What Defines Different Modes of Snake Locomotion?

Animals move in diverse ways, as indicated in part by the wide variety of gaits and modes that have been described for vertebrate locomotion. Much variation in the gaits of limbed animals is associated with changing speed, whereas different modes of snake locomotion are often associated with moving on different surfaces. For several decades different types of snake locomotion have been categorized as one of four major modes: rectilinear, lateral undulation, sidewinding, and concertina. Recent empirical work shows that the scheme of four modes of snake locomotion is overly conservative. For example, during aquatic lateral undulation, the timing between muscle activity and lateral bending changes along the length of the snake, which is unlike terrestrial lateral undulation. The motor pattern used to prevent sagging while bridging gaps also suggests that arboreal lateral undulation on narrow surfaces or with a few discrete points of support has a different motor pattern than terrestrial lateral undulation when the entire length of the snake is supported. In all types of concertina locomotion, the distance from the head to the tail changes substantially as snakes alternately flex and then extend different portions of their body. However, snakes climbing cylinders with concertina exert forces medially to attain a purchase on the branch, whereas tunnels require pushing laterally to form an anchoring region. Furthermore, different motor patterns are used for these two types of concertina movement. Some snakes climb vertical cylinders with helical wrapping completely around the cylinder, whereas all other forms of concertina bend regions of the body alternately to the left and right. Current data support rectilinear locomotion and sidewinding as being distinct modes, whereas lateral undulation and concertina are best used for defining categories of gaits with some unifying similarities. Partly as a result of different motor patterns, I propose recognizing five and four distinct types of lateral undulation and concertina, respectively, resulting in a total of 11 distinct gaits previously recognized as only four.