Berberis hayatana Mizushima (1954: 31)

Yu, Chih-Chieh & Chung, Kuo-Fang, 2014, Systematics of Berberis sect. Wallichianae (Berberidaceae) of Taiwan and Luzon with description of three new species, B. schaaliae, B. ravenii, and B. pengii, Phytotaxa 184 (2), pp. 61-99 : 80

4. Berberis hayatana Mizushima (1954: 31) . Type:— TAIWAN. Taihoku (Taipei), Taiheizan ( Taipingshan ), Ratogun ( Lo-Tong ), 17 May 1917, Hayata s.n. (holotype TI!) ( Fig. 10A–C ).

Heterotypic synonym: — Berberis formosana Li (1952: 41) , nom. illeg. non B. formosana Ahrendt (1941: 24) . Type:— TAIWAN. Mountains near Muroroahu, Taihoku-syu, 17 Jul 1932, Suzuki 7258 (holotype TAI-156164!, isotype PH-66417!).

Evergreen shrub 0.5–1.6 m tall. Mature stems brown to reddish-brown, subterete, not verruculose. Spines 3-fid, concolorous, 0.6–2.3 cm. Leaves subsessile; leaf blade elliptic to narrowly-elliptic, abaxially green sometimes pruinose, adaxially dull dark green; 2.1–6.9 × 0.7–2.1 cm, thin-leathery; midvein abaxially raised and adaxially impressed, lateral veins slightly raised, the secondary veins pinnate, jointly looped if present; base cuniform, margins sometimes slightly revolute and remotely spinose with spinules of 3–11 with 2.5–7.0 mm apart on each side, apex acute or acuminate. Inflorescence a fascicle, 4–7-flowered. Bracts absent. Pedicel pale green, 0.6–1.7 cm. Bracteoles absent or 2, 3, red, triangular, 1.5 mm × 1 mm. Flowers bright yellow. Sepals in 2 whorls, outer sepals yellow with reddish-tinged ovate 3.5 × 2 cm, inner sepals yellow obovate 4.5 × 3 mm. Petals obovate, 4.5 × 3 mm, base clawed with a pair of ovoid nectaries close to each other, apex incised. Stamens pale yellow ca. 3 mm, anther connective of stamen distinct, apex truncate. Pistil 4 mm long. Ovules 2 or 4. Berries black, ellipsoid ca. 7 × 4 mm, not pruinose, estylose.

Distinguishing features: Berberis hayatana was synonymized under B. mingetsensis in Lu & Yang (1996) and Yang et al. (1997); however, it can be easily distinguished from the latter by its unique dull adaxial leaf surfaces and generally shorter pedicels (0.6–1.7 vs. 1.3–2.8 cm). In areas of the northern Taiwan where winter northeastern monsoon are strong, B. hayatana can be found as low as 700 m. Notably, some individuals of B. hayatana from extreme humid environments (e.g., Yaungyang Lake, Lalashan) possess leaves with narrow-elliptic or lanceolate leaves with weakly developed spinose leaf margin.

Phenology: — Flowering February–May; Fruiting September–to February of the following year.

Distribution & habitat: — Berberis hayatana is relatively common in the understories of wet broad-leaved and coniferous and broad-leaved mixed forests from 700 to 2200 m in the northern Syue Mountain Range and northeastern Central Mountain Range ( Fig. 6B ).

Chinese name: va氏小ª

Proposed IUCN conservation status: —Nearly Threatened. Berberis hayatana is common in mountain areas of the northeast Taiwan, usually growing into large colonies. Although abundant, populations distributed in lower elevation are potentially threatened by human disturbance. Therefore we propose a provisional IUCN category of NT for the species ( IUCN 2012).

Notes: Berberis hayatana Mizushima was described by Mizushima (1954) to replace B. formosana H.L. Li , a later homonym of B. formosana Ahrendt.

Additional specimens examined: — TAIWAN. Taipei: Peichatienshan, 15 December 1972, Kuo 2609 (TAI); Lupeishan , 1400 m, 1 March 1989, Hsieh 687 (TAI); Ayushan, 1000–1400 m, 13 September 2001, Chung 4626 (TAIF); Dapolushan, 1400 m, 15 March 2009, Yu 263 (TAI); Nanchatienshan, 2200 m, 29 February 2009, Yu 261 (TAI) . Yilan: Chilanshan, 2 April 1984, Shih 310 (TAIF); Yuanyang Lake , 1600–1700 m, 10 January 1995, Shen 324 (HAST), 2100 m, 1 December 1 2008, Yu 190 (TAI); Taipingshan, 2000–2200 m, 27 March 2008, Yang 1528 (HAST), 2200 m, 3 April 1988, Kao 10685 (TAI); along Chilan No. 100 Forest Road, 1720 m, 28 October 2008, Huang 3551 (HAST); Songlohu, 700–1300 m, 28 March 2009, Lu 18081 (HAST); Tongshan, 2000 m, 9 April 2009, Yu 311 (TAI); Taipingshan, 2000–2200 m, 27 March 2008, Yang 1528 (HAST); Siatonshan, 2000 m, 3 April 2009, Yu 309 (TAI); at Taipingshan, along Taiwn Beech National Trail, 1900 m, 28 June 2011, Peng et al. 23110 (HAST) . Hsinchu: Chenhsibao , 1850–2200 m, July 1999, Wu 1819 (HAST); Litungshan, 1500–1900 m, 21 July 2002, Lu 4517 (TAIF); Sichiousihshan, 2000 m, 18 December 2008, Yu 252 (TAI) . Taoyuan: route from Meikueihsimoshan to Papokulushan , 1840 m, 11 April 2002, Leong 2880 (HAST); Pafu History Road, 1650 m, 27 February 2008, Huang 3277 (HAST); Lalashan, 18 March 2006, Lu 11513 (HAST) .

Figures

FIGURE 6. Distributions of Berberis sect. Wallichianae of Taiwan and Luzon. A. Major mountain ranges in Taiwan, with possible type localities of B. aristatoserrulata and B. brevisepala. NR: Nature Preserve. MWH: Major Wildlife Habitat. Mts: Mountain Range. B. Distributions of B. aristatoserrulata, B. hayatana, and B. pengii. C. Distributions of B. chingshuiensis, B. mingetsensis, and B. nantoensis. D. Distributions of B. kawakamii, B. ravenii, and B. schaaliae. E. Distributions of B. brevisepala and B. tarokoensis. F. Distribution of B. barandana in Luzon Island of the Philippines.

FIGURE 10. Berberis hayatana Mizush. (A–C) and B. mingetsensis Hayata (D–H). A. Flowering branch, showing the dull upper surface of leaves. B. Flowers, front view. C. Fruits. D. Leaves, waxy-shining adaxially. E. Leaves, pruinose abaxially. F. A solitary flower and a highly reduced umbel with two flowers, showing the long pedicels and bracts. G. Flower, front view. H. Berry. A, B: Lalashan; C: Tamenshan; D–H: Mienyueh.A, B, D–G: courtesy of Chi-Kai Yang.