Two new species and a new combination of Neotropical Sapotaceae ANDERSON ALVES-ARAÚJO1,2
AND
MARCCUS ALVES2
Programa de Pós-Graduação em Biologia Vegetal – PPGBV/UFPE, CEP 50670–901, Recife-PE, Brazil; e-mail:
[email protected] 2 Laboratório de Morfo-Taxonomia Vegetal, Departamento de Botânica, Universidade Federal de Pernambuco – UFPE, CEP 50670–901, Recife-PE, Brazil 1
Abstract. This paper provides descriptions and illustrations of two new species of Sapotaceae (Pradosia longipedicellata and Chromolucuma apiculata) from Bahia, Brazil, a new combination (Chromolucuma congestifolia), and a new identification key for Chromolucuma. Pradosia longipedicellata is distinguished from other congeners by its clustered fascicles of flowers on pedicels 2–3 cm long at the apices of the shoots, and a corolla that is 7–8 mm long. Chromolucuma apiculata is distinguished by glabrous shoots and leaves, short petioles (10–15 mm long), and stipules 5–8 mm long with apiculate apices, while its close relative C. congestifolia has a lower leaf surface with trichomes usually restricted to the midrib and stipules 3–6 mm long with acute apices. Due to intense deforestation in the Amazonian and the Atlantic forests, preliminary IUCN Red List assessments are provided. Pradosia longipedicellata and Chromolucuma congestifolia are proposed the IUCN status Vulnerable, while the data for C. apiculata are still insufficient (Data Deficient). Key Words: Brazil, Chrysophylloideae, conservation, diversity, Neotropics. Resumo. São descritas e ilustradas duas novas espécies de Sapotaceae (Pradosia longipedicellata e Chromolucuma apiculata), ocorrentes no estado da Bahia, Brasil, além de uma nova combinação (Chromolucuma congestifolia) e uma nova chave de identificação para Chromolucuma. Pradosia longipedicellata é reconhecida entre as espécies do gênero pela presença de fascículos agrupados no ápice dos ramos, flores longo-pediceladas (2–3 cm compr.) e corola 7-8 mm compr. Chromolucuma apiculata é distinta pelos seus ramos e folhas glabros, pecíolos mais curtos (10–15 mm long) e estípulas (5–8 mm long) com ápices apiculados, enquanto que C. congestifolia tem face abaxial foliar geralmente com tricomas restritos à nervura central e estípulas mais curtas (3–6 mm long) com ápices agudos. Dada à intensa devastação das florestas Amazônica e Atlântica, uma prévia classificação segundo as normas da IUCN Red List é proposta para as espécies. Pradosia longipedicellata e Chromolucuma congestifolia são consideradas como pertencentes ao status da IUCN como Vulneráveis, enquanto que dados sobre C. apiculata ainda são insuficientes para tal proposta (Dados Deficientes).
Tropical America is one of the regions with the highest species diversity of angiosperms in the world (Mittermeier et al., 1999; Myers et al., 2000), which also is where the sapodilla family, Sapotaceae, has its greatest diversity (Pennington, 2004). One of the most important centers of diversification for the family in the Neotropics is the Amazonian forest (Pennington
1991, 2006; Anderberg & Swenson, 2003; Swenson & Anderberg, 2005). However, other biomes, such as the Atlantic forest, also show a high species richness. The Atlantic forest, most of which is in Brazil, is considered one of the richest biogeographic regions in the world in terms of plant species diversity and endemism, but it has been heavily impacted by human
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development (Mittermeier et al., 1999; Myers et al., 2000). According to Myers et al. (2000), the Atlantic forest is a hotspot of diversity, and for this reason it is considered one of the highest priority areas for conservation. In most of northeastern Brazil, the Atlantic forest is restricted to a strip of vegetation along the coast with an average elevation of up to 100 m (Velloso et al., 1991). Sapotaceae consists of 58 genera and about 1250 species in subtropical and tropical areas of the world (Pennington, 1991; Govaerts et al., 2001; Swenson et al., 2007a, b). For the New World, Pennington (1990) monographed the family and later provided a generic classification for the family (Pennington, 1991). Recent phylogenetic studies have demonstrated that Pennington’s concepts of genera do not always correspond with natural groups (Anderberg & Swenson, 2003; Bartish et al., 2005; Swenson et al., 2007a, 2008). Subsequently, Swenson and Anderberg (2005) provided a new subfamilial classification, recognizing Chrysophylloideae, Sapotoideae, and Sarcospermatoideae, and also indicated that genera such as Chrysophyllum and Pouteria sensu Pennington (1990, 1991) are polyphyletic (Bartish et al., 2005; Swenson et al., 2007a, 2008). While carrying out a taxonomic survey of Pouteria in the Atlantic forest, new species of Pouteria (Alves-Araújo & Alves, in press), Pradosia, and Chromolucuma have been discovered. This paper aims to describe two novelties, one in each of the latter two genera, both of which are exclusively Neotropical and belong in the Chrysophylloideae (Swenson & Anderberg, 2005; Swenson et al., 2008). Pradosia is a genus of 23 species of trees and geoxylic shrubs, all confined to tropical America, except for the African species P. spinosa (Ewango & Breteler, 2001), which probably belongs to Pouteria (Swenson et al., 2008). Pradosia is characterized by spirally arranged leaves, usually eucamptodromous leaf venation, and the absence of stipules. The flowers are bisexual with exserted stamens and no staminodes. The fruit is different from most other genera in the family, because it develops into a one-seeded drupe with a thin cartilaginous endocarp (Pennington, 1991). The presence of a drupe is the most
important character to distinguish Pradosia from Elaeoluma, which has a berry. Chromolucuma is a small genus of two tree species in South America (Pennington, 1990). Two important diagnostic characters are yellow latex and persistent stipules. The inflorescence is axillary and bears unisexual, pedicellate flowers. The stamens are included in the corolla, and staminodes are always present (Pennington, 1991). Presence of stipules is an important feature for both Chromolucuma and Ecclinusa, but the latter genus has white latex (not yellow), sessile flowers, and lacks staminodes. Recent phylogenetic analyses, based on molecular and morphological data, indicate that Chromolucuma is related to some species of Pouteria (Swenson et al., 2008); however, only one species of Chromolucuma was included in this study. Although flowers of the species described below are still unknown, the presence of yellow latex, persistent stipules, and a distinct pedicel give us a reliable indication that it is a member of Chromolucuma. Pilz (1981) described Pouteria congestifolia and called attention to the presence of stipules and yellow latex, which are uncommon for the genus. Despite this fact, Pennington (1990, 1991) kept the species in Pouteria when he recognized Chromolucuma. Hence, we here transfer P. congestifolia to Chromolucuma and provide a new identification key for the genus. New Species Pradosia longipedicellata Alves-Araújo & M. Alves, sp. nov. Type: Brazil. Bahia: Una, Reserva Biológica do Mico-LeãoDourado, BA-001 Km 46, 15º09’S, 39º05’W, 9 Mar 1993 (fl, fr), J. G. Jardim, A. M. Amorim, S. C. de Sant’Ana, E. B. dos Santos & J. L. Lage 92 (holotype: CEPEC; isotype: NY). (Figs. 1A–D, 2A–C, 3) Pradosiae brevipedi similis, sed habitu arboreo, 10– 20 m alto, ramis juvenibus sericeis, pilis candidis vel auratis, lenticellis absentibus, foliis 7–13 cm longis, chartaceis, ellipticis vel ovatis, petiolis 1–1.5 cm longis, inflorescentiis floribus 2–4, pedicellis 1.8–3 cm longis, calyce et corolla 5–6 mm longis, staminibus 5–6 mm longis, ovario loculis 5–7, stylis 5–6 mm longis imprimis differt.
Trees 10–20 m tall, shoot without lenticels, tomentulose with whitish to golden trichomes. Leaves spirally arranged, ovate to
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FIG. 1. A–D. Pradosia longipedicellata. A. Habit with details of lower leaf indument. b Outer (above) and inner (below) sepals. C. Corolla and stamens. D. Gynoecium. E–H. Chromolucuma apiculata. E. Habit. F. Stipules from the outside (left) and inside (right). G. Fruit. H. Seed, view of testa (left) and seed scar (left). (A–D, from Jardim et al. 92, CEPEC; E–H from Voeks 88, CEPEC.)
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FIG. 2. A–C. Pradosia longipedicellata. A. Holotype (Jardim et al. 92, CEPEC). B. Inflorescence and flowers of living material (photo by J. G. Jardim). C. Inflorescence of the voucher (Jardim et al. 92, CEPEC). D–E. Chromolucuma apiculata. D. Holotype (Voeks 88, CEPEC). E. Stipule (arrow).
elliptic, 7–12 × 4.8–6 cm, chartaceous, both surfaces tomentulose with whitish to golden trichomes, upper surface sometimes shiny, base cuneate, apex acute, margin flat; venat-
ion eucamptodromous, midrib slightly sunken on the upper surface; petiole 9–14 mm long, channeled, tomentulose with whitish to golden trichomes. Inflorescences at apices of
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criteria (IUCN, 2001, 2008), P. longipedicellata is assigned a preliminary threat status of Vulnerable (VU – A1cd,B1). Phenology.─Pradosia longipedicellata has been recorded with flowers and fruits almost throughout the year (March to November). Etymology.─The name of this species refers to the long pedicel of the flower.
FIG. 3. Location of the Atlantic forest, together with known distribution of Pradosia longipedicellata (circles), Chromolucuma apiculata (square), and C. congestifolia (triangles).
short branches; fascicles in clusters, umbelliform, 2–4 flowers (5-merous) per fascicle; pedicels 2–3 cm long, tomentulose with whitish to golden trichomes. Sepals 5–6 mm long, the outer slightly shorter than the inner, ovate, tomentulose on the outer surface with whitish to golden trichomes, glabrous on the inner surface, apex acute to ± rounded, margin entire. Corolla cup-shaped, white, 7–8 mm long, tube ca. 2 mm long; lobes 5–6 mm long, ovate, margin entire, apex truncate to rounded, glabrous. Stamens ca. 6 mm long, fixed in the middle of the tube; filaments glabrous, ca. 4 mm long, upper half deflexed, whitish; anthers 1.8– 2 mm long, glabrous. Ovary 5–7-locular, 5– 6 mm long, ferruginous pilose; style 5–6 mm long, glabrous, green; stigma simple. Fruit 1seeded, 3–4 cm long, ellipsoid, style persistent in fruit, villous to tomentulose of ferruginous trichomes; seeds laterally compressed, 20– 22 mm long; testa smooth, shiny, brownish; seed scar 18–20 mm long, 2–3 mm wide. Distribution and conservation status.─Pradosia longipedicellata is currently known only from the coastal Atlantic forest of southern Bahia in northeastern Brazil (Fig. 3). It has a narrow distribution with seven known subpopulations, often near cocoa plantations. It is a naturally rare species with few individuals in each location. Following the IUCN Red List
Additional specimens examined. BRAZIL. Bahia: Una, Reserva Biológica do Mico-Leão-Dourado, 15º09'S, 39º05'W, 9 Nov 1993 (fl, fr), A. Amorim et al. 1424 (CEPEC, NY); Marambaia, 15 Jul 1995 (fr), A. M. Amorim et al. 6067 (CEPEC); Estrada Ubaitaba-Maraú, 5 Sep 1999 (fr), A. M. Carvalho et al. 6730 (CEPEC, MO, NY); Una, Estação Experimental do CEPLAC, 24 Oct 1980 (fr), A. Rylands 48–1980 (CEPEC); Ilhéus, Fazenda Guanabara, 16 Oct 1980 (fr), L. A. Mattos-Silva et al. 1166 (CEPEC); Estrada Itacaré-Serra Grande, Maraú, 12 Jan 1967 (fr), R. P. Belém 3079 & R. S. Pinheiro (CEPEC); Itacaré, 9 May 1968 (fr), R. P. Belém 3507 (CEPEC); Estrada Ubaitaba-Maraú, 24 May 1990 (fl), T. S. Santos et al. 4554 (CEPEC); Una, Reserva Biológica do Mico-Leão-Dourado, 15º09'S, 39º05'W, 20 Sep 1998 (fr), S. C. Sant'Ana et al. 668 (CEPEC, MO, NY).
Pradosia longipedicellata is recognized by its clustered fascicles at the apices of short shoots and pedicels 2–3 cm long. However, there are other species of Pradosia that have flowers with long pedicels, such as P. brevipes (Pierre) T. D. Penn., P. colombiana (Standl.) T. D. Penn., and P. ptychandra (Eyma) T. D. Penn. (Pennington, 1990). Pradosia longipedicellata is easily distinguished from P. ptychandra, which has shorter sepals (~2 mm long) and a shorter corolla (~5 mm long), and from P. colombiana, which has longer petioles (20–45 mm) and a shorter corolla (~3 mm). The flowers of P. longipedicellata resemble that of P. brevipes, but the first is a tree that lacks lenticels on the shoots, and it is only known from the Atlantic forest. In contrast, P. brevipes is a geoxylic subshrub that usually has shoots with lenticels, and it is known from the Brazilian Cerrado. In other words, P. longipedicellata and P. brevipes differ in habit and are allopatric. Chromolucuma apiculata Alves-Araújo & M. Alves, sp. nov. Type: Brazil. Bahia: Ilhéus, Fazenda Barra do Manguinho, Rodovia BA-001 Km 11, 14º47’S, 39º02'W, 31 Jan 1985 (fr), R. Voeks 88 (holotype: CEPEC). (Figs. 1E–H, 2D–E, 3)
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Chromolucumae rubriflorae affinis, a qua lenticellis praesentibus, stipulis lanceolatis, ubi juvenibus tomentosis, ubi adultis glabris, apice apiculato, foliis coriaceis vel crassis, glabris, petiolis 10–15 mm longis, fructibus 15– 18 mm longis, seminibus 15–16 mm longis, testis laevibus et nitidis imprimis differt.
Trees 10–12 m tall with whitish bark and yellow latex; shoots glabrous with lenticels; stipules 5–8 mm long, lanceolate, first tomentose with ferruginous trichomes, glabrescent; apex apiculate. Leaves spirally arranged, oblanceolate, 16–21 x 5.5– 7.5 cm, coriaceous, glabrous, lower surface glaucous, base cuneate, apex acute, margin slightly revolute; petiole 10–15 mm long, not channeled, glabrous. Fascicles at apices of branches, axillary, 1–2-flowered; pedicel 10–18 mm, glabrous. Sepals ca. 4 mm long, ovate, glabrous to glabrescent on the outer surface, glabrous on the inner surface, apex obtuse, margin entire, sometimes ciliate. Flowers unknown. Fruit 1-seeded, 25–30 mm long, spherical to ellipsoid, glabrous, purple; seeds 15–18 mm long, slightly laterally compressed; testa smooth, shiny, brownish; seed scar 14–16 mm long, ca. 4 mm wide. Distribution and conservation status.─ Chromolucuma apiculata is known only from the coastal Atlantic forest of Bahia, Brazil (Fig. 3). This is the first record of the genus occurring outside the Amazonian region and is an example of a disjunct distribution between the Amazonian and Atlantic forests. The species is only known from the type collection and we refrain from giving it a preliminary conservation assessment, and instead classify it as Data Deficient (DD). Phenology.─Fruiting in January; flowering period unknown. Etymology.─This species is named for the characteristic apiculate stipules. Among Chromolucuma, C. apiculata is the only species that has young shoots and leaves that are completely glabrous, along with short petioles (10–15 mm long), and stipules that are 5–8 mm long and apiculate. New Combination Chromolucuma congestifolia (Pilz) AlvesAraújo & M. Alves, comb. nov. Pouteria congestifolia Pilz, Ann. Missouri Bot.
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Gard. 68: 191. 1981. Type: Panamá. Coclé: El Valle de Antón, [16 Mar 1946] (fl), P. H. Allen 3426 (holotype: MO; isotypes: BM, F, G). (Fig. 3) Distribution and conservation.─Chromolucuma congestifolia is known from the mountain region between Costa Rica and Panama, and the Amazonian forest in Brazil and French Guiana (Fig. 3). Because of human impact, increasingly more forest is fragmented, especially in the Brazilian Amazon. Ten populations of C. congestifolia are known, three of which are in protected areas. Considering this information, C. congestifolia is assigned a preliminary threat status of Vulnerable (VU – A1c,B1,B2c). Phenology.─Chromolucuma congestifolia has been recorded with flowers and fruits from February to September. Additional specimens examined. COSTA RICA. Alajuela: Reserva Biológica Monteverde, Laguna de Minor Vargas, 10°18'36"N, 84°42'36"W, 14 Sep 1989 (fl), E. Bello 1269 (MO); Laguna de Poco Sol, 10°21'N, 84°39'36"W, 30 Sep 1989 (fl), E. Bello 1300 (MO); Reserva Biológica de San Ramon, 10º18’N, 84º34’W, 30 May–1 Jun 1986 (fl), B. Hammel et al. 15262 (MO, CR); without exact locality, 21 Feb 1984 (fr), T. D Pennington & L. J. Poveda 11546 (K). Cartago: Paraiso, 09° 44'05"N, 83°46'42"W, 14 Mar 2000 (fl), L. Acosta & V. H. Ramírez 627 (MO). Heredia: North of Quebrada Tigre from Finca El Plástico, 10º18’N, 84º02’W, 14 Feb 1986 (fl), M. H. Grayum & P. Sleeper 6494 (MO). PANAMA. Chiriquí: Along Quebrada de Arena, 12 Mar 1982 (fr), S. Knapp et al. 4068 (MO). Veraguas: Near Cerro Tute-Arizona, above Santa Fe and Alto de Piedra, 08°30'N, 81°10'W, 5 Feb 1988 (fl), G. McPherson 12050 (MO). FRENCH GUIANA. Saul, Eaux Claires, 9 Feb 1993, (fr), T. D. Pennington et al. 13840 (K). BRAZIL. Amazonas: Manaus-Itacoatiara, Reserva Florestal Ducke, km 26, 02º53’S, 59º58’W, 1976, J. A. Souza s.n. (INPA); Manaus-Itacoatiara, Reserva Florestal Ducke, km 26, 02º53’S, 59º58’W, 22 Aug 1997, P. A. C. L. Assunção 627 (INPA, K).
Among Chromolucuma species, C. congestifolia is distinguished by its shorter stipules (3–6 mm long) and a lower leaf surface that usually has long trichomes restricted to the midrib. In contrast, C. baehniana has a sericeous lower leaf surface, and C. rubriflora and C. apiculata have glabrous leaves. For more detailed information about Chromolucuma species, see the following key.
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Key to the species of Chromolucuma 1. Lower leaf surface sericeous; shoots without lenticels (Guiana, Venezuela). . . . . . . . . C. baehniana 1. Lower leaf surface glabrous (except for midrib); shoots with lenticels. 2. Leaves chartaceous; stipules 2–4 cm long; pedicels 4–8 cm long (Venezuela, northern Brazil) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. rubriflora 2. Leaves coriaceous; stipules
