Video filmed by a camera mounted on the head of a Northern Goshawk (Accipiter gentilis) was used to study how the raptor used visual guidance to pursue prey and land on perches. A combination of novel image analysis methods and numerical simulations of mathematical pursuit models was used to determine the goshawk's pursuit strategy. The goshawk flew to intercept targets by fixing the prey at a constant visual angle, using classical pursuit for stationary prey, lures or perches, and usually using constant absolute target direction (CATD) for moving prey. Visual fixation was better maintained along the horizontal than vertical direction. In some cases, we observed oscillations in the visual fix on the prey, suggesting that the goshawk used finite-feedback steering. Video filmed from the ground gave similar results. In most cases, it showed goshawks intercepting prey using a trajectory consistent with CATD, then turning rapidly to attack by classical pursuit; in a few cases, it showed them using curving non-CATD trajectories. Analysis of the prey's evasive tactics indicated that only sharp sideways turns caused the goshawk to lose visual fixation on the prey, supporting a sensory basis for the surprising frequency and effectiveness of this tactic found by previous studies. The dynamics of the prey's looming image also suggested that the goshawk used a tau-based interception strategy. We interpret these results in the context of a concise review of pursuit-evasion in biology, and conjecture that some prey deimatic 'startle' displays may exploit tau-based interception.KEY WORDS: Pursuit-evasion, Predator-prey, Avian vision, Northern Goshawk, Visual guidance, Sensory ecology, Accipiter gentilis, Looming, Antipredator behavior, Startle effect
INTRODUCTIONMany problems in the study of animal behavior require an integrated biomechanical and sensory ecology approach that considers the organism's locomotor and perceptual capabilities, its sensory cues and the dynamics of its behavioral responses (Fernández-Juricic, 2012;Martin, 2012). Animal-borne video methods (Rutz and Troscianko, 2013) now make it possible to measure the visual cues received by organisms in the field, revealing new information about, for example, how they move through their environment (BBC, 2009), interact with conspecifics (Takahashi et al., 2004) and use tools (Rutz et al., 2007). Recent studies have used the stable video recorded by cameras mounted on the heads of birds (headcams) to explore the visual strategies used by falcons chasing prey (Kane and Zamani, 2014) and peafowl detecting model predators (Yorzinski and Platt, 2014). Here, we report using headcam video to explore for the first time pursuit-evasion and landing behavior in the Northern Goshawk, Accipiter gentilis (Linnaeus 1758) (hereafter, goshawk), a large diurnal raptor (Fig. 1). Biologically derived models have inspired robotic algorithms for swarming, following and collision avoidance (Mischiati and Krishnaprasad, 2012;Srinivasan, 2011), and the goshawk is of special inte...