| # The Families of Angiosperms |

L. Watson and M.J. Dallwitz

### Caryocaraceae Szyszyl.

Alternatively Rhizobolaceae.

Including Simabaceae Horan. (p.p.).

Habit and leaf form. Trees (mostly), or shrubs (or undershrubs). Leaves persistent; alternate (Anthodiscus), or opposite (Caryocar); leathery; petiolate; compound; digitately ternate, or bipinnate (to 5-foliolate). Lamina palmately veined; cross-venulate. Leaves stipulate, or exstipulate. Stipules when present, caducous. Lamina margins sub entire, or serrate to dentate. Domatia occurring in the family; manifested as hair tufts.

Leaf anatomy. The leaf lamina dorsiventral, or centric. Extra-floral nectaries present (on the young leaf margins and stipules). Hydathodes commonly present. Mucilaginous epidermis present, or absent. Stomata present; mainly confined to one surface (abaxial); anomocytic, or anisocytic, or paracytic. Hairs present (infrequent), or absent. Adaxial hypodermis present, or absent. The mesophyll with sclerenchymatous idioblasts; without crystals.

Axial (stem, wood) anatomy. Young stems with solid internodes. Secretory cavities absent. Cork cambium present; initially superficial. Nodes multilacunar. Primary vascular tissues in a cylinder, without separate bundles. Internal phloem absent. Cortical bundles absent. Medullary bundles absent. Secondary thickening developing from a conventional cambial ring.

The wood diffuse porous. The vessels small, or medium, or large; solitary, radially paired, and in radial multiples. The vessel end-walls oblique; simple, or scalariform and simple. The vessels without vestured pits; without spiral thickening. The axial xylem with tracheids; without vasicentric tracheids; with fibre tracheids, or without fibre tracheids; with libriform fibres; including septate fibres (rarely), or without septate fibres. The fibres without spiral thickening. The parenchyma apotracheal, or paratracheal. The secondary phloem not stratified. ‘Included’ phloem absent. The wood not storied. Tyloses present, or absent.

Reproductive type, pollination. Unisexual flowers absent. Plants hermaphrodite. Pollination chiropterophilous (in Caryocar).

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’; in racemes (these sometimes condensed). The ultimate inflorescence units racemose. Inflorescences terminal. Flowers ebracteate; large; calyptrate (Anthodiscus), or not calyptrate; regular. Free hypanthium present to absent (the stamens ‘subperigynous’).

Perianth with distinct calyx and corolla; 10 (usually), or 12; 2 whorled; isomerous. Calyx 5(–6); 1 whorled; gamosepalous; regular; imbricate, or open in bud. Corolla 5(–6); 1 whorled; polypetalous, or gamopetalous (basally). Corolla lobes markedly longer than the tube. Corolla calyptrate (in Anthodiscus), or not calyptrate; imbricate; regular.

Androecium 50–200 (‘very many’). Androecial members branched (?); maturing centrifugally; free of the perianth; coherent; 1 adelphous, or 5 adelphous (shortly connate basally, into a ring or five bundles alternating with the corolla members). The androecial bundles when bundled, alternating with the corolla members. Androecium exclusively of fertile stamens, or including staminodes (the inner filaments sometimes without anthers). Staminodes internal to the fertile stamens. Stamens 50–200 (‘very many’); polystemonous; oppositisepalous. Anthers dorsifixed; sub versatile; dehiscing via longitudinal slits. Pollen grains aperturate; (2–)3(–6) aperturate; (syn) colporate, or rugate.

Gynoecium 4–20 carpelled. Carpels isomerous with the perianth to increased in number relative to the perianth. The pistil 4–20 celled. Gynoecium syncarpous; synovarious; superior. Ovary 4–20 locular (equalling G). Gynoecium stylate. Styles 4–20; free; apical. Stigmas 4–20. Placentation axile. Ovules 1 per locule; ascending (Hutchinson, Thonner); orthotropous to anatropous; bitegmic.

Fruit fleshy, or non-fleshy; indehiscent, or a schizocarp (then leathery). Mericarps 4–20 (?). Fruit usually a drupe (the fleshy mesocarp sometimes poisonous, sometimes edible). The drupes with separable pyrenes (‘mericarps’). Seeds thinly endospermic, or non-endospermic. Embryo well differentiated. Cotyledons 2 (small). Embryo with an enlarged, oily and proteinaceous, spirally twisted hypocotyl, and inflexed cotyledons.

Physiology, phytochemistry. Not cyanogenic. Arbutin absent.

Geography, cytology. Neotropical. Tropical. Tropical America.

Taxonomy. Subclass Dicotyledonae; Crassinucelli (?). Dahlgren’s Superorder Theiflorae; Theales. Cronquist’s Subclass Dilleniidae; Theales. APG III core angiosperms; core eudicot; Superorder Rosanae; fabid. APG IV Order Malpighiales.

Species 25. Genera 2; Anthodiscus, Caryocar.

Illustrations. • Caryocar glabrum: Hutchinson. • Anthodiscus obovatus: Fl. Bras. 12 (1858). • Caryocar villosum as C. butyrosum, and Anthodiscus trifoliatus: Lindley.

We advise against extracting comparative information from the descriptions. This is much more easily achieved using the DELTA data files or the interactive key, which allows access to the character list, illustrations, full and partial descriptions, diagnostic descriptions, differences and similarities between taxa, lists of taxa exhibiting or lacking specified attributes, distributions of character states within any set of taxa, geographical distribution, genera included in each family, and classifications (Dahlgren; Dahlgren, Clifford, and Yeo; Cronquist; APG). See also Guidelines for using data taken from Web publications.

Cite this publication as: ‘Watson, L., and Dallwitz, M.J. 1992 onwards. The families of flowering plants: descriptions, illustrations, identification, and information retrieval. Version: 8th June 2026. delta-intkey.com’.

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