Pleistocene effects on North American songbird evolution

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Abstract
Recent studies have used comparisons of mitochondrial DNA (mtDNA) sequence divergence among populations and species to test existing hypotheses about avian evolution during the Pleistocene epoch. In 1998, Avise and Walker concluded that the Pleistocene was an important time for avian evolution, including the initiation of phylogeographic separations and the completion of speciation events that began in the Pliocene. The study implied that these conclusions conflicted with the study, in 1997, by Klicka and Zink, which concluded that most species pairs previously thought to have originated in the past 250 000 years were much older. The two studies are complementary in the sense that Avise and Walker dealt primarily with phylogeographic (intraspecific) separations. Furthermore, Klicka and Zink concentrated on the inception of divergences whereas the Avise and Walker focused on the timing of the completion of speciation. To accomplish this, Avise and Walker analysed ‘phylogroups’, geographically coherent subsets of biological species in which mtDNA haplotypes exhibit reciprocal monophyly. The study used the average interphylogroup mtDNA distance (0.027), calibrated at 2% per million years, to conclude that speciation required on average one million years to complete. Hence, speciation events begun in the Late Pliocene would have been completed in the mid– to late Pleistocene. Although we appreciate the extended nature of the speciation process and Avise and Walker's insightful attempt to estimate its duration, we conclude that their value was an overestimate by a factor of two. In particular we question whether phylogroups can be used in the novel evolutionary role that Avise and Walker envisioned, because of the vagaries of taxonomic practices and lack of consensus regarding species concepts. To extend their analysis of intraspecific, phylogeographic separations, we compiled previously analysed and newly available data for divergence times for North American songbird (order Passeriformes) phylogroups. More than 80% were initiated at least one million years ago, which is inconsistent with the late Pleistocene origins model previously rejected by Klicka and Zink mentioned above. Although some divergence events can be traced to the late Pleistocene, the significance of the distribution must be judged with reference to a null model. Whether the Pleistocene was a profound time for avian phylogeographic differentiation is at present unknown.

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