Corresponding author: Jun Wen (
Academic editor: Hanno Schaefer
The
The
The morphological differences among the species mostly concern pubescence and color on the abaxial leaf surface, teeth on the leaf margin, and the raceme length and density (
About 350 herbarium specimens from A, BM, CAS, CDB, E, GH, IBSC, K, KUN, L, MO, NY, PE and US were examined. We also conducted field studies in Fujian, Gansu, Hubei, Sichuan, Xizang and Zhejiang provinces of China. We herein provide a description of the
Trees or shrubs, deciduous. Winter buds ovoid, with several scales. Stipules caduceus, margin glandular at least on the lower part. Branchlet of first year’s growth pubescent. Leaves alternate, simple; leaf blade abaxially glabrous to tomentose; leaf margin toothed, lower part with a few to many glandular teeth, teeth simple, irregularly doubly serrate or incised-serrate. Inflorescence a terminal raceme, 8–20 flowered. Hypanthium pubescent. Perianth segments usually 10, narrowly triangular, caduceus, pubescent, not differentiated into sepals and petals. Stamens 20–45. Style slender, glabrous. Ovary 1- to rarely 2-locular, glabrous. Drupe ovoid, glabrous, dark purple to black.
Four species distributed in temperate regions of the Himalaya and eastern to western China. The morphology-based species delimitation will be tested by molecular data in our future work. At present, our available molecular data are congruent with our delimitation (
1a | Leaf blade pubescent to villous on the abaxial surface, or at least pubescent on veins | 2 |
2a | Leaf blade abaxially densely pubescent to rusty tomentose, lower part of margin densely with glandular teeth, bract and stipule margin glandular |
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2b | Leaf blade abaxially pubescent or sometimes only pubescent on the veins, lower part of leaf margin only with a few glandular teeth near the base |
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1b | Leaf blade glabrous on abaxial surface or with tufts of pubescence on lateral vein axils | 3 |
3a | Leaf margin with fewer (fewer than 15) glandular teeth near the base, pubescence on abaxial vein axils usually not conspicuous, leaf base acute to rarely subcordate |
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3b | Leaf margin with many (more than 20) glandular teeth at the lower 1/3 of the margin near the base, conspicuously with tufts of hairs on abaxial vein axils, leaf base subcordate to cordate |
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India. Sikkim: temperate, 8–10000 ft, bearing flowers and fruits, J. D. Hooker s.n. (lectotype: K!, here designated; isolectotype: K!).
Trees (2–) 3.5–10 m tall. Branches purple, slightly puberscent; branchlets of first year’s growth densely pubescent. Winter buds purplish brown, ovoid; scales to 3–23 × 3–12 mm, broadly to narrowly ovate, outside brown pubescent. Stipules lanceolate, 12–25 × 2–5.5 mm, membranaceous, caduceus, margin with glandular and fine teeth, glandular teeth at the lower 1/3–1/2, apex acuminate to acute, base rounded. Petiole 2–4.5 mm, densely brownish pubescent. Leaves ovate-oblong, elliptic to ovate, 5.5–13.5 × 2.7–6 cm, abaxially light green, densely pubescent, adaxially dull green and pubescent along the veins; margin doubly irregularly serrate to serrulate at the upper 2/3, glandularly serrulate at the lower 1/3, teeth at the margin sharp; apex acuminate, base subcordate to broadly cuneate; secondary veins 13–15 on either side of midvein. Racemes 3.5–6.5 cm, axis densely pubescent, (8–) 10–18-flowered; bracts broadly lanceolate, narrowly ovate, 6–14 × 2–5 mm, membranaceous, caduceus, pubescent on both surfaces, margin sparsely with glandular teeth. Pedicel 2.5–5 mm at anthesis, densely pubescent. Hypanthium campanulate, 3–4 × 5–7 mm, densely brownish pubescent outside, glabrous inside. Perianth segments 10, narrowly triangular, 1.5–2 × 1–1.3 mm, caduceus, pubescent. Stamens 30–40, 4–7.5 mm long; filament cream white, 3.5–7 mm; anthers oblong, pale yellow, 0.5–0.6 × 0.4–0.5 mm. Ovary glabrous, 1- or occasionally 2-locular, sometimes developing into twin fruits as shown in
Bhutan, Nepal, N India, N Myanmar and W China
Forest. Fl Apr-May; fr May-Aug; 2000–3500 m.
China. Sichuan: western Sichuan, southeast of Tachien-lu, 6–9000 ft, May 1908, E.H. Wilson 909 (holotype: A!, specimen barcode 00026559; isotypes: A!, 2 sheets, E!, K!, US!).
Shrubs to trees 1.5–10 (–15) m tall. Branches purple to dark purple, slightly pubescent; branchlets of first year’s growth densely pubescent. Winter buds purple brown, ovoid; scales to 3–20 × 3–12 mm, broadly to narrowly ovate, pubescent to slightly so on the outside surface, margin entire to slightly glandular on the upper scales. Stipules lanceolate, 12–22 × 2–6 mm, membranaceous or herbacious, caduceus, margin often with glandular teeth, especially on the lower part of the margin, apex acuminate to acute, base rounded or truncate. Petioles 2–4 mm, densely pubescent. Leaves elliptic to ovate, 4.5–14 × 2.5–6 cm, abaxially light green, pubescent or densely so especially on veins, adaxially dull green, sparsely pubescent with scattered short hairs and/or along the veins; margin mainly doubly irregularly serrate to doubly serrullate, glandularly serrulate at the lower part of the base with 2–15 glandular fine teeth; apex acute, attenuate or acuminate, base broadly cuneate to occasionally subcordate; secondary veins 16–20 on each side of midvein. Racemes 2.5–5 cm, 10–18-flowered; bracts broadly lanceolate to narrowly ovate, 5–12 × 1.5–4.5 mm, membranaceous, often caduceus, margin sparsely with glandular teeth, more or less pubescent on both surfaces. Pedicel 2–4 mm at anthesis, pubescent to densely so. Hypanthium campanulate, 3–4.5 × 3–6 mm, densely brownish pubescent outside, glabrous inside. Perianth segments 10, narrowly triangular to oblong-lanceolate, 1.8–2.4 × 0.8–1.1 mm, caduceus, densely pubescent. Stamens 25–35, 5–7.5 mm long; filaments 5–7 mm; anthers oblong, light yellow, 0.5–0.6 × 0.4–0.5 mm. Ovary glabrous, 1- or rarely 2-locular. Style slender, 7–9 mm long. Drupe ovoid, 7–10 × 4–5.5 mm, glabrous, dark purple to black.
Western to central China.
Forests. Fl Apr-May; fr Jun-Sep; 1800–3800 m.
China. Hubei: western Hubei, Hsing-Shan Hsien, bush 6–20 ft, woods, 4–6000 ft, flower greenish, May 1907, fl, E.H.Wilson 2850 (lectotype: A!, here designated, specimen barcode 00026557; isolectotypes: E!, K!, US!).
Shrubs to trees 1.5–6 m tall. Branches dark purple, glabrous; branchlets of first year’s growth sparsely pubescent at the very young part, then glabrescent. Winter buds ovoid to narrowly or broadly so, scales 3–15 × 3–8 mm, several, imbricate, pubescent on the outer scales, but glabrous or nearly so on the inner scales, margin more or less ciliate. Stipules linear to lanceolate, 7–17 x 1–4.5 mm, membranaceous, slightly pubescent to glabrous, margin glandularly ciliate at least on the lower part. Petiole to 2–6 mm, slightly pubescent, glabrescent. Leaves ovate, elliptic to broadly so, 3.5–16 × 1.3–7.5 cm, abaxially pale green, glabrous or often pubescent in lateral vein axils, adaxially glabrous; margin doubly irregularly serrate, with 1–5 glandularly serrulate teeth at the base; apex acute, attenuate or acuminate, base acute to rounded; lateral veins 14–16 on each side of midvein. Racemes 1.5–5.5 cm, with 8–15 flowers; bracts lanceolate to narrowly triangular, 4–5 × 1–2 mm, nearly glabrous, with glandular teeth at margin. Pedicel 1.5–4 (–6) mm long, pubescent. Hypanthium campanulate, 2.5–6 × 4–7 (–9) mm, slightly pubescent to glabrescent on the outer surface, glabrous on the inner surface. Perianth segments 10, slightly unequal, narrowly triangular to lanceolate, 1.5–3 × 0.9–1.2 mm, slightly pubescent to glabrescent. Stamens 20–30, 4–7 mm long; filaments 3.5–6.5 mm; anthers oblong, 0.25–0.35 × 0.2–0.25 mm. Ovary glabrous, 1– or rarely 2–locular (see E.J.Palmer 130, CAS). Style slender, 4–8 mm long. Drupe 6–8 × 5–6 mm, glabrous, dark purple to black.
Eastern, central to western China.
Forests. Fl Apr – Jun; fr late May–Jul; 1300–3700 m.
China. Yunnan: Gongshan Xian, Gongshan, on the way from Qingnatong to Anwalong, 3100 m, small tree 4 m tall, in the valley in shrublands, 31 May 1979, fl, flowers white, common, Lujiang Expedition 790292 (holotype: KUN!; isotype: KUN!).
Small trees 4–8 m tall. Branches purple, shiny glabrous; branchlets of first year’s growth pubescent. Winter buds purplish brown, ovoid; scales to 4–20 × 3–15 mm, ovate, outside brown pubescent but glabrescent. Stipules lanceolate to broadly so, 12–25 × 2–8 mm, membranaceous, margin with glandular teeth, apex acuminate to acute. Petiole 2.5–5 mm, brownish pubescent. Leaves oblong, elliptic to ovate, 5–13 × 2–6 cm, abaxially light green, nearly glabrous, only pubescent on lateral vein axils, adaxially dull green and glabrous, margin doubly irregularly serrate at the upper 2/3, glandularly serrulate at the lower 1/3, teeth at the margin sharp, apex acuminate, base subcordate to broadly cuneate; secondary veins 20–24 on either side of midvein. Racemes 4–6 cm, brown pubescent, 12–16 flowered; bracts lanceolate to ovate, 4.5–6 × 2.5–3.4 mm, membranaceous, margin entire to serrate. Pedicel 2.5–4 mm at anthesis, densely brown-pubescent. Hypanthium campanulate, 4–6 × 4–9 mm, brownish pubescent outside, glabrous inside. Perianth segments 10, narrowly triangular to lanceolate, 1.5–3.2 × 0.7–1 mm, caduceus, pubescent. Stamens 25–45, 5.5–7 mm long; filaments 5–7 mm; anthers oblong, 0.3–0.35 × 0.25–0.3 mm. Ovary glabrous, 1-locular. Style slender, 7–9 mm long. Drupe ovoid, glabrous, 8–9 × 5–6.5 mm, dark purple to black.
Western China, Bhutan, Myanmar, Nepal, and northern India.
Shady valleys and forests. Fl. Mar–Jul; fr. Jun–Jul; 2100–3500 m.
This species is named after the mountain range, the Gaoligong mountains or known as Gongshan, where this species was first recognized.
India. Mishmi Hills, Griffith s.n. (holotype: K!, K000396854).
As noted by
This study was supported by NSF Award number 0515431, the Smithsonian Endowment Grant Program, and the Small Grants Program of the National Museum of Natural History of the Smithsonian Institution. Field assistance was provided by C.-X. Fu, Z.-L. Nie, Y.-X. Qiu, T.-S. Yi and S.-L. Zhou. We thank A, BM, CAS, CDB, E, GH, IBSC, K, KUN, L, MO, NY, PE and US for permission to examine their specimens, either through loans or during visits. We thank Kai Wen for providing the illustrations, Sue Lutz for herbarium assistance, and Larry Dorr for discussions on nomenclature and historical references.