Tiger beetles pursue prey using a proportional control law with a delay of one half-stride

Tiger beetles are fast diurnal predators capable of chasing prey under closed-loop visual guidance. We investigated this control system using statistical analyses of high-speed digital recordings of beetles chasing a moving prey dummy in a laboratory arena. Correlation analyses reveal that the beetle uses a proportional control law in which the angular position of the prey relative to the beetle's body axis drives the beetle's angular velocity with a delay of about 28 ms. The proportionality coefficient or system gain, 12 s(-1), is just below critical damping. Pursuit simulations using the derived control law predict angular orientation during pursuits with a residual error of about 7°. This is of the same order of magnitude as the oscillation imposed by the beetle's alternating tripod gait, which was not factored into the control law. The system delay of 28 ms equals a half-stride period, i.e. the time between the touch down of alternating tripods. Based on these results, we propose a physical interpretation of the observed control law: to turn towards its prey, the beetle on average exerts a sideways force proportional to the angular position of the prey measured a half-stride earlier.