Flying in reverse: kinematics and aerodynamics of a dragonfly in backward free flight.

In this study, we investigated the backward free flight of a dragonfly, accelerating in a flight path inclined to the horizontal. The wing and body kinematics were reconstructed from the output of three high-speed cameras using a template-based subdivision surface reconstruction method, and numerical simulations using an immersed boundary flow solver were conducted to compute the forces and visualize the flow features. During backward flight, the dragonfly maintained an upright body posture of approximately 90° relative to the horizon. The upright body posture was used to reorient the stroke plane and the flight force in the global frame; a mechanism known as 'force vectoring' which was previously observed in manoeuvres of other flying animals. In addition to force vectoring, we found that while flying backward, the dragonfly flaps its wings with larger angles of attack in the upstroke (US) when compared with forward flight. Also, the backward velocity of the body in the upright position enhances the wings' net velocity in the US. The combined effect of the angle of attack and wing net velocity yields large aerodynamic force generation in the US, with the average magnitude of the force reaching values as high as two to three times the body weight. Corresponding to these large forces was the presence of a strong leading edge vortex (LEV) at the onset of US which remained attached up until wing reversal. Finally, wing-wing interaction was found to enhance the aerodynamic performance of the hindwings (HW) during backward flight. Vorticity from the forewings' trailing edge fed directly into the HW LEV to increase its circulation and enhance force production.