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The families of flowering plants

L. Watson and M.J. Dallwitz

Berberidaceae Juss.

Including Nandinaceae Horan. (non J.G. Agardh); excluding Nandinaceae J.G. Agardh, Leonticaceae, Podophyllaceae.

Habit and leaf form. Shrubs (mostly, usually with distinct ‘long-’ and ‘short-’ shoots, the tissues commonly coloured yellow with berberine), or herbs (more or less, Epimedium). Mesophytic, or xerophytic. Commonly heterophyllous (the long-shoot leaves often spiny or transformed into spines), or not heterophyllous. Leaves persistent, or deciduous; small to large; alternate; spiral; flat; ‘herbaceous’, or leathery, or modified into spines (especially on the long-shoots, the spines simple or branched); petiolate; non-sheathing; simple, or compound, or simple and compound; when compound, unifoliolate (the ‘simple’ form of Berberis perhaps representing this), or ternate, or pinnate; when pinnate, imparipinnate. Lamina when simple, dissected, or entire; commonly spinose; pinnately veined; cross-venulate. Leaves stipulate (the stipules minute or vestigial), or exstipulate. Stipules when present, intrapetiolar; caducous. Lamina margins entire, or dentate (usually prickly). Leaf development not ‘graminaceous’.

Leaf anatomy. Stomata mainly confined to one surface (abaxial); anomocytic. Hairs present, or absent (infrequent); eglandular (simple unicellular or uniseriate), or glandular (very rarely, but glandular hairs with a uniseriate stalk and ellipsoidal head recorded in Epimedium). Minor leaf veins without phloem transfer cells (Berberis, Epimedium, Mahonia).

Axial (stem, wood) anatomy. Young stems with solid internodes. Pith more or less homogeneous (the peripheral cells tending to be thicker-walled). Cork cambium present; initially deep-seated, or initially superficial. Nodes tri-lacunar, or multilacunar (e.g. Epimedium, Mahonia). Primary vascular tissues comprising a ring of bundles, or consisting of scattered bundles (in herbaceous and semi-herbaceous forms); collateral. Secondary thickening absent (Epimedium, Vancouveria), or developing from a conventional cambial ring. Primary medullary rays wide.

The wood of Berberis and Mahonia ring porous, or semi-ring porous, or diffuse porous. The vessels very small; commonly clustered, or in radial multiples. The vessel end-walls scalariform, or simple. The vessels with spiral thickening. The axial xylem with libriform fibres; occasionally including septate fibres (in Berberis), or without septate fibres. The parenchyma absent; wood partially storied (VPI), or not storied. Tyloses absent.

Reproductive type, pollination. Plants hermaphrodite. Floral nectaries present. Nectar secretion from the perianth, or from the androecium (from the ‘honey-leaves’ of the perianth, which are interpretable as staminodial). Pollination entomophilous; mechanism conspicuously specialized (the stamens springing violently upwards when their irritable bases are touched).

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’; in cymes, or in racemes, or in fascicles. The ultimate inflorescence units cymose, or racemose. Inflorescences racemes, fascicles and cymes. Flowers small, or medium-sized; regular; (2–)3(–4) merous; cyclic; polycyclic (usually of 6–7(-9) cycles). Floral receptacle not markedly hollowed (short). Free hypanthium absent. Hypogynous disk absent.

Perianth with distinct calyx and corolla; (8–)12(–15); free; 4–6(–7) whorled; isomerous. Calyx usually interpreted as (4–)6; 2 whorled (alternatively interpretable as bracts); polysepalous; regular; not persistent; imbricate. Corolla 6–12; 2–4 whorled (the outer two without nectaries, sometimes interpreted as petaloid K, the inner two or three whorls when present nectariferous, alternatively interpretable as staminodes); polypetalous; imbricate; regular; yellow, or orange.

Androecium 6. Androecial members free of the perianth; free of one another; 2 whorled (3+3). Androecium exclusively of fertile stamens, or including staminodes (if the inner petals are so interpreted). Staminodes if interpreted as such, 3–9; external to the fertile stamens. Stamens usually 6 (usually the same number as the nectariferous C); alternisepalous (i.e. usually oppositipetalous); opposite the corolla members. Anthers adnate; non-versatile; dehiscing by longitudinal valves; extrorse (the two valves posterior, but usually moving upwards and turning around, orientating the adhering pollen towards the centre of the flower); tetrasporangiate. Endothecium developing fibrous thickenings. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral. Anther wall initially with more than one middle layer. Tapetum glandular. Pollen without viscin strands. Pollen grains aperturate; 3 aperturate; spiraperturate; 2-celled.

Gynoecium ostensibly 1 carpelled (fancifully interpretable as 3). Carpels reduced in number relative to the perianth. The pistil 1 celled. Gynoecium monomerous (with no convincing evidence of pseudomonomery?); ostensibly of one carpel; superior. Carpel shortly stylate; apically stigmatic; 1–6 ovuled. Placentation marginal to basal. Stigmas wet type, or dry type; papillate, or non-papillate; Group II type, Group III type, and Group IV type. Ovules ascending; arillate, or non-arillate; hemianatropous, or anatropous; bitegmic; crassinucellate. Outer integument contributing to the micropyle. Embryo-sac development Polygonum-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; not proliferating; large. Synergids pear-shaped. Endosperm formation nuclear. Embryogeny onagrad.

Fruit fleshy. The fruiting carpel indehiscent; baccate. Seeds endospermic. Endosperm oily. Cotyledons flat. Embryo achlorophyllous (2/8); straight.

Seedling. Germination phanerocotylar, or cryptocotylar.

Physiology, phytochemistry. C3. C3 physiology recorded directly in Berberis, Mahonia. Sugars transported as sugar alcohols + oligosaccharides + sucrose (Mahonia). Not cyanogenic (with the segregate families excluded). Alkaloids present (usually), or absent. Berberine present (?always, colouring the tissues yellow). Arbutin absent. Iridoids not detected. Proanthocyanidins present, or absent; when present, cyanidin, or delphinidin. Flavonols present, or absent; when present, quercetin, or kaempferol and quercetin, or quercetin and myricetin. Ellagic acid absent (6 species, 3 genera). Aluminium accumulation not found. Sieve-tube plastids S-type.

Geography, cytology. Temperate to sub-tropical. Widespread in the North Temperate, tropical mountains, South America.

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.

Species 575. Genera 4; Berberis, Epimedium, Mahonia, Vancouveria.

General remarks. Nandinaceae J.G. Agardh, Leonticaceae and Podophyllaceae are close to Berberidaceae, but the compiled descriptions (q.v.) show enough differences to defend retaining them as segregate families for practical purposes (including keys).

Economic uses, etc. Edible berries from Berberis spp. (barberry), Mahonia aquifolium (‘Oregon grape’).

Illustrations. • Le Maout and Decaisne: Berberis vulgaris. • Le Maout and Decaisne: Berberis, Epimedium. • Berberis darwinii: Hook. Ic. Pl. 7–8 (1844). • Berberis dealbata: Bot. Reg. 1750, 1836. • Berberis coriaria: Bot. Reg. 46, 1841. • Berberis empetrifolia: Bot. Reg. xxvi, 27 (1840). • Berberis vulgaris: B. Ent., 1831. • Berberis umbellata: Bot. Reg. 1844, 44. • Epimedium alpinum: Eng. Bot. 52, 1863. • Epimedium grandiflorum: as E. macranthum, Bot. Reg. 1906 (1836). • Epimedium grandiflorum var. violaceum: as E. violaceum, Bot. Reg. xxvi, 43 (1840). • Mahonia aquifolium: as Berberis aquifolium, Bot. Reg. 1425, 1831. • Mahonia nervosa: as Berberis glumacea, Bot. Reg. 1426, 1831. • Mahonia pallida: as Berberis pallida, Bot. Reg. 1844, 16. • Mahonia pinnata: Bot. Reg.702, 1823. • Mahonia tenuifolia: as Berberis tenuifolia, Bot. Reg. 1844, 26.

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: 15th April 2018.’.