Berbers brevisepala Hayata (1913: 14)

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 : 76-78

2. Berbers brevisepala Hayata (1913: 14) . Type :— TAIWAN. Mt. Morrison (in protologue), “Central Mountains” (translated from Japanese) on the type, 15 April 1910, U. Mori s.n. (holotype TI-02620!) ( Fig. 8 ).

Heterotypic synonym:— Berbers alpicola Schneider (1939: 253) . Type :— TAIWAN. Ari-san to Mt. Morrison, bushy side of torrent, 3666 m, 24 October 1918, Wilson 10952 (holotype A-00038721!, isotype B-100365257, BM-000559458!, K-000644916!, US-00103858)

Evergreen shrub or occasionally small tree-like shrub 0.5–2 m tall. Mature stems brown or greyish, subterete, inconspicuous verruculose. Spines 3-fid, concolorous, 1.1–2.2 cm. Leaves subsessile; leaf blade obovate or elliptic, abaxially pale green sometimes pruinose, adaxially shiny green; 2.3–5.4 × 0.6–1.7 cm, leathery; midvein abaxially raised and adaxially impressed, lateral veins slightly raised, the secondary veins pinnate, jointly looped if present; base cuniform, margins slightly revolute, sparsely spinose with 2–7 spinules with 3–8 mm apart on each side, apex attenuate or mucronate. Inflorescence a fascicle, 3–5-flowered. Bracts absent. Pedicel pale green, 0.5–1.3 cm. Bracteoles usually 2, yellow or sometimes reddish tinged, triangular, 1.5 × 1 mm. Flowers bright yellow. Sepals in 4 or 5 whorls, outermost sepals of both 4- and 5-whorled flowers yellow with reddish-tinge triangular 1.5–2 × 1–1.5 mm, for flowers with 5- whorled sepals, the 4-whorled sepals yellow sometimes with reddish-tinge ovate 2.5 × 1.5 mm, the third-whorled sepals of both 4- and 5-whorled flowers yellow ovate 3.5 × 2.5 mm, sepals of the innermost two whorls yellow obovate 4–6 × 2.5–4 mm. Petals obovate, 4.5 × 3.5 mm, base clawed with a pair of ovoid nectaries, apex incised or acutely emarginated. Stamens pale yellow ca. 3.5 mm, anther connective of stamen distinct, apex truncate or slightly apiculate. Pistil 4 mm long. Ovules 3, 4 or 6. Berries black, ellipsoid ca. 5 × 3.5 mm, not pruinose, estylose, or occasionally up to 0.9 mm long.

Distinguishing features: —Given the wide distributional and elevational ranges of the species, it is not surprising that B. brevisepala is highly variable both among populations and within an individual plants, as shown by the type specimen ( Fig. 8C ). At the upper distributional limit (i.e., 3700 m), plants of B. brevisepla tend to possess smaller leaves ( Fig. 8A ) and such individuals were previously identified as B. alpicola (see Notes). Berberis brevisepala was often misidentified as B. kawakamii , or more frequently as B. nantoensis when reproductive organs are absent. However, B, brevisepala differs from B. kawakamii obviously by its outer ovate sepals (v.s. narrow-triangular or rarely linear sepals in B. kawakamii ) and from B. nantoensis by its much longer pedicels and the numbers of whorls of sepals.

Phenology: — Flowering May–August; Fruiting August–November.

Distribution & habitat: — Berberis brevisepala grows in subalpine meadows and the understory of coniferous forests throughout the major high mountain systems of Taiwan from 2100 to 3700 m ( Fig. 6E ). It is usually a small shrub less than 1 m tall; however, individuals more than 2 m tall can be found in wind-sheltered sites such as wet basins or stream valleys.

Chinese name:—¾山小ª

Proposed IUCN conservation status: —Nearly Threatened. Berberis brevisepala is common in subalpine and alpine regions along major high mountains in Taiwan, growing into small colonies in its habitats. Although most high mountain areas are under protection in Taiwan, the high mountain ecosystem is currently threatened by global climate change. We therefore propose a provisional IUCN category of NT for the species ( IUCN 2012).

Notes: — Berberis brevisepala was described based on Mori s.n. 1910, a specimen represented by leafy branches and fascicles of flowerless pedicels or early developing fruits. In the protologue of the species, no information regarding its floral (“ Flores non visi ”) and fruit morphology was provided ( Hayata 1913). The species status of B. brevisepala had long been controversial; the species had been synonymized under B. kawakamii or expanded to included B. alpicola ( Table 2). Because of the absence of flowers and fruits, the type specimen of B. brevisepala (Mori s.n. 1910) could not be included into our multivariate statistical analyses. Our initial attempt to visit the type locality was also hindered by the fact that the information given in the protologue (Mt. Morrison) is not found on the collection label which records only the ‘Central Mountains’. By analyzing Mori’s travel log ( Mori 2000), we identified Danda Major Wildlife Habitat (Danda MWH; Fig. 6A & 6E ) as most likely type locality of B. brevisepala and this conjecture was confirmed when plants possessing similar leaves to the type specimens were located in this area in April 2014.

With plants collected from Danda MWH (Guanmen Expedition-Harber & Yu 9, 10, 11) included, multivariate statistical analyses identified one grouping (Subgroup 5a) composed of specimens assignable to both B. brevisepala and B. alpicola (type included). Because no attributes could consistently separate B. brevisepala from B. alpicola , the latter species is synonymized under the former. Under this new circumscription, B. brevisepala has a wide distributional and elevational ranges and plants occurring in the upper distributional limits (ca. 3700 m), previously known as B. alpicola , tend to have much smaller leaves ( Fig. 8A ). Additionally, it was also noted in the protolouge of B. alpicola that ‘immature’ berries are stylose ( Schneider 1939, 1941) and some cultivated plants of ‘ B. alpicola ’ in U.K. also possess stylose berries with short style (Julian Harber, pers. comm.). However, close examination of the specimens from type locality [e.g., Yang 3618 (TNM)] indicated that all mature berries were estylose.

Additional specimens examined: — TAIWAN. Yilan: Chialohu ( Lake Chialo ), 2260 m, 11 May 2002, Huang 839 (HAST); near Fatushan, 2750 m, 10 August 2008, Yu 112 (TAI); Wulanshan, 2100 m, 16 August 2008, Yu 107 (TAI); on east-southern range of Nanhutashan, 2600 m, 12 August 2008, Yu 116 (TAI) . Hsinchu: Tapachienshan, 2600–3300 m, 15 May 2003, Lee 2532 (TAIF); near Kelayehshan, 3000 m, 6 April 2008, Yu 49 (TAI) . Taichung: the east peak of Mt. Syue , 3100 m, 19 July 2009, Huang 13084 (HAST) . Nantou: Shalihsiensi Forest Road, 2550–2680 m, 5 May 1998, Wang & Tsai 3169 (IBSC, TNM); Wuchieh Logging Trail near Shili River , 2200 m, 8 September 2008, Yu 140 (TAI); Hohuanshan, 3100 m, 23 December 2008, Yu 224 (TAI); near Antungchushan, 3000 m, 3 July 2009, Yu 414 (TAI) . Chiayi: Nitakayama ( Yushan ), 3300 m, 27 October 1918, Kanehira and Sasaki s.n. (TAIF); Peiyun Lodge to Tatachia Saddle, 3450– 2600 m, 11 August 1987, Yang 3618 (TNM), 3400 m, Peiyun Lodge, 17 May 1991, Lin s.n. (TNM), 8 May 1993, Chiu 1830 (TNM), 5 September 1995, Chen 1398 (TNM), 12 April 2009, Yu 434 (TAI) . Kaohsiung: Kuanshanlingshan, 24 August 1987, Kuoh 12484 (TNM), 5 April 1988, Kuoh 14236 (TNM); trail to Takuanshan , 2940 m, 20 May 1992, Wang 1077 (HAST); on the way from campsite to the top of Kuanshan, 3026–3150 m, 16 May 1995, Wang 1096 (HAST); near Kuanshan, 3100–3300 m, 11 October 1996, Liou 407 (TAIF) . Pingtung: the last water resource plot along the trail to Peitawushan , 2700 m, 8 May 2009, Yu 304 (TAI) . Taitung: Siangyang Lodge , 2200 m, 23 November 1987, Chen 713 (HAST) . Hualien: Tonkuran River, 15 April 1910, Mori s.n. (TAIF); Yangtoushan , 2800 m, 13 October 2008, Yu 162 (TAI); Rontaiwen Mts. , 2900 m, 12 April 2014, Guanmen Expedition-Harber & Yu 9, 10, 11 (TAI) .

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 8. Berberis brevisepala Hayata. A. Branches. B. Flowering branch. C. Leaves in the envelope of type specimen. D. Habitat and habit. E. Flowering branches. F. Flowers, exterior view. G. Flowers, exterior view, showing the sepals. H. Flowers. I. Flowers, exterior view, showing the sepals. J. Berries. A, B, F, G: Peiyun Lodge, type locality of B. alpicola, courtesy of Bo-Chang Chen; D, E: Guanmenshan; H: cultivated plant collected from Mt. Syue; I: Beitawushan; J: Shinkangshan.