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Multiscale Partitioning of Small Mammal β Journal of Biogeography (J. Biogeogr.) (2016) 43, 1412–1424 ORIGINAL Multiscale partitioning of small mammal ARTICLE b-diversity provides novel insights into the Quaternary faunal history of Qinghai–Tibetan Plateau and Hengduan Mountains Zhixin Wen1, Qisen Yang1,*, Qing Quan1,2, Lin Xia1, Deyan Ge1 and Xue Lv1,2 1Key Laboratory of Zoological Systematics and ABSTRACT Evolution, Institute of Zoology, Chinese Aim To assess the validity of four hypothesized drivers (Quaternary climate, Academy of Sciences, Beijing 100101, China, 2 niche conservatism, contemporary climate, spatial configuration) of small College of Life Science, University of Chinese – Academy of Sciences, Beijing 100049, China mammal beta diversity in the Qinghai Tibetan Plateau (QTP) and the Hengduan Mountains (HDM). Location QTP and HDM of China. Methods We partitioned the beta diversity of small mammals in QTP and HDM into the spatial turnover and nestedness components at the regional (longitudinal/latitudinal zones) and grid (1° 9 1°) scales. Regional beta diver- sity was evaluated by calculating the multiple-site dissimilarities and the dis- tance-dissimilarity relationships. We examined the relative effects of geographical distance, environmental difference, habitat diversity, geographical isolation and Quaternary climate stability on the beta diversity patterns. Results The overall beta diversity in all longitudinal/latitudinal zones of both regions was primarily driven by spatial turnover, longitudinal nestedness pat- terns were almost non-existent in QTP. Turnover was stronger in the latitudi- nal direction of QTP and in the longitudinal direction of HDM, which corresponded to the general topography of each region. At the grid scale, higher turnover was primarily concentrated in mountainous areas. Turnover was highly correlated with geographical distance and environmental difference in both regions, and geographical isolation was another strong predictor of turnover in HDM. Habitat diversity independently explained most of the varia- tion in nestedness of HDM. Main conclusions Spatial turnover is the primary cause of the small mammal beta diversity in QTP and HDM. Three non-exclusive mechanisms including the historic effect of past glaciation, contemporary climate and spatial configu- ration of the landscape might act in combination to shape the beta diversity patterns in QTP and HDM, particularly the directional patterns. Our results challenge the prevailing view that the current distribution of QTP fauna is pri- marily explained by westward post-glacial recolonization, and support the alter- native idea that QTP retained considerable refugia and even centres of origin during the Quaternary glaciations. *Correspondence: Qisen Yang, Institute of Keywords Zoology, Chinese Academy of Sciences, No. 1 Contemporary climate, glaciations, Hengduan Mountains, nestedness, Qing- Beichen West Road, Chaoyang District, Beijing hai–Tibetan Plateau, Quaternary climatic oscillations, recolonization, small 100101, China E-mail: [email protected] mammals, spatial configuration, spatial turnover 1412 http://wileyonlinelibrary.com/journal/jbi ª 2016 John Wiley & Sons Ltd doi:10.1111/jbi.12706 Beta diversity partitioning reveals Quaternary mammalian faunal history turnover by promoting speciation and endemism, the glacial INTRODUCTION extinction and post-glacial recolonization due to the large Since the seminal work of Whittaker (1960), ecologists have oscillations in past climates may cause prominent nested recognized three components of species diversity (alpha, beta subset patterns in the composition of species (Dobrovolski and gamma diversity). In the broadest definition, beta diver- et al., 2012). As a consequence of this decomposition sity is generally applied to any measure of variation in the approach, the historical scenarios of how beta diversity pat- identities of species among sites (Anderson et al., 2011). terns formed in some taxa (e.g. global patterns of freshwater Therefore, an understanding of the patterns and underlying fishes and amphibians: Leprieur et al., 2011 and Baselga mechanisms of beta diversity is crucial to explain many eco- et al., 2012; European mammals: Svenning et al., 2011; New logical phenomena, from metacommunity dynamics (Soini- World vertebrates: Dobrovolski et al., 2012; and western nen et al., 2007a,b; Heino, 2013) to the origin and Mediterranean butterflies: Dapporto et al., 2014) were finely distribution of global biodiversity (Buckley & Jetz, 2008). detailed. To date, only a small number of studies have Because of the significance of beta diversity in a wide array focused on East Asia (e.g. Srinivasan et al., 2014), which of theoretical issues, in recent decades the increase in the contains two major biogeographical regions (Palearctic and number of studies exploring the deterministic and stochastic Oriental) and is the origin of many extant animals (Mayr, drivers of beta diversity has been dramatic (Shmida & Wil- 1963). Moreover, most previous studies on the partitioning son, 1985; Soininen et al., 2007a,b; Qian & Ricklefs, 2012), of beta diversity were conducted at a single scale. Therefore, with different definitions and measures of beta diversity con- multiscale examinations of the patterns of beta diversity are stantly being proposed (e.g. Tuomisto, 2010a,b; Anderson required to obtain more thorough insights into the mecha- et al., 2011). nisms behind (Barton et al., 2013). Among the theories used to explain beta diversity patterns, The Qinghai–Tibetan Plateau (QTP) and the Hengduan the theories on dispersal limitations (i.e. limited dispersal Mountains (HDM) are two adjacent regions located in abilities of species and geographical and environmental barri- south-west China, and HDM is a world hotspot of biodiver- ers to dispersal) and niche filtering have received the most sity (Mittermeier et al., 1998). Although QTP and HDM are attention in previous empirical studies. This attention indi- close geographically, the Quaternary climatic and faunal his- cates that both mechanisms play a significant role in struc- tories are thought to be largely different. For the diversity of turing beta diversity, but that the relative contributions may QTP, the prevailing view is that mass extinction of species vary substantially among taxa and regions (e.g. Buckley & had occurred several times in most regions of the plateau Jetz, 2008; Qian & Ricklefs, 2012). Nevertheless, historical during the Last Glacial Maximum (LGM) and other glacia- processes are equally important for their influence on pat- tions and that, a narrow zone at the eastern edge of QTP terns of beta diversity, but far fewer studies have explored with HDM served as a glacial refugium and sources of west- these mechanistic processes (but see Dobrovolski et al., 2012; ward interglacial and post-glacial recolonization (Zhang, Gavilanez & Stevens, 2013; Kubota et al., 2014). This smaller 2002; Qu et al., 2010). However, for HDM, most researchers number of studies is most probably because disentangling maintain that this large mountainous region harboured the roles of current and historical factors remains a difficult many climatically suitable habitats in low elevational areas task given that climatic conditions co-vary, and species typi- throughout the Quaternary. Thus, these refugia retained a cally follow more than one evolutionary path. large number of species from QTP and were the centres of Recently, with the methodological advance in the parti- subsequent speciation (Chen et al., 2010). Additionally, the tioning of beta diversity, this situation has changed; the total spatial configuration of the landscape also differs between beta diversity can be additively decomposed into two com- the two regions, which is primarily evident in the orientation ponents, which represent the pure spatial turnover and nest- of the mountain ranges (QTP: longitudinal orientation; edness (Baselga, 2010). A typical spatial turnover pattern HDM: latitudinal orientation). Based on these different his- occurs when the species at one site are replaced by different torical scenarios and spatial configurations of the landscape, species at another site (Gaston & Blackburn, 2000), with dif- we expected different patterns of spatial turnover and nested- ferent pools of species constituting the sources of the indi- ness of the taxa between QTP and HDM, particularly for vidual assemblages. Spatial turnover can be attributed to those in the latitudinal and longitudinal directions. Alterna- many causes, including dispersal limitations and niche filter- tively, if niche conservatism (Wiens & Donoghue, 2004) con- ing along climatic gradients (Qian et al., 2009; Dapporto stitutes a primary mechanism for the distribution of small et al., 2014). In contrast to turnover, nestedness is a pattern mammals in QTP and HDM, a nestedness pattern should be in which the biota present in one site are a strict subset of evident along the latitudinal gradient of both regions because the biota that occur at another more species-rich site, which of the failure of species to colonize the high-latitude regions may result from interspecific variation in environmental tol- (Svenning et al., 2011). erance and habitat nestedness (Ulrich et al., 2009). More- In this study, we partitioned the beta diversity of non- over, the historical drivers that lead to spatial turnover and volant small mammals (hereafter referred to as small mam- nestedness are different. Whereas the relatively stable climatic mals, including the orders Erinaceomorpha, Soricomorpha,
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