Locomotory transition from water to sand and its effects on undulatory kinematics in sand lances (Ammodytidae).

Sand lances, fishes in the genus Ammodytes, exhibit a peculiar burrowing behavior in which they appear to swim rapidly into the substrate. They use posteriorly propagated undulations of the body to move in both water, a Newtonian fluid, and in sand, a non-Newtonian, granular substrate. In typical aquatic limbless locomotion, undulations of the body push against water, which flows because it is incapable of supporting the static stresses exerted by the animal, thus the undulations move in world space (slipping wave locomotion). In typical terrestrial limbless locomotion, these undulations push against substrate irregularities and move relatively little in world space (non-slipping wave locomotion). We used standard and X-ray video to determine the roles of slipping wave and non-slipping wave locomotion during burrowing in sand lances. We find that sand lances in water use slipping wave locomotion, similar to most aquatic undulators, but switch to non-slipping waves once they burrow. We identify a progression of three stages in the burrowing process: first, aquatic undulations similar to typical anguilliform locomotion (but without head yaw) push the head into the sand; second, more pronounced undulations of the aquatic portion of the body push most of the animal below ground; third, the remaining above-ground portion of the body ceases undulation and the subterranean portion takes over, transitioning to non-slipping wave locomotion. We find no evidence that sand lances use their body motions to fluidize the sand. Instead, as soon as enough of the body is underground, they undergo a kinematic shift and locomote like terrestrial limbless vertebrates.