A Monograph of Neotropical Potalia Aublet (Gentianaceae: Potalieae)

Lena Struwe; Victor A. Albert

According to the ongoing treatment of the Rubiaceae for the Flora of the Guianas, this family is represented in the region (Guyana, Suriname, French Guiana) by about 80 genera and 470 species. In the Guayana Shield, the Rubiaceae is the third most diversified family, with about 750 species (after the Leguminosae, with 1032 species, and the Orchidaceae, with 1020 species). In addition, it is found in practically all habits (herbs, shrubs, treelets, trees, lianas) and habitats (forests, savannas, tepuis, etc.), and is an important floristic component, both in terms of diversity and frequency. A preliminary assessment is made, in order to compare the local floristic composition with those of surrounding regions (northern South America, Amazon Basin, Central America, Antilles).

According to recent molecular phylogenies, the tribe Sipaneeae is recognized as a separate tribe, and positioned within the subfamily Ixoroideae. The Sipaneeae, as here delimited, include the genera Chalepophyllum, Dendrosipanea, Limnosipanea, Maguireothamnus, Neblinathamnus, Neobertiera, Pteridocalyx, Sipanea, Sipaneopsis and Steyermarkia (with a total of about 50 species). Several genera of this group are small shrubs, and the woody habit is shown to be basal in the Sipaneeae. Sipanea and Limnosipanea were shown to be monophyletic and not sister taxa, indicating that the herbaceous habit evolved at least two times in the tribe. The tribe has its main center of diversity in the Guayana Shield, with 9 genera and 47 species, with a few species in Central America (3 genera and 3 species), and a few species on the Brazilian Shield (2 genera and 5 species). Following these results, three projects are currently undergoing: 1) a complete monographic treatment, with key, generic and specif...

Symbolanthus (Gentianaceae, Helieae) is a genus of small trees, shrubs, and herbs distributed in South America (the central and northern Andes and the Guayana Highlands) and southern Central America. To discover the pattern of Symbolanthus evolution and to assess the relationships between Symbolanthus and the closely related Wurdackanthus of the Guayana Highlands (Venezuela, Brazil) and the Lesser Antilles (St. Vincent), we performed cladistic analyses on members of both taxa. Molecular data (5S-NTS rDNA sequences) were gathered from individual plant samples, mainly from herbarium specimens. 5S-NTS sequence data did not resolve internal relationships well in Symbolanthus and show that the genus may be a recently and rapidly diverging clade. However, these results indicate that Wurdack- anthus is either paraphyletic to Symbolanthus or that W. argyreus is nested within a Symbolanthus clade. Furthermore, W. frigidus may be the basalmost species of the Wurdackanthus-Symbolanthus clade. ...

Systematic Botany (2004), 29(3): pp. 670–701 q Copyright 2004 by the American Society of Plant Taxonomists A Monograph of Neotropical Potalia Aublet (Gentianaceae: Potalieae) LENA STRUWE1,3 and VICTOR A. ALBERT2 Dept. of Ecology, Evolution, and Natural Resources, Rutgers University—Cook College, 237 Foran Hall, 59 Dudley Road, New Brunswick, New Jersey 08901; 2 The Natural History Museums and Botanical Garden, University of Oslo, P.O. Box 1172 Blindern, NO-0318 Oslo, Norway 3 Author for correspondence (struwe@aesop.rutgers.edu) 1 Communicating Editor: James F. Smith ABSTRACT. The first revision of the neotropical genus Potalia (Gentianaceae: Potalieae, formerly Loganiaceae) is presented. This genus was formerly considered monotypic, with Potalia amara being the only recognized species. New morphological and anatomical studies show that Potalia resinifera should be resurrected and that nine newly discovered species should be recognized; one endemic to Central America (P. turbinata), two endemic to Chocó, Colombia (P. chocoensis and P. crassa), two endemic to white-sand areas along Rio Negro and its tributaries (P. elegans and P. maguireorum; Brazil, Colombia, and Venezuela), and the last two endemic to white-sand areas in the western Amazon basin (P. coronata and P. yanamonoensis; Peru, Brazil and Colombia). Potalia amara sensu stricto is restricted to the Guianas and northeast Brazil, whereas P. resinifera is widespread in lowland Amazonia and the Andean foothills. Morphological differences between the species are primarily in fruit and inflorescence morphology, but there are also differences in plant size, leaf shape, and corolla color. Potalia resinifera shows the greatest infraspecific variation. Phylogenetic relationships between Potalia and its close relatives Anthocleista from Africa and Fagraea from tropical Asia-Australia-Pacifics are discussed. The widespread pharmacological uses of Potalia species by indigenous peoples of the Neotropics are reviewed and vernacular names are listed. RESUMEN. Se presenta la primera revisión del género Neotropical Potalia (Gentianaceae: Potalieae, anteriormente Loganiaceae). Este género se considero monotı́pico en el pasado, siendo Potalia amara la única especie reconocida. Sin embargo, nuevos estudios morfológicos y anatómicos muestran que Potalia resinifera debe resucitar y que se deben reconocer nueve especies, una endémica de Centro América (Potalia turbinata), dos endémicas del Chocó, Colombia (P. chocoensis y P. crassa), tres endémicas de áreas con arenas blancas a lo largo del Rı́o Negro y sus tributarios (P. elegans y P. maguireorum; Brasil, Colombia y Venezuela), y las últimas dos endémicas de áreas con arenas blancas al oeste de la cuenca del Rı́o Amazonas (P. coronata y P. yanamonoensis; Perú, Brasil y Colombia). Potalia amara sensu stricto se restringe a las Guianas y al noreste de Brasil, mientras que P. resinifera se encuentra ampliamente distribuida en las áreas bajas de la Amazonia y en el pie de monte andino. Las diferencias morfológicas entre las especies se encuentran básicamente en el fruto y en la morfologı́a de las inflorescencias, pero adicionalmente se encuentran diferencias en el tamaño de las plantas, la forma de las hojas y el color de la corola. Potalia resinifera es la especie que muestra mayor variación infraespecı́fica. Se discuten las relaciones filogenéticas entre Potalia y sus parientes mas cercanos, Anthocleista de Africa, y Fagraea del tropico AsiáticoAustraliano-Pacı́fico. Se revizan los ampliamente conocidos usos farmacológicos que las comunidades indigenas del Neotropico dan a las especies de Potalia y se presenta una lista de nombres comunes. RESUMO. A primeira revisão do gênero neotropical Potalia (Gentianaceae: Potalieae, previamente considerada como Loganiaceae) é apresentada. Esse gênero foi considerado a princı́pio monotı́pico, sendo Potalia amara a única espécie reconhecida. Entretanto novos estudos morfológicos e anatômicos mostram que Potalia resinifera deve ser revalidada e que nove espécies recém-descobertas devem ser reconhecidas; uma endêmica à América Central (P. turbinata), duas endêmicas a Chocó, Colômbia (P. chocoensis e P. crassa), duas endêmicas aos terrenos com solo de areia branca (areias quartzosas) ao longo do Rio Negro e seus afluentes (P. elegans e P. maguireorum; Brasil, Colômbia e Venezuela), e duas endêmicas aos terrenos com solo de areia branca da região oeste da Bacia Amazônica (P. coronata e P. yanamonoensis; Peru, Brasil e Colômbia). Potalia amara sensu stricto restringe-se às Guianas e ao nordeste do Brasil, enquanto P. resinifera apresenta-se amplamente distribuı́da nas planı́cies da Amazônia e nas encostas dos Andes. As principais diferenças entre as espécies encontram-se na morfologia dos frutos e das inflorescências, havendo também diferenças no tamanho da planta, forma da lâmina foliar e cor da corola. Potalia resinifera apresenta grande variação infraespecı́fica. São discutidas as relações filogenéticas entre Potalia e dois gêneros próximos, Anthocleista da África e Fagraea da Ásia-Austrália-Pacı́fico. O difundido uso farmacológico das espécies de Potalia por povos indı́genas dos neotrópicos é revisado e os nomes populares são listados. Potalia Aubl. has long been known as a broad-ranging, yet monospecific South and Central American genus of the angiosperm family Loganiaceae (Leeuwenberg and Leenhouts 1980). Morphological, molecular, and phytochemical investigations (Bureau 1856; Fosberg and Sachet 1980; Jensen 1992) of Gentianales taxa have shown that Potalia, together with its relatives Anthocleista R. Br. and Fagraea Thunb. (of tribe Potalieae sensu Leeuwenberg and Leenhouts, 1980), is better considered a member of Gentianaceae (Struwe and Albert 1997; Struwe et al. 1994). Following this work, Po- talia is now classified in subtribe Potaliinae of tribe Potalieae in Gentianaceae (Struwe et al. 2002). Molecular and morphological data strongly support African/Malagasy Anthocleista as the sister group of Potalia (Frasier et al., 2002; Struwe et al., 2002). Anthocleista and Potalia share the unusual characteristic among Gentianales of having an 8 to 16(–24)-merous corolla and androecium with a 4-merous calyx. Anthocleista and Potalia are closely related morphologically but geographically separated by the Atlantic. The closest relative of these two genera, Fagraea (from Asia, Austra- 670 2004] STRUWE & ALBERT: POTALIA (GENTIANACEAE) 671 FIG. 1. Distribution map of Potalia amara (G), Potalia chocoensis (n), Potalia coronata (s), Potalia crassa (●), Potalia elegans (D), Potalia maguireorum (l) Potalia turbinata (C), and P. yanamonoensis (▫). lia, and the Pacific), has strictly pentamerous flowers, and all three genera bear fleshy berries, an unusual feature in Gentianaceae. Species of Potalia are widely distributed as understory rain forest shrubs and small trees throughout low and middle elevations in northern South America, FIG. 2. occurring southwards to ca. 158 S (Figs. 1, 2). Additionally, Potalia spp. are also known from the Colombian Chocó (Forero and Gentry 1989) and Panama to Costa Rica in Central America (Blackwell 1968; Woodson and Schery 1942). The terminal inflorescences of Potalia have conspicuously yellow to orange branches Distribution map of Potalia resinifera (●). 672 SYSTEMATIC BOTANY bearing flowers with four decussate sepals, ca. 1 cm long, yellow, green, or white corollas, and 8–10 corolla lobes and stamens. Earlier concepts of Potalia as a monotypic genus have obscured considerable morphological heterogeneity. Our revisionary studies have shown that several species can be discerned, and that Potalia amara, the type species, is restricted to the Guianas and northeastern Brazil (Fig. 1). In this revision we accept nine species of Potalia. Two of these were previously known; the type species, Potalia amara, described by Aublet (1775; Fig. 3.), and P. resinifera, described by von Martius (1826–27) from Rio Negro, Brazil, and now in large areas in Amazon Basin and neighboring areas. Four newly found species are endemic to nutrientpoor, white-sand areas. Of these, Potalia elegans and P. maguireorum are largely endemic to the western Guayana Shield along the Rio Negro and its tributaries (Struwe and Albert 1998b), and P. coronata and P. yanamonoensis are endemic to eastern Peru (especially around Iquitos, Loreto) and southeastern Colombia as well as westernmost Amazonas, Brazil. Another new species, Potalia turbinata, is restricted to rainforests of Panama and Costa Rica. Endemic to the lowland rainforests of Chocó, Colombia, are Potalia chocoensis and P. crassa, each known from only one and two collections, respectively. All seven of the newly recognized species and Potalia amara are discrete, well-defined entities, whereas Potalia resinifera is polymorphic. Potalia is also the pharmacologically most important gentian genus in South America, and the uses and vernacular names of the species are summarized here. MATERIALS AND METHODS Pressed and liquid preserved material was used for detailed morphological and anatomical studies. Approximately 800 sheets were studied from the following herbaria: COL, ECON, F, GB, GH, IAN, INPA, K, MG, MO, NY, QCA, QCNE, S, UPS, US, VEN (abbreviations following Holmgren et al. 1990), and determinations are listed in Appendix 1. Field studies of living plants of Potalia amara were also performed by VAA in French Guiana in 1995 (see Notes under Potalia amara). Measurements given in keys and descriptions are based on dried material. If the key is used for field identification, it should be kept in mind that fruits and corollas of gentians often shrink up to 10% while drying. Vernacular names were collected from herbarium sheets and literature. TAXONOMIC HISTORY Potalia and its type species, P. amara, were first described by Aublet (1775) based on material from French Guiana (Fig. 3). Until recently, the only other species described was Potalia resinifera, published by von Martius (1826–27) based on material collected along Rio Negro in Brazil. Potalia resinifera was accepted by Don (1837–38) but since then has been treated as a synonym of P. amara. Indeed, the distinction of Potalia elegans with its arborescent habit, coriaceous [Volume 29 leaves, long and gracile inflorescences, non-ringed and globose fruits from P. amara sensu lato was first suggested only in 1980, by Fosberg and Sachet, and confirmed by Struwe and Albert (1998b). Potalia amara differs from P. elegans in having shorter stature, more chartaceous leaves, shorter inflorescences, and ringed, obconical fruits. Fosberg and Sachet (1980) cited specimens of Potalia elegans (Schultes & López 9339 and Schultes et al. 18127; cf. Struwe and Albert 1998b) in their argument supporting a gentianaceous position for Loganiaceae tribe Potalieae (cf. Struwe et al. 1994; Struwe and Albert 1997). Further consideration of morphological variation among the extensive herbarium collections of ‘‘Potalia amara’’ have led to the additional new species presented here. Potalia was initially included in Gentianaceae by Jussieu (1791), but was made the type genus of the new family Potaliaceae by von Martius (1826–27). Potaliaceae also included Anthocleista and Fagraea. Only three years later, Bartling (1830) transferred all three genera to Loganiaceae, a family that had been described by von Martius (1826–27) at the same time as Potaliaceae. Subsequently, Potalia has most often been included in Loganiaceae (e.g., Endlicher [1841], Bentham and Hooker [1873–76], and Solereder [1892–95]). Exceptions were the treatments that included Potalia again in Potaliaceae by Lindley (1836), Don (1837–38), as well as more recently, Hutchinson (1973). Baillon (1888) was alone in including Potalia in Solanaceae. Potalia was also included in the major work on Loganiaceae edited by Leeuwenberg (1980), where Loganiaceae reached its widest circumscription ever. In 1856, Bureau already had transferred Potalia back to Gentianaceae in his pharmacological thesis—a primarily anatomical and chemosystematic work that apparently did not reach many contemporary botanists. Except for one note on an herbarium sheet of Potalia amara from K (Voiteau s.n.), we have seen very few references to Bureau’s conclusions. Over 100 years later, Fosberg and Sachet (1974, 1980) again suggested that Potalia and associated genera in the Potalieae should be transferred to Gentianaceae, a conclusion accepted by Takhtajan (1987). It was later shown with cladistic analyses of morphological and molecular data that Loganiaceae were highly para- and polyphyletic and that Anthocleista, Fagraea, and Potalia should be placed in Gentianaceae (Bremer et al. 1994; Struwe et al. 1994). This was followed in the new family classification presented by Struwe and Albert (in Struwe et al. 1994), and also by Takhtajan (1997). Furthermore, sequencing of the trnL intron and matK gene have shown that Potalieae are deeply embedded inside a tropical clade of gentians (Thiv et al. 1999; Struwe et al. 2002). In the current classification of Gentianaceae, Anthocleista, Fagraea, and Potalia form subtribe Potaliinae in the recircumscribed tribe Potal- 2004] STRUWE & ALBERT: POTALIA (GENTIANACEAE) 673 FIG. 3. Potalia amara Aubl. A. flowering branch (Mori et al. 24074, NY). B, C. (Mori et al. 24073, NY). B. young branch with interpetiolar sheaths. C. young flower bud on inflorescence with floral bracts. D. flower from above (from photo in the field, Mori et al. 23865, NY). E, F, G. (Skog & Feuillet 7067, US). E. opened flower. F. anther at anthesis. G. gynoecium. H–J. (Mori et al. 24073, NY). H. fresh fruit. I. longitudinal and cross section of fruit. J. seed. K. dried fruit (Mori et al. 7236, NY). 674 SYSTEMATIC BOTANY ieae (Struwe et al. 2002; classification also available at www.rci.rutgers.edu/;struwe/gentnet). Also included in tribe Potalieae are the monotypic subtribe Lisianthiinae, based on Lisianthius P. Browne, and the pantropical subtribe Faroinae (including, e.g., Enicostema Blume, Faroa Welw., and Neurotheca Salisb. ex Benth.). TAXONOMIC TREATMENT POTALIA Aubl., Hist. Pl. Guian. 1: 394, t. 151. 1775. Nicandra Schreb., Gen. Pl. 1: 283. 1789, non Adans. 1763, nom. illeg. (homotypic synonym, and later homonym of Nicandra Adans.). TYPE: Potalia amara Aubl. Fig. 3. Shrubs or trees, monopodial or sparsely branched 2–12 m tall, glabrous, without (teste Klinge s.n.) or with small amounts of yellowish (teste Oldeman 247), clear (teste Knapp & Mallett 7092) or milky white (teste Soejarto et al. 3977) latex. Root a thickened primary taproot with thinner secondary roots (Archer 7651). Stems terete with leaves conspicuously crowded at branch apices, with light-colored wood and soft and separating pith, often becoming hollow in the middle; nodes with low, prominent and persistent interpetiolar ocreas, bases of young leaves often with large amounts of resin secreted by the colleters. Leaves decussate, simple, petiolated, entire, 180–1050 3 28–210 mm, narrowly elliptic to obovate and oblanceolate, widest at middle or above, often very large, dark green to yellowish or olive green, lighter below, coriaceous to chartaceous, base long-attenuate with the lamina decurrent into the petiole, the apex obtuse to acute (rarely truncate and sometimes with a short acumen); petioles up to 82 mm long; margins flat or slightly recurved; venation pinnate with many straight or slightly arcuate secondary veins, the midrib prominently raised below. Inflorescences terminal, sometimes overtopped by a lateral vegetative shoot, a dichasial cyme, 32–310 mm long, up to 50-flowered, up to 6 3 branched; branches bright yellow to orange (seldom pale green in Potalia elegans), not thickened at the nodes; pedicels stout, sometimes slightly bent, 7–30 mm long; bracts and floral bracts small, persistent, scale-like, about two times as long as wide. Flowers actinomorphic, erect. Calyx 4-merous, divided to the base, 3–11 3 3–10 mm, campanulate, usually yellow or rarely green, white, or orange, coriaceous and woody, tightly appressed to the corolla, persistent in fruit; lobes decussate, convex, the outer pair slightly larger than the inner pair, usually hyalinemargined, often rupturing by the developing fruit, erect to spreading or sometimes recurved in fruit, the apex obtuse; calyx bud apex obtuse. Corolla 8–10-mer- [Volume 29 ous, 7–20 mm long, tubular with a narrow lower part below the insertion of the stamens and a wider upper part, white, cream, yellow, or green (tube often paler than lobes), thick and coriaceous, deciduous in fruit; lobes lanceolate, sometimes with yellow margins, right-contorted in bud, erect or slightly spreading at anthesis; corolla bud apex tapering (acute) to rounded (obtuse). Stamens 8–10, isomerous with the corolla, inserted in the middle of the corolla tube, included in the corolla mouth; filaments very short, of equal length, completely fused into a very short (ca. 1 mm long) staminal tube; anthers free, linear, 3–9 mm long, sagittate, often with a small sterile apical appendage, longitudinally dehiscent, introrse; pollen in monads (tetrads in Potalia maguireorum), the exine smooth, porate; pores protruding. Gynoecium bicarpellate, included in the corolla, 5–9 mm long, the placentation axile with a thin septum; ovary slightly contracted at the base and with a ca. 1 mm high, basal glandular disk, upper part sterile; ovules numerous; style 2–5 mm long, filiform, deciduous in fruit; stigma capitate, globose, ca. 1 mm wide. Fruit a fleshy berry, globose, turbinate, elliptic, ovoid, to obovoid, 6–19 3 6–18 mm, green when fresh, brown to yellow when dry, often with a thickened horizontal ring around the middle of the fruit (Potalia amara, P. chocoensis, P. coronata, P. resinifera, P. turbinata, and P. yanamonoensis), the apex acute, truncate to obtuse, often consisting of sterile tissue (best seen in cross-sections). Seeds elliptic, flattened, not winged, red, brown, or black; hilum distinct, in the middle of one side; testa reticulate, pitted. Distribution. This genus is distributed in forests of Central America (Costa Rica, Panama) and tropical South America, more specifically along the Andes and its foothills (only east of the Andes except for Chocó, Colombia), Amazonia, and on the lower elevations of the Guayana Shield. Potalia grows from sea level in Central America to 1100 meters elevation in the Colombian Andes. For distribution maps, see Figs. 1, 2. Number of Species. Nine species. Etymology. Aublet (1775) did not provide an explanation for the derivation of the name Potalia, but it has been proposed that the name derives from the Latin word potare, to drink (Kuntze 1891; Struwe and Albert 1998a). Aublet (1775) noted that infusions and decoctions from this plant are used against diseases. Key to the species of Potalia. This key is based on measurements and investigations of herbarium material. Note that fresh leaves are often fleshy, but either chartaceous or coriaceous when dry, and corollas and fruits can shrink up to 10–15% when drying. All measurements in the key are based on dried material. 1. Leaves elliptic, broadest in the middle; corolla dark green (S. Venezuela, NW. Brazil) . . . . . . . . . . . . . . . . . 6. P. maguireorum 1. Leaves oblanceolate to obovate-spathulate, broadest towards the apex; corolla white, cream, yellow to orange and/or green (never completely green), the lobes often with yellow margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2004] STRUWE & ALBERT: POTALIA (GENTIANACEAE) 675 2. Inflorescence over 10 cm long; leaves with inconspicuous and relatively straight secondary veins that often appear wrinkled when dry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Fruit crown-shaped (truncate-coronate) with a massive horizontal ring above the sepals (Colombia, Peru, W. Brazil) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. P. coronata 3. Fruit globose to ellipsoid, without a horizontal ring (Colombia, Venezuela) . . . . . . . . . . . . . . . . . . . . . . 5. P. elegans 2. Inflorescence less than 10 cm long (rarely longer); leaves with conspicuous secondary veins that usually arc towards the leaf apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Fruit without a thickened ring (Colombia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. P. crassa 4. Fruit with a conspicuous (or rarely inconspicuous) thickened ring . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5. Corolla concolorous white or greenish white (rarely with green or yellow markings); fruit turbinate to globose, the sterile apical part less distinct and often confluent with a thickened but indistinct ring surrounding the fruit . . 6 6. Sepals recurved or loosely appressed when fruits mature; mature fruit wider than long (Panama, Costa Rica) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. P. turbinata 6. Sepals erect and appressed when fruits mature; mature fruit about as long as wide or longer (French Guiana, NE. Brazil) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. P. amara 5. Corolla green (rarely yellow), lobe margins and lower tube sometimes yellow (annotations referring to yellow and orange corollas [or flowers] may mistakenly refer to the calyx, which, along with the inflorescence axis, is usually colored a brilliant yellow-orange); fruit ovoid to obpyriform, often acorn-shaped (sometimes globose and/or rhomboid), the sterile apical part distinctly protruding with a thickened ring above the sepals . . . . . . . . . . . . . . 7 7. Fruit over 18 mm long with a sterile apical part longer than Ï of length of fruit (Fig. 4a–b; Colombia); leaves thick and coriaceous when dry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. P. chocoensis 7. Fruit less than 18 mm long, with a sterile apical part at most Ï of total length of fruit; leaves thin and chartaceous to slightly coriaceous when dry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 8. Horizontal ring of fruit undulating; calyx lobes in fruit 8–10 mm long (Fig. 9l–m; Peru) . . 9. P. yanamonoensis 8. Horizontal ring of fruit straight or inconspicuous; calyx lobes in fruit 4–8(-10) mm long (Figs. 9e–f, h, 10g–h, and 4d–e, g; widespread in Bolivia, Brazil, Colombia, Ecuador, Peru, Venezuela) . . . . . . . . . 7. P. resinifera 1. POTALIA AMARA Aubl., Hist Pl. Guian. 1: 394; 3: t. 151. 1775.—Nicandra amara (Aubl.) Gmel. in Forsyth, Bot. Nom.: 248. 1794.—TYPE: FRENCH GUIANA. Aublet s.n. (holotype BM!, photo NY!, US!). Fig. 3. Trees and shrubs, 0.5–2(–5) m tall. Stems up to 50 mm in diameter, below inflorescences 6–8 mm in diameter, the bark slightly fissured, brown to greyish brown or blackish; internodes short or long, 15–240 mm long, 2–ca. 6 times as long as wide. Leaves (265–)430–620 3 88–193 mm, obovate to oblanceolate, widest above middle, chartaceous to slightly coriaceous, usually thin when dry, dull when dry, green to olive green above, lighter and yellowish below, base attenuate, apex acute to acuminate; petioles up to 48 mm long, 4–5 mm in diameter; margins not thickened, not or slightly revolute; venation with secondary veins straight to arcuate (arcuate especially towards the tip of the leaf), ca. 90 degree angle to the midrib, the midrib prominent below, when dry the lamina usually not sinuous along the secondary veins. Inflorescences compact, not long-stalked, up to 40-flowered, 50–110 mm long, up to 110 mm wide; branches straight, the first order of inflorescence branches 4–10(–28) mm long, the second order 18–38 mm long and third order 9–21 mm long; pedicels short, 8–18(–30) mm long; bracts and floral bracts 2–3 mm long. Calyx lobes 3–6 3 3–6 mm, yellow, orange, or white, rarely with green tips, becoming loosely appressed in fruit, rarely slightly spreading, and not distinctly enlarged (then 5–6 3 4–6 mm). Corollas 9–10-merous, 10–13 mm long, the lower part 3–4 mm wide, upper part 4–6 mm wide, yellow, white, or cream (over-all green corollas never reported and probably not present, apex of corolla bud reported once as green [teste Cowan 38651]); lobes narrowly elliptic, 3–5 3 1–2 mm, 1/4–1/3 as long as the entire corolla, erect to slightly curved inwards at anthesis forming a small entrance to the corolla, yellow, cream, or white, the margins concolorous with the rest of the corolla (slightly lighter in herbarium specimens). Anthers 3–4 mm long, greenish-white to green (teste Mori et al. 15785). Fruits globose to elliptic, 11–15 3 10–15 mm, 2–2.5 times as long as calyx, green when immature and fresh (also recorded as reddish-green, teste Irwin et al. 48460), dark brown when dry, with a horizontal slightly thickened ring, apex obtuse (sometimes acute when dry, not crowned by a small persistent remnant of the style base. Seeds flattened, dark brown. Illustrations and Descriptions. Original description and illustration in ‘‘Histoire des plantes de la Guiane Françoise’’ (Aublet 1775: vol. 1: p. 394, vol. 3: plate 151) and the facsimile ‘‘The ethnobotany of Aublet’s histoire des plantes de la Guiane Françoise (1775)’’ (Plotkin et al. 1991: fig. 53). Also treated in ‘‘General System’’ (Don 1837–38: vol 4: 67) and in the monograph of Loganiaceae by Leeuwenberg (1980; illustrated in the chapter by Hallé [1980: fig. 3.10.2] and Leeuwenberg and Leenhouts [1980: fig. 2.29 pro parte]). Distribution. Potalia amara is distributed in French Guiana and in the easternmost parts of Amapá in northeastern Brazil (Fig. 1). The species may occur in Suriname as well, but we have seen no collections from that area, and the single collection we have seen from Guyana can only be determined with some uncertainty. This species grows in lowland and middle elevation non-flooded rainforests at 70–700 m elevation, some- 676 SYSTEMATIC BOTANY [Volume 29 FIG. 4. Potalia chocoensis Struwe & V. Albert and P. resinifera Mart. A–C. Potalia chocoensis Struwe & V. Albert (Forero et al. 8886, COL). A. fruiting branch. B. fruit at maturity, also in longitudinal section. C. seeds. D. Potalia resinifera, fruit (Philipson 2330, S). E, F. Potalia resinifera (Smith 1286, MO). E. inflorescence and infructescence on monopodially branching stem. F. flower at anthesis. G. Potalia resinifera. Fruit, also in longitudinal section (Vásquez & Jaramillo 5659, MO). 2004] STRUWE & ALBERT: POTALIA (GENTIANACEAE) TABLE 1. Phenology of Potalia species. Black boxes mark the months when flowers or fruits have been collected for each species. times also in secondary forests, along roads and trails, on river banks, and often on lateritic soil. Potalia amara seems to occur scattered in the understory, being a rather common but still occasional plant in the forest. Phenology. Potalia amara has been found flowering during June to December, and fruiting during all months except January (Table 1). Vernacular Names. Anilapouie (French Guiana), Bokokini (French Guiana), Mavévé-grand-bois (French Guiana), and pau de cobra (Brazil; Table 2). Etymology. Potalia amara was named by Aublet after its bitter taste (Aublet 1775; Plotkin et al. 1991), which is almost certainly caused by the seco-iridoid compounds characteristic of gentians’ bitterness (Jensen and Shripsema 2002). Notes. Potalia amara is a patchily distributed but locally abundant understory shrub in the Saül region of central French Guiana (V. A. Albert, pers. obs.; cf. Mori et al. 1997). Plants no more than ca. 1.5 m tall were found mainly in disturbed habitats such as along trails or roadsides and in light gaps in the rainforest. These were mostly monopodial, but branching or artificial cutting in previous years would lead to limited branching in some individuals. Fresh leaves have the color and fleshiness of spinach, although they are much stronger. The fleshiness trait is not retained after drying; herbarium specimens typically have chartaceous leaves. Flowering stage was synchronous over all individuals observed. Over two weeks of casual observation, corollas in flower buds never opened more than a couple of millimeters (Fig. 3d). The bright yelloworange inflorescences and calyces contrasted strongly with the dark green leaves, providing a striking floral display easily seen from a distance. Despite the visibility of flowering specimens, no pollinators were observed. Indeed, fruit set appeared to be extremely low, as no more than two fruits were observed among ap- 677 proximately 20 mature individuals examined. The fruits were found on plants in flower but on earlier growth (the same fruiting cycle can be seen in some Potalia resinifera specimens). Representative Specimens Examined. BRAZIL. AMAPÁ: Serra do Navio, Rio Amapari, slopes of Chumbo Ore Body, Cowan 38349 (NY); Forest on hill behind Oiapoque Airfield, Cowan 38651 (NY); immediately east of Colônia Agrı́cola do Oiapoque, about 4 kilometers north of mouth of Cricoú River, Irwin et al. 47515 (NY); Rio Oiapoque, virgin forest along road between Oiapoque and Clevelândia, Maguire et al. 47101 (NY); Camaipi, EMBRAPA reserve and vicinity, Mori et al. 15785 (NY); Mun. de Oiapoque, BR 156, 109 km SSE of Oiapoque on road between Oiapoque and Calçoene, Mori et al. 17205 (NY); Rio Araguari, Pires et al. 50587 (IAN, NY); Rio Araguari, along road between Porto Platon and Macapá, Pires et al. 51094 (NY); coastal region, Colônia do Torrão, Pires & Cavalcante 52675 (IAN, MG, NY). FRENCH GUIANA. St. Elie, SE of ORSTOM Biological Station at Forest Concession, Acevedo-Rdgz & Grimes 4929 (K, NY, US); Layon Saül vers Cambrousse, Allorge 380 (NY); Fl. Sinnamary, rive droite, 500 m E saut Aristide, de Granville et al. 186 (NY); Montagne Bellevue de l’Inini, de Granville et al. 7965 (MO); Montagne Cacao, Hallé 1145 (US); Rio Oiapoque, 1 kilometer south of Piedra Alice, Irwin et al. 47626 (NY); Rio Oiapoque, about 1 km. south of mouth of Riv. Yaroupi, Irwin et al. 48460 (NY); Rio Oiapoque, lower and middle slopes of Mt. Alikene, near Riv. Camopi, about 15 km. from its confluence with Rio Opapoque, Irwin 48605 (NY); Lieux de Récoltes, Rivière Tampoc, Lemoine 7904 (NY); Saül, in vicintiy of village and airport, Marshall & Rombold 120 (NY); Saül, near geographic center of the department of Guyane Française, Tracé Roche Bateau, Mori & de Granville 8829 (NY); Saül, Monts La Fumée, Mori & Boom 14816 (NY); Saül and vicinity, Route de Bélizon, 0–4 km N of Eaux Claires and third log bridge, Mori & Gracie 23865 (NY); Sentier Botanique, at Crique Tortue, Mori et al. 24091 (NY); Haute Approuague, crique Sable, crique Calebasse, Oldeman 247 (NY); tête saut Ouasseye, Oldeman 2609 (NY); Vallée encaissée de la crique Parépou, sur des collines à environ 18 km de son confluent, Oldeman 2853 (NY); Flats rive G. Yaroupi à saut Tainoua, Oldeman 3087 (NY); Cogneau, à 2 km de La Mirande, Ile de Cayenne, Oldeman 3171 (NY); Montagne de Kaw region, summit plateau, near Camp Caiman, Skog et al. 7067 (US); Saül Region, Saut Mais, 17 km E of Saül, Skog et al. 7234 (US). GUYANA. Basin of Kuyuwini River (Essequibo tributary), about 150 miles from mouth, Smith 2526 (NY). 2. Potalia chocoensis Struwe & V. Albert, sp. nov.— TYPE: COLOMBIA. Chocó: Hoya del Rio Atrato, 678 [Volume 29 SYSTEMATIC BOTANY TABLE 2. Alphabetical list of vernacular names. All vernacular names found in literature and on herbarium sheets during the scope of this study are listed here. The language is listed here when indicated on the label. The symbol ‘‘*’’ denotes approximate species determination due to insufficient or ambiguous herbarium material. The symbol ‘‘**’’ indicates records listed by Duke and Vasquez (1994) for Potalia amara in their book on Amazonian ethnobotany, however, Potalia amara in our concept does not occur in this area, only P. coronata, P. elegans, P. maguireorum, and P. resinifera. Vernacular name Species páo-de-cobre sacha curarina sacha curarina P. resinifera P. aff. resinifera* P. sp.** P. amara P. sp.** P. amara P. resinifera P. aff. resinifera* P. resinifera P. aff. resinifera* P. sp.** P. resinifera P. coronata P. coronata P. resinifera P. resinifera P. coronata P. resinifera P. sp.** P. resinifera P. aff. resinifera* P. aff. resinifera* P. aff. coronata* P. coronata P. coronata P. elegans P. sp.* P. aff. resinifera* P. resinifera P. resinifera P. resinifera P. resinifera P. elegans P. elegans P. resinifera P. maguireorum P. amara P. aff. resinifera* P. aff. resinifera* P. elegans P. elegans P. elegans P. resinifera P. resinifera P. amara, P. maguireorum, P. resinifera P. resinifera P. coronata P. resinifera sacha mangua shin-wee’ temblador tremblador turarina sacha yampak yanapa P. P. P. P. P. P. P. ahuaramacha amargo Andrés anabi anilapouie awuibiden bokokini boyomemo caspisenango chá-de-bugre chañetsopan chin-wee’ chirosenango chocho cu’yacono’le cuararina curare curarina curarina curarina curarina sacha curarina sacha hembre curarina sacha macho fé-ril goyongorz gúiongorı̈ hoja de gavilan itanóconi jaisoldao kararái kúranai ma-mee-mhoy’-da mapiguari martiguaja martiguaje matiguaja matugua mavévé-grand-bois ócaji cappuva okajı́ kahpuú palo de lagarto mato palo de rana palo de temblador palu huacu pao-de-cobra pau-de-cobra sp.** resinifera elegans sp.** resinifera resinifera resinifera Country: language Bolivia: Chácobo Colombia Brazil French Guiana: Oyampi Brazil: Makú French Guiana: Taki-Taki Ecuador: Quichua Peru Brazil Peru: Amuesha ? Colombia: Puinave Peru Colombia Peru Ecuador: Quichua Ecuador: Quichua Peru Brazil, Ecuador: Quichua, Peru Colombia Peru: Quichua Peru: Quichua (& Spanish?) Peru: Quichua (& Spanish?) Colombia: Kofán Colombia Colombia: Witoto Venezuela Colombia: Andoque Colombia: Emberá Peru Peru Colombia: Kuripako Colombia: Tuc Colombia Colombia Colombia Brazil French Guiana: Créole Peru Peru: Bora Venezuela Venezuela Venezuela Ecuador: Quichua Brazil Brazil Brazil Peru: Quichua Colombia: Quichua, Ecuador: Quichua, Peru: Quichua ‘‘Amazonia’’: Quichua Colombia: Puinave Venezuela Brazil Peru: Quichua Ecuador: Shuar Ecuador: Quichua 2004] STRUWE & ALBERT: POTALIA (GENTIANACEAE) Beté, 50–60 m alt, alrededores de la población, arbusto de 2 m, frutos verdes, 4 Apr 1982, E. Forero, Jaramillo, & A. Forero 8886 (holotype COL!; isotype MO!). Fig. 4a–c. Potalia resiniferae Mart. similis, sed fructu grandior (. 18 mm longo) differt. Treelet, 2–3 m tall. Stems at least up to 17 mm in diameter, below inflorescences ca. 10 mm in diameter, bark type unknown, internode length unknown. Leaves ca. 500 3 ca. 120–140 mm, broadly obovate, widest above middle, coriaceous, dull when dry, dull dark green above, dark yellowish below when dry, base long-attenuate, apex acuminate; petioles up to ca. 30 mm long, ca. 6 mm in diameter; margins slightly revolute, slightly thickened; venation with secondary veins slightly arcuate, diverging from the midvein at ca. 45–80 degree angle, the midrib prominent below. Inflorescences compacted, up to at least 7-flowered, ca. 60 mm long, up to ca. 70 mm wide; branches straight, not curved, inflorescence branches at least up to 15 mm long; pedicels short, 8–10 mm long; bracts and floral bracts 4–5 mm long. Flowers unknown. Calyx lobes in fruit 10–12 3 9–12 mm, other sepals without membranaceous margin, inner pair with 1 mm membranaceous margin, tightly appressed. Fruits ovate, 18– 21 mm long, 10–12 mm wide, ca. 3 times as long as calyx, green when fresh, brown when dry, with distinct horizontal thickened ring below the middle of the fruit, apex obtuse, not crowned by a small persistent remnant of the style base. Distribution. Potalia chocoensis is only known from the lowlands around Beté in Chocó (Colombia; Fig. 1). Phenology. Potalia chocoensis has been found flowering in October according to herbarium labels (flowering specimen not seen by us, this duplicate is presumably present in a Colombian herbarium). Fruiting specimens were collected in the months of April, August, and October (Table 1). Etymology. This species is named chocoensis after its distribution in Chocó, Colombia. Notes. This species has fruits with the most strongly developed sterile apices, with the fertile part of the fruit being contained in the lower 1/3 of the fruit. For a discussion on the presence of species with large fruits and coriaceous leaves in Chocó, see Notes under Potalia crassa. Representative Specimens Examined. COLOMBIA. CHOCÓ: Mun. de Bojayá, corregimiento La Loma, arboretum de CONIF en la margen derecha del Rı́o Bojayá, Espina & Mosquera 2230 (COL); Mun. de Quibdó, corregimiento de Guayabal, Rı́o Hugón, Garcia et al. 35 (COL). 3. Potalia coronata Struwe & V. Albert, sp. nov.—TYPE: BRAZIL. Amazonas: Municı́po de Atalaia do Norte, Rio Javari, localidade Estirão do Equador a 3 679 km da estrada do campo de pouso, mata de terra firme, solo argiloso, 48 339 S 718 409 W, arbusto até 2 m de altura, frutos com cálice amarelado, imaturos verdes, 2–18 Jan 1989, C. A. Cid Ferreira et al. 9908 (holotype NY!; isotype INPA). Fig. 5. Potalia eleganti Struwe & V. Albert similis, sed fructibus annulo horizontali supra sepala coronatis diversa. Trees, treelets, and shrubs, 2–20 m tall. Stems up to 250 mm in diameter, below inflorescences 10–14 mm in diameter, the bark fissured and uneven, brown; internodes rather short, 12–35 mm long, 1–3 times as long as wide. Leaves 510–1050 3 121–208 mm, oblanceolate, widest above middle, coriaceous, dull and conspicuously yellowish when dry, green above, lighter below, base long-attenuate, apex acute to acuminate; petioles up to 82 mm long, 7–11 mm in diameter; margins thickened, revolute; venation with secondary veins straight except slightly arcuate at the very apex, the midrib prominent below, when dry the lamina becomes sinuous along the secondary veins. Inflorescences elongated, long-stalked, up to ca. 50-flowered, 75–310 mm long, up to 265 mm wide; lower inflorescence branches straight, upper branches often slightly bent and/or curved upwards, the first order of inflorescence branches 22–95 mm long, the second order (22–)45–105 mm long and third order 10–48 mm long; pedicels short, 7–13 mm long; bracts and floral bracts 2–3 mm long. Calyx lobes 4–5 3 4–5 mm, yellow to white, tightly appressed in fruit and enlarged (then 6– 10 3 7–9 mm). Corollas 9–10-merous, 11–17 mm long, the lower part 3–4 mm wide, upper part 6–7 mm wide, green and yellow; lobes lanceolate-elliptic, 4–6 3 2–3 mm, 1/3–1/2 as long as the entire corolla, erect, green with lighter, yellow-colored margins. Anthers 6–9 mm long, greenish-white (teste Pipoly et al. 12142). Fruits obovate, 10–12 3 9–16 mm, 1.5–2 times as long as calyx, when fresh yellow (teste Cid Ferreira et al. 9908), greenish-yellow, green, or reddish (teste Idrobo 8554), yellow or light brown when dry, reported to be secondarily dehiscent by Idrobo 8554 but no dehiscent fruits ever seen on sheets, with a distinct horizontal thickened ring giving the fruit a coronate shape, apex strongly truncate with small acute apex, crowned by a small persistent remnant of the style base. Seeds flattened, pitted, not winged, brown, or black and red on same plant (teste Idrobo 8554). Distribution. Potalia coronata is endemic to the lowlands of western Amazonia (Fig. 1). It grows from Rio Javari on the Brazilian-Peruvian border, throughout Loreto, Peru, and to southern Colombia (Amazonas, Caquetá, Putumayo, and Vaupés). It grows at 90 to 250 meters elevation mainly in primary or secondary rain forests, sometimes inundated, and commonly on sandy soils (often white-sands) or, more rarely, on terra firme. 680 SYSTEMATIC BOTANY [Volume 29 FIG. 5. Potalia coronata Struwe & V. Albert. A. flowering branch (Cid Ferreira et al. 9908, NY). B. inflorescence (Rimachi Y. 6983, US). C- F. (van der Werff et al. 10254, MO). C. young flower. D. opened corolla. E. calyx with gynoecium (corolla removed) at anthesis. F. anther. G, H, I. (Vasquez & Soto 11921, MO). G. mature fruit. H. longitudinal and cross section of fruit. I. seed with papillate testa (close-up at the upper right). 2004] STRUWE & ALBERT: POTALIA (GENTIANACEAE) Phenology. Potalia coronata has been found flowering during February to March and May to November. Fruiting specimens have been collected in January to March and May to June (Table 1). Vernacular Names. Chocho (Colombia), curarina (Peru), cu’yacono’le (Peru), goyongorz (Colombia), gúiongorı̈ (Colombia), fé-ril (Colombia), sacha curarina (Peru; Table 2). Etymology. Potalia coronata is named after its coronate fruits, reminiscent of a king’s crown. The crown appears to be formed during fruit growth by pressure from the sepals, which remain tightly appressed throughout fruit development. Notes. Similar crown-shaped fruits are also known from Potalia yanamonoensis (distributed around the Iquitos region, Loreto, Peru), but P. coronata plants are readily distinguished by their thicker and larger leaves, longer inflorescences, and non-undulate fruit rings. The ring-bearing fruits of Potalia resinifera are also similar but this species differs in having smaller fruits with a relatively shorter sterile apical part. Representative Specimens Examined. BRAZIL. AMAZONAS: Rio Javari, behind Palmeiras Army Post, Lleras et al. P16992 (INPA, NY). COLOMBIA. AMAZONAS: Trocha Umañajedó, del Rı́o Apaporis al Rı́o Miritiparaná y Rı́o Miritiparaná, Garcia-Barriga 14470 (COL); Rı́o Igará-Paraná, La Chorrera, Idrobo 8049 (COL). CAQUETÁ: Rı́o Peneya, Pto. Tokio, Idrobo 8554 (COL). PUTUMAYO: Rı́o Putumayo, Puerto Ospina y alrededores, Schultes 3430 (GH). VAUPÉS: Comisario de Vaupés, Rı́o Piraparan, environs of catholic mission of San Miguel, trail from airstrip to the Cañon Colorado, Davis 133 (GH, K); Riberas del Rı́o Inı́rida, alrededores del sitio llamado ‘‘Morichal’’, cerca de la boca del Rı́o Papunáua, Fernández 2225 (COL). PERU. LORETO: Maynas Prov., Tamishiyacu, Ayala et al. 2867 (K, MO, NY); Iquitos, Rı́o Nanay, Mishana, Ayala et al. 3869 (MO, NY); near Base Araguana, upper Rı́o Mazán, c. due N. of Santa Maria de Nanay, Gentry & Revilla 16563 (MO); Rı́o Samiria, Gentry et al. 38039 (MO); Allpahuayo, Km 20 Iquitos-Nauta, ca. 35 km southwest of Iquitos, Gentry et al. 56013 (MO); Mishuyacu, near Iquitos, Klug 102 (NY, US); Iquitos and vicinity, Rı́o Nanay, trail to Mamon from Lopuna, Martin et al. 1721 (K); Iquitos, Estación Experimental IIAP, Alpahuayo, 21 km al S de Iquitos, Pipoly et al. 12142 (MO); Iquitos, Rı́o Nanay, Caserio Mishana, trocha hasta la Quebrada de Yarana mas abajo del Caserio de Chamchama, Rimachi 1223 (NY); Iquitos, tracha del caserio del Varillal, Rimachi 6983 (US); Iquitos, Carretera del caserio de Puerto almendras, Rimachi 7530 (US); Indiana, Rı́o Amazonas, Quebrada de Yanayacu, Bombonaje, Rimachi 8994 (US); Caserı́o Mishana, Rı́o Nanay, Campamento No. 1, Ruı́z & Murphy 200, (K, MO, NY); Carretera IQ-Nauta, Km. 32, Vásquez & Jar- 681 amillo 7883 (MO); Quebrada Paparo, afluente derecho del Rı́o Manati, Vásquez & Jaramillo 11640 (NY); Requena Prov., Jenaro Herrera, Rı́o Ucayali, Gentry et al. 56333 (MO); Ucayali Prov., Sapuena, Jenaro Herrera (CDJH); Vásquez & Soto 11921 (MO); Rı́o Javari, 2 hours above Rı́o Javarizinho, Prance et al. 24099 (INPA, MO, NY); Rı́o Napo, Negro Ureo, Martin & Lau-Cam 1312 (ECON); Rı́o Gueppi, tributary of Rı́o Putumayo, northernmost tip of Peru, on border with Ecuador, Gentry et al. 21923 (MO). 4. Potalia crassa Struwe & V. Albert, sp. nov.—TYPE: COLOMBIA. Chocó: Carretera (en construccı́on) Yuto-Lloró, 1 km de Yuto, bosque, alt. 70 m, arbolito de 3.5 m, frutos verdes, 9 Sep 1976, E. Forero & R. Jaramillo 2698 (holotype COL!; isotypes MO!, NY!). Fig. 6. Potalia eleganti Struwe & V. Albert similis sed inflorescentia breviori fructibusque majoribus, a P. amara Aubl. floribus et fructibus majoribus diversa. Small trees, ca. 3.5 m tall. Stems below inflorescences 8–10 mm in diameter, mature stems and bark unknown. Leaves 445–780 mm long, 130–180 mm wide, oblanceolate, widest above middle, very coriaceous, dull and yellowish when dry, green above, lighter below, base long-attenuate, apex acuminate; petioles up to 86 mm long, ca. 6 mm in diameter; margins slightly thickened, slightly revolute; venation with secondary veins straight to slightly arcuate, becoming arcuate at the very apex, the midrib prominent below, when dry the lamina is not sinuous along the secondary veins. Infructescences compact, short-stalked, at least up to 10-flowered, at least up to 55 mm long, at least up to 105 mm wide; infructescence branches straight, 5–8 mm long; pedicels short, 14–18 mm long; bracts and floral bracts 3–4 mm long (description based on incomplete material, only partial infructescences known). Flowers unknown. Calyx lobes in fruit 9–10 mm long, 9–10 mm wide, presumably yellow, tightly appressed. Fruits globose, 18–19 3 18–22 mm, 2–2.5 times as long as calyx, green when fresh (teste Forero & Jaramillo 2698), dark brown when dry, without a distinct horizontal thickened ring (cf. Fig. 7c with thickened pericarp in cross-section), apex obtuse. Distribution. This species is known from only one collection from low elevation (ca. 70 m) forests around Yuto in Chocó, Colombia (Fig. 1; Forero and Gentry 1989). The only other Potalia taxon known from Chocó is Potalia chocoensis, which differs from P. crassa by having fruits with conspicuously thickened rings (Fig. 4b). Phenology. The only collection of Potalia crassa was fruiting and collected during September (Table 1). Etymology. The species epithet crassa is derived from the thick-walled fruits (Fig. 6c). Notes. The characteristics of this species are poorly known and no complete inflorescences/infructesc- 682 SYSTEMATIC BOTANY [Volume 29 FIG. 6. Potalia crassa Struwe & V. Albert (Forero & Jaramillo 2698, NY [branch], MO [fruit]). A. branch. B. infructescence. C. longitudinal section of fruit. D. seed. 2004] STRUWE & ALBERT: POTALIA (GENTIANACEAE) 683 FIG. 7. Potalia elegans Struwe & V. Albert. A. flowering branch (Schultes et al. 18034, US). B. older stem (Clark 7163, NY). C– F. (Schultes et al 18034, US). C. flower at anthesis. D. opened corolla. E. anther. F. gynoecium at anthesis. G. infructescence (Schultes et al. 18127, GH). H, I. (Clark 7163, NY). H. mature fruit, also in longitudinal section. I. seed. 684 SYSTEMATIC BOTANY ences, flower buds, or open flowers are known. However, the characteristic thick and large fruits in combination with the large and coriaceous leaves mark this as a distinct species. Potalia crassa is the species with the largest fruits in Potalia. This is one of three species without a ring on the fruit, the others being Potalia elegans and P. maguireorum. This species differs in several vegetative and fruit characters from both of these Venezuelan/Brazilian species. Primarily the fruits are larger and with a thicker wall. The leaves of Potalia maguireorum are narrower and elliptic, whereas this species has typical larger, obovate leaves (as found in P. elegans). Potalia crassa has fruits that are distinctly longer and wider than P. elegans, and the yellow coloring of the lower side of the leaves, calyx, and inflorescence branches that are common in P. elegans are not present in P. crassa. Interestingly enough, the species with the next-largest fruits, Potalia chocoensis, is also an endemic of Chocó. This tendency for large-fruited species in Chocó was also remarked upon by Gentry (1986) for species in Arecaceae, Humiriaceae, and Myristicaceae, and could be related to the higher prevalence of mammaldispersed (compared to wind- and bird-dispersed) fruits that are found in Chocó. There is a tendency for taxa from this area to have unusually large and sclerophyllous leaves as well (Gentry 1986), which also agrees well with the features of Potalia chocoensis and P. crassa. 5. POTALIA ELEGANS Struwe & V. Albert, Harvard Papers in Botany 3: 185, fig. 2.—TYPE: COLOMBIA. Vaupés: Rı́o Negro, San Felipe and vicinity (below confluence of Rı́o Guainı́a and Rı́o Casiquiare), alt. about 600 feet, 18 509 S, 678 09 W, shrub, height 6 ft., flowers greenish yellow, 25 Oct 1952, R. E. Schultes, R. E. D. Baker, & I. Cabrera 18034 (holotype US!; isotype GH!). The collector R. E. D. Baker was inadvertently omitted from the citation of the type collection in the protolog (cf., Struwe and Albert 1998b). Fig. 7. Trees, treelets, or rarely shrubs, 2–12 m tall. Stems up to 150 mm in diameter (teste Williams 14352), below inflorescences (6–)10–17 mm in diameter, the bark fissured, greyish-brown; internodes short, 8–30(–45) mm long, 1–2 (–4) times as long as wide. Leaves 600– 1000 3 105–210 mm, oblanceolate to obovate, widest above middle, coriaceous, dark green above, paler below, dull when dry, base long-attenuate, apex acute to acuminate; petioles up to 60 mm long, 5–10 mm in diameter; margins entire or minutely crenate, thickened, sometimes revolute (only at the very edge); venation with secondary veins inconspicuous on both sides, the midrib prominent and triangular below, when dry the lamina becomes sinuous along the secondary veins. Inflorescences elongated, long-stalked, [Volume 29 up to ca. 50-flowered, (145–)170–290 mm long, up to 240 mm wide; branches often slightly bent or curved upwards, the first order of inflorescence branches usually very short, 7–25 mm long, the second order (55-) 90–165 mm long and third order 20–100 mm long; pedicels short, 8–20 mm long; bracts and floral bracts 3 mm long. Calyx lobes 7–9 3 5–6 mm, yellow to white, sometimes green, becoming loosely appressed in fruit and enlarged (9–11 3 7–10 mm), apex rounded. Corollas 9–10-merous, 11–15 mm long, the lower part 3–4 mm wide, upper part 5–6 mm wide, green and/or yellow (rarely whitish); lobes oblong, 6–8 3 2– 3 mm, 1/5–1/4 as long as the entire corolla, erect at anthesis, green with lighter margins. Anthers 5–7 mm long, slightly bent, with a small sterile apex. Fruits globose, 10–13 3 10–13 mm, ca. 2 times as long as calyx, green when fresh and filled with a clear, sticky liquid (teste Clark & Maquirino 7840), yellow to brown and dull when dry, without a horizontal thickened ring, apex acute, crowned by a small persistent remnant of the style base. Seeds flattened, cream-colored when young, dark brown when older, not winged, with hilum in the middle of one of the sides; testa pitted. Illustrations and Descriptions. Original description in Struwe and Albert (1998b: 185–189, fig. 2. Note that a NY sheet was erroneously listed as the source; the type material is located at GH and US). The species was also treated in ‘‘Flora of the Venezuelan Guayana’’, vol. 5 (Struwe and Albert in Struwe et al., 1999: 511, fig. 441). Distribution. Potalia elegans is known principally from lowland, evergreen, seasonally flooded forests, savannas, and Amazonian caatinga of the Casiquiare peneplains (upper Orinoco lowlands), Venezuela (Huber 1995) at 50–200 m elevation (Fig. 1). It occurs also in Colombia on scattered white-sand areas up to 500 m elevation in the states of Amazonas, Guainı́a, and Vaupés, areas that are also part of the Guayana Shield (Fig. 1). Only two collections are known from the Brazilian part of Rı́o Vaupés (one from terra firme forest). The Casiquiare region is approximately bounded by the Rı́o Orinoco to the northeast and the Rı́o Guainı́a-Rı́o Negro to the southwest. The type locality, San Felipe, lies just across the Rı́o Negro in the small flange of Colombia bordered by Venezuela to the east and Brazil to the west. Known Venezuelan localities beyond the Orinoco include the flanks of Cerro Yapacana, Rı́o Cunucunuma (which runs between Cerros Huachamacari and Duida), and La Esmeralda (at the base of Cerro Duida). No specimens of Potalia elegans are known from the Rı́o Ventuari basin (just northeast of Cerro Yapacana) nor from the poorly-collected river drainages between Sierra Parima and Sierra de la Neblina. Additionally, no specimens of Potalia elegans are known from Brazil, despite the proximity of San Car- 2004] STRUWE & ALBERT: POTALIA (GENTIANACEAE) los de Rı́o Negro and San Felipe. As seen on the distribution map (Fig. 1), there appear to be at least two disjunct populations of Potalia elegans on the western Guayana Shield, one eastern and one western. Future collections in the above and neighboring regions may establish whether these approximate range limits represent sampling artifacts or local endemicity. Phenology. Potalia elegans has been found flowering during February to March, May to June, and August to December. Fruiting specimens have been collected between October to June (Table 1). Vernacular Names. Hoja de gavilan (Venezuela), martiguaja (Colombia), martiguaje (Colombia), palo de lagarto mato (Venezuela), palo de mato (Venezuela), palo de rana (Venezuela), palo de temblador (Venezuela), temblador (Venezuela; Table 2). Etymology. The species epithet of Potalia elegans refers to the slender and graceful cymose inflorescences that are typical for the species and stand in contrast to the shorter and compressed ones borne by, e.g., P. amara and P. resinifera. Notes. Some specimens from the Rı́o Apoporis region (e.g., Cuatrecasas 6852, Schultes & Cabrera 17337) have shorter, more condensed inflorescences (all detached except Schultes & Cabrera 13726, which appears to have aborted most of its flower buds), but leaf and fruit characteristics are typical for Potalia elegans. The globose fruit of Potalia elegans does not become crownshaped during development as in P. coronata despite its similarly tightly appressed sepals. This would seem to indicate that a strong lateral fruit growth pressure (i.e., against the rigid sepals), apparently characteristic of Potalia coronata and other species, is absent from P. elegans, which also lacks the sterile fruit apex characteristic of most other Potalia species. Representative Specimens Examined. BRAZIL. AMAZONAS: Vaupés, Jauareté, Rio Negro, de Lemos Fróes 21221 (G, IAN, US). BRAZIL. AMAZONAS: Rio Uaupés, Taracúa, margem esquerda do rio, Cavalcante 746 (MG); Iauareté, margem do Rio Uaupés, próximo a front. Brasil-Colômbia, Ribeiro 1020 (MG). COLOMBIA. AMAZONAS: Rı́o Apaporis, raudal de Jirijirimo, Bernal et al. 1245 (COL); A la orilla del Rı́o Yarı́, Rı́o Yarı́, cerca de la desembocadura de la quebrada El Mochilero, Galeano et al. 1103 (COL). AMAZONAS-VAUPÉS: Rı́o Apaporis, Jino-gojé, entre los Rı́os Piraparaná y Popeyaká, Garcia-Barriga 14337 (US); Rı́o Apaporis, entre el Rı́o Pacoa y el Rı́o Kananari, Soratama, Schultes & Cabrera 13726 (US); Rı́o Vaupés, Mitú y alrededores, Schultes & Cabrera 13972 (GH). VAUPÉS: Mitú (selva del Vaupés), Cuatrecasas 6852 (US); Rı́o Negro, San Felipe (El Castillo), below confluence of Rı́os Guainı́a and Casiquiare, Schultes & López 9339 (IAN, NY, US); Rı́o Piraparaná (tribunary of Rı́o Apoporis), headwaters of Caño Teemeeña, Lobo Igarapé, Schultes 685 & Cabrera 17337 (GH, S, US); Rı́o Vaupés, Mitú and vicinity, Cerro Mitú, Schultes & Cabrera 17919 (GH); Schultes 22718 (ECON); Rı́o Negro, San Felipe and vicinity (below confluence of Rı́o Guainı́a and Rı́o Casiquiare), Schultes et al. 18127 (GH, US). VENEZUELA. AMAZONAS: Dpto. Atabapo, Rı́o Cunucunuma, Raudal Mapaco (Majako Shodö en lengua Ye’kuana), Fernandez 7797 (MO); Playa Alta, Maguire et al. 29354 (NY); Rı́o Orinoco, Cerro Yapacana, edge of Savanna no. 3, Maguire et al. 30807 (NY); northwest base of Cerro Yapacana, Maguire & Wurdack 34557 (NY); Planicie Anegadiza entre el Caño Cotúa y Cerro Yapacana, Marin 671 (MO); Terreños arenosos y húmedos a orillas del Rı́o Orinoco en los alrededores de San Fernando, Molina. & Barkley 18.V.177 (US); Rı́o Caname, a 1 km de la confluencia con el Rı́o Atabapo, Yanez 15 (MO); Dpto. Atures, Rı́o Sipapo, 2 km. abajo de la boca del Rı́o Guayapo, Carnevali et al. 1849 (VEN); Dpto. Casiquiare, Rı́o Casiquiare, Laguna del Paciba y sus alrededores, Colella et al. 2021 (NY); Capihuara, Casiquiare, en selvas rebalseras, Williams 15725 (VEN); Dpto. Rı́o Negro, 1–4 km south of Yavita, Berry et al. 5688 (NY); San Carlos de Rı́o Negro, ca. 20 km S of confluence of Rı́o Negro and Brazo Casiquiare, Clark 7163 (NY); IVIC Main study site, 4.3 km NNE on Solano road, Clark & Maquirino 7840 (MO); Klinge s.n. (NY); Liesner 3800 (MO, VEN); Mamurividi, laja along the Rı́o Pasimoni, Davidse & Miller 26717 (MO, NY); IVIC Study Area, 4 km east of San Carlos de Rı́o Negro, Liesner 3800 (MO, VEN); 0–2 km NE of San Carlos de Rı́o Negro, Liesner 8474 (MO); rock outcrop and forest around it, Mamurividi, Zona 10 Estacion Climatologica of Ministerio del Ambiente on Rı́o Pasimoni, Liesner 17197 (MO); Medio Casiquiare, desde la piedra La Esterita arriba del Raudal Cabrúa hasta boca del Pasiba, Stergios & Aymard 7642 (MO, NY). 6. POTALIA MAGUIREORUM Struwe & V. Albert, Harvard Papers in Botany 3: 189, fig 3.—TYPE: BRAZIL. Amazonas: Rio Negro, Rio Marı́e, ca. 30 km above mouth, village Macobeta, moist forest by river, unbranched treelet 3 m high, calyx yellow, petals green, ‘‘matugua’’, 9 Sep 1979, K. Kubitzki, C. E. Calderón, & H.-H. Poppendieck 79–216 (holotype NY!; isotypes INPA!, MG!, US!). Fig. 8. Shrubs or treelets 1–3 m tall. Stems up to 50 mm in diameter, below inflorescences 5–10 mm in diameter, the bark gray, fissured (fide Williams 15737); internodes in upper part of stems short, 12–19(–45) mm long, 1– 3(–8) times as long as wide. Leaves 180–360 3 28–79 mm, narrowly elliptic to oblanceolate, widest at or slightly above middle, coriaceous, dull when dry, green above, lighter below, base attenuate, apex bluntly acute, petioles up to 48 mm long, 2–4 mm in diameter; margins slightly revolute, thickened; venation with straight secondary veins at about 90 degree angle to 686 SYSTEMATIC BOTANY [Volume 29 FIG. 8. Potalia maguireorum Struwe & V. Albert. A. flowering branch (Maguire et al. 41657, NY). B. inflorescence branch (Kubitzki et al. 79–216, NY). C. young stem (Stergios & Aymard 9201, NY). D–G. (Kubitzki et al. 79–216, NY). D. flower. E. opened corolla. F. calyx and gynoecium (corolla removed). G. anther. H–J. (Stergios & Aymard 9201, NY). H. part of infructescence. I. fruit, also in longitudinal section. J. seed. 2004] STRUWE & ALBERT: POTALIA (GENTIANACEAE) midrib, the midrib prominently triangular below, when dry the lamina becomes sinuous along the secondary veins. Inflorescences rather compacted, not conspicuously long-stalked, up to 34-flowered, 32–135 mm long, up to 140 mm wide; branches orange, sometimes slightly bent or curved upwards, the first order of inflorescence branches 3–25 mm long, the second order 15–58 mm long and third order 5–28 mm long; pedicels not distinctly short, 9–13 mm long; bracts and floral bracts 2–3 mm long. Calyx lobes 4–5 3 4–5 mm, pale yellow, loosely appressed in fruit and not enlarged. Corollas 8–9-merous, 7–8 mm long, the lower part ca. 3 mm wide, upper part 5–6 mm wide, dark green; lobes elliptic, 2–4 3 ca. 2 mm, ca. 1/4 as long as the entire corolla, erect at anthesis, dark green and with lighter margins. Anthers 4–6 mm long, slightly bent, with a small sterile apex. Style ca. 3 mm long; stigma capitate, very small. Fruits globose, 6–10 3 6– 12 mm, 2–4 times as long as calyx, green when fresh, brown when dry, without a horizontal thickened ring, apex bluntly acute, not crowned by a small persistent remnant of the style base. Seeds flattened, dark brown, not winged, with hilum in the middle of one of the sides; testa pitted. Illustrations and Descriptions. Original description in Struwe and Albert (1998b: 189–191, fig. 3); Flora of the Venezuelan Guayana, vol. 5 (Struwe and Albert in Struwe et al., 1999: 515, fig. 442). Distribution. White-sand savannas and forests (Amazon caatinga) in the lowlands (100–200 m elevation) surrounding the upper Rio Negro and its tributaries in Brazil and Venezuela as well as Rı́o Casiquiare and its tributaries towards Neblina in southernmost Venezuela (Fig. 1). Phenology. Potalia maguireorum has been found flowering during September and October. Fruiting specimens have been collected in February to April, June, and October (Table 1). Vernacular Names. Matugua (Brazil, fide Kubitzki et al. 79–216) and pau-de-cobra (Brazil, fide Ramos et al. 688; Table 2). Etymology. This species honors Bassett Maguire and his wife Celia, prominent explorers and botanists of The New York Botanical Garden. One of the few specimens of this species was collected by them as well. Notes. The fruits of Potalia maguireorum are similar to those of P. elegans in that the fruit wall is equally thickened all around, with no ring or crown shape forming due to lateral fruit growth against rigidly appressed sepals. The two species are readily distinguished, however, by leaf shape (elliptic-lanceolate for this species), corolla color (dark green), and fruit size (up to 12 mm in diameter). Representative Specimens Examined. BRAZIL. AMAZONAS: Rio Negro, Praia Grande, mata de varzea, 687 igapo arenoso, sujeito a inundacao temporaria, Ramos et al. 688 (INPA); Basin of Rio Negro, Rio Uneiuxi, 200– 300 km. above mouth, Prance et al. 15552 (INPA, NY, US). VENEZUELA. AMAZONAS: Rio Negro, Anavilhanas, igapó, Goulding 2208 (MG); Dpto. Casiquiare, Caño ‘‘San Miguel’’, Aymard 9265 (NY); Rios Pacimoni, Yatua, Casiquiare, sabanita on right bank of Rı́o Pacimoni 12 km. below mouth of Rı́o Yatua, Maguire et al. 41657 (NY); Bajo Chimoni, Stergios & Aymard 9201 (MO, NY); caño Pendare, Capihuara, Alto Casiquiare, Williams 15737 (US, VEN). 7. POTALIA RESINIFERA Mart., Nov. Gen. Sp. Pl. 2: 90, t. CLXX. 1826. Potalia amara var. resinifera (Mart.) Progel in Mart., Fl. Bras. 6(1): 268. 1868.—TYPE: BRAZIL. Amazonas: Habitat in sylvis ad Barra do Rio Negro, Proviniciae Rio Negro, K. F. P. von Martius s.n. (holotype: M!). Several unnumbered Martius sheets are present at M and one at G, but the remaining ones are not considered isotypes. Figs. 4d–g, 9, and 10a–j. Shrubs or treelets, 0.5–3(–8, teste Pipoly & Murillo 15519) m tall. Stems up to 20(–150; teste Pipoly & Murillo 15519) mm in diameter, below inflorescences 4–11 mm in diameter, the bark fissured, brown; internodes short to long, 16–130 mm long, 1–15 times as long as wide. Leaves 270–800 3 72–190 mm, broadly oblanceolate to obovate, widest above middle, chartaceous (less often coriaceous), glossy to dull when dry, green above, lighter below, base long-attenuate, apex acute to acuminate; petioles up to 42 mm long, 3–6 mm in diameter; margins flat to slightly revolute, not thickened; venation with secondary veins usually arcuate, diverging from the midvein usually at 50–80 degree angle, the midrib prominent below, when dry the lamina sometimes becomes sinuous along the secondary veins. Inflorescences compacted, not long-stalked, up to ca. 40-flowered, 25–76 mm long, up to 116 mm wide; branches usually straight, not curved, the first order of inflorescence branches 3–9 mm long, the second order 5–44 mm long and third order 8–25 mm long; pedicels short, 8–20 mm long; bracts and floral bracts 2–5 mm long. Calyx lobes 4–7 3 4–5 mm, yellow or orange, usually becoming loosely spreading to recurved in fruit and often enlarged (in fruit 4–8(–10) 3 4–7 mm). Corollas 9–10-merous, 11–20 mm long, the lower part 2–4 mm wide, upper part 5–8 mm wide, yellow and green (rarely basally orange); lobes ovate, 2–3 3 ca. 2 mm, ca. 1/3 as long as the entire corolla, slightly to distinctly spreading at anthesis, green with lighter (yellow) margins. Anthers 5–7 mm long. Fruits globose, 8–17 3 11–16 mm, ca. 2 times as long as calyx, green, yellow to orange when fresh, brown when dry, with a distinct horizontal thickened ring (rarely less 688 SYSTEMATIC BOTANY [Volume 29 FIG. 9. Potalia resinifera Mart. (Zaruchii & Barbosa 5801, MO). A. flowering branch. B. young stem with ocrea. C. flower at anthesis. D. opened corolla and gynoecium. E. anther. F. infructescence. G. dry fruit. H. fruit at maturity, also in longitudinal section. I. seed. 2004] STRUWE & ALBERT: POTALIA (GENTIANACEAE) 689 FIG. 10. Potalia resinifera Mart. and P. yanamonoensis Struwe & V. Albert. A- D. Potalia resinifera, typical form (Cid & Lima 3533, NY). A. flowering branch. B. flower at anthesis. C. opened corolla with gynoecium. D. anther. E. Potalia resinifera, fruit (Prance et al. 25571, MO). F, G. Potalia resinifera (Ducke 364, S). F. mature fruit, also in longitudinal section. G. seed. H–J. Potalia resinifera (Henderson et al. 217, NY). H. mature fruit. I. longitudinal and cross section of fruit. J. seed. K. Potalia yanamonoensis Struwe & V. Albert, flower (Vasquez & Jaramillo 553, MO). L, M. Potalia yanamonoensis (Vasquez et al. 5132, MO). L. fruit at maturity. M. fruit with removed calyx, also in longitudinal section. 690 SYSTEMATIC BOTANY developed), apex acute to obtuse, rarely crowned by a small persistent remnant of the style base. Illustrations and Descriptions. This species was treated in ‘‘A general history of dichlamydeous plants’’, vol. 4 (Don 1837–38: 68), in ‘‘Flora of Peru’’ (MacBride 1959: 269; as Potalia amara), was illustrated in Leeuwenberg and Leenhouts (1980: fig. 2.29 pro parte; as P. amara), and in ‘‘Flora of the Venezuelan Guayana’’, vol. 5 (Struwe and Albert in Struwe et al. 1999: 515, fig. 514). Distribution. Potalia resinifera is the most widespread of the species of Potalia in the Neotropics. It occurs commonly in the understory of forests at 90– 1250 m elevation from Andean and Amazonian Colombia, southern Venezuela, Amazonian Brazil, Andean Ecuador, and southwards to Amazonian and Andean Peru and Bolivia (Fig. 2), and possibly also Guyana, from where only one non-fruiting specimen is known. It reaches its highest elevational range in Huanuco and Cuzco, Peru, and Caquetá, Colombia. Potalia resinifera is found in wet, tropical, primary, secondary, or disturbed forests, inundated or not, ‘‘varzea’’ and ‘‘campinarana’’ forests, mountain crests, river banks, gallery forests, and roadsides. It grows on substrates such as clay or humus-rich soils, terra firme, laterite, red oxisoils, white-sand, and fine, recent sediments. In the southernmost part of its distribution, Potalia resinifera also has been found in semi-deciduous gallery forests (teste Nee 41199), riverine forests, savanna margins, and on savannas with scattered, brushy ant and termite mounds (‘‘pampa termitera’’, teste Guillén & Choré 2440 and Nee 41278). Phenology. Potalia resinifera appears to be flowering and fruiting year-round. Flowering and fruiting specimens have been found during every month of the year (Table 1). Vernacular Names. Ahuaramacha (Bolivia), Amargo Andrés (Colombia [appr. 5 approximate species determination only]), anabi (Macbride, 1959), boyomemo (Ecuador: Quichua), caspisenango (Peru [appr.]), cháde-bougri (Brazil), Chañetsopan (Peru [appr.]), chirosenango (Peru), cuararina (Ecuador: Quichua), curarina (Brazil, Ecuador, Peru), curarina sacha (Peru), curarina sacha hembre (Peru [appr.]), curare (Ecuador: Quichua), sacha macho (Peru [appr.]), jaisoldao (Colombia [appr.]), kararái (Peru), kúranai (Peru), mapiguari (Colombia), ma-mee-mhoy’-da (Colombia), matiguaja (Colombia), ócaji cappuva (Peru [appr.]), okajı́ kahpuú (Peru [appr.]), palu huacu (Ecuador: Quichua), pao-de-cobra (Brazil), pau-de-cobra (Brazil), sacha curarina (Colombia, Ecuador, Peru), shin-wee’ (Colombia), turarina sacha (Peru), yampak (Ecuador: Shuar), yanapa (Ecuador: Quichua; Table 2). Etymology. This species was named by von Martius (1826–27) after the abundant resins secreted by the colleters in the leaf axils. Notes. We have found no evidence that this species [Volume 29 has been accepted by any botanist after Don (1837–38; except for our recent treatment in Struwe et al., 1999). Don, however, correctly described some of the distinctive characteristics that separate the two species known at the time: Potalia amara by its yellow (-white) corollas and P. resinifera by its green corollas with yellow margins. Progel (1868) included Potalia resinifera as a subspecies under P. amara, and discussed the possibility that it was indeed a separate species. Nevertheless, Potalia resinifera is an extremely variable taxon. Leaf texture varies from coriaceous to chartaceous (the most common), but the greatest differences are in fruit shape. Most specimens of Potalia resinifera have strongly to moderately ringed fruits (e.g., Figs. 10e, h, 9h); the rings appear to be formed by fruit growth against sepals, which are initially rigid during fruit development. The sepals often reflex during fruit development (e.g., Fig. 10h), but rigidly appressed sepals are also known, with ringed or even crown-shaped fruits resulting (Figs. 10l, m). A number of Potalia resinifera specimens lack a readily discernable fruit ring (e.g., Fig. 10f), especially when the mature fruit is bloated with seeds. Due to this broad variation among cardinal characters for the other Potalia species, the present circumscription of P. resinifera is not entirely satisfactory. The reason for this great range of morphological variation is not clear. From the phylogenetic analysis (Frasier et al., 2002), this species appears to be one of the younger ones and also the one that has conquered the most habitat and soil types. In fact, this species is the only one that can be considered ‘weedy’. The current distribution and morphological variation can have been affected by paleoclimatic cooling and drying events that reshaped the tropical rainforests of South America, causing repeated and restricted gene flow among meta-populations. The most similar species appear to be Potalia chocoensis and P. yanamonoensis, both of which are poorly known. Several collections from Sierra de Macarena (Meta, Colombia; Philipson 2330 and Philipson et al. 2060) show some intermediate traits between Potalia resinifera and P. chocoensis, but differ from P. chocoensis in having smaller, less robust fruits and thinner leaves. These collections fit into the overall variation seen in the rather variable P. resinifera and show most similarities to specimens from Amazonas (Venezuela), Napo (Ecuador) and Antioquia (Colombia). No specimen seen from Antioquia has fully mature fruits, so determinations for these collections are preliminary. Representative Specimens Examined. BOLIVIA. BENI: Prov. of Vaca Diez, 25 km E of Riberalta on the road to Guayaramerı́n, Solomon 6388 (MO, NY). LA PAZ: Prov. Iturralde, Siete Cielos, Rı́o Manupare, Solomon 16938 (MO). PANDO: Prov. Madre de Dios, Jatata, en las riveras del Rı́o Manupare, en partiendo Cam- 2004] STRUWE & ALBERT: POTALIA (GENTIANACEAE) pamento de la Western Geophysical, Rueda 997 (MO). SANTA CRUZ: Prov. Velasco, campamento LA TOLEDO, Guillén & Choré 2440 (MO). BRAZIL. ACRE: Highway Abunã to Rio Branco, km 242–246, vicinity of Campinas, Forero et al. 6365 (K, NY, S, US). AMAZONAS: Manaus, Estrada do Aleixo, silva non inundati, Ducke 1766 (A, K, IAN, MG, NY, US). GOIÁS: Igarapé Ucuricaca, região de Araguantins, de Oliveira 1816 (IAN). MARANHÃO: S of Km 157 of BR316, Araguaná, Daly et al. D216 (MG); Rio Alto Turiaçu, Barraquinha, Jangoux & Bahia 142 (MG). MATO GROSSO: Rio Aripuanã, at the base of Salto dos Dardanelos, west side of river, Berg et al. P18494 (NY). PARÁ: BR 163, Cuiabá-Santarém Highway, vicinity of Igarapé Kazuo, km 1234, Prance et al. 25571 (K, MO, NY, S, US). RONDÔNIA: Basin of Rio Madeira, Km. 214– 215 Madeira-Mamoré railroad near Abunã, Prance et al. 5971 (K, NY, US). COLOMBIA. AMAZONAS: Mun. de Leticia, Parque Nacional Natural Amacayacu, Trocha de Matamatá a Amacayacu, Pipoly & Murillo 15519 (MO). ANTIOQUIA: Lugar, Vera, camino al Rı́o Segovia (Segovia Antioquia), Renterı́a et al. 1704 (COL, MO, NY). CAQUETÁ: Cordillera Oriental, vertiente oriental, Sucre, Quebrada de La Calaña, Cuatrecasas 9202 (US). CORDOBA: Mun. de Tierralta, bosques entre los Rı́os Esmeralda y Sinú, 2 km arriba de la confluencia, Bernal et al. 1153 (COL). META: Sierra de la Macarena, Cano Cierve, Phillipson et al. 2060 (COL, S, US). NARIÑO: Mun. de Tumaco, Curay, Madrigal et al. 785 (NY). VAUPÉS: Rı́o Apaporis, entre el Rı́o Pacoa y el Rı́o Kananari, Soratama, Schultes & Cabrera 12716 (NY, US). VICHADA: Parque Nacional Natural ‘‘El Tuparro’’, along Rı́o Orinoco at old site of Maipures, Zarucchi & Barbosa 3550 (MO). ECUADOR. MORONA-SANTIAGO: Vieja Cordillera de Cucutú, km 16 on Mendez-Morona road paralleling the Rı́o Namangoza, Dorr & Valdespino 6314 (NY). NAPO: Cantón Aguarico, Parque Nacional Yasunı́, Lagunas de Garza Cocha, Cerón & Gallo 4927 (MO). PASTAZA: Puerto Sarayacu, Lugo S. 3900 (NY). SANTIAGOZAMORA: west of the Rı́o Upano, along the Rı́o Chupiangas, Prieto ChuP-15 (NY, US). GUYANA. Rupununi District, Kuyuwini Landing, Kuyuwini River, Jansen-Jacobs et al. 2567 (NY). PERU. AMAZONAS: 1 km. de La Poza, banda oeste del Rı́o Santiago, Peña 41 (MO). CUZCO: Prov. de la Convencion, along Rı́o Mapituriani, a tributary of the Rı́o Apurimac, Wasshausen & Encarnación 680 (US). HUÁNUCO: Dist. Puerto Inca, carretera marginal ca. 14 km from a point across the Rı́o Pachitea from Puerto Inca, Smith 1286 (MO). JUNı́N: Puerto Yessup, Killip & Smith 26234 (NY, US). LORETO: Prov. Maynas, Pucaurquillo, Pevas, Ayala et al. 2741 (MO). MADRE DE DIOS: Tambopata, along Rı́o Tambopata, Gentry et al. 45720 (MO). PASCO: Prov. Oxapampa, Central Selva, Palcazu Valley, Iscozacin, above PEPP project Camp. SAN MAR- 691 TIN: Prov. Mariscal Caceres, 12 km W of Tocache Nuevo on Carretera marginal, Gentry et al. 25612 (MO). UCAYALI: Prov. Coronel Portillo, bosque Von Humboldt, Young 951 (MO). VENEZUELA. AMAZONAS: Cerro Neblina base camp in Rı́o Mawarinuma, Gentry & Stein 46816 (MO). 8. Potalia turbinata Struwe & V. Albert, sp. nov.— TYPE: COSTA RICA. Limon: Cordillera de Talamanca Ridge separating Quebrada Cañabral from Rı́o Barbilla, and slope leading down to the latter, 108 029 N, 838 269 W, 200–400 m., erect, unbranched shrub ca. 2 m tall in disturbed primary forest, lvs coriaceous, rather flexible, yellow-green below, midrib quadrangular-raised above, yellow-green, fruits presently green, 4 Sep 1988, M. Grayum, Herrera, & Robles 8753 (holotype MO!). Fig. 11. Potalia amarae Aubl. similis sed fructibus dorsiventraliter complanatis sepalisque fructeris plerumque recurvis vel laxe appressis diversa. Shrubs, rarely trees, up to 4.5 m tall. Stems up to 14 mm in diameter, below inflorescences 8–12 mm in diameter, old bark not seen; internodes short or long, 9– 120 mm long, 1–ca. 10 times as long as wide. Leaves 385–665 3 152–180 mm, oblanceolate, widest above middle, chartaceous to coriaceous, dull when dry, green above, yellow-green below, base attenuate, apex obtuse to slightly acute; petioles up to 21 mm long, 5– 6 mm in diameter; margins not conspicuously thickened, slightly revolute; venation with secondary veins prominent, the midrib triangular and prominent below, when dry the lamina is not sinuous along the secondary veins. Inflorescences compacted, short-stalked, up to 30-flowered, 64–95 mm long, up to 115 mm wide; branches straight, the first order of inflorescence branches 10–12 mm long, the second order 20–24 mm long and third order 8–11 mm long; pedicels short, 9– 12 mm long; bracts and floral bracts 1–2 mm long. Calyx lobes 5–6 3 4–8 mm, yellow to white (rarely green, teste Hammel 13716), becoming loosely or distinctly spreading in fruit and not enlarged. Corollas 9– 10-merous, 9–12 mm long, the lower part ca. 3 mm wide, upper part ca. 5–7 mm wide, white, yellow, pale green, yellow-green, or green; lobes narrowly elliptic, ca. 4 3 ca. 1.5 mm, ca. 1/3 as long as the entire corolla, slightly spreading at anthesis, green (or sometimes yellow?) with narrow lighter-colored margins. Anthers ca. 3–4 mm long. Fruits turbinate, 10–14 3 11–18 mm, green when fresh, brown when dry, with a thickened horizontal ring, apex obtuse to slightly pointed, not crowned by a small persistent remnant of the style base. Illustrations and Descriptions. This species was treated as Potalia amara in floras of Central America, such as ‘‘Flora of Costa Rica’’ (Standley 1938: 921), ‘‘Flora of Panama’’ (Blackwell 1968; p. 405, fig. 4), and ‘‘Man- 692 SYSTEMATIC BOTANY [Volume 29 FIG. 11. Potalia turbinata Struwe & V. Albert. A. fruiting branch (Grayum et al. 8753, MO). B. inflorescence (Hammel 13716, MO). C–F. (Stevens et al. 25009, MO). C. flower at anthesis. D. opened corolla. E. anther. F. gynoecium. G, H. (Stevens & Montiel 24489, MO). G. fruit at maturity, also in longitudinal section. H. seed. I. dry fruit (Grayum et al. 8753, MO). 2004] STRUWE & ALBERT: POTALIA (GENTIANACEAE) ual de la Flora de Costa Rica’’’s web site (http:// www.mobot.org/manual.plantas/019239/S019286.html; the photo provided at this web site for ‘‘Potalia amara’’ might be P. turbinata, however, no locality or voucher information is given). Distribution. Potalia turbinata occurs sporadically in eastern Costa Rica (Alajuela, Cartago, Heredia, Limon) and Panama (Bocas del Toro; Fig. 1). It grows in evergreen, wet primary forests, swamp forests, along roadsides, disturbed areas, or in cloud forests on mountain slopes or flat areas at 5–350 meters elevation. Phenology. Potalia turbinata has been found flowering during March to May, and fruiting in May, July to September, and December (Table 1). Vernacular Names. No vernacular names are known for Potalia turbinata. Etymology. Potalia turbinata is named after its distinctly shaped fruits, which are reminiscent of turbans. Notes. The fruit rings of Potalia turbinata probably develop only under early pressure against the sepals as these organs are not at all strongly appressed against the fruits at maturity. The chartaceous leaves and obconical fruits (when mature and fresh; see Fig. 11g) are similar to those of Potalia amara, but the strongly dorsiventrally flattened and turbinate fruits of P. turbinata are still quite distinct. Additionally, the sepals of Potalia amara may extend all the way to the fruit ring, whereas the sepals of P. turbinata are shorter and do not extend to the ring (see Figs. 3h, k and 11g, i). All specimens of Potalia we have examined from Central America have the above fruit characteristics. Representative Specimens Examined. COSTA RICA. HEREDIA: Finca La Selva, the OTS Field Station on the Rı́o Puerto Viejo just E of its junction with Rı́o Sarapiquı́, Jacobs 3009 (MO). LIMON: Above unnamed tributary of Rı́o Siquirres, Dodge et al. 6076 (GH, MO 2 sheets); Limon, 7 km SW of Bribri, Gómez et al. 20322 (MO); Cerro Colonel, E of Rı́o Zapote, along and above new road within 1 km of Rı́o Colorado, Stevens & Montiel 24309 (MO); Cerro Colonel, E of Laguna Danto, Stevens & Montiel 24489 (MO); along shore of Laguna de Enmedio, NNE of Barra del Colorado, Stevens et al. 25009 (MO). PANAMA. BOCAS DEL TORO: Along oleoducto road between continental divide and Chiriqui Grande, Hammel 13716 (MO); Chiriqui Lagoon, on Cayo Agua, McPherson 11464 (MO); vicinity of Chiriqui Lagoon, Fish Creek Mts., von Wedel 2369 (GH, MO, US). 9. Potalia yanamonoensis Struwe & V. Albert, sp. nov.—TYPE: PERU. Loreto: Prov. Maynas, Yanamono (Rı́o Amazonas), 38 309 S, 728 509 W, alt. 106 m, bosque primario, arbusto, 2 m, flores amarillentas, 25 Jun 1984, R. Vásquez et al. 5132 (holotype NY!; isotype MO!). Fig. 10k–m. 693 Potalia resinifera Mart. similis, sed fructu cum annulo undulato differt. Treelet, 1.5–3 m tall. Stems up to ca. 13 mm in diameter, below inflorescences ca. 10 mm in diameter, the bark slightly fissured, light brown; internodes short to long, ca. 38–100 mm long, ca. 4–10 times as long as wide. Leaves ca. 340–430 3 120–180 mm, broadly oblanceolate to obovate, widest above middle, chartaceous, somewhat glossy when dry, brownishyellowish when dry, lighter below, base long-attenuate, apex acute to acuminate; petioles up to 40 mm long, 4–6 mm in diameter; margins flat, not thickened; venation with secondary veins arcuate, diverging from the midvein usually at 50–80 degree angle, the midrib prominent below, when dry the lamina sometimes becomes sinuous along the secondary veins. Inflorescences compacted, not long-stalked, up to ca. 30-flowered, ca. 100 mm long, up to ca. 90 mm wide; branches usually straight, not curved, the first order of inflorescence branches ca. 10 mm long, the second order ca. 10–30 mm long and third order ca. 12–25 mm long; pedicels short, ca. 5–10 mm long; bracts and floral bracts ca. 3 mm long. Calyx lobes 4–5 3 ca. 4 mm, yellow, usually becoming loosely appressed in fruit and often enlarged (in fruit 8–10 3 7–8 mm). Corollas 9–10-merous, 10–13 mm long, the lower part ca. 3–4 mm wide, upper part ca. 5–7 mm wide, yellowish or green and yellow; lobes ovate, ca. 5 mm long, ca. 3 mm wide, ca. 1/3 as long as the entire corolla, barely spreading at anthesis, green with lighter (yellow) margins. Anthers ca. 5 mm long. Fruits globose, 8–12 mm long, 8–10 mm wide, ca. 1–1.5 times as long as calyx, of unknown color when fresh, light brown when dry, with a distinct undulating, horizontal thickened ring, apex obtuse, not crowned by a small persistent remnant of the style base. Distribution. Potalia yanamonoensis is only known from three collections from Yanamono, Peru, which is close to Iquitos and the mouth of Rı́o Napo in Loreto (Fig. 1). It grows in lowland primary forests at ca. 100 m elevation. The type locality, Yanamono, was one of the sites for Al Gentry’s classic biodiversity inventories, and one of the greatest densities of woody species recorded was found there (ca. 300 species per hectare; Gentry 1988a, b). Phenology. Potalia yanamonoensis has been collected with flowers in October and with fruits in June (Table 1). Etymology. This species is named after its narrow distribution around Yanamono (Loreto, Peru). Notes. Potalia yanamonoensis is morphologically similar to P. resinifera, especially with regard to flower and fruit morphology. The main difference is the undulation of the fruit ring in Potalia yanamonoensis. Representative Specimens Examined. PERU. LORETO: Prov. Maynas, Quebrada Yanomono, Explorama 694 SYSTEMATIC BOTANY tourist camp, Rı́o Amazonas above mouth of Rı́o Napo, Gentry et al. 27505 (MO); Yanamono tourist camp, 50 miles NE of Iquitos, 38 309 S, 728 509 W, alt. ca. 106 m, bosque primario, arbusto 2 m, flores amarillo-verdoso, 18 Oct 1980, Vásquez & Jaramillo 553 (MO). DISCUSSION Vegetative Morphology. HABIT. Potalia contain species that are up to 12 meters tall, monopodially branched, single stemmed shrubs or trees (Hallé 1980: fig. 3.4.3a–d). The leaves are characteristically crowded at the branch apex, which is terminated by an inflorescence or infructescence (e.g., Figs. 3a, 11a). This is similar to the appearance of arborescent species of the sister genus Anthocleista, which differs by most species being ‘‘candelabra’’ branched (Hallé 1980: fig. 3.5; Leeuwenberg 1961). More highly branched plants rarely occur in Potalia, and several unbranched stems from a tight basal clump have also been reported (teste Nee 41278). Like most Gentianaceae and all Potaliinae (Anthocleista, Fagraea, and Potalia), all organs of this genus are glabrous. The presence of latex, rare in the family, has been recorded in a few cases but not consistently (see genus description). STEMS AND WOOD. The stems of Potalia are always terete and the bark becomes fissured with age (Fig. 7b). They have light-colored, moderately hard wood with a soft and separating pith, causing the stems to become hollow in the center. The anatomical features of the wood are very similar among Anthocleista, Fagraea, and Potalia, as reviewed by Mennega (1980). LEAVES. Fresh leaves are usually thick, fleshy, and sometimes brittle. When dry, they vary from thin-chartaceous (typically Potalia amara, P. turbinata, P. yanamonoensis) to very thick and coriaceous (P. chocoensis, P. coronata, P. crassa, and P. elegans). Potalia exhibits a mixed leaf venation with pinnate secondary veins that are usually free from each other in the lower part of the leaves but connected at the apex (camptodromous and brochiodromous venation, respectively). The primary vein is conspicuous in all species, with a raised and triangular midrib below, whereas the secondary veins are inconspicuous to invisible in Potalia coronata, P. crassa, and P. elegans. COLLETERS. As in most Gentianaceae, as well as in other families of the Gentianales, colleters are present in the axils of the leaves (e.g., Robbrecht 1988). These multicellular glands secrete a resin (thus the name Potalia resinifera), which is particularly abundant at young branch tips. STIPULES. Interpetiolar structures are common in Gentianaceae, despite the family lacking ordinary stipules (interpetiolar stipules occur in the related families Gelsemiaceae, Loganiaceae, and Rubiaceae). Potalia has conspicuous, but small, persistent, interpetiolar sheaths at the nodes, which are similar to the ones [Volume 29 found in Macrocarpaea Gilg and Chorisepalum Gleason & Wodehouse. Such sheaths are also found in some Fagraea spp. (Leenhouts 1963), but not in Anthocleista, which has species with axillary scales and similar structures (Leeuwenberg 1961). Most genera in the Gentianaceae have only a small, inconspicuous interpetiolar line. Reproductive Morphology. INFLORESCENCES. The inflorescences are up to ca. 50-flowered, terminal, and regularly branched dichasial cymes (rarely thyrsoid with a central stalk), sometimes with some of the terminal flowers aborted (cf. Fig. 5b). The inflorescence branches as well as the pedicels and peduncles are conspicuously bright yellow to orange (seldom pale green in Potalia elegans). Anthocleista and most species of Fagraea also have dichasial cymes, but not with this branch color (Leenhouts 1963; Leeuwenberg 1961). FLOWERS. The flowers of Anthocleista and Potalia are unique among Gentianaceae in having tetramerous calyces and supermerous corollas and androecia. Potalia has 8–10 corolla lobes, whereas Anthocleista has 9–16(– 24) corolla lobes (Leeuwenberg 1961; see also photo in van Wyk and van Wyk 1997). A similar case is found in the distantly related Chorisepalum, which also has a tetramerous calyx, but a six-merous corolla and androecium (Ewan 1947; Maguire 1981; Struwe et al. 1999). Fagraea, the most closely related genus to Anthocleista and Potalia (Struwe et al. 2002), differs in having pentamerous calyces and corollas. The reason for this doubling of corolla lobe primordia in Anthocleista and Potalia is unknown, but SEM studies of corolla primordia in Anthocleista amplexicaulis show an initial formation of four lobe primordia at each corner of a square-shaped flower primordium, with additional lobes forming later, midway down the sides of the square, between the initial four primordia (Struwe 2002). CALYX. The calyces are always tetramerous and divided to the base. The calyx gives a decussate impression with 2 opposing outer and 2 opposing inner sepals, but the sepals are derived from one whorl and this is an example of imbricate aestivation. The inner pair of lobes is slightly smaller than the outer pair. The calyces are similar to those of Anthocleista and most Fagraea in their coriaceous texture and rounded, obtuse lobes. The calyx lobes are persistent in fruit, and the inner lobes often become ruptured by the developing fruit. The lobes are usually erect and tightly appressed to the fruit, but can be spreading or sometimes recurved in Potalia amara, P. resinifera, and P. turbinata. COROLLA. The corolla color varies from white, cream, yellow to green, or combinations of these often with a paler staminal-corolline corolla tube and corolla lobe margins. Green corollas are not known from any other genus in Potalieae, but occur in tribe Helieae 2004] STRUWE & ALBERT: POTALIA (GENTIANACEAE) (e.g., Chelonanthus alatus (Aubl.) Pulle and Rogersonanthus arboreus (Britton) Maguire & B.M.Boom; Struwe et al. 2002). The texture of the corolla is thick and coriaceous as in Anthocleista and most Fagraea (Struwe and Albert 1997). The corolla lobes are lanceolate and erect to slightly spreading at anthesis (Fig. 3e), but never recurved as in some Anthocleista (Leeuwenberg 1961). ANDROECIUM. The stamens are isomerous with the corolla and not exserted from the corolla mouth (as opposed to Anthocleista stamens [Leeuwenberg 1961: fig. 1.5]). In an earlier paper we hypothesized that the thickened ring below the stamens in the Fagraea berteroana group represented a case homologous to the fused filaments of Anthocleista and Potalia (Struwe and Albert 1997). The Fagraea berteroana group was consequently placed as the sister group to Anthocleista and Potalia in this morphology-based cladistic study. However, recent molecular data show that the broadened tribe Potalieae exhibits extensive variation of corolline appendages and filaments (cf. in Enicostema and Neurotheca; Struwe et al. 2002). Each anther has a small, sterile, apical appendage (Fig. 9e), a character that is also found in the Malagasy species of Anthocleista (Leeuwenberg 1961: fig. 1.7) and in species of Helieae (Struwe et al. 1997). POLLEN. The pollen of Potalia has been described by Punt (1978, 1980) and Nilsson (2002). All investigated species have 3-porate pollen of Potalia-type that are released in monads, except for P. maguireorum, which bears tetrads (Nilsson 2002). The pollen is very similar to that of Anthocleista (Punt 1980), but not to other Gentianaceae (Nilsson 2002). GYNOECIUM. The upper part of the gynoecium is sterile and globose before fruit development (Fig. 3g) and often forms a sterile apex in the mature fruits (Fig. 10f). However, Potalia elegans and P. maguireorum lack a conspicuous, sterile apex in the mature fruits (Figs. 7h, 8i). Potalia has bilocular ovaries, which appear to have axile placentation. This is an unusual feature in Gentianaceae, in which there is great placental variation and predominantly parietal placentation (Lindsey 1940). The often deeply inrolled and bilobed placentas can secondarily fuse in the middle to give rise to secondarily axile placentation (e.g., in Anthocleista, Fagraea, and many genera of the Helieae; Leeuwenberg 1961: fig. 9.10; Leeuwenberg and Leenhouts 1980; Struwe et al. 1997, 2002). Many taxa also have basally bilocular ovaries that become unilocular with parietal placentation towards the apex. Therefore, multiple and conflicting views on ovary morphology can be found in the literature, some clearly deriving from confusion as to how to interpret the inrolled and superficially bilobed placentas (each of which is derived ontogenetically from the ovule-bearing tissues of separate carpels; cf. Gopal Krishna and Puri 1962: figs. 79–81, 94). Plasticity in carpel fusion has caused considerable con- 695 fusion when interpreting the number of cells in the ovary and type of placentation in Gentianaceae, especially in Potalia and Anthocleista. The ovaries of Potalia were originally said to be 3celled (Aublet 1775; Schreber 1789), but reports range from 2-celled (von Martius 1826–27; Standley 1938), 2– 4-celled (Blackwell 1968), 4-celled at the base and 2celled at the apex (Leeuwenberg and Leenhouts 1980; MacBride 1959; Solereder 1892–95), to ‘‘4-carpellate’’ (Woodson and Schery 1942). The situation in Anthocleista is similar with reports of 1–4-celled (Leeuwenberg 1972, 1984; Leeuwenberg and Bamps 1979), 2celled (Coates Palgrave 1991), 2 or spuriously 4-celled (Bruce and Lewis 1960), 4-celled at the base and 2celled at the apex (Solereder 1892–95), or even 4-celled ovaries (Leeuwenberg 1983; Leeuwenberg and Leenhouts 1980). All individuals of Potalia that we have investigated have a bilocular ovary with placentation that at least appears to be axile. Future anatomical studies of immature and mature fruits of Potalia will show if the apparently axile, simple placentation is the result of strongly intruding parietal placentas or might instead be true axile placentation. In Gentianaceae, true axile placentation throughout the entire ovary is otherwise known only from tribe Exaceae, a relatively basal lineage within the family some phylogenetic distance away from the Potalieae (Gilg 1895; Struwe et al. 2002). There are also no signs of extensive postgenital fusion in the mature fruits of Anthocleista, Fagraea, or Potalia, and the ovaries instead appear to be congentially fused (L. Struwe, unpubl.). Postgenital fusion of carpels is the more prevalent condition in Gentianaceae (cf. Aripuana Struwe, Maas, & V. Albert; Struwe et al. 1997). The stigma is small and capitate as in many Fagraea species (except for the Fagraea berteroana group; Struwe and Albert 1997) and Anthocleista (Leenhouts 1963; Leeuwenberg 1961). The tendency towards bilobed stigmas, as seen in Anthocleista amplexicaulis, for example, has not been observed in Potalia. FRUIT. Fleshy fruits are also uniformly present in Anthocleista and Fagraea, although a few species of Fagraea have secondarily dehiscent fleshy fruits. The fruits of Potalia are smaller (6–19 3 6–18 mm) than the fruits of Anthocleista, whereas Fagraea includes species with both smaller (e.g., F. fragrans Roxb. and F. umbelliflora Gilg & Benedict) and larger fruits (e.g., F. auriculata Jack). The fruits of Potalia are green when fresh and become yellow to brown when dry. The upper sterile part of the gynoecium (see discussion above) develops into a large, sterile, thickwalled apical part of the fruit (e.g., Fig. 3i) in all Potalia species except P. elegans and P. maguireorum (Figs. 7h, 8i). This is best seen in longitudinal sections of the fruits. In contrast, fruits of Anthocleista and Fagraea 696 SYSTEMATIC BOTANY have evenly thick pericarp. Additionally, most species of Potalia have a sterile thickened ring around the middle of the fruit, which is especially well developed in P. coronata, some P. resinifera, P. turbinata, and P. yanomonoensis. This ring is interpreted to result from mechanical constraints imposed on the developing fruit by the sterile apical parts as well as the persistent and tightly appressed calyx. The outer surface of the pericarp is smooth and often glossy. SEEDS. Studies with scanning electron microscopy show that the testa is reticulate with narrow furrows leading to a central pit in each testa cell (Bouman et al. 2002: fig. 5.15g). The hilum is distinct and centered on one of the flattened sides, as in Anthocleista (Leeuwenberg 1961: fig. 1.10–11). No wings are present. Phylogeny and Intergeneric Relationships. Potalia, together with Anthocleista and Fagraea, belongs to subtribe Potaliinae in tribe Potalieae, which is one of six major clades/tribes in Gentianaceae (Struwe et al. 2002). This subtribe corresponds to tribe Potalieae-Loganiaceae in the circumscription used by Leeuwenberg and Leenhouts (1980). Anthocleista and Potalia are well supported as a monophyletic group (Struwe and Albert 1997; Struwe et al. 2002) and are similar morphologically, even though the genera have been kept separate for over 175 years. Brown (1818) distinguished the African/Malagasy Anthocleista from Aublet’s (1775) American Potalia (amara) based on woody versus herbaceous habit and 4- rather than 2-celled ovaries. The first trait is simply erroneous, and the second is disputable (see discussion under Reproductive Morphology). Rightly cautious in accepting the placentation difference, Bentham (1857: 75) stated that ‘‘had the genus Anthocleista not been already established, I should certainly have considered it as a second species of Potalia.’’ Another difference between Anthocleista and Potalia that has been raised is the ‘‘large, sterile, globular structure between the style and the ovary [of the latter genus]. . . .’’ (Bruce 1955: 45; Fig. 3g). Leeuwenberg and Leenhouts (1980) used the ventricose style of Potalia as a key characteristic, and we agree that this difference exists in all but three species, but it is not always obvious. Additionally, Punt (1978) noted that pollen of Potalia amara differs from six Anthocleista species examined in having distinctly thicker and more protruding pores and costae. This was confirmed in a later study of pollen from six Potalia species as ‘‘rims beset with numerous rounded to elongated protruberances’’ (Nilsson 2002: 460; note that the figures of pollen of Potalia amara [fig. 4.38] and Anthocleista nobilis G. Don [fig. 4.31] are erroneously identical). The morphological investigations of Potalia presented here show that there are additional good morphological characters that distinguish Potalia from Anthocleista, e.g., pedicel color (yellow vs. green), interpetio- [Volume 29 lar sheaths (present vs. absent), corolla lobe number (8–10 vs. 9–16), fruit size (less than 2 cm long vs. larger), and the presence of a sterile apex and ring on the fruit (in all but three species; present vs. absent). Anthocleista was found to be paraphyletic to Potalia in an earlier, preliminary, study based on non-molecular characters (Struwe and Albert 1997). In that work, we discussed the possibility for combining these two genera (the name Potalia would then have priority). However, since molecular data now support a sister group relationship between two monophyletic Anthocleista and Potalia genera (Frasier et al. 2002; Struwe et al. 2002), in addition to the above-mentioned morphological differences, there is now strong evidence for keeping Anthocleista and Potalia as separate genera based on both phylogeny and practicality. Pollination and Dispersal. Little is known about the pollination and fruit dispersal of Potalia. Flowers of Potalia elegans in Venezuela have been reported to be visited by hummingbirds (teste Davidse & Miller 26744). Potalia fruits and seeds are also likely dispersed by birds, but no observations have been reported. Ethnobotany and Phytochemistry. Gentians are well-known for their pharmacological properties worldwide and Potalia is no exception. Potalia species are widely used by indigenous peoples in South America as herbal remedies and have an abundance of vernacular names (Table 2; Schultes and Raffauf 1990; Duke and Vasquez 1994). They are considered particularly powerful plants (Schultes and Raffauf 1990). Common uses are against snake bites (thereby the indigenous name pao-de-cobra—‘‘the snake branch’’ or ‘‘the snake stick’’), inflammations, and fevers. The use against snake bites, as well as other poisonous animals such as sting rays and ants (Duke and Vasquez 1994; Schultes and Raffauf 1990), has been reported for Potalia coronata from Peru (teste Ruı́z & Murphy 200), P. elegans from Colombia (teste Schultes 22718), and for P. resinifera from Brazil, Colombia, Ecuador, and Peru (Schultes and Raffauf 1990; Friedman et al. 1993; e.g., teste Archer 7651, Schultes & Cabrera 12621, and Schunke V. 10675). The effect is reportedly that it ‘‘calms the body and eliminates the pain’’ (teste Plowman et al. 6803). Fresh leaves of Potalia resinifera are applied to the area of the bite or an unheated infusion of the leaves or a leaf, bark, or root decoction are drunk (e.g., teste Archer 7651, Daly et al. 4359, Hahn & Tredwell 56, and Schunke 2845, 3610, and 10712). The vernacular names of Potalia elegans (e.g., palo de temblador, ‘‘the trembling branch’’) also suggest a neurostimulatory effect. In Peru the root and resin of Potalia resinifera are used to kill parasites (teste Salick 7371). In their review of Amazonian ethnobotany, Duke and Vasquez (1994) reported that Potalia amara (in the earlier wider species circumscription) had widespread indigenous uses. Potalia amara in the current strict 2004] STRUWE & ALBERT: POTALIA (GENTIANACEAE) sense does not occur in this area and the listed uses should now be associated with one or several of the Amazonian Potalia species, i.e., P. coronata, P. elegans, P. maguireorum, and P. resinifera. Duke and Vasquez (1994) list uses against syphilis, urethritis, and other veneral diseases, as well as ophtalmia. A strong tea of Potalia is used to treat poisoning from cassava (Manihot esculenta) and other plants (Plotkin et al. 1991), and the plant is used as an emetic in food poisoning and as a laxative (Duke and Vasquez 1994; Schultes and Raffauf 1990). It is also used against dental cavities and to soothe body aches and pains. Aublet (1775) remarked that leaves and young twigs of Potalia amara were used as a treatment for venereal diseases, and in larger doses, as an emetic. An infusion of the older bark of Potalia elegans has been reported to be used as a laxative (teste Williams 14352), and use against ulcers has also been reported (teste Carnevali et al. 1849). Potalia resinifera and P. coronata have been used to treat eye inflammations (Martius 1826–27), for example among the Barasana Indians in Vaupés, Colombia (teste Davis 133). In the province of Pasco (Peru) the root and resin of Potalia resinifera is used to treat rheumatism and anemia (teste Salick 7371), crying babies are bathed in an infusion from the plant (teste Salick 7240), and the herb is also rubbed on back aches (teste Salick 7095). In Napo, Ecuador, skin problems are treated with boiled and mashed leaves of Potalia resinifera that are applied to the affected areas (teste Jaramillo 9053). Potalia resinifera is considered a shamanistic and spiritual plant with medical properties in Cordoba, Colombia (teste Bernal et al. 1153). Only two reports of the plants being used for food have been found—apparently the Chácobo Indians in Benı́ (Bolivia) eat the fruits (teste Boom 4498 and Boom 4639). The species epithet ‘‘amara’’ rightly describes the bitterness of all plant parts, which is most likely due to accumulated seco-iridoids (Bisset 1980a, b; Jensen 1992; Jensen and Schripsema 2002). An extract of a bulk collection of Potalia amara yielded the greatest pergram tissue weight of the gentian-specific seco-iridoid swertiamarin ever encountered (S. R. Jensen, pers. comm.; Jensen and Shripsema 2002). These iridoids and other pharmacologically active compounds such as xanthones are also known from African/Malagasy species of Anthocleista and Asian/Australian Fagraea, which also have many native pharmacological uses (Bisset 1980a, b; Boiteau and Allorge-Boiteau 1993; Motley, in press). Schultes and Raffauf (1990) reported that Potalia also contains squalene, saturated fat, and methyl esters. Detailed phytochemical screening of Potalia remains to be done, and as Schultes and Raffauf (1990) wrote, ‘‘in view of the extraordinarily use of this plant for medicinal purposes, it is incredible that practically no chemical examination has been made.’’ 697 ACKNOWLEDGEMENTS. We thank Gerardo Aymard, Paul Berry, Søren Rosendal Jensen, Carol Gracie, Jason Grant, Andrew Henderson, Scott Mori, and Gustavo Romero for material and helpful information, the late Rupert Barneby for help with Latin diagnoses, Ken Cameron and Mats Gustafsson for comments on earlier versions of the manuscript, Nora Martinez for translation of some ethnobotanical information, Douglas Daly for help with location of collection localities in Brazil, BM and Chris Humphries for a photo of the holotype of Potalia amara, Cynthia Frasier for excellent help with the distribution maps and other practical things, two anonymous reviewers for helpful comments on a previous version of this manuscript and the following persons for help with translation and geographic information: Maria Fernanda Calı́o, Rocio Cortes, Guilhem Mansion, and Jim Smith. The herbaria and their curators that provided loans and additional information are also gratefully acknowledged (COL, ECON, F, GB, GH, IAN, INPA, K, MG, MO, NY, QCA, QCNE, S, SP, SPF, UPS, US, VEN). The illustrations were provided by Bobbi Angell. This research was funded by the The Cullman Program for Molecular Systemtics Studies, The New York Botanical Garden (Bronx, USA), Rutgers UniversityCook College (New Brunswick, NJ, USA), and the UddenbergNordingska Foundation (Kalmar, Sweden). VAA thanks the University of Oslo, Norway, for support during the final stages of manuscript preparation. LITERATURE CITED AUBLET, M. F. 1775. Histoire des plantes de la Guiane Françoise, vol. 1. London & Paris: P.-F. Didot. BAILLON, H. 1888. Solanacées. Histoire des plantes, vol. 10. Paris: Librairie Hachette & Co. BARTLING, F. G. 1830. Ordines naturales plantarum corumque characteres et affinitates. Göttingen: Dieterich. BENTHAM, G. 1857. Notes on Loganiaceae. 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Contributions toward a flora of Panama. VI. Collections chiefly by H. von Wedel in Bocas del Toro: Loganiaceae. Annals of the Missouri Botanical Garden 29: 364. APPENDIX 1 Index to collectors and material. Data are presented in the following sequence: Collection, Species, Country, Herbarium. Acevedo-Rdgz & Grimes 4929, P. amara, French Guiana, K, NY, US; Allorge 380, P. amara, French Guiana, NY; Amaral et al. 218, P. resinifera, Brazil, NY; Archer 7651, P. resinifera, Brazil, IAN; Asplund 14732, P. resinifera, Peru, G, S, US; Asplund 14616, P. resinifera, Peru, S; Asplund 19797, P. resinifera, Peru, NY; Assumpcão & Duarte s.n., P. resinifera, Brazil, US; Aublet / von Rohr s.n., P. amara (type), French Guiana, BM ; Ayala 2071, P. resinifera, Peru, MO; Ayala et al. 2741, P. resinifera, Peru, MO; Ayala et al. 2867, P. coronata, Peru, K, MO, 699 NY; Ayala et al. 3869, P. coronata, Peru, MO, NY; Ayala & Criollo 3993, P. resinifera, Peru, NY; Aymard 9265, P. maguireorum, Venezuela, NY. Bahia 162, P. resinifera, Brazil, MG, NY; Baldwin 3235, P. resinifera, Venezuela, US; Balick & Allon 1017, P. resinifera, Peru, ECON; Bastos et al. 2329, P. resinifera, Brazil, MG; Berg et al. P18494, P. resinifera, Brazil, NY; Berg & Steward P19917, P. resinifera, Brazil, NY; Berlin 1935, P. resinifera, Peru, GH; Bernal et al. 1153, P. resinifera, Colombia, COL; Bernal et al. 1245, P. elegans, Colombia, COL; Berry et al. 5688, P. elegans, Venezuela, NY; Black 25965, P. amara, Brazil, IAN; Black 49–8232, P. amara, Brazil, IAN; Black 51–13902, P. resinifera, Brazil, IAN; Black 51–13953, P. resinifera, Brazil, IAN; Black 54– 17664, P. amara, French Guiana, IAN; Boom 4498, P. resinifera, Bolivia, GH, NY, US; Boom 4639, P. resinifera, Bolivia, NY; Boom 4998, P. resinifera, Bolivia, US; Boom et al. 8541, P. resinifera, Brazil, NY. Callejas et al. 5288, P. aff. resinifera, Colombia, NY; Carnevali et al. 1849, P. elegans, Venezuela, VEN; Cavalcante 746, P. elegans, Brazil, MG; Cavalcante & Silva 1603, P. resinifera, Brazil, MG; Cerón 653, P. resinifera, Ecuador, MO, QCNE; Cerón 883, P. resinifera, Ecuador, MO, QCNE; Cerón et al. 2061, P. resinifera, Ecuador, NY, QCNE; Cerón & Cerón 4496, P. resinifera, Ecuador, MO, QCNE; Cerón & Gallo 4927, P. resinifera, Ecuador, MO, QCNE; Cerón & Gallo 5024, P. resinifera, Ecuador, MO, QCNE; Cerón & Gallo 5143, P. resinifera, Ecuador, MO, QCNE; Cerón et al. 5567, P. resinifera, Ecuador, MO, QCNE; Cerón 6359, P. resinifera, Ecuador, MO, QCNE; Cid Ferreira 9343, P. resinifera, Brazil, NY; Cid Ferreira et al. 8477, P. resinifera, Brazil, NY; Cid Ferreira et al. 9908, P. coronata (type), Brazil, INPA, NY; Cid & Nelson 2752, P. resinifera, Brazil, NY; Cid & Lima 3533, P. resinifera, Brazil, MG; Clark 7163, P. elegans, Venezuela, NY; Clark & Maquirino 7840, P. elegans, Venezuela, MO; Colella et al. 2021, P. elegans, Venezuela, NY; Cowan 38349, P. amara, Brazil, NY; Cowan 38395, P. amara, Brazil, NY; Cowan 38438, P. amara, Brazil, IAN, NY; Cowan 38651, P. amara, Brazil, NY; Croat 17690, P. resinifera, Peru, MO; Croat 18090, P. resinifera, Peru, MO; Croat 18337, P. resinifera, Peru, MO; Croat 58678, P. resinifera, Ecuador, MO, QCA; Cuatrecasas 6852, P. elegans, Colombia, US; Cuatrecasas 9202, P. resinifera, Colombia, US. da Costa Neto et al. 200, P. resinifera, Brazil, MG; Daly et al. D216, P. resinifera, Brazil, MG; Daly et al. 1542, P. resinifera, Brazil, MG, NY; Daly et al. 4359, P. resinifera, Brazil, NY; Daly et al. 8388, P. resinifera, Brazil, NY; Daly et al. 8462, P. resinifera, Brazil, NY; Daly et al. 8652, P. resinifera, Brazil, NY; Daly et al. 8912, P. resinifera, Brazil, NY; Davidse & Miller 26717, P. elegans, Venezuela, MO, NY; Davidse & Miller 26744, P. elegans, Venezuela, MO; Davis 133, P. coronata, Colombia, GH, K; de Granville et al. 186, P. amara, French Guiana, NY; de Granville et al. 7965, P. amara, French Guiana, MO; de Lemos Fróes 20531, P. resinifera, Brazil, IAN, NY; de Lemos Fróes 21221, P. elegans, Brazil, G, IAN, US; de Lemos Fróes 25541, P. resinifera, Brazil, IAN; de Lemos Fróes 25718, P. amara, Brazil, IAN; de Lemos Fróes 26185, P. resinifera, Brazil, IAN; de Lemos Fróes 28087, P. resinifera, Brazil, IAN; de Lemos Fróes & Addison 28653, P. elegans, Brazil, IAN; de Lemos Fróes 28849, P. elegans, Brazil, IAN; de Oliveira 1816, P. resinifera, Brazil, IAN; Delascio & Guánchez 11025, P. resinifera, Venezuela, VEN; Dias et al. 253, P. resinifera, Brazil, MG; Dodge et al. 6076, P. turbinata, Costa Rica, GH, MO; Dorr & Valdespino 6314, P. resinifera, Ecuador, NY, QCE; Ducke 364, P. resinifera, Brazil, S; Ducke 1766, P. resinifera, Brazil, A, IAN, K, MG, NY, US. Espina & Mosquera 2230, P. chocoensis, Colombia, COL. Fariñas et al. 601, P. elegans, Venezuela, NY, VEN; Fernández 2225, P. coronata, Colombia, COL; Fernandez 7797, P. elegans, Venezuela, MO; Forero & Jaramillo 2698, P. crassa (type), Colombia, COL, MO, NY; Forero et al. 6365, P. resinifera, Brazil, K, NY, S, US; Forero et al. 8886, P. chocoensis (type), Colombia, COL, ; Foster et al. 605, P. resinifera, Bolivia, G. 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