10Undulatory swimming in flowing media like water is well-studied, but little is known about loco-11 motion in environments that are permanently deformed by body-substrate interactions like snakes in 12 sand, eels in mud, and nematode worms in rotting fruit. We study the desert-specialist snake Chion-13 actis occipitalis traversing granular matter and find body inertia is negligible despite rapid transit 14 and speed dependent granular reaction forces. New surface resistive force theory (RFT) calculation 15 reveals how this snakes wave shape minimizes memory effects and optimizes escape performance 16 given physiological limitations (power). RFT explains the morphology and waveform dependent 17 performance of a diversity of non-sand-specialist, but overpredicts the capability of snakes with 18 high slip. Robophysical experiments recapitulate aspects of these failure-prone snakes and elucidate 19 how reencountering previously remodeled material hinders performance. This study reveals how 20 memory effects stymied the locomotion of a diversity of snakes in our previous studies [Marvi et 21 al, Science, 2014] and suggests the existence of a predictive model for history-dependent granular 22 physics. 23
I. INTRODUCTION
24Movement is critical to the survival of many organisms and a necessary ability in robots used in fields like medicine 25 [1], search and rescue [2], and extraterrestrial exploration [3]. This complex phenomenon emerges from the interplay 26 between a locomotor's internal body shape changes and the physics of the surroundings. Successful locomotion 27 thus depends on the execution of self-deformations which generate appropriate reaction forces from the terrain; a 28 relationship which can be further complicated if the body motion permanently changes the state of the substrate. 29 Much of our knowledge of terrestrial locomotion is in the regime of rigid materials where the terrain is not affected 30 by passage of the animal [4-7], leading to the development of robots which are effective on hard ground [8-10].31 Little is known about locomotion in non-rigid materials which are plastically deformed by the movement of animals 32 or robots, leaving tracks or footprints. Deformable terrains inhabit a spectrum from materials like water which 33 continuously flow toward the undisturbed, zero shear state [11] to those which are remodeled by the interaction, like 34 sand. Most work on motion in yielding materials has been on systems in which disturbances dissipate (fluids) [12-14]; 35 the impact of soft material hysteresis on locomotion is not well-understood [15]. 36 At one end of the spectrum, where deformations are short-lived [16], small fluid swimmers like the nematode 37 Caenorhabditis elegans [17], spermatazoa [18], and bacteria in water [19] and macroscale frictional-fluid swimmers like 38 the sandfish lizard and shovel-nosed snake moving subsurface through sand [5] use the resistance of the surrounding 39 material to the motion of their body shape changes to propel themselves. The fluid in these...