Three new species of Dendropanax (Araliaceae) from Bahia, Brazil PEDRO FIASCHI Fiaschi, P. (Herba´rio Andre´ Maurı´cio Vieira de Carvalho, CEPEC/CEPLAC, Rodovia Ilhe´us-Itabuna, Km 22, Itabuna, Bahia, C.P. 7, 45600-970, Brazil; e-mail:
[email protected]). Three new species of Dendropanax (Araliaceae) from Bahia, Brazil. Brittonia 57: 240–247. 2005.—Three new species of Brazilian Dendropanax from the rain forests of Bahia state are described and illustrated. Dendropanax amorimii, endemic to the rain forests near Boa Nova, in southern Bahia, is characterized by compound umbellate inflorescences with elongated primary branches each having a proximal whorl of bracts, and by leaves with welldeveloped tertiary venation. Dendropanax bahiensis, endemic to the Atlantic rain forests of southern Bahia, is distinguished by its large and membranous leaves with reddish schizogenous glands on the abaxial surface and by the reduced and branched inflorescences with short-pedicellate flowers and large fruits. Dendropanax geniculatus, an endemic species from the ‘‘matas de grota˜o’’ of Serra da Chapadinha, Chapada Diamantina, is characterized by simple and usually geniculate inflorescences bearing long-pedicellate flowers and fruits. Key words: taxonomy.
Araliaceae, Atlantic rain forests, Dendropanax, southern Bahia,
Fiaschi, P. (Herba´rio Andre´ Maurı´cio Vieira de Carvalho, CEPEC/CEPLAC, Rodovia Ilhe´us-Itabuna, Km 22, Itabuna, Bahia, C.P. 7, 45600-970, Brazil; e-mail:
[email protected]). Three new species of Dendropanax (Araliaceae) from Bahia, Brazil. Brittonia 57: 240–247. 2005.—Treˆs espe´cies novas de Dendropanax de florestas ombro´filas do estado da Bahia sa˜o descritas e ilustradas. Dendropanax amorimii, endeˆmica de florestas ombro´filas das proximidades de Boa Nova, na regia˜o sul da Bahia, e´ caracterizada por infloresceˆncias do tipo umbela composta com ramos prima´rios alongados com um verticilo proximal de bra´cteas e folhas com a nervac¸a˜o tercia´ria bastante desenvolvida. Dendropanax bahiensis, endeˆmica de florestas ombro´filas do sul da Bahia, e´ caracterizada pelas folhas grandes e membrana´ceas com glaˆndulas esquizo´genas avermelhadas visı´veis na face abaxial e pelas infloresceˆncias curtas e ramificadas com flores curtopediceladas e frutos grandes. Dendropanax geniculatus, endeˆmica das ‘‘matas de grota˜o’’ da Serra da Chapadinha, Chapada Diamantina, e´ caracterizada pelas infloresceˆncias simples e geralmente geniculadas com flores e frutos longo-pedicelados.
Dendropanax Decne. & Planch. comprises 92 species from tropical and subtropical Asia and Central and South America. The majority of these (ca. 65 spp.) occur in the Americas, were the genus is best represented in the rain forests of northwestern South America, with secondary centers in south-
ern Mesoamerica, Jamaica, and Eastern Brazil (Frodin & Govaerts, 2004). Most of the Brazilian species of Dendropanax are endemic to the Atlantic rain forests of the Eastern coast, where more than 10 species occur, some of which are, as yet, undescribed.
Brittonia, 57(3), 2005, pp. 240–247. q 2005, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.
ISSUED: 29 September 2005
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FIASCHI: DENDROPANAX (ARALIACEAE)
Species of Dendropanax are glabrous plants with simple or palmatilobed leaves that are often 3-nerved at the base and have schizogenous glands and small intrapetiolar stipules; the inflorescences are simple or more or less compoundly arranged umbels of unjointed, pedicellate 5–9-merous flowers with valvate petals, glandular and versatile anthers, and fruits that sometimes have a distinct stylar column (Cannon & Cannon, 2001; Hutchinson, 1967). A recent phylogenetic analysis using ITS sequence data initially suggested that Dendropanax is related to Hedera (Wen et al., 2001), but Plunkett et al. (2004) have since shown this relationship to be uncertain, placing Dendropanax within the broader ‘‘Asian Palmate clade.’’ In addition to this uncertain position within Araliaceae, Lowry et al. (2004) suggested that the genus may not be monophyletic and could thus require changes in its circumscription. As a result of recent collections made in the state of Bahia by the staff of the Mata Atlaˆntica Nordeste and the Chapada Diamantina projects, and the subsequent analysis of specimens available at the herbaria ALCB, CEPEC, HUEFS, and SPF, three new species of Dendropanax were found. These species are here described and illustrated, and some of their ecological and phylogenetic affinities are discussed. Dendropanax amorimii Fiaschi, sp. nov. (Fig. 1) TYPE: BRAZIL. Bahia: Mun. Boa Nova, Fazenda Sa˜o Jose´, 8.8 km E de Boa Nova, na estrada para Dario Meira, a 1.4 km do ramal a` esquerda 148239450S, 408089460W, 850 m, 7 Jan 2001 (fl), A. M. Amorim, J. G. Jardim & F. S. Junchum 3589 (HOLOTYPE: CEPEC; ISOTYPES: K, MBM, NY, SPF). Species nova a D. geniculato Fiaschi inflorescentiis ramosis, ramis primariis 6–8, elongatis, bracteis verticillatis in dimidio proximali, basibus laminarum cuneatis vel attenuatis, apicibus plerumque obtusis vel rotundatis differt.
Shrubs or treelets 2.5–6 m high. Young branches 3.5–4.5 mm diam., striate longitudinally. Stipule ca. 2 mm long, entire.
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Leaves spirally alternate, sometimes subopposite, crowded on the terminal portion of the branches or separated by internodes to ca. 3 cm long; petiole 0.5–6.5 cm long, slightly compressed laterally, adaxially plane or slightly canaliculate, thickness uniform or thicker at apex and/or base; blade horizontal, plane, papery, discolorous, 3.7– 14.7 cm long, 1.4–6.4 cm wide, elliptic to slightly obovate, symmetric or slightly asymmetric, the apex obtuse to rounded, sometimes mucronate, the base cuneate to attenuate, the margin entire, plane or slightly revolute; venation acrodromous suprabasal, imperfect, brochidodromous, the main vein prominent on both surfaces, more so on abaxial one, the secondary veins in 6–8 pairs, slightly prominent on abaxial surface; the basal pair diverging from midvein at ca. 1.5 cm from blade base, distinct, the angle of divergence 258–508, other pairs with angle of divergence 508– 608, the intersecondary veins clearly thinner than secondary ones, the tertiary veins and reticulation hardly to easily visible and prominent on abaxial surface. Inflorescence terminal, erect, the main axis up to ca. 1.5 mm long; the primary branches 6–8, 4.5– 11 cm long, umbellately or sub-racemosely arranged, erect to sub-horizontal, with whorled bracts 1.5–4.5 cm from the base, the apex thickened. Terminal inflorescence units with 25–30 flowers grouped in umbels. Flowers greenish, the pedicel 0.7–1.8 cm long; hypanthium 2–3 mm long, longer than corolla; calyx lobes evident; petals 5, triangular, 2.1–2.4 mm long, 1.1–1.7 mm wide, the apex cucullate; stamens 5, white, the filaments 1.8–2.2 mm long, the anthers versatile, ca. 1.2 mm long, ca. 1.2 mm wide, dorsally glandular at the apex; styles 0.5–0.7 mm long. Fruits unknown. Distribution and ecology. This species appears to be endemic to a single patch of wet forest near Boa Nova, in southern Bahia. Although both wet and dry forests occur in the area, the species was found only in the former. Etymology. The specific epithet honors Andre´ M. Amorim, who has made many excellent collections of Araliaceae. Additional specimen examined: BRAZIL. BAHIA:
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FIG. 1. Dendropanax amorimii.A. Flowering branch. B. Leaf undersurface. C. Detail of leaf undersurface. D. Floral bud. E. Flower. F. Petal, adaxial view. G. Stamen, adaxial view. H. Stamen, abaxial view. I. Transverse section of ovary. (from Amorim 3589, CEPEC)
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FIASCHI: DENDROPANAX (ARALIACEAE)
Boa Nova, Fazenda Sa˜o Jose´, 8.8 km E de Boa Nova, na estrada para Dario Meira, a 1.4 km do ramal a` esquerda 148239450S, 408089460W, 850 m, 8 Mar 2003 (st), Sant’Ana et al. 1088 (CEPEC).
Dendropanax amorimii resembles D. geniculatus with respect to its leaf undersurfaces, which have distinct tertiary venation and lack schizogenous glands. It differs from D. geniculatus mainly by its inflorescences, which are composed of 6–8 umbellately arranged (vs. simple) primary branches with a whorl of bracts in the proximal half, and by the leaf blades, which have an obtuse to rounded (vs. acute to acuminate) apex and a cuneate to attenuate (vs. usually obtuse to rounded) base. Dendropanax amorimii shares with D. langsdorfii (Marchal) Frodin and D. trilobus (Marchal) Frodin the presence of a whorl of bracts in the proximal half of the primary branches, but differs from these species mainly by its slightly obovate (vs. elliptic) leaves with obtuse to rounded (vs. attenuate to acuminate) apex, indistinguishable (vs. visible) schizogenous dots, and the hypanthium larger (vs. shorter or with the same length) than corolla. Dendropanax bahiensis Fiaschi, sp. nov. (Fig. 2) TYPE: BRAZIL. Bahia: Mun. Itajuı´pe, Distrito de Unia˜o Queimada, ca. 4 km na estrada para Pimenteiras. Ca. 148399S, 398289W, 29 Jan 2003 (fr), P. Fiaschi, S. C. Sant’Ana & J. L. Paixa˜o 1259 (HOLOTYPE: CEPEC; ISOTYPES: HUEFS, MBM, MO, NY, K, SPF). Species nova a D. exili (Toledo) Jung fructibus maioribus (9–10 mm longis et 10–10.5 mm latis) et breviter pedicellatis (usque ad 5 mm longis), basibus laminarum obtusis vel rotundatis, interdum cuneatis, apicibus acutis vel breviter acuminatis differt.
Shrubs or sub-shrubs 0.5–3.5 m high. Young branches 3–5 mm diam. Stipule reduced, to ca. 1 mm long, entire. Leaves spirally alternate, sometimes sub-opposite, crowded on the terminal portion of the branches or separated by internodes to ca. 6 cm long; petiole 1.5–9.5 cm long, slightly compressed laterally, adaxially plane or slightly canaliculate, of uniform diameter or thicker at apex and/or base; blade horizon-
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tal, plane, membranous, slightly discolorous, 10–27.8 cm long, 3–12.5 cm wide, elliptic to slightly obovate, symmetric or slightly asymmetric, the apex short to longacuminate, the base cuneate to obtuse, the margin entire or with up to 4 pairs of teeth (each less than 1 mm long) in the distal half, plane; venation acrodromous suprabasal, imperfect, brochidodromous, the main vein prominent on both surfaces, more so on abaxial one, the secondary veins in 7–11 pairs, prominent only on the abaxial surface, the basal pair diverging from midvein at ca. 0.5 cm from blade base, distinct, the angle of divergence 308–408, the other pairs with the angle of divergence 608–708, the intersecondary veins sometimes present, clearly thinner than secondary ones, the tertiary veins and reticulation hardly visible on the abaxial surface. Inflorescence terminal, erect, the main axis 2–3 cm long; primary branches 5–10, 1–3.5 cm long, racemosely arranged, erect, lacking whorled bracts, the apex thickened. Terminal inflorescence units with ca. 20 flowers grouped in umbels. Flowers greenish, the pedicel up to 2 mm long (expanding to 3– 5 mm long in fruit); hypanthium ca. 1.2 mm long, shorter than corolla; calyx lobes evident, acuminate; petals 5, ovate, 1.2–1.6 mm long, 0.8–1.1 mm wide, the apex cucullate; stamens 5, white, the filaments ca. 1.5 mm long, the anthers versatile, ca. 0.7 mm long, 0.7 mm wide, dorsally glandular at apex; styles ca. 0.4 mm long. Fruits spheroid to obloid, 5-lobed when dry, 9–10 mm long, 10–10.5 mm wide, the stylar column ca. 1 mm long; pyrenes 5, ca. 8 mm long, 4 mm wide. Distribution and ecology. Dendropanax bahiensis is endemic to wet forests in the vicinity of Ilhe´us, in the Southern Bahia region. Despite its rather widespread distribution in the ‘‘cocoa region’’ of southern Bahia, where recent collecting efforts have been made, this species remains poorly collected and, thus, is presumably rare. Etymology. The specific epithet makes reference to the fact that this species is presumably endemic to the state of Bahia. Additional specimens examined: BRAZIL. BAHIA: Almadina, Rod. Almadina-Ibitupa˜, entrada a 7 km,
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FIG. 2. Dendropanax bahiensis. A. Fruiting branch. B. Leaf. C. Detail of leaf undersurface. D. Detail of the leaf margin. E. Floral bud. F. Flower. G. Stamen, adaxial view. H. Stamen, abaxial view. I. Fruit. J. Transverse section of fruit. (A, C, I, J: Fiaschi et al. 1259, CEPEC; B, D: Carvalho & Sant’Ana 4355, CEPEC; E,F,G,H: Jardim et al. 1025, CEPEC)
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FIASCHI: DENDROPANAX (ARALIACEAE)
Serra do Sete Paus, ca. 12 km da entrada, Faz. Cruzeiro do Sul, 148449060S, 398419460W, 578 m, 28 Feb 1997, Jardim et al. 1025 (CEPEC); Ibirapitanga, 22 km N of Itamarati on BR 101, then 6.8 km E on road to Embratel Tower. Reserva Municipal Cachoeira do Pau. 13853927909S, 398279339W, 690 m, 19 Mar 2003, Thomas et al. 13457 (CEPEC, NY, SPF); Ilhe´us, Mata da Esperanc¸a, parte W, acesso rodovia Ilhe´us/Itabuna, Banco da Vito´ria, 148469300S, 398059000W, 30 Jan 2000, Jardim et al. 2589 (CEPEC, NY, SPF); Itajuı´pe, Distrito de Unia˜o Queimada, ca. 4 km na estrada de Unia˜o Queimada para Pimenteira, 9 Feb 1994, Carvalho & Sant’Ana 4355 (CEPEC).
Together with D. exilis (Toledo) Jung and other as yet undescribed species from Espı´rito Santo state, these species form a group characterized by a shrubby habit, leaves with schizogenous dots visible on the abaxial surface, reduced inflorescences (simple or branched), and small and shortpedicellate greenish flowers. This group is endemic to the interior of the Atlantic rain forests of eastern Brazil, ranging from Bahia to Santa Catarina states. Dendropanax bahiensis can be distinguished from D. exilis by its larger leaves (10–27.8 3 3–12.5 cm vs. 5–15 3 1.5–3.5 cm), inflorescences with more densely grouped primary branches (peduncle up to 1 cm long vs. usually more than 1 cm long), shorter pedicels (to ca. 2 mm vs. 3–4 mm), and larger fruits (9–10 3 10–10.5 mm vs. ca. 5 3 5 mm). Dendropanax geniculatus Fiaschi, sp. nov. (Fig. 3) TYPE: BRASIL. Bahia. Lenc¸o´is, Serra da Chapadinha, mata de grota˜o. Ca. 128279S, 418269W, 900 m, 23 Feb 2003, P. Fiaschi, A. Rapini & Lı´cio 1338, fr. (HOLOTYPE: CEPEC; ISOTYPES: HUEFS, K, MBM, NY, SPF). Species nova a D. monogyno (Vell.) Seem. foliis chartaceis, lamina elliptica subtus non punctata, nervis tertiariis atque reticulo subtus manifestis; a D. amorimii Fiaschi inflorescentiis simplicibus geniculatis, basibus laminarum obtusis vel rotundatis interdum cuenatis, apicibus acutis vel acuminatis differt.
Trees or hemi-epiphytes to ca. 10 m high. Young branches 2–3 (–5) mm diam., striate longitudinally. Stipule reduced, to ca. 1.5 mm long, entire. Leaves spirally alternate, sometimes sub-opposite, crowded on the terminal portion of the branches or separat-
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ed by internodes to ca. 2 cm long; petiole 0.5–2.5(–5) cm long, laterally compressed, usually thicker at the base; blade horizontal, plane, papery, slightly discolorous, 3.9–14 cm long, 2.2–6.5 cm wide, elliptic to slightly ovate or obovate, symmetric to slightly asymmetric, the apex acute to acuminate, mucronate, the base obtuse, rarely cuneate or rounded, the margin entire, slightly revolute, sometimes with as many as two pairs of small teeth (each less than 1 mm long) in the distal third; venation acrodromous suprabasal, imperfect, brochidodromous, the main vein prominent on both surfaces, more so on abaxial one, the secondary veins 6–9 pairs, prominent only on the abaxial surface, the basal pair diverging from midvein at ca. 0.5 cm from blade base, sometimes indistinct, the angle of divergence 408–508, other pairs with angle of divergence 608–708, the inter-secondary veins similar to secondary ones, the tertiary veins and reticulation evident on the abaxial surface. Inflorescence terminal, or pseudo-lateral due to the subsequent growth of a new vegetative branch, the main axis 0.5–1 cm long; the primary branch 1, geniculate or rarely sub-erect, 4.2–8 cm long, purplish to blackish when mature, lacking whorled bracts. Terminal inflorescence units with 25–30 flowers grouped in umbels. Flowers not seen. Fruits obloid, 5–6-lobed when dry, ca. 6.5 mm long, 8.5 mm wide, the stylar column 1–1.2 mm long; pyrenes 5(6), 4.5–5.5 mm long, ca. 3 mm wide. Distribution and ecology. Dendropanax geniculatus has to date been found only in the ‘‘grota˜o’’ forests of Serra da Chapadinha, in the vicinity of Lenc¸o´is, in the interior of the state of Bahia. Although these forests have a more western distribution than the main body of the coastal rain forests, they were recently included in the Mata Atlaˆntica phytogeographic unit based on floristic similarities (Veloso et al., 1991). Etymology. The specific epithet refers to the position of the inflorescence, which tends to be perpendicular to the axis of growth of the fertile branch, although the angle between the primary branch of the inflorescence and the vegetative branch sometimes does not exceed 308, as can be seen in the illustration here presented.
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FIG. 3. Dendropanax geniculatus. A. Fruiting branch. B. Leaf. C. Detail of leaf undersurface. D. Fruit. E. Transverse section of fruit. (A–C: Gasson et al. PCD5878, CEPEC; D, E: Fiaschi et al. 1338, CEPEC)
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Additional specimens examined: BRAZIL. BAHIA: Lenc¸o´is, Serra da Chapadinha, Chapadinha, 128279000S, 418269000W, 910 m, 21 Feb 1995, Melo et al. PCD1670 (ALCB, CEPEC, SPF); Gasson et al. PCD5878 (ALCB, CEPEC, HUEFS, SPF); 128279000S, 418259000W, 700 m, Giulietti & Funch PCD1574 (ALCB, CEPEC).
Dendropanax geniculatus shares with D. monogynus (Vell.) Seem. simple and geniculate inflorescences, but clearly differs by several leaf features, such as the elliptic blade with acuminate apex (vs. usually ovate blade with acute apex), the presence of evident tertiary venation (vs. inconspicuous), and the absence of clearly evident schizogenous glands on the abaxial surface. It differs from D. amorimii Fiaschi by having simple inflorescences (sometimes with two shorter lateral branches) and leaves with an acute to acuminate (vs. obtuse to rounded) apex and a usually obtuse (vs. cuneate to attenuate) base. To date, there is only one available flowering collection of D. geniculatus (Gasson et al. PCD5878, ALCB), but this bears only immature flower buds on two short branches lateral to the primary branch of the inflorescence. In the other material examined, lateral branches are absent but leave a scar when shed, thus rendering the infructescences simple. Acknowledgments I am grateful to Jose´ Lima da Paixa˜o and Se´rgio Cardoso de Sant’Ana for their help during the field work, to Andre´ M. Amorim for his excellent collections of Dendropa-
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nax, to Jackie Kallunki for her revision of the English text, Latin diagnosis, and helpful comments, and to Greg Plunkett and an anonymous reviewer for their suggestions. This work was funded by the Mata Atlaˆntica Nordeste project, a scientific collaboration between CEPLAC, UESC, and NYBG. Literature Cited Cannon, M. J. & J. F. M. Cannon. 2001. Araliaceae. Pages 189–192. In: W. D. Stevens, C. U. Ulloa, A. Pool & O. M. Montiel, editors. Flora of Nicaragua. Missouri Botanical Garden, St. Louis. Frodin, D. G. 1995. Neotropical Montane Araliaceae: an overview. Pages 421–430. In: S. P. Chrchill, H. Baslev, E. Forero & J. L. Luteyn, editors. Biodiversity and conservation of neotropical Montane forests. The New York Botanical Garden,Bronx, New York. ——— & R. Govaerts. 2004. World checklist and bibliography of Araliaceae. The Royal Botanic Gardens, Kew. Hutchinson, J. 1967. The genera of flowering plants (Angiospermae). Vol. 2. Clarendon Press, Oxford. Lowry, II, P. P., G. M. Plunkett & J. Wen. 2004. Generic relationships in Araliaceae: looking into the crystal ball. S. African J. Bot. 70: 382–392. Plunkett, G. M., J. Wen & P. P. Lowry, II. 2004. Infrafamilial classifications and characters in Araliaceae: insights from the phylogenetic analysis of nuclear (ITS) and plastid (trnL-trnF) sequence data. Plant Syst. Evol. 245: 1–39. Veloso, H. P., A. L. R. R. Filho & J. C. A. Lima. 1991. Classificac¸a˜o da vegetac¸a˜o brasileira, adaptada a um sistema universal. IBGE, Departamento de Recursos Naturais e Estudos Ambientais, Rio de Janeiro. Wen, J., G. M. Plunkett A. D. Mitchell & S. J. Wagstaff. 2001. The evolution of Araliaceae: a phylogenetic analysis based on ITS sequences of nuclear ribosomal DNA. Syst. Bot. 26(1): 144–167.
