The existence and strength of higher order interactions is sensitive to environmental context.

Abstract

One strategy for understanding the dynamics of any complex system, such as a community of competing species, is to study the dynamics of parts of the system in isolation. Ecological communities can be decomposed into single species, and pairs of interacting species. This reductionist strategy assumes that whole-community dynamics are predictable and explainable from knowledge of the dynamics of single species and pairs of species. This assumption will be violated if higher order interactions (HOIs) are strong. Theory predicts that HOIs should be common. But it is difficult to detect HOIs, and to infer their long-term consequences for species coexistence, solely from short-term data. I conducted a protist microcosm experiment to test for HOIs among competing bacterivorous ciliates, and test the sensitivity of HOIs to environmental context. I grew three competing ciliate species in all possible combinations at each of two resource enrichment levels, and used the population dynamic data from the one- and two-species treatments to parameterize a competition model at each enrichment level. I then compared the predictions of the parameterized model to the dynamics of the whole community (three-species treatment). I found that the existence, and thus strength, of HOIs was environment dependent. I found a strong HOI at low enrichment, which enabled the persistence of a species that would otherwise have been competitively excluded. At high enrichment, three-species dynamics could be predicted from a parameterized model of one- and two-species dynamics, provided that the model accounted for nonlinear intraspecific density dependence. The results provide one of the first rigorous demonstrations of the long-term consequences of HOIs for species coexistence, and demonstrate the context dependence of HOIs. HOIs create difficult challenges for predicting and explaining species coexistence in nature.