Multiple environmental determinants of regional species richness and effects of geographic range size

Understanding patterns of species richness at broad geographic extents remains one of the most challenging yet necessary scientific goals of our time. Many hypotheses have been proposed to account for spatial variation in species richness; among them, environmental determinants have played a central role. In this study, we use data on regional bat species richness in the New World to study redundancy and complementarity of three environmental hypotheses: energy, heterogeneity and seasonality. We accomplish this by partitioning variation in species richness among components associated with unique and combined effects of variables from each hypotheses, and by partitioning the overall richness gradient into gradients of species with varying breadths of geographic distribution. These three environmental hypotheses explain most variation in the species richness gradient of all bats, but do not account for all positive spatial autocorrelation at short distances. Although environmental predictors are highly redundant, energy and seasonality explain different and complementary fractions of variation in species richness of all bats. On the other hand, heterogeneity variables contribute little to explain this gradient. However, results change dramatically when richness is estimated for groups of species with different sizes of geographic distribution. First, the amount of variation explained by environment decreases with a decrease in range size; this suggests that richness gradients of small‐ranged species can not be explained as easily as those of broadly distributed species, as has been implied by analyses that do not consider differences in range size among species. Second, the relative contribution of environmental predictors to explained variation also changes with change in range size. Seasonality and energy are good predictors of species with broad distributions, but they loose almost all explanatory power for richness of species with small ranges. In contrast, heterogeneity, which is a relatively poor predictor of richness of species with large ranges, becomes the main predictor of richness gradients of species with restricted distributions. This suggests that range size is a different dimension on which heterogeneity and other environmental characteristics are complementary to each other. Our results suggest that determinants of species richness gradients might be complex, or at least more complex than many studies have previously suggested.     

Butterfly,spider, and plant communities in different land-use types in Sardinia,Italy

Butterfly, spider, and plant species richness and diversity were investigated in five different land-use types in Sardinia. In 16 one-hectare plots we measured a set of 15 environmental variables to detect the most important factors determining patterns of variation in species richness, particularly endemicity. The studied land-use types encompassed homogeneous and heterogeneous shrublands, shrublands with tree-overstorey, Quercus forest and agricultural land. A total of 30 butterfly species, among which 10 endemics, and 50 spider (morpho)species, were recorded. Butterfly and spider community composition differed according to land-use type. The main environmental factors determining diversity patterns in butterflies were the presence of flowers and trees. Spiders reacted mainly to habitat heterogeneity and land-use type. Traditional land-use did not have adverse effects on the diversity of butterflies, spiders, or plants. The number of endemic butterfly species per treatment increased with total species richness and altitude. Butterfly and spider richness did not co-vary across the five land-use types. Butterflies were, however, positively associated with plant species richness and elevation, whereas spiders were not. Conclusively, butterflies did not appear to be good indicators for spider diversity and species richness at the studied sites.     

Environmental correlates of leguminosae species richness in Mexico: Quantifying the contributions of energy and environmental seasonality

Explaining species richness patterns is a central issue in ecology, but a general explanation remains elusive. Environmental conditions have been proposed to be important drivers of these patterns, but we still need to better understand the relative contribution of environmental factors. Here, we aim at testing two environmental hypotheses for richness gradients: energy availability and environmental seasonality using diversity patterns of the family Leguminosae across Mexico. We compiled a data base of 502 species and 32,962 records. After dividing Mexico into 100 × 100 km grid cells, we constructed a map of variation in species richness that accounts for heterogeneity in sampling effort. We found the cells with the highest species richness of legumes are in the Neotropical region of Pacific coastal and southern Mexico, where the legume family dominates the tropical rain forests and seasonally dry tropical forests. Regression models show that energy and seasonality predictors can explain 25% and 49% of the variation in richness, respectively. Spatial autocorrelation analysis showed that richness has a strong spatial structure, but that most of this structure disappears when both energy and seasonality are used to account for richness gradient. Our study demonstrates multiple environmental conditions contribute complementarily to explain diversity gradients. Moreover, it shows that in some regions, environmental seasonality can be more important than energy availability, contradicting studies at coarser spatial scales. More basic taxonomic and floristic work is needed to help describe patterns of diversity for many groups to allow for testing the underlying mechanisms responsible for diversity gradients. Abstract in Spanish is available with online material.     

中国蚂蚁丰富度地理分布格局及其与环境因子的关系

物种丰富度分布格局及其形成机制的研究对于生物多样性保护具有重要意义。为了了解中国蚂蚁物种丰富度分布格局,利用中国省级尺度蚂蚁物种分布数据和环境信息,结合GIS和数理统计方法,探讨蚂蚁物种丰富度的地理分布格局与环境因子之间的关系。研究结果表明:(1)蚂蚁丰富度随纬度增加呈逐渐递减趋势,但缺乏显著的经度梯度。丰富度最高的地区主要集中在南方省份,我国北方、西北干旱区和青藏高原北部地区丰富度较低;(2)简单线性回归分析表明,能量、水分和季节性因素中,影响蚂蚁物种丰富度最强的因子分别为最冷月均温(TEMmin)(R2adj=0.532)、年均降水量(PREC)(R2adj=0.376)和年温度变化范围(TEMvar)(R2adj=0.539),而单个生境异质性因子对蚂蚁物种丰富度的影响均不显著;(3)最优模型由年均温(TEM)、海拔变化范围(ELErange)和年温度变化范围(TEMvar)组成,能够解释68.4%的蚂蚁丰富度地理分异。鉴于海拔变化范围更多地反映与温度相关的生境异质性,因此温度是限制中国蚂蚁分布的最重要因素。另外,分析结果还表明,海南、贵州、江西、四川、安徽和山西等6省蚂蚁区系调查最不充分,是未来发现蚂蚁新分布的热点地区。     

Grasshopper and butterfly local congruency in grassland remnants

Grasshoppers could be considered as appropriate ecological indicators for grasslands owing to their sensitive response to environmental features. However, if grasshoppers are a good ecological indicator, they must (i) also represent other taxa, and (ii) provide additional information over straight measurement of environmental variables. To assess this, we compared the congruence of species richness patterns of grasshoppers with butterflies and environmental variables in two areas with extensive ecological networks (ENs). ENs are landscape-scale remnants of corridors and nodes of natural habitat running throughout a transformed, usually agricultural, landscape. Species richness of grasshoppers and butterflies did not differ among reference and EN sites, but guild composition differed significantly. While ENs adequately conserved overall diversity of these two groups, they were utilized preferentially by small-sized grasshoppers and shrub and tree-feeding butterflies. Reference sites had significantly more graminivorous and intermediate-mobility grasshopper species, as well as more butterfly species with widespread distribution, herbaceous dicot feeders and those with no recorded association to forest edges. Nevertheless, grasshopper and butterfly species richness’ were highly correlated. These results were similar across geographic areas, despite the fact that the areas differed significantly in their overall richness and species composition. Although there were some specific significant correlations between environmental variables and diversity, none of the variables could adequately replace use of the insect assemblage for bioindication. We conclude that grasshopper species and guild richness are representative of the butterfly assemblage, and provide information which is not sufficiently clear when utilizing only environmental variables.     

Ecological biogeography of Mexican bats: the relative contributions of habitat heterogeneity,beta diversity,and environmental gradients to species richness and composition patterns

Mexico has higher mammalian diversity than expected for its size and geographic position. High environmental hetero geneity throughout Mexico is hypothesized to promote high turnover rates (β‐diversity), thus contributing more to observed species richness and composition than within‐habitat (α) diversity. This is true if species are strongly associated with their environments, such that changes in environmental attributes will result in changes in species composition. Also, greater heterogeneity in an area will result in greater species richness. This hypothesis has been deemed false for bats, as their ability to fly would reduce opportunities for habitat specialization. If so, we would expect no significant relationships between 1) species composition and environmental variables, 2) species richness and environmental heterogeneity, 3) β‐diversity and environmental heterogeneity. We tested these predictions using 31 bat assemblages distributed across Mexico. Using variance partitioning we evaluated the relative contribution of vegetation, climate, elevation, horizontal heterogeneity (a variate including vegetation, climate, and elevational heterogeneity), spatial variation (lat‐long), and vertical hetero geneity (of vegetation strata) to variation in bat species composition and richness. Variation in vegetation explained 92% of the variation in species composition and was correlated with all other variables examined, indicating that bats respond directly to habitat composition and structure. Beta‐diversity and vegetational heterogeneity were significantly correlated. Bat species richness was significantly correlated with vertical, but not horizontal, heterogeneity. Nonetheless, neither horizontal nor vertical heterogeneity were random; both were related to latitude and to elevation. Variation in bat community composition and richness in Mexico were primarily explained by local landscape heterogeneity and environmental factors. Significant relationships between β‐diversity and environmental variation reveal differences in habitat specialization by bats, and explain their high diversity in Mexico. Understanding mechanisms acting along environmental or geographic gradients is as important for understanding spatial variation in community composition as studying mechanisms that operate at local scales.     

Does herbivore diversity depend on plant diversity? The case of California butterflies

It is widely believed that the diversity of plants influences the diversity of animals, and this should be particularly true of herbivores. We examine this supposition at a moderate spatial extent by comparing the richness patterns of the 217 butterfly species resident in California to those of plants, including all 5,902 vascular plant species and the 552 species known to be fed on by caterpillars. We also examine the relationships between plant/butterfly richness and 20 environmental variables. We found that although plant and butterfly diversities are positively correlated, multiple regression, path models, and spatial analysis indicate that once primary productivity (estimated by a water-energy variable, actual evapotranspiration) and topographical variability are incorporated into models, neither measure of plant richness has any relationship with butterfly richness. To examine whether butterflies with the most specialized diets follow the pattern found across all butterflies, we repeated the analyses for 37 species of strict monophages and their food plants and found that plant and butterfly richness were similarly weakly associated after incorporating the environmental variables. We condude that plant diversity does not directly influence butterfly diversity but that both are probably responding to similar environmental factors.     

Butterfly species and traits associated with selectively logged forest in Borneo

Logging can significantly change the structure of rainforest communities. To better understand how logging drives this change, butterflies and environmental variables were assessed within both unlogged and logged forest in Indonesian Borneo. In the whole dataset, we found local environmental variables and geographic distance combined captured 53.1% of the variation in butterfly community composition; 29.6% was associated with measured local environmental variables, 13.6% with geographic distance between sites, and 9.9% with covariation between geographic distance and environmental variables. The primary axis of variation in butterfly community composition represented a disturbance gradient from unlogged to logged forest. Subsequent axes represented gradients influenced by variables such as canopy cover and total tree density. There were significant associations between environmental variables and geographic range and larval host plant use of species. Specifically, butterflies using trees as larval host plants and those with distributions limited to Borneo were more likely to be present in unlogged forest. By contrast, species that tended to be more abundant in logged forest were those with widespread distributions and those using lianas and grasses as larval host plants. The results of this study highlight the importance of environmental variables and disturbance, e.g., selective logging, in structuring rainforest community diversity. Moreover, they confirm how species traits, such as larval food use and geographic distributions can determine patterns of species abundance following environmental change.     

Latitudinal diversity gradients in bryophytes and woody plants: Roles of temperature and water availability

It remains unclear whether the latitudinal diversity gradients of micro- and macro-organisms are driven by the same macro-environmental variables. We used the newly completed species catalog and distribution information of bryophytes in China to explore their spatial species richness patterns, and to investigate the underlying roles of energy availability, climatic seasonality, and environmental heterogeneity in shaping these patterns. We then compared these patterns to those found for woody plants. We found that, unlike woody plants, mosses and liverworts showed only weakly negative latitudinal trends in species richness. The spatial patterns of liverwort richness and moss richness were overwhelmingly explained by contemporary environmental variables, although explained variation was lower than that for woody plants. Similar to woody plants, energy and climatic seasonality hypotheses dominate as explanatory variables but show high redundancy in shaping the distribution of bryophytes. Water variables, that is, the annual availability, intra-annual variability and spatial heterogeneity in precipitation, played a predominant role in explaining spatial variation of species richness of bryophytes, especially for liverworts, whereas woody plant richness was affected most by temperature variables. We suggest that further research on spatial patterns of bryophytes should incorporate the knowledge on their ecophysiology and evolution.     

Butterfly species richness in the north-west Mediterranean Basin: the role of natural and human-induced factors

Aim We developed a model enabling us to evaluate the contribution of both natural and human‐related factors to butterfly species richness in Catalonia, a Mediterranean area that harbours one of the most diverse butterfly faunas in Europe. Location The study was carried out in Catalonia (north‐east Iberian Peninsula), a region of 31,930 km² lying between the Pyrenees, the Ebro depression and the Mediterranean sea. Methods Data from the Catalan Butterfly Monitoring Scheme were used to assess butterfly species richness from 55 transects spread all over the region. Three groups of environmental variables likely to affect the presence of butterfly species were calculated, above all from geographic information system data: (1) climatology and topography, (2) vegetation structure and (3) human disturbance. Because climatic and topographic variables are expected to be strongly correlated, we first performed a principal component analysis (PCA) to create a summarizing factor that would account for most of the variance within this set of variables. Subsequently, a backward stepwise multiple regression was performed in order to assess the effects of environmental factors on butterfly species richness. Results A total of 131 species were detected in the monitoring transects, representing 75.7% of the butterfly fauna known from Catalonia. Mean species richness per transect and per year was 41.4, although values varied greatly among sites (range: 14–76.8). The final regression model explained more than 80% of the total variance, which indicated a strong association between butterfly species richness and the studied environmental factors. The model revealed the very important contribution of climatic and topographic variables, which were combined into a single factor in the PCA. In contrast to what has been found in other, more northerly countries, species richness was negatively correlated with temperature and positively correlated with rainfall, except for extreme cold and wet conditions. This may be a consequence of the predictably adverse effects of the Mediterranean summer drought on herbivorous insects, and the fact that the limits of distribution of many butterflies correlate well with climatic variables. Human disturbance (defined as the proportion of urban and agricultural landscape cover in buffer areas of 5 km around the transect sites) was the second most important predictor for species richness. We found that a significant decrease in species numbers was associated with an increase in human pressure, a finding that indicates that not only building development, but also modern‐day agricultural practices are detrimental to the conservation of Mediterranean butterflies. Surprisingly, vegetation variables had an almost negligible effect on butterfly species richness. Main conclusions Our findings strongly indicate that the current motors of global change will have a negative effect on Mediterranean butterfly assemblages. First, changes in land‐use are transforming and fragmenting the landscape into an inhospitable and less permeable matrix for butterflies. Secondly, the negative correlation between species richness and temperature will lead to a predictable loss of diversity over the coming years, as predicted in the most plausible scenarios of climate change. Considering the high butterfly richness characterizing the Mediterranean Basin, this future trend will pose a serious threat to biodiversity.     

Urbanization promotes the loss of seasonal dynamics in the semi-natural grasslands of an East Asian megacity

The biodiversity of agricultural landscapes has been noticeably affected by rapid urbanization. Although many studies have examined species diversity per unit area (alpha diversity), knowledge about the patterns of species turnover (beta diversity) in urban areas remains limited. Furthermore, most beta diversity studies have focused on spatial heterogeneity; however, losses of temporal heterogeneity resulting from urbanization remain limited. In this study, we examined how urbanization is associated with decreases in the seasonal heterogeneity of species composition, which could be used as an indicator of the loss of seasonality by ecologists and policy makers aiming to conserve biodiversity. We investigated (1) changes in species richness based on seasonal averages (alpha diversity) and (2) the seasonal turnover of species composition (beta diversity) for flowering plants and butterflies along a rural-urban gradient in semi-natural grasslands. The response variables were alpha and beta diversity for flowering plants and butterflies, and the explanatory variables were urban areas within a 1-km radius of the center of each site. Increasing urban area caused both the seasonal alpha and beta diversity of flowering plants and butterflies to decline. These results supported the homogenization hypothesis for the seasonality of plants and butterflies in semi-natural grasslands of dominant urban areas in East Asia. Future studies should focus on investigating how urbanization is causing both declines in seasonality and changes in the spatial heterogeneity of species composition and associated biodiversity loss. Ecologists and policy makers should focus on developing strategies to halt the loss of temporal biological heterogeneity to maintain biodiversity.     

Butterfly diversity and historical land cover change along an altitudinal gradient

Land cover and climate change are both major threats for biodiversity. In mountain ecosystems species have to adapt to fragmented habitats and harsh environmental conditions but so far, altitudinal effects in combination with land cover change have been rarely studied. The objective of this study was to determine the effects of altitude and historical land cover change on butterfly diversity. We studied species richness patterns of butterflies occuring in wetlands and other open habitats along an altitudinal gradient in a low mountain region (340–750 m a.s.l., Bavaria, Germany) with drastic loss of open habitats within the last 40–60 years. We recorded in 27 sites a total of 4,523 individuals of 49 butterfly species and five species of burnet moths. Species richness peaked at mid elevation and increased with patch size. Land cover change was most pronounced at high altitudes, but neither current open habitats, nor the historical loss of open habitats affected the species richness of butterflies. Neither open land specialized butterflies nor generalist and forest species were significantly affected by the loss of open habitats. However, increasing forest area in high altitudes reduces possible refuge open habitats for butterflies at their thermal distribution limits. This could lead to extinction of such butterfly species when temperatures further rise due to global warming.     

宁夏贺兰山自然保护区蝴蝶群落多样性及其环境影响因子

宁夏贺兰山自然保护区蝴蝶群落多样性及其与环境因素的关系,2017年5-9月采用样线法对贺兰山东麓6类生境和不同干扰类型10条样线的蝴蝶群落结构及其多样性季节动态进行调查。共记录蝴蝶5科36属45种,蛱蝶科Nymphalidae的属和物种数最多,为17属19种;凤蝶科Papilionidae最少,仅1属1种。菜粉蝶Pieris rapae、云粉蝶Pontia daplidice、斑缘豆粉蝶Colias erate和小檗绢粉蝶Aporia hippia是该地区的优势种,个体数量分别占总个体数的11.76%、11.63%、11.21%和10.17%。不同生境样线优势类群和常见类群不同。蝴蝶的栖息地偏好与寄主植物有关,蝴蝶的生境分布类型可分为生境广布型、湿润平原型、荒漠半荒漠草原型和山地森林型。蝴蝶群落Shannon-Wiener多样性和丰富度指数以灰榆疏林草地生境最高,优势度最低。各物种在生境内的季节变化趋势与不同生境植被生长季节相关,高峰期为7-8月。不同调查时间蝴蝶的优势种和常见种不同。物种数以7月份调查最多,有33种,占全年调查总物种数的73.33%;5月份调查最少,有20种。蝴蝶群落Shannon-Wiener多样性和丰富度指数以8月份最大,5月份最小。蝴蝶成虫发生类型分为全年发生型、春季型、夏季型和夏秋季型。不同生境和季节发生的优势种可以作为对生境状况进行评估的指示类群。采用CCA分析物种分布与微环境因子的关系,海拔对蝴蝶物种多样性分布格局有显著影响。蝴蝶丰富度与海拔、温度、风速显著正相关。适度干扰有利于蝶类多样性增加,较强的人为干扰会影响蝶类栖息环境,降低蝶类多样性。因此,生境差异性和干扰与蝴蝶群落的物种多样性密切相关,维持贺兰山垂直植被带的生境异质性和保持适度干扰是保护蝴蝶多样性的关键。     

A test of multiple hypotheses for the species richness gradient of South American owls

Many mechanisms have been proposed to explain broad scale spatial patterns in species richness. In this paper, we evaluate five explanations for geographic gradients in species richness, using South American owls as a model. We compared the explanatory power of contemporary climate, landcover diversity, spatial climatic heterogeneity, evolutionary history, and area. An important aspect of our analyses is that very different hypotheses, such as history and area, can be quantified at the same observation scale and, consequently can be incorporated into a single analytical framework. Both area effects and owl phylogenetic history were poorly associated with richness, whereas contemporary climate, climatic heterogeneity at the mesoscale and landcover diversity explained ca. 53% of the variation in species richness. We conclude that both climate and environmental heterogeneity should be retained as plausible explanations for the diversity gradient. Turnover rates and scaling effects, on the other hand, although perhaps useful for detecting faunal changes and beta diversity at local and regional scales, are not strong explanations for the owl diversity gradient.     

Geographic variations of seasonality and coexistence in communities: The role of diversity and climate

One of the most conspicuous and widely analyzed patterns in ecology is the latitudinal gradient in species richness. Over the 200 years since its recognition, several hypotheses have accumulated in order to account for spatial variations in diversity. Geographic variations in seasonality have been repeatedly proposed as a determinant of community richness. However, the geographic structure of community seasonality has not yet been analyzed. In the present work we evaluated three hypotheses that account for variations in the temporal structuring of communities: first, environmental seasonality determines community seasonality; second, community richness determines its degree of structuring; and third, the presence of an increase in species segregation with latitude, reflected in a pattern of species negative co‐occurrence. The hypotheses were evaluated using path analysis on 29 amphibian communities from South America, connecting latitude, environmental conditions, diversity, seasonality, and coexistence structure – nestedness and negative co‐occurrence – within communities. Latitude positively affects community seasonality through an increase in temperature seasonality, but a weak negative direct effect suggests that other variables not considered in the model – such as the strength of biotic interactions – could also be involved. Both latitude and diversity (directly and indirectly) determine an increase in negative co‐occurrence and nestedness. This suggests that groups of species that are mutually nested in time are internally segregated. Further, the strength of this structure is determined by community diversity and latitude. Temporal structuring of a community is associated with latitude and diversity, pointing to the existence of a systematic change in community organization far beyond, but probably interrelated, with the recognized latitudinal trend in richness. The available information and analysis supported the three hypotheses evaluated.     

Bird diversity patterns in Neotropical temperate farmlands: The role of environmental factors and trophic groups in the spring and autumn

Productivity, habitat heterogeneity and environmental similarity are of the most widely accepted hypotheses to explain spatial patterns of species richness and species composition similarity. Environmental factors may exhibit seasonal changes affecting species distributions. We explored possible changes in spatial patterns of bird species richness and species composition similarity. Feeding habits are likely to have a major influence in bird–environment associations and, given that food availability shows seasonal changes in temperate climates, we expect those associations to differ by trophic group (insectivores or granivores). We surveyed birds and estimated environmental variables along line‐transects covering an E‐W gradient of annual precipitation in the Pampas of Argentina during the autumn and the spring. We examined responses of bird species richness to spatial changes in habitat productivity and heterogeneity using regression analyses, and explored potential differences between seasons of those responses. Furthermore, we used Mantel tests to examine the relationship between species composition similarity and both the environmental similarity between sites and the geographic distance between sites, also assessing differences between seasons in those relationships. Richness of insectivorous birds was directly related to primary productivity in both seasons, whereas richness of seed‐eaters showed a positive association with habitat heterogeneity during the spring. Species composition similarity between assemblages was correlated with both productivity similarity and geographic proximity during the autumn and the spring, except for insectivore assemblages. Diversity within main trophic groups seemed to reflect differences in their spatial patterns as a response to changes between seasons in the spatial patterns of food resources. Our findings suggest that considering different seasons and functional groups in the analyses of diversity spatial pattern could contribute to better understand the determinants of biological diversity in temperate climates.     

Maximized sampling of butterflies to detect temporal changes in tropical communities

There are few papers describing long-term fluctuations and general patterns of temporal diversity in butterfly assemblages in the Neotropical region. The present paper presents a long-term study on the variation in richness and composition of butterflies in a fragment of semi-deciduous forest in Southeastern Brazil, and examines the viability of using maximized butterfly transect counts as a methodology to rapidly and adequately access the local characteristics of butterfly communities. Based on the eight annual standard lists, 518 species in six butterfly families were recorded, representing 74 % of the total butterfly fauna known from the study site. Hesperiidae was the richest family (248 species), followed by Nymphalidae (154), Lycaenidae (49), Riodinidae (29), Pieridae (26), and Papilionidae (12). The accumulation curves show that 8 years of sampling were not enough to result in stable species totals for all butterfly families, especially Hesperiidae and Lycaenidae, which are still increasing in number of species. A great similarity in species composition was observed among all the years (54 %). Comparing the similarity between two standard lists at different time intervals (from 1 to 8 years), a clear pattern of increasing dissimilarity was observed in most families. Our results show that the maximized sampling method is effective in revealing temporal patterns of diversity across several years and could be valuable in monitoring temporal variation in butterfly assemblages for conservation purposes, since the obtained standard lists can be successfully compared to temporal patterns over large periods of time.     

Environmental effects on vertebrate species richness: testing the energy, environmental stability and habitat heterogeneity hypotheses

Background

Explaining species richness patterns is a central issue in biogeography and macroecology. Several hypotheses have been proposed to explain the mechanisms driving biodiversity patterns, but the causes of species richness gradients remain unclear. In this study, we aimed to explain the impacts of energy, environmental stability, and habitat heterogeneity factors on variation of vertebrate species richness (VSR), based on the VSR pattern in China, so as to test the energy hypothesis, the environmental stability hypothesis, and the habitat heterogeneity hypothesis.

Methodology/Principal Findings

A dataset was compiled containing the distributions of 2,665 vertebrate species and eleven ecogeographic predictive variables in China. We grouped these variables into categories of energy, environmental stability, and habitat heterogeneity and transformed the data into 100×100 km quadrat systems. To test the three hypotheses, AIC-based model selection was carried out between VSR and the variables in each group and correlation analyses were conducted. There was a decreasing VSR gradient from the southeast to the northwest of China. Our results showed that energy explained 67.6% of the VSR variation, with the annual mean temperature as the main factor, which was followed by annual precipitation and NDVI. Environmental stability factors explained 69.1% of the VSR variation and both temperature annual range and precipitation seasonality had important contributions. By contrast, habitat heterogeneity variables explained only 26.3% of the VSR variation. Significantly positive correlations were detected among VSR, annual mean temperature, annual precipitation, and NDVI, whereas the relationship of VSR and temperature annual range was strongly negative. In addition, other variables showed moderate or ambiguous relations to VSR.

Conclusions/Significance

The energy hypothesis and the environmental stability hypothesis were supported, whereas little support was found for the habitat heterogeneity hypothesis.     

Can stochastic geographical evolution re‐create macroecological richness–environment correlations?

Aim Richness gradients are frequently correlated with environmental characteristics at broad geographic scales. In particular, richness is often associated with energy and climate, while environmental heterogeneity is rarely its best correlate. These correlations have been interpreted as evidence in favour of environmental determinants of diversity gradients, particularly energy and climate. This interpretation assumes that the expected‐by‐random correlation between richness and environment is zero, and that this is equally true for all environmental characteristics. However, these expectations might be unrealistic. We investigated to what degree basic evolutionary/biogeographical processes occurring independently of environment could lead to richness gradients that correlate with environmental characteristics by chance alone. Location Africa, Australia, Eurasia and the New World. Methods We produced artificial richness gradients based on a stochastic simulation model of geographic diversification of clades. In these simulations, species speciate, go extinct and expand or shift their distributions independently of any environmental characteristic. One thousand two hundred repetitions of this model were run, and the resulting stochastic richness gradients were regressed against real‐world environmental variables. Stochastic species–environment relationships were then compared among continents and among three environmental characteristics: energy, environmental heterogeneity and climate seasonality. Results Simulations suggested that a significant degree of correlation between richness gradients and environment is expected even when clades diversify and species distribute stochastically. These correlations vary considerably in strength; but in the best cases, environment can spuriously account for almost 80% of variation in stochastic richness. Additionally, expected‐by‐chance relationships were different among continents and environmental characteristics, producing stronger spurious relationships with energy and climate than with heterogeneity. Main conclusions We conclude that some features of empirical species–environment relationships can be reproduced just by chance when taking into account evolutionary/biogeographical processes underlying the construction of species richness gradients. Future tests of environmental effects on richness should consider structure in richness–environment correlations that can be produced by simple evolutionary null models. Research should move away from the naive non‐biological null hypotheses that are implicit in traditional statistical tests.