| # The Families of Angiosperms |

L. Watson and M.J. Dallwitz

### Leonticaceae (Spach) Airy Shaw

~ Berberidaceae, Podophyllaceae.

Habit and leaf form. Glaucous herbs. Plants succulent, or non-succulent. Perennial; with a basal aggregation of leaves; rhizomatous, or tuberous. Leaves alternate; ‘herbaceous’, or fleshy; compound; pinnate, or palmate, or bipinnate, or multiply compound (2–3 pinnate); when pinnate, imparipinnate. Lamina pinnately veined. Leaves stipulate. Stipules intrapetiolar (on the petiole); caducous.

Leaf anatomy. The leaf lamina dorsiventral. Stomata present; anomocytic (at least in Leontice).

Axial (stem, wood) anatomy. Secondary thickening absent.

The vessel end-walls simple.

Reproductive type, pollination. Unisexual flowers absent. Plants hermaphrodite. Floral nectaries present. Nectar secretion from the perianth (the corolla often nectariform). Pollination entomophilous.

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’; in racemes (or compound racemes). The ultimate inflorescence units racemose. Inflorescences simply or compoundly racemose. Flowers more or less regular; 3 merous; cyclic; pentacyclic, or polycyclic. Free hypanthium absent. Hypogynous disk absent.

Perianth with distinct calyx and corolla; 9–15; 3–5 whorled; isomerous, or anisomerous. Calyx 3–9 (the inner members often petaloid); 1–3 whorled; polysepalous; imbricate. Corolla 6; 2 whorled; polypetalous (often nectariform); yellow.

Androecium 6; exclusively of fertile stamens (unless the nectariferous petals are interpreted as staminodes). Stamens 6; isomerous with the perianth; alternisepalous; opposite the corolla members. Anthers dehiscing by longitudinal valves; extrorse. Endothecium developing fibrous thickenings. Microsporogenesis simultaneous. Pollen grains aperturate; 3 aperturate; colpate.

Gynoecium ostensibly 1 carpelled. Carpels reduced in number relative to the perianth. The pistil 1 celled. Gynoecium monomerous (or pseudomonomerous?); ostensibly of one carpel; superior. Carpel shortly stylate; apically stigmatic (the stigma small or plicate-dilated); 2–8 ovuled. Placentation basal. Ovules ascending; anatropous; bitegmic; pseudocrassinucellate. Outer integument contributing to the micropyle. Embryo-sac development Polygonum-type. Endosperm formation nuclear.

Fruit bladdery, non-fleshy. The fruiting carpel dehiscent (or gaping above, or the walls evanescent), or indehiscent. Seeds endospermic; drupelike, large (often blue). Embryo straight.

Physiology, phytochemistry. C3. C3 physiology recorded directly in Leontice. Anatomy non-C4 type (Leontice). Cyanogenic.

Geography, cytology. Holarctic. Temperate. North temperate.

Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgren’s Superorder Ranunculiflorae; Ranunculales. Cronquist’s Subclass Magnoliidae; Ranunculales. APG III core angiosperms; peripheral eudicot; Superorder Ranunculanae. APG IV Order Ranunculales (as a synonym of Berberidaceae).

Species 14. Genera 4; Leontice, Bongardia, Caulophyllum, Gymnospermium.

General remarks. Seemingly differing cryptically from Berberidaceae sensu stricto (q.v.) in being cyanogenic, in the colpate pollen and the pseudocrassinucelate ovules, as well as in the non-fleshy fruit.

Illustrations. • Gymnospermium albertii (as Leontice): Bot. Mag. 112 (1886). • Gymnospermium altaicum (as Leontice): Bot. Mag. 60 (1833). • Carpel, fruit and seed (Caulophyllum). • Bongardia chrysogonum, as B. rauwolfii: Bot. Mag. 102 (1876).

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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