Multi-scale tarsal adhesion kinematics of freely-walking dock beetles

In this experimental study, living dock beetles are observed during their free upside-down walk on a smooth horizontal substrate. Their weight is balanced by the adhesion of hairy structures present on their tarsomeres. The motions involved in the attachment and detachment of these structures were characterized by simultaneously imaging the beetle from the side at the body scale, and from the top at the scale of a single tarsal chain. The observed multi-scale three-dimensional kinematics of the tarsi is qualitatively described, then quantified by image processing and physically modelled. A strong asymmetry is systematically observed between attachment and detachment kinematics, in terms of both timing and directionality.