Dominique Gravel

Ecophylogenetics: advances and perspectives

Ecophylogenetics can be viewed as an emerging fusion of ecology, biogeography and macroevolution. This new and fast‐growing field is promoting the incorporation of evolution and historical contingencies into the ecological research agenda through the widespread use of phylogenetic data. Including phylogeny into ecological thinking represents an opportunity for biologists from different fields to collaborate and has provided promising avenues of research in both theoretical and empirical ecology, towards a better understanding of the assembly of communities, the functioning of ecosystems and their responses to environmental changes. The time is ripe to assess critically the extent to which the integration of phylogeny into these different fields of ecology has delivered on its promise. Here we review how phylogenetic information has been used to identify better the key components of species interactions with their biotic and abiotic environments, to determine the relationships between diversity and ecosystem functioning and ultimately to establish good management practices to protect overall biodiversity in the face of global change. We evaluate the relevance of information provided by phylogenies to ecologists, highlighting current potential weaknesses and needs for future developments. We suggest that despite the strong progress that has been made, a consistent unified framework is still missing to link local ecological dynamics to macroevolution. This is a necessary step in order to interpret observed phylogenetic patterns in a wider ecological context. Beyond the fundamental question of how evolutionary history contributes to shape communities, ecophylogenetics will help ecology to become a better integrative and predictive science.

Accounting for dispersal and biotic interactions to disentangle the drivers of species distributions and their abundances

Although abiotic factors, together with dispersal and biotic interactions, are often suggested to explain the distribution of species and their abundances, species distribution models usually focus on abiotic factors only. We propose an integrative framework linking ecological theory, empirical data and statistical models to understand the distribution of species and their abundances together with the underlying community assembly dynamics. We illustrate our approach with 21 plant species in the French Alps. We show that a spatially nested modelling framework significantly improves the models performance and that the spatial variations of species presence-absence and abundances are predominantly explained by different factors. We also show that incorporating abiotic, dispersal and biotic factors into the same model bring new insights to our understanding of community assembly. This approach, at the crossroads between community ecology and biogeography, is a promising avenue for a better understanding of species co-existence and biodiversity distribution.

A road map for integrating eco-evolutionary processes into biodiversity models

The demand for projections of the future distribution of biodiversity has triggered an upsurge in modelling at the crossroads between ecology and evolution. Despite the enthusiasm around these so-called biodiversity models, most approaches are still criticised for not integrating key processes known to shape species ranges and community structure. Developing an integrative modelling framework for biodiversity distribution promises to improve the reliability of predictions and to give a better understanding of the eco-evolutionary dynamics of species and communities under changing environments. In this article, we briefly review some eco-evolutionary processes and interplays among them, which are essential to provide reliable projections of species distributions and community structure. We identify gaps in theory, quantitative knowledge and data availability hampering the development of an integrated modelling framework. We argue that model development relying on a strong theoretical foundation is essential to inspire new models, manage complexity and maintain tractability. We support our argument with an example of a novel integrated model for species distribution modelling, derived from metapopulation theory, which accounts for abiotic constraints, dispersal, biotic interactions and evolution under changing environmental conditions. We hope such a perspective will motivate exciting and novel research, and challenge others to improve on our proposed approach.

Experimental niche evolution alters the strength of the diversity-productivity relationship

The relationship between biodiversity and ecosystem functioning (BEF) has become a cornerstone of community and ecosystem ecology and an essential criterion for making decisions in conservation biology and policy planning. It has recently been proposed that evolutionary history should influence the BEF relationship because it determines species traits and, thus, species’ ability to exploit resources. Here we test this hypothesis by combining experimental evolution with a BEF experiment. We isolated 20 bacterial strains from a marine environment and evolved each to be generalists or specialists. We then tested the effect of evolutionary history on the strength of the BEF relationship with assemblages of 1 to 20 species constructed from the specialists, generalists and ancestors. Assemblages of generalists were more productive on average because of their superior ability to exploit the environmental heterogeneity. The slope of the BEF relationship was, however, stronger for the specialist assemblages because of enhanced niche complementarity. These results show how the BEF relationship depends critically on the legacy of past evolutionary events.

Trophic theory of island biogeography

MacArthur and Wilsons Theory of Island Biogeography (TIB) is among the most well-known process-based explanations for the distribution of species richness. It helps understand the species-area relationship, a fundamental pattern in ecology and an essential tool for conservation. The classic TIB does not, however, account for the complex structure of ecological systems. We extend the TIB to take into account trophic interactions and derive a species-specific model for occurrence probability. We find that the properties of the regional food web influence the species-area relationship, and that, in return, immigration and extinction dynamics affect local food web properties. We compare the accuracy of the classic TIB to our trophic TIB to predict community composition of real food webs and find strong support for our trophic extension of the TIB. Our approach provides a parsimonious explanation to species distributions and open new perspectives to integrate the complexity of ecological interactions into simple species distribution models.

The dissimilarity of species interaction networks

In a context of global changes, and amidst the perpetual modification of community structure undergone by most natural ecosystems, it is more important than ever to understand how species interactions vary through space and time. The integration of biogeography and network theory will yield important results and further our understanding of species interactions. It has, however, been hampered so far by the difficulty to quantify variation among interaction networks. Here, we propose a general framework to study the dissimilarity of species interaction networks over time, space or environments, allowing both the use of quantitative and qualitative data. We decompose network dissimilarity into interactions and species turnover components, so that it is immediately comparable to common measures of β-diversity. We emphasise that scaling up β-diversity of community composition to the β-diversity of interactions requires only a small methodological step, which we foresee will help empiricists adopt this method. We illustrate the framework with a large dataset of hosts and parasites interactions and highlight other possible usages. We discuss a research agenda towards a biogeographical theory of species interactions.

The influence of interspecific interactions on species range expansion rates

Ongoing and predicted global change makes understanding and predicting species’ range shifts an urgent scientific priority. Here, we provide a synthetic perspective on the so far poorly understood effects of interspecific interactions on range expansion rates. We present theoretical foundations for how interspecific interactions may modulate range expansion rates, consider examples from empirical studies of biological invasions and natural range expansions as well as process-based simulations, and discuss how interspecific interactions can be more broadly represented in process-based, spatiotemporally explicit range forecasts. Theory tells us that interspecific interactions affect expansion rates via alteration of local population growth rates and spatial displacement rates, but also via effects on other demographic parameters. The best empirical evidence for interspecific effects on expansion rates comes from studies of biological invasions. Notably, invasion studies indicate that competitive dominance and release from specialized enemies can enhance expansion rates. Studies of natural range expansions especially point to the potential for competition from resident species to reduce expansion rates. Overall, it is clear that interspecific interactions may have important consequences for range dynamics, but also that their effects have received too little attention to robustly generalize on their importance. We then discuss how interspecific interactions effects can be more widely incorporated in dynamic modeling of range expansions. Importantly, models must describe spatiotemporal variation in both local population dynamics and dispersal. Finally, we derive the following guidelines for when it is particularly important to explicitly represent interspecific interactions in dynamic range expansion forecasts: if most interacting species show correlated spatial or temporal trends in their effects on the target species, if the number of interacting species is low, and if the abundance of one or more strongly interacting species is not closely linked to the abundance of the target species.

Inferring biotic interactions from proxies

Inferring biotic interactions from functional, phylogenetic and geographical proxies remains one great challenge in ecology. We propose a conceptual framework to infer the backbone of biotic interaction networks within regional species pools. First, interacting groups are identified to order links and remove forbidden interactions between species. Second, additional links are removed by examination of the geographical context in which species co-occur. Third, hypotheses are proposed to establish interaction probabilities between species. We illustrate the framework using published food-webs in terrestrial and marine systems. We conclude that preliminary descriptions of the web of life can be made by careful integration of data with theory.

Inferring food web structure from predator–prey body size relationships

1. Current global changes make it important to be able to predict which interactions will occur in the emerging ecosystems. Most of the current methods to infer the existence of interactions between two species require a good knowledge of their behaviour or a direct observation of interactions. In this paper, we overcome these limitations by developing a method, inspired from the niche model of food web structure, using the statistical relationship between predator and prey body size to infer the matrix of potential interactions among a pool of species. 2. The novelty of our approach is to infer, for any species of a given species pool, the three species-specific parameters of the niche model. The method applies to both local and metaweb scales. It allows one to evaluate the feeding interactions of a new species entering the community. 3. We find that this method gives robust predictions of the structure of food webs and that its efficiency is increased when the strength of the body-size relationship between predators and preys increases. 4. We finally illustrate the potential of the method to infer the metaweb structure of pelagic fishes of the Mediterranean sea under different global change scenarios.

Source and sink dynamics in meta‐ecosystems

We present a theory extending the source-sink concept with an ecosystem perspective. We analyze a model for meta-ecosystem dynamics in a heterogeneous environment to study how the spatial flows of materials such as inorganic nutrients and nutrients sequestered into producers, herbivores, and detritus affect the community dynamics. We show that spatial flows of an inorganic nutrient (direct nutrient flow) and organic matter (indirect nutrient flow) through detritus, producer, or herbivore compartments can reverse the source-sink dynamics of a local ecosystem. More precisely, the balance between such direct and indirect nutrient flows determines the net direction of nutrient flows between two ecosystems of contrasted productivities. It allows a sink to turn into a source and vice versa. This effect of nutrient flows on source and sink dynamics is robust to the ecosystem structure (with and without herbivores) and to specific ecosystem compartments contributing to nutrient flows (primary producers, herbivores, or detritus). Ecosystems in distinct localities thus interact together with the structure at one place influencing that of the other. In meta-ecosystems, the source-sink dynamics of an organism is not only constrained by its dispersal from the source to the sink, but also by the fertility and community composition in the neighborhood responsible for spatial flows of nutrients and energy. The meta-ecosystem perspective provides a powerful theoretical framework to address novel questions in spatial ecosystem ecology.