Ghillean T. Prance & Cynthia Sothers
Royal Botanic Gardens, Kew, UK.
Trees and shrubs. Leaves simple, entire, alternate; stipules small and caducous to large and persistent; lamina often coriaceous, with lower surface glabrous or hairs simple. Inflorescence racemose, paniculate or cymose; flowers bracteate and usually 2-bracteolate. Flowers actinomorphic to zygomorphic, hermaphrodite, markedly perigynous; receptacle short to elongate, sometimes gibbous at the base; disc always present, forming a lining to the receptacle or an annular or shortly tubular structure at its mouth; calyx lobes 5, imbricate, often unequal, erect or reflexed; petals 5, occasionally absent, commonly unequal, imbricate, usually caduceus; stamens 2-100 (-300), inserted on margin or surface of disc, or basally adnate to it, forming a complete circle or unilateral, all fertile or some without anthers and then often reduced to small staminodes; filaments filiform, free, connate at the base or ligulately connate, included to far-exserted; anthers small, dorsifixed, longitudinally dehiscent, glabrous; gynoecium of 1-3 carpels, gynobasic, usually with only 1 carpel fully developed, attached to the base, middle or mouth of the receptacle-tube, sessile or with a short gynophore, pubescent or villous; carpel unilocular with 2 ovules or falsely bilocular with 1 ovule in each compartment; style filiform, arising from the receptacle at the base of the carpel(s); stigma distinctly or indistinctly 3-lobed; ovules erect, epitropous, with micropyle directed towards base. Fruit a fleshy to dry drupe; endocarp various, thick or thin, fibrous or bony, often with a special mechanism for seedling escape, often densely hairy inside; seed 1, rarely 2, erect, almost exalbuminous; cotyledons planoconvex, fleshy, sometimes ruminate.
Notes on delimitation
- Originally classified as a subfamily of the Rosaceae, presently elevated to family level. Molecular sequence data place it in Eurosids I, order Malpighiales, sister to the Trigoniaceae, Euphroniaceae and Dichapetalaceae. The Chrysobalanaceae as presently circumscribed are monophyletic.
Distribution in the Neotropics
From SE USA, the Caribbean and Mexico to southern Brazil and Paraguay. 422 species in 8 genera.
- Acioa Aubl.: tropical northern South America.
- Chrysobalanus L.: Mexico, Central America, the Caribbean and South America, mainly coastal apart from 1 species.
- Couepia Aubl. and Hirtella L.: widespread throughout Neotropics, including Caribbean (Hirtella only).
- Exellodendron Prance: central and eastern tropical South America.
- Licania Aubl.: from SE USA and widespread throughout Neotropics, including Caribbean.
- Maranthes Blume: Costa Rica, Panama and Nicaragua.
- Parinari Aubl.: Central America and throughout South America.
Distinguishing characters (always present)
- Gynobasic ovary; stipules (although often caducous); pair of glands on petiole or base of leaf; wood has a high content of silica (grainy to the touch); many species exude red resin (but only from main trunk); fruit a drupe (1-seeded).
Other important characters
- Receptacle-tube forming a short or elongate tube (mainly Couepia, Hirtella, Maranthes), stamens often unilateral, ovary at mouth of receptacle-tube, leaf lower surface often with some type of indumentum (hairs always simple) lanate, strigose, appressed, sometimes glabrous.
Key differences from similar families
- Gynobasic ovary (with style arising from base of ovary) distinguishes it from the Dichapetalaceae, and other families with stipules and foliar glands, fruit a drupe often woody (except for Hirtella which are all fleshy), epicarp often hairy.
Number of genera
Eight genera and number of corresponding species:
- Acioa (3-4)
- Chrysobalanus (3)
- Couepia (71)
- Exellodendron (5)
- Hirtella (105)
- Licania (215)
- Maranthes (1)
- Parinari (18).
Useful tips for generic identification
- Number of stamens, floral symmetry and ovary placement (Licania and Chrysobalanus sub-actinomorphic, ovary at base of receptacle-tube; Parinari few stamens 6-10, zygomorphic, ovary at side or mouth of receptacle-tube; Couepia 15-300 stamens, zygomorphic,ovary at mouth of receptacle-tube; Hirtella 3-9 stamens, zygomorphic,ovary at mouth of receptacle-tube; Maranthes Exellodendron Acioa usually fused filaments (one species with free filaments), leaf glabrous and coriaceous.
- Stamen attachment to disc (Licania and Chrysobalanus in a complete circle; Couepia in a complete circle or unilateral, Parinari, Acioa and Hirtella unilateral.
- Parinari: leaf lower surface lanate and with hair-filled stomatal cavities and secondaries closely spaced, pair of glands on petioles; fruit large and woody.
- Hirtella: c. 7 species with and domatia at base of leaf; leaves with glands, stipules.
Notable genera and distinguishing features
- Acioa: only neotropical genus with fused stamens, forming a ligule (except A. edulis Prance).
- Parinari: leaf venation (parallel secondaries), stomatal cavities on abaxial leaf surface, large foliaceous caducous stipules.
- Hirtella: a few species have ant domatia at the base of the leaf blade.
- All taxa are native and endemic.
- Two taxa occur outside the Neotropics: Chrysobalanus icaco L.and Parinari excelsa Sabine.
- Cultivated taxa include Neocarya macrophylla (Sabine) Prance ex F. White and Maranthes corymbosa Blume.
- Couepia, Acioa and Exellodendron are endemic genera to the Neotropics.
- The Chrysobalanaceae has many species with edible fruit, used locally, as well as hard resistant wood also used locally (not commercialized due to high silica content).
- Fruit are dispersed mainly by mammals.
- Two species are known to be bat-pollinated and dispersed (Couepia).
Prance, G.T. (1972). Chrysobalanaceae. Fl. Neotrop. Monogr. 9: 1-409.
Prance, G.T. (1989). Chrysobalanaceae. Fl. Neotrop. Monogr. 9S: 1- 267.
Prance, G.T. & Sothers, C.A. (2003). Chrysobalanaceae I: Chrysobalanus to Parinari. Species Plantarum: Flora of the World Part 9: 1-319.
Prance, G.T. & Sothers, C.A. (2003). Chrysobalanaceae II: Acioa to Magnistipula. Species Plantarum: Flora of the World Part 10: 1-268.
Prance, G.T. & White, F. (1988). The genera of Chrysobalanaceae. Philos. Trans. B320 (1197): 1-184.
How to cite
Prance, G.T. & Sothers, C. (2009). Neotropical Chrysobalanaceae. 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/Chrysobalanaceae.htm.
Click images to enlarge
Couepia grandiflora © Peter Gasson, RBG, Kew.
Couepia paraensis © Denise Sasaki, Programa Flora Cristalino.
Slash of Hirtella gracilipes © Denise Sasaki, Programa Flora Cristalino.
Flowering branch of Hirtella racemosa © A Gallo, Programa Flora Cristalino.
Fruiting branch of Hirtella racemosa © Denise Sasaki, Programa Flora Cristalino.
Trunk of Licania brittoniana © Denise Sasaki, Programa Flora Cristalino.
Budding branch of Licania gardneri © Denise Sasaki, Programa Flora Cristalino.
Fruiting branch of Licania polita © Denise Sasaki, Programa Flora Cristalino.
Flowering branch of Licania sclerophylla © Denise Sasaki, Programa Flora Cristalino.
Parinari excelsa © Denise Sasaki, Programa Flora Cristalino.