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Smart wing rotation and trailing-edge vortices enable high frequency mosquito flight

Nature volume 544, pages 9295 (06 April 2017) | Download Citation

This article has been updated


Mosquitoes exhibit unusual wing kinematics; their long, slender wings flap at remarkably high frequencies for their size (>800 Hz)and with lower stroke amplitudes than any other insect group1. This shifts weight support away from the translation-dominated, aerodynamic mechanisms used by most insects2, as well as by helicopters and aeroplanes, towards poorly understood rotational mechanisms that occur when pitching at the end of each half-stroke. Here we report free-flight mosquito wing kinematics, solve the full Navier–Stokes equations using computational fluid dynamics with overset grids, and validate our results with in vivo flow measurements. We show that, although mosquitoes use familiar separated flow patterns, much of the aerodynamic force that supports their weight is generated in a manner unlike any previously described for a flying animal. There are three key features: leading-edge vortices (a well-known mechanism that appears to be almost ubiquitous in insect flight), trailing-edge vortices caused by a form of wake capture at stroke reversal, and rotational drag. The two new elements are largely independent of the wing velocity, instead relying on rapid changes in the pitch angle (wing rotation) at the end of each half-stroke, and they are therefore relatively immune to the shallow flapping amplitude. Moreover, these mechanisms are particularly well suited to high aspect ratio mosquito wings.

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  • 05 April 2017

    The y-axis units were corrected in Fig. 3.


  1. 1.

    , , & A role for acoustic distortion in novel rapid frequency modulation behaviour in free-flying male mosquitoes. J. Exp. Biol. 219, 2039–2047 (2016)

  2. 2.

    & Flapping wing aerodynamics: from insects to vertebrates. J. Exp. Biol. 219, 920–932 (2016)

  3. 3.

    , , & Leading-edge vortices in insect flight. Nature 384, 626–630 (1996)

  4. 4.

    & Dragonfly flight: novel uses of unsteady separated flows. Science 228, 1326–1329 (1985)

  5. 5.

    , , , & Visualising the flow around insect wings. Phys. Fluids 14, S4 (2002)

  6. 6.

    , , , & Dragonfly flight: free-flight and tethered flow visualizations reveal a diverse array of unsteady lift-generating mechanisms, controlled primarily via angle of attack. J. Exp. Biol. 207, 4299–4323 (2004)

  7. 7.

    , & Smoke visualization of free-flying bumblebees indicates independent leading-edge vortices on each wing pair. Exp. Fluids 46, 811–821 (2009)

  8. 8.

    & Unconventional lift-generating mechanisms in free-flying butterflies. Nature 420, 660–664 (2002)

  9. 9.

    , & Leading-edge vortex lifts swifts. Science 306, 1960–1962 (2004)

  10. 10.

    , & Aerodynamics of the hovering hummingbird. Nature 435, 1094–1097 (2005)

  11. 11.

    et al. Bat flight generates complex aerodynamic tracks. Science 316, 894–897 (2007)

  12. 12.

    , & A CFD-informed quasi-steady model of flapping wing aerodynamics. J. Fluid Mech. 783, 323–343 (2015)

  13. 13.

    & Aeromechanics of passive rotation in flapping flight. J. Fluid Mech. 660, 197–220 (2010)

  14. 14.

    The aerodynamics of hovering insect flight. V.A vortex theory. Phil. Trans. R. Soc. Lond. B 305, 115–144 (1984)

  15. 15.

    , & Wing rotation and the aerodynamic basis of insect flight. Science 284, 1954–1960 (1999)

  16. 16.

    & The aerodynamic effects of wing rotation and a revised quasi-steady model of flapping flight. J. Exp. Biol. 205, 1087–1096 (2002)

  17. 17.

    , & Near- and far-field aerodynamics in insect hovering flight: an integrated computational study. J. Exp. Biol. 211, 239–257 (2008)

  18. 18.

    , & The aerodynamics of hovering flight in Drosophila. J. Exp. Biol. 208, 2303–2318 (2005)

  19. 19.

    , , & Harmonic convergence in the love songs of the dengue vector mosquito. Science 323, 1077–1079 (2009)

  20. 20.

    ., ., ., & Short-amplitude high-frequency wing strokes determine the aerodynamics of honeybee flight. Proc. Natl Acad. Sci. USA 102, 18213–18218 (2005)

  21. 21.

    & The mechanics of flight in the hawkmoth Manduca sexta. I. Kinematics of hovering and forward flight. J. Exp. Biol. 200, 2705–2722 (1997)

  22. 22.

    Integrated modeling of insect flight: From morphology, kinematics to aerodynamics. J. Comput. Phys. 228, 439–459 (2009)

  23. 23.

    & The effects of artificial wing wear on the flight capacity of the honey bee Apis mellifera. J. Insect Physiol. 65, 27–36 (2014)

  24. 24.

    , & The effect of aspect ratio on the leading-edge vortex over an insect-like flapping wing. Bioinspir. Biomim. 10, 056020 (2015)

  25. 25.

    , & Photogrammetric reconstruction of high-resolution surface topographies and deformable wing kinematics of tethered locusts and free-flying hoverflies. J. R. Soc. Interface (2009)

  26. 26.

    , & Operation of the alula as an indicator of gear change in hoverflies. J. R. Soc. Interface (2011)

  27. 27.

    & Paddles and rakes: fluid-flow through bristled appendages of small organisms. J. Theor. Biol. 129, 17–39 (1987)

  28. 28.

    & A fluid-structure interaction model of insect flight with flexible wings. J. Comput. Phys. 231, 1822–1847 (2012)

  29. 29.

    & Energy-minimizing kinematics in hovering insect flight. J. Fluid Mech. 582, 153–168 (2007)

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The authors were supported by the EPSRC (EP/H004025/1 and EP/M003698/1), BBSRC (BB/J001244/1). R.J.B. was supported by an EPSRC Career Acceleration Fellowship. S.M.W. was supported by a Royal Society University Research Fellowship. The work reported in this paper was funded by the Autonomous Systems Underpinning Research (ASUR) programme under the auspices of the Defence Science and Technology Laboratory (Dstl), UK Ministry of Defence. The authors acknowledge useful discussions with I. Russell and G. Gibson, P. Simoes for rearing the mosquitoes, and F. Albert-Davie and M. Inglis for assistance during raw data collection. The authors thank G. Taylor for the loan of four high-speed cameras purchased on European Research Council (ERC) grant 204513, and H. Liu for the permission to use the simulator and surface pressure distribution of the fruit fly wing.

Author information


  1. Structure and Motion Laboratory, Royal Veterinary College, University of London, Hatfield AL9 7TA, UK

    • Richard J. Bomphrey
    • , Toshiyuki Nakata
    •  & Nathan Phillips
  2. Graduate School of Engineering, Chiba University, 1-33, Yayoi-cho, Inage-ku, Chiba-shi, Chiba 263-8522 Japan

    • Toshiyuki Nakata
  3. Department of Zoology, University of Oxford, Oxford OX1 3PS, UK

    • Simon M. Walker


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R.J.B. and S.M.W. conceived the experimental design; N.P. and S.M.W. designed and constructed the apparatus and N.P. led the data collection; all authors contributed to data collection; S.M.W. processed the raw data to extract detailed kinematics; T.N. performed the CFD simulations; N.P., T.N. and R.J.B. collected and processed the PIV data; R.J.B. drafted the manuscript; all authors contributed to data interpretation and manuscript preparation.

Competing interests

The authors declare no competing financial interests.

Corresponding author

Correspondence to Richard J. Bomphrey.

Reviewer Information Nature thanks S. Swartz and the other anonymous reviewer(s) for their contribution to the peer review of this work.

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Extended data

Supplementary information


  1. 1.

    Wing stroke cycle

    Video showing the experimental apparatus, raw data, wing geometry routine, kinematics, vortex wake (using isosurfaces of the Q-criterion), and pressure distribution and instantaneous flow fields at key instants (t1-t5) throughout the wing stroke cycle.

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