| Species exist in complex biotic environments, engaging in a variety of antagonistic and cooperative interactions. While these interactions are generally recognized to be context-dependent, varying in outcome in the presence of other interactions, studies tend to focus on each interaction in isolation. One of the main classes of species interaction is mutualism, in which partner species gain a net benefit from their interaction. However, mutualisms are beset by a variety of species that can reduce or even eliminate the benefits of mutualism through exploitation of and competition for the resources and services offered by mutualists. These exploiter species potentially threaten the ecological stability of mutualisms and may alter selection on mutualistic traits. Thus, understanding the ecology and evolution of mutualisms requires consideration of interactions with exploiter species. In this dissertation, I investigated the effects of exploiter species on mutualisms between plants and pollinators using a combination of eco-evolutionary modeling, optimization theory, and behavioral studies. Using two adaptive dynamics models of coevolution in exploited pollinating seed parasite mutualisms, I found that exploiters reduce mutualist densities and select for more parasitic mutualists. Nevertheless, the models demonstrate that intraspecific competition for host resources and host defense of those resources restrict the ecological conditions that lead to extinction of the mutualism, as well as the chances of evolution to extinction. Thus, exploiters are unlikely to be the threat to mutualisms that has been assumed previously. On the other hand, in another type of exploitation, exploitative predators may pose a greater threat to investment in mutualism than has been presumed. Through both optimal foraging theory and behavioral experiments on bumble bees, I found that the risk from ambush predators can change pollinator floral preferences when predators preferentially use high-quality flowers to locate their prey. This research suggests that predators of mutualists may have important top-down effects and that further research is needed to investigate the effects of exploitative predators on selection on mutualist traits. |
