Coordinating tiny limbs and long bodies: geometric mechanics of diverse undulatory lizard locomotion

Although typically possessing four limbs and short bodies, lizards have evolved a diversity of body plans, from short-bodied and fully-limbed to elongate and nearly limbless. Such diversity in body morphology is hypothesized as adaptations to locomotion cluttered terrestrial environments, but the mode of propulsion -- e.g., the use of body and/or limbs to interact with the substrate -- and potential body/limb coordination remain unstudied. Here, we use biological experiments, a geometric theory of locomotion, and robophysical experiments to comparatively and systematically investigate such dynamics in a diverse sample of lizard morphologies. Locomotor field studies in short-limb, elongated lizards (Brachymeles) and laboratory studies of full-limbed lizards (Uma scoparia and Sceloporus olivaceus) and a limbless laterally undulating organism (Chionactis occipitalis) reveal that the body wave dynamics can be described by a combination of traveling and standing waves; the ratio of the amplitudes of these components is inversely related to limb length. We use geometric theory to analyze and explain the wave dynamics and body-leg coordination observations; the theory predicts that leg thrust modulates the body weight distribution and self-propulsion generation mechanism, which in turn facilitates the choice of body waves. We test our hypothesis in biological experiments by inducing the use of traveling wave in stereotyped lizards by modulating the ground penetration resistance, as well as in controlled non-biological experiments involving an undulating limbed robophysical model. Our models could be valuable in understanding functional constraints on the evolutionary process of elongation and limb reduction in lizards, as well as advancing robot designs.