Systematic Botany (2012), 37(4): pp. 978–982 © Copyright 2012 by the American Society of Plant Taxonomists DOI 10.1600/036364412X656554
Spirotheca elegans (Malvaceae: Bombacoideae), a New Species from Bahia, Brazil Jefferson Guedes de Carvalho-Sobrinho,1 Marlon Caˆmara Machado, and Luciano Paganucci de Queiroz Herba´rio HUEFS, Universidade Estadual de Feira de Santana, Avenida Universita´ria s/n, Novo Horizonte, 44036-900, Feira de Santana, Bahia, Brazil. 1 Author for correspondence (
[email protected]) Communicating Editor: Benjamin van Ee Abstract—Spirotheca elegans, a new species of Malvaceae subfamily Bombacoideae from the state of Bahia, Brazil, is described and illustrated. It is considered to be endemic to an area of Caatinga vegetation bordering the Chapada Diamantina mountain range, in the central region of the state. Notes on its distribution, ecology, phenology, and comments on its conservation status are given. Furthermore, a discussion of the relationship of the new taxon to other species of Spirotheca and an updated key are provided.
Spirotheca Ulbr. (Malvaceae) is a genus of trees that is noteworthy among the Bombacoideae for often displaying a strangler habit, the plants start life as epiphytes, grow in the canopies of host trees, send down masses of adventitious roots which may gradually envelop the host tree and eventually replace it. It occurs in moist mid- to highelevation forests from Nicaragua, Costa Rica, and Panama through the western Andean countries to Bolivia, and in coastal forests of eastern and southeastern Brazil (Gibbs and Alverson 2006). The genus is diagnosed by anthers with four bisporangiate thecae arrayed as two superposed pairs, and these anthers are tightly spiralled around the style in the bud stage until anthesis, conferring the genus its name (Gibbs and Alverson 2006). Also, the staminal tube of some species have a distinct swelling, a feature which is used to distinguish species in the genus. The precise phylogenetic placement of Spirotheca is still unresolved. It can either be more closely related to Bombax L. and Pachira quinata ( Jacq.) W. S. Alverson, or to Ceiba Mill., sharing with all of these taxa a spiny trunk (Duarte et al. 2011). Spirotheca and Ceiba also share an androecium of five stamens and androecial units with two or four thecae, differing from the remaining multistaminate, monothecate representatives of the core Bombacoideae (Alverson et al. 1999; Baum et al. 2004). The morphological similarities between Ceiba and Spirotheca are reflected in their taxonomic history, considering that Ulbrich (1914) transferred Ceiba rivieri Decne. to his new genus Spirotheca. Gibbs and Alverson (2006) accepted five species and one variety: Spirotheca awadendron Fern. Alonso, S. mahechae Fern. Alonso, S. michaeli Cuatrec., S. rivieri (Decne.) Ulbr., S. rivieri var. passifloroides (Cuatrec.) P. E. Gibbs & W. S. Alverson, and S. rosea (Seem.) P. E. Gibbs & W. S. Alverson. All species of Spirotheca published to date occur in moist forests. Efforts carried out to monitor populations of Bombacoideae in semi-arid and wet areas of eastern Brazil led us to collect sterile material of a striking tree at first thought to belong to Ceiba due to the presence of spines on the trunk and articulated leaflets. Complete material covering all the phenological phases of this tree allowed us to recognize it as a new species of Spirotheca, the first of the genus found to inhabit seasonally dry tropical forest vegetation (SDTF) sensu Pennington et al. (2000), locally known as Caatinga, from the state of Bahia, Brazil. It is described and illustrated herein.
Taxonomic Treatment Spirotheca elegans Carv.-Sobr., M. Machado & L. P. Queiroz, sp. nov.—TYPE: BRAZIL. Bahia: Iraquara, estrada para Gruta da Pratinha, logo apo´s a vila Lagoa de Santa Rita, 12 210 03.500 S, 41 340 42.000 W, 691 m, 18 April 2011 (lf, fl, fr), J. G. de Carvalho-Sobrinho & M. Machado 3055 (holotype: HUEFS; isotypes: RB, SPF).
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Similar to Spirotheca rivieri var. passifloroides (Cuatrec.) P. E. Gibbs & W. S. Alverson in its small flowers, but differing by the downward slanting flowers, white winged petals, doubly swollen staminal tube and anthers with smaller thecae. Free-standing tree to 12 m, crown with sparse, remote branches, usually leafless when flowering; trunk densely covered with woody, conical to umbonate spines up to 20 mm long, or more rarely unarmed, terminal branches often unarmed; bark grayish with lighter-colored striations. Stipules linear, 1–2 mm long, caducous. Leaves palmately compound, clustered toward the branch tips; petioles pulvinate, slender, 15–60 1 mm; petiolules absent; leaflets (3–)4–5, 20– 45 8–20 mm, membranaceous, glabrous, upper surface lustrous green, lower surface matte-green, oblong to narrowly obovate, slightly conduplicate, base cuneate, apex slightly retuse, obtuse or acuminate, often mucronate, margins entire, midrib conspicuous on both surfaces, secondary venation inconspicuous. Inflorescences fasciculate, arising from the branch tips, with 1–3(–5) flowers; pedicels 20–25 mm long; flower buds linear-oblong, basal third swollen, ascending to erect. Flowers ca. 45 mm long, slanting downwards; receptacle 3–4 mm long, with a whorl of 5–8 glands; calyx cupuliform, 4–5 6–8 mm, sericeous internally, glabrous externally, inconspicuously 5–apiculate, caducous in fruit; petals 40–45 7–8 mm (to 10 mm at the wing), linear-oblong, markedly reflexed at anthesis, longitudinally asymmetrical, outer margin straight, inner margin crispate and membranous, provided with a wing located midway up from base, proximal 10–12 mm of each petal concave, greenish-yellow, fleshy, then becoming white, flushed pink towards the proximal fleshy part, with minute, dense, tufted trichomes on both surfaces, glabrescent next to the wing, wing ca. 12 mm long, crispate, folded in bud, in open flower located 3–4 mm above the reflexion point of petal; staminal tube 23–25 mm long, with dense, tufted trichomes, glabrous at the ends, proximal two-thirds greenish-yellow, distal one-third pinkish, doubly swollen with a lower, pentagonal, cupuliform portion, ca. 3 6 mm, and an upper obclavate portion, ca. 16 4 mm, apex
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Keywords—Caatinga, distribution, ecology, endemism, taxonomy.
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CARVALHO-SOBRINHO ET AL.: NEW BRAZILIAN SPECIES OF SPIROTHECA
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dividing into five free, pinkish staminal filaments ca. 6 mm long, declinate at anthesis, anthers extrorse, both upper and lower thecae pairs 3–4 mm long; ovary conical, densely pubescent, with moniliform trichomes, 5-locular, style terminal, slender, pinkish, projecting ca. 14 mm beyond the staminal tube, declinate at anthesis, stigma 5-lobed, reddish. Capsule 30–65 30–35 mm, obovoid, rarely spheroid, apex rounded, 5-valvate, valves glabrous, kapok abundant, white. Seeds 6– 7 5–6 mm, pyriform, brownish, alveolate in SEM, hilum region often prominent. Pollen in monads, amb triangular, ca. 30 mm, 3-colporate, planaperturate, exine reticulate. Figures 1–3. Additional Specimens Examined—BRAZIL. Bahia: Mun. Anage´, ca. 5 km na estrada para a barragem, 14 380 15.100 S, 41 100 06.500 W, 402 m, 18 Sep 2010 (fr), J. G. de Carvalho-Sobrinho et al. 2868 (HUEFS); 14 380 15.100 S, 41 100 06.500 W, 402 m, 12 Mar 2011 (lf, fl), J. G. de Carvalho-Sobrinho & M. Machado 2964 (HUEFS); 14 380 15.100 S, 41 100 06.500 W, 400 m, 20 May 2011 (fr), J. G. de Carvalho-Sobrinho & M. Machado 3094 (HUEFS); 14 380 15.100 S, 41 100 06.500 W, 400 m, 16 Jul 2011 (fr), J. G. de Carvalho-Sobrinho 3181 (HUEFS). Mun. Iraquara, Olhos D’a´gua, 12 200 6000 S, 41 340 4000 W, 7 Feb 2008 (lf), A. F. Venaˆncio & F. Esteves 11 (ALCB). Mun. Paramirim, localidade
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de Tabuinha, ca. 16 km de Caturama, 13 180 12.800 S, 42 120 24.100 W, 668 m, 14 Mar 2011 (lf, fl), J. G. de Carvalho-Sobrinho & M. Machado 2977 (HUEFS).
Etymology—The specific epithet is a reference to the exquisite pinkish-white flowers of this species and to the small, delicate leaves. Habitat, Distribution, and Ecology—Spirotheca elegans has only been found growing as a tall, free-standing tree, and the epiphytic strangler habit reported for the other species of the genus has not been observed. It is the only species of the genus inhabiting a SDTF area, regarded as endemic to the vegetation bordering the Chapada Diamantina mountain range, in the central region of Bahia state, occurring at elevations from about 400 to about 700 m (Fig. 4). Queiroz and Lavin (2011) identified a radiation of species of Coursetia DC. confined to this area, and suggested that small patches of Caatinga may harbor distinctive biotas. This seems to be the case of the vegetation surrounding the Chapada Diamantina. This area has been more intensely botanized in the last 15 yr and several new taxa, probably endemic to this area, have been described, for example, Tabaroa L. P. Queiroz, G. P.
Fig. 1. Spirotheca elegans. A. Branch with leaves. B. Detail of trunk densely covered with woody spines. C. Terminal branch with spines. D. Twoflowered fascicle. E. Petal, showing the winged inner margin. F. Androecium. G. Fruit. [A from Carvalho-Sobrinho & Machado 3055; B–F from CarvalhoSobrinho & Machado 2964; and G from Carvalho-Sobrinho & Machado 3094]. Drawing by Pe´tala G. Ribeiro.
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Fig. 2. Spirotheca elegans. A. Individual at the type locality (Carvalho-Sobrinho & Machado 3055). B. Leaves of Spirotheca elegans (right) and S. rivieri (left), both from material cultivated at UEFS. C. Flower at anthesis and bud (Carvalho-Sobrinho & Machado 2977). D. Xylocopa bee, indicated with a white arrow, visiting a flower at anthesis in Paramirim, Bahia. E. Fruits (Carvalho-Sobrinho 3181). Bars: A: 1 m; B–C, E: 1 cm.
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Fig. 3. Scanning electron micrographs of Spirotheca elegans. A. Tufted trichomes from the staminal tube. B. Pollen grain. C. Seed. D. Seed detail showing the alveolate surface. [A–B from Carvalho-Sobrinho & Machado 2964; C–D from Carvalho-Sobrinho 3181].
Fig. 4.
Known localities of Spirotheca elegans in Bahia, Brazil.
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located uppermost, being at an appropriate position for pollination when the bee lands in the flower. Indeed, the bees were observed moving up and down the staminal tube, and as they did so they most likely brushed the anthers and stigma with the back of their abdomens, thus possibly effecting pollination. A further indication that these bees are the pollinators of S. elegans is that the upper staminal tube is thickened near the apex (obclavate) and quite floccose on its proximal portion. This could be an adaptation to facilitate the landing of bees, giving them a foothold while at the same time protecting the style within the staminal tube from damage. Symptomatically, the upper staminal tube of senescent flowers is usually quite damaged in S. elegans, suggesting repeated visitation by bees. Taxonomic Discussion—Spirotheca elegans appears to be closest to S. rivieri, which possesses similar small leaves and flowers, in particular to S. rivieri var. passifloroides, which also has a staminal tube with a swollen portion, above which the tube becomes constricted. However, S. elegans differs from S. rivieri due to the stronger and more numerous spines on the trunk, the membranaceous leaflets ( chartaceous leaflets), the downward slanting flowers ( erect flowers), the white flushed pink color of the petals ( uniformly brilliant red) which are asymmetrical lengthwise ( symmetrical petals) with a wing and a crispate margin ( wing absent and entire margins), the yellowish doubly swollen staminal tube ( red, not swollen or singly swollen tube), and the smaller anthers with upper and lower thecae 3–4 mm long ( upper thecae 6–8 mm long and lower thecae 7–11 mm long). Moreover, the two species appear to be adapted to different pollinators
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Lewis & M. Wojciech. (Queiroz et al. 2010), Aeschynomene soniae G. P. Lewis (Lewis 1993), Senegalia santosii (G. P. Lewis) Seigler & Ebinger, and Coursetia caatingicola L. P. Queiroz (Leguminosae; Queiroz and Lavin 2011). Conservation Status—The known populations of Spirotheca elegans consist of few individuals, six were observed at Anage´, one at Paramirim, and five at Iraquara, although no exhaustive survey was made at the locations. The species as a whole, although distributed over a relatively large area, seems to occur only in small patches of suitable habitat that are quickly disappearing due to logging and forest clearing for cultivation and cattle. Additionally, none of the known populations of this species are located in protected areas. Spirotheca elegans is therefore regarded as endangered (EN) according to criteria B2ab(ii,iii), C2a(i) of the IUCN red list (IUCN 2010). Pollination—Spirotheca elegans flowers from March to April and fruits from May to September. The flower morphology of S. elegans is distinctive and appears to be adapted to bee pollination, unlike the putative close relative S. rivieri, which seems adapted to pollination by perching birds (Rocca and Sazima 2008), hummingbirds, and woodpeckers (Rocca et al. 2006). Spirotheca rivieri has erect flowers with wide, open tubes making access to nectar easy for birds. In contrast, the flowers of S. elegans slant downwards and birds would have to hang upside-down or nearly so to reach the nectar. Moreover, the flower tube of S. elegans is narrower and occluded by the cupuliform lower swollen portion of the staminal tube, blocking access to nectar. The bases of the petals are fleshy, and would require a relatively strong animal to move them aside so that the nectar can be reached. Carpenter bees (Xylocopa sp.) were observed busily visiting the flowers of S. elegans at Paramirim and Iraquara. The bees landed on the staminal tube, probed for nectar, and then moved on to repeat the process on another flower. The anthers and stigma in S. elegans are spatially separated and radially disposed, with anthers and stigma facing the petals, suggesting a tube-brush functional type. Moreover, the stigma is
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as discussed above. Both S. elegans and S. rivieri occur in Bahia but are not sympatric since S. elegans occurs in Caatinga vegetation of the central region of the state and S. rivieri is known only from humid coastal forests. Another tree species of Bombacoideae which occurs in semiarid region of Bahia, Ceiba rubriflora Carv.-Sobr. & L. P. Queiroz, resembles S. elegans in its stoutly spiny trunk, 5-foliolate leaves, small, elliptic leaflets with entire margins, deciduous calyx, and reflexed petals. However, C. rubriflora can be promptly distinguished by the red flowers and larger dimensions of the calyx, petals, anthers, and capsules. Furthermore, C. rubriflora has a lobed calyx, obovate petals, staminal tube swollen only at base, sinuous and dithecate anthers, and oblong fruits (Carvalho-Sobrinho and Queiroz 2008). Spirotheca elegans also resembles S. mahechae from Colombia in that both species have small leaves and pinkish flowers. However, S. elegans differs from it in the swelling on the staminal tube, by the shorter petals (40–45 mm 75–85 mm), staminal tube (23–25 mm 45–47 mm) and free staminal filaments (ca. 6 mm 25–28 mm), and smaller thecae (3–4 mm 15–18 mm). Spirotheca elegans differs from the remaining species
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of the genus by its smaller flowers and leaves, and by the calyx which is caducous (Fig. 1G, 2E), but persistent in the other species according to Gibbs and Alverson (2006). Moreover, Spirotheca elegans has a particular kind of trichome reported and illustrated here for the first time for the genus. The tufted trichomes displayed on its staminal tube (Fig. 3A) and on both surfaces of the petals are similar to those found by Carvalho-Sobrinho et al. (2009) on the abaxial surface of Pseudobombax Dugand petals. In summary, Spirotheca elegans is a remarkable species, both morphologically and ecologically different from its congeners, possibly with a pollination syndrome distinct from that of its closest allies, and also apparently endangered. Extensive surveys of the known localities to estimate population size and of similar areas to verify the extent of occurrence of the species are thus necessary. In addition, an investigation of the reproductive biology of this species is highly desirable to best determine the floral visitors and pollinators, so that the pollination system of the species can be compared to those of related species as a way of better understanding the evolution of the genus as a whole.
Expanded Key to Species of SPIROTHECA based on Gibbs and Alverson (2006) 1.
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1.
Petals 40–55 mm, brilliant red or pinkish-white throughout anthesis (Brazil) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Flowers erect; petals brilliant red, symmetrical; staminal tube red, not swollen or solely swollen at base; anthers with upper thecae 6–8 mm long and lower thecae 7–11 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. rivieri 2. Flowers slanting downwards; petals pinkish-white, asymmetrical, with a wing and a crispate margin; staminal tube yellowish, doubly swollen; anthers with upper and lower thecae 3–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. elegans Petals 60–120 mm, initially whitish green, becoming pinkish salmon, and finally dark reddish (Central America, Andean South America) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Staminal tube 45–47 mm, lacking an articulation, more or less uniformly sericeous below, sparsely so above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Petals ca. 120 mm; leaflets 70–160 30–56 mm, oblanceolate with an acuminate apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. michaeli 4. Petals 75–85 mm; leaflets 40–50 18–22 mm, narrowly obovate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. mahechae 3. Staminal tube 12–45 mm, with a distinct articulation, densely grey tomentose below, variably and usually sparsely sericeous above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5. Leaflets glabrous or glabrate, sometimes with sparse hairs or papillose or sparsely glandular-scurfy below (i.e. on the abaxial surface) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. rosea 5. Leaflets densely stellate-tomentose below . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. awadendron
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Acknowledgments. The authors would like to thank Dr. Marcello Moreira from Universidade Estadual do Sudoeste da Bahia, Vito´ria da Conquista, Bahia, for drawing our attention and taking one of us (MM) to the area where this new species grows (Anage´, Bahia); to Dr. Vinı´cius Antonio de Oliveira Dittrich (UFJF) and to Maria do Carmo M. Iannaccone for providing information and photographs of Spirotheca rivieri; to Leilton Damascena for preparing the map; to Fundac¸a˜o Osvaldo Cruz (FIOCRUZ) for the use of the scanning electron microscope, and Aline C. da Mota for technical guidance; Pe´tala G. Ribeiro for the line drawings, and to two anonymous reviewers for their suggestions to the first version of the manuscript. Fieldwork was supported by Graduate Program in Botany (PPGBot/UEFS) and Fapesb (APP0006/2011). JGCS and MM were supported by doctorate fellowships, respectively from CAPES and CNPq. LPQ was supported by a PQ-1C fellowship from CNPq (300811/2010-1). Literature Cited Alverson, W. S., B. A. Whitlock, R. Nyffeler, C. Bayer, and D. A. Baum. 1999. Phylogeny of the core Malvales: evidence from ndhF sequence data. American Journal of Botany 86: 1474–1486. Baum, D. A., S. D. Smith, A. Yen, W. S. Alverson, R. Nyffeler, B. A. Whitlock, and R. L. Oldham. 2004. Phylogenetic relationships of Malvatheca (Bombacoideae and Malvoideae; Malvaceae sensu lato) as inferred from plastid DNA sequences. American Journal of Botany 91: 1863–1871. Carvalho-Sobrinho, J. G. and L. P. Queiroz. 2008. Ceiba rubriflora (Malvaceae: Bombacoideae), a new species from Bahia, Brazil. Kew Bulletin 63: 649–653. Carvalho-Sobrinho, J. G., F. A. R. dos Santos, and L. P. Queiroz. 2009. Morfologia dos tricomas das pe´talas de Pseudobombax Dugand (Malvaceae, Bombacoideae) e seu significado taxonoˆmico. Acta Botanica Brasilica 23: 929–934.
Duarte, M. C., G. L. Esteves, M. L. F. Salatino, K. C. Walsh, and D. A. Baum. 2011. Phylogenetic analyses of Eriotheca and related genera (Bombacoideae, Malvaceae). Systematic Botany 36: 690–701. Gibbs, P. E. and W. S. Alverson. 2006. How many species of Spirotheca (Malvaceae s. l., Bombacoideae)? Brittonia 58: 245–258. IUCN. 2010. Guidelines for using the IUCN red list categories and criteria. Version 8.1. Prepared by the Standards and Petitions Subcommittee in March 2010. ( http://intranet.iucn.org/webfiles/doc/SSC/RedList/ RedListGuidelines.pdf ). [accessed 09/10/2011]. Lewis, G. P. 1993. Two new taxa of Aeschynomene (Leguminosae: Papilionoideae) from Brazil. Kew Bulletin 49: 93–97. Pennington, R. T., D. E. Prado, and C. A. Pendry. 2000. Neotropical seasonally dry forests and Quaternary vegetation changes. Journal of Biogeography 27: 261–273. Queiroz, L. P. de, G. P. Lewis, and M. Wojciechowski. 2010. Tabaroa, a new genus of Leguminosae tribe Brongniartieae from Brazil. Kew Bulletin 65: 189–203. Queiroz, L. P. de and M. Lavin. 2011. Coursetia (Leguminosae) from eastern Brazil: the monophyly of three caatinga-inhabiting species is revealed by nuclear ribosomal ITS/5.8S sequence data. Systematic Botany 36: 69–79. Rocca, M. A. and M. Sazima. 2008. Ornithophilous canopy species in the Atlantic rain forest of southeastern Brazil. Journal of Field Ornithology 79: 130–137. Rocca, M. A., M. Sazima, and I. Sazima. 2006. Woody woodpecker enjoys soft drinks: the blond-crested woodpecker seeks nectar and pollinates canopy plants in south-eastern Brazil. Biota Neotropica 6: 1–9. Ulbrich, E. 1914. Bombacaceae. Pp. 156–166 in Plantae Uleanae novae vel minus cognitae. ed. Pilger R. Notizblatt des Botanischen Gartens und Museums zu Berlin-Dahlem 6: 143–179.
