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1. Interspecific variation in bristle number on forewings of tiny insects does not influence clap-and-fling aerodynamics

   https://doi.org/10.1242/jeb.239798  

   Kasoju, Vishwa T.; Moen, Daniel S.; Ford, Mitchell P.; Ngo, Truc T.; Santhanakrishnan, Arvind (September 2021, Journal of Experimental Biology)

   null (Ed.)

   ABSTRACT Miniature insects must overcome significant viscous resistance in order to fly. They typically possess wings with long bristles on the fringes and use a clap-and-fling mechanism to augment lift. These unique solutions to the extreme conditions of flight at tiny sizes (<2 mm body length) suggest that natural selection has optimized wing design for better aerodynamic performance. However, species vary in wingspan, number of bristles (n) and bristle gap (G) to diameter (D) ratio (G/D). How this variation relates to body length (BL) and its effects on aerodynamics remain unknown. We measured forewing images of 38 species of thrips and 21 species of fairyflies. Our phylogenetic comparative analyses showed that n and wingspan scaled positively and similarly with BL across both groups, whereas G/D decreased with BL, with a sharper decline in thrips. We next measured aerodynamic forces and visualized flow on physical models of bristled wings performing clap-and-fling kinematics at a chord-based Reynolds number of 10 using a dynamically scaled robotic platform. We examined the effects of dimensional (G, D, wingspan) and non-dimensional (n, G/D) geometric variables on dimensionless lift and drag. We found that: (1) increasing G reduced drag more than decreasing D; (2) changing n had minimal impact on lift generation; and (3) varying G/D minimally affected aerodynamic forces. These aerodynamic results suggest little pressure to functionally optimize n and G/D. Combined with the scaling relationships between wing variables and BL, much wing variation in tiny flying insects might be best explained by underlying shared growth factors. 

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2. The aerodynamics of flying snake airfoils in tandem configuration

   https://doi.org/10.1242/jeb.233635  

   Jafari, Farid; Holden, Daniel; LaFoy, Roderick; Vlachos, Pavlos P.; Socha, John J. (July 2021, Journal of Experimental Biology)

   null (Ed.)

   ABSTRACT Flying snakes flatten their body to form a roughly triangular cross-sectional shape, enabling lift production and horizontal acceleration. While gliding, they also assume an S-shaped posture, which could promote aerodynamic interactions between the fore and the aft body. Such interactions have been studied experimentally; however, very coarse models of the snake's cross-sectional shape were used, and the effects were measured only for the downstream model. In this study, the aerodynamic interactions resulting from the snake's posture were approximated using two-dimensional anatomically accurate airfoils positioned in tandem to mimic the snake's geometry during flight. Load cells were used to measure the lift and drag forces, and flow field data were obtained using digital particle image velocimetry (DPIV). The results showed a strong dependence of the aerodynamic performance on the tandem arrangement, with the lift coefficients being generally more influenced than the drag coefficients. Flow field data revealed that the tandem arrangement modified the separated flow and the wake size, and enhanced the lift in cases in which the wake vortices formed closer to the models, producing suction on the dorsal surface. The downforce created by the flow separation from the ventral surface of the models at 0 deg angle of attack was another significant factor contributing to lift production. A number of cases showing large variations of aerodynamic performance included configurations close to the most probable posture of airborne flying snakes, suggesting that small postural variations could be used to control the glide trajectory. 

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

   https://doi.org/10.1098/rsif.2025.0061  

   Sikandar, Usama Bin; Aiello, Brett R; Sponberg, Simon (August 2025, Journal of The Royal Society Interface)

   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. 

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4. First lift-off and flight performance of a tailless flapping-wing aerial robot in high-altitude environments

   https://doi.org/10.1038/s41598-023-36174-5  

   Tsuchiya, Shu; Aono, Hikaru; Asai, Keisuke; Nonomura, Taku; Ozawa, Yuta; Anyoji, Masayuki; Ando, Noriyasu; Kang, Chang-kwon; Pohly, Jeremy (December 2023, Scientific Reports)

   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. 

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5. Investigation of chordwise functionally graded flexural rigidity in flapping wings using a two-dimensional pitch–plunge model

   https://doi.org/10.1088/1748-3190/ac8f05  

   Reade, Joseph; Jankauski, Mark (October 2022, Bioinspiration & Biomimetics)

   Abstract Insect wings are heterogeneous structures, with flexural rigidity varying one to two orders of magnitude over the wing surface. This heterogeneity influences the deformation the flapping wing experiences during flight. However, it is not well understood how this flexural rigidity gradient affects wing performance. Here, we develop a simplified 2D model of a flapping wing as a pitching, plunging airfoil using the assumed mode method and unsteady vortex lattice method to model the structural and fluid dynamics, respectively. We conduct parameter studies to explore how variable flexural rigidity affects mean lift production, power consumption and the forces required to flap the wing. We find that there is an optimal flexural rigidity distribution that maximizes lift production; this distribution generally corresponds to a 3:1 ratio between the wing’s flapping and natural frequencies, though the ratio is sensitive to flapping kinematics. For hovering flight, the optimized flexible wing produces 20% more lift and requires 15% less power compared to a rigid wing but needs 20% higher forces to flap. Even when flapping kinematics deviate from those observed during hover, the flexible wing outperforms the rigid wing in terms of aerodynamic force generation and power across a wide range of flexural rigidity gradients. Peak force requirements and power consumption are inversely proportional with respect to flexural rigidity gradient, which may present a trade-off between insect muscle size and energy storage requirements. The model developed in this work can be used to efficiently investigate other spatially variant morphological or material wing features moving forward. 

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