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Studying the genetic differentiation in a unique geographical area contributes to understanding the process of speciation. Here, we explore the spatial genetic structure and underlying formation mechanism of two congeneric small mammal species (Apodemus draco and A. chevrieri), which are mainly distributed in the mountains surrounding the lowland Sichuan Basin, southwest China. We applied a set of comparative phylogeographical analyses to determine their genetic diversification patterns, combining mitochondrial (Cytb and COI) and nuclear (microsatellite loci) markers, with dense sampling throughout the range (411 A. draco from 21 sites and 191 A. chevrieri from 22 sites). Moreover, we performed three complementary statistical methods to investigate the correlation between genotype and geographical and environmental components, and predicted the potential suitable distributional range under the present and historical climate conditions. Our results suggest that both species have experienced allopatric differentiation and admixture in historical periods, resulting in a ring-shape diversification, under the barrier effect of the Sichuan Basin. We infer that the tectonic events of the Qinghai-Tibetan Plateau and climatic oscillations during the Quaternary played an important role on the genetic divergence of the two species by providing environmental heterogeneity and geographical variation. Our study reveals a case of two sympatric small mammals following a ring-shaped diversification pattern and provides insight into the process of differentiation.

Strong correlations between species diversity and climate have been widely observed, but the mechanism underlying this relationship is unclear. Here, we explored the causes of the richness–climate relationships among passerine birds in China by integrating tropical conservatism and diversification rate hypotheses using path models. We found that assemblages with higher species richness southwest of the Salween–Mekong–Pearl River Divide are phylogenetically overdispersed and have shorter mean root distances (MRDs), while species-rich regions northeast of this divide (e.g., north Hengduan Mountains–south Qinling Mountains) are phylogenetically clustered and have longer MRDs. The results of the path analyses showed that the direct effect of climatic factors on species richness was stronger than their indirect effects on species richness via phylogenetic relatedness, indicating that neither tropical conservatism nor diversification rate hypotheses can well explain the richness–climate relationship among passerines in China. However, when path analyses were conducted within subregions separately, we found that the tropical conservatism hypothesis was well supported in the southwestern Salween–Mekong–Pearl River Divide, while the diversification rate hypothesis could explain the richness–climate relationship well in the northeastern divide. We conclude that the diversity patterns of passerines in different subregions of the Eastern Himalayas-Mountains of Southwest China may be shaped by different evolutionary processes related to geological and climatic histories, which explains why the tropical conservatism or diversification rate hypothesis alone cannot fully explain the richness–climate relationships.

The ecological gradient–morphological variation (‘ecomorph’) relationship has long interested ecologists and evolutionary biologists, but it is applied far less frequently than genetic differentiation in cryptic species detection and species identification. With integrative methods, we revise taxonomic uncertainties in the Orientallactaga sibirica complex (OSC), with 298 sequence specimens and 469 voucher specimens from 138 localities covering nearly the entire distribution of the OSC. Phylogenetic relationships are assessed by Bayesian inference and maximum likelihood using two mitochondrial and nine nuclear genes. We use species-delimitation approaches to divide and validate the ‘candidate species’. We evaluate correlations between ecological divergence and phylogenetic splits, and visualize geographical patterns of morphological variation. The OSC is divided into four phylogenetic groups, the Ognevi, Altay, Bogda and Sibirica groups, and the OSC exhibits a significant ecomorph relationship and ecological divergence pattern. Morphological variations not only follow the general regularity under a large gradient of ecological factors, but are also closely related to the local environment/habitat. We suggest considering the comprehensive ecomorph relationship to identify species. Molecular analyses reveal that the OSC more easily forms deeply divergent lineages in the foothills and this differentiation depth may be related to mountain system size.

Understanding how biodiversity components are related under different environmental factors is a fundamental challenge for ecology studies, yet there is little knowledge of this interplay among the biotas, especially small mammals, in karst mountain areas. Here, we examine the elevation patterns of the taxonomic diversity (TD), phylogenetic diversity (PD), and functional diversity (FD) of small mammals in a karst mountain area, the Wuling Mountains, Southwest China, and compare these patterns between taxa (Rodentia and Eulipotyphla) and scales (broad‐ and narrow‐range species). We also disentangle the impacts of the human influence index, net primary productivity (NPP), normalized difference vegetation index (NDVI), annual precipitation (AP), and annual mean temperature (AMT) on these three facets of biodiversity by using structural equation modeling. We recorded a total of 39 small mammal species, including 26 rodents and 13 species of the order Eulipotyphla. Our study shows that the facets of biodiversity are spatially incongruent. Net primary productivity has a positive effect on the three facets for most groups, while the effect of the NDVI is negative for TD and PD in most groups. AMT temperature and AP have negative effects on FD and PD, whereas TD is dependent on the species range scale. The human influence index effect on TD and PD also depends on the species range scale. These findings provide robust evidence that the ecological drivers of biodiversity differ among different biotas and different range scales, and future research should use multifacet approach to determine biodiversity conservation strategies., We investigate elevational patterns of three diversity metrics for small mammals and relate them to climate, productivity, vegetation, and human impact. We find ecological distinct groups of small mammals were unevenly affected by environmental and anthropogenic factors.

The Niviventer fulvescens species complex (NFSC), a group of abundant and taxonomically ambiguous rodent taxa, is distributed from Southeast Asia to south-eastern China. We combined molecular and morphological datasets to clarify the species composition and variation of the NFSC. Our phylogenetic analyses, using molecular data, recovered eight genetic lineages in the NFSC, including a novel, distinct lineage from Jilong, Tibet, China, which is described as a new species, N. fengi sp. nov. The species status of N. fengi is supported by a species delimitation analysis, and it is morphologically distinguished from other members of the NFSC by its greyish dorsal fur, soft hairs covering the whole body and a hairy tail. NFSC species bearing well-developed spines are found at lower elevations. A comprehensive taxonomic revision of the NFSC within China is provided, represented by five species: N. cremoriventer, N. fulvescens, N. huang, N. mekongis comb. nov. and N. fengi. A further study of this species complex, including samples from Southeast Asia, is needed.

In this century, China has sustained unparalleled economic development, leading to exponentially growing investments in scientific research. Yet, the demand for research-funding is large and tracing the current knowledge is a key step to define priority research topics. In this same span, studies on bats in China have uncovered an overlooked diversity and revealed novelties in bats’ evolutionary history and life-history aspects. All this 21st-century knowledge, however, is scattered and a large part is concealed from most of the international scientific community in Mandarin-language articles. Here, we summarize the post-millennium (2000–2017) research on bats in China and point out trends and future directions based on neglected topics, groups, and regions. In addition, we provide an up-to-date list of bat species in China. We retrieved 594 publications related to bats in China, nearly half were written in Mandarin. At least 147 bat species are present in China, which places it among the most bat-rich countries in the world. There was a significant positive trend on the number of publications, from 12.5 annual average in 2000–2005 to 46.5 in recent years, reflecting the Chinese economic-scientific development in this century. We found marked taxonomic and spatial biases. Half of the studies in this century focused on Rhinolophus, Myotis, and Hipposideros, and the southern and eastern provinces were the most studied. Systematic/taxonomy and Ecology were the predominant topics post-millennium, whereas only 10 articles have clear conservation-driven goals. Our review shows that the majority of studies were focused on the least concern, cave-dweller species, and on bat-rich provinces. Future projects should address the effects of human-modified landscapes on bat community to define proper conservation actions. We discuss some priority actions and projects that will help to enhance bat protection in China.

Mice of the genus Apodemus are widely distributed across Eurasia. Several species of this genus are hosts of important zoonotic diseases and parasites. The evolutionary history and dispersal routes of these mice remain unclear and the distribution of these species in China was poorly explored in previous studies. We here investigate the divergence times and historical geographical evolution of Apodemus and study the taxonomy of species in China by integrating molecular and morphological data. The crown age of this genus is dated to the Late Miocene, approximately 9.84 Mya. Western and Central Asia were inferred as the most likely ancestral area of this genus. Moreover, we recognize nine living species of Apodemus in China: Apodemus uralensis, A. agrarius, A. chevrieri, A. latronum, A. peninsulae, A. draco, A. ilex, A. semotus and A. nigrus sp. nov., the last from the highlands (elevation > 1984 m) of Fanjing Mountain in Guizhou Province and Jinfo Mountain in Chongqing Province. This new species diverged from A. draco, A. semotus and A. ilex approximately 4.53 Mya. The discovery of A. nigrus highlights the importance of high mountains as refugia and ‘isolated ecological islands’ for temperate species in south-eastern China.

Rats of the Niviventer niviventer species complex (NNSC) are among the most abundant small mammals in Southeast Asia and China. More than 20 species or subspecies have been reported in the scientific literature; however, this species complex remains taxonomically ambiguous. With extensive sampling and integrated information from molecular and morphological data, we examined genetic divergences and morphological differences among species within the NNSC. Analyses of molecular voucher specimens revealed that the reported geographic ranges of most NNSC species need revision. Morphological analyses demonstrated that substantial differences exist among these species. Niviventer confucianus, the NNSC species with the largest number of subspecies, showed the greatest intraspecific morphological variation. The taxonomic revision presented here establishes 2 new taxa as independent species. This species complex in China now includes 8 species: N. bukit, N. confucianus, N. coninga, N. culturatus, N. lotipes, N. niviventer, 1 new species, N. gladiusmaculus sp. nov., and a new combination N. pianmaensis comb. nov.Version of Record, first published online October 30, 2018, with fixed content and layout in compliance with Art. 8.1.3.2 ICZN.

Mountainous landscapes are characterized by strong spatial heterogeneity, leading to increasing geographical isolation and decreasing area with elevation. Consequently, the colonization rate decreases from the low to high elevation zone, while the extinction rate shows the opposite. Due to such changes, we test whether (1) species occur at a declining number of sites (mountains) and have a less positive abundance–occupancy relationship (AOR) in a higher elevation zone; (2) a lower proportion of rare large-bodied species (less resistant to extinction) and a more positive abundance–body mass relationship (ABR) emerge in a higher elevation zone. Using the data of small mammals from 20 elevational gradients in the Mountainous Region of Southwest China, we compared the AORs and ABRs among the low, middle and high elevation zones. The AOR and ABR were fitted with linear and polynomial regression models. We compared endemic ratios among the different elevation zones. The AOR was best characterized by a linear model and positive in all elevation zones. Its slope decreased from the low to high elevation zone. The quadratic model performed the best in fitting the ABR in each zone. When fitted with linear models, both the R2 and slope of the ABR increased towards the high elevation zone. The endemic ratios were significantly higher in the middle and high elevation zones. Both the AOR and ABR in mountainous landscapes are strongly elevation-dependent. The increasing geographical isolation and decreasing area with elevation can have a high impact on macroecological patterns and processes.

Rodents are the major remaining mammals in many terrestrial ecosystems after the historical loss of megafauna and large-bodied taxa. Niviventer confucianus is a dominant habitat generalist in natural forests in most of China. It is also recorded as an important vector of diverse zoonotic diseases. Here, three mitochondrial and one nuclear DNA fragments were sequenced from samples covering most of the species range to study intraspecific genetic diversification and demographic history. Molecular voucher specimens of N. confucianus revealed that its assumed distribution range has been overestimated because of the hitherto unrecognized separation from parapatric species. Phylogenetic inferences recognized three geographically delimited intraspecific lineages that diverged at approximately 1.28 and 0.68 Mya. Hengduan Mountains, the east margin of Qinghai Tibetan Plateau, and the mountains surrounding Sichuan Basin were recognized as the major geographical barriers. Demographic analysis revealed dramatic population growth in southwest, central, and northern China in the late Pleistocene, but only slight growth in Yunnan/Tibet. The population boom apparently coincided with the reduction of predation and competition from the loss of megafauna in the late Pleistocene. Distributional ranges were inferred to be fairly stable through the late Quaternary glacial-interglacial climatic oscillations, possibly enabled by the species’ seed hoarding behavior and wide climatic tolerance. The demographic history of N. confucianus suggests that these rodents directly profited from the loss of megafauna, while their most recent increases potentially led to the proliferation of zoonotic disease by this species.

Background Understanding whether species’ elevational range is shifting in response to directional changes in climate and whether there is a predictable pattern in that response is one of the major challenges in ecology. However, so far very little is known about the distributional responses of subtropical species to climate change, especially for small mammals. In this study, we examined the elevational range shifts at three range points (upper and lower range limits and abundance-weighted range centre) of rodents over a 30-year period (1986 to 2014–2015), in a subtropical forest of Southwest China. We also examined the influences of four ecological traits (body mass, habitat breadth, diet and daily activity pattern) on the upslope shifts in species’ abundance-weighted range centres. Results Despite the warming trend between 1986 and 2015, the 11 rodent species in analysis displayed heterogeneous dynamics at each of the three range points. Species which have larger body sizes and narrower habitat breadths, show both diurnal and nocturnal activities and more specialized dietary requirements, are more likely to exhibit upslope shifts in abundance-weighted range centres. Conclusions Species’ distributional responses can be heterogeneous even though there are directional changes in climate. Our study indicates that climate-induced alleviation of competition and lag in response may potentially drive species’ range shift, which may not conform to the expectation from climate change. Difference in traits can lead to different range dynamics. Our study also illustrates the merit of multi-faceted assessment in studying elevational range shifts. Electronic supplementary material The online version of this article (doi:10.1186/s12898-017-0128-x) contains supplementary material, which is available to authorized users.

Species distributed along mountain slopes, facing contrasting habitats in short geographic scale, are of particular interest to test how ecologically based divergent selection promotes phenotypic and genetic disparities as well as to assess isolation‐by‐environment mechanisms. Here, we conduct the first broad comparative study of phenotypic variation along elevational gradients, integrating a large array of ecological predictors and disentangling population genetic driver processes. The skull form of nine ecologically distinct species distributed over a large altitudinal range (100–4200 m) was compared to assess whether phenotypic divergence is a common phenomenon in small mammals and whether it shows parallel patterns. We also investigated the relative contribution of biotic (competition and predation) and abiotic parameters on phenotypic divergence via mixed models. Finally, we assessed the population genetic structure of a rodent species (Niviventer confucianus) via analysis of molecular variance and FST along three mountain slopes and tested the isolation‐by‐environment hypothesis using Mantel test and redundancy analysis. We found a consistent phenotypic divergence and marked genetic structure along elevational gradients; however, the species showed mixed patterns of size and skull shape trends across mountain zones. Individuals living at lower altitudes differed greatly in both phenotype and genotype from those living at high elevations, while middle‐elevation individuals showed more intermediate forms. The ecological parameters associated with phenotypic divergence along elevation gradients are partly related to species' ecological and evolutionary constraints. Fossorial and solitary animals are mainly affected by climatic factors, while terrestrial and more gregarious species are influenced by biotic and abiotic parameters. A novel finding of our study is that predator richness emerged as an important factor associated with the intraspecific diversification of the mammalian skull along elevational gradients, a previously overlooked parameter. Population genetic structure was mainly driven by environmental heterogeneity along mountain slopes, with no or a week spatial effect, fitting the isolation‐by‐environment scenario. Our study highlights the strong and multifaceted effects of heterogeneous steep habitats and ecologically based divergent selective forces in small mammal populations.

Background South China encompasses complex and diverse landforms, giving rise to high biological diversity and endemism from the Hengduan Mountains to Taiwan Island. Many species are widely distributed across South China with similar disjunct distribution patterns. To explore the causes of these disjunct distribution patterns and their genetic consequences, we investigated the endemic species Père David’s Chinese Vole (Eothenomys melanogaster) by integrating geological and ecological factors. We analysed the genetic structure and divergence time of E. melanogaster based on fast-evolving mitochondrial and nuclear markers using Bayesian trees and coalescent species tree approaches. Historical scenarios of distribution range and demography were reconstructed based on spatial interpolations of genetic diversity and distance, extended Bayesian skyline plots, phylogeographic diffusion analysis, and ecological niche modelling (ENM) during different periods. We also assessed the relationships between geographical distance/ecological vicariance and genetic distance (isolation by distance, IBD; isolation by environment, IBE). Results The genetic analysis revealed three deeply divergent clades—Southeast, Southwest and Central clades, centred on the Wuyi Mountains, the Yunnan-Guizhou Plateau (YGP) and the mountains around the Sichuan Basin, respectively—that have mostly developed since the Pleistocene. IBD played an important role in early divergence, and geological events (sedimentation of plains and linking of palaeo-rivers) and IBE further reinforced genetic differentiation. ENM shows the importance of suitable habitats and elevations. Conclusions Our results suggest that the primary cause of the disjunct distribution in E. melanogaster is the high dependence on middle-high-altitude habitat in the current period. Mountains in the occurence range have served as “sky islands” for E. melanogaster and hindered gene flow. Pleistocene climatic cycles facilitated genetic admixture in cold periods and genetic diversification in warm periods for inland clades. During cold periods, these cycles led to multiple colonization events between the mainland and Taiwan and erased genetic differentiation. Electronic supplementary material The online version of this article (10.1186/s12862-018-1168-3) contains supplementary material, which is available to authorized users.

Aim To assess the validity of four hypothesized drivers (Quaternary climate, niche conservatism, contemporary climate, spatial configuration) of small 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° × 1°) scales. Regional beta diversity was evaluated by calculating the multiple-site dissimilarities and the distance-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 patterns were almost non-existent in QTP. Turnover was stronger in the latitudinal 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 variation 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 configuration 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 primarily explained by westward post-glacial recolonization, and support the alternative idea that QTP retained considerable refugia and even centres of origin during the Quaternary glaciations.

Understanding the mechanisms that govern the spatial patterns of species turnover (beta diversity) has been one of the fundamental issues in biogeography. Species turnover is generally recognized as strong in mountainous regions, but the way in which different processes (dispersal, niche, and isolation) have shaped the spatial turnover patterns in mountainous regions remains largely unexplored. Here, we explore the directional and elevational patterns of species turnover for nonvolant small mammals in the Hengduan Mountains of southwest China and distinguish the relative roles of geographic distance, environmental distance, and geographic isolation on the patterns. The spatial turnover was assessed using the halving distance (km), which was the geographic distance that halved the similarity (Jaccard similarity) from its initial value. The halving distance was calculated for the linear, logarithmic, and exponential regression models between Jaccard similarity and geographic distance. We found that the east–west turnover is generally faster than the south–north turnover for high‐latitudinal regions in the Hengduan Mountains and that this pattern corresponds to the geographic structure of the major mountain ranges and rivers that mainly extend in a south–north direction. There is an increasing trend of turnover toward the higher‐elevation zones. Most of the variation in the Jaccard similarity could be explained by the pure effect of geographic distance and the joint effects of geographic distance, environmental distance, and average elevation difference. Our study indicates that dispersal, niche, and isolation processes are all important determinants of the spatial turnover patterns of nonvolant small mammals in the Hengduan Mountains. The spatial configuration of the landscape and geographic isolation can strongly influence the rate of species turnover in mountainous regions at multiple spatial scales.

Medium- and large-bodied mammals (MLM) are of global interest of conservation due to their indispensable role in ecosystem functioning and sensitivity to human activities. Conservationists regard MLMs’ species richness as an important indicator of the conservation effectiveness of nature reserves (NR). However, species richness alone is unable to detect how much of the species pool is realized within local communities, nor the different consequences of loss of species having varying importance in ecosystem functioning on the communities. Here, we simultaneously studied the completeness and defaunation indices (both are useful in informing conservation practices) of MLMs in 33 NRs of Sichuan Province, Southwest China. We evaluated the main correlate (area, habitat heterogeneity, energy, water availability, anthropogenic impact) of each index using Pearson’s correlations and generalized least squares regressions. The effects of species’ body size, trophic guild and protection status on the defaunation index were also assessed. The results revealed a significant negative relationship between the two indices across the studied NRs. The NRs with larger elevation ranges (greater habitat heterogeneity) tended to have a more complete MLM fauna and less defaunated than NRs with smaller elevation ranges, and NRs with higher mean annual temperature were also more defaunated. At the species level, large-bodied protected herbivores were more defaunated than other MLM groups. Based on the findings, we argue that habitat heterogeneity and temperature should be the key considerations during NR designing of Sichuan Province to conserve MLM biodiversity, and large-bodied protected herbivores are the priority taxa in further conservation practices.

The underlying mechanisms that allow the Hengduan Mountains (HDM), the terrestrial biodiversity centre of China, to harbour high levels of species diversity remain poorly understood. Here, we sought to explore the biogeographic history of the endemic rat, Niviventer andersoni species complex (NASC), and to understand the long-term persistence of high species diversity in this region. In contrast to previous studies that have proposed regional refuges in eastern or southern of the HDM and emphasized the influence of climatic oscillations on local vertebrates, we found that HDM as a whole acted as refuge for the NASC and that the historical range shifts of NASC mainly occurred in the marginal regions. Demographic analyses revealed slight recent population decline in Yunnan and south-eastern Tibet, whereas of the populations in Sichuan and of the entire NASC were stable. This pattern differs greatly from classic paradigms of temperate or alpine and holarctic species. Interestingly, the mean elevation, area and climate of potential habitats of clade a (N. excelsior), an alpine inhabitant, showed larger variations than did those of clade b (N. andersoni), a middle-high altitude inhabitant. These species represent the evolutionary history of montane small mammals in regions that were less affected by the Quaternary climatic changes.

The Mongolian Plateau (MP), which is situated in the interior of Asia and possesses a typical continental climate, experienced harsh climatic conditions during the Quaternary glacial fluctuations. Although these events likely had huge impacts on the local animal populations, the current effects have hardly been explored. To investigate whether the MP supported a refugium along an oceanic-continental gradient (ROCG), and whether this refugium was glacial or interglacial, we investigated the demographic and phylogeographic history of an endemic mammal species, the desert hamster Phodopus roborovskii. We reconstructed the demographic variation, the phylogeographic diffusion, and modelled the potential habitat during historical periods. The genetic diversity in the MP was the highest among all the localities, and the MP was a suitable habitat throughout the modelled historical periods. A phylogeographic diffusion analysis emphasized the importance of the MP as the centre of origin, preservation and spread for P. roborovskii. The homogeneous landscape provided the opportunity for a wide gene flow, which resulted in low resolution of the phylogenetic relationships. Moreover, P. roborovskii was favoured by the interglacial condition, with both its demographical and geographical ranges expanded within the interglacial periods. The range variation from the Last Glacial Maximum to the current condition reflects a distinct longitudinal shift, while both ranges largely contracted from that of the Last Interglacial. Our results support that the MP served as a refugium and spread centre for P. roborovskii during the Quaternary climate fluctuations. The interglacial expansion and the longitudinal shifts highlighted the important effects of precipitations on the distribution range of species adapted to arid and semi-arid during glacial oscillations.