Non-planar snake gaits: from Stigmatic-starts to Sidewinding

Of the vast variety of animal gaits, one of the most striking is the non-planar undulating helical motion of a sidewinder. But non-planar helical gaits are not limited to sidewinders. Here we report a new non-planar gait used as an escape strategy in juvenile anacondas (Eunectes notaeus). In the S(tigmatic)-start, named for its eponymous shape, transient locomotion arises when the snake writhes and bends out of the plane while rolling forward about its midsection without slippage. We present a mathematical model for an active non-planar filament that interacts anisotropically with a frictional substrate to quantify our observations and show that locomotion is due to a propagating localized pulse of a topological quantity, the link density. A phase diagram as a function of scaled body size and muscular torques shows that relatively light juveniles are capable of S-starts but heavy adults are not, consistent with our experimental observations. We further show theoretically that a periodic sequence of S-starts naturally leads to sidewinding. All together, our characterization of a novel escape strategy in snakes using non-planar gaits highlights the role of topology in locomotion, provides a phase diagram for gait feasibility as a function of body size, and shows that the S-start forms the fundamental kernel underlying sidewinding.