Effects of Enrichment on Three-Level Food Chains with Omnivory

Although omnivory (the consumption of resources from more than one trophic level) is widespread, this fundamental limitation to the applicability of food chain theory to real communities has received only limited treatment. We investigated effects of enrichment (increasing carrying capacity, K, of the resource) on a system consisting of a resource (R), an intermediate consumer (N), and an omnivore (P) using a general mathematical model and tested the relevance of some of its predictions to a laboratory system of mixed bacteria (=R) and the ciliates Tetrahymena (=N) and Blepharisma (=P). The model produced six major predictions. First, N may facilitate or inhibit P. Enrichment may revert the net effect of N on P from facilitation to inhibition. Second, along a gradient of K, up to four regions of invasibility and stable coexistence of N and P may exist. At the lowest K, only R is present. At somewhat higher K, N can coexist with R. At intermediate K, either N and P coexist, or either consumer excludes the other depending on initial conditions. At the highest K, N may be excluded through apparent competition and only R and P can coexist. The pattern of persistence of Tetrahymena and Blepharisma along an enrichment gradient conformed fairly well to the scenario allowing coexistence at intermediate K. Third, for stable equilibria of the omnivory system, R always increases and N always decreases with K. The abundances of bacteria and Tetrahymena were suggestive of such a pattern but did not allow a strict test because coexistence occurred at only one level of enrichment. Fourth, an omnivore can invade an R-N system at a lower K than an otherwise identical specialist predator of N. Fifth, an omnivore can always invade a food chain with such a specialist predator. Sixth, over ranges of K where both omnivory systems and otherwise identical three-level food chains are feasible, N is always less abundant in the omnivory system, whereas the relative abundances of R and P in omnivory systems compared to food chains may change with K. It is thus possible that total community biomass at a given K is lower in an omnivory system than in a food chain. Both the model and the experimental results caution that patterns of trophic-level abundances in response to enrichment predicted by food chain theory are not to be expected in systems with significant omnivory.