Effects of perch diameter and incline on the kinematics, performance and modes of arboreal locomotion of corn snakes (Elaphe guttata).

Animals moving through arboreal habitats face several functional challenges, including fitting onto and moving on cylindrical branches with variable diameters and inclines. In contrast to lizards and primates, the arboreal locomotion of snakes is poorly understood, despite numerous snake species being arboreal. We quantified the kinematics and performance of corn snakes (Elaphe guttata) moving on seven cylinders (diameters 1.6-21 cm) with five inclines (horizontal, +/-45 degrees and +/-90 degrees) and through horizontal tunnels of corresponding widths. When perches were inclined at either 45 degrees or 90 degrees , snakes were unable to move uphill or downhill on the larger diameters. None of the locomotion on perches conformed to any previously described mode of limbless locomotion. On horizontal and uphill perches snakes performed a variant of concertina locomotion with periodic stopping and gripping. When moving downhill, snakes often slid continuously while grasping the perch to reduce their speed. Mean forward velocity decreased both with increased incline and with increased perch diameter, contrary to the beneficial effect of increased diameter on the speeds of lizards. Both tunnel width and perch diameter had widespread and similar effects on kinematics. When perches and tunnels were narrower, the snakes had more lateral bends at shallower angles. The numerous effects of perch diameter on kinematics and the similarity to tunnel concertina locomotion emphasize the importance of fit as a limitation in arboreal locomotion of snakes. However, the slower speeds on horizontal perches compared to tunnels also suggest that balance and grip may further limit locomotor performance.