Neotropical Malvaceae (Malvoideae)

Bente B. Klitgård, Sara L. Edwards, Nicola Biggs & Sue Frisby

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 cacao where it was first observed).  Flowers with epicalyx present or absent;  sepal aestivation valvateandrogynophore present or absent; stamens 5-many, in five groups (fundamentally obdiplostemonous).  Fruit a capsule, berry, or schizocarp.

Subfamily Malvoideae (Malvaceae s.s.)

Habit: herbs, shrubs or trees.  Leaves alternate; 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; ovary superior, syncarpous, 1-40-locular, styles usually branched.  Fruit a loculicidal capsule or schizocarp with variously ornamented mericarps, rarely a berrySeeds solitary to numerous, pubescent or glabrous; endosperm oily.

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.

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.

Distinguishing characters (always present)

 

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.

    Key to the Latin American Genera of Malvaceae (Subfamily Malvoideae)

http://www.kew.org/science/tropamerica/malvaceae.htm

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.

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.

How to cite

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