Dynamics of the vortex wakes of flying and swimming vertebrates.

The vortex wakes of flying and swimming animals provide evidence of the history of aero- and hydrodynamic force generation during the locomotor cycle. Vortex-induced momentum flux in the wake is the reaction of forces the animal imposes on its environment, which must be in equilibrium with inertial and external forces. In flying birds and bats, the flapping wings generate lift both to provide thrust and to support the weight. Distinct wingbeat and wake movement patterns can be identified as gaits. In flow visualization experiments, only two wake patterns have been identified: a vortex ring gait with inactive upstroke, and a continuous vortex gait with active upstroke. These gaits may be modelled theoretically by free vortex and lifting line theory to predict mechanical energy consumption, aerodynamic forces and muscle activity. Longer-winged birds undergo a distinct gait change with speed, but shorter-winged species use the vortex ring gait at all speeds. In swimming fish, the situation is more complex: the wake vortices form a reversed von Kármán vortex street, but little is known about the mechanism of generation of the wake, or about how it varies with speed and acceleration or with body form and swimming mode. An unresolved complicating factor is the interaction between the drag wake of the flapping fish body and the thrusting wake from the tail.