Berberis kawakamii Hayata (1911a: 24–25)

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 : 80-82

5. Berberis kawakamii Hayata (1911a: 24–25) . Type :— TAIWAN. Monte Morrison, ad 9000 ped. alt., Oct 1906, T. Kawakami 1941 (holotype TI-02622!) ( Fig. 11A–C ).

Heterotypic synonyms:— Berberis formosana Ahrendt (1941: 24) . Type:— TAIWAN. Prov. Kagi, Arisan, 25 October 1918, Wilson 10910 ” (holotype BM-001015554!, isotypes A-00038750!, K-000644915!, US-00956032, n.v., image seen); B. kawakamii var. formosana (Ahrendt) Ahrendt (1961: 65) .

Evergreen shrub or small tree-like shrub 0.5–3 m tall. Mature stems greyish or brownish-yellow, inconspicuously verruculose. Spines 3-fid, concolorous, 0.8–2.7 cm. Leaves subsessile; leaf blade elliptic, elliptic-obovate, oblong, oblong-lanceolate, lanceolate, abaxially pale green, adaxially green; 2.9–7.2 ×0.8–2.0 cm, leathery; midvein abaxially raised and adaxially impressed, lateral veins slightly raised, the secondary veins pinnate, jointly looped; base cuniform or attenuate, margins remotely spinose with spinules of 5–15 with 2.5–6.0 mm apart on each side, apex acuminate. Inflorescence a dense, congested fascicle of 6–12 flowers. Bracts absent. Pedicel red, 0.3–1.2 cm. Bracteoles absent. Flowers yellow. Sepals in 3 whorls, outer sepals yellow with reddish-tinged narrowly-triangular or rarely linear 3–6 × 1 mm, middle sepals yellow triangularly-oblong 5 × 2 mm, inner sepals yellow narrowly obovate 5.5 × 2.5 mm. Petals narrowly-oblong 5 × 1.5 mm, base clawed with a pair of ovoid nectaries very close to each other, apex incised with obtuse lobes. Stamens pale yellow ca. 3 mm, anther connective of stamen distinct, apex truncate. Pistil 4 mm long usually with reddish-tinged. Ovules 2 or 3. Berries black, ellipsoid ca. 5 × 3.5 mm, sometimes slightly pruinose, stylose ca. 0.5–1 mm.

Distinguishing features: The combination of triangular-oblong outer sepals and a densely fascicled inflorescence makes this an easily distinguished species ( Fig. 1L & 11B ). Early literatures often allied B. kawakamii with the Filipino B. barandana ( Hayata 1908, Kawakami 1910, 1911), such as those by Merrill (1923) who stated that “ I failed to find any reason for distinguishing the Formosan B. kawakamii Hayata. ” However, these two species can easily be differentiated by the shape of outer sepals and as noted above molecular analysis places them in different clades ( Fig. 5 ).

Phenology: — Flowering March–May, August; Fruiting January, April–July, August–December.

Distribution & habitat: — Berberis kawakamii is very well represented in herbaria of Taiwan and is undoubtedly the commonest species of sect. Wallichianae in Taiwan, occurring throughout all major mountain systems north of Beinanchushan from 1600 to 3100 m ( Fig. 6A & D ). Across its wide distributional ranges, B. kawakamii is found in various habitats, ranging from subalpine and alpine meadows, montane coniferous forests, to coniferous and broad-leaved mixed cloud forests and exhibits substantial local morphological variation in both height and leaf shapes and size.

Chinese name: fi灣小ª

Proposed IUCN conservation status: —Nearly Threatened. Berberis kawakamii is common throughout major high mountains of Taiwan, usually growing into large colonies. However, the high mountain ecosystem of Taiwan is potentially threatened by the effect of global climate change, we therefore propose a provisional IUCN category of NT for the species ( IUCN 2012).

Notes: Berberis kawakamii var. formosana (Ahrendt) Ahrendt was first synonymized under B. kawakamii Hayata by Li (1963), a treatment that is confirmed by our multivariate statistical analyses. The name ‘ Berberis miyabei Kawakami’ in sched. was discovered on a specimen of B. kawakamii with the label ‘ T. Kawakami s.n. (1907) & U. Mori s.n. (1907)’ in the Herbarium of the University of Tokyo (TI). Accompanied with the specimen, there is also a typed description of B. miyabei Kawakami in sched. by Takiya Kawakami, to who B. kawakamii was attributed to.

Additional specimens examined: — TAIWAN. Yilan: Nanhu River , 2800–3000 m, 13 May 1989, Leu 24993 (TAIF); Szuyuanyakou, 1600 m, 5 April 2008, Lu 15630 (HAST); Wulanshan, 2400 m, 12 August 2008, Yu 117 (TAIF) . Hsinchu: Chiuchiushanchuang (“99 Lodge ”), 2694 m, 6 September 1993, Huang 45 (HAST); Kelayehshan, 3000 m, 6 April 2008, Yu 50 (TAI) . Miaoli: Leshan-Kuanwu , 2000–2300 m, 24 July 1987, Wang & Yang 4685 (TAI) . Taichung: Anmashan , 2200 m, 22 April 2004, Wang 7290 (TNM); Chica Lodge to 369 Lodge, 2900 m, 12 November 12 1989, Her 14 (TNM), Chika Lodge, 2600 m, 18 August 2008, Yu 99 (TAI); Taoshan, 3300 m, October 1981, Ou s.n. (TNM); en route from Wuling Lodge to Taoshan, 2600 m, August 1988, Peng 12018 (HAST); Tienchih-Nengkaopeifeng, 3100 m, 16 June 1996, Chiu 3433 (HAST); along trail from Tienluanchih to Hohuanpeifeng, 2950 m, May 2002, Huang 1089 (HAST) . Nantou: Chichia Lake , 2800–2900 m, 3 September 1998, Chen 321 (TAI); Pass Nengkao, 14 August 1955, Hsii 19 (TAI); Hohuanshan, 26 April 1985, Lu 15953 (TAIF); roadside, Pilushan, 2600–2900 m, 25 July 1990, Lin & Lin s.n. (TNM); Kuankao-Chunyangchinkuang, 2500–2900 m, 8 July 1993, Yang 5287 (TAI); Chuntashan, 3175 m, 8 March 1999, Liou 1147 (TAIF); Chilaishan shelter, 2700–3000 m, 30 April 2004, Chung 1013 (TAI); Puli, 18 March 2009, Yang 6203 (TNM); Hsiluantashan, 2850 m, 11 April 2009, Yu 399 (TAI) . Chiayi: Alishan-Chushan , 2300 m, 22 March 1985, Huang 2584 (TAI); Tatashan, 2400–2663 m, 23 September 2000, Kuo 98 (HAST); Tatakaanpu- Yushanchienfu, 3100 m, 11 November 1990, Wu & Lai s.n. (TNM); Yushan, 3300 m, 13 April 1996, Yang 4684 (TAIF) . Kaohsiung: trail to Takuanshan , 2690 m, 20 May 1992, Wang 1074 (HAST); on the way from Chinching Bridge to a campsite by the trail to Kuanshan, 2500–2700 m, 17 May 1995, Wang 1122 (HAST); Kuanshanlingshan, 2900–3175 m, 12 August 2002, Lee 244 (TAIF); Shunyunshan, 2500 m, 31 March 2003, Lu 5727 (TAIF); Kuanshan, 3050 m, 2 February 2008, Yu 9 (TAI) . Taitung: South Cross Highway , 2600 m, 30 March 1996, Lu 25003 (TAIF) . Hualien: Yuanfeng adjacent to Prov. Rd. 14, 2700–2800 m, 28 March 1994, Chen 469 (HAST); near Liwuchushan, 3000 m, 21 November 2004, Yang 804 (TNM); Yangtoushan, 2900 m, 12 October 2008, Yu 158 (TAI); Chukushan, 2200 m, 27 January 2012, Yu 693 (TAI) .

Figures

FIGURE 1. Diagnostic morphological attributes. A. Simple-veined leaf, with prominent midvein and pinnate, jointly looped secondary veins (B. brevisepala). B. Complex-veined leaf, secondary veins are pinnate, jointly looped and multi-festooned, and the tertiary veins are weakly reticulated (B. pengii). C. Simple-veined leaf with dull and silky-luster adaxial surface (B. hayatana). D. Complex-veined leaf with waxy-shinning adaxial surface (B. mingetsensis). E. Leaf specimen of B. hayatana, showing the dull adaxial surface when desiccated. F. Leaf specimen of B. brevisepala, showing the simple-vein type and waxy-shinning adaxial surface when desiccated. G. Brownish mature stem (B. hayatana). H. Purplish-red mature stem (B. ravenii). I. Bracts (B. mingetsensis). J. Bracteoles (white arrow) and outermost sepals (black arrow) in B. brevisepala. K.A dense, congested fascicle inflorescence with invisible pedicels (B. nantoensis). L. A congested fascicle inflorescence with long pedicels in B. kawakamii, showing the narrowly triangular-oblong outermost sepals (black arrow). M. The round (ovate or obovate) outermost or first whorl of sepals (B. tarokoensis). N. Ellipsoid, stylose berry (B. tarokoensis). O. Globose or nearly globose, estylose berries (B. pengii). A, C, D, N, and O: courtesy of Chi-Kai Yang.

FIGURE 5. The best ML tree from the analysis of three concatnated chloroplast markers. Numbers adjacent to the nodes are ML bootstrap supports (BS) and BI posterior clade probabilities (PP). The well supported clades are denoted by thick line (BS> 50 and PP> 0.6).

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 11. Berberis kawakamii Hayata (A–C) and B. barandana S. Vidal (D–H). A. Flowering branches. B. Flowers, showing the reddish, narrow-triangular outer sepals. C. Stylose, ellipsoid fruits. D. Habitat and habit. E. Fruiting branches. F. Flower, exterior view, showing the bracteoles, and outermost sepals. G. Flower, front view, with parts of the petals removed. H. Flowering branch, showing the pair of bracts.A, B: Hohuanshan; C: Liwuchushan, courtesy of Chi-Kai Yang; D–H: Mt. Santo Thomas, Luzon, Philippines.