A BIOLOGY OF DRAGONFLIES ASPECTS OF ZOOLOGY SERIES A BIOLOGY OF CRUSTACEA J. Green, B.Sc., Ph.D. INSECT SOUNDS P. T. Haskell, B.Sc., Ph.D. SOIL ANIMALS D. Keith McE. Kevan, B.Sc., Ph.D., A.I.C.T.A., F.R.S.Edin. A BIOLOGY OF DRAGONFLIES Philip S. Corbet, D.Sc., Ph.D. e FRONTISPIECE A BIOLOGY OF DRAGONFLIES by PHILIP S. CORBET, D.Sc., Ph.D. Entomology Research Institute, Research Branch, Canada Department of Agriculture, Ottawa tt A young male Anax imperator settled away from water during its period of H. F. & G. WITHERBY LTD. maturation. The glistening wings and light blue abdomen show that it is probably 5 Warwick Court, London, W.CW.C.1 between two and six days old. The time is late May in southern England, and the branch of Pinus sylvestris, to which the dragonfly is clinging, has bluish-green older leaves, a closed female cone, and golden-stemmed young shoots. The surrounding heathland comprises Pinus, Betula and Calluna growing on plateau gravel and Tertiary clay, an association much favoured by Anax imperator. From an original watercolour drawing by Ralph Thompson First published in 1962 by H. F. & G. WITHERBY LTD 5 Warwick Court, London, W.C.1 To my wife H. F. & G. WITHERBY LTD. 1962 All rights reserved PRINTED IN GREAT BRITAIN BY NORTHUMBERLAND PRESS LIMITED GATESHEAD ON TYNE ' Another thing worthy of observation in these creatures is, that they must hunt and seek after their food flying in the air, for which purpose nature has given them two large eyes, which make almost the whole head, and are like glittering mother-of-pearl. They have moreover four remarkable membranous silver-coloured wings, with which, as with oars, they can turn themselves like swallows with a prodigious velocity to all parts of the air. Mouffet, who erroneously asserted, that these crea- PREFACE tures were produced from rotten bull-rushes, speaks however very ration- ally of them, when he says: " They form one of the greatest beauties in Tilts book is concerned with the ecology of dragonflies, and in all nature, being superior by far to all art."' J. Swammerdam, c. 1675 particular with their physiology and behaviour. I have surveyed existing knowledge in these fields and considered it in relation to ' Dragonflies occur in all parts of the earth. Some species are found even the problems of adaptation which dragonflies encounter. While within the Arctic Circle, though, of course, they become more abundant doing this, I have taken the opportunity to indicate certain prob- the nearer we approach to the tropics' lems towards which research might profitably be directed in future. W. Harcourt Bath, 1890 Although dragonflies are primarily tropical insects, previous accounts of their biology have had to be based on the specialised, ' There are . .. other (Hawaiian) species the nymphs of which live under very different circumstances. These have given up their aquatic and therefore less typical, forms in temperate regions. Here I have life, and live hidden at the bases of the leaves of a liliaceous plant- been able to incorporate the observations which have been made on Astelia veratroides. .. These nymphs . differ in some points from tropical species, and thus reveal more clearly some of the factors those which frequent the water; they are shorter and stouter, and much more sluggish, and the caudal appendages are very short and thick, which determine patterns of behaviour in different environments differing therein greatly from some of the aquatic species, the appendages and at different latitudes. My main purpose, then, has been to of which form beautiful tracheal gills.' present an integrated ecological study of the group as a whole. R. C. L. Perkins, 1897 In reviewing the literature bearing on the subjects discussed, I have made no special attempt to cover work published after August Gynacantha nervosa seems essentially crepuscular in its flight. Where the species occurs abundantly the numbers on the wing and in sight at 196o, although several later publications which came to my notice once, the mobile active flight, and the rapidly coming darkness of the have plant Astelia. Specific facts or observations mentioned in tropical night combine to form a scene to fire the imagination. .. Sud- the text without an authority are from my own unpublished work. denly, a large brown dragonfly goes with undulating flight circling about a thatched but and at once the garden and path are alive with I wish to take this opportunity of acknowledging the help I have interweaving forms. received while preparing this book. E. B. Williamson, 1923 My main debt of gratitude is to my wife, who has assisted with every important aspect of the work, from the observation and cap- ' The age of specimens taken in the field cannot be determined with ture of dragonflies in the field to the typing of the manuscript. In accuracy, and if studies are to be made upon the development of glands and their physiological relations to reproduction, longevity and other particular I wish to thank her for many helpful discussions and for functional conditions, individuals must be reared from emergence to criticism of the text. reproductive maturity at least and possibly to normal death.' I am much indebted to Dr. M. A. Lieftinck and Dr. T. R. E. A. D. Whedon, 1942 Southwood for having criticised the original draft of the manu- script, and to Dr. Charles E. Jenner and Dr. Merle E. Jacobs for ' Copulation time seemed to depend on the male, for the same female (of having commented on Chapters IV and VII respectively. Although Perithemis tenera) would copulate long with a long-copulating male or briefly with a short-copulating male. The duration was often prolonged in I have benefited greatly from their criticisms, these authorities do shade or when the air temperature was below 25°C. Stop-watch timing of not necessarily agree with all the views advanced in this book. I am 623 undisturbed copulations of 25 males in sunlight at temperatures of 3o-32°C. showed the mean duration to be 17.5 seconds.' very grateful to my mother, Mrs. Irene Corbet, for having critically Merle E. Jacobs, 1955 read through the whole of the original draft. It is a pleasure for me to be able to thank several other friends ix X A BIOLOGY OF DRAGONFLIES and colleagues who have assisted in various ways. Mr. Ralph Thompson generously provided the drawing for the Frontispiece. Mr. A. E. Gardner and Dr. M. A. Lieftinck made available to me their published and unpublished drawings, and gave timely help by loaning notes and publications. Mr. S. Beaufoy made available photographs at short notice. Mr. D. E. Kimmins advised on the CONTENTS copying of figures. Mr. C. P. Barker gave help in the field. Several librarians gave courteous assistance, in particular Miss A. Wright of the Royal Entomological Society of London, and Mr. Stanley M. Page McDonald of the Concord Library, Massachussetts, who sent me an extract from the paper by R. H. Howe (1917), the only reference Preface ix cited I have been unable to see in the original. Mr. Zoltan Lukàcs Introductory Note xv translated the paper by L. Biro (1884). Miss Cynthia Longfield and Mr. E. C. G. Pinhey named large numbers of African dragonflies Chapter for me. Frau Hortense Wenner allowed my wife and me the use I HABITAT SELECTION AND OVIPOSITION of her beautiful house by Lake Constance, where the first draft of the manuscript was written during September and October 196o. By II THE EGG STAGE 3 1 his consideration and help. Mr. Antony Witherby has made my III THE LARVAL STAGE : GENERAL task of preparing this book a very pleasant one. 47 Most of the text figures derived from the publication of other IV GROWTH. METAMORPHOSIS AND EMER- authors or myself have been redrawn, as stated, but those repro- GENCE 90 duced here in their original form have been acknowledged in the V ADULT LIFE : GENERAL 120 manner `(From Smith, 1958)'. For permission to reproduce such illustrations I am indebted to the publishers of the following : VI ADULT FEEDING BEHAVIOUR 1 47 The Entomologist for Figures 6o, 62 and 63; Internationale Revue VII REPRODUCTIVE BEHAVIOUR 156 der gesamten Hydrobiologie for Figures 42-48; The Journal of Animal Ecology for Figures 83, 86-89, 96 and 104; Miscellaneous VIII DISPERSAL 183 Publications of the Museum of Zoology, University of Michigan IX EVOLUTIONARY PERSPECTIVE 198 for Figures 97 and 98; Nova Guinea (new series) for Figures 77-8o; The Proceedings of the Royal Entomological Society of London for References 207 Figures 3o, 31, 35 and 36; The Proceedings of the South London Supplementary References 2 Entomological and Natural History Society for Figures 14-16 and 33 o; Tombo (Acta Odonatologica) for Figure 55; and Treubia for Author Index 2 34 Figures 9, 65-70 and 72. The photographs for Plates V and VI were Odonata Index 238 first published by William Collins Sons and Co., Ltd. in Dragon- flies (1960). The picture on the front of the book-jacket, which por- Subject Index 2 43 trays a larva of Archilestes californica, was redrawn from the original by Kennedy (1915a). PHILIP S. CORSET. Entebbe, Uganda, April, 1962. xi LIST OF PLATES Frontispiece Young male adult of Anax imperator settled away from water. Facing page I Anax imperator. Eggs attached to a leaf; and a final instar larva feeding. 40 II Stages in the emergence of Libellula depressa. III Anax imperator emerging on an unusual support. Between pages IV Normal emergence of Anax imperator. Adults about 104 four hours old, just before departing on the maiden and flight.