Sanicula lamelligera Hance (1878: 11)

Li, Hui-Min & Song, Chun-Feng, 2022, Taxonomic studies on the genus Sanicula (Apiaceae) from China (I): The identity of S. orthacantha var. pumila and S. pengshuiensis, Phytotaxa 532 (2), pp. 114-138 : 134-136

Sanicula lamelligera Hance (1878: 11) . Figs. 1‒7 , 9‒13 , 17A .

Type:— CHINA, Guangdong, North River , March 1877, T.L. Bullock s.n. (Herb. Hance 19942) [K000697286!, lectotype designated by Pimenov (2017: 204), isolectotype BM000944682!]. Fig. 1 A-upper part, 1B-lower part .

= Sanicula satsumana Maximowicz (1886: 47) .

Type:— JAPAN, Satsuma (= the western half region of Kagoshima Prefecture on the island of Kyūshū ), Kago-sima (= Kagoshima), October 1883, Y. Tashiro s.n. (holotype LE01029618!). Fig. 2 .

= Sanicula yunnanensis Franchet (1894: 108) .

Type:— CHINA, Yunnan, Zhaotong, along the Yang-tze-kiang River , Kéou-pa-tan, March 29 1882, J.M. Delavay 453 [P03226688!, lectotype designated by Pimenov (2017), isolectotypes A00589886!, NY00406259!, P03226687!). Fig. 3 .

= Sanicula orthacantha var. pumila Boissieu (1906: 421) , syn. nov.

Type:— CHINA, Kouy-Tchéou (= Guizhou), Pin-fa in southwestern Guiding county, 1902, P.J. Cavalerie & J.J. Fortunat 1721 (P03226692!, lectotype designated here, isolectotypes K!). Remaining syntypes: same locality, July 12 1900, É. Bodinier 2718 (P03226683!, P03226685!). Fig. 4 .

= Sanicula ichangensis Wolff (1913: 54) .

Type:— CHINA, Hubei, Yichang , October 1887, A. Henry 3525a [K!, lectotype designated by Pimenov (2017: 204); isolectotype P03226671!]. Remaining syntype: Same locality, October 1887, A. Henry 3526a (BM!, K!). Fig. 5A, B .

= Sanicula orthacantha var. longispina Wolff (1913: 55) .

Type:— CHINA, Hubei, Yichang , March 16 1900, E.H. Wilson 156 [K!, lectotype designated by Pimenov (2017), isolectotypes E00000710!, US03074345!]. Fig. 5C, D .

= Sanicula lamelligera var. wakayamensis (Masamune) Murata (1962: 195) . S. wakayamensis Masamune (1941: 340) .

Type:— Japan, Honsyū, Kamegawa, Kaisô-gun, Wakayamken, Septermber 22 1940, Y. Syȏzi s.n. (TAI084140! lectotype designated here, isolectotypes TAI084139!, TAI084141!, TAI084142!). Fig. 6 .

= Sanicula pengshuiensis Sheh & Liu (1991: 469) , syn. nov.

Type:— CHINA, Chongqing, Pengshui, Hanjia town , 500 m, April 11 1979, S.X.Tan 106 (holotype SM715005741!, isotypes NAS00082800!, SM715005739!). Fig. 7A, B .

Description:—Perennial. Rhizome short, woody, bearing fibrous roots, occasionally with nodes. Stems slender and erect, few branched above the middle, 8‒40 cm tall. Basal leaves 3‒30, long petiolate; petioles (2‒)5‒20(‒28) cm long; blade glabrous on both surfaces, sometimes purplish red abaxially, (0.8‒) 2‒7 cm long, (1.5‒) 4‒15 cm broad, cordatesuborbicular, ternate, segments distinctly petiolate, the median segments cuneate-obovate to rhombic, obtuse to acute and more or less trilobed at the apex, the lateral segments oblique, bilobed to the middle or only notched or rarely entire, the margins crenate-serrate with spinose teeth. Upper stem leaves reduced, 1 or 2, trisect or undivided, bractlike, linear-lanceolate to obovate-lanceolate, ca. 5 mm long, shortly petiolate or sessile. Inflorescence 1- or 2- to severalbranched; involucrate bracts 3‒6, trisect or linear-lanceolate, ca. 2 mm long; rays of the umbels 1‒7, ca. 3.5 mm long; involucellate bracteoles 4‒6, linear to lanceolate, ca. 0.6 mm long. Umbellules 4‒7-flowered. Staminate flowers 3‒6 per umbellule, pedicels ca. 1.5 mm long, petals white, pinkish or bluish purple. Fertile flower 1 per umbellule, sessile; calyx teeth linear to lanceolate, 0.3‒1 mm long; styles 1.5‒5 mm long, sessile, 1.7‒7 times longer than the calyx teeth. Mericarps are oblong-ovoid to ovoid, ca. 2.5 mm long, ca. 2 mm broad, variably spiculate-lamellate, comblike. Vittae 1 under each rib and 2 on commissure.

Distribution and habitat:— Sanicula lamelligera is widely distributed in central, eastern, southern, and southwestern China and southern Japan ( Fig. 19 ). It grows in forests on mountain slopes or along ravines and streams at elevations of 450‒1600 m above sea level.

Phenology:—Flowering from late February to June; Fruiting from March to October.

Additional specimens examined:— CHINA. Anhui: Qimen, K. Yao 10293 (NAS); Shexian, Anhui Exped. 1521 (NAS), Anhui Wild Plant Exped. 1780 (NAS, PE); Xiuning, Anhui Wild Plant Exped. 2235 (NAS). Chongqing: Chengkou, T.L. Dai 100226 (PE), T.L. Dai 100521 (PE); Hechuan, Hechuan Exped. 740 (SM), Hechuan Exped. 821 (SM); Pengshui, H.M. Li, X. Zhang & Y.S. Zhang 1129 (NAS), H.M. Li, X. Zhang & Y.S. Zhang 1131 (NAS), S.X. Tan 1259 (NAS, SM); Qianjiang, Anonymous 131 (SM); Shizhu, Anonymous 589 (NAS, SM), H.M. Li, X. Zhang & Y.S. Zhang 1126 (NAS); Wulong, Z.Y. Liu 182668 (PE); Wushan, T.P. Wang 10603 (PE, WUK), K.H. Yang 57629 (IBK, PE), K.H. Yang 57731 (IBK, PE); Wuxi, P.Y. Li 1950 (NAS, WUK); Xiushan, Anonymous 359 (SM); Zhongxian, A.G. Luo 1210 (SM). Fujian: Jiangle, Longxishan Exped. 1004 (PE); Yongtai, Anonymous s.n. (PE). Guangdong: Lechang, W.T. Tsang 20972 (NAS, PE, US); Lianshan, P.C. Tan 58270 (NAS); Xinyi, Z. Huang 32084 (PE); Zengcheng, W.T. Tsang 20145 (NAS, PE); Zijin, C.F. Wei 120979 (NAS, PE). Guangxi: Luocheng, Luocheng Exped. 451225130428001LY (GXMG, IBK); Yongfu, K.J. Yan & J.Y. Huang 46733 (GXMI). Guizhou: Guiyang, Sun Yat-sen Univ. Biol. Depart. 50221 (PE); Huishui, G.M. Li 7 (ZY); Libo, Q.H. Chen 2154 (HGAS), Q.H. Chen 2294 (HGAS); Pingtang, Guizhou Exped. 292 (HGAS); Qinglong, C.Z. Dang 557 (HGAS); Suiyang, P.C. Tsong 485 (KUN, PE). Hubei: Badong, H.C. Chow 207 (N, NAS, PE), T.P. Wang 10938 (NAS, PE, WUK); Wufeng, D.G. Zhang et al. 34 (HHE); Xianfeng, Y.S. Zhang 1121 (NAS); Xingshan, H.M. Li & Y.S. Zhang 1085 (NAS), H.M. Li & Y.S. Zhang 1087 (NAS), H.M. Li & Y.S. Zhang 1088 (NAS); Xuan’en, H.M. Li, Y.S. Zhang & Y.M. Yi 1079 (NAS), H.M. Li, Y.S. Zhang & Y.M. Yi 1098 (NAS); Yichang, exact locality unknown, A. Henry 1304 (K, P), A. Henry 3525 (E, K); Yiling, P.X. Dong 21 (PE), Z. Zhen 68 (PE); Zigui, J.Q. Zhang 234 (PE). Hunan: Guzhang, P.W. Sweeney & D.G. Zhang PWS2694 (JIU), D.G. Zhang LL20130303035 (JIU), D.G. Zhang y090807072 (JIU); Huayuan, K.G. Li & C. Zhang HY20160314_0006 (JIU); Jishou, D.G. Zhang 130227002 (JIU), D.G. Zhang YH080308998 (JIU); Longshan, D.K Tian, Y. Xiao & Y. Chen LS-1077 (CSH); Sangzhi, Anonymous 230 (PE); Shimen, Hupingshan Exped. 217 (PE); Xinning, L.B. Luo 471 (P); Xinshao, Z.Z. Li 42 (NAS); Yizhang, L.H. Liu 631 (IBK); Yongshun, D.G. Zhang zdg140506246 (JIU). Jiangxi: Dexing, Anonymous 11583 (NAS), M.B. Deng 87090 (NAS); Fengxin, S.L. Liu, K. Yao & X.Z. Wang 1270 (KUN, NAS, PE, WUK); Fenyi, S.H. Lai 44 (LBG, PE), H.M. Li, Y.S. Zhang & Y. Xu 1107 (NAS); Fuliang, Q.H. Li & C. Chen 606 (LBG, PE); Shangyou, Jiangxi Exped. 68 (PE); Wuyuan, Q.H. Li & C. Chen 389 (PE). Macau: exact locality unknown, M. Callery 44 (P). Sichuan: Baoxing, Baopu Exped. 78-0003 (SM); Chongzhou, Anonymous 79-005 (SM); Dachuan, Daxian Exped. 262 (SM); Dazhu, Anonymous 225 (SM); Dujiangyan, D.E. Boufford & B. Bartholomew 24200 (HAST, US), D.Z. Fu 87‒0186 (WCSBG), K.C. Kuan, J.W. Wang & C.L. Li 781 (CDBI, PE), X.Q. Li 63 (CDBI, NAS), M.L. Sheh 83640 (NAS), M.L. Sheh 88639 (NAS), C. Zhang 20060167 (WCSBG), C. Zhang 20060184 (WCSBG), C. Zhang & L. Yu 2006238 (WCSBG), Z.L. Zhao 87‒0093 (WCSBG); Ebian, Ebian Exped. 189 (SM), Ebian Exped. 356 (SM), Sichuan Med. Team 1080 (SM); Emei, S.Y. Chen et al. 3006 (NAS, PE), W.C. Cheng 10821 (KUN), T.Y. Cheo & G.J. Xu 9 (NAS), C.Y. Chiao, C.S. Fan & P.Y. Tan 384 (E, HAST, N), H.C. Chow 7592 (NAS), W.P. Fang 2625 (E, PE, US), W.P. Fang 7431 (PE), W.P. Fang 12715 (PE), W.P. Fang 12749 (PE), C.H. Hsiung, S.S. Chang & H.L. Tsiang 30089 (NAS, PE), K.C. Kuan, J.W. Wang & C.L. Li 717 (CDBI, PE), K.C. Kuan, J.W. Wang & C.L. Li 1710 (CDBI, PE), H.M. Li & W. Zhou 1040 (NAS), H.M. Li & Y.S. Zhang 1152 (NAS), X.M. Lu et al. 2008050920 (CDCM), X.B. Peng 6009 (IBK, PE), Sichuan Med. Exped. 12013 (NAS), S.L. Sun 2 (TIE), S.L. Sun 34 (TIE), S.L. Sun 1417 (NAS, TIE), S.L. Sun 1450 (NAS, TIE), H.L. Tsiang, S.S. Chang & C.H. Hsiung 37107 (NAS, PE), E.H. Wilson 4927 (K, P), K.H. Yang 54534 (KUN, NAS, PE), Yang, Zhang & Chen 2008111308 (CDCM); Gongxian, Anonymous 203 (SM), K.C. Kuan, J.W. Wang & C.L. Li 781 (CDBI, PE); Hongya, Hongya Exped. 260 (SM); Kaijiang, Kaijiang Exped. 26 (SM); Leibo, M.Y. He & Q.S. Zhao 116727 (CDBI), C.T. Kuan 8984 (NAS, PE), Z.M. Tan & M.Y. He 121131 (CDBI), K.P. Yin & Q.H. Chen 124 (CDBI), Q.S. Zhao & M.Y. He 120920 (CDBI), Q.S. Zhao & Z.J. Zhao 121205 (CDBI); Linshui, Anonymous 250 (SM); Lushan, D.Y. Peng 47644 (CDBI); Meigu, Sichuan Veg. Exped. 13586 (CDBI, NAS); Muchuan, Muchuan Exped. 484 (SM); Pingshan, R.H. Shan 1399 (NAS, PE), Sichuan Econ. Plant Exped. 674 (CDBI, KUN, PE, SM), Sichuan Econ. Plant Exped. 749 (CDBI, PE), Sichuan Econ. Plant Exped. 884 (PE), T.T. Yu 2958 (PE); Qingshen, Sichuan Econ. Plant Exped. 5155 (PE); Tianquan, K.L. Chu 2299 (NAS, PE), D.Y. Peng 45446 (CDBI), D.Y. Peng 45562 (CDBI), D.Y. Peng 45616 (CDBI); Wenchuan, Maoxian Exped. 631 (SM), Sichuan Veg. Exped. 8416 (CDBI, NAS); Xingwen, Xingwen Exped. 77‒470 (SM); Xuyong, Anonymous 1169 (SM); Xuzhou, Yibin Exped. 206 (SM); Yingjing, Yingjing Exped. 78‒0148 (SM); Yucheng, Ya’an Exped. 5 (SM). Taiwan: Chiayi, Y.F. Chen 5120 (TAI); Hsinchu, Y. Simada 3399 (TAI); Hualien, P.F. Lu 13994 (HAST), T. Nakamura 4591 (TAI); Taipei, Y.T. Chen 2250 (TAI), K.F. Chung, Y.C. Kao & S.H. Wu 1115 (HAST), C.C. Hsu 6704 (TAI), B.W. Huang 42 (HAST), Y.Y. Huang & J.C. Chen 420 (HAST), C.I. Huang, J.C. Chen & J.H. Pak 1895 (HAST), T. Ito s.n. (TAI), M.T. Kao 3607 (TAI), M.T. Kao 3630 (TAI), Y.C. Kao & S.H. Lai 299 (HAST), Y. Kudo 889 (TAI), H.C. Lee & M.T. Kao 3936 (TAI), S. Sasaki s.n. (TAI), H. Simada 107 (TAI), H. Simada 991 (TAI), Y. Simada 1419B (NTUF), S. Suzuki 6413 (P), S. Suzuki 10236 (TAI), S. Suzuki s.n. (TAI); Yilan, C.I. Huang 6887 (HAST), C.I. Huang & M.L. Weng 22035 (HAST), C.C. Liao & C.C. Chen 1794 (HAST), T.Y. Liu et al. 164 (HAST), T.Y. Liu, C.C. Wang & T.T. Liu 227 (HAST), C.I. Peng 11332 (HAST), C.I. Peng et al. 13817 (HAST), C.I. Peng et al. 15404 (HAST), C.I. Peng 16263 (HAST), S. Suzuki 103 (TAI). Yunnan: Zhaotong, Delavay 4900 (P), R.P. Maire s.n. (P). Zhejiang: Kaihua, Zhejiang Plant Resour. Exped. 26213 (NAS); Lin’an, H.M. Li & L. Zhao 1115 (NAS); Longquan, S.Y. Zhang 4488 (AU, PE, WUK); Ningbo, Anonymous 8177 (NAS), Zhejiang Mus. Exped. 3125 (IBK, WUK); Qingyuan, F.G. Zhang 5081 (ZM); Suichang, Chang et al. 5207 (ZM), Chang et al. 5228 (ZM), Chang & Li 5335 (ZM); Taishun, Anonymous 8108 (ZM), Li et al. 535 (ZM), P. Li LP0804009 (CSH); Tiantai, Faber 279 (K, P); Wencheng, Zhejiang Med. Exped. 3093 (ZM); Yueqing, HZ Herbarium 2436 (HZ), Zhejiang Mus. Exped. 2894 (ZM); Zhenhai, Anonymous s.n. (PE).

JAPAN. Ryukyu: Iriomote Island, M. Furuse 5470 (PE).

Lectotypification:— Sanicula orthacantha var. pumila was described on the basis of two collections, namely P.J. Cavalerie & J.J. Fortunat 1721 and É. Bodinier 2718. We have been able to trace four specimens of these collections at K and P. All these specimens match well the protologue and are the syntypes as Boissieu (1906) did not designate a type for S. orthacantha var. pumila . According to ICN Art. 9.11 ( Turland et al. 2018), here we designate P03226692 of P.J. Cavalerie & J.J. Fortunat 1721 ( Fig. 4A ), which is well preserved and bears the annotation “ S. orthacantha var. pumila ” in Boissieu’s hand, as lectotype.

Sanicula wakayamensis was described based on a single collection, namely Y. Syȏzi s.n. We have been able to trace four specimens of this collection at TAI. All these specimens match well the protologue and are the syntypes as Masamune (1941) did not designate type for S. wakayamensis . According to ICN Art. 9.11 ( Turland et al. 2018), here we designate TAI084140 of Y. Syȏzi s.n. ( Fig. 6A ), which is well preserved with intact flowers, as lectotype.

Notes:— Pimenov (2017) designated K000697286 of the single collection, namely T.L. Bullock s.n. (Herb. Hance 19942) ( Fig. 1 A-part), as the lectotype for S. lamelligera , which matches the original Latin description perfectly and is well-preserved, so here we accept his lectotypification.

Sanicula tuberculata was described on the basis of C. Wilford 906 (GH00589885, K000697277, LE01002000) from Chusan in south Korea ( Maximowicz 1867). It is also distributed in China (Heilongjiang) and southern Japan (Sheh & Philleppe 2005). After carefully checking the type material, we found that S. tuberculata is distinctly different from S. lamelligera by having fertile flowers number 3‒5 per umbellule, and tubercles fruits (vs. fertile flowers number 1 per umbellule, and piculate-lamellate fruits). Thus, we consider the relationship between S. tuberculata and S. lamelligera is fairly distant, and not so closely related as stated by Hance (1878) and Maximowicz (1887).

Hiroe (1979) placed Sanicula petagnioides Hayata (1908: 103) in synonymy with S. lamelligera . After checking the type material of S. petagnioides , we considered that this name should be treated as a synonym of S. orthacantha . We will deal with the identity of S. petagnioides in detail elsewhere.

In the protologue of Sanicula lamelligera and S. penghshuiensis , the authors did not mention the number of vittae in fruit. Hiroe (1979) reported that S. lamelligera has 5 vittae, while Sheh & Phillippe (2005) noted that S. pengshuiensis has 1 vittae under each rib and 2 vittae on commissure. However, Liu (2002) reported that there are 1 vittae under each rib and 2 vittae on commissure in S. lamelligera . Thus, S. lamelligera and S. penghshuiensis are indistinguishable from each other in the character of vittae.

Figures

FIGURE 1. Lectotype (A-upper part) and isolectotype (B-lower part) sheets of Sanicula lamelligera. Inset showing the fruits with calyx teeth and styles.

FIGURE 2. Holotype sheet of Sanicula satsumana (= S. lamelligera).

FIGURE 3. Lectotype and isolectotype sheets of Sanicula yunnanensis (= S. lamelligera).

FIGURE 4. Lectotype (A), isolectotype (B) and remaining syntype (C, D) sheets of Sanicula orthacantha var. pumila (= S. lamelligera).

FIGURE 5. Lectotype (A-lower part), isolectotype (B) and remaining syntype (A-upper part) sheets of Sanicula ichangensis (= S. lamelligera); lectotype (C) and isolectotype (D) sheets of S. orthacantha var. longispina (= S. lamelligera).

FIGURE 6. Lectotype (A) and isolectotype (B‒D) sheets of Sanicula lamelligera var. wakayamensis (= S. lamelligera). A. Inset showing the spiculate-lamellate fruits with calyx teeth.

FIGURE 7. Holotype (A), isotype (B), paratype (C) and isoparatype (D) sheets of Sanicula pengshuiensis (= S. lamelligera). B. Inset a, the fruits with calyx teeth and styles; inset b, folded blade edge. C. Inset a, the shallowly bilobed lateral segments of the basal leaf; inset b, notched margin in the lateral segments of the basal leaf.

FIGURE 9. Specimens of Sanicula lamellligera [China, Guangdong, Shaoguan, Lechang, near to North River, W.T. Tsang 20972 (NAS, PE, US)]. A. Inset showing umbel and the spiculate-lamellate fruits with calyx teeth and styles. B. Inset a showing umbels, inset b showing spiculate-lamellate fruits with calyx teeth and styles. C. Inset showing umbels. D. Inset showing spiculate-lamellate fruits.

FIGURE 10. Sanicula lamelligera in the wild (China, Hubei, Yichang, Xingshan). A. Habitat and Habit. B. Leaves (left: adaxial surface; right: abaxial surface). C. Portion of inflorescence (side view). D. Portion of inflorescence (top view). E. Involucrate bracts. F. Umbellule (side view). G. Staminate flowers (side view). H. Fertile flower with fruit. I. Mericarp.

FIGURE 11. Sanicula lamelligera in the wild (China, Chongqing, Pengshui, Hanjia town, the type locality of S. pengshuiensis).A. Habitat. B. Habit. C. Leaves (left: adaxial surface; right: abaxial surface). D. Portion of inflorescence (side view). E. Portion of inflorescence (top view). F. Involucrate bracts. G. Umbellule (side view). H. Staminate flowers (side view). I. Fertile flower with fruit. J. Mericarp.

FIGURE 12. Sanicula lamelligera in the wild (China, Chongqing, Shizhu, Shiliu town). A. Habitat. B. Habit. C. Leaves (left: adaxial surface; right: abaxial surface). D. Portion of inflorescence (side view). E. Portion of inflorescence (top view). F. Involucrate bracts. G. Umbellule (side view). H. Staminate flowers (side view). I. Fertile flower with fruit. J. Mericarp.

FIGURE 13. Plant individuals in three populations of Sanicula lamelligera, showing the variation in height, the number of basal leaves, the length of petioles, and the inflorescences character.A. China, Chongqing, Pengshui, Hanjia town, the type locality of S. pengshuiensis. B. China, Chongqing, Shizhu, Shiliu town. C. China, Hubei, Yichang (the type locality of S. ichangensis and S. orthacantha var. longispina), Xingshan.

FIGURE 17. One sheet (A) of Sanicula lamelligera and the other sheet (B) of S. caerulescens. Note: the sheet (A) was identified as S. pengshuiensis on the determination slip by Sheh & Liu (1986).

FIGURE 19. Distribution of Sanicula lamelligera (●).