Understanding the dependence of species interaction strengths on environmental factors and species diversity is crucial to predict community dynamics and persistence in a rapidly changing world. Nontrophic (e.g. predator interference) and trophic components together determine species interaction strengths, but the effects of environmental factors on these two components remain largely unknown. This impedes our ability to fully understand the links between environmental drivers and species interactions. Here, we used a dynamical modelling framework based on measured predator functional responses to investigate the effects of predator diversity, prey density, and temperature on trophic and nontrophic interaction strengths within a freshwater food web. We found that (i) species interaction strengths cannot be predicted from trophic interactions alone, (ii) nontrophic interaction strengths vary strongly among predator assemblages, (iii) temperature has opposite effects on trophic and nontrophic interaction strengths, and (iv) trophic interaction strengths decrease with prey density, whereas the dependence of nontrophic interaction strengths on prey density is concave up. Interestingly, the qualitative impacts of temperature and prey density on the strengths of trophic and nontrophic interactions were independent of predator identity, suggesting a general pattern. Our results indicate that taking multiple environmental factors and the nonlinearity of density-dependent species interactions into account is an important step towards a better understanding of the effects of environmental variations on complex ecological communities. The functional response approach used in this study opens new avenues for (i) the quantification of the relative importance of the trophic and nontrophic components in species interactions and (ii) a better understanding how environmental factors affect these interactions and the dynamics of ecological communities.

Department of Ecosystem Biology Faculty of Science University of South Bohemia 370 05 České Budějovice Czech Republic

Department of Ecosystem Biology Faculty of Science University of South Bohemia 370 05 České Budějovice Czech Republic Laboratory of Aquatic Insects and Relict Ecosystems Biology Centre CAS Institute of Entomology 370 05 České Budějovice Czech Republic

Department of Ecosystem Biology Faculty of Science University of South Bohemia 370 05 České Budějovice Czech Republic Laboratory of Aquatic Insects and Relict Ecosystems Biology Centre CAS Institute of Entomology 370 05 České Budějovice Czech Republic Unité Mixte de Recherche 5174 'Evolution et Diversité Biologique' Centre National de la Recherche Scientifique Université de Toulouse Institut de Recherche pour le Développement École Nationale de Formation Agronomique BP 22687 31326 Castanet Tolosan France

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