Neotropical Ochnaceae s.l. (incl. Quiinaceae)

Pedro Fiaschi*, Cláudio Nicoletti de Fraga** and Kikyo Yamamoto***

*Universidade de São Paulo, São Paulo, Brasil,** Jardim Botânico do Rio de Janeiro, Brazil, and ***Universidade Estadual de Campians (UNICAMP) 

Description

Small to large trees, shrubs (rarely scandent), or herbs, glabrous, glabrescent or pilose. Leaves simple, alternate (subf. Ochnoideae), opposite or whorled (subf. Quiinoideae), sometimes pinnatifid or imparipinnately compound (Rhytidanthera and Touroulia), sessile or usually short-petiolate; stipules intra- or interpetiolar, entire, awned or divided into setae, persistent or caducous; margins usually crenate, serrate, denticulate, sometimes entire or ciliate (Luxemburgia); venation craspedodromous, brochidodromous or eucamptodromous (especially Ouratea s.l.); secondary veins usually running parallel, at least in the proximal half, a few well-spaced or numerous and closely arranged (Clusia-like); tertiary veins parallel or reticulate. Inflorescences terminal or axillary, generally compound, thyrsoid, botryoid or diplobotryoids, rarely reduced to a single flower (sometimes in Sauvagesia); bracts caducous or persistent. Flowers actinomorphic or zygomorphic due to the lateral position of the androecium and gynoecium (Cespedesia, Luxemburgia, Philacra and Wallacea); uni- or bisexual, the plants hermaphroditic, androdioecious, or dioecious (subf. Quiinoideae); pedicels articulated. Sepals (2-)4-5(-12), usually free, imbricate, sometimes ciliate or glandular-ciliate. Petals (3-)5(-8), free, usually yellow, sometimes whitish, reddish or pink. Androgynophore rarely present (Adenarake). Androecium of 5-25(->300) stamens, equally distributed in the receptacle or laterally grouped, sometimes surrounded by 1-2 cycles of filamentous or petaloid staminodes which may form a corona (Sauvagesia); anthers usually basifixed, sometimes dorsifixed (subf. Quiinoideae), dehiscing through longitudinal slits or 1-2 pores. Ovary superior, 2-10(-14)-carpellate, syncarpous or rarely apocarpous (Froesia), secondarily free in Ouratea; ovules 1-many per locule, placentation parietal, axilary or basal; one or more styles (then each per carpel), terminal, gynobasic in Ouratea; stigma punctate, slightly lobed or subpeltate. Fruits capsules or berries, sometimes schizocarps with 1-10 fleshy mericarps sitting on a reddish fleshy carpophore (Ouratea). Seeds one to many; endosperm present or absent; embryo straight or curved, cotyledons thick or small and delicate, equal to very unequal in size.

Notes on delimitation

  • Recent classification systems have included Quiinaceae (and the Seychelles endemic Medusagynaceae) in the Ochnaceae (APG 2009). This view is here adopted. Whenever necessary the Ochnaceae s.s. is referred to as subf. Ochnoideae, and the Quiinaceae s.s. as subf. Quiinoideae [Stevens 2001 (onwards)].
  • Sauvagesiaceae has been earlier recognized as a family on its own, because of the flowers with 2-5 fused carpels, staminodes present, and seeds with endosperm. We agree that Sauvagesiaceae is better regarded as a subfamily of Ochnaceae (Amaral 1991).

Distribution in the Neotropics

  • Adenarake, Blastemanthus, Philacra, Poecilandra, and Tyleria - endemic to the Guayana Shield in northern South America.
  • Elvasia - Mesoamerican and South American rainforests, especially in the Amazon Basin, the Guianas and the Brazilian Atlantic forests.
  • Froesia and Touroulia are restricted to northern South America, mostly in the Amazonian forests.
  • Godoya and Rhytidanthera are restricted to the Andean forests, from Colombia to Bolivia.
  • Krukoviella is endemic to the western Amazon, in Brazil (Amazonas), Peru and Ecuador (Zamora-Chinchipe).
  • Lacunaria - most species in the Amazonian lowland forests. Lacunaria crenata (Tul.) A.C. Sm. also in the Brazilian Atlantic forests, and L. panamensis (Standl.) Standl. endemic to the South central American rainforests.
  • Luxemburgia - Brazil. Most species endemic to the Espinhaço Range in Minas Gerais; a few species in Goiás, Bahia, and in granitic outcrops in Espírito Santo and Rio de Janeiro states.
  • Quiina - most species in the Amazon Basin. Distribution ranging from Belize and Jamaica to Bolivia and southern Brazil.
  • Perissocarpa - northern South America, at high-elevation areas (600-1900m) in the Andes, Coastal Cordillera of Venezuela, and the western portion of the Guayana Shield.
  • Sauvagesia - most species either endemic to white-sand forests and remnant areas of the Guayana Shield in Venezuela, Colombia, Brazil and Guyana, or to campos along the Espinhaço Range in Minas Gerais and Bahia, Brazil; yet, S. erecta has worldwide distribution.
  • Wallacea - white-sand forests in southeastern Colombia, southern Venezuela and northern Brazil.

Distinguishing characters (always present)

Other important characters

  • The petioles are usually short, resembling a pulvinus.
  • Leaf margin is often crenate or serrate, sometimes shallowly denticulate, ciliate or smooth.
  • Inflorescences are usually paniculate, with showy flowers often with yellow petals (subf. Ochnoideae).
  • 2-15 fused carpels with a common style (subf. Ochnoideae s.s.) or with as many styles as the number of carpels (subf. Quiinoideae). Free carpels only in Froesia (subf. Quiinoideae).

Number of genera

19 native genera in the Neotropics:

  • Adenarake Maguire & Wurdack (2 spp.)
  • Blastemanthus Planch. (2 spp.)
  • Cespedesia Goudot [1 spp., C. spathulata (Ruiz. & Pav.) Planch.]
  • Elvasia DC. (14 spp.)
  • Froesia Pires (5 spp.)
  • Godoya Ruiz & Pav. (~ 5 spp.)
  • Krukoviella A.C.Sm. [1 spp., K. disticha (Tieghem) Dwyer]
  • Lacunaria Ducke (10 spp.)
  • Luxemburgia A. St.-Hil. (19 spp.)
  • Ouratea Aubl., including Gomphia Schreb., Kaieteurea Dwyer and Plicouratea Tiegh. (~ 300 spp.)
  • Perissocarpa Steyerm. & Maguire (3 spp.)
  • Philacra Dwyer (4 spp.)
  • Poecilandra Tul. (2 spp.)
  • Quiina Aubl. (~ 35 spp.)
  • Rhytidanthera (Planch.) Tieghem (5 spp.)
  • Sauvagesia L., including Lavradia Vell. (~ 40 spp.)
  • Tyleria Gleason, including Adenanthe Maguire, Steyerm. & Wurdack (15 spp.)
  • Touroulia Aubl. (2 spp.)
  • Wallacea Spruce ex Benth. & Hook.f. (3 spp.)

Useful tips for generic identification

  • Adenarake have glandular bracts, flowers with an androgynophore, and 15 staminodes, the 5 inner ones enclosing the ovary and the fertile stamens.
  • Blastemanthus has leaves with numerous close-parallel secondary veins, flowers with 8-12 sepals, 3-carpellate and 3-locular ovary with numerous ovules per locule.
  • Cespedesia, Godoya and Rhytidanthera have fimbriate appendages at the proximal and adaxial surface of bracts and sepals, 5-carpelate pistils with intrusive T-shaped placentae and sessile stigmas, and winged seeds (Dwyer 1944a). These genera also do not have staminodes. Godoya and Rhytidanthera have 5 unequal sepals, and the latter is the only genus of Ochnaceae s.s. having imparipinnate leaves.
  • Elvasia and Perissocarpa have leaves with numerous close-parallel secondary veins, and undulate margins with minute teeth, uniovulate carpels and indehiscent fruits.
  • Froesia has bisexual flowers with an apocarpous gynoecium, glabrous seeds, pinnately-compound leaves and deeply divided stipules.
  • Krukoviella comprises scandent shrubs with reticulate tertiary venation, and stamens dehiscing by a single terminal pore.
  • Lacunaria has verticillate (rarely opposite) simple leaves with well-developed stipules and parallel tertiary veins, unisexual flowers without remnants of the other floral type (dioecious plants), and 4-14-locular ovaries.
  • Luxemburgia and Philacra have zygomorphic flowers at the androecium and gynoecium levels, with the stamens grouped around one side of the pistil.
  • Poecilandra, Tyleria and Wallacea have leaves with numerous close-parallel secondary veins, and flowers with (2)3-carpellate and unilocular ovary with numerous ovules per locule.
  • Ouratea has very frequently a unique venation (eucamptodromous), with the secondary veins curving strongly near the margin and continuing almost as submarginal veins, and short intersecondary veins paralleling the basal portion of secondaries (Gentry 1996); but sometimes, the intersecondary veins are obliquely reticulate (see Maguire & Steyermark, 1989); stipules are usually early caducous; when persistent, it may be an important species diagnostic feature.
  • Quiina has opposite leaves with intersecondary veins and plumose-reticulate tertiary veins, and unisexual flowers (androdioecious plants), predominantly with a 2-locular ovary.
  • Sauvagesia are usually herbs or subshrubs with awned stipules, and white, reddish or pink flowers; staminodes are common, in 1-2 series, sometimes forming a corona.
  • Touroulia - are trees with pinnate leaves, the leaflets with numerous closely spaced and more or less parallel running tertiary veins, unisexual flowers (androdioecious plants), syncarpous gynoecium, and villous seeds.

Notable genera and distinguishing features

  • Ouratea - the fruits have a fleshy and usually reddish 'carpophore' (derived from the enlargement of the basal portion of the carpels) that carries 1-10 one-seeded and usually erect blackish mericarps; a small group of Amazonian species (e.g., O. decagyna and O. discophora) has a 'carpophore' with mericarps horizontally arranged.
  • Sauvagesia, Tyleria, Wallacea and Elvasia kollmannii Fraga & Saavedra have white or reddish petals, contrasting with the majority of the Ochnaceae s.s., which have yellow petals.

Status

  • The abovementioned genera are mostly endemic to the Neotropics (Sauvagesia has a few Paleotropical species). A few species of Ochna L. are cultivated in the Neotropics.

Important literature

Amaral, M.C.E. 1991. Phylogenetische Systematik der Ochnaceae. Bot. Jahrb. Syst. 113: 105-196.

APG. 2009. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Botanical Journal of the Linnean Society 161: 105-121.

Cardoso, D.B.O.S. & Conceição, A.A. 2008. A new acicular -leaved species of Sauvagesia (Ochnaceae) from Chapada Diamantina, Bahia, Brazil. Brittonia 60(4): 305-309.

Chacon, R.G.; Yamamoto, K. & Cavalcanti, T.B. 2003. Ochnaceae. In: Cavalcanti, T.B. & Ramos, A.E. (orgs.) Flora do Distrito Federal, Brasil. v.3. Distrito Federal, Embrapa Recursos Genéticos e Biotecnologia. Pp. 207-225.

Dwyer, J.D. 1943. The taxonomy of the monogeneric Tribe Elvasieae (Ochnaceae). Bulletin of the Torrey Botanical Club 70(1): 42-49.

Dwyer, J.D. 1944a. A discussion of the Ochnaceous genus Fleurydora A. Chev. and the allied genera of the Luxemburgieae. Bulletin of the Torrey Botanical Club 71(2): 175-178.

Dwyer, J.D. 1944b. Philacra, a new genus of the Ochnaceae. Brittonia 5: 124-127.

Dwyer, J.D. 1945. The taxonomy of the genus Sauvagesia (Ochnaceae). Bulletin of the Torrey Botanical Club 72(6): 521-540.

Dwyer, J.D. 1946. The taxonomy of Godoya R. and P., Rhytidanthera van Tieghem, and Cespedezia Goudot (Ochnaceae). Lloydia 9: 45-61.

Dwyer, J.D. 1951. The genus Luxemburgia (Ochnaceae). Lloydia 14: 82-97.

Dwyer, J.D. 1964. The taxonomy of Lavradia Vell. (Ochnaceae). Bulletin du Jardin botanique de l'État a Bruxelles 34(4): 507-518.

Dwyer, J.D. 1965. The Amazonian genus Wallacea Spruce ex Hook. f. (Ochnaceae). Bulletin du Jardin botanique de l'État a Bruxelles 35(1): 85-90.

Eichler, A. 1871. Sauvagesiaceae. In: Martius, C.F.P. & Eichler, A.G. (eds). Flora brasiliensis. Monachii, Frid. Fleischer 13(1): 398-419, tab. 81-85.

Engler, A. 1876. Ochnaceae. In: Martius, C.F.P., Eichler, A.G. & Urban, I. (eds). Flora brasiliensis. Monachii, Frid. Fleischer 12(2): 297-366, tab. 62-77.

Engler, A. 1888. Guttiferae et Quiinaceae. In: Martius, C.F.P., Eichler, A.G. & Urban, I. (eds) Flora brasiliensis. Monachii, Frid. Fleischer pp. 12(1): 381-486, tab. 109-110.

Farron, C. 1963. Contribuition à la taxonomie des Ourateae Engl. Bulletin Societe Botanique de Suisse 73: 196-217.

Feres, F. 2001. O gênero Luxemburgia A. St.-Hil. (Ochnaceae) - Revisão taxonômica e estudo cladístico. Dissertação de Mestrado, Universidade de Campinas.

Fraga, C.N. & Feres, F. 2007. Luxemburgia mysteriosa (Ochnaceae), a new species From the Atlantic Rain Forest of Espírito Santo, Brazil. Harvard Papers in Botany 12(2): 405-408.

Fraga, C.N. & Saavedra, M.M. 2006. Three new species of Elvasia (Ochnaceae) from the Brazilian Atlantic Forest, with an emended key for Subgenus Hostmannia. Novon 16: 483-489.

Gentry, A.H. 1996. A Field Guide to the Families and Genera of Woody Plants of North west South America: (Colombia, Ecuador, Peru). University of Chicago Press, Chicago.

Gleason. 1931. Botanical results of the Tyler-Duida Expedicion. Bulletin of the Torrey Botanical Club 58: 277-506.

Guimarães, E.F. & Pereira, J.M.C. 1966. Ochnaceae no Estado da Guanabara. Rodriguésia 25(37): 59-65.

Harley, R.M., A.M. Giulietti & K.R.B. Leite. 2005. Two new species and a new record of Sauvagesia (Ochnaceae) in the Chapada Diamantina of Bahia, Brazil. Kew Bulletin 60: 571-580.

Maguire, B. & Steyermark, J. 1989. Ouratea (Ochnaceae) in Guayana and adjacent Amazonian hylea. Botany of Guayana Highland. XIII. Memoirs of the New York Botanical Garden 51: 56-102.

Macbride, J.F. 1956. Flora of Peru: Ochnaceae. Field Museum of Natural History-Botany 13(3a/2): 686-697.

Rocha, A.E.S. & Secco, R.S. 2004. Uma sinopse de Lacunaria Ducke. Acta Amazonica 34: 425-433.

Sastre, C. 1981. Ochnacées nouvelles du Brésil. Bulletin du Jardin Botanique National de la Bélgique 51 (3/4): 347-413.

Sastre, C. 1988. Studies on the Flora of the Guianas 34. Synopsis generis Ouratea Aublet (Ochnaceae). Bulletin du Muséum National d'Histoire Naturelle, Paris 4e. sér., 10, section B, Adansonia 1: 47-67.

Sastre, C. 1995a. Novelties in the neotropical genus Ouratea Aublet (Ochnaceae). Novon 5: 193-200.

Sastre, C. 1995b. Ochnaceae. In: Stannard, B.L.; Harvey, Y.B. & Harley, R.M. (eds.) Flora of the Pico das Almas: Chapada Diamantina-Bahia, Brazil. Royal Botanic Gardens, Kew.  Pp. 519-522.

Sastre, C. 1997. Uma espécie nova de Sauvagesia L.(Ochnaceae) do campo rupestre do Estado de Goiás. Boletim de Botânica da Universidade de São Paulo 16: 71-73.

Sastre, C. 2001. New Ouratea species (Ochnaceae) from Venezuela and adjacent countries. Novon 11: 105-118.

Sastre, C. 2003. Ochnaceae. In: Steyermark J.A., Berry, P.E., Yatskievych, K. & Holst, B.K. (eds.) Flora of the Venezuelan Guayana, vol. 7. Missouri Botanical Garden, Saint Louis.  Pp. 124-161

Sastre, C. 2004. Une nouvelle espèce d'Ouratea (Ochnaceae) du Venezuela. Adansonia ser. 3, 29(1): 77-91.

Sastre, C. 2005. Une nouvelle espèce d'Ouratea (Ochnaceae) de l'Amazonie Brésilienne. Adansonia ser. 3, 27(1): 85-88.

Sastre, C. 2006. Deux nouvelles espèces d'Ouratea (Ochnaceae) des Guyanes. Adansonia sér. 3, 28(1) : 119-127.

Sastre, C. 2007. Six nouvelles espèces d'Ouratea (Ochnaceae) des Guyanes. Adansonia ser. 3, 29(1): 77-91.

Sastre, C., Whitefoord, C. & Knapp, S. 1999. A new species of Elvasia (Ochnaceae) from Mesoamerica with discussion of subgeneric classification and phytogeography. Novon 9: 253-256.

Sastre, C. & Offroy, B. 2009. Description de trois nouveaux Ouratea L. (Ochnaceae) du Paraguay, de Bolivie et d'Équateur. Considérations taxonomiques, nomenclaturales et biogéographiques sur les espèces affi nes d'O. superba Engl. Adansonia ser. 3, 31(1): 89-101.

Schneider, J.V. 1998. El género Quiina (Quiinaceae), con especial referencia a las especies de Venezuela. Acta Botánica Venezuélica 21: 1-74.

Schneider, J.V. & Zizka, G. 2004. Quiinaceae. In: Berry, P.E., Yatskievych, K. & Holst, B.K. (eds.) Flora of the Venezuelan Guayana, vol. 8. Missouri Botanical Garden, Saint Louis. Pp. 393-407.

Schneider, J.V., Swenson, U. & Zizka, G. 2002. Phylogenetic reconstruction of the Neotropical family Quiinaceae (Malpighiales) based on morphology with remarks on the evolution of an androdioecious sex distribution. Ann. Missouri Bot. Gard. 89: 64-76.

Schneider, J.V., Swenson, U., Samuel, R., Stuessy, T. & Zizka, G. 2006. Phylogenetics of Quiinaceae (Malpighiales): evidence from trnL-trnF sequence data and morphology. Plant Syst. Evol. 257: 189-203.

Steyermark, J.A. & Bunting, G.S. 1975. Revision of the genus Froesia (Quiinaceae). Brittonia 27(2): 172-178.

Steyermark. J. & Maguire, B. 1984. Ochnaceae. In: Steyermark, J.A. Flora of the Venezuelan Guayana-I. Annals of the Missouri Botanical Gardens 71(1): 319-323.

Tieghem, P. Van. 1902. Sur les Ochnacées. Annales Sciences Naturelles Botanique, sér. 8e, 16: 161-416.

Wallnöffer, B. 1998. A revision of Perissocarpa Steyerm. & Maguire (Ochnaceae). Ann. Naturhist. Mus. Wien 100 B: 683-707.

Woodson, R.E., Schery, R.W. & Dwyer, J.D. 1967. Flora of Panama. Part VI. Family 119. Ochnaceae. Annals of the Missouri Botanical Garden 54(1): 25-40.

Yamamoto, K. 1989. Morfologia, anatomia e sistemática do genero Ouratea Aublet (Ochnaceae) : levantamento preliminar das características de importância taxonômica e avaliação das classificações vigentes. Dissertação de Mestrado, UNICAMP.

Yamamoto, K. 1995a. Estudos taxonômicos sobre Ouratea parviflora (DC.) Baill (Ochnaceae) e especies afins ocorrentes em floresta atlântica nas regiões sudeste e sul do Brasil. Tese de Doutorado, UNICAMP.

Yamamoto, K. 1995b. Ouratea hatschbachii (Ochnaceae): Uma nova espécie de Grão-Mogol, Estado de Minas Gerais. Boletim de Botânica da Universidade de São Paulo 14: 33-37.

Yamamoto, K. & Sastre, C. 2004. Flora de Grão Mogol, Minas Gerais: Ochnaceae. Boletim de Botânica da Universidade de São Paulo 22(2): 343-348.

Yamamoto, K., Chacon, R.G., Proença, C., Cavalcanti, T.B., and Graciano-Ribeiro, D.A. 2008. A distinctive new species of Ouratea (Ochnaceae) from the Jalapão Region, Tocantins, Brazil. Novon 18(3): 397-404.

Zappi, D.C. & Lucas, E. 2002. Sauvagesia nitida Zappi & E. Lucas (Ochnaceae): a new species from Catolés, Bahia, NE Brazil. Kew Bulletin 57(3): 711-717.

Zizka, G. & Schneider, J.V. 1999. The genus Touroulia Aubl. (Quiinaceae). Willdenowia 29: 227-234.

How to cite

Fiaschi, P., Nicoletti de Fraga, C. & Yamamoto, K. (2010). Neotropical Ochnaceae s.l. (incl. Quiinaceae). 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/Ochnaceae_s.l._(incl._Quiinaceae).htm.

Click images to enlarge


Flower of Elvasia kollmannii © Cláudio Nicoletti de Fraga, Instituto de Pesquisas Jardim Botânico do Rio de Janeiro.



Branches with inflorescences of Luxemburgia mysteriosa © Cláudio Nicoletti de Fraga, Instituto de Pesquisas Jardim Botânico do Rio de Janeiro.



Inflorescence of Luxemburgia damazioana ©  Gustavo Shimizu, Universidade Estadual de Campinas, Campinas.



Flowers of Ouratea acuminata © Cláudio Nicoletti de Fraga, Instituto de Pesquisas Jardim Botânico do Rio de Janeiro.



Inflorescence of Ouratea aff. hatschbachii © Abel A. Conceição, Universidade Estadual de Feira de Santana.



Infructescence of Ouratea aff. spruceana © Pedro Fiaschi, Universidade de São Paulo, São Paulo.



Flower and buds of Ouratea castaneifolia © Gustavo Shimizu, Universidade Estadual de Campinas, Campinas.



Flowers showing detail of ovaries of Ouratea caudata © Cláudio Nicoletti de Fraga, Instituto de Pesquisas Jardim Botânico do Rio de Janeiro.



Infructescence of Ouratea discophora © Fernanda Antunes Carvalho, Instituto Nacional de Pesquisas da Amazônia (INPA-CPBO).



Habit with inflorescences of Ouratea floribunda © Gustavo Shimizu, Universidade Estadual de Campinas, Campinas.



Flower of Ouratea humilis © Gustavo Shimizu, Universidade Estadual de Campinas, Campinas.



Flower of Ouratea sp. © Cláudio Nicoletti de Fraga, Instituto de Pesquisas Jardim Botânico do Rio de Janeiro.



Flowers of Ouratea sp. © Abel A. Conceição, Universidade Estadual de Feira de Santana.



Inflorescence of Poecilandra retusa © Pedro Fiaschi, Universidade de São Paulo, São Paulo.



Branch with inflorescence of Quiina glaziovii © Wm. Wayt Thomas, The New York Botanical Garden, Bronx, New York.



Branch with flowers of Sauvagesia erecta © Cláudio Nicoletti de Fraga, Instituto de Pesquisas Jardim Botânico do Rio de Janeiro.



Flower of Sauvagesia erecta © Abel A. Conceição, Universidade Estadual de Feira de Santana.



Flower of Sauvagesia erecta (Guyana) © Mac H. Alford, University of Southern Mississippi.



Inflorescence of Sauvagesia nitida © Abel A. Conceição, Universidade Estadual de Feira de Santana.



Plant with inflorescences of Sauvagesia oliveirae © Abel A. Conceição, Universidade Estadual de Feira de Santana.



Flowers of Sauvagesia sp. © Abel A. Conceição, Universidade Estadual de Feira de Santana.



Infructescence of Sauvagesia sp. © Abel A. Conceição, Universidade Estadual de Feira de Santana.



Branch with inflorescences of Wallacea sp. © Mário Terra, Instituto Nacional de Pesquisas da Amazônia (INPA-CPBO); Projeto Flórula de São Gabriel da Cachoeira.