Intermediate predation pressure leads to maximal complexity in food webs

Responses of ecosystems to modifications of their environmental conditions are usually considered in terms of biodiversity or function. Maybe because they represent a hidden part of ecosystems, responses of ecological interactions are rarely studied. A more comprehensive view of the processes underlying the restructuring of food webs under environmental gradients appears crucial to understand how ecosystems functionalities are altered. We address this general issue in an experiment where trophic interactions are reorganized under a gradient of top-down effects (predation pressure) and bottom-up effects (nutrient availability). Unimodal relationships of species diversity are pervasive in ecology; we extend this principle to food-web topology: in our study, most topological descriptor values peak at intermediate predation intensity. The same unimodal pattern holds for network complexity, measured by entropy and scaled entropy (a measure independent of species diversity). Moreover, food web complexity is maximized at higher fish abundance when nutrient availability is increased. We infer that whereas in absence of top predators, a mechanism of competitive exclusion takes place, and indirect facilitation process underlies systems with a moderate predation pressure. These results highlight the importance of the dynamic reorganization of trophic links in response to bottom-up and top-down effects. Interaction between bottom-up and top-down forces argue for multifactorial studies of ecological effects.