The wake of hovering flight in bats

Hovering means stationary flight at zero net forward speed, which can be achieved by animals through muscle powered flapping flight. Small bats capable of hovering typically do so with a downstroke in an inclined stroke plane, and with an aerodynamically active outer wing during the upstroke. The magnitude and time history of aerodynamic forces should be reflected by vorticity shed into the wake. We thus expect hovering bats to generate a characteristic wake, but this has until now never been studied. Here we trained nectar-feeding bats, Leptonycteris yerbabuenae, to hover at a feeder and using time-resolved stereoscopic particle image velocimetry in conjunction with high-speed kinematic analysis we show that hovering nectar-feeding bats produce a series of bilateral stacked vortex loops. Vortex visualizations suggest that the downstroke produces the majority of the weight support, but that the upstroke contributes positively to the lift production. However, the relative contributions f...