Divergence and adaptation in Beringian birds

Abstract

Beringia is a high-latitude hotspot of avian divergence and speciation. The unique biogeography of Beringia impacted avian speciation in two ways: through the cyclic appearance of the land bridge between the Asian and North American continents and through glacial refugia. These cyclic processes repeatedly split and connected avian populations, alternately reducing and increasing opportunities for gene flow between populations. In this thesis, I examine how this dynamic system impacted Beringian avian taxa using population genomic analyses. First, I examine broad patterns of divergence and gene flow across 11 lineages of birds using ultraconserved elements (UCEs), which are a multi-locus subsampling of the nuclear genome. These bird lineages contain two or more sister taxa at the population, subspecies, or species level that were likely impacted by the Bering land bridge and/or by glacial refugia. I tested models that provided key demographic information, such as population size, gene flow, and divergence time estimates. Demographic modeling showed gene flow in all cases at a wide range of rates between pairwise comparisons, and all inferred models included a divergence event during the Quaternary. Next, I focus on one species, the Song Sparrow (Melospiza melodia), in the Beringian part of its range. Five subspecies of the Song Sparrow reside in southern Alaska, from the Aleutian Islands to southeast Alaska, and have a wide range of body sizes. Using whole- genomic sequencing and morphology, I examine the phenotypic and genomic differences in these subspecies. I quantified the morphological differences, showing that the western subspecies are significantly larger than the eastern subspecies. I then determined that two candidate genes are under positive selection in the most isolated subspecies, M. m. maxima. Finally, I reconstructed a phylogeny and found that M. m. maxima is sister to the other M. melodia subspecies. These results highlight how the unique biogeography of Beringia impacted the generation of avian diversity in the region.