Adaptive colonization across a parasitism-mutualism gradient.

Adaptive colonization is a process wherein a colonizing population exhibits an adaptive change in response to a novel environment, which may be critical to its establishment. To date, theoretical models of adaptive colonization have been based on single-species introductions. However, given their pervasiveness, symbionts will frequently be co-introduced with their hosts to novel areas. We present an individual-based model to investigate adaptive colonization by hosts and their symbionts across a parasite-mutualist continuum. The host must adapt in order to establish itself in the novel habitat, and the symbiont must adapt to track evolutionary change in the host. First, we classify the qualitative shifts in the outcome that can potentially be driven by non-neutral effects of the symbiont-host interaction into three main types: parasite-driven co-extinction, parasite release, and mutualistic facilitation. Second, we provide a detailed description of a specific example for each type of shift. Third, we disentangle how the interplay between symbiont transmissibility, host migration, and selection strength determines: (a) which type of shift is more likely to occur and (b) the size of the interaction effects necessary to produce it. Overall, we demonstrate the crucial role of host and symbiont dispersal scales in shaping the impacts of parasitism and mutualism on adaptive colonization.