Gerardo A. Aymard C.1 & Lisa M. Campbell2 Caraipa Aublet
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CARAIPA ANDINA (CLUSIACEAE), A NEW SPECIES FROM THE VENEZUELAN ANDES, AND ITS BIOGEOGRAPHICAL IMPLICATIONS Gerardo A. Aymard C.1 & Lisa M. Campbell2 ABSTRACT (Caraipa andina (Clusiaceae), a new species from the Venezuelan Andes, and its biogeographical implications) Caraipa andina from La Fundación, in the western Andes of Venezuela (Táchira State) is described, illustrated, and its morphological relationship with an allied species is discussed. This new species is morphologically related to C. heterocarpa Ducke, but it differs by the leaf blade shape, size, and dense abaxial papillae; fewer- flowered inflorescence; and shorter pedicel, flower buds, petals, and filaments. Phytogeographical information about the La Fundación area is presented. This species represents the first report of this genus in the Venezuelan Andes. Key words: Amazonia, Estado Táchira, Guayana, Guttiferae, sandstone outcrops. RESUMO (Caraipa andina (Clusiaceae), uma nova espécie para os Andes da Venezuela e suas implicações biogeográficas) Caraipa andina foi descoberta em La Fundación, nos Andes Ocidentais da Venezuela (Estado de Táchira) e é aqui descrita e ilustrada, discutindo-se suas afinidades morfológicas com as espécies próximas. Morfologicamente relacionada com C. heterocarpa Ducke, a nova espécie diferencia-se pelo tamanho, forma da lâmina foliar e face abaxial densamente papilosa; inflorescência paucifloras; pedicelos, botões florais, pétalas e filamentos mais curtos. Informações sobre a fitogeografia do local são apresentadas. Esta espécie representa o primeiro registro deste gênero nos Andes Venezuelanos. Palavras-chave: Amazônia, Estado Táchira, Guayana, Guttiferae, afloramentos de arenito. Caraipa Aublet comprises about 30 densifolia, C. punctulata Ducke and C. species of small to large trees or shrubs, richardiana Cambess.) are found in the wet distributed in tropical South America and gallery forests in the Venezuelan Llanos (Gustafsson et al. 2002; Stevens 2007). The (Aymard 2003; Aymard & González 2006). genus has been variously treated as Bonnetiaceae, Caraipa llanorum Cuatr. subsp. llanorum Theaceae, and is presently included in and C. savannarum Kub. form ecologically Clusiaceae (Kubitzki 2007). The genus is interesting communities of dense trees called characterized by an indumentum of usually saladillales. These communities are very stellate hairs; alternate leaves with tertiary common in the vast eolic, seasonally flooded veins perpendicular to the secondary veins; savannas (the Llanos) that extend from stamens with free filaments and bearing an northeastern Colombia through western apical gland; and capsular fruits. Venezuela (Schargel & Aymard 1992; Huber Species occur in moist forest formations et al. 2006). The genus is rare in the Andes, (lowland, montane slopes, gallery, flooded, and and currently is known only by several Amazonian caatinga) and shrubby savannas, collections of C. punctulata Ducke from with the largest number of species in the Colombia (Antioquia and Norte de Santander Amazon Basin and on the Guiana Shield. departments), and reported herein for However, a few taxa occur outside of this region, Venezuela. A revision of the genus was such as the two subspecies of C. densifolia published by Kubitzki (1978; as Bonnetiaceae), Martius in southeastern and central Brazil. who reconized 21 species and three morphological Three species (C. densifolia Martius subsp. groups; however, these groups did not Artigo recebido em 11/2007. Aceito para publicação em 04/2008. 1Corresponding author: UNELLEZ-Guanare, Programa de R. N. R., Herbario Universitario (PORT), Mesa de Cavacas, Estado Portuguesa, Venezuela 3350. [email protected] 2The New York Botanical Garden, Bronx, NY 10458-5126, U.S.A. [email protected] 394 Aymard C., G. A. & Campbell, L. M. accommodate all of the species. Kubitzki (1978) with microscopic epicuticular wax, secondary pointed out two taxonomically useful leaf veins in 8–10 pairs, impressed adaxially, characters to separate the species: phyllotaxy prominent abaxially. Inflorescence axillary, lax spiral vs. distichous, and the cells of the abaxial panicles, 10–15 cm long, 12–16-flowered; the surface bullate or clavate, with the cuticle minutely rachis and pedicels covered by simple and wrinkled and scaly vs. cells of the abaxial stellate trichomes; pedicels 5–10 mm long, surface not protruding, and the cuticle mostly bracts and bracteoles not seen. Flower buds smooth. Since then, five new species have been globose, 4–6 cm long before anthesis; sepals described (Kubitzki 1987; Vásquez-Martínez, ovate, ca. 2 mm long, ca. 2 mm wide, stellate 1991) and others suggested (Kubitzki & Holst pubescent and covered with translucent dots 1998; see also Stevens 2007). Additionally, the abaxially, adpressed yellow pubescent genus has been treated for the Flora of the adaxially, margins ciliate; petals oblong, 6–8 Venezuelan Guayana (Kubitzki & Holst mm long, 3–5 mm wide, puberulent abaxially, 1998), and two local florulas in the Amazonian glabrous adaxially; stamens 80–100, filaments region of Colombia and Peru (Vásquez- 5–6 mm long, anthers ca. 0.5 mm long, oblong, Martínez 1997; Rudas & Prieto 2005). introrse, connective distally widened and This contribution increases to fifteen the topped by a gland; ovary ca. 2 mm long, number of Caraipa species known from tomentose, trichomes yellow; style ca. 3 mm Venezuela. This new species represents the long, glabrous, stigma trilobed. Fruits not seen. first record of this genus in the Venezuelan Distribution and habitat: Caraipa andina Andes, and was discovered during herbarium appears to be restricted to the moist montane work for a phytogeographic study being forests near La Fundación, Táchira state, conducted in the area in which it was collected. Venezuela. Phenology: Collected with flowers in July. Caraipa andina Aymard & L. M. Campb. The ranked phyllotaxis, with leaves sp. nov. Fig. 1 glabrous on both surfaces, cells of their lower Type: VENEZUELA. TÁCHIRA: Dtto. surface not protruding, and with a smooth Uribante, empresa Las Cuevas, near La cuticle, and lax panicles more 4 cm long, relate Fundación, 08º50’N, 71º47’W, ca. 900 m, this new species morphologically to Caraipa 7.VII.1983, H. van der Werff & A. González heterocarpa, a species known from a few 5013 (Holotype: PORT; Isotype: MO, n.v.; specimens, with restricted distribution in low NY; VEN). elevation flooded forests in the Amazon basin Species Caraipa heterocarpae Ducke of Brazil and Venezuela (Kubitzki 1978; proxima, sed differt: foliis oblongis, 6–9 cm Kubitzki & Holst 1998). However, C. andina longis, subtus dense papillatis, differs from that species by its leaf blades inflorescencia 12–16-flora, pedicellis 5–10 oblong, 6–9 cm long, densely papillate on the mm longis, gemmis floralibus 4–6 longis, abaxial surface (vs. leaf blades lanceolate- sepalis ovatis, ca. 2 mm longis, intus adpress ovate or lanceolate, 10–17 cm long, not papillate luteo-pubescentibus, petalis 6–8 longis, on the abaxial surface), inflorescence with 12– filamentis 5–6 longis, ovario dense luteo- 16 flowers, pedicels 5–10 mm long (vs. glandulosis trichomatibus obtecto. inflorescence with 20–40 flowers, pedicels 12– Tree, 20 m tall. Young branches and 16 mm long), flower buds 4–6 cm long; sepals branchlets smooth, glabrous. Leaves distichous, ovate, ca. 2 mm long, adpressed yellow petiole 6–8 mm long, stout, glabrous; blades pubescent adaxially, petals 6–8 mm long, and coriaceous, 6–9 cm long, 3–5 cm wide, base filaments 5–6 mm long (vs. flower buds 7–8 rounded, apex acute to rounded, margins slightly cm long; sepals obtuse, 3–4 mm long, slightly revolute, glabrous on both surfaces with sparse strigose adaxially, petals ca. 10 mm long, and translucent dots, abaxial epidermal cells papillate, filaments 10–12 mm long). Rodriguésia 59 (2): 393-398. 2008 Caraipa andina 395 1 cm b 5 cm a Figure 1 – Caraipa andina – a. branch showing the inflorescence and flowers; b. detail of an inflorescence branch. (van der Werff 5013). Phytogeography: Species of Caraipa are The only known locality of Caraipa distributed almost exclusively in the Guayana andina, La Fundación, is located in the western region and Amazon Basin, and are rarely portion of the Cordillera Mérida, on sandstone encountered in the tropical Andes. The first outcrops belonging to the Formación Aguardiente report of the genus in this region was C. (Notestein et al. 1944; Salvador 1961a, b). The colombiana Ewan, a species described from vegetation of this area is relatively well known a collection from forests on the slopes of the because of the extensive collections made by R. Cordillera Central in Colombia (Ewan 1951). Liesner, J. A. Steyermark and H. van der Werff. Kubitzki (1978) treated this species as a Liesner (pers. comm.) observed that the floristic synonym of C. punctulata Ducke, a mostly composition of the vegetation associated with lowland wet forest species, that he considered these sandstone outcrops includes several species to also be distributed on mountain slopes in that are known from the Guiana Shield, and the Venezuela and Colombia; notably, he did not Amazon Basin floras, such as Philodendron mention that this collection (J. Cuatrecasas atabapoense G. S. Bunting (Araceae), Bonnetia 13335, F, US) would have represented the only paniculata Spruce ex Benth. (Bonnetiaceae), known occurrence of Caraipa in the Andes. Vriesia duidae (L. B. Sm.) Gouda (Bromeliaceae), Intensive botanical exploration in the Colombian Licania latifolia Benth. (Chrysobalanacae), Andes during