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Hovering flight helps facilitate feeding, pollination, and courtship. Observed only in smaller flying animals, hover kinematic characteristics are diverse except for the decreasing flapping frequency with the animal size. Although studies have shown that these wing patterns enable distinct unsteady aerodynamic mechanisms, the role of flapping frequency scaling remains a source of disagreement. Here we show that negative allometry of the flapping frequency is required to sustain body attitude during hovering, consistent with experimental data of hovering animals, from fruit flies to hummingbirds, reported in the literature. The derived scaling model reveals that the lift coefficient and reduced frequency remain invariant with mass, enabling leading-edge vortex formation and wake-capture for a wide range of fliers to hover. 

Abstract Flapping flight of animals has captured the interest of researchers due to their impressive flight capabilities across diverse environments including mountains, oceans, forests, and urban areas. Despite the significant progress made in understanding flapping flight, high-altitude flight as showcased by many migrating animals remains underexplored. At high-altitudes, air density is low, and it is challenging to produce lift. Here we demonstrate a first lift-off of a flapping wing robot in a low-density environment through wing size and motion scaling. Force measurements showed that the lift remained high at 0.14 N despite a 66% reduction of air density from the sea-level condition. The flapping amplitude increased from 148 to 233 degrees, while the pitch amplitude remained nearly constant at 38.2 degrees. The combined effect is that the flapping-wing robot benefited from the angle of attack that is characteristic of flying animals. Our results suggest that it is not a simple increase in the flapping frequency, but a coordinated increase in the wing size and reduction in flapping frequency enables the flight in lower density condition. The key mechanism is to preserve the passive rotations due to wing deformation, confirmed by a bioinspired scaling relationship. Our results highlight the feasibility of flight under a low-density, high-altitude environment due to leveraging unsteady aerodynamic mechanisms unique to flapping wings. We anticipate our experimental demonstration to be a starting point for more sophisticated flapping wing models and robots for autonomous multi-altitude sensing. Furthermore, it is a preliminary step towards flapping wing flight in the ultra-low density Martian atmosphere. 

The long-range migration of monarch butterflies, extended over 4000 km, is not well understood. Monarchs experience varying density conditions during migration, ranging as high as 3000 m, where the air density is much lower than at sea level. In this study, we test the hypothesis that the aerodynamic performance of monarchs improves at reduced density conditions by considering the fluid–structure interaction of chordwise flexible wings. A well-validated, fully coupled Navier–Stokes/structural dynamics solver was used to illustrate the interplay between wing motion, aerodynamics, and structural flexibility in forward flight. The wing density and elastic modulus were measured from real monarch wings and prescribed as inputs to the aeroelastic framework. Our results show that sufficient lift is generated to offset the butterfly weight at higher altitudes, aided by the wake-capture mechanism, which is a nonlinear wing–wake interaction mechanism, commonly seen for hovering animals. The mean total power, defined as the sum of the aerodynamic and inertial power, decreased by 36% from the sea level to the condition at 3000 m. Decreasing power with altitude, while maintaining the same equilibrium lift, suggests that the butterflies generate lift more efficiently at higher altitudes.