MO). Distrito Fed- eral: sabana de Monte, 15 Sep 1940, Vogl 708 (F, GH). Merida: arriba Santo Domingo, Aug 1958, Ariste- guieta 3268 (K, NY). Miranda: Quebrada de Las Co- madres, near Las Mostazas, Nov 1934, Allart 230 (NY, US). Monagas: El Paramo, NE Las Delicias, Apr 1945, Steyermark 62029 (F, MO). Sucre: Peninsula de Paria, Steyermark & Liesner 120954 (MO).

GUYANA. East Coast, Water Conservancy, SE of Georgetown, 25 Nov 1919, Hitchcock 16875 (GH); Kaieteur National Park, 25 Jan 1987, Pipoly & Ghar- barran 9962 (MO); forest of the Agric. Exp. Sta., Gro- ningen, Samuels 345 (GH).

SURINAME. Potaro-Siparuni Region, 12 Oct 1987, Kvist et al. 230 (F); Paramaribo, near Agric. Exp. Sta., 2 Jun 1921, Lanjouw 49 (GH).

FRENCH GUIANA. Cayenne: Broadway 376 (GH, US); Montagne des Chevaux, 27 Jul 1987, Gentry & Morawetz 63181 (MO).

ECUADOR. Bolivar: Balsapampa, Jun 1969, Har- ling et al. 9670 (F, MO). Cafiar: ca. 77 km ESE of Guayaquil, 22 Jun 1976, King & Garvey 6871 (CAS, MO, US). El Oro: 48.5 km ENE of La Avanzada to Pinias, 4 Sep 1979, Stuessy & Nesom 5873 (OS). Es- meraldas: Quininde to Esmeraldas, Holm-Nielsen et al. 7070 (F). Guayas: vic. of Naranjito, Camp 3601, 6-7 Jun 1945 (OS); Milagro, 30 Jun-2 Jul 1923, Hitchcock 20261 (GH, NY); Naranjal, jct. with rd. to Guayaquil, 25 Feb 1976, King & Garvey 7005 (CAS, MO, US). Imbabura: Lita, Acosta Solfs 12285 (F). Loja: 24 km S of Catamayo, rd. to Cariamanga, King & Almeda 7951 (CAS, MO). Los Rios: Hacienda Cle- mentina on Rio Pita, Asplund 5389 (F); Cant6n Vinces, Km 70 Quevedo-Palenque, 2 Oct 1976, Dodson & Gentry 9817 (OS). Manabi: 13 km E of Alajuela, rd. to Quevedo, 24 Jul 1979, Stuessy et al. 4919 (OS). Napo: San Jos6 de Payamino, 40 km of Coca, Sept 1982, Irvine 363 (OS). Tungurahua: La Merced, Val- ley of Rio Pastaza, 10 Jul 1939, Asplund 7614 (A, F); bridge over Rfo Blanco, 30 Jul 1977, Stuessy & Jansen 4954 (OS). Zamora-Chichipe: near Zamora, 1 Apr 1966, Knight 711 (WIS).

BRAZIL. Acre: Rio Branco-Serra Madureira rd., 33 km from Rio Branco, 29 Sep 1980, Lowrie et al. 265 (GH). Amazonas: Rio Branco, Campus Univer- sitario, 13 May 1980, Coelho et al. 1738 (F). Sao Pau- lo: Rio Solimoes, Barrio de Odorio, 17 Aug 1973, Lle- ras et al. P1 7376 (F, K); Rio Branco, below Serra Sa- bang, 16-18 Dec 1954, Maguire & Maguire 40336 (NY).

BORNEO: Kalimaritan Selatan, near Martaoura, 24 Feb 1979, Murata et al. 4451 (A); South Kalimaritan along rd. from Simpomg, 10 Nov 1996, Kessler et al. 1772 (A).

JAVA. Nirmala Estate, Kalimun area, Balgooy & Wiriadinata 2903 (A); Batavia, ad ripas fluminis Tji- liwong, Schiffner 2819 (K); Gunnong Salak, Sinclair 10052 (K).

SUMATRA. Toba: Toetoepan, 4-11 Nov 1933, Boeea 5901 (A); vic. of Tomoean Dolok, Asahan, 10- 15 Jun 1936, Boeea 9055 (A); Riau Prov., Tigapulu Mts., 27 Nov 1988, Burley et al. 2000 (A).

MAURITIUS. Perrier Nature Reserve near Mare aux Vacoas, Lorence 1852 (K).

Clibadium surinamense is distinguished by ovate to ovate-lanceolate leaves with well marked, reticulate, strongly hispidulous venation of the abaxial surfaces. Cli- badium surinamense is the most common species of the genus and is broadly distributed from Nicaragua to Ecuador. Size, shape, and pubescence of the leaves show considerable variation. The variety macrophyllum was described by Steyermark (1953) to include Venezuelan specimens with very large leaves (e.g., 17 X 10 cm wide, Steyermark 56567; 20 x 11 cm wide, Bernardi 361). Large leaves are found in plants scattered along the Andean Cordil- lera, without any particular distributional pattern. Clibadium villosum, described by Bentham (1845) from Colombian material, is clearly an excessively tomentose variant of C. surinamense. Lanceolate leaves also occur (C. lanceolatum Rusby), as well as densely strigose leaves (C. lehmannianum 0. E. Schulz). Such variation is scattered among individuals mainly at higher elevations.

Since Schulz (1912) synonymized C. sylve,s tre with C. surinamense, confusion has existed with the application of the names vurinamense and asperum. My observations of the type specimens of Baillie- ria 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 sheet labelled Baillie- ria svylvestris corresponds to the type of B. asperca, which clearly falls within C. surinamense.

Clibadium surinamense is morphologically close to C. peruvianum; the former differs by having 5-veined phyllaries, 10- 14 staminate florets, 4-lobed staminate corollas, and villose cypselae. Clibadium surinamense is distributed from Nicaragua to Ecuador; C. peruvianum is restricted to Peru, Bolivia, and Argentina.

CLIBADIUM PERUVIANUM Poepp. ex DC., Nov. Gen. Sp. Plant. 3: 1. 1845. TYPE: PERU, Amazonas, 1832, Poeppig 3.5 (HOLOTYPE: BM, photos: GH, OS; iso- TYPES: BM; P, photo: OS; G-DC, IDC microfiche 28.925:1.1).


Clibadium remotiflorum 0. E. Schulz, Bot. Jahrb. Syst. 46: 621. 1912. TYPE: BOLIVIA. Cochabam- ba, 1891, Bang 1203 (LECTOTYPE, here designated: K).

Clibadium heterotrichum S. F Blake, Contr. Gray Herb. 3 (52): 3. 1917. TYPE: BOLIVIA. La Paz: Polo-Polo, near Coroico, Oct-Nov 1912, Buch- tien s.n. (HOLOTYPE: B; photos: GH, OS; ISOTYPES:

K; MO, photo: OS). Clibadium psilogynum S. F Blake, J. Wash. Acad.

Sci. 21: 329-330. 1931. TYPE: PERU. Cuzco: Quispicanchi, Marcapata Valley, near Chilechile, 21 Feb 1929, Weberbauer 7864 (HOLOTYPE: US; ISOTYPES: F, GH).

Clibadium vargasianum H. Rob., Wrightia 6: 46. 1979. TYPE: PERU. Madre de Dios: Manu, Car- b6n Salvaci6n, 24 Nov 1965, Vargas 16928 (HO-

LOTYPE: US; photos: CAS, F, GH).

Shrubs, 2-3 m tall; stems striate, purplish with green spotting, branches densely strigose, hairs 1 mm long. Leaves petiolate; blades ovate-lanceolate to ovate, 8-10 X 3- 7 cm; apices acute, bases attenuate (round- cuneate), margins finely irregularly serrate, 3-veined, veins yellowish, strigose-pubescent on both surfaces, hairs 0.5-1 mm long; petioles 1-3 cm long, strigose, hairs 1 mm long. Capitulescence paniculiform, terminal, with capitula short-pedicellate. Capit- ulum subglobose, 5 mm high, 4 mm diam.; phyllaries 3, broadly ovate, 4.5-5 X 3.5-4 mm, apices acute, 7-veined, margins ciliate on distal one-third, strigose-pubescent on abaxial side, hairs 0.5 mm long, light-green with fine green dark lines. Pistillate florets 5, subtended by ovate paleae, 4-4.5 X 3.5- 4 mm, 5-veined, apices acute, strigose, hairs 0.5 mm, margins distal half ciliate; corollas white, tubulate, 3 mm long, 1 mm diam., 3- lobed, lobes deltoid, 0.5 mm long, apices scarcely ciliate; styles 2 mm long, branches 1 mm long. Staminate florets 19, epaleaceous; corollas tubulate, white, 3 mm long, 1 mm diam., 5-lobed, lobes 0.5 mm long, scarcely ciliate at apex; anthers black, 1.5 mm long; style 1 mm long, undivided, ovaries 1.5-2 mm long, 1-1.5 mm diam., glabrous to scarcely pubescent at apex, hairs 0.2-0.5 mm long. Cypselae subglobose, black, 2 X 1.5 mm, glabrous, slightly pa- pillose on distal one-third. Chromosome number unknown.

Distribution, ecology, and phenology. Peru, Bolivia, and Argentina; growing in secondary lowland tropical wet forests,

open and grazed selva, riverbanks, and sea- sonally inundated areas; 100-600(-1200) m; flowering throughout the year.

Selected specimens examined: PERU. Amazonas: Bagua, ca. 5 km S La Peca, 27 Oct 1978, Barbour 4299 (F). Ayacucho: Ocarrapa, 10-11 May 1929, Kil- lip & Smith 22450 (LP). Cuzco: Paucartambo, Atala- ya, Vargas 13983 (US). Huinuco: Pachitea, Honoria, Bosque Nacional de Iparia, 23 Feb 1967, Schunke 1672 (F, OS, US). Junin: ca. 18-24 km N of San Ra- m6n, 20 Dec 1978, Dillon & Turner 1454 (F, MO, OS). Loreto: Maynas, camino a Santa Maria, Sep 1978, Ayala 501 (MO); Rio Putumayo, at mouth of Rio Zubineta, Oct-Dec 1931, Klug 2348 (A, F, GH, MO, NY, US); vic. of Iquitos, 22 Jul 1972, Croat 18301 (OS). Madre de Dios: Tambopata, Puerto Mal- donado, 28 May 1980, Barbour 5386 (F); Rio Alto Madre de Dios, between Shintuya and Boca Manu, 26 Oct 1979, Gentry el al. 27259 (F, MO). Pasco: Oxa- pampa, 2-4 km N of Mallampampa, 22 Jan 1984, Smith & Canne 5830 (F). San Martin: San Martin, near Tarapoto on trail to Juan Guerra, 25-27 Aug 1937, Belshaw 3316 (GH, LL, MO, NY); Juanjui, Sha- paja, 8 Oct 1977, Johns & Ramfrez 239 (OS). Santa Cruz: Cant6n Buenavista, 29 Jun 1916, Steinbach 2786 (A).

BOLIVIA. Arce: Reis Bolivia, June 1886, Rusby 2145 (NY); 7.4 km E of Emborozd on rd. to Bermejo, Jan 1983, Solomon 10025 (MO). Chapare: Cocha- bamba, Locotal, 2 Feb 1929, Steinbach 9018 (GH, MO). Murillo: La Paz, valley of the Rio Zongo, 22 Apr 1982, Solomon 7515 (MO). Pando: Nicolks Sud- rez, ca. of Cobija on Rio Acre, 7 Jan 1983, Fernandez- Casas & Susanna 8048 (MO).

ARGENTINA. Jujuy: Ledesma, Rio Agua Negra, Logname & Cuezzo 7534c (LP). Tucuman: 4 May 1985, Solomon 13577 (MO); Famaillk, Quebrada de Lules, Feb 1921, Venturi 1180 (GH; LP).

Clibadium peruvianum is easily recognized by the yellowish veins on the leaf abaxial surfaces; striate and green and purple spotted stems; dark green, striated phyllaries; and open panicles. Capitula and the capitulescences are as in C. surinamense but the leaves are like those of C. sylvestre. Clibadium peruvianum is closely related to C. surinamense; the former differs by its strigose 7-veined phyllaries, 5 pistillate and 19 staminate florets, 5-lobed staminate corollas, and glabrous cypselae vs. 5-veined phyllaries, 3-5 pistillate and 10-14 staminate florets, 4-lobed staminate corollas, and villous cypselae in C. surinamense. Cliba- dium surinamense ranges from Nicaragua to Ecuador; C. peruvianum is found in northern Argentina, Bolivia, and Peru.

Clibadium psilogynum was based on material collected from the Cuzco region of


Peru. The three type specimens represent one extreme of morphological variation observed in C. peruvianum involving larger leaves (15 cm), irregularly serrate margins, fewer staminate florets, and, sometimes, 1 or 2 staminate florets each subtended by a small pale.

Clibadium heterotrichum was based on the harsh spreading pubescence of the stems and larger capitula (6 mm). My examination of many specimens indicates that the pubescence and floral variations of C. heterotrichum grade into those of C. peruvianum and vitiate specific recognition.

CLIBADIUM ARBOREUM Donn. Sm., Bot. Gaz. 14: 26. 1889. TYPE: GUATEMALA. Alta Verapaz: Pansamala, Jun 1886, von Tiirckheim 929 (HOLOTYPE: US; ISOTYPES: GH; K; NY-2; US, photos: OS, US).

Clibadium donnell-smithii J. M. Coult., Bot. Gaz. 16: 98. 1891. TYPE: GUATEMALA. Guatemala, Feb 1890, Smith 2347 (HOLOTYPE: US; ISOTYPE:

GH, 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. (HOLOTYPE: US-1038790; ISOTYPE: GH).

Clibadium oligandrum S. F Blake, Brittonia 2: 342. 1937. TYPE: GUATEMALA. Quezaltenango: Vol- can Zunil, 3 Aug 1934, Skutch 927 (HOLOTYPE: A; ISOTYPES: BM, photos: F, GH, NY, OS, US; GH).

Shrubs or small trees, to 6 m tall; stems densely branched, branches densely pilose, hairs 1 mm long. Leaves petiolate; blades ovate to broadly ovate, 10-20 X 8-18 cm, membranaceous, apices acuminate or long acuminate, bases more or less rounded and then cuneate, 3-veined far above the base, margins serrate; adaxial surface scabrous, abaxial surface densely short pilose, hairs 0.5 mm long. Capitulescences paniculiform, with small or large cluster of capitula, leafly at base. Capitula subglobose, sessile or shortly pedicellate, 5 mm high, 3.5 mm diam.; phyllaries 3(-4), broadly ovate, striate, 5 X 4-5 mm, 5-7 veined, apices acute or slightly obtuse to obtusely pointed, strigose, hairs 1 mm long, margins ciliate on distal one-third. Pistillate florets 3-6, subtended by broadly ovate paleae, 4.5 X 4 mm wide, 5-veined, strigose, hairs 0.5 mm long, apices obtuse, scarcely ciliate; corollas white, 2.5 mm long, 1 mm diam., 4-

lobed, lobes 0.2 mm long, apices scarcely pubescent; styles 2 mm long, branches 1.5 mm long, tips recurved. Staminate florets 4-10, epaleaceous; corollas white, 2-3 mm long, tubes 1 mm long, expanded into a campanulate throat, 0.5 mm diam., 5-lobed, lobes deltoid, 1 mm long, apices scarcely hirsute; ovaries 1.5 mm long, 0.5 mm diam., hirsute at distal one-third, hairs 0.5 mm long. Cypselae blackish, obovoid, 2 X 1.5 mm, hirsute at apex, hairs 0.5 mm long. Chromosome number, n = 16 (Stuessy & Arriagada, 1993).

Distribution, ecology, and phenology. Mexico, Guatemala, Belize, Honduras, and Nicaragua; on Pacific slopes, in humid dense forests in stream-valleys associated with Carpinus, Magnolia, Podocarpus, and Quercus, damp or wet secondary forest along roadsides; sea level to 2500 m; flowering from (October-) April to June (-No- vember).

Selected specimens examined: MEXICO. Chiapas: Mun. Jitotol, 7 mi N of Jitotol, 28 Aug 1966, Breed- love 15423 (F, LL, NY); Mun. Motozintla de Mendoza, near Niquivil, 16 Dec 1976, Breedlove 42714 (MO); Mun. Ocozocoautla de Espinosa SW of the Presa de Malpaso, 5 Dec 1967, Shilom Ton 3311 (F, NY); Mun. Palenque, 6-12 km S of Palenque on rd. to Ocosingo, 27 Jul 1972, Breedlove 26504 (MO); Mun. San Pedro Chenalho near the Colonia Choro, 8 Jun 1967, Shilom Ton 2496 (NY); Mun. Solosuchiapa, 3-5 km above Solosuchiapa along rd. to Tapilula, 26 Jul 1972, Breed- love 26410 (MO). Jalisco: 11-12 mi S of Talpa de Allende, 23-25 Nov 1960, McVaugh 21414 (LL, NY). Mexico: Ejido Lazaro Cardenas, 100 m del puente del ferrocarril, 16 Sep 1979, Cowan 2427 (MO). Oaxaca: S of Valle Nacional and ca. 109 rd. miles NNE of Oaxaca, 9 Oct 1962, Cronquist 9631 (GH, NY); 11.8 mi W of Tuxtepec, on rd. to Ojitldn, 16 Nov 1976, Stuessy & Gardner 4574 (OS). Puebla: 4 km NE of Villa Judrez, 27 Jun 1969, Marcks & Marcks 826 (WIS). Veracruz: Estaci6n Biol6gica de los Tuxtlas, 8 Sep 1971, Calzada 500 (F, MO); rd. from Coyame to Tabanca, 5 Aug 1965, Cruz 286 (LL); Barranca de Teoxolo near Jalapa, 5 Jul 1908, Pringle 10802 (F, GH, LL).

GUATEMALA. Alta Verapaz: ca. 6 km NE of Panz6s, 20 Jul 1977, Croat 41579 (CAS, MO). Chi- maltenango: Quisach6, 5-6 Jan 1939, Standley 62042 (US). Escuintla: Finca Monterrey, S slope of Volcan de Fuego, 5 Feb 1939, Standley 64559 (F, GH). Izabal: Puerto M6ndez, on Toquela rd., Feb 1890, Donnell- Smith 2347 (GH). Peten: La Libertad and vic., 28 Jul 1934, Aguilar 322 (MO, NY). Quezaltenango: S of San Martin Chile Verde on rd. to Colomba, 27 Jan 1941, Standley 85097 (F). Quiche: Finca Chaita, Zona Reyna, 29 Nov 1934, Skutch 1797 (GH); Retalhuleu: ca. 27 km W of Mazatenango, 16 Jun 1976, King &


Renner 7020 (CAS, MO). San Marcos: Finca El Por- venir along Rio Cabds above Potrero Matasan, 12 Mar 1940, Steyermark 37622 (F). Solola: Volcan Atitlan, 11 Jun 1942, Steyermark 47342 (F). Suchitepequez: Volcan Santa Clara, between Finca El Naranjo and upper slopes, 23 May 1942, Steyermark 46626 (F, GH).

BELIZE. Belize: 1.5 mi S of Mile 22 on Western Highway, 4-5 Jun 1973, Croat 23863 (OS). Cayo: Rio Frfo Caves near Augustine, 2 Sep 1970, McDaniel 14482 (F); Hummingbird Hwy., E of Over-the-Top Camp, Sep 1980, White.foord 2421 (F). Cortez: Mon- tania Santa Ana along Rfo Santa Ana, 6 Dec 1950, Molina 3601 (GH). Olancho: Izabal, 6-8 km S of Mo- desto Mendez, 12 Jun 1970, Harmon 2552 (GH). To- ledo: Columbia Forest Reserve, ca. 1-2 mi N of en- trance, 11 Jun 1973, Croat 24155 (OS).

HONDURAS. Cortez: along Rio Santa Ana, 6 Dec 1950, Molina 3601 (F, US). Santa Barbara: 24 km E of Santa Rita de Copan, 18 Aug 1970, Harmon & Dry- er 4031 (MO, NY).

NICARAGUA. Granada: Volcan Mombacho, I May 1975, Atwood A161 (OS).

Clibadium arboreum is the northernmost species of the genus and is characterized by being densely hirsute with soft, yellowish hairs on its leaves. It can be also distinguished by its broadly ovate leaves with finely serrate margins, rounded bases, and acuminate apices. The leafly paniculiform capitulescence is mostly wider than high and most of the young branches irregularly exceed the central (terminal) branch.

CLIBADIUM ARMANII (Balb.) Sch. Bip. ex 0. E. Schulz, Linnaea 30:180. 1859.

Eupatorium armani Balb., Hort. Taurin. Stirp. 1. 27. t.6. 1810. TYPE: Brazil, locality and date unknown, Armani s.n. (HOLOTYPE: TO, not located). [The plate in the protologue, however, is adequate to understand this taxon].

Clibadium rotundifolium DC., Prodr. 5: 505. 1836. TYPE: Brazil. Bahia: Dec 1832, Blanchet 632 (LECTOTYPE, here designated: NY).

Shrubs, 3-4 m tall; stems with branches terete, scabrid-hispid, hairs 1 mm long. Leaves petiolate; blades broadly ovate, edges recurved, 4-7 X 3-5 cm, subcoriaceous, apices acuminate and mucronate, bases rounded or sometimes cordate, margins mucronate serrate, pinni-nerved, 5-7-veined at the base, pale green on both faces, hispidulous, hairs 1 mm long; petioles 3-10 mm long, scabrid-hispid, hairs 1 mm long. Ca- pitulescences thyrsoid. Capitula ovoid, sessile, 5-6 mm high, 4 mm diam.; phyllaries 3, coriaceous, concave, broadly ovate, 4.5 X 3.5-4 mm; 5-veined; whitish green, api-

ces shortly acuminate to mucronate; strigose, hairs 1 mm long. Pistillate florets 4, subtended by broadly ovate paleae, 4 X 3.5 mm; 5-veined; apices short-acuminate; strigose, hairs 0.5 mm long; corollas white, tubular, 3.5 mm long, 1.5 mm diam., 3-lobed, lobes irregularly deltoid, 0.3 mm long, apices scarcely ciliate. Staminate florets 5-8, epaleaceous, corollas white, tubular, 4.5 mm long, 2 mm diam., 5-lobed, lobes deltoid, 0.8-1 mm long, apices scarcely ciliate; ovaries 2 mm long, 1-1.5 mm diam., hirsute, hairs 1 mm long. Cypselae broadly obovoid, 3 X 2-2.5 mm, apices hirsute, hairs 1 mm long. Chromosome number, n = 16 (Robinson et al., 1981).

Distribution, ecology, and phenology.- Brazil and Paraguay; in gallery forests, disturbed areas, on sandy soil, and at edges of wet savannas; 500-1600 m; flowering from November to March.

Selected specimens examined: Brazil. Bahia: Mun. Ilh6us, Rodovia Uruquca, 1 Jul 1965, Belem & Aguiar 1289 (NY); Mun. Santa Cruz de Cabralia, 16 Jul 1981, Brito & da Vinha 12 (US); Mun. Itabuna, 9 Jul 1979, King & Mori 8000 (MO, US); Mun. Ilheus, 4 km nr. Distrito Industrial, 10 Jul 1979, King & Silva 8008 (MO, US). Brasilia: Taguatinga Norte, 9 Mar 1980, Chagas & Silva 252 (CAS); area do Cristo Redentor, 5 Feb 1990, da Silva & Alvarenga 936 (US). Distrito Federal: Cachoeira Piripiripau, ca. 15 km S of Pla- naltina, 20 Feb 1970, Irwin et al. 26443 (F). Gokis: Chapada dos Veadeiros ca. 29 km N of Alto Paraiso, 9 Mar 1973, Anderson 6739 (F, MO, NY, US); Serra dos Pirineus, 20 km N Corumba de Goias, 9 Feb 1966, Irwin et al. 18743 (DS). Mato Grosso do Sul: Mun. Rio Brilhante Fazenda Bela Vista, Aug 1971, Hatsch- bach 26091 (UC); Campo Grande, 25 Jan 1979, Ca- brera & Zardini 29986 (OS). Minas Gerais: Brejo ca. 35 km W of Montes Claros rd. to Agua Boa, 23 Feb 1969, Irwin et al. 23772 (F, GH, MO, NY, US); Mun. Belo Horizonte, Brejo, 28 Jul 1942, Magalhaes 3183 (US); Mun. Betim, Fazenda do Cabuf, Feb 1945, Wil- liams 5119 (GH). Parana: Ibarar6, 20 Jan 1915, Du- sin 16424 (F, NY); Mun. Ibaiti, Guai, 15 Jan 1971, Hatschbach 25957 (UC); Mun. Senges, Fazenda Mo- rungava Rio Funil, 19 Jan 1965, Smith et al. 14842 (NY, US). Rio de Janeiro: Corva Grande, Itaguai, 26 Jun 1957, Duarte 4464 (US). Sao Paulo: Parque do Estado, 12 Jan 1966, Coleman 265 (US); Mun. Itape- tininga, Estacao Experimental de Itapatininga, Instituto Forestal, Giannotti 4567 (F); Jaragua, 17 Mar 1907, Usteri 16711 (NY).

PARAGUAY. Alto Parana: Oct 1909, Fiebrig 5815 (GH, US). Amambay: Sierra de Amambay, Aug 1907, Rojas 10053 (A, F); Parque Nacional Cerro Cord, 5 Jan 1988, Zardini et al. 4049 (US). San Pedro: Co- lonia Guayaibi, 20 km N of San Estanislao, 26 Nov 1968, Krapovickas et al. 14274 (UC).


Clibadium armanii has characteristic pale-green, hispidulous, subcoriceous leaves, whitish-green phyllaries, and stems and petioles tinged slightly red. Petioles are 3-10 mm long; blades broad-obovate; base more or less attenuate, rounded, or sometimes cordate; and margins regularly serrate with minute mucros. The midrib and secondary veins are conspicuously yellowish. 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), 5-7-veined, broadly ovate phyllaries, and 5-8 staminate florets with corollas scarcely ciliate at the apices of the lobes.

Clibadium armanii, however, does show variation across its distributional range. Some specimens have attenuate leaf bases (e.g., Eiten & Eiten 5880); others have more rounded to cordate bases (e.g., Kra- povickas & Cristobal 34351, whence the specific epithet of Clibadium rotundifolium was derived). Some specimens have very small leaves (2 X 1.5 cm) with a typical deltoid shape (King & Almada 8205); others have lanceolate leaves (Irwin et al. 12394); still others have large (7 X 6 cm) proximal leaves with attenuate bases and smaller leaves along the peduncles (Harley et al. 11249). There is also variation in the typical mucronate dentate margin: it is irregularly serrate to slightly erose in Eiten & Eiten 5880.

CLIBADIUM L. sect. OSWALDA (Cass.) DC., Prodr. 5: 505. 1836.

Oswalda Cass. In F Cuvier, Dict. Sci. Nat. 59. 322. 1829. TYPE: Clibadium sylvestre (Aubl.) Baill.

Leaves ovate-lanceolate; capitulescences thyrsoid; capitula paleaceous; phyllaries 7- 1 1-veined; pistillate florets 4-14, corollas 3-lobed, subtending paleae 5-7-veined; staminate florets 5-22, staminate corollas 3-, 4-, or 5-lobed, with paleae 3-veined.

CLIBADIUM LEIOCARPUM Steetz in Seem., Bot. voy. Herald 152. 1853. TYPE: PAN- AMA. Veraguas: volcano of Chiriqui, 7000 ft, Feb 1849, Seamann 1591 (HO- LOTYPE: K; photos: F, GH, OS, US; iso- TYPE: BM; photos: F, OS, US).

Clibadium schulzii S. F Blake, Contr. U.S. Natl. Herb. 22: 602. 1924. TYPE: COSTA RICA. San Jose: thickets at Copey, 1800 m, Mar 1898, Ton- duz 11915 (HOLOTYPE: US; ISOTYPES: GH, US).

Clibadium leiocarpum Steetz var. strigosum S. F Blake, J. Wash. Acad. Sci. 27: 382. 1937. TYPE:

COSTA RICA. San Jose: Cerro de Piedra Blanca, above Escazu, 31 Jan 1924, Standley 32593 (HO-

LOTYPE: US).

Shrubs, 2-5 m tall; stems usually densely branched, striate-angulate, 4-10 cm at base, strigose to densely pilose-tomentose, hairs to 1 mm long. Leaves petiolate; blades lanceolate to ovate, 5-10 X 2-10 cm, apices acuminate, bases narrowly attenuate, margins serrate; both faces hirsute-tomentose to strigose, hairs to 1 mm long; petioles 2-6 cm long, 1-3 mm diam., tomentose to tomentose-lanate, hairs 1 mm long. Capitu- lescences thyrsoid, with < 100 capitula; peduncles 0.5-1.0 mm long, densely strigose. Capitula radiate, 5 mm high, 3-3.5 mm diam.; phyllaries 3, suborbicular to broadly ovate, 3.5-4.5 X 3-4 mm, 8-9-veined, abaxially glabrous, apices acute, margins ciliate. Pistillate florets 7-9, subtended by ovate paleae, 3.5-4 X 3-3.5 mm, glabrous, scarcely ciliate at apex, 7-veined; corollas white, 1.5 mm long, 0.5 mm diam., glabrous, 3-lobed, lobes 0.4 mm long; styles 2 mm long, branches 0.5-0.8 mm long. Sta- minute florets 10-15, subtended by lanceolate paleae (occasionally 1-2 central disc florets epaleaceous), 2.5 X 1-1.5 mm, 3- veined; corollas white, 2.5-3 mm long, throats 2 mm long, 1 mm diam., 4-lobed, lobes 0.5 mm long, apices strigillose, hairs 0.3 mm long; anthers black, 1.5 mm long; styles undivided; ovaries 2 mm long, 1 mm diam., glabrous. Cypselae 1.5-1.7 X 1 mm, black, glabrous or sparsely pilose at apex. Chromosome number, n = 16 (Stuessy & Arriagada, 1993).

Distribution, ecology, and phenology. Nicaragua, Costa Rica, and Panama; in secondary wet forests, common along roadsides and on disturbed slopes; 1400-2800 m; flowering from December to September.

Selected specimens examined: NICARAGUA. Gra- nada: Volcan Mombacho, Plan de Las Flores, 17 Jan 1980, Araquistain & Moreno 655 (F), 7 Jan 1983, Gri-

jalva et al. 2109 (OS), 31 Aug 1983, Miller et al. 1422 (F), 3 Feb 1977, Neill 1365 (MSU).

COSTA RICA. Alajuela: between Atenas and Oro-


tina, a few km W Atenas toward Cerro del Aguacate, 10 Sep 1991, Arriagada 405 (OS). Cartago: San Fran- cisco 10 km after Paraiso on rd. to Turrialba, 21 Aug 1990, Arriagada 146 (OS). Heredia: from Pueblo Bar- va to San Miguel on rt 9, Km 23, 28 Aug 1990, Arria- gada 162 (OS). Puntarenas: Cordillera de Talamanca, Cotonsito, along rd. to Sitio Coto Brus, 3-4 Sep 1983, Davidse 24607 (CR); 6 km S of San Vito de Java at Finca Las Cruces, 11 Feb 1971, Gillis & PlowmGan

10162 (GH, MSU, OS). San Jose: W slope Volcan Irazui, Finca Alto Pizote, 14 Sep 1991, Arriagada 415 (OS).

PANAMA. Chiriqui: E slope of Volcan de Chiriqui, WNW of Boquete, 19 Nov 1975, Davidse & D'Arcv 10211 (OS); valley of the upper Rfo Chiriquf Viejo, vic. of Monte, 23 Jun-13 Jul 1935, Seibert 245 (GH, NY); Bajo Grande, 1-3 km E of town of Cerro Punta, 24 Feb 1974, Nee 9983 (OS).

Clibadium leiocarpum is distinguished from other species of the genus by its typically glabrous phyllaries (rarely strigillose or ciliate at the apex), black and glabrous cypselae, 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 because of more copious pubescence. C. Ieiocarpum var. strigosum was based on specimens (e.g., Standley 23593) having petioles and blades strigose and cypselae (sometimes) sparsely villose at apex. In my opinion, C. schultzii does not warrant formal recognition because the 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, Stuessy & Gardner 4456; Heredia, Weston et al. 5624).

Clibadium leiocarpum has C. sylvestre as its closest relative; the latter differs by strigose, obtuse phyllaries, which are whitish- green when dried (vs. brownish-green in C. leiocarpum), by a typically thyrsoid capitulescence (vs. paniculiform), and by the presence of an unusual rudimentary pappus of two short awns. Moreover, Clibadium leiocarpum has a longer flowering season than C. sylvestre, and is adapted to higher elevations (1400-2800 m vs. 0-600(-1600) m). Clibadium leiocarpum also lacks the acetylenic compounds isolated from C. sylvestre that are responsible for fish-poisoning (Arriagada, 1995a).

CLIBADIUM SYLVESTRE (Aubl.) Baill., Hist. P1. 8: 307. 1886.

Baillieria sylvestris Aubi., Hist. P1. Guianae 2: 807. 1775. TYPE: GUYANA, 1762-64 [Urban, 1903], Aublet s.n. (HOLOTYPE: BM; photos: GH, OS).

Clibadiurn 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).

Clibadium vargclsii DC., Prodr. 5: 506. 1836. TYPE:

VENEZUELA. Caracas, 1829, Vargas 13 (HO-

LOTYPE: G-DC, IDC microfiche 28.925:1.7; photo: A).

Bailleria barbasco H.B.K., Nov. gen. sp. pl. 4: 226. 1818 [folio ed.J. TYPE: VENEZUELA. Misiones del Orinoco, in umbrosis Javitae, 30 Apr-7 May 1800 [Sandwith, 1968], Humboldt & Bonpland s.n. (HOL.OTYPE: P, photos: F, GH, OS; ISOTYPE: P, photo: OS).

Clibadium badieri (DC.) Griseb., Fl. Brit. W.I. 368. 1864. TYPE: DOMINICA, locality unknown, 1832, Imray 145 (HOLOTYPE: K, photos: A, OS). Clibadium barbasco (H.B.K.) DC., Prodr. 5: 507.

Clibadium strigillosum S. E Blake, Contr. Gray Herb. 52: 4. 1917. TYPE: PERU. Huaca: Rio Hua- llaja, 1835, Mathews 1360 (LECTOTYPe, here designated: BM; ISOLECTOTYPES: GH, photos: GH, OS; K 2, photo: OS; P, photos: F, GH, MO, OS).

Clibadium latifolium Rusby, Desc. S. Amer. P1. 150. 1920. TYPE: COLOMBIA. Magdalena: in open places in alluvial forest on banks of River Buri- taca, 2 miles from the sea, 28 Sep 1898, Smith 2014 (HOLOTYPE: NY; ISOTYPES: F, photo: US; K, photo: OS; P, photo: OS; US, photo: US-3).

Clibadium appressipilum S. E Blake, Contr. U.S. Natl. Herb. 22: 600. 1924. TYPE: PANAMA. Da- rien: Boca de Cupe, 13 Apr 1908, Williams 698 (HOLOTYPE: US; ISOTYPE: NY, photo: OS).

Clibadium caudatum S. E Blake, Contr. U.S. Natl. Herb. 22: 600. 1924. TYPE: PANAMA. Canal Zone: alluvial bottom near Bohio, 12 Feb 1912, Maxon 4767 (HOLOTYPE: US).

Shrubs, 2-4 m tall; stems strigillose, hairs 0.1 mm long. Leaves petiolate; blades ovate to ovate-lanceolate; 8-10 X 4-12 cm, apices acuminate to caudate, bases attenuate, margins serrate, pauciserrate, dentate, or crenate-serrate; both faces strigillose (hairs usually more abundant along main veins), hairs 1 mm long; petioles 1-5 cm long, to 2 mm diam., strigillose, hairs 1 mm long. Capitulescences thyrsoid, ca. 100 capitula at end of leafy twigs; peduncles 1-2 mm long. Capitula radiate, 4-5 mm high, 3-4 mm diam.; involucres cupulate; phyllaries 3(-4), ovate to obovate, 3-4 X 2-3 mm, 7-11-veined, abaxially strigose, hairs 0.5 mm long. Pistillate florets 5-9, sub-


tended by ovate paleae, 3-4 X 2-3 mm, 7- veined, abaxially shortly strigose, margins shortly ciliate on distal one-third; corollas 1.5-2 mm long, 0.3-4 mm diam., 3-lobed, 0.4 mm long, styles 2-2.5 mm long, branches 1 mm long; reduced pappus of two short awns. Staminate florets 9-22, subtended by lanceolate paleae, 2 mm X 1 mm, 3-veined; corollas 2.5 mm long, throats 2 mm long, 4-lobed, lobes 0.5 mm long, tubes 0.3 mm diam.; anthers 1.4 mm long, styles 4 mm long, ovaries 1.2 mm long, villous (frequently only the distal one- third). Cypselae 2 X 1.5 mm, villose at apex (distal one-third). Chromosome number unknown.

Distribution, ecology, and phenology. Costa Rica, Panama, Lesser Antilles, Co- lombia, Venezuela, British Guyana, Suri- nam, French Guiana, Ecuador, Peru, and Brazil; in tropical rain forests, disturbed forests with secondary vegetation, along roadsides, also in disturbed soils, commonly cultivated; 200-600(-1600) m; flowering from May to September, occasionally as late as January.

Selected specimens examined: COSTA RICA. Car- tago: 6 miles S of Cartago on Rt. 2, 8 Nov 1976, Stuessy & Gardner 4493 (OS). Puntarenas: Herra- dura, 2 km N Rio Cania Blanca, 17 Aug 1990, Arria- gada 132 (OS); Quepos, Parque Nacional Manuel An- tonio, 28 Aug 1990, Arriagada 165 (OS).

PANAMA. Canal Zone: Barro Colorado Island, W side of Orchid Island, 21 Sep 1970, Croat 12283 (MO, NY). Chiriqui: N of Gualaca on Rfo Chiriquf, Antonio 2881 (CAS). Cocle: vic. of El Valle, N rim, Allen 224 (GH); on way to Nombre de Dios, above rd. 18 km past Sardinilla, 2 Aug 1974, Croat 26090 (MO). Co- lon: 1 mile E from Puerto Pil6n along dirt rd. to Maria Chiquita, 7 Sep 1967, Correa & Haines 234 (F, MO). Darien: vic. of Yape, 4 Oct 1938, Allen 857 (NY); near helipad at Camp Hydro on Rio Morti, 15 Mar 1968, Duke 15409 (OS). Panama: Rio Canina, near Jenine, 24 Sep 1961, Duke 3828 (CAS, MO); 5 km N of Pan. Am. Hwy. at El Llano, 10 Nov 1973, Nee 7898 (MO). San Blas: Ustopo, Oct 1975, D'Arcy 9494 (MO). Veraguas: Mountains S of Azuero Peninsula, 29 Oct 1978, Hammel 5453 (MO).

DOMINICA. Carib Territory, Windblew Hill, 11 Jan 1994, Higgins 105 (GH); Botanic Garden at Roseau, Aug 1937, Hodge 901 (GH); Carib Trail from Salybia to Hatton Garden, 30 Apr 1940, Hodge 3195 (GH).

MARTINIQUE. Ridge of Morne Rouge, 13 Mar 1979, Howard 18887 (A).

SAINT VINCENT. Mount Brisbane, 20-21 May 1947, Morton 5946 (GH).

GRENADA. Saint Andrew, vic. Of Birch Grove, 30

Oct-11 Dec 1957, Proctor 17113 (A); Saint George, Black Forest, Broadway s.n. (F).

COLOMBIA. Amazonas: Leticia, ca. 1.5 hr walk E, ca. 4 mi of Muroodoo, 5 Jul 1969, McDaniel et al. 11799 (F, FSU, MO); RIo Amazonas, Schultes & L6- pez 10400 (US). Bolivar: Mun. San Jacinto, Cerro Maco, 6 Aug 1985, Zarucchi & Cuadros 3988 (US). Caqueta: Rio Caqueta, La Pedrera, Schultes 5879 (ECON). Choc6: Pizarro, Pie de Pepe-Barrecul rd., 17 Nov 1985, Espina 1930 (MO, US); Ungufa, 9 Jun 1976, Forero et al. 1984 (COL). Magdalena: Funda- ci6n, 3 mi N Santa Rosa, 5 Aug 1971, Romero-Cas- taneda 11I55 (F, MO, NY). Nariiio: El Palmar, 25 Oct 1944, Sneidern 4522 (A, GH, LL, MO, NY, US). Pu- tumayo: Puerto Lim6n, 16 Dec 1968, Plowman 2153 (A, ECON, US). Santander: La Motilonia, frontera Colombo-Venezolana, Hoya del Rfo Oro, 15-20 May 1965, Garcfa-Barriga & Lozano 18284 (GH). Valle: 2 km W of Queremal on gravel rd. to Buenaventura, 1 Aug 1979, Stuessy & Funk 5737 (OS, WIS). Vau- pes: Rfo Piraparia, 3 Sep 1952, Schultes & Cabrera 17153 (GH).

VENEZUELA. Anzoategui: Mendocero, orillas RIo Chive, Pittier 15059 (US). Bolivar: Confluencia Ca- run, Rio Paragua, 1 Mar 1958, Cardona-Puig 2848 (NY). Carabobo: La Cumaca near San Diego, 31 Dec 1938, Alston 5905 (BM, NY, US). Merida: entre Santo Domingo-MWrida y La Mucuy, 8-12 Jan 1963, Aris- teguiet(I 4932 (F). Monagas: near Capirito, 6 Mar 1948, Tarnayo 3520 (MO). Tachira: along Quebrada Aguia AZul, S of El Reposo, Steyermark & Liesner 118741 (MO). Territorio Federal Amazonas: Ata- bapo, near Culebra on RIo Cunucunuma, 23 Feb 1985, Lies,w>r 17877 (MO).

BRITISH GUYANA. Northwest Dist.: Waini River, 3-18 Apr 1923, De La Cruz 3679 (GH). Pomeroon Dist.: Kabakaburi, 10-15 Feb 1923, De La Cruz 3289 (F, GH, MO).

SURINAME. Sandrij I, Archer 2767 (US); Carolina and vic., Archer 2893 (US); Konashen-area, Essequibo River, Saparimo, Jansen-Jacobs et al. 1819 (MO); Pa- ramaribo, Agric. Exp. Stat., Samuels s.n. (MO).

FRENCH GUIANA. Inini, Comune de Saul, Region de Saul, 1974, 18 Sep 1979, Capus 59 (MO), 22 Jul 1982, Fournet 217 (MO), Mar 1978, Moretti 896 (MO), I I Nov 1984, Riera 929 (MO).

ECUADOR. Los Rios: Pichilingue, Acosta Solis 10771 (F). Morona Santiago: basin of Rio Morona, Limb(uch 120 (CAS, MO). Napo: confluence of Qui- wado and Tiwaeno rivers, 11 Apr 1981, Davis & Yost 924 (ECON, F). Pastaza: Cabeceras Chiquita at RIo Bobonaza, 26 Jul 1980, Ollgaard et al. 35322 (MO). Pichincha: 20 km W of Santo Domingo de Los Co- lorados, 26 Oct 1961, Cazalet & Pennington 5169 (NY). Zamora-Chichipe: Pangui, 14 June 1978, Hart 1567 (A, OS).

PERU. Amazonas: Quebrada Chichijam, RIo Ca- nepa, 24 May 1973, Ancuash 461 (MO); confluence of Icikiti and Canepa, 18 Dec 1972, Berlin 625 (MO). Huanuco: Tingo MarIa-Aguaytia rd., Gentry et al. 36138 (MO); Pachitea, Honoria, Bosque Nacional de Iparia, 5 Dec 1966, Schunke 1320 (COL, OS, US). Junin: Huacapistana, on RIo Chanchamayo between Tarma and San Ram6n, 27 Jun 1976, Fosberg 56191


(MO, US). Loreto: on the Purus River at the village of San Bernardo, 1978, Rutter & Barozo-Ferreyra s.n. (OS); Quebrada Shanuce above Yurimaguas, I I Jul 1972, Croat 18069 (MO). San Martin: Lamas, N of San Antonio, 21 Oct-4 Nov 1937, Belshaw 3577 (CAS, F, GH, LL, MO, NY, US, WIS); Bolz6n de Utchya, 7 Oct 1977, Johns & Ramirez 202 (OS).

Brazil. Amazonas: Rio Branco, B6a Vista, 6 Sep 1943, Ducken 1323 (F, GH, US); Carafia, Vaup6s, Rio Negro, 30 Oct 1945, Lemos Fr6es 21289 (US); Ya- nomano Indian village, 22 Aug 1975, Prance & Ramos 23617 (MO, US). Para: Belem, near Instituto Agro- nomico do Norte, Archer 8249 (US); Tapana, near Para, Killip & Smith 30232 (NY, US).

Clibadium sylvestre is one of the three most broadly distributed species in the genus (C. eggersii and C. surinamense are the others). Morphological 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) to 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, John- ston 1069). These characters do not correlate well with geography, however, and hence vitiate varietal recognition.

Clibadium sylvestre has its closest relative in C. leiocarpum. It differs by the strigose, white-green (when dried) phyllaries with obtuse apices (vs. non-strigose, brown- green phyllaries with acute apices in C. leiocarpum), by a typically paniculiform capitulescence (vs. thyrsoid), villose cypselae (vs. glabrous), and also by a rudimentary pappus of two short awns. Clibadium sylvestre is adapted to lower elevations (200-600(-1600) m), whereas C. leiocarpum occurs higher (1400-2800 m). Cliba- dium sylvestre seems to be the only species producing acetylenic compounds, which are responsible for fish-poisoning (Arriagada, 1995).

Blake (1924) described Clibadium appressipilum to include only Panamanian el- ements; his taxon falls clearly within the variation observed in C. sylvestre. The type of C. appressipilum has phyllaries 9-11- veined and C. sylvestre usually has phyllaries 7-1 1-veined; the paleae subtending pis-

tillate florets are always 7-veined (vs. 5-7- 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 over- lapping distributions, the documented use of C. appressipilum as a fish poison (Allen 857, NY), also supports its inclusion within C. sylvestre.

Blake (1924, p. 601), when describing Clibadium appressipilum, indicated that the nearest ally of his species was C. caudatum (also suggested by Robinson, 1979b): "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 affinities were based.

Earlier workers (Schulz, 1912; Blake, 1917; Stuessy, 1975) mentioned a possible confusion in the application of the name Clibadium asperum with C. sylvestre, also described by Aublet in the same publica- tion, where both names were then called species of Baillieria. Schulz (1912) synonymized C. sylvestre with C. surinamense. My 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. There- fore, the sheet labelled as Baillieria aspera corresponds to the type of Baillieria sylvestris, and vice versa.

CLIBADIUM ACUMINATUM Benth., Bot. voy. Sulphur 5: 114. 1844. TYPE: COSTA RICA. Cocos Island, 3-7 Apr 1838, Bar- clay 2179 (LECTOTYPE, here designated: BM. ISOLECTOTYPES: K-2, photo: OS; US; MO). Clibadium parviceps S. F Blake, Contr. U.S. Natl.

Herb. 22: 598. 1924. TYPE: VENEZUELA. Ara- gua: Colonia Tovar, 1856-1857, Fendler 1967 (HOLOTYPE: GH; ISOTYPES: K, photo: OS; US, photos: F, US).

Clibadium sneidernii Cuatrec., Revista Acad. Co- lomb. Ci. Exact. 9: 240. 1954. TYPE: COLOM- BIA. Narifio: El Palmar (Cordillera Occidental,


vertiente occidental), 25 Oct 1944, von Sneidern 4522 (HOLOTYPE: F, photo: US).

Clibadium pediculatum Aristeg. in T Lasser, Fl. Venezuela 10: 393. 1964. TYPE: VENEZUELA. Merida: La Carbonera, 24 Sep 1956, Aristeguieta 2479 (HOLOTYPE: VEN; ISOTYPES: NY, US).

Shrubs, to 5 m tall, woody at base; stems with branches slender, striate, strigose to densely strigose, hairs 1 mm long. Leaves petiolate; blades ovate to ovate-lanceolate; membraneous; 6-20 X 3-1 0 cm; 3-5- veined at base, apices acuminate to largely acuminate, bases acute or attenuate, margins serrate, pauciserrate, serrulate or crenate-serrate, teeth usually mucronate; adaxially dark green, slightly strigose, hairs 0.5 mm long, abaxially pale green, scarcely pilose or shortly pilose mostly on the main veins, hairs 1 mm long; petioles slender, 1.5 to 5 cm long, strigose, hairs 1 mm long. Capitulescences thyrsoid, trichotomously branched. Capitula subglobose, 4-5 mm high, 3-4 mm diam., sessile or shortly pedicellate; phyllaries 3, suborbicular-orbicular, 3-4 X 2.5-3.5 mm, 7-veined, apices acute, abaxially greenish, strigose, hairs 0.5 mm long, margins shortly ciliate at distal one- third. Pistillate florets 4-6, subtended by ovate paleae, 3.5 X 2.5 mm, 7-veined, apices acute, strigose, hairs 0.5 mm long, margins scarcely ciliate at distal one-third; corollas white, 3 mm long, 1 mm diam., glabrous, 3-lobed, lobes irregularly deltoid, 0.4 mm long, scarcely pilose at apex; styles 1.5 mm long, branches 1 mm long. Staminate fiorets 5-10, subtended by lanceolate paleae (occasionally few central florets without paleae), 3 X 1-1.5 mm, glabrous, margins distal one-third shortly ciliate, 3-veined; corollas white, tubulate, 3 mm long, 1 mm diam., 5-lobed, lobed deltoid, 0.5 mm long, scarcely pilose at apex; anthers black, 1 mm long; ovaries 1.5 mm long, 0.8 mm diam., scarcely pilose at distal apex. Cypselae obovoid, rounded at base, 2 X 1.7 mm, pilose at distal one-third, hairs 1 mm long. Chro- mosome number unknown.

Distribution, ecology, and phenology. Costa Rica (Cocos Island), Colombia, and Venezuela; mostly on edges of secondary tropical wet forests; 300-1300 m; flowering from March to December.

Selected specimens examined: COSTA RICA. Pun-

tarenas: Isla del Coco, 20 Jun 1997, Rojas 3637 (MO).

COLOMBIA. Nariiio: El Palmar, 25 Oct 1944, von Sneidern 4522 (F).

VENEZUELA. Aragua: Altos del Choroni, Badillo 1945 (F); Peninsula Paraguana, Cerro Santa Ana, 15 Dec 1964, Breteler 4286 (MO). Distrito Federal: entre Aguas Negras-El Junquito, Badillo 2069 (F), 16 Jun 1984, Davidse & Miller 28067 (MO). Merida: Tovar, alrededores de Zea, Badillo 6457 (F). Miranda: Parque Nacional Guatopo, Nee 17658 (F); Cerros del Bachiller, S of Santa Cruz, Steyermark & Davidse 116730 (MO). Trujillo: alrededores de la Via Flor de Patria-Bocon6, Perez-Levis 79 (F).

Clibadium acuminatum is most closely related to C. micranthum. Both species have strigose ovate to ovate-lanceolate leaves; thyrsoid capitulescences; sub-orbicular, 7-veined, acute phyllaries; pistillate florets subtended by 7-veined paleae; and staminate florets subtended by 3-veined paleae. Clibadium acuminatum differs from C. micranthum in having strigose phyllaries (vs. pilose), 4-6 (vs. 10-14) pistillate florets, 5-10 (vs. 10-11) staminate florets, 5- lobed (vs. 3-lobed) corollas, and pilose (vs. glabrous) cypselae.

Clibadium acuminatum was considered by Blake (1924) as closely related to C. parviceps based on general morphology, except for minor differences in number and shape of phyllaries and paleae and by more acuminate leaf apices. He suggested this relationship based on the examination of a single specimen of C. acuminatum (Stew- ard 326) from the type locality. He noticed similarity in general morphology but the specimen had only 2 narrow (vs. 3) more ovate phyllaries observed in C. parviceps. After having studied the same material (Stewaird 326) and the type specimens of C. acuminatum and C. parviceps, I have observed capitula of C. acuminatum that have 2 or 3 ovate phyllaries (2 phyllaries are less frequently found, however). The character states of both species overlap to such extent that reducing C. parviceps to synonymy seems appropriate.

Four syntypes of Clibadium acuminatum collected by Barclay have been located. The two specimens at K and the one at BM are not in very good condition and therefore they do not show the diagnostic characteristics of the species very well. The one at MO (ex BM), here chosen as lectotype, is


the best-preserved specimen and has all of the diagnostic characteristics.

CLIBADIUM MICRANTHUM 0. E. Schulz, Bot. Jahrb. Syst. 46: 625. 1912. TYPE: PERU, 1780, Ruiz 3 (HOLOTYPE: B destroyed; photos: F, GH, MO).

Branched shrub, 2 m tall; stems with branches striate, pilose hairs 0.5-1 mm long. Leaves petiolate; blades ovate to ovate-lanceolate, membraneous, 8-15 X 4- 8 cm, apices largely acuminate, bases attenuate, margins shortly serrate to serrulate; adaxial surface glabrous to shortly strigose, hairs 0.5 mm long, abaxial surface glabrous; petioles 1-2.5 cm long, strigose, hairs 1 mm long. Capitulescences thyrsoid, trichotomously divided, branches slightly striate, strigose. Capitula subglobose, 3 mm high, 2 mm diam., sessile or shortly pedicellated; phyllaries 3, ovate to suborbicular, 3 X 2-2.5 mm, apices acute, abaxially short pilose, hairs 0.3 mm long, margin shortly ciliate on distal one-third, 7-veined. Pistil- late florets 10-14, subtended by ovate to oblong-ovate paleae, 3 X 2 mm, 7-veined, shortly pilose, hairs 0.4 mm long, margins shortly ciliate on distal one-third; corollas white, glabrous, 1.5 mm long, 0.5 mm diam., 3-lobed, lobes 0.3 mm long, glabrous; styles 1.5 mm long, branches 0.5 mm long. Staminate florets 10-1 1, subtended by lanceolate paleae, 2.5 X 1-1.5 mm, 3-veined; corollas tubulate, white, 1.5 mm long, 0.5 mm diam., 3-lobed, lobes irregularly deltoid, 0.4 mm long, glabrous; anthers black, 1 mm long; ovaries 1.5 mm long, 0.5 mm diam., glabrous. Cypselae obovoid, 1.5-2 X 1-1.4 mm, rounded at base, glabrous (exceptionally scarcely pilose at apex). Chromosome number unknown.

Distribution, ecology, and phenology. Peru and Bolivia; in secondary vegetation, disturbed areas; 700-1700 m; flowering from April to December.

Selected specimens examined: PERU. Amazonas: Valle del Rio Santiago, Quebrada Caterpiza, 14 Dec 1979, Tunqui 335 (F), 28 Dec 1979, 496 (F). Cuzco: Quispicanchis, Inambari, Vargas 15472 (US). Hua- nuco: Leoncio Prado, Hermilio Valdizan, cerca de La Divisora, Rimachi 5006 (CAS). Junin: Schunke Ha- cienda, above San Ram6n, 8-12 Jun 1929, Killip & Smith 24822 (F, GH, LP, NY, US); Chanchamayo, Fon-

do Romero, Pampatigre, above Santa Ana, 7 Mar 1985, Stein & Todzia 2340 (MO, OS). Loreto: Iquitos, Williams 8205 (LP); Puerto Legufa, 24 Jul 1929, Killip & Smith 26834 (LP, NY). Madre de Dios: Iberia, vic. Rio Tahuamanu, Seibert 2022 (LP).

BOLIVIA. La Paz: Yungas: Polo-Polo bei Coroico, Oct-Nov 1912, Buchtien 230 (F, NY); Maripi, Apr 1886, Rusby 2144 (GH, NY, US); near Yungas, Apr 1886, Rusby 2146 (GH, NY, US).

Clibadiumn micranthum is characterized by small capitula (3 mm high), arranged in loosely cymose-thyrsoid capitulescences. It is closely related to C. acuminatum. Cli- badium micranthum has pilose phyllaries (vs. strigose in C. acuminatum), 10-14 (vs. 4-6) pistillate florets, 10-i1 staminate florets with corollas 3-lobed (vs. 5-10 and 5- lobed), and glabrous (vs. pilose) cypselae.

Clibadium micranthum is superficially similar to C. microcephalum. They differ clearly in capitulum structure and organization. Clibadium nmicrocephalum has glabrous, obtuse phyllaries, 3 pistillate florets with 4-lobed corollas, 3 epaleaceous staminate florets with 5-lobed corollas, and pilose cypselae.

Doubtful and Excluded Names

Clibadium agreste Mart. ex Baker in Fl. Bras. (Mart.) 6(3): 157. 1873. = Ichthyo- there agreste Baker, Fl. Bras. (Mart.) 6(3): 157. 1884. nom. nud. pro syn.

Clibadium angustifolium Mart. ex Baker in Fl. Bras. (Mart.) 6(3): 154. 1884. nom. nud. pro syn. = Ichthyothere linearis (Benth.) Baker, Fl. Bras. (Mart.) 6(3): 154. 1884 (Latreillea linearis Benth., Ann. Nat. Hist. 2: 110. 1830).

Clibadium commelinoides (Less.) DC., Prodr. 5:506. 1836. TYPE: Brazil. May- puri, 1836, Schomburgk 247 (HOLOTYPE: K, photo: OS; ISOTYPE: BM, photo: OS; K, photo: OS). = Ichthyothere cunabi Mart. Buchn. Rep. Pharm. 35: 195. 1830.

Clibadium foetidum Alemand ex L., Mant. P1. 294. 1771. nom. nud.

Clibadium hispidulum (C. Wright ex Gri- seb.) M. G6mez & Molina, Anal. Soc. Esp. Hist. Nat. 19: 276. 1890. Lantanop- sis hispidula C. Wright ex Griseb., Mem.


Amer. Acad. Arts, 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). = Lantanop- sis hispidula C. Wright ex Griseb.

Clibadium neriifolium (Kunth) DC., Prodr. 5: 507. 1836. Baillieria neriifolia Kunth., Nov. gen. sp. pl. [folio ed.] 4: 227. 1818. TYPE: VENEZUELA Federal District: Si- Ila de Caracas, 21 Nov 1799-7 Feb 1800, Humboldt & Bonpland s.n. (HOLOTYPE: P). = Espeletia neriifolia (Kunth) Sch.Bip. In: H. A. Weddell, Chlor. And. 1: 67. 1856.

Clibadium pedunculosum (Rich.) DC., Prodr. 5: 507. 1836. TYPE: FRENCH GUIANA, Cayenne, 1792, Leblond 339 (LECTOTYPE: G, photos: F, MO). = Rien- courtia pedunculosa (Rich.) Pruski, Brit- tonia 50: 478-479. 1998.

Clibadium schomburgkii Sch. Bip. in R. Schomburgk, Reis. Br. Guiana 940. 1849. nom. nud.

Acknowledgments

First and foremost, I thank Tod E Stuessy, for his invaluable help and constant encouragement. Numerous friends and colleagues have provided help and assis- tance 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 out- standing Colombian synantherologist, who shared with me his botanical and historical expertise on the Colombian Compositae; thanks go also Jose L. Panero, for sharing with me important information concerning the Andean Heliantheae; Javier Francisco- Ortega, for his valuable comments on no- menclatural problems; Hugo Valdebenito and Fausto Sarmiento, for obtaining valuable information about types at QPSL; and to John Pruski, for sharing with me information regarding the nomenclature and eth- nobotanical uses of some species of Cli- badium. I thank the curators of the follow- ing herbaria from which I borrowed speci-

mens 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. Ap- preciation is expressed to Eric Lamont and two anonymous reviewers for their valuable suggestions to this paper and to Thomas Za- noni for his editorial precision and expertise to enhance the manuscript. The author wants to thanik Tod Stuessy for allowing the use of his illustrations and to the Missouri Botanical Garden Press for granting per- mission for its use. Financial support was provided by the National Science Founda- tion (INT-9017695), The Garden Club of America, The Tinker Foundation, and The Ohio State University Janice C. Beatley Herbarium Award.

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Numerical List of Taxa

Clibadium subg. Paleata Arriagada Clibadium sect. Eggersia Arriagada

1. C. eggersii 2. C. leptophyllum 3. C. divaricatum

Clibadium sect. Trixidium DC. 4. C. terebinthinaceum 5. C. erosum

Clibadium subg. Clibadium Clibadium sect. Glomerata Arriagada

6. C. anceps 7. C. glomeratum 8. C. trianae 9. C. sessile

10. C. microcephalum 11. C. frontinoense 12. C. zarucchii 13. C. rhytydophyllum 14. C. sprucei

Clibadium sect. Grandifolia Arriagada 15. C. grandifolium 16. C. cordatum 17. C. manabiense 18. C. glabrescens 19. C. pentaneuron

Clibadium sect. Clibadium 20. C. laxum 21. C. sodiroi 22. C. surinamense 23. C. peruvianum 24. C. arboreum

25. C. armanii Clibadium sect. Oswalda (Cass.) DC.

26. C. leiocarpum 27. C. sylvestre 28. C. acuminatum 29. C. micranthum

List of Exsiccatae

Acosta-Solis, M. 9723(10); 10771(27); 12285(4).

Aguilar H., M. 322(24). Alain (Brother Alain). 9422(5). Allart, A. 230(4). Allen, P. H. 118(26); 224(27); 711(26);

857(27); 1341(26); 1998(15); 3621(15); 4592(27); 4653(26); 4970(9); 5272(24); 5637(7).

Almeda, F 2987(6); 3145(4); 3304(4); 3515(26); 4023(1); 4033(3); 4171(7).

Alston, A. H. G. 5210(4); 5324(4); 5905(27); 7901(3).

Ancuash, E. 461(27). Anderson, W. R. 3301(4); 6739(25);

9603(25). Andre, E. s.n.(22). Andrews, L. M. 3-15(5). Antonio, T. 1136(15); 1146(6); 2224(27);

2881(27); 5041(6). Aranda, M. 32(4). Araque, J. 18C740(15). Araquistain, M. 655(26); 1157(26);

2440(3); 3050(22).


Arbainsyah, A. 1028(22). Archer, W. A. 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); 3415(18); 8249(27).

Aristeguieta, L. 2479(28); 3268(22); 4932(27).

Armani, A. s.n.(25). Arriagada, J. 131(22); 132(27); 133(27);

134(27); 135b(6); 135c(27); 136(26); 137(26): 140(26); 142(22); 143(22); 146(26); 147(26); 149(22); 150(22); 151(22); 152(7); 153(7); 154(7); 155(22); 156(22); 157(22); 159(22); 160(26); 161(26); 162(26); 163(22); 164(22); 165(27); 304(22); 308(26); 309(26); 310(22); 400(26); 402(26); 403(26); (404(22); 405(26); 406(15); 407(26); 408(6); 409(26); 410(26); 411(26); 412(1); 413(18); 415(26); 417(26).

Asplund, E. 5389(22); 7614(22); 10544(22).

Atwood, J. T 161(24). Aublet, J. s.n.(22); s.n(27). Austin, D. 4097(27). Ayala, F 501(23); 3104(23). Badillo, V. M. 1945(28); 2069(28);

6457(28). Baeza, S. 212/23(22). Bailey, L. H. 228(22); 252(22). Bakes, R. J. 33(22). Balgooy, M. M. 2903(22). Balick, M. 2622(24). Balslev, H. 10341(10); 10652(22). Bang, A. M. 361(23); 1203(23). Barbosa, C. 33(27); 6646(16). Barbour, P. 4299(23); 4386(23); 5386(23). Barclay, A. 2179(28); 3299(8). Barringer, K. A. 3211(22); 3365(26);

4003(6). Beard, P. 1157(5); 1360(5). Belem, R. P. 1258(25); 1289(25). Belshaw, C. M. 3316(23); 3577(27). Benavides, S. 4688(19). Benitez, C. 2517(28). Bensnam, R. 140(27). Berlin, B. 625(27); 984(27); 1633(23). Bernardi, L. 361(22). Berry, P 917(22). Biolley, P. 7399(1).

Blackwell, W. 2749(15). Blake, S. F 7354(1). Blanchet, J. A. 632(25). Blum, K. 487(22); 2662(6). Boeea, R. S. 5901(22); 9055(22). Boom, B. 6864(5). Borsboom, J. 12293(22). Box, H. E. 25(22). Brant, A. E. 1378(19). Breckon, D. 3082(5). Breedlove, D. E. 10145(24); 15423(24);

26410(24); 26504(24); 42714(24). Brenes, A. M. 3817(6); 5608(22);

5977(26); 6262(22); 6335(6); 17317(22); 19292(22); 20586(6).

Breteler, F 4286(28). Bridgeman 55(22). Bristan C. 1110(1). Bristol, M. L. 519(19). Brito, D. 12(25). Britton, N. L. 166(4); 179(5); 357(4);

554(4); 626(22); 663(22); 693(22); 1466(22); 1812(22); 2132(5); 2245(22); 2810(4); 3579(4); 3976(4); 5420(5); 5530(5); 6264(5); 6454(5); 7586(5); 10135(5).

Broadway, W. E. (s.n.)(24); 376(22); 634(22); 4234(22); 9826(22).

Brown, S. s.n.(27); 5530(5). Buchtien, 0. s.n(23); 134(23); 230(29);

5622(23). Burch, D. G. 4656(6). Burger, W. C. 4744(7); 5144(22); 5735(7);

6582(26); 8753(6); 9010(7); 9048(6); 9406(6).

Burley, J. S. 2000(22). Burnham, S. 114(24). Bussey, M. G. 534(26); 825(1). Cabrera, A. L. 29986(25). Callejas, R. 2285(4); 6610(15); 6842(12). Calzada, J. I. 500(24); 4716(24). Camp, W. H. E-884(21); E-1484(10); E-

3305(22); E-3601(22); E-3826(20). Capus, P. G. 59(27). Cairdenas, M. 1160(23) Cardona-Puig, F 264(27); 2848(27). Carleton, M. 458(1). Carlson, M. C. 3405(15). Cazzalet, P. C. 5169(27); 5188(1). Cedillo T, R. 288(24). Ceron, C. E. 2648(1). Chacon, A. 832(22). Chagas, J. 252(25).


Chambers, K. L. 2738(5); 2756(5); 2770(5).

Chaparro, M. L. 81(1). Charpin, A. 13274(8). Chavarria, M. 143(1). Churchill, H. 4429(1); 5485(6). Clark, H. L. 7674(27). Clark, J. 3545(13). Claussen, P. 666(25); 1314(25). Cochrane, T S. 6287(9); 6289(9). Coelho, L. 1739(22). Coleman, R. A. 265(25). Contreras, J. L. 2873(24); 7026(24);

9951(24). Cooper, G. P. 170(5). Cordoba, K. 21(27). Correa A., M. D. s.n.(26); 115(27);

234(27); 2781(26); 3349(15). Cowan, C. P. 3980(24); 3924(24);

4031(24). Cowan, R. S. 1418(4); 2427(24). Crawford, D. 1243(24). Croat, T. B. 6267A(22); 6242(22);

10005(1); 10478(26); 10715(22); 10929(22); 11364(27); 12283(27); 13372(15); 13518(20); 15520(27); 16014(26); 17465(23); 18301(23); 18069(27); 18142(23); 18301(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); 39712(24); 40283(24); 41579(24); 41776(24); 42897(22); 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, A. 9631(24). Crosby, G. T. 21(22). Crosby, M. R. 1045(4). Cruz, E. 286(24). Cuatrecasas, J. 61(8); 6112(22); 8182(8);

8422(19); 11521(8); 11596(19); 11994- A(22); 12086(22); 12798(8); 13696(16); 14004(22); 14378(16); 15025(15); 15298(15); 16683(15); 16941(2); 17534(22); 18653(22); 21514(1); 21660(19); 22034(19); 22478(22); 22163(22); 23808(22); 23932(15);

24084(27); 27121(22); 27504(22); 27601(15); 27639(8); 27933(22).

Cuevas, L. 3271(24). Daniel (Brother Daniel). 3986(22);

4205(19). D'Arcy, W. C. 6537(26); 6611(6);

6333(27); 6590(27); 9494(27); 10221(26); 10322(15); 10911(9); 12255(27); 12294(15); 15860(26); 15904(15); 15970(6); 16230(15).

Davidse, G. 4748(22); 5649(22); 10221(26); 24607(26); 28067(28).

Davidson, M. E. 141(26); 954(26); 6717(1).

Davis, E. W. 789(23); 924(27). De la Cruz, J. S. 1228(27); 1972(27);

3229(27); 3289(27); 3459(22); 3679(27).

De la Ossa, J A. s.n.(27). Delessert, J. 5625(25). Denslow, J. S. 79-104(1); 407(22);

2276(22); 2383(1); 2642(1). Devia, W. 624(16); 977(22). DeWolf, G. P. 332(22). Diaz-Piedrahita, S. 47(19); 181(4);

650(23). Diers, R. 564(23). Dillon, M. 0. 1428(29); 1454(23). Dodge, C. W. 5603(7); 6288(22). Dodson, C. H. 1849(14); 6385(20);

7235(1); 7589(20); 9817(22); 12703(22).

Donnell-Smith, J. 2347(24). Dressier, R. L. 50(24); 4798(1). Duarte, A. P. 4464(25); 10100(25). Ducken, A. 1323(27). Dudley, T D. 10091(29); 11411(29). Dugand, A. 2485(27); 3014(22); 3854(22). Duke, J. A. 3828(27); 4838(22); 6088(3);

9886(15); 10171(27); 11470(3); 11918(3); 11927(27); 12116(6); 12282(27); 12516(27); 13778(1); 15487(1); 15409(27); 16117(7).

Duque J., J. M. 4030(22). Dusen, P. 16424(25). Duss (Brother Duss). 316(5); 2491(5). Dwyer, J. D. 596(26); 1019(22); 1256(22);

5111(15); 5128(15); 7197(22); 8066(27); 8669(28); 8730(22); 8799(22); 8844(6); 9496(1); 10969(24); 11198(1); 11415(24); 11988(27); 14400(24).

Ebinger, J. 362(22).


Echeverry, R. 2322(4). Eggers, H. F 619(5); 3057(22); 5460(22);

13057(22); 15309(1). Egler, N. A. 24614(22). Eiten, G. 5880(25). Ekman, E. L. 168(4); H-3854(22);

8771(4). Erlanson, C. 0. 561(27). Ernst, W. R. 1088(5); 2035(5). Espina, J. 1144(27); 1930(27); 1991(15);

3668(27). Eyerdam, W. 24694(23); 24802(23). Faber-Langendoen, J. 1856(19). Fay, L. 789(24); 2182(1). Feisinger, J. 812093(6); 822182(26). Fendler, A. 484(22); 640(22); 1967(28). Fernaindez, A. 232(15). Fernaindez, 0. 1934(27). Fernandez-Casas, J. 8048(23). Ferreira, A. 3358(23); 4515(27); 6805(3). Fiebrig, K. 5815(25). Fleming, M. 22(22). Folsom, J. P. 2422(6); 3648(6);

4564A(27); 4753(6); 5378(6); 5813(15). Forero, E. 361(27); 725(1); 1767(27);

1984(27); 2257(16); 2279(19); 2347(16); 2824(22); 3312(15); 3517(16); 3871(15); 4163(2); 4622(27); 5139(27); 5418(1); 5476(15); 5527(2); 5540(15); 5871(16); 6853(15); 7594(16).

Fosberg, F R. 21619(22); 22969(20); 56191(27).

Foster, R. B. 1282(22); 9315(1). Fournet, J. P. 217(27). Fredholm, A. 3355(22). Froes, R. de L. 1738(27); 23310(27). Fuchs, H. P. 21728(15). Funk, V. 10654(22). Gadner, G. 3277(25). Garcia, E. E. 232(27). Garcia-Barriga, H. 7716(22); 12772(22);

15802(27); 18284(27); 18325(22). Garganta, M. 853(22). Gehriger, W. 361(22). Gentle, P. H. 1541(24); 8251(1). Gentry, A. H. 948(22); 1541(27);

1737(27); 3034(15); 5938(26); 9036A(19); 9594(27); 9811(27); 12409(1); 14657(22); 15002(22); 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).

Giannotti, E. 4567(25). Gibson, A. C. 2512(24). Gill, R. C. 44(27). Gilli, A. 113(17). Gillis, W. T. 10162(26). Gines, B. 4481(22). Glaziou, A. 9484(25); 9489(25). Gleason, H. A. 286(27); 619(27). Gomez, L. 23246(6). Gomez-Laurito, J. 2160(1); 5515(7);

6419(15); 8599(26); 8740(1); 9378(26). Gomez-Pompa, A. 1546(24); 3920(24);

4507(24); 4517(24); 4527(24); 4564(24).

Gonza'lez, J. 3931(24). Graham, S. A. 298(26). Grant, M. L. 9759(22). Grasshoff, J. 91(7); 100(6). Grayum, M. H. 2700(22); 9041(22). Grijalva, A. 1180(22); 2109(26);

2509(26); 2859(22). Grubb, P. 1. 715(22). Guainchez, F J. 2088(22). Guilding, L. s.n.(5). Guillemin, J. B. 411(25). Gutierrez, G. s.n.(27); 753(27); 944(22);

170122(1); 170657(22). Guzmain, M. 701(22); 2018(22). Haber, W. A. 737(6); 835(26); 1189(26);

5362(6). Hagen, W. von. 14(27); 441(5). Hahn, W. 195(27). Hamilton, C. W. 715(26); 1391(15);

2729(15); 3656(26); 3857(26). Hammel, B. E. 1691(15); 2599(6);

2621(15); 3891(15); 4072(22); 4353(22); 5453(27); 9240(15); 14039(22).

Hampshire, R. J. 492(1); 976(15). Hansen, B. 7586(24); 9422(5). Harley, R. M. 11081(25); 11249(25);

11295(25); 21874(25). Harling, G. 9410(17); 9670(22); 9680(20);

10124(10); 10153(21); 11525(20); 11876(22); 12363(13).

Harlow, J. R. 2(27).


Harmon, W. E. 2552(24); 2553(24); 2826(24); 4031(24).

Harris, J. A. C15413(4). Harris, W. H. 3183(4); 8674(22); 9463(4);

9886(22); 10299(22); 15157(4). Hart, J. A. 1567(27). Hartman, R. L. 3870(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).

Hartweg, K. T. 1139(22). Hassler, F 5625(25). Hatheway, W. H. 1391(26). Hatschbach, G. 25957(25); 26091(25);

34993(25); 43850(25); 46174(25). Haught, 0. L. 1914(22); 2047(22);

2124(22); 4678(15); 6488(8). Hayes, S. s.n.(22); 61(27). Hayworth, D. 342(26). Hendrix, C. 149(23). Heringer, E. P 2784(25); 7560(25). Herrera, G. 939(22). Hespenheide, H. A. 533(5); 1119(4). Higgins, J. 105(27). Hill, S. R. 89(1); 22115(5). Hinton, G. B. 14631(24); 14698(24). Hitchcock, A. S. s.n.(4); 16875(4);

17021(22); 17543(27); 17658(27); 20259(22); 20261(22); 20430(20); 21873(10).

Hodge, W. H. 735(5); 736(5); 901(27); 1037(5); 1201(5); 1396(5); 1462(5); 1870(5); 2241(5); 2380(5); 3195(27); 6675(22).

Holm, R. W. 7(22); 121(7). Holton, I. F 346(22). Hood, R. 977(22). Howard, R. A. 11824(5); 11911(5);

13622(4); 17533(5); 19387(5). Huft, M. J. 1621(15). Humboldt, F W. s.n.(27). Hunnewell, F W. 20003(22). Hutchinson, P. C. 3721(1). Idrobo, J. M. 69(22). Imray, J. 145(27). Irvine, G. C. 104(27); 363(22). Irwin, H. S. 393(22); 9642(25);

11209(25); 11381(25); 12674(25); 16352(25); 18743(25); 23772(25); 25432(25); 25681(25); 26443(25); 29507(25); 32216(25).

Jansen, R. 547(22).

Jansen-Jacobs, M. J. 1819(27). Jaramillo M., R. 4030(22); 6700(3);

7559(20); 7797(17). Jativa, C. D. 103(22); 804(20); 2102(20). Jeffrey, C. 100(16). Jimenez M., A. s.n.(l); 1031(1); 1269(6);

2117(1); 2224(6). Johns, D. 146(27); 202(27); 239(23). Johnston, I. M. 1069(27). Jones, S. 9144(29). Jorgenisen, P. 4811(25). Judd, W. 525(5). Judziewicz, E. J. 4581(27). Juncosa, A. 1733(3). Kayap, R. 1460(27). Kennedy, H. 1910(7). Kessler, P. 772(22); 822(22). Khan, R. 450(6); 587(26). Killip, E. P. 5144(15); 5268(15);

5995(22); 8342(22); 8976(22); 12103(22); 12120(22); 20783(22); 22450(23); 22810(23); 22860(23); 22916(23); 23034(23); 23550(29); 24049(23); 24180(23); 24822(29); 25068(23); 25494(28); 26305(23); 26474(23); 26834(29); 27061(23); 27263(23); 30332(27); 31232(27); 33061(22); 33575(15); 33867(22); 33982(8); 34462(22); 34815(22); 35123(24); 37165(28); 37592(22); 38862(6); 39842(19).

King, R. M. 875(20); 5257(22); 5754(19); 5764(19); 5780(22); 5791(22); 5814(22); 5824(22); 5852(22); 5875(22); 6305(5); 6396(5); 6560(17); 6567(20); 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, J. H. 94(26); 393(22); 1523(27).

Kirsch, J. R. 114(26). Klug, G. 2348(23); 3335(23). Knapp, S. D. 1276(22); 1532(26);

2401(27); 3101(1); 3207(27); 3273(27); 5633(6); 5825(22).

Knight, D. s.n.(5); 711(4). Koch, J. 5028(7). Koie, M. E. 4880(2). Koptur, S. 177(27).


Koster, H. 49(22). Krapovickas, A. 14274(25); 34351(25). Krukoff, B. A. 1738(27); 5212(27). Kuntze, C. E. 0. s.n.(23); s.n.(250);

452(5); 1327(22); 1967(8). Kvist, L. P. 230(22). Lagenheim, K. 3264(22). Langlasse, E. 7(15). Lanjouw, J. 12056(22). Lasser, T. 1446(22). Lawrance, A. E. 135(15). Lazor, R. 2983(22). Lehmann, F C. 855(19); 1256(19);

5200(22); 9056(22). Lellinger, D. B. 448(27). Lemos-Froes, R. 21289(27). Lent, R. W. 218(26); 435(3); 860(7);

1127(6); 2895(1). Leon (Hermano Leon). 660(3); 10880(4). Lewis, H. 596(26); 975(1); 1745(6);

2082(15); 3382(27); 3434(1). Liesner, R. L. 1494(24); 1713(7); 2098(1);

2815(7); 17877(27); 24447(22); 25925(27).

Limbach, H. 120(27). Little, E. L. 8114(22); 13064(5);

13554(5); 16320(5). Lleras, E. P17376(22). Logname, C. 7399(23); 7534c(23). Lorentzen, R. A. 3792(20). Lowrie, S. R. 265(22). Lugo, S. H. 152(22); 224(22); 228(22);

763(14); 1027(22); 1343(22); 1360(27); 1371(22); 1396(22); 1703(22); 2052(27); 2481(1); 2926(22); 3890(22); 3986(22); 4554(10); 4605(21); 4941(22).

Lundell, C. L. 20044(24). Luteyn, J. L. 11197(14); 1447(22);

1638(15); 4833(4). Maas, P. J. M. 2021(2). MacDaniels, L. H. 860(24); 27433(27). Macedo, A. 1771(25). Madison, M. T. 4940(20). Madrifian, S. 237(22). MacRae, J. 35(23); 118(3). Magalhaes, H. 3183(25). Maguire, B. 22707(22); 22956(22);

33603(22); 40227(22); 40336(22). Maias 144(22). Marcks, B. 826(24). Marles, J. 33(27). Mairquez, J. M. 152(24).

Martin, B. 61(27). Martinez, C. G. 128(24); 652(24);

1725(24). Masson, F s.n.(5). Mathews, A. 1360(27). Mathias, M. 5056(27); 5191(20). Matuda, E. 2422(24); 2754(24); 4059(24). Maxon, W. R. 197(4); 1323(4); 4767(27);

6505(22); 6960(22); 7033(22). Maxwell, R. 217(24). McDaniel, S. T. 2366(27); 11799(27);

14482(24); 17089(23). McDonagh, P 165(1). McDowell, T 715(22). McPherson, G. D. 7529(6); 11073(15);

13396(19). McVaugh, R. 21414(24). Meacham, C. A. 2(8). Medina, E. 249(19). Mejia, A. 86(22). Mexia, Y. 6208(22); 6254(27); 6255(1);

6506(27); 6595(18); 6668(18); 6678(27); 6696(20); 6915a(18); 6946(21); 6949(21); 7024(1); 7110A(18); 7256(27); 7272(1); 8467(20).

Miller, J. S. 902(22); 1221(22); 1422(26). Molina, R. A. 2413(22); 3148(22);

3601(24); 11863(22); 12260(24); 14147(22); 15070(22); 15860(24); 17508(6); 20029(26); 30573(22).

Monsalve, M. 624(19); 1601(19). Monteiro, 0. P 3100(25). Moore, L. 142(23). Moreno, P 9165(26); 10313(22);

10613(22); 13072(1); 14989(1); 15504(26); 15098(1); 17744(22); 23298(22).

Moreti, C. 896(27). Mori, S. A. 3860(15); 5590(1); 5897(15);

6164(1); 13643(25); 13719(25). Morley, B. D. 480(4); 493(4). Morton, C. V. 5946(27). Murata, G. 4451(22). Murillo, M. T 560(22). Nee, M. 3780(22); 6963(24); 7139(27);

7898(27); 8176(4); 8890(22); 9854(1); 9983(26); 17658(28); 18792(24); 24914(24); 26315(24).

Neill, D. A. 97(1); 1365(26); 2920(26); 3578(1); 4207(1); 7394(22); 8525(21).

Nelson, E. B. 2728(1); 3109(24);


3558(24); 4078(22); 4653(22); 4830(1); 5059(22).

Nevling, I. 1683(25). Nichols, E. 497(22). Nicolson, D. H. 2025(27); 4098(5). Nowicke, J. W. 3434(1). Nunes, E. 116(25). Oliver, R. L. 2385(27); 3667(1). Ollgaard, B. 35322(27). Olsen, J. 596(22). Opler, P. 1602(1). Orcutt, C. R. 2738(4); 3113(4); 3324(4). Otero, J. L. 586(5). Panero, J. 3039(20). Peck, M. E. 238(24). Pelaez, G. 510(19). Pennell, F W. 1721(22); 1771(22);

1975(8); 3239(22); 3898(22); 4219(22); 5194(22); 7217(22); 8322(22); 8704(22); 14035(29).

Pereira, E. 4950(25); 8443(25); 9064(25). Perez-Arbelaez, E. 6142(15); 6441(22). Perez-Levis 79(28). Persaud, R. 45(27); 341(22). Peters, C. 121(27). Philcox, D. 3292(25). Philipson, W. R. 2407(8). Piper, C. V. 5534(22). Pipoly, J. J. 1079(26); 1999(1); 3759(1);

6005(1); 6204(1); 9962(4). Pittier, H. F s.n.(1); 749(22); 1211(8);

2228(22); 3136(26); 3798(22); 4157(27); 4730(27); 7880(22); 7893(22); 8656(15); 8964(27); 9664(4); 11068(22); 11280(15); 11290(1); 13596(22); 15059(27); 15061(22); 16068(15).

Plowman, T. C. 2153(27); 6487(23); 6852(27); 11647(23); 12080(22).

Poeppig, E. F 35(23); 1611(22). Porter, D. M. 4651(15); 4785(1). Poveda, L. 1508(7); 4096(22); 4135(1);

4138(15). Prance, G. T. 10529(27); 11151(27);

15555(27); 23617(27). Prescott, G. W. s.n.(21); 404(21); 405(1). Pringle, C. G. 10802(24). Prior, R. C. s.n.(4). Proctor, G. R. 170(5); 6864(4); 8313(4);

10341(4); 17113(27); 19116(5); 19605(5); 21495(22); 32215(5).

Purpus, C. A. 3705(24); 10888(24). Purseglove, J. W. s.n.(22).

Puttemans, A. 209(25). Ramirez, D. 59(23). Ramos, J. 1599(15). Ratcliff, J. 10(24). Raven, P H. 19974(24); 20863(7);

21977(6); 22048(22). Redher, A. s.n.(4). Renteria, E. 1831(22). Revilla, J. 2446(23). Rimachi, Y. M. 3877(3); 5006(29);

5992(23); 7400(23). Rimbach, A. 700(14). Robles, E. L. 1722(22). Rodriguez, J. 169(22). Rojas, T. 3637(28); 10053(25). Romero-Castanieda, R. 2974(3); 6144(27);

6254(27); 6689(22); 11155(27). Rosario, A. J. 38(22). Rosas, M. 836(24). Rossbach, G. B. 3254(22). Rothery, H. C. s.n.(22). Rowlee, W. W. 59(22). Ruiz, P 3(29). Ruiz-Teran, L. 1139(18). Rusby, H. H. 2144(29); 2145(23);

2146(29). Rutter, J. s.n.(27). Saga'stegui, A. 5733(27). Salazar, F s.n.(24). Salvoza, F M. 839(22). Samuels, J. A. 345(22); s.n.(27). Sainchez, D. 213(11); 510(15). Sandino, J. 171(22). Santos, T. S. 1938(25). Sargent, C. B98(5). Sastre, C. H. 2626(5). Sauleda, R. P. 8784(5). Schipp, W. A. 972(1). Schmalzel, R. 1023(4); 1207(27);

1571(26); 1989(22). Schubert, B. J. 419(5). Schultes, R. E. 5120(22); 5879(27);

8965(27); 10400(27); 13955(22); 17153(27); 17861(15); 22245(27).

Schunke, V. J. 310(27); 1320(27); 1471(23); 1672(23); 2959(23); 6629(27); 7786(3); 9350(29); 11260(23).

Scolnik, R. 224(8). Seaman, F C. 88(1); 1591(26). Seibert, R. J. 245(26); 519(22); 559(27);

2022(29). Shafer, J. A. 485(5); 3430(5).


Sharp, A. J. 45808(24). Shemluck, M. 292(27). Shilom-Tom, A. 2496(24); 2829(24);

3311(24); 4116(24); 6660(24). Sieber, F W. 71(22); 617(5). Silva, A. 193(27); 936(25); 2168(25). Silverstone, P. A. 1254(22). Sinclair, J. 10052(22). Sintenis, P. 4183(5). Skutch, A. F 927(24); 1797(24); 2657(6);

2733(22); 3260(6); 3320(26); 3873(1); 3937(22); 4402(21); 4486(1); 5217(1); 5271(1).

Smith, A. C. s.n.(22); 153(26); 2182(27); 10245(5); 10530(5); 12855(23).

Smith, H. H. 327(22); 355(22); 536(27); 627(5); 1230(27); 2014(27).

Smith, J. F 494(22). Smith, L. B. 14842(25); 15415(25). Smith, S. F 538(23); 1601(25); 1698(23);

5830(23); 6614(15). Sneidern, K. 349(22); 1540(22); 4522(18);

5167(22); 5168(1). Sodiro, A. 21/3(1); 22/1(8); 22/2(21); 22/

3(21). Soejarto, D. D. 2715(1). Solange, de V. A Pessoa. 9(22). Solomon, J. C. 7515(23); 10025(23);

12945(23). Soukup, J. 2260(3). Spellman, D. L. 1960(24). Spruce, R. 2136(22); 4522(3); 5826(14). Standley, P. C. 32593(26); 33652(6);

34088(26); 41894(26); 44639(1); 48375(22); 48507(1); 49762(7); 62042(24); 64559(24); 85097(24); 86786(24).

Stehle, H. 86(5); 1679(5); 5806(5); 6408(27).

Stein, B. 2340(29). Steinbach, J. 2358(23); 2786(23);

9018(23). Stern, W. L. 80(26); 236(27); 300(27);

720(22); 1019(6); 1066(26); 1079(26); 1084(26); 7201(22).

Stevens, W. D. 753(18); 1104(24); 1390a(24); 4816(15); 4928(22); 9625(22); 10659(22); 10836(26); 12330(22); 13310(1); 13986(26); 13986(6); 14136(6); 15140(22); 15401(22); 17702(22); 18186(20); 19799(22); 21078a(1); 22445(22); 23951(1); 24140(1); 24606(1).

Stevenson, J. R. 1558(5). Steyermark, J. 37622(24); 41670(24);

45106(24); 46626(24); 47342(24); 55928(24); 56567(25); 56956(28); 56890(22); 58625(22); 62029(25); 62437(22); 91075(22); 116730(28); 118392(28); 118648(28); 118741(27); 120954(25); 127470(22); 131743(4).

Stimson, W. R. 1533(5); 1571(5); 5140(22); 5274(22).

Straw, R. M. 2441(23). Stuessy, T F 4451(22); 4460(22);

4461(22); 4462(26); 4463(22); 4465(1); 4479(26); 4489(22); 4493(27); 4494(26); 4504(22); 4507(27); 4509(27); 45 17(6); 4518(6); 4519(7); 4534(15); 4540(22); 4574(24); 4863(13); 4871(13); 4877(10); 4896(13); 4902(20); 4911(1); 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); 5510(22); 5524(22); 5528(8); 5571(22); 5578(8); 5595(19); 5604(22); 5606(18); 5623(8); 5627(8); 5641(22); 5680(8); 5709(19); 5725(15); 5727(22); 5737(27); 5742(15); 5748(22); 5770(8); 5775(8); 5782(27); 5783(22); 5814(10); 5816(10); 5817(21); 5822(10); 5823(10); 5856(22); 5867(20); 5870(20); 5873(22); 5897(20); 5901(20); 5902(19); 12318(20); 12352(20); 12311(13); 12337(8); 12347(8); 12353(8); 12355(8); 12362(10); 12363(10); 12365(10); 12369(10); 12370(21); 12400(14).

Sullivan, J. R. 181(1); 328(15); 375(6); 433(22); 4492(7).

Swartz, 0. P s.n.(4). Sydow, H. 240(13); 807(21). Systma, K. 1606(27); 1803(6). Tamayo, F 3520(27). Tate, G. H. H. 477(27); 577(14). Tate, R. 191(15). Taylor, J. 11836(26). Taylor, N. 318(4). Terborgh, J. W. 346(5). Tillet, S. S. 636-64(24). Todzia, C. A. 1721(l); 2549(6). Tomas (Brother Tomas). 175(19).


Tonduz, A. s.n.(6); 7330(7); 8892(22); 11479(1); 11915(26).

Toro, R. A. 154(22); 526(22); 1127(22). Torres, C. 5917(24). Triana, J. J. 1310(22); 1316(22); 1317(8). Tun-Ortiz, R. 1359(24); 1996(24);

2468(24). Tunqui, S. 335(29); 496(29). Turckheim, H. von 929(24); 4148(1);

7903(24). Turner, B. L. 789(23); 15024A(26);

15030(22); 15042(26). Tyson, E. L. 919(22); 1411(22); 3422(1);

4122(22); 5525(27); 6286(22); 6312(22); 6317(22); 4122(22).

Underwood, L. M. 3367(22); 3403(22). Urban, 0. 4183(5). Uribatan, S. J. 171(27). Uribe, U. L. 2696(8); 4098(19); 4340(22);

4917(19); 5715(22); 5873(22); 6310(19); 6957(22).

Usteri, A. 16711(25). Utley, A. 105(7); 2306(26); 5158(26);

6059(22). Vargas, C. C. 13(27); 292(22); 13983(23);

15472(29); 16928 (23); 18619(23); 18788(23).

Velarde-Nunez, 0. 2466(29). Venturi, S. 1180(23). Vickers, E. W. 210(27). Viegas, A. 3803(25). Vogl, C. 708(22). Wachenheim, G. s.n.(4). Wagner, M. 98(5); 116(5); 970(5). Warming, E. s.n.(25). Warner, R. H. 76(1). Wasshausen, D. C. 416(5). Weberbauer, A. 7864(23). Webster, G. L. 5102(22); 12198(6);

13352(5); 16782(27); 21911(7); 25014(25); 25014(25).

Weddel, H. 2161(1). Werling, L. 84(20). Weston, A. 4550(7); 4479(26); 3420(26);

5624(26); 3001(20); 4714(26); 4407(22).

Wetmore, R. H. 66(22); 180(22). Whetzel, H. 216(28). White, P. 181(26). White, J. W. 14(1); 740(22). Whitefoord, C. 2421(24); 5143(5). Whitlock, B. 181(26). Widgren, J. F 166(22).

Wilbur, R. L. 10575(6); 10661(22); 10678(6); 10884(26); 12084(22); 13423(1); 13669(26); 13701(26); 15271(26); 22349(6); 22460(6); 22785(7); 24258(26); 70279(22).

Williams, R. S. 284(23); 328(23); 468(23); 641(23); 698(27); 1929(23); 1998(23); 3344(23); 4304(23); 4967(26); 5119(25); 5122(4); 5198(25); 5655(23); 6684(23); 7363(27); 8205(29); 10478(28); 12241(25); 14474(24); 14621(l);(25); 22029(22); 20263(15); 26903(24); 28141(26); 28560(26); 28923(26); 42259(25).

Wilson, P. 150(1). Woodson, R. E. 212(26); 658(9); 731(22);

836(22); 908(26); 1359(22); 1408(22); 1743(22).

Woronow, G. 4788(22). Woytkowski, E 376(23); 5018(27);

8310(1); 35185(27). Wullschagel, H. 291(22); 1324(27). Wurdack, J. J. 2224(23); 35740(22). Yuncker, T. G. 8699(1); 17624(22);

18827(4). Zak, V. 1194(17); 1318(17); 2881(19);

3454(20). Zardini, E. 4049(25); 4185(25). Zarucchi, J. L. 1304(27); 2429(22);

3988(27); 5036(15); 5286(22); 5443(22); 5566(8); 5674(12); 7097(19).

Zaruma, J. 66(22).

Index to Names for Species of Clibadium

Accepted names are in boldface and synonyms are in italics. Each synonym is followed by the number of the accepted species name used in this index.

Bailleria aspera Aubl. 25 Bailleria barbasco H.B.K. 26 Bailleria sylvestris AubI. 26

1. Clibadium acuminatum Benth. Clibadium alatum H. Rob. 13 Clibadium alexandri Griseb. 27

2. Clibadium anceps Greenm. Clibadium appressipilum S. F Blake 26

3. Clibadium arboreum Donn.Sm. 4. Clibadium armanii (Balb.) Sch.Bip.

Clibadium asperum (Aubl.) DC. 25


Clibadium badieri Griseb. 26 Clibadium barbasco (H.B.K.) DC. 26 Clibadium caracasanum DC. 25 Clibadium caudatum S. F Blake 26 Clibadium chocoense Cuatrec. 7 Clibadium congestum Cuatrec. 28

5. Clibadium cordatum Cuatrec. 6. Clibadium divaricatum S. F Blake

Clibadium donnell-smithii J.M. Coult. 3

7. Clibadium eggersii Hieron. 8. Clibadium erosum (Sw.) DC.

Clibadium fragiferum Griseb. 8 9. Clibadium frontinoense Df'az &

Arriagada Clibadium funkiae H. Rob. 19

10. Clibadium glabrecens S. F Blake Clibadium grande S. F Blake 11

11. Clibadium grandifolium S. F Blake 12. Clibadium glomeratum Greenm.

Clibadium harlingii H. Rob. 21 Clibadium havanense DC. 26 Clibadium heterotrichum S. F Blake 20 Clibadium latifolium Rusby 26 Clibadium lanceolatum Rusby 25

13. Clibadium laxum S. F Blake Clibadium lehmannianum O.E. Schulz 25

14. Clibadium leiocarpum Steetz Clibadium leiocarpum var. strigosum S. F Blake 14

15. Clibadium leptophyllum Cuatrec. 16. Clibadium manabiense H. Rob.

Clibadium mexiae S. F Blake 23 17. Clibadium micranthum 0. E.

Schulz 18. Clibadium microcephalum S. F

Blake Clibadium napoense H. Rob. 10 Clibadium oligandrum S. F Blake 3 Clibadium pacificum Cuatrec. 11 Clibadium pallidum Diels 23 Clibadium parviceps S. F Blake 1 Clibadium pediculatum Aristeg. 1

19. Clibadium pentaneuron S. F Blake 20. Clibadium peruvianum Poepp. Ex

DC.

Clibadium pileorubrum Cuatr. 19 Clibadium pilonicum Stuessy 2 Clibadium pittieri Greenm. 7 Clibadium pittieri f. phrixium Greenm. 7 Clibadium polygynum S. F Blake 7 Clibadium psilogynum S. F Blake 20 Clibadium pueblanum S. F Blake 3 Clibadium remotifiorum O.E. Schulz 20 Clibadium rimachii H. Rob. 6

21. Clibadium rhytidophyllum Diels Clibadium rotundifolium DC. 4 Clibadium sarmentosum Cuatrec. 19 Clibadium scandens Cuatrec. 19 Clibadium schultzii S. F Blake 14

22. Clibadium sessile S. F Blake Clibadium sneidernii Cuatrec. 1

23. Clibadium sodiroi Hieron. 24. Clibadium sprucei S. F Blake

Clibadium strigillosum S. F Blake 26 Clibadium subauriculatum Stuessy 22 Clibadium subsessilifolium Hieron. 28

25. Clibadium surinamense L. Clibadium surinamense var. macrophyllum Steyerm. 25 Clibadium sychnocephalum S. E Blake 28

26. Clibadium sylvestre (Aubl.) Baill. 27. Clibadium terebinthinaceum (Sw.)

DC. Clibadium terebinthinaceum subsp. colombiense Cuatrec. 11

28. Clibadium trianae (Hieron.) S. F Blake Clibadium trinitatis DC. 25 Clibadium vargasii DC. 26 Clibadium vargasianum H. Rob. 20 Clibadium villosum Benth. 25 Clibadium zakii H. Rob. 19

29. Clibadium zarucchii H. Robin. Orsinia eupatoria DC. 4 Oswalda baillerioides Cass. 25 Trixis aspera Sw. 25 Trixis erosa Sw. 8 Trixis terebinthinacea Sw. 27

revision of the genus clibadium (asteraceae, heliantheae)

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