Genetics and ecological speciation.

Species originate frequently by natural selection. A general mechanism by which this occurs is ecological speciation, defined as the evolution of reproductive isolation between populations as a result of ecologically-based divergent natural selection. The alternative mechanism is mutation-order speciation in which populations fix different mutations as they adapt to similar selection pressures. Although numerous cases now indicate the importance of ecological speciation in nature, very little is known about the genetics of the process. Here, we summarize the genetics of premating and postzygotic isolation and the role of standing genetic variation in ecological speciation. We discuss the role of selection from standing genetic variation in threespine stickleback (Gasterosteus aculeatus), a complex of species whose ancestral marine form repeatedly colonized and adapted to freshwater environments. We propose that ecological speciation has occurred multiple times in parallel in this group via a "transporter" process in which selection in freshwater environments repeatedly acts on standing genetic variation that is maintained in marine populations by export of freshwater-adapted alleles from elsewhere in the range. Selection from standing genetic variation is likely to play a large role in ecological speciation, which may partly account for its rapidity.