Bot. Bull. Acad. Sin. (1995) 36: 253-257

Yang and Compton— The genus Cimicifuga in Taiwan

Additional remarks on Ranunculaceae in Taiwan (4) The genus Cimicifuga L. ex Wernisch.

T.Y. Aleck Yang1,3 and James A. Compton2

1 Division of Collection and Research, National Museum of Natural Science, Taichung, Taiwan, Republic of China

2Department of Botany, Plant Science Laboratory, University of Reading, Whiteknights, Reading, Berkshire,

RG6 2AS, England, U.K.

(Received April 10, 1995; Accepted August 1, 1995)

Abstract. The present work is a taxonomic study of Cimicifuga L. ex Wernisch. (Ranunculaceae) in Taiwan. The morphological and palynological characters of one species, C. simplex (DC.) Wormsk. ex Turcz., are presented. The taxonomic description and a distribution map of this taxon in Taiwan are given.

Keywords: Cimicifuga; Cimicifuga simplex; Morphology; Palynology; Ranunculaceae; Taiwan; Taxonomy.


There are approximately eighteen species of Cimicifuga worldwide. The genus Cimicifuga was not recorded in the account of Ranunculaceae in the Flora of Taiwan (Liu and Hsieh, 1976). Ou (1976) reported the presence of Cimicifuga simplex on Taiwan, based on a herbarium sheet collected as early as 1925 by Japanese botanists from Nantou Hsien. In 1984, the first author of the present paper re-collected C. simplex from Taichung Hsien. Subsequently, a few other collections were made of this apparently rare species from the central mountain ranges of this island (see below). In our paper, we present morphological and palynological studies and document the distribution of C. simplex in Taiwan. A line drawing is provided to facilitate recognition of this species. Nomenclature of this species follows the treatment of Compton and Jury (1995).

Materials and Methods

The materials were taken from living plants and herbarium collections. Scanning electron photomicrographs were made with a Bausch & Lomb Nanolab 2100 (National Chung-Hsing University, Taichung) and a Hitachi S2300 (Tunghai University, Taichung).

Morphological Characters

Three largest mature leaflets of living specimens collected in the wild were chosen for examination. Herbarium specimens or dehydrated fresh materials were coated with gold for SEM study. The leaf surface, stomatal complex, trichomes, and carpels were observed and described.

Palynological Characters

The anthers were dissected from fresh flowers with a pair of fine forceps and a mounted needle. They were then placed on double-sided cellotape on a stub under a dissecting microscope. The anther fragments were drawn to one side, leaving the pollen grains together in the middle. The pollen grains were coated with gold for about 3 min, then SEM observations were made at magnifications from 3,000 to 15,000.

Results and Discussion

Morphological Characters

The surfaces of the leaflets on the adaxial side can be defined as striate (Figure 1A and B) and smooth (Figure 1C) on the abaxial side. The shape of the stomatal complexes is elliptic and the size is 35.2_44.8 µm long and 29.3_32.8 µm wide. The outer stomatal rim is raised with fine, radiating striations (Figure 1C). The trichomes are linear, filiform simple hairs, with both long and short types, and the surface of the trichomes is finely striate (Figure 1D).

The stomatal complexes are anomocytic in the Ranunculaceae—there are no distinct subsidiary cells—and the genus Cimicifuga is hypostomatic, with a high percentage of stomata occurring on the abaxial surface. Sometimes, stomata can be found in reproductive organs, such as the carpel, but there they are smaller (Figure 1E).

Palynological Characters

Grains are tricolpate (Figure 1F and G); the size in polar view is 26.8_29.2 µm long, and in equatorial view is 26.9_27.8 µm long; the apocolpum diameter is 8.4_10.6 µm and colpus width is 3.2_4.5 µm; the colpus end is acute; the colpus membrane is spinulose. The value of

3Corresponding author.

Botanical Bulletin of Academia Sinica, Vol. 36, 1995

Figure 1. SEM micrographs of Cimicifuga simplex. A_C, surface of the leaflets: A and B, adaxial side; C, abaxial side; D, surface of the trichome on the follicle; E, surface of carpel, arrow showing the stoma; F_H, tricolpate pollen grains. F, polar view; G, equatorial view; H, irregular perforation. (All from Yang et al. 416)

Yang and Compton— The genus Cimicifuga in Taiwan

P.V./E.V is 99.6_105; the shape is oblate spheroidal to prolate spheroidal; irregular perforations are present (Figure 1H); the exine is 1_2 µm thick, sexine and nexine are almost equal to each other.

Kumazawa (1936) divided the family Ranunculaceae into four pollen types, viz., type O, which is inaperturate; type I, which is tricolpate with occasionally 6 to 8 colpi; type II, which is polypantocolpate, and type III, which is polypantoporate. The genus Cimicifuga has type I pollen, the majority of which are tricolpate. Wang et al. (1993) and Xi and Ning (1993) examined members of tribe Cimicifugeae in China. Wang et al. recognized that pollen grains of C. simplex had numerous spines and perforations, and Xi and Ning indicated that all grains in the genus Cimicifuga were tricolpate. The second author of the present paper, however, has found species with pantocolpate and pantoporate grains. Xi and Ning's description of C. simplex collected in mainland China was similar to that of those collected in Taiwan, except the size in polar view (29.6_34.8 µm) was bigger than that of those in Taiwan.

Taxonomic Treatment

Cimicifuga simplex (DC.) Wormsk. ex Turcz., Bull. Soc. Imp. Naturalistes Moscou 15(1): 87. 1842; Komarov, Act. Hort. Petrop. 22: 301. 1903; Sugawara, Ill. Fl. Saghal. 2: 883, t.412. 1939; Hsiao, Fl. Reip. Pop. Sin. 27: 97. 1979.—Actaea cimicifuga var. simplex DC., Prodr. 1: 64. 1824.—C. foetida var. simplex (DC.) Regel, Tent. Fl.-Ussur. 13. 1861.—C. simplex var. ramosa Maxim. ex Franch. & Sav., Enum. Pl. Jap. 1: 13. 1875, nom. illeg.—Actaea simplex (DC.) Prantl, Bot. Jahrb. Syst. 9: 246. 1887.—Thalictrodes simplex (DC.) Kuntze, Revis. Gen. Pl. 1: 4. 1891.—C. simplex (DC.) var. typica Nakai, Bot. Mag. (Tokyo) 30: 145. 1916.—C. ramosa Nakai, Cat. Sem. Spor. Hort. Bot. Univ. Imp. Tokyo: 15. 1932, nom. illeg.

Cimicifuga foetida var. intermedia Regel, Reis. Ostsib. 1(1): 122. 1862.—C. simplex var. intermedia (Regel) Nakai, Bot. Mag. (Tokyo) 30: 146. 1916.—C. cimicifuga var. intermedia (Regel) Graebn. & P. Graebn., Syn. Mitteleur. Fl. 5(2): 632. 1926.

Cimicifuga ussuriensis Oett., Trudy Bot. Sada Imp. Yur'evsk. Univ. 6: 138. 1906.

Cimicifuga taquetii H. Lév., Repert. Spec. Nov. Regni Veg. 9(11): 448_449. 1911.

Figures 2 and 3

A perennial deciduous woody shrub, ca. 100_150 cm tall. Rhizome strong, black. Stem round, shallowly sulcate, glabrous or nearly so below inflorescence. Leaves bi- or tripinnately triternate, chartaceous; petioles ca. 7.5_30 cm long, sparsely pubescent, glabrous or nearly so, with an extended membrane sheathing the base, sparsely pubescent. Leaflets 9_27 or more, terminal leaflet ca. 35_85 mm long, 20_55 mm wide, broadly ovate, ovate, elliptic or obovate; apex acuminate; base obtuse, truncate or acute;

3-lobed; margin serrate or coarsely serrate; glabrous or nearly so on adaxial surface, sparsely pubescent on abaxial surface at least along nerves and margin; mainveins 1_3, nerves flat or sunken on adaxial side and elevated or flat on abaxial side; petiolulate. Inflorescence racemose, usually simple, or with 2 or 3 or more short basal branchlets, terminal or axillary; pedicels ca. 3_8 mm long, elongating after anthesis, ca. 5_13 mm long, albovelutinous; bracts chartaceous, glabrous; bracteoles 3, subulate, glabrous. Flowers ca. 5_8 mm diameter, white. Sepals 4, ca. 3.5_5 mm long, 2_3 mm wide, broadly ovate-elliptic, glabrous on both sides except with very sparsely pubescent margins. Stamens numerous, ca. 3.5_7.5(_10) mm long, glabrous; anthers oblong, with lateral longitudinal dehiscence; connective shorter than the pollen sacs; filaments linear, with 1 central nerve. Staminodes 2, rarely 3, ca. 4.6_6.5 mm long, 2_3 mm wide, cupuliform, broadly elliptic or ovate, glabrous on both sides, apex bilobed, nectary at the base, stipe with three nerves. Carpels 2_8, ca. 3_5 mm long, pubescent, except style glabrous; ovaries long stipitate, ca. 0.8_1.5 mm long. Fruits ca. 70_130 mm long, follicles oblong, dark brown, sparsely pubescent; stipes elongating after anthesis, ca. 2_4 mm long. Seeds 4_8, elliptical, brown, ca. 1.3_2 mm long, with extended membranaceous scales from the sides.

Distribution. Distributed around southwest, central and northeast China (Sichuan, Hubei, Shaanxi, Gansu, Inner Mongolia, Hebei, Henan, Liaoning, Jilin, and Heilongjiang provinces), Mongolia, eastern Siberia, Korea and south to Taiwan. In Taiwan, it is represented in the Central Mountains ca. 2,200_3,200 m altitude in open lands or forest margins. Flowering late July to early October; fruiting late August to November.

Additional Specimen Examined. MAINLAND CHINA. JILIN: Mt. Changbaishan, 1800 m, B.B. Wan & Chow 81065 (K). HENAN: Lushi Hsien, Laochungshan, K.M.

Figure 2. Latitudinal and altitudinal distribution of Cimicifuga simplex in Taiwan.

Botanical Bulletin of Academia Sinica, Vol. 36, 1995

Liou 5350 (K). HUBEI: A. Henry 6073 (K), E.H. Wilson 1629 (K), 1675 (K), 2592 (K); Ichang, A. Henry 4903 (K), 4935 (K). SICHUAN: Omei, Mt Omei, E.H. Wilson 4713 (K). TAIWAN. MIAOLI HSIEN: Mt. Chunghsueshan, 2400 m, T.C. Huang 10286 (TAI), 2500 m, T.Y. Yang & Huang et al. 416 (HAST, K, RNG, TAI, TNU, TUNG, National Museum of Natural History, Taichung, Taiwan), Yang & Yang 5878 (TAI, TUNG, National Museum of Natural History, Taichung, Taiwan). NANTOU HSIEN: Mt. Nenkaochushan, Nov 1925, Y. Saito & Kikuchi, s.n.

(TCF). KAOHSIUNG HSIEN: South Cross Highway, B. & S. Wynn-Jones 343 (priv. herbarium), 2728 m, B. & S. Wynn-Jones 1486 (priv. herbarium). HUALIEN HSIEN: Mt. Chilaishan, 2870 m, K.F. Chung 46 (TAI), Mt. Chilaichushan, Nov 1925, Y. Saito & Kikuchi s.n. (TCF). KOREA: N. Kankyo, E.H. Wilson 8984 (K).

Notes. Cimicifuga simplex is distinguished by the sometimes simple, elongate terminal raceme on the inflorescence, the white, cupuliform staminodia with deeply bifid

Figure 3. Cimicifuga simplex (DC.) Wormsk. ex Turcz. A, habit; B, central bracteole; C, lateral bracteole; D, sepal; E, staminode; F, stamen; G, immature carpel; H, fruit; I, pubescence on the fruit; J, seed.

Yang and Compton— The genus Cimicifuga in Taiwan

lobes, the three short deltoid-acute bracteoles subtending the pedicels and long stipitate carpels, which elongate after anthesis. All other Asian species of Cimicifuga possess short stipitate carpels, paniculate inflorescence, or if simple, then virgate with entire or emarginate staminode lobes.

Acknowledgments. We thank the herbarium curators who loaned us specimens for study. Thanks are also due to Prof. V.C. Yang (Tunghai University), who permitted T.Y. Aleck Yang to use SEM for morphological and palynological studies, and Mr. G.Z. Song (TAI), who printed the micrographs for this paper.

Literature Cited

Compton, J.A. and S.L. Jury. 1995. Lectotypification of Cimicifuga simplex (Ranunculaceae) and some synonyms. Taxon 44: 401_404.

Hsiao, P.K. 1979. Cimicifuga L. In Flora of China Edit. Comm. (eds.), Flora Reipublicae Popularis Sinicae, vol. 27. Science Press, Beijing, pp. 93_103. (in Chinese)

Kumazawa, M. 1936. Pollen grain morphology in Ranunculaceae, Lardizabalaceae and Berberidaceae. Jap. J. Bot. 8: 19_46, Plates II_VI.

Liu, T.S. and C.F. Hsieh. 1976. Ranunculaceae. In H.L. Li, T.S. Liu, T.C. Huang, T. Koyama, and C.E. DeVol (eds.), Flora of Taiwan, vol. 2. Epoch Publ. Co., Taipei, pp. 476_513.

Ou, C.H. 1976. Contributions to the dicotyledonous plants of Taiwan (I). Quart. J. Chin. Forest. 9(2): 117_126. (in Chinese with English abstract)

Wang, X.Q., D.Y. Hong, and Z.Y. Li. 1993. A study on pollen and seed coat in the tribe Cimicifugeae and some allied genera (Ranunculaceae). Cathaya 5: 131_149.

Xi, Y.Z. and J.C. Ning. 1992. Pollen morphology and its taxonomic significance of tribe Cimicifugeae. Yushania 10: 45_60. (in Chinese with English abstract)