The relationship between mammal faunas and climatic instability since the Last Glacial Maximum: A Nearctic vs. Western Palearctic comparison

Climate has played a key role in shaping the geographic patterns of biodiversity. The imprint of Quaternary climatic fluctuations is particularly evident on the geographic distribution of Holarctic faunas, which dramatically shifted their ranges following the alternation of glacial-interglacial cycles during the Pleistocene. Here, we evaluate the existence of differences between climatically stable and unstable regions – defined on the basis of climatic change velocity since the Last Glacial Maximum – in the geographic distribution of several biological attributes of extant terrestrial mammals of the Nearctic and Western Palearctic regions. Specifically, we use a macroecological approach to assess the dissimilarities in species richness, range size, body size, longevity and litter size of species that inhabit regions with contrasting histories of climatic stability. While several studies have documented how the distributional ranges of animals can be affected by long-term historic climatic fluctuations, there is less evidence on the species-specific traits that determine their responsiveness under such climatic instability. We find that climatically unstable areas have more widespread species and lower mammal richness than stable regions in both continents. We detected stronger signatures of historical climatic instability on the geographic distribution of body size in the Nearctic region, possibly reflecting lagged responses to recolonize deglaciated regions. However, the way that animals respond to climatic fluctuations varies widely among species and we were unable to find a relationship between climatic instability and other mammal life-history traits (longevity and litter size) in any of the two biogeographic regions. We, therefore, conclude that beyond some biological traits typical of macroecological analyses such as geographic range size and body size, it is difficult to infer the responsiveness of species distributions to climate change solely based on particular life-history traits.     

Climate and mammalian life histories

Mammals display considerable geographical variation in life history traits. To understand how climatic factors might influence this variation, we analysed the relationship between life history traits – adult body size, litter size, number of litters per year, gestation length, neonate body mass, weaning age and age at sexual maturity – and several environmental variables quantifying the seasonality and predictability of temperature and precipitation across the distribution range of five terrestrial mammal groups. Environmental factors correlated strongly with each other; therefore, we used principal components analysis to obtain orthogonal climatic predictors that could be used in multivariate models. We found that in bats, primates and even‐toed ungulates adult body size tends to be larger in species inhabiting cold, dry, seasonal environments, whereas in carnivores and rodents a smaller body size is characteristic of warm, dry environments, suggesting that low food availability might limit adult size. Species inhabiting cold, dry, seasonal habitats have fewer, larger litters and shorter gestation periods; however, annual fecundity in these species is not higher, implying that the large litter size of mammals living at high latitudes is probably a consequence of time constraints imposed by strong seasonality. On the other hand, the number of litters per year and annual fecundity were greater in species inhabiting environments with higher seasonality in precipitation. Lastly, we found little evidence for specific effects of environmental variability. Our results highlight the complex effects of environmental factors in the evolution of life history traits in mammals. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 719–736.     

Extinction and replacement among predatory mammals in the North American late Eocene and Oligocene: Tracking a paleoguild over twelve million years

Changes in taxonomic and morphologic diversity within the paleoguild of predatory mammals were explored within a mammalian chronofauna spanning a twelve million year interval, from the latest Eocene to the end of the Oligocene of North America (36–24 ma). The timespan encompassed a modest extinction event among terrestrial mammals (circa 34 ma) followed by a period of relative stability. Morphological diversity was assessed with estimates of body mass, relative tooth size, and tooth shape. Principal component, nearest‐neighbor, and minimum‐spanning‐tree analyses were used to compare morphological diversity and species packing within predator paleoguilds in the mid‐Chadronian (37–34 ma), Orellan (34–32), Whitneyan (32–29.5), and early Arikareean (29.5–24) land mammal ages. Species richness of predators throughout the interval was relatively constant, fluctuating between 15 and 18 total taxa. Moreover, despite significant differences in taxonomic composition and a modest extinction event among terrestrial mammals, morphological diversity within the paleoguild was very similar in the Chadronian and Orellan. In the Whitneyan and especially the early Arikareean, the diversity of feeding adaptations among species declined slightly, largely due to the loss of several highly specialized meat‐eaters (creodonts, nimravids) and the addition of small omnivores (canids).     

Effects of neutrality and productivity on mammal richness and evolutionary history in Australia

Explaining how heterogeneous spatial patterns of species diversity emerge is one of the most fascinating questions of biogeography. One of the great challenges is revealing the mechanistic effect of environmental variables on diversity. Correlative analyses indicate that productivity is associated with taxonomic, phylogenetic, and functional diversity of communities. Surprisingly, no unifying body of theory have been developed to understand the mechanism by which spatial variation of productivity affects the fundamental processes of biodiversity. Based on widely discussed verbal models in ecology about the effect of productivity on species diversity, we developed a spatially explicit neutral model that incorporates the effect of primary productivity on community size and confronted our model's predictions with observed patterns of species richness and evolutionary history of Australian terrestrial mammals. The imposed restrictions on community size create larger populations in areas of high productivity, which increases community turnover and local speciation, and reduces extinction. The effect of productivity on community size modeled in our study causes higher accumulation of species diversity in productive regions even in the absence of niche‐based processes. However, such a simple model is not capable of reproducing spatial patterns of mammal evolutionary history in Australia, implying that more complex evolutionary mechanisms are involved. Our study demonstrates that the overall patterns of species richness can be directly explained by changes in community sizes along productivity gradients, supporting a major role of processes associated with energetic constraints in shaping diversity patterns.     

Relationships between body size and geographical range size among Australian mammals: has human impact distorted macroecological patterns?

Extinction and artificial reduction in the size of geographical ranges of many species have occurred extensively across the globe because of human activities. In particular, Australian mammals have suffered heavily in the last two hundred years, with the highest number of reported cases of mammal extinctions anywhere. In the present study, we investigated the extent to which human impact has affected contemporary macroecological patterns in Australian terrestrial mammals. After examining patterns relating to body size and range size among the contemporary mammal fauna, we removed the effects of the last two hundred years of human impact by exploring patterns in the pre‐European assemblage. This permitted us to determine whether contemporary macroecological patterns are distortions of pre‐European patterns. In contrast to the expected pattern of a significant positive relationship between body size and range size, our results showed no significant association for the complete fauna in both cross‐species and phylogenetic analyses, even when data were corrected for species extinctions and range reductions. Analyses within families and among species with the same dietary strategy revealed three significant positive relationships (Macropodidae, Peramelidae, and herbivores) and one significant negative relationship (insectivores) within the contemporary assemblage that disappeared when the pre‐European assemblage was analysed. A positive relationship also emerged in the pre‐European assemblage for the Vombatidae that was not apparent in the contemporary fauna. Thus, correcting for human impact revealed important distortions among contemporary macroecological relationships that have been brought about by human‐induced range reduction and extinction. These findings not only provide further evidence that the Australian continent presents a unique and valuable opportunity with which to test the generality of macroecological patterns, but they also have important ramifications for the analysis and interpretation of contemporary macroecological datasets.     

Diversity,extinction, and threat status in Lagomorphs

A quarter of all lagomorphs (pikas, rabbits, hares and jackrabbits) are threatened with extinction, including several genera that contain only one species. The number of species in a genus correlates with extinction risk in lagomorphs, but not in other mammal groups, and this is concerning because the non‐random extinction of small clades disproportionately threatens genetic diversity and phylogenetic history. Here, we use phylogenetic analyses to explore the properties of the lagomorph phylogeny and test if variation in evolution, biogeography and ecology between taxa explains current patterns of diversity and extinction risk. Threat status was not related to body size (and, by inference, its biological correlates), and there was no phylogenetic signal in extinction risk. We show that the lagomorph phylogeny has a similar clade‐size distribution to other mammals, and found that genus size was unrelated to present climate, topography, or geographic range size. Extinction risk was greater in areas of higher human population density and negatively correlated with anthropogenically modified habitat. Consistent with this, habitat generalists were less likely to be threatened. Our models did not predict threat status accurately for taxa that experience region‐specific threats. We suggest that pressure from human populations is so severe and widespread that it overrides ecological, biological, and geographic variation in extant lagomorphs.     

Biases,gaps, and opportunities in mammalian extinction risk research

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Mammals of Australian islands: factors influencing species richness

Distribution patterns of indigenous non-volant terrestrial mammals on 257 Australian islands were examined in relation to environmental parameters and the effects of human-induced disturbance during prehistoric and historic times on island species numbers. Species occurrence for individual species, for taxonomic and trophic groups, and for all species together was related to environmental parameters using regression analysis and the extreme-value function model. Patterns of occurrence were examined separately within three major biogeographic regions derived by pattern analysis. The number of species known to have occurred on these islands during historic times was adequately predicted from area alone. No statistically significant improvement in predicted species number was gained by including island elevation, mean annual rainfall, isolation from the mainland or the number of potentially competing species present on the island. Similarly, no single factor other than area was found to influence consistently the presence of individual species. We conclude that the occurrence of indigenous non-volant terrestrial mammal species on these islands indicates a relictual rather than equilibrial fauna. Visitation by Aboriginal people during prehistoric times did not significantly increase mammal extinctions on islands. Examination of patterns of species richness for a given area on a regional basis showed that islands in and around Bass Strait and Tasmania (Bass Region) were the most species-rich, islands off the northern coasts were slightly less rich, and islands off the south western coasts had fewest species. This is in contrast to the usual latitudinal gradient in species richness patterns. However, islands off the northern and eastern coasts had an overall greater number of different species. When considered in relation to the number of different species of mammals occurring within each region, islands of a given size in Bass Region typically bore a higher proportion of this species pool than other regions. The Bass Region was found to be particularly rich in macropoid herbivores and dasyurid carnivores and insectivores. Analyses indicated that there is a very strong relationship between the presence of exotics as a whole and the local extinction of native mammals. Many mammal species formerly widespread on the Australian mainland are now restricted totally to islands (nine species) or are threatened with extinction on the mainland and have island populations of conservation significance (ten species). In all, thirty-five islands protect eighteen taxa of Australian threatened mammals. The land-use and management of these islands is of considerable importance to nature conservation. The introduction of exotic mammals to these islands should be prevented; any introductions that occur should be eradicated immediately.     

Biogeographic structure of the northeastern Pacific rocky intertidal: the role of upwelling and dispersal to drive patterns

Identifying the environmental conditions that drive biogeographic structure remains a major challenge of biogeography, evolutionary ecology and increasingly, conservation biology. Here, we use multivariate classification trees to assess the biogeographic structure of northeast Pacific (～ 26–58°N) rocky intertidal species (406 species of algae and invertebrates) from 102 field sites. Random forest analyses are used to assess the importance of 29 environmental variables, encompassing a broad range of potential drivers, to predict biogeographic structure. Analyses are repeated for species with different larval dispersal capabilities and by broad taxonomic categories (invertebrates and algae). Results show that overall biogeographic structure is in general agreement with classic classification schemes, but patterns are variable among species with different larval dispersal capabilities. Random forest models show a very high fit (pseudo r² > 0.94) and indicate that biogeographic structure can be predicted by a relatively modest subset of variables. Upwelling related variables are the best overall predictors of biogeographic structure (nutrient concentrations, sea‐surface temperature, upwelling/downwelling seasonal switch index), but the relative importance of predictors is geographically variable and top predictors are dependent on the type of larval dispersal. Upwelling related variables are more important to predict biogeographic structure for invertebrates with lower‐medium dispersal capabilities and algae, whereas species with high larval dispersal (planktotrophic) are better predicted by a different subset of variables (i.e. salinity, precipitation seasonality). Our results lend support to the influence of coastal upwelling in structuring biogeographic patterns and highlight the potential for climate change‐induced alterations of upwelling regimes to profoundly affect biodiversity at biogeographic scales.     

Climatic niche conservatism and ecological opportunity in the explosive radiation of arvicoline rodents (Arvicolinae,Cricetidae)

Climatic niche conservatism shapes patterns of diversity in many taxonomic groups, while ecological opportunity (EO) can trigger rapid speciation that is less constrained by the amount of time a lineage has occupied a given habitat. These two processes are well studied, but limited research has considered their joint and relative roles in shaping diversity patterns. We characterized climatic and biogeographic variables for 102 species of arvicoline rodents (Arvicolinae, Cricetidae), testing the effects of climatic niche conservatism and EO on arvicoline diversification as lineages transitioned between biogeographic regions. We found that the amount of time a lineage has occupied a precipitation niche is positively correlated with diversity along a precipitation gradient, suggesting climatic niche conservatism. In contrast, shift in diversification rate explained diversity patterns along a temperature gradient. Our results suggest that an indirect relationship exists between temperature and diversification that is associated with EO as arvicoline rodents colonized warm Palearctic environments. Climatic niche conservatism alone did not fully explain diversity patterns under density‐dependence, highlighting the additional importance of EO‐related processes in promoting the explosive radiation in arvicoline rodents and shaping diversity pattern among biogeographic regions and along climatic gradients.     

Did impacts, volcanic eruptions, or climate change affect mammalian evolution?

In recent years, it has become popular to attribute faunal change and mass extinction to impacts, volcanic eruptions, or climatic change. How well do these supposed causes compare to the excellent record of Cenozoic life, especially that of fossil mammals? Two different Cenozoic mammal diversity curves were compared, and important climatic, volcanic, and impact horizons were examined in detail. In no case is there a strong correlation between impacts, eruptions, or climatic events and any episode of mammalian turnover. On the contrary, most of the known impact, eruption, and climatic events of the Cenozoic occurred during intervals of faunal stability. Conversely, episodes of high turnover and faunal change among Cenozoic mammals correlate with no known extrinsic causes. Apparently, extrinsic environmental factors such as impacts, eruptions, and climate change have a minimal effect, and intrinsic biological factors must be more important.     

Defining climate's role in ecosystem evolution: Clues from late quaternary mammals

Mammalian communities alter their taxonomic composition through time as the species composing them change their biogeographic range, become extinct, or evolve into new species. When taxonomic compositions change through these processes, inevitably the links between taxa and communities change too, resulting in evolution from one ecosystem into the next. Late Quaternary examples suggest that on a timescale encompassing a few thousand to a few hundred thousand years (the “multi‐millennial timescale"), climatic change is perhaps the most important driver of ecosystem evolution because it periodically forces biogeographic changes and extinction. Climatic change over this timescale, which essentially slips between “geological time”; and “ecological time”;, is not very closely in phase with population‐level evolution of a species analyzed for this study, the meadow vole Microtus pennsylvanicus; therefore climatic oscillations on the multi‐millennial timescale may not stimulate speciation much. Instead, speciation may contribute to ecosystem evolution independent of climatic change and over a longer time scale.     

Introduced cats Felis catus eating a continental fauna: inventory and traits of Australian mammal species killed

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Body size and activity times mediate mammalian responses to climate change

Model predictions of extinction risks from anthropogenic climate change are dire, but still overly simplistic. To reliably predict at‐risk species we need to know which species are currently responding, which are not, and what traits are mediating the responses. For mammals, we have yet to identify overarching physiological, behavioral, or biogeographic traits determining species' responses to climate change, but they must exist. To date, 73 mammal species in North America and eight additional species worldwide have been assessed for responses to climate change, including local extirpations, range contractions and shifts, decreased abundance, phenological shifts, morphological or genetic changes. Only 52% of those species have responded as expected, 7% responded opposite to expectations, and the remaining 41% have not responded. Which mammals are and are not responding to climate change is mediated predominantly by body size and activity times (phylogenetic multivariate logistic regressions, P < 0.0001). Large mammals respond more, for example, an elk is 27 times more likely to respond to climate change than a shrew. Obligate diurnal and nocturnal mammals are more than twice as likely to respond as mammals with flexible activity times (P < 0.0001). Among the other traits examined, species with higher latitudinal and elevational ranges were more likely to respond to climate change in some analyses, whereas hibernation, heterothermy, burrowing, nesting, and study location did not influence responses. These results indicate that some mammal species can behaviorally escape climate change whereas others cannot, analogous to paleontology's climate sheltering hypothesis. Including body size and activity flexibility traits into future extinction risk forecasts should substantially improve their predictive utility for conservation and management.     

Climatic niche conservatism and the evolutionary dynamics in species range boundaries: global congruence across mammals and amphibians

Aim Comparative evidence for phylogenetic niche conservatism – the tendency for lineages to retain their ancestral niches over long time scales – has so far been mixed, depending on spatial and taxonomic scale. We quantify and compare conservatism in the climatic factors defining range boundaries in extant continental mammals and amphibians in order to identify those factors that are most evolutionarily conserved, and thus hypothesized to have played a major role in determining the geographic distributions of many species. We also test whether amphibians show stronger signals of climatic niche conservatism, as expected from their greater physiological sensitivity and lower dispersal abilities. Location Global; continental land masses excluding Antarctica. Methods We used nearly complete global distributional databases to estimate the climatic niche conservatism in extant continental mammals and amphibians. We characterized the climatic niche of each species by using a suite of variables and separately investigate conservatism in each variable using both taxonomic and phylogenetic approaches. Finally, we explored the spatial, taxonomic and phylogenetic patterns in recent climatic niche evolution. Results Amphibians and mammals showed congruent patterns of conservatism in cold tolerance, with assemblages of escapee species (i.e. those escaping most from the climatic constraints of their ancestors) aggregated in the North Temperate Zone. Main conclusions The relative strength of climatic niche conservatism varies across the variables tested, but is strongest for cold tolerance in both mammals and amphibians. Despite the apparent conservatism in this variable, there is also a strong signal of recent evolutionary shifts in cold tolerance in assemblages inhabiting the North Temperate Zone. Our results thus indicate that distribution patterns of both taxa are influenced by both niche conservatism and niche evolution.     

The influence of past and present climate on the biogeography of modern mammal diversity

Within most terrestrial groups of animals, including mammals, species richness varies along two axes of environmental variation, representing energy availability and plant productivity. This relationship has led to a search for mechanistic links between climate and diversity. Explanations have traditionally focused on single mechanisms, such as variation in environmental carrying capacity or evolutionary rates. Consensus, though, has proved difficult to achieve and there is growing appreciation that geographical patterns of species richness are a product of many interacting factors including biogeographic history and biological traits. Here, we review some current hypotheses on the causes of gradients in mammal richness and range sizes since the two quantities are intimately linked. We then present novel analyses using recent datasets to explore the structure of the environment-richness relationship for mammals. Specifically, we consider the impact of glaciation on present day mammalian diversity gradients. We conclude that not only are multiple processes important in structuring diversity gradients, but also that different processes predominate in different places.     

Variation in behavioral traits of two frugivorous mammals may lead to differential responses to human disturbance

Human activities can lead to a shift in wildlife species’ spatial distribution. Understanding the specific effects of human activities on ranging behavior can improve conservation management of wildlife populations in human‐dominated landscapes. This study evaluated the effects of forest use by humans on the spatial distribution of mammal species with different behavioral adaptations, using sympatric western lowland gorilla and central chimpanzee as focal species. We collected data on great ape nest locations, ecological and physical variables (habitat distribution, permanent rivers, and topographic data), and anthropogenic variables (distance to trails, villages, and a permanent research site). Here, we show that anthropogenic variables are important predictors of the distribution of wild animals. In the resource model, the distribution of gorilla nests was predicted by nesting habitat distribution, while chimpanzee nests were predicted first by elevation followed by nesting habitat distribution. In the anthropogenic model, the major predictors of gorilla nesting changed to human features, while the major predictors of chimpanzee nesting remained elevation and the availability of their preferred nesting habitats. Animal behavioral traits (body size, terrestrial/arboreal, level of specialization/generalization, and competitive inferiority/superiority) may influence the response of mammals to human activities. Our results suggest that chimpanzees may survive in human‐encroached areas whenever the availability of their nesting habitat and preferred fruits can support their population, while a certain level of human activities may threaten gorillas. Consequently, the survival of gorillas in human‐dominated landscapes is more at risk than that of chimpanzees. Replicating our research in other sites should permit a systematic evaluation of the influence of human activity on the distribution of mammal populations. As wild animals are increasingly exposed to human disturbance, understanding the resulting consequences of shifting species distributions due to human disturbance on animal population abundance and their long‐term survival will be of growing conservation importance.     

Tree species distribution in temperate forests is more influenced by soil than by climate

Knowledge of the ecological requirements determining tree species distributions is a precondition for sustainable forest management. At present, the abiotic requirements and the relative importance of the different abiotic factors are still unclear for many temperate tree species. We therefore investigated the relative importance of climatic and edaphic factors for the abundance of 12 temperate tree species along environmental gradients. Our investigations are based on data from 1,075 forest stands across Switzerland including the cold‐induced tree line of all studied species and the drought‐induced range boundaries of several species. Four climatic and four edaphic predictors represented the important growth factors temperature, water supply, nutrient availability, and soil aeration. The climatic predictors were derived from the meteorological network of MeteoSwiss, and the edaphic predictors were available from soil profiles. Species cover abundances were recorded in field surveys. The explanatory power of the predictors was assessed by variation partitioning analyses with generalized linear models. For six of the 12 species, edaphic predictors were more important than climatic predictors in shaping species distribution. Over all species, abundances depended mainly on nutrient availability, followed by temperature, water supply, and soil aeration. The often co‐occurring species responded similar to these growth factors. Drought turned out to be a determinant of the lower range boundary for some species. We conclude that over all 12 studied tree species, soil properties were more important than climate variables in shaping tree species distribution. The inclusion of appropriate soil variables in species distribution models allowed to better explain species' ecological niches. Moreover, our study revealed that the ecological requirements of tree species assessed in local field studies and in experiments are valid at larger scales across Switzerland.