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Saxifragaceae Flora of China 8: 269–452. 2001. SAXIFRAGACEAE 虎耳草科 hu er cao ke Pan Jintang (潘锦堂)1, Gu Cuizhi (谷粹芝 Ku Tsue-chih)2, Huang Shumei (黄淑美 Hwang Shu-mei)3, Wei Zhaofen (卫兆芬 Wei Chao-fen)4, Jin Shuying (靳淑英)5, Lu Lingdi (陆玲娣 Lu Ling-ti)6; Shinobu Akiyama7, Crinan Alexander8, Bruce Bartholomew9, James Cullen10, Richard J. Gornall11, Ulla-Maj Hultgård12, Hideaki Ohba13, Douglas E. Soltis14 Herbs or shrubs, rarely trees or vines. Leaves simple or compound, usually alternate or opposite, usually exstipulate. Flowers usually in cymes, panicles, or racemes, rarely solitary, usually bisexual, rarely unisexual, hypogynous or ± epigynous, rarely perigynous, usually biperianthial, rarely monochlamydeous, actinomorphic, rarely zygomorphic, 4- or 5(–10)-merous. Sepals sometimes petal-like. Petals usually free, sometimes absent. Stamens (4 or)5–10 or many; filaments free; anthers 2-loculed; staminodes often present. Carpels 2, rarely 3–5(–10), usually ± connate; ovary superior or semi-inferior to inferior, 2- or 3–5(–10)-loculed with axile placentation, or 1-loculed with parietal placentation, rarely with apical placentation; ovules usually many, 2- to many seriate, crassinucellate or tenuinucellate, sometimes with transitional forms; integument 1- or 2-seriate; styles free or ± connate. Fruit a capsule or berry, rarely a follicle or drupe. Seeds albuminous, rarely not so; albumen of cellular type, rarely of nuclear type; embryo small. About 80 genera and 1200 species: worldwide; 29 genera (two endemic), and 545 species (354 endemic, seven introduced) in China. During the past several years, cladistic analyses of morphological, chemical, and DNA data have made it clear that the recognition of the Saxifragaceae sensu lato (Engler, Nat. Pflanzenfam. 18a: 74–226. 1928) is untenable. Among the angiosperm families, Saxifragaceae sensu lato may in fact represent the most extreme example of a polyphyletic assemblage. For example, recent analyses of DNA sequence data indicate that these taxa represent at least ten separate evolutionary lines, many of which are only distantly related to one another (Morgan & Soltis, Ann. Missouri Bot. Gard. 80: 631–660. 1993; Soltis & Soltis, Amer. J. Bot. 84: 504–522. 1997). Furthermore, very large molecular phylogenetic analyses of hundreds of angiosperms indicate that these separate lineages are distributed among four of the six traditionally recognized subclasses of dicotyledons (Savolainen et al., Syst. Biol. 49: 306–362. 2000; Soltis et al., Nature 402: 402–404. 1999). These recent studies have also greatly clarified how this phylogenetically diverse assemblage should be divided into families and treated taxonomically (see The Angiosperm Phylogeny Group (APG), Ann. Missouri Bot. Gard. 85: 531–553. 1998). Recent studies of DNA sequence data have clarified both the circumscription and affinities of a narrowly defined Saxifragaceae (Saxifragaceae sensu stricto) and Hydrangeaceae (Soltis et al., Amer. J. Bot. 82: 504–514. 1995; Savolainen et al., loc. cit.; Soltis et al., loc. cit. 1999). Saxifragaceae sensu stricto should consist only of Saxifragoideae, a group of about 30 herbaceous genera. Members of Saxifragaceae sensu stricto from the Chinese flora include Astilbe, Astilboides, Bergenia, Chrysosplenium, Mitella, Mukdenia, Oresitrophe, Rodgersia, Saxifraga, Tanakaea, Tiarella, and the recently described Saniculiphyllum. Close relatives of Saxifragaceae sensu stricto include Itea, Penthorum, and Ribes. These genera, the sole members of Iteoideae, Penthoroideae, and Ribesioideae, respectively, are also best treated in separate families: Iteaceae, Penthoraceae, and Grossulariaceae (see APG, loc. cit.). These taxa, as well as several others, such as Crassulaceae, are basal to a large assemblage of taxa, most of which were traditionally placed in Rosidae. Sequence data also indicate that Parnassia (the sole member of the Parnassioideae) is a more derived member of the rosid alliance, most closely related to Brexia and Lepuropetalon (also part of Saxifragaceae sensu lato) and Celastraceae. Parnassia and Lepuropetalon should be placed in Parnassiaceae with Brexia part of an expanded Celastraceae (APG, loc. cit.). Both morphological and molecular data indicate that Hydrangeoideae and Escallonioideae are, in contrast, allied with taxa traditionally placed in Asteridae. Hydrangeoideae are a well-defined, monophyletic lineage that should be treated as Hydrangeaceae. In China they include Cardiandra, Decumaria, Deinanthe, Deutzia, Dichroa, Hydrangea, Kirengeshoma, Philadelphus, Pileostegia, Platycrater, and Schizophragma, and are closely allied with families such as Cornaceae, Loasaceae, and Nyssaceae. Escallonioideae appear to be polyphyletic, and this group of approximately 14 genera is in need of thorough study. Members of this subfamily are allied with several different lineages of higher asterids. Polyosma, the only member of Escallonioideae in China, appears closely allied with Caprifoliaceae (Xiang & Soltis in Boufford & Ohba, Sino-Japanese Flora: its Characteristics and Diversification, 1998). Nevertheless, in the present account, the Saxifragaceae are retained in the sense of FRPS, using the same sequence of genera and indicating the subfamilies in the key below, in order to facilitate comparison with that flora. 1 Herbarium, North-western Plateau Institute of Biology, Chinese Academy of Sciences, 57 Xiquan Street, Xining, Qinghai 810001, People’s Republic of China. 2 Herbarium, Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China. 3 Herbarium, Department of Taxonomy, South China Institute of Botany, Chinese Academy of Sciences, Wushan, Guangzhou, Guangdong 510650, People’s Republic of China. 4 Herbarium, Department of Taxonomy, South China Institute of Botany, Chinese Academy of Sciences, Wushan, Guangzhou, Guangdong 510650, People’s Republic of China. 5 Herbarium, Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China. 6 Herbarium, Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China. 7 Department of Botany, National Science Museum, Tsukuba, Ibaraki, 305-0005, Japan. 8 Herbarium, Royal Botanic Garden Edinburgh, 20a Inverleith Row, Edinburgh EH3 5LR, Scotland, United Kingdom. 9 Botany Department, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118-4599, U.S.A. 10 Stanley Smith (UK) Horticultural Trust, Cory Lodge, P.O. Box 365, Cambridge CB2 1HR, England, United Kingdom. 11 Biology Department, University of Leicester, Leicester LE1 7RH, England, United Kingdom. 12 Museum of Evolution, Botany Section (Fytoteket), Uppsala University, Norbyvägen 16, SE-752 36, Uppsala, Sweden. 13 University Museum, University of Tokyo, Hongo 7-3-1, Tokyo 113-0033, Japan. 14 Ownbey Herbarium, Department of Biology, Washington State University, Pullman, Washington 99164-4309, U.S.A. Flora of China 8: 269–452. 2001. The genus Changiodendron R. H. Miao (Acta Sci. Nat. Univ. Sunyatseni 34: 65. 1995) and its single species, C. guangxiense R. H. Miao (loc. cit.: 66), was described from Guangxi (Napo Xian) and stated to belong to the Iteaceae (i.e., Iteoideae). However, Peng (Acta Bot. Yunnan. 18: 299– 300. 1996) demonstrated that C. guangxiense is a synonym of Sabia parviflora Wallich (Sabiaceae). Pan Jin-tang. 1992. Saxifragaceae (1) [Penthoroideae, Saxifragoideae]. In: Pan Jin-tang, ed., Fl. Reipubl. Popularis Sin. 34(2): 1–309; Hwang Shu-mei, Wei Chao-fen, Lu Ling-ti, Ku Tsue-chih & Jin Shu-ying. 1995. Saxifragaceae (2) [Parnassioideae, Hydrangeoideae, Escallonioideae, Iteoideae, Ribesioideae]. In: Lu Ling-ti & Hwang Shu-mei, eds., Fl. Reipubl. Popularis Sin. 35(1): 1–406. 1a. Plants woody or herbs; leaves opposite or alternate, rarely subverticillate or cespitose. 2a. Stamens as many as sepals; leaves usually alternate, opposite or subopposite in Polyosma. 3a. Leaves opposite or subopposite; fruit a 1-seeded berry (Escallonioideae) ............................. 27. Polyosma 3b. Leaves alternate; fruit a capsule or many-seeded berry. 4a. Leaves never lobed; stipules present, linear; sepals not petal-like; petals narrow; ovary 2- loculed; fruit a capsule (Iteoideae) .............................................................................................. 28. Itea 4b. Leaves often palmately lobed; stipules absent, spines or prickles sometimes present; sepals usually petal-like; petals usually scalelike; ovary 1-loculed; fruit a berry (Ribesioideae) ....... 29. Ribes 2b. Stamens ca. 2 × as many as sepals, sometimes more; leaves usually opposite or verticillate, alternate or fascicled in Cardiandra (Hydrangeoideae). 5a. Herbs; leaf blade palmately lobed; stamens 3 × as many as petals .................................. 16. Kirengeshoma 5b. Shrubs, scandent shrubs, or herbs; leaf blade not palmately lobed; stamens numerous or 2 × as many as petals. 6a. Filaments subulate, flat, apex 2-dentate or 2-lobed; shrubs; calyx lobes never enlarged and petaloid. 7a. Leaves often stellate hairy; petals 5; stamens 10(–15); capsule 3–5-valved, loculicidally dehiscent ........................................................................................................................ 17. Deutzia 7b. Leaves not stellate hairy; petals 4; stamens 20–40; capsule 4-valved, loculicidally or septicidally dehiscent ............................................................................................. 18. Philadelphus 6b. Filaments
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