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思霁夏
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1 2005, 13 (4): Biodiversity Science doi: /biodiv http: // * 1 (, ) 2 (, ) : 166, , 24 54, 19, G-F ( ), ; Jaccard 19, :,,,, ( ), :,,, A large-scale pattern in species diversity of reptiles in the Yangtze River Basin Xiaodong Yu 1, Tianhong Luo 1, Qiang Dai 2, Yuming Wu 1, Hongzhang Zhou 1* 1 Institute of Zoology, Chinese Academy of Sciences, Beijing Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu Abstract: We synthesized information on reptile biodiversity in the Yangtze River Basin. We documented 166 species that had been recorded and described from the basin. There are 3 orders, 18 families and 68 genera. Of these, 24 species are endemic and 54 endangered. Since the distribution patterns of terrestrial reptiles are determined by the deep rivers and high mountains to a great extent, we divided the Yangtze River Basin into 19 sub-regions. Except the headwater of the basin, the other 18 sub-regions show similar values in species richness and G-F index. However, the proportion of endemic species decreased gradually from the headwater to the estuary of the basin with the gradient of elevation. Based on the species distribution in 19 sub-regions (Jaccard similarity), cluster analysis was used to analyze the similarity of reptiles in the 19 sub-regions. The 19 sub-regions were clustered into five groups: (1) The headwaters of the basin, (2) Hengduan Mountains and Yunnan Plateau, (3) Sichuan Basin and Qinling-Dabashan Mountains, (4) Guizhou Plateau, Jiangnan hills, the Two-Lake Plain and the delta of the Yangtze River Basin, (5) Poyang Lake Plain, lower reaches of the basin, and Huaiyang Mountains (from Hanjiang River to Dabieshan Mountain). This grouping reflects the environmental characteristics of the total basin and the three large topographic platforms of the Chinese mainland. Key words: reptile, species diversity, large-scale pattern, Yangtze River Basin,, 11 ( / ), 6300 km, , : ; : : (G ) (KSCX3-IOZ-01) (NSFC-J ) * Author for correspondence. zhouhz@ioz.ac.cn

2 4 : km 2, 19%, 3000 km,, 900 km, 5000 m 20 m, (, 1999),,, (, 1990),,,, (Chen et al., 2002; Fu et al., 2003, 2004),,,,, (1999), 11, 4300 km, km 2,,, ( ) ( ),, ( ),,, ( ),, 955 km, km 2,,,, ( ), km 2,,, ( ) ( 1),, (11 ) (, 1999),, 19 ( 1): (HY) (JY) (YD) (SJ) (DM) (SB) (JW) (MJ) (JH) (HQ) (WY) (YX) (HD) (LP) (XG) (GW) (PP) (NLM) (SLM)( 1) 1.2, ( 1) (, 1956;, 1977;, 1977;, 1980;, 1985;, 1989;, 1989;, 1992;, 1997;, 1998, 2003;, 1998, 1999;, 1998;, 1999),,,, ( 1) (CSIS) (, 2004), 3 : (1),,, (Mauremys

3 300 Biodiversity Science 13 1 ( ) ( ) Fig. 1 Sketch map of the Yangtze River Basin (, River and tributary;, Geographic sub-regions)

4 4 : 301 guangxiensis) (Calotes jerdoni) (Draco maculates) (Gekko swinhonis) (Takydromus amurensis) (Cyclophiops multicinctus) (Lycodon aulicus) (Sibynophis collaris) (Bungarus fasciatus) (Gloydius intermedius);(2),,, (Trimeresurus xiangchengensis) (Naja naja);(3),, (Japalura grahami) (Gekko scabridus) (Pareas stanleyi) (Scincella schmidti) (Opisthotropis kuatunensis) (Ermia mangshanensis) ( ), (, 2004) (CSIS) (, 2004) (, 2004), G-F (, 1999): (1) F (D F ): k, D Fk = n i= 1 ln pi pi p i = s ki / S k, S k k, s ki k i, n k F D F = m k= 1 DFk m (2) G (D G ): D G = p j= 1 ln qj qj q j = s j / S, S, s j j, p (3) G-F : D G-F 1 D G /D F G-F :(1), G-F ;(2) G-F 0 1 (Ludwig & Reynolds, 1988),,, (0 1 ), Jaccard 19, ; 1 (within-groups) (between-groups) (nearest neighbor) (furthest neighbor),, SPSS(1997) , ( 1);, 1, ; 18,, 10%;, 46, 1/3, 1.2%, 101,, 4.2%( 1) 1 ( ) Table 1 Faunal composition (order) of reptiles in the Yangtze River Basin, in China, and in the world Order Yangtze River Basin * China* Crocodiliformes Testudoformes Squamata * World* Lacertilia Serpentes * (Vol. 1, 2, 3) * Data are cited from Fauna Sinica Reptilia (Vol. 1, 2, 3)

302 Biodiversity Science 13 2 ( ) Fig. 2 Faunal composition (family) of reptiles in the Yangtze River Basin. Others denote the families less than 2% of all species 5%, ; 2% 5%, ; 1 2, 1%, ( 2) 2.

5 302 Biodiversity Science 13 2 ( ) Fig. 2 Faunal composition (family) of reptiles in the Yangtze River Basin. Others denote the families less than 2% of all species 5%, ; 2% 5%, ; 1 2, 1%, ( 2) , 14.46%; 37.35%, 62, 42.47%( 2) ( ), ( 3) 9,, 3, (Python molurus) ( ) (Alligator sinensis) (Pelochelys bibroni); 6, (Cuora yunnanensis) (Geoemyda spengleri) (Manouria impressa) (Caretta caretta) (Palea steindachneri) (Pelodiscus sinensis) , 32.53%,, 1/3 ( 2);, 42.59%,, 18.52% 9.26%, 2 3, 8 ( ) 1 ( 4) 2.3 5, (2 ) (25 ), 55, ;, 77, ( 5), (1 ) (6 ),, 12 13,, 10 ; (SJ), 18,, 1 ( 6) 3 : G 2 Table 2 Number of endemic species, China endemic species, and endangered species of reptiles to the Yangtze River Basin Order Endemic species to Yangtze River Basin Endemic species to China Yangtze River Basin * China* * Endangered species* Yangtze River Basin Crocodiliformes Testudoformes Squamata Lacertilia Serpentes Total * * Data are cited from China Species Red List China

4 : 303 3 Fig. 3 Percentage of endemic species of terrestrial reptiles in the Yangtze River Basin. The sub-region codes are the same as those in Fig. 1., (0.0679) (0.7661), G (1.55) (1.44) (1.18) (0.

6 4 : Fig. 3 Percentage of endemic species of terrestrial reptiles in the Yangtze River Basin. The sub-region codes are the same as those in Fig. 1., (0.0679) (0.7661), G (1.55) (1.44) (1.18) (0.6318) (0.7657) 1.0, (2.1566) 2.0, , F G, ( 3.0), (7.3652), G-F (0.8361), , 2.4 Jaccard 19 4 ( ) Fig. 4 Faunal composition (family) of endangered reptiles in the Yangtze River Basin

7 304 Biodiversity Science 13 5 Fig. 5 Species richness of terrestrial reptiles in different regions in the Yangtze River Basin. The sub-region codes are the same as those in Fig Fig. 6 Endangered terrestrial reptile species richness in different sub-regions in the Yangtze River Basin 3 166, 41%, 14.46%, 31% G-F,, 19, : 4 ;,,,,, (, 1999;, 1992)

8 4 : G-F Table 3 G-F index of terrestrial reptile species in different sub-region of the Yangtze River Basin Sub-region G G-index (D G ) F F-index (D F ) G-F G-F index (D G-F ) HY JY YD SJ DM MJ SB JW HQ JH WY YX HD LP XG GW PP SLM NLM The sub-region codes are the same as those in Fig. 1 7 Fig. 7 Dendrogram of 19 sub-regions based on Jaccard similarity of the terrestrial reptile distribution. The sub-region codes are the same as those in Fig. 1.

9 306 Biodiversity Science 13 ( ),, 3000 m,,, ;,,, ( ),, m,,,,,,,,, ( ) ( ),,, m,, ;, ( ),, 500 m,,, ;, ( ),, 100 m,, ; ( ),, 3 5 m,,, m,,,,,,, m m ( ),,, ;,,,,,,,,,,,,,, ;,,,,,,,,,,, ( 7) (HY), ; 2, (Phrynocephalus vlangalii) (JY YD) (SJ),,,,, ; 90, 23% (DM MJ SB) (HQ JH),,,

10 4 : 307 ; 88,, 10% (JW WY) (YX XG GW) (LP) (SLM), ; 123, 28% (HD) (PP) (NLM), ; 53, 68,, ;,,,,,,,,,,,,,, ;,,,,,, Changjiang Hydrological Committee of Hydrology Ministry( )(1999) Atlas of the Yangtze River Basin( ), pp China Map Press, Beijing.(in Chinese) Chen BH( )(1992) The Amphibian and Reptilian Fauna of Anhui( ), pp Anhui Science & Technology Publishing House, Hefei.(in Chinese) Chen Y, Chen Y, He D(2002) Biodiversity in the Yangtze river fauna and distribution of fishes. Journal of Ichthyology, 42, Chu XL( )(1989) Yunnan Province Annals(Vol. 6): Yunnan Animal Annals( ). The People s Press of Yunnan, Kunming.(in Chinese) Fu C, Wu J, Chen J, Wu Q, Lei G(2003) Freshwater fish biodiversity in the Yangtze River basin of China: patterns, threats and conservation. Biodiversity and Conservation, 12, Fu C, Wu J, Wang X, Lei G, Chen J(2004) Patterns of diversity, altitudinal range and body size among freshwater fishes in the Yangtze River basin, China. Global Ecology and Biogeography, 13, Herpetological Department, Sichuan Biological Research Institute( )(1977) A survey of reptiles in Xizang Autonomous Region, with faunal analysis and description of new forms. Acta Zoologica Sinica( ), 23, (in Chinese) Hu BQ( ), Huang MH( ), He SX( ), Wei KJ( )(1956) Serpentes Fauna of Zhejiang( ), pp Science Press, Beijing.(in Chinese) Hu SC( )(1977) Amphibia-Reptilia of Xizang( ). Science Press, Beijing.(in Chinese) Jiang ZG( ), Ji LQ( )(1999) Avian-mammalian species diversity in nine representative sites in China. Chinese Biodiversity( ), 7, (in Chinese with English abstract) Li DH( )(1989) Qinghai Fauna Economica( ). Qinghai People s Press, Xining.(in Chinese) Ludwig JA, Reynolds JF (1988) Statistical Ecology, pp Wiley, New York. Ren ME( ), Bao HS( )(1992) Natural Regions of China and Their Exploitation( ). Science Press, Beijing.(in Chinese) Shi BN( ), Zhao EM( )(1980) Sichuan Fauna Economica( ). Sichuan Publishing House of Science and Technology, Chengdu.(in Chinese) SPSS Inc(1997). SPSS Base 7.5 for Windows User s Guide. SPSS Inc, Chicago. The Law of Wild Animal Protection of People s Republic of China( )(1989) The List of Wild AnimalsUnder National Protection( ).(in Chinese) Wang S( ), Xie Y( )(2004) China Species Red List (Vol. 1): Red List( : ), pp Higher Education Press, Beijing.(in Chinese) Wu L( ), Li DJ( ), Liu JC( )(1985) The Reptilian Fauna of Guizhou ( ), pp Guizhou People s Press, Guiyang.(in Chinese)

11 308 Biodiversity Science 13 Xie Y( ), Wang S( ), He FQ( ), Zhao EM( ) (2004) China Species Information Service (CSIS) ( ). Zeng XC( )(1990) Fishery Resources of the Yangtze River Basin( ). Marine Press, Beijing.(in Chinese) Zhang MW( ), Zong Y( ), Ma JF( )(1998) Fauna Sinica (Reptilia 1): General Accounts of Reptilia, Testudoformes and Crocodiliformes( :,, ), pp Science Press, Beijing.(in Chinese) Zhang RZ( )(1999) Zoogeography of China( ). China Forestry Publishing House, Beijing. (in Chinese) Zhao EM( )(1998) China Red Data Book for Endangered Animals (Vol. 2): Amphibia & Reptilia.( : ). Science Press, Beijing.(in Chinese) Zhao EM( )(2003) The Colour Handbook of the Reptiles of Sichuan( ). China Forestry Publishing House, Beijing. (in Chinese) Zhao EM( ), Huang MH( ), Zong Y( ) (1998) Fauna Sinica (Reptilia 3): Squamata (Serpentes).( : ), pp Science Press, Beijing.(in Chinese) Zhao EM( ), Yang DT( )(1997) The Series of the Scientific Expedition to the Hengduan Mountains, Qinghai-Xizang Plateau: Amphibians and Reptiles of the Hengduan Mountains Region( : ). Science Press, Beijing.(in Chinese) Zhao EM( ), Zhao KT( ), Zhou KY( )(1999) Fauna Sinica (Reptilia 2): Squamata (Lacertilia) ( : ), pp Science Press, Beijing.(in Chinese) ( : : )

12 4 : Appendix 1 List and distribution of reptiles in the Yangtze River Basin Species HY JY YD SJ DM MJ SB JW HQ JH WY YX HD LP XG GW SLM PP NLM Testudoformes Platysternidae Platysternon megacephalum c, d, [3,10] + + Emydidae Chinemys megalocephala b, d C. reevesii c, d, [3,10,13,23] Cuora aurocapitata b, c, d + C. flavomarginata c, [2,3,10] C. pani b, c, d, [19] + + C. yunnanensis b, c, d + Geoemyda spengleri c, d, [10] + Mauremys mutica c, d, [23] Ocadia sinensis c, d + + Sacalia bealei b, c, d S. quadriocellata c, d + + Testudinidae Manouria impressa c, d, [27] + Cheloniidae Caretta caretta c, d + + Trionychidae Palea steindachneri c, d + Pelochelys bibroni c, d + + P. maculatus a,b, d + Pelodiscus sinensis c, d, [9,10] Crocodiliformes Alligatoridae ( ) Alligator sinensis b, c, d, [1,17,22] + + Squamata Agamidae Acanthosaura lepidogaster + Calotes emma + C. versicolor + + Japalura dymondi b J. flaviceps b J. grahami a, b + J. micangshanensis a, b + + J. splendida [9] J. szechwanensis [25] + + +

13 310 Biodiversity Science 13 1 ( ) Appendix 1 (continued) Species HY JY YD SJ DM MJ SB JW HQ JH WY YX HD LP XG GW SLM PP NLM J. varcoae a, b, [11] + J. yunnanensis + Phrynocephalus vlangalii b + Gekkonidae Gekko chinensis + + G. hokouensis G. japonicus [9,10] G. scabridus b G. subpalmatus b G. taibaiensis a, b + Hemidactylus bowringii + + Hemiphyllodactylus yunnanensis [4,28] + Scincidae Ateuchosaurus chinensis b + Eumeces capito b E. chinensis b, [9,10] E. elegans [9] E. liui a, b + + E. tunganus a, b + + Scincella barbouri a, b, c + S. doriae S. modesta b S. monticola b S. potanini a, b S. reevesii S. schmidti a, b + S. tsinlingensis b Sphenomorphus incognitus + + S. indicus [10] Tropidophorus hainanus + Lacertidae Eremias argus Platyplacopus intermedius a, b P. kuehnei b + Takydromus septentrionalis T. sexlineatus [10] T. wolteri Dibamidae

14 4 : ( ) Appendix 1 (continued) Species HY JY YD SJ DM MJ SB JW HQ JH WY YX HD LP XG GW SLM PP NLM Dibamus bourreti c + Anguidae Ophisaurus gracilis c + + O. harti c Typhlopidae Ramphotyphlops braminus [9] Boidae Python molurus c, [26] + Xenopeltidae Xenopeltis hainanensis c + + Colubridae Achalinus ater b + A. jinggangensis a, b, c + A. meiguensis a, b, [7] A. rufescens A. spinalis [9,10] Ahaetulla prasina + + Amphiesma atemporalis + + A. boulengeri b A. craspedogaster b, [9,10] A. johannis a, b A. metusia a, c + A. modesta + + A. octolineata b A. optata [25] A. popei b + + A. sauteri A. stolata Boiga kraepelini b, [14] B. multomaculata Calamaria pavimentata C. septentrionalis Coluber spinalis Cyclophiops major [15] Dinodon flavozonatum D. rufozonatum [6,10,15] Elaphe anomala c E. bimaculata b

15 312 Biodiversity Science 13 1 ( ) Appendix 1 (continued) Species HY JY YD SJ DM MJ SB JW HQ JH WY YX HD LP XG GW SLM PP NLM E. carinata c, [6,9] E. dione [13] E. frenata c, [24] E. mandarina c, [13] E. perlacea a, b, c E. porphyracea E. prasina c + + E. rufodorsata E. taeniura c, [10,13] Enhydris chinensis c, d, [9] E. plumbea c, d + + Lycodon fasciatus [14] L. ruhstrati [6,14] Macropisthodon rudis b Oligodon chinensis [10] O. cinereus + O. formosanus O. kunmingensis a, b, c, [12] + O. lacroixi c + O. lungshenensis b, c + + O. multizonatum b, c + + O. ningshanensis a, b, c + O. ornatus c, [24] Opisthotropis balteata + O. kuatunensis + O. lateralis + + O. latouchii Pareas boulengeri [6] P. chinensis [16] P. hamptoni c + P. stanleyi b Plagiopholis blakewayi [24] + + P. styani c, [14] Psammodynastes pulverulentus Pseudoxenodon bambusicola P. karlschmidti P. macrops [9] P. stejnegeri c

16 4 : ( ) Appendix 1 (continued) Species HY JY YD SJ DM MJ SB JW HQ JH WY YX HD LP XG GW SLM PP NLM Ptyas korros c P. mucosus c, [9] Rhabdophis leonardi R. nigrocinctus c + R. nuchalis [8,9] R. pentasupralabialis a, b R. subminiatus R. tigrinus [9,10,15] Rhynchophis boulengeri c + Sibynophis chinensis [6,9,10] Sinonatrix aequifasciata b S. annularis b S. percarinata [9,10] Xenochrophis piscator [10] Zaocys dhumnades b, c, [6,10,15] Z. nigromarginatus c Elapidae Bungarus multicinctus c, [10] Calliophis kelloggi [4] C. macclellandi [5] Naja naja c Ophiophagus hannah c, [24] Viperidae Azemiops feae c Deinagkistrodon acutus c, [9,10] Ermia mangshanensis a, b, c + Gloydius brevicaudus c, [18] G. strauchii c, [18,21] Ovophis monticola [5,10] Protobothrops jerdonii [9,13,20] P. mucrosquamatus Trimeresurus albolabris + + T. stejnegeri T. xiangchengensis a, b T. yunnanensis + + Total * : a: ; b: ; c: ( ); d: ; *, ; Note: a, endemic species to Yangtze River Basin; b, endemic species to China; c, endangered species (cited from China Species Red List ); d, aquatic species; * Because the habitat distribution of terrestrial and aquatic reptiles are significantly different, the distribution data for the aquatic species based on the sub-regions of terrestrial species characteristics here are only partly comparable to the terrestrial species distribution; The figure in the square bracket represents the reference recording the information of the species distribution.

17 314 Biodiversity Science 13 [1] Chen BH( ), Hua ZH( ), Li BH( )(1985) Alligator sinensis( ), pp Anhui Science & Technology Publishing House, Hefei.(in Chinese) [2] Chen BH( )(1979) Ecological data of the turtle Cuora flavomarginata. Chinese Journal of Zoology( ), 14(1), (in Chinese) [3] Deng XJ( ), Ye YY( )(1998) Reptile fauna and zoogeographical division in Hunan Province. Sichuan Journal of Zoology( ), 17(2), (in Chinese) [4] Deng XJ( ), Ye YY( ), Yang DD( )(1998) Two reptile records new to Hunan Province. Sichuan Journal of Zoology( ), 17(2), 53.(in Chinese) [5] Feng XY( )(1986) Two new records of venomous snakes from Gansu. Acta Herpetologica Sinica( ), 5(2), 158.(in Chinese) [6] Gao ZF( )(1989) Fauna and ecological distribution of amphibians and reptiles in Anxian County. Sichuan Journal of Zoology( ), 8(4), 40.(in Chinese) [7] Gao ZF( )(1991) Discovery of Achalinus meiguensis in Anxian County, Sichuan. Sichuan Journal of Zoology( ), 10, 33.(in Chinese) [8] Gu XH( )(1987) Discovery of Rhabdophis nuchalis(baulenger) in Pingwu County in Sichuan. Sichuan Journal of Zoology( ), 6(3), 38.(in Chinese) [9] Herpetological Department, Sichuan Institute of Biology( )(1976) A survey of reptiles in western Hubei Province. Materials for Herpetological Research, Chengdu: Sichuan Institute of Biology( : ), 3, (in Chinese) [10] Herpetological Department, Sichuan Institute of Biology( )(1976) Preliminary list of reptiles of Hunan Province and their geographical distribution. Materials for Herpetological Research, Chengdu: Sichuan Institute of Biology( : ), 3, (in Chinese) [11] Hu SC( ), Zhao EM( ), Liu CC( ) (1973) A survey of amphibians and reptiles in Kweichow Province, including a herpetofaunal analysis. Acta Zoologica Sinica( ), 19, (in Chinese) [12] Kou ZT( ), Wu JY( )(1993) A new species of Oligodon from Yunnan(Serpentes: Colubridae). Acta Zootaxonomica Sinica ( ), 18, (in Chinese) [13] Li C( ), Li SQ( ), Wang YZ( )(2000) diversity and origin of herpetological fauna from southern Gansu Province, China. Zoological Research( ), 21, (in Chinese) [14] Luo J( ), Gao HY( ), Zhou YY( )(2004) Species diversity and protection of reptiles in Chongqing Municipality. Sichuan Journal of Zoology( ), 23(3), [15] Nie LW( ), Guo CW( ), Wu XB( )(1995) The LDH isozyme page analysis of four tissues of five species in Colubridae. Zoological Research( ), 16, (in Chinese) [16] Rao DQ( ), Yang DT( )(1992) Phylogenetic systematics of Pareinae(Serpents) of southeastern Asia and adjacent islands with relationship between it and the geology changes. Acta Zoologica Sinica( ), 38, (in Chinese) [17] Ross JP(1992) Crocodile Conservation Action. IUCN, Gland, Switzerland. [18] Shen X( ), Zhou KY( ), Wang YQ( )(1999) RAPD analysis of pit-vipers of the genus Agkistrodon in China. Acta Zoologica Sinica( ), 45, (in Chinese) [19] Song MT( )(1984) A new species of the turtle genus Cuora(Testudoformes: Testudinidae). Acta Zootaxonomica Sinica( ), 9, (in Chinese) [20] Song MT( )(1985) Reproductive data of Trimeresurus jerdonii. La Animala Mondo( ), 2(3-4), (in Chinese) [21] Song MT( ), Chen FG( )(1985) A new subspecies of pit-viper in Qin-Ling, Shaanxi, China. La Animala Mondo( ), 2(2), (in Chinese) [22] Wang RP( )(1998) The current status of Alligator sinensis Fauvel and its conservation. Sichuan Journal of Zoology( ), 17(2), (in Chinese) [23] Wang YH( ), Yang JS( ), Chen BH( )(1988) Variations of horny shields on the Mauremys mutica and Chinemys reevesii. Sichuan Journal of Zoology( ), 7(2), (in Chinese) [24] Zhao EM( ), Liu XB( ), Kang SH( )(1986) Five snake species new to Sichuan. Acta Herpetologica Sinica( ), 5(2), 157.(in Chinese) [25] Zhao EM( ), Hu SC( )(1966) Three new species of reptiles from Szechwan. Acta Zootaxonomica Sinica( ), 3, (in Chinese) [26] Zhao EM( )(1987) The distribution of Python molurus bivittatus Schlegel over Sichuan Province being reconfirmed. Acta Herpetologica Sinica( ), 6(3), 78.(in Chinese) [27] Zhao XB( )(1986) Geochlone impressa(günther) discovered from Shaoyang, Hunan. Acta Herpetologica Sinica( ), 5(4), 313.(in Chinese) [28] Zhou KY( ), Liu YZ( ), Yang GP( )(1981) Three new subspecies of Hemiphyllodactylus yunnanensis(boulenger) from China (Lacertiformes: Gekkonidae). Acta Zootaxonomica Sinica( ), 6, (in Chinese)

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