1. Malvaceae Juss.

    1. This family is accepted.

[FTEA]

Malvaceae, Bernard Verdcourt & Geoffrey Mwachala. Pavonia, B Verdcourt; Kosteletzkya, OJ Blanchard Jr.; Gossypium, P Fryxell & B Verdcourt. Flora of Tropical East Africa. 2009

Habit
Annual or perennial herbs, shrubs or less often trees, usually with stellate or lepidote indumentum, mucilaginous cells etc in the parenchymatous tissues and fibrous stems or bark
Leaves
Leaves alternate, sessile to petiolate, usually stipulate, simple, unlobed to deeply and often palmately divided, the margins entire to serrate; venation often palmate
Flowers
Flowers regular, hermaphrodite or rarely unisexual, solitary or in cymose inflorescences; epicalyx bracts often present; calyx-lobes (3–)5, sometimes united at base, valvate; petals 5, free but often adnate to the base of the staminal tube, imbricate, contorted or convolute, usually obliquely obovate and with a retuse to emarginate apex
Stamens
Stamens 5–numerous, the filaments joined into a tube for most of their length; anthers dehiscing by slits; pollen spinulose
Ovary
Ovary superior with (1–)5–many locules with as many or twice as many style-branches as carpels.
Style
Style-branches partly free or almost completely to completely connate; placentation axile with 1–numerous ovules per locule
Fruits
Fruits mostly loculicidal capsules or schizocarpic, less often indehiscent berries or samaras.
Note
The circumscription of genera in the tribe Hibisceae (e.g., Abelmoschus, Hibiscus, Kosteletzkya, Pavonia, Urena), is based on morphological features that are not corroborated by phylogenetic evidence. Pfeil, Brubaker, Craven & Crisp have published on the phylogeny of this group in  Systematic Botany 27 (2): 333–350 (2002) and came to the conclusion that these groups are not monophyletic. They state that nomenclatural upheavals in and around Hibiscus are almost inevitable. Specifically, Abelmoschus and Fioria (in the FTEA treatment treated as Roifia) cannot be maintained as separate and will have to sink in Hibiscus; or Hibiscus will have to be divided into more than ten genera. In a second article, Pfeil & Crisp discuss ‘What to do with Hibiscus’ (Australian Systematic Botany 18: 49–60, 2005) and confirm inclusion of Abelmoschus and Fioria in Hibiscus, and add Kosteletzkya, Pavonia and Urena to Hibiscus synonymy. Sadly, Dr Verdcourt is seriously ill at the moment and unable to change his treatment accordingly, or to comment on these developments. I (HB) have therefore decided to let the current FTEA treatment stand, rather than implement as yet incomplete, though scientifically convincing, new findings. The editors are grateful to David Mabberley (RBG Kew) and Lyn Craven (Australian National Herbarium, CSIRO) for drawing attention to this recent work and providing references. Lyn Craven also comments: “Now that we have the benefit of DNA technologies we can devise better classifications but there are many groups in which we are not quite there yet, and Malvaceae is one of these (the infrageneric classification of Hibiscus in particular will take some considerable work)”
[FZ]

Malvaceae, A. W. Exell. Flora Zambesiaca 1:2. 1961

Habit
Herbs, shrublets, shrubs or small trees, usually with stellate hairs or bristles, sometimes aculeate, more rarely lepidote
Leaves
Leaves alternate, usually petiolate, often palmately divided
Stipules
Stipules present
Flowers
Flowers bisexual (rarely unisexual), actinomorphic
Epicalyx
Epicalyx present or absent
Calyx
Calyx (3–4) 5-lobed, truncate or occasionally 5- or 10-toothed; lobes valvate
Corolla
Petals 5, free but often slightly adnate to the staminal tube, contorted or imbricate
Androecium
Stamens numerous, united in, a tube surrounding the style; anthers 1-thecous
Gynoecium
Ovary superior, (1) 2-multilocular (carpels rarely in vertical rows); style simple at the base, often branched towards the apex, branches the same number as or twice as many as the carpels; ovules 1-numerous in each loculus; placentation axile
Fruits
Fruit a loculicidally dehiscent capsule or composed of follicles, achenes or pseudo-achenes arranged around a central columella, or indehiscent and woody or fleshy
Seeds
Seeds usually with some endosperm; cotyledons often folded
[NTK]

Klitgård, B.B., Edwards, S.L., Biggs, N. & Frisby, S. (2011). Neotropical Malvaceae (Malvoideae).

Morphology
Description

Malvaceae s.l.

Habit: shrubs, trees (herbs).  Leavesalternate or two-ranked, stipulate; leaf margin toothed or entire; venationpalmate or 3-nerved.  Inflorescences made up of cymose units (bicolor units, named after Theobroma cacao where it was first observed).  Flowers with epicalyx present or absent;  sepalaestivationvalvate;  androgynophore present or absent; stamens 5-many, in five groups (fundamentally obdiplostemonous).  Fruita capsule, berry, or schizocarp.

Subfamily Malvoideae (Malvaceae s.s.)

Habit: herbs, shrubs or trees.  Leavesalternate; simple to dissected, ovate to lanceolate or cordate; venation 3-5-nerved from the base, margins entire, crenate, or serrate; stipules present, often falling early; indumentum usually of stellate hairs or simple, glandular or lepidote hairs; sometimes with extrafloral nectaries.  Flowers usually bisexual, actinomorphic, axillary or terminal, solitary or in fasciculate panicles, racemes or spikes; epicalyx present or absent; calyx of 5 sepals fused basally; corolla of 5 petals, these usually clawed, adnate to base of the stamens; stamens few to many, filaments fused into a tube; ovarysuperior, syncarpous, 1-40-locular, styles usually branched.  Fruit a loculicidal capsule or schizocarp with variously ornamented mericarps, rarely a berry.  Seeds solitary to numerous, pubescent or glabrous; endosperm oily.

Distribution
Distribution in the Neotropics
  • The family Malvaceae s.l. includes 243 genera, and about 4225 species and is largely tropical to temperate.  Of these 129 genera and 1900-2200 species are native to the Neotropics.
  • Subfamily Malvoideae is almost cosmopolitan with 111-115 genera and 1800-2000 species, throughout the warm temperate and temperate zones worldwide, but mostly in the New World where 78 genera and about 1200-1400 species are present. 

Tribe Kydieae

  • Kydia Roxb.:  two species, native from Pakistan to China, but one species naturalized in South America.

Tribe Hibisceae

  • Abelmoschus Medik.:  15 species, endemic to the Old World, but a few species introduced and cultivated in the Neotropics.
  • Anotea (DC.) Kunth:  one species endemic to Mexico.
  • Hibiscus L.:  about 200 species almost worldwide, mostly tropical and subtropical.
  • Kosteletzkya C. Presl:  17 species worldwide, mostly tropical America and Africa, in the New World from southern U.S.A. to Colombia and Ecuador; sometimes considered synonymous with Hibiscus.
  • Malachra L.:  eight to ten species, pantropical, but centered in northern South America.
  • Malvaviscus Fabr.:  four species from southern U.S.A. and south to Peru and Brazil. Widely naturalised worldwide.
  • Pavonia Cav.:  about 250 species, ca. 225 of these in the New World from southern U.S.A. to Argentina and in the Caribbean Islands.
  • Peltaea (C. Presl)Standl.:  16 species, endemic to the New World, from tropical Paraguay and north to Mexico, and in the Caribbean Islands.
  • Phragmocarpidium Krapov.:  one species from central Brazil.
  • Rojasimalva Fryxell:  one species in Venezuela.
  • Talipariti Fryxell:  22 species worldwide with some species present in Latin America; sometimes considered synonymous with Hibiscus.
  • Urena L.:  six to eight species, pantropical, some species introduced into temperate areas.
  • Wercklea Pittier & Standl.:  13 species, endemic to the Neotropics: the Caribbean Islands, Costa Rica to Colombia and Ecuador; sometimes considered synonymous with Hibiscus.

Tribe Gossypieae

  • Cienfuegosia Cav.:  25 species, with subgenus Cienfuegosia in Africa and subgenus Articulata in the New World from Florida and Texas to the Caribbean Islands, and south to northern Argentina.
  • Gossypium L.:  50 species, pantropical - subgenus Houzingenia (13 species in the Neotropics, mostly Mexico), subgenus Gossypium (14 species mainly Africa, but also Neotropics), subgenus Karpas (5 species endemic to the Neotropics); and cultivated almost worldwide.
  • Hampea Schltdl.:  20 species, endemic to the Neotropics from Mexico to Colombia.
  • Thespesia Sol. ex Correa:  17 species, in tropical and South Africa, Arabia, the Indian Ocean Islands, tropical Asia, Australia, the Caribbean Islands and coastlines throughout the tropics.

Tribe Malveae

  • Abutilon Mill.:  about 160 species, mostly Neotropical.
  • Acaulimalva Krapov.:  19 species, in the high Andes, from Venezuela and Colombia, to Bolivia and northwestern Argentina.
  • Akrosida Fryxell & Fuertes:  one species from Brazil.
  • Alcea L.:  about 60 species from the Middle East and southwestern Asia with one species Alcea rosea L. introduced and widely cultivated in the Neotropics.
  • Allosidastrum (Hochr.) Krapov., Fryxell & D.M. Bates ex Fryxell:  four species from Mexico and the Caribean Islands to Brazil and Bolivia.
  • Allowissadula D.M. Bates:  nine species in southern Texas and central Mexico.
  • Andimalva J.A. Tate:  four species, one from Chile, the remaining from California, Baja California, and Mexico.
  • Anoda Cav.:  24 species, mostly Mexican, a few in the U.S.A., and one in South America.
  • Bakeridesia Hochr.:  about 20 species in Mexico and northern Central America, one disjunct in Venezuela, Colombia and Ecuador, and several in Brazil.
  • Bastardia Kunth:  three species widely scattered throughout the Neotropics.
  • Bastardiastrum D.M. Bates:  eight species in western Mexico.
  • Bastardiopsis (K. Schum.)Hassl.:  ten species from Venezuela, Colombia and Ecuador to Paraguay and Argentina.
  • Batasimalva Fryxell:  four species, three in Mexico and one in Venezuela.
  • Billieturnera Fryxell:  one species from SE Texas and NE Mexico.
  • Bordasia Krapov.:  one species from Paraguay.
  • Briquetia Hochr.:  five species from Mexico to Paraguay and Argentina.
  • Callirhoe Nutt.:  nine species from central U.S.A. to northeastern Mexico.
  • Calyculogygas Krapov.:  one species endemic to Uruguay.
  • Calyptraemalva Krapov.:  one species endemic to southern Brazil.
  • Corynabutilon (K. Schum.) Kearney:  six species from temperate Chile and Argentina.
  • Cristaria Cav.:  74 species from Chile, southern Peru and Bolivia.
  • Dendrosida Fryxell:  seven species from Mexico, Colombia and Venezuela.
  • Dirhamphis Krapov.:  two species, one in Mexico and one in Bolivia and Paraguay.
  • Eremalche Greene:  three species, from southern California and northern Baja California.
  • Fryxellia D.M. Bates:  one species in Texas, U.S.A. and Coahuila, Mexico.
  • Fuertesimalva Fryxell:  14 species, Andean (Venezuela to Chile and Argentina) and Mexico.
  • Gaya Kunth:  33 species mostly from Mexico, but also in the Caribbean Islands, the U.S.A., and one in South America.
  • Herissantia Medik.:  six species from tropical America, and one species H. crispa (L.) Brizicky is pantropical.
  • Hochneutinera Krapov.:  two species, one from Mexico and one from Paraguay and Argentina.
  • Horsfordia A. Gray:  four species in Arizona and California, U.S.A. and Sonora and Baja California, Mexico.
  • Kearnemalvastrum D.M. Bates:  two species from Mexico south to Colombia.
  • Krapovickasia Fryxell:  four species, one in Texas, U.S.A. and Mexico, and three from Peru to Uruguay.
  • Lecanophora Speg.:  five species in Argentina.
  • Lavatera L.:  four species are indigenous to the offshore islands of California and Baja California.
  • Malacothamnus Greene:  11 species from California and northern Baja California
  • Malva L.:  about 40 species in Europe, Africa, and the Middle and Far East. A few species have been introduced to the New World and have become naturalised, some as garden ornamentals, some as weeds; six species in Ecuador and Mexico.
  • Malvastrum A. Gray:  15 species from North, Central and South America, Australia, and one species adventive elsewhere.
  • Malvella Jaub. & Spach:  four species, three in western U.S.A. and Mexico, one in Peru to Uruguay, and one disjunct in the Mediterranean.
  • Meximalva Fryxell:  two species from southern U.S.A. to central Mexico.
  • Modiola Moench:  one pantropical species.
  • Modiolastrum K. Schum.:  five species in Brazil, Paraguay, Bolivia, Argentina, and Uruguay.
  • Monteiroa Krapov.:  eight species in southern Brazil and northern Argentina.
  • Neobaclea Hochr.:  one species from Argentina.
  • Neobrittonia Hochr.:  one species from Mexico to Panama.
  • Nototriche Turcz.:  about 100 Andean species from Ecuador to Chile and Argentina.
  • Palaua Cav.:  15 species in coastal Peru and Chile.
  • Periptera DC.:  five species in western Mexico and possibly also in Guatemala.
  • Phymosia Ham.:  eight species in Mexico and Guatemala and the Caribean Islands.
  • Pseudabutilon R.E. Fr.:  19 species from U.S.A. to Argentina.
  • Rhynchosida Fryxell:  two species, in Texas, U.S.A., northern Mexico, Bolivia and Argentina.
  • Robinsonella Rose & Baker f.:  15 species from Mexico to Costa Rica.
  • Sida L.:  about 100 species, pantropical, extending into temperate zones worldwide.
  • Sidalcea A. Gray:  20 species from western U.S.A. and northwestern Mexico.
  • Sidasodes Fryxell & Fuertes:  two species from Colombia, Ecuador, and Peru.
  • Sidastrum Baker f.:  eight species from Mexico and the Caribbean Islands to Argentina.
  • Spirabutilon Krapov.:  one species from eastern Brazil.
  • Sphaeralcea A.St.-Hil.:  about 40 species disjunct between temperate North and temperate South America.
  • Tarasa Phil.:  about 30 species, mainly Andean (Peru to Argentina and Chile), with two species in Mexico.
  • Tetrasida Ulbr.:  two species from Ecuador and Peru.
  • Wissadula Medik.:  26 species from Texas, U.S.A. to Argentina, and a few species extending to Africa.
Diagnostic
Distinguishing characters (always present)
  • Leaves alternate.
  • Stipules present, but often caducous.
  • Leaves simple, sometimes dissected.
  • Indumentum usually of stellate hairs or simple, glandular or lepidote hairs.
  • Ovarysuperior.
  • Fruit a loculicidal capsule or schizocarp with variously ornamented mericarps, rarely a berry.

 

Key differences from similar families

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

  • Brownlowioideae - sepals fused into a campanulate or urceolate tube; an androgynophore is always absent; stamens are many, free or fasciculate.
  • Bombacoideae - are usually stout trees with bottle-shaped and/or trunks armed with prickles.
  • Byttnerioideae - an epicalyx is always absent; the petals are free from the androecium; the stamens are free.
  • Cochlospermaceae -  the anthers dehisce via pores.
  • Grewioideae - an epicalyx is always absent; the petals are usually yellow or white; the stamens are free, sometimes fasciculate.
  • Helicteroideae - an epicalyx is always present; stamens 10-30.
  • Sterculioideae - petals are always absent; an androgynophore is usually present; the stamen filaments are free; the ovaries are apocarpous.
  • Turneraceae - stipules and epicalyx are always absent; two glands are often present at the leaf base; the stamens are free.
Useful tips for generic identification

Genera in Malvoideae

  • In the Neotropics an epicalyx is present in 38 and absent in 46 genera.
  • The number of styles per flower is also a good genus-level character, it varies from 1 to 45.
  • Only 15 genera have capsules and 3 have berries; the remaining have schizocarpous fruits.
  • The number of mericarps per schizocarp varies from 2 to 40.

General Description
Status

Native and introduced.

General notes
  • Abelmoschus esculentus (L.) Moench (= okra or ladies fingers) - is introduced, cultivated and used as a vegetable.
  • Abutilon - several species are widely cultivated as ornamentals.
  • Hibiscus - some species are ornamental e.g. H. rosa-sinensis L. and H. syriacus L.
  • Talipariti tiliaceum (L.) Fryxell - widely distributed along tropical shores.
Notes on delimitation

The past 15 years have seen an explosion in the number of 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 delimitations among the families previously recognised within the clade.  Recent outcomes of these papers include two reclassifications of the clade.  One is by Bayer & Kubitzki (2002), who 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. The infrafamiliar classification adopted here follows that of Bayer & Kubitzki (2002).

Malvoideae (alternatively Malvaceae) is unchanged from previous taxonomic frameworks.

Literature
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., 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.

Areces B., F. & Fryxell, P.A. 2007.  Malvaceae. In: Greuter, W. & Rankin R., R. (eds.). Flora de la República de Cuba, fasc. 13, pp. 1-228. A.R. Gantner Verlag KG, Ruggell.

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

Bayer, C., Fay, M.F., Bruijn, A.Y. de, Savolainen, V., Morton, C.M., Kubitzki, K., Alverson, W.S. & Chase, M.W. 1999.  Support for an expanded family concept of Malvaceae within a recircumscribed order Malvales: a combined analusis of plastid atpB and rbcL DNA sequences.  Botanical Journal of the Linnean Society 129: 267-303.

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

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.

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.

Cheek, M.R. 2007. Malvaceae. In: Heywood, V.H., Brummitt, R.K., Culham, A & Seberg, O. (eds.). Flowering Plant Families of the World, pp. 201-202. Royal Botanic Gardens Kew, Richmond.

Dorr, L.J. 2008.  Malvaceae. In: Hokche, O., Berry, P.E. & Huber, O. (eds.). Nuevo Catálogo de la Flora Vascular de Venezuela, pp. 458-464. Fundación Instituto Botánico de Venezuela Dr. Tobias Lasser, Caracas.

Fryxell, P.A. 1988.  Malvaceae of Mexico. Systematic Botany Monographs vol. 25, pp. 522.  The American Society of Plant Taxonomists, Ann Arbor.

Fryxell, P.A. 1989.  Malvaceae. In: Howard, R.A. (ed.). Flora of the Lesser Antilles: Leeward and Windward vol. 5, pp. 199-263. Jamaica Plain, Arnold Arboretum, Harvard University.

Fryxell, P.A. 1990.  Malvaceae. In: Breedlove, D.E. (ed.). Flora of Chiapas vol 3, pp. 1-90. California Academy of Sciences.

Fryxell, P.A. 1992a.  Malvaceae. In: Harling, G. & Andersson, L. (eds.). Flora of Ecuador 44 (118), pp. 1-141. Department of Systematic Botany, University of Gothenburg.

Fryxell, P.A. 1992b.  Malvaceae. In: Gomez-Pampa, A. (ed.). Flora de Veracruz, fasc. 68, pp. 1-255. Instituto de Ecología, Xalapa.

Fryxell, P.A. 1993a.  Malvaceae. In: Rzedowski, J. & Calderón de Rzedowski, G. Flora del Bajio y de regiones adyacentes fasc. 16, pp. 1-175. Instituto de Ecología, Xalapa.

Fryxell, P.A. 1993b.  Malvaceae. In: Flora del Valle de Tehuacan-Cuitcatlan fasc. 1, pp. 1-87. Universidad Nacional Autonomia de Mexico, Ciudad de Mexico.

Fryxell, P.A. 1997.  The American genera of Malvaceae II. Brittonia 49: 204-269.

Fryxell, P.A. 1999.  Pavonia cavanilles (Malvaceae). Flora Neotropica Monograph 76: 1-284.  New York Botanical Garden Press, New York.

Fryxell, P.A. 2001a.  Malvaceae. In: Berry, P.E., Yatskievych, K. & Holst, B.K. (eds.). Flora of the Venezuelan Guyana vol. 6, pp. 186-219.  Missouri Botanical Garden Press, St. Louis.

Fryxell, P.A. 2001b.  Malvaceae. In: Stevens, W.D., Ulloa U., C., Pool, A. & Montiel, O.M. (eds.). Flora de Nicaragua tomo II, pp. 1293-1322. Monographs in Systematic Botany from the Missouri Botanical Garden vol. 85.  St. Louis.

Fryxell, P.A. 2004.  Malvaceae. In: Smith, N., Mori, S.A., Henderson, A., Stevenson, D.W. & Heald, S.V. (eds.). Flowering Plants of the Neotropics, pp. 232-235. Princeton University Press, Princeton.

Fryxell, P.A. 2007.  Malvaceae. In: Hammel, B.E., Grayum, M.H., Herrera, C. & Zamora, N. (eds.). Manual de Plantas de Costa Rica vol. 6, pp. 313-373. Missouri Botanical Garden Press, St. Louis.

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.

Jørgensen, P.M. 1999.  Malvaceae. In: Jørgensen, P.M. & León-Yánez, S. (eds.). Catalogue of the Vascular Plants of Ecuador, pp. 548-554. Missouri Botanical Garden Press, St. Louis.

Krapovickas, A. 1965.  Malvaceae. In: Cabrera, A.L. (ed.). Flora de la Província de Buenos Aires vol. 4, pp. 169-220. I.N.TA. y Facultad de Ciencias Naturales y Museo de la Universidad Nacional de La Plata, Buenos Aires.

Krapovickas, A. 2008.  Malvaceae. In: Zuloaga, F.O., Morrone, O. & Belgrano, M.J. (eds.). Catálogo de las Plantas Vasculares del Cono Sur vol. 3, pp. 2463-2520. Missouri Botanical Garden Press, St. Louis.

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

Marticorena, A. 2005.  Malvaceae. In: Marticorena, C. & Rodrígues, R. (eds.). Flora de Chile vol. 2(3), pp. 22-119. Universidad de Concepción, Concepción.

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., Brubaker, C.L., Craven, L.A. & Crisp, M.D. 2002.  Phylogeny of Hibiscus and tribe Hibisceae (Malvaceae) using chloroplast DNA sequences of ndhF and the rpl16 intron. Syst. Bot. 27: 333-350.

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.

Pool, A. & Brako, L. 1993. Malvaceae. In: Brako, L. & Zarucchi, J.L. (eds.). Catalogue of the Flowering Plants and Gymnosperms of Peru, pp. 643-664. Monographs in Systematic Botany from the Missouri Botanical Garden vol. 45. St. Louis.

Robyns, A. 1966.  Malvaceae, family 115. In: Woodson, R.E. & Schery, R.W. (eds.) Flora of Panama. Ann. Missouri Bot. Gard. 52: 497-578.

Stevens, P. F. (2001 onwards). Angiosperm Phylogeny Website. Version 9, June 2008 (visited 9th Feb. 2011). http://www.mobot.org/MOBOT/research/APweb/.

Tate, J.A., Aguilar, J.F., Wagstaff, S.J., La Duke, J.C., Slotta, T.A.B. & Simpson, B.B. 2005. Phylogenetic relationships within the tribe Malveae (Malvaceae, subfamily Malvoideae) as inferred from ITS sequence data. American J. Bot. 92: 584-602.

[FWTA]

Malvaceae, Hutchinson and Dalziel. Flora of West Tropical Africa 1:2. 1958

Habit
Herbs often with fibrous stems or rarely shrubs; hairs usually stellate or lepidote
Leaves
Leaves alternate, often palmately nerved or divided; stipules present
Flowers
Flowers actinomorphic, hermaphrodite or rarely unisexual
Calyx
Sepals valvate, with or without an epicalyx of bracteoles
Corolla
Petals 5, free from each other but often adnate at the base to the staminal column, contorted or imbricate
Androecium
Stamens numerous, monadelphous; anthers 1-celled
Gynoecium
Ovules on axile placentas Ovary syncarpous, rarely of 1 carpel, rarely the carpels in vertical rows; style one
Fruits
Fruit a capsule or breaking into separate compartments
Seeds
Seeds usually with some endosperm and straight or curved embryo, the cotyledons often folded
[FTEA]

Bombacaceae, H. Beentje (East African Herbarium). Flora of Tropical East Africa. 1989

Habit
Trees
Leaves
Leaves alternate, petiolate, simple (not in East Africa) or digitate; stipules deciduous
Inflorescences
Inflorescence cymose or racemose, or flowers solitary, occasionally cauliflorous
Flowers
Flowers bisexual, regular, 5-merous
Calyx
Calyx truncate or lobed, often (but not in East Africa) with an epicalyx
Corolla
Petals free, contorted in bud
Androecium
Stamens 5-many, free or with the filaments united in their lower part, often in bundles; anthers ± medifixed, 1-thecous (in Flora area); pollen smooth
Gynoecium
Ovary superior, 2-10-locular, with 2-many ovules on the inner angle of each locule; style simple, with a capitate or lobed stigma
Fruits
Fruit a capsule, or indehiscent
Seeds
Seeds glabrous, often enclosed in the woolly hairs or mealy pulp of the endocarp; endosperm little or none; cotyledons plicate, or less often flat or contorted
[FWTA]

Bombacaceae, Hutchinson and Dalziel. Flora of West Tropical Africa 1:2. 1958

Habit
Trees sometimes very large
Leaves
Leaves simple or digitate, alternate, often lepidote; stipules deciduous
Flowers
Flowers hermaphrodite, large and showy
Calyx
Calyx closed and valvate in bud, often with an epicalyx
Corolla
Petals often elongated, sometimes absent
Androecium
Stamens numerous free or united into a tube; anthers reniform to linear, 1-celled; pollen smooth
Gynoecium
Ovary superior, 2–5-celled; style simple, capitate or lobed; ovules 2 or more on the inner angle of each cell
Fruits
Capsule loculicidally dehiscent or not dehiscent, the valves rarely falling away
Seeds
Seeds often embedded in hairs from the wall of the fruit, with little or no endosperm and flat or contorted or plicate cotyledons
[NTK]

Klitgård, B.B. (2013). Neotropical Malvaceae (Bombacoideae).

Morphology
Description

Malvaceae s.l.

Habit: shrubs, trees (herbs).  Leavesalternate or two-ranked, stipulate; leaf margin toothed or entire; venationpalmate 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; sepalaestivationvalvate; 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 climbers.  Leavesalternate; palmately lobed or palmate, rarely simple (Matisia and Quararibea); venationpalmate or 3-nerved, margins entire, rarely dentate; stipules present, often falling early; indumentum, if present, stellate or rarely lepidote.  Inflorescences of few-flowered, axillary clusters or panicles, solitary or rarely in leaf-opposing cymes.  Flowersbisexual, actinomorphic; epicalyx conspicuous, of 2-3 bracts, usually present; calyx tubular, 5-lobed fused for ½ its length; corollacontorted, 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; ovarysuperior, (2-)5(-8)-locular; style with short stylar branches.  Fruit a many-seeded, dehiscent, often ligneouscapsule with densely hairy endocarp (e.g. kapok) or indehiscent, juicy, 1-5-seeded drupe, rarely winged.  Seeds usually glabrous, sometimes arillate or winged.

General Description
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).

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.
Distribution
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.
Diagnostic
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 locule — Uladendron
11.  Leaves not lobed, margin entire or dentate; ovules several per locule — 12

12.  Leaves cordate, venation clearly palmate; trichomes lepidote — Septotheca
12.  Leaves elliptic to ovate, venation pinnate or faintly palmate; trichomes stellate — Huberodendron

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 sheath — Chiranthodendron

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 pericarp — Cavanillesia
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; indumentumlepidote — Phragmotheca
23.  Thecae not septate; indumentum never lepidote — Matisia

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.
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.
Other important characters
  • Trees, often buttressed or bottle-shaped and with or without stout prickles.
  • Leaves palmately lobed or palmate, rarely simple (Matisia and Quararibea)
  • Indumentum, if present, then usually of stellate, rarely lepidote trichomes.
  • Fruit often a ligneouscapsule with hairy endocarp (kapok), or indehiscent, juicy drupe, rarely winged.
Literature
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.

[FZ]

Bombacaceae, H. Wild. Flora Zambesiaca 1:2. 1961

Habit
Trees, sometimes very large; bark often smooth
Leaves
Leaves alternate, petiolate, simple or digitate; stipules deciduous
Flowers
Flowers bisexual, actinomorphic, large and showy
Calyx
Calyx tubular, truncate, shortly toothed or lobed (lobes valvate in bud), or rarely deeply 5-lobed with slightly imbricate lobes, often subtended by an epicalyx
Corolla
Petals free, often elongated, sometimes absent
Androecium
Stamens 15–?, free or filaments united into a tube below or in phalanges of 2–3; anthers 1-thecous, reniform or linear; pollen smooth
Gynoecium
Ovary superior, 2–10-locular, with axile placentas; ovules 2 or more per loculus; style simple; stigma capitate or lobed
Fruits
Fruit a woody capsule or indehiscent
Seeds
Seeds embedded in a powdery matrix or often in hairs on the wall of the fruit; endosperm scanty or absent; cotyledons flat or contorted or plicate
[FTEA]

Tiliaceae, C. Whitehouse, M. Cheek, S. Andrews & B. Verdcourt. Flora of Tropical East Africa. 2001

Habit
Trees, shrubs, subshrubs or less often annual or perennial herbs; stellate or lepidote indumentum very often present; bark and cortex with mucilage cells
Leaves
Leaves alternate or rarely opposite or subopposite, simple, sometimes cordate at the base, entire or sometimes ± deeply lobed, rarely subdigitate, often palmately lobed at base; stipules ± deciduous or absent, sometimes divided, rarely leafy
Inflorescences
Inflorescences various in structure, mostly cymose, axillary or terminal or less often lateral, leaf-opposed or borne on the old wood; bracts and bracteoles usually present, sometimes united to form an involucre, or large and foliaceous forming a wing (in Tilia )
Flowers
Flowers regular, hermaphrodite or less often unisexual; androgynophore often present; sepals (2–)4–5(–7), valvate, free or rarely joined, rarely accrescent; petals 4–5(–7), contorted, imbricate or valvate or occasionally absent, free or rarely connate, often with a basal internal glandular area
Androecium
Stamens usually numerous, often some reduced to staminodes; filaments free or all joined in 4–10 phalanges; anthers basi- or dorsifixed, dehiscing by pores or longitudinal slits; disc often present either extrastaminal or bearing the stamens
Gynoecium
Ovary superior (save in one genus), syncarpous or very rarely apocarpous, (1–)2–5(–10)-locular, often deeply lobed; placentation axile, or parietal when unilocular; ovules 1–many; style usually 1; stigma entire or lobed
Fruits
Fruits either baccate, drupaceous, nut-like or dehiscent often winged capsules or capsule-like but indehiscent or separating into follicles, the surface smooth or covered with prickles or spines
Seeds
Seeds 1–many, often one per locule, often winged, sometimes arillate; albumen usually present
[FZ]

Tiliaceae, H. Wild. Flora Zambesiaca 2:1. 1963

Habit
Small trees, lianes, shrubs, or annual or perennial herbs, often stellately hairy; leaves usually alternate, simple or rarely digitate, entire, toothed, or lobed; stipules usually small and deciduous
Inflorescences
Inflorescence usually cymose, with the cymes often leaf-opposed, sometimes in corymbs or panicles, usually axillary, sometimes terminal
Flowers
Flowers actinomorphic, usually bisexual
Calyx
Sepals 5 or sometimes 2–4, free or occasionally connate, usually valvate
Corolla
Petals free, equalling the number of sepals and alternating with them, rarely absent, often with a glandular claw or appendage at the base
Androecium
Stamens 4-? (usually ?), often on a raised torus or androgynophore, free or connate at the base, all fertile or the outer ones sterile; anthers dehiscing longitudinally or by terminal pores
Gynoecium
Style entire or lobed at the apex or the stigmas almost free Ovary superior, 2–10-locular, with 1 to many ovules per loculus
Fruits
Fruit a dry or somewhat fleshy drupe or a schizocarp, 2–10-locular or 1-locular by abortion, sometimes transversely septate between the seeds
Seeds
Seeds with endosperm
[FWTA]

Tiliaceae, Hutchinson and Dalziel. Flora of West Tropical Africa 1:2. 1958

Habit
Trees or shrubs, sometimes herbs, often clothed with stellate hairs
Leaves
Leaves usually alternate, simple; stipules paired or absent
Flowers
Flowers mostly cymose, actinomorphic, mostly hermaphrodite
Calyx
Sepals valvate
Corolla
Petals free or absent, contorted, imbricate or valvate
Androecium
Stamens usually numerous, free or rarely connate into 5–10 bundles; anthers 2-celled
Gynoecium
Ovary syncarpous or very rarely apocarpous, superior, 2–10-celled; style usually simple; ovules on axile placentas
Fruits
Fruit baccate or drupaceous or variously dehiscent
Seeds
Seeds sometimes hairy, mostly with copious endosperm and usually straight embryo
[NTK]

Klitgård, B.B. (2013). Neotropical Malvaceae (Byttnerioideae).

Morphology
Description

Malvaceae s.l.

Habit: shrubs, trees (herbs).  Leavesalternate or two-ranked, stipulate; leaf margin toothed or entire; venationpalmate 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; sepalaestivationvalvate; androgynophore present or absent; stamens 5-many, in five groups (fundamentally obdiplostemonous).  Fruita capsule, berry, schizocarp.

Subfamily Byttnerioideae (Byttneriaceae)

Habit: usually small shrubs, less often trees, lianas or herbs; bark highly fibrous.  Leavesalternate or two-ranked; simple, rarely palmate (Herrania); venationpalmate or 3-nerved, margins often serrate; stipules present, often falling early; indumentum, if present, stellate.  Inflorescencesaxillary, leaf-opposed or rarely terminal thyrses, sometimes reduced to solitary flowers or cauliflorous fascicles (Theobroma and Herrania).  Flowersbisexual, actinomorphic; epicalyx usually absent; calyx of 5 sepals fused for less than ½ their length; corolla of 5 free petals, these often cupped or hooded and with a strap-like appendage, basally free from the androecium, sometimes completely reduced (species of Hermannia, Melochia, Waltheria); androgynophore absent; stamens 5 to 10, rarely more, fused into short tube or fascicles, anthers usually dithecal, rarely trithecal; ovarysuperior, (1-3)5-locular; stylesimple.  Fruit a many-seeded, indehiscent, fleshyberry, dehiscentcapsule or schizocarp with 1-many-seeded mericarps.  Seeds usually glabrous, sometimes arillate or pubescent.

General Description
Notes on delimitation

The past 15 years have seen an explosion in the number of 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 delimitations among the families previously recognised within the clade.  Recent outcomes of these papers include two reclassifications of the clade.  One is by Bayer & Kubitzki (2002), who 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.  The infrafamiliar classification adopted here follows that of Bayer & Kubitzki (2002).

Byttnerioideae (alternatively Byttneriaceae):  its members were formerly included in the Sterculiaceae, but molecular, morphological (presence of cupped petals and absence of androgynophore) and biogeographic data joined these genera and resolved Grewioideae as its sister, thus separating them from the reduced Sterculiaceae/Sterculioideae.

General notes
  • Herrania lemniscata (Schomb.) R.E.Schult. the sweet-sour pulp around the seeds is edible, tasting similar to Theobroma pulp (Millken & Albert 1999).
  • Theobroma cacao and relatives are used to make chocolate.  Chocolate is more than just a delicacy: evidence suggests that eating between 46 and 105g of chocolate a day can have a moderate effect on lowering blood pressure (Ried et al. 2012).
  • Cocoa has also been used for an array of medicinal purposes. Unfermented cocoa seeds and the seed coat are used to treat a variety of ailments, including diabetes, digestive and chest complaints. Cocoa powder, prepared from fermented cocoa beans, is used to prevent heart disease. Cocoa butter is taken to lower cholesterol levels, although its efficacy is unclear.  It is also used widely in foods and pharmaceutical preparations, as well as being used as a rich moisturiser for the skin.
  • The crushed shells of cocoa beans are used as an alternative to peat mulch. Mulches are layered on to the soil surface to suppress weeds, conserve moisture, improve its visual appearance and minimize erosion. Not only does this make good use of cocoa-shell, which is a by-product of the chocolate industry, but it also helps reduce the use of peat.
  • For further information see http://www.kew.org/plants-fungi/Theobroma-cacao.htm.
Status

Native.

Distribution
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 Byttnerioideae is pan- and subtropical and consists of 27 genera and c. 650 species; widely distributed in Africa and Latin America with eight genera and 280-300 species in the Neotropics.  About half the genera are restricted to Australasia.  Some groups are mainly found in humid tropical forest, while others are restricted to drier habitats.

  • Ayenia L.:  about 70 species from southern North America to Argentina.
  • Byttneria Loefl.:  pantropical with more than 130 species, about 80 of which are neotropical.
  • Guazuma Mill.:  three species from Mexico to NE Argentina and West Indies.
  • Herrania Goudot:  about 17 species in tropical Central and South America.
  • Hermannia L.:  More than 100 spp., majority in the Old World (South Africa, Madagascar, Australia), four spp. in southern North America and adjacent Mexico (H. inflata Link & Otto, H. palmeri Vasey & Rose, H. pauciflora S. Watson, and H. texana A. Gray)
  • Melochia L.:  less than 50 species, mostly neotropical, a few in the Palaeotropics - some widely distributed weeds (Chocolate weed - M. corchorifolia L).
  • Rayleya Cristóbal:  one species (R. bahiensis Cristóbal) from Brazil.
  • Theobroma L.:  22 species - in tropical rain forests of Central and South America.
  • Waltheria L.:  about 50 to 60 species, mostly neotropical with centres of diversity in Mexico and Brazil (W. indica L. native to the New World, but a pantropical weed).
Diagnostic
Other important characters
  • Usually small shrubs, less often trees, lianas or herbs.
  • Indumentum, if present, usually of stellate, rarely lepidote trichomes.
  • Stamens antepetalous, 5-10, solitary or in groups of 2-3(-6) with the filaments fused within groups.
  • Fruit a berry, capsule, rarely schizocarp.
Distinguishing characters (always present)
  • Leaves simple (except palmate in Herrania).
  • Stipules present, but often early caducous.
  • Leaves often distichous with plagiotropic branching.
  • Epicalyx absent.
  • Petals usually cupped or hooded, often with apical appendage.
  • Androgynophore absent.
  • Anthers dithecal.
Useful tips for generic identification

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

1.  Stamens usually more numerous than sepals — 2
1.  Stamens usually as many as sepals — 4

2.  Petal appendages bifid; staminodes short — Guazuma
2.  Petal appendages entire; staminodes prominent — 3

3.  Leaves digitately compound; trees without plagiotropic side branches — Herrania
3.  Leaves simple or rarely lobed; trees with plagiotropic side branches — Theobroma

4.  Staminodes absent or rudimentary, never lianas, usually shrubs or herbs, rarely trees — 5
4.  Staminodes present, never herbs, usually shrubs or lianas, rarely trees — 6

5.  Ovary 5-locular — 6
5.  Ovary 1-locular — Waltheria

6.  Carpels antepetalous; ovarial locules usually with 2 ascending ovules — Melochia
6.  Carples aneepalous; ovarial locules with several more or less horizontal locules — Hermannia

7.  Anthers trithecal, androgynophore present, usually long and prominent — Ayenia
7.  Anthers dithecal, androgynophore usually absent, rarely present —8

8.  Shoots often spinose; androgynophore absent — Byttneria (throughout Latin America)8.  Shoots not spinose; androgynophoreprominent — Rayleya (Bahia, Brazil)

Key differences from similar families

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

  • Brownlowioideae - sepals fused into a campanulate or urceolate tube; stamens many, free or fasciculate.
  • Bombacoideae - usually stout trees with bottle-shaped and/or trunks armed with prickles; leaves usually palmate/palmately lobed; sepals fused into a tube; petals adnate to androecium; style branched.
  • Cochlospermaceae - anthers dehisce via pores.
  • Grewioideae - petals usually yellow or white, clawed, often with hairy basal nectaries; stamens numerous, free, sometimes grouped in antesepalous fascicles.
  • Helicteroideae - calyx tubular; epicalyx always present; petals clawed; androgynophore usually present; stamens 10-30; ovaries usually apocarpous, except Ungeria and Reevesia.
  • Malvoideae - epicalyx is present in ½ and absent in ½ the Neotropical genera; petals adnate to the androecium; stamens fused into a staminal column.
  • Sterculioideae - petals always absent; androgynophore usually present; stamen filaments free; ovaries apocarpous.
Literature
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., 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.

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., Fay, M.F., Bruijn, A.Y. de, Savolainen, V., Morton, C.M., Kubitzki, K., Alverson, W.S. & Chase, M.W. 1999.  Support for an expanded family concept of Malvaceae within a recircumscribed order Malvales: a combined analusis of plastid atpB and rbcL DNA sequences.  Botanical Journal of the Linnean Society 129: 267-303.

Bayer, C. & Kubitzki, K. 2002.  Malvaceae, Byttnerioideae. In: K. Kubitzki, & C. Bayer (eds). The Families and Genera of Vascular Plants vol. V, pp. 241-247. Springer-Verlag, Berlin.

Bornstein, A.J. 1989. Sterculiaceae. In: Howard, R.A. (ed.). Flora of the Lesser Antilles: Leeward and Windward vol. 5, pp. 272-292. 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.

Cheek, M.R. 2007.  Byttneriaceae, p. 76. In: V.H. Heywood, R.K. Brummitt, A. Culham, & 0. Seberg (eds). Flowering Plant Families of the World, pp. 241-247. Royal Botanic Gardens, Kew.

Cristóbal, C.L. 1960.  Revisión del género "Ayenia". Opera Lilloana 4: 1-230.

Cristóbal, C.L. 1976.  Estudio taxonomico del genero Byttneria Loefling (Sterculiaceae).  Bonplandia 4: 3-428.

Cristóbal, C.L. 1981.  Rayleya, nueva Sterculiaceae de Bahia - Brasil. Bonplandia 5: 43-50.

Cristobal, C.L. 1998. Sterculiaceae fam. 178.  In: Hunziker, A.T. (ed.).  Flora fanerogamica Argentina.  Fasciculo 57, 32pp.  Proflora, Cordoba.

Cristóbal, C.L. 2001.  Sterculiaceae. In: Stevens, W.D., Ulloa U., C., Pool, A. & Montiel, O.M. (eds.). Flora de Nicaragua tomo III, pp. 2428-2437. Monographs in Systematic Botany from the Missouri Botanical Garden vol. 85.  St. Louis.

Cristóbal, C.L. 2007.  Sterculiaceae de Paraguay, I.  Ayenia, Byttneria, Guazuma, Helicteres, Melochia y Sterculia. Bonplandia 16(1-2): 5-142.

Cristóbal, C.L., Saunders, J.G., & Berry, P.E. 2005.  Sterculiaceae.  In: Berry, P.E., Holst, B.K. & Yatskievych, K. (eds.).  Flora of the Venezuelan Guyana vol. 9, pp. 248-280.  Missouri Botanical Garden, St. Louis.

Cristóbal, C.L. & Saunders, J.G. 2008.  Sterculiaceae. In: Zuloaga, F.O., Morrone, O. & Belgrano, M.J. (eds.). Catálogo de las Plantas Vasculares del Cono Sur vol. 3, pp. 3055-3064. Missouri Botanical Garden Press, St. Louis.

Door, L.J. 2002.  Sterculiaceae. 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. 700-706.  Memoirs of the New York Botanical Garden 76, part 2.  New York Botanical Garden Press.

Door, L.J. & Fryxell, P.A. 1999.  Sterculiaceae. In: Jørgensen, P.M. & León-Yánez, S. (eds.). Catalogue of the Vascular Plants of Ecuador, pp. 918-922. Missouri Botanical Garden Press, St. Louis.

Fryxell, P.A. 2004.  Sterculiaceae. In: N. Smith, S.A. Mori, A. Henderson D.W. Stevenson & S.V. Heald (eds). Flowering Plants of the Neotropics, pp. 360-362. Princeton University Press, Princeton.

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.

Goldberg, A. 1967.  The genus Melochia (Sterculiaceae). Contr. U.S. Nat. Herb. 34(5): 191-363.

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

Klitgård, B.B., Grace, O. & Tredwell, E. (2010 onwards).  Theobroma cacao (cocoa tree) - species page. http://www.kew.org/plants-fungi/Theobroma-cacao.htm

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

Merello, M.C., Cristóbal, C.L. & Dorr, L.J. 1993. Sterculiaceae. In: Brako, L. & Zarucchi, J.L. (eds.). Catalogue of the Flowering Plants and Gymnosperms of Peru, pp. 1137-1142. Monographs in Systematic Botany from the Missouri Botanical Garden vol. 45. St. Louis.

Milliken, W. & Albert, B. 1999.  Yanomami: A Forest People. Royal Botanic Gardens, Kew.

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.

Ried K, Sullivan TR, Fakler P, Frank OR, Stocks NP. 2012. Effect of cocoa on blood pressure. Cochrane Database of Systematic Reviews Issue 8. Art. No.: CD008893. DOI: 10.1002/14651858.CD008893.pub2.

Rose, J.N. 1897.  A Synopsis of the American Species of Hermannia. Contrib. US National Herbarium 5(3): 130-131.

Ruiz, E. 2005.  Sterculiaceae. In: Marticorena, C. & Rodrígues, R. (eds.). Flora de Chile vol. 2(3), pp. 20-21. Universidad de Concepción, Concepción.

Robyns, A. & Cuatrecasas. 1964.  Sterculiaceae, family 117. In: Woodson, R.E. & Schery, R.W. (eds.) Flora of Panama part VI. Ann. Missouri Bot. Gard. 51: 69-107.

Rondon, J.B., Cumana C., L.J. 2005.  Revision taxonomica del genero Theobroma (Sterculiaceae) en Venezuela.  Acta Bot. Venezuel. 28: 113-133.

Saunders, J.G. 1993.  Four new distylous species of Waltheria (Sterculiaceae) and a key to the Mexican and Central American species and species groups. Syst. Bot. 18: 356-376.

Saunders, J.G. 1995.  Systematics and Evolution of Waltheria (Sterculiaceae-Hermannieae). Ph.D. diss. The University of Texas at Austin. 854 pp.

Saunders, J. G. 2005. New species of Waltheria (Hermannieae, Byttnerioideae, Malvaceae) from Paraguay, Argentina, and Venezuela, and two new records for Paraguay. Darwiniana 43(1-4): 201-211.

Silva, C.R.S. & Figueira, A. 2005.  Phylogenetic analysis of Theobroma (Sterculiaceae) based on Kunitz like trypsin inhibitor sequences.  Plant Systematics & Evolution 250: 93-104.

Stevens, P. F. (2001 onwards).  Angiosperm Phylogeny Website. Version 9, June 2008 (visited 6th Feb. 2013). http://www.mobot.org/MOBOT/research/APweb/.

Whitlock, B.A. & Baum, D.A. 1999.  Phylogenetic relationships of Theobroma and Herrania (Sterculiaceae) based on sequences of the nuclear gene vicilin. Systematic Botany 24(2): 12-138.

Whitlock, B.A., Bayer, C. & Baum, D.A. 2001.  Phylogenetic Relationships and Floral Evolution of the Byttnerioideae ("Sterculiaceae" or Malvaceae s.l.) Based on Sequences of the chloroplast Gene, ndhF. Systematic Botany 26(2): 420-437.

Whitlock, B.A. & Hale, A.M. 2011.  The Phylogeny of Ayenia, Byttneria, and Rayleya (Malvaceae s.l.) and Its Implications for the Evolution of Growth Forms. Systematic Botany 36(1): 129-136.

[FTEA]

Sterculiaceae, Martin Cheek & Laurence Dorr; Nesogordonia, Laurence Dorr, Lisa Barnett. Flora of Tropical East Africa. 2007

Habit
Monoecious or polygamous, evergreen or deciduous trees and shrubs, rarely herbs or climbers
Indumentum
Indumentum stellate, often mixed with simple hairs, rarely with peltatelepidote scales
Stem
Stems sometimes exuding mucilage when wounded, fibrous
Leaves
Leaves alternate, simple, elliptic to oblanceolate or ± orbicular and then often digitately lobed, venation palmate or pinnate, reticulate; petiole usually swollen and often kneed at base and apex
Stipules
Stipules linear to lanceolate, usually caducous. Flowers actinomorphic, hypogynous, bisexual, perianth biseriate, calyx 5-lobed or of 5 free sepals (spathaceous in Mansonia) valvate
Corolla
Corolla imbricate, with 5 free often clawed petals, or (genera 1–6) flowers unisexual, perianth uniseriate, the perianth 5-lobed, valvate.
Androgynophore
Androgynophore well-developed, inconspicuous or absent.
Androecium
Androecium uniseriate, stamens (4–)5–20, all fertile or with some filiform staminodes, the filaments united into a long or short tube, or free or, (genera 1–6) anthers subsessile in a ± globose head on an androgynophore; less usually androecium biseriate, the inner whorl being staminodes, usually petaloid, the outer whorl fertile.
Anthers
Anthers dithecal, dehiscing by slits, rarely by apical pores, extrorse.
Gynoecium
Gynoecium syncarpous, locules (1–)5, placentation axile, ovules 2–numerous or apocarpous (genera 1–7), carpels 5(–numerous, then spiralled) cohering, separating in fruit; styles as many as carpels, cohering and appearing single, with 3–5 apical branches
Fruits
Fruit syncarpous, then loculicidally, rarely septicidally ( Byttneria) dehiscent, rarely indehiscent and then berry-like ( Theobroma), if apocarpous, the fruitlets ventrally dehiscent along the placenta, then woody or leathery; or indehiscent and fleshy, leathery or papery
Seeds
Seeds 1-numerous, rarely winged or arillate, endosperm present or absent, cotyledons flat or folded, thin or fleshy.
Note
Byttneriaceae (genera 12–16) An alternative approach is to unite all these families in a ‘Super-Malvaceae’ and to recognise them at the subfamily level instead (Bayer & Kubitzki, Genera and Families of Vascular Plants 5 (2001)). This seems the less desirable solution since it treats Malvaceae as a dustbin family and creates greater taxonomic instability at the family level than the solution above. Helicteraceae (genus 7) Pentapetaceae (genera 8–11) Sterculiaceae (genera 1–6 of this volume) Brownlowiaceae (Christiana and Carpodiptera, earlier published in FTEA Tiliaceae, 2001) Sparrmanniaceae (the rest of the genera in FTEA Tiliaceae, 2001) Bombacaceae (as published in FTEA Bombacaceae, 1989) Malvaceae (in preparation for FTEA) Mainly a tropical family but several species in the subtropics. About 67 genera and 1500 species. The division of Sterculiaceae from Tiliaceae using this division has long been problematic, as illustrated by Hutchinson’s provision of a key to the genera of the combined families (Hutchinson 1967, The Genera of Flowering Plants 2). Recent molecular work (e.g. Baum et al., Harvard Pap. Bot. 3: 315–330 (1998) and Bayer et al., Bot. J. Linn. Soc. 129: 267–303 (1999)) has shown that some groups of traditional Sterculiaceae are undoubtedly more closely related to some parts of traditional Tiliaceae than they are to the rest of the family. While Malvaceae and Bombacaceae can be maintained ± in the traditional sense (although Durio and its SE Asian relatives require removal from the last), there is no doubt that this is not feasible regarding Tiliaceae and Sterculiaceae, which need to be broken up into several units, all of which are morphologically distinct and have previously been recognised as families. Core Malvales can thus be expanded from the traditional four to ten newly circumscribed families. So far as FTEA is concerned these are as follows:
[FWTA]

Sterculiaceae, Hutchinson and Dalziel. Flora of West Tropical Africa 1:2. 1958

Habit
Trees or shrubs, often with soft wood, or very rarely herbs, mostly with stellate hairs
Leaves
Leaves alternate, simple or digitately compound; stipules usually present
Flowers
Flowers variously arranged, hermaphrodite or unisexual, actinomorphic
Calyx
Sepals valvate, mostly partly connate or rarely spathaceous
Corolla
Petals 5, or absent, contorted imbricate, often hooded
Androecium
Stamens free or connate into a column, sometimes with staminodes; anthers 2-celled
Gynoecium
Ovary superior, of 2–12 united carpels or of one carpel; ovules on axile placentas; style simple or rarely the styles free to the base
Fruits
Fruit various
Seeds
Seeds with or without endosperm and straight or curved embryo
[FZ]

Sterculiaceae, H. Wild. Flora Zambesiaca 1:2. 1961

Habit
Herbs, shrubs or trees, almost invariably with stellate hairs, sometimes with simple hairs intermingled
Leaves
Leaves usually alternate, simple and pinnately or palmately nerved, entire to toothed or lobed, or digitately compound; stipules usually present
Flowers
Flowers actinomorphic, bisexual, unisexual or polygamous
Calyx
Calyx often persistent, more or less deeply divided into 5 or rarely 3 or 4 or 6 valvate lobes, rarely splitting irregularly into 2 valves, or sepals quite free
Corolla
Petals usually 5, hypogynous, free or adhering to the staminal column, contorted in bud, or small and scale-like or absent
Androecium
Stamens 5–?, with filaments united into a tube or free or confluent at the apex of an androphore, sometimes with staminodes alternating with the stamens or with fascicles of stamens or with an inner ring of staminodes; anthers 2-thecous and opening outward by slits or occasionally by apical pores
Gynoecium
Ovary superior, sessile or stipitate, 2–5-locular or with 2–5 ± coherent carpels which separate in fruit, rarely 10–12-locular or reduced to a single carpel; styles simple or equalling the number of loculi
Fruits
Fruit various but often a dehiscent capsule or of woody or membranous follicles or of indehiscent fruiting carpels, occasionally winged
Seeds
Seeds 1-co, sometimes winged; endosperm present or absent; cotyledons flat or folded, thin or fleshy
[FTEA]
Use
Many species are cultivated as ornamentals or for fibres and as vegetables.

Images

Malvaceae Juss. appears in other Kew resources:

First published in Gen. Pl. [Jussieu] 271. 1789 [4 Aug 1789] (1789)

Accepted by

  • APG IV (2016) http://dx.doi.org/10.1111/boj.12385

Sources

Flora Zambesiaca
Flora Zambesiaca
[A] http://creativecommons.org/licenses/by-nc-sa/3.0

Flora of Tropical East Africa
Flora of Tropical East Africa
[B] http://creativecommons.org/licenses/by-nc-sa/3.0

Flora of West Tropical Africa
Flora of West Tropical Africa
[C] http://creativecommons.org/licenses/by-nc-sa/3.0

Kew Names and Taxonomic Backbone
The International Plant Names Index and World Checklist of Selected Plant Families 2018. Published on the Internet at http://www.ipni.org and http://apps.kew.org/wcsp/
[D] © Copyright 2017 International Plant Names Index and World Checklist of Selected Plant Families. http://creativecommons.org/licenses/by/3.0

Neotropikey
Milliken, W., Klitgard, B. and Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics.
[E] http://creativecommons.org/licenses/by/3.0