© 2016 Elsevier Ltd. When a more complex, functionally integrated entity emerges from the association of simpler, initially independent entities, a major evolutionary transition has occurred. Transitions that result from the association of different species include the evolution of the eukaryotic cell and some obligate mutualisms. Recent studies are revolutionizing our understanding of how these intimate interspecific associations come to be, revealing how and to what extent each partner contributes to the relationship, and how partners mediate conflict. Here, we review work on the evolution of mutualistic symbioses in the context of transitions in individuality and highlight how a better mechanistic understanding of the ecological drivers of host-symbiont interdependencies can help elucidate the evolutionary path to symbiotic organismality.