Abstract
BACKGROUND: Skulls serve many functions and as a result, are subject to many different evolutionary pressures. In squamates, many fossorial species occupy a unique region of skull morphospace, showing convergence across families, due to modifications related to head-first burrowing. As different substrates have variable physical properties, particular skull shapes may offer selective advantages in certain substrates. Despite this, studies of variation within burrowers have been limited and are typically focused on a single origin of fossoriality. We focused on seven skink genera (Acontias, Typhlosaurus, Scelotes, Sepsina, Feylinia, Typhlacontias, and Mochlus; 39 sp.) from southern Africa, encompassing at least three independent evolutions of semi-fossoriality/fossoriality. We used microCT scans and geometric morphometrics to test how cranial and mandibular shape were influenced by phylogenetic history, size, and ecology. We also qualitatively described the skulls of four species to look at variation across phylogenetic and functional levels, and assess the degree of convergence. RESULTS: We found a strong effect of phylogenetic history on cranial and mandibular shape, with size and substrate playing secondary roles. There was a clear gradient in morphospace from less specialized to more specialized burrowers and burrowers in sand were significantly different from those in other substrates. We also created an anatomical atlas for four species with each element described in isolation. Every bone showed some variation in shape and relative scaling of features, with the skull roofing bones, septomaxilla, vomer, and palatine showing the most variation. We showed how broad-scale convergence in traits related to fossoriality can be the result of different anatomical changes. CONCLUSIONS: Our study used geometric morphometrics and comparative anatomy to examine how skull morphology changes for a highly specialized and demanding lifestyle. Although there was broad convergence in both shape and qualitative traits, phylogenetic history played a large role and much of this convergence was produced by different anatomical changes, implying different developmental pathways or lineage-specific constraints. Even within a single family, adaptation for a specialized ecology does not follow a singular deterministic path.