Systematic Botany (2014), 39(1): pp. 260–267© Copyright 2014 by the American Society of Plant TaxonomistsDOI 10.1600/036364414X678224Date of publication 02/05/2014

A New Deciduous Species of Pachira (Malvaceae: Bombacoideae) from a SeasonallyDry Tropical Forest in Northeastern Brazil

Jefferson G. Carvalho-Sobrinho,1,3 William S. Alverson,2 Aline C. Mota,1

Marlon C. Machado,1 and David A. Baum2

1Programa de Pos-Graduacao em Botanica, Universidade Estadual de Feira de Santana, Avenida Transnordestina s/n,Novo Horizonte, Feira de Santana, Bahia, 44036-900, Brazil.

2Department of Botany, Birge Hall, University of Wisconsin-Madison, Madison, Wisconsin 53706, U. S. A.3Author for correspondence (jef.sobrinho@gmail.com)

Communicating Editor: Leslie Goertzen

Abstract—Pachira (Malvaceae: Bombacoideae) is distributed predominantly in northern South America and is the largest genus in thesubfamily, including about 50 species. Ongoing studies of the systematics of neotropical Bombacoideae and field work in northeastern Brazilled to the discovery of a new species, herein described and illustrated. Pachira moreirae is known from a single population that is endemic tocaatinga vegetation (a type of seasonally dry tropical forest) in the state of Bahia. The new species is leafless during its flowering period, acondition commonly seen in some genera of Bombacoideae but rare in Pachira, a genus that mostly comprises evergreen species. We provide adescription of the morphology of this new species and notes on its distribution, phenology, and conservation status. A phylogenetic analysisusing ITS data shows that P. moreirae is a member of a Brazilian extra-Amazonian clade of Pachira that also includes P. glabra and P. retusa.

Keywords—Bombacaceae, Caatinga, ITS, phylogeny, taxonomy.

Pachira Aubl. (Malvaceae: Bombacoideae) is a Neotropicalgenus with the majority of its species found in northernSouth America (Alverson et al. 1999; Bayer et al. 1999; Baumet al. 2004). As currently circumscribed, it is the largestgenus in the subfamily, with ca. 50 species (Alverson 1994;Fernandez-Alonso 1998, 2003a, 2003b, 2003c; Robyns 1963;Tropicos 2012).Many of the species now placed in Pachira were assigned

previously to Bombacopsis Pittier, Pochota Ram. Goyena, orRhodognaphalopsis A. Robyns (Alverson 1994; Alverson andSteyermark 1997; Fernandez-Alonso 1998, 2003a). However,molecular phylogenetic data (Duarte et al. 2011; Carvalho-Sobrinho unpubl. data) do not support segregation of thesegenera, and justify the broad conception of Pachira adoptedhere. These molecular analyses place Pachira in a well-supported clade with Eriotheca Schott. & Endl., a SouthAmerican genus with 24 species. Whereas ITS and trnL-Fdata suggest that Pachira is embedded in a paraphyleticEriotheca, matK data supports the reciprocal monophyly ofthese two genera (Duarte et al. 2011). This suggests thateither Eriotheca should be lumped into Pachira (as proposedby Duarte et al. in review), or that pending a more compre-hensive molecular study with additional taxa, it should besplit into two or more genera.Pachira (inclusive or exclusive of Eriotheca, but excluding

Pachira quinata [Alverson and Duarte in prep.]) is composedof unarmed, small- to medium-sized trees with articulateleaflets; deciduous calyces; glandular receptacles; filamentsorganized in phalanges; and monothecate, oblong anthers.The fruits are usually woody capsules of either one oftwo types: fruits with abundant kapok and small seeds(a few millimeters in diameter), or fruits with sparse kapokand larger seeds (2–6 cm in diameter). The seeds are striateand often angular (Robyns 1963).In Brazil, the genus is represented by 18 species (Duarte

2012), which are mostly evergreen and found in Amazonianwhite-sand forests (“campinas” and “campinaranas”,Anderson 1981) and riparian forests. Until the present, onlyfour Brazilian species were known to occur outside of theAmazonian region: Pachira calophylla (K. Schum.) Fern. Alonso,

P. glabra Pasq., P. retusa (Mart.) Fern. Alonso, andP. endecaphylla (Vell.) Carv.-Sobr. Pachira calophylla is endemicto the Atlantic Forest in southeastern Brazil; P. endecaphylla iswidespread throughout extra-Amazonian semi-deciduousforests (Carvalho-Sobrinho et al. 2013a); P. glabra is oftenfound in cultivation, mostly in eastern Brazil (J. G. Carvalho-Sobrinho, pers. obs.) where it is probably native, although itsoriginal distribution area has been attributed to the Africancontinent (Robyns 1960); and P. retusa is the only speciesknown to occur naturally in caatinga vegetation (Andrade-Lima 1981; APNE 2001; Queiroz et al. 2006), a type ofseasonally dry tropical forest (sensu Pennington et al.2000) in northeastern Brazil. This last species also inhabitsarenaceous-quartzitic rock outcrops in the Espinhaco moun-tain range in the states of Bahia and Minas Gerais.

Ongoing studies of neotropical Bombacoideae, includinganalysis of collections in herbaria and in-situ monitoringof populations, has resulted in the discovery of taxonomicnovelties from Brazil (Carvalho-Sobrinho and Queiroz 2008,2010; Carvalho-Sobrinho et al. 2012; Carvalho-Sobrinho 2013;Carvalho-Sobrinho et al. 2013a) and Bolivia (Carvalho-Sobrinho et al. 2013b), and revealed the importance of inten-sive field work in advancing the systematics of Bombacoideae.Here we describe a new species of Pachira from a small patchof caatinga in the state of Bahia. This new species is describedand illustrated, with notes on its distribution, ecology, phe-nology, phylogeny, and conservation status.

Materials and Methods

This study was based on an analysis of specimens of Pachira s.l. fromthe following herbaria: ALCB, BHCB, CEN, CEPEC, CVRD, F, HAMAB,HRB, HUEFS, IAC, IBGE, INPA, MBM, MBML, NY, R, RB, SJRP, SP, SPF,UB, UEC, US, VIC, and WIS (Thiers 2013). The distribution map wasprepared using ArcMap GIS 9.3 (ESRI 2008).

We used a scanning electron microscope to examine the abaxial sur-face of the leaflets and the external surface of the floral calyces. Materialwas not pre-treated, and was stub-mounted and sputter-coated with gold(Balzers SCD 050). It then was examined and photographed with a JEOL6390LV at Fundacao Oswaldo Cruz in the city of Salvador, Bahia.

For phylogenetic analysis, DNA was extracted from the newspecies and Pachira retusa, and the two new ITS sequences generated

260

(GenBank numbers KF477294 and KF477293, respectively) were alignedby eye to a published data matrix for Bombacoideae (Duarte et al. 2011).All characters were equally weighted, alignment gaps were treated asmissing data, and Rhodognaphalon schumannianum A. Robyns was usedas the outgroup.

Maximum parsimony analyses were performed in PAUP* 4.0b10(Swofford 2002) using heuristic searches (10,000 replicates with randomaddition) with TBR branch swapping and saving 20 trees per replicate.Bootstrap support (BS) was estimated using 2,000 replicates with 10 simpleaddition sequences using TBR, keeping up to 20 trees per replication.

Bayesian analyses were implemented in MrBayes v.3.1.2 (Huelsenbeckand Ronquist 2001). The best available substitution model was determinedas nst = 6 = gamma in MrModeltest 2.3 (Nylander 2004). Two independentMCMC runs were conducted, each composed of four chains that ran for3,000,000 generations with sampling every 1,000 generations. To calculateposterior probabilities (PP), trees sampled during the burn-in stage of eachrun were discarded, and the remaining trees were used to calculate a 50%majority rule consensus tree. The phylogenetic data matrix is available inNexus format from TreeBASE (study number S14561).

Taxonomic Treatment

Pachira moreirae Carv.-Sobr. & W. S. Alverson, sp. nov. —TYPE: BRAZIL. Bahia, Anage, ca. 5 km na estradapara a barragem, 14�38015.100S, 41�10006.500W, 402 m,12 March 2011 (lf), J. G. de Carvalho-Sobrinho &M. Machado 2963 (holotype: HUEFS; isotypes: RB, SP).

Similar to Pachira glabra Pasq. and P. retusa (Mart.) Fern.Alonso in having mucronate leaflets, the androeciumwith ca.250 stamens, and large seeds (ca. 20 mm long), but differingfrom the former by its obovate to obcordate (vs. oblong toelliptic) leaflets that are truncate to emarginate (vs. acute),shorter petals (120 mm vs. 145–200 mm long); from thelatter, it differs mainly by its longer petioles (100–115 vs.up to 40 mm long), wider petals (15 vs. 9 mm width), andthe presence of stellate trichomes on leaflets and calyces(vs. absence of stellate trichomes).

Trees, to 4 m tall, deciduous when flowering; branchesglabrous. Leaves palmately compound, (5–)6–7-foliolate;petioles 100–115 mm long, furrowed, pulvinate at both ends(distinctly swollen when fresh); leaflets articulate, caducous,pulvinate, sessile, papyraceous, brochidodromous, midribprominent abaxially, secondary veins 8–12 per side, incon-spicuous, with golden stellate trichomes on both surfaces,obovate to obcordate, the apex truncate to emarginate,mucronate, mucron to 4 mm long, the base cuneate to atten-uate, sometimes asymmetric on basal leaflets, margin entire,basal leaflets 70–104 +36–55 mm, distal leaflets 105–120 +50–70 mm, length-to-width ratio (L/W) 1.5–2.4, glabrescentwith trichomes mainly on midrib and margins; stomataanomocytic, ca. 20 mm long, numerous on abaxial surface.Flowers axillary, solitary; pedicels 10 mm long, dark greenwhen fresh, bracteoles caducous; flower buds linear-oblong.Flowers 120 mm long, receptacles 4 mm long, dark greenwhen fresh with a whorl of five orange glands on outersurface; calyces 15 + 13 mm, cupuliform to campanulate,L/W 0.7–1.1, truncate, 5-apiculate, caducous before fruitdehiscence, densely covered by stellate trichomes without,yellowish green when fresh, brownish when dried, on innersurface tomentose distally and sericeous on the basal half;petals 120 + 15 mm, green outwardly and pale greeninwardly when fresh, linear, apex rounded, tomentose withstellate and simple trichomes on both surfaces; stamens 250–300, staminal tube 25–36 mm long, glabrous, originating10 phalanges each ca. 10 mm long below free filaments

45–55 mm long, anthers 3 mm long, dorsifixed, oblong;ovary 4 mm long, subglobose, sparsely sericeous (trichomessimple), style velutinous, proximal one-third with stellatetrichomes, stigma not seen. Capsules (60) 80–120 + 80–130 mm, woody, globose, sometimes apiculate, outwardlyglabrous, green, with sparse, whitish-brown to straw coloredkapok, the columella entire, marcescent. Seeds 20–24 mm,angulate, glabrous, striate, pale brown. Figures 1–5.

Additional Specimens Examined—BRAZIL. Bahia: Mun. Anage,estrada para a barragem, 14�38015.100S, 41�10006.500W, 402 m, 20 May 2011(fr), J. G. de Carvalho-Sobrinho & M. Machado 3091 (HUEFS); 14�38015.100S,41�10006.500W, 402 m, 16 July 2011 (lf, fr), J. G. de Carvalho-Sobrinho 3180(HUEFS); 14�38015.100S, 41�10006.500W, 402 m, 13 Oct 2012 (fl), D. Cardoso &M. Machado 3256 (HUEFS).

Etymology—The specific epithet honors Dr. MarceloMoreira from Universidade Estadual do Sudoeste da Bahia,Vitoria da Conquista, for drawing the attention of one of us(MCM) to the area of caatinga in the municipality of Anage,in the state of Bahia, where we have found this new speciesof Pachira and a new species of Spirotheca Ulbrich (Carvalho-Sobrinho et al. 2012), and for providing material for DNAanalysis from several species of Bombacoideae cultivated inhis arboretum.Habitat, Distribution, and Phenology—Pachira moreirae

appears to be endemic to caatinga vegetation in the state ofBahia, in clay soils at ca. 400 m in altitude. Caatinga harbors ahigh diversity of Bombacoideae, including several endemics,e.g. Ceiba glaziovii (Kuntze) K. Schum. (Gibbs and Semir 2003),C. rubriflora Carv.-Sobr. & L. P. Queiroz, Pseudobombax calcicolaCarv.-Sobr. & L. P. Queiroz, Pseudobombax parvifolium Carv.-Sobr. & L. P. Queiroz, Pseudobombax simplicifolium A. Robyns,and Spirotheca elegans Carv.-Sobr. et al. (Carvalho-Sobrinhoand Queiroz 2008; Carvalho-Sobrinho and Queiroz 2010;Carvalho-Sobrinho et al. 2012). Evidently, Pachira moreirae isnever locally sympatric with Pachira retusa (Fig. 1).Monitoring of the population at the type locality from

September 2010 to October 2012 allowed us to obtain collec-tions in various phenological stages. These had leaves inMarch 2011, fruits and leaves from May to July 2011, andcopious numbers of flower buds and some initial flowers(but no leaves) in October 2012. The species appears to syn-chronize its flowering into a short period of time (cf. big-bangphenology, Gentry 1974).Conservation Status—Pachira moreirae is known only from

small patches of suitable habitat at the type locality, which isnot located in a protected area. These patches are quicklydisappearing due to logging as well as forest clearingfor agriculture and cattle. Thus, P. moreirae should be consid-ered to be critically endangered (CR) according to criteriaB1ab(ii, iii)+2ab(ii,iii) of the IUCN red list (IUCN 2010).Micromorphology—The abaxial surface of the leaflets of

Pachira moreirae exhibits a thick, rugose cuticle with supra-stomatal chambers over numerous anomocytic stomata asdo P. glabra and P. retusa (Fig. 4). Other neotropicalBombacoideae in seasonally dry tropical forests have similarfeatures, e.g. Eriotheca roseorum (Cuatrec.) A. Robyns, and dif-fer strikingly from wet forest species such as E. longipedicellata(Ducke) A. Robyns and E. surinamensis (Uittien) A. Robynsthat have a smooth, apparently thin cuticle, and fewer sto-mata (Duarte 2010: 96).Phylogenetic Relationships—Phylogenetic analysis of ITS

sequences using either parsimony or Bayesian inferenceplaced Pachira moreirae within the Pachira s. l. clade of Duarteet al. (2011), nested in a well-supported clade that also

2014] CARVALHO-SOBRINHO ET AL.: NEW DECIDUOUS SPECIES OF PACHIRA 261

includes two other Brazilian extra-Amazonian species ofPachira, P. glabra and P. retusa (PP = 0.99, BS = 88%; Fig. 5).Pachira moreirae is especially close to P. glabra, differingfrom that species by one confirmed 3-bp indel and one sub-stitution. It shares with P. glabra and P. retusa mucronateleaflets, an androecium with ca. 250 stamens, globose, gla-brous capsules, and seeds ca. 20 mm long.

The three species in this Brazilian extra-Amazonian cladeusually have globose capsules that are externally glabrousand contain relatively large seeds with modest amounts ofkapok. Although we could not obtain satisfactory ITSsequences from additional Brazilian extra-Amazonian species,one of these species, P. endecaphylla, appears as sister to aclade with P. glabra, P. moreirae, and P. retusa in an analysis

Fig. 1. Distribution map of the five Brazilian extra-Amazonian species of Pachira including the new species P. moreirae.

262 SYSTEMATIC BOTANY [Volume 39

Fig. 2. Pachira moreirae. A. Branch with leaves. B. The swollen petiole apex and articulate, pulvinate leaflets. C. Flower at anthesis. D. Staminal tubewith phalanges. E. Ovary and style. F. Immature fruit. G. Detached valve and striate seed surrounded by kapok. A–B. Carvalho-Sobrinho & Machado 2963.C–E. Cardoso & Machado 3256. F–G. Carvalho-Sobrinho 3180.

2014] CARVALHO-SOBRINHO ET AL.: NEW DECIDUOUS SPECIES OF PACHIRA 263

Fig. 3. Photos of Pachira moreirae. A. Leaf 6-foliolate. B. Flower at anthesis. C. The swollen petiole apex and articulate, pulvinate leaflets. D. Calyxand receptacle with glands. E. Young 5-foliolate leaf with a tomentose indumentum. F. Fruits before dehiscence and caducous leaflets. G. Detachedvalve and striate seed surrounded by kapok. Bars: A: 2 cm, B–E: 1 cm. A, C. Carvalho-Sobrinho & Machado 2963. B, D, E. Cardoso & Machado 3256. F, G.Carvalho-Sobrinho 3180.

264 SYSTEMATIC BOTANY [Volume 39

of nuclear ribosomal DNA data from the external transcriberspacer (ETS) region (Carvalho-Sobrinho et al. unpubl. data).

Taxonomic Discussion—Pachira moreirae is morphologi-cally distinct within the Brazilian extra-Amazonian clade.From P. glabra, it can be distinguished by its obovate toobcordate leaflets (vs. oblong to elliptic) that are truncate toemarginate (vs. acute), its shorter petals (120 mm vs. 145–200 mm long), and deciduous foliage when flowering(vs. evergreen). From P. retusa, the only other caatingaspecies, it can be distinguished by its (5–)6–7-foliolate leaves

(vs. 3–4[–5]-foliolate leaves), which are densely tomentose(vs. glabrous) when young (Fig. 3D), its longer petioles(100–115 vs. up to 40 mm long), a petiole apex that isstrongly swollen (vs. slightly swollen), a cupuliform to cam-panulate calyx with a L/W of 0.7–1.1 (vs. tubuliform andL/W of 1.6), wider petals (15 vs. 9 mm width), andthe presence of stellate trichomes on leaflets and calyces(vs. absence of stellate trichomes; Fig. 4). It is also unique inits entire (vs. inconspicuously crenulate) leaflets that arewider distally (50–70 vs. 13–38 mm). Finally, P. moreirae is

Fig. 4. Micromorphology of Pachira moreirae and allies. Pachira moreirae, A–C. A. Abaxial surface of the leaflet with stellate trichomes. B. Detail ofstomata on abaxial surface of the leaflet. C. External surface of the calyx with stellate trichomes. Pachira retusa, D. Detail of stomata on abaxial surfaceof the leaflet. Pachira glabra, E–F. E. Abaxial surface of the leaflet lacking trichomes. F. Detail of stomata on abaxial surface of the leaflet. [A–B fromCarvalho-Sobrinho 3180; C from Cardoso 3256; D from Queiroz 7629; E–F from Carvalho-Sobrinho 2900]. Bars: A, C, E: 100 mm; B, D, F: 20 mm.

2014] CARVALHO-SOBRINHO ET AL.: NEW DECIDUOUS SPECIES OF PACHIRA 265

deciduous and flowers when leafless during a short periodat the end of the dry season (October), in contrast withP. retusa, which is evergreen and flowers throughout theyear (J. G. Carvalho-Sobrinho pers. obs.), despite occurringin the same caatinga biome. These phenological differences

suggest ecological or edaphic specialization that, in combi-nation with the several morphological traits, support therecognition of P. moreirae and P. retusa as distinct entitiesat species rank. The following key can be used to separatethe five Brazilian extra-Amazonian species of Pachira:

1. Leaflets petiolulate, petiolule 10–25 mm long, cuspidate; pedicel 25–45 mm long; staminal tube 15–20 mm long,stamens 350–400 (wet forests in the southeastern Atlantic Forest) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. calophylla

1. Leaflets sessile, acute, truncate to emarginated; pedicel 10–20(–25) mm long; staminal tube 25–95 mm long,stamens 150–300 (seasonally dry tropical forests) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. Leaves 7–11-foliolate, leaflets to 30 mm wide; staminal tube 70–95 mm long; kapok grayish, abundant;seeds more than 150, ca. 6 mm diam. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. endecaphylla

2. Leaves 3–6(–7)-foliolate, leaflets 36–70 mm wide; staminal tube 25–47 mm long; kapok whitish-brown to straw colored,sparse; seeds 10–30, ca. 20 mm diam. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3. Leaflets oblong to elliptic, acute; petals 145–200 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. glabra3. Leaflets obovate to obcordate, truncate to emarginate; petals 65–120 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. Plants evergreen while flowering; brachyblasts often present; leaflets inconspicuously crenulate,13–38 mm wide distally; petiole to 40 mm long, apex slightly swollen; calyx tubuliform,L/W 1.6; petals 6–9 mm wide; leaflets and calyces lacking stellate trichomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. retusa

4. Plants leafless while flowering; brachyblasts absent; leaflets entire, 50–70 mm distally, petiole 100–115 mm long,apex strongly swollen; calyx cupuliform to campanulate, L/W 0.7–1.1; petals 15 mm wide;leaflets and calyces with stellate trichomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. moreirae

Fig. 5. Phylogram showing the phylogenetic placement of the newly described Pachira moreirae in the Pachira s.l. clade based on sequence data fornuclear marker ITS. All nodes indicated have >50% bootstrap support. Numbers above the branches represent posterior probability (PP) and numbersbelow these branches are bootstrap values (BS).

266 SYSTEMATIC BOTANY [Volume 39

Acknowledgments. We thank Petala Gomes Ribeiro for her linedrawings, Dr. Domingos Cardoso for climbing individuals of the newspecies to collect flowers (with MCM), Dr. Luciano Paganucci de Queiroz,curator of the herbarium HUEFS, Dr. Cassio van den Berg and Dr. ReyjanePatrıcia Oliveira, coordinators of the Laboratory of Plant Systematics(LAMOL/UEFS), and the Fundacao Osvaldo Cruz (FIOCRUZ) for theuse of the scanning electron microscope. Fieldwork was supportedby the Graduate Program in Botany (PPGBot/UEFS) and FAPESB(APP0006/2011). JGCS was supported by doctoral fellowships from theCoordenacao de Aperfeicoamento de Pessoal de Nıvel Superior (CAPESprocess BEX 5415/13-6"); ACM and MCM were supported by doctoratefellowships from the Conselho Nacional de Desenvolvimento Cientıficoe Tecnologico (CNPq).

Literature Cited

Alverson, W. S. 1994. New species and combinations of Catostemma andPachira (Bombacaceae) from the Venezuelan Guayana. Novon 4: 3–8.

Alverson, W. S. and J. A. Steyermark. 1997. Bombacaceae. Pp. 496–527. inFlora of the Venezuelan Guayana. Vol. 3. eds. P. E. Berry, B. K. Holst,and K. Yatskievych. St. Louis: Missouri Botanical Garden Press.

Alverson, W. S., B. A. Whitlock, R. Nyffeler, C. Bayer, and D. A. Baum.1999. Phylogeny of the core Malvales: evidence from ndhF sequencedata. American Journal of Botany 86: 1474–1486.

Anderson, A. B. 1981. White-sand vegetation of Brazilian Amazonia.Biotropica 13: 199–210.

Andrade-Lima, D. 1981. The caatingas dominium. Revista Brasileira deBotanica 4: 149–153.

APNE. 2001. Seminario de Planejamento Ecorregional da Caatinga,1ª etapa. (http://www.plantasdonordeste.org/Livro/sumario.htm).[accessed 03/02/2013].

Baum, D. A., S. D. Smith, A. Yen, W. Alverson, R. Nyffeler, B. A. Whitlock,and R. L. Oldham. 2004. Phylogenetic relationships of Malvatheca(Bombacoideae and Malvoideae; Malvaceae sensu lato) asinferred from plastid DNA sequences. American Journal of Botany91: 1863–1871.

Bayer, C., M. Fay, A. de Bruijn, V. Savolainen, C. Morton, K. Kubitzki,W. Alverson, and M. Chase. 1999. Support for an expanded familyconcept of Malvaceae within a recircumscribed order Malvales: acombined analysis of plastid atpB and rbcL DNA sequences. BotanicalJournal of the Linnean Society 129: 267–303.

Carvalho-Sobrinho, J. G. 2013. A new species of Eriotheca (Malvaceae:Bombacoideae) from Espırito Santo, eastern Brazil. Phytotaxa108: 49–53.

Carvalho-Sobrinho, J. G. and L. P. Queiroz. 2008. Ceiba rubriflora: a newspecies from Bahia, Brazil. Kew Bulletin 63: 649–653.

Carvalho-Sobrinho, J. G. and L. P. Queiroz. 2010. Three new speciesof Pseudobombax (Malvaceae, Bombacoideae) from Brazil. Novon20: 13–20.

Carvalho-Sobrinho, J. G., M. C. Machado, and L. P. Queiroz. 2012.Spirotheca elegans (Malvaceae: Bombacoideae), a new species fromBahia, Brazil. Systematic Botany 37: 978–982.

Carvalho-Sobrinho, J. G., L. P. Queiroz, and L. J. Dorr. 2013a. DoesPseudobombax have prickles? Assessing the enigmatic speciesPseudobombax endecaphyllum (Malvaceae: Bombacoideae). Taxon62: 814–818.

Carvalho-Sobrinho, J. G., L. P. Queiroz, and W. S. Alverson. 2013b.Reinterpretation of the nomenclatural type of Pseudobombaxheteromorphum (Malvaceae: Bombacoideae) reveals an overlookednew species from Bolivia. PhytoKeys 21: 53–61.

Duarte, M. C. 2010. Analise filogenetica de Eriotheca Schott. & Endl. e generosafins (Bombacoideae, Malvaceae) e estudo taxonomico de Eriothecano Brasil. Ph. D. thesis. Sao Paulo: Instituto de Botanica de SaoPaulo. (http://capesdw.capes.gov.br/capesdw/resumo.html?idtese=20102533120013001P4). [accessed 08/02/2013].

Duarte, M. C. 2012. Pachira in Lista de Especies da Flora do Brasil. JardimBotanico do Rio de Janeiro. (http://floradobrasil.jbrj.gov.br/2012/FB023584). [accessed 18/12/2012].

Duarte, M. C., G. L. Esteves, M. L. F. Salatino, K. C. Walsh, and D. A.Baum. 2011. Phylogenetic analyses of Eriotheca and relatedgenera (Bombacoideae, Malvaceae). Systematic Botany 36: 690–701.

ESRI. 2008. ArcMap GIS, version 9.3. Redlands, California: EnvironmentalSystems Research Institute.

Fernandez-Alonso, J. L. 1998. Novedades taxonomicas, nomenclaturalesy corologicas en el genero Pachira Aubl. (Bombacaceae). Anales delJardin Botanico de Madrid 56: 305–314.

Fernandez-Alonso, J. L. 2003a. Bombacaceae neotropicae novae vel minuscognitae VI. Novedades en los generos Cavanillesia, Eriotheca, Matisiay Pachira. Revista de la Academia Colombiana de Ciencias Exactas. Fısicasy Naturales 27: 25–37.

Fernandez-Alonso, J. L. 2003b. Bombacaceae neotropicae novae vel minuscognitae VII. Redescubrimiento de Pachira pulchra Planch. & Lind.ex Triana & Planch. Caldasia 25: 65–71.

Fernandez-Alonso, J. L. 2003c. Bombacaceae neotropicae novae vel minuscognitae VIII. Una nueva e inusual especie de Pachira Aubl. deColombia. Anales del Jardin Botanico de Madrid 60: 220–222.

Gentry, H. A. 1974. Flowering phenology and diversity in tropicalBignoniaceae. Biotropica 6: 64–68.

Gibbs, P. and J. Semir. 2003. A taxonomic revision of the genusCeiba Mill. (Bombacaceae). Anales del Jardin Botanico de Madrid60: 259–300.

Huelsenbeck, J. P. and F. Ronquist. 2001. MRBAYES: Bayesian inferenceof phylogenetic trees. Bioinformatics 17: 754–755.

IUCN. 2010. Guidelines for using the IUCN red list categories andcriteria. Version 8.1. Prepared by the Standards and Petitions Sub-committee in March 2010. (http://intranet.iucn.org/webfiles/doc/SSC/RedList/RedListGuidelines.pdf). [accessed 09/10/2011].

Nylander, J. A. A. 2004. MrModeltest, v.2.3. Program distributedby the author. Uppsala, Sweden: Evolutionary Biology Centre,Uppsala University.

Pennington, R. T., D. E. Prado, and C. A. Pendry. 2000. Neotropicalseasonally dry forests and Quaternary vegetation changes. Journal ofBiogeography 27: 261–273.

Queiroz, L. P., A. A. Conceicao, and A. M. Giulietti. 2006. Nordeste semi-arido: caracterizacao geral e lista das fanerogamas. Pp. 15–359.in Diversidade e caracterizacao das fanerogamas do semi-arido brasileiro.Vol. 1. eds. A. M. Giulietti and L. P. de Queiroz. Recife: AssociacaoPlantas do Nordeste.

Robyns, A. 1960. Contribution a 1’etude monographique du genreBombax s. l. - I. B. glabra (Pasq.) A.Robyns, comb. nov. Bulletin duJardin Botanique de l’Etat a Bruxelles 30: 473–484.

Robyns, A. 1963. Essai de monographie du genre Bombax L. s. l.(Bombacaceae). Bulletin du Jardin Botanique de l’Etat a Bruxelles33: 1–315.

Swofford, D. L. 2002. PAUP* phylogenetic analysis using parsimony(*and other methods). version 4.10b. Sunderland: Sinauer Associates.

Thiers, B. 2013. Index Herbariorum: A global directory of public herbariaand associated staff. New York Botanical Garden’s Virtual Herbarium.(http://sweetgum.nybg.org/ih/). [accessed 07.02.2013].

Tropicos. 2012. Tropicos.org. Missouri Botanical Garden. Available at(http://www.tropicos.org/Name/3701652). [accessed 18.10.2012].

2014] CARVALHO-SOBRINHO ET AL.: NEW DECIDUOUS SPECIES OF PACHIRA 267