F I S H and F I S H E R I E S , 2014, 15, 209–230 Mapping China’s freshwater fishes: diversity and biogeography Bin Kang1, Junming Deng2, Yunfei Wu3, Liqiang Chen4, Jie Zhang5, Haiyuan Qiu6, Yin Lu4 & Daming He4 1Asian International Rivers Center, Yunnan University, Kunming, 650091, China; 2College Animal Science & Technology, Yunnan Agriculture University, Kunming, 650201, China; 3Fisheries College, Ocean University of China, Qingdao, 266003, China; 4Yunnan Key Lab of International Rivers and Transboundary Eco-security, Kunmin, 650091, China; 5Institute of Zoology, Chinese Academy of Sciences, Beijing, 100083, China; 6Fujian Monitoring Center of Geological Environment, Fuzhou, 3550001, China Abstract Correspondence: Freshwater fishes worldwide face an array of threats from dam construction, river Bin Kang, Asian International Rivers fragmentation, pollution, over-exploitation and interactions with introduced spe- Center, Yunnan cies. Such impacts are especially prevalent in densely populated countries undergo- University, 650091 ing rapid development, and their effects are felt most strongly where regional fish Kunming, China diversity is high. We reviewed studies of the distribution of freshwater fish species Tel.: +86-871- throughout China to map a biogeographical pattern and ascertain the environmen- 5033118 Fax: +86-871- tal factors contributing to this distribution. We then linked this information to 5034577 identify geographic areas to be recommended as the focus of conservation efforts. E-mail: bink A total of 920 species in 302 genera, 54 families and 21 orders were recorded. [email protected] Among the recorded species, 73% were Cypriniformes and 12% were Siluriformes. Cyprinidae was the most dominant family with 473 species, followed by Balitoridae Received 20 Mar with 157 species. The administrative division of the biogeography of China’s fresh- 2012 water fishes consisted of nine regions, including the Qinghai-Tibetan Plateau Accepted 16 Nov Region, Oriental Region, Northwest Region, South Region, Loess Plateau Region, 2012 Heilongjiang Region, Upper Yangtze Region, 3H Plain Region and Middle-Lower Yangtze Plain Region. The river system was the primary factor in determining Chi- na’s freshwater fish biogeography. Under stepwise regression analysis, river dis- charge was found to be the most influential factor in determining richness, followed by population size and net primary productivity. The higher level of fish endemicity and sensitivity to environmental change led to the identification of Southwest China and the higher areas of Qinghai-Tibetan Plateau as the primary areas to be considered for fish conservation and potential natural reserves. Keywords Cyprinidae, discharge, distribution, geological event, NPP, river system Introduction 210 Materials and methods 212 Study area 212 Fish data 212 Environmental factors 212 Geographical data 212 © 2013 John Wiley & Sons Ltd DOI: 10.1111/faf.12011 209 Freshwater fishes of China B Kang et al. Climatic data 213 Biological data 213 Statistical analysis 213 Richness and endemism 213 Cluster analysis 214 Regression analysis 214 Results 214 Diversity 214 Biogeography 214 Qinghai-Tibetan Plateau Region 214 Oriental Region 215 North-west Region 215 South Region 216 Loess Plateau Region 216 Heilongjiang Region 216 Upper Yangtze Region 217 The 3H Plain Region 217 Middle-Lower Yangtze Plain Region 217 Relationship between richness and environmental factors 217 Discussion 217 Diversity 217 Richness and environment 220 Factors affecting Chinese freshwater fish biogeography 220 Qinghai-Tibetan Plateau Region 220 Oriental Region 221 North-west Region 221 South Region 221 Loess Plateau Region 222 Heilongjiang Region 222 Upper Yangtze Region 223 3H Plain Region (see earlier for the derivation of this label) 223 Middle-Lower Yangtze Plain Region 223 Conclusions from biogeographical region delineation 224 The possible future tendency of fish biogeography 225 Conclusion 226 Acknowledgements 226 References 226 Supporting Information 229 et al. 2010; Hugueny et al. 2011). To mitigate Introduction such changes, fisheries management should aim Presently, the worldwide biota is entering into a at robust and functional ecosystems to reach a sixth mass extinction (Dirzo and Raven 2003; sustainable use of fisheries resources. Spatial-tem- Wake and Vredenburg 2008; Barnosky et al. poral distribution data, combined with the applica- 2011). Moreover, fishes, which comprise the larg- tion of biogeographical principles and theories, est group of vertebrates, are now experiencing a need to be clarified and updated. The general and marked change at a global scale (Casey and Myers fundamental characteristics of freshwater fish 1998; Pauly et al. 2003; Xenopoulos et al. 2005). fauna, for example, spatial patterns and distribu- Thus, the geographic map of fishes on this planet tion–environment relationship at global (Oberdorff needs to be redrawn (Sala et al. 2000; Butchart et al. 1995; Lev^eque et al. 2008) and continental 210 © 2013 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 15, 209–230 Freshwater fishes of China B Kang et al. scales (Oberdorff et al. 1999; Reyjol et al. 2007; (1985) found eight faunistic complexes that Kang et al. 2009) have been clearly delineated. explained China’s freshwater fish fauna. Although China’s freshwater fish fauna, which is charac- Shi (1985) provided lists of the typical fish species terized by the dominance of Cyprinidae, includes a in each complex, he did not identify the geographic part of the Oriental subregion of the Indian Region distribution patterns in detail; instead, he worked (south and south-west parts of China) and a part of them out in mostly unorganized combinations of the Central Asia subregion of the Palearctic Region plain/mountain and south/north. (north, east and middle parts of China) (Rainboth In contrast, Liu and Chen (1998) divided China’s 1991; Reshetrukov and Shakirova 1993; Jayaram freshwater fish fauna into six types based on extant 1999). Several ichthyologists have designated Cyprinidae. Their conclusion was primarily based China as the ‘centre of dispersal’ of the world’s on fish biology and ignored the inclusion of climate freshwater fishes (Darlington 1957; Wu 1964, and ecological factors. Thus, the divisions that Liu 1977; Menon 1987; Banarescu 1992). and Chen (1998) formed were vague, with many A 400-million-year-old fossil fish Psarolepis romeri, overlapping areas. With continuous improvements Incertae sedis, considered as the origin of bony in the data and statistical analysis methods on fish fishes, was discovered in Southwest China (Yu species, a more detailed study on the structure of 1998; Zhu et al. 1999). The current distribution of China’s freshwater fish fauna is now possible. China’s freshwater fish fauna formed in the late Freshwater fishes disperse in response to climate Tertiary to early Quaternary during the uplift of the or topographical changes, as well as to evolution- Qinghai-Tibetan Plateau and the global climate ary history and geological events (Olden et al. cooling. During periods of glaciations, fish species 2010). A species-discharge hypothesis (Welcomme retreated southwards, which favoured frequent 1979; Bronmark€ et al. 1984) similar to the species– land connections between the south-eastern main- area curve in terrestrial systems, which implies that land of China and western Pacific islands. In the fish species richness increases logarithmically with interglacial periods, species spread back to the river discharge, is generally used to explain the bio- north, resulting in the formation of dispersal species geography of fish spatial distribution. The species– and relict species (Zakaria-Ismail 1994; Chang energy hypothesis (Wright 1983), which predicts et al. 1996; Chang and Chen 2000). Alternate gla- that species richness is positively correlated with cial and interglacial changes also enabled fish energy availability, also gained attention in biogeo- exchanges among sections of large rivers, and these graphical studies (Guegan et al. 1998; Vor€ osmarty€ species differentiated in the upper, middle and lower et al. 2010). In addition, other factors (e.g. latitudi- reaches under the pressure of habitat change. nal zonation, longitudinal process, habitat hetero- Since the 1950s, ichthyologists have analysed geneity, contemporary climate, physicochemical China’s freshwater fish fauna at different levels, variability) have been considered to have marginal mainly by considering the differences in climate importance in shaping the spatial patterns of fish and drainage, as well as by empirical appraisal of fauna (Hurlbert et al. 1972; Oberdorff et al. 1997; fish distribution. Zhang (1954) first divided Chinese Leprieur et al. 2008; Araujo et al. 2009; Pracheil freshwater fishes into five divisions (Appendix S1). et al. 2009). Anthropogenic factors such as over- According to palaeogeohistory and ecological distri- exploitation, pollution, dam construction and river bution information on 767 China’s freshwater spe- fragmentation have significant effects on the abun- cies and subspecies from 209 genera, 33 families dance and distribution of fishes (Allan et al. 2006; and 13 orders, Li (1981) divided the fish fauna into Dudgeon et al. 2006). Integration of information five divisions and 21 subdivisions (Appendix S2). about these factors in China and in other parts of Pan et al. (1985) reanalysed the fish data from the the world can provide insights into the catalysts of study by Li (1981) at a subregional level using prime conservation problems and