Human pressures predict species’ geographic range size better than biological traits

Geographic range size is the manifestation of complex interactions between intrinsic species traits and extrinsic environmental conditions. It is also a fundamental ecological attribute of species and a key extinction risk correlate. Past research has primarily focused on the role of biological and environmental predictors of range size, but macroecological patterns can also be distorted by human activities. Here, we analyse the role of extrinsic (biogeography, habitat state, climate, human pressure) and intrinsic (biology) variables in predicting range size of the world's terrestrial mammals. In particular, our aim is to compare the predictive ability of human pressure vs. species biology. We evaluated the ability of 19 intrinsic and extrinsic variables in predicting range size for 4867 terrestrial mammals. We repeated the analyses after excluding restricted‐range species and performed separate analyses for species in different biogeographic realms and taxonomic groups. Our model had high predictive ability and showed that climatic variables and human pressures are the most influential predictors of range size. Interestingly, human pressures predict current geographic range size better than biological traits. These findings were confirmed when repeating the analyses on large‐ranged species, individual biogeographic regions and individual taxonomic groups. Climatic and human impacts have determined the extinction of mammal species in the past and are the main factors shaping the present distribution of mammals. These factors also affect other vertebrate groups globally, and their influence on range size may be similar as well. Measuring climatic and human variables can allow to obtain approximate range size estimations for data‐deficient and newly discovered species (e.g. hundreds of mammal species worldwide). Our results support the need for a more careful consideration of the role of climate change and human impact – as opposed to species biological characteristics – in shaping species distribution ranges.     

Correlates of species richness in mammals: body size, life history, and ecology

We present the most extensive examination to date of proposed correlates of species richness. We use rigorous phylogenetic comparative techniques, data for 1,692 mammal species in four clades, and multivariate statistics to test four hypotheses about species richness and compare the evidence for each. Overall, we find strong support for the life-history model of diversification. Species richness is significantly correlated with shorter gestation period in the carnivores and large litter size in marsupials. These traits and short interbirth intervals are also associated with species richness in a pooled analysis of all four clades. Additionally, we find some support for the abundance hypotheses in different clades of mammals: abundance correlates positively with species richness in primates but negatively in microchiropterans. Our analyses provide no evidence that mammalian species richness is associated with body size or degree of sexual dimorphism.     

Patterns of life-history traits in open-nesting palearctic passerines as a function of the climatic variation of their ranges

Latitude, a surrogate of climatic conditions, is commonly used in the examination of life-history variation. However, the climatic mechanisms underlying latitudinal life-history variation have only rarely been tested. Here, we test whether the number of climates to which species are subjected in their ranges predicts geographical life-history variation. In particular, we examine whether eurytopic species, the range of which covers more climates, show different reproductive effort to stenotopic species, which are distributed over climatically more homogeneous environments. We examined female body mass, egg mass controlled for female body mass, clutch size and the number of breeding attempts per season for 34 sedentary and short-distance migratory passerine species of the Western Palearctic. For each species, we assessed how many climate zones extend over the species' wintering and breeding ranges. We found that avian body mass, and also clutch size, significantly increases with the number of climatic zones extended over the species' wintering range. In turn, species whose breeding ranges span more climates show more breeding attempts per season. Whereas the mass of a single egg declines, clutch size increases with increasing climatic variation in breeding ranges. Our study suggests that the level of climatic variation over species' ranges during and outside the breeding season might be responsible for variation in life-history traits in open-nesting Western Palearctic passerines.     

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.     

Geographical variation in the 'bottom-up' control of diversity: fleas and their small mammalian hosts

Aim We searched for signs of the ‘bottom‐up’ diversity effect in the association between fleas (Siphonaptera) and their small mammalian hosts (Rodentia, Insectivora and Lagomorpha). We asked (1) whether a strong dependence of flea species richness on host species richness is characteristic for both Palaeoarctic and Nearctic realms; (2) if yes, whether the ratio of host species per flea species along the host diversity gradient is similar between the Palaeoarctic and Nearctic; and (3) whether factors other than host species richness (i.e. geographical position, climate and landscape) might better explain variation in flea species richness than host species richness. Location The study used previously published data on species richness of fleas and their small mammalian hosts from 26 Palaeoarctic and 19 Nearctic regions. Methods We regressed the number of flea species on the number of small mammal species across regions, separately for Palaeoarctic and Nearctic realms, using both non‐transformed data as well as data corrected for the confounding effects of host sampling effort and sampling area. To test whether flea species richness is determined by external factors unrelated to the host, we used stepwise multiple regressions of flea species richness against host species richness and parameters describing the geographical position, climate and relief of a region. Results When non‐transformed data were analysed, flea species richness was positively correlated with host species richness in both the Palaeoarctic and Nearctic, although the slopes of the two regressions differed significantly. After removal of the confounding effects of host sampling effort and sampling area, Palaeoarctic flea species richness remained strongly positively correlated with host species richness, whereas in the Nearctic, flea species richness appeared to be completely independent of host species richness. Results of the multiple regressions using corrected data demonstrated that in the Palaeoarctic, flea species richness was correlated with both the number of host species and the mean altitude of the region, whereas in the Nearctic, flea species richness only tended to be weakly correlated with latitude (however, this correlation turned out to be non‐significant after Bonferroni correction). Main conclusions We found evidence of bottom‐up control of flea diversity in the Palaeoarctic regions only, and not in the Nearctic. We explore several potential explanations for the different patterns observed in the two biogeographical realms, including differences in (1) levels of host specialization, (2) history of host–parasite associations and (3) landscape effects on flea diversification. We conclude that these factors combine to create different macroecological patterns in different biogeographical realms, and that diversity is not governed by the same forces everywhere.     

A Review for Life-history Traits Variation in Frogs Especially for Anurans in China

Environmental variation can promote differentiation in life-history traits in species of anurans. Increased environmental stress usually results in larger age at sexual maturity, older mean age, longer longevity, slower growth, larger body size, and a shift in reproductive allocation from offspring quantity to quality, and a stronger trade-off between offspring size and number. However, previous studies have suggested that there are inconsistent geographical variations in life-history traits among anuran species in China. Hence, we here review the intraspecific patterns and differences in life-history traits(i.e., egg size, clutch size, testes size, sperm length, age at sexual maturity, longevity, body size and sexual size dimorphism) among different populations within species along geographical gradients for anurans in China in recent years. We also provide future directions for studying difference in sperm performance between longer and shorter sperm within a species through transplant experiments and the relationships between metabolic rate and brain size and life-history.     

Species are not most abundant in the centre of their geographic range or climatic niche

The pervasive idea that species should be most abundant in the centre of their geographic range or centre of their climatic niche is a key assumption in many existing ecological hypotheses and has been declared a general macroecological rule. However, empirical support for decreasing population abundance with increasing distance from geographic range or climatic niche centre (distance–abundance relationships) remains fairly weak. We examine over 1400 bird, mammal, fish and tree species to provide a thorough test of distance–abundance relationships, and their associations with species traits and phylogenetic relationships. We failed to detect consistent distance–abundance relationships, and found no association between distance–abundance slope and species traits or phylogenetic relatedness. Together, our analyses suggest that distance–abundance relationships may be rare, difficult to detect, or are an oversimplification of the complex biogeographical forces that determine species spatial abundance patterns.     

Broad‐scale geographic patterns in body size and hind wing development of western Palaearctic carabid beetles(Coleoptera: Carabidae)

Research into large‐scale ecological rules has a long tradition but has received increasing attention over the last two decades. Whereas environmental, especially climatic, influences on the geographic distribution of species traits such as body size are well understood in mammals and birds, our knowledge of the determinants and mechanisms which shape spatial patterns in invertebrate traits is still limited. This study analyzes macroecological patterns in two traits of the highly diverse invertebrate taxon of carabid beetles: body size and hind wing development, the latter being directly linked to species’ dispersal abilities. We tested for potential impacts of environmental variables (spatial, areal, topographic and climate‐related) representing both contemporary conditions and historical processes on large‐scale patterns in the two traits. Regression models revealed hump‐shaped relationships with latitude for both traits in the categories 1) all species, 2) widespread and 3) endemic (restricted‐range) species: body size and the proportion of flightless species increased from northern towards southern Europe and then decreased towards North Africa. The shared and independent influence of environmental factors was analyzed by variation partitioning. While contemporary environmental productivity and stability (represented by measures of ambient energy and water energy balance) had strong positive relationships with carabid body size, patterns in hind wing development were most notably influenced by topography (elevation range). Regions with high elevation range and low historical climate variability (since the last ice age), which likely offer long‐term stable habitats (i.e. glacial refugia), coincide with regions with high proportions of flightless species. Thus geographic patterns in carabid traits tend to be formed not only by recent climate but also by dispersal and historical climate and processes (i.e. glaciations and postglacial colonization).     

Decline of Mammal Species Diversity Along the Yungas Forest of Argentina

The southernmost extension of tropical Andean–Amazonian forests protrudes into the arid to semiarid habitats in northwestern Argentina. We analyzed the decline of species richness of forest mammals along these Yungas forests. In particular, we tested whether species decline is due to a general latitudinal effect and whether the drop of species from the assemblages was independent from species' traits and environmental variables. For these tests, we estimated the geographic range edges of 39 species of forest mammal and estimated the species richness with bands of 30'. First, we compared the slope of the decline of species richness from north to south (5.8 species/degree) with the decline expected from the latitudinal species gradient (1–3 species/degree). The decline in species richness of forest mammals along the Yungas was significantly steeper than expected. Second, with a null model assuming a random drop of mammal species we derived confidence limits for the expected species richness and number of range edges within the bands. None of the forest mammals reached the tip of the forest, in marked contrast to nonforest mammals. More range edges than expected from the null model fell within the bands 23°30'–24°00' S and a band near the tip of the Yungas indicating a nonrandom drop of species. The correlation between vulnerability traits and range edges suggests that processes associated with the availability of resources influence the drop of species. We propose that a suite of macroecological attributes interacting with a decrease in habitat quality determines the pattern of species richness in the Yungas forest.     

Interspecific patterns of species richness, geographic range size, and body size among New World venomous snakes

Many higher taxa exhibit latitudinal gradients in species richness, geographic range size, and body size. However, these variables are often interdependent, such that examinations of univariate or bivariate patterns alone may be misleading. Therefore, I examined latitudinal gradients in, and relationships between, species richness, geographic range size, and body size among 144 species of New World venomous snakes [families Elapidae (coral snakes) and Viperidae (pitvipers)]. Both lineages are monophyletic, collectively span 99° of latitude, and are extremely variable in body size and geographic range sizes. Coral snakes exhibit highest species richness near the equator, while pitviper species richness peaks in Central America. Species – range size distributions were strongly right-skewed for both families. There was little support for Bergmann's rule or Rapoport's rule for snakes of either family, as neither body size nor range size increased significantly with latitude. However, range area and median range latitude were positively correlated above 15° N, indicating a possible "Rapoport effect" at high northern latitudes. Geographic range size was positively associated with body size. Available continental area strongly influenced range size. Comparative (phylogenetically-based) analyses revealed that shared history is a poor predictor of range size variation within clades. Among vipers, trends in geographic range sizes may have been structured more by historical biogeography than by macroecological biotic factors.     

川西平原大足鼠的种群生态学 III.繁殖

在川西平原利用标志重捕、夹捕解剖和半自然条件下笼养的方法,提供了1989～1995年大足鼠种群雄性和雌性个体处于繁殖状态的比例、胎次数、每胎幼仔数、幼年个体的补充和发育等有关繁殖的基本资料。结果显示,大足鼠极灵活的繁殖策略 提高了这个物种幼年个体的存活,使该物种在可预见而频繁的周期性变化的农田环 境中能够保持相对稳定的种群。大足鼠的繁殖特征主要有：处于繁殖状态的雄性比例较大（平均0.491,95%置信区间0.451和0.534）,怀孕雌性的比例较小（平均0.227,95％置信区间0.213和0.301）,较高和相对稳定的繁殖能力（每年4胎,每胎8.245±0.452只）,雌性幼仔较慢的生长和性成熟（4月）,以及幼年个体加入种群与春季的小麦和秋季的水稻成熟相同步。这些繁殖特征使得大足鼠的繁殖活动集中在有利的时间。结果,虽然伴随繁殖活动雌雄性的个体质量下降,但仍能在短暂的时间里恢复到原来的水平。将大足鼠的胎仔数与一些关于中国啮齿动物繁殖参数地理变异 的研究进行了分析比较,发现后者提供的资料尚不足以支持中国所有啮齿动物都有 "胎仔数随纬度增加"的过分简单的一般结论。每一个物种种群都有自己的最优胎 仔数,但最优胎仔数不一定随纬度而增加。我们认为在大尺度上研究如胎仔数这样的生活史性状时,生活史多样性的提法比单一性状的过分简单的一般化更好。     

Seeing the forest for the trees: partitioning ecological and phylogenetic components of Bergmann's rule in European Carnivora

Comparative methods have commonly been applied in macroecological research. However, few methods exist to map and analyze phylogenetic variation in geographical space. Here we develop a general analytical framework to partition the phylogenetic and ecological structures of macroecological patterns in geographic space. As an example, we apply the framework to evaluate interspecific patterns of body size geographic variation (Bergmann's rule) in European Carnivora. We model the components of variance attributable to ecological and phylogenetic effects, and to the shared influence of both factors. Spatial patterns in the ecological component are stronger than those in the original body size data. More importantly, the magnitude of intraspecific body size patterns (as measured by the correlation coefficient between body size and latitude) is significantly correlated with the ecological component across species, providing a unified interpretation for Bergmann's rule at multiple levels of biological hierarchy. This approach provides a better understanding of patterns in macroecological traits and allows improved understanding of their underlying ecological and evolutionary mechanisms.     

Predictors of mammal species richness in KwaZulu-Natal,South Africa

The management of multi-functional landscapes warrants better knowledge of environment-richness associations at varying disturbance levels and habitat gradients. Intensive land-use patterns for agricultural purposes lead to fragmentation of natural habitat resulting in biodiversity loss that can be measured using landscape metrics to assess mammalian richness. Since carnivores and herbivores are likely to show different responses to disturbance, we calculated carnivore, non-carnivore, and total mammal species richness from camera surveys using a first order Jackknife Estimator. Richness was compared along a habitat gradient comprising coastal forest, Acacia thicket, and highland in KwaZulu-Natal, South Africa. We used standardized OLS regression models to identify climatic and disturbance variables, and landscape metrics as predictors of species richness. The estimated total and non-carnivore species richness were highest in coastal forest, while carnivore species richness was highest in highland followed by coastal forest and Acacia thicket. Average monthly maximum temperature was a significant predictor of all richness groups, and precipitation of the wettest month and isothermality determined total and non-carnivore species richness, respectively. These climatic variables possibly limit species distribution because of physiological tolerance of the species. Total mammal richness was determined by mean shape (+) and habitat division (−) while diversity (+) and patch richness (−) explained carnivore species richness. Mean shape index (+) influenced non-carnivore richness. However, habitat division and patch richness negatively influenced total mammal richness. Though habitat patch size and contiguity had a weak positive prediction, these metrics demonstrated the importance of habitat connectivity for maintaining mammal richness. The identification of these climatic and landscape patterns is important to facilitate future landscape management for mammal conservation in forest-mosaics.     

Bergmann's rule and the geography of mammal body size in the Western Hemisphere

Aim To describe the geographical pattern of mean body size of the non‐volant mammals of the Nearctic and Neotropics and evaluate the influence of five environmental variables that are likely to affect body size gradients. Location The Western Hemisphere. Methods We calculated mean body size (average log mass) values in 110 × 110 km cells covering the continental Nearctic and Neotropics. We also generated cell averages for mean annual temperature, range in elevation, their interaction, actual evapotranspiration, and the global vegetation index and its coefficient of variation. Associations between mean body size and environmental variables were tested with simple correlations and ordinary least squares multiple regression, complemented with spatial autocorrelation analyses and split‐line regression. We evaluated the relative support for each multiple‐regression model using AIC. Results Mean body size increases to the north in the Nearctic and is negatively correlated with temperature. In contrast, across the Neotropics mammals are largest in the tropical and subtropical lowlands and smaller in the Andes, generating a positive correlation with temperature. Finally, body size and temperature are nonlinearly related in both regions, and split‐line linear regression found temperature thresholds marking clear shifts in these relationships (Nearctic 10.9 °C; Neotropics 12.6 °C). The increase in body sizes with decreasing temperature is strongest in the northern Nearctic, whereas a decrease in body size in mountains dominates the body size gradients in the warmer parts of both regions. Main conclusions We confirm previous work finding strong broad‐scale Bergmann trends in cold macroclimates but not in warmer areas. For the latter regions (i.e. the southern Nearctic and the Neotropics), our analyses also suggest that both local and broad‐scale patterns of mammal body size variation are influenced in part by the strong mesoscale climatic gradients existing in mountainous areas. A likely explanation is that reduced habitat sizes in mountains limit the presence of larger‐sized mammals.     

Pleistocene climatic changes drive diversification across a tropical savanna

Spatial responses of species to past climate change depend on both intrinsic traits (climatic niche breadth, dispersal rates) and the scale of climatic fluctuations across the landscape. New capabilities in generating and analysing population genomic data, along with spatial modelling, have unleashed our capacity to infer how past climate changes have shaped populations, and by extension, complex communities. Combining these approaches, we uncover lineage diversity across four codistributed lizards from the Australian Monsoonal Tropics and explore how varying climatic tolerances interact with regional climate history to generate common vs. disparate responses to late Pleistocene change. We find more divergent spatial structuring and temporal demographic responses in the drier Kimberley region compared to the more mesic and consistently suitable Top End. We hypothesize that, in general, the effects of species’ traits on sensitivity to climate fluctuation will be more evident in climatically marginal regions. If true, this points to the need in climatically marginal areas to craft more species‐(or trait)‐specific strategies for persistence under future climate change.     

Life history, ecology and parasite community structure in Soviet birds

The paper describes an investigation of parasite richness in relation to host life history and ecology using data from an extensive survey of helminth parasites (cestodes, trematodes and nematodes) in Soviet birds. Correlates of parasite richness (number of parasite species per host species) were sought among 13 life-history variables, 13 ecological variables and one non-biological variable (number of host individuals examined) across a sample of 158 species of host. A statistical method to control for the effects of phylogenetic association was adopted throughout. Parasite richness correlates positively with the number of hosts examined (sample size) in all three parasite groups. Positive correlations (after controlling for the effects of sample size) were also found between host body weight and parasite richness for trematodes and nematodes, but not for cestodes.
A number of ecological variables were associated with parasite richness. However, when the effects of sample size and body weight were controlled for, only a single significant correlation (an association between trematode richness and aquatic habitat) remained. Similarly, a number of significant correlates of parasite richness were found among the life-history variables examined. Though several of these were robust to the confounding effects of sample size, all could be explained by the co-variation between life-history traits and body weight among the host species under investigation.     

Diversity of mammals in the tropical–temperate Neotropics: hotspots on a regional scale

The tropical–temperate interface of the southern Neotropics harbours an interdigitating array of biomes (Puna, Monte, Chaco, Yungas). This topographic and climatically complex region needs urgent conservation efforts, as it is being transformed by human activities at an accelerating pace. We analyse georeferenced field records of mammal species in northwestern Argentina (provinces: Catamarca, Jujuy, Salta, Tucuman) in order to define biodiversity hotspots on the basis of 0.5°× 0.5° grid cells within northwestern Argentina according to total richness of mammal species, richness of megaspecies (species above 10 kg), and endemic species (species restricted to Argentina or neighbouring countries with shared biomes). The mammal fauna of northwestern Argentina is fairly well known (176 species). The biomes differ considerably in species richness (Puna low, Yungas high) and species composition. We found no significant difference between endemic and non-endemic species regarding cell occupancy or body size. Cell occupancy was not correlated to body size. Across grids, species richness, number of megaspecies as well as richness of endemics are all correlated to sampling effort. More than 50% of the species in the region are restricted to one or two biomes. Overall, the species turn-over between biomes in northwestern Argentina is high. Using a simple algorithm we identified 10 grid cells which covered 90% of the total number of recorded species, and contrast them with the protected areas. While the Puna and Yungas biomes are rather well protected, the arid and semiarid Monte and Chaco are in need of urgent attention in biodiversity conservation.     

Are species lists derived from modeled species range maps appropriate for macroecological studies? A case study on data from BIEN

Because survey-based species lists for fine-scale areas are generally lacking, ecologists have been using species lists derived from modeled range maps in macroecological studies. Here I evaluate whether it is appropriate to use species lists derived from modeled range maps in macroecological studies. I compared species lists derived from about 90,000 range maps in the Botanical Information and Ecology Network (BIEN) for over 180 geographic regions (12 broad regions covering the whole of the New World; 172 smaller regions, i.e. Taxonomic Databases Working Group units) with those representing complete or nearly complete species lists obtained from the literature for the regions. I determined the completeness of native species composition and proportion of false presences and exotic (non-native) species in species lists derived from range maps in BIEN. I determined whether errors in species lists derived from modeled range maps affect macroecological inferences by comparing the relationships of α- and β-diversity with geographic and climatic variables between different datasets (i.e. species lists derived from range maps versus complete or nearly complete species lists derived from the literature). The completeness for native species lists derived from range maps was 60–74%, depending on spatial scale. On average, 65% of the species in a species list derived from BIEN for a geographic area are false presences or non-native species to the area. The relationships of α- and β-diversity derived from the BIEN dataset with geographic and climatic variables differ significantly from those derived from complete species lists. This study shows that the results of a macroecological study based on species lists derived from modeled range maps may be substantially incorrect due to high percentages of missing native species, and high numbers of non-native species and false presences. Thus, it may not be appropriate to use species lists derived from modeled range maps in the macroecological studies that require complete or nearly complete species lists.