Drinking in snakes: kinematic cycling and water transport.

Snakes are purported to drink by sucking water into their mouths and then compressing the oral cavity to force water into the oesophagus. Video recordings of drinking behaviour in 23 snakes representing 14 species from three families, combined with simultaneous recordings of water volumes consumed, show that all the snakes vary widely in the amount of water taken in when drinking. This variation is not correlated with kinematic events. Kinematic recordings and indirect measurements of water flow suggest that moving water into the mouth can be decoupled from the processes that move water into the oesophagus and that, infrequently, water may continue flowing into the mouth during both opening (suction) and closing (presumed compression) of the mouth. Drinking in snakes is not a simple, stereotyped behaviour. Different snake species differ in both drinking kinematics and water inflow patterns. Vertical excursions of the mandible are smallest in booids and larger, but highly variable, in different viperids and colubrids. Cyclic movements of the tongue seen in booids are not evident in viperids or colubrids. All the snakes usually take in water at rates far below their potential maximum rate. Although drinking is apparently achieved by suction, a single model cannot explain all water movement patterns in snakes. At a practical level, functional morphological studies of drinking in snakes (and possibly many other animals) must demonstrate that fluid flow actually correlates with kinematic events. Without such an empirical demonstration, interpretation of other measurements (pressure, movement, etc.) is unlikely to produce meaningful models.