Rules for the Leg Coordination of Dung Beetle Ball Rolling Behaviour.

Dung beetles can perform a number of versatile behaviours, including walking and dung ball rolling. While different walking and running gaits of dung beetles have been described in previous literature, little is known about their ball rolling gaits. From behavioural experiments and video recordings of the beetle Scarabaeus (Kheper) lamarcki, we analysed and identified four underlying rules for leg coordination during ball rolling. The rules describe the alternation of the front legs and protraction waves of the middle and hind legs. We found that while rolling a ball backwards, the front legs are decoupled or loosely coupled from the other legs, resulting in a non-standard gait, in contrast to previously described tripod and gallop walking gaits in dung beetles. This provides insight into the principles of leg coordination in dung beetle ball rolling behaviour and its underlying rules. The proposed rules can be used as a basis for further investigation into ball rolling behaviours on more complex terrain (e.g., uneven terrain and slopes). Additionally, the rules can also be used to guide the development of control mechanisms for bio-inspired ball rolling robots.