Transitions to and from the air are critical for aerial locomotion and likely shaped the evolution of flying animals. Research on take-off demonstrates that legs generate greater body accelerations compared with wings, and thereby contribute more to initial flight velocity. Here, we explored coordination between wings and legs in two species with different wingbeat styles, and quantified force production of these modules during the final phase of landing. We used the same birds that we had previously studied during take-off: zebra finch (Taeniopygia guttata, N=4) and diamond dove (Geopelia cuneata, N=3). We measured kinematics using high-speed video, aerodynamics using particle image velocimetry, and ground-reaction forces using a perch mounted on a force plate. In contrast with the first three wingbeats of take-off, the final four wingbeats during landing featured ~2 times greater force production. Thus, wings contribute proportionally more to changes in velocity during the last phase of landing compared with the initial phase of take-off. The two species touched down at the same velocity (~1 m s −1 ), but they exhibited significant differences in the timing of their final wingbeat relative to touchdown. The ratio of average wing force to peak leg force was greater in diamond doves than in zebra finches. Peak ground reaction forces during landing were ~50% of those during take-off, consistent with the birds being motivated to control landing. Likewise, estimations of mechanical energy flux for both species indicate that wings produce 3-10 times more mechanical work within the final wingbeats of flight compared with the kinetic energy of the body absorbed by legs during ground contact.KEY WORDS: Zebra finch, Taeniopygia guttata, Diamond dove, Geopelia cuneata, Locomotion, Hindlimb, Forelimb, Particle image velocimetry, PIV
INTRODUCTIONLanding allows the transition from the air to the ground or other substrate. This phase is fundamental to flight, but is often overlooked in studies compared with take-off or cruising flight. Landing places a unique selective pressure upon aerial animals, because a controlled deceleration and descent must be accomplished to avoid injury (Bonser, 1999;Paskins et al., 2007). Animals that do not decelerate properly risk broken wings or legs, or damaged integument: collisions often lead to mortality (Klem, 1990 birds, and even the engineering of autonomous flying vehicles. Herein, we explored the coordination of force production between both legs and wings during landing in birds.Several lines of evidence indicate that birds carefully control their landing and generally seek to minimize their kinetic energy at touchdown (Warrick et al., 2002) even if some species are eminently capable of targeting in fast flight [for example, a peregrine falcon stooping on a pigeon at relatively fast speed (Alerstam, 1987)]. During landing, birds modulate both wing and tail kinematics to decelerate prior to arriving at the perch (Berg and Biewener, 2010). The distal muscles of the wing, associ...