Compliant legs enable lizards to maintain high running speeds on complex terrains.

Substrate variations are likely to constrain animal performance in natural environments, as running over complex terrains challenges the dynamic stability of the body differently in each step. Yet, being able to negotiate complex terrains at top speed is a strong advantage for animals that have to deal with predators and evasive prey. Little is known on how animals negotiate such terrain variability at high speed. We investigated this in fast-running lizards, by measuring their 3D kinematics using four synchronised high-speed video cameras (325 Hz) on an adaptable racetrack. This racetrack was covered with four different substrates, representing increasing levels of terrain complexity. We found that the lizards deal with this complexity gradient by gradually adopting more erect parasagittal leg postures. Legs in a more-erect position are more compliant and are therefore highly adjustable on complex terrains. Additionally, the lizards stabilise their head, which facilitates vestibular and visual perception. Together, compliant legs and head stabilisation enable the lizards to minimise movements of the body centre of mass, even when running on highly irregular terrains. This suggests that the head and the centre of mass are the priority targets for running on uneven terrains. As a result, running performance (mean forward speed) decreases only slightly, and only on the most challenging substrate under investigation.