The multiple meanings of omnivory influence empirical, modular theory and whole food web stability relationships

The persistence of whole communities hinges on the presence of select interactions which act to stabilize communities making the identification of these keystone interactions critical. One potential candidate is omnivory, yet theoretical research on omnivory thus far has been dominated by a modular theory approach whereby an omnivore and consumer compete for a shared resource. Empirical research, however, has highlighted the presence of a broader suite of omnivory modules. Here, we integrate empirical data analysis and mathematical models to explore the influence of both omnivory module (including classic, multi-resource, higher level, mutual predation and cannibalism) and omnivore-resource interaction type on food web stability. We use six classic empirical food webs to examine the prevalence of the different types of omnivory, a multi-species consumer-resource model to determine the stability of these different kinds of omnivory within a module context, and finally extend these models to a 50 species, whole food web model to examine the influence of omnivory on whole food web persistence. Our results challenge the concept that omnivory is broadly stabilizing. In particular, we demonstrate that the impact of omnivory depends on the type of omnivory being examined with multi-resource omnivory having the largest correlation with whole food web persistence. Moreover, our results highlight that we need to exercise caution when scaling modular theory to whole food web theory. Cannibalism, for example, was the most persistent and stable omnivory module in the modular theory analysis, but only demonstrated a weak correlation with whole food web persistence. Lastly, our results demonstrate that the frequency of omnivory modules are more important for whole food web persistence than the frequency of omnivore-resource interactions. Together, these results demonstrate that the role of omnivory often depends both on the type of omnivory being examined and the food web within which it is nested. In whole food web models, omnivory acts less as a keystone interaction, rather, specific types of omnivory, particularly multi-resource omnivory, act as keystone modules. Future work integrating module and whole food web theory is critical for resolving the role of key interactions in food webs.