Three new species in the neotropical genus Clidemia - Springer Link

Three new species in the neotropical genus Clidemia (Melastomataceae: Miconieae) RICARDO KRIEBEL1,2

AND

FRANK ALMEDA3

1

The New York Botanical Garden, 200th St. & Southern Blvd., Bronx, NY 10458, USA; e-mail: [email protected] 2 Instituto Nacional de Biodiversidad, Apdo. 22-3100, Santo Domingo, Heredia, Costa Rica 3 Department of Botany, California Academy of Sciences, 55 Concourse Drive, Golden Gate Park, San Francisco, CA 94118, USA; e-mail: [email protected]

Abstract. We here propose and describe three new species of Clidemia: C. aguilarii from Costa Rica and Panama; C. aurantiaca from Costa Rica, Panama, and Ecuador; and C. subpeltata, a Costa Rican endemic. Phenological notes, distribution maps, line drawings, color images of the live plants in the wild, and discussions comparing the new species to their presumed closest relatives are provided. Key Words: Clidemia, Melastomataceae, Costa Rica, Panama, Ecuador. Resumen. Aquí proponemos y describimos tres especies nuevas de Clidemia: C. aguilarii de Costa Rica y Panamá; C. aurantiaca de Costa Rica, Panamá, y Ecuador; y C. subpeltata, endémica de Costa Rica. Se incluyen notas sobre su fenología, mapas de su distribución, ilustraciones, fotografías a color de plantas silvestres, así como discusiones comparando las nuevas especies con sus parientes aparentemente más cercanos.

Several new species of Clidemia D. Don have been described from Costa Rica and Panama in recent years. Specimens of many of these new taxa are mainly from remote undercollected areas (Almeda, 1984, 1989, 2003, 2004). Interestingly, more new species continue to appear from readily accessible localities in Costa Rica like Tapantí and Braulio Carrillo National Parks, Cabo Blanco Absolute Reserve, and El Rodeo Protected Zone. Several specimens of the three species proposed in this paper were gathered by collectors and tourists at the entrance or exit of the commonly visited trails in these protected areas. The above pattern suggests that although our knowledge of the Mesoamerican Melastomataceae has grown immensely in recent years (Almeda, in press), more novelties will continue to be discovered in this, one of the most diverse families of flowering plants.

Clidemia, a berry-fruited genus of suffrutescent herbs or shrubs (rarely scandent or epiphytic) with over 180 species, ranges throughout tropical America (Almeda, 2004). Of the total species described to date, close to 30% (56 spp.) are found in Costa Rica and Panama combined, and 11% (20 spp.) are restricted to both countries. In this paper we propose the following new species: Clidemia subpeltata, so far endemic to Costa Rica; Clidemia aguilarii, endemic to the pacific coast of Costa Rica to west Panama; and Clidemia aurantiaca, known from Costa Rica, Panama, and Ecuador. These new species are only tentatively placed in Clidemia, since the genus is likely not monophyletic (Michelangeli et al., 2004), and generic delimitations have yet to be sorted out. The new taxa have axillary flowers as is typical of Clidemia and Ossaea DC., but differ from Ossaea in that their petal apex is

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rounded and not acuminate. Nevertheless, petal apex has been shown not to be phylogenetically informative in the closely related Leandra Raddi, where it was previously considered to be a diagnostic character (Martin et al., 2008). Clidemia aurantiaca in particular is very similar to the Central American species of Ossaea sharing staminal connectives appendages that are gland edged dorso-basally as well as costate fruit. Ongoing phylogenetic studies in the Miconieae will help elucidate the appropriate placement of these taxa. Clidemia aguilarii Kriebel & Almeda, sp. nov. Type: Costa Rica. Putarenas: Cantón de Golfito, along gravel road to microwave tower above Golfito, 0.7 m above main road, 08°39′N, 83°11′W, 15 Sep 1987, T. Croat 67598 (holotype: CAS; isotype: MO, n.v.). (Figs. 1, 2C, D) Ramuli teretes, sicut petioli, foliorum venae primariae et secundariae abaxiales et inflorescentiae axes, dense pilis asperis induti. Lamina 4–23×3–11.5 cm elliptica vel ovata apice acuta vel acuminata basi longe decurrens, 5−7(−9)-nervata. Inflorescentia primum terminalis demum lateralis, bracteolis 0.5 mm longis; flores 4-meri, calycis lobis interioribus 0.25 mm longis undulatis, dentibus exterioribus triangularibus 0.1–0.25 mm longis. Stamina isomorpha glabra, antherarum thecis 1.5–2× 0.5 mm oblongis in porum apicalem ventraliter inclinatum contractis; connectivum vix (0.1 mm) prolongatum, dente dorsi-basali 0.25 longo, glandulis marginato. Ovarium 4-loculare et omnino inferius apice dense glandulis inconspicuis indutum. Bacca aurantiaca.

Subshrub or shrub 0.4−2(−3) m, the terete upper internodes and branchlets, petioles, primary and secondary abaxial leaf veins, and inflorescence axes covered with a dense indument of stipitate or subsessile clavate asperous hairs. Leaves of a pair somewhat unequal in size, 4–23×3–11.5 cm, elliptic to ovate, 5−7(−9)-plinerved, abruptly narrowed at the base and narrowly decurrent along the petiole, apex acute to acuminate, the margins ciliolate-denticulate, glabrous adaxially except for subsessile clavate asperous hairs on the primary veins towards the base, sparsely and deciduously resinous-glandular on the higher order veins and blade surface abaxially. Inflorescence a modified pseudolateral dichasium 2–5 cm long, mostly divaricately and laxly branched from the base; flowers 4merous on pedicels 0.25–0.5 mm long; the

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bracteoles subulate, persistent, mostly 0.5 mm long. Hypanthium bluntly 8-costate, deciduously resinous-glandular in flower. Calyx vaguely undulate, ca. 0.25 mm long, the exterior teeth bluntly triangular, 0.1– 0.25 mm long. Petals densely papillose on both surfaces with a projecting infra-apical, abaxial tooth, obovate-oblong, 0.75–1.25× 0.75–1 mm. Filaments 1–1.5 mm long, anthers 1.5–2×0.5 mm long, linear-oblong, yellow, tapering to a ventrally inclined apical pore; connective barely prolonged dorsobasally into a gland-edged spur ca. 0.25 mm long. Ovary 4-locular, completely inferior, apex somewhat depressed, minutely and deciduously resinous-glandular puberulent. Style ca. 4–5 mm long, straight, glabrous; stigma punctiform. Berry ± compressed, initially white and turning bright orange when mature, 4–5×3 mm when dry. Seeds 0.5 mm long, the testa smooth, the raphe extending for the entire length of the seed. Distribution.—Clidemia aguilarii is known from the Pacific slope of Costa Rica south to western Panama, at 0–600 m, where it grows in the understory of primary or secondary rain forest, and along road banks and rocky streams (Fig. 3). In Costa Rica, it is widely distributed, ranging from the Nicoya Peninsula on the northern Pacific coast, through the central Pacific lowlands in El Rodeo Protected Ciudad Colón Zone, south to the Osa Peninsula; it is protected in the Cabo Blanco Absolute Reserve, Curú Wildlife Refuge, El Rodeo Protected Zone, and Carara, Manuel Antonio and Corcovado National Parks. In Panama it is restricted to Veraguas Province (including Coiba Island) where it is known from a few specimens. Phenology.—Plants with flowers and fruits have been collected from June through December. Etymology.—The specific epithet is dedicated to Costa Rican botanist Reinaldo Aguilar who has studied the flora of Costa Rica, concentrating on the Osa Peninsula, where he now resides, for over 15 years. Additional specimens examined. COSTA RICA. Guanacaste: Nandayure, Pilas de Bejuco, Finca de Abel Rodríguez, 09°51′38′′N, 85°21′33′′W, 13 Jul 1994, A. Estrada & A. Rodríguez 10 (CR, INB, MO); Cantón de Hojancha, Finca Dyalá, Puerto Carrillo, bosque situado en el nacimiento de la Quebrada Zapotal, 09°52′10′′N,

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FIG. 1. Clidemia aguilarii. A. Habit. B. Detail of the branchlet indument. C. Flower at anthesis. D. Stamen. E. Infrutescence detail. F. Dried fruit. G. Seeds. (Drawn from the holotype.)

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FIG. 2. A. Flowering branch of C. aurantiaca. B. Fruiting branch of C. aurantiaca (Kriebel 4607; CR, INB). C. Flower of C. aguilarii. D. Infructescences of C. aguilarii (Kriebel et. al 5072; INB). E. Leaf base of C. subpeltata. F. Flower of C. subpeltata (Kriebel & Solano 3643; CR, CAS, INB, MO).

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FIG. 3. Distributions of the closely related species Clidemia aguilarii and C. quinquenervia.

85°28′15′′W, 26 Dec 1988, L. Flores & C. Herrera 9 (INB); Nandayure, Bejuco, Pilas de Bejuco, 09°50′52′′N, 85°21′27′′W, 10 Mar 1999, A. Rodríguez & P. Hurtado 4599 (CR, INB, MO). Puntarenas: Reserva Forestal Golfo Dulce, Los Mogos, Golfito, 08°45′30′′N, 83°22′ 30′′W, 5 Sept 1991, R. Aguilar 335 (CAS, CR, INB, MO); P. N. Corcovado, Estación La Leona, junto a Río Madrigal, 08°27′00′′N, 83°29′30′′W, 19 Jan 1993, R. Aguilar 1603 (CR, INB, MO); Parque Nacional Corcovado, Península de Osa, Estación Sirena, Sendero a Río Claro, 14 Oct 1993, R. Aguilar 2529 (CAS, INB, MO); P.N. Corcovado, Península de Osa, Estación Sirena, Alrededores de Sendero Ollas, 08°28′50′′N, 83°35′ 30′′W, 6 Feb 1994, R. Aguilar 3076 (CR, INB, MO); P. N. Corcovado, Península de Osa, Estación Sirena, Sendero Pavo, 08°28′N, 83°35′W, 23 July 1994, R. Aguilar 3507 (CAS, CR, INB, MO); P.N. Corcovado, sendero a Río Claro, 08°28′50′′N, 83°35′30′′W, 12 Sept 1998, R. Aguilar 5542 (CR, INB); disturbed sites above Golfito along the road to the television tower, 16 Jul 1977, F. Almeda et al. 3092, 3094 (CAS, CR); R. N. A. Cabo Blanco, Península de Nicoya, Estación San Miguel, límite de la reserva, 09°35′42′′N, 85°07′13′′W, 10 Feb 1997, F. Alvarado 115 (CR, INB, MO); Cantón de Nicoya, Reserva Absoluta Cabo Blanco, Estación Cabo Blanco, 09°35′N, 85°06′W, 4 Nov 1991, U. Chavarría 305 (CAS, INB, MO); R.N.A. Cabo Blanco, Sendero Central, 09°35′00′′N, 85°07′00′′W, 17 Dec 1993, A. Fernández et al. 1315 (CR, INB, MO); P.N. Corcovado, Estación Sirena, 08°28′N, 83°35′W, 26 Oct 1990, G. Fonseca 9 (CR, INB, MO); P. N. Corcovado, alrededores de la Estación Sirena, 08°28′50′′N, 83°35′30′′W, 14 Apr 1995, B. Gamboa & A. Picado 139 (CR, INB); Cantón de Quepos, Punta Catedral, ca. 7 km SE of Quepos, 09° 22.5′N, 84°09′W, 20 Aug 1985, M. Grayum & P. Sleeper 5896 (CAS, CR, MO); R. N. A. Cabo Blanco, Península de Nicoya, Estación San Miguel, ca. 2 km S. de Malpaís,

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09°35′40′′N, 85°07′50′′W, 18 Jan 1996, B. Hammel et al. 20091 (CR, INB, MO); Parque Nacional Manuel Antonio, 09°23′N, 84°09′W, 3 Aug 1990, P. Harmon 136 (CAS, CR, INB, MO); P.N. Corcovado, Estación Sirena, 08°27–30′N, 83°33–38′W, 11 Feb 1988, C. Kernan 109 (INB); P.N. Corcovado, Península de Osa, Ollas trail, 08°27–30′N, 83°33–38′W, 26 Jul 1988, C. Kernan & P. Phillips 717 (CAS, CR, INB); P.N. Corcovado, Península de Osa, ridges above Río Claro, 08°28′N, 83°35′W, 24 Nov 1981, S. Knapp & J. Mallet 2209 (CAS); Cantón de Golfito, Jiménez, bosques alrededor del Albergue Río Nuevo, 08°29′50′′N, 83°23′ 20′′W, 1 Jan 2005, R. Kriebel et al. 5072 (INB); Garabito, Cuenca del Tulín, Villa Lapas, 09°45′30′′N, 84°36′50′′W, 29 Dec 2002, R. Kriebel 2164 (CR, INB); P.N. Corcovado, Península de Osa, Estación Sirena, 08° 28′N, 83°35′W, 15 Jun 1990, G. Maass 37 (CAS, CR, INB, MO); P.N. Corcovado, Península de Osa, Estación Sirena, 08°28′N, 83°36′W, 1 Oct 1990, J.C. Saborío 78 (CAS, CR, INB, MO); Nicoya Peninsula, canyons and ridges towards Punta Georgia, Punta Blanca trail, 23 Aug 1995, A. Sanders et al. 17706 (CAS, USJ); Aguirre, Manuel Antonio, Fincas Naturales en frente del Hotel Sí Como No, 09°24′50′′N, 84°09′26′′W, 29 Nov 2003, D. Solano 647 (CR, INB, MO); Cantón de Garabito, Reserva Biológica Carara, Sendero Quebrada BonitaBijagual, 09°46′N, 84°16′W, 8 Dic 1989, R. Zuñiga & Q. Jiménez 18 (CAS, CR, INB). San Jose: Puriscal, 09°51′ 02′′N, 84°19′07′′W, 22 Dec 2002, J. Bustamante 351 (CR, INB, MO); Cantón de Turrubares, Valle de Tárcoles, S.E. de San Luis, entre Quebrada Zorrillal y Cerro San Luis, 09°50′20′′N, 84°28′40′′W, 20 Dec 1995, B. Hammel & J. González 20033 (CR, INB, MO); Puriscal, Parque Nacional La Cangreja, 09°41′17′′N, 84° 22′09′′W, 21 Dic 2002, R. Kriebel & J. Larraguivel 2029 (CR, INB); Cantón de Puriscal, Z. P. La Cangreja, Cerros de Puriscal, Falda W Cerro Cangreja, Santa Gertrudis, Cabeceras Río Negro, 09°43′25′′N, 84°22′41′′W, 27 Jun 1996, J. F. Morales et al. 5434 (CR, INB, MO); Cantón de Mora, Ciudad Colón, Zona Protectora El Rodeo, 09° 54′N, 84°16′W, 4 Aug 1992, G. Varela 241 (CAS, INB, MO). PANAMA: Veraguas: Dto. De Montijo, Isla de Coiba, campamento de La Salina, subiendo hacia Río Negro, 21 Oct 1995, M. Ballesteros et al. 1345 (CAS, PMA); Mountains of southern Azureo Peninsula Los Santos and Veraguas Provinces, near proposed road from El Cortezo (Prov. Los Santos), to Arenas, (Prov. Veraguas), 0–5 km SW of El Cortezo, 29 Oct 1978, B. Hammel 5461 (CAS, MO).

Specimens of Clidemia aguilarii have been confused with and annotated as Ossaea quinquenervia (Mill.) Cogn. or Clidemia quinquenervia (Mill.) Almeda in the past. Both species are part of a closely related group of taxa that also includes C. coronata Gleason. These three taxa share a similar indument of asperous, clavate hairs, densely papillose-furfuraceous petals, anther connectives that are glandularappendaged dorso basally, a completely inferior ovary, and a smooth seed coat. Almeda (2004)

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commented recently on the differences between C. coronata and C. quinquenervia involving vegetative characters such as leaf venation (5–7 nerved in C. coronata vs. 5–9 plinerved in C. quinquenervia), leaf margins (ciliate and ± entire vs. ciliolate-denticulate) and blade base (not decurrent vs. conspicuously decurrent on the petiole), as well as differences in the shape of the anther connective appendages (rounded and entire vs. spur-like and gland-edged). To the above differences we can add the presence of setulose exterior calyx teeth in C. quinquenervia that are lacking in C. coronata. In the recently published key to the Costa Rican species of Clidemia (Almeda et al., 2007), C. aguilarii would key out to couplet 23 because of the 4-merous flowers and 4locular ovary, but differing from C. ombrophila in the lack of leaf domatia, an abruptly acuminate leaf apex, and basally fused bracteoles that form a collar. Then it would key out to couplet 25 which includes both species discussed above, C. coronata and C. quinquenervia which are distingushed in the key by different leaf characteristics. Clidemia aguilarii is very similar to C. quinquenervia in that they both share the strongly decurrent leaf base and gland-edged connective appendage. Clidemia aguilarii differs in lacking setulose calyx teeth, and having 4merous flowers and a 4-locular ovary (versus 5-merous and 5-locular C. quinquenervia). The most striking difference are the bright orange mature berries of C. aguilarii (versus purple-black berries in C. quinquenervia). Regarding their distribution, the new species is restricted to the Pacific slope of Costa Rica and western Panama; C. coronata is endemic to Costa Rica where it is very rare and has been collected on both slopes (e.g., Tarrazú on the Pacific slope, and Turrialba on the Caribbean slope); C. quinquenervia is the most widely distributed of the three taxa, ranging from Honduras south through Central America to Colombia, Venezuela, and Ecuador (Almeda, 2004). In Costa Rica and Panama, the latter is known only from the Caribbean slope (Fig. 3). Clidemia aurantiaca Almeda & Kriebel, sp. nov. Type: Costa Rica. Limón: Cantón de Pococí, P.N. Braulio Carrillo, Estación Quebrada González, sendero Las Palmas, 10°09′50′′N, 83°56′24′′W, 400–500 m, 6

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Jun 2003, R. Kriebel 3342 (holotype: INB; isotypes: CAS, CR, INB, MO, PMA, NY, US). (Figs. 2A, B, 4) Ramuli teretes, sicut petioli, foliorum venae primariae et secundariae abaxiales, inflorescentiae axes hypanthiaque dense pilis asperis induti. Lamina 4.9–17.5×2.4–6.3 cm elliptica apice gradatim acuminata vel caudato-acuminata basi acuta vel attenuata, 3–5-nervata. Inflorescentia in foliorum superiorum axillis vel lateralis in ramulis infra folia plerumque oriunda 1–1.5 cm longa multiflora, bracteolis 0.3–0.75×0.3–0.5 mm; flores 4-meri, calycis lobis interioribus 0.75–1.25×0.5–0.75 mm, triangularibus, dentibus exterioribus 1–1.25 mm longis, subulatis, late patentibus. Stamina isomorpha glabra, antherarum thecis 1.25×0.5 mm oblongis poro dorsaliter inclinato; connectivum vix (0.1 mm) prolongatum, dente dorsi-basali 0.5 mm longo, glandulis 0.1 mm longis marginato. Ovarium 4-loculare, 2/3 inferius, apice moderate glandulis inconspicuis indutum. Bacca aurantiaca.

Shrub or small tree, 1.5–3.5 m tall; uppermost terete internodes, vegetative buds, petioles, elevated primary and secondary veins on abaxial foliar surface, bracts and bracteoles, pedicels, hypanthia, calyx lobes and calyx teeth moderately to copiously covered with asperous hairs. Leaves of a pair ± equal to somewhat unequal in length; petioles 0.3–1.3 cm; blade 4.9–17.5×2.4– 6.3 cm, membranaceous, elliptic, apex acuminate to long-acuminate, base acute to attenuate, margin essentially entire, 3–5nerved, adaxially with sparsely distributed asperous hairs on young leaves, glabrous with age, tertiary veins copiously beset with a white granulose indument abaxially. Inflorescence a cluster of pseudofasciculate cymes in the axils of the upper leaves or lateral, usually arising on branchlets below the leaves and mainly on defoliated nodes, ca. 1–1.5 cm long; bracts and bracteoles 0.3–0.75 mm long and 0.3–0.5 mm wide at the base, triangular. Flowers 4-merous, sessile. Hypanthia (at anthesis) subcylindric to narrowly campanulate, 2–2.5 mm long. Calyx tube obsolete; calyx lobes 0.75–1.25 mm long and 0.5– 0.75 mm wide at the base, triangular, typically reflexed, calyx teeth 1–1.25 mm long, subulate, widely spreading. Petals 2–2.75×1– 1.5 mm, glabrous, translucent-white, ovateoblong, conspicuously reflexed at anthesis and with a minute and inconspicuous subapical dorsal tooth. Stamens 8, isomorphic; filaments ca. 1.5 mm long, glabrous; anthers

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FIG. 4. Clidemia aurantiaca. A. Habit. B–C. Detail of the abaxial foliar veinlet indument. D. Flower at anthesis. E. Stamens. F. Defoliated node with fasciculate infrutescence. G. Mature fruit. H. Seeds. (Drawn from the holotype.)

ca. 1.25×0.5, white, linear-oblong, widest apically, emarginate at the apex with a dorsally inclined pore; connective prolonged dorso-basally into a deflexed, ± triangular, gland-edged appendage ca. 0.5 mm long and also somewhat prolonged and gland-edged

but unappendaged ventro-basally. Ovary 4locular, 2/3 inferior, the apex consisting of a shallow bowl-like depression with a slightly raised perimeter, moderately but inconspicuously glandular-puberulent. Style ca. 5 mm long, glabrous; stigma punctiform. Berry ca.

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4×4.5 mm, globose, bright orange at maturity, prominently 8-costate. Seeds 0.4–0.6 mm long, yellow-brown, ± triangular in outline, rugulate, somewhat angulate, with a foot-like projection at the narrow end. Distribution.—Clidemia aurantiaca is known from undisturbed rain forest understory on the Caribbean slope of the Tilarán, Central and Talamanca mountain ranges of Costa Rica, the provinces of Bocas del Toro and Veraguas in Panama, and in the province of Cotopaxi, Ecuador, north west of El Corazón at 250–900 m (Fig. 5). Phenology.— This species has been collected in flower from June through August and in fruit from July through February. Etymology.— The specific epithet refers to the orange berries of the new species. Additional specimens examined. COSTA RICA. Alajuela: Cantón de Guatuso, P.N. Volcán Tenorio, cuenca del Río Frío, Estación El Pilón, sendero hacia catarata, 10°42′57′′N, 84°59′32′′W, 800 m, 13 Jun 2000, J. L. Chaves 500 (CR, INB, MO); Cantón de Guatuso, P. N. Volcán Tenorio, Estación El Pilón, sendero Misterioso, 10°42′16′′N 84°59′32′′W, 700–800 m, 17 May 2004, R. Kriebel et al. 4542 (CAS, CR, INB, MO, NY, US); Cantón de Guatuso, P.N. Volcán Tenorio, entrada La Carmela a salir a la estación, 10°43′29′′N 84°59′ 56′′W, 600 m, 19 May 2004, R. Kriebel et al. 4607 (CR, INB), 4608 (CR, INB, MO); Cantón de San Ramón, R.B. Alberto Manuel Brenes, curso de plantas, 10°13′13′′N, 84°35′52′′W, 800–900 m, 11 Aug 2002, R. Kriebel et al. 898 (CR, INB, MO); Cantón de San Ramón, R.B. Alberto Manuel Brenes, curso de plantas, 10°13′13′′N, 84°35′52′′W, 800–900 m, 11 Aug 2002, R. Kriebel et al. 924 (CAS, CR, INB, MO, NY, US); S of volcano along Rio Agua Caliente from base of observatory trail (ca. 10° 26′ 39′′ N 84° 42′ 62′′ S) to a point ca. 1 km upstream, 11 Apr 1991, F. O. Smith et al. 10872 (NY); Cantón de San Ramón, along road from San Ramón northward threw Balsa, ca. 13.8 km north of bridge over Quebrada Volio and ca. 4.6 km N of bridge over (apparently) Río Balsa, at small stream, 10°12′ N 84°31′ W, 29 Aug 1979, W.D. Stevens 13794 (CAS, MO). Limon: Cantón de Pococí, P. N. Braulio Carrillo, Estación Quebrada González, sendero Las Palmas, 10°09′20′′N, 83°56′30′′W, 400–500 m, 6 Jun 2003, R. Kriebel 1441 (CR, INB). San Jose: P. N. Braulio Carrillo, La Montura, 1100 m, 25–30 Jul, 1982, 1100 m, C. Todzia 2004 (NY). PANAMA. Bocas del Toro: Fortuna Dam area, Pipeline road off Chiriquí Grande road at Continental divide, 2–8 miles from divide point, 25 Jun 1986, W. G. D′Arcy 16400 (CAS). Veraguas: Valley of the Río Dos Bocas: 11–13 km beyond Agriculture School at Santa Fé, 25 Jul 1974, T.B. Croat 25764 (CAS, NY); on the road between Alto Piedra (above Santa Fé) and Calovebora, primary forest along road, 29 Aug 1974, T.B. Croat 27371 (CAS, NY); along road between Escuela Agrícola Alto Piedra and Calevebora, 15.6 km northwest of Santa

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Fé, primary forest along trail to Santa Fé, steep forested hill east of river, 31 Aug 1974, T. B. Croat 27667 (CAS); 11 km from Escuela Agrícola Alto Piedra, 20 Dic 1974, S. Mori & J. Kallunki 3811 (CAS); 8.8 km from Escuela Agrícola Alto Piedra, 25 Feb 1975, S. Mori & J. Kallunki 4852 (CAS). ECUADOR. Cotopaxi: Río Guapara: ±20 km NW El Corazón, monsoon forrest, 19–24 Jun 1967, B. Sparre 17268 (S, US).

Clidemia aurantiaca is characterized by its dense indument of asperous hairs, tertiary veins on the abaxial foliar suface copiously covered with a white granulose indument, fascicle-like short inflorescences mainly on defoliated nodes, 4-merous sessile flowers, connective prolonged dorso-basally into a deflexed glandedged appendage and also somewhat prolonged and gland-edged but unnappendaged ventrobasally, and costate orange berry. In the recently published key to the Costa Rican species of Clidemia (Almeda et al., 2007), C. aurantiaca would key out to couplet 31 which differentiates Clidemia densiflora (Standl.) Gleason and C. evanescens Almeda. These three species share the poorly or undeveloped inflorescence branching, 4-merous flowers, anthers with a dorsally inclined pore, connective that is prolonged dorso-basally into a gland-edged tooth, glandular-puberulent ovary apex, and orange,

FIG. 5. Distributions of Clidemia aurantiaca and C. subpeltata.

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costate fruit. Of these two species, C. aurantiaca is most similar to C. densiflora from which it differs in having hairs on stem apex, petioles and hypanthia usually less than 0.25 mm long, tertiary venation on the abaxial leaf surface copiously beset with a white granulose indument, petioles generally 1 cm long, and inflorescences mainly on defoliated nodes with the axis to 1.1 cm long (vs. hairs on stem apex, petioles and hypanthia usually 0.5–1 mm, tertiary venation on the abaxial leaf surface glabrous to brownishgranulose or brownish-resinose, petioles generally 0.5 cm long, and inflorescences mainly on foliated nodes with the axis absent). There is an additional perhaps new species in this complex from the province of Esmeraldas, Ecuador represented by the two following specimens: B. Sparre 18048 (S, US) and D. Rubio & C. Quelal 1931 (MO, QCNE). This entity is similar to C. aurantiaca because of its pseduo-fasciculate inflorescence, but differs in lacking the white-hyaline papillae on the tertiaty veins of the abaxial foliar surface and also has fimbriate scales on the torus, which are lacking in C. aurantiaca. We have postponed the description of the Esmeraldas species until additional specimens are collected. Clidemia subpeltata Kriebel & Almeda, sp. nov. Type: Costa Rica. Cartago: Cantón de Paraíso: Parque Nacional Tapantí, al lado de quebrada en la salida del Sendero Arboles Caídos, 9°48′18′′N, 83°57′12′′W, 1200 m, 8 Aug 2003, R. Kriebel & D. Solano 3643 (holotype: INB; isotypes: CAS, CR, MO, NY, US). (Figs. 2E, F, 6) Ramuli sulcato-quadrangulati, sicut petioli, foliorum venae primariae et secundariae abaxiales, inflorescentiae axes, bracteae et bracteolae pilis pinoideis vel pilis stellatis et asperis modice vel dense induti. Lamina 8.1– 16×4.2–7.5 cm elliptica vel ovata apice acuminata basi rotundata vel subcordata et subpeltata, 3–5-nervata. Inflorescentia in axillis vel lateralis in ramulis infra folia plerumque oriunda 1.5–4 cm longa multiflora, bracteolis 0.5–1 mm longis; flores 4(5)-meri; calyx primum in gemma apiculata clausus demum in lobos irregulares persistentes [this word ‘persistent’not in the English text] ruptus, dentibus calycis exterioribus 0.25–0.5 mm lineari-oblongis. Stamina isomorpha glabra, antherarum thecis 1.25×0.5 mm flavis oblongis poro truncato vel ventraliter subinclinato; connectivum nec prolongatum nec appendiculatum. Ovarium 4-loculare et omnino inferius, apice glabrum.

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Shrub or small tree, 1.5–3 m tall; uppermost quadrisulcate internodes, vegetative buds, petioles, elevated primary and secondary veins on abaxial foliar surface, inflorescence rachis, bracts and bracteoles moderately to copiously covered with pinoid hairs intermixed or replaced by sessile stellate and asperous hairs. Leaves of a pair equal to somewhat unequal in length; petioles 0.9–3 cm long; blade membranaceous to chartaceous, 8.1–16×4.2–7.5 cm, elliptic to elliptic-ovate, apex acuminate, base rounded to subcordate and generally subpeltate, margin subentire to denticulate or crenulate, 3–5-plinerved with the innermost pair of primary veins diverging from the midvein 0.8–1.4 cm above the blade base, adaxially glabrous in mature leaves, abaxially stellate on tertiary and higher order veins. Inflorescences laxly branched dichasia in the axils of the upper leaves or lateral, usually arising on branchlets below the leaves and paired mainly at defoliated nodes, 1.5–4 cm long; bracts and bracteoles 0.5–1 mm long, subulate. Flowers 4(5)-merous, on pedicels 1–2 mm long that lengthen to 4 mm in fruit. Hypanthia (at anthesis)1.5–2 mm long, narrowly campanulate to urceolate, apically elevated into a neck-like collar, sparsely stellate. Calyx fused in bud, shortly apiculate and rupturing into 4 irregular, broadly rounded hyaline lobes 0.25–0.75 mm long and 0.5–0.75 mm wide at the base, the exterior calyx teeth 0.25–0.5 mm long, linearoblong. Petals 2.5–3×2.5–3 mm, glabrous, white, ovate to orbicular, conspicuously reflexed at anthesis. Stamens 8(–10), isomorphic; filaments 1.5 mm long, glabrous; anthers 1.25× 0.5 mm, yellow, linear-oblong, laterally compressed and deeply channeled between the thecae, truncate to slightly emarginate distally with a truncate to somewhat ventrally inclined pore; connective simple. Ovary 4-locular, completely inferior, the apex elevated into a 0.1 mm glabrous collar. Style 5 mm long, glabrous; stigma truncate to capitellate. Berry 3–4×3– 4 mm, reportedly white when mature (F. Almeda & T. Daniel 7040). Seeds 0.4–0.6 mm long, yellow-brown, ± triangular in outline, rugulate. Distribution.—Clidemia subpeltata is endemic to Costa Rica where it is known from the Caribbean slope of the Guanacaste and Talamanca mountain ranges at (100) 900– 1200 m. Clidemia subpeltata is a very rare

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species known from few collections; in the Guanacaste mountain range for example, it is known from the single specimen cited from that area. (Fig. 5).

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Phenology.—Collected in flower and fruit in June, August, and September. Etymology.—The specific epithet refers to the subpeltate leaf blade base of this species.

FIG. 6. Clidemia subpeltata. A. Habit. B. Subpeltate leaf base. C. Abaxial foliar surface indument. D. Floral bud. E. Flower at anthesis. F. Hypanthial indument. G. Stamens. H. Immature fruit. I. Seeds. (Drawn from the holotype.)

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Additional specimens examined. COSTA RICA. Cartago: Cantón de Paraíso, Parque Nacional Tapantí, al lado de quebrada en la salida del Sendero Arboles Caídos, 9°48′18′′N 83°57′12′′W, 1200 m, 27 Sept 2002, R. Kriebel et al. 1389 (CAS, CR, INB, MO); near bridge over Rio Aguiares, 1/2 km above Santa Cruz, 1500 m, 17 Jul 1965, R. W. Lent 659 (F). Guanacaste: Macizo Miravalles, Estación Cabro Muco, 10°42′45′′N 85°09′ 21′′W, 1100 m, 2 Oct. 2003, J. A. Azofeifa 76 (CR, INB, MO). Limon: Cuenca del Río Estrella, R. B. Hitoy Cerere, sendero en alrededores de la estación, 9°40′25′′N 83°01′35′′W, 100–200 m, 12 Jun 1999, A. Rodríguez et al. 4921 (CR, INB, MO); Rainforest slopes of Cerro Skopte west of Río Siori and Río Coén about 7 km beyond Coroma, 450–700 m, 19 Feb 1992, F. Almeda & T. Daniel 7040 (CAS, CR).

Clidemia subpeltata is distinguished by its complex indument that includes pinoid, sessile stellate, and asperous hairs on various parts, leaf blades that are mostly elliptic-ovate, with the base rounded to subcordate and generally subpeltate and conspicuously 3–5-plinerved, laxly branched dichasial inflorescences mainly on defoliated nodes but also on foliated nodes, 4-merous flowers, calyx that is fused in bud and then rupturing irregularly into 4 lobes, broadly rounded hyaline lobes, conspicuous linearoblong calyx teeth and yellow, unappendaged laterally compressed anther thecae, and 4locular ovary. Among congeners Clidemia subpeltata appears to be related to the widespread C. septuplinervia Cogn. Both species share inflorescences which are borne mainly on defoliated nodes, 4-merous flowers with conspicuous subulate calyx teeth, unappendaged anther thecae and 4-locular ovary. Other less significant similarities between the two are their conspicuously plinerved leaves and pedicels that can lengthen to 4 mm in fruit. They can be easily be differentiated because C. septuplinervia has 7–9-plinerved leaves with a long decurrent base, cymose inflorescences, ovate calyx lobes with glandular-setose calyx teeth, white anther thecae (drying yellow) with a dorsally inclined pore and a glandular-puberulent collar on the ovary apex (vs. 5(−7)-plinerved leaves with the base rounded to subcordate and generally subpeltate, dichasial inflorescences, irregular calyx lobes with non glandular-setose calyx teeth, yellow anther thecae with a truncate pore and glabrous ovary apex in C. subpeltata). In Mesoamerica, there are

more than 20 species of Clidemia with 4merous flowers. Of these, only C. subpeltata has a fused calyx in bud that ruptures into irregular hyaline lobes. This character is more common in the genus Miconia, and three species of 4-merous Miconias in Mesoamerica have this distinctive calyx, one of which, M. calocoma Almeda, seems to be similar to C. subpeltata. Both species share a similar indument type, 5(−7)-plinerved leaves, 4merous flowers with fused calyx and linearoblong calyx teeth, anther thecae that are yellow, linear-oblong, laterally compressed and deeply channeled between the thecae, truncate to slightly emarginate distally with a truncate to somewhat ventrally inlined pore, simple connective, 4-locular ovary and a truncate to capitellate stigma. The two species are easily differentiated because M. calocoma has petioles 5–10 mm long, leaf-blade margins that are undulate to undulate-dentate, terminal or pseudolateral paniculiform inflorescences in which the ultimate branches terminate in multiflowered, congested glomerules, and pedicels 0.5 mm long. Their distribution is also different. Although both species are presently known from the Caribbean slope of Costa Rica, M. calocoma occurs in northeastern part of the country south to the OTS La Selva Field Station and Pandora, Limón, at 20–220 m, whereas C. subpeltata occurs on the mountain ranges of Guanacaste and Talamanca, at 900–1200 m, with southernmost populations that occur at 100–700 m in the Hitoy Cerere Biological Reserve region. The specimens from the more southern populations (Almeda & Daniel 7040, Rodríguez et al. 4921) differ from typical C. subpeltata in the narrower leaves to 5.3 cm wide, which are less evidently subpeltate. Acknowledgments We wish to thank Beth Guy and Silvia Troyo for their illustrations. William Burger and an anonymous reviewer provided valuable comments on the manuscript. Daniel Santamaria, Frank Gonzalez, and Isabel Pérez kindly provided assistance in processing and locating herbarium specimens at INB. Kriebel also also thanks Daniel Solano and Manuel Zumbado for assistance and company during field work. We

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also thank the Lakeside Foundation of the California Academy of Sciences for additional financial support. Literature Cited Almeda, F. 1984. New and noteworthy additions to the Melastomataceae of Panama. Proceedings of the California Academy of Sciences 43(17): 269–282. ———. 1989. Five new berry-fruited species of Tropical American Melastomataceae. Proceedings of the California Academy of Sciences 46(5): 137–150. ———. 2003. Chromosome Cytology and Taxonomy of the Red Goblet-Flowered Species of Clidemia (Melastomataceae: Miconieae) in Central and South America. Novon 13: 161–169. ———. 2004. Novelties and nomenclatural adjustments in the neotropical genus Clidemia (Melastomataceae: Miconieae). Proceedings of the California Academy of Sciences 55: 89–124.

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———. in press. Melastomataceae. In G. Davidse, M. Sousa-Sánchez, S. Knapp & F. Chiang (editores generales), Flora Mesoamericana 4(1). ———, R. Kriebel & G. Umaña. 2007. Melastomataceae. Pages 394–574 in B.E. Hammel, M.H. Grayum, C. Herrera, and N. Zamora, eds. Manual de Plantas de Costa Rica. Vol. 6. Dicotiledóneas (Haloragaceae-Piperaceae). Monographs in Systematic Botany from the Missouri Botanical Garden Vol. 111. Martin, C. V., D. P. Little, R. Goldenberg & F. A. Michelangeli. 2008. A phylogenetic evaluation of Leandra (Miconieae, Melastomataceae): a polyphyletic genus where the seeds tell the story, not the petals. Cladistics 24(3): 315– 327. Michelangeli, F. A., D. S. Penneys, J. Giza, D. Soltis, M. H. Hills & J. D. Skean Jr. 2004. A preliminary phylogeny of the tribe Miconieae (Melastomataceae) based on nrITS sequence data and its implications on inflorescence position. Taxon 53 (2): 279–290.