Thalictrum fortunei Moore (1878: 130)

Zeng, You-Pai, Yuan, Qiong & Yang, Qin-Er, 2022, Clarification of morphology and distribution of Thalictrum fortunei (Ranunculaceae) from China, with reduction of T. fortunei var. bulbiliferum to its synonymy, Phytotaxa 561 (2), pp. 121-137 : 123-133

Thalictrum fortunei Moore (1878: 130) . Figs. 1‒8 .

Type:— CHINA. Zhejiang: Ningpo (= Ningbo), C.W. Everard s.n. (lectotype, designated by Yuan et al. (2018), K000694061 (the lower specimen!); isolectotype K000694060!). Remaining syntypes:— CHINA. Zhejiang: without precise locality, R. Fortune 28 (BM000559555!, K000959812!, K000959813!, P00147391!, P00147392!, P00147393!, P00852558!). Figs. 1 , 2 .

= T. argyi Léveillé & Vaniot in Léveillé (1906: 504).

Type:— CHINA. Kiang-Sou (= Jiangsu): Gong-Chan, C. d’Argy s.n. (holotype E00386260!; isotypes DAO000001581!, GH00038598!) .

= T. fortunei var. bulbiliferum Chen & Tian in Chen et al. (2005: 281), syn. nov.

Type:— CHINA. Jiangsu: Jurong, Baohua Shan , 23 May 2004, B. Chen 523001 (holotype N, not seen; isotype PE01458769!). Fig. 3 .

Description:—Perennial herbs. Roots fibrous, distally thicker, up to 3 mm in diameter. Stem 20‒60 cm tall, usually basally branched, distally glandular-pubescent, with bulbils borne in leaf axils at late flowering and fruiting stage. Leaves 2‒3-ternate; blade triangular, 5‒15 cm long and broad; leaflets ovate, broadly ovate, obovate, or suborbicular, 0.5‒ 4 cm long, 0.6‒ 5 cm broad, papery, abaxially glandular-pubescent when tender, both sides glabrous when mature, green on adaxial side, pale green on abaxial side, base rounded or subcordate, apex obtuse, 3-lobate; lobes crenatedentate; veins slightly submerged adaxially, raised abaxially; petiole slender, 1.5‒ 6 cm long; stipule membranous, ovate-orbicular, margin torn. Inflorescence a compound cyme, few-flowered, corymbiform. Pedicels 0.5‒ 2 cm long, glandular-pubescent. Flowers bisexual, erect. Sepals 4, elliptic or obovate, caducous, 3‒4.5 mm long, 2‒2.5 mm broad, white or tinged with purplish, glabrous or abaxially very sparsely glandular-pubescent. Stamens 40‒60, 5‒7 mm long; filaments narrowly clavate, 4‒6 mm long, whitish or purplish; anthers oblong, 1‒1.2 mm long, ca. 0.6 mm broad, apex obtuse, whitish. Carpels 3‒6, sessile, 1.2‒ 2 mm long; ovaries fusiform, shallowly ribbed, ca. 1 mm long, subglabrous; styles 0.6‒ 1 mm long, slightly recurved at apex; stigmas conspicuous, linear, ca. 0.5 mm long. Achenes sessile, 3.5‒ 5 mm long, sparsely glandular-pubescent; body fusiform, profoundly ribbed; styles persistent, recurved at apex.

Phenology:—Flowering from March to May; fruiting from April to June.

Distribution and habitat:— Thalictrum fortunei is currently known to occur in southern Anhui, eastern Hubei, northern Jiangxi, southern Jiangsu and Zhejiang provinces, China ( Fig. 9 ). The previous records of its distribution in Fujian province ( Lin & Zhao 1985, Wang 2000) and in Henan province ( Anonymous 1981) should have resulted from misidentifications, and thus are doubtful (see below). This species grows in forests, along streams, on damp rocks in ravines or on slopes at elevations of 10‒600 m above sea level.

Additional specimens examined:— CHINA. Anhui: Huaining, Anonymous 3060 (NAS, barcode unavailable); Nanling, L.H. Li 340223120422008 (PE01981467), L.H. Li 340223120422098 (PE01981468); Shitai, M.P. Teng et al. 116 (NAS00187677, P00147360); Tongling, Z.H. Zhang s.n. (PE00471333); Wuhu, J.L. Bullock s.n. (BM, barcode unavailable); Xuancheng, F. Courtois 25026 (NAS00187678). Hubei: Yangxin, M.H. Ma 507 (PE00471342). Jiangsu: Jurong, B. Chen 044135 (PE01900654), B. Chen 045245 (PE01900655), B. Chen 403008 (PE01458770), S. Chen & M.P. Teng 139 (NAS00110622), S. Chen & M.P. Teng 178 (NAS00110623), J.J. Gong 560 (NAS00110624), K.C. Kuan 150 (PE00470855, PE00471337), M.P. Teng 2919 (NAS00110626), C.L. Tso 139 (IBSC0090698, NAS00110618, NAS00110630, PE00471338), Z.Y. Wang & B.C. Wu SCSB-JS0078 (KUN0579864, KUN1232993); Precise locality unknown, R.H. Shan 201 (NAS00110628); Suzhou, H.T. Chang 475 (PE00471339), C.C. Chen 935 (NAS00110617, NAS00110627), P.P. Ling & S.S. Sun 68 (IBSC0090287, KUN0690810, NAS00110619, NAS00110629, PE00471334, SZ00092188), C.T. Ting & Y.C. Wang 219 (NAS00110609, NAS00110633); Wuxi, W.X. Wu 5041 (NAS00110608); Yixing, Anonymous 8 (NAS00110615), Anonymous 6856 (NAS00110621), Anonymous 20169 (NAS00110625), C.Y. Luh 506 (NAS00110620, NAS00110631), W.X. Wu 8557 (NAS00110604). Jiangxi: Jiujiang, C.M. Tan 11130A (JJF00012911); Lushan, K.K. Tsoong 4289 (PE00470857), K.K. Tsoong 4290 (PE00470856, PE00470858, PEY0003108), M.K. Wang 146 (LBG00013428, SZ00092187); Xiushui, M.K. Wang 236 (IBSC0090288). Zhejiang: Daishan, Anonymous465 (ZM NH0006317), X.Y.Ye CTD201305031 (CSH0032037), X.Y.Ye DSD13014 (CSH0033892); Jiande, Anonymous 4277 (ZM NH0006324), L. Hong s.n. (HHBG HZ008565); Jinhua, V. King 40 (NAS00187657, PE00470579); Ningbo, E. Faber 1550 (K, barcode unavailable), W. Hancock 24 (K, barcode unavailable), Y.Y. Ho 849 (NAS00187146, NAS00187147), K.K. Tsoong s.n. (PE00470572), K.K. Tsoong 942 (PE00470574); Pan’an, F.G. Zhang DPS-841 (ZM NH0006322); Xianju, R.C. Ching 1673 (BM, barcode unavailable, E, barcode unavailable, K, barcode unavailable, P00147359); Xinchang, X.Y. Ye XC201304077 (CSH0032140); Zhoushan, Y.K. Bi et al. BYK3868 (CSH0034447); Zhuji, S. Chen 66 (NAS00187145, NAS00187148). Without locality, Anonymous 239 (PE00470859), Anonymous 1079 (PE00471122, PE00471123).

Notes:—Our molecular systematic studies based on nrDNA ETS and ITS and cpDNA trn V- ndh C sequence data of 115 species, with 84 from China ( Zeng 2022), indicate that Thalictrum fortunei is most closely related to T. ramosum Boivin (1945: 115), a species widely distributed in China (Chongqing, Guangxi, Guizhou, Hubei, Hunan, Sichuan, Yunnan). Morphologically these two species are somewhat similar in habit and share an array of other characters, including caducous sepals, narrowly clavate filaments, apically recurved styles and sessile, fusiform achenes.

Because of their superficial morphological similarity and somewhat overlapping geographical distribution, Thalictrum fortunei has been sometimes confused with T. fargesii . Some specimens of the latter species from Anhui, Henan, Jiangxi and Zhejiang provinces were misidentified as T. fortunei ( Fig. 10 ). As shown in Figures 5‒7 , 11 , 12 , T. fortunei is similar to T. fargesii in having 2‒3-ternate leaves, usually broadly ovate leaflets, few-flowered inflorescence, clavate filaments of stamens, and fusiform achenes, but differs by the presence (vs. absence) of bulbils in leaf axils at late flowering and fruiting stage, caducous (vs. more or less persistent) sepals ( Figs. 5L , 6K , 7I , 11F , 12G ), more numerous (40‒60 vs. 15‒30) stamens ( Figs. 5L , 6K , 7I , 11F , 12G ), and apically slightly recurved (vs. straight) styles ( Figs. 5O , 6N , 7K , 11G , 12J ) (also see Table 1 ). Thalictrum fortunei has been recorded to occur in southern Anhui, southern Fujian, southern Henan, eastern Hubei, southern Jiangsu, northern Jiangxi and Zhejiang provinces, China ( Anonymous 1972, 1976, 1981, 1982, Wang & Wang 1979, Lin & Zhao 1985, Wang 1987, 2000, 2004, 2018, Lin 1992, Fu 2001, Fu & Zhu 2001, Deng & Ye 2013, Wang & Liu 2016, Xie 2016, Yuan et al. 2018, Zhang 2021), and T. fargesii has been recorded to occur in Chongqing, Gansu, Guizhou, Henan, Hubei, Shaanxi and Sichuan ( Anonymous 1976, 1981, 2014, Wang & Wang 1974, 1979, Wang 1980, 2018, Li 1986, Xie 2016), but actually the former occurs in southern Anhui, eastern Hubei, southern Jiangsu, northern Jiangxi and Zhejiang, very likely not in Fujian and Henan, whereas the latter occurs in Anhui, Chongqing, Gansu, Guizhou, Henan, Hubei, Hunan, Jiangxi, Shaanxi, Sichuan and Zhejiang. We have not as yet seen any specimens of T. fortunei from Fujian and Henan. Some specimens from Anhui, Jiangxi and Zhejiang have been correctly identified as T. fargesii or T. pallidum Franchet (1885: 187) (a synonym of T. fargesii ) ( Fig. 13 ). Li et al. (2016) correctly cited some specimens from Jiangxi and Zhejiang under T. fargesii .

Thalictrum argyi had long been reduced to the synonymy of T. javanicum Blume (1825: 2) until it was placed in synonymy with T. fortunei by Yuan et al. (2018). We have checked the type material of T. argyi and here we accept that decision. Judging from the fact that T. argyi was described on the basis of material from Jiangsu, the treatment by Yuan et al. (2018) is also more justifiable. According to Zeng et al. (2021), T. javanicum is distributed only in Indonesia (Java), not in China.

Figures

FIGURE 1. Lectotype (A-in part, the lower specimen), isolectotype (B-in part, the lower specimen) and syntype (B-in part, the upper two specimens) sheets of Thalictrum fortunei.

FIGURE 2. Syntype sheets of Thalictrum fortunei.

FIGURE 3. Isotype sheet of Thalictrum fortunei var. bulbiliferum (= T. fortunei) (insets a, b: bulbils).

FIGURE 4. Selected specimens of Thalictrum fortunei from Baohua Shan in Jurong county, Jiangsu province, China, the exact type locality of T. fortunei var. bulbiliferum (= T. fortunei). A. B. Chen 044135 (PE). B. B. Chen 045245 (PE); insets a, b: bulbils.

FIGURE 5. Thalictrum fortunei var. bulbiliferum (= T. fortunei) in the wild (Baohua Shan in Jurong county, Jiangsu province, China, the exact type locality). A. Habitat. B, C. Habit. D. Roots. E. Portion of stem. F. Bulbils. G. Leaf (left: adaxial side; right: abaxial side). H. Leaflet (above: adaxial side; below: abaxial side). I. Portion of abaxial side of leaflet. J. Portion of petiole. K. Portion of pedicel. L. Flower with sepals having fallen off. M. Sepal (left: adaxial side; right: abaxial side). N. Stamens. O. Carpels. P. Aggregate fruit (immature). Q. Achenes (immature). Photographed by Y.P. Zeng.

FIGURE 6. Thalictrum fortunei in the wild (Ningbo in Zhejiang province, China, the type locality). A. Habitat. B. Habit. C. Roots. D. Portion of stem. E. Bulbils. F. Leaf (left: adaxial side; right: abaxial side). G. Leaflet (above: adaxial side; below: abaxial side). H. Portion of abaxial side of leaflet. I. Portion of petiole. J. Portion of pedicel. K. Flower with sepals having fallen off. L. Sepal (left: adaxial side; right: abaxial side). M. Stamens. N. Carpels. O. Aggregate fruit (immature). P. Achenes (immature). Photographed by Y.P. Zeng.

FIGURE 7. Thalictrum fortunei moved from Tongling in Anhui province to the Nanjing Botanical Garden Mem. Sun Yat-Sen in Nanjing, Jiangsu province, China. A. Habit. B. Portion of stem. C‒E. Bulbils. F. Leaf (adaxial side). G. Leaf (abaxial side). H. Portion of pedicel. I. Flower with sepals having fallen off. J. Stamens. K. Carpels. L. Aggregate fruit (immature). M. Achenes (immature). Photographed by Y.P. Zeng.

FIGURE 8. Selected specimens of Thalictrum fortunei with bulbils borne in leaf axils (arrowed).A. China, Jiangsu, Yixing, W.X. Wu 8557 (NAS) (inset: bulbils). B. China, Jiangxi, Lushan, K.K. Tsoong 4289 (PE) (inset: bulbils). C. China, Jiangxi, Lushan, K.K. Tsoong 4290 (PE) (inset: bulbils). D. Duplicate (PE) (inset: bulbils).

FIGURE 9. Distribution of Thalictrum fortunei (●). Black arrow indicates the type locality of T. fortunei var. bulbiliferum (= T. fortunei), i.e. Jurong county in Jiangsu province, China; red arrow indicates the type locality of T. fortunei, i.e. Ningbo in Zhejiang province, China.

FIGURE 10. Selected specimens of Thalictrum fargesii previously misidentified as T. fortunei. A. China, Anhui, Huangshan, M.P. Teng & K. Yao 79184 (NAS) (inset a: flower; inset b: aggregate fruit). B. China, Henan, Xinyang, C.F. Zhang 0074 (HENU). C. China, Jiangxi, Lushan, C.Y. Wu et al. L-143 (KUN) (inset: flowers). D. China, Zhejiang, Lin’an, Y.Y. Ho 21730 (PE) (inset a: flower; inset b: aggregate fruit).

FIGURE 11. Thalictrum fargesii in the wild (Tianquan county in Sichuan province, China). A. Habitat. B. Habit. C. Portion of stem. D. Leaf (left: adaxial side; right: abaxial side). E. Leaflet (left: adaxial side; right: abaxial side). F. Flower. G.Aggregate fruits (immature). H. Achenes (immature). Photographed by Y.P. Zeng.

FIGURE 12. Thalictrum fargesii in the wild (Pingwu county in Sichuan province, China). A. Habitat. B. Habit. C. Roots. D. Portion of stem. E. Leaf (left: adaxial side; right: abaxial side). F. Leaflet (left: adaxial side; right: abaxial side). G. Flower. H. Sepal (left: adaxial side; right: abaxial side). I. Stamens. J. Carpels. K. Aggregate fruit (immature). L. Achenes (immature). Photographed by Y.P. Zeng.

FIGURE 13. Selected specimens of Thalictrum fargesii. A. China, Anhui, Huoshan, M.P. Teng & K. Yao 81519 (NAS). B. China, Anhui, Jinzhai, K. Yao 9033 (NAS). C. China, Jiangxi, Lushan, H. Migo s.n. (NAS). D. China, Zhejiang, Lin’an, Y.Y. Ho 21730 (NAS).