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      Flight of the dragonflies and damselflies

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          Abstract

          This work is a synthesis of our current understanding of the mechanics, aerodynamics and visually mediated control of dragonfly and damselfly flight, with the addition of new experimental and computational data in several key areas. These are: the diversity of dragonfly wing morphologies, the aerodynamics of gliding flight, force generation in flapping flight, aerodynamic efficiency, comparative flight performance and pursuit strategies during predatory and territorial flights. New data are set in context by brief reviews covering anatomy at several scales, insect aerodynamics, neuromechanics and behaviour. We achieve a new perspective by means of a diverse range of techniques, including laser-line mapping of wing topographies, computational fluid dynamics simulations of finely detailed wing geometries, quantitative imaging using particle image velocimetry of on-wing and wake flow patterns, classical aerodynamic theory, photography in the field, infrared motion capture and multi-camera optical tracking of free flight trajectories in laboratory environments. Our comprehensive approach enables a novel synthesis of datasets and subfields that integrates many aspects of flight from the neurobiology of the compound eye, through the aeromechanical interface with the surrounding fluid, to flight performance under cruising and higher-energy behavioural modes.

          This article is part of the themed issue ‘Moving in a moving medium: new perspectives on flight’.

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          Most cited references105

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          Wing rotation and the aerodynamic basis of insect flight.

          The enhanced aerodynamic performance of insects results from an interaction of three distinct yet interactive mechanisms: delayed stall, rotational circulation, and wake capture. Delayed stall functions during the translational portions of the stroke, when the wings sweep through the air with a large angle of attack. In contrast, rotational circulation and wake capture generate aerodynamic forces during stroke reversals, when the wings rapidly rotate and change direction. In addition to contributing to the lift required to keep an insect aloft, these two rotational mechanisms provide a potent means by which the animal can modulate the direction and magnitude of flight forces during steering maneuvers. A comprehensive theory incorporating both translational and rotational mechanisms may explain the diverse patterns of wing motion displayed by different species of insects.
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            Leading-edge vortices in insect flight

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              The aerodynamics of insect flight.

              The flight of insects has fascinated physicists and biologists for more than a century. Yet, until recently, researchers were unable to rigorously quantify the complex wing motions of flapping insects or measure the forces and flows around their wings. However, recent developments in high-speed videography and tools for computational and mechanical modeling have allowed researchers to make rapid progress in advancing our understanding of insect flight. These mechanical and computational fluid dynamic models, combined with modern flow visualization techniques, have revealed that the fluid dynamic phenomena underlying flapping flight are different from those of non-flapping, 2-D wings on which most previous models were based. In particular, even at high angles of attack, a prominent leading edge vortex remains stably attached on the insect wing and does not shed into an unsteady wake, as would be expected from non-flapping 2-D wings. Its presence greatly enhances the forces generated by the wing, thus enabling insects to hover or maneuver. In addition, flight forces are further enhanced by other mechanisms acting during changes in angle of attack, especially at stroke reversal, the mutual interaction of the two wings at dorsal stroke reversal or wing-wake interactions following stroke reversal. This progress has enabled the development of simple analytical and empirical models that allow us to calculate the instantaneous forces on flapping insect wings more accurately than was previously possible. It also promises to foster new and exciting multi-disciplinary collaborations between physicists who seek to explain the phenomenology, biologists who seek to understand its relevance to insect physiology and evolution, and engineers who are inspired to build micro-robotic insects using these principles. This review covers the basic physical principles underlying flapping flight in insects, results of recent experiments concerning the aerodynamics of insect flight, as well as the different approaches used to model these phenomena.
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                Author and article information

                Journal
                Philos Trans R Soc Lond B Biol Sci
                Philos. Trans. R. Soc. Lond., B, Biol. Sci
                RSTB
                royptb
                Philosophical Transactions of the Royal Society B: Biological Sciences
                The Royal Society
                0962-8436
                1471-2970
                26 September 2016
                26 September 2016
                : 371
                : 1704 , Theme issue ‘Moving in a moving medium: new perspectives on flight’ compiled and edited by Steven J. Portugal and Emily L. C. Shepard
                : 20150389
                Affiliations
                [1 ]Structure and Motion Laboratory, Department of Comparative Biomedical Sciences, Royal Veterinary College , North Mymms, Hatfield AL9 7TA, UK
                [2 ]Graduate School of Engineering, Chiba University , 1-33, Yayoi-cho, Inage-ku, Chiba-shi, Chiba 263-8522, Japan
                [3 ]Department of Biology, Lund University , Ecology Building, 223 62 Lund, Sweden
                [4 ]Howard Hughes Medical Institute , Janelia Research Campus, 19700 Helix Drive, Ashburn, VA 20147, USA
                Author notes

                One contribution of 17 to a theme issue ‘ Moving in a moving medium: new perspectives on flight’.

                Author information
                http://orcid.org/0000-0002-4748-0510
                Article
                rstb20150389
                10.1098/rstb.2015.0389
                4992713
                27528779
                c266f1da-3609-4cdf-9381-74dde5073c35
                © 2016 The Authors.

                Published by the Royal Society under the terms of the Creative Commons Attribution License http://creativecommons.org/licenses/by/4.0/, which permits unrestricted use, provided the original author and source are credited.

                History
                : 6 May 2016
                Funding
                Funded by: Vetenskapsr, http://dx.doi.org/10.13039/501100004359;
                Award ID: 2013-4838
                Funded by: Engineering and Physical Sciences Research Council, http://dx.doi.org/10.13039/501100000266;
                Award ID: EP/HH04025/1
                Funded by: Biotechnology and Biological Sciences Research Council, http://dx.doi.org/10.13039/501100000268;
                Award ID: BB/J001244/1
                Categories
                1001
                25
                14
                60
                133
                Articles
                Research Article
                Custom metadata
                September 26, 2016

                Philosophy of science
                odonata,flight,biomechanics,aerodynamics,prey capture,performance
                Philosophy of science
                odonata, flight, biomechanics, aerodynamics, prey capture, performance

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