Aerial shaking performance of wet Anna's hummingbirds.

External wetting poses problems of immediate heat loss and long-term pathogen growth for vertebrates. Beyond these risks, the locomotor ability of smaller animals, and particularly of fliers, may be impaired by water adhering to the body. Here, we report on the remarkable ability of hummingbirds to perform rapid shakes in order to expel water from their plumage even while in flight. Kinematic performance of aerial versus non-aerial shakes (i.e. those performed while perching) was compared. Oscillation frequencies of the head, body and tail were lower in aerial shakes. Tangential speeds and accelerations of the trunk and tail were roughly similar in aerial and non-aerial shakes, but values for head motions while perching were twice as high when compared with aerial shakes [corrected] . Azimuthal angular amplitudes for both aerial and non-aerial shakes reached values greater than 180° for the head, greater than 45° for the body trunk and slightly greater than 90° for the tail and wings. Using a feather on an oscillating disc to mimic shaking motions, we found that bending increased average speeds by up to 36 per cent and accelerations of the feather tip up to fourfold relative to a hypothetical rigid feather. Feather flexibility may help to enhance shedding of water and reduce body oscillations during shaking.