This study reports on experiments on falcons wearing miniature videocameras mounted on their backs or heads while pursuing flying prey. Videos of hunts by a gyrfalcon (Falco rusticolus), gyrfalcon (F. rusticolus)/Saker falcon (F. cherrug) hybrids and peregrine falcons (F. peregrinus) were analyzed to determine apparent prey positions on their visual fields during pursuits. These video data were then interpreted using computer simulations of pursuit steering laws observed in insects and mammals. A comparison of the empirical and modeling data indicates that falcons use cues due to the apparent motion of prey on the falcon's visual field to track and capture flying prey via a form of motion camouflage. The falcons also were found to maintain their prey's image at visual angles consistent with using their shallow fovea. These results should prove relevant for understanding the co-evolution of pursuit and evasion, as well as the development of computer models of predation and the integration of sensory and locomotion systems in biomimetic robots. KEY WORDS: Predation, Pursuit-evasion, Avian vision, Falcon, Motion camouflage INTRODUCTIONHow predators track and capture their prey poses a fundamental problem in animal behavior that combines sensory perception, neural computation and locomotion. Empirical studies in combination with computational modeling (Pais and Leonard, 2010;Reddy et al., 2006;Srinivasan and Davey, 1995) have identified pursuit strategies used by insects (Olberg, 2012), bats (Ghose et al., 2006), dogs (Shaffer et al., 2004), fish (Lanchester and Mark, 1975) and humans (Fajen and Warren, 2004;McBeath et al., 1995). However, no empirical studies have addressed how falcons and other birds pursue flying prey, mostly due to the difficulty of recording their 3D flight trajectories in the field. The pursuit strategies used by birds have evolved in the context of their unique flight capabilities, as well as their need to pursue rapid, erratically moving prey in complex environments. Understanding their methods for tracking and following rapidly moving objects should provide inspiration for the design of unmanned aerial vehicles (UAVs) and other biomimetic robots.Stereoscopic video methods successfully used to study flocking (Ballerini et al., 2008a;Cavagna et al., 2010;Cavagna et al., 2008) are challenging to apply to this problem because of the wide geographic areas covered and the rapid, unpredictable motion of both predator and prey. However, bird-mounted sensors offer new opportunities for this field. While miniaturized, bird-mounted GPS sensors have been used to study navigation, flocking energetics and decision making in bird flocks (Biro et al., 2006;Nagy et al., 2010;Steiner et al., 2000;Usherwood et al., 2011), the 10 Hz data collection rate and 0.3 m spatial resolution of the GPS are of limited use in the present context. By contrast, miniaturized bird-mounted cameras have proved effective in studying the aerodynamics of avian flight (Carruthers et al., 2007;Gillies et al., 2011) and bird ...
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