Angles and waves: intervertebral joint angles and axial kinematics of limbed lizards, limbless lizards, and snakes

Segmentation gives rise to the anterior-posterior axis in many animals, and in vertebrates this axis comprises serially arranged vertebrae. Modifications to the vertebral column abound, and a recurring, but functionally understudied, change is the elongation of the body through the addition and/or elongation of vertebrae. Here, we compared the vertebral and axial kinematics of the robustly limbed Fire skink (Riopa fernandi) representing the ancestral form, the limbless European glass lizard (Ophisaurus apodus), and the Northern water snake (Nerodia sipedon). We induced these animals to traverse through channels and peg arrays of varied widths and densities, respectively, using high-speed X-ray and light video. We found that even though the snake had substantially more and shorter vertebrae than either lizard, intervertebral joint angles did not differ between species in most treatment levels. All three species decreased the amplitude and wavelength of their undulations as channels narrowed and the lizard species increased wave frequency in narrower channels. In peg arrays, both lizard species decreased wave amplitude, while the snake showed no differences. All three species maintained similar wavelengths and frequencies as peg density increased in most cases. Our results suggest that amplitude is decoupled from wavelength and frequency in all three focal taxa. The combination of musculoskeletal differences and the decoupling of axial kinematic traits likely facilitates the formation of different undulatory waves, thereby allowing limbless species to adopt different modes of locomotion.