Although species within Lagomorpha are derived from a common ancestor, the distribution range and body size of its two extant groups, ochotonids and leporids, are quite differentiated. It is unclear what has driven their disparate evolutionary history. In this study, we compile and update all fossil records of Lagomorpha for the first time, to trace the evolutionary processes and infer their evolutionary history using mitochondrial genes, body length and distribution of extant species. We also compare the forage selection of extant species, which offers an insight into their future prospects. The earliest lagomorphs originated in Asia and later diversified in different continents. Within ochotonids, more than 20 genera occupied the period from the early Miocene to middle Miocene, whereas most of them became extinct during the transition from the Miocene to Pliocene. The peak diversity of the leporids occurred during the Miocene to Pliocene transition, while their diversity dramatically decreased in the late Quaternary. Mantel tests identified a positive correlation between body length and phylogenetic distance of lagomorphs. The body length of extant ochotonids shows a normal distribution, while the body length of extant leporids displays a non-normal pattern. We also find that the forage selection of extant pikas features a strong preference for C3 plants, while for the diet of leporids, more than 16% of plant species are identified as C4 (31% species are from Poaceae). The ability of several leporid species to consume C4 plants is likely to result in their size increase and range expansion, most notably in Lepus. Expansion of C4 plants in the late Miocene, the so-called ‘nature’s green revolution’, induced by global environmental change, is suggested to be one of the major ‘ecological opportunities’, which probably drove large-scale extinction and range contraction of ochotonids, but inversely promoted diversification and range expansion of leporids.
Th e biodiversity of non-volant small mammals along an extensive subtropical elevational gradient was studied for the fi rst time on Gongga Mountain, the highest mountain in Hengduan Mountain ranges in China, located in one of the 25 global biodiversity hotspots. Non-volant small mammals were replicate sampled in two seasons at eight sampling sites between 1000 and 4200 m elevation on the eastern slope of Gongga Mountain. In all, 726 individual small mammals representing 25 species were documented in 28 800 trap nights. Th e species richness pattern for non-volant small mammals along the elevational gradients was hump-shaped with highest richness at mid-elevations. However, diff erent richness patterns emerged between endemic and non-endemic species, between larger-ranged and smaller-ranged species and between rodents and insectivores. Temperature, precipitation, plant species richness and geometric constraints (middomain eff ect) were most signifi cant in explaining species richness patterns. Based on the analysis of simple ordinary least squares (OLS) and stepwise multiple regressions, the overall richness pattern, as well as the pattern of insectivores, endemic species and larger-ranged species showed strong correlation with geometric constraint predictions. However, nonendemic species richness was more strongly correlated with temperature, while rodent richness was correlated with plant species richness. Our study shows that no single key factor can explain all richness patterns of non-volant small mammals. We need to be cautious in summarizing a general richness pattern of large species groups (e.g. small mammals or mammals) from species in smaller groups having diff erent ecological distributions and life histories. Elevational richness patterns and their driving factors for small mammals are more likely dependent on what kind of species we study.
The observed patterns and underlying mechanisms of elevational beta-diversity have been explored intensively, but multi-dimensional comparative studies remain scarce. Herein, across distinct beta-diversity components, dimensions and species groups, we designed a multi-faceted comparative framework aiming to reveal the general rules in the observed patterns and underlying causes of elevational beta-diversity. We have found that: first, the turnover process dominated altitudinal patterns of species beta-diversity ( β sim > β sne ), whereas the nestedness process appeared relatively more important for elevational trait dissimilarity ( β funcsim < β funcsne ); second, the taxonomic turnover was relative higher than its phylogenetic and functional analogues ( β sim > β phylosim / β funcsim ), conversely, nestedness-resultant trait dissimilarity tended to be higher than the taxonomic and phylogenetic measures ( β funcsne > β sne / β phylosne ); and third, as elevational distance increased, the contradicting dynamics of environmental filtering and limiting similarity have jointly led the elevational patterns of beta-diversity, especially at taxonomic dimension. Based on these findings, we infer that the species turnover among phylogenetic relatives sharing similar functional attributes appears to be the main cause of shaping the altitudinal patterns of multi-dimensional beta-diversity. Owing to the methodological limitation in the randomization approach, currently, it remains extremely challenging to distinguish the influence of the neutral process from the offset between opposing niche-based processes. Despite the complexities and uncertainties during species assembling, with a multi-dimensional comparative perspective, this work offers us several important commonalities of elevational beta-diversity dynamics.
The recently described trait‐based approach is becoming widely popular for a mechanistic understanding of species coexistence. However, the greatest challenge in functional analyses is decomposing the contributions of different ecological and evolutionary processes (e.g., niche‐based process, neutral process, and evolutionary process) in determining trait structure. Taking rodents (Rodentia) in the Hengduan Mountains as our study model, we aim to (1) quantify the vertical patterns of functional structure for head–body length (HL), tail/body ratio (TR), animal component in diet (ACD), and all traits; (2) disentangle the relative importance of different assembly processes (environment, space, and phylogeny) in structuring trait dispersion; and (3) assess the feasibility of Bergmann's rule and Allen's rule along elevational gradient. Our results have suggested that the vertical functional structure pattern varied across these three traits, indicating distinct functional roles in the community assembly process. These nonrandom vertical patterns of HL, TR, and terminal ACD have demonstrated these traits were dominated by different ecological process along environmental gradient. In variance partitioning, high proportion of the spatial variations in trait dispersion was explained by environmental and spatial models, which have provided supporting strong evidence for niche‐based and neutral processes in leading species coexistence. Although the three traits all exhibited apparent phylogenetic signals, phylogenetic relationship within community failed to predict the spatial variations of functional dispersion, confirming the enormous inference of phylogenetic signals in predicting trait structure. By assessing the vertical patterns of HL and TR at order and family levels, we argued that functional adaptation along an environmental gradient is a surrogate of series of complex processes (e.g., environmental filtering, interspecific interaction, and neutral dispersal) acting on multiple functional axes, which results in inconsistence with the empirical rules along elevational gradient.
The idea that a positive abundance-range size relationship (ARR) is pervasive in nature has been challenged by recent studies focused on montane and island vertebrate assemblages. However, because some of these studies used species' local abundance and regional or global range size in examining the ARRs, the negative and neutral trends reported are questionable. Here, by relating species' mean abundance along elevational gradients to elevational range size, we examined the ARRs of non-flying small mammals on three subtropical mountains of southwest China. We also examined the relationship between mean abundance and elevational range centre (reflecting species' elevational distribution) on each mountain, and compared the elevational range centre and mean abundance between endemic and non-endemic species as they may have been subjected to different intensities of historical (e.g. geographical isolation and colonization) and ecological (e.g. ecological specialization) processes. The results show significantly positive relationship between mean abundance and elevational range size on each mountain. We also observed a consistent positive relationship between mean abundance and elevational range centre, probably due to the stronger local specialization of mid-and high-elevation species, lower species richness at higher elevations, and increasing extinction rate of small-ranged less abundant species towards higher elevations. A novel finding of our study is that endemic species show higher elevational range centres and higher mean abundance than non-endemic species on each mountain, which is most likely driven by the increasing geographical isolation with elevation and the higher degree of ecological specialization for endemic species. Measuring abundance and range size at the same spatial scale is a key prerequisite to evaluate the ARRs of montane small mammals.
Q. (2015). Molecular phylogeny and the underestimated species diversity of the endemic white-bellied rat (Rodentia: Muridae: Niviventer) in Southeast Asia and China. -Zoologica Scripta, 44, 475-494.The white-bellied rat, Niviventer, is a genus endemic to Southeast Asia and China. However, the interspecific phylogenetic relationships and species diversity of this genus remain poorly understood. In the present study, single and multi-locus analyses were performed. Phylogenetic reconstruction on Cytochrome b (512 individuals, including data from Genbank) revealed five major clades with approximately 35 operational taxonomic units (OTUs), a number twice the existing taxonomy. The first clade (N. langbianis species group) was the earliest diverged. The second clade (N. fulvescens species group) diverged in Southeast Asia, the south and lower altitude regions of the Hengduan Mountains, and Southeast China. The third clade (the N. eha species group) is endemic to high altitudes in Northwest Yunnan and the central region of Himalaya. The fourth clade (the N. andersoni species group), is mainly confined to alpine regions of the Hengduan Mountains. The fifth clade (N. confucianus species group) is mainly distributed in the north and higher altitude regions of eastern Himalaya, the Hengduan Mountains and Taiwan, with the complex also invading central and northern China. Results from the combined dataset of four genes (Cytochrome b, Cytochrome oxidase subunit I, the D-loop sequence of the mitochondrial genome and the first exon of the nuclear interphotoreceptor retinoid binding protein) for 82 representative individuals from China generally coincide with the result of the single gene, with 12 OTUs identified. These results provide a preliminary framework for the existing classification of this highly diversified genus. The divergence time of Niviventer based on the four gene topology was dated to the late Miocene~6.41 Ma. Significant differences were detected in the general body form changes among these units based on voucher specimens. Moreover, geometric morphometric analysis of the cranium shape of voucher specimens indicated significant differences among five major species groups. Shape divergence of the cranium among several OTUs within the N. confucinaus complex is also significant. Our results provide further evidence for rapid and highly underestimated diversification of Niviventer both in genetics and morphology.
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°9 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.
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