The development of distinct regions in the amniote vertebral column results from somite formation and Hox gene expression, with the adult morphology displaying remarkable variation among lineages. Mammalian regionalization is reportedly very conservative or even constrained, but there has been no study investigating vertebral count variation across Amniota as a whole, undermining attempts to understand the phylogenetic, ecological, and developmental factors affecting vertebral column variation. Here, we show that the mammalian (synapsid) and reptilian lineages show early in their evolutionary histories clear divergences in axial developmental plasticity, in terms of both regionalization and meristic change, with basal synapsids sharing the conserved axial configuration of crown mammals, and basal reptiles demonstrating the plasticity of extant taxa. We conducted a comprehensive survey of presacral vertebral counts across 436 recent and extinct amniote taxa. Vertebral counts were mapped onto a generalized amniote phylogeny as well as individual ingroup trees, and ancestral states were reconstructed by using squared-change parsimony. We also calculated the relationship between presacral and cervical numbers to infer the relative influence of homeotic effects and meristic changes and found no correlation between somitogenesis and Hox-mediated regionalization. Although conservatism in presacral numbers characterized early synapsid lineages, in some cases reptiles and synapsids exhibit the same developmental innovations in response to similar selective pressures. Conversely, increases in body mass are not coupled with meristic or homeotic changes, but mostly occur in concert with postembryonic somatic growth. Our study highlights the importance of fossils in large-scale investigations of evolutionary developmental processes.omitogenesis, somatic growth, and Hox gene expression are primary factors driving the formation of the vertebrate body axis (1, 2). Somitogenesis occurs via the rhythmic budding of embryonic somites from the anterior part of the presomitic mesoderm, until a point when the latter has shrunk to such a degree that no further somites can be formed (3). Periodic somite formation is controlled by a molecular oscillator or "segmentation clock" (4), whereas the speed of the clock is highly variable across different vertebrate lineages. For example, the segmentation clock in snakes ticks much faster than in a mouse, resulting in a higher number of relatively smaller somites (3). Because the ossified vertebrae are derived from the embryonic somites through resegmentation (1), vertebral numbers can provide insights into the speed of the segmentation clock and the pattern of somitogenesis. In addition, the relative size of vertebrae relative to the number of vertebrae provides information about the rate of somatic growth (5, 6).Besides the marked variation in the number and sizes of vertebrae, the vertebrate body axis is also highly regionalized. In tetrapods, this regionalization is expressed in the formatio...