Head stabilisation in fast running lizards.

The cyclic patterns of terrestrial animal locomotion are frequently perturbed in natural environments. The terrain can be complex or inclined, the substrate can move unexpectedly and animals can misjudge situations. Loosing stability due to perturbations increases the probability of capture by predators and decreases the chance of successful prey capture and winning intraspecific battles. When controlled corrective actions are necessary to negotiate perturbations, animals rely on their exteroceptive and proprioceptive senses to monitor the environment and their own body movements. The vestibular system in the inner ear perceives linear and angular accelerations. This information enables gaze stabilisation and the creation of a stable, world-bound reference frame for the integration of the information of other senses. During locomotion, both functions are known to be facilitated by head stabilisation in several animals with an erect posture. Animals with a sprawled body posture, however, undergo very large body undulations while running. Using high speed video recordings, we tested whether they nevertheless stabilise their head during running, and how this is influenced by perturbations. We found that running Acanthodactylus boskianus lizards strongly stabilise their head yaw rotations when running on a flat, straight runway: the head rotation amplitude is only 4.76±0.99°, while the adjacent trunk part rotates over 27.0±3.8°. Lateral head translations are not stabilised (average amplitude of 7.4±2.0mm). When the lizards are experimentally perturbed by a large and unexpected lateral substrate movement, lateral translations of both the head and the body decrease (on average by 1.52±0.81mm). At the same time, the rotations of the head and trunk also decrease (on average by 1.62°±7.21°). These results show that head stabilisation intensifies because of the perturbation, which emphasises the importance of vestibular perception and balance in these fast and manoeuvrable animals.