Cyrille Violle

Phylogenetic limiting similarity and competitive exclusion

One of the oldest ecological hypotheses, proposed by Darwin, suggests that the struggle for existence is stronger between more closely related species. Despite its long history, the validity of this phylogenetic limiting similarity hypothesis has rarely been examined. Here we provided a formal experimental test of the hypothesis using pairs of bacterivorous protist species in a multigenerational experiment. Consistent with the hypothesis, both the frequency and tempo of competitive exclusion, and the reduction in the abundance of inferior competitors, increased with increasing phylogenetic relatedness of the competing species. These results were linked to protist mouth size, a trait potentially related to resource use, exhibiting a significant phylogenetic signal. The likelihood of coexistence, however, was better predicted by phylogenetic relatedness than trait similarity of the competing species. Our results support phylogenetic relatedness as a useful predictor of the outcomes of competitive interactions in ecological communities.

The emergence and promise of functional biogeography

Understanding, modeling, and predicting the impact of global change on ecosystem functioning across biogeographical gradients can benefit from enhanced capacity to represent biota as a continuous distribution of traits. However, this is a challenge for the field of biogeography historically grounded on the species concept. Here we focus on the newly emergent field of functional biogeography: the study of the geographic distribution of trait diversity across organizational levels. We show how functional biogeography bridges species-based biogeography and earth science to provide ideas and tools to help explain gradients in multifaceted diversity (including species, functional, and phylogenetic diversities), predict ecosystem functioning and services worldwide, and infuse regional and global conservation programs with a functional basis. Although much recent progress has been made possible because of the rising of multiple data streams, new developments in ecoinformatics, and new methodological advances, future directions should provide a theoretical and comprehensive framework for the scaling of biotic interactions across trophic levels and its ecological implications.

A global meta‐analysis of the relative extent of intraspecific trait variation in plant communities

Recent studies have shown that accounting for intraspecific trait variation (ITV) may better address major questions in community ecology. However, a general picture of the relative extent of ITV compared to interspecific trait variation in plant communities is still missing. Here, we conducted a meta-analysis of the relative extent of ITV within and among plant communities worldwide, using a data set encompassing 629 communities (plots) and 36 functional traits. Overall, ITV accounted for 25% of the total trait variation within communities and 32% of the total trait variation among communities on average. The relative extent of ITV tended to be greater for whole-plant (e.g. plant height) vs. organ-level traits and for leaf chemical (e.g. leaf N and P concentration) vs. leaf morphological (e.g. leaf area and thickness) traits. The relative amount of ITV decreased with increasing species richness and spatial extent, but did not vary with plant growth form or climate. These results highlight global patterns in the relative importance of ITV in plant communities, providing practical guidelines for when researchers should include ITV in trait-based community and ecosystem studies.

Community assembly along a soil depth gradient: contrasting patterns of plant trait convergence and divergence in a Mediterranean rangeland

Summary 1. Understanding how environmental factors drive plant community assembly remains a major challenge in community ecology. The strength of different assembly processes along environmental gradients, such as environmental filtering and functional niche differentiation, can be quantified by analysing trait distributions in communities. While environmental filtering affects species occurrence among communities, functional divergence or convergence is strongly related to species abundances within communities, which few studies have taken into account. We examine the trait-mediated effect of these two processes along a stress-resource gradient. 2. We measured species abundances and the distributions of eight traits related to vegetative and regenerative phases in plant communities along a gradient of soil depth and resource availability in Mediterranean rangelands. We quantified environmental filtering, defined as a local restriction of trait range, and trait divergence, based on abundance-weighted trait variance, using a two-step approach with specifically designed null models. 3. Communities presented a clear functional response to the soil gradient, as evidenced by strong trends in community-weighted trait means. We detected environmental filtering of different traits at both ends of the gradient, suggesting that, contrary to widespread expectations, trait filtering may not necessarily be the result of abiotic filtering under harsh conditions but could likely also result from biotic interactions in productive habitats. 4. We found marked shifts in trait abundance distributions within communities along the gradient. Vegetative traits (e.g. leaf dry matter content) diverged on shallow soils, reflecting the coexistence of distinct water- and nutrient-use strategies in these constrained habitats and converged with increasing soil resource availability. By contrast, regenerative traits (e.g. seed mass) tended to diverge towards deeper soils, while plant reproductive heights diverged all along the gradient. 5. Synthesis: Our study highlights how the combination of abundance data with traits capturing different functional niches is critical to the detection of complex functional responses of plant communities to environmental gradients. We demonstrate that patterns of trait divergence and filtering are strongly contingent on both trait and environment such that there can be no expectation of a simple trend of increasing or decreasing functional divergence along a gradient of resource availability.

Venation networks and the origin of the leaf economics spectrum

The leaf economics spectrum describes biome-invariant scaling functions for leaf functional traits that relate to global primary productivity and nutrient cycling. Here, we develop a comprehensive framework for the origin of this leaf economics spectrum based on venation-mediated economic strategies. We define a standardized set of traits - density, distance and loopiness - that provides a common language for the study of venation. We develop a novel quantitative model that uses these venation traits to model leaf-level physiology, and show that selection to optimize the venation network predicts the mean global trait-trait scaling relationships across 2548 species. Furthermore, using empirical venation data for 25 plant species, we test our model by predicting four key leaf functional traits related to leaf economics: net carbon assimilation rate, life span, leaf mass per area ratio and nitrogen content. Together, these results indicate that selection on venation geometry is a fundamental basis for understanding the diversity of leaf form and function, and the carbon balance of leaves. The model and associated predictions have broad implications for integrating venation network geometry with pattern and process in ecophysiology, ecology and palaeobotany.

Functional trait space and the latitudinal diversity gradient

Significance We present a conceptual framework for testing theories for the latitudinal gradient of species richness in terms of variation in functional diversity at the alpha, beta, and gamma scales. We compared ecological community theory with large-scale observational data of tree species richness and functional diversity. We found that the patterns of functional trait diversity are not consistent with any one theory of species diversity. These conflicting results indicate that none of the broad classes of biodiversity theory considered here is alone able to explain the latitudinal gradient of species diversity in terms of functional trait space. The processes causing the latitudinal gradient in species richness remain elusive. Ecological theories for the origin of biodiversity gradients, such as competitive exclusion, neutral dynamics, and environmental filtering, make predictions for how functional diversity should vary at the alpha (within local assemblages), beta (among assemblages), and gamma (regional pool) scales. We test these predictions by quantifying hypervolumes constructed from functional traits representing major axes of plant strategy variation (specific leaf area, plant height, and seed mass) in tree assemblages spanning the temperate and tropical New World. Alpha-scale trait volume decreases with absolute latitude and is often lower than sampling expectation, consistent with environmental filtering theory. Beta-scale overlap decays with geographic distance fastest in the temperate zone, again consistent with environmental filtering theory. In contrast, gamma-scale trait space shows a hump-shaped relationship with absolute latitude, consistent with no theory. Furthermore, the overall temperate trait hypervolume was larger than the overall tropical hypervolume, indicating that the temperate zone permits a wider range of trait combinations or that niche packing is stronger in the tropical zone. Although there are limitations in the data, our analyses suggest that multiple processes have shaped trait diversity in trees, reflecting no consistent support for any one theory.

Competition, traits and resource depletion in plant communities

Although of primary importance to explain plant community structure, general relationships between plant traits, resource depletion and competitive outcomes remain to be quantified across species. Here, we used a comparative approach to test whether instantaneous measurements of plant traits can capture both the amount of resources depleted under plant cover over time (competitive effect) and the way competitors perceived this resource depletion (competitive response). We performed a large competition experiment in which phytometers from a single grass species were transplanted within 18 different monocultures grown in a common-garden experiment, with a time-integrative quantification of light and water depletion over the phytometers’ growing season. Resource-capturing traits were measured on both phytometers (competitive response traits) and monocultures (competitive effect traits). The total amounts of depleted light and water availabilities over the season strongly differed among monocultures; they were best estimated by instantaneous measurements of height and rooting depth, respectively, performed when either light or water became limiting. Specific leaf area and leaf water potential, two competitive response traits measured at the leaf level, were good predictors of changes in phytometer performance under competition, and reflected the amount of light and water, respectively, perceived by plants throughout their lifespan. Our results demonstrated the relevance of instantaneous measures of plant traits as indicators of resource depletion over time, validating the trait-based approach for competition ecology.

Intraspecific trait variability mediates the response of subalpine grassland communities to extreme drought events

Climate change is expected to increase the magnitude and the frequency of extreme climatic events such as droughts. Better understanding how plant communities will respond to these droughts is a major challenge. We expect the response to be a shift in functional trait values resulting from both species turnover and intraspecific trait variability, but little research has addressed the relative contribution of both components. We analysed the short-term functional response of subalpine grassland communities to a simulated drought by focusing on four leaf traits (LDMC: leaf dry matter content, SLA: specific leaf area, LNC: leaf nitrogen concentration and LCC: leaf carbon concentration). After evaluating species turnover and intraspecific variability separately, we determined their relative contribution in the community functional response to drought, reflected by changes in community-weighted mean traits. We found significant species turnover and intraspecific variability, as well as significant changes in community-weighted mean for most of the traits. The relative contribution of intraspecific variability to the changes in community mean traits was more important (42-99%) than the relative contribution of species turnover (1-58%). Intraspecific variability either amplified (for LDMC, SLA and LCC) or dampened (for LNC) the community functional response mediated by species turnover. We demonstrated that the small contribution of species turnover to the changes in community mean LDMC and LCC was explained by a lack of covariation between species turnover and interspecific trait differences.Synthesis. These results highlight the need for a better consideration of intraspecific variability to understand and predict the effect of climate change on plant communities. While both species turnover and intraspecific variability can be expected following an extreme drought, we report new evidence that intraspecific variability can be a more important driver of the short-term functional response of plant communities.

Experimental demonstration of the importance of competition under disturbance

Ecologists have long recognized the roles of competition and disturbance in shaping ecological communities, and the combinatorial effects of these two factors have been the subject of substantial ecological research. Nevertheless, it is still unclear whether competition remains as an important structuring force in habitats strongly influenced by disturbance. The conventional belief remains that the importance of competition decreases with increasing disturbance, but limited theory suggests otherwise. Using protist communities established in laboratory microcosms, we demonstrate that disturbance does not diminish the importance of competition. Interspecific competition significantly increased rates of species extinction over a broad disturbance gradient, and increasing disturbance intensities increased, rather than decreased, the tempo of competitive exclusion. This community-level pattern is linked to the species-level pattern that interspecific competition led to most frequent extinctions of each species at the highest level of disturbance that the species can tolerate. Consequently, despite a strong tradeoff between competitive ability and disturbance tolerance across the competing species, species diversity generally declined with disturbance. The consistent structuring role of competition throughout the disturbance gradient underscores the need to understand competitive interactions and their consequences even in highly disturbed habitats.

A common genetic basis to the origin of the leaf economics spectrum and metabolic scaling allometry

Many facets of plant form and function are reflected in general cross-taxa scaling relationships. Metabolic scaling theory (MST) and the leaf economics spectrum (LES) have each proposed unifying frameworks and organisational principles to understand the origin of botanical diversity. Here, we test the evolutionary assumptions of MST and the LES using a cross of two genetic variants of Arabidopsis thaliana. We show that there is enough genetic variation to generate a large fraction of variation in the LES and MST scaling functions. The progeny sharing the parental, naturally occurring, allelic combinations at two pleiotropic genes exhibited the theorised optimum ¾ allometric scaling of growth rate and intermediate leaf economics. Our findings: (1) imply that a few pleiotropic genes underlie many plant functional traits and life histories; (2) unify MST and LES within a common genetic framework and (3) suggest that observed intermediate size and longevity in natural populations originate from stabilising selection to optimise physiological trade-offs.