Wake patterns of the wings and tail of hovering hummingbirds

Abstract

The flow fields of slowly flying bats and faster-flying birds differ in that bats produce two vortex loops during each stroke, one per wing, and birds produce a single vortex loop per stroke. In addition, the circulation at stroke transition approaches zero in bats but remains strong in birds. It is unknown if these difference derive from fundamental differences in wing morphology or are a consequence of flight speed. Here, we present an analysis of the horizontal flow field underneath hovering Anna’s hummingbirds (Calypte anna) to describe the wake of a bird flying at zero forward velocity. We also consider how the hummingbird tail interacts with the wake generated by the wings. High-speed image recording and analysis from three orthogonal perspectives revealed that the wing tips reach peak velocities in the middle of each stroke and approach zero velocity at stroke transition. Hummingbirds use complex tail kinematic patterns ranging from in phase to antiphase cycling with respect to the wings, covering several phase shifted patterns. We employed particle image velocimetry to attain detailed horizontal flow measurements at three levels with respect to the tail: in the tail, at the tail tip, and just below the tail. The velocity patterns underneath the wings indicate that flow oscillates along the ventral–dorsal axis in response to the down- and up-strokes and that the sideways flows with respect to the bird are consistently from the lateral to medial. The region around the tail is dominated by axial flows in dorsal to ventral direction. We propose that these flows are generated by interaction between the wakes of the two wings at the end of the upstroke, and that the tail actively defects flows to generate moments that contribute to pitch stability. The flow fields images also revealed distinct vortex loops underneath each wing, which were generated during each stroke. From these data, we propose a model for the primary flow structures of hummingbirds that more strongly resembles the bat model. Thus, pairs of unconnected vortex loops may be shared features of different animals during hovering and slow forward flight.