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Estimates of circulation and gait change based on a three-dimensional kinematic analysis of flight in cockatiels (Nymphicus hollandicus) and ringed turtle-doves (Streptopelia risoria)
Tyson L. Hedrick, Bret W. Tobalske, Andrew A. Biewener


Birds and bats are known to employ two different gaits in flapping flight, a vortex-ring gait in slow flight and a continuous-vortex gait in fast flight. We studied the use of these gaits over a wide range of speeds (1-17 ms-1) and transitions between gaits in cockatiels (Nymphicus hollandicus) and ringed turtle-doves (Streptopelia risoria) trained to fly in a recently built, variable-speed wind tunnel. Gait use was investigated via a combination of three-dimensional kinematics and quasi-steady aerodynamic modeling of bound circulation on the distal and proximal portions of the wing. Estimates of lift from our circulation model were sufficient to support body weight at all but the slowest speeds (1 and 3 ms-1). From comparisons of aerodynamic impulse derived from our circulation analysis with the impulse estimated from whole-body acceleration, it appeared that our quasi-steady aerodynamic analysis was most accurate at intermediate speeds (5-11 ms-1). Despite differences in wing shape and wing loading, both species shifted from a vortex-ring to a continuous-vortex gait at 7 ms-1. We found that the shift from a vortex-ring to a continuous-vortex gait (i) was associated with a phase delay in the peak angle of attack of the proximal wing section from downstroke into upstroke and (ii) depended on sufficient forward velocity to provide airflow over the wing during the upstroke similar to that during the downstroke. Our kinematic estimates indicated significant variation in the magnitude of circulation over the course the wingbeat cycle when either species used a continuous-vortex gait. This variation was great enough to suggest that both species shifted to a ladder-wake gait as they approached the maximum flight speed (cockatiels 15 ms-1, doves 17 ms-1) that they would sustain in the wind tunnel. This shift in flight gait appeared to reflect the need to minimize drag and produce forward thrust in order to fly at high speed. The ladder-wake gait was also employed in forward and vertical acceleration at medium and fast flight speeds.

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