SecOnD eDITIOn Dragonflies and damselflies are conspicuous insects: many are large and brightly coloured. They are also valuable indicators of environmental wellbeing. d A detailed knowledge of the dragonfly fauna is therefore an important basis r for decisions about environmental protection and management. aT H This comprehensive guide to the Australian dragonfly fauna covers eight g e families of dragonflies and 10 families of damselflies, comprising the O c 113 genera and 333 species found in Australia. It has been updated with THe cOmpleTe field guide TO O newly identified species and revised family names to reflect new world n consensus systematics. Stunning full-colour images and distribution maps m f dragOnflies are accompanied by identification keys for adults as well as larvae, which lp are often used as bait for freshwater fish. ile e T This second edition of The Complete Field Guide to Dragonflies of Australia se also includes illustrations by Albert Orr, one of the most renowned dragonfly O f Of ausTralia illustrators. The extraordinary diversity of dragonflies will interest entomologists i e and amateur naturalists alike. fl d a g uu SecOnD eDITIOn si d T günther Theischinger and John Hawking e r T aO colour illustrations by albert Orr l i a Günther Theischinger has had a lifelong interest in aquatic insects. He has published more than 300 scientific papers, and over the years has described more than 60 new species and several new genera of Australian dragonflies. T h He has been a visiting scientist at the Australian national Insect collection e in canberra, and is a Research Associate of the Australian Museum and a i s Visiting Fellow at the Smithsonian Institution, USA. c h i John Hawking is an Honorary Research Fellow at La Trobe University and n former invertebrate ecologist at the Murray Darling Research centre, Wodonga. g e He studied the ecology of dragonfly larvae for his Master’s degree and has r , since published greatly on their ecology, taxonomy and conservation. H a Albert Orr is a professional entomologist, writer, illustrator, consultant and scientific w editor. He is a retired lecturer in ecology and invertebrate zoology, still actively k researching. His main expertise is with butterflies, moths and dragonflies. in g a n d O r r THE COMPLETE FIELD GUIDE TO DRAGONFLIES OF AUSTRALIA SECOND EDITION Also available from CSIRO Publishing: The Complete Field Guide to Butterflies of Australia, Second Edition, by Michael F. Braby THE COMPLETE FIELD GUIDE TO DRAGONFLIES OF AUSTRALIA SECOND EDITION Günther Theischinger and John Hawking Colour illustrations by Albert Orr © Text Günther Theischinger and John Hawking 2021 © Colour illustrations Albert Orr 2021 All rights reserved. Except under the conditions described in the Australian Copyright Act 1968 and subsequent amendments, no part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means, electronic, mechanical, photocopying, recording, duplicating or otherwise, without the prior permission of the copyright owner. Contact CSIRO Publishing for all permission requests. The authors and illustrator assert their moral rights, including the right to be identified as a creator. A catalogue record for this book is available from the National Library of Australia. ISBN: 9781486313747 (pbk) ISBN: 9781486313754 (epdf) ISBN: 9781486313761 (epub) How to cite: Theischinger G, Hawking J, Orr A (2021) The Complete Field Guide to Dragonflies of Australia. 2nd edn. CSIRO Publishing, Melbourne. Published by: CSIRO Publishing Locked Bag 10 Clayton South VIC 3169 Australia Telephone: +61 3 9545 8400 Email: [email protected] Website: www.publish.csiro.au Front cover: (top, left to right) Austroaeschna ingrid, Eusynthemis ursula, Austropetalia annaliese (photos by Günther Theischinger); (bottom) Diphlebia coerulescens (photo by Albert Orr). Set in Minion 9.5/12 Cover design by James Kelly Printed in China by Leo Paper Products Ltd CSIRO Publishing publishes and distributes scientific, technical and health science books, magazines and journals from Australia to a worldwide audience and conducts these activities autonomously from the research activities of the Commonwealth Scientific and Industrial Research Organisation (CSIRO). The views expressed in this publication are those of the author(s) and do not necessarily represent those of, and should not be attributed to, the publisher or CSIRO. The copyright owner shall not be liable for technical or other errors or omissions contained herein. The reader/user accepts all risks and responsibility for losses, damages, costs and other consequences resulting directly or indirectly from using this information. The paper this book is printed on is in accordance with the standards of the Forest Stewardship Council® and other controlled material. The FSC® promotes environmentally responsible, socially beneficial and economically viable management of the world’s forests. Nov20_01 Foreword to first edition In today’s crowded world humans are imposing increasing pressures on the pre-existing fauna and flora. Freshwater habitats have become especially vulnerable to such pressures and protection of their biota is now an urgent priority for people and agencies concerned with nature conservation. To an increasing extent conservationists are becoming aware of the vital role that insects play in maintaining the balance of nature. Large, diurnal, conspicuous animals like dragonflies serve as excellent ambassadors for insects. At the same time they are valuable indicators of healthy freshwater habitats and so have an important role to play in aiding conservation and conservation awareness. The Australian dragonfly fauna, comprising more than 300 species, is dauntingly large for the novice to tackle without an authoritative identification manual. An important first step towards this goal was the publication, in 1991, of the book The Australian Dragonflies by Tony Watson, Günther Theischinger and Hilda Abbey, which became an essential reference for Australian odonatologists. Still needed was a user-friendly field guide. The authors of this book deserve the gratitude and congratulations of entomologists and conservationists alike for having met this need by producing this excellent book. Apart from giving field observers much pleasure and satisfaction, it will provide an invaluable tool in the armoury of those charged with compiling reliable fauna lists needed for the conservation of aquatic habitats and their occupants. I wish this attractive book the success it richly deserves. Philip S Corbet University of Edinburgh, UK Foreword to second edition Since the middle of last century, interest in dragonflies by professionals and non- professionals alike has increased enormously throughout the globe. Whereas formerly scientists focussed their interest on the structure and behaviour of these ‘living fossils’, that have remained essentially unchanged since Upper Palaeozoic times, nowadays the role of dragonflies in biotic communities and their significant capacity as indicators of freshwater health has become a key issue. No serious nature conservation research today would omit the critical consideration of the effects of environmental degradation on the regional dragonfly fauna. Just as no poetry can be written without the author having a command of words, in biological research a knowledge of species is absolutely vital. To this end the present book will serve as an indispensable tool throughout Australia. First published in 2006, this second edition has been significantly revised and updated. The frequent reprinting of the first edition within a short space of time speaks for the importance of and demand for this definitive work. The scientific profile of its authors is a firm guarantee of the excellence of the treatment, based on the most recent views of odonate classification. The taxa described or recorded from Australia after 2006 are added, bringing the number of the hitherto known Australian species close to the 350 mark. All species are concisely described and the information on their habitats and larvae is supplied. The adults and the known larvae are keyed in a user-friendly style. The quality of the text is enhanced with artwork by Dr Albert George Orr, one of the greatest dragonfly illustrators of our time. Many of the dragonfly portraits from the previous edition are also replaced by new, higher-quality photographs. This new edition represents a milestone in our knowledge of Australian dragonflies and will remain a standard reference for a long time to come. Bastiaan Kiauta Past-Editor of Odonatologica Xanthagrion erythroneurum. Preface and acknowledgments to first edition Dragonflies, with their kaleidoscope of colours, exquisite symmetry and incredible flying abilities, are one of the most eminently watchable kinds of animals. They have featured in folklore over time and their aesthetic beauty still fascinates us today. There has always been much interest in the group – in the past mainly with scientists, but more recently also with amateur watchers. Through recent major studies we now have a reasonable understanding of the Australian dragonflies and their larvae, best known as ‘mud-eyes’ and often used as bait in freshwater fishing. The introductory chapter of this guide deals briefly with the biology and ecology of the order Odonata. This is followed by the species accounts, which should allow identification of individual species by diagnostic notes, figures, distribution maps and photographs. The photographs and diagrams are not to scale. At the end of the book are illustrated keys for families, genera and some species, as well as diagnostic keys for larvae which are, however, based on characters of the final instar and are less reliable for earlier instars. There is also an illustrated glossary of descriptive terms. A number of people greatly supported us by sending particular photographs for publication and/or by allowing us to pick photographs from their collection. They are: Steve Axford, Graham Brown, Klaus Buck, Max Carpenter, Peter Chew, Sid Dunkle, Günther Fleck, Andre Günther, Ego Guiotto, Karlie Hawking, Antoine van der Heijden, Glenn Hoye, Steve Jacobs, Kathie Le Busque, Leonard Müller, Norman Müller, Dennis Paulson, David Rentz, Ulrich Röder, Deniss Reeves, Steve Richards, Netta Smith, Tanya Whiteway and Keith Wilson. For help with proofreading we are deeply indebted to Sid Dunkle, Ian Endersby, Leonard Müller, Deniss Reeves and Neville Yates. We wish to thank Dr P Scanes, Dr T Pritchard and Dr C Koop of the New South Wales Department of Environment and Conservation, and Dr D Mitchell, Dr T Hillman and Dr B Gawne of the Murray-Darling Freshwater Research Centre for their support. Special thanks go to Nick Alexander who never lost track during his navigation through difficult terrain. It is he who should be particularly credited if the book is found appealing. Günther Theischinger John Hawking Preface and acknowledgments to second edition The first edition of The Complete Field Guide to Dragonflies of Australia was published in 2006, and three reprints (2010, 2012, 2016) followed without any significant changes, additions or amendments. The opportunity was then offered and taken to present a new edition with changes, additions or amendments where it appeared timely and appropriate. As the fundamental aims of the book have not changed, namely to provide an authoritative but usable reference for the identification of dragonflies from Australia, this update is based as closely as possible on the first edition. There were, however, in the last 14 years several significant advances in the knowledge of dragonflies world-wide and in Australia that necessitated updating and revising this field guide. Other significant recent research is listed in the ‘References and further reading’ section. New studies by various authors on the morphology of dragonfly adults and larvae, cladistics and lately molecular phylogeny paved the way for a multi-authored consensus on the higher classification of dragonflies based on the principles of stability (usage should change as little as possible) and monophyly of family group taxa (Dijkstra et al. 2013). This arrangement, complemented by Kalkman & Theischinger (2013) and Dijkstra et al. (2014), is generally followed in this edition. Thus the order of family group taxa in Australia is at least temporarily, and for better or worse, brought in line with that followed in the rest of the world. Details and differences of this classification to the previously used classification are given in the section ‘Species guide’. A molecular phylogeny and classification of Anisoptera is now also available (Carle et al. 2015), but its findings seem too premature to alter the consensus arrangement, which is still the most widely used scheme. Since the first edition there have been several additions to the Australian fauna. These include an isostictid genus not previously known from Australia but recently found in tropical Queensland (Theischinger 2009), as well as seven newly described species, six from eastern Australia (Theischinger 2008b, 2013, 2018, 2019, 2020; Theischinger & Burwell 2017) and one supposed from photographic evidence to occur in Australia (Rowe 2019). Name and taxonomic status for some taxa were changed (e.g. Peters 2006). De Baar et al. (2010), reporting on the fauna of Dauan Island, an Australian Island very close to New Guinea, listed two spectacular dragonfly species hitherto not recorded from Australia. Kalkman & Orr (2012) discussed the Australian monsoon tropics as a barrier for exchange of dragonflies between New Guinea and Australia. Theischinger & Endersby (2009) published a book on the Australian fauna, including keys for identification of adults and larvae at species level, distribution dot maps for all species and references to species of conservation concern. Important particularly for studying Australia’s northernmost dragonflies, the fauna of New Guinea, Maluku and the Solomon