BERBERIDACEAE [Draft]

小檗科  xiao bo ke

Ying Junsheng (应俊生 Ying Tsun-shen)[1]; David E. Boufford[2], Anthony R. Brach[3], Julian Harber[4]

Herbs, perennial, or shrubs, rarely small trees, evergreen or deciduous, sometimes rhizomatous or tuberous. Stems with or without spines. Leaves alternate, opposite, or basal, simple, or 1–3 Χ pinnately or 2–3 Χ ternately compound; stipules present or absent; venation pinnate or palmate. Inflorescences terminal or axillary, racemes, spikes, umbels, cymes, or panicles, or flowers fascicled or solitary. Flowers pedicellate or sessile, bisexual, radially symmetric; bracteoles or bracts present or absent. Perianth usually 2- or 3-merous, rarely absent. Sepals 6–9, often petaloid, distinct, in 2 or 3 whorls. Petals 6, distinct, flat, hooded, pouched, or spurred; nectary present or absent. Stamens 6, opposite petals; anthers 2-celled, dehiscing by valves or longitudinal silts. Ovary superior, apparently 1-carpellate; ovules numerous, rarely solitary; placentation marginal or appearing basal; style present or absent, sometimes persistent in fruit as a beak. Fruit a berry, capsule, follicle, or utricle. Seeds 1 to numerous, sometimes arillate; endosperm abundant.

Seventeen genera and ca. 650 species: mainly in the north temperate zone and on subtropical mountains; 11 genera (1 endemic) and 320 species in China.

The family Berberidaceae presents an interesting, distinctive biogeography. The two species of Achlys are disjunctly distributed between eastern Asia and North America, with one in Japan and Korea and one along the west coast of North America. Caulophyllum and Diphylleia each contain three species; the former has one species in eastern Asia and two in North America, the latter has two in eastern Asia and one in North America. Sinopodophyllum and Plagiorhegma (if maintained separate from Podophyllum and Jeffersonia, respectively) are endemic to eastern Asia, but their closest relatives, Podophyllum and Jeffersonia, are widely disjunct in eastern North America. Epimedium has a wide, fragmented range from Japan westward to Algeria in Eurasia; the nearest relative, Vancouveria, is endemic to the Pacific Northwest of North America. Dysosma, Nandina, and Ranzania are endemic to China or to eastern Asia.

Ying Tsun-shen. 2001. Berberidaceae. In: Ying Tsun-shen, ed. Fl. Reipubl. Popularis Sin. 29: 50–343.

Podophyllum cavaleriei H. Lιveillι = Pilea panzhihuaensis C. J. Chen, A. K. Monro & L. Chen (Urticaceae; see Novon 17: 24. 2007).

1a.    Shrubs.

2a.    Leaves 2 or 3 Χ pinnately compound; margins of leaflets entire; anthers dehiscing by longitudinal slits; placentation parietal ......................................................................................  1. Nandina

2b.    Leaves simple or pinnately compound; margins of leaflets toothed; anthers dehiscing by 2 apically hinged valves; placentation basal.

3a.    Leaves simple; branches spiny ...................................................................  2. Berberis

3b.    Leaves pinnately compound; branches without spines ................................  3. Mahonia

1b.    Perennial herbs.

4a.    Leaves simple; petals without nectary.

5a.    Flowers solitary.

6a.    Rhizomes stout; flowers appearing before leaves; fruit a berry, indehiscent  4. Sinopodophyllum

6b.    Rhizomes slender; flowers appearing with the leaves; fruit a dehiscent capsule  5. Plagiorhegma

5b.    Flowers 2 or more, in fascicles, umbels or cymes.

7a.    Inflorescences fascicles or umbels; leaves 3–9-parted or -lobed; seeds numerous  6. Dysosma

7b.    Inflorescences terminal cymes or umbelliform; leaves 2-parted; seeds 2–4  7. Diphylleia

4b.    Leaves compound (except in few species of Epimedium); petals with nectary.

8a.    Fruit resembling a berry, actually a naked seed at maturity; inflorescences compound cymes             11. Caulophyllum

8b.    Fruit capsules; seeds contained within fruit; inflorescences racemes or panicles.

9a.    Plants from rhizomes; margins of leaflets toothed; petals 4, often spurred  8. Epimedium

9b.    Plants from tubers; margin of leaflets entire; petals 6, nectary-like, spurs absent.

10a.  Tubers producing 1 or rarely 2 leaves per year; fruit a dehiscent capsule; seeds with arils          9. Gymnospermium

10b.  Tubers producing 2–5 leaves per year; fruit an achene, globose, utricular; seeds without arils  10. Leontice

 

1. NANDINA Thunberg, Nov. Gen. Pl. 1: 14. 1781.

南天竹属  nan tian zhu shu

Ying Junsheng (应俊生 Ying Tsun-shen)[5]; David E. Boufford[6], Anthony R. Brach[7]

Shrubs, evergreen, without rhizomes. Leaves alternate, 2 or 3 Χ pinnately compound, estipulate; petioles and petiolules swollen at base; leaflets entire, venation pinnate. Inflorescences terminal or axillary panicles of dozens to hundreds of flowers. Flowers bisexual, 3-merous; bracteoles present. Sepals numerous, spirally arranged. Petals 6, larger than sepals, basally without nectaries. Anthers dehiscing by longitudinal slits; pollen grains prolate, 3-polyporate, exine distinctly reticulate. Ovary ellipsoid; placentation submarginal; style short; stigma entire or crisped. Fruit berries, globose. Seeds 1–3; aril absent. 2n = 20.

One species: endemic to eastern Asia.

Nandina is occasionally treated as a separate family, Nandiaceae (Takhtajan, A., 1997. Diversification and classification of flowering plants. Columbia University Press, New York) or subfamily Nandinoideae (Loconte, H. 1993. Berberidaceae. Pp. 147–151, in Kubitzki, K., Rohwer, J. G., & Bittrich, V. (eds), The Families and Genera of Vascular Plants. II. Flowering Plants: Dicotyledons, Magnoliid, Hamamelid and Caryophyllid Families. Springer, Berlin).

1. Nandina domestica Thunberg, Fl. Jap. 147. 1784.

南天竹  nan tian zhu

Nandina domestica var. linearifolia C. Y. Wu ex S. Y. Bao.

Shrubs, evergreen, erect, to 3 m tall, glabrous, few branched. Branchlets reddish when young. Leaves 30–50 cm; leaflets nearly sessile; blade of leaflet elliptic or elliptic-lanceolate, 2–10 Χ 0.5–2 cm, weakly leathery, glabrous, base cuneate, apex acuminate. Inflorescences erect, 20–35 cm. Flowers cream or white. Sepals ovate-triangular or ovate-oblong, 1–4 mm. Petals oblong, ca. 4.2 Χ 2.5 mm, apex obtuse. Stamens ca. 3.5 mm; anther connective prolonged. Ovary locules 1; ovules 1–3. Carpopodium 4–8 mm. Berry red or purplish, 6–8 mm in diam. Seeds grayish or brownish, oblate-spheroidal. Fl. Mar–Jun, fr. May–Nov. 2n = 20*.

Streamsides in montane forests, roadsides, thickets; below 1000 m. Anhui, Fujian, Guangdong, Guangxi, Guizhou, Henan, Hubei, Hunan, Jiangsu, Jiangxi, ?Shaanxi, Shandong, Shanxi, Sichuan, Yunnan, Zhejiang [India, Japan (probably not native; SE North America, the West Indies, and South America (Peru)].

2. BERBERIS Linnaeus, Sp. Pl. 1: 330. 1753.

小檗属  xiao bo shu

Ying Junsheng (应俊生 Ying Tsun-shen); David E. Boufford, Julian Harber

Shrubs evergreen or deciduous. Branches gray, usually terete or becoming so in age, glabrous or tomentose, spinose or not; mature stems yellow, brown, red, purple, or black, young shoots usually green; spines simple or usually 3–5-fid. Leaves on short shoots, simple, alternate, usually petiolate. Inflorescences solitary, fascicled flowers, racemes, umbels, or panicles. Flowers 3-merous; bractlets usually 3, caducous, scale-like. Flowers pale, greenish, or shades of yellow (except red and white in B. bicolor, and partially purple in B. sanguinea). Sepals 6(or 9), rarely 3 or 12. Petals 6, base nectariferous. Stamens opposite petals; anthers dehiscing by valves; pollen grains subspherical, exine reticulate. Ovary symmetrically club-shaped; ovules 1–12, rarely to 15, subbasal; styles short. Fruit a berry, usually red, dark red, purple, or black, globose, ellipsoid, oblong, ovoid, or obovoid. Seeds 1–10, light tan to red-brown or black; aril absent.

About 400–450 species: widespread in the northern hemisphere (especially diverse in the Himalayas and eastern Asia) and extending into Africa (one species south of the equator), temperate and Andean Latin America (the other center of diversity in Central and South America); ca. 174 species in China (154 endemic).

Many species of the genus are grown as ornamental shrubs, and used for medicinal purposes.

There are four interrelated limitations to this treatment. First, a lack of complete information about individual species; of the 174 species listed here, 21 have incomplete or no information about flowers, 29 have incomplete or no information about fruit, and 7 have incomplete or no information about either. Secondly, a significant number of species are only known from the type or a very small number of collections, a few from collections made so long ago that the species concerned could even be extinct. Thirdly, many species vary considerably in leaf size, shape, margin, and occasionally in color. This characteristic is particularly marked in many evergreen species from Series Wallichianae and can include significant variation even between leaves on the same plant. In all cases, the extent of this variation awaits detailed population studies. Fourthly, the question of natural hybridization. Studies from other parts of the world show this to be a significant issue (e.g., for SW Asia, see Browicz, K. & J. Zielinski. In: K. H. Rechinger, Flora des Iranischen Hochlandes und der umrahmenden Gebirge, 1975; for Chile and Argentina, see L. R. Landrum, Ann. Missouri Bot. Gard. 86: 793–834, 1999). It is well known among horticulturalists that species of Berberis from China brought together artificially in cultivation hybridize freely (including hybridization between evergreen and deciduous species), but no study of hybridization in the wild in China is known to us.

The limitations of this treatment are especially marked in relation to species endemic to Xizang.

 

Key 1

1a.     Shrub deciduous ..........................................................................................................  Key 3

1b.     Shrub evergreen; leaves leathery or thickly papery.

2a.    Inflorescence 1-flowered or a fascicle; style not persistent, or persistent and short; leaves leathery (ser. Wallichianae) .......................................................................................................  Key 2

2b.    Inflorescence a subumbel, raceme, subraceme, or fascicle; style persistent or not; leaves leathery or thickly papery ...................................................................................................................  Key 3

 

Key 2 (Berberis ser. Wallichianae)

1a.     Spines on stems mostly absent.

2a.    Shrub to 0.6 m; branches with wart-like hairs ............................................  1. B. dasyclada

2b.    Shrub to 2 m; branches glabrous ...........................................  2. B. insignis var. incrassata

1b.     Spines on stems present, (1–)3-fid.

3a.    Inflorescence always or mostly 1-flowered.

4a.     Pedicels 18–40 mm.

5a.     Mature stems dark red; pedicels 18–25 mm ..............................  3. B. chrysosphaera

5b.     Mature stems yellow; pedicels 20–40 mm ..........................................  4. B. asmyana

4b.     Pedicels to 10 mm.

6a.     Pedicels 3–4 mm; leaves lanceolate or oblanceolate, to 6.5 cm .........  5. B. paraspecta

6b.     Pedicels 4–10 mm; leaves to 3 cm.

7a.    Leaves oblong-lanceolate to lanceolate; ovules 3 or 4 ...................  6. B. candidula

7b.    Leaves elliptic or ovate-elliptic; ovules 4–6 ................................  7. B. verruculosa

3b.    Inflorescence a fascicle.

8a.     Fascicle mixed with racemes ................................................................  8. B. centiflora

8b.     Fascicle not mixed with racemes.

9a.     Mature stems purplish black or deep purple.

10a.   Mature stems purplish black; leaves entire or inconspicuously 1–3-spinose-serrulate  9. B. ziyunensis

10b.   Mature stems deep purple; leaves 8–15-spinose-serrulate .....  10. B. photiniaefolia

9b.     Mature stems purplish brown, reddish brown, or brown.

11a.   Mature stems purplish brown.

12a.   Leaves abaxially pruinose, oblong-elliptic, elliptic, or broadly elliptic, margin (3–)5–9-spinose-serrulate ...........................................................................  11. B. hsuyunensis

12b.   leaves abaxially not pruinose, margin entire or 2–6-spinose-serrate.

13a.   Leaves elliptic, margin entire ......................................  12. B. pingshanensis

13b.   Leaves linear-lanceolate, oblong-lanceolate, or narrowly elliptic, margin 2–6-spinose-serrate, occasionally entire .................................................  13. B. triacanthophora

11b.   Mature stems reddish brown or brown, grayish yellow, yellow, or brownish yellow.

14a.   Mature stems reddish brown or brown.

15a.   Leaf margin conspicuously revolute.

16a.   Leaves adaxially dull, lanceolate; young shoots sometimes purple when dry; pedicels 3–4 mm ....................................................................  16. B. grodtmanniana

16b.   Leaves adaxially very shiny, abaxially pruinose.

17a.   Leaves narrowly obovate or lanceolate; pedicels 1–2 mm  14. B. nantoensis

17b.   Leaves lanceolate; pedicels to 20 mm .........................  15. B. taliensis

15b.   Leaf margin not conspicuously revolute.

18a.   Ovules 1.

19a.   Leaves to 13 cm, oblong-elliptic, elliptic, elliptic-lanceolate, or lanceolate, margin (8–)20–35-spinose-serrate on each side ....................  17. B. deinacantha

19b.   Leaves to 4 cm, oblong, oblong-lanceolate, or elliptic, margin 3–7-spinose on each side ...............................................................................  18. B. hayatana

18b.   Ovules to 2 or more.

20a.   Ovules 10–14 .........................................................  23. B. calliantha

20b.   Ovules to 5.

21a.   Ovules to 2.

22a.   Ovules 1 or 2; leaves oblong-elliptic, to 15 cm, margin 10–25-spinose-serrate on each side ...................................................  19. B. sargentiana

22b.   Ovules 2; leaves oblong-lanceolate, ovate, or rhombic-elliptic, to 9 cm, margin closely (7–)15–40-spinulose-serrulate on each side  20. B. aristato-serrulata

21b.   Ovules to 5.

23a.   Leaf margin 4–7-spinose-serrulate on each side, blade elliptic-oblanceolate, narrowly lanceolate, or oblong, to 4 cm; ovules 2–4  21. B. jiangxiensis

23b.   Leaf margin 7–18(–25)-spinose-serrate on each side, blade elliptic or broadly elliptic, occasionally ovate, to 8 cm; ovules 3–5  22. B. xanthoclada

14b.   Mature stems grayish yellow, yellow, or brownish yellow.

24a.   Ovules 1 or 2.

25a.   Leaves abaxially pruinose.

26a.   Leaves elliptic or elliptic-lanceolate, 6–10 Χ 1.2–2 cm, margin flat  26. B. wangii

26b.   Leaves oblong-elliptic or oblong-lanceolate, 1.5–4.5 cm, margin revolute.

27a.   Leaves dull light green, oblong-elliptic or oblong-lanceolate, 1.5–3.5(–4.5) Χ 0.3–0.5(–0.8) cm, margin conspicuously revolute ..........  24. B. replicata

27b.   Leaves shiny green, oblong-elliptic, 2–4 Χ 1–1.5 cm, margin slightly revolute         25. B. nemorosa

25b.   Leaves not abaxially pruinose.

28a.   Leaves linear-oblong, 3–8 Χ 0.3–0.4 cm .......................  27. B. insolita

28b.   Leaves narrowly elliptic, elliptic, elliptic-lanceolate, oblong-elliptic, elliptic-oblanceolate, lanceolate, oblong, ovate, obovate, or oblanceolate.

29a.   Leaves to 5 cm long.

30a.   Leaf margin 12–16-spinose-serrate on each side, blade elliptic or oblong, 2–5 Χ 1–2 cm ............................................................  30. B. silvicola

30b.   Leaf margin 3–6-spinulose-serrulate or 4–8-spinose-serrate on each side.

31a.   Leaves lanceolate, 1–2 Χ 0.2–0.35 cm, margin 3–6-spinulose-serrulate on each side .......................................................  28. B. davidii

31b.   Leaves ovate, 1.2–3 Χ 0.8–1.7 cm, margin 4–8(–12)-spinose-serrate on each side ..........................................................  29. B. petrogena

29b.   Leaves to 8 cm long or longer.

32a.   Leaves to 8 cm long.

33a.   Leaves yellowish green, thickly leathery.

34a.  Leaf margin minutely 6–15(–18)-spinose-serrate on each side, blade elliptic, oblanceolate, or obovate, 2.5–6 Χ 1–1.5 cm  31. B. cavaleriei

34b.   Leaf margin coarsely 4–12-serrulate on each side, blade oblong-elliptic or elliptic, (3–)5–8 Χ 1.6–2.5 cm ...........  32. B. bergmanniae

33b.   Leaves adaxially deep green and shiny, or dull green, lanceolate, elliptic-lanceolate, or elliptic.

35a.   Leaves adaxially deep green, shiny, lanceolate or elliptic-lanceolate, 3–7 Χ 1–1.8 cm, margin 15–30(–40)-spinose-serrate on each side .....  33. B. fallaciosa

35b.   Leaves adaxially dull green, elliptic, narrowly elliptic, or lanceolate, 3–8 Χ 0.7–2 cm ......................................................  34. B. levis

32b.   Leaves longer than 8 cm.

36a.   Ovules 1.

37a.   Leaf margin 35–60-spinose-serrate on each side, blade elliptic-oblanceolate or elliptic-lanceolate, 4–10 Χ 1.5–2.5 cm, often castaneous, adaxially with close very distinct impressed reticulation  38. B. ferdinandi-coburgii

37b.   Leaf margin 5–27-spinose-serrate on each side.

38a.   Leaves 4–12 Χ 1–1.5 cm, linear-lanceolate, margin closely 10–20-spinose-serrate on each side, sometimes entire; funicles 3–6 Χ longer than ovules .........................................  35. B. sublevis

38b.   Leaves to 3.5 cm wide.

39a.   Mature berry white pruinose; leaves elliptic, lanceolate, oblanceolate, or obovate, 4–10 Χ 1–3 cm, adaxially with lateral veins slightly visible, margin (5–)10–20-spinose-serrate on each side  36. B. julianiae

39b.   Mature berry not pruinose; leaves lanceolate or elliptic-lanceolate, rarely ovate-lanceolate, 3–12 Χ 0.7–3.5 cm, adaxially with lateral veins obvious or slightly visible, margin 10–24(–27)-spinose-serrate on each side ...................................  37. B. vernalis

36b.   Ovules 1 or 2.

40a.   Ovules 1 or 2; leaves elliptic, 5–9.5 Χ 1–1.8 cm, adaxially with close, very distinct reticulation .................................  39. B. dumicola

40b.   Ovules 2.

41a.   Shrub to 1 m tall; leaves elliptic or elliptic-lanceolate, 7–10 Χ 2–2.5 cm; flowers 3–6, red and white ....................  40. B. bicolor

41b.  Shrub to 3 m tall; leaves lanceolate or oblong-elliptic, 3–10 Χ 0.7–2.5 cm; flowers 3–10, yellow ........................  41. B. atrocarpa

24b.   Ovules 2 or 3, or more than 3.

42a.   Ovules 2 or 3.

43a.   Leaves abaxially pruinose.

44a.   Ovules 3; leaves narrowly oblong-elliptic or oblong-lanceolate, 2.5–6 Χ 0.6–1.5 cm, margin 2–7-spinose-serrate on each side; inflorescence 2–12-flowered; pedicels 10–25 mm ..............................................................  44. B. taronensis

44b.   Ovules 2 or 3; leaves to 8 Χ 2.5 cm, margin 2–10-spinose-serrate on each side.

45a.   Leaves oblong-oblanceolate, elliptic-oblanceolate, or narrowly oblong-elliptic, 2–8 Χ 0.8–2.5 cm, margin above middle 2–10-spinose-serrate on each side, occasionally entire; inflorescence 4–14-flowered; pedicels 7–18 mm    42. B. chingii

45b.   Leaves elliptic, 3–7 Χ 1.3–2.5 cm, margin 3–10-spinose-serrate on each side; inflorescence 5–8-flowered; pedicels 15–20 mm  43. B. jingfushanensis

43b.   Leaves abaxially not pruinose.

46a.   Leaves to 8 cm long.

47a.   Leaf margin entire, sometimes slightly undulate, rarely 1- or 2-spinose-serrate on each side ....................................................  45. B. malipoensis

47b.   Leaf margin 2–20-spinose-serrate or 7–12-spinose-serrulate on each side.

48a.   Leaf margin 7–12-spinose-serrulate on each side, blade ovate to oblong-ovate, 3.3–5.5 Χ 1–1.8 cm; inflorescence 3–15-flowered; pedicel 7–20 mm           46. B. delavayi

48b.   Leaf margin 2–20-spinose-serrate.

49a.   Pedicels 4–10 mm; leaves oblanceolate, elliptic-oblanceolate, or elliptic, 3–7 Χ 0.8–1.6(–2.5) cm, margin 2–8(–14)-spinose-serrate on each side, occasionally entire; inflorescence (2–)4–12-flowered  47. B. fujianensis

49b.   Pedicels 10–30 mm; inflorescence 3–6-flowered; pedicel 10–15 mm; leaves elliptic, narrowly elliptic, or oblong, adaxially dark green, 2–6.5 Χ 0.8–1.6 cm, margin 13–20-spinose-serrate on each side  48. B. guizhouensis

46b.   Leaves longer than 8 cm.

51a.   Sepals in 3 whorls; leaves ovate-elliptic, oblong-elliptic, or lanceolate, 3.5–9 Χ 1–4 cm, adaxially dull gray-green or blue green ..  53. B. lempergiana

51b.   Sepals in 2 whorls.

52a.   Leaf margin 24–50(–60)-spinose-subserrulate on each side, blade narrowly elliptic, 7–20 Χ 1–3.5 cm ...........................  52. B. acuminata

52b.   Leaf margin 4–18-spinose-serrate on each side.

53a.   Leaves lanceolate or elliptic-lanceolate, 3–9 Χ 1.2–2.2 cm, margin 8–15-spinose-serrate on each side; inflorescence 2–6-flowered  49. B. subacuminata

53b.   Leaves ovate-elliptic, oblong, or oblong-obovate.

54a.   Leaves ovate-elliptic, very shiny, 6–10 Χ 2–4 cm, margin 8–15-spinose-serrate on each side ...........................  50. B. simulans

54b.   Leaves oblong, oblong-elliptic, or oblong-obovate, rigidly leathery, 3.5–10 Χ 1–1.5 cm, (4–)6–10(–18)-spinose-serrate on each side              51. B. soulieana

42b.   Ovules more than 3.

55a.   Dwarf shrub to 60 cm; leaves elliptic to oblong-elliptic, 1–2.5 Χ 0.7–1 cm      54. B. alpicola

55b.   Shrubs taller than 60 cm.

56a.   Leaves to 4 cm long, elliptic-lanceolate, 1.2–3.2 Χ ca. 0.9 cm; ovules (2 or)3 or 4 .......................................................................  55. B. griffithiana

56b.   Leaves to 7 cm or longer.

57a.   Leaves to 7 cm long.

58a.   Ovules to 4.

59a.   Flowers (8–)10–20; leaves elliptic to obovate, rigidly leathery, abaxially pruinose or not; petiole almost absent; ovules 2 or 3(or 4)  57. B. pruinosa

59b.   Flowers 1–6.

60a.   Leaves abaxially white pruinose, elliptic to ovate-elliptic; petiole 3–10 mm; flowers 3–6; ovules 2–4 ....................  56. B. coxii

60b.   Leaves not pruinose.

61a.   Leaves oblong-lanceolate, petiole almost absent; inflorescence (1- or)2–4-flowered; ovules 2–4 ...............  58. B. fallax

61b.   Leaves oblong-elliptic or elliptic-lanceolate, petiole short; inflorescence 2–6-flowered; ovules 3 or 4  59. B. phanera

58b.   Ovules to more than 4.

62a.   Flowers partially purple; leaves linear-lanceolate, 1.5–6 Χ 0.3–0.8 cm; ovules 3(–5) ........................................  60. B. sanguinea

62b.   Flowers yellow.

63a.   Sepals in 2–4 whorls; pedicels 6–30 mm.

64a.   Pedicels 6–12 mm; sepals in 2–4 whorls; ovules (1 or)2–5            64. B. kawakamii

64b.   Pedicels 15–30 mm; sepals in 3 whorls; ovules 4–8  63. B. hookeri

63b.   Sepals in 2 whorls.

65a.   Leaves abaxially white pruinose, margin 3–5-spinose-serrate on each side; young shoots and spines white pruinose; ovules 4 or 5           61. B. lijiangensis

65b.   Leaves abaxially glaucous-pruinose at first, finally greenish, margin 8–15-spinose-serrate on each side; young shoots and spines not pruinose; ovules 4–6 .......................  62. B. impedita

57b.   Leaves longer than 7 cm.

66a.   Leaf margin entire or 1–5-spinose-subserrulate on each side, blade oblong-lanceolate or oblong-elliptic, 4–10 Χ 1.5–2.5 cm  65. B. holocraspedon

66b.   Leaf margin more than 5-spinose-serrulate on each side.

67a.   Sepals in 3 whorls; leaves ovate-lanceolate to elliptic-lanceolate, 3.5–11 Χ 0.4–2.2 cm, adaxially dull or gray-green; inflorescence 3–8(–15)-flowered; pedicels 10–35(–40) mm; ovules 4 or 5  69. B. gagnepainii

67b.   Sepals in 2 whorls.

68a.   Leaves adaxially dull gray-green, lanceolate, 5–11 Χ 1–2 cm; pedicels 14–35 mm; ovules 2–4 ..........................  66. B. veitchii

68b.   Leaves adaxially bright green or mid-green; pedicels 5–13 mm.

69a.   Inflorescence 6–10-flowered; pedicels 5–10 mm; leaves lanceolate or broadly lanceolate, 6–9 Χ 1–1.5 cm, adaxially bright green; ovules 4 .....................................................  67. B. lubrica

69b.   Inflorescence 2–5-flowered; pedicels 7–13 mm; leaves very narrowly lanceolate, 4.5–12 Χ 0.8–1.5(–2) cm, adaxially mid-green; ovules (1–)3 or 4 ....................  68. B. yingjingensis

 

Key 3

1a.     Shrub evergreen, semi-evergreen, or deciduous; inflorescence 1-flowered, a fascicle, umbel, subumbel, subfascicle, subraceme, raceme, or panicle.

2a.    Shrub evergreen; leaves leathery or thickly papery.

3a.     Inflorescence 1-flowered or a fascicle; style not persistent, or persistent and short; leaves leathery (ser. Wallichianae) ..................................................................................................  Key 2

3b.     Inflorescence a subumbel, raceme, subraceme, or fascicle; style persistent or not; leaves leathery or thickly papery.

4a.     Inflorescence a raceme or umbellate subraceme; style persistent.

5a.    Leaves obovate or obovate-elliptic; inflorescence a raceme; seeds 4  73. B. fengshanensis

5b.    Leaves lanceolate, narrowly obovate, or obovate; inflorescence a raceme or an umbellate subraceme; ovules 1 or 2 .............................................................  74. B. potanini

4b.     Inflorescence a fascicle or subumbel; leaves leathery and style to 2 mm, or leaves thickly papery and style not persistent.

6a.    Inflorescence a subumbel, sometimes with few fascicled flowers at base; leaves obovate; seeds 4 or 5 ......................................................................................  72. B. pulangensis

6b.    Inflorescence a fascicle.

7a.     Mature stems grayish yellow; leaves elliptic, oblong-obovate, or obovate, thickly and rigidly leathery; inflorescence (4–)15–25-flowered; style persistent, to 2 mm  70. B. asiatica

7b.     Mature stems pale red; leaves obovate or obovate-elliptic, thickly papery; inflorescence 2–4-flowered; style not persistent .............................................  71. B. obovatifolia

2b.    Shrub semi-evergreen; leaves papery or thinly leathery.

8a.     Inflorescence 1-flowered ...............................................................  75. B. tsangpoensis

8b.     Inflorescence more than 1-flowered.

9a.     Inflorescence a fascicle, corymb, subraceme, raceme, or subumbel.

10a.   Mature stems reddish brown; inflorescence a fascicle, stalked corymb, subraceme, or raceme     76. B. wilsoniae

10b.   Mature stems pale yellow; inflorescence a fascicle, racemose subfascicle, or subumbel  77. B. heteropsis

9b.     Inflorescence a raceme or panicle.

11a.   Inflorescence a raceme.

12a.   Mature stems pale yellow; leaves to 1.1 cm; inflorescence a raceme  78. B. sabulicola

12b.   Mature stems reddish brown; leaves to 2.5 cm; inflorescence a fascicle, stalked corymb, subraceme, or raceme ...........................................................  76. B. wilsoniae

11b.   Inflorescence a panicle.

13a.   Mature stems purplish red; inflorescence 20–30-flowered ...  81. B. jiulongensis

13b.   Mature stems pale yellow or yellow-brown.

14a.   Mature stems very pale yellow; inflorescence 15–30-flowered  79. B. xanthophlaea

14b.   Mature stems yellow-brown; inflorescence 30–100-flowered  80. B. polyantha

1b.     Shrub deciduous.

15a.   Inflorescence 1-flowered or flowers paired.

16a.   Inflorescence 1-flowered but sometimes paired; mature stems reddish purple, dark red, or purple.

17a.   Ovules 3; pedicels (10–)15–30 mm ..................................................  97. B. muliensis

17b.   Seeds or ovules 6–12.

18a.   Seeds 6; fruiting pedicels 13–18 mm ........................................  98. B. platyphylla

18b.   Ovules 7–12; pedicels 25–50(–55) mm .......................................  99. B. capillaris

16b.   Inflorescence 1-flowered.

19a.   Mature stems yellowish brown or yellow.

20a.   Mature stems pale yellowish brown, spines 3–9-fid, partly foliaceous; ovules 5–8  85. B. sibirica

20b.   Mature stems yellow, spines simple or 3(or 5)-fid.

21a.   Ovules/seeds 1 ..............................................................  82. B. mianningensis

21b.   Ovules/seeds 4–11.

22a.   Ovules/seeds 4 ...............................................................  83. B. multicaulis

22b.   Ovules/seeds 6–11 ...............................................................  84. B. daiana

19b.   Mature stems red, reddish brown, reddish purple, or purple.

23a.   Mature stems dark reddish brown, spines simple, 3-fid, or 5-fid; ovules 6–10  86. B. angulosa

23b.   Mature stems red, reddish purple, or purple.

24a.   Dwarf shrub no taller than 1 m.

25a.   Plant to ca. 30 cm tall; spines 5–7(–9)-fid; ovules 6 or 7  87. B. erythroclada

25b.   Plant to 40–100 cm tall; spines 3(–5)-fid; ovules 2–5.

26a.   Plant to 50 cm tall; ovules 2 or seeds 4.

27a.   Plant to 40 cm tall; ovules 2 ...................................  88. B. minutiflora

27b.   Plant to 50 cm tall; seeds 4 ..................................  89. B. qiaojiaensis

26b.   Plant to 100 cm tall; ovules 3–5.

28a.   Plant less than 1 m tall; spines (3–)5-fid; ovules 3–5 ......  90. B. tsarica

28b.   Plant to 1 m tall; spines 3-fid; ovules 3 ...................  91. B. crassilimba

24b.   Shrub taller than 1 m.

29a.   Pedicels 1–3 mm; ovules 4–6 ......................................  95. B. campylogyna

29b.   Pedicels 6–55 mm; ovules 2–4 or 6–11.

30a.   Pedicels 13–55 mm; ovules 2, or 7 or 8.

31a.   Pedicels 13–25 mm; ovules 2 .......................  92. B. pseudothunbergii

31b.   Pedicels 25–55 mm; ovules 7 or 8 ...............................  94. B. ludlowii

30b.   Pedicels 6–15 mm; ovules 3 or 4, or 6–11.

32a.   Ovules 6–11; pedicels 8–15 mm ...............................  93. B. temolaica

32b.   Ovules 3 or 4; pedicels 6–15 mm .........................  96. B. dictyophylla

15b.   Inflorescence a fascicle, umbel, raceme, or panicle.

33a.   Inflorescence a fascicle.

34a.   Mature stems pale gray or grayish yellow.

35a.   Mature stems pale gray; leaves oblanceolate or elliptic, margin entire or 1–4-spinose-serrate on each side; inflorescence (1–)3- or 4-flowered ......................  100. B. reticulinervis

35b.   Mature stems pale grayish yellow; leaves oblanceolate or obovate-elliptic, margin closely 9–16-spinulose-serrulate on each side; inflorescence 5–8-flowered ................  101. B. yui

34b.   Mature stems pale yellow, pale yellow-brown, reddish yellow, red, red-brown, or purple.

36a.   Mature stems pale yellow or pale yellow-brown, or reddish yellow.

37a.   Mature stems pale yellow-brown; leaves oblong or obovate-oblong, margin 2–14-spinose-serrate on each side, occasionally entire; inflorescence (1- or)2–5-flowered; pedicels 12–22 mm         105. B. diaphana

37b.   Mature stems yellow or reddish yellow; leaf margin 15–40-spinulose-serrate on each side, or 1- or 2-spinose-serrate on each side, or entire.

38a.   Leaf margin closely 15–40-spinulose-serrulate on each side, blade oblong-orbicular; mature stems yellow; inflorescence 2–5-flowered; pedicels 15–30 mm  104. B. circumserrata

38b.   Leaf margin entire or 1- or 2-spinose-serrate on each side, blade obovate or oblanceolate; mature stems pale yellow or reddish yellow.

39a.   Leaves obovate, margin entire; mature stems pale yellow; inflorescence a fascicle or subfascicle, 2–6-flowered; pedicels 6–12(–16) mm ........  102. B. hobsonii

39b.   Leaves narrowly obovate or oblanceolate, margin 1- or 2-spinose-serrate on each side or entire; mature stems pale yellow or reddish yellow; inflorescence 1- or 2(–4)-flowered; pedicels 2–5 mm .....................................................  103. B. lhapsoensis

36b.   Mature stems red, pale or dark red, red-brown, purplish red, or dark purple.

40a.   Mature stems red, pale or dark red, or red-brown.

41a.   Mature stems dark red or red-brown; leaves obovate or obovate-lanceolate, margin 4–7-spinose-serrate on each side or entire; inflorescence (1- or)2–5(or 6)-flowered; pedicels 12–25(–30) mm ............................................................  108. B. morrisonensis

41b.   Mature stems pale red or red; leaves elliptic or oblanceolate; pedicels 6–12 mm.

42a.   Mature stems pale red; leaf margin entire or remotely 1–5-spinose-serrate on each side; inflorescence 1(–3)-flowered; pedicels 6–12 mm ...  106. B. campylotropa

42b.   Mature stems red; leaf margin entire, occasionally 1- or 2-spinose-serrate on each side; inflorescence 3- or 4-flowered; pedicels 8–11 mm ......  107. B. nullinervis

40b.   Mature stems purplish red or dark purple.

43a.   Mature stems purplish red; leaves entire; inflorescence 5–10-flowered, rarely a subfascicled raceme; pedicels 12–17 mm ..............................  109. B. trichohaematoides

43b.   Mature stems dark purple; leaf margin spinose-serrate; pedicels 2–5 mm or 20–30 mm.

44a.  Leaves linear-oblanceolate or linear-oblanceolate, margin 1–3-spinose-serrate on each side, occasionally entire; inflorescence 3–6-flowered; pedicels 2–5 mm; ovules 3–5  110. B. ulicina

44b.   Leaves oblong-obovate or elliptic, margin 5–12-spinose-serrate on each side; inflorescence 2–4-flowered; pedicels 20–30 mm; ovules 7–11 ...........  111. B. aemulans

33b.   Inflorescence an umbel, raceme, or panicle.

45a.   Inflorescence an umbel.

46a.   Leaf petiole ca. 1 cm; mature stems reddish brown; leaves obovate or obovate-lanceolate, to 5 cm ..............................................................................................  114. B. umbratica

46b.   Leaf petiole almost absent; mature stems purplish red or purplish black.

47a.   Mature stems purplish red; leaves oblong or elliptic, to 5 cm; pedicels 5–9 mm  112. B. gilungensis

47b.   Mature stems purplish black; leaves broadly obovate, occasionally orbicular, to 5.5 cm; pedicels 10–17 mm ......................................................................  113. B. multiserrata

45b.   Inflorescence a subfascicle, subumbel, or subraceme, raceme, or panicle.

48a.   Inflorescence a subfascicle, subumbel, or subraceme.

49a.   Mature stems grayish yellow; inflorescence to 3 cm, 3–5(–7)-flowered; sepals in 3 whorls      115. B. johannis

49b.   Mature stems pale yellow, yellowish brown, reddish or purplish brown.

50a.   Mature stems pale yellow; sepals in 2 whorls.

51a.   Ovules 1 or 2.

52a.   Ovules 2; leaves elliptic-obovate; inflorescence to 1.5 cm; pedicels 1–2 mm         116. B. arido-calida

52b.   Ovules 1 or 2; inflorescence to 3 cm; pedicels 4–6 mm.

53a.   Leaves narrowly oblong-obovate, margin entire; inflorescence to 3 cm; pedicels 6–12 mm ....................................................  117. B. papillifera

53b.   Leaves ovate or obovate, margin closely 7–15-spinose-serrate on each side; inflorescence to 3 cm; pedicels 4–6 mm .....  118. B. dictyoneura

51b.   Ovules 3 or 4.

54a.   Inflorescence 5–7-flowered, to 3 cm; pedicels 5–10 mm; ovules 4  119. B. kartanica

54b.   Inflorescence 4–15-flowered, to 6(–9) cm; pedicels 10–18(–23) mm; ovules 3 or 4 ...........................................................................  120. B. tischleri

50b.  Mature stems yellowish brown, reddish or purplish brown, red, or purple; sepals in 1–3 whorls.

55a.   Mature stems yellowish brown.

56a.   Ovules 2.

57a.   Leaves obovate, margin 5–12-spinose-serrate on each side  121. B. tomentulosa

57b.   Leaves narrowly oblanceolate, margin entire ......  122. B. lepidifolia

56b.   Ovules or seeds 4–6.

58a.   Ovules 4–6; leaves oblanceolate or narrowly obovate, margin entire, sometimes 1–4-spinose on each side; inflorescence to 3 cm, 3–5-flowered  123. B. bimbilaica

58b.   Seeds 4 or 5; leaves obovate, margin entire, rarely 1–5-spinose on each side; inflorescence to 4.5 cm, 5–8-flowered .....................  124. B. elliotii

55b.   Mature stems reddish or purplish brown, red, or purple.

59a.   Mature stems pruinose.

60a.   Mature stems and spines purple, glaucous, pruinose  125. B. leptoclada

60b.   Mature stems dark red, white pruinose, spines orange  126. B. calcipratorum

59b.   Mature stems not pruinose.

61a.   Leaves 2 cm long or less.

62a.   Shrubs to 2 m; leaves oblanceolate or narrowly elliptic, abaxially not pruinose; sepals in 3 whorls .................................  131. B. pseudoamoena

62b.   Shrubs to 1 m or higher.

63a.   Shrubs to 1 m.

64a.   Leaves narrowly obovate-elliptic or narrowly elliptic, abaxially pruinose ..............................................................  127. B. amoena

64b.   Leaves obovate or oblanceolate, abaxially not pruinose  128. B. atroviridis

63b.   Shrubs to 1.5 m.

65a.   Spines simple .................................  129. B. integripetala

65b.   Spines 3(or 5)-fid .............................  130. B. derongensis

61b.   Leaves more than 2 cm long.

66a.   Ovules 4–6; spines red; leaves obovate-elliptic, to 6 cm  132. B. heteropoda

66b.   Ovules fewer than 4.

67a.   Petiole to more than 5 mm.

68a.   Mature stems dark red or grayish brown; leaf petiole 2–8 mm or almost absent ...............................................  135. B. thunbergii

68b.   Mature stems purple; leaf petiole to 25 mm.

69a.   Petiole 1–15 mm; sepals in 1 whorl  133. B. mouillacana

69b.   Petiole 5–25 mm; sepals in 2 whorls  134. B. silva-taroucana

67b.   Petiole almost absent or to 5 mm.

70a.   Berry densely blue or violet pruinose  136. B. humido-umbrosa

70b.   Berry not or slightly pruinose.

71a.   Ovules 1 ........................................  137. B. moloensis

71b.   Ovules 2 or 3.

72a.   Spines to 12 mm ..........................  138. B. zayulana

72b.   Spines longer than 12 mm.

73a.   Leaves abaxially shiny ........  139. B. hemsleyana

73b.   Leaves abaxially not shiny.

74a.   Sepals in 3 whorls .........  140. B. yunnanensis

74b.   Sepals in 2 whorls.

75a.   Leaves adaxially dull dark green, margin entire  141. B. lecomtei

75b.   Leaves adaxially bright green, margin often 1–4-spinose-serrate on each side .  142. B. tsarongensis

48b.   Inflorescence a raceme or panicle.

76a.   Inflorescence a panicle.

77a.   Panicle congested or corymbose; mature stems pale yellow.

78a.   Panicle congested; panicle to 2.5 cm, 10–30-flowered .  167. B. aggregata

78b.   Panicle corymbose; panicle to 7 cm, 10–24-flowered .....  168. B. concolor

77b.   Panicle not congested or corymbose; mature stems pale yellow or brown, or reddish or purplish brown.

79a.   Mature stems pale yellow or pale brown ...........................  169. B. prattii

79b.   Mature stems reddish or purplish brown.

80a.   Sepals in 3 whorls.

81a.   Panicle 5–14 cm; leaves ovate to elliptic, sometimes oblong-lanceolate     170. B. francisci-ferdinandi

81b.   Panicle 2.5–5 cm; leaves obovate ........................  171. B. sherriffii

80b.   Sepals in 2 whorls.

82a.   Ovules 1; panicle to 9 cm ......................................  174. B. taylorii

82b.   Ovules 2–5; panicle to 6 cm or 7.5(–16 cm).

83a.   Ovules 2; panicle to 6 cm ........................  172. B. oblanceifolia

83b.   Ovules 3–5; panicle to 7.5(–16) cm .................  173. B. gyalaica

76b.   Inflorescence a raceme.

84a.   Raceme spike-like; pedicels to 4 mm.

85a.   Spike less than 2 cm long; mature stems dark purplish; leaves margin entire, blade obovate or narrowly obovate; raceme to 1.5 cm, often with a few fascicled flowers at base        143. B. batangensis

85b.   Spike more than 2 cm long; leaf margin spinose-serrate, spinulose-serrate, or entire.

86a.   Mature stems yellow or brown.

87a.   Leaves lanceolate, margin 15–40-spinose-serrate on each side; mature stems yellow ..........................................................................  144. B. salicaria

87b.   Leaves elliptic, obovate, or oblong-elliptic, margin 20–40-spinose-serrate on each side; mature stems yellow or pale brown ........  145. B. brachypoda

86b.   Mature stems reddish or purple.

88a.   Mature stems purple; leaves oblanceolate or spatulate-oblanceolate, margin entire .............................................................................  148. B. vernae

88b.   Mature stems red-brown to purple or purplish red; leaf margin spinulose-serrulate or entire, or spinose-serrate.

89a.   Mature stems red-brown to purple; leaves elliptic, margin entire or 2–9-spinulose-serrulate on each side .....................................  146. B. gilgiana

89b.   Mature stems purplish red; leaves lanceolate or obovate-lanceolate, margin (2–)9–15-spinose-serrate on each side .................  147. B. purdomii

84b.   Raceme not spike-like; pedicels longer than 4 mm.

90a.   Mature stems pale yellow or yellow, brownish, red, or purple.

91a.   Mature stems pale yellow or yellow.

92a.   Raceme not compound at base.

93a.   Leaves oblong-rhombic; raceme to 5 cm, 3–15-flowered, inflorescence sometimes subumbellate; ovules 1 ................  152. B. virgetorum

93b.   Leaves oblanceolate or oblong-obovate; raceme to 12 cm, 4–8-flowered; ovules 1 or 2 ..............................................................  153. B. pallens

92b.   Raceme sometimes compound at base.

94a.   Raceme to 7 cm, 6–12-flowered; pedicels 4–15 mm; ovules 1 or 2      151. B. mekongensis

94b.   Raceme to 4.5 cm; ovules 2–5.

95a.   Raceme to 2.8 cm, 9–15-flowered; pedicels 3–5 mm; ovules 2(–5)             149. B. racemulosa

95b.   Raceme to 4.5 cm, 6–15-flowered; pedicels 4–12 mm; ovules 3 or 4           150. B. sichuanica

91b.   Mature stems brownish, red, or purple.

96a.   Mature stems brownish.

97a.   Mature stems pale yellowish brown; leaf margin 12–25-spinose-serrate on each side; inflorescence a raceme, sometimes compound at base; ovules 3              154. B. tianshuiensis

97b.   Mature stems red-brown, purplish brown, pale yellow, or yellowish gray; inflorescence a raceme.

98a.   Mature stems pale red-brown or purplish brown.

99a.   Mature stems pale red-brown, pale yellow, or yellowish gray; leaves 15–60-spinulose-serrulate on each side; raceme to 10 cm  155. B. amurensis

99b.   Mature stems stramineous to purplish brown; leaves entire; raceme 8–30-flowered, to 7 cm .....................................  156. B. poireti

98b.   Mature stems reddish brown.

100a. Leaves elliptic or obovate-elliptic, 1.5–3(–6) cm, petiole 4–15 mm; raceme to 6 cm, 10–20-flowered ......................  157. B. henryana

100b. Leaves obovate, suborbicular, oblong-elliptic, or broadly elliptic, 3–6(–8) cm, petiole 10–40 mm; raceme to 20 cm, (10–)15–50(–75)-flowered            158. B. dasystachya

96b.   Mature stems red or purple.

101a. Mature stems dark red, purplish red, or purple; raceme sometimes compound at base.

102a. Raceme to 3.5 cm, 3–10-flowered; leaves obovate or obovate-elliptic          159. B. tenuipedicellata

102b. Raceme to 10 cm, 10–40-flowered.

103a. Raceme to 10 cm, 20–40-flowered; leaves elliptic or oblong-obovate      162. B. jamesiana

103b. Raceme to 7.5 cm, 10–24-flowered; leaves elliptic, elliptic-lanceolate, or obovate.

104a. Leaves elliptic or elliptic-lanceolate; raceme to 6 cm, 10–20-flowered .......................................................  160. B. dielsiana

104b. Leaves obovate; raceme to 7.5 cm, often verticillate, 16–24-flowered ........................................................  161. B. agricola

101b. Mature stems purplish red; raceme not compound at base.

105a. Leaves 6–14-spinulose-serrulate on each side ..  163. B. dubia

105b. Leaves entire.

106a. Leaves oblanceolate .......................  166. B. kongboensis

106b. Leaves ovate, obovate, or oblong.

107a. Leaves orbicular-ovate or orbicular-obovate; berry red, ovoid-globose ...................................................  164. B. nummularia

107b. Leaves obovate or oblong; berry purple-red, obovoid or oblong           165. B. integerrima

 

Berberis ser. Wallichianae D. F. Chamberlain & C. M. Hu, Notes Roy. Bot. Gard. Edinburgh 42(3): 544. 1985.

??vernacular name

Berberis subsect. Wallichianae C. K. Schneider, Bull. Herb. Boissier, sιr. 2, 5: 400. 1905.

Leaves evergreen, thinly or thickly leathery. Flowers pale, greenish, or shades of yellow (except red and white in B. bicolor, and partially purple in B. sanguinea), solitary, paired, or in fascicles (except inflorescence both fascicled and racemose in B. centiflora). Fruit always black, dark blue, or dark purple. Style absent or very short.

Eighty three species; 69 in China (62 endemic). Four with no or incomplete descriptions of flowers, eight with no or imperfect descriptions of fruit, three with no or imperfect descriptions of either flowers or fruit.

A western and southern extension of the series takes in Nepal, India (Sikkim and NE), Bhutan, Myanmar, Vietnam, Philippines, and Indonesia.

Chamberlain & Hu (1985) noted that this series ‘is a well defined natural group of species, the circumscription of which is not disputed.’

1. Berberis dasyclada Ahrendt, J. Bot., Lond. Suppl., 12. 1941.

??vernacular name

Berberis insignis J. D. Hooker & Thompson var. zeleica Ahrendt; B. sanei T. Husain, B. Datt, A. Garg & R. R. Rao, syn. nov.

Shrubs, evergreen, to 60 cm tall. Mature stems very dark reddish brown, terete, pubescent with wart-like hairs; spines absent. Petiole almost absent or 2–3 mm; leaf blade abaxially shiny, very pale yellow-green, adaxially shiny, dark green, narrowly elliptic, 4–9(–11.5) Χ 1.2–2.6(–3.5) cm, thickly leathery, both surfaces not pruinose, midvein raised abaxially, impressed adaxially, lateral veins scarcely visible abaxially, margin coarsely 8–12(–18)-aristate-dentate on each side, apex subacuminate. Inflorescence a fascicle, 3–6(–13)-flowered; flowers golden yellow; bracts lanceolate, ca. 3.5 Χ 3 mm, apex acuminate. Pedicel 10–12 mm, slender; bractlets lanceolate, ca. 4 Χ 1.25 mm, apex acuminate. Sepals in 3 whorls; outer sepals elliptic, ca. 5.5 Χ 3 mm, apex obtuse; median and inner sepals obovate-elliptic, ca. 7 Χ 6 mm. Petals obovate, ca. 4.5 Χ 4 mm, base clawed, glands separate and lanceolate, apex entire. Stamens ca. 4.25 mm; anther connective produced. Ovules 3(–5). Berry unknown. Fl. May, fr. unknown.

Pine and Rhododendron forests; 2700–3600 m. SE Xizang-Assam border region [NE India].

Berberis insignis var. zeleica and B. sanei are both from Assam and have occasional 3-fid spines but otherwise appear to be indistinguishable from B. dasyclada.

2. Berberis insignis J. D. Hooker & Thomson subsp. incrassata (Ahrendt) D. F. Chamberlain & C. M. Hu, Notes Roy. Bot. Gard. Edinburgh 42: 537. 1985.

球果小檗  qiu guo xiao bo

Berberis incrassata Ahrendt, Gard. Chron., ser. 3, 105: 371. 1939; B. incrassata var. bucahwangensis Ahrendt; B. incrassata var. fugongensis S. Y. Bao.

Shrubs, evergreen, 1–2 m tall. Mature stems dark red, terete, glabrous; spines absent. Petiole 2–4 mm; leaf blade abaxially shiny, pale yellow-green, adaxially dark grayish green, elliptic to elliptic-lanceolate, 5–16 Χ 2–6 cm, leathery, both surfaces not pruinose, midvein raised abaxially, impressed adaxially, lateral veins visible on both surfaces, base cuneate, margin coarsely 12–24-aristate-dentate on each side, apex acuminate. Inflorescence a fascicle, (4–)8–15-flowered. Pedicel 10–24 mm, slender, toward apex slightly thickened, glabrous. Sepals in 3 whorls; outer sepals ovate, ca. 4 Χ 3 mm, apex acute; median sepals ovate-elliptic, ca. 6 Χ 5 mm, apex obtuse; inner sepals obovate, ca. 7 Χ 6 mm. Petals obovate, ca. 6 Χ 4 mm, base cuneate, glands separate, apex entire. Stamens ca. 4 mm; anther connective truncate. Ovules 5–7. Berry purplish black, not pruinose, subglobose, 6–7 Χ 5–6 mm; style persistent, conspicuous. Fl. Apr–May, fr. Aug–Oct.

Thickets, broad-leaved forests, mixed forests, bamboo forests, cliffs; 1200–2400 m. Xizang, Yunnan [Myanmar].

The typical subspecies occurs across the Himalayas from Nepal to NE India. It is possible that population studies across this range could cast doubt on there being a separate infraspecific incrassata.

3. Berberis chrysosphaera Mulligan, Bull. Misc. Inform. Kew 1940: 78. 1940.

黄球小檗  huang qiu xiao bo

Shrubs evergreen, 30–60 cm tall. Mature stems dark red, very sparsely verruculose; internodes 1.5–3 cm; spines 3-fid, concolorous, 1–2 cm, sulcate. Petiole almost absent; leaf blade adaxially very shiny, dark green, narrowly elliptic or obovate-elliptic, 1.5–4 Χ 0.4–1 cm, leathery, abaxially pruinose, finely papillose, veins not visible on both surfaces, base attenuate, margin conspicuously revolute, 5–12-spinose-serrate on each side, apex acuminate. Flowers 1. Pedicel 18–25 mm. Sepals in 3 whorls; outer sepals ovate, ca. 4 Χ 3 mm, apex acute; median sepals obovate-elliptic, ca. 7 Χ 6 mm; inner sepals ca. 10 Χ 8 mm. Petals obovate, ca. 8 Χ 6 mm, glands oblong, apex slightly emarginate. Stamens ca. 5 mm; anther connective slightly prolonged, truncate. Ovules 9–12. Berry black, slightly pruinose, ovoid, ca. 10 Χ 6 mm; style not persistent, or persistent and strict. Fl. May, fr. Oct–Nov.

* Granite cliffs in valleys, forest understories; 2700–3000 m. SE Xizang (Rongto Valley, Zayu).

4. Berberis asmyana C. K. Schneider in Sargent, Pl. Wilson. 1: 357. 1913.

直梗小檗  zhi geng xiao bo

Shrubs evergreen, to 1.2 m tall. Mature stems yellow, sulcate, sparsely black verruculose; spines weak, 3-fid, concolorous, 2–6(–9) mm, terete. Petiole almost absent; leaf blade abaxially pale green, adaxially shiny, deep green, elliptic or obovate-elliptic, 1–3 Χ 0.6–1 cm, thinly leathery, abaxially not pruinose, midvein markedly raised abaxially, impressed adaxially, secondary and reticulate veins inconspicuous on both surfaces, base cuneate, margin occasionally entire, otherwise 1–3-spinose-serrate on each side, apex acute. Flowers 1(or 2). Pedicel 20–40 mm, slender, glabrous; bractlets ovate, ca. 1.8 mm. Sepals in 2 whorls; outer sepals ovate, ca. 3 Χ 2.5 mm; inner sepals obovate-orbicular, ca. 5 Χ 4.5 mm. Petals obovate, ca. 4.5 Χ 3.2 mm, base clawed, glands very close together, apex slightly emarginate. Stamens ca. 3.2 mm; anther connective truncate. Ovules 4 or 5, sessile. Berry (immature) green, slightly pruinose, ellipsoid, (6.5–)7–8 Χ 4–5 mm; style not persistent. Fl. May–Jun, fr. Jul–Sep.

* Mountain thickets, grassy slopes; 3000–3200 m. Sichuan.

This species appears to be only known from the type E. H. Wilson 2873 (K) and one other specimen S. Y. Hu 946 (A), both collected near Baoxing.

5. Berberis paraspecta Ahrendt, J. Linn. Soc., Bot. 57: 47. 1961.

鸡脚连  ji jiao lian

Shrubs, evergreen, to 2 m tall. Mature stems yellow, sulcate, glabrous; spines 3-fid, concolorous, 2.5–3.5 cm, stout. Petiole almost absent; leaf blade lanceolate or oblanceolate, 3–6.5 Χ 0.7–1.2 cm, subleathery, midvein raised abaxially, complanate or slightly impressed adaxially, lateral veins inconspicuous abaxially, conspicuous adaxially, base cuneate, margin 7–15-spinose-serrate on each side, apex acute or shortly acuminate. Inflorescence 1-flowered, otherwise unknown. Fruiting pedicel 3–4 mm. Berry (immature) blue pruinose, ovoid, ca. 6 Χ 4 mm; style not persistent. Fl. and fr. unknown.

* Forest understories; ca. 2700 m. Yunnan.

This species is known only from the type specimen with two fruit, collected near Lijiang. It is distinguished from other evergreen species found in this part of Yunnan by its solitary flowers and very short pedicels. However, it remains a possibility that this is an atypical specimen of a fascicled species.

6. Berberis candidula C. K. Schneider, Bull. Herb. Boissier, sιr. 2, 5: 402. 1905.

单花小檗  dan hua xiao bo

Berberis hookeri Lemaire var. candidula C. K. Schneider, Ill. Handb. Laubholzk. 1: 303. 1904; B. laojunshanensis T. S. Ying, syn. nov.

Shrubs, evergreen, densely compact, ca. 1 m tall. Mature stems yellow, verruculose; spines 3-fid, concolorous, 1–1.5 cm, subterete. Petiole very short or almost absent; leaf blade adaxially very shiny, dark green, oblong-lanceolate to lanceolate, 1.2–2.6(–3) Χ 0.4–1 cm, thickly leathery, abaxially white pruinose, reticulate veins inconspicuous on both surfaces, base cuneate, margin conspicuously revolute, 3- or 4(–7)-spinose-serrate on each side, apex acuminate, aristate. Inflorescence 1(–4)-flowered; pedicels 4–10 mm, glabrous. Sepals in 3 whorls; outer sepals yellowish red, oblong-ovate, ca. 4 Χ 2 mm, apex acute; median sepals oblong-obovate, ca. 7 Χ 5 mm; inner sepals obovate, ca. 10 Χ 8 mm. Petals obovate, ca. 8 Χ 6 mm, base cuneate, glands ovate, apex entire. Stamens ca. 5 mm; anther connective prolonged, truncate. Ovules 3 or 4. Berry purplish black, thinly pruinose, ellipsoid, 8–9 Χ 4–5 mm; style not persistent. Fl. Apr–May, fr. Jun–Sep.

* Montane trailsides, scrub thickets; 1200–3000 m. Chongqing, NW Hubei, E Sichuan.

The type of Berberis candidula — Farges s.n. is a cultivated specimen grown from seed collected at Tchen Kou Tin, Chongqing. This has solitary flowers although cultivated plants supposedly grown from cuttings from the type sometimes occasionally have 2- or 3-flowered fascicles. E. H. Wilson 2880 (A) also cited by Schneider was collected near Fan Xian in NW Hubei and also has solitary flowers. The type collection and only known specimen of B. laojunshanensis was collected at Laojunshan, also in NW Hubei. It has the same leaves as B. candidula and four ovules but a 3- or 4-flowered fascicled inflorescence and smaller flowers. Population studies are needed to determine whether this should be regarded as a separate variety of B. candidula.

Berberis candidula is on the 2007 IUCN Red List of Threatened Species.

7. Berberis verruculosa Hemsley & E. H. Wilson, Bull. Misc. Inform. Kew 1906: 151. 1906.

疣枝小檗  you zhi xiao bo

Shrubs, evergreen, to 1 m tall. Mature stems brownish yellow, terete, densely verruculose; young shoots yellow-green, densely pubescent and verruculose; spines pale yellow, 1–2 cm, adaxially subsulcate. Petiole almost absent; leaf blade abaxially grayish green, adaxially shiny, dark green, ovate-elliptic or elliptic, 1–2 Χ 0.6–1.1 cm, leathery, abaxially pruinose, sometimes becoming epruinose on old leaves, densely papillose, midvein conspicuously raised abaxially, impressed adaxially, lateral veins 3 or 4 pairs, reticulate veins inconspicuous on both surfaces, base cuneate, margin slightly revolute, 2–4-spinose-serrate on each side, apex acute. Inflorescence 1-flowered. Pedicel 4–10 mm, glabrous. Sepals in 3 whorls; outer sepals ovate, ca. 4 Χ 3 mm; median sepals ovate, ca. 6 Χ 5 mm; inner sepals obovate, ca. 10 Χ 8 mm. Petals elliptic or obovate, 5.5–6 Χ ca. 3 mm, base cuneate, glands separate, apex emarginate or retuse with rounded lobes. Stamens ca. 3.5 mm; anther connective slightly prolonged, rounded. Ovules 4–6. Berry purplish blue to black, pruinose, oblong-ovoid, 10–12 Χ 6–7 mm; style not persistent. Fl. May–Jun, fr. Jul–Sep.

* Thickets, forests, rocky slopes; 1300–3200 m. Gansu, ?Shaanxi, Sichuan, ?Yunnan.

The type collection is from near Kangding, W Sichuan.

8. Berberis centiflora Diels, Notes Roy. Bot. Gard. Edinburgh 5: 167. 1912.

多花大黄连刺  duo hua da huang lian ci

Berberis pruinosa Franchet var. centiflora (Diels) Handel-Mazzetti.

Shrubs, evergreen, 0.6–1.5 m tall. Mature stems pale yellow, terete, glabrous, not verruculose; spines 3-fid, concolorous, ca. 1 cm, slender. Petiole 1–4 mm; leaf blade abaxially pale green, adaxially shiny, yellow-green, oblong-obovate or oblanceolate, 3–7 Χ 1–3 cm, leathery, midvein obviously raised abaxially, slightly impressed adaxially, lateral veins conspicuous abaxially, reticulate veins inconspicuous on both surfaces, base cuneate, margin sometimes slightly revolute, 10–20-spinose-serrate on each side, apex obtuse, apiculate. Inflorescence a fascicle, (15–)20–30-flowered, often mixed with 2- or 3-flowered racemes, 5–6 cm. Pedicel 15–25 mm, glabrous, not pruinose; bractlets ca. 2 Χ 1.5 mm. Sepals in 2 whorls; outer and inner sepals obovate, equal in size, ca. 5.5 Χ 4 mm. Petals obovate, ca. 6.5 Χ 5 mm, base clawed, glands separate and lanceolate, apex slightly retuse. Stamens ca. 5 mm; anther connective prolonged, shortly apiculate. Ovules 4 or 5. Berry unknown. Fl. Jun, fr. unknown.

* Open situations in pine and Rhododendron forests; 2700–3300 m. NW Yunnan.

This species is apparently known only from the type specimen collected on the eastern flank of Dali Range. The presence of flowering racemes might indicate that this is a natural hybrid possibly between Berberis pruinosa and a deciduous Berberis species.

9. Berberis ziyunensis P. K. Hsiao & Z. Y. Li, Acta Bot. Yunnan. 21: 30. 1999.

紫云小檗  zi yun xiao bo

Shrubs, evergreen, 1.5–2.5 m tall. Mature stems purplish black, shiny, terete; spines 1–3-fid or absent, pale yellow, 5–6 mm, very weak. Petiole 2–3 mm; leaf blade abaxially pale yellowish green, adaxially deep green, elliptic or narrowly elliptic, occasionally oblanceolate, 4–11 Χ 1–3 cm, thickly leathery, abaxially slightly pruinose, midvein conspicuously raised abaxially, impressed adaxially, lateral veins inconspicuous on both surfaces, base cuneate, margin slightly revolute, entire or inconspicuously 1–3-spinulose-subserrulate on each side, apex acuminate or acute. Inflorescence a fascicle, 4–10(–25)-flowered; pedicels purplish brown when dry, 15–20 mm, glabrous; bractlets triangular-ovate, ca. 2.2 Χ 1 mm. Outer sepals ovate, ca. 3 Χ 3 mm; inner sepals obovate, ca. 4 Χ 3 mm. Petals obovate, ca. 3.5 Χ 3 mm, base not clawed, glands separate, apex entire. Stamens ca. 2.3 mm; anther connective truncate. Ovules 3, shortly funiculate. Berry (immature) reddish, ellipsoid, ca. 4 Χ 2 mm; style persistent, short. Fl. Mar–Apr, fr. unknown.

* Thickets under mixed forests; 1000–1300 m. SW Guizhou.

This taxon is known only from the type collection from Baiyun, Ziyun.

10. Berberis photiniaefolia C. M. Hu, Bull. Bot. Res., Harbin 6(2): 4. 1986.

石楠小檗  shi nan xiao bo

Shrubs, evergreen, to 2 m tall. Mature stems deep purple, terete, glabrous; internodes 3–6 cm; spines 3-fid, concolorous, 1–2 cm. Petiole very short; leaf blade adaxially shiny, green, lanceolate, 4–8.5 Χ 1.5–2.8 cm, leathery, abaxially glaucous, not pruinose, midvein distinctly raised abaxially, slightly impressed adaxially, lateral veins conspicuous abaxially, slightly raised adaxially, reticulate veins conspicuous abaxially, inconspicuous adaxially, base cuneate, margin flat, 8–15-spinose-serrate on each side, apex acuminate. Inflorescence a fascicle, (2–)4–8-flowered; pedicels 10–15 mm; bractlets triangular-ovate. Sepals in 2 whorls; outer sepals ovate-oblong, 3–3.5 mm; inner sepals obovate or suborbicular, ca. 4 Χ 3.2 mm. Petals obovate, ca. 3 mm, base clawed, glands separate, apex subentire. Stamens ca. 2.5 mm; anther connective truncate. Ovules 3 or 4. Berry bluish black, slightly pruinose, ovoid-ellipsoid, 7–8 Χ 4–5 mm; style not persistent. Fl. unknown, fr. Aug.

* Mountain summits; ca. 1000 m. E and NE Guangdong.

The type collection is from Zijin Xian, NE Guangdong.

11. Berberis hsuyunensis P. K. Hsiao & W. C. Sung, Acta Phytotax. Sin. 12: 388. 1974.

叙永小檗  xu yong xiao bo

Shrubs, evergreen, 1–2 m tall. Mature stems purplish brown, slightly angulate; internodes 5–6 cm; spines 3-fid, orange-brown, to 1 cm, slender. Petiole 2–3 mm; leaf blade abaxially yellow-green, adaxially green, oblong-elliptic, elliptic, or broadly elliptic, 6–9 Χ 3–5 cm, thickly leathery, abaxially pruinose, midvein and lateral veins conspicuously raised abaxially, slightly raised adaxially, reticulate veins visible adaxially, base cuneate, margin 5–9-spinose-serrate on each side, or above middle 3–5-spinose-serrate on each side, apex acute. Inflorescence a fascicle, 30–50-flowered. Pedicel 1.5–2 cm; bractlets triangular, ca. 1 Χ 1 mm. Sepals in 2 whorls; outer sepals triangular-ovate, ca. 3 Χ 2.5 mm; inner sepals broadly ovate, ca. 4 Χ 4 mm, apex obtuse. Petals obovate, ca. 4.5 Χ 3.5 mm, base clawed, with separate glands, apex slightly emarginate to subrounded. Stamens ca. 2 mm; anther connective prolonged, truncate. Ovules 5 or 6, shortly funiculate or subsessile. Berry black, pruinose, subglobose, ca. 7 Χ 6 mm; style not persistent. Fl. May–Jun, fr. Jul–Sep.

* Forests, streamsides; 1200–1600 m. Sichuan, NE Yunnan.

The type collection is from Xuyong Xian, C Sichuan.

12. Berberis pingshanensis W. C. Sung & P. K. Hsiao, Acta Phytotax. Sin. 12: 387. 1974.

屏山小檗  ping shan xiao bo

Shrubs, evergreen, to 2 m tall. Mature stems purplish brown, sulcate, glabrous; spines 3-fid, concolorous, ca. 5 mm. Petiole 2–3 mm; leaf blade abaxially green, adaxially deep green, elliptic, 8–17 Χ 3.5–7 cm, subleathery, abaxially not pruinose, midvein conspicuously raised abaxially, impressed adaxially, lateral and reticulate veins slightly raised abaxially, lateral veins slightly raised adaxially, base broadly cuneate or subrounded, margin entire, apex acute. Inflorescence a fascicle, 6–14-flowered. Pedicel purplish brown, 15–30 mm, glabrous; bractlets ovate-triangular, ca. 1 mm. Sepals in 2 whorls; outer sepals broadly elliptic, ca. 4 Χ 3 mm; inner sepals suborbicular or reniform-orbicular, ca. 4 Χ 4–5 mm. Petals suborbicular, base with glands separate, apex entire or obtusely emarginate. Stamens ca. 2 mm; anther connective truncate. Ovules 2, funicles ca. as long as ovules. Berry black, not or slightly pruinose. Fl. May–Jun, fr. Jun–Jul.

* Trailsides, sides of rocks; ca. 800 m. S Sichuan.

This species is known only from the type collection from Pingshan Xian on the border with NE Yunnan.

13. Berberis triacanthophora Fedde, Bot. Jahrb. Syst. 36(Beibl. 82): 43. 1905.

芒齿小檗  mang chi xiao bo

Shrubs, evergreen, 1–2 m tall. Mature stems reddish purple, terete, scarcely verruculose; spines 3-fid, concolorous, 1–2.5 cm. Petiole almost absent; leaf blade abaxially gray-green, adaxially shiny, deep green, linear-lanceolate, oblong-lanceolate, or narrowly elliptic, 2–6 Χ 0.25–0.8 cm, leathery, abaxially papillose, sometimes slightly pruinose, midvein raised abaxially, lateral and reticulate veins inconspicuous on both surfaces, base cuneate, margin 2–6-spinose-serrate on each side, occasionally entire, apex acuminate or acute, aristate. Inflorescence a fascicle, 2–6-flowered. Pedicel 15–30 mm, glabrous. Sepals in 3 whorls; outer sepals ovate-orbicular, ca. 2 Χ 1.8 mm; median sepals ovate, ca. 3.5 Χ 2.5 mm, apex acute; inner sepals obovate, ca. 5 Χ 4 mm, apex obtuse. Petals obovate, ca. 4 Χ 3 mm, base cuneate, glands separate, oblong, apex emarginate. Stamens ca. 2 mm; anther connective prolonged, truncate. Ovules 2 or 3. Berry blue-black, slightly pruinose, ellipsoid, 6–8 Χ 4–5 mm. Fl. May–Jun, fr. Jun–Oct.

* Mixed forests; 500–2100 m. ?Guizhou, W Hubei, ?Hunan, S Shaanxi, E Sichuan.

The type collection is from Changyang, W Hubei.

14. Berberis nantoensis C. K. Schneider, Repert. Spec. Nov. Regni Veg. 46: 252. 1939.

??vernacular name

Berberis densifolia Byhouwer (1928), not Rusby (1920).

Shrubs, evergreen, compact, 1.5–2 m tall. Mature stems brown, subterete or sulcate; spines sometimes absent, 3-fid, concolorous, 0.8–1.2 cm, weak, abaxially sulcate. Petiole almost absent; leaf blade abaxially pale green, adaxially green, narrowly obovate, 1–3 Χ 0.5–1 cm, leathery, abaxially often pruinose at first, both surfaces with obscure lateral veins, base attenuate or cuneate, margin conspicuously revolute especially when dry, 1–4 spinose-serrate usually only on upper ½ of each side. Inflorescence a dense, congested fascicle, 6–10(–18)-flowered; flowers bright yellow; pedicels 1–2 mm; bractlets yellow with red markings, oblong-ovate, ca. 2 Χ 1.5 mm, apex acute. Sepals in 2 whorls; outer sepals elliptic-obovate, ca. 6 Χ 3.5 mm, apex rounded; inner sepals narrowly elliptic, ca. 5 Χ 2.5 mm, apex rounded. Petals broadly obovate, ca. 4.5 Χ 3 mm, apex rounded, entire or slightly incised, base cuneate, glands separate, elliptic. Stamens ca. 3 mm; anther connective prolonged, truncate. Ovules 3–5. Berry dark purplish, pruinose, ellipsoid, 5–7 Χ 4–5 mm; style persistent, short. Fl. May, fr. Nov–Dec.

* Forest margins on mountain slopes; 2200–3000 m. Taiwan.

The type collection is from Mt. Kiraishui. The flower description used here is from a cultivated plant grown by one of us (Harber) from seeds of BSWJ 6926 collected at Yushan at 2815 m elevation in 1999.

15. Berberis taliensis C. K. Schneider, Repert. Spec. Nov. Regni Veg. 46: 252. 1939.

大理小檗  da li xiao bo

Shrubs, evergreen, to 1.2 m tall. Mature stems erect, reddish brown, stout, sulcate, scarcely verruculose; spines 3-fid, concolorous, 0.8–2 cm, adaxially conspicuously sulcate. Petiole almost absent; leaf blade adaxially shiny, dark green, narrowly elliptic or oblanceolate, 1.2–3 Χ 0.3–0.5 cm, leathery, abaxially pruinose, midvein raised abaxially, impressed adaxially, lateral and reticulate veins very inconspicuous adaxially, base cuneate, margin conspicuously revolute, 1–3(or 5- or 6)-spinose-serrate on each side, occasionally entire, apex acuminate, mucronate. Inflorescence a fascicle, 2–5-flowered. Pedicel brownish gray, (12–)15–20 mm; bractlets ovate, ca. 3 mm. Sepals in 2 whorls; outer sepals oblong-elliptic, ca. 4.5 Χ 3 mm; inner sepals broadly elliptic, ca. 6 Χ 3.5–4 mm. Petals obovate, ca. 5.5 Χ 3.5–4 mm, base cuneate, slightly clawed, glands separate, apex entire, slightly emarginate. Stamens ca. 3.5 mm; anther connective not prolonged, truncate. Ovules 4, sessile. Berry black-purple, white pruinose, oblong, 8–12 Χ ca. 5 mm; style not persistent. Fl. May–Jul, fr. May–Jul.

* Pinus forests, scrub on valley slopes; 3000–4000 m. Yunnan (Dali area).

16. Berberis grodtmanniana C. K. Schneider, Oesterr. Bot. Z. 67: 32. 1918, [‘grodtmannia’].

安宁小檗  an ning xiao bo

Berberis grodtmannia var. flavoramea C. K. Schneider, syn. nov.; B. jinshajiangensis X. H. Li, syn. nov.; B. micropetala T. S. Ying (1999), not C. K. Schneider (1939), syn. nov.

Shrubs, evergreen, 1–3 m tall. Mature stems reddish brown, stout, very sulcate, not verruculose; young shoots sometimes purple when dry; spines 3-fid, 1–3 cm, stout, adaxially terete or flat. Petiole almost absent; leaf blade abaxially slightly shiny, pale green, adaxially dull, lanceolate, 3–6(–14) Χ 0.5–1.2(–2.4) cm, leathery, abaxially not pruinose, midvein conspicuously raised abaxially, impressed adaxially, lateral and reticulate veins inconspicuous abaxially, slightly visible adaxially, base cuneate, margin conspicuously revolute, especially on upper leaves, minutely 7–12(–15)-spinose-serrate on each side, apex acute or acuminate, aristate. Inflorescence a fascicle, 5–10-flowered; flowers bright yellow. Pedicel (1–)3–4 mm; bractlets upper part reddish, lanceolate, ca. 2 Χ 1 mm, apex acute. Sepals in 2 whorls; outer sepals ovate or ovate-elliptic, sometimes with faint reddish stripe, ca. 4 Χ 2 mm; inner sepals obovate or obovate-elliptic, ca. 5 Χ 2–4 mm. Petals obovate, 4.5–5 Χ 2.5–3 mm, apex emarginate or entire. Stamens ca. 3 mm; anther connective slightly prolonged, truncate. Ovules 1–3, sessile. Berry blue, not pruinose, ellipsoid, 8–9 Χ ca. 4 mm; style persistent, short. Fl. Apr–May, fr. May–Aug.

* Forests, thickets, trailsides, runlet sides; 1900–3500 m. Sichuan, Yunnan.

The type collections of Berberis grodtmanniana and its synonyms are from the border area of NW Yunnan and SW Sichuan. The type collection of var. flavoramea has somewhat lighter mature stems, but this seems too trivial a difference to justify a separate variety. Cultivated plants grown by one of us (Harber) from EDHC 97063 and BSWJ 7889 from seed collected respectively from Langshou Shan, Yunnan in 1998 and Huili, Sichuan in 2000 have lower leaves that are slightly shiny and non-revolute when in shade.

17. Berberis deinacantha C. K. Schneider, Repert. Spec. Nov. Regni Veg. 46: 259. 1939.

壮刺小檗  zhuang ci xiao bo

Berberis deinacantha C. K. Schneider var. valida C. K. Schneider, syn. nov.; B. jingguensis G. S. Fan & X. W. Li, syn. nov.; B. valida (C. K. Schneider) C. K. Schneider; B. wuliangshanensis C. Y. Wu ex S. Y. Bao, syn. nov.

Shrubs, evergreen, 1–3 m tall. Mature stems reddish brown, terete or slightly angulate, black verruculose, glabrous; semi-mature shoots sometimes purple when dry; spines downcurved, 3-fid, concolorous, 2.5–5 cm, stout, adaxially sulcate. Petiole 2–5 mm; leaf blade abaxially yellow-green, adaxially dark green, oblong-elliptic, elliptic, elliptic-lanceolate, or lanceolate, 3–13 Χ 1.5–3.5 cm, leathery, both surfaces not pruinose, midvein obviously raised abaxially, obviously impressed adaxially, lateral veins slightly raised abaxially, lateral veins 12–17 pairs, reticulate veins prominent on both surfaces, base cuneate, margin sometimes slightly revolute, (8–)20–35-spinose-serrate on each side, apex obtuse, aristate. Inflorescence a fascicle, (3–)6–15(–20)-flowered; flowers deep yellow. Pedicel (6–)12–15(–20) mm. Sepals in 2 whorls; outer sepals ovate or suborbicular, ca. 4 Χ 3.5 mm, apex rounded, obtuse; inner sepals obovate, obovate-orbicular, or elliptic, 5.5(–7) Χ ca. 5 mm. Petals oblong-obovate or obovate, ca. 4.5 Χ 3 mm, base not clawed, with separate glands, apex slightly emarginate or entire. Stamens 3–4.5 mm; anther connective prolonged, slightly retuse. Ovules 1. Berry purplish black, not or slightly pruinose, ellipsoid, 6–8 Χ ca. 4 mm; style not persistent. Fl. May, fr. Nov.

* Thickets, forests; 1700–3100 m. ?Guizhou, SW Sichuan, NE, NW, and W Yunnan.

Li Xin Hua (J. Trop. Subtrop Bot. 16(2): 176–178. 2008) reduced Berberis jingguensis to a synonym of B. wuliangshanensis while noting the protolog’s description of the former as a vine up to 7 m tall was likely to be the result of mistaken recording in the field. The types of both taxa have distinctly purple stems and spines. However, observations by one of us (Harber) of the green semi-mature stems of a cultivated plant of B. deinacantha grown from AC 1010 from seed collected near Zhaotong in NE Yunnan in 1995 show that such stems and their spines can dry to the exact color of those of the holotypes of B. jingguensis and B. wulangshanensis.

18. Berberis hayatana Mizushima, Misc. Rep. Res. Inst. Nat. Resourc. Tokyo, No. 35: 31. 1954.

南湖小檗  nan hu xiao bo

Berberis formosana H. L. Li, J. Wash. Acad. Sci. 42: 41. (1952), not Ahrendt (1941).

Shrubs, evergreen, to 1 m tall. Mature stems brownish red, very sulcate, not or scarcely verruculose; spines 3-fid, concolorous, 1–1.7 cm, weak, abaxially sulcate. Petiole almost absent or short; leaf blade abaxially slightly shiny, green, adaxially dull green, sometimes slightly glaucous, oblong, oblong-lanceolate, or elliptic, 2–4 Χ 0.6–1.2 cm, leathery, abaxially not pruinose, midvein obviously raised abaxially, slightly impressed adaxially, lateral veins 5–7 pairs, inconspicuously raised abaxially, slightly impressed adaxially, base attenuate or cuneate, margin sometimes slightly revolute, 3–7-spinose-serrate on each side, apex acute, aristate. Inflorescence a fascicle, 2–7-flowered; flowers bright yellow. Pedicel red, 8–10 mm, slender, glabrous; bractlets brownish red, triangular-ovate, ca. 1.5 mm, apex acute. Sepals in 2 whorls; outer sepals broadly ovate or obovate-elliptic, ca. 5 mm, apex acute; inner sepals obovate-elliptic or obovate, ca. 5 mm, apex rounded. Petals obovate, ca. 4 mm, base attenuate, glands separate, apex emarginate. Stamens ca. 2.5 mm; anther connective slightly prolonged, truncate. Ovules 1, funiculate. Berry black, pruinose, ellipsoid or ovoid, 4–6 Χ ca. 2 mm; style not persistent. Fl. Apr, fr. Nov–Dec.

* Thickets, mixed conifer and broad-leaved forests on mountain ridges and slopes; 1500–2500 m. N Taiwan.

19. Berberis sargentiana C. K. Schneider in Sargent, Pl. Wilson. 1: 359. 1913.

刺黑珠  ci hei zhu

Berberis recurvata Ahrendt.

Shrubs, evergreen, 1–3 m tall. Mature stems light reddish brown, terete, sometimes black verruculose; young shoots often bright red; internodes 3–6 cm; spines 3-fid, light brown, (2–)3–6 cm, stout, adaxially sulcate. Leaves subsessile; leaf blade abaxially pale green, adaxially slightly shiny, mid-green, oblong-elliptic, 4–15 Χ 1.5–6.5 cm, thickly leathery, midvein conspicuously raised abaxially, impressed adaxially, lateral and reticulate veins prominent abaxially, lateral veins slightly raised and reticulate veins indistinct adaxially, base cuneate, margin 10–25-spinose-serrate on each side, apex acute. Inflorescence a fascicle, 4–10-flowered; flowers pale greenish yellow. Pedicel 10–20 mm; bractlets red, ca. 2 Χ 2 mm. Sepals in 3 whorls; outer sepals with a red band along middle, ovate, ca. 3.5 Χ 3 mm, apex subacute; median sepals rhombic-elliptic, ca. 5 Χ 4.5 mm; inner sepals obovate, ca. 6.5 Χ 5 mm. Petals obovate, ca. 6 Χ 4.5 mm, base cuneate, with contiguous, orange glands, apex emarginate with rounded lobes. Stamens ca. 4.5 mm; anther connective truncate. Ovules 1 or 2. Berry black, not pruinose, oblong or oblong-ellipsoid, 6–8 Χ 4–6 mm; style not persistent. Fl. Apr–May, fr. Jun–Nov.

* Thickets, trailsides, bamboo forests, streamsides of forest understories; 700–2100 m. W Hubei, ?Hunan.

20. Berberis aristato-serrulata Hayata, Icon. Pl. Formosan. 3: 13. 1913.

密齿小檗  mi chi xiao bo

Berberis bicolor auct non Levi Mizushima; B. chingshuiensis Shimizu, syn. nov.; B. mingetsensis Hayata.

Shrubs, evergreen. 1–1.5 m tall. Mature stems reddish brown, slender, sulcate; internodes 5–6 cm; spines 3-fid, concolorous, to 15 mm, slender. Petiole 2–4 mm; leaf blade abaxially pale green, adaxially shiny, green, oblong-lanceolate, ovate, or rhombic-elliptic, (2.5–)5–9 Χ 0.8–2.5 cm, thinly leathery or papery, abaxially sometimes slightly glaucous, midvein conspicuously raised abaxially, reticulate veins on both sides, raised abaxially, base attenuate or cuneate, margin slightly revolute, closely (7–)15–40-spinulose-serrulate on each side, apex acute, aristate. Inflorescence a fascicle, (4–)10–15-flowered; flowers greenish yellow. Pedicel 6–10 mm; bractlets oblong-ovate or ovate, 1.25–2.5 mm, apex acuminate. Sepals in 2 or 3 whorls; outer sepals triangular-oblong, ca. 3 mm, apex acuminate; median and inner sepals obovate, 4–5.5 Χ ca. 4 mm, apex rounded. Petals obovate-spatulate or broadly obovate, ca. 5 Χ 2.5 mm, basal glands ca. 1 mm, apex rounded, emarginate to incised. Ovules 2; stipes ca. as long, or to 2 Χ as long as ovules. Berry dark blue-black, ovoid to ellipsoid, 4–7 Χ 2–4 mm; style persistent, short. Fl. Apr, fr. unknown.

* Alpine thickets, scrub; 2000–3000 m. Taiwan.

Berberis chingshuiensis was mistakenly included as a synonym of B. kawakamii in the first edition of the Flora of Taiwan.

21. Berberis jiangxiensis C. M. Hu, Bull. Bot. Res., Harbin 6(2): 9. 1986.

江西小檗  jiang xi xiao bo

Berberis jiangxiensis var. pulchella C. M. Hu, syn. nov.

Shrubs, evergreen, ca. 1 m tall. Mature stems dark brown, very sulcate, not verruculose; spines 3-fid, concolorous, 1–2.5 cm, subterete. Petiole 1–3 mm; leaf blade adaxially shiny, green, elliptic-oblanceolate, narrowly lanceolate, or oblong, 1.5–4 Χ 0.5–1.2 cm, thinly leathery, abaxially pruinose, midvein obviously raised abaxially, slightly impressed adaxially, lateral veins conspicuous abaxially, 2–4 pairs, obviously raised adaxially, reticulate veins inconspicuous abaxially, base attenuate, margin 4–7-spinose-serrate on each side, apex acute. Inflorescence a fascicle, (2- or)3- or 4-flowered. Pedicel 6–10 mm, slender or stout; bractlets ovate, ca. 3 Χ 2 mm, apex acuminate. Sepals in 2 whorls; outer sepals ovate-elliptic, ca. 4 mm; inner sepals broadly elliptic, ca. 4.5 mm. Petals obovate, ca. 3.5 mm, base shortly clawed, glands very close together, apex emarginate, lobes rounded, obtuse. Stamens ca. 3 mm; anther connective truncate. Ovules 2–4. Berry blue-black, slightly pruinose, ellipsoid, ca. 9 Χ 5.5 mm; style persistent, short. Fl. Apr–Jun, fr. Jul–Sep.

* Trailsides, rock crevices, sides of rocks, forests, runlet sides; 1500–1800 m. Jiangxi.

The type collection of Berberis jiangxiensis is from Wu Gong Shan, Anfu Xian, W Jiangxi, that of var. pulchella from Nanfeng Xian, E Jiangxi.

22. Berberis xanthoclada C. K. Schneider, Repert. Spec. Nov. Regni Veg. 46: 261. 1939.

梵净小檗  fan jing xiao bo

Shrubs, evergreen, 1–2.5 m tall. Mature stems reddish brown, obviously sulcate, glabrous; spines 3-fid, pointing downward, orange-brown, 1–3 cm, abaxially sulcate. Petiole 2–4 mm; leaf blade abaxially yellow-green, adaxially shiny, dark green, elliptic or broadly elliptic, occasionally ovate, 4–8 Χ 1.5–3 cm, thinly leathery, abaxially occasionally slightly pruinose, midvein conspicuously raised abaxially, obviously impressed adaxially, lateral veins slightly raised abaxially, slightly conspicuous adaxially, reticulate veins indistinct on both surfaces, base attenuate, margin 7–18(–25)-spinose-serrate on each side, apex acute. Inflorescence a fascicle, 2–6-flowered. Pedicel reddish, 7–13 mm; bractlets lanceolate, 4–5 Χ 1.5–2 mm. Sepals in 2 whorls; outer and inner sepals obovate-oblong, 6–10 Χ 2.5–5 mm, apex obtuse. Petals obovate, 4–5 Χ 2.2–3 mm, base clawed, with separate glands, apex emarginate. Stamens 2–5 mm; anther connective prolonged, truncate. Ovules 3–5, shortly funiculate. Berry black, not pruinose, ellipsoid, 8–9 Χ 5–6 mm; style persistent, short. Fl. Apr–May, fr. Jun–Oct.

* Thickets, forests, bamboo groves, scrub; 1300–2600 m. Guangxi, Guizhou.

The type collection is from Fan Ching Shan (Lei Shan), SE Guizhou.

23. Berberis calliantha Mulligan, Gard. Chron., ser. 3, 97: 394. 1935.

??vernacular name