Predicting coexistence in species with continuous ontogenetic niche shifts and competitive asymmetry.

A longstanding problem in ecology is whether structured life cycles impede or facilitate coexistence between species. Theory based on populations with only two discrete stages in the life-cycle indicates that for two species to coexist, at least one must shift its niche between stages and each species must be a better competitor in one of the niches. However, in many cases, niche shifts are associated with changes in an underlying continuous trait like body size and we have few predictions concerning conditions for coexistence for such a widespread form of ontogenetic development. We develop a framework for analyzing species coexistence based on Integral Projection Models (IPMs) that incorporates continuous ontogenetic changes in both the resource niche and competitive ability. We parameterize the model using experimental data from Trinidadian guppies and show how niche shifts and competitive symmetries impact species coexistence. Overall, our results show that the effects of competition on fitness depend upon trait-mediated niche-separation, trait-mediated competitive asymmetry in the part of the niche that is shared across body sizes, and the sensitivity of fitness to body size. Interactions among these processes generate multiple routes to coexistence. We discuss how our modeling framework expands results from two-stage models to mutli-stage or continuous stage models and allows for deriving predictions that can be tested in populations displaying continuous changes in niche use and competitive ability.