[Show abstract][Hide abstract] ABSTRACT: AimTo reconstruct the historical assembly of the eudicot flora of Mediterranean sierras by examining compositional (CBD), phylogenetic (PBD) and functional (FBD) beta diversity between elevational belts among disjunct mountain ranges (sierras), and relating these measures of turnover to environmental and geographical distances.LocationBaetic ranges, Andalusia, southern Spain.Methods
We compiled eudicot species and subspecies (‘entities’) checklists for three elevational belts within each of eight sierras of Andalusia (‘sites’) and tested for non-random patterns of PBD and FBD of all entities and of endemic entities separately among sites between and within sierras. Multiple regression on distance matrices was used to determine the respective contribution of climate, lithology and geographical distance to CBD, PBD and FBD. Finally, we decomposed PBD into the turnover and nestedness components of beta diversity, and quantified the phylogenetic diversity (PD) within sites.ResultsThe observed PBD and FBD among elevational belts within sierras for all entities were generally higher than expected based on their respective null distributions, whereas CBD among elevational belts within sierras was similar or even lower than between sierras. In contrast, the observed PBD and FBD for endemics were non-significant in most of the comparisons. Temperature-related variables best explained patterns of CBD, PBD and FBD for all entities, whereas lithology and geographical distance were the main drivers of endemic CBD. The observed PBD among elevational belts within sierras was mainly attributable to differences in PD rather than ‘true’ turnover.Main conclusionsThere is strong structuring of plant lineages along elevational gradients in the Baetic range, probably due to habitat filtering acting on life forms and character syndromes that show strong phylogenetic signal. The differentiation of the endemic flora that contributed to the emergence of this western Mediterranean biodiversity hotspot was probably driven by geographical isolation and/or by repeated specialization to contrasting lithologies.
Journal of Biogeography 08/2014; · 4.86 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: We investigate patterns of phylogenetic diversity in relation to species diversity for European birds, mammals and amphibians, to evaluate their congruence and highlight areas of particular evolutionary history. We estimate the extent to which the European network of protected areas (PAs) network retains interesting evolutionary history areas for the three groups separately and simultaneously.
Phylogenetic (QEPD) and species diversity (SD) were estimated using the Rao's quadratic entropy at 10' resolution. We determined the regional relationship between QEPD and SD for each taxa with a spatial regression model and used the tails of the residuals (QERES) distribution to identify areas of higher and lower QEPD than predicted. Spatial congruence of biodiversity between groups was assessed with Pearson's correlation. A simple classification scheme allowed building a convergence map where a convergent pixel equalled to a QERES value of the same sign for the 3 groups. This convergence map was overlaid to the current PAs network to estimate the level of protection in convergent pixels and compared it to a null expectation built on 1000 randomization of PAs over the landscape.
QERES patterns across vertebrates show a strong spatial mismatch highlighting different evolutionary histories. Convergent areas represent only 2.7% of the Western Palearctic, with only 8.4% of these areas being covered by the current PAs network while a random distribution would retain 10.4% of them. QERES are unequally represented within PAs: areas with higher QEPD than predicted are better covered than expected, while low QEPD areas are undersampled.
Patterns of diversity strongly diverge between groups of vertebrates in Europe. Although Europe has the world's most extensive PAs network, evolutionary history of terrestrial vertebrates is unequally protected. The challenge is now to reconcile effective conservation planning with a contemporary view of biodiversity integrating multiple facets.
Diversity and Distributions 06/2014; 20(6):674-685. · 6.12 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Evolutionary adaptation is a key driver of species’ range dynamics. Understanding the factors that affect rates of adaptation at range margins is thus crucial for interpreting and predicting changes in species’ ranges. The spatial structure of environmental conditions is one of the determinants of whether and how quickly adaptations occur. However, while landscape structures at range edges are typically complex, most theoretical work has so far focused on relatively simple environmental geometries.Using an individual-based allelic model, we explore the effects of different landscape structures on the rate of adaptation to novel environments and investigate how these structures interact with the genetic architecture of the trait governing adaptation and the dispersal capacity of the considered species. Generally, we find that rapid adaptation is favored by a good match between the coarseness of the trait's genetic architecture (many loci of small effects versus few loci of large effects) and the coarseness of the landscape (abruptness of transitions in environmental conditions). For example, in rugged landscapes, adaptation is quicker for genetic architectures with few loci of large effects, while for shallow gradients the opposite is true. Moreover, dispersal capacities affect the rate of adaptation by modulating the ‘apparent coarseness’ of the landscape: a gradient perceived as smooth by species with limited dispersal capacities appears rather steep for highly dispersive ones. We also find that the distribution of evolving phenotypes strongly depends on the interplay of landscape structure and dispersal capacities, ranging from two distinct phenotypes for most rugged landscapes, over the co-occurrence of an additional third phenotype for highly dispersive species, to the whole range of phenotypes on smooth gradients.By identifying basic factors that drive the fixation probability of newly arising beneficial mutations, we hope to further broaden the understanding of evolutionary adaptation at range margins and, hence, species’ range dynamics.
[Show abstract][Hide abstract] ABSTRACT: Abstract Recent debate on whether climatic niches are conserved through time has focused on how phylogenetic niche conservatism can be measured by deviations from a Brownian motion model of evolutionary change. However, there has been no evaluation of this methodological approach. In particular, the fact that climatic niches are usually obtained from distribution data and are thus heavily influenced by biogeographic factors has largely been overlooked. Our main objective here was to test whether patterns of climatic niche evolution that are frequently observed might arise from neutral dynamics rather than from adaptive scenarios. We developed a model inspired by neutral biodiversity theory, where individuals disperse, compete, and undergo speciation independently of climate. We then sampled the climatic niches of species according to their geographic position and showed that even when species evolve independently of climate, their niches can nonetheless exhibit evolutionary patterns strongly differing from Brownian motion. Indeed, climatic niche evolution is better captured by a model of punctuated evolution with constraints due to landscape boundaries, two features that are traditionally interpreted as evidence for selective processes acting on the niche. We therefore suggest that deviation from Brownian motion alone should not be used as evidence for phylogenetic niche conservatism but that information on phenotypic traits directly linked to physiology is required to demonstrate that climatic niches have been conserved through time.
The American Naturalist 05/2014; 183(5):573-84. · 4.55 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Biological invasions can transform our understanding of how the interplay of historical isolation and contemporary (human-aided) dispersal affects the structure of intraspecific diversity in functional traits, and in turn, how changes in functional traits affect other scales of biological organization such as communities and ecosystems. Because biological invasions frequently involve the admixture of previously isolated lineages as a result of human-aided dispersal, studies of invasive populations can reveal how admixture results in novel genotypes and shifts in functional trait variation within populations. Further, because invasive species can be ecosystem engineers within invaded ecosystems, admixture-induced shifts in the functional traits of invaders can affect the composition of native biodiversity and alter the flow of resources through the system. Thus, invasions represent promising yet under-investigated examples of how the effects of short-term evolutionary changes can cascade across biological scales of diversity. Here, we propose a conceptual framework that admixture between divergent source populations during biological invasions can reorganize the genetic variation underlying key functional traits, leading to shifts in the mean and variance of functional traits within invasive populations. Changes in the mean or variance of key traits can initiate new ecological feedback mechanisms that result in a critical transition from a native ecosystem to a novel invasive ecosystem. We illustrate the application of this framework with reference to a well-studied plant model system in invasion biology and show how a combination of quantitative genetic experiments, functional trait studies, whole ecosystem field studies and modeling can be used to explore the dynamics predicted to trigger these critical transitions.
Ecology and Evolution 04/2014; 4(7):899-910. · 1.66 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: The origins of ecological diversity in continental species assemblages have long intrigued biogeographers. We apply phylogenetic comparative analyses to disentangle the evolutionary patterns of ecological niches in an assemblage of European birds. We compare phylogenetic patterns in trophic, habitat and climatic niche components.
From polygon range maps and handbook data we inferred the realized climatic, habitat and trophic niches of 405 species of breeding birds in Europe. We fitted Pagel's lambda and kappa statistics, and conducted analyses of disparity through time to compare temporal patterns of ecological diversification on all niche axes together. All observed patterns were compared with expectations based on neutral (Brownian) models of niche divergence.
In this assemblage, patterns of phylogenetic signal (lambda) suggest that related species resemble each other less in regard to their climatic and habitat niches than they do in their trophic niche. Kappa estimates show that ecological divergence does not gradually increase with divergence time, and that this punctualism is stronger in climatic niches than in habitat and trophic niches. Observed niche disparity markedly exceeds levels expected from a Brownian model of ecological diversification, thus providing no evidence for past phylogenetic niche conservatism in these multivariate niches. Levels of multivariate disparity are greatest for the climatic niche, followed by disparity of the habitat and the trophic niches.
Phylogenetic patterns in the three niche components differ within this avian assemblage. Variation in evolutionary rates (degree of gradualism, constancy through the tree) and/or non-random macroecological sampling probably lead here to differences in the phylogenetic structure of niche components. Testing hypotheses on the origin of these patterns requires more complete phylogenetic trees of the birds, and extended ecological data on different niche components for all bird species.
Global Ecology and Biogeography 04/2014; 23(4):414-424. · 7.22 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Climate and land cover changes are important drivers of the plant species distributions and diversity patterns in mountainous regions. Although the need for a multifaceted view of diversity based on taxonomic, functional and phylogenetic dimensions is now commonly recognized, there are no complete risk assessments concerning their expected changes. In this paper, we used a range of species distribution models in an ensemble-forecasting framework together with regional climate and land cover projections by 2080 to analyze the potential threat for more than 2500 plant species at high resolution (2.5 × 2.5 km) in the French Alps. We also decomposed taxonomic, functional and phylogenetic diversity facets into α and β components and analyzed their expected changes by 2080. Overall, plant species threats from climate and land cover changes in the French Alps were expected to vary depending on the species’ preferred altitudinal vegetation zone, rarity, and conservation status. Indeed, rare species and species of conservation concern were the ones projected to experience less severe change, and also the ones being the most efficiently preserved by the current network of protected areas. Conversely, the three facets of plant diversity were also projected to experience drastic spatial re-shuffling by 2080. In general, the mean α-diversity of the three facets was projected to increase to the detriment of regional β-diversity, although the latter was projected to remain high at the montane-alpine transition zones. Our results show that, due to a high-altitude distribution, the current protection network is efficient for rare species, and species predicted to migrate upward. Although our modeling framework may not capture all possible mechanisms of species range shifts, our work illustrates that a comprehensive risk assessment on an entire floristic region combined with functional and phylogenetic information can help delimitate future scenarios of biodiversity and better design its protection.
[Show abstract][Hide abstract] ABSTRACT: Despite the recognized joint impact of climate and land cover change on facets of biodiversity and their associated functions, risk assessments have primarily evaluated impacts on species ranges and richness. Here we quantify the sensitivity of the functional structure of European avian assemblages to changes in both regional climate and land cover. We combine species range forecasts with functional-trait information. We show that species sensitivity to environmental change is randomly distributed across the functional tree of the European avifauna and that functionally unique species are not disproportionately threatened by 2080. However, projected species range changes will modify the mean species richness and functional diversity of bird diets and feeding behaviours. This will unequally affect the spatial structure of functional diversity, leading to homogenization across Europe. Therefore, global changes may alter the functional structure of species assemblages in the future in ways that need to be accounted for in conservation planning.
[Show abstract][Hide abstract] ABSTRACT: Aim Phylogenetic diversity patterns are increasingly being used to better understand the role of ecological and evolutionary processes in community assembly. Here, we quantify how these patterns are influenced by scale choices in terms of spatial and environmental extent and organismic scales. LocationEuropean Alps.
Methods We applied 42 sampling strategies differing in their combination of focal scales. For each resulting sub-dataset, we estimated the phylogenetic diversity of the species pools, phylogenetic α-diversities of local communities, and statistics commonly used together with null models in order to infer non-random diversity patterns (i.e. phylogenetic clustering versus over-dispersion). Finally, we studied the effects of scale choices on these measures using regression analyses.
Results Scale choices were decisive for revealing signals in diversity patterns. Notably, changes in focal scales sometimes reversed a pattern of over-dispersion into clustering. Organismic scale had a stronger effect than spatial and environmental extent. However, we did not find general rules for the direction of change from over-dispersion to clustering with changing scales. Importantly, these scale issues had only a weak influence when focusing on regional diversity patterns that change along abiotic gradients.
Main conclusions Our results call for caution when combining phylogenetic data with distributional data to study how and why communities differ from random expectations of phylogenetic relatedness. These analyses seem to be robust when the focus is on relating community diversity patterns to variation in habitat conditions, such as abiotic gradients. However, if the focus is on identifying relevant assembly rules for local communities, the uncertainty arising from a certain scale choice can be immense. In the latter case, it becomes necessary to test whether emerging patterns are robust to alternative scale choices.
Global Ecology and Biogeography 01/2014; · 7.22 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: We still have limited understanding of the contingent and deterministic factors that have fostered the evolutionary success of some species lineages over others. We investigated how the interplay of intercontinental migration and key innovations promoted diversification of the genus Androsace.
Mountain ranges and cold steppes of the Northern Hemisphere.
We reconstructed ancestral biogeographical ranges at regional and continental scales by means of a dispersal-extinction-cladogenesis analysis using dated Bayesian phylogenies and contrasting two migration scenarios. Based on diversification analyses under two frameworks, we tested the influence of life form on speciation rates and whether diversification has been diversity-dependent.
We found that three radiations occurred in this genus, at different periods and on different continents, and that life form played a critical role in the history of Androsace. Short-lived ancestors first facilitated the expansion of the genus' range from Asia to Europe, while cushions, which appeared independently in Asia and Europe, enhanced species diversification in alpine regions. One long-distance dispersal event from Europe to North America led to the diversification of the nested genus Douglasia. We found support for a model in which speciation of the North American-European clade is diversity-dependent and close to its carrying capacity, and that the diversification dynamics of the North American subclade are uncoupled from this and follow a pure birth process.
The contingency of past biogeographical connections combined with the evolutionary determinism of convergent key innovations may have led to replicated radiations of Androsace in three mountain regions of the world. The repeated emergence of the cushion life form was a convergent key innovation that fostered radiation into alpine habitats. Given the large ecological similarity of Androsace species, allopatry may have been the main mode of speciation.
Journal of Biogeography 10/2013; 40(10):1874-1886. · 4.86 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Species distribution models (SDMs) have been frequently employed to forecast the response of alpine plants to global changes. Efforts to model alpine plant distribution have thus far been primarily based on a correlative approach, in which ecological processes are implicitly addressed through a statistical relationship between observed species occurrences and environmental predictors. Recent evidence, however, highlights the shortcomings of correlative SDMs, especially in alpine landscapes where plant species tend to be decoupled from atmospheric conditions in micro-topographic habitats and are particularly exposed to geomorphic disturbances. While alpine plants respond to the same limiting factors as plants found at lower elevations, alpine environments impose a particular set of scale-dependent and hierarchical drivers that shape the realized niche of species and that require explicit consideration in a modelling context. Several recent studies in the European Alps have successfully integrated both correlative and process-based elements into distribution models of alpine plants, but for the time being a single integrative modelling framework that includes all key drivers remains elusive. As a first step in working toward a comprehensive integrated model applicable to alpine plant communities, we propose a conceptual framework that structures the primary mechanisms affecting alpine plant distributions. We group processes into four categories, including multi-scalar abiotic drivers, gradient dependent species interactions, dispersal and spatial-temporal plant responses to disturbance. Finally, we propose a methodological framework aimed at developing an integrated model to better predict alpine plant distribution.
[Show abstract][Hide abstract] ABSTRACT: Metacommunity theories attribute different relative degrees of importance to dispersal, environmental filtering, biotic interactions and stochastic processes in community assembly, but the role of spatial scale remains uncertain. Here we used two complementary statistical tools to test: (1) whether or not the patterns of community structure and environmental influences are consistent across resolutions; and (2) whether and how the joint use of two fundamentally different statistical approaches provides a complementary interpretation of results.
Grassland plants in the French Alps.
We used two approaches across five spatial resolutions (ranging from 1 km × 1 km to 30 km × 30 km): variance partitioning, and analysis of metacommunity structure on the site-by-species incidence matrices. Both methods allow the testing of expected patterns resulting from environmental filtering, but variance partitioning allows the role of dispersal and environmental gradients to be studied, while analysis of the site-by-species metacommunity structure informs an understanding of how environmental filtering occurs and whether or not patterns differ from chance expectation. We also used spatial regressions on species richness to identify relevant environmental factors at each scale and to link results from the two approaches.
Major environmental drivers of richness included growing degree-days, temperature, moisture and spatial or temporal heterogeneity. Variance partitioning pointed to an increase in the role of dispersal at coarser resolutions, while metacommunity structure analysis pointed to environmental filtering having an important role at all resolutions through a Clementsian assembly process (i.e. groups of species having similar range boundaries and co-occurring in similar environments).
The combination of methods used here allows a better understanding of the forces structuring ecological communities than either one of them used separately. A key aspect in this complementarity is that variance partitioning can detect effects of dispersal whereas metacommunity structure analysis cannot. Moreover, the latter can distinguish between different forms of environmental filtering (e.g. individualistic versus group species responses to environmental gradients).
Journal of Biogeography 08/2013; 40(8):1560-1571. · 4.86 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Around the world, the human-induced collapses of populations and species have triggered a sixth mass extinction crisis, with rare species often being the first to disappear. Although the role of species diversity in the maintenance of ecosystem processes has been widely investigated, the role of rare species remains controversial. A critical issue is whether common species insure against the loss of functions supported by rare species. This issue is even more critical in species-rich ecosystems where high functional redundancy among species is likely and where it is thus often assumed that ecosystem functioning is buffered against species loss. Here, using extensive datasets of species occurrences and functional traits from three highly diverse ecosystems (846 coral reef fishes, 2,979 alpine plants, and 662 tropical trees), we demonstrate that the most distinct combinations of traits are supported predominantly by rare species both in terms of local abundance and regional occupancy. Moreover, species that have low functional redundancy and are likely to support the most vulnerable functions, with no other species carrying similar combinations of traits, are rarer than expected by chance in all three ecosystems. For instance, 63% and 98% of fish species that are likely to support highly vulnerable functions in coral reef ecosystems are locally and regionally rare, respectively. For alpine plants, 32% and 89% of such species are locally and regionally rare, respectively. Remarkably, 47% of fish species and 55% of tropical tree species that are likely to support highly vulnerable functions have only one individual per sample on average. Our results emphasize the importance of rare species conservation, even in highly diverse ecosystems, which are thought to exhibit high functional redundancy. Rare species offer more than aesthetic, cultural, or taxonomic diversity value; they disproportionately increase the potential breadth of functions provided by ecosystems across spatial scales. As such, they are likely to insure against future uncertainty arising from climate change and the ever-increasing anthropogenic pressures on ecosystems. Our results call for a more detailed understanding of the role of rarity and functional vulnerability in ecosystem functioning.
[Show abstract][Hide abstract] ABSTRACT: Functional variability (FV) of populations can be decomposed into three main features: the individual variability of multiple traits, the strength of correlations between those traits and the main direction of these correlations, the latter two being known as 'phenotypic integration'. Evolutionary biology has long recognized that FV in natural populations is key to determining potential evolutionary responses, but this topic has been little studied in functional ecology.Here we focus on the arctico-alpine perennial plant species Polygonum viviparum L.. We used a comprehensive sampling of seven functional traits in 29 wild populations covering the whole environmental niche of the species. The niche of the species was captured by a temperature gradient, which separated alpine stressful habitats from species-rich, competitive sub-alpine ones. We seeked to assess the relative roles of abiotic stress and biotic interactions in shaping different aspects of functional variation within and among populations, that is, the multi-trait variability, the strength of correlations between traits, and the main directions of functional trade-offs.Populations with the highest extent of functional variability were found in the warm end of the gradient whereas populations exhibiting the strongest degree of phenotypic integration were located in sites with intermediate temperatures. This could reveal both the importance of environmental filtering and population demography in structuring FV. Interestingly, we found that the main axes of multivariate functional variation were radically different within and across population.Although the proximate causes of FV structure remain uncertain, our study presents a robust methodology for the quantitative study of functional variability in connection with species' niches. It also opens up new perspectives for the conceptual merging of intraspecific functional patterns with community ecology.
[Show abstract][Hide abstract] ABSTRACT: Plants affect the spatial distribution of soil microorganisms, but the influence of the local
abiotic context is poorly documented. We investigated the effect of a single plant species, the cushion plant Silene acaulis, on habitat conditions, and microbial community. We col- lected soil from inside (In) and outside (Out) of the cushions on calcareous and siliceous cliffs in the French Alps along an elevation gradient (2,000–3,000 masl). The composition of the microbial communities was assessed by Capillary-Electrophoresis Single Strand Con- formation Polymorphism (CE-SSCP). Univariate and multivariate analyses were conducted to characterize the response of the microbial beta-diversity to soil parameters (total C, total N, soil water content, N−NH4 , N−NO3 , and pH). Cushions affected the microbial communities, modifying soil properties. The fungal and bacterial communities did not respond to the same abiotic factors. Outside the cushions, the bacterial communities were strongly influenced by bedrock. Inside the cushions, the bacterial communities from both types of bedrock were highly similar, due to the smaller pH differences than in open areas. By contrast, the fungal communities were equally variable inside and outside of the cush- ions. Outside the cushions, the fungal communities responded weakly to soil pH. Inside the cushions, the fungal communities varied
strongly with bedrock and elevation as well as increases in soil nutrients and water content.
Furthermore, the dissimilarities in the microbial communities between the In and Out habitats increased with increasing habitat modification and environmental stress.
Our results indicate that cushions act as a selective force that counteracts the influence of the bedrock and the resource limitations on the bac- terial and fungal communities by buffering soil pH and enhancing soil nutrients. Cushion plants structure microbial communities, and this effect increases in stressful, acidic and nutrient-limited environments.
Frontiers in Microbiology 03/2013; 4(64):1-14. · 3.90 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Dispersal is a key determinant of a population's evolutionary potential. It facilitates the propagation of beneficial alleles throughout the distributional range of spatially outspread populations and increases the speed of adaptation. However, when habitat is heterogeneous and individuals are locally adapted, dispersal may, at the same time, reduce fitness through increasing maladaptation. Here, we use a spatially explicit, allelic simulation model to quantify how these equivocal effects of dispersal affect a population's evolutionary response to changing climate. Individuals carry a diploid set of chromosomes, with alleles coding for adaptation to non-climatic environmental conditions and climatic conditions, respectively. Our model results demonstrate that the interplay between gene flow and habitat heterogeneity may decrease effective dispersal and population size to such an extent that substantially reduces the likelihood of evolutionary rescue. Importantly, even when evolutionary rescue saves a population from extinction, its spatial range following climate change may be strongly narrowed, that is, the rescue is only partial. These findings emphasize that neglecting the impact of non-climatic, local adaptation might lead to a considerable overestimation of a population's evolvability under rapid environmental change.
Philosophical Transactions of The Royal Society B Biological Sciences 01/2013; 368(1610):20120083. · 6.23 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Predicting how and when adaptive evolution might rescue species from global change, and integrating this process into tools of biodiversity forecasting, has now become an urgent task. Here, we explored whether recent population trends of species can be explained by their past rate of niche evolution, which can be inferred from increasingly available phylogenetic and niche data. We examined the assemblage of 409 European bird species for which estimates of demographic trends between 1970 and 2000 are available, along with a species-level phylogeny and data on climatic, habitat and trophic niches. We found that species' proneness to demographic decline is associated with slow evolution of the habitat niche in the past, in addition to certain current-day life-history and ecological traits. A similar result was found at a higher taxonomic level, where families prone to decline have had a history of slower evolution of climatic and habitat niches. Our results support the view that niche conservatism can prevent some species from coping with environmental change. Thus, linking patterns of past niche evolution and contemporary species dynamics for large species samples may provide insights into how niche evolution may rescue certain lineages in the face of global change.
Philosophical Transactions of The Royal Society B Biological Sciences 01/2013; 368(1610):20120091. · 6.23 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Questions: Traditional null models used to reveal assembly processes from functional diversity patterns are not tailored for comparing different spatial and evolutionary scales. In this study, we present and explore a family of null models that can help disentangling assembly processes at their appropriate scales and thereby elucidate the ecological drivers of community assembly. Location: French Alps. Methods: Our approach gradually constrains null models by: (1) filtering out species not able to survive in the regional conditions in order to reduce the spa-tial scale, and (2) shuffling species only within lineages of different ages to reduce the evolutionary scale of the analysis. We first tested and validated this approach using simulated communities. We then applied it to study the func-tional diversity patterns of the leaf–height–seed strategy of plant communities in the French Alps. Results: Using simulations, we found that reducing the spatial scale correctly detected a signature of competition (functional divergence) even when environ-mental filtering produced an overlaying signal of functional convergence. How-ever, constraining the evolutionary scale did not change the identified functional diversity patterns. In the case study of alpine plant communities, investigating scale effects revealed that environmental filtering had a strong influence at larger spatial and evolutionary scales and that neutral processes were more important at smaller scales. In contrast to the simulation study results, decreasing the evolutionary scale tended to increase patterns of func-tional divergence. Conclusion: We argue that the traditional null model approach can only iden-tify a single main process at a time and suggest to rather use a family of null models to disentangle intertwined assembly processes acting across spatial and evolutionary scales.
Journal of Vegetation Science 01/2013; 24:853-864. · 2.82 Impact Factor