Winged flying animals, such as insects, are superior to similar-sized flying robots in terms of flight performance, particularly in unstable and unpredictable natural environments. Unlike flying robots, such as drones, which generate aerodynamic forces by rotary or fixed wings, insects utilize flapping wings to generate and control aerodynamic forces (Floreano and Wood, 2015;Phan and Park, 2019;Shyy et al., 2013). Using flapping motion, insects dynamically change their wing kinematics, such as the angle of attack. Therefore, their flight relies heavily on various unsteady aerodynamic mechanisms, such as leading-edge vortex (LEV), rotational forces, wake capture, and clap-and-fling (Sane, 2003). Furthermore, controlling the wingbeat amplitude and rotational timing affects the generation of aerodynamic force and torque through the synergetic effects of unsteady aerodynamic mechanisms (Dickinson et al., 1999). Therefore, wing motion adjustment is considerably important for efficient lift generation and flight control during disturbances.Insects achieve flapping motion by rhythmic contraction of the flight muscles. Flight muscles are categorized into two anatomically, physiologically, and functionally distinct groups: direct muscles inserted into the wing base, and indirect muscles attached to the thoracic exoskeleton (Dickinson and Tu, 1997). Dragonflies, which move their wings using direct flight muscles, can control all four wings independently at varying frequencies, amplitudes, and angles of attack when maneuvering (Svidersky et al., 2008). A number of other insects with indirect flight muscles, including