Structure et dynamique des communautés multi-espèces : le rôle de l’espace

This thesis is a study of the role of space in the organization and dynamics of multi-species ecological communities. Two weaknesses can be identified from previous theoretical studies concerned with the spatial dimension of ecological communities: the scarcity of multi-species models based on a spatially explicit representation of space, and the lack of attention toward positive interspecific interactions, such as mutualism, despite the recognition of their ubiquity in ecological systems. This thesis explores this problematic by adopting a theoretical framework based on complex system theory and statistical mechanics. Following this approach, ecological communities can be viewed as complex systems whose global properties emerge from the local interactions between the organisms that composed them, and between the organisms and their environment.             The first objective of this thesis is to develop a multi-species metacommunity model which is spatially explicit, individual-based, and centered on a general interspecific interaction web containing exploitation, competition as well as mutualism. In this model, local communities are created by an assembly process whereby species are drawn from a regional pool. Population growth is restricted by a carrying capacity and its dynamics is driven by simple reproduction and dispersal mechanisms acting at the level of single individual. These mechanisms depend on the biotic and abiotic conditions of the local communities and their effect varies with species, time and space. The second objective of this thesis is to determine the impact of an increasing spatial connectivity on the dynamics, and structural and functional properties of this metacommunity. More precisely, we set out to evaluate different community properties under changes in the level of species dispersal: i) the similarity in local community composition and its patterns of spatial correlations, ii) the local and regional diversity and the local species abundance, iii) the local and regional biomass, productivity and dynamical stability, and iv) the structure of the local interaction webs. These properties are examined under two spatial schemes. First, we employ a homogeneous environment, and second we employ a heterogeneous environment whereby the carrying capacity of local communities evolves along a gradient.             In general, our results reveal that spatially distributed ecological communities are extremely sensitive to the modes and levels of species dispersal. Their spatiotemporal dynamics as well as their structural and functional properties can undergo profound changes in the form of significant transitions under slight changes of the level of dispersal. These changes are also highlighted by the emergence of spatiotemporal patterns in the spatial distribution of the populations, which are characteristics of phase transition generally observed in physical systems.             The metacommunity presents two dynamical regimes. In the first regime, corresponding to weak levels of species dispersal, the assembly dynamics promotes the emergence of species-poor but stable communities made of abundant and strongly mutualistic species. The metacommunity has a high regional diversity since weakly connected communities conserve a distinct assemblage of species. On the other hand, in the second regime, corresponding to strong dispersal rates, regional diversity decreases at the benefit of an increase in local diversity. Local communities are more productive but their stability is reduced due to the important migration of individuals. This regime is also characterized by assemblages containing a richer diversity of interspecific interactions. These results suggest that an augmentation in the level of species dispersal permits organisms to couple local communities together which increases local coexistence and promotes the organization of richer and more complex ecological communities.             Finally, our results suggest that mutualism is fundamental to the organization and persistence of ecological communities. Mutualistic species dominate in habitats characterized by a restricted carrying capacity and serve as ecological engineer by facilitating the establishment of competitors, predators and opportunists which benefit from their presence.

Keyphrases: Metacommunity, Spatially explicit model, Dispersal, Mutualism, Biodiversity, Complex systems, Phase transitions