| # The genera of Leguminosae-Caesalpinioideae and Swartzieae |

L. Watson and M.J. Dallwitz

### Copaifera L.

Copaiba Adans., Copaiva Jacq.; including Pseudosindora Symington

Type species: C. officinalis (Jacqu.) L.

Habit and leaf form. Trees, or shrubs; unarmed.

Phyllotaxy distichous, or spiral. The leaves compound; pinnate; imparipinnate, or paripinnate. The leaflets many per leaf to few per leaf; opposite or sub-opposite, or alternate; petiolulate; with markedly twisted petiolules, or without noticeably twisted petiolules; pinnately veined, symmetrical or nearly so; with a predominant ‘midrib’; with a strong, continuous marginal nerve. Stipules absent or early caducous or very inconspicuous in mature leaves; membranous. Stipels absent.

Inflorescence and floral morphology. The inflorescences unbranched, or branched; when branched, of racemose units; simple racemes, or panicles. The flowers distichous, or not distichous (rarely: Pseudosindora). Bracts absent at anthesis. Bracteoles present; small, not enclosing the flower buds; absent at anthesis; not valvate; free.

The flowers hermaphrodite; not pentamerous throughout; departing from pentamery in the calyx and in the corolla, or in the calyx, in the corolla, and in the androecium (rarely); white or green. Hypanthium absent, the androecium hypogynous. The perianth comprising distinct calyx and corolla (rarely), or exclusively sepaline. Calyx 4; covering the rest of the flower in bud; polysepalous; more or less regular; members imbricate (but subvalvate). Corolla present (rarely), or absent; if present,vestigial, 1–4; including greatly reduced members; when present, polypetalous, or monopetalous. Petals sessile. Disk present and conspicuous. The androecium comprising about 7–15 members (?); members all free of one another; members all more or less equal in length, or markedly unequal; comprising only fertile stamens. Fertile stamens 10, or 8–13 (rarely). Anthers attached well above the base of the connective; dehiscing longitudinally. Ovary sessile or subsessile, or stipitate; free (on the disc). Stigma conspicuously dilated, or not dilated. Ovules few.

Fruit, seed and seedling. Fruit a two-valved pod (almost round, flattened and shortly beaked); not drupaceous; without markedly twisting or enrolling valves; not noticeably winged; not becoming woody. The mature valves with conspicuous, prominent, raised venation, or without prominent venation; conspicuous venation not predominantly longitudinal. Seeds non-endospermic; arillate; with a straight or slightly oblique radicle; amyloid-positive. Cotyledons flat; epigeal.

Transverse section of lamina. Leaves with conspicuous phloem transfer cells in the minor veins. Druses absent from the mesophyll. Mesophyll secretory cavities (gland-dots) common; epithelium-lined. Adaxial hypodermis absent. Leaf girders common (the veins transcurrent). Laminae dorsiventral. Mesophyll without unaligned fibres or sclereids. Minor veins mainly with abundant accompanying fibres.

Leaf lamina epidermes. Epidermal crystals present; prisms. Simple unbranched hairs common; smooth. No compound or branched eglandular hairs seen. Capitate glands not seen. Hooked hairs not seen. Cassieae-type leaf pseudo-glands not seen. Expanded and embedded hair-feet absent. Basally bent hairs present, or absent. Adaxial: Adaxial interveinal epidermal cell walls straight in optical section, or markedly sinuous in high-focus optical section; conspicuously pitted, or not conspicuously pitted. Stomata adaxially very rare. Abaxial: Abaxial stomata predominantly paracytic. Abaxial epidermis not papillate. Abaxial interveinal epidermal cell walls markedly sinuous in high-focus optical section; conspicuously pitted in optical section; staining normally with safranin.

Wood anatomy. Wood without septate fibres; not storied; with normal intercellular canals; without traumatic canals. Intervascular pits very small, or medium to large.

Pollen ultrastructure. Tectum punctate; smooth punctate, or puncticulate. Length of colpi greater than one half pole to pole distance (without a margocolpus).

Cytology. Basic chromosome number, x = 12. 2n = 24.

Species number and distribution. About 45 species. Tropical America, tropical and southern Africa.

Tribe. Detarieae; Detarieae sensu stricto clade of Bruneau et al. (2008).

Comments. Widely cultivated.

Miscellaneous. Illustrations: • Copaifera langsdorfii and C. martii: Fl. Brasiliensis 15 (1870). • Copaifeera officinalis: Baillon, Histoire des Plantes 2 (1870). • Copaifera mildbraedii and C. religiosa: Aubréville, Flore du Gabon (1968). • C. langsdorfii (as Copaiba): Nat. Pflanzenfam. III (1894). • C. baumiana and Guibourtia coleosperma: Baum (1903). • Pseudosindora (Copaifera) palustris leaflet: abaxial epidermis.

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, and classification. See also Guidelines for using data taken from Web publications.

Cite this publication as: ‘Watson, L., and Dallwitz, M.J. 1993 onwards. The genera of Leguminosae-Caesalpinioideae and Swartzieae: descriptions, illustrations, identification, and information retrieval. In English and French. Version: 4th August 2019. delta-intkey.com’.

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