Bot. Bull. Acad. Sin. (2002) 43: 313-327

Shui et al. — Synopsis of the Chinese species of Begonia

Synopsis of the Chinese species of Begonia (Begoniaceae), with a reappraisal of sectional delimitation

Yu-Min Shui¹, Ching-I Peng^2,*, and Cheng-Yih Wu¹

¹Kunming Institute of Botany, The Chinese Academy of Sciences, Kunming 650204, P.R. China

²Institute of Botany, Academia Sinica, Nankang, Taipei 115, Taiwan

(Received July 12, 2000; Accepted July 21, 2002)

Abstract. Nine sections of Begonia are recognized and illustrated based on the 150 species known from the Chinese region. They are sect. Coelocentrum Irmsch., sect. Petermannia (Klotzsch) A. DC., sect. Alicida C.B. Clarke, sect. Diploclinium (Wight) A. DC., sect. Reichenheimia (Klotzsch) A. DC., sect. Parvibegonia A. DC., sect. Platycentrum (Klotzsch) A. DC., sect. Sphenanthera (Hassk.) Warb. and sect. Leprosae (T.C. Ku) Y.M. Shui (combination made here). Section Leprosae is characterized by the trilocular, wingless, clavate, berry-like fruits. Based on a detailed analysis of fruits and placentae, the sectional placement of many species of Chinese Begonia is realigned. Relationships among the sections are discussed: sect. Coelocentrum is likely to represent one of the most ancient groups in the genus Begonia; sect. Diploclinium, a complex group, is related to several sections; sect. Sphenanthera is an entomochoric and entomophilous group; sect. Leprosae is intermediate between sections Sphenanthera and Diploclinium. Begonia tessaricarpa C.B. Clarke (sect. Sphenanthera), from Xizang, is documented here as a new record for China.

Keywords: Begonia; China; Sections; Taxonomy.

Introduction

Begonia L. is a genus of ca. 1,400 species, comprising nearly all the species of Begoniaceae (Smith et al., 1986). Plants of Begonia usually grow in moist, tropical and subtropical areas. As plants of much horticultural interest, they have been studied by many botanists notably De Candolle (1859, 1864); Clarke (1879) and Warburg (1894) in the 19^th century. It was not until 1925 that E. Irmscher reviewed the former works and classified world Begonia into 65 sections: 34 sections in America, 13 in Africa, and 18 in Asia. Barkley (1972) and Baranov and Barkley (1974) adopted Irmscher's system with little modification. Smith et al. (1986), despite making a significant effort to provide an illustrated guide to Begonia, did not attempt to delimit sections. More recently, Doorenbos et al. (1998) reviewed Begonia, and classified the genus into 63 sections (29 in America, 16 in Africa, and 18 in Asia). Doorenbos et al. (1998) considered sect. Begonia is not American-Asiatic in its distribution but strictly American (mainly in the West Indies and Brazil), which is followed here.

Irmscher (1939) was the first to classify the 51 species of Begonia known from China at that time into 6 sections, Begoniastrum, Diploclinium, Sphenanthera, Reichenheimia, Platycentrum and Coelocentrum. Yü (1948), the first Chinese botanist to study Begonia systematically, divided the 30 species of Begonia in southwestern China into five sections: Sphenanthera, Begoniastrum, Platycentrum, Parvibegonia, and Reichenheimia. Liou (1983) placed 50 species from China into five sections: Coelocentrum, Sphenanthera, Begoniastrum, Platycentrum and Reichenheimia. More recently, Ku (1999a) recognized 139 species from China and divided them among 6 sections: Coelocentrum, Begonia, Reichenheimia, Sphenanthera, Pleiothece and Platycentrum. A detailed comparison of these studies reveals discrepancies and uncertainties in the sectional assignment of a number of species. According to our studies, the confusion resulted mainly from errorneous observations or a lack of observations altogether on the characters of fruits and placentae, the most important diagnostic characters for delimiting sections. For example, Begonia brevicaulis T.C. Ku (1997) placed in sect. Reichenheimia when initially published, is a taxonomic synonym of Begonia gulinqingensis S.H. Huang & Y.M. Shui (1994) in sect. Diploclinium. It is therefore of both scientific interest and pragmatic merit to clearly delimit the sections of Chinese Begonia.

*Corresponding author. Tel: 886-2-2789-9590 ext 404; Fax: 886-2-2789-1623; E-mail: bopeng@sinica.edu.tw

Botanical Bulletin of Academia Sinica, Vol. 43, 2002

Key to the Sections of Chinese Begonia (Based Mainly on Chinese Material)

1. Fruit dehiscent, capsular; tepals pinkish or red, rarely white.

2. Placentae parietal Sect. 1. Coelocentrum Irmsch.

2. Placentae axile.

3. Ovary 3-locular.

4. Placentae bifid or multifid.

5. Flower solitary, axillary Sect. 2. Petermannia (Klotzsch) A. DC.

5. Flowers many, at summit of stems and branches.

6. Herbs less than 10 cm tall; tepals less than 0.5 cm long; fruit wall papery, dehiscent irregularly Sect. 3. Alicida C.B. Clarke

6. Herbs or subshrubs 20-200 cm tall; tepals 0.5-1.5 cm long; fruit wall usually hard, dehiscent regularly along dorsal sutures Sect. 4. Diploclinium (Wight) A. DC.

4. Placentae undivided Sect. 5. Reichenheimia (Klotzsch) A. DC.

3. Ovary 2- locular.

7. Plant herbaceous, tuberous, mostly deciduous; fruit dehiscent irregularly on fruit surface, the wings subequal Sect. 6. Parvibegonia A. DC.

7. Plant herbaceous or shrubby, rhizomatous, mostly evergreen; fruit dehiscent along sutures, abaxial wing conspicuously elongate Sect. 7. Platycentrum (Klotzsch) A. DC.

1. Fruit tardily dehiscent, berry-like, bursting very late after maturation; tepals white, rarely pinkish.

8. Fruit clavate, without horns or wings; ovary 3-locular; herbs; stems creeping; plant monoecious Sect. 8. Leprosae (T.C. Ku) Y.M. Shui

8. Fruit turbinate, with indistinct horns or wings; ovary 3-4 (-7)-locular; herbs or subshrubs; stems creeping or erect; plant monoecious (dioecious when ovary 4- or more locular) Sect. 9. Sphenanthera (Hassk.) Warb.

Sectional Delimitation of Chinese Begonia

Sect. 1. Coelocentrum Irmsch., Mitt. Inst. Allg. Bot. Hamburg 10: 533. 1939; Liou, Iconogr. Cormophyt. Sin. Suppl. 2: 532, 538. 1983, in clavi; Wu and Ku, Acta Phytotax. Sin. 35(1): 43, fig. 25. 1997; Doorenbos et al., The Sections of Begonia 84. 1998; Shui and Huang, Acta Bot. Yunnan. 21(1): 21, fig. 8: 1-14. 1999; Ku, Acta Phytotax. Sin. 37(3): 285. 1999 & Fl. Reipubl. Popularis Sin. 52(1): 127, 402. 1999; Shui, Acta Phytotax. Sin. 40(4): 374. 2002.—LECTOTYPE SPECIES: Begonia porteri H. Lév. & Vaniot (designated by Barkley and Baranov, 1972).

Herbs, monoecious, epipetric, rhizomatous creeping, acaulescent. Leaves mostly broadly ovate, rarely peltate, palmately veined. Inflorescence terminal, tepals red; staminate flowers: tepals 4, anthers yellow; carpellate flowers: tepals 3-5, styles 3, stigmatic surface a continual helical band with 2 coils. Ovary unilocular; placentae 3, parietal, bifid from apical to middle part of ovary but axile and undivided at base (Figures 1, 2). Capsule nodding, usually with 3 subequal wings, dehiscent.

Cytology. 2n = 30 (B. masoniana Irmsch.: Legro and Doorenbos, 1969).

Distribution and Habitat. About 18 species in China (Guangxi, SE Yunnan and S Guizhou) and Vietnam (Tonkin), seven of which published after 1995. In China, all species grow on limestones between 300 and 1,300 m elevation.

Species Found in China:

1 B. biflora T.C. Ku

2 B. bonii Gagnep. [previously assigned to sect. Reichenheimia (Dorrenbos et al., 1998; Ku, 1999; Golding and Wasshausen, 2002)]

3 B. cirrosa L.B. Sm. et al.

4 B. daxinensis T.C. Ku

5 B. filiformis Irmsch. [sect. Reichenheimia (Irmscher, 1939; Liou, 1983; Dorrenbos et al., 1998; Ku, 1999; Golding and Wasshausen, 2002)]

6 B. guangxiensis C.Y. Wu

7 B. lanternaria Irmsch.

8 B. luzhaiensis T.C. Ku

9 B. masoniana Irmsch.

10 B. morsei Irmsch.

11 B. obliquefolia S.H. Huang & Y.M. Shui

12 B. ornithophylla Irmsch.

13 B. porteri H. Lév. & Vaniot

14 B. pseudodryadis C.Y. Wu [previously assigned to sect. Platycentrum (Wu and Ku, 1995; Dorrenbos et al., 1998; Golding and Wasshausen, 2002)]

15 B. setuloso-peltata C.Y. Wu

16 B. umbraculifolia Y. Wan & B.N. Chang

Shui et al. — Synopsis of the Chinese species of Begonia

17 B. yishanensis T.C. Ku

18 B. zhengyiana Y.M. Shui (Shui, 2002b)

Sect. 2. Petermannia (Klotzsch) A. DC., Ann. Sci. Nat. Bot. Ser. 4. 11: 128. 1859; Doorenbos et al., The Sections of Begonia 148. 1998.—Petermannia Klotzsch, Monatsber. Königl. Preuss. Akad. Wiss. Berlin 124. 1854. —Sect. Begonia Ser. Begonia, pro parte, Fl. Reipubl. Popularis Sin. 52(1): 145, and B. hainanensis Chun & F. Chun, in clavi, 157. 1999.—LECTOTYPE SPECIES: Begonia cumingiana (Klotzsch) A. DC. (designated by Barkley and Baranov, 1972) (º Petermannia cumingiana Klotzsch).

Herbs or subshrubs, monoecious, terrestrial; stem with a creeping base. Leaves pinnately veined. Flowers solitary, axillary, tepals red; staminate flowers unknown; carpellate flowers: tepals 5, styles 3, bifid, stigmatic surface with 2 coils. Placentae bifid from summit to the base of ovary. Capsules nodding, 3-locular, with 3 subequal wings (Figure 3).

Cytology. 2n = 30 (seven species, e.g., B. isoptera Dryand. ex J.E. Sm.: Legro and Doorenbos, 1969, 1971, 1973), 44 (three species, e.g., B. brevirimosa Irmsch.: Legro and Doorenbos, 1969, 1971).

Distribution and Habitat. Over 193 species in SE Asia, mainly Malaysia (Doorenbos et al., 1998); only one species, Begonia hainanensis Chun & F. Chun, in China (endemic to Hainan Province, on wooded valley cliffs, ca. 300 m)

Species Found in China. Begonia hainanensis Chun & F. Chun [previously assigned to sect. Begoniastrum (Liou, 1983) and sect. Begonia ( Ku, 1999)]

Sect. 3. Alicida C.B. Clarke, Fl. Brit. Ind. 2: 637. 1879. Wu and Ku, Acta Phytotax. Sin. 33(3): 252, fig. 2. 1995; Doorenbos et al., The Sections of Begonia 64. 1998.—LECTOTYPE SPECIES: Begonia alicida C.B. Clarke, "alaecida" (designated by Barkley and Baranov, 1972).

Herbs, monoecious, terrestrial, rarely epiphytic (B. peii C.Y. Wu), erect; stem usually less than 10 cm tall; leaves palmately veined. Inflorescence terminal; staminate flowers: tepals 4, red, filaments fused up to 1/2, anthers yellow; carpellate flowers: tepals 3 or 4, red, styles 2 (rarely 3), fused at base, bifid or half-moon shaped band without coils. Placentae axile, bifid at middle of ovary, unknown at summit and the base of ovary. Fruit suberect, 3-locular, irregularly dehiscent, with 3 subequal wings (Figure 4).

Cytology. Unknown.

Distribution and Habitat. Four species in Burma and China. Only one species, Begonia péii C.Y. Wu, in China, occurring on limestone rocks at ca. 1,000 m in Xishuangbanna, southern Yunnan.

Species Found in China. Begonia péii C.Y. Wu [previously assigned to sect. Parvibegonia (Doorenbos et al., 1998; Golding and Wasshausen, 2002) and sect. Begonia (Ku, 1999)]

Sect. 4. Diploclinium (Lindl.) A. DC., Ann. Sci. Nat. Bot. Ser. 4. 11: 129. 1859; Dorrenbos et al., The Sections of Begonia 90. 1998. —Diploclinium Lindl., Veg. Kingd. 319. 1846. —Sect. Diploclinium and Trilobaria Irmsch., Mitt. Inst. Allg. Bot. Hamburg 6: 354. 1927. —Sect. Begonia Ser. Diploclinium (Lindl.) S.H. Huang & Y.M. Shui ex T.C. Ku, Fl. Reipubl. Popularis Sin. 52(1): 145, in clavi, 128, 152, pl. 31: 1-5. 1999.

Sect. Begonia (excluding type), sensu Wu and Ku, Acta Phytotax. Sin. 33(3): 251, 269, 254, figs. 1, 4, 5, 7, 8, 9, 10, 15. 1995 & Acta Phytotax. Sin. 35(1): 50, fig. 30. 1997; Shui and Huang, Acta Bot. Yunnan. 21(1): 15, fig. 3:1-9. 1999; Ku, Fl. Reipubl. Popularis Sin. 52(1): 128, 144, in clavi. 1999.—Sect. Begoniastrum A. DC., Ann. Sci. Nat. Bot. Ser. 4. 11: 123. 1859; Yü, Bull. Fan. Mem. Inst. Biol. 1(2): 115. 1948; Liou, Iconogr. Cormophyt. Sin. Suppl. 2: 533. 1983.—LECTOTYPE SPECIES: Begonia grandis Dryand. (designated by Doorenbos et al., 1998)(= Diploclinium evansianum (Andrews) Lindl.).

Herbs or subshrubs, monoecious, terrestrial, seldom epiphytic, erect, acaulous or stems decumbent, occasionally with a tuberous base and axillary bulbils. Inflorescence usually terminal, rarely axillary, oblique in some species, tepals pink; staminate flowers: tepals 4 (rarely 2), anthers yellow, connective scarcely protruded; carpellate flowers: tepals 4 or 5 (rarely 2 or 3), styles 3, style-branches often twisted, stigma forming a continual helical band with 1 or 2 coils. Placentae axile, bifid or cleft at upper and middle part of ovary, rarely parietal at the upper part (Begonia wangii T.T. Yü; Figure 5A), either bifid or undivided at the ovary base. Capsules erect or nodding, 3-locular, with 3 equal or unequal wings (Figures 5-9).

Cytology. 2n = 22 (B. cyclophylla Hook. f.: Legro and Doorenbos, 1971; B. fimbristipula Hance: Doorenbos et al., 1998; B. picta Sm.: Legro and Doorenbos, 1969); 26 (B. acaulis Merr. & Perry: Legro and Doorenbos, 1969; B. fenicis Merr.: Oginuma and Peng, 2002), 28 (B. grandis Dryand. var. evansiana (Andrews) Irmsch.: Legro and Doorenbos, 1969), 30 (B. cavaleriei H. Lév.: Tian et al., 2002), 32 (B. subnummularifolia Merr.: Legro and Doorenbos, 1971), 36 (B. ravenii C.I Peng & Y.K. Chen: Peng et al., 1988; Oginuma and Peng, 2002), 38 (B. taiwaniana Hayata: Peng and Chen, 1991; Oginuma and Peng, 2002), 44 (B. nigritarum (Kamel) Steud.: Doorenbos et al., 1998), 52 (B. lukuana Y.C. Liu & C.H. Ou: Oginuma and Peng, 2002), 56 (B. fenicis Merr.: Kokubugata and Madulid, 2001).

Distribution and Habitat. About 160 species in Asia, as far north as 45° N latitude. Forty-five species in China, growing on thick humus in forest along rivers, or occurring on rock faces and/or tree trunks, often with tuberous stem-base, occasionally with thin and elongate rhizomes creeping on ground surface; elev. ca. 300-3,600 m.

Species Found in China:

1 B. acutitepala K.Y. Guan & D.K. Tian [previously assigned to sect. Begonia (Guan and Tian, 2000; Golding and Wasshausen, 2002)]

Botanical Bulletin of Academia Sinica, Vol. 43, 2002

651) and a paratype (Y.M. Shui Lunan 44) of B. guishanensis S.H. Huang & Y.M. Shui, which was published a year earlier. Thus, we reduce B. rhodophylla C.Y. Wu to the synonym of B. guishanensis S. H. Huang & Y.M. Shui.

16 B. gulinqingensis S.H. Huang & Y.M. Shui [as sect. Begonia (Ku, 1999)]

—B. brevicaulis T.C. Ku, Acta Phytotax. Sin. 35(1): 53, fig. 31. 1997, non B. brevicaulis A. DC. [previously assigned to sect. Reichenheimia (Wu and Ku, 1997; Dorrenbos et al., 1998)]

—B. sinobrevicaulis T.C. Ku [previously assigned to sect. Reichenheimia (Ku, 1999; Golding and Wasshausen, 2002)], syn. nov.

Our careful examination of all the Begonia collections from Malipo County, Yunnan, where both holotypes of B. sinobrevicaulis and B. gulinqingensis were collected, as well as additional field explorations to the type locality revealed that the placentae of B. sinobrevicaulis are axial and bifid in each locule. Thus, it clearly belongs to sect. Diploclinium. Furthermore, morphological comparison of B. sinobrevicaulis and B. gulinqingensis suggests that the two are conspecific.

17 B. hymenocarpa C.Y. Wu, as sect. Begonia (Wu and Ku, 1995; Ku, 1999)

18 B. imitans Irmsch. [previously assigned to sect. Begoniastrum (Irmscher, 1939); sect. Begonia (Ku, 1999)]

19 B. josephii A. DC. [previously assigned to sect. Begoniastrum (Liou, 1983); sect. Begonia (Ku, 1999)]

20 B. labordei H. Lév. [previously assigned to sect. Begoniastrum (Irmscher, 1927, 1939; Yü, 1948; Liou, 1983); sect. Begonia (Ku, 1999)]

21 B. lukuana Y.C. Liu & C.H. Ou [previously assigned to sect. Begonia (Ku, 1999)]

22 B. malipoensis S.H. Huang & Y.M. Shui [previously assigned to sect. Begonia (Ku, 1999)]

23 B. miranda Irmsch. [previously assigned to sect. Begoniastrum (Irmscher, 1951; Liou, 1983); sect. Begonia (Ku, 1999)]

24 B. morifolia T.T. Yü [ previously assigned to sect. Begoniastrum (Yü, 1948; Liou, 1983); sect. Begonia (Ku, 1999)]

—B. anceps Irmsch. [previously assigned to sect. Begoniastrum (Irmscher, 1951); sect. Begonia (Ku, 1999)], syn. nov.

According to Prof. C.Y. Wu (pers. comm.), B. anceps is separable from B. morifolia by having less deeply serrate leaves. However, leaf shapes tend to be variable and the two species are hardly distinguishable otherwise. Begonia anceps Irmsch. is hereby reduced to a synonym of B. morifolia T.T. Yü.

25 B. muliensis T.T. Yü (1948) [previously assigned to sect. Begoniastrum (Ku, 1999); sect. Begonia (Ku, 1999)]

2 B. alveolata Yü [previously assigned to sect. Begoniastrum (Yü, 1948; Liou, 1983); sect. Begonia (Ku, 1999)]

—Begonia pingbienensis C.Y. Wu [previously assigned to sect. Begonia (Wu and Ku, 1995; Ku, 1999)], syn. nov.

A detailed comparison of the holotypes of B. pingbienensis (H.T. Tsai 61724) and B. alveolata (C.W. Wang 82780), both from Pingbien, Yunnan, reaveals that the two species are synonymous.

3 B. arboreta Y.M. Shui (Shui, 2002a)

4 B. asperifolia Irmsch. [previously assigned to sect. Begoniastrum (Irmscher, 1927, 1939; Yü, 1948; Liou, 1983); sect. Begonia (Ku, 1999)]

5 B. cavaleriei H. Lév. [previously assigned to sect. Begoniastrum (Liou, 1983); sect. Begonia (Ku, 1999)]

6 B. cehengensis T.C. Ku [previously assigned to sect. Begonia (Wu and Ku, 1997; Ku, 1999)]

7 B. clavicaulis Irmsch. [previously assigned to sect. Begoniastrum (Irmscher, 1939; Yü, 1948; Liou, 1983) and sect. Begonia (Ku, 1999)]

8 B. coptidimontana C.Y. Wu [previously assigned to sect. Alicida ("Alaecida", Wu and Ku, 1995); sect. Begonia (Ku, 1999)]

9 B. dentatobracteata C.Y. Wu ["B. dentato-bracteata", previously assigned to sect. Begonia (Wu and Ku, 1995; Ku, 1999)]

10 B. fenicis Merr. [previously assigned to sect. Begonia (Ku, 1999)]

11 B. fimbristipula Hance [previously assigned to sect. Begoniastrum (Irmscher, 1939; Yü, 1948; Liou, 1983); sect. Begonia (Ku, 1999)]

12 B. fordii Irmsch. [previously assigned to sect. Begoniastrum (Irmscher, 1939); sect. Begonia (Ku, 1999)]

13 B. glechomifolia C.M. Hu ex C.Y. Wu & T.C. Ku[previously assigned to sect. Begonia (Wu and Ku, 1995; Ku, 1999)]

14 B. grandis Dryand. [previously assigned to sect. Begoniastrum (Irmscher, 1939; Yü, 1948); sect. Begonia (Ku, 1999)]

—B. sinensis A. DC. [previously assigned to sect. Begoniastrum (Irmscher, 1927; Liou, 1983)]

—B. evansiana Andrews [previously assigned to sect. Begoniastrum (Liou, 1983)]

15 B. guishanensis S.H. Huang & Y.M. Shui

—B. rhodophylla C.Y. Wu [previously assigned to sect. Begonia (Wu and Ku, 1995; Ku, 1999)], syn. nov.

A careful examination of the holotypes of both B. guishanensis and B. rhodophylla, both from Lunan County, Yunnan, reveals that they are synonymous. In fact, in the protologue of B. rhodophylla Wu and Ku (1995) cited as paratypes the holotype (Y.M. Shui B91-

Shui et al. — Synopsis of the Chinese species of Begonia

26 B. nymphaeafolia T.T. Yü [previously assigned to sect. Reichenheimia (Yü, 1948; Barkley, 1972)]

27 B. obsolescens Irmsch. [previously assigned to sect. Coelocentrum (Irmscher, 1951; Liou, 1983; Ku, 1999; Golding and Wasshausen, 2002); sect. Begonia (Shui and Huang, 1999)]

—B. fengii T.C. Ku [previously assigned to sect. Platycentrum (Wu and Ku, 1995); sect. Begonia (Ku, 1999)], syn. nov.

We have examined an isotype of B. fengii (K.M. Feng 22990, KUN), which is the same as B. obsolescens in aspects, characters of fruits, indumentum, etc. Types of both B. fengii and B. obsolescens were collected from Malipo Xian, Yunnan. In our opinion, B. fengii is merely a diminutive individual of B. obsolescens and should be reduced to a synonym of the latter species.

28 B. peltatifolia H.L. Li [previously assigned to sect. Reichenheimia? (Dorrenbos et al., 1998; Golding and Wasshausen, 2002); sect. Begonia (Ku, 1999)]

29 B. picta Sm. [previously assigned to sect. Begonia (Ku, 1999)]

30 B. ravenii C.I Peng & Y.K. Chen [previously assigned to sect. Begonia (Ku, 1999)]

31 B. rongjiangensis T.C. Ku [previously assigned to sect. Begonia (Wu and Ku, 1995; Ku, 1999)]

32 B. rotundilimba S.H. Huang & Y.M. Shui [previously assigned to sect. Begonia (Ku, 1999)]

33 B. ruboides C.M. Hu ex C.Y. Wu & T.C. Ku [previously assigned to sect. Begonia (Wu and Ku, 1995; Ku, 1999)]

34 B. setifolia Irmsch. [previously assigned to sect. Platycentrum (Irmscher, 1939; Barkley et al., 1972; Liou, 1983; Dorrenbos et al., 1995; Golding and Wasshausen, 2002) and sect. Begonia (Ku, 1999)]

—B. tsaii Irmsch. [previously assigned to sect. Platycentrum (Irmscher, 1951; Barkley et al., 1972; Dorrenbos et al., 1998; Ku, 1999; Golding and Wasshausen, 2002)], syn. nov.

We have examined an isotype of B. tsaii Irmsch. (H.T. Tsai 62727, PE). Although this specimen was badly pressed, the long, red setae on the upper surfaces of leaves and long crisped hairs on the petioles are immediately indicative of B. setifolia Irmsch. In addition, the first author's field study and collection for many years in SE Yunnan where both `taxa' co-occur strongly suggests that B. tsaii Irmsch. is conspecific with B. setifolia Irmsch.

35 B. sinofloribunda Dorr [B. floribunda T.C. Ku, non B. floribunda Carrière, previously assigned to sect. Platycentrum (Wu and Ku, 1997; Ku, 1999)]

36 B. sino-vietnamica C.Y. Wu [previously assigned to sect. Begonia (Wu and Ku, 1997; Ku, 1999)]

37 B. summoglabra T.T. Yü [previously assigned to sect. Begoniastrum (Yü, 1948), sect. Reichenheimia (Liou, 1983) and sect. Begonia (Ku, 1999)]

38 B. taiwaniana Hayata [previously assigned to sect. Platycentrum (Barkley and Baranov, 1972), sect. Begoniastrum (Liou, 1983) and sect. Begonia (Ku, 1999)]

39 B. taliensis Gagnep. [previously assigned to sect. Begoniastrum (Irmscher, 1927, 1939; Liou, 1983), sect. Reichenheimia (Yü, 1948; Barkley et al., 1972) and sect. Begonia (Ku, 1999)]

40 B. wangii T.T. Yü [previously assigned to sect. Reichenheimia (Yü, 1948; Barkley et al., 1972), sect. Begoniastrum (Liou, 1983) and sect. Begonia (Ku, 1999)]

41 B. wenshanensis C.M. Hu ex C.Y. Wu & T.C. Ku [previously assigned to sect. Begonia (Wu and Ku, 1995; Ku, 1999)]

42 B. xingyiensis T.C. Ku [previously assigned to sect. Begonia (Wu and Ku, 1995; Ku, 1999)]

43 B. xishuiensis T.C. Ku [previously assigned to sect. Begonia (Wu and Ku, 1995; Ku, 1999)]

44 B. yui Irmsch. ["yuii", previously assigned to sect. Begoniastrum (Irmscher, 1951; Liou, 1983) and sect. Begonia (Ku, 1999)]

45 B. yunnanensis H. Lév. [previously assigned to sect. Begoniastrum (Irmscher, 1939; Yü, 1948; Liou, 1983) and sect. Begonia (Ku, 1999)]

As the most polymorphic section in Asia (Doorenbos et al., 1998), sect. Diploclinium is not only similar to the New World sections Begonia and Knesebeckia (Klotzsch) A. DC., but also related to several other sections in Asia. Further studies of sect. Diploclinium are desirable.

Sect. 5. Reichenheimia (Klotzsch) A. DC., Prodr. 15(1): 385. 1864; Liou, Iconogr. Cormophyt. Sin. Suppl. 2: 533, 537. 1983; Wu and Ku, Acta Phytotax. Sin. 35(1): 53, fig. 31. 1997; Doorenbos et al., The Sections of Begonia 170. 1998; Ku, Fl. Reipubl. Popularis Sin. 52(1): 128, 129, in clavi, 197. 1999.—Reichenheimia Klotzsch, Abh. Königl. Akad. Wiss. Berlin `1854':174. 1855 & Begoniac. 54. 1855.—LECTOTYPE SPECIES: Begonia tenera Dryand. (designated by Barkley and Baranov, 1972) (=Reichenheimia thwaitesii (Hook.) Klotzsch).

Herbs, monoecious, epipetric, stemless, rarely erect, mostly deciduous with a tuberous stem-base, rarely rhizomatous. Inflorescence terminal, tepals pink, rarely white; staminate flowers: tepals 4, rarely 2, anthers yellow, filaments free or fused at base; carpellate flowers: tepals 2-5, styles 3, mostly persistent, fused at base, shortly bifid, stigma forming a continual helical band with 1 coil. Placentae axile and undivided from the top to the bottom of ovary, rarely bifid at the upper part. Capsule 3-locular, wings 3, subequal, the wings rarely different (B. lithophila C.Y. Wu) or lacking (B. wilsonii Gagnep.) (Figures 10-12).

Cytology. 2n = 30 (B. morelii Irmsch.: Legro and Doorenbos, 1973), 32 (B. floccifera Bedd.: Sharma and Bhattacharyya, 1961; Legro and Doorenbos, 1971), 34 (B. goegoensis N.E. Br.: Legro and Doorenbos, 1969), 44 (B. nuri Irmsch.: Legro and Doorenbos, 1973).

Botanical Bulletin of Academia Sinica, Vol. 43, 2002

Figures 1-11. Fruits and placentae of representative species of Begonia from China* (illustrated by L. Wang). Sect. Coelocentrum: 1, B. obliquifolia (Shui B92-20, YUNU), 2, B. cirrosa (Shui B92-64, YUNU); Sect. Petermannia: 3, B. hainanensis (modified from Chun and Chun, 1939); Sect. Alicida: 4, B. péii (P. J. Pei 59-9976, KUN); Sect. Diploclinium: 5, B. wangii (Shui B91-307, YUNU), 6, B. cavaleriei (Shui B91-287, YUNU), 7, B. guishanensis (Shui B91-651, YUNU), 8, B. coptidimontana (Shui B91-595, YUNU), 9, B. obsolescens (Shui B91-120, YUNU); Sect. Reichenhaimia: 10, B. parvula (Shui B91-628, YUNU), 11, B. lithophila (Shui B91-650, YUNU). *In each figure the lateral view of a fruit is designated by A; the upper, middle and basal cross-section of an ovary is designated by B, C, and D, respectively.

Shui et al. — Synopsis of the Chinese species of Begonia

Figures 12-21. Fruits and placentae of representative species of Begonia from China (illustrated by L. Wang) (cont.). Sect. Reichenhaimia: 12, B. wilsonii (G.H. Yang 57191, KUN); Sect. Parvibegonia: 13, B. discreta (C. W. Wang 79447, PE); Sect. Platycentrum: 14, B. oreodoxa (Shui 90-42, YUNU), 15, B. psilophylla (Shui B91-409, YUNU); Sect. Leprosae: 16, B. leprosa (modified from Hooker, 1900), 17, B. longicarpa (D. K. Tian 9727, KUN); Sect. Sphenanthera: 18, B. crassirostris (Shui B91-610, YUNU), 19, B. balansana (Shui B91-21, YUNU), 20, B. tetragona (Shui 12924, KUN), 21, B. ceratocarpa (Shui B92-06, YUNU). *In each figure the lateral view of a fruit is designated by A, and the upper, middle and basal cross-section of an ovary is designated by B, C, and D, respectively.

Distribution and habitat. About 46 species in SE Asia and E Asia as far as India. In China, about 5 species, ranging from SW China to SE China, mostly in warm, tropical secondary forests at 300-2,600 m.

Species Found in China:

1 B. chingii Irmsch.

2 B. henryi Hemsl.

3 B. lithophila C.Y. Wu [previously assigned to sect. Begonia (Wu and Ku, 1995; Ku, 1999) and sect. Diploclinium (Dorrenbos et al., 1998)]

4 B. parvula H. Lév. & Vant.

5 B. wilsonii Gagnep. [previously assigned to sect. Begoniastrum (Irmscher, 1939) and sect. Diploclinium (Dorrenbos et al., 1998; Golding and Wasshausen, 2002)]

Botanical Bulletin of Academia Sinica, Vol. 43, 2002

Sect. 6. Parvibegonia A. DC., Ann. Sci. Nat. Bot. Ser. 4. 11: 136. 1859; Doorenbos et al., The Sections of Begonia 170. 1998.—LECTOTYPE SPECIES: Begonia martabanica A. DC. (designated by Barkley, 1972).

Herbs tuberous, rarely stemless, monoecious; leaves palmately veined. Inflorescence terminal, tepals less than 0.5 cm long, red; staminate flowers: tepals 4, filaments fused to a column, anthers yellow; carpellate flowers: tepals 4 or 5, styles 2 (-3), often persistent, stigma kidney or moon-shaped, ovary 2(-3)-locular. Placentae bifid at the middle and upper part of ovary, unknown at the base. Capsule suberect, with 3 unequal wings and papery walls, bursting in an irregular fashion (Figure 13).

Cytology. 2n = 56 (B. crenata Dryand.: Legro and Doorenbos, 1971).

Distribution and habitat. Twenty-nine species in SE Asia and E Asia as far as India. Only one species, B. discreta Craib., in China, on rocks in thickets in Xishuangbanna, southern Yunnan.

Species Found in China. Begonia discreta Craib. [previously assigned to sect. Diploclinium (Barkley and Baranov, 1972; Dorrenbos et al., 1998; Golding and Wasshausen, 2002); as sect. Begonia (Ku, 1999)]

Sect. 7. Platycentrum (Klotzsch) A. DC., Ann. Sci. Nat. Bot. Ser. 4. 11: 134. 1859; Yü, Bull. Fan. Mem. Inst. Biol. 1(2): 122. 1948; Irmscher, Notes Roy. Bot. Gard. Edinburgh 21: 39. 1951; Barkley, Buxtonian 1 (Suppl. 4): 12. 1972; Liou, Iconogr. Cormophyt. Sin. Suppl. 2. 533, 536. 1983; Wu and Ku, Acta Phytotax. Sin. 33: 265. 1995; Acta Phytotax. Sin. 35: 54. 1997; Dorrenbos et al., The Sections of Begonia 155. 1998; Shui and Huang, Acta Bot. Yunnan. 21: 16. 1999; Ku, Fl. Reipubl. Popularis Sin. 52: 129, 212. 1999.—Platycentrum Klotzsch, Abh. Königl. Akad. Wiss. Berlin `1854': 243. 1855; Begoniac. 123. 1855.—Sect. Parvibegonia auct non A. DC, as to B. houttuynioides Yü, Bull. Fan. Mem. Inst. Biol. 1(2): 125. 1948.—LECTOTYPE SPECIES: Begonia xanthina Hook. (designated by Barkley and Baranov, 1972)(ºPlatycentrum xanthinum (Hook.) Klotzsch).

Herbs or subshrubs, monoecious, terrestrial, seldom epiphytic, creeping or erect, with thin or thick rhizomes; leaves palmate-veined or pinnate-veined, unlobed or lobed, rarely palmate-foliolate (B. hemsleyana Hook. f.). Inflorescence terminal or axillary; tepals pink, rarely yellow (B. flaviflora H. Hara) or green (B. polytricha C.Y. Wu); staminate flowers: tepals 4, filaments free, anthers yellow, rarely red (B. purpureofolia S.H. Huang & Y.M. Shui, B. polytricha C.Y. Wu), connectives occasionally projecting; carpellate flowers: tepals 5, rarely 3 (consistent in B. austrotaiwanensis Y.K. Chen & C.I Peng), 4 (consistent in B. oreodoxa Chun & F. Chun ex C.Y. Wu & Ku) or 6(-10) [occasional in B. formosana (Hayata) Masam.], styles 2, rarely 3 (B. versicolor Irmsch.), not persistent, bifid, stigma of 2 or 3 coils. Ovary 2-locular. Placentae axile, bifid or multi-fid from apical to basical part of ovary, rarely parietal at upper part of ovary (B. oreodoxa Chun & F. Chun ex C.Y. Wu & Ku). Capsule nodding, wings 3, unequal, the largest of which

considerably elongating downwards, nearly always without ventral sutures between the large and smaller wings (Figures 14-15).

Cytology. 2n = 22 (B. rex Retz.; B. laciniata Roxb.: Sharma et al., 1960; B. rex Retz.: Tian et al., 2002; B. cathayana Hemsl., B. circumlobata Hance, B. hemsleyana Hook. f., B. palmata D. Don ("B. laciniata Roxb."), etc.: Legro et al., 1969; B. hemsleyana Hook.: Tian et al., 2002; B. versicolor Irmsch.: Legro et al., 1971, Tian et al., 2002; B. sikkimensis A. DC.: Legro et al., 1973; B. palmata D. Don.: Peng et al., 1991; Oginuma and Peng, 2002; B. palmata D.Don var. browingiana (Champ. ex Benth.) J. Golding & C. Kareg.: Tian et al., 2002, 20 (B.cathayana Hemsl.: Legro et al., 1969), 36 (B. austrotaiwanensis Y.K. Chen & C.I Peng: Peng and Chen, 1990; Oginuma and Peng, 2002), 38 (B. austrotaiwanensis Y.K. Chen & C.I Peng: Peng and Chen, 1990; Oginuma and Peng, 2002; B. chitoensis T.S. Liu & M.J. Lai: Oginuma and Peng, 2002), 44 (several cultivars of B. rex Retz.: Sharma et al., 1960; B. venusta Ridl.: Legro et al., 1971), 60 (B. formosana (Hayata) Masam.: Peng and Sue, 2000; Oginuma and Peng, 2002), 64 (B. formosana (Hayata) Masam.: Oginuma and Peng, 2002).

Distribution and habitat. About 110 species from India and central China to SE Asia. Sixty-three species in China, occurring S of Yangtze River and SE Tibet and in Taiwan. These species are either terrestrial on thick humus, epiphytic on trees, or epipetric on rock faces or crevices, at 100-2,200 m.

Species Found in China:

1 B. algaia L.B. Sm. & Wassh.

2 B. augustinei Hemsl. [previously assigned to sect. Begoniastrum (Yü, 1948)]

—B. menglianensis Y.Y. Qian, syn. nov.

A study of the protologue of B. menglianensis (Qian, 2001) immediately revealed the overall similarity between B. menglianensis and B. augustinei except that the former is more glabrous. However, we were able to examine an isotype of B. menglianensis, Y.Y. Qian 3110 (KUN), which showed that the upper surfaces of leaves are densely covered with short hairs. Thus we feel confident that B. menliangensis should be synonymized under B. augustinei.

3 B. austrotaiwanensis Y.K. Chen & C.I Peng

4 B. baviensis Gagnep.

5 B. brevisetulosa C.Y. Wu

6 B. cathayana Hemsl.

7 B. chishuiensis T.C. Ku

8 B. chitoensis T.S. Liu & M.J. Lai [previously assigned to sect. Begonia (Ku, 1999)]

9 B. chuniana C.Y. Wu [previously assigned to sect. Parvibegonia (Golding and Wasshausen, 2002)]

10 B. circumlobata Hance

11 B. cucurbitifolia C.Y. Wu

12 B. daweishanensis S.H. Huang & Y.M. Shui

Shui et al. — Synopsis of the Chinese species of Begonia

13 B. dielsiana E. Pritz.

14 B. digyna Irmsch.

15 B. discrepans Irmsch.

16 B. dryadis Irmsch.

17 B. duclouxii Gagnep. [previously assigned to sect. Reichenheimia (Yü, 1948)]

18 B. edulis H. Lév.

19 B. emeiensis C.M. Hu ex C.Y. Wu & T.C. Ku

20 B. flaviflora H. Hara

21 B. formosana (Hayata) Masam.

—B. tarokoensis M.J. Lai, syn. nov.

22 B. forrestii Irmsch.

23 B. gagnepainiana Irmsch.

24 B. gungshanensis C.Y. Wu

25 B. hatacoa Buch.-Ham. ex D. Don

26 B. hekouensis S.H. Huang

27 B. hemsleyana Hook. f.

28 B. houttuynioides T.T. Yü [previously assigned to sect. Parvibegonia (Yü, 1948; Barkley and Baranov, 1972)]

29 B. howii Merr. & Chun [previously assigned to sect. Diploclinium? (Dorrenbos et al., 1998; Golding and Wasshausen, 2002)]

30 B. kouytcheouensis Guillaumin [previously assigned to sect. Reichenheimia (Barkley and Baranov, 1972)]

31 B. lacerata Irmsch.

32 B. laminariae Irmsch. [previously assigned to sect. Begoniastrum (Liou, 1983)]

33 B. limprichtii Irmsch.

34 B. lipingensis Irmsch.

35 B. longanensis C.Y. Wu

36 B. longialata K.Y. Guan & D.K. Tian

37 B. macrotoma Irmsch.

38 B. maguanensis S.H. Huang & Y.M. Shui

39 B. manhaoensis S.H. Huang & Y.M. Shui

40 B. megalophyllaria C.Y. Wu

41 B. mengtzeana Irmsch.

42 B. nantoensis M.J. Lai & N.J. Chung

43 B. oreodoxa Chun & F. Chun

44 B. palmata D. Don

—B. randaiensis Sasaki (cf. Lai, 1979)

45 B. paucilobata C.Y. Wu

46 B. pedatifida H. Lév.

47 B. polytricha C.Y. Wu

48 B. psilophylla Irmsch.

49 B. purpureofolia S.H. Huang & Y.M. Shui

50 B. reflexisquamosa C.Y. Wu

51 B. repenticaulis Irmsch.

52 B. rex Putz.

53 B. rockii Irmsch.

54 B. rubropunctata S.H. Huang & Y.M. Shui

55 B. scitifolia Irmsch.

56 B. sikkimensis A. DC.

57 B. smithiana T.T. Yü ex Irmsch.

58 B. subhowii S.H. Huang

59 B. truncatiloba Irmsch.

60B. tsoongii C.Y. Wu

61 B. versicolor Irmsch.

62 B. villifolia Irmsch. [previously assigned to sect. Begoniastrum (Liou, 1983)]

63 B. yingjiangensis S.H. Huang [previously assigned to sect. Parvibegonia (Golding and Wasshausen, 2002)]

Sect. 8. Leprosae (T.C. Ku) Y.M. Shui, comb. nov.—Sect. Begonia Subsect. Crassirostris T.C. Ku Ser. Leprosae T.C. Ku, Fl. Reipubl. Popularis Sin. 52(1): 128, 144, 149, 401. 1999.—Sect. Parvibegonia auct. non A. DC, as to B. leprosa Hance, Yu, Bull. Fan. Mem. Inst. Biol. 1(2): 125. 1948.—Sect. Sphenanthera (Hassk.) Warb. pro parte, as to B. leprosae Hance: Irmscher, Mitt. Inst. Allg. Bot. Hamburg 10: 556. 1939; Yü, Bull. Fan. Mem. Inst. Biol. 1(2): 125. 1948; Liou, Iconogr. Cormophyt. Sin. Suppl. 2:536, 541. 1983, in clavi; Guan and Tian, Acta Bot. Yunnan. 22(2): 129, 2000.—Sect. Reichenheimia (Klotzsch) A. DC. Subsect. Wilsonianae T.C. Ku Ser. Cylindricae T.C. Ku, Fl. Reipubl. Popularis Sin. 52(1): 128, 197, 401. 1999, syn. nov.—TYPE SPECIES: Begonia leprosa Hance (1883).

Herbs monoecious, terrestrial, with slender creeping rhizomes, without erect stems, leaves broadly ovate, oblique at base, palmately nerved, glaberous on upper surface, tomentose beneath. Inflorescencecymose, shorter than petiole, tepals white, rarely bisexual (occasional in B. leprosa Hance); staminate flowers: tepals 4, filaments fused at base, anthers oblong and yellow; carpellate flowers: tepals 3-5, styles 3, bifid, stigma 2-coiled. Ovary 3-locular. Placentae axile and bifid from apical to basical part of ovary. Fruit not dehiscent, berry-like, pendulous, thick-walled, clavate, without horns or wings (Figures 16-17).

Sect. Leprosae (T.C. Ku) Y.M. Shui is similar to sect. Sphenanthera A. DC. in the berry-like fruit. Fruit of the latter, however, is turbinate and with horns or wings. Sect. Leprosae is also similar to sect. Mezierea (Goudich.) Warb. [in Africa] in fruit shape, but the latter has parietal or septal placentae (Doorenbos et al., 1998). It may be that sect. Leprosae is a transitional section between Asia and Africa.

Cytology. 2n = 20 (B. longicarpa K.Y. Guan & D.K. Tian: Tian et al., 2002)

Distribution and habitat. Three species in southern China, one of which also found in Vietnam (Tonkin). In China (Guangdong, Guangxi and Yunnan), on moist soil at river banks or on rocks nearby a limestone cave in tropical forests, at 30-300 m altitude.

Botanical Bulletin of Academia Sinica, Vol. 43, 2002

Species Found in China:

1 B. cylindrica D.R. Liang & X.X. Chen [previously assigned to sect. Reichenheimia (Ku, 1999)]

2 B. leprosa Hance [previously assigned to sect. Sphenanthera (Irmscher, 1939; Barkley and Baranov, 1972), sect. Parvibegonia A. DC and sect. Begonia (Ku, 1999)]

—B. bretschneideriana Hemsl. [previously assigned to sect. Sphenanthera (Irmscher, 1939)]

3 B. longicarpa K.Y. Guan & D.K. Tian [previously assigned to sect. Sphenanthera (Guan and Tian, 2000)]

Sect. 9. Sphenanthera (Hassk.) Warb. in Engler and Prantl, Nat. Pflanzenfam. 3(6a): 138, 141. 1894; Yü, Bull. Fan. Mem. Inst. Biol. 1(2): 114. 1948; Liou, Iconogr. Cormophyt. Sin. Suppl. 2: 533, 535. 1983; Doorenbos et al., The Sections of Begonia 194. 1998; Shui and Huang, Acta Bot. Yunnan. 21(1): 11,1999; Ku, Fl. Reipubl. Popularis Sin. 52(1): 128, 206. 1999.—Sect. Pleiothece T.C. Ku, Fl. Reipubl. Popularis Sin. 52(1): 129, 211, 401. 1999, syn. nov. —Sphenanthera Hassk., Versl. Kon. Akad. Wetensch. 4: 139. 1855.—LECTOTYPE SPECIES: Begonia robusta Blume (designated by Dorrenbos et al., 1998) (º Sphenanthera robusta (Blume) Hassk. ex Klotzsch ).

Herbs or subshrubs, terrestrial, monoecious with 3-locular ovary or dioecious with 4 or more locules; stems creeping, or erect with short and thick rhizomes; leaves auriculate, very obliquely palmately veined. Inflorescence axillary, short and congested; tepals white or pink; flowers of the dioecious species fragrant; staminate flowers: tepals 4, sometimes to 6 cm long, anthers yellow; carpellate flowers: tepals 4 or 5, styles 3-7, deciduous, free, deeply bifid, stigma forming a continual helical band with multiple coils. Ovary 3-7- locular (often variable within the same individual). Placentae bifid from bottom to apex of fruit, rarely unclefted at base. Fruits pendulous or recurved, berry-like, turbinate, thick-walled, the walls wingless or with obscure wings or horns, bursting very late or irregularly after falling (Figures 18-21).

Cytology. 2n = 20 (B. ceratocarpa S.H. Huang & Y.M. Shui: Tian et al., 2002), 22 (B. inflata C.B. Clarke: Legro and Dorrenbos, 1969; B. roxburghii A. DC.: Legro and Dorrenbos, 1971; B. hayatae Gagnep.: "B. aptera Blume", Peng and Su, 2000, Oginuma and Peng, 2002; B. silletensis subsp. mengyangensis M.C. Tebbitt & K.Y. Guan: "B. mengyangensis subsp. mengyangensis M.C. Tebbitt & K.Y. Guan", Tian et al., 2002), and 88 (possibly in B. robusta Blume: Dorrenbos et al., 1998).

Distribution and habitat. About 27 species from India to Indochina as far as Malaysia, Indonesia. Eleven species in China, distributed mainly in tropical valleys of Taiwan, Fujian, Guangdong, Hainan, Guangxi, Guizhou, Yunnan and Tibet, usually found on deep humus soil at forest margin, at 60-1,300 m altitude.

Species Found in China:

1 B. acetosella Craib

2 B. balansana Gagnep. [previously assigned to sect. Pleiothece T.C. Ku (Ku, 1999)]

—B. handelii Irmsch.

—B. xinyiensis T.C. Ku [previously assigned to sect. Begonia (Wu and Ku, 1995; Ku, 1999); sect. Diploclinium? (Dorrenbos et al., 1998; Golding and Wasshausen, 2002)], syn. nov.

The type (Z. Huang 31748, PE) of B. xinyiensis is a plant with a staminate inflorescence. No additional specimens were available for examination by Wu and Ku (1995). Thus, important characters such as female flowers and fruits were not described in the protologue of B. xinyiensis. However, diagnostic characters of B. xinyiensis include a unisexual inflorescence, ovate bracts and elongate, prostrate rhizomes, which is characteristic of B. balansana. Furthermore, type of B. xinyiensis was collected from southern Guangdong, which is within the range of widespread B. balansana. In our opinion, the two species are conspecific.

3 B. ceratocarpa S.H. Huang & Y.M. Shui

4 B. crassirostris Irmsch. [previously assigned to sect. Begonia (Ku, 1999)]

5 B. hayatae Gagnep. [previously assigned to sect. Begonia (Ku, 1999)]

6 B. inflata C.B. Clarke

7 B. lancangensis S.H. Huang

8 B. prostrata Irmsch.

9 B. silletensis (A. DC.) C.B. Clarke ssp. mengyangensis M.C. Tebbitt & K.Y. Guan (Tebbitt and Guan, 2002)

10 B. tessaricarpa C. B. Clarke

The species, previously known from India (Assam), is a new record for China. It is documented here for the first time. China. Xizang, Medog County, Genbala hill, under forests, herb, 650 m, with fruits, 28 Nov 1992, ETM 1720 ( KUN 0372087, KUN 0372086).

11 B. tetragona Irmsch.

Natural Hybrids Between Members of Different Sections

1. Between Members of sect. Platycentrum and sect. Diploclinium.

Begonia ×buimontana Yamamoto, assigned to sect. Begonia (Ku, 1999).

The hybridity and parentage of B. ×buimontana Yamamoto were documented thoroughly by Peng and Chen (1991). Detailed analyses of morphology, flowering habit, pollen stainability, chromosome number, meiotic chromosome behavior and geographical distribution suggest that B. ×buimontana Yamamoto represents a natural hybrid between B. palmata D. Don [sect. Platycentrum] and B. taiwaniana Hayata [sect. Diploclinium] in Taiwan.

2. Between members of sect. Platycentrum and sect. Sphenanthera.

Shui et al. — Synopsis of the Chinese species of Begonia

Begonia ×taipeiensis C.I Peng (Peng and Sue, 2000). Based on a comparison of morphology, geographical distribution, pollen stainability, seed set, cytological observation, and experimental crosses, Peng and Su (2000) concluded that B. ×taipeiensis represents F₁ progeny from natural hybridization between B. formosana (Hayata) Masam. [sect. Platycentrum] and B. hayatae Gagnep. ["B. aptera Blume", sect. Sphenanthera] in Taiwan. Data from experimental crosses and molecular studies further confirmed unidirectional hybridization in B. ×taipeiensis, i.e., formation of the natural hybrid B. ×taipeiensis occurs via pollen transfer from B. aptera to the maternal species, B. formosana (Peng and Chiang, 2000; Chiang et al., 2001).

Relationships of the Sections of Chinese Begonia

Section Coelocentrum is likely to be the most ancient group among Asian Begonia because of its parietal placentae (Reitsma, 1983). Section Coelocentrum is closely related to some species in sect. Diploclinium, such as Begonia wangii T.T. Yü and B. peltatifolia H.L. Li. Members of sect. Diploclinium have parietal placentae near the apex of the ovary (Figure 5B). Members of both sections grow on rocks in limestone areas, which provide a refuge to many seed plants (Ying et al., 1993; Shui et al., 1999) in China (SE Yunnan, S Guizhou, SE Guangxi) and Vietnam (Tonkin). Plants of Begonia sect. Coelocentrum may well be a group in this refuge with ancient characters. By way of prolongation and fusion of placentae from the middle of ovary to top (Gauthier, 1950, 1957; Jin and Wang, 1994), sect. Diploclinium seems to have evolved from sect. Coelocentrum into a large group in Asia.

Sect. Diploclinium, a complex group, is related to many other sections. Sect. Platycentrum has apparently derived from sect. Diploclinium by the complete degeneration of one of the three ovarian cells. Transitional species include Begonia versicolor Irmsch., with 3 stigmas, and B. obsolescens Irmsch. as well as B. gulinqingensis S.H. Huang & Y.M. Shui, which has one degenerating ovarian cell (Shui and Huang, 1999). Plants of sect. Platycentrum have speciated profusely in tropical and subtropical mountains. Sect. Reichenheimia is probably evolved from a tuberous and stemless group of sect. Diploclinium through the fusion of placentae. Begonia labordei H. Lév. and B. guishanensis S.H. Huang & Y.M. Shui (Huang and Shui, 1994) are representative of transitional species in being deciduous and having tuberous stem-base, characters of sect. Reichenheimia that adapt to seasonally dry or cold areas. Sections Petermannia, Parvibegonia and Alicida are probably also related to sect. Diploclinium with bifid axile placentae, but the number of species of the three sections in China are not sufficient to reveal their relationship with plants of sect. Diploclinium. Further studies may reveal the need of dividing sect. Diploclinium and sect. Petermannia, two large and divergent sections in Asia (Doorenbos et al., 1998).

Sect. Sphenanthera is a peculiar group. The fragrant tepals, unisexual inflorescence and berry-liked fruits sug

gest that plants of this section have evolved into an entomochoric and entomophilous group. Sect. Leprosae is similar to sect. Sphenanthera in having berry-like fruits, but the occasional occurrence of bisexal flowers in B. leprosa (sect. Leprosae) (Hance, 1883) implies its primitiveness. It is possible that sect. Leprosae represents an intermediate section between sections Sphenanthera and Diploclinium.

Alphabetical List of Chinese Begonia Species with Sectional Assignment

The Chinese species of Begonia are listed below with reference to their sectional placement. Asterisks (*) indicate names accepted in the account of Chinese Begonia by Ku (1999a) that we place in synonymy with species to the right of the equal (=) sign.

1 B. acetosella Craib, sect. Sphenanthera

2 B. acutitepala K.Y. Guan & D.K. Tian, sect. Diploclinium

3 B. algaia L.B. Sm. & Wassh., sect. Platycentrum

4 B. alveolata T.T. Yü, sect. Diploclinium

*B. anceps Irmsch. = B. morifolia T. T. Yü

5 B. arboreta Y. M. Shui, sect. Diploclinium (Shui, 2002a)

6 B. asperifolia Irmsch., sect. Diploclinium

7 B. augustinei Hemsl., sect. Platycentrum

8 B. austrotaiwanensis Y.K. Chen & C.I Peng, sect. Platycentrum

9 B. balansana Gagnep., sect. Sphenanthera

10 B. baviensis Gagnep., sect. Platycentrum

11 B. biflora T.C. Ku, sect. Coelocentrum

12 B. bonii Gagnep., sect. Coelocentrum

13 B. brevisetulosa C.Y. Wu, sect. Platycentrum

14 B. ×buimontana Yamam. = B. palmata D. Don [sect. Platycentrum] × B. taiwaniana Hayata [sect. Diploclinium]

15 B. cathayana Hemsl., sect. Platycentrum

16 B. cavaleriei H. Lév., sect. Diploclinium

17 B. cehengensis T.C. Ku, sect. Diploclinium

18 B. ceratocarpa S.H. Huang & Y.M. Shui, sect. Sphenanthera

19 B. chingii Irmsch., sect. Reichenheimia

20 B. chishuiensis T.C. Ku, sect. Platycentrum

21 B. chitoensis T.S. Liu & M.J. Lai, sect. Platycentrum

22 B. chuniana C.Y. Wu, sect. Platycentrum

23 B. circumlobata Hance, sect. Platycentrum

24 B. cirrosa L.B. Sm. & Wassh., sect. Coelocentrum

25 B. clavicaulis Irmsch., sect. Diploclinium

26 B. coptidimontana C.Y. Wu, sect. Diploclinium

27 B. crassirostris Irmsch. sect. Sphenanthera

28 B. cucurbitifolia C.Y. Wu, sect. Platycentrum

Botanical Bulletin of Academia Sinica, Vol. 43, 2002

29 B. cylindrica D.R. Liang & X.X. Chen, sect. Leprosae

30 B. daweishanensis S.H. Huang & Y.M. Shui, sect. Platycentrum

31 B. daxinensis T.C. Ku, sect. Coelocentrum

32 B. dentato-bracteata C.Y. Wu, sect. Diploclinium

33 B. dielsiana E. Pritz., sect. Platycentrum

34 B. digyna Irmsch., sect. Platycentrum

35 B. discrepans Irmsch., sect. Platycentrum

36 B. discreta Craib, sect. Parvibegonia

37 B. dryadis Irmsch., sect. Platycentrum

38 B. duclouxii Gagnep., sect. Platycentrum

39 B. edulis H. Lév., sect. Platycentrum

40 B. emeiensis C. M. Hu, sect. Platycentrum

*B. fengii T. C. Ku = B. obsolescens Irmsch.

41 B. fenicis Merr., sect. Diploclinium

42 B. filiformis Irmsch., sect. Coelocentrum

43 B. fimbristipula Hance, sect. Diploclinium

44 B. flaviflora H. Hara, sect. Platycentrum

*B. floribunda T.C. Ku = B. sinofloribunda Dorr

45 B. fordii Irmsch., sect. Diploclinium

46 B. formosana (Hayata) Masam., sect. Platycentrum

47 B. forrestii Irmsch., sect. Platycentrum

48 B. gagnepainiana Irmsch., sect. Platycentrum

49 B. glechomifolia C.M. Hu ex C.Y. Wu & T.C. Ku, sect. Diploclinium

50 B. grandis Dryand., sect. Diploclinium

51 B. guangxiensis C.Y. Wu, sect. Coelocentrum

52 B. guishanensis S.H. Huang & Y.M. Shui, sect. Diploclinium

53 B. gulinqingensis S.H. Huang & Y.M. Shui, sect. Diploclinium

54 B. gungshanensis C.Y. Wu, sect. Platycentrum

55 B. hainanensis Chun & F. Chun, sect. Petermmania

*B. handelii Irmsch. = B. balansana Gagnep.

56 B. hatacoa Buch.-Ham. ex D. Don, sect. Platycentrum

57 B. hayatae Gagnep., sect. Sphenanthera

58 B. hekouensis S.H. Huang, sect. Platycentrum

59 B. hemsleyana Hook. f., sect. Platycentrum

60 B. henryi Hemsl., sect. Reichenheimia

61 B. houttuynioides Yü, sect. Platycentrum

62 B. howii Merr., sect. Platycentrum

63 B. hymenocarpa C. Y. Wu, sect. Diploclinium

64 B. imitana Irmsch., sect. Diploclinium

65 B. inflata C. B. Clarke, sect. Sphenanthera

66 B. josephii A. DC., sect. Diploclinium

67 B. kouy-tcheouensis Guillaumin, sect. Platycentrum

68 B. labordei H. Lév., sect. Diploclinium

69 B. lacerata Irmsch., sect. Platycentrum

70 B. laminariae Irmsch., sect. Platycentrum

71 B. lancangensis S.H. Huang, sect. Sphenanthera

72 B. lanternaria Irmsch., sect. Coelocentrum

73 B. leprosa Hance, sect. Leprosae

74 B. limprichtii Irmsch., sect. Platycentrum

75 B. lipingensis Irmsch., sect. Platycentrum

76 B. lithophila C.Y. Wu, sect. Reichenhaimia

77 B. longanensis C.Y. Wu, sect. Platycentrum

78 B. longialata K.Y. Guan & D. K. Tian, sect. Platycentrum

79 B. longicarpa K.Y. Guan & D. K. Tian, sect. Leprosae

*B. longiciliata C.Y. Wu = B. rex Putz.

80 B. lukuana Y.C. Liu & C.H. Ou, sect. Diploclinium

81 B. luzhaiensis T.C. Ku, sect. Coelocentrum

82 B. macrotama Irmsch., sect. Platycentrum

83 B. maguanensis S.H. Huang & Y.M. Shui, sect. Platycentrum

84 B. malipoensis S.H. Huang & Y.M. Shui, sect. Diploclinium

85 B. manhaoensis S.H. Huang & Y.M. Shui, sect. Platycentrum

86 B. masoniana Irmsch., sect. Coelocentrum

87 B. megalophyllaria C.Y. Wu, sect. Platycentrum

*B. menglianensis Y.Y. Qian = B. augustinei Hemsl.

88 B. mengtzeana Irmsch., sect. Platycentrum

89 B. miranda Irmsch., sect. Diploclinium

90 B. morifolia T.T. Yü, sect. Diploclinium

91 B. morsei Irmsch., sect. Coelocentrum

92 B. muliensis T.T. Yü, sect. Diploclinium

93 B. nantoensis M.J. Lai & N.J. Chung, sect. Platycentrum

94 B. nymphaeifolia T.T. Yü, sect. Diploclinium

95 B. obliquefolia S.H. Huang & Y.M. Shui, sect. Coelocentrum

96 B. obsolescens Irmsch., sect. Diploclinium

97 B. oreodoxa Chun & F. Chun ex C.Y. Wu & T.C. Ku, sect. Platycentrum

98 B. ornithophylla Irmsch., sect. Coelocentrum

99 B. palmata D. Don, sect. Platycentrum

100 B. parvula H. Lév. & Vant., sect. Reichenheimia

101 B. paucilobata C.Y. Wu, sect. Platycentrum

102 B. pedatifida H. Lév., sect. Platycentrum

103 B. péii C.Y. Wu, sect. Alicida

104 B. peltatifolia H.L. Li, sect. Diploclinium

105 B. picta Sm., sect. Diploclinium

*B. pingbienensis C.Y. Wu = B. alveolata T.T. Yü

106 B. polytricha C.Y. Wu, sect. Platycentrum

107 B. porteri H. Lév. & Vaniot, sect. Coelocentrum

108 B. prostrata Irmsch., sect. Sphenanthera

Shui et al. — Synopsis of the Chinese species of Begonia

109 B. pseudodryadis C.Y. Wu, sect. Coelocentrum

110 B. psilophylla Irmsch., sect. Platycentrum

111 B. purpureofolia S.H. Huang & Y.M. Shui, sect. Platycentrum

*B. randaiensis Sasaki = B. palmata D. Don

112 B. ravenii C.I Peng & Y.K. Chen, sect. Diploclinium

113 B. reflexisquamosa C.Y. Wu, sect. Platycentrum

114 B. repenticaulis Irmsch., sect. Platycentrum

115 B. rex Putz., sect. Platycentrum

*B. rhodophylla C.Y. Wu = B. guishanensis S.H. Huang & Y.M. Shui

116 B. rockii Irmsch., sect. Platycentrum

117 B. rongjiangensis T.C. Ku, sect. Diploclinium

118 B. rotundilimba S.H. Huang & Y.M. Shui, sect. Diploclinium

119 B. ruboides C.M. Hu ex C.Y. Wu & T.C. Ku, sect. Diploclinium

120 B. rubropunctata S.H. Huang & Y.M. Shui, sect. Platycentrum

121 B. scitifolia Irmsch., sect. Platycentrum

122 B. setifolia Irmsch., sect. Diploclinium

123 B. setuloso-peltata C.Y. Wu, sect. Coelocentrum

124 B. sikkimensis A. DC., sect. Platycentrum

125 B. silletensis (A. DC.) C.B. Clarke subsp. mengyangensis M.C. Tebbitt & K.Y. Guan, sect. Sphenanthera (Tebbit and Guan, 2002)

*B. sinobrevicaulis T.C. Ku = B. gulinqingensis S.H. Huang & Y.M. Shui

126 B. sinofloribunda Dorr, sect. Diploclinium

127 B. sino-vietnamica C. Y. Wu, sect. Diploclinium

128 B. smithiana T. T. Yü ex Irmsch., sect. Platycentrum

129 B. subhowii S. H. Huang, sect. Platycentrum

130 B. summoglabra T. T. Yü, sect. Diploclinium

131 B. ×taipeiensis C.I Peng = B. formosana (Hayata) Masam. [sect. Platycentrum] × B. hayatae Gagnep. ["B. aptera Blume", sect. Sphenanthera] (Peng and Su, 2000)

132 B. taiwaniana Hayata, sect. Diploclinium

133 B. taliensis Gagnep., sect. Diploclinium

*B. tarokoensis M.J. Lai = B. formosana (Hayata) Masam.

134 B. tessaricarpa C.B. Clarke, sect. Sphenanthera

135B. tetragona Irmsch., sect. Sphenanthera

136 B. truncatiloba Irmsch., sect. Platycentrum

*B. tsaii Irmsch. = B. setifolia Irmsch.

137B. tsoongii C.Y. Wu, sect. Platycentrum

138 B. umbraculifolia Y. Wan & B.N. Chang, sect. Coelocentrum

139 B. versicolor Irmsch., sect. Platycentrum

140 B. villifolia Irmsch., sect. Platycentrum

141 B. wangii T.T. Yü, sect. Diploclinium

142 B. wenshanensis C.M. Hu ex C.Y. Wu & T.C. Ku, sect. Diploclinium

143 B. wilsonii Gagnep., sect. Reichenheimia

144 B. xingyiensis T.C. Ku, sect. Diploclinium

*B. xinyiensis T.C. Ku = B. balansana Gagnep.

145 B. xishuiensis T.C. Ku, sect. Diploclinium

146 B. yingjiangensis S.H. Huang, sect. Platycentrum

147 B. yishanensis T.C. Ku, sect. Coelocentrum

148 B. yui Irmsch., "yuii", sect. Diploclinium

149 B. yunnanensis H. Lév., sect. Diploclinium

150 B. zhengyiana Y.M. Shui, sect. Coelocentrum (Shui, 2002b)

Acknowledgments. We thank Professors Su-Hua Huang (YUNU) and De-Zhu Li (KUN) for helpful suggestions oo the manuscript, and Dr. Hai-Ning Qin (PE) and Professor Kai-Yun Guan for providing useful literature. We are indebted to Ms. Wen-Hong Chen and Hui-Chun Hsiau for technical assistance and Miss Ling Wang for the handsome illustration. The authors are grateful for detailed reviews of the manuscript by Drs. David E. Boufford and Thomas G. Lammers.This study was supported in part by research grants from National Science Council and Academia Sinica, Taiwan, to Ching-I Peng.

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