Conservation of Rib Skeleton Regionalization in the Homoplastic Evolution of the Snake-like Body Form in Squamates

Squamates have independently evolved an elongate, limb-reduced body form numerous times. This transition has been proposed to involve either changes to regulatory gene expression or downstream modification of target enhancers to produce a homogeneous, deregionalized axial skeleton. Analysis of vertebral morphology has suggested that regionalization is maintained in snake-like body forms, but morphological variation in the other primary component of the axial skeleton, the dorsal ribs, has not been previously examined. We quantified rib morphology along the anterior-posterior axis in limbed and snake-like squamates to test different regionalization models. We find that the relative position of regional boundaries remains consistent across taxa of differing body types, including in the homoplastic evolution of snake-like body forms. The consistent retention of regional boundaries in this primaxial domain is uncorrelated with more plastic abaxial region markers. Rather than loss of regions, rib shape at the anterior and posterior of the axis converges on those in the middle, resulting in axial regions being distinguishable by allometric shape changes rather than by discrete morphologies. This complexity challenges notions of deregionalization, revealing a nuanced evolutionary history shaped by shared functions.