Undulatory swimming in sand: subsurface locomotion of the sandfish lizard

The desert-dwelling sandfish (Scincus scincus) moves within dry sand, a material that displays solid and fluidlike behavior. High-speed x-ray imaging shows that below the surface, the lizard no longer uses limbs for propulsion but generates thrust to overcome drag by propagating an undulatory traveling wave down the body. Although viscous hydrodynamics can predict swimming speed in fluids such as water, an equivalent theory for granular drag is not available. To predict sandfish swimming speed, we developed an empirical model by measuring granular drag force on a small cylinder oriented at different angles relative to the displacement direction and summing these forces over the animal movement profile. The agreement between model and experiment implies that the noninertial swimming occurs in a frictional fluid.