47 (5) : , Fei, Ye and Li. Maeda et al., 1990 ; Matsui, 1994 ;, and Yu) O. k ; Fei et al., 2001), Hu) (, 1961) (Cladistics) PAU P

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1 47 (5) : , 2001 A cta Zoologica S inica 3 ( : ) (, ), PAU P , 1, 4 ; ;, ;, ;, ;, ; O dorrana Ranidae, 21, 16, 5 (, 1990 ; Maeda et al., 1990 ; Matsui, 1994 ;, and Yu) O. k uangw uensis ( Liu and 1999 ; Fei et al., 2001), Hu) O. livida (Blyth) O., l ungshengensis (Liu and Hu) O. m argare2, tae (Liu) O. nasuta Fei, Ye and Li O. schm ackeri (Boettger ) O. (, 1961) sw inhoana (Boulenger) O. tiannanensis, 7612 %, ( Yang and Li) O. versabilis (Liu and Hu) O. w uchuangensis (Xu) ; (Cladistics) PAU P ( Rana japanica Guenther R. omei montis Ye and Fei), (Liem, 1970 ;, 18 (, taxa), 60, ; Emerson et al., 1993 ; Macey et al., 1998) ; Darwin (1859) 112,, (, 2000) , ( ), (outgroup analy2 sis) (polarity), , :, O dorrana andersonii (Boulenger), (Maddison et al., 1984) O. anl ungensis (Liu and Hu) PAU P311 ( ), O. exiliversabilis Fei, Ye and Li O., Kluge (1969), , grahami (Boulenger) O. hai nanensis Fei, Ye and Li O. hejiangensis ( Deng 1,, 9 ( Emerson et al., 1993) (globally parsimonious method), 3 (No ),, 60, :

2 5 : ( : ) Table 1 Matrix of characters for 16 species of Odorrana and 2 species of Rana 529 Taxon R. japanica Characters and character states Plesio2morphy Apo2morphy Missing characters R. omei montis O. margaratae O. grahami O. andersonii O. hai nanensis O. kuangw uensis O. w uchuanensis O. hejiangensis O. tiannanensis O. anl ungensis O. schmackeri O. l ungshengensis O. swinhoana O. livi da O. exiliversabilis O. versabilis O. nasuta : (Character states coded with 0 are plesiomorphic states) apomorphic states) 9 : (Character states coded with 9 are missing value) 1, 2 : ( Character states coded with 1 or 2 are, 211 ( SWAP = TBR) MUL2 PARS, 71 63, 2 (the consistency index, CI) 01507, 1

3 sis; 1 16 Fig11 Phylogenetic relationships among the 16 species of Odorrana in China, 2 O. hainanensis, 3 O., 16 m argaretae O. grahami O. andersonii,,, : ; ; ; ; 113 1,,,, ; ( ) ; 3 4 ;,, 1,, (, 1990 ;, 1993), O. grahami O. andersonii,, 212 ;, 16 4, 4 ( ) : (1) : O dorrana m argaretae O. O. hainanensis 3 andersonii O. grahami O. hainanensis ; (2), O. andersonii : O. kuangw uensis, O. w uchuanen2,, (3) : O. hejiangensis O. tianna2 O. hainanensis ( ),, nensis O. anlungensis O. schmackeri O. l ung2,, shengensis O. sw inhoana ; (4) : 3 ( 1) O. livida O. exiliversabilis O. versabilis O. O. hainanensis nasuta,

4 5 : ( : ) ( O., O. nasuta O. versabilis 2 kuangw uensis O. w uchuanensis ),,,,,,,,,,,, ; O. nasuta,,,, :, 1, 4, 3, O. kuangw uensis ( ), O. w uchuanensis 311,, 21, %,, ( O. hejiangensis 15 %, O. tiannanensis O. anlungensis O. schmackeri O. l ungshengensis O. sw inhoana),,,,, :,,, E, 4 33 N,, O. hejiangensis, O. sw in2 hoana, 16,, 3 ( ),, ( O., tiannanensis O. anl ungensis),,, : ; ( O. schm ackeri O. l ungshengensis),, m argaretae; ; O. sw inhoana, kuangw uensis ;, l ungensis2l ungshengensis2hejiangensis2schm ackeri ;,,, : nasuta2versabilis2ex iliversabilis2 ( O. livida O. exiliv2,,, 312 hai nanensis2andersonii2graham i2 : w uchuanensis2 : tiannanensis2an2 livida ( 2), : andersonii 2 ersabilis O. versabilis O. nasuta), kuangw uensis 2schm ackeri 2livida :, 3,,, O. livida,,,,,,,,, 3,, 313,, 1,,, 3 1,, O. exiliversabilis,, 531

5 Fig12 The substitution among related species and four species groups of the genus Odorrana, 8, 50 % ;, ; 4,, O. m argaretae O. anl ungensis ; O. livida O. schm ackeri O. versabilis O. l ungshengensis, O. m argaretae O. w uchuanensis,,,,,,,, O. gra2 : 29 (1) 0 : ; 1 : ; (2) hami, O. andersonii 0 : ; 1 : ; (3) / ;, 0 : 1 ; 1 : 2 ; (4) 0 : ; 1 : ; (5) O. sw inhoana O. versabilis O. nasuta 0 : ; 1 : ; (6) 0 : O. exiliversabilis ; 1 : ; (7) 0 : ; 1 : ; (8) SVL / 0 : 110 ; 1 : 113 ; 2 : 117 ; (9) 0 : ; 1 : ; 2 :,,, ; (10) 0 : ; 1 : ; 2 : ; (11) 0 : ; 1 : ; 2 : ; (12) 0 :, ; 1 : ; (13) 0 : ; 1 : ; (14), 0 : : 525 ; 1 : : 424 ; 2 : (Liu, 1950 ; : 323 ; (15) 0 : ; 1 : ;, 1965 ;, 1979 ;, 1982) (16) 0 : ; 1 : ; (17) 0 : ; 1 : 314 ; (18) 0 : 1 ; 1 : 2 ; 2 3 ; (19) 1 0 : No ; 1 : No. 6 ; (20) 1, 0 : ; 1 : ; (21) 0 : ;, 1 : ; (22) 0 : ;,, 1 : ; (23) 0 :, ; 1 : ; 2 :, ; (24), 0 : ; 1 : ; (25) 0 :, T ; 1 : T ; (26) 0 : ; 1 :, ; (27) 0 : ; 1 : ; (28) 0 : ; 1 : ; (29) 0 : :, ; 1 :,

6 5 : ( : ) 533 ( References) Emerson, S. B. and D. Berrigan 1993 Systematics of southeast Asian ranids : multiple origins of voicelessness in the subgenus L i m nonectes ( Fitzinger). Herpetologica 49 (1) : Fei, L., C. Y. Ye and Y. Z. Huang 1990 Key to Chinese Amphibia. Chongqing : Chongqing Branch, Science and Technology Literature Pub2 lishing House, [,, :, ] Fei, L., C. Y. Ye and C. Li 2001 Descriptions of two new species of the Genus Odorrana in China (Anura : Ranidae). Acta Zootaxonomica Si nica 26 (1) : Kluge, A. G. and J. S. Farris 1969 Quantitative phyletics and the evolution of anurans. Syst. Zool. 18 : Li, C., C. Y. Ye and L. Fei 2001 Taxonomic studies of Odorrana andersonii in China (Anura : Ranidae). Acta Zootaxonomica Si nica 26 (2) : Liem, S. S Liu, C. Z The morphology, systematics and evolution of the Old world treefrogs (Rhacophoridae and Hyperoliidae). Zool. Mem. 57 : Taxonomic studies of Megophryinae in China. In : Collected Papers of Thirty Anniversary of The China Zoological Society. Bei2 jing : Science Press, 197. [ :. :, 197. ] Liu, C. Z. and S. Q. Hu 1961 Tailless Amphibians of China. Beijing : Science Press, [, :, ] Liu, C. C Amphibians of western China. Zool. Mem. 2 : 1 400, Plates 10. Macey, J. R., J. A. Schulte and A. Larson 1998 Phylogenetic relationships of toads in the B uf o buf o species group from the eastern escarpment of the Tibetan Plateau : a case of vicariance and dispersal. Molecular Phylogenetics and Evol ution 9 (1) : Maddison, W. P., M. J. Donoghue and D. R. Maddison 1984 Outgroup analysis and Parsimony. Syst. Zool. 33 (1) : Maeda, N. and M. Matsui 1990 Frogs and Toads of Japan. Tokyo : Bun2ichi Sogo Shuppan, Matsui, M. Okada, Y A taxonomic study of the Rana nari na complex, with description of three new species. Zool. J. L i nn. Soc. 111 : Fauna Japanica Anura (Amphibia). Tokyo : Electrical Engineer College Press, Xu, N., G. Wei and D. J. Li 1992 Study on the phylogenetic relationship between species of genus Oreolalax (Amphibia : Pelobatidae). Acta Herpetologica Si nica 1 2 : : ] Xu, B. C. [,, The migration a basic theory of Darwinπs evolutionism. Acta Palaeontologica Si nica 39 (2) : [ (2) : ] Ye, C. Y., C. Li and L. Fei 1999 Study on the phylogenetic relationships between species of Odorrana in China. In : Zoological Studies in Chi2 na. Beijing : China Forestry Publishing House, [,, 1999 ( ). :. :, ] Ye, C. Y., L. Fei and S. Q. Hu 1993 Rare and Economic Amphibians of China. Chengdu : Sichuan Publishing House of Science and Technology, [,, :, ] Zhang, R. Z Natural Geography of China Zoo2geography. Beijing : Science Press, [ :, ] Zheng, Z. X Classification of Vertebrata. Beijing : China Agriculture Publishing House, [ :, ]

7 ( Abstract) PHYLOGENY OF GENUS ODO R RANA ( AMPHIBIA : RANIDAE) IN CHINA 3 YE Chang2Yuan FEI Liang ( Chengdu Instit ute of Biology, The Chi nese Academy of Sciences, Chengdu , Chi na) We st udied t he inter2species relationships and biogeograp hy of t he genus O dorrana. 29 morp hological char2 acters of 16 species of O dorrana were studied with PAU P311, their geographic distribution were analyzed by migration t hesis. The cladogram shown t hat t he O dorrana species were of monop hyleticly origin and could be divided into four groups. andersonii group : O dorrana m argaretae, O. andersonii, O. grahami, O. hainangensis : kuangw uensis group : O. kuangw uensis, O. w uchuanensis. schm ackeri group : O. hejiangensis, O. tiannanensis, O. anl ungensis, O. schm ackeri, O. l ungshengensis, O. sw inhoana. livi da group : O. livi da, O. ex iliversabilis, O. versabilis, O. nasuta. The inter2species relationships show that andersonii group had the most amount plesiomorphy, so they were suggested as the most primitive species in the genus O dorrana. Because the most primitive species are distributed in Mt. Hengduan and Plateau of western Yunnan, t hus Mt. Hengduan and Plateau of western Yunnan are probably t he original regions of O dorrana. Eight species, accounting for 50 % of t he total number of O dorrana species in China, were discov2 ered in Guizhou Plateau, and all four species groups had representative species in t he area. Therefore, t his area was t he t ransition zone of odor f rogs migrated f rom western China to sout heastern China. Accordingly, t he Guizhou Plateau is probably t he center of diversification of O dorrana. The livi da group wit h more apomorp hic characters is dist ributed in eastern, sout hern part s and islands of China, t he group is evolutionary species2group in the genus. Along the distributed regions, the substitution of species within a species2group was found from south to north ( see Fig12), the substitution among species2groups was found from west to east, andersonii group2k uangw uensis group2schm ackeri group2livi da group. The primitive species dist ribute in sout hwestern part of China, and t he evolutionary species in eastern, sout heastern part s and islands of China. The livi da group is more closely related to schm ackeri group than to kuangw uensis group, and is more distantly to ander2 sonii group. These relationships among species groups are coincided very well wit h t heir dist ributional pattern. Key words O dorrana, Ranidae, Amphibia, Phylogeny, Origin and evolution 3 This project is supported by the National Natural Science Foundation of China (No ), the grant for Systematic and Evolutionary Bi2 ology, CAS., and the Knowledge Innovation Program of the Chinese Academy of Sciences