The most effective way to avoid intense inter- and intra-specific competition at the dung source, and to increase the distance to the other competitors, is to follow a single straight bearing. While ball-rolling dung beetles manage to roll their dung balls along nearly perfect straight paths when traversing flat terrain, the paths that they take when traversing more complex (natural) terrain are not well understood. In this study, we investigate the effect of complex surface topographies on the ball-rolling ability of Kheper lamarcki. Our results reveal that ball-rolling trajectories are strongly influenced by the characteristic scale of the surface structure. Surfaces with an increasing similarity between the average distance of elevations and the ball radius cause progressively more difficulties during ball transportation. The most important factor causing difficulties in ball transportation appears to be the slope of the substrate. Our results show that, on surfaces with a slope of 7.5 deg, more than 60% of the dung beetles lose control of their ball. Although dung beetles still successfully roll their dung ball against the slope on such inclinations, their ability to roll the dung ball sideways diminishes. However, dung beetles do not seem to adapt their path on inclines such that they roll their ball in the direction against the slope. We conclude that dung beetles strive for a straight trajectory away from the dung pile, and that their actual path is the result of adaptations to particular surface topographies.

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