The axial skeleton of Tiktaalik roseae.

The axial columns of the earliest limbed vertebrates show distinct patterns of regionalization as compared to early tetrapodomorphs. Included among their novel features are sacral ribs, which provide linkage between the vertebral column and pelvis, contributing to body support and propulsion by the hindlimb. Data on the axial skeletons of the closest relatives of limbed vertebrates are sparce, with key features of specimens potentially covered by matrix. Therefore, it is unclear in what sequence and under what functional context specializations in the axial skeletons of tetrapods arose. Here, we describe the axial skeleton of the elpistostegalian Tiktaalik roseae and show that transformations to the axial column for head mobility, body support, and pelvic fin buttressing evolved in finned vertebrates prior to the origin of limbs. No atlas-axis complex is observed; however, an independent basioccipital-exoccipital complex suggests increased mobility at the occipital vertebral junction. While the construction of vertebrae in Tiktaalik is similar to early tetrapodomorphs, its ribs possess a specialized sacral domain. Sacral ribs are expanded and ventrally curved, indicating likely attachment to the expanded iliac blade of the pelvis by ligamentous connection. Thus, the origin of novel rib types preceded major alterations to trunk vertebrae, and linkage between pelvic fins and axial column preceded the origin of limbs. These data reveal an unexpected combination of post-cranial skeletal characters, informing hypotheses of body posture and movement in the closest relatives of limbed vertebrates.

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.