The conservation status of African vertebrates is unrelated to environmental and spatial patterns in their geographic ranges

Statistical predictions of the impact of climate change on biodiversity assume that the environmental and spatial characteristics of contemporary species’ distributions reflect the conditions needed for their continued and prolonged existence. Here we explore this assumption by testing whether a species’ threatened status is associated with the amount of variation in its distribution range attributable to environmental and spatial patterns. Using a variation partitioning approach, we decomposed variation in the distribution ranges of 4423 vertebrate species in sub-Saharan Africa into components attributable exclusively to environmental variables (E|S), exclusively to spatial variables (S|E) or to the collinearity between environmental and spatial variables (E∩S). We found that species’ threatened status was unrelated to E|S, S|E or E∩S variation components, but that unexplained variation was higher for species threatened with extinction. This suggests that spatio-environmental patterns in species’ ranges likely underestimate the overall extinction threat caused by climate change. We also found clear geographic patterns in the strength of E|S, S|E or E∩S that differed amongst biogeographical regions, but no component was over- or underrepresented in the present-day protected area network. While there may be benefits to tailoring protected area expansion to differences between biogeographical regions, this should aim to incorporate species-specific information wherever possible.     

Environmental variability and biogeography: the relationship between bathymetric distribution and geographical range size in marine algae and gastropods

Aims Gradients of environmental variability have been proposed to explain spatial variation in patterns of geographical range size. We explore this relationship in NE Pacific algae and NW Atlantic gastropods by using the characteristics of species’ bathymetric distributions as a proxy for tolerance of environmental variability. Location NE Pacific and NW Atlantic. Methods Data on species bathymetric and geographical distributions were compiled from the literature. Results For both algae and gastropods, species that inhabit highly seasonal, shallow depth zones have broader latitudinal ranges, and occupy more biogeographical provinces, than species that live in more temporally stable, deeper zones. Furthermore, species that tolerate spatial variability along the bathymetric axis, i.e. those that occur in multiple depth zones, have broader geographical ranges than species restricted to fewer depth zones. Main conclusions Within‐range environmental variability, through both space and time, is predictive of large geographical ranges for marine algae and gastropods. Analysis of species distributions across perpendicular gradients (e.g. depth and latitude) is a powerful approach to discerning the mechanisms that govern biogeographical patterns, and provides easily obtainable broad‐brush predictions regarding the biogeographical outcomes of global change.     

Spatial autocorrelation analysis allows disentangling the balance between neutral and niche processes in metacommunities

One of the most popular approaches for investigating the roles of niche and neutral processes driving metacommunity patterns consists of partitioning variation in species data into environmental and spatial components. The logic is that the distance decay of similarity in communities is expected under neutral models. However, because environmental variation is often spatially structured, the decay could also be attributed to environmental factors that are missing from the analysis. Here, we use a spatial autocorrelation analysis protocol, previously developed to detect isolation‐by‐distance in allele frequencies, to evaluate patterns of species abundances under neutral dynamics. We show that this protocol can be linked with variation partitioning analyses. Moreover, in an attempt to test the neutral model, we derive three predictions to be applied both to original species abundances and to abundances predicted by a pure spatial model species abundances will be uncorrelated; Moran's I correlograms will reveal similar short‐distance autocorrelation patterns; an increasing degree of non‐neutrality will tend to generate patterns of correlation among abundances within groups of species with similar correlograms (i.e. within species with neutral and non‐neutral dynamics). We illustrate our protocol by analyzing spatial patterns in abundance of 28 terrestrially breeding anuran species from Central Amazonia. We recommend that researchers should investigate spatial autocorrelation patterns of abundances predicted by pure spatial models to identify similar patterns of spatial autocorrelation at short distances and lack of correlation between species abundances. Therefore, the hypothesis that spatial patterns in abundances are primarily due to pure neutral dynamics (rather than to missing spatiallystructured environmental factors) can be confirmed after taking environmental variables into account.     

Range size patterns in European freshwater trematodes

While patterns in geographic range sizes in free‐living species have received much attention, little is known on the corresponding patterns in parasites. For the first time, we report on patterns in geographic range sizes and dimensions of endoparasites, using published species lists of freshwater trematodes in 25 biogeographical regions of Europe. In general, the range sizes of trematodes showed a typical hollow curve frequency distribution, with most species having small ranges. Contrary to expectations, there were no differences in range sizes among trematodes using hosts with high (birds) and limited dispersal capacity (e.g. fish). This suggests that the well known importance of host dispersal capacity for parasite dispersal at small spatial scales is overridden by other factors on larger scales. Regression analyses and Rohde plots showed that the relationship between the latitudinal centre and trematode range size was hump‐shaped in all host groups except for reptiles, for which it was linear. Most of the variation fell within the expectations given by null models, suggesting that the patterns mainly result from the geographic properties of the European continent and the biogeographical regions. Finally, trematode ranges tended to stretch more in east‐west than in north‐south directions, indicating dispersal barrier effects for parasite faunas, probably resulting from the geographical idiosyncrasies of the European continent.     

Biogeographical patterns of rocky intertidal communities along the Pacific coast of North America

Aim Our aim in this paper is to present the first broad‐scale quantification of species abundance for rocky intertidal communities along the Pacific coast of North America. Here we examine the community‐level marine biogeographical patterns in the context of formerly described biogeographical regions, and we evaluate the combined effects of geographical distance and environmental conditions on patterns of species similarity across this region. Location Pacific coast of North America. Methods Data on the percentage cover of benthic marine organisms were collected at 67 rocky intertidal sites from south‐eastern Alaska, USA, to central Baja California Sur, Mexico. Cluster analysis and non‐metric multidimensional scaling were used to evaluate the spatial patterns of species similarity among sites relative to those of previously defined biogeographical regions. Matrices of similarity in species composition among all sites were computed and analysed with respect to geographical distance and long‐term mean sea surface temperature (SST) as a measure of environmental conditions. Results We found a high degree of spatial structure in the similarity of intertidal communities along the coast. Cluster analysis identified 13 major community structure ‘groups’. Although breaks between clusters of sites generally occurred at major biogeographical boundaries, some of the larger biogeographical regions contained several clusters of sites that did not group according to spatial position or identifiable coastal features. Additionally, there were several outliers – sites that grouped alone or with sites outside their region – for which localized features may play an important role in driving community structure. Patterns of species similarity at the large scale were highly correlated with geographical distance among sites and with SST. Importantly, we found community similarity to be highly correlated with long‐term mean SST while controlling for the effects of geographical distance. Main conclusions These findings reveal a high degree of spatial structure in the similarity of rocky intertidal communities of the north‐east Pacific, and are generally consistent with those of previously described biogeographical regions, with some notable differences. Breaks in similarity among clusters are generally coincident with known biogeographical and oceanographic discontinuities. The strong correlations between species similarity and both geographical position and SST suggest that both geography and oceanographic conditions have a large influence on patterns of intertidal community structure along the Pacific coast of North America.     

Understanding global patterns in amphibian geographic range size: does Rapoport rule?

Aim Species geographic ranges are the ‘fundamental units’ of macroecology. Range size is a major correlate of extinction risk in many groups, and is also critical in studies of biotic responses to climate change. Despite this, there is a lack of studies exploring the role of environmental, historical and anthropogenic processes in determining large‐scale patterns in range size. We perform the first global analysis of putative drivers of range size variation in any group, choosing amphibians as our study taxon. Our aims are to disentangle the many hypothesized causes of range size variation and evaluate support for ‘Rapoport's rule’, the observation that range size correlates with latitude. Location Global. Methods We develop a global map of gridded median range size using the International Union for Conservation of Nature (IUCN) distribution maps. From this we perform spatial and non‐spatial regressions to explore relationships between range size and nine hypothesized variables in six biogeographic realms. We use information‐theoretic model selection to compare multiple competing variables, simultaneously evaluating the relative support for each one. Results Current climate – environmental water and energy, and temperature seasonality – is consistently highly ranked in spatial and non‐spatial analyses. Human impacts and other environmental measures (topographic and landscape complexity, effective area, climate extremes) show mixed support, and glacial history is consistently unimportant. Our findings add further evidence to the view that Rapoport's rule is a regional, not global, phenomenon. Main conclusions The primary importance of temperature seasonality may explain why Rapoport's rule is largely restricted to northern latitudes, as this is where seasonality is most pronounced. More generally, the dominance of contemporary climate in our analyses (even when accounting for space) has stark implications for the future status of amphibians. Changes in climate will almost certainly interact with the anthropogenic processes already threatening a third of amphibians globally, with the effects being most keenly felt by species with a restricted range.     

Geographic variation in genetic and demographic performance: new insights from an old biogeographical paradigm

The ‘centre–periphery hypothesis’ (CPH) is a long‐standing postulate in ecology that states that genetic variation and demographic performance of a species decrease from the centre to the edge of its geographic range. This hypothesis is based on an assumed concordance between geographical peripherality and ecological marginality such that environmental conditions become harsher towards the limits of a species range. In this way, the CPH sets the stage for understanding the causes of distribution limits. To date, no study has examined conjointly the consistency of these postulates. In an extensive literature review we discuss the birth and development of the CPH and provide an assessment of the CPH by reviewing 248 empirical studies in the context of three main themes. First, a decrease in species occurrence towards their range limits was observed in 81% of studies, while only 51% demonstrated reduced abundance of individuals. A decline in genetic variation, increased differentiation among populations and higher rates of inbreeding were demonstrated by roughly one in two studies (47, 45 and 48%, respectively). However, demographic rates, size and population performance less often followed CPH expectations (20–30% of studies). We highlight the impact of important methodological, taxonomic, and biogeographical biases on such validation rates. Second, we found that geographic and ecological marginality gradients are not systematically concordant, which casts doubt on the reliability of a main assumption of the CPH. Finally, we attempt to disentangle the relative contribution of geographical, ecological and historical processes on the spatial distribution of genetic and demographic parameters. While ecological marginality gradients explain variation in species' demographic performance better than geographic gradients, contemporary and historical factors may contribute interactively to spatial patterns of genetic variation. We thereby propose a framework that integrates species' ecological niche characteristics together with current and past range structure to investigate spatial patterns of genetic and demographic variation across species ranges.     

Can environmental constraints determine random patterns of plant species co-occurrence?

Plant community ecologists use the null model approach to infer assembly processes from observed patterns of species co‐occurrence. In about a third of published studies, the null hypothesis of random assembly cannot be rejected. When this occurs, plant ecologists interpret that the observed random pattern is not environmentally constrained – but probably generated by stochastic processes. The null model approach (using the C‐score and the discrepancy index) was used to test for random assembly under two simulation algorithms. Logistic regression, distance‐based redundancy analysis, and constrained ordination were used to test for environmental determinism (species segregation along environmental gradients or turnover and species aggregation). This article introduces an environmentally determined community of alpine hydrophytes that presents itself as randomly assembled. The pathway through which the random pattern arises in this community is suggested to be as follows: Two simultaneous environmental processes, one leading to species aggregation and the other leading to species segregation, concurrently generate the observed pattern, which results to be neither aggregated nor segregated – but random. A simulation study supports this suggestion. Although apparently simple, the null model approach seems to assume that a single ecological factor prevails or that if several factors decisively influence the community, then they all exert their influence in the same direction, generating either aggregation or segregation. As these assumptions are unlikely to hold in most cases and assembly processes cannot be inferred from random patterns, we would like to propose plant ecologists to investigate specifically the ecological processes responsible for observed random patterns, instead of trying to infer processes from patterns.     

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

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

The role of environmental and spatial processes in structuring native and non‐native fish communities across thousands of lakes

Quantifying the role of spatial patterns is an important goal in ecology to further understand patterns of community composition. We quantified the relative role of environmental conditions and regional spatial patterns that could be produced by environmental filtering and dispersal limitation on fish community composition for thousands of lakes. A database was assembled on fish community composition, lake morphology, water quality, climatic conditions, and hydrological connectivity for 9885 lakes in Ontario, Canada. We utilized a variation partitioning approach in conjunction with Moran's Eigenvector Maps (MEM) and Asymmetric Eigenvector Maps (AEM) to model spatial patterns that could be produced by human‐mediated and natural modes of dispersal. Across 9885 lakes and 100 fish species, environmental factors and spatial structure explained approximately 19% of the variation in fish community composition. Examining the proportional role of spatial structure and environmental conditions revealed that as much as 90% of the explained variation in native species assemblage composition is governed by environmental conditions. Conversely on average, 67% of the explained variation in non‐native assemblage composition can be related to human‐mediated dispersal. This study highlights the importance of including spatial structure and environmental conditions when explaining patterns of community composition to better discriminate between the ecological processes that underlie biogeographical patterns of communities composed of native and non‐native fish species.     

Community assembly is a race between immigration and adaptation: eco‐evolutionary interactions across spatial scales

Both ecological and evolutionary mechanisms have been proposed to describe how natural communities become assembled at both regional and biogeographical scales. Yet, these theories have largely been developed in isolation. Here, we unite these separate views and develop an integrated eco‐evolutionary framework of community assembly. We use a simulation approach to explore the factors determining the interplay between ecological and evolutionary mechanisms systematically across spatial scales. Our results suggest that the same set of ecological and evolutionary processes can determine community assembly at both regional and biogeographical scales. We find that the importance of evolution and community monopolization effects, defined as the eco‐evolutionary dynamics that occur when local adaptation of early established immigrants is fast enough to prevent the later immigration of better pre‐adapted species, are not restricted to adaptive radiations on remote islands. They occur at dispersal rates of up to ten individuals per generation, typical for many species at the scale of regional metacommunities. Dispersal capacity largely determines whether ecological species sorting or evolutionary monopolization structure metacommunity diversity and distribution patterns. However, other factors related to the spatial scale at which community assembly processes are acting, such as metacommunity size and the proportion of empty patches, also affect the relative importance of ecology versus evolution. We show that evolution often determines community assembly, and this conclusion is robust to a wide range of assumptions about spatial scale, mode of reproduction, and environmental structure. Moreover, we found that community monopolization effects occur even though species fully pre‐adapted to each habitat are abundant in the metacommunity, a scenario expected a priori to prevent any meaningful effect of evolution. Our results strongly support the idea that the same eco‐evolutionary processes underlie community assembly at regional and biogeographical scales.     

Niche versus neutrality in structuring the beta diversity of damselfly assemblages

1. Differences among communities in taxonomic composition – beta diversity – are frequently expected to result from taxon‐specific responses to spatial variation in ecological conditions, through niche partitioning. Such process‐derived patterns are in sharp contrast to arguments from neutral theory, where taxa are ecologically equivalent and beta diversity results primarily from dispersal limitation. 2. Here, we compared beta diversity among assemblages of damselflies (Odonata: Zygoptera), for which previous experiments have shown that niche differences maintain genera within a community, but patterns of relative abundance for species within each genus are shaped primarily by neutral dynamics. 3. Using null‐model and ordination‐based methods, we find that both genera and (in contrast to neutral theory) species assemblage composition vary across the landscape in a deterministic fashion, shaped by environmental and spatial factors. 4. While the observed patterns in species composition conflict with theory, we suggest that this a result of weak ecological filters acting to produce spatial variation in assemblages of ecologically similar species undergoing ecological drift within communities. Such patterns are especially likely in systems of relatively weak dispersers like damselflies.     

Disentangling the effects of geographic distance and environmental dissimilarity on global patterns of species turnover

Aim To distinguish the effects of geographic distance and environmental dissimilarity on global patterns of species turnover in four classes of terrestrial vertebrates (mammals, birds, reptiles and amphibians). Location Six hundred and sixty terrestrial ecoregions across the globe. Methods We calculated species turnover between each pair of ecoregions, using the Jaccard index (J). We selected seven variables to quantify environment in each ecoregion, and subjected the environmental values to a principal components analysis. For each realm, we applied multiple regression analysis relating the natural logarithm of the Jaccard index (lnJ) to geographic distance alone and in combination with differences in the environment variables measured as principal components (PC). We used partial correlations to partition variance in lnJ between unique contributions of distance and environmental PC scores, the covariation between distance and environment, and unexplained variance. To examine the latitude and species turnover relationship, we regressed lnJ on latitude with distance between ecoregions being included as a covariate. Results The natural logarithm of the Jaccard index (lnJ) decreased significantly with increasing geographic distance for all vertebrate classes in each zoogeographic realm, and the slopes of the relationships per 1000 km ranged from ?0.251 to ?1.043. With environmental differences included in the analysis, both geographic distance and environmental differences were substantial predictors of lnJ for every combination of taxon and realm. On average, the unique contribution of geographic distance to variation in species turnover between ecoregions was about 1.4 times that of the environmental differences between ecoregions. Species turnover generally decreased with increasing latitude when controlling for geographic distance. The value of lnJ for each vertebrate class was highly and positively correlated with those of the other vertebrate classes. Main conclusions Our analyses suggest that both dispersal‐based and niche‐based processes have played important roles in determining faunal similarities among vertebrate assemblages at the spatial scale examined. Furthermore, reptiles and amphibians exhibited greater distance‐independent faunal heterogeneity among ecoregions and greater turnover among ecoregions with respect to geographic and environmental distance than birds and mammals.     

Host plant associations and geography interact to shape diversification in a specialist insect herbivore

Disentangling the processes underlying geographic and environmental patterns of biodiversity challenges biologists as such patterns emerge from eco‐evolutionary processes confounded by spatial autocorrelation among sample units. The herbivorous insect, Belonocnema treatae (Hymenoptera: Cynipidae), exhibits regional specialization on three plant species whose geographic distributions range from sympatry through allopatry across the southern United States. Using range‐wide sampling spanning the geographic ranges of the three host plants and genotyping‐by‐sequencing of 1,217 individuals, we tested whether this insect herbivore exhibited host plant‐associated genomic differentiation while controlling for spatial autocorrelation among the 58 sample sites. Population genomic structure based on 40,699 SNPs was evaluated using the hierarchical Bayesian model entropy to assign individuals to genetic clusters and estimate admixture proportions. To control for spatial autocorrelation, distance‐based Moran's eigenvector mapping was used to construct regression variables summarizing spatial structure inherent among sample sites. Distance‐based redundancy analysis (dbRDA) incorporating the spatial variables was then applied to partition host plant‐associated differentiation (HAD) from spatial autocorrelation. By combining entropy and dbRDA to analyse SNP data, we unveiled a complex mosaic of highly structured differentiation within and among gall‐former populations finding evidence that geography, HAD and spatial autocorrelation all play significant roles in explaining patterns of genomic differentiation in B. treatae. While dbRDA confirmed host association as a significant predictor of patterns of genomic variation, spatial autocorrelation among sites explained the largest proportion of variation. Our results demonstrate the value of combining dbRDA with hierarchical structural analyses to partition spatial/environmental patterns of genomic variation.     

Complementary strengths of spatially-explicit and multi-species distribution models

Species distribution models (SDMs) project the outcome of community assembly processes – dispersal, the abiotic environment and biotic interactions – onto geographic space. Recent advances in SDMs account for these processes by simultaneously modeling the species that comprise a community in a multivariate statistical framework or by incorporating residual spatial autocorrelation in SDMs. However, the effects of combining both multivariate and spatially-explicit model structures on the ecological inferences and the predictive abilities of a model are largely unknown. We used data on eastern hemlock Tsuga canadensis and five additional co-occurring overstory tree species in 35 569 forest stands across Michigan, USA to evaluate how the choice of model structure, including spatial and non-spatial forms of univariate and multivariate models, affects ecological inference about the processes that shape community composition as well as model predictive ability. Incorporating residual spatial autocorrelation via spatial random effects did not improve out-of-sample prediction for the six tree species, although in-sample model fit was higher in the spatial models. Spatial models attributed less variation in occurrence probability to environmental covariates than the non-spatial models for all six tree species, and estimated higher (more positive) residual co-occurrence values for most species pairs. The non-spatial multivariate model was better suited for evaluating habitat suitability and hypotheses about the processes that shape community composition. Environmental correlations and residual correlations among species pairs were positively related, perhaps indicating that residual correlations were due to shared responses to unmeasured environmental covariates. This work highlights the importance of choosing a non-spatial model formulation to address research questions about the species–environment relationship or residual co-occurrence patterns, and a spatial model formulation when within-sample prediction accuracy is the main goal.     

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

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

Adding energy gradients and long‐distance dispersal to a neutral model improves predictions of Madagascan bird diversity

Macroecological patterns are likely the result of both stochastically neutral mechanisms and deterministic differences between species. In Madagascar, the simplest stochastically neutral hypothesis – the mid‐domain effects (MDE) hypothesis – has already been rejected. However, rejecting the MDE hypothesis does not necessarily refute the existence of all other neutral mechanisms. Here, we test whether adding complexity to a basic neutral model improves predictions of biodiversity patterns. The simplest MDE model assumes that: (1) species' ranges are continuous and unfragmented, (2) are randomly located throughout the landscape, and (3) can be stacked independently and indefinitely. We designed a simulation based on neutral theory that allowed us to weaken each of these assumptions incrementally by adjusting the habitat capacity as well as the likelihood of short‐ and long‐distance dispersal. Simulated outputs were compared to four empirical patterns of bird diversity: the frequency distributions of species richness and range size, the within‐island latitudinal diversity gradient, and the distance‐decay of species compositional similarity. Neutral models emulated empirical diversity patterns for Madagascan birds accurately. The frequency distribution of range size, latitudinal diversity gradient, and the distance‐decay of species compositional similarity could be attributed to stochastic long‐distance migration events and zero‐sum population dynamics. However, heterogenous environmental gradients improved predictions of the frequency distribution of species richness. Patterns of bird diversity in Madagascar can broadly be attributed to stochastic long‐distance migration events and zero‐sum population dynamics. This implies that rejecting simple hypotheses, such as MDE, does not serve as evidence against stochastic processes in general. However, environmental gradients were necessary to explain patterns of species richness and deterministic differences between species are probably important for explaining the distributions of narrow‐range and endemic species.     

Beta‐diversity in temperate and tropical forests reflects dissimilar mechanisms of community assembly

Site‐to‐site variation in species composition (β‐diversity) generally increases from low‐ to high‐diversity regions. Although biogeographical differences in community assembly mechanisms may explain this pattern, random sampling effects can create this pattern through differences in regional species pools. Here, we compared assembly mechanisms between spatially extensive networks of temperate and tropical forest plots with highly divergent species pools (46 vs. 607 species). After controlling for sampling effects, β‐diversity of woody plants was similar and higher than expected by chance in both forests, reflecting strong intraspecific aggregation. However, different mechanisms appeared to explain aggregation in the two forests. In the temperate forest, aggregation reflected stronger environmental correlations, suggesting an important role for species‐sorting (e.g. environmental filtering) processes, whereas in the tropics, aggregation reflected stronger spatial correlations, more likely reflecting dispersal limitation. We suggest that biogeographical differences in the relative importance of different community assembly mechanisms contribute to these striking gradients in global biodiversity.     

Non-random patterns in the Yellowstone ecosystem: inferences from mammalian body size, order and biogeographical affinity

Aim Our aim was to investigate how the environment, species characteristics and historical factors at the subcontinental scale affect patterns of diversity. We used the assembly of the Yellowstone biota over the past 10,000 years as a natural experiment for investigating the processes that generate a modern non‐volant mammal species pool. Location The data represent species from throughout North America with special attention to the non‐volant mammals of Yellowstone National Park, USA. Methods We used digitized range maps to determine biogeographical affinity for all non‐volant mammals in the Rocky Mountains, Deserts and Great Plains biogeographical regions of North America. This biogeographical affinity, along with taxonomic order and body size class, was used to test whether non‐random patterns exist in the assemblage of Yellowstone non‐volant mammals. These characteristics were also used to investigate the strength of non‐random processes, such as habitat or taxon filtering, on particular groups of species or individual species. Results Our results indicated that the Yellowstone fauna is composed of a non‐random subset of mammals from specific body size classes and with particular biogeographical affinities. Analyses by taxonomic order found significantly more Carnivora from the Rocky Mountains region and significantly fewer Rodentia from the Deserts region than expected from random assembly. Analyses using body size classes revealed deviations from expectations, including several significant differences between the frequency distribution of regional body sizes and the distribution of those species found within Yellowstone. Main conclusions Our novel approach explores processes affecting species pool assembly in the Yellowstone region and elsewhere, and particularly identifies unique properties of species that may contribute to non‐random assembly. Focusing on the mechanisms generating diversity, not just current diversity patterns, will assist the design of conservation strategies given future environmental change scenarios.     

A Dated Phylogeny Complements Macroecological Analysis to Explain the Diversity Patterns in Geonoma (Arecaceae)

Integrating phylogenetic data into macroecological studies of biodiversity patterns may complement the information provided by present‐day spatial patterns. In the present study, we used range map data for all Geonoma (Arecaceae) species to assess whether Geonoma species composition forms spatially coherent floristic clusters. We then evaluated the extent to which the spatial variation in species composition reflects present‐day environmental variation vs. nonenvironmental spatial effects, as expected if the pattern reflects historical biogeography. We also examined the degree of geographic structure in the Geonoma phylogeny. Finally, we used a dated phylogeny to assess whether species richness within the floristic clusters was constrained by a specific historical biogeographic driver, namely time‐for‐diversification. A cluster analysis identified six spatially coherent floristic clusters, four of which were used to reveal a significant geographic phylogenetic structure. Variation partitioning analysis showed that 56 percent of the variation in species composition could be explained by spatial variables alone, consistent with historical factors having played a major role in generating the Geonoma diversity pattern. To test for a time‐for‐diversification effect, we correlated four different species richness measures with the diversification time of the earliest large lineage that is characteristic of each cluster. In support of this hypothesis, we found that geographic areas with higher richness contained older radiations. We conclude that current geographic diversity patterns in Geonoma reflect the present‐day climate, but to a larger extent are related to nonenvironmental spatial constraints linked to colonization time, dispersal limitation, and geological history, followed by within‐area evolutionary diversification. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .