Kinematics and aerodynamics of avian upstrokes during slow flight

Slow flight is extremely energetically costly per unit time, yet highly important for takeoff and survival. However, at slow speeds it is presently thought that most birds do not produce beneficial aerodynamic forces during the entire wingbeat: instead they fold or flex their wings during upstroke, prompting the long-standing prediction that the upstroke produces trivial forces. There is increasing evidence that the upstroke contributes to force production, but the aerodynamic and kinematic mechanisms remain unknown. Here, we examined the wingbeat cycle of two species: the diamond dove (Geopelia cuneata) and zebra finch (Taeniopygia guttata), which exhibit different upstroke styles - a wingtip-reversal and flexed-wing upstroke, respectively. We used a combination of particle image velocimetry and near-wake streamline measures alongside detailed 3D kinematics. We show that during the middle of the wingtip-reversal upstroke, the hand-wing has a high angular velocity (15.3 +/- 0.8 deg ms(-1)) and translational speed (8.4 +/- 0.6 m s(-1)). The flexed-wing upstroke, in contrast, has low wingtip speed during mid-upstroke. Instead, later in the stroke cycle, during the transition from upstroke to downstroke, it exhibits higher angular velocities (45.5 +/- 13.8 deg ms(-1)) and translational speeds (11.0 +/- 1.9 m s(-1)). Aerodynamically, the wingtip-reversal upstroke imparts momentum to the wake, with entrained air shed backward (visible as circulation of 14.4 +/- 0.09 m(2) s(-1)). In contrast, the flexed-wing upstroke imparts minimal momentum. Clap and peel in the dove enhances the time course for circulation production on the wings, and provides new evidence of convergent evolution on time-varying aerodynamic mechanisms during flapping in insects and birds.