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3. Mahonia Nuttall, Gen. N. Amer. Pl. 1: 211. 1818.
十大功劳属 shi da gong lao shu
Authors: Junsheng Ying, David E. Boufford & Anthony R. Brach
Odostemon Rafinesque.
Shrubs or small trees, evergreen, 0.3-8 m tall. Spines absent. Leaves imparipinnate, alternate, sessile or petiolate; petiole to 14 cm; leaflets 3-41; lateral leaflets usually sessile; terminal leaflet sessile or petiolulate; margins of leaflets entire, variously toothed, or with coarse or fine serrations. Inflorescence terminal, of (1-)3-18-fascicled simple or branched racemes or panicles, 3-35 cm, subtended by leafletlike bracts. Pedicel 1.5-24 mm, subtending bract shorter or longer than pedicel. Flowers yellow, with 3 whorls of sepals and 1 whorl of petals, with or without glands at base of petals. Anther connective not prolonged, apiculate or conspicuously prolonged. Ovary ellipsoid; ovules 1-7; styles absent or to 3 mm, persistent on mature fruit. Fruit berries, bluish or black, often glaucous. Seeds 1-7.
About 60 species: mainly in E and SE Asia, also in W North America, Central America, and W South America; 31 species (27 endemic) in China; six additional species (five endemic) are insufficiently known.
Most of the Chinese wild populations of Mahonia have been extirpated, probably as a result of over-collecting for medicinal use, and the plants now being described are probably individuals that vary only in minor ways from the relatively few specimens that exist in herbaria. Ahrendt (J. Linn. Soc., Bot. 57: 1-410. 1961) recognized two "groups" in Mahonia: "Orientales" and "Occidentales." All of the Asian species of Mahonia, plus a single North American species from the Pacific Northwest, M. nervosa (Pursh) Nuttall, are in the "Orientales." See the paper by J. Y. Wu, H. N. Qin, D. Y. Xue, and K. X. Zhou, "Study on seed morphology of Mahonia (Berberidaceae)" (Guihaia 30: 155-160. 2010).
Insufficiently known species
- Mahonia annamica Gagnepain, Bull. Soc. Bot. France 55: 84. 1908.
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Berberis annamica (Gagnepain) Laferrière.
This species was described from Vietnam and reported for Guangxi. We have not seen specimens from Guangxi and are unsure of its placement.
- Mahonia bijuga Handel-Mazzetti, Symb. Sin. 7: 331. 1931.
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This species was described from Sichuan ("Lose-schan s von S of Ningyüen," 3300 m), based on H. v. Handel-Mazzetti 1430 (?W), but we have not seen the type.
- Mahonia dulongensis H. Li, Ann. Bot. Fenn. 46: 469. 2009.
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This species was recently described from Yunnan (Gongshan, Dulong Jiang, 1620 m, 5 Nov 2005), based on Gaoligongshan Biodiversity Survey 21834 (holotype, KUN; isotypes, CAS, E).
- Mahonia hypoleuca Takeda, Notes Roy. Bot. Gard. Edinburgh 6: 238. 1917.
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Berberis nivea (C. K. Schneider) Laferrière; Mahonia nivea C. K. Schneider.
Both Mahonia hypoleuca and M. nivea are based on A. Henry 9863, from Mengtze, Yunnan, which is without flowers and fruit and the only collection known.
- Mahonia jingxiensis J. Y. Wu, M. Ogisu, H. N. Qin & S. N. Lu, Bot. Stud. (Taipei) 50: 487. 2009.
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This species was recently described from Guangxi (Jingxi Xian, Sanhe Xiang, Niansi Tun, ca. 500 m, 23 Oct 2007), based on J. Y. Wu & M. Ogisu 1023 (holotype, PE; isotype, IBK).
- Mahonia lushuiensis T. S. Ying & H. Li, Ann. Bot. Fenn. 46: 472. 2009.
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This species was recently described from Yunnan (Lushui County, Luzhang Township, 325 m, 19 May 2005), based on Gaoligongshan Biodiversity Survey 24522 (holotype, KUN; isotypes, CAS, E).
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| 1 |
Inflorescences panicles |
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(2) |
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Inflorescences racemes |
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(4) |
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| 2 (1) |
Petals retuse; pedicels 6-11 mm, bracts 2-3 mm; teeth per upper leaflet 4-11; ovules 5 or 6. |
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1 M. bracteolata |
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Petals emarginate or incised; pedicels 2.5-3 mm or 13-24 mm, bracts 1-2 mm or 4.5-7 mm; teeth per upper leaflet 1-5; ovules 1-4 |
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(3) |
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| 3 (2) |
Petals emarginate; inflorescences 25-35 cm; leaflets 2 or 3 pairs, abaxially pruinose; pedicels 13-24 mm. |
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2 M. gracilipes |
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Petals incised; inflorescences 11-18 cm; leaflets 5-9 pairs, abaxially not pruinose; pedicels 2.5-3 mm. |
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3 M. monyulensis |
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| 4 (1) |
Inflorescence of branched racemes |
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(5) |
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Inflorescence of unbranched racemes |
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(7) |
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| 5 (4) |
Petals incised; leaflets 12-20 pairs; ovules 2 or 3. |
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4 M. oiwakensis |
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Petals emarginate; leaflets 4-9 pairs; ovules 4-7 |
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(6) |
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| 6 (5) |
Bracts longer than pedicels; fruit obovoid, style less than 0.5 mm; leaf blade basally cuneate. |
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5 M. calamicaulis |
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Bracts shorter than to slightly longer than pedicels; fruit globose, style 2-3 mm; leaf blade basally oblique, rounded. |
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6 M. duclouxiana |
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| 7 (4) |
Petals entire |
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(8) |
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Petals emarginate or incised |
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(10) |
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| 8 (7) |
Petioles 3.5-14 cm; leaflets entire or with 1-3 inconspicuous teeth near apex. |
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9 M. shenii |
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Petioles 0.5-2.5 cm; leaflets with teeth |
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(9) |
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| 9 (8) |
Leaflets remote, not contiguous, apex acuminate or caudate-acuminate, margin with 9-23 teeth on each side; anther connective not prolonged; fruit globose. |
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7 M. taronensis |
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Leaflets contiguous, apex acute, margin with 2-7 teeth on each side; anther connective prolonged; fruit obovoid. |
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8 M. subimbricata |
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| 10 (7) |
Leaves petiolate, petiole 2.5-9 cm |
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(11) |
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Leaves subsessile or petiole less than 2 cm |
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(16) |
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| 11 (10) |
Inner sepals lanceolate or obovate; anther connective prolonged |
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(12) |
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Inner sepals oblong or elliptic; anther connective not prolonged |
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(13) |
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| 12 (11) |
Leaflets subrhombic or elliptic, margin with 2-4 teeth on each side; inner sepals obovate. |
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10 M. breviracema |
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Leaflets lanceolate, margin 8-20-spinose-serrate on each side; inner sepals lanceolate. |
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11 M. setosa |
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| 13 (11) |
Inflorescence 1- or 2-fascicled racemes; leaflets ovate or ovate-elliptic, basally truncate. |
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12 M. decipiens |
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Inflorescence 4-10-fascicled racemes; leaflets basally cuneate |
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(14) |
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| 14 (13) |
Leaflets 2 or 3 pairs, oblong-ovate or elliptic, shortly petiolate. |
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13 M. paucijuga |
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Leaflets 2-9 pairs, lanceolate to elliptic-lanceolate or elliptic, sessile |
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(15) |
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| 15 (14) |
Leaflets 2-5 pairs; pedicels ca. as long as bracts; glands of petals distinct. |
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14 M. fortunei |
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Leaflets 6-9 pairs; pedicels much longer than bracts; glands of petals distinct or sometimes indistinct. |
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15 M. eurybracteata |
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| 16 (10) |
Leaflets entire. |
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16 M. microphylla |
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Leaflets spinose-serrate |
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(17) |
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| 17 (16) |
Leaflets lanceolate, elliptic-lanceolate, or ovate-oblong |
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(18) |
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Leaflets oblong, ovate, broadly elliptic, or rhombic |
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(19) |
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| 18 (17) |
Leaflets 6-9 pairs, margin 3-9-spinose-serrate on each side; base of petals with glands. |
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15 M. eurybracteata |
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Leaflets 8-12 pairs, margin 35-65-spinulose-serrulate on each side; base of petals without glands. |
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17 M. leptodonta |
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| 19 (17) |
Veins on adaxial surface of leaflets reticulate, obviously raised, margin of leaflets 2-11-spinose-serrate on each side. |
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18 M. retinervis |
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Veins on adaxial surface of leaflets reticulate, flat or inconspicuous, margin of leaflets 2-16-dentate on each side |
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(20) |
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| 20 (19) |
Abaxial surface of leaflets pruinose; fruit 10-12 mm in diam. |
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19 M. bealei |
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Abaxial surface of leaflets yellowish green, not pruinose; fruit less than 10 mm in diam |
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(21) |
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| 21 (20) |
Floral bracts longer than pedicels |
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(22) |
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Floral bracts shorter than to ca. as long as pedicels |
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(25) |
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| 22 (21) |
Racemes 3-9 per fascicle, 5-9 cm; apex of petals slightly emarginate |
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(23) |
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Racemes 8-15 per fascicle, 7-19 cm; apex of petals incised |
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(24) |
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| 23 (22) |
Racemes 5-9 per fascicle; fruit not pruinose; leaflets serrate, with 4-7(-11) teeth on each side. |
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20 M. longibracteata |
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Racemes 3-5 per fascicle; fruit pruinose; leaflets spinose-serrate, with 10-16 teeth on each side. |
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21 M. polyodonta |
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| 24 (22) |
Anther connective prolonged; ovules 2-6; leaf blade adaxially dark green and shiny. |
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22 M. napaulensis |
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Anther connective not prolonged; ovules 2; leaf blade adaxially green. |
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23 M. cardiophylla |
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| 25 (21) |
Margin of leaflets 1-3(-6)-dentate on each side. |
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24 M. nitens |
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Margin of leaflets 4-10-dentate on each side |
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(26) |
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| 26 (25) |
Leaflets markedly overlapping |
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(27) |
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Leaflets separate or sometimes contiguous |
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(28) |
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| 27 (26) |
Inflorescence 3-6 racemes; anther connective prolonged; leaflets 8-18 pairs. |
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25 M. conferta |
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Inflorescence 9 or 10 racemes; anther connective not prolonged; leaflets 5-7 pairs. |
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26 M. imbricata |
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| 28 (26) |
Pedicels shorter than floral bracts; anther connective prolonged, rounded; petals 1.8-2 mm wide. |
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31 M. hancockiana |
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Pedicels longer than or ca. as long as floral bracts; anther connective not prolonged, truncate; petals 2-2.8 mm wide |
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(29) |
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| 29 (28) |
Fruit globose or pyriform, style not persistent; petiolar glands indistinct; leaflets 8-13 pairs. |
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27 M. bodinieri |
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Fruit ovoid, obovoid, or ellipsoid, persistent style short; petiolar glands conspicuous; leaflets 4-9 pairs |
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(30) |
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| 30 (29) |
Pedicels ca. as long as or slightly longer than floral bracts; leaf blade shiny, apically acute, cuspidate. |
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28 M. sheridaniana |
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Pedicels markedly longer than floral bracts; leaf blade apically acute or acuminate, sometimes aristate |
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(31) |
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| 31 (30) |
Leaflets 4-6 pairs, margin 2-4-dentate on each side; pedicels 6-7 mm; ovules 4-7. |
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29 M. japonica |
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Leaflets 5-9 pairs, margin 2-9-spinose-serrate on each side; pedicels 2.5-4 mm; ovules 2. |
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30 M. fordii |
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List of lower taxa
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