Contrasting environmental and regional effects on global pteridophyte and seed plant diversity

Pteridophytes (ferns and fern‐allies) represent the second‐largest group of vascular plants, but their global biogeography remains poorly studied. Given their functional biology, pteridophytes are expected to show a more pronounced relation to water availability and a higher dispersal ability compared to seed plants. We test these assertions and document the global pattern of pteridophyte richness across 195 mainland and 106 island regions. Using non‐spatial and spatial simple and multiple regression models, we analyze geographic trends in pteridophyte and seed plant richness as well as pteridophyte proportions in relation to environmental and regional variables. We find that pteridophyte and seed plant richness are geographically strongly correlated (all floras: r=0.68, mainland: r=0.82, island floras: r=0.77), but that the proportions of pteridophytes in vascular plant floras vary considerably (0–70%). Islands (mean=15.3%) have significantly higher proportions of pteridophytes than mainland regions (mean=3.6%). While the relative proportions of pteridophytes on islands show a positive relationship with geographic isolation, proportions in mainland floras increase most strongly along gradients of water availability. Pteridophyte richness peaks in humid tropical mountainous regions and is lowest in deserts, arctic regions, and on remote oceanic islands. Regions with Mediterranean climate, outstanding extra‐tropical centres of seed plant richness, are comparatively poor in pteridophytes. Overall, water‐energy variables and topographical complexity are core predictors of both mainland pteridophyte and seed plant richness. Significant residual richness across biogeographic regions points to an important role of idiosyncratic regional effects. Although the same variables emerge as core predictors of pteridophyte and seed plant richness, water availability is clearly a much stronger constraint of pteridophyte richness. We discuss the different limitations of gametophytes and sporophytes that might have limited the ability of pteridophytes to extensively diversify under harsh environmental conditions. Our results point to an important role of taxon‐specific functional traits in defining global richness gradients.     

Disentangling the relative effects of ambient energy,water availability,and energy–water balance on pteridophyte species richness at a landscape scale in China

Understanding what factors generate geographic variation in species richness is a fundamental goal of ecology and biogeography. Water and energy are considered as the major environmental factors influencing large-scale patterns of species richness, but their roles vary among taxa and regions. Pteridophytes are an ideal group of organisms for examining the relationship between species richness and their environment because the distribution of pteridophytes is usually in equilibrium with contemporary climate to a greater degree than those of seed plants and most terrestrial vertebrates partly due to the lightness of their spores, which is highly capable of long-distance dispersal by wind, and partly due to their single-spore reproduction strategy. Using correlation and regression analyses and structural equation modeling technique, we examine the relationship of pteridophyte species richness in 151 localities from across China with environmental factors representing energy, water, and energy–water balance. We found that pteridophyte species richness is correlated to water availability more strongly than to ambient energy. Furthermore, we found that of all environmental variables considered, energy–water balance has played the most important role in regulating pteridophyte species richness gradients in China.     

Multi-extent analysis of the relationship between pteridophyte species richness and climate

Aim To determine the relationship between the distribution of climate, climatic heterogeneity and pteridophyte species richness gradients in Australia, using an approach that does not assume potential relationships are spatially invariant and allows for scale effects (extent of analysis) to be explicitly examined. Location Australia, extending from 10° S to 43° S and 112° E to 153° E. Method Species richness within 50 × 50 km grid cells was determined using point distribution data. Climatic surfaces representing the distribution and availability of water and energy at 1 km and 5 km cell resolutions were obtained. Climate at the 50 km resolution of analysis was represented by their mean and standard deviation in that area. Relationships were assessed using geographically weighted linear regression at a range of spatial bandwidths to investigate scale effects. Results The parameters and the predictive strength of all models varied across space at all extents of analysis. Overall, climatic variables representing water availability were more highly correlated to pteridophyte richness gradients in Australia than those representing energy. Their variance in cells further increased the strength of the relationships in topographically heterogeneous regions. Relationships with water were strong across all extents of analysis, particularly in the tropical and subtropical parts of the continent. Water availability explained less of the variation in richness at higher latitudes. Main conclusions This study brings into question the ability of aspatial and single‐extent models, searching for a unified explanation of macro‐scaled patterns in gradients of diversity, to adequately represent reality. It showed that, across Australia, there is a positive relationship between pteridophyte species richness and water availability but the strength and nature of the relationship varies spatially with scale in a highly complex manner. The spatial variance, or actual complexity, in these relationships could not have been demonstrated had a traditional aspatial global regression approach been used. Regional scale variation in relationships may be at least as important as more general relationships for a true understanding of the distribution of broad‐scale diversity.     

Pteridophyte richness, climate and topography in the Iberian Peninsula: comparing spatial and nonspatial models of richness patterns

Aim To describe the spatial variation in pteridophyte species richness; evaluate the importance of macroclimate, topography and within‐grid cell range variables; assess the influence of spatial autocorrelation on the significance of the variables; and to test the prediction of the mid‐domain effect. Location The Iberian Peninsula. Methods We estimated pteridophyte richness on a grid map with c. 2500 km² cell size, using published geocoded data of the individual species. Environmental data were obtained by superimposing the grid system over isoline maps of precipitation, temperature, and altitude. Mean and range values were calculated for each cell. Pteridophyte richness was related to the environmental variables by means of nonspatial and spatial generalized least squares models. We also used ordinary least squares regression, where a variance partitioning was performed to partial out the spatial component, i.e. latitude and longitude. Coastal and central cells were compared to test the mid‐domain effect. Results Both spatial and nonspatial models showed that pteridophyte richness was best explained by a second‐order polynomial of mean annual precipitation and a quadratic elevation‐range term, although the relative importance of these two variables varied when spatial autocorrelation was accounted for. Precipitation range was weakly significant in a nonspatial multiple model (i.e. ordinary regression), and did not remain significant in spatial models. Richness is significantly higher along the coast than in the centre of the peninsula. Main conclusions Spatial autocorrelation affects the statistical significance of explanatory variables, but this did not change the biological interpretation of precipitation and elevation range as the main predictors of pteridophyte richness. Spatial and nonspatial models gave very similar results, which reinforce the idea that water availability and topographic relief control species richness in relatively high‐energy regions. The prediction of the mid‐domain effect is falsified.     

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

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

Iberian–Balearic fern regions and their explanatory variables

This article delineates the compositional regions present in the Iberian–Balearic fern flora and compares these regions to previously proposed biogeographic units. It also assesses the extent to which environmental variables could explain the regions and the fern species richness gradients found within them. A combination of 40 previously published and new maps were used to compile the distribution of 123 pteridophytes on a 50 × 50 km UTM grid. Cluster analysis of the resulting 257 squares was used to classify 10 regions based on fern species assemblages. Discriminant function analysis identified the environmental variables that best explained these fern composition regions. Using generalized linear models; the number of species in each square was regressed against topography, climate, geology, environmental diversity, land use and spatial variables within each region. Two main latitudinal pteridophyte zones can be recognized in the Iberian Peninsula. These two zones are longitudinally subdivided into two sub zones. The 10 regions established significantly differ both in species richness and influential environmental variables. Climatic variables discriminate the most among regions, followed by topography, heterogeneity and geology. Pteridophyte richness varies, with richer areas being located along the coast and the main mountain ranges and the poorest areas being in the central plateaus and some north eastern and south western river basins. Species richness variation in Iberia is positively correlated with altitude range, precipitation, maximum altitude and area with siliceous soils. It is negatively correlated with the total annual days of sun, however. The fact that species richness is explained by different variables within each of the 10 regions indicates that the specific factors determining the spatial distribution of species richness vary from region to region. Some coastal regions are poorly explained by the model, and display a negative correlation with the selected causal factors. This finding suggests that persistent historic effects might play a local role in determining species assemblages in these regions. An erratum to this article can be found at     

Mesoscale Gradients of Herb Richness and Abundance in Central Amazonia1

There are few hypotheses to explain local understory diversity patterns. There is a consensus that climate and soil fertility affect understory density and diversity at large scales, but few studies addressed the mechanisms controlling density and diversity locally. Here, I examine patterns of abundance and diversity of three understory herb groups along gradients of soil nutrients and topography at the mesoscale (64 km²) in a wet tropical forest, and possible factors causing them. Herb richness, diversity, density, and cover were measured in fifty‐nine 250 × 2 m plots systematically distributed over Reserva Ducke, Manaus. Herb groups responded differently to environmental gradients. Whereas density and cover of pteridophytes increased with altitude and slope, Marantaceae density and cover decreased. Density of sedges increased with altitude, but did not vary with slope. Density and cover of Marantaceae and sedges but not pteridophytes increased with the soil cation content. Pteridophyte richness increased with slope whereas Marantaceae richness decreased, richness of both groups increased with cation content. Diversity increased with altitude for Marantaceae and decreased for pteridophytes. Some of these patterns agree with what is expected from herbs, such as the greater abundance of Marantaceae and sedges in flat and low altitude plots, where water availability is higher and probably also light, and the greater richness of Marantaceae and pteridophytes in higher nutrient plots. The unexpected results of higher abundance and richness of pteridophytes in slopes, instead of in bottomlands, suggest that biotic or litter‐mediated controls may be important to set these patterns.     

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

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

Relationships between spatial environmental heterogeneity and plant species diversity on a limestone pavement

No empirical studies have examined the relationship between diversity and spatial heterogeneity across unimodal species richness gradients. We determined the relationships between diversity and environmental factors for 144 0.18 m² plots in a limestone pavement alvar in southern Ontario, Canada, including within-plot spatial heterogeneity in soil depth, microtopography and microsite composition. Species richness was unimodally related to mean soil depth and relative elevation. Microsite heterogeneity and soil depth heterogeneity were positively correlated with species richness, and the richness peaks of the unimodal gradients correspond to the maximally spatially heterogeneous plots. The best predictive models of species richness and evenness, however, showed that other factors, such as ramet density and flooding, are the major determinants of diversity in this system. The findings that soil depth heterogeneity had effects on diversity when the effects of mean soil depth were factored out, and that unimodal richness peaks were associated with high spatial heterogeneity in environmental factors represent significant contributions to our understanding of how spatial heterogeneity might contribute to diversity maintenance in plant communities.     

Vegetation heterogeneity and diversity in flat and mountain landscapes of Patagonia (Argentina)

Abstract. We studied floristic and diversity patterns and their environmental controls in two landscapes of contrasting topography in the Patagonian steppe. The analyses were focused on the effects of water availability gradients and landscape configuration on plant species distribution and coexistence. Floristic variation was investigated using Correspondence Analysis. The relationship between floristic and environmental variation was analyzed using Canonical Correspondence Analysis and correlation tests. We explored diversity patterns by relating spatial distance to floristic dissimilarities. The floristic gradient was determined by shrub and grass species and was related to precipitation in the flat area, and to precipitation, elevation and potential radiation in the mountain area. Site species richness increased with water availability in both areas. Mean site species richness and species turnover in space was higher in the mountain than in the flat area. Landscape species richness and floristic gradients were more concentrated in the mountain than in the flat area. In contrast to shrubs and grasses, forb species distributions were uncoordinated and probably independent of any environmental gradient. Our results suggest (1) that landscape configuration affects species composition and diversity through its direct effect on abiotic environmental heterogeneity, and (2) that the environmental controls of the community composition vary depending on the plant functional type considered.     

Shifts in trait means and variances in North American tree assemblages: species richness patterns are loosely related to the functional space

One of the key hypothesized drivers of gradients in species richness is environmental filtering, where environmental stress limits which species from a larger species pool gain membership in a local community owing to their traits. Whereas most studies focus on small‐scale variation in functional traits along environmental gradient, the effect of large‐scale environmental filtering is less well understood. Furthermore, it has been rarely tested whether the factors that constrain the niche space limit the total number of coexisting species. We assessed the role of environmental filtering in shaping tree assemblages across North America north of Mexico by testing the hypothesis that colder, drier, or seasonal environments (stressful conditions for most plants) constrain tree trait diversity and thereby limit species richness. We assessed geographic patterns in trait filtering and their relationships to species richness pattern using a comprehensive set of tree range maps. We focused on four key plant functional traits reflecting major life history axes (maximum height, specific leaf area, seed mass, and wood density) and four climatic variables (annual mean and seasonality of temperature and precipitation). We tested for significant spatial shifts in trait means and variances using a null model approach. While we found significant shifts in mean species’ trait values at most grid cells, trait variances at most grid cells did not deviate from the null expectation. Measures of environmental harshness (cold, dry, seasonal climates) and lower species richness were weakly associated with a reduction in variance of seed mass and specific leaf area. The pattern in variance of height and wood density was, however, opposite. These findings do not support the hypothesis that more stressful conditions universally limit species and trait diversity in North America. Environmental filtering does, however, structure assemblage composition, by selecting for certain optimum trait values under a given set of conditions.     

Water–energy,land‐cover and heterogeneity drivers of the distribution of plant species richness in a mountain region of the European Alps

Aim To evaluate the relative importance of water–energy, land‐cover, environmental heterogeneity and spatial variables on the regional distribution of Red‐Listed and common vascular plant species richness. Location Trento Province (c. 6200 km²) on the southern border of the European Alps (Italy), subdivided regularly into 228 3′ × 5′ quadrants. Methods Data from a floristic inventory were separated into two subsets, representing Red‐Listed and common (i.e. all except Red‐Listed) plant species richness. Both subsets were separately related to water–energy, land‐cover and environmental heterogeneity variables. We simultaneously applied ordinary least squares regression with variation partitioning and hierarchical partitioning, attempting to identify the most important factors controlling species richness. We combined the analysis of environmental variables with a trend surface analysis and a spatial autocorrelation analysis. Results At the regional scale, plant species richness of both Red‐Listed and common species was primarily related to energy availability and land cover, whereas environmental heterogeneity had a lesser effect. The greatest number of species of both subsets was found in quadrants with the largest energy availability and the greatest degree of urbanization. These findings suggest that the elevation range within our study region imposes an energy‐driven control on the distribution of species richness, which resembles that of the broader latitude gradient. Overall, the two species subsets had similar trends concerning the relative importance of water–energy, land cover and environmental heterogeneity, showing a few differences regarding the selection of some predictors of secondary importance. The incorporation of spatial variables did not improve the explanatory power of the environmental models and the high original spatial autocorrelation in the response variables was reduced drastically by including the selected environmental variables. Main conclusions Water–energy and land cover showed significant pure effects in explaining plant species richness, indicating that climate and land cover should both be included as explanatory variables in modelling species richness in human‐affected landscapes. However, the high degree of shared variation between the two groups made the relative effects difficult to separate. The relatively low range of variation in the environmental heterogeneity variables within our sampling domain might have caused the low importance of this complex factor.     

Fern and lycophyte diversity in the Pacific Northwest: Patterns and predictors

Recent floristic efforts in the Pacific Northwest (PNW) have now made it possible to characterize the broad‐scale patterns of fern and lycophyte diversity across this large and geologically‐complex region of western North America. The physiography of the PNW has been developing for over 200 million years, but Pleistocene glaciation‐induced migrations and recolonizations have strongly influenced the assembly of the flora. With the high dispersal potential of fern and lycophyte spores, the distribution patterns of pteridophytes may be representative of habitat suitability more than dispersal constraints. Our objective was to describe the biodiversity of pteridophytes in the PNW, determine the spatial distribution of that biodiversity in terms of phylogenetic diversity, identify centers of regional endemism, explore the correlations between biodiversity and environmental variables, and infer possible influences of past glaciation on the pteridophyte flora. We obtained presence‐only distribution data from two online databases. A phylogenetic tree was constructed using chloroplast DNA sequence data from GenBank. We used the Biodiverse software package to estimate and map phylogenetic diversity and phylogenetic endemism across the PNW, and to identify those regions of the PNW where diversity was higher or lower than expected in comparison to randomization models. Environmental correlates of diversity were identified using principal components analysis with bioclimatic data from WorldClim.org, and we used Maxent to predict habitat suitability for species under past and future climate conditions. We found evidence for the influence of past glaciations and glacial refugia on the patterns of pteridophyte diversity, that moisture availability and cold temperatures are strongly correlated with patterns of genus richness, phylogenetic diversity, and phylogenetic endemism. We infer that the topographic complexity of the region may be driving the assembly of the pteridophyte flora indirectly by influencing climate and precipitation patterns.     

Plant species diversity and environmental heterogeneity: spatial scale and competing hypotheses

Questions: What is the observed relationship between plant species diversity and spatial environmental heterogeneity? Does the relationship scale predictably with sample plot size? What are the relative contributions to diversity patterns of variables linked to productivity or available energy compared to those corresponding to spatial heterogeneity? Methods: Observational and experimental studies that quantified relationships between plant species richness and within‐sample spatial environmental heterogeneity were reviewed. Effect size in experimental studies was quantified as the standardized mean difference between control (homogeneous) and heterogeneous treatments. For observational studies, effect sizes in individual studies were examined graphically across a gradient of plot size (focal scale). Relative contributions of variables representing spatial heterogeneity were compared to those representing available energy using a response ratio. Results: Forty‐one observational and 11 experimental studies quantified plant species diversity and spatial environmental heterogeneity. Observational studies reported positive species diversity‐spatial heterogeneity correlations at all points across a plot size gradient from ～1.0 × 10^?1 to ～1.0 × 10¹¹ m², although many studies reported spatial heterogeneity variables with no significant relationships to species diversity. The cross‐study effect size in experimental studies was not significantly different from zero. Available energy variables explained consistently more of the variance in species richness than spatial heterogeneity variables, especially at the smallest and largest plot sizes. Main conclusions: Species diversity was not related to spatial heterogeneity in a way predictable by plot size. Positive heterogeneity‐diversity relationships were common, confirming the importance of niche differentiation in species diversity patterns, but future studies examining a range of spatial scales in the same system are required to determine the role of dispersal and available energy in these patterns.     

Environmental correlates of mammal species richness in South America: effects of spatial structure, taxonomy and geographic range

Although some consensus exists regarding the positive synergism between energy and heterogeneity in increasing species diversity, the role of environmental variability remains controversial. We examine how these factors interact to explain spatial variation in mammal species richness in South America. After taking into account the effects of spatial autocorrelation and area, elevation variability and energy mainly drive spatial variation in mammal species richness. The effect of environmental variability is less important. When different taxonomic groups of mammals are analyzed separately, three ways emerge whereby energy and heterogeneity interact to promote species richness. Heterogeneity may have no effect on species richness, habitat heterogeneity and energy availability contribute independently to species richness, or heterogeneity increases in importance with an increase in energy availability. The partition of species into range size quartiles shows that habitat heterogeneity and temporal instability in the resource supply account for the species richness pattern in the narrowest- ranging species. Habitat heterogeneity is significant also for intermediate ranging species but not for the widest-ranging species. Energy alone drives the species richness pattern in the latter species. The interplay between ecology and biogeographic history may ultimately explain these differences given that narrow- and wide-ranging species show distinct biogeographic patterns, and different taxonomic groups also unequally represent them.     

Regional enrichment of local assemblages is robust to variation in local productivity, abiotic gradients, and heterogeneity

Theory predicts that the effects of regional richness on the richness of local communities may depend on the productivity, resource availability, and/or heterogeneity of local sites. Using the wetland plant communities of 50 independent streams as 'regions', we tested whether: (1) local richness in 1-m² quadrats and 50-m stream segments was positively related to regional richness, even after environmental influences were considered; and (2) the effect of regional richness would interact with the effects of biomass, soil moisture, and/or heterogeneity on local richness. In models that explained up to 88% of variation in local richness, we found that richness at both local scales was positively related to regional richness, and that regional richness did not interact with any of the environmental gradients that also shaped local richness. We conclude that species availability from the regional pool may consistently enrich local communities, even while other constraints on local richness operate.     

Predicting patterns of plant species richness in megadiverse South Africa

Using new tools (boosted regression trees) in predictive biogeography, with extensive spatial 23 distribution data for >19 000 species, we developed predictive models for South African plant species richness patterns. Further, biome level analysis explored possible functional determinants of country‐wide regional species richness. Finally, to test model reliability independently, we predicted potential alien invasive plant species richness with an independent dataset. Amongst the different hypotheses generally invoked to explain species 30 diversity (energy, favorableness, topographic heterogeneity, irregularity and seasonality), results revealed topographic heterogeneity as the most powerful single explanatory variable for indigenous South African plant species richness. Some biome‐specific responses were observed, i.e. two of the five analyzed biomes (Fynbos and Grassland) had richness best explained by the “species‐favorableness” hypothesis, but even in this case, topographic heterogeneity was also a primary predictor. This analysis, the largest conducted on an almost exhaustive species sample in a species‐rich region, demonstrates the preeminence of topographic heterogeneity in shaping the spatial pattern of regional plant species richness. Model reliability was confirmed by the considerable predictive power for alien invasive species richness. It thus appears that topographic heterogeneity controls species richness in two main ways: firstly, by providing an abundance of ecological niches in contemporary space (revealed by alien invasive species richness relationships) and secondly, by facilitating the persistence of ecological niches through time. The extraordinary richness of the South African Fynbos biome, a world‐renowned hotspot of biodiversity with the steepest environmental gradients in South Africa, may thus have arisen through both mechanisms. Comparisons with similar regions of the world outside South Africa are needed to confirm the generality of topographic heterogeneity and favorableness as predictors of plant richness.     

Host trait preferences and distribution of vascular epiphytes in a warm-temperate forest

To illustrate the ecological factors and process leading to the observed diversity patterns of vascular epiphytes, we examined the effect and importance of host tree traits on epiphyte richness and spatial aggregation of epiphytes. The study was conducted in warm-temperate forest in Japan. The recorded host traits were diameter, height, species, habitat topography, and growth rate, and we analyzed the effects and importance of these traits on three species groups: total epiphytic species, epiphytic orchid species, and epiphytic pteridophyte species. Diameter and species of host trees had the greatest influence on epiphytes and their magnitudes were roughly similar in all species groups. Growth rate and topography were less important than host size and species. Growth rate had a negative effect on all three groups, and topography was important for pteridophytes. Epiphyte richness did not exhibit clear spatial aggregation. Our results suggest that size, stability, and quality of the host are equally important in determining epiphyte colonization.     

The biogeography and filtering of woody plant functional diversity in North and South America

Aim In recent years evidence has accumulated that plant species are differentially sorted from regional assemblages into local assemblages along local‐scale environmental gradients on the basis of their function and abiotic filtering. The favourability hypothesis in biogeography proposes that in climatically difficult regions abiotic filtering should produce a regional assemblage that is less functionally diverse than that expected given the species richness and the global pool of traits. Thus it seems likely that differential filtering of plant traits along local‐scale gradients may scale up to explain the distribution, diversity and filtering of plant traits in regional‐scale assemblages across continents. The present work aims to address this prediction. Location North and South America. Methods We combine a dataset comprising over 5.5 million georeferenced plant occurrence records with several large plant functional trait databases in order to: (1) quantify how several critical traits associated with plant performance and ecology vary across environmental gradients; and (2) provide the first test of whether the woody plants found within 1° and 5° map grid cells are more or less functionally diverse than expected, given their species richness, across broad gradients. Results The results show that, for many of the traits studied, the overall distribution of functional traits in tropical regions often exceeds the expectations of random sampling given the species richness. Conversely, temperate regions often had narrower functional trait distributions than their smaller species pools would suggest. Main conclusion The results show that the overall distribution of function does increase towards the equator, but the functional diversity within regional‐scale tropical assemblages is higher than that expected given their species richness. These results are consistent with the hypothesis that abiotic filtering constrains the overall distribution of function in temperate assemblages, but tropical assemblages are not as tightly constrained.     

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.