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References on Dragonfly Flight & Biomechanics

A Web Page by Roy J. Beckemeyer

Last Updated 12 May 2006                                          

Annotations and comments are the opinions of Roy Beckemeyer and are provided for anyone beginning the study of flight in dragonflies.

  1. Alexander, D.E., 1982, Studies on flight control and aerodynamics in dragonflies, PhD Dissertation, Duke University [One of the few studies of flight control and maneuvering.  See also the next two references and the works by Ruppell.]
  2. Alexander, D.E., 1984, Unusual phase relationships between the forewings and hindwings in flying dragonflies, J. Exp. Biol., 109:379-383
  3. Alexander, D.E., 1986, Wind tunnel studies of turns by flying dragonflies, J. Exp. Biol., 122:81-98
  4. Anholt, B.R., J.H. Marden, & D.M. Jenkins, 1991, Patterns of mass gain and sexual dimorphism in adult dragonflies (Insecta: Odonata), Can. J. Zool., 69:1156-1163  [One of the first studies to investigate sexual dimorphism in Odonata from a functional morphology and ecological/behavioral point of view.  See other works by Marden and his colleagues as well.]
  5. Arnold, J.W., 1964, Blood circulation in insect wings, Mem. Entomol. Soc. Canada, No. 38
  6. Azuma, A.  2006. The Biokinetics of Flying abnd Swimming.  Second Edition. American Institute of Aeronautics and Astronautics.  Blackberg, VA.  xxxvii + 518 pp.
  7. Azuma, A., S. Azuma, I. Watanabe, & T. Furuta, 1985, Flight mechanics of a dragonfly, J. Exp. Biol., 116:79-107 [Application of an unsteady aerodynamics approach, the local circulation method, to compute air forces on dragonfly wings in flapping flight.  Includes interaction with the shed wake. Also see the next paper, Sato & Azuma, 1997,  and Okamoto et al., 1996]
  8. Azuma, A., M. Okamoto, & K. Yasuda, 2001, Aerodynamic characteristics of wings at low Reynolds Number, pp. 341-398 (Ch. 17) In Mueller, T.J. (Ed.), Fixed and flapping wing aerodynamics for Micro Air Vehicle applications, Vo. 195, Progress in Astronautics and Aeronautics, American Institute of Aeronautics and Astronautics, Reston, Virginia
  9. Azuma, A., & T. Watanabe, 1988, Flight performance of a dragonfly, J. Exp. Biol., 137:221-252
  10. Beckemeyer, R. J., 1999, Measurements of total fresh mass for some species of Odonata from the Great Plains of the United States, Notulae odonatologicae, 5(3):35-36
  11. Beckemeyer, R.J., 2004, Notes on the behavior and mechanics of scooping oviposition in Libellula composita (Hagen) (Anisoptera: Libellulidae), Odonatologica,  33(1):11-23.
  12. Beckemeyer, R.J., 2004, Measurements of total fresh mass for some species of Odonata from Kansas and Missouri, United States,, Notulae odonatologicae, 6(3):33-34.
  13. Bernáth, B.,  J. Gál, & G Horváth, 2004, Why is it worth flying at dusk for aquatic insects? Polarotactic water detection easiest at low solar elevation angles, J. Exp. Biol., 207:755-765 [Not really a flight mechanics paper, but one on polarization, that sheds some light (sorry) on crepuscular flying.]
  14. Birch, J.M., W.B. Dickinson, & M.H. Dickinson.  2004.  Force production and flow structure of the leading edge vortex on flapping wings at high and low Reynolds numbers, J. Exp. Biol., 207(7):1063-1072.
  15. Bried, J.T., L.W. Bennett & G.N. Ervin.  2005.  Live mass and length - mass allometry of adult odonates collected in east-central Mississippi, United States.  Odonatologica, 34(2):111-122.
  16. Brodsky, A.K., 1994, The evolution of flight, Oxford Science Publications, 229 pp.  [A functional-morphological approach to the study of insect evolution.  Scattered references to Odonata throughout, but especially see Section 6.3.]
  17. Burnside, C.A., & J.V. Robinson, 1995, The functional morphology of caudal lamellae in coenagrionid (Odonata: Zygoptera) damselfly larvae, Zool. J. Linn. Soc., 114:155-171
  18. Chadwick, L.E., 1940, The wing motion of the dragonfly, Bull. Brooklyn. Ent. Soc., 35:109-112  [Probably the first accurate depiction of the kinematics of wing motion in Odonata.  Concise, and a minor classic.]
  19. Clark, H.W., 1940, The adult musculature of the anisopterous dragonfly thorax (Odonata, Anisoptera), J. Morphology, 67:523-565
  20. Coelho, J.R., 1997, Sexual size dimorphism and flight behavior in cicada killers, Sphecius speciosus, Oikos, 79:371-375 [Ok, not dragonflies, but definitely a paper of interest dealing as it does with size dimorphism and flight in an insect that carries other dead insects around - search on the author's name for other similar papers]
  21. Coelho, J.R., & J. Hoagland, 1995, Load-lifting capacities of three species of yellowjackets (Vespula) foraging on honey-bee corpses, Funcitonal ecology, 9:171-174
  22. Combes, S.A., and T. L. Daniel, 2003, Flexural stiffness in insect wings.  I. Scaling and the influence of wing venation.  Journal of Experimental Biology, 206(17):2679-2987 [Covers a number of insect taxa, including some Odonata.  The third paper by these authors, although it deals with a moth, is a natural continuation of their research and contains information applicable as well to Odonata.]
  23. Combes, S.A., and T. L. Daniel, 2003, Flexural stiffness in insect wings.  II. Spatial distribution and dynamic wing bending.  Journal of Experimental Biology, 206(17):2989-2997
  24. Combes, S.A., and T. L. Daniel, 2003, Into thin air: contributions of aerodynamic and inertial-elastic forces in wing bending oin the hawkmoth Manduca sexta,  Journal of Experimental Biology, 206(17):2999-3006
  25. Davies, D.A.L., 2003, How did Pterosaur soar?, Notulae odonatologicae, 6(2):13-14
  26. deEguileor, M., R. Valvassori, & G. Lanzsavecchia, 1980, Discontinuity of sarcoplasmic reticulum in the mid-sarcomere region in flight muscle of dragonflies, Tissue and Cell, 12(4):749-759
  27. Dickinson, M.H., & K.G. Gőtz, 2993, Unsteady performance of model wings at low Reynolds numbers, J. Exp. Biol., 174:45-64 [One of a series of important papers by Dickinson and associates dealing with the mechanics of lift generation in pronating and supinating insect wings.  He works mainly with Diptera: Drosophila, but many of the observations can be extrapolated or applied somewhat directly to the mechanics of Odonata flight.  See the next paper in particular.]
  28. Dickinson, M.H., F.-O. Lehamnn, & S.P. Sane, 1999, Wing rotation and the aerodynamic basis of insect flight, Science, 284:1954-1960
  29. Dillon, M.E., & R. Dudley, 2004, Allometry of maximum vertical force production during hovering flight of neotropical orchid bees (Apidae: Euglossini), Journal of Experimental Biology, 207(3):417-425 [Not dragonflies, but allometry, max force production, and techniques that are applicable to dragonflies and damselflies.]
  30. Dudley, R., 2000, The biomechanics of insect flight: form, function, evolution, Princeton Univ. Press, Princeton, N.J.  [Comprehensive and definitive summary of the state of the art through the late 1990's.  Indispensable.]
  31. Dunham, M., 1993, Changes in mass, fat content, and water content with growth in adult Pachydiplax longipennis (Odonata: Libellulidae), Can. J. Zool., 71:1470-1473
  32. Edgar, P.K., & J.R. Coelho, 2000, Load-lifting constraints on provisioning and nest building in the carpenter wasp, Monobia quadriens L. (Hymenoptera: Eumenidae), J. Hymen. Res., 9(2):370-376
  33. Ennos, A.R., 1988, The importance of torsion in the design of insect wings, J. Exp. Biol., 140:137-160
  34. Fitzhugh, G. H. and Marden, J. H., 1997,  Maturational changes in troponin T expression, Ca2+-sensitivity and twitch contraction kinetics in dragonfly flight muscle,  200:1473-1482
  35. Fried, C.S., & M.L. May, 1983, Energy expenditure and food intake of territorial male Pachydiplax longipennis (Odonata: libelllidae), Ecological Entomology, 8:283-292
  36. Frye, M.A., & R.M. Olberg, 1995, Visual receptive field properties of feature detecting neurons in the dragonfly, J. Comp. Physiol. A, 177:569-576
  37. Gibo, D.L., 1981, Some observations on slope soaring in Pantala flavescens (Odonata: Libellulidae), J. N.Y. Ent. Soc., LXXXIX(3):184-187
  38. Gorb, S., The skeleton-muscle organization of the head fixation system in odoantes and its evolutionary implications: a comparative study, Petalura, 1(1):1-17
  39. Gorb, S.N., Serial elastic elements in the damselfly wing: mobile vein joints contain resilin, Naturwissenschaften, 86:552-555 [Shows that the flexible veins in damselfly wings, first noted by Newman in 1982, contain a rubber-like protein, resilin. ] 
  40. Gorb, S., 2001, Attachment devices of insect cuticle, Kluwer Academic Pub., 305 pp.
  41. Grabow, K., & G. Rüppell, 1995, Wing loading in relation to size and flight characteristics of European Odonata, Odonatologica, 24(2):187-218
  42. Grandi, M., 1947, Gli scleriti ascellari degli Odonati, loro morfologia e miologia comparate, Boll. de..'Istituto di Entomologia della Univ. di Bologna, 41:254-278 [A study of the axillary sclerite morphology and myology in some Odonata.]
  43. Grodnitsky, D. L., 1999, Form and function of insect wings: the evolution of biological structures, Johns Hopkins University Press, Baltimore and London, 261 pp.  [Somewhat confusing at first in taxonomic terms, as it uses and promotes the Russian school's (A. P. Rasnitsyn's) view of insect systematics. Material on Odonata (but see "Libellulida" in the book's index) scattered through the book.]
  44. Hamdani, H., & M. Sun, 2000, Aerodynamic forces and flow structures of an airfoil in some unsteady motions at small Reynolds Numbers, Acta Mechanica, 145:173-187 [A series of papers - see also Lan & Sun and Hamdani & Sun - that use implicit numerical algorithms to solve the Navier-Stokes equations for large scale motions of airfoils at low Reynolds Numbers, in attempts to model the experimental results of Dickinson et al for Diptera, and of Luttges for Odonata.  Reprints obtained from Dr. Sun Mao.]
  45. Hardy, H.T., Jr., 1965, The effect of sunlight and temperature on theposture of Perithemis tenera (Odonata), Proc. Oklahoma Acad. Sci. for 1965: 41-45
  46. Harrison, J. F. and Lighton, J. R. B., 1998,  Oxygen-sensitive flight metabolism in the dragonfly Erythemis simplicicollis,  201:1739-1744
  47. Hass, F., S. Gorb, & R.J. Wootton.  2001.  Elastic joints in dermapteran hind wings: materials and wing folding.  Arthropod Structure and Devleopment.  29:137-146. [OK, it's not Odonata, but I put it here because it relates to elastic joints in wings, which are present in Odonata.]
  48. Hatch, G., 1966, Structure and mechanics of the dragonfly pterothorax, Annals, Entomol. Soc. America, 59(4):702-714
  49. Heinrich, B., & T.M. Casey, 1978, Heat transfer in dragonflies: 'fliers' and 'perchers', J. Exp. Biol., 74:17-36
  50. Ishizawa, N., 1988, Morphological differences in a dragonfly, Sympetrum frequens Selys, with relation to the centre of gravity, Yosegaki, 51:856-863 [In Japanese, with English summary]  [To my knowledge, the first author to note the difference in ratio of hind wing to forewing area in female and male Odonata and the associated difference in center of mass.]
  51. Ishizawa, N., 1994, Thermoregulation of dragonflies of Sympetrum species in the lowlands in midsummer, Gekkan-Mushi, 281:13-17
  52. Ishizawa, N., 1995, Observations on migrating Sympetrum frequens Selys (Libellulidae, Odonata), Gekkan-Mushi, 293:22-27
  53. Ishizawa, N., 1996, Thermoregulation in dragonflies of Sympetrum, Nature & Insects, 31(8):18-22
  54. Ishizawa, N., 1998, Thermoregulation in Sympetrum frequens (Selys), with notes on other Sympetrum species (Anisoptera: Libellulidae), Odonatologica, 27(3):317-334
  55. Jurzitza, G., 1973, Zwei Flugtypen bei Paarungsradern von Libellen, Odonatologica, 2(4):329-332
  56. Kesel, A.B., 2000, Aerodynamic characteristics of dragonfly wing sections compared with technical airfoils, J. Exp. Biol., 203:3125-3135  [Functioning of the corrugated airfoils of dragonfly wings is studied at near real Reynolds numbers (8,000-10,000), and she shows that angles of attack of greater than 0ş are required for the negative pressure in the corrugation valleys on both the dorsal and ventral sides of the wing to yield a net negative pressure on the dorsal side.  For earlier work on aerodynamics of corrugated wings, see Newman et al, 1977, Rudolph, 1978, and Okamoto et al, 1996.]
  57. Kesel, A.B., U. Philippi, & W. Nachtigall, 1998, Biomechanical aspects of the insect wing: an analysis using the finite element method, Computers in Biology and Medicine, 28:423-437  [One of the first finite element models of a dragonfly wing - Aeshnidae: Aeshna cyanea.]
  58. Kliss, M., C. Somps, & M.W. Luttges, 1989, Stable vortex structures: a flat plate model of dragonfly hovering, J. theor. Biol., 136:209-228 
  59. Kreuz, P., A.B. Kesel, H. Vehoff, A. Fery, & W. Arnold, 1999, The insect wing membrane - a challenge for materials science, http://www.medizin.uni-halle.de/biomechanik/acas99/abstracts/kreuz.htm (Downloaded Sept. 2002).
  60. Lan, S., & M. Sun, 2001, Aerodynamic force and flow structures of two airfoils in flapping motions, Acta Mechanica Sinica (English Series), 17(4):310-331
  61. Lan, S., & M. Sun, 2001, Aerodynamic properties of a wing performing unsteady rotational motions at low Reynolds Number, Acta Mechanica, 149:145-147
  62. MacNeil, N., 1968, On the possible function of the pterostigma, The Entomologist, pp. 60-63
  63. Marden, J.H., 1987, Maximum lift production during takeoff in flying animals, J. Exp. Biol., 130:235-258
  64. Marden, J.H., 1989, Bodybuilding dragonflies: costs and benefits of maximizing flight muscle, Physiol. Zool., 62(2):505-521  [A very important work showing the growth in weight of male and female Libellulidae: Plathemis lydia, and the respective investment in either abdomen (female) or flight muscle (male) mass with maturity.]
  65. Marden, J.H., 1995, Large-scale changes in thermal sensitivity of flight performance during adult maturation in a dragonfly, J. Exp. Biol., 198:2095-2102
  66. Marden, J.H., G.H. Fitzhugh, & M.R. Wolf, 1998, From molecules to mating success: integrative biology of muscle maturation in a dragonfly, Amer. Zool., 38:528-544  [What appears to be a landmark paper in integrating ecological, organismal, and molecular biology into a study of dragonfly flight. ]
  67. Marden, J.H., M.G. Kramer, & J. Frisch, 1996, Age-related variation in body temperature, thermoregulation and activity in a thermally polymorphic dragonfly, J. Exp. Biol., 199:529-535
  68. Marden, J.H., & R.A. Rollins, 1994, Assessment of energy reservesw by damselflies engaged in aerial contests for mating territories, Anim. Behav., 48:1023-1030
  69. Marden, J.H, & J.K. Waage., 1990, Escalated damselfly territorial contests are energetic wars of attrition, Anim. Behav., 39:954-959
  70. May, M.L., 1976, Thermoregulation and adaptation to temperature in dragonflies (Odonata: Anisoptera), Ecol. Monog., 46:1-32 [An incredible body of work related to biomechanics and energetics in Odonata that is required reading.  An excellent way to prepare for a program of research in thermoregulation or energetics would be to work through all of May's papers.]
  71. May, M.L., 1976, Warming rates as a function of body size in periodic endotherms, J. Comp. Physiol., 111:55-70
  72. May, M.L., 1978, Thermal adaptations of dragonflies, Odonatologica, 7(1):27-47
  73. May, M.L., 1979, Energy metabolism of dragonflies (Odonata: Anisoptera) at rest and during endothermic warm-up, J. Exp. Biol., 83:79-94
  74. May, M.L., 1981, Wingstroke frequency of dragonflies (Odonata: Anisoptera) in relation of temperature and body size, J. Comp. Physiol., 144:229-240
  75. May, M.L., 1981, Allometric analysis of body and wing dimensions of male Anisoptera, Odonatologica, 10(4):279-291
  76. May, M.L., 1982, Heat exchange and endothermy in Protodonata, Evolution, 36(5):1051-1058
  77. May, M.L., 1984, Energetics of adult Anisoptera, with special reference to feeding and reproductive behavior, Adv. Odonatol., 2:95-116
  78. May, M.., 1986, A preliminary investigation of variation in temperature among body regions of Anax junius (Drury) (Anisoptera: Aeshnidae), Odonatologica, 15(1):119-128
  79. May, M.L., 1991, Dragonfly flight: power requirements at high speed and acceleration, J. Exp. Biol., 158:325-342 [A particularly nice combination of field observations and measurements with aerodynamic performance theory.  He determines that dragonflies apparently "routinely, albeit for short intervals, operate near their maximum sustainable power output during flight".]
  80. May, M.L., 1995, Simultaneous control of head and thoracic temperatures by the Green Darner dragonfly Anax junius (Odonata: Aeshnidae), J. Exp. Biol., 198:2373-2384
  81. May, M.L., 1995, Dependence of flight behavior and heat production on air temperature in the Green Darner dragonfly Anax junius (Odonata: Aeshnidae), J. Exp. Biol., 198:2385-2392
  82. Miller, P.L., 1962, Spiracle control in adult dragonflies (Odonata), J. Exp. Biol., 39:513-535
  83. Mittelstaedt, H., 1950, Physiologie des Gleichgewichtssins bei Fliegenden Libellen, Zeitschrift fur vergleichende Physiologie, 32:422-463 [Aspects of the flight control system in Odonata.  A ladmark paper.]
  84. Misutani, A., J.S. Chahl, & M.V. Srinivasan, 2003, Motion camouflage in dragonflies, Nature, 423:604
  85. Nachtigall, W.,  2003, Insektenflug: Konstruktionsmorphologie, Biomechanik, Flugverhalten, Springer, Berlin, xiii+482 pp.  [Nachtigall's magnum opus, a treatment of the whole of insect flight.  Magnificent, but get out the old German scientific dictionary and grammar.]
  86. Needham, J.G., & M.H. Anthony, 1903, The skewness of the thorax in the Odonata, J. New York Entomol. Soc. 11(3):117-125 + plate 8  
  87. Neville, A.C., 1960, Aspects of flight mechanics in anisopterous dragonflies, J. Exp. Biol., 37:631-656  [Details of the flapping mechanics in Anisoptera, with microscopic looks at the functioning of the muscles and axillary sclerites.]
  88. Newman, B.G., S.B. Savage, & D. Schouella, 1977, Model tests on a wing section of an Aeschna Dragonfly, pp.445-477 in Pedley, T.J., Editor, Scale effects in animal locomotion, Academic Press, London
  89. Newman, D.J.S.,  1982, The functional wing morphology of some Odonata, PhD Thesis, University of Exeter [An important and groundbreaking study that covers nearly every aspect of structural/mechanical importance in flight - center of mass, wing construction including various vein intersection types, wing kinematics, etc.  It is too bad that only a few short excerpts from this work ever saw publication in the open literature (see next two references).  Invaluable!]
  90. Newman, D.J.S., & R.J. Wooton, 1986, An approach to the mechanics of pleating in dragonfly wings, J. Exp. Biol., 125:361-372
  91. Newman, D.J.S., & R.J. Wooton, 1988, The role of the fulcroalar muscle in dragonfly flight, Odonatologica, 17(4):401-408
  92. Norberg, R.A., 1972, The pterostigma of insect wings as an inertial regulator of wing pitch, J. Comp. Physiol., 81:9-22
  93. Norberg, R.A., 1975, Hovering flight of the dragonfly Aeschna juncea L., kinematics and aerodynamics, pp. 763-781 in Wu, T. Y.-T., C.J. Brokaw, & C. Brennen, Editors, Swimming and flying in nature. Vol. 2, Plenum Press, NY
  94. Okamoto, M., K. Yasuda, & A. Azuma, 1996, Aerodynamic characteristics of the wings and body of a dragonfly, J. Exp. Biol., 199:281-294
  95. Olberg, R.M., A.H. Worthington, & K.R. Venator, 2000, Prey pursuit and interception in dragonflies, J. Comp. Physiol. A, 186: 155-162
  96. Olesen, J., 1972, The hydraulic mechanism of labial extension and jet propulsion in dragonfly nymphs, J. Comp. Physiol., 8153-55
  97. Peckham, M., & D.C.S. White, 1991, Mechanical properties of demembrenated flight muscle fibres from a dragonfly, J. Exp. Biol., 159:135-147
  98. Pfau, H.K., 1986, Untersuchungen zur Konstruktion, Funktion und Evolution des Flugapparates der Libellen (Insecta, Odonata), Tijdschrift voor Entomologie, 129(3):35-123  [Another magnum opus on functional morphology by Pfau.  He also studied the genitalia of Odonata.  Not easy going for non-German readers, but a very important work that also reviews the work of others who have studied the mechanics of muscle and sclerite functioning in wing flapping.  Pages 118-120 contain a short English summary of the results of the paper.]
  99. Pfau, H.K.  2000.  Erasipteron larischi Pruvost, 1933, Eugeropteron lunatum Riek, 1984 und die Evoultion der Verstellpropeller-Flugel der Libellen.  Mitteilungen der Schweizerischen Entomologischen Gesellschaft. Bulletin de la Societe Entomologique Suisse.  73:223-263. [Functional morphology in the earliest known Carboniferous dragonflies.]
  100. Pringle, J.W.S., 1957, Insect flight, Cambridge University Press, Cambridge  [An excellent primer on insect flight (but certainly not simplistic).  Includes a couple of pages that briefly recount some of Mittelstaedt's important work on the head/prothorax system as a stabilization mechanism in Odonata.]
  101. Pritchard, G., 1986, The operation of the labium in larval dragonflies, Odonatologica, 15(4):451-456
  102. Reavis, M.A., & M.W. Luttges, 1988, Aerodynamic forces produced by a dragonfly, Paper No. AIAA-88-0330, Am. Inst. Aeronautics & Astronautics
  103. Riek, E.F., & J. Kukalova-Peck, 1984, A new interpretation of dragonfly wing venation based upon Early Upper Carboniferous fossils from Argentina (Insecta: Odonatoidea) and basic character states in pterygote wings, Can. J. Zool., 62:1150-1166
  104. Roberts, S.P., J.F. Harrison, & R. Dudley.  2004.  Allometry of kinematics and energetics in carpenter bees (Xylocopa varipuncta) hovering in variable-density gasses.  Journal of Experimental Biology.  207(6):993-1004.
  105. Rudolph, R., 1976, Preflight behaviour and the initiation of flight in tethered and unrestrained dragonfly, Calopteryx splendens (Harris) (Zygoptera: Calopterygidae), Odonatologica, 5(1):59-64
  106. Rudolph, R., 1976, Some aspects of wing kinematics in Calopteryx splendens (Harris) (Zygoptera: Calopterygidae), Odonatologica, 5(2):119-127
  107. Rudolph, R., 1976, Die aerodynamische Eigenschaften von Calopteryx splendens (Harris) (Zygoptera: Calopterygidae), Odonatologica, 5(4):383-386
  108. Rudolph, R., 1978, Aerodnamic properties of Libellula quadrimaculata L. (Anisoptera: Libellulidae), and the flow around smooth and corrugated wing section models during gliding flight, Odonatologica, 7(1):49-58
  109. Ruffieux, L., J.-M. Elouard, & M. Sartori.  1998. Flightlessness in mayflies and its relevence to hypotheses on the origin of insect flight.  Proceedings of the Royal Society of London.   B.  265:2135-2140.
  110. Rüppell, G., 1985, Kinematic and behavioural aspects of flight of the male Banded Agrion, Calopteryx (Agrion) splendens L., pp. 195-204 in Gewecke, M., & G. Wendler,Editors, Insect locomotion, Paul Parey, Berlin [Rüppell is a master of flight photography and kinematic analysis of free flight in Odonata. The following (1989) paper is particularly important, as it summarizes the major flight styles of the major groups of Odonata. A suite of papers that deserve and require careful study on the part of serious students of dragonfly flight.]
  111. Rüppell, G., 1989, Kinematic analysis of symmetrical flight manoeuvers of Odonata, J. Exp. Biol., 144:13-42
  112. Rüppell, G., & D. Hilfert, 1993, The flight of the relict dragonfly Epiophlebia superstes (Selys) in comparison with that of modern Odonata (Anisozygoptera: Epiophlebiidae), Odonatologica, 22(3):295-309
  113. Rüppell, G., & D. Hilfert: Videotapes of films of Calopterygidae in flight: "Jewelwings" (49 minutes) and "Life history of C. haemorrhoidalis" (20 minutes).  [Courtesy of Dr. Rüppell, these are marvelous films, with more to see each time they are viewed.]
  114. Saharon, D., & M.W. Luttges, 1988, Visualization of unsteady separated flow produces by mechanically driven dragonfly wing kinematics model, Paper No. AIAA-88-0569, Am. Inst. Aeronautics & Astronautics
  115. Saharon, D., & M.W. Luttges, 1989, Dragonfly unsteady aerodynamics: The role of the wing phase relationships in controlling the produced flows, Paper No. AIAA-89-0832, Am. Inst. Aeronautics & Astronautics
  116. Sane, S.P., 2003, Review: The aerodynamics of insect flight,  Journal of Experimental Biology, 206:4191-4208. [The most recent and up-to-date summary of insect aerodynamics.]
  117. Sargent, W.D., 1937, The internal thoracic skeleton of the dragonflies (Odonata: Suborder Anisoptera), Annals Entomol. Soc. America, 30:81-93 + plates 1,2
  118. Sato, M., & A. Azuma, 1997, The flight performance of a damselfly Ceriagrion melanurum Selys, J. Exp. Biol., 200:1765-1779
  119. Savage, S.B., B.G. Newman, & D. T.-M. Wong, 1979, The role of vortices and unsteady effects during the hovering flight of dragonflies, J. Exp. Biol., 83:59-77
  120. Schilder, R.J. & J. H. Marden, 2004, A hierarchical analysis of the scaling of force and power production by dragonfly flight motors, J. Exp. Biol., 207:767-776
  121. Shelly, T.E., 1982, Comparative behavior of light- versus shade-seeking adult damselflies in alowland Neotropical forest (Odonata: Zygoptera), Physiol. Zool., 55(4):335-343
  122. Simmons, P., 1977, The neuronal control of dragonfly flight.  I. Anatomy, J. Exp. Biol., 71:123-140
  123. Simmons, P., 1977, The neuronal control of dragonfly flight.  II. Physiology, J. Exp. Biol., 71:141-155
  124. Singer, F., 1987, A physiological basis of variation in postcopulatory behaviour in a dragonfly Sympetrum obtrususm, animal Behaviour, 35(5):1575-1577
  125. Smith, M.J.C., P.J. Wilkins, & M.H. Williams, The advantages of an unsteady panel method in modeling the aerodynamic forces on rigid flapping wings,  1996, J. Exp. Biol., 199:1073-1083
  126. Snodgrass, R.E., 1909, The thorax of insects and the articulation of the wings, No. 1687, Proceedings of the U.S. National Museum, Vol. 36:511-595, Plates 40-69 [Includes material on Odonata.]
  127. Somps, C., & M. Luttges, 1985, Dragonfly flight: novel uses of unsteady separated flows, Nature, 228:1326-1329 [Also see Kliss et al.  These investigators studied the use of unsteady vortex interactions in dragonfly flight, mainly the interaction of shed vortices and the wing surfaces.]
  128. Sotavalta, O., 1954, On the thoracic temperature of insects in flight, Annales Zoologici Societatis Zoologicae Botanicae Fennicae 'Vanamo', 16(8):1-21
  129. Song, D., H. Wang, L. Zeng, & C. Yin, 2001, Measuring the camber deformation of a dragonfly wing using projected comb fringe, Review of Scientific Instruments, 72(5):2450-2454 [One of a series of articles about non-contact methods of measuring wing deformation patterns.  Thanks to Dr. Zeng for providing me with reprints of her papers.  See also entries under Zeng.]
  130. Srygley, R.B., & R. Dudley.  174.  Correlations of the position of center of body mass with butterfly escape tactics.  Journal of Experimental Biology.  174:155-166.
  131. Strickland, E.H., 1946, Adult Odonata as classroom material, Annals, Entomol. Soc. America, 39:28-32 [The first one to note that the center of mass in Odonata is behind the wing bases (see also D.J.S. Newman's thesis).]
  132. Sturtevant, M.A., & E. Bier.  1995. Analysis of the genetic hierarchy guiding wing vein development in Drosophila.  Development.  121:785-801.  [Although dealing with Diptera, the paper sheds light on the genetic control of wing vein development that should have wide application in the Insecta.]
  133. Sun, M., & H. Hamdani, 2001, A study on the mechanism of high-lift generation by an airfoil in unsteady motion at low Reynolds Number, Acta Mechanica Sinica (English Series), 17(2):97-114
  134. Sun, M., & S.L. Lan.  2004.  A computational study of the aerodynamic forces and power requirements of dragonfly (Aeschna juncea) hovering.  Journal of Experimental Biology. 207(11):1887-1901.
  135. Sunada, S., L. Zeng, & K. Kawachi, 1998, The relationship between dragonfly wing structure and torsional deformation, J. Theor. Biol., 193:39-45
  136. Tanaka, Y., & M. Hisada, 1980, The hydraulic mechanism of the predatory strike in dragonfly larvae, J. Exp. Biol., 88:1-19
  137. Taylor, G.K., 2001, Mechanics and aerodynamics of insect flight control, Biol. Rev., 76:449-471
  138. Taylor, G.K., & A. L. R. Thomas.  2002.  Animal flight dynamics. II. Longitudinal stability in flapping flight.  Journal of Theoretical Biology.  214:351-370.
  139. Taylor, G.K., & A. L. R. Thomas. 2003. Corrigendum "Animal flight dynamics. II. Longitudinal stability in flapping flight." [Journal of Theoretical Biology (2002) 214:351-370]. Journal of Theoretical Biology.  221:671.
  140. Taylor, G.K., & A. L. R. Thomas.  2003.  Dynamic flight stability in the desert locust Schistocerca gregaria.  Journal of Experimental Biology.  206:2803-2839.
  141. Thomas, A.L.R., & G. K. Taylor.  2001.  Animal flight dynamics.  I. Stability in gliding flight.  Journal fo Theoretical Biology. 212:399-424.
  142. Tracy, C.R., B.J. Tracy, & D.S. Dobkin, 1979, The role of posturing in behavioral thermoregulation by Black Dragons (Hagenius brevistylus Selys; Odonata), Phyziol. Aool., 565-571
  143. Vogt, F.D., & B. Heinrich, 1983, Thoracic temperature variations in the onset of flight in dragonflies (Odonata: Anisoptera), Physiol. Zool., 56(2):236-241
  144. Wakeling, J.M., 1993, Dragonfly aerodynamics and unsteady mechanisms: a review, Odonatologica, 22(3):319-334
  145. Wakeling, J.M., & C.P. Ellington, 1997, Dragonfly flight.  I. Gliding flight and steady-state aerodyamic forces, J. Exp. Biol., 200:543-556 [A "definitive" look at Odonata flight.  Three parts (see below).]
  146. Wakeling, J.M., & C.P. Ellington, 1997, Dragonfly flight.  II. Velocities, accelerations, and kinematics of flapping flight, J. Exp. Biol., 200:557-582
  147. Wakeling, J.M., & C.P. Ellington, 1997, Dragonfly flight.  III. Lift and power requirements, J. Exp. Biol., 200:583-600
  148. Wang, H.W., L. Zeng, H. Liu, & C. Yin, 2003, Measuring wing kinematics, flight trajectory and body attitude during forward flight and turning maneuvers in dragonflies, J. Exp. Biol., 206(4):745-757 [Uses a photographic technique to track kinematics in free flight.  A very important technique that will yield much important information on insect flight.]
  149. Wang, Z. Jane, 2000, Vortex shedding and frequency selection in flapping flight, J. Fluid Mech., 410:323-341
  150. Wang, Z. Jane, 2000, Two dimensional mechanism for insect hovering, Phys. Rev. Lett., 85(10):2035-2038  [A 2-d Navier-Stokes solver is used to show that even in two dimensions vortex shedding can generate enough lift for hovering flight.  The phase relationship between pitching and plunging motions is critical in properly timing the leading and trailing edge vortex shedding.  Thanks to Dr. Wang for supplying copies of her papers.]
  151. Wang, Z.J., J.M. Birch, & M.H. Dickinson, 2004, Unsteady forces and flows in low Reynolds number hovering flight: two-dimensional computations vs robotic wing experiments, Journal of Experimental Biology, 207(1):449-460 [Important paper - another step along the path to understanding what's going on here!]
  152. Weis-Fogh, T., 1964, Functional design of the tracheal system of flying insects as compared with the avian lung, J. Exp. Biol., 41:207-227  [Both this paper and the following one use dragonflies and desert locusts (Orthoptera) as models.] 
  153. Weis-Fogh, T., 1964, Diffusion in insect wing muscle, the most active tissue known, J. Exp. Biol., 41:229-256
  154. Weis-Fogh, T., 1967, Respiration and tracheal ventilation in locusts and other flying insects, J. Exp. Biol., 47:561-587 [Includes various species of Aeshnidae: Aeshna as experimental animals.]
  155. Whitten, J.M., 1962, Homolgy and development of insect wing tracheae, Annals, Entomol. Soc. America, 288-295 [Even though she does not deal with Odonata in this paper, it is of importance because she refutes the basic argument used by Comstock and Needham in their analysis of the wing venation in Odonata.  Important to an understanding of the homology and evolution of wing venation.]
  156. Wootton, R.J., 1979, Function, homology and terminology in insect wings, Syst. Entomol., 4:81-93 [Wootton, in his work with his students and colleagues, has basically established the whole discipline of interpreting wing structure in terms of flight functional morphology.  Indispensible.]
  157. Wootton, R.J., 1981, Support and deformability in insect wings, J. Zool. Lond., 193:447-468
  158. Wootton, R.J., 1990, The mechanical design of insect wings, Scientific American, Nov. 1990: 114-120
  159. Wootton, R.J., 1991, The functional morphology of the wings of Odonata, Adv. in Odonatology, 5:153-169
  160. Wootton, R.J., 1992, Functional morphology of insect wings, Ann. Rev. Entomol. 37:113-140
  161. Wootton, R.J., 2002, Design, function and evolution in the wings of holometabolous insects, Zoologica Scripta, 31(1):31-40
  162. Wootton, R.J., 2002, Insect wings: the world's smallest, smartest airfoils, Biologist, 49(3):97-100
  163. Wootton, R.J., & C.R. Betts, 1986, Homology and function in the wings of Heteroptera, Systematic Entomology, 11:389-400
  164. Wootton, R.J., & A. R. Ennos, 1989, The implications of function on the origin and homologies of the dipterous wing, Systematic Entomology, 14:507-520
  165. Wootton, R.J., R.C. Herbert, P.G. Young, & K.E. Evans, 2003, Approaches to the structural modeling of insect wings, Philosphical Transactions of the Royal Society of London, B, 358:1577-1587
  166. Wootton, R.J., & J. Kukalova-Peck, 2000, Flight adaptations in Palaeozoic Palaeoptera (Insecta), Biol. Rev., 75:129-167
  167. Wootton, R.J., J. Kukalova-Peck, D.J.S. Newman, & J. Muzon, 1998, Smart engineering in the mid-Carboniferous: how well could Palaeozoic dragonflies fly?, Science, 282:749-751
  168. Wu, J.H., & M. Sun.  2004.  Unsteady aerodynamic forces of a flapping wing.  J. Exp. Biol., 207(7):1137-1150.
  169. Zeng, L., 1996, High-resolution method for measuring the torsional deformation of a dragonfly wing by combining a displacemnt probe with an acousto-optic deflector, Optical engineering, 35(2):507-513
  170. Zeng, L., 1996, Two-color compensation method for measuring unsteady vertical force of an insect in a wind tunnel, Meas. Sci. Technol., 7:515-519
  171. Zeng, L., 2000, Optical methods for measuring the natural frequencies and torsional deformations of a dragonfly wing, Internet article: http://www.jst.go.jp/jst/erato/erato-symp-j/95Yo1/9501-TXT/koenl-07-E.html

 

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