Neotropical Malvaceae (Bombacoideae)

Bente B. Klitgård

Royal Botanic Gardens, Kew, U.K.

Description

Malvaceae s.l.

Habit: shrubs, trees (herbs).  Leaves alternate or two-ranked, stipulate; leaf margin toothed or entire; venation palmate or 3-nerved.  Inflorescences made up of cymose units (bicolor units, named after Theobroma bicolor where it was first observed).  Flowers with epicalyx present or absent; sepal aestivation valvate; androgynophore present or absent; stamens 5-many, in five groups (fundamentally obdiplostemonous).  Fruit a capsule, berry, schizocarp.

Subfamily Bombacoideae (Bombacaceae)

Habit: usually trees, often buttressed and/or with a bottle-shaped trunk, and/or with chunky spines, rarely shrubs or climbersLeaves alternate; palmately lobed or palmate, rarely simple (Matisia and Quararibea); venation palmate or 3-nerved, margins entire, rarely dentate; stipules present, often falling early; indumentum, if present, stellate or rarely lepidoteInflorescences of few-flowered, axillary clusters or panicles, solitary or rarely in leaf-opposing cymes.  Flowers bisexual, actinomorphic; epicalyx conspicuous, of 2-3 bracts, usually present; calyx tubular, 5-lobed fused for ½ its length; corolla contorted, 5 free petals attached basally to the androecium; stamens 5 to numerous (up to 1,000), filaments fused into a tube or in antepetalous fascicles, anthers usually monothecal, straight, reniform or spirally contorted; ovary superior, (2-)5(-8)-locular; style with short stylar branches.  Fruit a many-seeded, dehiscent, often ligneous capsule with densely hairy endocarp (e.g. kapok) or indehiscent, juicy, 1-5-seeded drupe, rarely wingedSeeds usually glabrous, sometimes arillate or winged.

Notes on delimitation

The past 15 years have seen an explosion in the number phylogenetic and taxonomic studies of the core Malvales clade (e.g. La Duke & Doebley 1995; Alverson et al. 1998, 1999; Bayer et al. 1999; Baum et al. 2004; Nyffeler et al. 2005; Pfeil et al. 2005; García et al. 2009).  These studies tackle the arbitrary and inconsistent limitations among the families previously recognised within the clade.  Recent outcomes of these papers are two reclassifications of the clade.  One is by Bayer & Kubitzki (2002) in which they subsumed the previously recognized families Bombacaceae, Brownlowiaceae, Byttneriaceae, Dombeyaceae, Grewiaceae (~ more or less Sparrmanniaceae), Helicteraceae (~ more or less Durionaceae), Malvaceae s.s., Pentapetaceae, Sterculiaceae and Tiliaceae as subfamilies under Malvaceae s.l.  The other is by Cheek (2007) who recognised ten families inside the core Malvaceae clade.  The APG III system (2011) adopted the subfamily approach which is also followed in this treatment.  In Latin America there are representatives of the following Malvaceae subfamilies:  Bombacoideae, Brownlowioideae, Byttnerioideae, Grewioideae, Helicteroideae, Malvoideae, Sterculioideae, and Tilioideae.

Bombacoideae (alternatively Bombacaceae) is unchanged, except the exclusion of the Asian tribe Durioneae in the Durionaceae or Helicteroideae, and the transfer of Fremontodendron Coville and Pentaplaris L.O.Williams & Standl. from Sterculiaceae and Tiliaceae, tribe Brownlowieae, respectively.  In recent phylogenetic studies, the Malvoideae is resolved as sister to the Bombacoideae with some intermediate genera (e.g. Pentaplaris and Matisia).

Distribution in the Neotropics

The family Malvaceae s.l. includes 243 genera and about 4,225 species which are largely tropical and temperate.  Of these 129 genera and 1,900-2,200 are native to the Neotropics.

Subfamily Bombacoideae is mostly present in the New World tropics and contains 27 genera (c. 250 species); a few of those genera (Adansonia L., Bombax L., Camptostemon Mast., and Lagunaria (DC.) Rchb.), containing c. 19 species, are restricted to the Old World tropics.

  • Aguiaria Ducke:  monospecific (A. excelsa Ducke) restricted to the Brazilian Amazon.
  • Bernoullia Oliv.:  two-three species from Mexico to Colombia.
  • Catostemma Benth.:  about 10-15 species from northern South America.
  • Cavanillesia Ruiz & Pav.:  three-four species from Panama to Brazil in the East and Peru in the West.
  • Ceiba Mill.:  about 12-20 spp. from tropical America.
  • Chiranthodendron Sessé ex Larreat.:  monospecific (C. pentadactylon Larreat.) from Mexico and Guatemala.
  • Eriotheca Schott & Endl.:  about 20 species in the Neotropics.
  • Fremontodendron Coville:  two-three species from NW Mexico to adjacent SW USA.
  • Gyranthera Pittier:  two species, G. dariensis Pittier from Panama and G. caribensis Pittier from Venezuela.
  • Huberodendron Ducke:  four-five species from Costa Rica extending South to Brazil in the East and Bolivia in the West.
  • Matisia Bonpl.:  about 30 species in tropical America.
  • Neobuchia Urb.:  monospecific (N. paulinae Urb.) restricted to Haiti.
  • Ochroma Sw.:  monospecific (O. pyramidale (Cav. ex Lam.) Urb.) from tropical America.
  • Pachira Aubl.:  about 50 species (including Bombacopsis Pittier recently synonymized under Pachira), mostly neotropical, but with six species in tropical Africa.
  • Patinoa Cuatrec.:  four species from Panama through Colombia to Brazil and Peru.
  • Pentaplaris L.O.Williams & Standl.:  three species from Costa Rica, Ecuador, Bolivia, and Peru.
  • Phragmotheca Cuatrec.:  eleven species from Panama to Peru.
  • Pseudobombax Dugand:  about 20 species from Mexico, through Central America to South America.
  • Quararibea Aubl.:  neotropical, more than 20 species.
  • Scleronema Benth.:  five species from Venezuela, Guyana and Brazil.
  • Septotheca Ulbr.:  monospecific, S. tessmannii Ulbr. from Peru, Colombia and Brazil.
  • Spirotheca Ulbr.:  nine species from Panama to Peru in the West and Brazil in the East.
  • Uladendron Marc.-Berti:  monospecific, U. codesuri Marc.-Berti from Venezuela.

Distinguishing characters (always present)

  • Stipules present, but often caducous.
  • Epicalyx present, of 2-3 bracts.
  • Petal bases adnate to the stamen tube or filaments.
  • Androgynophore absent.
  • Stamens 5-1000, in a tube or fascicles.
  • Anthers monothecal, straight, reniform or spirally contorted.

Other important characters

Key differences from similar families

The families and subfamilies listed below differ from the Malvaceae subfamily Bombacoideae as follows:

  • Araliaceae - if trees, then not bottle-shaped nor with stout prickles; stipules absent; petioles often sheating the base of the leaf; flowers in umbellate units; stamens equal in number to the petals and free, anthers dithecal; fruits usually fleshy (rarely dry) drupes or berries, never with kapok.
  • Brownlowioideae - sepals fused into a campanulate or urceolate tube; an androgynophore is always absent.
  • Malvoideae - mostly shrubs and herbs without stout, bottle-shaped and/or prickled trunks.
  • Byttnerioideae - sepals free; epicalyx always absent; petals cupped or hooded with strap-like appendages, free from the androecium; style unbranched.
  • Cochlospermaceae - anthers dehiscing via pores.
  • Convolvulaceae - stipules absent; leaves pinnately veined; stamens 5.
  • Grewioideae - without stout, bottle-shaped or prickled trunk; epicalyx always absent; sepals free; petals usually yellow or white, often with hairy basal nectaries; stamens free, rarely fasciculate.
  • Helicteroideae - calyx tubular; petals free and clawed; androgynophore usually present; stamens 10-30; ovaries usually apocarpous, except Ungeria and Reevesia.
  • Sterculioideae - petals always absent; androgynophore usually present; the stamen filaments free; ovaries apocarpous.

Useful tips for generic identification

Key to the genera of Neotropical Malvaceae-Bombacoideae (modified from Bayer & Kubitzki, 2003)

1.  Leaves digitately compound (simple in Pseudobombax simplicifolium) … 2
1.  Leaves simple (3-5-foliolate in Catostemma digitatum) … 9

2.  Seeds winged, not embedded in kapok … 3
2.  Seeds not winged, embedded in kapok or endocarp velutinous … 4

3.  Anthers spirally twisted … Gyranthera
3.  Anthers not spirally twisted … Bernoullia

4.  Stamens 5-15 … 5
4.  Stamens more numerous … 7

5.  Trees; anthers 2(-3)-thecate … Ceiba
5.  Epiphytic stranglers or trees; anthers 4-thecate … 6

6.  Epiphytic stranglers; flowers reddish, South America … Spirotheca
6.  Tree; flowers whitish; Haiti … Neobuchtia

7.  Leaflets without basal suture … Pseudobombax
7.  Leaflets with basal suture … 8

8.  Flowers 1.5-5.5 cm long … Eriotheca
8.  Flowers 7-35 cm long … Pachira

9.  Fruits dehiscent … 10
9.  Fruits indehiscent, or with dehiscent outer and indehiscent inner spongy layer … 18

10.  Seeds winged … 11
10.  Seeds not winged … 18

11.  Leaves lobed, margin serrate; ovules 2 per loculeUladendron
11.  Leaves not lobed, margin entire or dentate; ovules several per locule … 12

12.  Leaves cordate, venation clearly palmate; trichomes lepidoteSeptotheca
12.  Leaves elliptic to ovate, venation pinnate or faintly palmate; trichomes stellateHuberodendron

13.  Sepals petaloid, keeled, imbricate; petals absent … 14
13.  Sepals not petaloid, not keeled, valvate; petals present … 15

14.  Shrubs; flowers yellow to orange; stamens held in a tubular staminal column … Fremontodendron
14.  Trees; flowers dark reddish; stamens held in a unilateral sheathChiranthodendron

15.  Leaf venation clearly palmate; fruits with copious kapok … Ochroma
15.  Leaf venation pinnate or faintly palmate; fruits without kapok … 16

16.  Ovary 5-locular; fruits with dehiscent outer and indehiscent, spongy inner layer; restricted to Brazilian Amazon … Aguiaria
16.  Ovary 2-3(-4)-locular; fruits indehiscent or tardily dehiscent … 17

17.  Staminal filaments thickened at apex; ovary 2-3(-4)-locular … Scleronema
17.  Staminal filaments not thickened at apex; ovary 3-locular … Catostemma

18.  Fruits winged … 19
18.  Fruits not winged … 20

19.  Ovary (3-)5-locular; wings formed by pericarpCavanillesia
19.  Ovary 2-locular; wings formed by accrescent sepals … Pentaplaris

20.  Locules many-ovulate; fruits with numerous, pubescent seeds … Patinoa
20.  Locules usually 2-ovulate; Fruits with 5 or less, glabrous seeds … 21

21.  Staminal tube short, filaments fused for less than half of their length … Scleronema
21.  Staminal filaments forming a long, apically lobed tube … 22

22.  Ovary 2-4-locular; lobes of staminal tube 4 mm or shorter … Quararibea
22.  Ovary 5-locular; lobes of staminal tube longer than 4 mm … 23

23.  Thecae septate; indumentum lepidotePhragmotheca
23.  Thecae not septate; indumentum never lepidoteMatisia

Status

Native.

General notes

  • Ceiba pentandra (L.) Gaertn. - economically important kapok producer; introduced into and cultivated in the Old World.
  • Chiranthodendron pentadactylon (the Devil's Hand Tree) has great potential as ornamental (see  http://www.strangewonderfulthings.com).
  • Ochroma pyramidale (Cav. ex Lam.)Urb. yields the extremely light balsa wood.
  • Phragmotheca, Matisia, Quararibea - the fruits of some species are fleshy, aromatic and edible.
  • Patinoa spp. - the pulpy fruits are edible or used locally as fish poison.

Important literature

Alverson, W.S., Karol, K.G., Baum, D.A., Chase, M.W., Swensen, S.M., McCourt, R. & Systma, K.J. 1998.  Circumscription of the Malvales and relationships to other Rosidae: Evidence from rbcL sequence data. American J. Bot. 85: 876-887.

Alverson, W.S. 1999.  Bombacaceae. In: Jørgensen, P.M. & León-Yánez, S. (eds.). Catalogue of the Vascular Plants of Ecuador, pp. 326-329. Missouri Botanical Garden Press, St. Louis.

Alverson, W.S. 2004.  Bombacaceae. In: Smith, N., Mori, S.A., Henderson, A., Stevenson, D.W. & Heald, S.V. (eds.). Flowering Plants of the Neotropics, pp. 55-58. Princeton University Press, Princeton.

Alverson, W.S. & Steyermark, J.A. 1997.  Bombacaceae.  In: Berry, P.E., Holst, B.K. & Yatskievych, K. (eds.).  Flora of the Venezuelan Guyana vol. 3, pp. 496-527.  Missouri Botanical Garden, St. Louis.

Alverson, W.S., Karol, K.G., Baum, D.A., Chase, M.W., Swensen, S.M., McCourt, R. & Systma, K.J. 1998.  Circumscription of the Malvales and relationships to other Rosidae: Evidence from rbcL sequence data. American J. Bot. 85: 876-887.

Alverson, W.S., Whitlock, B.A., Nyffeler, R., Bayer, C. & Baum, D.A. 1999.  Phylogeny of core Malvales: Evidence from ndhF sequence data. American J. Bot. 86: 1474-1486.

Alverson, W.S. & Mori, S.A 2002.  Bombacaceae. In: S.A. Mori, G. Cremers, C.A. Gracie, J.-J. de Granville, S.V. Heald, M. Hoff & J.D. Mitchell (eds.). Guide to the Vascular Plants of Central French Guiana, pp. 139-145.  Memoirs of the New York Botanical Garden 76, part 2.  New York Botanical Garden Press.

Baum, D.A., Smith, S.D., Yen, A., Alverson, W.S., Nyffeler, R., Whitlock, B.A., & Oldham, R.L. 2004.  Phylogenetic relationships of Malvatheca (Bombacoideae and Malvoideae; Malvaceae sensu lato) as inferred from plastid DNA sequences. American J. Bot. 91: 1863-1871.

Bayer, C. 1999.  The bicolor unit - homology and transformation of an inflorescence structure unique to core Malvales. Plant Syst. Evol. 214: 187-198.

Bayer, C. & Kubitzki, K. 2002.  Malvaceae: subfamily Bombacoideae. In: Kubitzki, K. & Bayer, C. (eds.). The Families and Genera of Vascular Plants vol. V, pp. 277-311. Springer-Verlag, Berlin.

Bornstein, A.J. 1989. Bombacaceae. In: Howard, R.A. (ed.). Flora of the Lesser Antilles: Leeward and Windward vol. 5, pp. 263-272. Jamaica Plain, Arnold Arboretum, Harvard University.

Bovini, M.G., Esteves, G. & Duarte, M.C. 2010. Malvaceae. In: Forzza, R.C. et al. (eds.). Catálogo de Plantas e Fungos do Brasil, pp. 1201-1227. Institúto de Pesquisas Jardím Botânico do Rio de Janeiro, Rio de Janeiro.

Brako, L. & Alverson, W.S. 1993. Bombacaceae. In: Brako, L. & Zarucchi, J.L. (eds.). Catalogue of the Flowering Plants and Gymnosperms of Peru, pp. 213-216. Monographs in Systematic Botany from the Missouri Botanical Garden vol. 45. St. Louis.

Cheek, M.R. 2007. Malvaceae. In: Heywood, V.H., Brummitt, R.K., Culham, A & Seberg, O. (eds.). Flowering Plant Families of the World, pp. 65. Royal Botanic Gardens Kew, Richmond.

Carranza G., E. & Blanco G., A. 2000.  Bombacaceae. In: Rzedowski, J. & Calderón de Rzedowski, G. Flora del Bajio y de regiones adyacentes fasc. 90, pp. 1-16. Instituto de Ecología, Xalapa.

García, P.E., Schönswetter, P., Aguilar, J.F., Feliner, G.N., & Schneeweiss, G.M. 2009.  Five molecular markers reveal extensive morphological homoplasy and reticulate evolution in the Malva alliance (Malvaceae). Mol. Phyl. Evol. 50: 226-239.

Gentry, A.H. & Alverson, W.S. 2001.  Bombacaceae. In: Stevens, W.D., Ulloa U., C., Pool, A. & Montiel, O.M. (eds.). Flora de Nicaragua tomo I, pp. 430-435. Monographs in Systematic Botany from the Missouri Botanical Garden vol. 85.  St. Louis.

Hinsley, S.R. 2013.  MALVACEAE info.  http://www.malvaceae.info/index.html (accessed 17/01/2013)

Jansen-Jacobs, M.J. 1986. Bombacaceae. In: Stoffers, A.L. & Lindeman, J.C. (eds.). Flora of Suriname III(1-2), pp. 277-282.  E.J. Brill, Leiden.

La Duke, J.C. & Doebley, J. 1995.  A chloroplast DNA based phylogeny of the Malvaceae. Systematic Botany 20: 259-271.

Nyffeler, R., Bayer, S., Alverson, S.A., Yen, A., Whitlock, B.A., Chase, M.W., Baum, D.A. 2005.  Phylogenetic analysis of the Malvadendrina clade (Malvaceae s.l.) based on plastid DNA sequences.  Organisms, Diversity & Evolution 5: 109-123.

Pfeil, B.E. & Crisp, M.D. 2005.  What to do with Hibiscus? A proposed nomenclatural resolution for a large and well known genus of Malvaceae and comments on paraphyly. Australian Syst. Bot. 18: 49-60.

Robyns, A. 1964.  Bombacaceae, family 116. In: Woodson, R.E. & Schery, R.W. (eds.) Flora of Panama. Ann. Missouri Bot. Gard. 51: 37-68.

Stevens, P. F. (2001 onwards). Angiosperm Phylogeny Website. Version 12, July 2012 (visited 18th Jan. 2013). http://www.mobot.org/MOBOT/research/APweb/.

Zuloaga, F.O., Morrone, O. & Belgrano, M.J. 2008.  Bombacaceae. In: Zuloaga, F.O., Morrone, O. & Belgrano, M.J. (eds.). Catálogo de las Plantas Vasculares del Cono Sur vol. 3, pp. 1627-1631. Missouri Botanical Garden Press, St. Louis.

How to cite

Klitgård, B.B. (2013). Neotropical Malvaceae (Bombacoideae). In: Milliken, W., Klitgård, B. & Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics. http://www.kew.org/science/tropamerica/neotropikey/families/Malvaceae_(Bombacoideae).htm.