Herbs or shrubs [trees], perennial, evergreen or deciduous, sometimes rhizomatous. Stems with or without spines. Leaves alternate, opposite, or fascicled, simple, 2-3-foliolate, or 1-3-pinnately or 2-3(-4)-ternately compound; stipules present or absent; venation pinnate or palmate. Inflorescences terminal or axillary, racemes, cymes, umbels (or umbel-like), spikes, or panicles, or flowers solitary or in pairs, flowers pedicellate or sessile. Flowers bisexual, inconspicuous or showy, radially symmetric; stipitate glands absent (except in Vancouveria); sepaloid bracteoles 0-9; perianth sometimes absent (Achlys), more frequently present, 2- or 3-merous, or sepals and petals intergrading (Nandis); sepals 6, distinct, often petaloid and colored, not spurred; petals 6-9, distinct, plane or hooded; nectary present; stamens 6; anthers dehiscing by valves or longitudinal slits; ovary superior, apparently 1-carpellate; placentation marginal or appearing basal; style present or absent, sometimes persistent in fruit as beak. Fruits follicles, berries, or utricles. Seeds 1-50, sometimes arillate; endosperm abundant; embryo large or small; mature seeds elevated on elongating stalk in Caulophyllum.

Distribution

Widespread, well represented in the north temperate zone.

Discussion

Genera 15, species ca. 650 (8 genera, 33 species in the flora).

Berberidaceae presents several interesting biogeographic features. Achlys is disjunct from western North America to east Asia with few morphologic differences between taxa. Diphylleia, Jeffersonia, and Podophyllum, each with a single eastern North American species, exhibit wide disjunctions to east Asia. Caulophyllum has three species, one in east Asia and two in the flora. Vancouveria is endemic to northwestern United States with nearest relations to Epimedium Linnaeus (H. Loconte and J. R. Estes 1989b; W. T. Stearn 1938), an exclusively Eastern Hemisphere genus.

Nandina, Berberis, Epimedium, and Podophyllum are cultivated.

The perianth of Berberidaceae is commonly composed of three distinct types of organs, but terminology for the organs varies from author to author. In our treatment, we refer to the small, outer parts as bracteoles (collectively forming a calyculus); the large, middle parts as sepals; and the innermost parts, which are commonly nectariferous, as petals. Some authors have referred to the bracteoles as outer sepals and to the petals as staminodes.

Selected References

Ernst, W. R. 1964. The genera of Berberidaceae, Lardizabalaceae, and Menispermaceae in the southeastern United States. J. Arnold Arbor. 45: 1-35.

Loconte, H. and J. R. Estes. 1989b. Phylogenetic systematics of Berberidaceae and Ranunculales (Magnoliidae). Syst. Bot. 14: 565-579.

Loconte, H. 1993. Berberidaceae. In: K. Kubitzki et al., eds. 1990+. The Families and Genera of Vascular Plants. 2+ vols. Berlin etc. Vol. 2, pp. 147-152.

Meacham, C. A. 1980. Phylogeny of the Berberidaceae with an evaluation of classifications. Syst. Bot. 5: 149-172.

Nowicke, J. W. and J. J. Skvarla. 1981. Pollen morphology and phylogenetic relationships of the Berberidaceae. Smithsonian Contr. Bot. 50: 1-83.

Ohwi, J. 1965. Flora of Japan (in English).... Washington.

Terebayashi, S. 1985. The comparative floral anatomy and systematics of the Berberidaceae. I. Morphology. Mem. Fac. Sci. Kyoto Univ., Ser. Biol. 10: 73-90.

Terebayashi, S. 1985b. The comparative floral anatomy and systematics of the Berberidaceae. II. Systematic considerations. Acta Phytotax. Geobot. 36: 1-13.

Lower Taxa

Achlys, Berberis, Caulophyllum, Diphylleia, Jeffersonia, Nandina, Podophyllum, Vancouveria

Illustrations

	Family ⠉	Taxon	Illustrator ⠉
	Berberidaceae Berberidaceae	Caulophyllum thalictroides Nandina domestica	John Myers John Myers
	Berberidaceae Berberidaceae Berberidaceae Berberidaceae	Berberis trifoliolata Berberis fendleri Berberis fremontii Berberis haematocarpa	John Myers John Myers John Myers John Myers
	Berberidaceae Berberidaceae Berberidaceae	Berberis piperiana Berberis dictyota Berberis repens	John Myers John Myers John Myers
... further results

Key

1	Shrubs (aerial stems sometimes very short in Berberis repens and B. pumila).	> 2
1	Perennial herbs.	> 3
2	Leaves 2-3-pinnately compound; margins of leaflets entire; inflorescences panicles.	Nandina
2	Leaves simple or 1-pinnately compound; if compound, margins of leaflets toothed; inflorescences racemes or umbels, or flowers solitary.	Berberis
3	Leaves simple, 2-7(-9)-parted; usually both basal and cauline leaves present.	> 4
3	Leaves compound; leaves all basal or (in Caulophyllum) all cauline.	> 5
4	Flowers 2 or more in cymes or umbels; leaves 2-parted, each segment 5-9-lobed; anthers dehiscent by 2 apically hinged valves; berries dark blue.	Diphylleia
4	Flowers solitary; leaves 5-7(-9)-parted, segments lobed or not; anthers dehiscent by longitudinal slits; berries yellow, orange, red, or purplish red.	Podophyllum
5	Leaves 2-3-foliolate; flowers solitary or in dense spikes.	> 6
5	Leaves 2-3-ternately compound; flowers in racemes, panicles, or compound cymes.	> 7
6	Leaves 3-foliolate; perianth absent; flowers in dense spikes; British Columbia to California.	Achlys
6	Leaves 2-foliolate; perianth present; flowers solitary; e North America (w to Minnesota and Iowa).	Jeffersonia
7	Petals well developed, 4-6 mm; ovary wall persisting and enclosing seeds in fruit; leaves all basal; California to Washington.	Vancouveria
7	Petals inconspicuous, 1-3 mm; ovary wall rupturing, seeds naked at maturity; leaves cauline; e North America (w to Manitoba, Nebraska, and Arkansas).	Caulophyllum