1. Chrysobalanaceae R.Br.

    1. This family is accepted.

[FZ]

Chrysobalanaceae, F. White. Flora Zambesiaca 4. 1978

Habit
Trees, shrubs or rhizomatous, geoxylic suffrutices
Wood
Wood always with abundant silica inclusions
Leaves
Leaves simple, entire, alternate, often coriaceous, usually with two glands at base of lamina or near apex of petiole
Stipules
Stipules small and caducous to large and persistent
Inflorescences
Inflorescence a cyme, panicle or raceme
Flowers
Flowers mostly (in our area always) bisexual, actinomorphic to zygomorphic, strongly perigynous
Receptacle
Receptacle-tube short to elongate, straight or curved, often gibbous at the base, always lined with nectariferous tissue which is extended at the throat as a short annular disk; throat at least partly blocked by long hairs
Calyx
Sepals 5, free, imbricate, often unequal, ascending or reflexed
Corolla
Petals 5, rarely absent (not in our area), sometimes unequal, imbricate, often caducous
Androecium
Stamens 2–100 or more, included or exserted, inserted in 1 or 2 rows on the margin of the disk or adnate to its abaxial surface, either all fertile and forming a complete circle or partly staminodial; filaments free or appearing connate at the base or (not in our area) ligulately connate; anthers small, 2-thecous, dehiscing longitudinally
Gynoecium
Ovary superior, basically of 3 carpels and gynobasic but usually with only 1 carpel fully developed, attached to base, middle or mouth of receptacle-tube, sessile or on a short gynophore, always hairy, each carpel 1-locular with 2 ovules, or 2-locular, owing to a false septum, with 1 ovule in each compartment; style filiform; stigma distinctly or indistinctly 3-lobed
Fruits
Fruit a dry or fleshy drupe; endocarp thick or thin, fibrous, granular or bony, often with a special mechanism for seedling escape, often densely hairy inside
Seeds
Seed erect, exalbuminous; cotyledons plano-convex, fleshy, sometimes ruminate
Germination
Germination hypogeal or epigeal; first leaves of seedling opposite or alternate
[NTK]

Prance, G.T. & Sothers, C. (2009). Neotropical Chrysobalanaceae.

Morphology
Description

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 .

Distribution
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.
Diagnostic
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 foliaceouscaducous stipules.
  • Hirtella: a few species have ant domatia at the base of the leaf blade.
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).
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.
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.
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.
General Description
General notes
  • 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).
Status
  • All taxa are native and endemic.
  • Two taxa occur outside the Neotropics: Chrysobalanusicaco L.and Parinariexcelsa Sabine.
  • Cultivated taxa include Neocaryamacrophylla (Sabine) Prance ex F. White and Maranthes corymbosa Blume.
  • Couepia, Acioa and Exellodendron are endemic genera to the Neotropics.
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).
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.
Literature
Important literature

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.

Images

Chrysobalanaceae R.Br. appears in other Kew resources:

First published in Narr. Exped. Zaire 433. 1818 [5 Mar 1818] (as "Chrysobalaneae") (1818)

Accepted by

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

Sources

Flora Zambesiaca
Flora Zambesiaca
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 2019. Published on the Internet at http://www.ipni.org and http://apps.kew.org/wcsp/
© 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.
http://creativecommons.org/licenses/by/3.0