Speciation in seabirds: why are there so many species…and why aren’t there more?

Abstract

Speciation—the multiplication of species through the evolution of barriers to reproduction between populations—plays a central role in evolution since it enables two or more populations to adapt and evolve independently. However, mechanisms of speciation are notoriously difficult to study and poorly understood. Seabirds provide useful models to investigate factors that may promote or inhibit speciation because their ecology and evolutionary genetics are relatively well understood. Here I review population genetic studies of seabirds to test the importance of six factors with the potential to disrupt gene flow enough to result in speciation. Over 200 studies, including over 100 species, have been published to date. Most show evidence of restrictions in gene flow. Physical (geographic) barriers to dispersal are clearly important: conspecific populations that are separated by large expanses of land or ice show evidence of restricted gene flow, and sister species often are separated by physical barriers to gene flow. However, many species of seabirds show evidence of restrictions in gene flow in the absence of physical barriers to dispersal. Study results indicate that differences in ocean regimes, nonbreeding distributions, foraging distributions during the breeding season, and breeding phenology also can disrupt gene flow enough to lead to speciation. Of these, physical isolation and differences in ocean regime appear to be the most important. Philopatry alone may be sufficient to result in reproductive isolation, but usually it acts in combination with other barriers to gene flow. The effects of many other potential influences on gene flow need to be investigated more thoroughly, including colony distribution/location, wind, interspecific interactions, environmental stability/variability, variation in phenotypic traits associated with mate choice (morphology, behaviour, vocalisations) and intrinsic (genomic) incompatabilities. Recent advances in genome sequencing, especially if used in combination with ecological tools such as geolocators and new methods for data interpretation, are opening exiting new avenues to test the importance of various behavioural, ecological, demographic and genomic factors in reducing or promoting gene flow and so affecting speciation.