Body mass explains characteristic scales of habitat selection in terrestrial mammals

Niche theory in its various forms is based on those environmental factors that permit species persistence, but less work has focused on defining the extent, or size, of a species' environment: the area that explains a species' presence at a point in space. We proposed that this habitat extent is identifiable from a characteristic scale of habitat selection, the spatial scale at which habitat best explains species' occurrence. We hypothesized that this scale is predicted by body size. We tested this hypothesis on 12 sympatric terrestrial mammal species in the Canadian Rocky Mountains. For each species, habitat models varied across the 20 spatial scales tested. For six species, we found a characteristic scale; this scale was explained by species' body mass in a quadratic relationship. Habitat measured at large scales best-predicted habitat selection in both large and small species, and small scales predict habitat extent in medium-sized species. The relationship between body size and habitat selection scale implies evolutionary adaptation to landscape heterogeneity as the driver of scale-dependent habitat selection.     

Scale-specific correlations between habitat heterogeneity and soil fauna diversity along a landscape structure gradient

Habitat heterogeneity contributes to the maintenance of diversity, but the extent that landscape-scale rather than local-scale heterogeneity influences the diversity of soil invertebrates—species with small range sizes—is less clear. Using a Scottish habitat heterogeneity gradient we correlated Collembola and lumbricid worm species richness and abundance with different elements (forest cover, habitat richness and patchiness) and qualities (plant species richness, soil variables) of habitat heterogeneity, at landscape (1 km²) and local (up to 200 m²) scales. Soil fauna assemblages showed considerable turnover in species composition along this habitat heterogeneity gradient. Soil fauna species richness and turnover was greatest in landscapes that were a mosaic of habitats. Soil fauna diversity was hump-shaped along a gradient of forest cover, peaking where there was a mixture of forest and open habitats in the landscape. Landscape-scale habitat richness was positively correlated with lumbricid diversity, while Collembola and lumbricid abundances were negatively and positively related to landscape spatial patchiness. Furthermore, soil fauna diversity was positively correlated with plant diversity, which in turn peaked in the sites that were a mosaic of forest and open habitat patches. There was less evidence that local-scale habitat variables (habitat richness, tree cover, plant species richness, litter cover, soil pH, depth of organic horizon) affected soil fauna diversity: Collembola diversity was independent of all these measures, while lumbricid diversity positively and negatively correlated with vascular plant species richness and tree canopy density. Landscape-scale habitat heterogeneity affects soil diversity regardless of taxon, while the influence of habitat heterogeneity at local scales is dependent on taxon identity, and hence ecological traits, e.g. body size. Landscape-scale habitat heterogeneity by providing different niches and refuges, together with passive dispersal and population patch dynamics, positively contributes to soil faunal diversity. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Resource competition and community response to fertilization: the outcome depends on spatial strategies

Decreases in plant species richness and shifts in community structure following fertilization are usually attributed to increasing light limitation. However, there is increasing evidence that light limitation alone does not account for all of the observed effects of fertilization on plant communities. We present a model of competition for a single, spatially heterogeneous resource that shows fertility-mediated changes in community structure without light competition. This model predicts that in a low-productivity spatially heterogeneous habitat, species that interact with the resource environment over small spatial scales may exclude species that experience the environment at larger spatial scales, even when the latter species are better resource competitors in a uniform environment (have a lower R*). Increasing overall habitat fertility under these conditions minimizes the effects of spatial heterogeneity on the species that forage at a larger spatial scale, resulting in changes in species dominance and the potential for species coexistence. This analysis suggests that considering differences in the spatial scales at which species interact with environmental heterogeneity may help explain observed changes in community structure following fertilization.     

Using animal movement paths to measure response to spatial scale

Animals live in an environment that is patchy and hierarchical. I present a method of detecting the scales at which animals perceive their world. The hierarchical nature of habitat causes movement path structure to vary with spatial scale, and the patchy nature of habitat causes movement path structure to vary throughout space. These responses can be measured by a combination of path tortuousity (measured with fractal dimension) versus spatial scale, the variation in tortuousity of small path segments along the movement path, and the correlation between tortuousities of adjacent path segments. These statistics were tested using simulated animal movements. When movement paths contained no spatial heterogeneity, then fractal D and variance continuously increased with scale, and correlation was zero at all scales. When movement paths contained spatial heterogeneity, then fractal D sometimes showed a discontinuity at transitions between domains of scale, variation showed peaks at transitions, and correlations showed a statistically significant positive value at scales smaller than patch size, decreasing to below zero at scales greater than patch size. I illustrated these techniques with movement paths from deer mice and red-backed voles. These new analyses should help understand how animals perceive and react to their landscape structure at various spatial scales, and to answer questions about how habitat structure affects animal movement patterns.     

Does macrophyte fractal complexity drive invertebrate diversity,biomass and body size distributions?

Habitat structure is one of the fundamental factors determining the distribution of organisms at all spatial scales, and vegetation is of primary importance in shaping the structural environment for invertebrates in many systems. In the majority of biotopes, invertebrates live within vegetation stands of mixed species composition, making estimates of structural complexity difficult to obtain. Here we use fractal indices to describe the structural complexity of mixed stands of aquatic macrophytes, and these are employed to examine the effects of habitat complexity on the composition of free-living invertebrate assemblages that utilise the habitat in three dimensions. Macrophytes and associated invertebrates were sampled from shallow ponds in southwest England, and rapid digital image analysis was used to quantify the fractal complexity of all plant species recorded, allowing the complexity of vegetation stands to be reconstructed based on their species composition. Fractal indices were found to be significantly related to both invertebrate biomass–body size scaling and overall invertebrate biomass; more complex stands of macrophytes contained a greater number of small animals. Habitat complexity was unrelated to invertebrate taxon richness and macrophyte surface area and species richness were not correlated with any of the invertebrate community parameters. The biomass–body size scaling relationship of lentic macroinvertebrates matched those predicted by models incorporating both allometric scaling of resource use and the fractal dimension of a habitat, suggesting that both habitat fractal complexity and allometry may control density–body size scaling in lentic macroinvertebrate communities.     

Direct and indirect effects of area,energy and habitat heterogeneity on breeding bird communities

Aim To compare the ability of island biogeography theory, niche theory and species–energy theory to explain patterns of species richness and density for breeding bird communities across islands with contrasting characteristics. Location Thirty forested islands in two freshwater lakes in the boreal forest zone of northern Sweden (65°55′ N to 66°09′ N; 17°43′ E to 17°55′ E). Methods We performed bird censuses on 30 lake islands that have each previously been well characterized in terms of size, isolation, habitat heterogeneity (plant diversity and forest age), net primary productivity (NPP), and invertebrate prey abundance. To test the relative abilities of island biogeography theory, niche theory and species–energy theory to describe bird community patterns, we used both traditional statistical approaches (linear and multiple regressions) and structural equation modelling (SEM; in which both direct and indirect influences can be quantified). Results Using regression‐based approaches, area and bird abundance were the two most important predictors of bird species richness. However, when the data were analysed by SEM, area was not found to exert a direct effect on bird species richness. Instead, terrestrial prey abundance was the strongest predictor of bird abundance, and bird abundance in combination with NPP was the best predictor of bird species richness. Area was only of indirect importance through its positive effect on terrestrial prey abundance, but habitat heterogeneity and spatial subsidies (emerging aquatic insects) also showed important indirect influences. Thus, our results provided the strongest support for species–energy theory. Main conclusions Our results suggest that, by using statistical approaches that allow for analyses of both direct and indirect influences, a seemingly direct influence of area on species richness can be explained by greater energy availability on larger islands. As such, animal community patterns that seem to be in line with island biogeography theory may be primarily driven by energy availability. Our results also point to the need to consider several aspects of habitat quality (e.g. heterogeneity, NPP, prey availability and spatial subsidies) for successful management of breeding bird diversity at local spatial scales and in fragmented or insular habitats.     

Cross-scale Habitat Structure Drives Fish Body Size Distributions on Coral Reefs

Despite a large number of studies focusing on the complexity of coral reef habitats and the characteristics of associated fish assemblages, the relationship between reef structure and fish assemblages remains unclear. The textural discontinuity hypothesis, which proposes that multi-modal body size distributions of organisms are driven by discontinuous habitat structure, provides a theoretical basis that may explain the influence of habitat availability on associated organisms. In this study we use fractal techniques to characterize patterns of cross-scale habitat complexity, and examine how this relates to body-depth abundance distributions of associated fish assemblages over corresponding spatial scales. Our study demonstrates that: (1) Reefs formed from different underlying substrata exhibit distinct patterns of cross-scale habitat complexity; (2) The availability of potential refuges at different scales correlates with patterns in fish body depth distributions, but habitat structure is more strongly related to the relative abundance of fish in the body depth modes, rather than to the number of modes; (3) As reefs change from coral- to algal-dominated states, the complexity of the underlying reef substratum may change, presenting a more homogenous environment to associated assemblages; (4) Individual fish body depth distributions may be multi-modal, however, these distributions are not static characteristics of the fish assemblage and may change to uni-modal forms in response to changing habitat condition. In light of predicted anthropogenic changes, there is a clear need to improve our understanding of the scale of ecological relationships to anticipate future changes and vulnerabilities.     

Bird Community Composition in a Shaded Coffee Agro-ecological Matrix in Puebla,Mexico: The Effects of Landscape Heterogeneity at Multiple Spatial Scales

This study examined the importance of habitat heterogeneity on the avian community composition, and investigated the scale at which species abundances respond to habitat variables. The study was conducted within a diverse landscape matrix of a shaded coffee region in Mexico. To detect at which characteristic spatial scale different species and foraging guilds respond most strongly we analyzed the effect of plot-, patch- and landscape-level variables at different spatial extent (i.e., different kilometer radii) on species composition and foraging guilds. We used redundancy analysis to identify species–environment correlations, and to identify predictor variables that best explained the bird community structure, quantified the influence of plot-, patch- and landscape-level variables on the bird community composition. In addition, we used the 4th-corner method to detect significant relationships between the dietary guilds and plot-, patch- and landscape-level variables. We recorded 12,335 individuals of 181 bird species; 105 bird species were recorded foraging within the shaded coffee plantations. We found that plot- and landscape-level variables significantly explained the bird community composition best across all scales, and were significantly correlated with the abundance of the dietary guilds. In contrast, patch-level variables were less important. Habitat composition variables (i.e., coffee, forest and agricultural area) were among the most important predictors. Canopy structure was more important than other vegetation structure variables in explaining dietary guild structure. Hence, the maintenance of a heterogeneous landscape with a high-quality matrix within an agro-ecological region enhances bird conservation.     

Plant species coexistence at local scale in temperate swamp forest: test of habitat heterogeneity hypothesis

It has been suggested that a heterogeneous environment enhances species richness and allows for the coexistence of species. However, there is increasing evidence that environmental heterogeneity can have no effect or even a negative effect on plant species richness and plant coexistence at a local scale. We examined whether plant species richness increases with local heterogeneity in the water table depth, microtopography, pH and light availability in a swamp forest community at three local spatial scales (grain: 0.6, 1.2 and 11.4 m). We also used the variance partitioning approach to assess the relative contributions of niche-based and other spatial processes to species occurrence. We found that heterogeneity in microtopography and light availability positively correlated with species richness, in accordance with the habitat heterogeneity hypothesis. However, we recorded different heterogeneity-diversity relationships for particular functional species groups. An increase in the richness of bryophytes and woody plant species was generally related to habitat heterogeneity at all measured spatial scales, whereas a low impact on herbaceous species richness was recorded only at the 11.4 m scale. The distribution of herbaceous plants was primarily explained by other spatial processes, such as dispersal, in contrast to the occurrence of bryophytes, which was better explained by environmental factors. Our results suggest that both niche-based and other spatial processes are important determinants of the plant composition and species turnover at local spatial scales in swamp forests.     

Body size affects the spatial scale of habitat–beetle interactions

We used abundance data and the program Focus to determine the spatial scale at which 31 species of longhorned beetles (Coleoptera: Cerambycidae) respond to forest habitat amount. We predicted that the spatial scale at which species respond would increase with body size, and that species using ephemeral larval habitat would respond at larger spatial scales than species using more stable larval habitat. We found that forest cover was a better measure of the amount of habitat for polyphagous species than for oligophagous species. Larger species of longhorned beetles responded to forest cover at larger scales. We did not find evidence that species using more ephemeral larval habitat conditions responded at larger scales than species developing in more stable habitat conditions. Our results highlight the importance of accurately describing habitat in studies of species–environment relationships. While scales of response may be species-specific, some generalizations across species are possible.     

The relationship between multi‐scale habitat heterogeneity and farmland bird abundance

It has recently been proposed that losses in farmland habitat heterogeneity may have been a primary driver of the profound declines exhibited by many farmland bird species in recent decades. However, it has yet to be demonstrated which facets of heterogeneity and what spatial scales are most important for birds. Here we analyse the relationship between abundance and features of landscape heterogeneity at three spatial scales (1, 9 and 25 km²) for 32 bird species commonly associated with farmland. Heterogeneity was quantified using three contrasting indices reflecting 1) the spatial mixing of land uses, 2) variation in field sizes and 3) the density of field boundaries. The spatial mixing of land‐uses explained, on average, the most variation in, and was most likely to be positively associated with, abundance at all spatial scales. The majority of species (66–75%, depending on the spatial scale) were more common in heterogeneous landscapes overall; however, migrants, those under a high level of conservation concern and farmland specialist species tended to be less abundant in more heterogeneous landscapes at all scales. Ground‐nesting species were also more likely to be found in more homogeneous habitats than non‐ground‐nesters, but only at the finest spatial scale. Relationships between abundance and heterogeneity were generally consistent across spatial scales; however, species of high conservation concern had more variable associations compared with other species. These results highlight a potential role for farmland habitat heterogeneity in determining the abundance of many farmland species but suggest that population responses to an increase in heterogeneity would not be unanimously positive and would probably have negative impacts on some species, notably those that are already threatened.     

Anthropogenic modification disrupts species co‐occurrence in stream invertebrates

The question of whether species co‐occurrence is random or deterministic has received considerable attention, but little is known about how anthropogenic disturbance mediates the outcomes. By combining experiments, field surveys and analysis against null models, we tested the hypothesis that anthropogenic habitat modification disrupts species co‐occurrence in stream invertebrates across spatial scales. Whereas communities in unmodified conditions were structured deterministically with significant species segregation, catchment‐scale conversion to agriculture and sediment deposition at the patch‐ or micro‐habitat scale apparently randomized species co‐occurrences. This shift from non‐random to random was mostly independent of species richness, abundance and spatial scale. Data on community‐wide life‐history traits (body size, dispersal ability and predatory habits) and beta‐diversity indicated that anthropogenic modification disrupted community assembly by affecting biotic interactions and, to a lesser extent, altering habitat heterogeneity. These data illustrate that the balance between predictable and stochastic patterns in communities can reflect anthropogenic modifications that not only transcend scales but also change the relative forces that determine species coexistence. Research into the effects of habitat modification as a key to understanding global change should extend beyond species richness and composition to include species co‐occurrence, species interactions and any functional consequences.     

Spatial variability of the soil seed bank in a heterogeneous ephemeral wetland system in semi-arid Australia

Soil seed banks are an important component of plant community diversity in ephemeral wetlands, allowing many species to persist through unpredictable periods of flood and drought. Spatial variation of extant vegetation in such habitats commonly reflects patterns of flood history and often varies predictably between broadly differing hydro-geomorphic habitat types. Here we investigate whether spatial variation of soil seed banks is similarly controlled by fluvial processes at this scale. Results are presented from a seedling emergence trial using samples collected from a range of habitat types, and at different scales within these, in the ephemeral Narran Lakes system in semi-arid Australia. Composition and structure of soil seed banks varied significantly between habitat types reflecting broad differences in flood frequency. As predicted, germinable seed abundance was found to be highest in intermediately flooded habitats. Variability in soil seed bank composition at a local scale was also found to be influenced by hydrology with greater spatial heterogeneity evident in the river channel as well as amongst the least frequently inundated riparian and floodplain habitats.     

里海西北部Caspian地区草原扩大与啮齿类群落结构的变化

于1997-1999年检测了俄联邦卡尔梅克(Kalmyki)共和国ChernieZemly半干旱草原由10个物种组成的啮齿类群落。该区域的植被特点是家畜大量减少后形成的恢复植被。我们的任务之一就是了解不同啮齿动物的生境需求,以预测不同物种对环境变化的特异性反应。我们也检测了群落空间结构、生态位参数和物种多样性。逐步回归分析表明,各种啮齿动物的空间分布仅部分地决定于已知环境因子(2-6个变量)。虽然方程式具有强显著性,但决定系数R2很低,不超过18%。对于大多数种类,这种结果可由最近草原扩大而来的生境异质性降低解释。物种生境选择条件和资源的明显低水平可能是由物种对环境变化的惯性反应造成的。判别函数分析和主分量分析结果表明,群落结构以及单个物种的生态位参数在不同年间并不稳定。空间分层结构在种间中等和低重叠的高物种多样性年度表现明显。多样性、生态位组成和生态位宽度分析结果表明,不同物种对环境条件和资源变化的反应具有不同的个性化方式。同时,物种对环境动态的明显个性化反应成为物种多样性定向变化的原因。限定空间和相对单一条件下的结果表明,啮齿类α多样性与生境结构复杂性呈正相关,而与生境生产力特征呈负相关。观察表明,卡尔梅克国草原扩大伴随着初级生产力的升高和生境异质性的降低。随着时间的推移会导致大多数草原和半干旱区域啮齿类多样性的降低,并形成动物区系核心.     

毛乌素沙地沙生半灌木群落的空间异质性

在使用 5m× 50 0 0 m样带进行调查获得的植被盖度数据的基础上 ,通过半方差分析和分形分析揭示了毛乌素沙化草地沙生半灌木群落植被盖度的空间变异特点。半方差分析揭示出植被盖度的多尺度变异和等级斑块结构 ,在取样范围内至少存在着 3个等级。分形分析得出毛乌素沙地沙生半木群落植被盖度的分维 D=1 .90 ,接近于 2 ,指示着大部分空间变异发生在较小的尺度上 ;分段分形分析得出不同尺度范围内的分维 ,反映了空间异质性的多尺度变异性。这一结果揭示了荒漠化过程引起的生境破碎化     

Effects of windthrow disturbance on a forest bird community depend on spatial scale

Community structure is expected to be affected by spatial heterogeneity in a landscape. We examined the spatial-scale-dependent effects of windthrow caused by a large typhoon on a forest bird community. Typhoon events of this magnitude are rare in Hokkaido, Japan, occurring only once or twice a century. To assess the “functional spatial scale” at which bird groups (community, species, body-size class, and foraging guild) specifically responded to landscape heterogeneity, the canopy gap rate (CGR, gap percentage) was evaluated at different spatial scales by varying the radius of a circular landscape sector from 100 to 500 m stepwise by 10 m. We then analysed bird community responses, in terms of species richness and abundance, to CGR. Bird species richness did not significantly depend on CGR. In contrast, abundance was significantly dependent on CGR in many groups (species, body-size class, and foraging guild). The guild-level response was clearer than the species-level response, which suggests that the integration and filtration of species traits by guild can reveal a clear response of bird abundance to the extent of canopy gaps. For example, the scale dependence of responses to disturbance clearly varied among body-size classes, where larger birds had larger functional spatial scales. These results reveal that different groups of organisms have different functional spatial scales at which they respond to habitat heterogeneity. Our results also suggest that monitoring only a small number of species could be misleading for conserving biodiversity at the landscape level.     

Spatial pattern of distribution of marine invertebrates within a subtidal community: do communities vary more among patches or plots?

Making links between ecological processes and the scales at which they operate is an enduring challenge of community ecology. Our understanding of ecological communities cannot advance if we do not distinguish larger scale processes from smaller ones. Variability at small spatial scales can be important because it carries information about biological interactions, which cannot be explained by environmental heterogeneity alone. Marine fouling communities are shaped by both the supply of larvae and competition for resources among colonizers—these two processes operate on distinctly different scales. Here, we demonstrate how fouling community structure varies with spatial scale in a temperate Australian environment, and we identify the spatial scale that captures the most variability. Community structure was quantified with both univariate (species richness and diversity) and multivariate (similarity in species composition) indices. Variation in community structure was unevenly distributed between the spatial scales that we examined. While variation in community structure within patch was usually greater than among patch, variation among patch was always significant. Opportunistic taxa that rely heavily on rapid colonization of free space spread more evenly among patches during early succession. In contrast, taxa that are strong adult competitors but slow colonizers spread more evenly among patches only during late succession. Our findings show significant patchiness can develop in a habitat showing no systematic environmental spatial variation, and this patchiness can be mediated through different biological factors at different spatial scales.     

Scale specific determinants of tree diversity in an old growth temperate forest in China

The major processes generating pattern in plant community composition depend upon the spatial scale and resolution of observation, therefore understanding the role played by spatial scale on species patterns is of major concern. In this study, we investigate how well environmental (topography and soil variables) and spatial variables explain variation in species composition in a 25-ha temperate forest in northeastern China. We used new variation partitioning approaches to discover the spatial scale (using multi-scale spatial PCNM variables) at which environmental heterogeneity and other spatially structured processes influence tree community composition. We also test the effect of changing grain of the study (i.e. quadrat size) on the variation partitioning results. Our results indicate that (1) species composition in the Changbai mixed broadleaf-conifer forest was controlled mainly by spatially structured soil variation at broad scales, while at finer scales most of the explained variation was of a spatial nature, pointing to the importance of biotic processes. (2) These results held at all sampling grains. However, reducing quadrat size progressively reduced both spatially and environmentally explained variance. This probably partly reflects increasing stochasticity in species abundances, and the smaller proportion of quadrats occupied by each species, when quadrat size is reduced. The results suggest that environmental heterogeneity (i.e. niche processes) and other biotic processes such as dispersal work together, but at different spatial scales, to build up diversity patterns.     

Experimental effects of habitat fragmentation on old-field canopy insects: community, guild and species responses

We examined the effects of habitat fragmentation on the species distributions, guild membership, and community structure of old-field insects using a fine-scale experimental approach. A continuous 1-ha goldenrod field was fragmented into four treatments that varied in both patch size and degree of isolation. Each treatment was replicated four times and arranged in a Latin square design. Canopy insects in fragmented patches were sampled with sweep nets during early and late summer 1995. The species richness of insects was significantly lower in fragmented than in unfragmented treatments during July, but was similar among treatments in September. Overall community abundance showed no treatment effect during either month. We also found significant row and column effects, suggesting there was spatial heterogeneity in species richness and abundance apart from treatment effects. Differences in species richness during July were primarily due to the loss of rare species in highly fragmented plots. Overall abundance was less responsive to community change because deletions of rare species in fragmented areas were not detected in abundance analyses. Four feeding guilds showed different responses to fragmentation: the species richness of sucking herbivores and the abundance of parasitoids were significantly reduced by fragmentation but predators and chewing herbivores were largely unaffected. Analyses of a subset of individual species within guilds suggest that the greater effects of fragmentation on sucking herbivores and parasitoids may be due to the degree of habitat specificity of guild members. The effects of small-scale habitat fragmentation were therefore detectable at the level of community, guild, and individual species. Changes in species richness, guild structure and species distributions were likely due to differential effects of habitat alteration on individual movements and patch selection rather than dispersal or demographic change. Nonetheless, the selective loss of rare species, differential guild effects and changes in species occupancy that we found in this small-scale experiment are also factors that are likely to operate in fragmented habitats over broader spatial scales. Received: 11 May 1998 / Accepted: 27 September 1998     

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.