Disentangling scale dependencies in species environmental niches and distributions

Understanding species’ responses to environmental conditions, and how these ­species–environment associations shape spatial distributions, are longstanding goals in ecology and biogeography. However, an essential component of species–environment relationships – the spatial unit, or grain, at which they operate – remains unresolved. We identify three components of scale‐dependence in analyses of species–environment associations: 1) response grain, the grain at which species respond most strongly to their environment; 2) environment spatial structure, the pattern of spatial autocorrelation intrinsic to an environmental factor; and 3) analysis grain, the grain at which analyses are conducted and ecological inferences are made. We introduce a novel conceptual framework that defines these scale components in the context of analyzing species–environment relationships, and provide theoretical examples of their interactions for species with various ecological attributes. We then use a virtual species approach to investigate the impacts of each component on common methods of measuring and predicting species–environment relationships. We find that environment spatial structure has a substantial impact on the ability of even simple, univariate species distribution models (SDMs) to recover known species–­environment associations at coarse analysis grains. For simulated environments with ‘fine’ and ‘intermediate’ spatial structure, model explanatory power, and the frequency with which simple SDMs correctly estimated a virtual species’ response to the simulated environment, dramatically declined as analysis grain increased. Informed by these results, we use a scaling analysis to identify maximum analysis grains for individual environmental factors, and a scale optimization procedure to determine the grain of maximum predictive accuracy. Implementing these analysis grain thresholds and model performance standards in an example east African study system yields more accurate distribution predictions, compared to SDMs independently constructed at arbitrary analysis grains. Finally, we integrate our conceptual framework with virtual and empirical results to provide practical recommendations for researchers asking common questions about species–environment relationships.     

Global patterns of specialization and coexistence in bird assemblages

Aim Increased specialization has been hypothesized to facilitate local coexistence and thus high species richness, but empirical evaluations of the richness–specialization relationships have been relatively scant. Here, we provide a first assessment of this relationship for terrestrial bird assemblages at global extent and from fine to coarse grains. Location World‐wide. Methods We use two indices of specialization that describe species‐level resource use: diet and habitat specialization. The relationship between richness and mean assemblage‐level specialization was independently assessed at realm, biome‐realm, 12,100 km² equal‐area grid cells and fine‐grained scales. To identify assemblages that are diverse relative to environmental conditions we: (1) applied quantile regressions, (2) statistically accounted for other environmental variables which may constrain richness, and (3) parsed the data according to the residuals of a model relating species richness to the environmental variables. Results Assemblage species richness increases with both measures of specialization at all scales. Statistically, richness appears constrained by levels of specialization, with the highest richness values only found in specialized assemblages. Richness is positively associated with specialization even after accounting for gradients in resource availability. Net primary productivity and assemblage specialization have complementary statistical effects on assemblage species richness. Contrary to expectations based on niche partitioning of local resources, the relationship between specialization and richness is steep even at coarse scales. Main conclusions The results demonstrate that for an entire clade, totalling > 9000 species, specialization and species richness are related, at least for diverse assemblages. The strong patterns observed across scales suggest that this relationship does not solely originate from (1) limits on coexistence in present‐day assemblages, or (2) increased specialization in richer assemblages imposed by species’ abilities to partition ecological space. Instead, regional‐scale influences on the species pool may determine much of the observed relationship between richness and specialization. Although causal attribution is not straightforward, these findings support the idea that, for the scale of our analysis, specialization may be related to the past origination of high‐diversity assemblages, rather than their contemporary assembly.     

Multi-scale analysis of plant species richness in Serengeti grasslands

Aim To assess scale dependence between environmental factors and plant species richness. Additionally, we aimed to identify the scales at which niche relations and habitat heterogeneity, as hypothesized by A. Shmida & M.V. Wilson (1985) Journal of Biogeography, 12 , 1–20, operate in the savanna grasslands that were the focus of this study. Location Savanna grassland plant communities of Serengeti National Park, Tanzania. Methods Plant species richness was sampled in 102 modified Whittaker plots and tested for associations with two climate factors, mean annual rainfall (MAP) and potential evapotranspiration (PET), and two landscape variables, plot aspect (ASP) and topographic variation (TOPO), using multiple regressions. Scale dependence was assessed by conducting regressions after altering three aspects of spatial scale: grain, extent and focus. Grain was altered by analysing plant richness at 1, 10, 10² and 10³ m²; extent was investigated by restricting the maximum distance between samples to 75, 100, 125 and 150 km; and focus was manipulated by averaging samples spatially according to geographical land regions. Within the context of our data, we assumed that niche relations were represented by climate factors and habitat heterogeneity by landscape factors. Results Across all 102 plots, plant species richness between 1 and 10² m² had a negative relation to PET and a weak positive relation to MAP. Plant species richness at 10³ m² had a positive association with TOPO and weaker associations with climate factors. ASP stayed in the model between grains of 10 and 10³ m², but had a very weak positive association with richness. When the focus was changed to land regions, associations between plant species richness and explanatory variables strengthened, but were not qualitatively different. At spatial extents of 75 and 100 km, PET was the strongest correlate of plant species richness across all spatial grains. At spatial extents ≥ 125 km, PET explained the majority of the model variance at spatial grains ≤ 10² m², whereas TOPO explained equal amounts or more of the model variance at spatial grains of 10³ m². Main conclusions Both climate and topographic variation explained plant species richness in Serengeti grasslands, but specific patterns depended on grain, extent and, to a lesser degree, focus. Consistent with the ideas of Shmida & Wilson (1985) , determinants of plant species richness shifted from niche relations to habitat heterogeneity between spatial grains of 1 and 10³ m², although this occurred only at relatively large spatial extents (≥ 150 km). Finally, the signs, strength and shape of plant species richness relationships in Serengeti closely match those that describe macro‐scale patterns of woody plant species richness across the entire African continent.     

Effect of sampling grain on patterns of species richness and turnover in Amazonian forests

Grain (size of sampling units) affects the spatial resolution at which ecological patterns can be observed and analysed, and potentially has an important effect on the results of broad‐scale studies on diversity gradients. Here we examine the effect of grain on patterns of species richness and turnover in lowland rainforests of western Amazonia (Peru and Colombia). We inventoried pteridophytes (ferns and lycophytes), melastomes (Melastomataceae) and palms (Arecaceae) in four line transects of 22–29 km length. Different grains were obtained by aggregating original 100‐m‐long sampling units into larger segments up to 19.2 km long. With any given grain and plant group, local species richness varied considerably both within and among transects, and a transect segment that was species‐rich with one grain could be relatively species‐poor with another. Which transect had the highest vs lowest mean species richness per sampling unit (α richness) differed among plant groups. It also varied to some degree with grain, as transects differed in how rapidly local species richness increased with increasing grain. Patterns of species turnover were more consistently correlated among plant groups than patterns of species richness were, and NMDS ordinations were rather similar with all grains and plant groups. Floristic heterogeneity within the Amazonian terra firme rainforest seems to contain a general compositional pattern that is sufficiently robust to be detectable with various sampling schemes, but patterns of species richness appear more case‐specific. Therefore, using one plant group as an indicator for patterns in other plant groups can be expected to work better for species turnover than for species richness.     

Spatial scale, abundance and the species-energy relationship in British birds

1. The spatial scale of analysis may influence the nature, strength and underlying drivers of macroecological patterns, one of the most frequently discussed of which is the relationship between species richness and environmental energy availability. 2. It has been suggested that species-energy relationships are hump-shaped at fine spatial grains and consistently positive at larger regional grains. The exact nature of this scale dependency is, however, the subject of much debate as relatively few studies have investigated species-energy relationships for the same assemblage across a range of spatial grains. Here, we contrast species-energy relationships for the British breeding avifauna at spatial grains of 1 km x 1 km, 2 km x 2 km and 10 km x 10 km plots, while maintaining a constant spatial extent. 3. Analyses were principally conducted using data on observed species richness. While survey work may fail to detect some species, observed species richness and that estimated using nonparametric techniques were strongly positively correlated with each other, and thus exhibit very similar spatial patterns. Moreover, the forms of species-energy relationships using observed and estimated species richness were statistically indistinguishable from each other. 4. Positive decelerating species-energy relationships arise at all three spatial grains. There is little evidence that the explanatory power of these relationships varies with spatial scale. However, ratios of regional (large-scale) to local (small-scale) species richness decrease with increasing energy availability, indicating that local richness responds to energy with a steeper gradient than does regional richness. Local assemblages thus sample a greater proportion of regional richness at higher energy levels, suggesting that spatial turnover of species richness is lower in high-energy regions. Similarly, a crude measure of temporal turnover, the ratio of cumulative species richness over a 4-year period to species richness in a single year, is lower in high-energy regions. These negative relationships between turnover and energy appear to be causal as both total and mean occupancy per species increases with energy. 5. While total density in 1 km x 1 km plots correlates positively with energy availability, such relationships are very weak for mean density per species. This suggests that the observed association between total abundance and species richness may not be mediated by population extinction rates, as predicted by the more individuals hypothesis. 6. The sampling mechanism suggests that species-energy relationships arise as high-energy areas support a greater number of individuals, and that random allocation of these individuals to local areas from a regional assemblage will generate species-energy relationships. While randomized local species-energy relationships are linear and positive, predicted richness is consistently greater than that observed. The mismatch between the observed and randomized species-energy relationships probably arises as a consequence of the aggregated nature of species distributions. The sampling mechanism, together with species spatial aggregation driven by limited habitat availability, may thus explain the species-energy relationship observed at this spatial scale.     

Patterns of beta diversity in Europe: the role of climate,land cover and distance across scales

Aim We test the prediction that beta diversity (species turnover) and the decay of community similarity with distance depend on spatial resolution (grain). We also study whether patterns of beta diversity are related to variability in climate, land cover or geographic distance and how the independent effects of these variables depend on the spatial grain of the data. Location Europe, Great Britain, Finland and Catalonia. Methods We used data on European birds, plants, butterflies, amphibians and reptiles, and data on British plants, Catalonian birds and Finnish butterflies. We fitted two or three nested grids of varying resolutions to each of these datasets. For each grid we calculated differences in climate, differences in land‐cover composition (CORINE) and beta diversity (β_sim, β_Jaccard) between all pairs of grid cells. In a separate analysis we looked specifically at pairs of adjacent grid cells (the first distance class). We then used variation partitioning to identify the magnitude of independent statistical associations (i.e. independent effects in the statistical sense) of climate, land cover and geographic distance with spatial patterns of beta diversity. Results Beta diversity between grid cells at any given distance decreased with increasing grain. Geographic distance was always the most important predictor of beta diversity for all pairwise comparisons at the extent of Europe. Climate and land cover had weaker but distinct and grain‐dependent effects. Climate was more important at relatively coarse grains, whereas land‐cover effects were stronger at finer grains. In the country‐wide analyses, climate and land cover were more important than geographic distance. Climatic and land‐cover models performed poorly and showed no systematic grain dependence for beta diversity between adjacent grid cells. Main conclusions We found that relationships between geographic distance and beta diversity, as well as the environmental correlates of beta diversity, are systematically grain dependent. The strong independent effect of distance indicates that, contrary to the current belief, a substantial fraction of species are missing from areas with a suitable environment. Moreover, the effects of geographic distance (at continental extents) and land cover (at fine grains) indicate that any species distribution modelling should take both environment and dispersal limitation into account.     

Broad‐scale ecological implications of ectothermy and endothermy in changing environments

Aim Physiology is emerging as a basis for understanding the distribution and diversity of organisms, and ultimately for predicting their responses to climate change. Here we review how the difference in physiology of terrestrial vertebrate ectotherms (amphibians and reptiles) and endotherms (birds and mammals) is expected to influence broad‐scale ecological patterns. Location Global terrestrial ecosystems. Methods We use data from the literature and modelling to analyse geographic gradients in energy use and thermal limits. We then compare broad‐scale ecological patterns for both groups with expectations stemming from these geographic gradients. Results The differences in thermal physiology between ectotherms and endotherms result in geographically disparate macrophysiological constraints. Field metabolic rate (FMR) is stable or decreases slightly with temperature for endotherms, while it generally increases for ectotherms, leading to opposing latitudinal gradients of expected FMR. Potential activity time is a greater constraint on the distributions of ectotherms than endotherms, particularly at high latitudes. Differences in the primary correlates of abundance and species richness for two representative taxonomic groups are consistent with the consequences of these basic physiological differences. Ectotherm richness is better predicted by temperature, whereas endotherm richness is more strongly associated with primary productivity. Finally, in contrast to endotherms, ectotherm richness is not strongly related to abundance. Main conclusions Differences in thermal physiology affect how organisms interact with and are constrained by their environment, and may ultimately explain differences in the geographic pattern of biodiversity for endotherms and ectotherms. Linking the fields of physiological and broad‐scale ecology should yield a more mechanistic understanding of how biodiversity will respond to environmental change.     

Recovery of Amphibian and Reptile Assemblages During Old‐Field Succession of Tropical Rain Forests

Conversion of tropical forests to agriculture affects vertebrate assemblages, but we do not know how fast or to what extent these assemblages recover after field abandonment. We addressed this question by examining amphibians and reptiles in secondary forests in southeastern Mexico. We used chronosequence data (12 secondary forests fallow for 1–23 yr and 3 old‐growth forest sites) to analyze successional trajectories and estimate recovery times of assemblage attributes for amphibians and reptiles. We conducted 6 surveys at each site over 14 mo (1200 person‐hours) and recorded 1552 individuals, including 25 species of amphibians and 36 of reptiles, representing 96 and 74 percent of the expected regional number of species, respectively. Abundance, species richness, and species diversity of amphibians increased rapidly with successional age, approaching old‐growth forest values in < 30 yr. Species richness and species diversity of reptiles reached old‐growth forest values in < 20 yr. By contrast, the abundance of reptiles and the assemblage composition of amphibians and reptiles recovered more slowly. Along the chronosequence, we observed more species replacement in reptile assemblages than in amphibian assemblages. Several species in the old‐growth forest were absent from secondary forests. Dispersal limitation and harsh conditions prevailing in open sites and early successional environments appear to preclude colonization by old‐growth forest species. Furthermore, short fallow periods and isolation of forest remnants lead to the formation of new assemblages dominated by species favored by human disturbances.     

Partitioning sources of variation in vertebrate species richness

Aim To explore biogeographic patterns of terrestrial vertebrates in Maine, USA using techniques that would describe local and spatial correlations with the environment. Location Maine, USA. Methods We delineated the ranges within Maine (86,156 km²) of 275 species using literature and expert review. Ranges were combined into species richness maps, and compared to geomorphology, climate, and woody plant distributions. Methods were adapted that compared richness of all vertebrate classes to each environmental correlate, rather than assessing a single explanatory theory. We partitioned variation in species richness into components using tree and multiple linear regression. Methods were used that allowed for useful comparisons between tree and linear regression results. For both methods we partitioned variation into broad‐scale (spatially autocorrelated) and fine‐scale (spatially uncorrelated) explained and unexplained components. By partitioning variance, and using both tree and linear regression in analyses, we explored the degree of variation in species richness for each vertebrate group that could be explained by the relative contribution of each environmental variable. Results In tree regression, climate variation explained richness better (92% of mean deviance explained for all species) than woody plant variation (87%) and geomorphology (86%). Reptiles were highly correlated with environmental variation (93%), followed by mammals, amphibians, and birds (each with 84–82% deviance explained). In multiple linear regression, climate was most closely associated with total vertebrate richness (78%), followed by woody plants (67%) and geomorphology (56%). Again, reptiles were closely correlated with the environment (95%), followed by mammals (73%), amphibians (63%) and birds (57%). Main conclusions Comparing variation explained using tree and multiple linear regression quantified the importance of nonlinear relationships and local interactions between species richness and environmental variation, identifying the importance of linear relationships between reptiles and the environment, and nonlinear relationships between birds and woody plants, for example. Conservation planners should capture climatic variation in broad‐scale designs; temperatures may shift during climate change, but the underlying correlations between the environment and species richness will presumably remain.     

Host diversity and environmental variables as determinants of the species richness of the parasitoids of leaf‐cutting ants

Aim Because of the obligatory relationship between endoparasitoids and their hosts, we presume that hosts exert strong selection pressure on parasitoids. One prediction is a positive relationship between host diversity and parasitoid richness. This relationship could be the product of resource availability which could lead to more opportunities for speciation, or could represent shared responses to the environment by both groups. Location Argentina and Paraguay. Methods We sampled a 1800‐km transect to test for a correlation between the richness of leaf‐cutting ant hosts and their phorid parasitoids. Regression models were used to assess if host and environmental variables could explain phorid species richness at nest, hectare and locality spatial scales. We used canonical correspondence analysis (CCA) to explore if there were similar responses of phorid species to particular host and environmental variables at different spatial scales, and partial CCA to separate the relative importance of both groups of variables. Results Phorid richness was positively correlated with host richness. Host richness/abundance accounted for 20–53% of the variation in parasitoid richness at the hectare and locality scales of analysis, with most of the variation accounted for by ant abundance. We were not able to assess the prediction at the nest scale as only one phorid species was found at most nests. Climatic variables did not explain phorid species richness once host variables were in the models. Partial CCA showed that host‐related variables accounted for most of the variance associated with phorid species ordination at the nest and hectare scales, but not at the largest grain, the locality, where climatic variables were more important. However, most phorid species did not show particular positions along the climatic gradient. Main conclusions The association between parasitoid richness and host richness and abundance, and the overall weak associations with environmental variables, suggest that these host variables are key factors influencing parasitoid speciation.     

Bird species richness is associated with phylogenetic relatedness,plant species richness,and altitudinal range in Inner Mongolia

Bird species richness is mediated by local, regional, and historical factors, for example, competition, environmental heterogeneity, contemporary, and historical climate. Here, we related bird species richness with phylogenetic relatedness of bird assemblages, plant species richness, topography, contemporary climate, and glacial‐interglacial climate change to investigate the relative importance of these factors. This study was conducted in Inner Mongolia, an arid and semiarid region with diverse vegetation types and strong species richness gradients. The following associated variables were included as follows: phylogenetic relatedness of bird assemblages (Net Relatedness Index, NRI), plant species richness, altitudinal range, contemporary climate (mean annual temperature and precipitation, MAT and MAP), and contemporary‐Last Glacial Maximum (LGM) change in climate (change in MAT and change in MAP). Ordinary least squares linear, simultaneous autoregressive linear, and Random Forest models were used to assess the associations between these variables and bird species richness across this region. We found that bird species richness was correlated negatively with NRI and positively with plant species richness and altitudinal range, with no significant correlations with contemporary climate and glacial–interglacial climate change. The six best combinations of variables ranked by Random Forest models consistently included NRI, plant species richness, and contemporary‐LGM change in MAT. Our results suggest important roles of local ecological factors in shaping the distribution of bird species richness across this semiarid region. Our findings highlight the potential importance of these local ecological factors, for example, environmental heterogeneity, habitat filtering, and biotic interactions, in biodiversity maintenance.     

Fine‐scale heterogeneity in beetle assemblages under co‐occurring Eucalyptus in the same subgenus

Aim Insect biodiversity is often positively associated with habitat heterogeneity. However, this relationship depends on spatial scale, with most studies focused on differences between habitats at large scales with a variety of forest tree species. We examined fine‐scale heterogeneity in ground‐dwelling beetle assemblages under co‐occurring trees in the same subgenus: Eucalyptus melliodora A. Cunn. ex Schauer and E. blakelyi Maiden (Myrtaceae). Location Critically endangered grassy woodland near Canberra, south‐eastern Australia. Methods We used pitfall traps and Tullgren funnels to sample ground‐dwelling beetles from the litter environment under 47 trees, and examined differences in diversity and composition at spatial scales ranging from 100 to 1000 m. Results Beetle assemblages under the two tree species had distinctive differences in diversity and composition. We found that E. melliodora supported a higher richness and abundance of beetles, but had higher compositional similarity among samples. In contrast, E. blakelyi had a lower abundance and species richness of beetles, but more variability in species composition among samples. Main conclusions Our study shows that heterogeneity in litter habitat under co‐occurring and closely related eucalypt species can influence beetle assemblages at spatial scales of just hundreds of metres. The differential contribution to fine‐scale alpha and beta diversity by each eucalypt can be exploited for conservation purposes by ensuring an appropriate mix of the two species in the temperate woodlands where they co‐occur. This would help not only to maximize biodiversity at landscape scales, but also to maintain heterogeneity in species richness, trophic function and biomass at fine spatial scales.     

Dietary guild composition and disaggregation of avian assemblages under climate change

Climate change is expected to cause geographic redistributions of species. To the extent that species within assemblages have different niche requirements, assemblages may no longer remain intact and dis‐ and reassemble at current or new geographic locations. We explored how climate change projected by 2100 may transform the world's avian assemblages (characterized at a 110 km spatial grain) by modeling environmental niche‐based changes to their dietary guild structure under 0, 500, and 2000 km‐dispersal distances. We examined guild structure changes at coarse (primary, high‐level, and mixed consumers) and fine (frugivores, nectarivores, insectivores, herbivores, granivores, scavengers, omnivores, and carnivores) ecological resolutions to determine whether or not geographic co‐occurrence patterns among guilds were associated with the magnitude to which guilds are functionally resolved. Dietary guilds vary considerably in their global geographic prevalence, and under broad‐scale niche‐based redistribution of species, these are projected to change very heterogeneously. A nondispersal assumption results in the smallest projected changes to guild assemblages, but with significant losses for some regions and guilds, such as South American insectivores. Longer dispersal distances are projected to cause greater degrees of disassembly, and lead to greater homogenization of guild composition, especially in northern Asia and Africa. This arises because projected range gains and losses result in geographically heterogeneous patterns of guild compensation. Projected decreases especially of primary and mixed consumers most often are compensated by increases in high‐level consumers, with increasing uncertainty about these outcomes as dispersal distance and degree of guild functional resolution increase. Further exploration into the consequences of these significant broad‐scale ecological functional changes at the community or ecosystem level should be increasingly on the agenda for conservation science.     

Context‐dependent functional dispersion across similar ranges of trait space covered by intertidal rocky shore communities

Functional diversity is intimately linked with community assembly processes, but its large‐scale patterns of variation are often not well understood. Here, we investigated the spatiotemporal changes in multiple trait dimensions (“trait space”) along vertical intertidal environmental stress gradients and across a landscape scale. We predicted that the range of the trait space covered by local assemblages (i.e., functional richness) and the dispersion in trait abundances (i.e., functional dispersion) should increase from high‐ to low‐intertidal elevations, due to the decreasing influence of environmental filtering. The abundance of macrobenthic algae and invertebrates was estimated at four rocky shores spanning ca. 200 km of the coast over a 36‐month period. Functional richness and dispersion were contrasted against matrix‐swap models to remove any confounding effect of species richness on functional diversity. Random‐slope models showed that functional richness and dispersion significantly increased from high‐ to low‐intertidal heights, demonstrating that under harsh environmental conditions, the assemblages comprised similar abundances of functionally similar species (i.e., trait convergence), while that under milder conditions, the assemblages encompassed differing abundances of functionally dissimilar species (i.e., trait divergence). According to the Akaike information criteria, the relationship between local environmental stress and functional richness was persistent across sites and sampling times, while functional dispersion varied significantly. Environmental filtering therefore has persistent effects on the range of trait space covered by these assemblages, but context‐dependent effects on the abundances of trait combinations within such range. Our results further suggest that natural and/or anthropogenic factors might have significant effects on the relative abundance of functional traits, despite that no trait addition or extinction is detected.     

Regional and environmental effects on the species richness of mammal assemblages

Aim Variation in species richness has been related to (1) environmental conditions (water, energy and habitat characteristics) and (2) regional differences (contingent historical events and regional particularities that result in differences between regional faunas acting at broad extents). Whereas climatic factors have been widely studied, the effects of regional differences are less often quantified. This work aims to characterize global trends in the species richness of mammal assemblages with respect to both current and historical influences. Location All terrestrial biogeographical realms except Antarctica. Methods Species richness in checklists from 224 sites distributed worldwide were investigated by partitioning the variation between a general set of habitat/climate factors, biogeographical regions, and their overlaps. Additional analyses studied the specific overlaps of region, water and energy. Data were also divided according to area to determine if the strength of these effects varies according to the size of sites. Results Environmental effects explained 38% of richness variation across all sites, whereas environmentally independent regional effects explained 11% and the overlap between region and environment explained 13%. Results were similar when only larger sites (between 1000 km² and 10,000 km²) were considered. However, the importance of the overlap between region and all environmental variables was greater in smaller sites (between 100 km² and 1000 km²). In contrast, the specific importance of water and energy variables and their overlap with region was greater in larger sites. The strength of the independent effect of region remained almost invariant regardless of the size of the sites studied. Main conclusions The relationship between species richness and climate varies with scale and among regions. Although environmental variables are the strongest correlates of richness, the unique history and physiographic characteristics of a region produce differences between the richness of mammal assemblages and their response to environmental gradients. The importance of environmental variables varies with scale: climatic gradients are more important at coarse grain (larger sites), possibly as a result of their effects on species ranges, whereas habitat type is more important at the smaller sites, where the importance of ecological interactions increases. Therefore, regional differences and the scale at which richness is measured should be taken into account when evaluating species richness–energy hypotheses.     

Co‐declining mammal–dung beetle faunas throughout the Atlantic Forest biome of South America

The millennial–scale evolutionary relationships between mammals and dung beetles have been eroded due to several drivers of contemporary biodiversity loss. Although some evidence of co‐decline has been shown for mammals and dung beetles at some Neotropical sites, a biome‐scale analysis for the entire Atlantic Forest of South America would strengthen our understanding of how relictual sets of mammal species can affect dung beetle co‐occurrences and co‐declines. We therefore collated hundreds of assemblages of both dung beetles and medium‐ to large‐bodied mammals throughout the world's longest tropical forest latitudinal gradient to examine to what extent mammal assemblages may exert a positive influence on dung beetle species composition and functional assembly, and whether this relationship is scale dependent. We also collated several climatic and other environmental variables to examine the degree to which they shape mammal–dung beetle relationships. The relationships between local mammal and dung beetle faunas were examined using regression models, variation partitioning, dissimilarity indices and ecological networks. We found a clear positive relationship between mammal and dung beetle species richness across this forest biome, indicating an ongoing process of mammal–dung beetle niche‐mediated co‐decline. We found a strong relationship between the species composition of both taxa, in which dung beetle species dissimilarity apparently track changes in mammalian dissimilarity, typically in 80% of all cases. Co‐variables such as phytomass and climatic variables also influenced mammal–dung beetle patterns of co‐decline along the Atlantic Forest. We conclude that dung beetle diversity and community assembly are shaped by the remaining co‐occurring mammal assemblages and their functional traits, and both groups were governed by environmental features. We emphasize that ecosystem‐wide effects of mammal population declines remain poorly understood both quantitatively and qualitatively, and curbing large vertebrate defaunation will ensure the persistence of co‐dependent species.     

Biome transitions as centres of diversity: habitat heterogeneity and diversity patterns of West African bat assemblages across spatial scales

It is widely accepted that species diversity is contingent upon the spatial scale used to analyze patterns and processes. Recent studies using coarse sampling grains over large extents have contributed much to our understanding of factors driving global diversity patterns. This advance is largely unmatched on the level of local to landscape scales despite being critical for our understanding of functional relationships across spatial scales. In our study on West African bat assemblages we employed a spatially explicit and nested design covering local to regional scales. Specifically, we analyzed diversity patterns in two contrasting, largely undisturbed landscapes, comprising a rainforest area and a forest‐savanna mosaic in Ivory Coast, West Africa. We employed additive partitioning, rarefaction, and species richness estimation to show that bat diversity increased significantly with habitat heterogeneity on the landscape scale through the effects of beta diversity. Within the extent of our study areas, habitat type rather than geographic distance explained assemblage composition across spatial scales. Null models showed structure of functional groups to be partly filtered on local scales through the effects of vegetation density while on the landscape scale both assemblages represented random draws from regional species pools. We present a mixture model that combines the effects of habitat heterogeneity and complexity on species richness along a biome transect, predicting a unimodal rather than a monotonic relationship with environmental variables related to water. The bat assemblages of our study by far exceed previous figures of species richness in Africa, and refute the notion of low species richness of Afrotropical bat assemblages, which appears to be based largely on sampling biases. Biome transitions should receive increased attention in conservation strategies aiming at the maintenance of ecological and evolutionary processes.     

Scale‐dependent processes of community assemblyin an African rift lake

1. Ecologists continue to debate whether the assembly of communities of species is more strongly influenced by dispersal limitations or niche‐based factors. Analytical approaches that account for both mechanisms can help to resolve controls of community assembly. 2. We compared littoral snail assemblages in Lake Tanganyika at three different spatial scales (5–25 m, 0.5–10 km and 0.5–27 km) to test whether spatial distance or environmental differences are better predictors of community similarity. 3. At the finest scale (5–25 m), snail assemblages shifted strongly with depth but not across similar lateral distances, indicating a stronger response to environmental gradients than dispersal opportunities. 4. At the two larger scales (0.5–27 km), both environmental similarity and shoreline distance between sites predicted assemblage similarity across sites. Additionally, canonical correspondence analysis revealed that snail abundances were significantly correlated with algal carbon‐to‐nitrogen ratio and wave energy. 5. Our results indicate that the factors governing assemblage structure are scale dependent; niche‐based mechanisms act across all spatial scales, whereas community similarity declines with distance only at larger spatial separations.     

Grids versus regional species lists: are broad-scale patterns of species richness robust to the violation of constant grain size?

Where distribution maps do not exist ecologists often use regional species lists to examine geographic patterns of species richness, despite the fact that inconsistent grain sizes across areas may complicate interpretation of the results. We compare patterns of species richness of European butterflies and dragonflies using regional species lists (varying grain size) and regular grids (constant grain size). We asked if species lists give results comparable to the gridded data when used in simple macroecological analysis of environmental correlates of species richness. We generated two equal-area grids (220 × 220 km and 440 × 440 km) to map the richness gradients and model species richness as a function of actual evapotranspiration (AET) and range in elevation. Then we used species checklists of 33 administrative regions of unequal sizes to construct the same environmental models while accounting for differences in area. Analysis of butterfly checklist data produced comparable results to the analysis of gridded data. In contrast, dragonfly checklist data had a distorted spatial pattern and much weaker associations with environmental variables than the gridded data. The robustness of checklist data appears to be variable, even within a single geographical region, and may not generate patterns congruent with those found using equal-area grids.