Flight kinematics of black-billed magpies and pigeons over a wide range of speeds
To investigate how birds that differ in morphology change their wing and body movements while flying at a range of speeds, we analyzed high-speed (60 Hz) video tapes of black-billed magpies (Pica pica) flying at speeds of 4-14 m s-1 and pigeons (Columba livia) flying at 6-20 m s-1 in a wind-tunnel. Pigeons had higher wing loading and higher-aspect-ratio wings compared with magpies. Both species alternated phases of steady-speed flight with phases of acceleration and deceleration, particularly at intermediate flight speeds. The birds modulated their wingbeat kinematics among these phases and frequently exhibited non-flapping phases while decelerating. Such modulation in kinematics during forward flight is typical of magpies but not of pigeons in the wild. The behavior of the pigeons may have been a response to the reduced power costs for flight in the closed wind-tunnel relative to those for free flight at similar speeds. During steady-speed flight, wingbeat frequency did not change appreciably with increasing flight speed. Body angle relative to the horizontal, the stroke-plane angles of the wingtip and wrist relative to the horizontal and the angle describing tail spread at mid-downstroke all decreased with increasing flight speed, thereby illustrating a shift in the dominant function of wing flapping from weight support at slow speeds to positive thrust at fast speeds. Using wingbeat kinematics to infer lift production, it appeared that magpies used a vortex-ring gait during steady-speed flight at all speeds whereas pigeons used a vortex-ring gait at 6 and 8 m s-1, a transitional vortex-ring gait at 10 m s-1, and a continuous-vortex gait at faster speeds. Both species used a vortex-ring gait for acceleration and a continuous-vortex gait or a non-flapping phase for deceleration during flight at intermediate wind-tunnel speeds. Pigeons progressively flexed their wings during glides as flight speed increased but never performed bounds. Wingspan during glides in magpies did not vary with flight speed, but the percentage of bounds among non-flapping intervals increased with speed from 10 to 14 m s-1. The use of non-flapping wing postures seemed to be related to the gaits used during flapping and to the aspect ratio of the wings. We develop an 'adverse-scaling' hypothesis in which it is proposed that the ability to reduce metabolic and mechanical power output using flap-bounding flight at fast flight speeds is scaled negatively with body mass. This represents an alternative to the 'fixed-gear' hypothesis previously suggested by other authors to explain the use of intermittent flight in birds. Future comparative studies in the field would be worthwhile, especially if instantaneous flight speeds and within-wingbeat kinematics were documented; new studies in the laboratory should involve simultaneous recording of wing kinematics and aerodynamic forces on the wing.
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