Hawkmoth flight performance in tornado-like whirlwind vortices

Vertical vortex systems such as tornadoes dramatically affect the flight control and stability of aircraft. However, the control implications of smaller scale vertically oriented vortex systems for small fliers such as animals or micro-air vehicles are unknown. Here we examined the flapping kinematics and body dynamics of hawkmoths performing hovering flights (controls) and maintaining position in three different whirlwind intensities with transverse horizontal velocities of 0.7, 0.9 and 1.2 m s(-1), respectively, generated in a vortex chamber. The average and standard deviation of yaw and pitch were respectively increased and reduced in comparison with hovering flights. Average roll orientation was unchanged in whirlwind flights but was more variable from wingbeat to wingbeat than in hovering. Flapping frequency remained unchanged. Wingbeat amplitude was lower and the average stroke plane angle was higher. Asymmetry was found in the angle of attack between right and left wings during both downstroke and upstroke at medium and high vortex intensities. Thus, hawkmoth flight control in tornado-like vortices is achieved by a suite of asymmetric and symmetric changes to wingbeat amplitude, stroke plane angle and principally angle of attack.