Body oscillations couple with wing flapping to reduce aerodynamic power in wild silk moth flight

Insects show diverse flight kinematics and morphologies reflecting their evolutionary histories and ecological adaptations. Many silk moths use low wingbeat frequencies and large wings to fly and display body oscillations. Their bodies pitch and bob periodically, synchronized with their wingbeat cycle. Similar oscillations in butterflies improve weight support and forward thrust while reducing flight power requirements. However, how instantaneous body and wing kinematics interact for these beneficial aerodynamic and power consequences is not well understood. We hypothesized that body oscillations affect aerodynamic power requirements by influencing wing rotation relative to the airflow. Using three-dimensional forward flight video recordings of four silk moth species and a quasi-steady blade-element aerodynamic model, we analysed the aerodynamic effects of body and wing kinematics. We find that the body pitch and wing sweep angles maintain a narrow range of phase differences, which enhances the angle of attack variation between each half-stroke due to increased wing rotation relative to the airflow. This redirects the aerodynamic force to increase the upward and forward components during the downstroke and upstroke, respectively, thus lowering overall drag without compromising weight support and forward thrust. Reducing energy expenditure is beneficial because many adult silk moths do not feed and rely on limited energy budgets.