Community lifespan, niche expansion and the evolution of interspecific cooperation.

Microbes live in dense and diverse communities where they deploy many traits that promote the growth and survival of neighbouring species, all the while also competing for shared resources. Because microbial communities are highly dynamic, the costs and benefits of species interactions change over the growth cycle of a community. How mutualistic interactions evolve under such demographic and ecological conditions is still poorly understood. Here, we develop an eco-evolutionary model to explore how different forms of helping with distinct fitness effects (rate-enhancing and yield-enhancing) affect the multiple phases of community growth, and its consequences for the evolution of mutualisms. We specifically focus on a form of yield-enhancing trait in which cooperation augments the common pool of resources, termed niche expansion. We show that although mutualisms in which cooperation increases partners growth rate are generally favoured at early stages of community growth, niche expansion can evolve at later stages where densities are high. Further, we find that niche expansion can promote the evolution of reproductive restraint, in which a focal species adaptively reduces its own growth rate to increase the density of partner species. Our findings suggest that yield-enhancing mutualisms are more prevalent in stable habitats with a constant supply of resources, and where populations typically live at high densities. In general, our findings highlight the need to integrate different components of population growth in the analysis of mutualisms to understand the composition and function of microbial communities.