Dietary Specialization and Habitat Shifts in a Clade of Afro-Asian Colubrid Snakes (Colubridae: Colubrinae)

Speciation through niche divergence often occurs as lineages of organisms colonize and adapt to new environments with novel ecological opportunities that facilitate the evolution of ecologically different phenotypes. In snakes, adaptive diversification may be driven by the evolution of traits relating to changes in their diets. Accordingly, habitatmediated differences in prey available to ancestral snakes as they colonized and occupied novel dynamic landscapes are likely to have been a strong selective agent behind the divergence and radiation of snakes across the globe. Using an ancestral reconstruction approach that considers the multivariate nature of ecological phenotypes while accounting for sampling variation between taxa, we explored how diet and macro-habitat use coevolved across a phylogeny of 67 species of Afro-Asian colubrine snakes. Our results show that the most recent common ancestor of this clade was likely a dietary generalist that occupied tropical forests in Asia. Deviations from this generalist diet to a variety of specialist diets each dominated by the utilization of single prey types repeatedly occurred as ancestral colubrines shifted from tropical forests to savanna and grassland habitats across Africa. We additionally found that dietary specialist species were on average smaller in maximum length than dietary generalists, congruent with established predator-size, prey-diversity dynamics in snakes. We speculate that adaptive divergence in ancestral colubrines arose as a result of a selective regime that favored diets comprised of terrestrial prey, and that partitioning of different prey types led to the various forms of dietary specialization evident in these lineages today. Our findings provide new insights into the ecological correlates associated with the evolution of diet in snakes, thereby furthering our understanding of the driving forces behind patterns of snake diversification.