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Shrub, generally rhizomed.
Stem: spreading to erect, branched, spiny or not, vine-like or not; inner bark, wood generally bright yellow; over-wintering bud scales deciduous or not.
Leaf: simple or pinnately compound, cauline, alternate, generally leathery, generally persistent; leaflets generally 3–11, ± round to lanceolate, generally spine-toothed.
Inflorescence: raceme, axillary or terminal.
Flower: sepals 9 in 3 whorls of 3; petals 6 in 2 whorls of 3, bases generally glandular; stamens 6; ovules 2–9, stigma ± spheric.
Fruit: berry, spheric to elliptic, generally purple-black.
± 600 species: temperate worldwide. (Latin: ancient Arabic name for barberry) [Kim 2004 J Plant Res 117:175–182] Roots often TOXIC: spines may inject fungal spores into skin. Contact with filament causes stamen to snap inward, possibly to deposit pollen on pollinator.
Unabridged references: [Moran 1982 Phytologia 52:221–226, for relationship between Berberis, Mahonia.]

Key to Berberis

1. Lvs simple

2. Lvs evergreen, with 6–8 coarse, generally terminal spines; petals orange ..... B. darwinii

2' Lvs deciduous, with >> 8, small, marginal spines; petals yellow ..... B. vulgaris

1' Lvs compound

3. Bud scales persistent among upper leaves, 15–45 mm, thick, lanceolate; leaflets ± palmately veined ..... B. nervosa

3' Bud scales generally deciduous, < 5 mm, thin, ovate to deltate; leaflets generally pinnately veined

4. Infl open, flowers generally < 10; petiole generally < 2 cm; terminal leaflets generally lanceolate, oblong, or narrowly elliptic — SW, D

5. Lflets generally 3, terminal sessile ..... B. harrisoniana

5' Lflets 3–9, terminal stalked

6. Lflets flat to ± wavy, not folded along midrib, serrate, spines generally > 8 per side, ± 1 mm ..... B. nevinii

6' Lflets wavy, generally folded along midrib, ± lobed, spines generally 3–8 per side, 1–4 mm

7. Terminal leaflet generally < 2 × width; berries yellow-red when fresh, ± glaucous, not inflated ..... B. higginsiae

7' Terminal leaflet generally > 2 × width; berries blue-black when fresh, heavily glaucous, not inflated or inflated only due to insect parasitism

8. Terminal leaflet generally lance-ovate, length generally < 3 × width; flowers 8–12; fruit yellow- to purple-red ..... B. fremontii

8' Terminal leaflet generally narrow-lanceolate, length generally > 3 × width; flowers 3–5; fruit red-brown to dark purple ..... B. haematocarpa

4' Infl dense, flowers > 10; petiole generally > 2 cm (0–3 cm in Berberis pinnata); terminal leaflets generally ovate to wide-elliptic

9. Lflet margin strongly wavy, thick, generally hard, spines 6–10(12) per side, 2–5 mm ..... B. aquifolium var. dictyota

9' Lflet margin wavy to flat, ± thin, not hard, spines generally > 10 per side (to 0 in Berberis pinnata), 1–2 mm

10. Petioles generally > 3 cm; leaflets generally 5–9

11. Sts ascending to erect, generally 1–2 m; leaflets generally shiny adaxially ..... B. aquifolium var. aquifolium

11' Sts spreading to erect, generally < 0.8 m; leaflets generally dull adaxially ..... B. aquifolium var. repens

10' Petioles generally 1–3 cm; leaflets generally 7–11 ..... B. pinnata

12. Lflet margins generally flat, spines generally < 1 mm; upper stems reclining to weakly erect; plant 2–4 m ..... subsp. insularis

12' Lflet margins generally wavy, spines generally > 1 mm; upper stems generally erect; plant generally < 2 m ..... subsp. pinnata

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Citation for the whole project: Jepson Flora Project (eds.) [year] Jepson eFlora, http://ucjeps.berkeley.edu/IJM.html [accessed on month, day, year]
Citation for an individual treatment: [Author of taxon treatment] [year]. [Taxon name] in Jepson Flora Project (eds.) Jepson eFlora, [URL for treatment]. Accessed on [month, day, year].
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