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1 2005, 13 (6): Biodiversity Science doi: /biodiv http: // * (, ) :, 378 ( ), ( ),, 269 ( ), 11, 29 ; 162 ( ) 69 ( ), 19,, G-F,, Jaccard 19, : (1),, (2),, (3),, :,,, Large-scale patterns in species diversity of fishes in the Yangtze River Basin Xiaodong Yu, Tianhong Luo, Hongzhang Zhou * Institute of Zoology, Chinese Academy of Sciences, Beijing Abstract: We synthesized the information on fish diversity in the Yangtze River Basin, documenting 378 species/subspecies that had been recorded and described from the basin, belonging to 14 orders, 32 families and 144 genera. Of these, 338 species/subspecies are freshwater fishes, of which 269 species/subspecies are from the Cypriniformes. Of the total, 11 species are migratory fishes, and 29 are species of brackish water of the estuary. A total of 162 species/subspecies are endemic to the river and 69 are threatened. We divided the Yangtze River Basin into 19 sub-basins. Except for the two sub-basins of the headwaters and the upper and middle reaches of Jinshajiang River, the other 17 sub-basins showed similar values in species richness and G-F diversity indices, although the values were higher in the upper reaches than in the middle and lower reaches. However, the proportion of endemic species decreased gradually from the headwater to the estuary of the basin in parallel with the gradient of elevation. Jaccard similarity analysis showed that the 19 sub-basins were clustered into three groups: (1) the headwaters and the upper and middle reaches of Jinshajiang River, located in the eastern Qinghai-Xizang Plateau and Hengduan Mountains; (2) the other sub-basins of the upper reaches, located in Western Sichuan Plateau, Yungui Plateau, Sichuan Basin, and Qinling-Daba Mountains; (3) the middle and lower reaches, belonging to the Huaiyang Mountains, Jiangnan Hills and the plain along the middle and lower reaches of the Yangtze River. This grouping reflects the environmental characteristics of the basins and the three large topographic platforms of the Chinese mainland. Key words: fish, species diversity, large-scale pattern, Yangtze River Basin : ; : : (G ) (KSCX3-IOZ-01) (NSFC-J ) * Author for correspondence. [email protected]

2 474 Biodiversity Science 13,,,, (, 1990),,, (Chen et al., 2002; Fu et al., 2003, 2004;, 1997;, 2001;, 2004),, ;,,,, (1999), 11, 4300 km, km 2 ;,, ( ) ( ),, ( ),,, ( ),, 955 km, km 2 ;, ;, ( ), 1045 km, km 2,,, ( ) ( ), (11 ),, 19 ( 1) (HY) (UMJS) (LJS) (YLS) (DS) (MTS) (JLS) (WS) (HS) (YS) (XS) (DLS) (PLS) (GS) (TLS) (UMS) (MMS) (LMS) (EY)( 1) 1.2 ( 1),,,,, (, 1976;, 1980;, 1981;, 1981;, 1982;, 1984a, b;, 1987;, 1987;, 1987;, 1988;, 1988;, 1989a, b;, 1989;, 1989, 1995;, 1990;, 1990;, 1991;, 1992;, 1993;, 1993;, 1994;, 1995;, 1998a, b;, 1998;, 1999;, 2000),,

3 6 : I II Fig.1 Sketch map of the Yangtze River Basin. I, River and tributary; II, Geographic regions (sub-basins) in this study,, (, 1990, 1993;, 1991;, 1995;, 1996;, 1996;, 1997; He,

4 476 Biodiversity Science ; Zhang, 2000; Froese & Pauly, 2005), 4 : (1),,, (Allanetta bleekeri) (Coilia ectenes) (Coreoperca whiteheadi) (Liza carinatus) (Mastacembelus aculeatus) (Oryzias latipes sinensis) (Schistura dabryi) (Schistura xiangxiensis) (Schizothorax (Racoma) labrosus) (Siniperca obscura) (Siniperca roulei) (2),, (Acrossocheilus jishouensis) (Brachymystax lenok) (Channa maculata) (Ctenogobius multimaculatus) (Hemiculterella wui) (Mastacembelus armatus) (Pseudohemiculter hainanensis) (Salanx ariakensis) (Schistura incerta) (Triplophysa (Triplophysa) pseudoscleroptera) (3),, (Acrossocheilus elongates) (A. parallens) (Gnathopogon taeniellus) (Hemibarbus longirostris) (Liobagrus anguillicauda) (Paracheilognathus meridianus) (Pareuchiloglanis feae) (Phoxinus lagowskii) (Protosalanx chinensis) (Pseudobagrus adiposalis) (Puntius semifasciolatus) (Semilabeo notabilis) (Sinibrama macrops) Triplophysa (Triplophysa) dalaica) (4),, (Siniperca loona) ( ), (, 1993;, 2004; Chen et al., 2002; Fu et al., 2003), (, 2004), G-F (, 1999): (1) F (D F ): k, n D Fk = i= 1 ln pi pi p i = s ki / S k, S k k, s ki k i, n k F m D F = k= 1 DFk m (2) G (D G ): p D G 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 or complete linkage),, SPSS (1997) ( ),

5 6 : ( ), 1/3, 11, 29 ( 1), , 71 ;, 37 35, 10 ; 11, 37, 10 ( 1),, 1/3 ;, 10, ( 1) 4, , 50.26% 16.14%, ; 17, 4.50%; , 5.29% 2.12%; 16 7, %; 6, 1.59% 23 47, 1 5, ( 1) , 42.86%, ; 265, 70.10%, 60.36%; 5 ( 2),, ; ( 2) 7,, 3, (Acipenser dabryanus) (A. sinensis) (Psephurus gladius); 4, (Hucho bleekeri) (Myxocyprinus asiaticus) (Trachidermus fasciatus) (Anguilla marmorata)(, 1989) (, 2004) 69, 18.25%, 1/4,,, 32,, 6, 3, 2, 1 Table 1 Faunal composition (Order) of fishes in the Yangtze River Basin * * Order Yangtze River Basin China World Cypriniformes Siluriformes Perciformes Salmoniformes Clupeiformes Tetrodontiformes Cypriodontiformes Mugiliformes 4 23 Acipenseriformes Anguilliformes Scorpaeniformes Pleuronectiformes Atheriniformes Synbranchiformes * (1993) * Data are cited from Chen (1993)., No data.

6 478 Biodiversity Science 13 2 ( ) Table 2 Faunal composition (Order) of endemic species, China endemic species and threatened species of fishes in the Yangtze River Basin Endemic species to *, Endemic species to China *, Threatened species Order Yangtze River Basin Yangtze River Basin China Yangtze River Basin China Cypriniformes Siluriformes Perciformes Salmoniformes Clupeiformes Tetrodontiformes Cypriodontiformes Mugiliformes Acipenseriformes Anguilliformes Scorpaeniformes Pleuronectiformes Atheriniformes Synbranchiformes Total * (1993) (2004) * Data are cited from Chen (1993); Data are cited from Wang and Xie (2004) 2 ( 1) Fig.2 Percentage of endemic species of fishes in the Yangtze River Basin. The sub-basin codes are the same as those in Fig ( 1) 2.3 3, (21 ), 80, 139,

7 6 : 479, ;, , ( 3),,, 20,, ( 4) 3 4, 5 ( ), 3, G, ( 1.0),, 2.0, G ;,, , F G, ( 1.2), ( 8.0), , G-F, ,, 2.4, Jaccard 19, 0.3, 3 ( 5):,,, ;,, 3 ( 1) Fig.3 Species richness of fishes in different sub-basins in the Yangtze River Basin. The sub-basin codes are the same as those in Fig ( 1) Fig.4 Species richness of the threatened fishes in different sub-basins of the Yangtze River Basin. The sub-basin codes are the same as those in Fig. 1.

8 480 Biodiversity Science 13 3 G-F Table 3 G-F index of fish species in different sub-basins of the Yangtze River Basin Sub-basin G G-index (D G ) F F-index (D F ) G-F G-F index (D G-F ) HY UMJS YLS DS LJS MTS JLS WS UMS HS MMS YS XS DLS GS PLS LMS TLS EY The sub-basin codes are the same as those in Fig. 1. ;,, 0.5, ( ) ( ), ( ), 338 ( ), 1/3; 43%, ; 1/4,, ;, 5 ( ),,, (, 1992;, 1999) ( ),,, ;,

9 6 : ( 1) Fig.5 Dendrogram of 19 sub-basins using the method of complete linkage (furthest neighbor) based on Jaccard similarity measure of fish distribution in the Yangtze River Basin. The sub-basin codes are the same as those in Fig. 1.,,,,,,,,, ( ),,,,,,,,,,, ;,,,,,,,,,,,,,,,,,

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13 6 : Appendix 1 List and distribution of fishes in the Yangtze River Basin Species HY UMJS LJS YLS DS MTS JLS WS UMS HS YS XS DLS GS PLS MMS LMS TLS EY Acipenseriformes Acipenseridae Acipenser dabryanus d, e A. sinensis d, e Polyodontidae Psephurus gladius b, d, e Salmoniformes Salmonidae Brachymystax lenok d + Hucho bleekeri a, b, d Salangidae Hemisalanx brachyrostralis b H. prognathus f + + Neosalanx andersoni f + N. oligodontis b N. taihuensis b Salanx ariakensis e + Clupeiformes Clupeidae Tenualosa reevesii d, e Engraulidae Coilia branchygnathus a, b, f C. ectenes e C. mystus e + + Setipinna taty f + Thrissa kammalensis f + Anguilliformes Anguillidae Anguilla japonica e A. marmorata d, e + Cypriniformes Cyprinidae Abbottina obtusirostris a, b A. rivularis Acheilognathus barbatulus A. barbatus b A. chankaensis b A. elongatus elongatus a, b, d + A. gracilis a, b

14 486 Biodiversity Science 13 1 ( ) Appendix 1 (continued) Species HY UMJS LJS YLS DS MTS JLS WS UMS HS YS XS DLS GS PLS MMS LMS TLS EY A. hypselonotus a, b A. macropterus b A. omeiensis a, b A. polylepis b A. tabira A. tonkinensis b Acrossocheilus fasciatus b A. jishouensis a, b + A. labiatus b + + A. monticola a, b A. yunnanensis b Anabarilius alburnops a, b, d + A. brevianalis a, b + A. liui chenghaiensis a, b + A. liui liui a, b, d + + A. polylepis a, b, d + A. qionghaiensis a, b, d + A. songmingensis a, b + A. xundianensis a, b + Ancherythroculter kurematsui a, b A. nigrocauda a, b A. wangi a, b Aphyocypris chinensis Aristichthys nobilis Atrilinea macrolepis a, b, d + A. roulei b, d + Barbodes laticeps b + B. polylepis a, b + Belligobio nummifer b B. pengxianensis a, b, d + Carassius auratus auratus Coreius guichenoti a, b C. heterodon b Ctenopharyngodon idellus Culter alburnus C. dabryi dabryi C. mongolicus elongatus a, b + C. mongolicus mongolicus C. mongolicus qionghaiensis a, b + C. oxycephaloides a, b

15 6 : ( ) Appendix 1 (continued) Species HY UMJS LJS YLS DS MTS JLS WS UMS HS YS XS DLS GS PLS MMS LMS TLS EY C. oxycephalus b Cultrichthys erythropterus Cyprinus (Cyp.) carpio C. (Cyp.) chilia b, d + C. (Cyp.) qionghaiensis a, b, d + C. (Mesocyprinus) micristius a, b, d + Discogobio brachyphysallidos b + D. yunnanensis b Distoechodon hupeinensis a, b, d + D. tumirostris b Elopichthys bambusa b Garra pingi pingi Gnathopogon herzensteini a, b G. imberbis a, b G. nicholsi a, b + Gobiobotia abbreviata a, b G. brevirostris b + G. filifer a, b G. meridionalis G. tungi b + + Gobiocypris rarus a, b, c, d + + Gymnocypris potanini firmispinatus a, b + + G. potanini ptanini b Gymnodiptychus pachycheilus b, d Hemibarbus labeo H. maculatus Hemiculter bleekeri b H. leucisculus H. tchangi a, b Hemiculterella sauvagei a, b H. wui b + + Herzensteinia microcephalus b + + Hypophthalmichthys molitrix Luciobrama macrocephalus d Megalobrama amblycephala a, b M. elongata a, b, d + + M. pellegrini a, b M. skolkovii Microphysogobio fukiensis b

16 488 Biodiversity Science 13 1 ( ) Appendix 1 (continued) Species HY UMJS LJS YLS DS MTS JLS WS UMS HS YS XS DLS GS PLS MMS LMS TLS EY M. kiatingensis b M. microstomus a, b + + M. tungtingensis a, b Mylopharyngodon piceus Ochetobius elongatus Onychostoma angustistomata a, b O. barbata b O. barbatula b + O. brevis a, b O. daduensis a, b + O. lini d + + O. macrolepis b O. rara b, d O. sima Opsariichthys bidens Parabramis pekinensis Paracanthobrama guichenoti a, b Paracheilognathus himantegus b P. imberbis b Parasinilabeo maculatus a, b + Percocypris pingi pingi a, b, d Phoxinus oxycephalus Platysmacheilus exiguus P. longibarbatus a, b, d + P. nudiventris a, b Procypris rabaudi a, b, d Pseudobrama simoni b Pseudogobio vaillanti b Pseudogyrincheilus procheilus a, b Pseudohemiculter dispar + P. hainanensis + P. kweichowensis a, b + Pseudolaubuca engraulis b P. sinensis Pseudorasbora elongata b, d + + P. parva Ptychobarbus chungtienensis chungtiensis a, b, d + + P. chungtienensis gezaensis a, b + + P. kaznakovi b Rectoris luxiensis a, b

17 6 : ( ) Appendix 1 (continued) Species HY UMJS LJS YLS DS MTS JLS WS UMS HS YS XS DLS GS PLS MMS LMS TLS EY R. mutabilis b + Rhinogobio cylindricus a, b R. hunanensis a, b R. typus b R. ventralis a, b Rhodeus fangi b R. lighti R. ocellatus Sarcocheilichthys davidi a, b S. kiangsiensis b S. nigripinnis S. parvus b S. sinensis sinensis Saurogobio dabryi S. dumerili b S. gracilicaudatus a, b S. gymnocheilus a, b S. xiangjiangensis b + + Schizopygopsis kialingensis a, b + S. malacanthus baoxingensis a, b + + S. malacanthus chengi a, b S. malacanthus malacanthus a, b Schizothorax (Racoma) davidi a, b S. (Racoma) griseus b, d S. (Racoma) kozlovi a, b S. (Racoma) labrosus a, b, d + S. (Racoma) longibarbus a, b, d + + S. (Racoma) microstomus a, b, d + S. (Racoma) ninglangensis a, b, d + S. (Racoma) parvus a, b, d + S. (Racoma) yunnanensis weiningensi a, b S. (Schizothorax) chongi a, b S. (Schizothorax) cryptolepis a, b + S. (Schizothorax) dolichonema b, d S. (Schizothorax) grahami a, b, d S. (Schizothorax) heterochilus a, b + S. (Schizothorax) prenanti a, b S. (Schizothorax) sinensis a, b S. (Schizothorax) wangchiachii a, b Sinibrama taeniatus a, b

18 490 Biodiversity Science 13 1 ( ) Appendix 1 (continued) Species HY UMJS LJS YLS DS MTS JLS WS UMS HS YS XS DLS GS PLS MMS LMS TLS EY S. wui b Sinilabeo longibarbatus a, b + S. rendahli a, b S. tungting a, b Sinocrossocheilus guizhouensis a, b, d + Sinocyclocheilus(S.) grahami a, b, d + S.(S.) multipunctatus b + Spinibarbus hollandi S. sinensis a, b Squalidus argentatus b S. nitens a, b S. wolterstorffi b Squaliobarbus curriculus Tor (Foliter) brevifilis brevifilis Toxabramis swinhonis b Xenocypris argentea X. davidi b X. fangi a, b, d X. microlepis X. yunnanensis a, b, d Xenophysogobio boulengeri a, b, c X. nudicorpa a, b, c Zacco chengtui a, b, d + + Z. platypus Catostomidae Myxocyprinus asiaticus b, d Cobitidae Botia (Sinibotia) superciliaris a, b B. reevesae b Cobitis macrostigma a, b C. rarus b + + C. sinensis C. taenia + Leptobotia elongata a, b, d L. guilinensis b + L. microphthalma a, b L. orientalis b + L. pellegrini b L. rubrilabris a, b L. taeniops a, b

19 6 : ( ) Appendix 1 (continued) Species HY UMJS LJS YLS DS MTS JLS WS UMS HS YS XS DLS GS PLS MMS LMS TLS EY L. tchangi tchangi a, b + L. tientaiensis hansuiensis a, b Misgurnus anguillicaudatus Nemacheilus obscurus + Parabotia banarescui a, b P. bimaculata a, b P. fasciata b P. maculosa b Paracobitis potanini a, b P. variegatus b P. wujiangensis a, b + + Paramisgurnus dabryanus b Schistura dabryi b S. fasciolata b + + S. incerta + S. xiangxiensis a, b + Sphaerophysa dianchiensis a, b, c, d + Triplophysa polyfasciata a, b + + T. (Triplophysa) angeli a, b T. (Triplophysa) anterodorsalis a, b + T. (Triplophysa) bleekeri a, b T. (Triplophysa) brevibarba a, b + + T. (Triplophysa) brevicauda b T. (Triplophysa) crassilabris a, b + T. (Triplophysa) daqiaoensis a, b + + T. (Triplophysa) grahami b T. (Triplophysa) leptosoma b T. (Triplophysa) markehenensis a, b T. (Triplophysa) microps T. (Triplophysa) ninglangensis a, b + T. (Triplophysa) obscura b + T. (Triplophysa) orientalis b T. (Triplophysa) pseudoscleroptera b + T. (Triplophysa) robusta b + T. (Triplophysa) rotundiventris b + T. (Triplophysa) stenura T. (Triplophysa) stewarti + T. (Triplophysa) stoliczkae T. (Triplophysa) tanggulaensis a, b + T. (Triplophysa) venusta a, b +

20 492 Biodiversity Science 13 1 ( ) Appendix 1 (continued) Species HY UMJS LJS YLS DS MTS JLS WS UMS HS YS XS DLS GS PLS MMS LMS TLS EY T. (Triplophysa) xichangensis a, b + T. (Triplophysa) xiqiensis a, b + T. (Triplophysa) yaopeizhii a, b + Yunnanilus caohaiensis a, b + + Y. longibulla a, b + Y. nigromaculatus a, b + + Y. pleurotaenia a, b + Y. sichuanensis a, b + + Homalopteridae Beaufortia liui a, b B. szechuanensis a, b Hemimyzon yaotanensis a, b, d Jinshaia abbreviata a, b, c J. sinensis a, b, c Lepturichthys fimbriata a, b Metahomaloptera omeiensis omeiensis a, b, c M. omeiensis hangshuiensis a, b, c Paraprotomyzon lungkowensis a, b P. multifasciatus b + Pseudogastromyzon fangi b + Sinogastromyzon hsiashiensis a, b + + S. sichangensis a, b S. szechuanensis a, b Vanmanenia pinchowensis b V. stenosoma b + + V. tetraloba + Siluriformes Bagridae Leiocassis crassilabris b L. longibarbus a, b, d + L. longirostris b L. tenuifurcatus b Mystus macropterus b Pelteobagrus eupogon a, b, d P. fulvidraco P. nitidus b P. vachelli Pseudobagrus analis a, b + + P. brevicaudatus b P. emarginatus b

21 6 : ( ) Appendix 1 (continued) Species HY UMJS LJS YLS DS MTS JLS WS UMS HS YS XS DLS GS PLS MMS LMS TLS EY P. medianalis a, b, d P. omeihensis a, b + P. ondan b + P. pratti b P. teaniatus b P. tenuis b P. truncatus b P. ussuriensis Amblycipitidae Liobagrus kingi a, b, d + L. marginatoides a, b + + L. marginatus a, b, d L. nigricauda a, b L. styani a, b + + Siluridae Silurus asotus S. mento b, d + S. meridionalis b Clariidae Clarias fuscus Sisoridae Euchiloglanis davidi a, b, d E. kishinouyei a, b, d Glyptothorax fukiensis fukiensis G. sinense sinense a, b Pareuchiloglanis anteanalis a, b P. sinensis a, b, d P. robusta a, b + P. sichuanensis a, b + + Atheriniformes Atherinidae Allanetta bleekeri f + + Cypriodontiformes Oryziatidae Oryzias latipes O. latipes sinensis d + Poeciliidae Gambusia affinis Hemirhamphidae Hyporhamphus intermedius

22 494 Biodiversity Science 13 1 ( ) Appendix 1 (continued) Species HY UMJS LJS YLS DS MTS JLS WS UMS HS YS XS DLS GS PLS MMS LMS TLS EY Synbranchiformes Synbranchidae Monopterus albus Mugiliformes Mugilidae Liza carinatus f + L. haematocheila f + Mugil cephalus f Polynemidae Eleutheronema tetradactylum f + Perciformes Belontiidae Macropodus chinensis M. opercularis Channidae Channa argus C. asiatica C. maculata + Serranidae Coreoperca whiteheadi + + Siniperca chuatsi S. kneri b S. obscura b, d S. roulei b, d S. scherzeri S. undulata b, d Eleotridae Bostrichthys sinensis f + Eleotris oxycephala f + Hypseleotris swinhonis Odontobutis obscuras f Gobiidae Acanthogobius elongata d, f + Chaeturichthys stigmatias f + Ctenogobius brunneus + + C. chengtuensis a, b, d C. cliffordpopei b, f C. giurinus C. multimaculatus b, d + C. shennongensis a, b, d + +

23 6 : ( ) Appendix 1 (continued) Species HY UMJS LJS YLS DS MTS JLS WS UMS HS YS XS DLS GS PLS MMS LMS TLS EY C. szechuanensis a, b, d Glossogobius giuris f + Lophiogobius ocellicauda f + Mugilogobius myxodermus d, f M. polylepis b, d, f + Odontamblyopus rubicundus f + + Synechogobius ommaturus f + Tridentiger trigonocephalus f + + Mastacembelidae Mastacembelus aculeatus M. armatus Sinobdella sinensis + Scorpaeniformes Cottidae Trachidermus fasciatus d, e Pleuronectiformes Cynoglossidae Cynoglossus gracilis f + Tetrodontiformes Tetraodontidae Takifugu coronoidus a, b, d, f + + T. flavidus d, f + T. obscurus e T. ocellatus f T. vermicularis f + T. xanthopterus f + 1 (a) (b) (c) (d) (e) (f) The sub-basin codes are the same as those in Fig. 1. The superscripts represent endemic species to Yangtze River Basin (a), endemic species to China (b), endemic genus to China (c); threatened species (d); migratory fishes (e), and fishes chiefly in brackish water of the estuary of the river (f), respectively.