title: Skeletal heterochrony is associated with the anatomical specializations of snakes among squamate reptiles.
abstract: Snakes possess a derived anatomy, characterized by limb reduction and reorganization of the skull and internal organs. To understand the origin of snakes from an ontogenetic point of view, we conducted comprehensive investigations on the timing of skeletal elements, based on published and new data, and reconstructed the evolution of the ossification sequence among squamates. We included for the first time Varanus, a critical taxon in phylogenetic context. There is comprehensive delay in the onset of ossification of most skeletal elements in snakes when compared to reference developmental events through evolution. We hypothesize that progressing deceleration accompanied limb reduction and reorganization of the snake skull. Molecular and morphological studies have suggested close relationship of snakes to either amphisbaenians, scincids, geckos, iguanids, or varanids. Likewise, alternative hypotheses on habitat for stem snakes have been postulated. Our comprehensive heterochrony analyses detected developmental shifts in ossification for each hypothesis of snake origin. Moreover, we show that reconstruction of ancestral developmental sequences is a valuable tool to understand ontogenetic mechanisms associated with major evolutionary changes and test homology hypotheses. The "supratemporal" of snakes could be homolog to squamosal of other squamates, which starts ossification early to become relatively large in snakes.
authors.affiliation: Paläontologisches Institut und Museum der Universität Zürich, Karl-Schmid-Strasse 4, 8006, Zürich, Switzerland; Museum für Naturkunde, Leibniz-Institut für Evolutions & Biodiversitätsforschung, an der Humboldt-Universität zu Berlin, Invalidenstraße 43, 10115, Berlin, Germany. i.werneburg@gmail.com.
authors.forename: Ingmar
authors.initials: I
authors.lastname: Werneburg
authors.forename: Marcelo R
authors.initials: MR
authors.lastname: Sánchez-Villagra
mesh_terms: D000818:Animals; D001842:Bone and Bones / Q000033:anatomy & histology*; D019143:Evolution, Molecular; D005580:Fossils / Q000033:anatomy & histology; D012911:Snakes / Q000033:anatomy & histology / Q000235:genetics*
keywords: Embryos; Varanus; fossils; serpentes; skeletogenesis; square change parsimony