Sanicula serrata H. Wolff

Li, Hui-Min, Zhou, Wei, Wu, Bao-Cheng, Ma, Xu-Dong, Wen, Jun & Song, Chun-Feng, 2025, Taxonomic studies on Sanicula L. (Apiaceae) from China (V): Revision of the Sanicula serrata H. Wolff complex: morphological delimitation from S. potaninii Bobrov, synonymization of allied taxa, and lectotypifications, PhytoKeys 265, pp. 71-104 : 71-104

1. Sanicula serrata H. Wolff in Engler, Pflanzenr. 61: 56 (1913).

Figs 1 , 3 , 5 , 6 , 7 , 8 , 12 , 14 , 17 锯叶变豆菜

= Sanicula tienmuensis R. H. Shan & Constance , Univ. Calif. Publ. Bot. 25: 23 (1951), syn. nov. Type: China. Zhejiang, Hangzhou, Lin’an, West Tianmushan , 30 April 1931, West Lake Museum 67 ( holotype: NAS 00082802 !; isotypes: ZMNH 0057084 !, ZMNH 0057085 - right part!) .

= Sanicula elongata K. T. Fu , Fl. Reipubl. Popularis Sin. 55 (1): 297 (1979), syn. nov. Type: CHINA. Shaanxi, Mei County, Laojunling , 17 June 1952, P. C. Kuo 1424 ( lectotype: IBK 00159684 ! designated here; isolectotypes: IBK 00159685 !, IBK 00159686 !, KUN 0465506 !, PE 01933863 !) .

= Sanicula tienmuensis var. pauciflora R. H. Shan & F. T. Pu , Acta Phytotax. Sin. 27 (1): 66 (1989), syn. nov.

≡ Sanicula pauciflora (R. H. Shan & F. T. Pu) B. N. Song & X. J. He , Frontiers Pl. Sci. (Online journal) 15–1351023: 12 (2024).

Type: CHINA. Sichuan, Luding County, Desui Town, Mount Tianchi , under forests or by streams, alt. 2,200 m, 1 May 1984, Y. L. Cao 115 ( lectotype: CDBI 0172308 ! designated here; isolectotype: CDBI 0172309 !) .

= Sanicula langaoensis B. N. Song, T. Ren & X. J. He , Frontiers Pl. Sci. (Online journal) 15–1351023: 13 (2024), syn. nov. Type: China. Shaanxi, Lan’gao County, in stream banks inmixed forests, 32°13'47.88"N, 108°53'45.18"E, alt. 1,496 m, 12 April 2023, B. N. Song, T. Ren & X. J. He SBN 2023041201 ( holotype: SZ!) .

Type.

China. • Hubei, Badong (= Patung), April 1900, E. H. Wilson 156 a ( lectotype: K 001325368 !, designated by Pimenov (2017); isolectotype: NY 00406258 - right lower part!, US 00126980 !). Fig. 1 .

Description. Perennial. Rhizome short and stout, roots fascicled, fleshy, with fibrous. Stem 1–4, slender and erect, several branched above the middle, 7–47 cm tall. Basal leaves 2–8, long petiolate; petioles 1.5–23 cm long; blade glabrous adaxially and abaxially, 1–7 cm long, 1.5–16.5 cm wide, rounded-cordate to cordate-pentagonal, tripartite to the base or trifoliate, the median segment cuneate-obovate to rhombic-cuneate, distinctly shallowly trilobed, the lateral segments oblique, usually bilobed below the middle or nearly to the base, the margins irregularly crenulate-serrate with sharply serrate teeth. Cauline leaves few, conspicuous, ternate, upper stem leaves subsessile or sessile, tripartite, resembling involucrate bract, (0.5 –) 1.5–3 cm long, (0.3 –) 1.3–2.5 (– 4.5) cm wide. Inflorescence 1–3 - branched; involucrate bract usually 2, trisect, 1–6 mm long; rays of the umbels 1–6, 2–13 mm long; involucellate bracteoles 4–6, oblong ovate, 0.2–0.8 mm long. Umbellules 1–8 - flowered, staminate flowers 0–7 per umbellule, pedicels 1.9–3.2 mm long, petals white, fertile flowers 1 per umbellule, sessile; calyx teeth oblong ovate, 0.5–0.9 mm long; styles 1.9–3.0 mm long, 2–5 times longer than the calyx teeth. Mericarps oblong-ovoid to ovoid, 1.8–2.6 mm long, 1.3–3 mm broad, tuberculate spiculate and occasionally with prickly spines on the top during later ripening. Vittae obsure.

Distribution.

Sanicula serrata is widely distributed in China ( Anhui, Gansu, Hubei, Shaanxi, Sichuan, Yunnan and Zhejiang) (Fig. 18 ).

Habitat.

This species grows on mountain slopes under forest or along ravine streams near rock or along the road under forest at elevations of 500–3,200 m above sea level.

Phenology.

Flowering from April to May, fruiting from April to June.

Etymology.

The epithet serrata is derived from Latin and refers to the serrated margins of the leaves.

Additional specimens examined.

Anhui Province, • Huoshan [Hoshan] County, Cangping Village , 31°10'29.9"N, 116°11'6.3"E, alt. 796 m, 10 April 2021, H. M. Li, Y. S. Zhang, Y. Xu 1110 ( NAS) ; • Huoshan County, Cangpingdagou , alt. 950 m, 27 March 1982, M. B. Deng 81269 ( NAS) ; • Huoshan County, Cangpingdagou, alt. 1,000 m, 9 June 1980, M. B. Deng, K. Yao 80870 ( NAS) ; • Jinzhai [Chin Chai] County, Baima Zhai Forest Farm , alt. 650 m, 19 April 1993, K. Y. Lang, et al. 930047 ( PE) ; • Jinzhai County, Baima Zhai Forest Farm , alt. 900 m, 19 May 1984, K. Yao 9027 ( NAS) ; • Jinzhai County, Baima Zhai Forest Farm , alt. 1,000 m, 17 May 1984, M. B. Deng 81743 ( NAS) ; • Jinzhai County, Baima Zhai Forest Farm , alt. 1,100 m, 14 May 1984, Anonymous 81722 ( NAS) ; • Jinzhai County, Baima Zhai Forest Farm , alt. 1,300 m, 14 May 1984, M. B. Deng 81721 ( NAS) ; • Jinzhai County, Baimazhai , alt. 1,050 m, 21 April 1985, X. S. Shen 218 ( PE) . Gansu Province, • Hui County, alt. 1,800 m, 30 June 1956, P. C. Kuo 3167 ( WUK) ; • Kang County, Changba Township , alt. 1,600 m, 23 April 1963, Y. Q. He, C. L. Tang 31 ( WUK) ; • Kang County, alt. 1,500 m, 3 May 1963, Y. Q. He, C. L. Tang 249 ( WUK) ; • Long’an City, Wudu District, Luotang Town, Yuezhao Township , alt. 2,400 m, 24 June 1959, Z. Y. Zhang 5687 ( LBG, WUK) ; • Min County, alt. 3,200 m, 16 June 1951, T. P. Wang 15279 ( KUN, NAS, WUK) ; • Tianshui County, Dangchuan , alt. 2,200 m, 22 June 1964, K. T. Fu 15559 ( WUK) ; • Tianshui County, 1,200 m, 16 April 1959, S. B. He 1048 ( WUK) ; • Wen County, Bikou Town , alt. 700 m, 24 March 1972, T. P. Wang 20442 ( WUK) . Hubei Province, • Badong County, Shennongxi , Y. S. Zhang LHM 1111 ( NAS) ; • Nanzhang County, Xueping Town , 29°48'07.4"N, 109°15'25.1"E, alt. 742 m, 16 May 2025, F. Tan LHM 1701 ( NAS) ; • Wufeng County, Xiaolong Village , 30°16'44.4"N, 110°41'20.4"E, alt. 1,657 m, 18 May 2025, F. Tan LHM 1700 ( NAS) ; • Baoji City, Weibin District, Jifengshan , alt. 1,650 m, 28 May 1977, Z. X. Hu, Y. H. Guo 313 ( WUK) ; • Baoji City, Mount Zhongyan , 34°15'10.9"N, 107°06'56"E, alt. 1,778 m, 3 June 2022, H. M. Li, C. F. Song 1207 ( NAS) ; • Chang’an County, 1956, Huanghe Exped. 360 ( KUN) ; • Feng County, Miaowangshan , alt. 2,400 m, 12 June 1977, K. T. Fu 17337 ( WUK) ; • Foping County, Donghe Village , alt. 2,100 m, 11 May 2008, S. F. Li, B. Li, Y. Zhang, et al. 10192 ( XBGH) ; • Foping County, Gaojiaba , alt. 1,800 m, 20 July 1952, K. T. Fu 5115 ( KUN, IBK, WUK) ; • Liuba County, Miaotaizi Forest Farm , 22 April 1982, Z. Y. Zhang 18454 ( WUK) ; • Liuba County, Sangyuanba Township , 33°42.720'N, 107°9.635'E, alt. 1,072–1,470 m, 28 May 2010, S. F. Li, B. Li, Y. Zhang, et al. 13530 ( XBGH) ; • Liuba County, 33°42'25.8"N, 107°10'12.4"E, alt. 1,366 m, 4 June 2022, H. M. Li, C. F. Song 1219 ( NAS) ; • Lueyang County, Miaogou , 33°17'11.4"N, 105°55'17.8"E, alt. 1,295 m, 5 June 2022, H. M. Li, C. F. Song 1224 ( NAS) ; • Lueyang County, Zengjiagou , 33°16'59.6"N, 105°55'19.4"E, alt. 1,380 m, 5 June 2022, H. M. Li, C. F. Song 1223 ( NAS) ; • Lueyang County, Zhangjiaba , alt. 1,300 m, 4 May 1957, The Fifth Forest Survey Team of the Ministry of Forestry 9 ( WUK) ; • Nanzhen County, Longdonggou , 32°44'56.8"N, 106°56'45.0"E, alt. 869 m, 6 June 2022, H. M. Li, C. F. Song 1227 ( NAS) ; • Nanzhen County, On the roadside from Xihe to Xiaoba , X. X. Hou 226 ( NAS) ; • Nanzhen County, Xiaoba , alt. 1,973 m, 20 April 1973, X. X. Hou 475 ( WUK) ; • Ningshan County, Guankou , alt. 1,800 m, 3 June 1959, J. Q. Xing 4388 ( NAS, WUK) ; • Ningshan County, Guankou Town , alt. 1,690 m, 30 May 1959, J. Q. Xing 4049 ( NAS, SZ, WUK) ; • Ningshan County, Guankou Town , alt. 1,925 m, 4 June 1959, J. Q. Xing 5661 ( SZ, WUK) ; • Ningshan County, Xunyangba , alt. 1,439 m, 13 May 2013, X. H. Tian, L. H. Wu, TianXH 823 ( KUN) ; • Ningshan County, Xunyangba, 24 April 1993, X. H. Tian, L. Zhang T 934021 ( PE) ; • Ningshan County, Zhifanggou , 33°29'48.9"N, 108°31'32.3"E, alt. 1,679 m, 9 June 2022, H. M. Li, C. F. Song 1239 ( NAS) ; • Pingli County, Baxian Town , 32°1.720'N, 108°19.690'E, alt. 1,900 m, 8 April 2004, Y. S. Chen, Z. H. Wu, B. Li et al. 275 ( WUK) ; • Zhen’an County, Lizha Forest Farm , alt. 1,830 m, 30 May 1973, X. X. Hou, Y. H. Guo 650 ( WUK) ; • Zhen’an County, Muwang Forest Farm , 33°23.712'N, 108°37.017'E, alt. 2,037 m, 9 June 2008, S. F. Li, B. Li, Y. Zhang, et al. 10475 ( XBGH) ; • Zhen’an County, Muwang Forest Farm , 33°23'43.4"N, 108°37'01.5"E, alt. 2,015 m, 8 June 2022, H. M. Li, C. F. Song 1232 ( NAS) ; • Zhenping County, Falong Village , 31°58'35.3"N, 109°23'02.4"E, alt. 1,637 m, 7 June 2022, H. M. Li, C. F. Song 1231 ( NAS) ; • Zhenping County, Shangzhu Township , alt. 1,710 m, 3 May 1989, G. Y. Xu 4641 ( WUK) ; • Zhenping County, 1970, Zhenping Pharma. Exped. 111 ( WUK) ; • Zhouzhi County, Xinkou , alt. 2,000 m, 13 June 1952, P. C. Kuo 1313 ( IBK, KUN, WUK) ; • Zhouzhi County, alt. 1,220 m, 25 April 1977, Z. X. Hu, Y. H. Guo 89 ( WUK) ; • Pingli County, alt. 1,820 m, 27 May 1959, P. Y. Li 2566 ( KUN, WUK) . Sichuan Province, • Hongya [Hung ya] County, Hongya Forest Farm, Mahuanggou , alt. 1,950 m, 27 June 1994, W. K. Bao, et al. 2485 ( CDBI) ; • Luding County, Desui Town, Tianchishan , 29°35'20.4"N, 102°11'16.5"E, alt. 2,355 m, 18 May 2021, H. M. Li, Y. S. Zhang 1167 ( NAS) ; • Luding County, Moxi Town , alt. 2,350 m, 8 April 1981, Sichuan Plant Exped. 25023 ( CDBI) ; • Luding County, Moxi Town , alt. 2,900 m, 8 June 1980, Q. Q. Wang, Z. A. Liu 22183 ( CDBI) ; • Nanjiang County, Guangming Township, Guangwushan , alt. 1,600 m, 5 June 1956, Sichuan-Da Econ. Plant Exped. 2675 ( KUN) ; • Nanjiang County, alt. 1,400–1,500 m, 5 June 1959, B. L. Chen 2558 ( KUN) ; • Shimian County, Caoke Township , alt. 2,300 m, 1978, Shimian Exped. 78–1149 ( SM) ; • Shimian County, alt. 1,850 m, 11 April 1955, C. J. Xie 39763 ( PE) ; • Shimian County, alt. 1,850 m, 11 April 1955, C. J. Xie 39763 ( PE, WUK) ; • Tianquan County, Erlangshan , alt. 1,550 m, 20 April 1956, D. P. He 43222 ( SZ) ; • Sichuan, Tianquan County, Erlangshan, alt. 1,700–2,050 m, 26 April 1986, T. Naito et al. 150 ( PE) ; • Tianquan County, Lianghe Township , alt. 1,600 m, 22 April 1956, D. P. He 42968 ( SZ) ; • Tianquan County, Xingou , alt. 1,700 m, 3 June 1959, Sichuan-Ya Econ. Plant Exped. 551 ( SM) ; • Tianquan County, Xingou Village , alt. 1,720 m, 1 April 1980, Y. B. Yang 21603 ( CDBI) ; • Tianquan County, alt. 1,300 m, 23 April 1956, D. P. He 43013 ( SZ) ; • Tianquan County, alt. 1,550 m, 21 April 1956, D. P. He 42940 ( SZ) ; • Tianquan County, alt. 1,800 m, 31 March 1953, X. L. Jiang 33663 ( PE, SZ) ; • Tianquan County, alt. 2,800 m, 6 May 1978, Tianquan Exped. 78–132 ( SM) ; • Tianquan County, 1936, K. L. Chiu 2300 ( NAS, PE) ; • Tianquan County, K. L. Chiu 2398 ( PE) ; • Wangcang County, Micangshan Nature Reserve Shiziba Conservation Station , 32°39'42.3"N, 106°32'48.2"E, alt. 1,651 m, 28 June 2011, Bashan Exped. 4815 ( PE) ; • Zhaojue County, Guli Town , alt. 1,280 m, 22 April 1979, Zhaojue Exped. 96 ( SM) ; • Sichuan, Zhaojue County, Guli Town , alt. 1,720 m, 24 April 1979, Zhaojue Exped. 133 ( SM) . Yunnan Province, • Dali City, Fengyandong , June 1941, H. C. Wang 804 ( PE) ; • Gongshan County, 28°24'06.7"N, 104°14'37.3"E, alt. 2,049 m, 1 April 2023, H. M. Li, J. W. Zhu, B. C. Wu 130 ( NAS) ; • Weixi County, Baimaluo , alt. 2,500 m, 2 June 1940, K. M. Feng 4366 ( KUN, PE) ; • Weixi County, Yezhi Town , alt. 2,700 m, 8 May 1982, Qinghai-Tibet Plateau Exped. 6330 ( KUN, PE) ; • Weixi County, alt. 2,600 m, 29 April 1940, K. M. Feng 3543 ( KUN, PE) ; • Weixi County, alt. 3,500 m, June 1935, C. W. Wang 63892 ( PE) . Zhejiang Province, • Anji County, Longwangshan , 30°24'29.44"N, 119°26'59.21"E, alt. 933 m, 16 April 2021, H. M. Li, L. Zhao 1114 ( NAS); • Anji County, Longwangshan, alt. 500 m, 14 March 1997, L. P. Yu, M. B. Deng 97025 ( PE); • Anji County, Longwangshan, alt. 600 m, 16 May 1995, M. B. Deng 90144 ( PE) ; • Anji County, Longwangshan, 3 May 1988, Anonymous 1718 ( ZM) ; • Anji County, Longwangshan, Qianrenqiao , alt. 1,100 m, 3 May 1987, Xu 281 ( ZM) ; • Anji County, Longwangshan, Xianrenqiao , alt. 550 m, 7 May 1997, Y. M. Fang, M. B. Deng 975152 ( PE) ; • Anji County, Xiaofeng Town , 1958, H. Y. Ho 24193 ( HZ, NAS, PE) ; • Hangzhou City, Lin’an District, West Tianmushan , 30°20'13"N, 119°26'2"E, alt. 777 m, 17 May 2021, H. M. Li, L. Zhao 1116 ( NAS) ; • Hangzhou City, Lin’an District, West Tianmushan , 30°20'13"N, 119°26'2"E, alt. 783 m, 20 April 2025, X. D. Ma 360 ( NAS) ; • Hangzhou City, Lin’an District, Tianmushan , alt. 580 m, 25 April 1957, H. Y. Ho 21100 ( HZ, NAS) ; • Hangzhou City, Lin’an District, West Tianmushan , alt. 800 m, 25 May 1957, Yuan et al. 4114 ( HHBG, NAS, WUK) ; • Hangzhou City, Lin’an District, West Tianmushan , 23 April 1936, H. Migo s. n. ( NAS) ; • Hangzhou City, Lin’an District, Anonymous 94 ( HZ) ; • Pan’an County, Dapanshan , 28°58'17.91"N, 120°31'17.05"E, alt. 1,104 m, 18 April 2021, H. M. Li, L. Zhao 1117 ( NAS) ; • Tiantai County, Huadingshan , 21 April 1955, T. Y. Cheo, J. S. Yue 978 ( NAS) .

Notes.

1) In the protologue of Sanicula serrata, Wolff (1913) indicated that the type collection, E. H. Wilson 156 a, was deposited at B. This specimen was most likely destroyed during World War II. Five duplicates of this collection are currently preserved at E, HBG, K, NY, and US, respectively, and are all in good condition. However, upon detailed examination of both herbarium material and living plants, we observed that the characteristics of the cauline leaves and fruits in specimens E 00000706, HBG 510909 , and the upper left portion of NY 00406258 correspond to S. orthacantha var. orthacantha ( Li et al. 2022) , rather than to S. serrata . Only specimens K 001325368, the lower right portion of NY 00406258, and US 00126980 conform to the original description of S. serrata . Pimenov (2017) designated the specimen at K (K 001325368) as the lectotype, and the specimens at E, HBG, NY, and US as isolectotypes. In accordance with ICN Article 9.3 ( Turland et al. 2025), we accept Pimenov’s (2017) lectotypification and confirm that the lower right portion of NY 00406258 and US 00126980 represent authentic duplicates and are hereby recognized as isolectotypes.

2) Hiroe (1979) classified Sanicula elata var. acaulis as a synonym of S. serrata . Upon reviewing the protologue of S. elata var. acaulis , we note that it provides only a brief description, plant 40–50 cm tall, with almost no stem; umbel dense and globose, with very short, angular, winged raylets; calyx lobes triangular, with no type material cited ( Franchet 1883). Although Liu (1997) endorsed Hiroe’s treatment, we have not been able to locate any associated specimens or additional corroborating literature. A more detailed assessment of S. elata var. acaulis will be presented elsewhere.

3) Sanicula elongata was described based on two samplings: P. C. Kuo 1424 and K. T. Fu 15559. Fu (in Liou 1979) cited the former collection as the type but did not designate a specific specimen as the holotype. We have traced four specimens referable to the former collections, preserved at KUN, IBK (Fig. 5 A – C ). All specimens match the original description and are considered syntypes, as Fu (in Liou 1979) did not designate a single type specimen. Although Pimenov (2017) indicated that the specimen P. C. Kuo 1424 at WUK was the holotype, he did so without examining the specimen, and we were unable to locate a duplicate sheet at WUK. Based on our investigation, the specimen at WUK likely corresponds to K. T. Fu 15559 (Fig. 5 D ). In accordance with ICN Article 9.11 ( Turland et al. 2025), we designate the specimen P. C. Kuo 1424 kept at IBK ( IBK 00159684 ; Fig. 5 A ) – which is complete and well preserved – as the lectotype.

4) Sanicula tienmuensis var. pauciflora was described based on a single collection, Y. L. Cao 115, which was incorrectly attributed to “ Y. J. Li ” in the protologue. We have identified two specimens of this collection at CDBI, both of which are well-preserved and consistent with the protologue. Since Shan and Pu (1989) did not designate a single type specimen, both specimens qualify as syntypes according to the discussion by McNeill (2014). In accordance with ICN Article 9.17 ( Turland et al. 2025), Pimenov’s (2017) reference to a CDBI sheet as the “ holotype ” is incorrect and should be amended to “ lectotype ” ( McNeill 2014). Consequently, we designate sheet CDBI 0172308 (Fig. 6 A ) as the lectotype and CDBI 0172309 (Fig. 6 B ) as the isolectotype.

Figures

Figure 1. Lectotype (upper left; A) and isolectotype (B) sheets of Sanicula serrata var. serrata. Arrows indicate fruits with squamose spines.

Figure 3. Holotype (A), isotype (B), paratype (C), and a duplicate of the paratype (D) sheets of Sanicula tienmuensis var. tienmuensis. Arrows indicate fruits with squamose spines.

Figure 5. Lectotype (A), isolectotypes (B, C), and paratype (D) sheets of Sanicula elongata. Arrows indicate fruits covered with densely squamose spines, sometimes ending in minute prickles.

Figure 6. Lectotype (A) and isolectotype (B) sheets of Sanicula serrata var. pauciflora.

Figure 7. Holotype sheet of Sanicula langaoensis.

Figure 8. Sanicula serrata var. serrata in the wild (China, Hubei, Badong County, Shennongxi, the type locality of S. serrata var. serrata). A. Habitat; B. Habit; C. Leaf (left: adaxial surface; right: abaxial surface); D. Portion of the inflorescence (top view); E. Portion of the inflorescence (side view), arrow indicating the involucral bract; F. Umbellule (side view), arrow indicating the involucellate bracteoles; G. Staminate flower (side view); H. Fertile flower with immature fruit; I. Mericarps, arrow indicating the calyx teeth. Photographed by Hui-Min Li.

Figure 12. Sanicula serrata var. serrata in the wild (China, Zhejiang, Hangzhou, Lin’an, West Tianmushan, the type locality of S. tienmuensis var. tienmuensis). A. Habitat and habit; B. Rhizome; C. Leaf (left: adaxial surface; right: abaxial surface); D Portion of the inflorescence (side view); E. Umbellule (side view), early fruiting; F. Umbellule (side view), late fruiting, arrow (a) indicating the involucellate bracteoles; arrow (b) indicating the calyx teeth; G. Staminate flower (side view); H. Mericarps. Photographed by Hui-Min Li.

Figure 14. Sanicula serrata var. serrata in the wild (Shaanxi, Baoji City, Mount Zhongyan, near the type locality of S. elongata). A. Habitat; B. Habit; C. Rhizome; D. Leaf (left: adaxial surface; right: abaxial surface); E. Portion of the inflorescence (side view); F. Portion of the inflorescence (side view), arrow indicating the involucral bract; G. Umbellule (side view), late fruiting, arrow (a) indicating the involucellate bracteoles; arrow (b) indicating the calyx teeth; H. Mericarps. Photographed by Hui-Min Li.

Figure 17. Sanicula serrata var. serrata in the wild (Sichuan, Luding, Desui Town, Mount Tianchi, the type locality of S. serrata var. pauciflora). A. Habitat; B. Habit; C. Leaf (left: adaxial surface; right: abaxial surface); D. Portion of the inflorescence (side view), arrow indicating the involucellate bracteoles; E. Staminate flower (side view); F. Mericarps, arrow indicating the calyx teeth. Photographed by Hui-Min Li.

Figure 18. Distribution of Sanicula serrata var. serrata (red circles) and S. potaninii (black circles).