Abstract
Hummingbirds outperform other birds in terms of aerial agility at low flight speeds. To reveal the key mechanisms that enable such unparalleled agility, we reconstructed body and wing motion of hummingbird escape manoeuvres from high-speed videos; then, we performed computational fluid dynamics modelling and flight mechanics analysis, in which the time-dependent forces within each wingbeat were resolved. We found that the birds may use the inertia of their wings to achieve peak body rotational acceleration around wing reversal when the aerodynamic forces were small. The aerodynamic forces instead counteracted the reversed inertial forces at a different wingbeat phase, thereby stabilizing the body from inertial oscillations, or they could become dominant and provide additional rotational acceleration. Our results suggest such an inertial steering mechanism was present for all four hummingbird species considered, and it was used by the birds for both pitch-up and roll accelerations. The combined inertial steering and aerodynamic mechanisms made it possible for the hummingbirds to generate instantaneous body acceleration at any phase of a wingbeat, and this feature is probably the key to understanding the unique dexterity distinguishing hummingbirds from other small-size flyers that solely rely on aerodynamics for manoeuvering.