Cicada (Tibicen linnei) steers by force vectoring

To change flight direction, flying animals modulate aerodynamic force either relative to their bodies to generate torque about the center of mass, or relative to the flight path to produce centripetal force that curves the trajectory. In employing the latter, the direction of aerodynamic force remains fixed in the body frame and rotations of the body redirect the force. While both aforementioned techniques are essential for flight, it is critical to investigate how an animal balances the two to achieve aerial locomotion. Here, we measured wing and body kinematics of cicada (Tibicen linnei) in free flight, including flight periods of both little and substantial body reorientations. It is found that cicadas employ a common force vectoring technique to execute all these flights. We show that the direction of the half-stroke averaged aerodynamic force relative to the body is independent of the body orientation, varying in a range of merely 20 deg. Despite directional limitation of the aerodynamic force, pitch and roll torque are generated by altering wing angle of attack and its mean position relative to the center of mass. This results in body rotations which redirect the wing force in the global frame and consequently change the flight trajectory.