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With the increase in the number of introduced species each year, biological invasions are considered as one of the most important environmental problems for native biodiversity. In invaded habitats, the establishment of exotic plant species depends on the abiotic and biotic environment. Herbivores and neighboring plants (native or exotic) comprise an important part of the latter. Herbivores cause trophic and non-trophic damage to focal plants, which respond to herbivory by varying their different traits quantitatively (e.g., growth rate and biomass changes) and qualitatively (e.g., variation in morphological and chemical defenses strategies affecting plant palatability). Neighboring plant species also affect functional traits and the fitness of focal plant species, thus herbivore effects on a focal plant could also depend indirectly on the palatability and defensive traits of the neighboring species inside the community. Here, in a first step toward the integration of associational susceptibility/resistance theories in the field of ecological invasion, we performed a microcosm experiment to consider the effects of an exotic crayfish on the growth rate, morphological traits and damage level of three macrophytes (two exotic, one native) growing in pairwise combinations. We found that (i) the response to herbivore presence and to neighboring species identity seemed to be species specific, and (ii) crayfish enhance the fragmentation rate of the two exotic macrophytes  Ludwigia grandiflora  and  Egeria densa  in the presence of the native macrophyte  Myriophyllum spicatum , which could indirectly facilitate their invasion success. Indeed, fragmentation can increase dispersal abilities of the exotic macrophytes considered in this study as they are able to generate new plants from their fragments. However, our results showed that the interaction herbivore-neighbor species was hardly significant. Our paper presents some first results on associational resistance/susceptibility and lays the foundation for developing a general framework that combines plant community ecology and biological invasion ecology to explain invasive species success.

frontiers in plant science

Response of Macrophyte Traits to Herbivory and Neighboring Species: Integration of the Functional Trait Framework in the Context of Ecological Invasions