Reinforcement can overcome gene flow during speciation in Drosophila.

Reinforcement, the strengthening of prezygotic reproductive isolation by natural selection in response to maladaptive hybridization [1-3], is one of the few processes in which natural selection directly favors the evolution of species as discrete groups (e.g., [4-7]). The evolution of reproductive barriers via reinforcement is expected to evolve in regions where the ranges of two species overlap and hybridize as an evolutionary solution to avoiding the costs of maladaptive hybridization [2,3,8]. The role of reinforcement in speciation has, however, been highly controversial because population-genetic theory suggests that the process is severely impeded by both hybridization [8-11] and migration of individuals from outside the contact zone [12,13]. To determine whether reinforcement could strengthen the reproductive barriers between two sister species of Drosophila in the face of these impediments, I initiated experimental populations of these two species that allowed different degrees of hybridization, as well as migration from outside populations. Surprisingly, even in the face of gene flow, reinforcement could promote the evolution of reproductive isolation within only five generations. As theory predicts, high levels of hybridization (and/or strong selection against hybrids) and migration impeded this evolution. These results suggest that reinforcement can help complete the process of speciation.