Field Guide to the Dragonflies of Britain and Europe [2 ed.] 1472943996, 9781472943996

Citation preview

Field Guide to the

Dragonflies of Britain and Europe including western Turkey and north-western Africa

second edition

GENERAL EDITOR KLAAS-DOUWE B DIJKSTRA EDITOR FOR SECOND EDITION ASMUS SCHRÖTER ILLUSTRATED BY RICHARD LEWINGTON

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BLOOMSBURY WILDLIFE Bloomsbury Publishing Plc 50 Bedford Square, London, WC1B 3DP, UK BLOOMSBURY, BLOOMSBURY WILDLIFE and the Diana logo are trademarks of Bloomsbury Publishing Plc First edition published in the United Kingdom in 2006 This edition published in 2020 This electronic edition published in 2020 by Bloomsbury Publishing Plc Copyright © Klaas-Douwe B Dijkstra and Asmus Schröter, 2020 Illustrations © Richard Lewington, 2020 Maps © Klaas-Douwe B Dijkstra and Asmus Schröter, 2020 Recommended citation: Dijkstra, K.-D.B., A. Schröter & R. Lewington. 2020. Field Guide to the Dragonflies of Britain and Europe. Second edition. Bloomsbury Publishing, London. Klaas-Douwe B Dijkstra and Asmus Schröter have asserted their right under the Copyright, Designs and Patents Act,1988, to be identified as Authors of this work For legal purposes the photographic credits on p. 331 constitute an extension of this copyright page All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, recording, or any information storage or retrieval system, without prior permission in writing from the publishers Bloomsbury Publishing Plc does not have any control over, or responsibility for, any third-party websites referred to in this book. All internet addresses given in this book were correct at the time of going to press. The author and publisher regret any inconvenience caused if addresses have changed or sites have ceased to exist, but can accept no responsibility for any such changes A catalogue record for this book is available from the British Library Library of Congress Cataloguing-in-Publication data has been applied for ISBN: HB: 978-1-4729-4399-6; PB: 978-1-4729-4395-8; ePDF: 978-1-4729-4397-2; ePUB: 978-1-4729-4398-9 Designed by D & N Publishing, Baydon, Wiltshire

To find out more about our authors and books visit www.bloomsbury.com and sign up for our newsletters Front cover: Violet Dropwing Trithemis annulata

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Contents Acknowledgements . . . . . . . . . . 4

Thoughts on field guides . . . . . 5

Introduction

6

Studying dragonflies . . . . . . . . 6 Dragonfly behaviour . . . . . . . . 8 Dragonfly occurrence . . . . . . . . 9 Map key . . . . . . . . . . . . . . . . . 12 Habitat . . . . . . . . . . . . . . . . . 12 Flight season . . . . . . . . . . . . . 14

Dragonfly names . . . . . . . . . . Dragonfly identification . . . . . Glossary . . . . . . . . . . . . . . . . . Identification of suborders, families and genera . . . . . . . . Larvae and exuviae. . . . . . . . . .

Regional guide to dragonflies Identification Zygoptera Damselflies Lestidae . . . . . . . . . . . . . . . . . . . . . Lestes and Chalcolestes . . . . . . . Sympecma . . . . . . . . . . . . . . . . . . Calopterygidae . . . . . . . . . . . . . . Calopteryx . . . . . . . . . . . . . . . . . . Euphaeidae . . . . . . . . . . . . . . . . . Epallage . . . . . . . . . . . . . . . . . . . . Platycnemididae . . . . . . . . . . . . . Platycnemis . . . . . . . . . . . . . . . . .

69 69 80 84 84 93 93 95 95

Appendices

21 36

38 68 69

Coenagrionidae . . . . . . . . . . . . 103 Ischnura . . . . . . . . . . . . . . . . . . . 103 Enallagma . . . . . . . . . . . . . . . . . . 117 Coenagrion . . . . . . . . . . . . . . . . 120 Erythromma . . . . . . . . . . . . . . . . 135 Pyrrhosoma . . . . . . . . . . . . . . . . 139 Ceriagrion . . . . . . . . . . . . . . . . . . 142 Nehalennia . . . . . . . . . . . . . . . . . 144 Pseudagrion . . . . . . . . . . . . . . . . 145

Anisoptera True Dragonflies Aeshnidae . . . . . . . . . . . . . . . . . 147 Aeshna . . . . . . . . . . . . . . . . . . . . 147 Anax . . . . . . . . . . . . . . . . . . . . . . 170 Brachytron . . . . . . . . . . . . . . . . . 182 Boyeria . . . . . . . . . . . . . . . . . . . . 184 Caliaeschna . . . . . . . . . . . . . . . . 186 Gomphidae . . . . . . . . . . . . . . . . 188 Gomphus and Stylurus . . . . . . . 188 Ophiogomphus . . . . . . . . . . . . . 202 Onychogomphus . . . . . . . . . . . . 204 Paragomphus . . . . . . . . . . . . . . . 216 Lindenia . . . . . . . . . . . . . . . . . . . 218 Cordulegastridae . . . . . . . . . . . . 220 Cordulegaster . . . . . . . . . . . . . . 220 Family affiliation uncertain . . . 232 Oxygastra . . . . . . . . . . . . . . . . . . 232 Macromiidae . . . . . . . . . . . . . . . . 234 Macromia . . . . . . . . . . . . . . . . . . 234 Corduliidae . . . . . . . . . . . . . . . . . 236 Cordulia . . . . . . . . . . . . . . . . . . . 236

14 15 17

147

Somatochlora . . . . . . . . . . . . . . 238 Epitheca . . . . . . . . . . . . . . . . . . . 250 Libellulidae . . . . . . . . . . . . . . . . . . 252 Libellula . . . . . . . . . . . . . . . . . . . . 252 Orthetrum . . . . . . . . . . . . . . . . . 259 Leucorrhinia . . . . . . . . . . . . . . . . 274 Sympetrum . . . . . . . . . . . . . . . . . 281 Crocothemis . . . . . . . . . . . . . . . . 301 Trithemis . . . . . . . . . . . . . . . . . . . 304 Brachythemis . . . . . . . . . . . . . . . 309 Diplacodes . . . . . . . . . . . . . . . . . 312 Selysiothemis . . . . . . . . . . . . . . . 313 Acisoma . . . . . . . . . . . . . . . . . . . 314 Pachydiplax . . . . . . . . . . . . . . . . 315 Pantala . . . . . . . . . . . . . . . . . . . . 316 Tramea . . . . . . . . . . . . . . . . . . . . 318 Rhyothemis . . . . . . . . . . . . . . . . 320 Urothemis . . . . . . . . . . . . . . . . . . 321 Zygonyx . . . . . . . . . . . . . . . . . . . 322

323

Photographic and mapping acknowledgements. . . . . . . . . . . . . . . . 331

Index

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Acknowledgements

4

General Editor’s and Artist’s acknowledgements for the first edition Too many colleagues to mention provided their knowledge and support. Steve Brooks and Graham Vick were supportive when the book was in its embryonic stages. Henri Dumont, Reinhard Jödicke, Vincent Kalkman, Andreas Martens, Ole Müller, Gert Jan van Pelt, Göran Sahlén, Frank Suhling and Hansruedi Wildermuth contributed as species authors. Some of them also provided regional texts, as did Andras Ambrús, Matjaž Bedjanič, Rafał Bernard, Tomo Bogdanović, Jean-Pierre Boudot, Paweł Buczyński, Steve Cham, Adolfo Cordero Rivera, Sónia Ferreira, Matti Hämäläinen, Otakar Holuša, Gilles Jacquemin, Geert De Knijf, Milen Marinov, Rüdiger Mauersberger, Brian Nelson, Maurizio Pavesi, Rainer Raab, Boudjéma Samraoui and Florian Weihrauch. Many of the above helped with the distribution maps, especially Adolfo, Jean-Pierre, Otakar, Paweł, Rafał, Rainer and Sónia, and the mapping team was strengthened with Elena Dyatlova, Miloš Jović and Cosmin Manci. Others provided specimens and photographic references for illustration, particularly Gilles, Graham and Reinhard, but also Graham Giles, Bob Kemp, Günther Peters and Wolfgang Schneider. Adrian Parr invested much time to discuss vernacular names. Access to the facilities of the National Museum of Natural History Naturalis in Leiden, where K-D received particular support from Jan van Tol, was crucial to the project. Finally, we must honour our publishers Andrew and Anne Branson for their initiative and commitment. We owe all of the above incredible thanks, especially Vincent, who was K-D’s indispensable touchstone and workhorse throughout the project. With such a large and expert team it was not always easy to agree on how this ‘next generation’ field guide, as we liked to call it, should take form. As one author put it after another discussion: ‘Editing: the final frontier… To boldly rewrite what everybody has written before…’ We hope to have ‘boldly rewritten’ something new, that will take many on exciting voyages of odonatological enterprise. K-D B Dijkstra and Richard Lewington Acknowledgements for the second edition In addition to the people listed above, we are obliged to the following people for support during various stages of the revision process: Vasil Ananian, Jörg Arlt, Phil Benstead, Natia Berdzenishvili, Magnus Billqvist, Angelika Borkenstein, Christophe Brochard, Andrea Corso, Cecilia Díaz Martínez, André Günther, Fons Peels, Jorge Pérez, Oz Rittner, Malte Seehausen, David Sparrow, Warwick Tarboton, and Santiago Teruel (of PAROTETS S.O.C.V.). K-D B Dijkstra and Asmus Schröter The following authors contributed genus and species texts: H J Dumont (Calopteryx), R Jödicke (Sympecma, Ischnura, Enallagma), V J Kalkman (Epallage, Pyrrhosoma, Ceriagrion, Anax, Onychogomphus, Paragomphus, Lindenia, Orthetrum, Brachythemis, Selysiothemis), A Martens (Platycnemis), G J van Pelt (Cordulegaster), G Sahlén (Leucorrhinia), F Suhling and O Müller, (Gomphus and Stylurus, Ophiogomphus) and H Wildermuth (Cordulia, Somatochlora, Epitheca, Oxygastra, Macromia). All others by K-D B Dijkstra and A Schröter.

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Thoughts on field guides

5

G Erythromma najas pair mating.

I got my first field guide, to the birds of Europe, at the age of nine. We lived in Egypt, however, so on my tenth birthday I began to write and draw my own book. Then, with no reference for them at all, I started to describe and name dragonflies at the age of 12. Later, I’d learn that my ‘blood-red dragonfly’ and ‘grass dragonfly’ were the male and female of just one species, now called the Broad Scarlet. Eight years after naming one ‘hairy dragonfly’ as a boy, my friends and I found the first Vagrant Emperor in the Netherlands. From that moment on, my passion for dragonflies was fully fledged. And so, when this book’s first edition came out in 2006, I felt I was fulfilling my destiny. Fourteen years later, the field guide (now available in five languages) remains my proudest achievement. Reaching about ten new users every day in its first ten years, it surfed on a surge of interest. The dragons themselves, meanwhile, advanced as summers warmed, or rebounded as habitats recovered, while others declined. Regional dragonfly societies flourished and published atlases, and by 2015 all of Europe’s species had been mapped. Six more species were found in the area covered, of which two were new to science. Most other continents now have guides or handbooks too, allowin and online platforms to amass tens of thousands of records each year. In the coming years, all of the approximately 6,300 known species will be assessed for the IUCN Red List of Threatened Species, and as such Odonata will be the first insect order to be completed worldwide. Few creatures represent fresh water – probably life’s most precious resource – as vividly as do dragonflies, which emerge from their watery habitats and enter our lives on land to drive the message home. I’m often envious of the attention received by insects as pollinators, or by fish in freshwater conservation, and while people often ask what dragonflies are good for, their greatest value is actually not their use to us. We admire them first for their beauty, not for the mosquitoes they may devour. And we protect them first for the sake of their health and that of their – and thus our own – environment. Only such an unconditional attitude towards all of nature may cure us of our destructive demands. Field guides initiate that conviction with one potent idea: species have identities to honour, just like people do. Each has a face, a name and a story. Every guide (whether a book, or its edified user) offers a route into life’s overwhelming diversity, allowing us to get to know other life forms and come to love the realms they inhabit. Effectively, through identification, species ‘become’ a second time, now not into being but into being seen. Thus, we gain awareness of an infinite universe of parallel worlds, each occupied by another species and all of which are humanity’s equal. To me, therefore, field guides are ultimately about all destinies. Biodiversity lets us see Earth’s crises of inequality at their true scale, as a force that affects everything. Does the power to command all living space also give the right to take it? Our challenge is not to answer that question, but to make it rhetorical. Perhaps one day a global field guide movement will emerge to drive the message home, one species – and one observer – at a time. Klaas-Douwe ‘K-D’ B Dijkstra

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Introduction Studying dragonflies

6

Time and place Although adult dragonflies may be found almost anywhere, at any time, certain places and times can provide especially good opportunities to see them. To understand where and when to look for them, some knowledge of their behaviour and ecology is necessary (see: Dragonfly behaviour, Dragonfly occurrence). Adult dragonflies are most easily found near fresh water, under warm conditions. This generalisation, however, requires some qualification. Although salt water is uninhabitable for dragonfly larvae, slightly brackish conditions are tolerated by a few species. Very cold and wild water, such as meltwater torrents, are avoided. Special habitats such as bogs, seepages and large rivers harbour specialised species, which may be rare or difficult to find. The richest assemblages of species are found at sunny and sheltered ponds, on lake shores, and on rivers and calm streams, especially where these have a variety of aquatic and waterside vegetation. Many adult dragonflies may also be found away from water while foraging. Rough meadows, woodland glades, forest edges, roadside verges and other places rich in insect prey are particularly good. Shy species, for instance members of the family Gomphidae, may be found more easily in such situations than at their breeding habitat. The best time to look for dragonflies is at midday, in summer, with calm, sunny weather. Different species, however, have different periods of adult activity (see: Flight season). In most of Europe, a minimum of one visit during the period May–June and another in July–August will be necessary to see adults of all the present species. Rain and wind are not conducive to observing dragonflies, although unusual weather conditions can sometimes result in interesting discoveries: an influx of vagrants or invasions of southern species is often associated with favourable wind directions. The majority of species are most active in direct sunshine, with some even leaving the breeding habitat when it clouds over. Nonetheless, there are many species, especially damselflies, that are also active in overcast weather, provided it is sufficiently warm. Although activity is greatest around midday, certain behaviour can merit an excursion at the start or close of the day. For example, many hawker species (Aeshnidae) hunt most actively from shortly before sunset until dark, while two mate predominantly around sunrise. Emergence, especially of anisopterans, is concentrated in the early hours of the morning. Watching and catching A pair of close-focus binoculars is essential in order to examine details in the field. Many people prefer the compact binoculars that fit into a breast pocket, but the better optics of a good pair of binoculars is well worth the investment. You should at least be able to focus up to just in front of your feet. However, many details cannot be seen even with good optical equipment, making a net indispensable (see: Dragonfly identification). Numerous species, especially in southern regions, can be identified reliably only at close quarters. The speed and agility of dragonflies demands a fairly large, light net – the type used for butterflies is generally suitable (net opening about 40–75cm wide, handle about 1–2m long). The net must be deep enough to fold closed, so the catch cannot escape. A net with a handle composed of loose segments or a telescopic rod is useful as the length can be varied. Dragonflies are best held with the wings folded together between the thumb and forefinger. Larger species (especially anisopterans) can be held at the thorax or legs, provided at least three legs (on one side) are grasped. Although most dragonflies are sturdy and not easily damaged, careless or prolonged handling (especially with sweaty fingers) may lead to the loss of legs or damage to the wing membrane. Tenerals are too soft to handle safely. Some people object to the capture of animals, including dragonflies, but most individuals fly off unharmed after capture if handled carefully. Numerous studies in which dragonflies were caught, marked and released have shown that this procedure does not increase mortality. The impact on populations is negligible, and on the individual itself Introduction

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is probably minimal. Whether one catches dragonflies is a personal choice, and it is always good to bear in mind that some may prefer to see dragonflies behaving freely, rather than close up in the hand. The conservation status of a species should also be considered (see: Status). The protected status of certain species, and wildlife as a whole, also differs between nations, as do laws and attitudes concerning land access and the capture or G Dutch dragonfly watchers focus in on a Coenagrion armatum in the collecting of dragonflies. Weerribben, shortly after the species’ rediscovery in the Netherlands.

7

Research and collecting Inspired by their beauty, mystery or dynamism, some may wish to develop an initial admiration of dragonflies into a more profound understanding of their distribution, behaviour, relationships or conservation. Yet surprisingly little is known about many aspects of these insects. There are huge gaps in the known distribution ranges, which are, moreover, changing (see: Range). In some regions, especially in south-eastern Europe, the taxonomic status of certain species is poorly known and demands critical examination of relevant characters. The status of most subspecies, in particular, is unclear (see: Scientific names, Subspecies). Besides establishing the correct names of dragonflies, such research may clarify how colonisations, extinctions and hybridisation have shaped our present-day dragonfly fauna (see: Range). The increased application of molecular techniques may be particularly useful in clarifying many of these issues. Aside from this more geographically oriented research, numerous challenges lie in ecological and behavioural studies. The life history of many species is poorly known, with simple questions, such as the length of larval development, still unanswered. Such answers may, for instance, explain how the rapid colonisation of new habitat by some species is physically possible, while others are vulnerable to change. Most research requires some handling of dragonflies and, in many cases, the collection of specimens. As with catching dragonflies (see: Watching and catching), it is important to know the purpose of collecting. Some species, especially in the south-east of the region covered by this guide, can be identified reliably only under magnification. In such cases, a collected specimen is the best way to document a record, for instance of a range extension. In many areas, collecting is not essential as proof of a record (photographs generally suffice), but may be a tool to familiarise oneself with a species. In any case, it is important to document special records, as it is always possible to overlook details in the field that cannot be reproduced later. If you are considering creating a collection, it is a good idea to contact a national or regional natural history museum, not only for advice, but also to ensure that material is preserved for posterity. Nowadays, most dragonflies are not pinned like butterflies, but kept in small plastic or paper envelopes. The most suitable preservation agent is acetone. Dragonflies that are soaked in acetone (which also kills them) for a night up to a maximum of 24 hours are dehydrated and dry quickly on exposure to air, limiting discoloration and decay. It should also be remembered that most of our species can be identified by their larval skin (exuvia), and collecting these is a less invasive alternative to collecting adults; they merely need to be dried. A contemporary argument for collecting is its application in the analysis of dragonfly genes. Complex questions about the age, origin and relationships of species, subspecies and populations can be answered only with sufficiently large samples, spread across a representative geographical range. It seems likely that geneticists will increasingly seek the help of dragonfly Introduction

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observers. Although DNA can be extracted from dry samples, its structure may deteriorate with time. Preservation in concentrated alcohol (70% or more) is more reliable. In most cases, a single leg provides enough material for molecular purposes. Extracting DNA from exuviae seems ideal, as it does not require killing or damaging individuals, but it probably works only with very fresh, correctly preserved samples.

8

Literature Many nations covered by this guide have regional societies, journals, mapping schemes, distribution atlases and/or handbooks. Altogether, hundreds of titles, in various languages, now exist on dragonflies. This is only a selection of books of either a general or comprehensive nature: Boudot, J-P & Kalkman, V J (editors) 2015 Atlas of the European dragonflies and damselflies. KNNV Publishing, Zeist. 381pp. Comprehensive reference to the taxonomy, ecology, conservation and regional and global distribution of British and European species. Boudot, J-P, Kalkman, V J, Azpilicueta Amorín, M, Bogdanovic, ´ T, Cordero Rivera, A, Degabriele, G, Dommanget, J L, Ferreira, S, Garrigós, B, Jovic, ´ M, Kotarac, M, Lopau, W, Marinov, M, Mihokovic, ´ N, Riservato, E, Samraoui, B & Schneider, W 2009 Atlas of the Odonata of the Mediterranean and North Africa. Libellula Supplement 9: 1–256. Complement to the European atlas, also treating the species from Turkey and North Africa covered by this guide. Brochard, C, Groenendijk, D, van der Ploeg, E & Termaat, T 2012 Fotogids Larvenhuidjes van Libellen. KNNV Publishing, Zeist. 320pp. Sublimely illustrated guide to north-western European exuviae. Brochard, C, & van der Ploeg, E 2014 Fotogids Larven van Libellen. KNNV Publishing, Zeist. 272pp (both in Dutch). Sublimely illustrated guide to north-western European larvae. Cham, S 2012 Field guide to the larvae and exuviae of British dragonflies. British Dragonfly Society, Peterborough. 152pp. Excellent alternative to Brochard et al. (2012) and Brochard & van der Ploeg (2014), for British species only. Corbet, P S 1999 Dragonflies: behaviour and ecology of Odonata. Harley Books, Colchester. 829pp. Most comprehensive and detailed scientific account of all aspects of the group’s life history yet written. Paulson, D 2019 Dragonflies and damselflies: A natural history. Ivy Press, Brighton. 224pp. Richly illustrated popular introduction to the world’s species. Suhling, F, Sahlén, G, Gorb, S, Kalkman, V J, Dijkstra, K-D B, & van Tol, J 2015 Order Odonata. In: Thorp, J & Rodgers D C (editors). Ecology and general biology. Thorp and Covich’s freshwater invertebrates. 4th edition. Academic Press: 893–932. General but detailed introduction to the order’s diversity, morphology and habits. Wildermuth, H, & Martens, A 2018 Die Libellen Europas. Quelle & Meyer, Wiebelsheim. 957pp (in German). Most detailed ecological and photographic account of the species found in Europe, from the Azores to the Urals.

Dragonfly behaviour The limitations of space and the focus on identification mean that this guide does not include an extensive section on the ecology and life cycle of Odonata. In the species texts, only diagnostic behaviour that may aid identification is provided, e.g. tandem oviposition in a genus where solitary oviposition is the rule. However, numerous informative introductions about dragonfly biology exist in print (see above), including the monumental scientific treatise by Corbet (1999), as well as online. There are three elementary facts of dragonfly biology that explain most peculiarities of their behaviour: 1. Dragonflies are amphibiotic Larvae live in water, adults on land and in the air. The larva sheds its skin several times underwater, allowing it to grow. When it is fully grown, the larva leaves the water, moults a final time, expands its wings and abdomen and, when these have hardened sufficiently, flies off as an adult dragonfly. This is known as emergence; dragonflies Introduction

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9

G Sympetrum danae pair mating. Dragonflies have a unique way of copulating, for which the males have specialised secondary genitalia.

do not pupate. The final larval skin, the exuvia, can be found at the waterside as proof of a completed life cycle at that site. 2. Dragonflies are obligate carnivores All species hunt, both as larvae and adults. Most prey is invertebrate, especially insects, but a large larva may, for instance, eat tadpoles or small fish. 3. Dragonflies have a unique mode of reproduction, with indirect insemination and delayed fertilisation Sperm is transferred by the male from the abdomen tip, where it is produced, to the secondary genitalia at his abdomen base, from where it is passed on to the female. Other animals either transfer sperm indirectly outside the body, or directly. Eggs are fertilised when they are laid, allowing males to remove sperm of rival males when they succeed in copulating with a mated female. These facts have many consequences for dragonfly behaviour, explaining such characteristic sights as the fierce defence of waterside territories by males (good larval habitat, to which potential mates are lured when their eggs have matured), feeding swarms (e.g. where flying ants emerge), the heart-shaped mating wheel (achieved when insemination from the secondary genitalia takes place) and the male hovering above the female as she oviposits (guarding her against rivals that may attempt to replace his sperm with their own).

Dragonfly occurrence The guide covers all species of Odonata that occur in the wild in Europe and Turkey west of Moscow and the line Samsun–Iskenderun, in north-western Africa up to the northern fringe of the Sahara, and on the islands of Cyprus, Madeira, the Canaries and the Azores. Species that have accidentally been introduced to our area, such as those imported with tropical plants as larvae, are not covered in the guide as none have been known to survive in the wild. Given the large geographic area covered in this guide, there is no room to discuss each regional detail of occurrence in space (e.g. distribution and habitat), time (flight period) and the interaction of both these factors (decline, invasions). For example, Aeshna juncea inhabits most types of standing water at higher altitudes and latitudes, but in the low-lying areas of central Europe it is largely confined to acidic water, such as bog lakes. The habitat and location influence temperature and food availability, which in turn determine larval growth and adult emergence. Such scenarios are too complex for compact species texts, in which we have tried Introduction

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to describe general characteristics and mention only broad regional variation. This complexity is exacerbated by the lack of information for many areas, and the degree of recent change that has been observed. The many remaining ‘white spots’ and the dynamics of dragonfly distribution make future research urgent and, while attempting to provide information that is as up to date as possible, we call on the users of the field guide to contribute to that research.

10

Status The abundance of a species in an area is susceptible to change in both the long and short term. Many species are known to have suffered as a result of the destruction and degradation of their habitats. One-fifth of the species covered in this field guide are listed either on the IUCN Red List of Threatened Species or the European Habitats Directive, and a similar number occur only in Europe and adjacent north-western Africa (see table opposite). Together, this means that, while two-thirds of our species are safe in their huge ranges stretching into Asia and Africa, almost a third deserve our direct conservation attention. Many additional species are included on national or regional red lists. While some species are threatened with decline, numerous species have expanded or consolidated their range in recent years. These are mostly southern species such as Erythromma viridulum, Anax imperator and Crocothemis erythraea, which have benefited from the warm summers since the 1990s. Their increase is often not gradual, but invasive. With suitable weather, many individuals of Aeshna affinis or Sympetrum fonscolombii may appear and breed in an area where previously they had been rare or unknown. Some more eastern species are also known to be eruptive (such as Sympetrum flaveolum) or expansive (S. pedemontanum) in their occurrence. While many species have suffered through habitat change, others such as Chalcolestes viridis and Orthetrum cancellatum may benefit from ponds or gravel pits created by humans. Range The current distributions of dragonflies in our region were shaped by the impact of the ice ages on their environment and the species’ capacities to colonise changing landscapes. Many dragonfly species were probably completely obliterated by the relatively severe glaciation in western Eurasia. As a consequence, our fauna is much poorer than that of other temperate regions such as Japan and North America. The fauna in most areas is relatively young, as the most recent glaciation ended only about 10,000 years ago. The most characteristic patterns are: 1. Southern species Although Europe was never entirely covered by ice, large areas were inhospitable to many species, and continued so for long periods after glaciation. These species survived glaciation where warm conditions persisted, recolonising as the landscape recovered. Such climatically favourable pockets probably lay near both ends of the Mediterranean Sea. Species with a distinctly south-western European range are thought to have survived at the western end – Boyeria irene, Macromia splendens and Oxygastra curtisii are the most charismatic examples. Epallage fatime and Caliaeschna microstigma just enter Europe in the Balkans, and must have survived somewhere at the eastern end; their closest relatives live in tropical Asia. Other species now range so widely that it is hard to trace their origin. 2. Northern species The species that were most tolerant of cold were able to expand their range when taiga and tundra dominated the European landscape, but became confined to high altitudes and latitudes as the world warmed up. Examples of such ‘boreo-alpine’ species are Aeshna caerulea and Somatochlora alpestris. 3. Eastern species These take a position between the southern and northern species, inhabiting an intermediate temperature range. They expanded from the east, and are associated with temperate woodlands and bogs. Aeshna grandis and the Leucorrhinia species are examples of dragonflies that are common in north-eastern Europe and Siberia, but peter out further west, often being confined to highlands. The fragmented fringes of their Introduction

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Species

End. Red Eur. Med. HD List

Lestes macrostigma L. numidicus

VU N

DD

N

EN

Sympecma paedisca Calopteryx exul C. haemorrhoidalis C. xanthostoma

Species

NT

Lindenia tetraphylla

DD

Onychogomphus assimilis

EN

IV

N

E+N

O. costae

E+N NT

E

O. flexuosus O. macrodon

P. latipes

E

O. uncatus

N

Ophiogomphus cecilia

Ceriagrion georgifreyi

VU

C. tenellum

E+N

Coenagrion caerulescens

E+N

C. hylas

CR

VU

II+IV

E

VU

VU

VU

C. mercuriale

E+N NT

NT

NT

C. ornatum

NT

II+IV

CR

CR EN

NT

VU

VU

VU

VU

11

E+N II+IV NT

Cordulegaster bidentata

E

C. boltonii

E+N

C. helladica

E

EN

EN

EN

C. heros

E

NT

NT

VU

IV

NT

II+IV

C. insignis

EN

NT

II

C. picta

VU

VU

II+IV

C. princeps

N

C. trinacriae

E

NT

NT

NT

II+IV

E

Oxygastra curtisii

E+N NT

NT

NT

II+IV

E+N

Macromia splendens

E

VU

VU

VU

II+IV

Somatochlora borisi

E

VU

VU

VU

C. syriacum

NT

Ischnura fountaineae

I. graellsii

NT

NT EN

Stylurus flavipes

VU

C. intermedium

I. genei

VU VU

O. boudoti

EN

Platycnemis acutipennis E

P. subdilatata

End. Red Eur. Med. HD List

NT VU

I. hastata

VU

I. intermedia

NT

NT

NT

Nehalennia speciosa

NT

NT

CR

Pyrrhosoma elisabethae E

CR

CR

CR

Aeshna viridis

NT

Anax immaculifrons

VU

Boyeria cretensis

E

B. irene

E+N

Gomphus graslinii

E

NT

G. lucasii

N

VU

G. pulchellus

E

G. simillimus

E+N

EN

EN

NT

Brachythemis fuscopalliata

IV

VU

NT

II+IV

VU

Leucorrhinia albifrons

EN

IV

L. caudalis

NT

IV

L. pectoralis

II+IV

Libellula pontica

EN

EN

VU

NT

NT

Orthetrum nitidinerve E+N

VU

Sympetrum depressiusculum

VU

VU

VU

NT

S. (striolatum) nigrifemur Zygonyx torridus

N

International conservation status of species Species are endemic (End.) to Europe (E), north-western Africa (N) or both (E+N). They are assessed for the IUCN Red List (2019) as Critically Endangered (CR), Endangered (EN), Vulnerable (VU), Near Threatened (NT) or Data Deficient (DD), either globally (Red List) or within Europe (Eur.). The inclusion of species in the EU’s Habitats Directive (HD) on Appendix II (species must be included in a national network of protected habitats) and/or IV (survival of national populations must be ensured) is also indicated, as is the Red List status in the Mediterranean Basin (Med.) for selected species.

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ranges may be explained by a combination of only partial colonisation, as well as numerous subsequent local extinctions. Although not all distribution patterns fall clearly into one of the above scenarios (some species may have survived in several areas), in general they explain most of what we see today. The distribution of the three Erythromma species, for instance, suggests that they each developed from a single ancestor that was isolated in three areas: in the south-west (E. lindenii), south-east (E. viridulum) and east (E. najas). Nowadays, these species overlap but cannot interbreed. A similar scenario fits the genus Platycnemis. Populations that diverged genetically in isolation, but are still able to interbreed to variable degrees where they meet, may explain the complexity of Calopteryx splendens subspecies.

12

Maps As a result of the dynamic nature of dragonfly ranges (see: Status) and the recent increase in recording intensity, we have attempted to be as up to date as possible in the species texts and distribution maps. Maps show the general range of species and are intended to facilitate the identification process. For any further details on distribution and precise regional occurrence of species we refer to the Atlas of the European dragonflies and damselflies (see: Literature p. 8). Nonetheless, great gaps in our knowledge remain. The most glaring blind spots are in Algeria (except the extreme northeast) and Russia, although large areas with low research intensity such as Fennoscandia, Spain, Romania, Belarus and Ukraine also require more data. Regions that are important for species of taxonomic or conservation concern, such as the southern Balkans, Turkey, Portugal and Poland, also need more work. Only the triangle formed between Ireland, Germany and Italy is really known well, but even there continuous monitoring is required as species decline or increase. With this in mind, it will be clear that the distribution Presence on the Azores maps in the guide are an estimate (A), Canary Islands (C) and of species’ ranges: data are still Madeira (M) is indicated insufficient for many areas, and here, as is the inclusion in some regional recording schemes the global Red List are only in a preliminary phase of and appendices of preparation. In those regions, the the EU Habitats presented ranges are sometimes Directive (HB) founded on expert judgement. (see p. 11). Thus, valuable new discoveries are possible everywhere. Map key Purple: main area of distribution; dots indicate either isolated populations or single records. Blue: range of non-overlapping close relative, e.g. Gomphus lucasii on G. simillimus map.

Habitat Considering the size of the region covered and the huge range of many species, it is difficult to provide comprehensive descriptions of each species’ habitat. Many species have very specific requirements at the edge of their range but are more tolerant towards the centre. Central to each species’ requirements is, of course, the water in which the larvae breed, but its surroundings are also important as a home to the adult dragonfly. A suitable pond may be devoid of larvae if forest for the adult’s shelter is not available nearby. The primary determinants of dragonfly habitats are, in approximate order of importance: Introduction

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13

G Chalcolestes viridis female ovipositing into a willow twig. Recognising the requirements of both the larva and adult is important in understanding a species’ habitat preferences.

1. Water motion: flowing vs standing The majority of species are either wholly confined to water with a current (streams, rivers) or to still waters (ponds, lakes). This strict dichotomy operates at a high taxonomic level. The families Platycnemididae and Gomphidae, for instance, are almost entirely restricted to running water, while Coenagrionidae and Libellulidae are strongly associated with still-water habitats. These differences may be determined by the higher concentration of dissolved oxygen and the different bottom substrates (e.g. stones, sand) that are associated with moving water; some river species find suitable conditions provided by the motion of the waves along lake shores. 2. Water permanence: temporary vs permanent Many species are intolerant of the desiccation of their habitat, or the temperature fluctuations that are associated with water-level changes. Others, with drought-resistant eggs or larvae, may profit from the warm (and predatorfree) conditions that prevail in shallow, ephemeral pools, stimulating rapid larval growth (e.g. Chalcolestes, Lestes, Sympetrum).

 A mass of Water-soldier (Stratiotes aloides) in a ditch in East Anglia, UK, where it is the main plant used for oviposition by Aeshna isoceles. On the continent this is the exclusive habitat of A. viridis.

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14

3. Vegetation The presence and structure of submerged, floating, emergent or waterside plants provides a complex array of microhabitats for roosting, perching, oviposition and larval survival. Few relationships between plants and dragonflies are as simple as the preference of Aeshna viridis for Water-soldier (Stratiotes aloides) swamps, or that of Nehalennia speciosa for swathes of fine-leaved sedges (Carex). 4. Water chemistry The trophic status and pH of waterbodies influence their bottom substrate, vegetation and water clarity. Extreme conditions such as acidified or eutrophic lakes often support large numbers of only a few species, while intermediate conditions may harbour more diverse assemblages of specialised species. Such chemical balances are highly sensitive to environmental change, making dragonflies of mesotrophic lakes among the most vulnerable in our area. Of course, these four factors, and the diverse variables they encompass, are intricately interrelated, as are the responses of dragonflies to them in their different life stages. Moreover, complex factors such as parasitism, predation and competition between dragonfly species for resources such as food further influence the suitability of a habitat for a species’ survival.

Flight season The life cycle of dragonflies, and thus the period in which the adults are on the wing, is determined largely by larval development. The adult dragonfly can emerge only once its larval development is complete. Larval growth rate is governed by water temperature and availability of food, which depend on habitat. As a result, flight periods differ regionally, but also annually. In the species texts, a general impression of seasonality is given, but it will always be possible to find adult individuals before or after the dates indicated. In temperate regions, the warm season is generally too short to allow the emergence of more than one generation each year, but there is a notable dichotomy between species in which the larvae grow to their maximal size before winter or after it. Once the water warms up in spring, species with mostly full-grown larvae can emerge in great numbers as soon as conditions are favourable. The period in which most adults appear is therefore short and early, on average. Such species are known as ‘spring species’. Larvae that are still small at the end of winter (or have yet to hatch from the egg) cannot emerge until their development is complete. The emergence of these so-called ‘summer species’ is spread out across a longer and, on average, later part of the warm season. Although most species have only a single abundance peak each year, some are most numerous in two distinct periods. There may be various reasons for this. Anax imperator can complete its life cycle in one year and behave like a summer species, or in two years like a spring species. Sometimes both types inhabit the same water. Sympecma species are unusual because they hibernate as adults; the adults die after breeding in spring, and the next generation emerges in late summer and autumn, hibernating over winter. In arid regions (e.g. northern Africa), breeding habitats may dry up in summer. Adults are seen there only in spring when they emerge, they aestivate elsewhere (e.g. in montane forests) and they then return in autumn when rains replenish their habitat. Aestivation is the opposite of hibernation, although siccatation (inactivity during the dry season) is a better term, as it is the unfavourable dry season between emergence and breeding that the adults must survive. Warm conditions may allow fast-growing species such as Sympetrum fonscolombii to have several successive generations in one year. Influxes of this migratory species into central Europe may be early enough in spring to allow the development of a local generation by the end of the summer. The perpetual succession of generations is possible in some species in hot areas, such as at the fringes of the Sahara.

Dragonfly names Some readers will prefer the familiarity of vernacular names, while others favour the international solidity of scientific nomenclature. We therefore provide both types of names. One must bear in mind, however, that any name is susceptible to interpretation and therefore to change. Introduction

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Scientific names Although the taxonomy of European Odonata is generally well resolved, some problems remain and there are often inconsistencies in the usage of scientific names. In recent literature and on the Internet, for instance, the alternative genus names Anaciaeschna for Aeshna isoceles, Lestes for Chalcolestes viridis and Gomphus for Stylurus flavipes are still frequent, as are incorrect spellings of the species names of A. isoceles, Cordulegaster bidentata and Sympetrum fonscolombii. This indicates a lack of consensus, generally brought on by the uncritical practice of those proposing and adopting name changes. Our choices for the affiliation of species to genera and whether certain subspecies are better treated as species are summarised in Appendix 1, while a checklist is provided in Appendix 3. Vernacular names Familiar names are instrumental in promoting dragonflies, and must be as applicable, compatible and usable as possible. The discussion on English names for European species has seen polarisation rather than consensus, producing a fragmentary and debated array. We saw the need to develop standardised international names for all species covered (see: Appendix 2). Sometimes these deviate from names used (or proposed) by the British Dragonfly Society, which have a more regional focus, and we provide those as alternatives. Alternative names for our species used in North America (NA) are also provided.

15

Dragonfly identification In this guide the illustrations form the basis for identifying species, and descriptions and diagnostic tables have been kept to a minimum, focusing on the diagnosis of a species and essential details of habitat, status and range. However, when identifying dragonflies, readers should be aware of their variability, especially in size, colour and markings. This range of variation cannot be covered fully in the artworks, or even in the text, but, despite this, dragonfly identification is not difficult. Once the observer knows the general differences between male and female or young and old individuals, it becomes easy to correct for variation that at first seems confusingly complex (see: Measurements, Field characters, Variation). A lavishly illustrated field guide invites a different usage than a traditional identification manual with dichotomous keys. We expect users to approach species identification mainly through the artwork, comparing the insect in the field with similar illustrations before consulting the text. Tables to genera and species have been designed as reminders of the main characters and groups of similar species, or as simplified tools to select the best way forward where identification is complex (e.g. with many similar species). The tables (or any identification key, for that matter) should not be seen as the rigid presentation of perfectly reliable characters. Such perfection is utopian, considering the variability and complexity of dragonfly characters. Because of this and because of the numerous ways in which characters are presented in the book, we advocate an ‘organic’ approach to species identification. Feel free to browse through the information provided and try different routes from seeing a dragonfly to naming it, e.g. via the artwork or by trying a diagnostic table first. The most diagnostic (and thus decisive) information is always presented in the species texts, rather than in the tables or the annotation accompanying the artwork.

Anatomy and terminology A little knowledge of how a dragonfly’s body works may help the observer to understand its topography. Built as aerial, visually inclined predators, dragonflies have large compound eyes to track their prey, powerful but flexible wings to give chase and strong, forward-directed legs to catch their prey. The male is equipped for the unique reproductive behaviour of Odonata (see: Dragonfly behaviour), with secondary genitalia on the underside of the abdomen base and claspers at its tip to hold the female. The female abdomen tip is either equipped with an ovipositor to insert eggs into plant tissues (Zygoptera and Aeshnidae), or a vulvar scale through which eggs are deposited into the water (all other Anisoptera). Introduction

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leading edge or costa

frons median space

postnodal cross-veins (Px)

pterostigma

Dragonfly anatomy

antenodal subnode cross-veins (Ax) node antehumeral stripes

arculus

Aeshna cyanea mature ?

discoidal field

S1

16

membranule

S2 IR3 vein (with forking)

auricle S3 anal triangle anal loop

cubital crossveins (Cux)

S4

medial supplemental vein (Mspl)

triangle

radial supplemental vein (Rspl)

S5 postocular spots

S6

S1

S7

Lestes barbarus mature ?

S8

S2 quadrilateral S3

thorax

S9 S10

antehumeral stripe middorsal carina

S4 upper appendage

humeral suture

lower appendage S5

pronotum prothorax

occipital triangle antenna

S6

ocelli (enclosing vertex)

Cordulegaster boltonii

interpleural suture

coxa

metapleural suture

S7

frons

metastigma

S8 postclypeus (compound) eye

anteclypeus mandible

S9

clypeus

S10 S10

labrum

upper appendage lower appendage

Leucorrhinia dubia mature ?

vertex frons

? abdomen tip

labrum

/ abdomen tip secondary genitalia

labium tibia (pl: tibiae)

appendage

femur (pl: femora)

tarsus (pl: tarsi)

S8

S9

S10

genital lobe

anterior lamina hamule

valve

sheath (valvifer) ovipositor

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endage

Glossary of terms, abbreviations and synonyms Abdomen Posterior portion of body, comprising ten segments and terminating in appendages. Accessory genitalia See: secondary genitalia. Aestivation Change in adult activity in order to survive summer, e.g. moving to cooler areas. Antonym: hibernation. See also: siccatation. Amber Yellowish or orangey colour typically seen at the wing base of many anisopterans. Amphibiotic Aquatic as a larva, but terrestrial in the adult stage. Anal loop Distinct field of cells in the anisopteran hindwing base; its shape is determined by a vein that loops around the field, starting close to the posterior corner of the triangle and ending close to the wing base. See text on identifying Corduliidae and relatives (p. 30). Anal triangle Triangular field of two or more cells next to the membranule in the hindwing base of many Anisoptera. Andromorph Presence of colours typical of mature male in female. Synonym: androchrome, homeomorph, homeochrome. Antonym: gynomorph. Anisopteran Pertaining to the suborder of true dragonflies (Anisoptera). Anteclypeus See: clypeus. Antehumeral stripes Pale stripes on the thorax in front of the humeral suture. See text on identifying Coenagrionidae (p. 24). Antenodal cross-veins Cross-veins that lie along the anterior wing border between node and base. Abbreviation: Ax, antenodals. See text on identifying Anisoptera (p. 26) and Libellulidae (p. 31). Anterior Lying at the front, i.e. towards the head. Antonym: posterior. Anterior lamina Transverse structure of secondary genitalia that lies in front of hamules. Apex Extreme tip. Antonym: base. Apical At or towards the tip. Antonym: basal. Appendages Extremities at the end of the abdomen. In male dragonflies there are upper and lower appendages, which are used to clasp the female on the head (Anisoptera) or the pronotum (Zygoptera) during mating and when in tandem. Synonym: Anal appendages. Appressed Pressed against the body. Arculus Thick, bracket-like cross-vein that lies centrally in wing near base. Two longitudinal veins branch off the arculus. Auricles Ear-like structures on sides of S2 in males of some Anisoptera.

Ax See: antenodal cross-veins. Basal At or towards the base. Antonym: apical. Bridge space An elongate triangular space on the basal side of the subnode. See text on identifying Libellulidae (p. 31). Carina Keel- or ridge-like structure on thorax or abdomen. Plural: carinae. Clypeus Middle portion of face, between frons and labrum, consisting of a lower/anterior (anteclypeus) and an upper/posterior part (postclypeus). Coloration Pattern of colours and markings. Converge Coming together terminally. Antonym: diverge. Costa Thick vein on leading edge of wing, running from base to tip. Coxa Segment of leg that connects it to the thorax (‘hip’). Plural: coxae. Cubital cross-veins Cross-veins in the wing between the triangle and the wing base. Abbreviation: Cux. See text on identifying Corduliidae (p. 30). Cux See: cubital cross-veins. Denticles Minute black teeth covering parts of the body. Denticulate Bearing denticles. Diapause A state of suspended development that may occur at one or more stages in the life cycle and that typically constitutes an anticipatory response to conditions unfavourable for uninterrupted development. Discoidal cell Conspicuous (group of) cells near wing base, known as the quadrilateral in Zygoptera and the triangle in Anisoptera (see those definitions). Discoidal field Field of cells that extends distally from the triangle towards the wing border. See text on identifying Libellulidae (p. 31). Distal Away from the body. Antonym: proximal. Diverge Going apart terminally. Antonym: converge. Dorsal On or towards the upperside. Antonym: ventral. Dorsal carina Central ridge over length of abdomen upperside. Dorsum Upperside, literally back. Antonym: venter. Eclipse Overlap of structures, so one is concealed by another when seen from a certain direction. Emergence The moment when the dragonfly larva leaves the water, sheds its skin and departs as a flying adult.

17

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18

Exuvia The shed larval skin. Plural: exuviae. Femur Long and relatively thick leg segment above the ‘knee’ (‘thigh’). Plural: femora. Foliation Leaf-like expansion on terminal abdomen segment (‘flaps’). Form Discrete alternative appearances of individuals within populations, especially in colour of females. Frons Dorsal part of face, i.e. the ‘nose’ or ‘forehead’. Fw Abbreviation for forewing. Genera See: genus. Genital lobes Ventral expansions of S2 that in secondary genitalia lie behind the hamules. Genus A category in which species are classified, e.g. Lestes sponsa belongs to the genus Lestes. Plural: genera. Gynomorph Presence of typical dull colours in female. Synonym: heterochrome, heteromorph, gynochrome. Antonym: andromorph. Hamules Grasping organs of secondary genitalia, most clear in Libellulidae where typically they consist of a hook (anterior) and a lobe (posterior). Humeral stripes Black stripes on the humeral suture of the thorax. See text on identifying Coenagrionidae (p. 24). Hw Abbreviation for hindwing. Interpleural stripes Black stripes on the thorax below the humeral suture. See text on identifying Coenagrionidae (p. 24). Iridescent Reflectively glistening. Jizz General appearance or first impression that one obtains of a species in the field. Labium Lip-like structure covering the mandibles on the underside of the head, i.e. the ‘underlip’. Labrum Lower portion of face, a lip-like structure shielding the mandibles from the front, i.e. the ‘upperlip’. Lateral Along the sides. Antonym: medial. Lateral carina Ridge over length of each side of abdomen, separating upper- and underside. Lower appendage(s) See: appendages. Synonym: inferior appendage(s), epiproct (in Anisoptera, which have one lower appendage), paraprocts (in Zygoptera, which have two). Maturation period The immature period before the adult dragonfly is sexually mature; also the time when some dragonflies disperse to new breeding grounds. Medial Along the middle. Antonym: lateral. Medial supplemental vein Longitudinal vein without clear beginning or end points that lies centrally in the anisopteran wing, roughly

below the node. Abbreviation: Mspl. See text on identifying Aeshnidae (p. 28). Median space Field in wing between base and arculus, which is free of cross-veins in all Anisoptera except some Aeshnidae. Membranule Roughly triangular opaque membrane (i.e. not transparent like the majority of the true wing membrane) on posterior side of the anisopteran wing base, largest in hindwing. Mesostigmal plate Dorsal region of the thorax directly behind the pronotum; its structure is distinctive for some zygopteran females. Metapleural stripes Black stripes on the metapleural suture, which is the most ventral and posterior suture on the thorax sides. Metastigma Respiratory opening visible as a spot on side of thorax, just in front of metapleural suture. Mid-dorsal carina Keeled central suture on front of thorax, separating left and right sides. Mspl See: medial supplemental vein. Node Kink or break in leading edge of wing, roughly midway between base and pterostigma. Ocellus Light-sensitive organ appearing as a small single-lens eye; three of these lie between and/or in front of the compound eyes. Plural: ocelli. Occipital triangle Dorsal, triangular portion of occiput (back of head) lying between the eyes in Anisoptera (except Gomphidae). Odonate A member of the insect order Odonata, i.e. a dragonfly or damselfly. Oviposition Egg-laying. Ovipositor Apparatus used to lay the eggs into plant tissue, located beneath the terminal abdominal segments in females of Zygoptera and Aeshnidae (other females have a vulvar scale). Postclypeus See: clypeus. Posterior Lying behind, i.e. towards the rear. Antonym: anterior. Postnodal cross-veins Cross-veins that lie along the anterior wing border between the node and pterostigma. Abbreviation: Px, postnodals. Postocular spots Paired, contrastingly coloured pale markings next to the eyes on the back of the head in many damselflies. See text on identifying Coenagrionidae (p. 24). Pronotal hindlobe See: pronotum. Pronotum A shield-like plate covering the top of the prothorax; the shape of the rear edge hindlobe is diagnostic in many damselflies, especially females. Prothorax Small separate portion of thorax behind the head, bearing the forelegs. Proximal Towards the body. Antonym: distal.

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Pruinescence See: pruinosity. Pruinosity A waxy grey or bluish bloom that develops on various parts of the body as the dragonfly matures; especially noticeable on some male libellulids. Synonym: pruinescence. Pseudopterostigma Replaces pterostigma in Calopteryx females; differs because it is not well marked and is crossed by veins. Pt See: pterostigma. Pterostigma Conspicuously thickened and often darkened area on leading edge of wings near wing tips, present in most odonates. Abbreviation: Pt. Px See: postnodal cross-veins. Quadrilateral Conspicuous four-sided cell (or series of cells) near the base of all wings in Zygoptera. See text on identifying Zygoptera (p. 22). Radial supplemental vein Longitudinal vein without clear beginning and end points, that lies centrally in apical half of the anisopteran wing. Abbreviation: Rspl. See text on identifying Aeshnidae (p. 28) and Libellulidae (p. 31). Rspl See: radial supplemental vein. S1 First abdominal segment S2–4 Second to fourth abdominal segment, etc. Secondary genitalia Apparatus for storage and transfer of sperm, located beneath the male’s abdomen base (S2). The male transfers sperm from his primary genitalia (in abdomen tip) to his secondary genitalia, enabling him to clasp (see: appendages) and inseminate the female at the same time. Synonym: accessory genitalia. Siccatation Change in adult activity, e.g. reduced sexual activity, in order to survive periods of drought. Sub- Prefix indicating proximity to a certain state, e.g. sub-basal is near but not exactly at the base. Subnode Oblique vein that runs down from the node. Subspecies Category in which discrete regional varieties of a species are classified.

Subtriangle Roughly triangular field of one or more cells that lies basal of the forewing triangle in some Anisoptera. See text on identifying Corduliidae (p. 30) and Libellulidae (p. 31). Sutures Fine grooves separating parts of the body, e.g. on the thorax sides and in the face. Tail-light Conspicuous patch of colour near end of abdomen. Tarsal claws Paired hooks at tips of legs. Tarsus Group of small terminal leg segments (‘foot’). Plural: tarsi. Teneral A newly emerged (‘fresh’) adult dragonfly, soft and shiny, without the full coloration of the mature adult. Thorax Middle portion of body, bearing legs and wings. Tibia Long and relatively thin leg segment below the ‘knee’ (‘shin’). Plural: tibiae. Triangle Conspicuous small triangular field of one or more cells near the base of all wings in Anisoptera. See text on identifying Anisoptera (p. 26), Corduliidae (p. 30) and Libellulidae (p. 31). Upper appendages See: appendages. Synonym: superior appendages, cerci. Venation The network of veins in the wings. Venter Underside, literally belly. Antonym: dorsum. Ventral On or towards the underside. Antonym: dorsal. Vertex Raised structure on the most dorsal part of head, enclosed by ocelli. Vulvar scale A lip-, spout- or funnel-like structure below S8 in females, along which the eggs leave the body, present in those species that do not oviposit into plant tissue. Vulvar spine Spine on the underside of S8 at the base of the ovipositor found in some damselflies. See text on identifying Coenagrionidae (p. 24). Zygopteran Pertaining to the suborder of damselflies (Zygoptera).

19

Sexes It is important to be able to determine the sex of a dragonfly, since males and females, even of the same species, often look very different. Males can best be separated by the presence of secondary genitalia. In most cases, these protrude distinctly below the male’s abdomen base (view abdomen from the side). Males also tend to be more colourful, with well-developed clasping appendages at the abdomen tip; females are generally plumper and have saw- or spout-like organs below the abdomen tip, which are used to lay eggs. In any work on dragonfly identification, a strong focus on males is inevitable: because they behave more conspicuously, the majority of individuals encountered are likely to be male. Moreover, many characters, such as pruinosity and bright colours (but also secondary genitalia), are male-specific. Other characters, such as venation and markings, apply equally well to females. Introduction

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mature ?

Characters of male and female dragonflies

Enallagma cyathigerum

male relatively colourful secondary genitalia below S2 mature / (blue form)

abdomen relatively thick in /

ovipositor below S9-10

Sympetrum sinaiticum

20 mature ?

male relatively colourful

secondary genitalia below S2

male appendages more strongly developed

/

ovipositor below S9-10 vulvar spine present in some damselflies

abdomen relatively short and thick in /

vulvar scale below S8-9

vulvar scale below S8-9

Measurements All given measurements are approximate and must be regarded as an indication of scale, not as a species’ total range. The lengths provided are: total (Tot), from head to abdomen tip; abdomen (Ab) only; and hindwing (Hw) length. Data often originate from varied (often limited or invalidated) sources. Furthermore, most species vary considerably in size. Because of this, and because size difference between sexes is limited, we have not provided separate values for males and females, except for the large Cordulegaster species, where differences are more substantial (see: Variation). Field characters As the title implies, this guide is intended for use in the field, since most characters are visible with the naked eye or through binoculars. Some field characters may seem vague or subjective, such as the species’ general hairiness or hue. Such characters describe the ‘jizz’ of a species – the general appearance or first impression in the field. Jizz often determines whether an individual requires closer inspection. With experience, the majority of species can be recognised in the field, simply by sight. Nonetheless, all users, especially inexperienced ones, are recommended to examine specimens in the hand (see: Watching and catching). The most reliable characters are often small and demand detailed examination (see below). In the ‘Field characters’ sections of the species texts we have attempted to compare each species with the most similar ones, frequently referring to other texts for more information. Hand characters Characters of general appearance (colour, markings) that are visible with the naked eye or with close-focus binoculars are treated as field characters (see above). Hand characters include more technical features, mainly sexual characters (male’s appendages and secondary genitalia, female’s pronotum and ovipositor) and wing venation. These often require slight magnification, in which case a 10× hand lens generally suffices. Many difficult characters of venation are required to separate families and genera. These are explained in the Glossary and the section ‘Identification of suborders, families and genera’. Introduction

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Variation In all species, size, colour and markings can vary depending on sex, age and origin. In the species texts, sexual dimorphism and size variation are generally covered under ‘Field characters’, but all other variation is treated under ‘Variation’. Individual variation is substantial, making it impossible to cover comprehensively in illustrations and descriptions. Most confusion originates from age-related variation. Generally bright colours (especially red) and pruinosity develop only as the adult dragonfly matures. All libellulids, for instance, appear yellowish or brownish with black markings at emergence. In many odonates, even the black markings are restricted at emergence. Similarly, wing colour, especially yellowish shades, is often age-dependent; some species lose the amber tones present at emergence, while others develop smoky wings with age. Some variations occur frequently, and have therefore been named. These may be regional in occurrence (see: Subspecies) or are found together within populations (see: Forms).

21

Subspecies In the past, numerous subspecies have been named, often on the basis of unstable characters (e.g. extent of black markings) and without geographic and genetic considerations (e.g. regional variation in blackness may merely reflect climatic gradients). Moreover, many areas and species are poorly known, and variation may even be greater than is currently recognised. We believe that many subspecies will not prove to be genetically distinct entities worthy of recognition once patterns of geographic variation in characters, especially of DNA, are analysed. For this guide, decisions on the inclusion of subspecies could not be taken consistently and have been left to the discretion of the authors. Generally, only diagnosable subspecies, which are reasonably discrete in their characters and geography, are included. Since precise range limits are generally poorly known (if they exist), subspecies are not marked on the distribution maps. Forms While males are rather uniform in their appearance within populations, females of many species can appear more different from one another, most importantly in the presence or absence of ‘male-like’ coloration. It is likely that some of these females, called andromorphs, have obtained their unusual coloration only with great age. However, in Coenagrionidae, female colour forms are genetically determined, independent of age. Clear examples are seen in the red damselfly genera Ceriagrion and Pyrrhosoma, where wholly red andromorph forms exist, as well as wholly black so-called gynomorphs and intermediate (part red, part black) forms. The distinction between andromorphs and gynomorphs can thus be gradual and is not always clear. This variation is taken a level further in the genus Ischnura, where there is both a genetic and a developmental component: discrete, brightly coloured (violet, pink, orange) teneral forms obtain brownish ‘female’ or blue ‘male’ colours with maturity (see Glossary for synonyms in confusing terminology of andro- and gynomorphs).

Identification of suborders, families and genera The insect order Odonata is traditionally divided into three suborders: Zygoptera, Anisoptera and Anisozygoptera. Zygoptera are known as damselflies, but both the Anisoptera in a strict sense and the Odonata as a whole are referred to as dragonflies. This confusing usage is especially prevalent in Europe; throughout the Americas, dragonflies are regarded more in their strict sense. For absolute clarity, members of the two suborders are often called ‘zygopterans’ and ‘anisopterans’, respectively; combined, they are ‘odonates’. Worldwide, the Zygoptera and Anisoptera each number more than 3,000 species, while the Anisozygoptera includes only one Japanese, one Himalayan, and two somewhat disputed Chinese species. These are most like anisopterans, but are named for their damsel-like wings. Separating the two suborders found in our region is the first step to identification. Usually the posture is enough: anisopterans are robust and rest with outstretched wings, while zygopterans are slender and rest with the wings folded together. However, there are exceptions to the Introduction

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Separating suborders of dragonflies and damselflies (Odonata) Hw base shape

Eyes

Wings at rest

Suborder

Similar to Fw base

Widely separated by head

Usually held shut

Zygoptera

Much wider than Fw base

Envelop head and often touch each other Spread out

22

Blue Featherleg Platycnemis pennipes – a typical zygopteran

Anisoptera

Four-spotted Chaser Libellula quadrimaculata – a typical anisopteran

rule: the Chalcolestes and Lestes species perch with half-open wings, as their vernacular name ‘spreadwings’ suggests. A multitude of characters distinguish the suborders, but wing shape and the position and size of the eyes are easiest to recognise (see below). This section provides the tools to separate the higher categories (suborders, families, genera) of Odonata covered in the guide. Characters are compared in tables, and difficult ones (e.g. of wing venation) are explained in the accompanying text, where they are in bold for quick reference. Many groups are easily distinguished by one or more diagnostic characters. These are generally unique features that separate them from other groups in the same table, e.g. if ‘absent postocular spots’ is diagnostic for one group, the other groups may be expected always to possess these spots, unless stated otherwise. Because there are frequently exceptions, even to diagnostic characters, words such as ‘often’ and ‘usually’ are used to indicate where exceptions are possible. Genera can also be identified by reading the genus texts, or by first scanning the artwork and then confirming a suspected identification with the tables.

Identifying Zygoptera All families have a combination of characters that facilitate swift recognition. Because Coenagrionidae, which includes the majority of damselflies encountered in the field, is a large and diverse group, it is easiest to exclude the four other families by their diagnostic features first. Two families with relatively few species are instantly separated from the three other ones by their large size and dense venation. The families can also be separated by the shape of the quadrilateral. This four-sided structure lies centrally near the wing base and is formed by a single cell in all families except Calopterygidae, where it is a narrow rectangular series of many cells. The anterior side of the quadrilateral in Coenagrionidae and Lestidae is so short that the cell appears like a skewed trapezium, or even almost like a triangle in Lestidae. In Euphaeidae and Platycnemididae, the anterior side is about as long as the posterior and therefore the quadrilateral is more or less rectangular.

Introduction

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numerous cross-veins (Ax)

A

wings with stalked bases almost free of cross-veins

B

2 crossveins (Ax) wings gradually tapered to base with many cross-veins

often with pigmented areas on wings

23

Separating families of damselflies (Zygoptera) A

Hw length

Ax in forewing

Wings

23–37mm

12 or more

Often coloured. Gradually narrowed Calopterygidae towards base, with many cross-veins Euphaeidae right up to point of attachment to body

Family

A

11–28mm

2

Always clear. With narrow stalk-like bases that are almost devoid of cross-veins

B

Lestidae Platycnemididae Coenagrionidae

A: Large damselflies Body

Pterostigmas

Ax in forewing

A

Metallic green to blue

Absent in ?, but / with whitish ‘pseudopterostigmas’ (crossed by veins)

18 or more

Calopterygidae: Calopteryx (p. 84)

Pale to dark, becoming grey pruinose

Very long in both sexes, whitish to black

12–14

Euphaeidae: Epallage (p. 93)

B: Small damselflies Diagnostic characters

A

Pt rectangular, longer than high, not diamond-shaped. Many cells in wings pentagonal, not the majority quadrangular. Two longitudinal veins branch off the vein connecting the arculus and subnode, about halfway down that vein. Body often metallic green and/or with bluish pruinosity

Lestidae

Tibiae feather-like: partly pale and often expanded, not dark and entirely thin. Head very wide (3× as wide as long) with pale band across vertex from eye to eye. Quadrilateral is rectangular, not a skewed trapezium

Platycnemididae: Platycnemis (p. 95)

None of the above, but the following combination is distinctive. Pt typically a Coenagrionidae diamond. Most cells in wings quadrangular. Quadrilateral is a skewed trapezium. Body rarely metallic green or pruinose. Head about 2× as wide as long, at most with pale band in front of vertex (on frons) or behind it (or with postocular spots there). Tibiae thin and often dark

Introduction

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very wide head

pterostigma rectangular

quadrilateral rectangular

most cells pentagonal

areas of bluish pruinosity often present

expanded feather-like tibiae

two longitudinal veins branch off vein connecting subnode and arculus

Platycnemididae

24 Lestidae

quadrilateral a skewed trapezium pterostigma diamond-shaped

Coenagrionidae

Separating genera of spreadwings (Lestidae) Pterostigmas

Body

Wings at rest

A

Equidistant from wing tips in Fw and Hw

Metallic green to bronze and/or bluish pruinose

Usually half-spread

Chalcolestes and Lestes (p. 69)

Clearly closer to tips in Fw

Buff marked with glossy brown

Always closed

Sympecma (p. 80)

Identifying Coenagrionidae Aside from the numerous southern libellulid genera, separating coenagrionid genera may give novice enthusiasts most identification problems. Mature males of most species are easily allocated to two groups, based on the presence or absence of postocular spots and bright red coloration, particularly of the eyes. Immature males, as well as females, may be more confusing, but the presence in Enallagma and Ischnura females of a vulvar spine on the underside of S8, at the base of the ovipositor (visible with a hand lens), prevents much confusion. Orange Ischnura females, for instance, may be mistaken for red damsels, but have large postocular spots and a distinct vulvar spine. Pseudagrion and Nehalennia each have only one species that fall outside these groups, but are highly localised in occurrence (see below). The genera of bluets and bluetails (table B) are often mistaken for one another. The thorax markings are informative in most cases. The antehumeral stripes are the pale ‘shoulder’ stripes that lie anteriorly on the thorax. They can be narrow (i.e. narrower than the black humeral stripes below) or wide (equally wide or wider than the humeral stripes). The interpleural stripes are short black stripes on the sutures below the humeral sutures. The appearance of these markings is variable in Ischnura, in part because of the diverse female colour forms; here, the vulvar spine may again aid identification. Introduction

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Separating genera of small damselflies (Coenagrionidae) Diagnostic characters

A

Genus

Postocular spots absent. Eyes and often body red. Note: females especially may have Ceriagrion A no red, but always lack both postocular spots and a vulvar spine Erythromma Pyrrhosoma Not as above. Note: all damselflies with two distinct postocular spots and without red colour belong here, as does any female with a vulvar spine

Coenagrion B Enallagma Erythromma Ischnura

Morocco only. Mature ? combines red face, postocular spots and pruinose body

Pseudagrion (p. 145)

Virtually exclusive to bogs with slender sedges, very local from the Alps north and east. Hw usually less than 15mm. Body bright metallic green. Rear of head bears a pale bow-shaped marking, rather than two distinct postocular spots

Nehalennia (p. 144)

25

A: Red and red-eyed damselflies Diagnostic characters

A

Legs and Pt reddish, not black. Frons with transverse ridge between antennae

Ceriagrion (p. 142)

? body not largely red, but dark marked with blue. / body with blue and green but never red tints

Erythromma (p. 135)

None of the above, but combination of red(dish) body (except in some /) and black legs distinctive

Pyrrhosoma (p. 139)

B: Bluet and bluetail damselflies Diagnostic characters

AnteInterhumeral pleural stripes stripes

? S2 upperside

/ vulvar A spine

Entire upperside S8 in ? black, not Wide blue. ? upper appendages longer than lowers and S10. / has a knob on both sides of the thorax, just behind the pronotum

Present

Black

Absent

Erythromma (p. 135)

Combination of narrow antehumeral stripes and presence of interpleural stripes is diagnostic in most cases

Narrow

Present

Blue

Absent

Coenagrion (p. 120)

/ combines torpedo-shaped black markings on abdomen with a vulvar spine. Diagnostic combination of thorax and S2 markings in ?

Wide

Absent

Blue

Present

Enallagma (p. 117)

Pt of Fw in ? and some / black and white, not uniformly coloured

Variable

Variable

Black

Present

Ischnura (p. 103)

Introduction

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Identifying Anisoptera

26

With a little practice, separation of the anisopteran families becomes second nature. In particular, the three families that are considered as more primitive (sharing certain features of wing venation) are easily distinguished by the eyes: either distinctly separated (Gomphidae), just touching (Cordulegastridae) or broadly confluent (Aeshnidae). As a group, they are most easily distinguished by the shape of the triangles, which is similar in the fore- and hindwings. In the Corduliidae and Libellulidae, the acute corner of forewing triangle points in a different direction, because the triangle is shifted a quarter-turn. Therefore, the long axis of the triangle is parallel to the long axis of the wing in the hindwing, but perpendicular to it in the forewing. Another feature of the ‘primitive’ families is seen in two of the many antenodal cross-veins, which are relatively thick and cross a longitudinal vein, i.e. their anterior and posterior sections are aligned. The other antenodal cross-veins are thinner, and the numerous cross-veins on both sides of that vein are not aligned. In the Corduliidae and Libellulidae, all antenodal cross-veins are of similar thickness and alignment. Males of all anisopterans, except Anax (Aeshnidae) and all Libellulidae, possess an anal triangle, a conspicuous triangular field at the hindwing base next to the membranule, which coincides with a pair of auricles, two ear-like structures on the sides of S2, and often with an angled posterior border of the hindwing. Behaviour is a good aid to diagnose anisopteran families, and is best observed in territorial males or foraging individuals. ‘Fliers’ patrol their hunting or breeding territory constantly, and seldom perch. They rest with a hanging posture, the abdomen raised slightly above vertical at most. ‘Perchers’ make flights from a perch, holding the body well above vertical (unless they are weak or teneral) and often even above horizontal. The genera Pantala, Tramea and Zygonyx are the only fliers among the Libellulidae, while all Corduliidae are fliers. Although Corduliidae and Libellulidae are easily separated by jizz (see artwork), structural differences are either subtle or restricted to males. This includes the diagnostic presence of long and low pale keels on the anterior side of the tibiae in corduliid males (not necessarily on all leg pairs).

Separating families of true dragonflies (Anisoptera) Diagnostic characters

? Anal triangle and auricles

Acute corner of forewing triangle points

Behaviour A

Eyes separated by ridge, not touching

Present

Outward

Percher

Gomphidae (p. 29)

Abdomen often dark with blue spots that are coloured by internal pigments, not pruinosity (i.e. brighter and cannot be scraped off). / with complete ovipositor

Present or absent

Outwards

Flier

Aeshnidae (p. 28)

Eyes only just touching, not widely confluent. / with long spike-like vulvar scale, extending well beyond abdomen tip

Present

Outwards

Flier

Cordulegastridae: Cordulegaster (p. 220)

Hind margin of eyes shortly but abruptly arched at about mid-height. Body often metallic green

Present

Backwards

Flier

Corduliidae (p. 30)

Abdomen often bluish pruinose or red

Absent

Backwards

Percher

Libellulidae (p. 31)

Introduction

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eyes broadly confluent

acute corner of forewing triangle points outwards

Anisopteran families

body often dark with bright spots

eyes just touching

two thick aligned antenodal cross-veins

numerous thinner unaligned antenodal cross-veins

27

Aeshnidae

eyes separate

body dark with yellow bands

Cordulegastridae auricle

anal triangle

Gomphidae

acute corner of forewing triangle points backwards

body often metallic green

Corduliidae

abdomen often pruinose or reddish

Libellulidae

Introduction

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28

Identifying Aeshnidae Separating the genera in this family is straightforward with some knowledge of wing venation. The absence of several (usually two to four) cross-veins in the median space of the wing bases is very apparent because it is in marked contrast to the other spaces at the wing base, which always have cross-veins. The IR3 fork, when present, can be found centrally in the apical half of the wings, as a prominent forking in a longitudinal vein between the radial supplemental vein and pterostigma. There may be one or more (1–5) rows of cells before the radial and medial supplemental veins (the rows lie between these veins and the longitudinal vein anterior to them). Whether the end of the radial supplemental vein is directed towards a part of the wing anterior of the tip (Anax) or posterior of it (other genera) is also informative. Although all aeshnid females lack auricles on S2 and angled hindwing bases, and most have abdomens that are not waisted near their base, the presence of the first character in males is diagnostic for Anax and that of the latter for Brachytron. Aeshnidae wings Images to scale. Examples of features listed in tables are shown. IR3 forked no cross-veins in median space of wing base

no cross-veins in median space of wing base

end of Rspl points towards wing tip

3–5 rows before the Rspl and Mspl

anal triangle and angled hindwing base present in ?

Aeshna

anal triangle absent and base of hindwing rounded also in ? cross-veins in median space of wing base

IR3 forked

Anax

IR3 not forked

double row of cells before the Rspl and Mspl

pterostigma very long no waist in ?

wing often darkened at tips anal triangle present in ?, but base of hindwing rounded

Brachytron

Boyeria cross-veins in median space of wing base single rows of cells before Mspl and Rspl pterostigma almost square

Caliaeschna

Introduction

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Separating genera of hawkers (Aeshnidae) Diagnostic characters

Cross-veins IR3 fork in median space of wing base

A Rows of cells above Rspl and Mspl

None of the following

Absent

Present

3–5

Aeshna (p. 147)

Thorax often uniform, not banded. ? Hw base Absent round, not angled, anal triangle and auricles on S2 absent. End of Rspl points between Pt and wing tip

Absent

4–5

Anax (p. 170)

Wings often darkened at tip

Present

Absent

2

Boyeria (p. 184)

? abdomen cylindrical near base, not waisted. ? Hw base rounded, but anal triangle and auricles present. Pt very long

Absent

Present

1 (2)

Brachytron (p. 182)

Pt only slightly longer than broad

Present

Present

1

Caliaeschna (p. 186)

29

Identifying Gomphidae Although the genera in this family are superficially alike, males can instantly be assigned to the correct genus by the shape of their appendages. Females may pose more problems. The genera Gomphus and Stylurus can be recognised by the narrow, pale central line running down the upperside of the abdomen, but due to variation some familiarity with this feature is required. The anal loop is a distinct field of one or a few cells, which lies between the triangle and anal triangle in the hindwing. If present, it interrupts a straight perpendicular vein that runs from the wing’s hind margin to the most posterior thick longitudinal vein in the wing base. Separating genera of clubtails (Gomphidae) Diagnostic characters

? upper appendages and branches of lower

Anal loop

A

Total length more than 65mm. Triangles in wings of more than one cell. Broad foliations on S7–8

Uppers much longer than lower and parallel, like fingers

Present

Lindenia (p. 218)

Broad foliations on S8–9

Uppers much longer than lower and parallel, with tips curved down like hooks

Absent

Paragomphus (p. 216)

Upperside of abdomen marked All similarly long and with pale line that is much diverging narrower than abdomen is broad

Absent

Gomphus and Stylurus (p. 188)

/ occiput with thorny horns

All similarly long and parallel

Present

Ophiogomphus (p. 202)

None of the above

All similarly long and arched towards each other, with tips meeting, like a grasper

Present

Onychogomphus (p. 204)

Introduction

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30

Identifying Corduliidae, Macromiidae and Oxygastra The family Corduliidae traditionally included genera that are related to Libellulidae, but possess a number of characters (often seen as primitive) that have been lost in that family. Not all genera are closely related, and Macromia and Oxygastra are considered to belong in other families. Separating genera is quite easy, especially in males, but three features of venation require explanation. The forewing triangle and subtriangle (the latter is the triangular field that lies basal of the triangle) can lack a cross-vein, thus being of one cell (1), or be split up into more cells (2–3) by one or more cross-veins. The number of cubital cross-veins in the hindwing is counted between the triangle and the wing base. The anal loop shape is determined by a vein that loops around a distinct field of cells in the hindwing base, starting close to the posterior corner of the triangle and ending close to the wing base. The enclosed area may be boot-shaped (elongate with a distinct heel and toe, as in all Libellulidae), sack-shaped (elongate without a ‘foot’) or not elongate but rather square. Corduliidae wings Images to scale. Examples of features listed in tables are shown. 3 cells in subtriangle and 2 in triangle few Ax (7 here)

Cordulia

1 Cux

Somatochlora

anal loop boot-shaped

2 Cux

anal loop boot-shaped

3 (sometimes 2) cells in subtriangle and triangle

1 cell in subtriangle and triangle

Oxygastra

2 Cux

Epitheca

anal loop sack-shaped

2 Cux 1 cell in subtriangle and triangle

anal loop boot-shaped

numerous Ax (15 here)

Macromia

3 Cux

anal loop rather square

Introduction

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Separating genera of emeralds (Corduliidae), cruisers (Macromiidae) and Orange-spotted Emerald (Oxygastra curtisii) Diagnostic characters

Face

Fw triangle Cux and in subtriangle Hw

Anal loop shape

A ? lower appendage central cleft

Hw base with large, dark brown spot. Body mainly brown, not metallic and/ or black. Abdomen tapers gradually from broad base to narrow tip

Largely yellow

3 (2)

Boot

Absent

Abdomen usually thickest at S5–7, rather than more basally or terminally

Marked yellow

2–3

2 (1)

Boot

Absent

Somatochlora (p. 238)

? lower appendage with upturned teeth not only at tip, but also just before it

All dark

2–3

1

Boot

Present

Cordulia (p. 236)

? S10 with high pale crest on upperside

All dark

1

2

Sack

Present

Oxygastra (p. 232)

Total length 70mm or more. 13–15 Ax in forewing, not 7–10

Largely yellow

1

3 or Square more

Absent

Macromia (p. 234)

2

Epitheca (p. 250)

31

Identifying Libellulidae Separating genera in this family relies mostly on wing venation, and may be difficult. Only the genera in bold in the table on the next page are widespread, and their identification is straightforward if two features are combined. (1) The possible blackish spot at the hindwing base is very dark brown and should not be confused with amber at the hindwing base, which may form a marked yellow to brown-orange patch covering a considerable part of the wing (e.g. coming near triangle). (2) The number of antenodal cross-veins in the forewing is counted along the anterior wing border between the base and node. The last (i.e. most distal) antenodal cross-vein in the forewing is closest to the node and can be complete like all other antenodal cross-veins, crossing a longitudinal vein, or incomplete, with only the anterior half present (indicated as ½). The numerous additional genera are found only in southern regions, or as vagrants, and mostly have just one or two species in our area. For convenience, they are split into four groups on the same criteria, and compared further in separate tables, where in most cases they are separated by a combination of venation features. The forewing discoidal field extends distally from the triangle and has a distinctive number of rows of cells at its base (1–4), and may narrow or widen towards the wing border. The forewing triangle can lack a cross-vein, thus being of one cell (1), or be split up into more cells (2–3) by one or two cross-veins. Basal of the forewing triangle lies the subtriangle, a roughly triangular field that may consist of one or more cells (1–6). There may be a single (1) or double (2) row of cells before the radial supplemental vein (Rspl) – the rows lie between this vein and the longitudinal vein anterior to it. Finally, the genus Libellula is characterised by having additional (two or more, rather than one) cross-veins in the bridge space, an elongate triangular space on the basal side of the oblique vein that runs down from the node. Always remember that venation characters are not written in stone, but in a flexible network of veins that may be altered in development. Exceptions to the stated characters can occur, as is shown by some variation in the tables, but also by the artwork in this book, which is based on real individuals and not on the taxonomist’s ideal of a species. Some examples are shown on p. 34. Introduction

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Separating genera of Libellulidae Blackish spot at Hw base

32

Absent

Present

8½–20 Ax in Fw, often two rows of cells above Rspl

A A Crocothemis Orthetrum Pantala Trithemis Zygonyx

A B Libellula Tramea Trithemis

6–8 Ax in Fw,* usually one row of cells above Rspl

A C Sympetrum Acisoma* Brachythemis Diplacodes* Pachydiplax Selysiothemis

A D Leucorrhinia Brachythemis Diplacodes* Pachydiplax Rhyothemis Urothemis

* Acisoma and Diplacodes occasionally have 8½ Ax.

A: Skimmers without blackish hindwing spot and with 8½ or more antenodal cross-veins in forewing Diagnostic characters

Amber at Hw base

Fw discoidal field

Rows of cells before Rspl

A

No black on legs

Marked

Widens

1

Crocothemis (p. 301)

Last Ax in Fw is complete. ? abdomen often Absent or becomes grey-blue pruinose faint

Widens

1–2

Orthetrum (p. 259)

Hw length 45mm or more. Patrols without perching and does not rest in horizontal position. Abdomen dark with 6–7 pale rings

Absent

Narrows

1–2

Zygonyx (p. 322)

Pt in Hw much smaller than in Fw. R3 very wavy. Two Cux in hindwing, not one. S4–5 with transverse ridges like that on S3

Faint

Narrows

2

Pantala (p. 316)

None of the above

Marked

Narrows

2 (1)

Trithemis (p.304)

B: Skimmers with blackish hindwing spot and 8½ or more antenodal cross-veins in forewing Diagnostic characters

Number of Ax in Fw

Fw discoidal field

A

Two or more cross-veins in bridge space of both wings

12–20

Widens

Libellula (p. 252)

Hw base marked with dark ‘keyhole’. Pt in Hw much smaller than in Fw. Two rows of cells subtending Mspl in Fw

10½–12½

Narrows

Tramea (p. 318)

Total length less than 40mm. Face dorsally often with metallic lustre

9½–12½

Narrows

Trithemis (p. 304)

Introduction

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C: Skimmers without blackish hindwing spot and with eight or fewer antenodal cross-veins in forewing Fw discoidal field

Fw triangle

Subtriangle

A

Vagrant from North America. 1 Face white, contrasting with dark metallic base of frons. Hw base yellow, crossed by two dark bars

3

2

3

Pachydiplax (p. 315)

Abdomen often red

½

3 (2–4)

2

3

Sympetrum (p. 281)

None of other stated characters

½

2

1

1–2

Diplacodes (p. 312)

Diagnostic characters

Last Ax in Fw

S4 with transverse ridge like that on S3. All wings may be extensively marked with brown away from their base. There may be two rows of cells above Rspl

½

3

1

1

Brachythemis (p. 309)

NW Africa only. Abdomen parsnip-shaped, black with fragmented whitish spots

½

2

1

1

Acisoma (p. 314)

Venation very pale, almost invisible

1

2

1

1

33

Selysiothemis (p.313)

D: Skimmers with blackish hindwing spot and eight or fewer antenodal cross-veins in forewing Diagnostic characters

Last Ax in Fw

Fw discoidal field

Fw triangle

Subtriangle

A

NW Africa only. Dark area covering basal third of Hw. Body entirely bronzy

½

3–4

2–3

4–6

Rhyothemis (p. 320)

Entire face white

1

3

2

3

Leucorrhinia (p. 274)

Vagrant from North America. Face white, contrasting with dark metallic base of frons. Hw base yellow, crossed by two dark bars

1

3

2

3

Pachydiplax (p. 315)

NW Africa only. ? abdomen becomes dark blue, not pale blue, pruinose

1

2

1

3

Urothemis (p. 321)

None of other stated characters

½

2

1

1–2

Diplacodes (p. 312)

S4 with transverse ridge like that on S3

½ (1)

3

1

1 (3)

Brachythemis (p. 309)

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Libellulidae wings Examples of features listed in tables are shown on only a selection of genera. Not to scale. last Ax incomplete, only anterior half present (total here: 8½ Ax) the complete distal Ax and 3-celled subtriangle of this individual are atypical of the genus

34 Brachythemis fuscopalliata

Acisoma forewing triangle and subtriangle each of one cell

Rspl subtends single row of cells forewing discoidal field of 3 rows at base

Brachythemis impartita

forewing discoidal field widens

Crocothemis

last Ax complete

forewing discoidal field of 2 rows at base

Diplacodes

Leucorrhinia

numerous Ax (an exceptional 20 here), all complete

Rspl subtends double row of cells

several (4 here) bridge cross-veins

Libellula

single bridge cross-vein

Orthetrum

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forewing subtriangle of 3 cells

forewing triangle of 2 cells

R3 very wavy

2 Cux in hindwing

Pachydiplax

Pantala

Rhyothemis

Selysiothemis

35

forewing discoidal field narrows

Sympetrum

Trithemis

Urothemis

Zygonyx

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Larvae and exuviae When we study dragonflies, most attention is given to the adult insects, as in this field guide. However, to truly understand a species’ ecology or follow its population trends, the larvae and exuviae (the larval skins left after the adult emerges) are much more informative. Indeed, most European species spend more time below the water surface than above it. The larvae, moreover, are ferocious predators like the adults, with diverse appearances and behaviour. All dragonfly and damselfly larvae have mouthparts modified to grasp passing prey with incredible speed, much like the chameleon’s tongue or mantis’s front legs. This labial mask is unique to the insect order Odonata, although the shape varies greatly between species and is therefore a useful character to separate them.

36

G Aeshna mixta.

G Macromia splendens.

All damselfly larvae have three lamellae at the end of their abdomen that function as external gills. These are often leaf-like but differ widely in size and shape, allowing for species identification.

G Enallagma cyathigerum.

G Calopteryx haemorrhoidalis. The larva of Epallage fatime is unique in Europe for its swollen lamellae and the filamentous gills below the abdomen segments, recalling a mayfly larva.

G Chalcolestes viridis.  Epallage fatime.

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The larvae of true dragonflies lack lamellae, having gills in the gut instead. They let water in through the anus for respiration, which can then be expelled with great force, propelling the larva forward.

37

G Aeshna crenata.

G Leucorrhinia caudalis.

G Cordulegaster helladica. After the fully grown larva has crawled out onto a rock, branch or other bankside structure, the larval skin bursts open and the adult emerges, as here in this Onychogomphus uncatus.

G Onychogomphus uncatus.

A freshly emerged adult (described as ‘teneral’), like this Epitheca bimaculata male, will rest for some time to harden out before flying off, leaving the larval skin (known as an ‘exuvia’) behind. Multiple exuviae are often found piled on top of each other at suitable emergence sites, as with Trithemis annulata here.  Epitheca bimaculata, male.  Trithemis annulata.

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Regional guide to dragonflies

38

This book aims to make it easier to identify dragonflies in the field, and also to enable people to encounter the dragonflies themselves by suggesting some good places to visit. The regional section is not intended as a detailed site guide, nor as a comprehensive discussion of biogeography. Instead, it sketches the local flavour of rich but sometimes elusive faunas and the character of the landscapes where they occur. The highlighted sites are only a fraction of the best habitats. Many are unknown and will yield new discoveries, and the most interesting sites may well be those that are not mentioned. Moldova, for instance, is not treated, simply because very little information is available. Currently, some 35 species are known from this country, with up to 20 more likely to be discovered. At least 84 species are known from the European part of the Russian Federation, but aside from Kaliningrad, an exclave on the Baltic Sea between Poland and Lithuania where 60 species occur, this vast area is poorly explored. In the south-east, for example, Mediterranean species such as Epallage fatime and Cordulegaster picta occur, and the only European populations of 2 3 4 12 the Asian species Sympecma gobica, 13 1 Coenagrion glaciale, Coenagrion 14 5 15 6 7 ecornutum, Somatochlora graeseri and 10 11 8 16 9 20 21 Sympetrum tibiale (not included in the 19 17 18 22 33 27 book) are also found. 26 28 29 23 25

24

30 35

39

40

41

31 34 32 36

42

37 38

Key and order of entries 1 Denmark 2 Norway 3 Sweden 4 Finland 5 Ireland 6 Great Britain 7 Netherlands 8 Belgium 9 Luxembourg 10 Germany 11 Poland 12 Estonia 13 Latvia 14 Lithuania 15 Belarus 16 Ukraine

17 France 18 Switzerland 19 Austria 20 Czech Republic 21 Slovakia 22 Hungary 23 Portugal 24 Spain 25 Italy 26 Slovenia 27 Croatia 28 Bosnia & Herzogovinia 29 Serbia 30 Montenegro 31 Kosovo

32 North Macedonia 33 Romania 34 Bulgaria 35 Albania 36 Greece 37 Turkey 38 Cyprus 39 Canary Islands, Maderia and the Azores 40 Morocco 41 Algeria 42 Tunisia

Denmark, Norway and Sweden Scandinavia contains a wide variety of habitats, from icy mountains and tundra in the Scandes, through vast moraine landscapes and down to fertile clay and sandy areas in the south. Most of the region is covered with boreal coniferous forest, where peatbogs constitute up to 30% of the area. Here, water is present everywhere, in bog pools, lakes and rivers. The southern part of the region is drier, as many lakes and wetlands in the agricultural regions have been drained. Many of the remaining lakes in the south are home to strong populations of a wide variety of species, many of which are threatened in other parts of Europe, such as Aeshna viridis, Leucorrhinia albifrons and L. pectoralis. A typical species composition in many Scandinavian waters is Lestes sponsa, Enallagma cyathigerum, Aeshna grandis, Cordulia aenea, Libellula quadrimaculata, Leucorrhinia dubia, Leucorrhinia rubicunda and Sympetrum danae. In forested areas, Coenagrion

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hastulatum and Aeshna juncea can be added to this list. The fauna of the rivers resembles that of the lakes, but Calopteryx virgo, C. splendens, Platycnemis pennipes and sometimes Gomphus vulgatissimus are characteristic of southern Scandinavian rivers. In forested areas, Onychogomphus forcipatus and Cordulegaster boltonii appear, as does C. virgo on its own. Forty-eight species are currently known from Norway, Bog in the tundra of Abisko National Park, Sweden. 59 from Denmark and 64 from Sweden, with southern species like Erythromma all five Leucorrhinia (albifrons, caudalis, dubia, viridulum, Anax imperator, Crocothemis erythraea pectoralis, rubicunda) may occur in the same lake. and Sympetrum fonscolombii expanding and more A beautiful but hard-to-find locality is expected to be found. Mårdsjön, near Norrtälje, in Uppland (Sweden). In the tundra, at or above the mountain birch A visit there can yield up to 17 species. In many forest level, few species are able to breed. Aeshna lakes in this area, Water-soldier (Stratiotes aloides) caerulea, Somatochlora alpestris and Leucorrhinia growth enables Aeshna viridis to breed, and rubicunda may be found in good numbers. The floating Sphagnum facilitates large populations elusive Somatochlora sahlbergi can be observed of Aeshna subarctica. Coenagrion johanssoni at palsa mire lakes around lake Dávvajávri often occurs with the latter species. Tåkern and north-east of Kiruna (Sweden) and along the Hornborgasjön, shallow, reed-filled ‘bird lakes’ coastal road west of Bugøynes on the southern in southern Sweden, are interesting habitats to shore of the Varanger Fjord (Norway). visit. Here, one can find large populations of Within the boreal forest, boggy areas normally species such as Coenagrion armatum, Aeshna support only four species: Aeshna caerulea, serrata and Somatochlora flavomaculata, as well A. juncea, Leucorrhinia dubia and L. rubicunda. as vast numbers of Sympetrum species late in Localities with some sedges growing among the season. Forty species have been observed the Sphagnum and with only relatively small in the Tåkern area alone, including Epitheca open water surfaces contain more species, bimaculata. A well-known locality for this rare such as Coenagrion hastulatum, C. johanssoni, species is Dagstorpsjön, north of Höör, in Scania Enallagma cyathigerum, Aeshna grandis, Libellula (Sweden). In this mesotrophic brown-water quadrimaculata and sometimes Cordulia aenea. lake, the population may number thousands A good starting point for those intending to of individuals in a good year. The best way to explore these northern habitats is Abisko, west observe them is by boat or canoe. of Kiruna (Sweden), where there are hiking paths A particularly interesting habitat is the coastal leading into the Sarek National Park. brackish waters of the Baltic Sea, perhaps best The western coast of Norway between explored in northern Uppland, near the small Trondheim and Bergen has a relatively mild village of Hållnäs. The land has risen by about Atlantic climate and some interesting small a centimetre per year since the last Ice Age, and bogs and lakes where Pyrrhosoma nymphula, small, shallow lakes are continually forming. Somatochlora alpestris and S. arctica occur, as Here, you can find Brachytron pratense, Aeshna well as dark populations of Sympetrum striolatum serrata and Orthetrum cancellatum. Other that are sometimes treated as the distinct species lakes in this area house large populations of S. nigrescens. Some bog lakes in south Sweden Somatochlora flavomaculata. The Baltic island of are home to large numbers of species. All six Öland is special for its alvar, a limestone plateau northern Coenagrion (armatum, hastulatum, with only a very thin layer of topsoil. Large johanssoni, lunulatum, puella and pulchellum) and populations of Orthetrum cancellatum, and up

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40

to six species of Sympetrum, are found in the waters here. A good point to start exploring is Möckelmossen, between Resmo and Stenåsa, where several footpaths lead into the area. Wetlands on Gotland often have good populations of Aeshna isosceles, and the rare Nehalennia speciosa was recently rediscovered here. Both Baltic islands have had an influx of species from the south and the east during the last ten years, such as Sympecma paedisca, Sympetrum fonscolombii, Erythromma A bog lake in Nuuksio National Park, Finland. viridulum and Anax imperator. marshes cover large areas, especially in the north. Lakes in the south of the region may contain The abundance of wetlands and suitable dragonfly Gomphus vulgatissimus in large numbers. habitats is stunning. Algunnen, in Småland (Sweden), and many Currently, 62 species are known, the latest lakes in the area of Ry, on Jutland (Denmark), additions being Aeshna affinis (2008), Anax are good habitats to explore. In the latter area imperator and Sympetrum pedemontanum (both more than 30 species are found, including strong 2010), Lestes virens and Sympetrum fonscolombii populations of Aeshna subarctica and Libellula fulva. Having recently colonised northern Jutland, (both 2011), Anax parthenope (2013), and Anax Aeshna serrata can be found at several clear lakes ephippiger and Stylurus flavipes (both 2014). No fewer than 25 species have been found north of with large reedbeds, such as Han Vejle. Rivers the Arctic Circle, and 16 species north of 68°30’N. well worth visiting include Susån at Naestved, Aeshna caerulea is a dominant species in Lapland. on Zealand (Denmark), and the two rivers Emån For the enthusiast with less specialised quarry, any and Alsterån, between Kalmar and Oskarshamn random drive along smaller roads in the interior of (Sweden). Here, Calopteryx splendens and Finland is likely to lead to good dragonfly habitats, C. virgo, Platycnemis pennipes, Pyrrhosoma nymphula, Gomphus vulgatissimus, Somatochlora such as small boggy ponds, bays of lakes or small streams. The south-eastern corner of the country metallica and Libellula fulva are found. In (north and east of Hamina) has the richest fauna northern Sweden, the rivers harbour mostly Calopteryx virgo, Onychogomphus forcipatus and (at least 45 species). Areas surrounding lakes Lohjanjärvi and Enäjärvi (50–80km west of Cordulegaster boltonii. Ophiogomphus cecilia is found in the Råne älv in the very far north. Some Helsinki) are especially recommended. The highest number of species is to be found in the second good areas for those wishing to explore these half of July, the most common species being northern rivers are found around lake Siljan in Coenagrion hastulatum. Dalarna (Dalecarlia), in the Klarälven area in The hope of seeing such rare species as Aeshna Värmland, or the Ljusnan area in Hälsingland. G Sahlén crenata, A. serrata and Somatochlora sahlbergi lures dragonfly enthusiasts to Finland. The first Finland two are quite easy targets. A. crenata can best The Finnish landscape is characterised by the be found along Salpausselkä ridge about 100km mixture of forest and water. The coniferous forests, north of Helsinki, where retreating glaciers left which cover two-thirds of the land area, are blocks of ice that formed deep lakes. Many interspersed with countless lakes, ponds, rivers and of these kettle lakes hold strong populations, streams. Lakes abound in central and south-eastern such as Santastenlammi on the road between Finland in particular, and to the south-west lies a Tammela and Loppi. Other recommended sites complex archipelago of small islands. Bogs and are 20–25km north-west from the centre of

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Helsinki in the Nuuksio uplands, where the species inhabits several small forest lakes with variably broad boggy borders, both inside and outside the boundaries of Nuuksio National Park. The best place to see A. crenata is perhaps the forest area between lakes Bodominjärvi and Nuuksion Pitkäjärvi, where lake Pikku Sorlampi in particular offers good observation conditions. The species emerges in early July and is on the wing until the first week of September. Other interesting boreal species in Nuuksio, where 35 species are known, include Coenagrion johanssoni, Aeshna subarctica, Leucorrhinia albifrons and L. caudalis. Aeshna serrata occurs along the entire southern and western coast. It breeds mainly in brackish water, in inlets with rich aquatic vegetation. It can easily be seen at Uutela, a small nature reserve to the east of central Helsinki. Other localities worth visiting can be found around Lupinlahti Bay, in Hamina, and between Tvärminne and Hanko. Interestingly, Orthetrum cancellatum, Sympetrum striolatum and the very rare Nehalennia speciosa are also largely restricted to brackish-water habitats in Finland. Somatochlora sahlbergi is best searched for on the fjell plateau south of Nuorgam (Utsjoki municipality), Finland’s northernmost village, being abundant at several deep lakes here just east of lake Skáidejávri. Laassavaara fjell, south-east of lake Kilpisjärvi (Enontekiö municipality) in the country’s extreme north-west, is another good place to find this rare species, which breeds here in several small to medium-sized mire lakes that support sedges and cottongrass. M Hämäläinen

Ireland Ireland’s climate is temperate due to the moderating influence of the surrounding Atlantic Ocean and the Gulf Stream. The whole island was covered by ice at the height of the last Ice Age, and consequently much of the present-day land surface is covered by glacial deposits, which impede drainage. The centre of the island is a low-lying limestone plain, with numerous wetlands in the form of raised bogs, fens and lakes. Upland areas (which reach only moderate altitudes) are mainly found around the periphery of the island. They experience significantly cooler, duller conditions and dragonflies are rare above 200m. Thirty-two species have been recorded in Ireland; 24 of these are resident species, but four species have not been recorded since the early

20th century. Anax imperator and Aeshna mixta have colonised since 2000. Dragonflies are most abundant in the natural fens and lakes of the central plain and the drumlin belt (low, whale-backed hills of glacial drift) in the northern half of the island. Between eight and ten species are found at many sites, with the best locations boasting 13 or 14. Aeshna grandis, Brachytron pratense and Coenagrion pulchellum are the most characteristic species. The speciality of these wetlands is Coenagrion lunulatum. Raised bogs were once common in the large river valleys and lake basins of the central plain, but most have been harvested for their peat. Intact raised bogs support only a few acid-water species (e.g. Aeshna juncea and Pyrrhosoma nymphula), but where bogs were cut by hand for fuel (a dying occupation today) diverse and interesting wetlands have been created. Good examples of this habitat can be seen at the Montiaghs Moss, near Lough Neagh, in Northern Ireland, and in the Boora Parklands, near Tullamore, Co Offaly. Brachytron pratense, Coenagrion pulchellum and Sympetrum danae are usually common on all these sites. In the south-west, the bogs and small lakes in the region of Killarney support the only Irish populations of Somatochlora arctica and Cordulia aenea. These are best seen in the Killarney National Park. Ephemeral wetlands in the karst regions of western Ireland, known as turloughs, are the most distinctively Irish dragonfly habitat. These are the principal habitat for Lestes dryas. Shallow, base-rich but naturally oligotrophic lakes are a feature of western districts, and are the main habitat of Orthetrum cancellatum in Ireland. B Nelson

41

Great Britain Comprising England, Scotland and Wales, Great Britain is on the northern edge of the distribution of a number of species that are common in continental Europe. In general, the species diversity diminishes as one travels north. There are greatly differing habitat types, from lowland grazing marshes close to or at sea level, through to moorland and highland pools that support some regional specialities. In recent years, an increasing number of migration events has resulted in the colonisation of new species from mainland Europe, as well as a northerly range expansion for others. Of particular note are Erythromma viridulum, which

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42

has colonised much of southeast England in the last two decades, and Lestes barbarus, which was first recorded in 2002 and now occurs along the east coast of England. Although there were only three records prior to 2009, Chalcolestes viridis is now established and not uncommon in south-eastern England, especially Essex and Suffolk. Coenagrion scitulum was deemed extinct in the 1950s, but was refound recently and proven to breed in 2010. The first record of Sympecma fusca (2008) may be the prelude to colonisation by a species Habitat of Coenagrion mercuriale in the New Forest, UK. that is currently expanding northward in mainland Europe. At least 58 species Despite its relative abundance on mainland are now recorded from Great Britain in total, the Europe, Gomphus vulgatissimus has a restricted latest addition being Somatochlora flavomaculata distribution in Britain. One of the best places (2018), although 13 species are not considered to observe the mass emergence of this species established yet. in early summer is along the River Thames, In southern England the most notable habitats, between Goring and Pangbourne. Libellula fulva with the highest number of species, are associated is another species restricted to a small number of with lowland heaths and bogs. Thursley river catchments in southern and eastern England. Common, near Guildford, has the highest On the River Arun, in Sussex, it can be found in number of recorded species (28) anywhere in some abundance, along with G. vulgatissimus and Great Britain. These include strong populations of Brachytron pratense. The appearance in recent Cordulia aenea, Ceriagrion tenellum, Orthetrum years of L. fulva in new areas suggests that this coerulescens, Somatochlora metallica and species is starting to increase its range. Sympetrum danae. A nationally important range Aeshna isoceles is a scarce species in Great of heathland and bog habitats can also be Britain, and is restricted to a small number of found in the New Forest, in Hampshire. Here, sites in Suffolk and Norfolk, where it is mostly the trampling of pond and stream margins by associated with Water-soldier (Stratiotes aloides). livestock creates a unique microhabitat that is The low-lying grazing marshes associated with favoured by Coenagrion mercuriale, and is one the Norfolk Broads, in eastern England, are the of its major strongholds in southern England. best places to see this species – it breeds in good At the Crockford Stream in the New Forest, numbers in Upton Fen, a Norfolk Wildlife Trust where high numbers can be found, a study using Reserve. The main strongholds for Lestes dryas marked individuals has been carried out to assess are also in eastern England. It can be particularly dispersal of this species. The Mynydd Preselli in abundant where it occurs, especially on ditches Pembrokeshire (south Wales) also support some on the grazing marshes of Essex and Kent. In of the strongest populations of this species, which Norfolk, it can be found breeding on shallow occurs on small streams and runnels draining the pingoes, ponds that were formed after the retreat moorland bogs, grazed by livestock. In contrast, of the last Ice Age – Thompson Common and it has also been found along some stretches of the nearby Frost’s Common, for example, hold the rivers Test and Itchen, draining the chalk significant numbers. in Hampshire, where it favours adjacent ditch Many British species have shown a northward systems on the grazing marshes. range expansion since around 2010. Leucorrhinia A small number of lowland river systems dubia has, however, suffered a contraction of its in England and Wales support scarce species. range. The most southerly site is now at Chartley

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Moss, in Staffordshire. This restricted-access site is possibly the best example of schwingmoor habitat anywhere in Britain, and supports a substantial population of this species, which breeds on pools that have dense floating masses of Sphagnum moss. The species can also be found in significant numbers at Whixall Moss, further north in Shropshire, and also at a few sites such as Scaleby Moss, in Cumbria. It is only in northern Scotland that the species becomes widespread. The upland areas of Scotland, while supporting a limited range of species, have significant populations of rare species. The lochans, in the Glen Affric area of north-west Scotland, hold important populations of Cordulia aenea, Somatochlora arctica and S. metallica, as well as Leucorrhinia dubia and Aeshna caerulea. The distribution of S. metallica is an enigma, being centred on two areas in Britain: one in Scotland and the other in southern England. Scotland is also significant for its isolated colonies of Coenagrion hastulatum, which occur at shallow pools, particularly in the area of Loch Garten, in the Spey valley in the Highlands. S Cham

The Netherlands

One of the flattest and lowest countries in Europe, the Netherlands is also one of the wettest. It includes the delta of the Rijn (Rhine) and Maas (Meuse), and large areas lie on clay and peat. The countless waterbodies are artificial: miles of ditches dug to drain the polders and choppy lakes that grew as they swallowed the remnants of exploited bogs. The green flatlands are demarcated in the west by coastal dunes, while moraines and sandy soils give the east a more continental character. Some 71 species are currently known, Anax ephippiger and Coenagrion scitulum being added in 1995 and 2003, respectively. A characteristic Dutch habitat is laagveen: fenlands created by peat-cutting. The alternation of petgaten (where peat was extracted) and legakkers (where peat was dried) creates a patchwork of forest, reedbeds and water, with marshy borders and rich aquatic vegetation. This mosaic harbours species rare in neighbouring countries. Coenagrion pulchellum, Erythromma najas, Brachytron pratense, Cordulia aenea and Libellula fulva are common. Fields of Water-soldier (Stratiotes aloides) are a typical element, harbouring locally large populations of Aeshna isoceles and A. viridis. An easy place to see laagveen is the Kromme

Rade, near Hilversum, a footpath flanked by beautiful ditches where named species are abundant. Leucorrhinia pectoralis is often seen. The Netherlands’ best dragonfly area, however, is undoubtedly the Weerribben (petgaten and legakkers are called weren and ribben here, hence the area’s name). All specialities can be found along the footpaths of the Woldlakebos, although they can also be sought out from the water by renting a boat in Ossenzijl or Kalenberg and exploring the marshes from there. The only Dutch sites for Coenagrion armatum are inaccessible, but the other local speciality, Sympecma paedisca, is common, as are all species mentioned above. Moreover, Somatochlora flavomaculata, Leucorrhinia caudalis and Sympetrum depressiusculum have become common in the last decade. The coastal dunes are a good place to witness dragonfly movements, with easterly winds in midsummer resulting in massive influxes of Aeshna mixta and Sympetrum species. Depending on water levels, the shallow valleys in Voornes Duin and at Hoek van Holland, for example, can harbour good numbers of Lestes barbarus and three other Lestes species, while Aeshna affinis, Sympetrum fonscolombii and S. meridionale are present (and may breed) in some years. Coenagrion scitulum is marching steadily up the coast and is expected to pass north of Rotterdam soon. The large rivers that shaped the Netherlands seem devoid of dragonflies, but most beaches along the Lek, Waal and IJssel, all branches of the Rhine, yield exuviae of Stylurus flavipes and Gomphus vulgatissimus. Absent for most of the 20th century, they recolonised the Dutch rivers in the late 1990s. Diverse areas on the Belgian border are Budel-Dorplein, near Weert, and the Plateaux, near Bergeyk. These are wonderful to explore, easily yielding 25 species in a weekend. Specialities are Somatochlora flavomaculata, and Sympetrum depressiusculum and S. pedemontanum. Investigating the Meinweg, near Roermond, is equally worthwhile. Nearby is the Roer, with the country’s richest riverine fauna, including all five Dutch gomphids and the strongest Dutch population of Ophiogomphus cecilia. Secluded bogs (veen) and heathy lakes (ven) in the pine forests of Drenthe harbour Lestes virens, Ceriagrion tenellum, Coenagrion lunulatum and Leucorrhinia rubicunda in good numbers, and Aeshna subarctica where there is floating Sphagnum. K-D B Dijkstra

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Belgium and Luxembourg

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Although it is one of the smallest European countries, Belgium has a diverse range of natural regions and habitats. The rather flat northern part is characterised by its many rivers and mostly artificial waterbodies. The north-east has poor sandy soils that are intersected by small river valleys and peaty soils. The Ardennes, in the south, is a heavily wooded hilly region, with many clear, sparkling rivers, and with peat moors on the highest tops. This is also the wettest and coldest part, with a continental character. Some 71 species are known, with Orthetrum albistylum added only in 2016. The Kempen (Campine region), in the northeast, has the greatest dragonfly diversity of the Low Countries, with up to 48 species recorded at a single locality. The best place to start a visit is Den Diel, at Mol-Postel, or the Hageven, in Neerpelt. This region, which extends into the Netherlands (see Plateaux, above), includes a mosaic of mesotrophic vennen (heathy lakes), extensive fish ponds and rivulets. Sympecma fusca, Brachytron pratense and Libellula fulva are some of the more interesting spring species, which in summer are replaced by Sympetrum depressiusculum, S. pedemontanum, Orthetrum coerulescens and Ceriagrion tenellum. Further to the west lie the heathlands of Kalmthout and Groot Schietveld at BrechtWuustwezel, which are best visited in spring, when Coenagrion lunulatum and Leucorrhinia rubicunda are abundant. Also worth exploring are the heathlands of the Teut, at Zonhoven, or the Zijpbeek, at Lanaken. Specialities here are Coenagrion hastulatum, Ceriagrion tenellum, Somatochlora arctica, S. flavomaculata and Orthetrum coerulescens. The oxbows typically found along the river Schelde (Scheldt) at the Damvalley near Ghent, or the Creek at Rupelmonde, hold Erythromma najas, Coenagrion pulchellum, Libellula fulva and Brachytron pratense. The Ardennes region is well known for its riverine dragonflies. The most promising rivers are the Ourthe, Lesse and Semois, with many populations of Onychogomphus forcipatus, Calopteryx splendens and Gomphus vulgatissimus. Along the Ourthe, between Barvaux and Durbuy, is the most northerly population of Oxygastra curtisii. In the upper reaches and along most of the tributaries, Calopteryx virgo and Cordulegaster boltonii are quite common. The most interesting high peat moors in the Ardennes are the Plateau des Tailles, near Baraque de

Fraiture, and the Fagnes of Spa-Malchamps. These are wonderful hiking areas. Specialities here are Aeshna subarctica, Somatochlora arctica and Coenagrion hastulatum. In the extreme south, locally called the Gaume, the fauna is unique. Here are found populations of Cordulegaster bidentata at crons (travertine calcareous deposits), e.g. at Buzenol; Coenagrion mercuriale, C. scitulum, Libellula fulva and Orthetrum brunneum in the valley of Laclaireau, between Buzenol and Ethe; a huge population of Somatochlora flavomaculata at Landbruch (Vance); and, with luck, some southern species such as Anax parthenope and Aeshna affinis at several ponds in the region (e.g. Etalle, Latour). The fauna of adjacent Luxembourg is like that of south-east Belgium. The grand-duchy’s and Germany’s only known population of Oxygastra curtisii occurs on the border river Our. G De Knijf

Germany Ranging from the North Sea and Baltic Sea to the northern Alps, Germany offers a wide variety of beautiful and interesting sites for dragonflies. There is an outstanding diversity of habitats: the major rivers Rhein (Rhine), Elbe, Oder and Donau (Danube) and their catchments; the northeastern lowland plains, peppered with lakes; low mountain ranges with springs, rivulets and bogs; and the glacial bog and moor region in the south. With some 81 recorded species (77 of which are considered established), Germany has a typical cross section of central European dragonflies. All dragonfly species are protected in Germany, and collecting specimens or even netting adults is prohibited without special permits. However, exuviae may be collected outside nature reserves. In former times, north-west Germany was especially rich in heathland and peatbogs. Despite widespread agriculture, several good sites for Coenagrion lunulatum, C. hastulatum, Aeshna subarctica and A. juncea still exist in the Lüneburger Heide (Lüneburg Heath), especially around Celle. In May, these habitats produce huge numbers of Leucorrhinia rubicunda and L. dubia. Because of their favourable microclimate, the bogs are often also inhabited by Ceriagrion tenellum, Lestes virens or Orthetrum coerulescens. The streams flowing into the Elbe and Aller rivers are rich in Ophiogomphus cecilia, Cordulegaster boltonii, Calopteryx virgo and Platycnemis pennipes. The local speciality is the strong but

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highly isolated population of Boyeria irene on the river Örtze, discovered in 2010. The species is found mainly between Hermannsburg and the Örtze’s confluence with the Aller, such as the areas just south of Evers and around the bridge east of Winsen. The oxbows in the floodplain of the Aller and the ditches in that of the Weser, around Bremen, are well known for their populations of Aeshna virdis and Coenagrion pulchellum. The Pleistocene lake landscapes of Mecklenburg and Brandenburg are ideal for dragonflies. About 5,000 lakes of different types, as well as pools and wetlands, form a 300kmlong network of habitats between Schwerin and Frankfurt/Oder. Sympecma fusca, Brachytron pratense, Aeshna isoceles, Anax parthenope and Libellula fulva are found along the reed belts of the lakes and channels. Together with Epitheca bimaculata, they can be observed during a boat excursion on the lakes around Templin. In the Schorfheide-Chorin UNESCO Biosphere Reserve, 40 dragonfly species can be recorded over a typical sunny weekend in June. Male Onychogomphus forcipatus can be found on beaches of the clear Gollinsee lake. Small shallow lakes, with rich submerged vegetation and swampy banks, have a dragonfly fauna of up to 38 species, including Leucorrhinia caudalis, L. pectoralis, Sympecma paedisca, Erythromma viridulum and Somatochlora flavomaculata. A typical, easily accessible site is Steisssee lake near Metzelthin, in the western Uckermark. Pools in peatbogs harbour populations of Leucorrhinia albifrons and Aeshna subarctica. For part of the 20th century, the Oder and Spree were the most important German refuges for Stylurus flavipes, along with Gomphus vulgatissimus and O. cecilia. Today, however, the Elbe is important for these gomphids. Aeshna affinis and Lestes barbarus breed in temporary waterbodies adjacent to the river. Populations of Aeshna viridis reproduce in masses of Watersoldier (Stratiotes aloides) growing in ditches and ponds in the former floodplain of the Havel, west of Berlin, and in peat-cutting areas in the Peene valley, near Anklam. Drainage ditches in the tributary of the river Unstrut, between Sangerhausen and Gotha, are good sites to look for Coenagrion mercuriale and C. ornatum, which are at the edge of their distribution here. In the high altitudes of Erzgebirge, Thüringer Wald and Harz, peatbogs are home to Aeshna

subarctica, Somatochlora alpestris and S. arctica. Interesting communities of dragonflies are developing in the former brown-coal mines in the region between Cottbus and Halle: Lestes virens, Ischnura pumilio, Orthetrum coerulescens, O. brunneum, Sympetrum fonscolombii, Leucorrhinia dubia and L. albifrons can be encountered at large new lakes, temporary waters and seepages. In Nordrhein-Westfalen (North RhineWestphalia), peatbogs provide some excellent sites. Good examples are the Heidemoore, small moorland ponds in between inland sanddunes, such as in the Wahner Heide, near Köln (Cologne), or in the Schwalm-Nette nature reserve, near Viersen. Schlatts are small lakes in western Niedersachsen (Lower Saxony) that developed where postglacial ice blocks were left behind. The best example are the Dianaseen, near Cloppenburg. Only a few relicts of the formerly huge peatbogs, the Dosen, have been preserved, as the Esterweger Dose in the Emsland. In these habitats, Lestes virens, Ceriagrion tenellum, Coenagrion lunulatum and Aeshna subarctica may have huge populations. Kuhlen (oxbows and swamps), such as the Fleuthkuhlen, near Geldern, are typical of the Niederrhein (Lower Rhine) valley. Brachytron pratense and Libellula fulva are specialities here. Good sites for Sympetrum depressiusculum are the Dülmener and Ahlhorner Fischteiche, both extensive pond systems for carp breeding that regularly dry out in winter. At the border of Rheinland-Pfalz (RhinelandPalatinate) with Luxembourg, the river Our harbours the only known German population of Oxygastra curtisii. The Oberrhein (Upper Rhine) lowland plains between Karlsruhe and Freiburg are a hotspot for Coenagrion mercuriale. Southern species such as Anax parthenope, Crocothemis erythraea and Orthetrum albistylum can easily be found here. Springs and rivulets on the western slopes of the Schwarzwald (Black Forest) are habitats for Cordulegaster bidentata and C. boltonii. The High Rhine between Bodensee (Lake Constance) and Basel is the only German breeding site for Gomphus simillimus, and Boyeria irene may be observed at the shores of the Bodensee. Near Ulm, Schmiechener See, a shallow seasonal lake, is a perfect breeding site for Lestes dryas and Sympetrum flaveolum, and in some years even for S. meridionale. Mittelfranken (Middle Franconia) is well known for its populations of Stylurus flavipes,

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arctica, S. flavomaculata, Leucorrhinia caudalis and L. pectoralis, and a diligent searcher may even be able to discover the tiny jewel of this region, the highly endangered Nehalennia speciosa. Ascending to the Alps, the rare Aeshna caerulea can be found at Sphagnum bogs high up around Oberstdorf. F Weihrauch, R Jödicke, A Martens and R Mauersberger

Poland, Estonia, Lithuania, Latvia, Belarus and Ukraine

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Wetland in Chiemgau, south Germany. Gomphus vulgatissimus and Ophiogomphus cecilia, which are found in larger rivers, such as the Regnitz. An outstanding large population of Coenagrion ornatum breeds in the ditches of the river Altmühl catchment, around Ansbach. Gravel pits and larger oxbows in the Donau (Danube) floodplain around Plattling offer the best chance to watch (with binoculars) restlessly patrolling males of Epitheca bimaculata, and to collect its impressive exuviae in May (if one is not deterred by the brambles). Sympetrum pedemontanum is a typical species of the gravel plain around München (Munich), with its ephemeral waters. Here, in ditches and brooks of the Dachauer Moos, a very large population of Coenagrion mercuriale also breeds, together with Orthetrum coerulescens. Numerous fens and moors, many of them nature reserves, are found in the broad, glacially formed belt bordering the Alps to the north, comprising the Allgäu, Werdenfelser Land, Oberland and Chiemgau regions, e.g. Murnauer Moos, Kirchsee or the lake region around Eggstätt and Seeon. Here, a number of specialities are found, including Sympecma paedisca, Coenagrion hastulatum, C. pulchellum, Aeshna isoceles, A. subarctica, Somatochlora

This huge region is dominated by lowlands, with large areas rich in postglacial features. The mountains are limited mainly to southern Poland and Ukraine (the Sudety Mountains and Carpathians). A continental climate predominates. The range of landscapes is extremely large, from boreal forests (taiga) in the north, through temperate forests, up to mountains in the south-west and temperate grasslands (steppe) in the south-east. The distinguishing feature of the region is the abundant and diverse primary water habitats, which are largely well preserved. The most valuable are: lakes of various trophic status (i.e. nutrient enrichment), situated mostly in a broad zone from north Belarus, through south-east Estonia, east Latvia, east and south Lithuania up to north and central Poland; bogs and waterbodies bounded by Sphagnum (with their abundance decreasing towards the south); fens and marshes, both in lake districts and river valleys; and numerous medium and large rivers, which are largely unregulated (less so in the south-west) and frequently associated with an intact array of river valley habitats. At least 80 dragonfly species are known from the region, Lindenia tetraphylla (Ukraine, 2013) and Pantala flavescens (Poland, 2016) being the most notable recent additions. The northern range limits of many Mediterranean species run through this region. Especially noteworthy is the rich and abundant eastern dragonfly fauna. The comparative rarity and declining populations of eastern species in west and central Europe makes this region an important refuge for them. Knowledge of the regional fauna is moderate in Poland, Lithuania, Latvia and Ukraine, and limited in Estonia and Belarus. Closer study of this region will be a major challenge for European odonatologists in the future. Acid waters characterised by Sphagnum moss are abundant in the boreal and temperate

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coniferous forests. In the lake areas, waterbodies and roads in the neighbourhood are also excellent in Sphagnum bogs and/or small forest lakes for this species. The lakes of central and north bounded by Sphagnum harbour Aeshna Poland and the Baltic States are prime habitats subarctica (regularly), Leucorrhinia albifrons for Libellula fulva. Lakes of a lower trophic status, (frequently) and Nehalennia speciosa (locally, and which are frequently rich in submerged, floating only where there is shallow water with slenderand emergent vegetation, especially sedges leaved sedges). Aeshna crenata and Coenagrion (Carex), support interesting Leucorrhinia species. johanssoni occur in these habitats only in the L. albifrons is widely distributed and locally northern part of the lake zone. At small lakes in fairly common in north Poland and the Baltic the peatbog channel 2km east of Golce (north States; L. caudalis is generally more scattered, Poland), numerous N. speciosa are the main but not scarce in the north. L. albifrons is a attraction, and A. subarctica and L. albifrons also characteristic species of spectacular ‘Lobelia lakes’ occur. Such habitats in the Labanoras Regional in north Poland. These are mostly slightly acid, Park and the forests to the south of it (east poor in nutrients and characterised by specific Lithuania) offer a similar fauna, but the local vegetation (Lobelia, Isoetes, Litorella), clear water speciality is A. crenata. In small lakes in northand low concentrations of calcium. Examples east Latvia and south-east Estonia, C. johanssoni are Czarnówek lake, north of Złocieniec, joins this group (but A. crenata is absent), with a or Bieszkowickie lake, near Bieszkowice. large population at Vinnora lake (1.2km southRemarkably, Sympetrum striolatum also occurs in west of the south-west corner of Kerigumäe lake, this rather boreal habitat. In June and early July, Estonia). Also worth visiting are the lakes Mazais sightings of L. caudalis are possible at certain Pai¸kis, along the road to Pskov, 3km south-east lakes, such as Łakie, 5km south of Goscim in west of Korneti, and Lidacis, between Garulis lake and Poland (together with Erythromma lindenii), and Cirgali (Latvia). the south-eastern corner of Šventas, in Gražute Peatbogs and mires to the south of the lake Regional Park of north-east Lithuania (a boat is districts, which are mainly small and in forest, needed at both lakes). Good lakes in south-east are often inhabited by Somatochlora arctica and Estonia are Külajärv, near Plaani (also with L. locally by N. speciosa. Both species are found albifrons), and Vaaba järv, near Kooraste. in south-east Poland, e.g. 1km south-east of Also characteristic of north-east Europe are Hamernia. A classic habitat of S. arctica, small waterbodies with Water-soldier (Stratiotes aloides) pools completely overgrown with Sphagnum, is and the associated Aeshna viridis. Oxbows and found 0.7km south of Majdan Kasztelanski. In other waterbodies north of Khvaensk (Hvoyensk) late spring, swarms of S. arctica may be spotted in Belarus, a viewing platform at the south-east south of Khvaensk (Hvoyensk) in Prypiatskii National Lake in Białowieža Forest, north-east Poland. Park (Belarus), and nearby Simanichy (Simonichi) bog is inhabited by Aeshna subarctica, Leucorrhinia albifrons and numerous Nehalennia speciosa. Forest lakes, frequently with floating vegetation, e.g. in the Drawienski National Park (north-west Poland), the Iława Lake District (north Poland), Golosiivski (Goloseevskie) lakes in Kyiv (Kiev) and Supii (Supoy) lake near Yahotyn (Ukraine), yield numerous exuviae of Epitheca bimaculata in May. With binoculars, adults may be seen flying over the lakes in late May and June; forest clearings

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shore of Łukie lake in Poleski National Park and peat excavations north of Dorohucza (east Poland), and the environs of Borne Sulinozo (north Poland) are all hotspots for this species. If you visit the first two localities in May, check the half-open vegetation of slender emergent plants for the elusive Coenagrion armatum. Lowland rivers in a large part of the region have notable populations of gomphids. Ophiogomphus cecilia, found on a wide range of rivers, is most easily seen on smaller watercourses such as those in the Drawienski National Park or the Grabia river (north and central Poland, respectively). The region is the centre of Stylurus flavipes occurrence in Europe. Its exuviae can be collected in large numbers on the banks of many larger lowland rivers, e.g. the Warta, Wisła, Pilica, Narew and Bug (Poland, even in cities such as Warszawa and Poznan); Nioman (Nemunas; Belarus, Lithuania); Prypiat (Pripyat; Belarus, Ukraine); Biarezina (Berezina; Belarus); and Dnipr (Dnepr) and Vorskla (Ukraine). The rivers and streams in the mountains and foothills of southern Crimea (such as the Alma river and watercourses in the environs of Simferopol) are home to populations of Calopteryx splendens with very small wing spots, sometimes treated as the endemic species C. taurica. In Crimea, it is also worth visiting the waterbodies in the steppe zone for the numerous Lestes macrostigma and Sympetrum meridionale. Special habitats harbour special species: Sympetrum pedemontanum inhabits small watercourses, such as ditches, streams and small rivers, frequently adjacent to or flowing through ponds and marshes. The ditches near the ponds in Chojno-Młyn (west Poland), the Neresla river and neighbouring ditches in Tykocin (north-east Poland), and stagnant waterbodies in a gravel pit in Matuizos (south Lithuania), can provide amazing encounters with this species. Aeshna serrata is found in brackish and fresh waters rich in reeds, which border (or are close to) the Baltic Sea: Saaremaa, an island opposite the Estonian mainland, is a good site. Binoculars are recommended to observe males patrolling along reeds in the sea’s bays and lagoons. They also hunt in forest clearings and edges. In addition to natural habitats, human-made waters are abundant in the region, being important for some species. In Poland, easily accessible sites include the peat excavations near Strzeszynskie lake and the Bogdanka stream

in Pozna (Leucorrhinia caudalis, L. pectoralis, L. albifrons), and 0.8km south-east of Zagłebocze, in the buffer zone of Poleski National Park, where a diligent searcher may find Nehalennia speciosa, Aeshna subarctica, L. albifrons, L. pectoralis and Sympecma paedisca. The last species is locally abundant in the region – in east Poland, for example, huge numbers may be observed in ponds in Zalesie Kanskie, near Rejowiec, or 3km south-west of Brus Stary, in the Poleski National Park. In ponds and gravel and sand pits (but also at natural waterbodies) in south-east Poland and Ukraine, Orthetrum albistylum is widely distributed, having expanded its range in recent times. The south of the region is also especially favourable for species with Mediterranean affinities. The best localities are in Ukraine, where large numbers of Aeshna affinis and Sympetrum meridionale may be encountered without effort. The broad environs of Kaniv, Baryshivka and Yahotyn (central and northern Ukraine) are recommended, also for O. albistylum. R Bernard and P Buczynski

France

France experiences a combination of Atlantic and Mediterranean climates and has diverse landscapes, ranging from sea level to 4,808m. As a consequence, the French dragonfly fauna includes 97 species of diverse origin. Ten species are restricted to south-west Europe, of which Platycnemis latipes, P. acutipennis, Gomphus graslinii and Macromia splendens are known only from the Iberian peninsula and France. A quarter of the species are largely Mediterranean in occurrence, some of them restricted to the Tyrrhenian Islands. Seven species have an African origin, and some of these have recently expanded their range to the north. Different regions support a distinct array of dragonflies. The lowland areas can be conveniently divided into the south (Aquitaine, Mediterranean and Corsica), north-west and central, and northeast mainland. Streams in the Aquitaine and the Mediterranean areas hold most of the south and south-west European specialities. Almost any river has species such as Calopteryx xanthostoma, Platycnemis latipes, P. acutipennis, Gomphus simillimus and Oxygastra curtisii, and the subspecies Calopteryx virgo meridionalis, sometimes in dense populations. Some of these species even colonise standing waters in great numbers. More restricted are two legally protected south-west European endemics, Gomphus graslinii and Macromia

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splendens. Good populations of the former are follow suit, being recorded for the first time at five confined to the region that extends between the places in 2017. Mediterranean and the Massif Central, from the In addition to many Mediterranean species and Rhône to the Gironde. Here, it will be found records of rare species such as Lindenia tetraphylla, without difficulty in slow- and moderate-flowing Brachythemis impartita, Selysiothemis nigra and sandy and stony rivers, where Boyeria irene is also Orthetrum trinacria, the true speciality of Corsica widespread. The best populations of Macromia is the Tyrrhenian endemic Ischnura genei. As in splendens are present in the same area, but this Sardinia and Sicily, it is widespread in all standing species is rare except in some spots in the and slow-flowing waters. The eastern species southern Massif Central and in the Languedoc Chalcolestes parvidens and Somatochlora lowlands, mainly either in naturally calm river meridionalis and the southern Paragomphus genei stretches or where the river is dammed (e.g. on also occur on Corsica, but they are very rare and the river Tarn between Millau and Albi). will be found only by chance or careful searching. On small herbaceous brooks, e.g. with The Loire catchment is the main dragonfly Water Mint (Mentha aquatica), Coenagrion hotspot in western and central France, with good mercuriale, Ceriagrion tenellum and Calopteryx habitats for gomphids. Stylurus flavipes and haemorrhoidalis are widespread, but Coenagrion Ophiogomphus cecilia are most common. The caerulescens is extremely localised. The subspecies best stretches for these two are on the Loire Cordulegaster boltonii immaculifrons can be and Vienne in the Indre-et-Loire department, found east of the Rhône on any small sandy the Loire in the Loiret department and the brook. Larger streams and rivers on the southern Loire and Allier in the Nièvre department. foothills of the Alps and Massif Central down to They can be found alongside other gomphids, the Mediterranean always have good populations such as Gomphus vulgatissimus, G. simillimus, of the subspecies Onychogomphus forcipatus G. pulchellus, the subspecies Onychogomphus unguiculatus, while O. uncatus prefers even more forcipatus forcipatus and, more scarcely, strongly flowing, almost torrential, rivers. O. uncatus. The Parc Naturel Régional de Standing waters are also characteristic of la Brenne, near Chateauroux, is popular with this warmest part of France. Small pools called visitors from northern Europe, but many of its lavognes were constructed on the calcareous numerous ponds are on private land. In addition plateaux to collect rainwater for livestock. They harbour River with shingle bank, Gorge du Tarn, Massif Central, France. good populations of Lestes virens, L. barbarus and Coenagrion scitulum. Along the Mediterranean coast, brackish waters support a vegetation of salt-tolerant plants such as glassworts (Salicornia). These so-called sansouires are widespread in the Camargue (the delta of the Rhône), and are the favoured habitat of Mediterranean and tropical species such as Lestes barbarus, L. macrostigma (found mainly by chance), Anax parthenope, A. ephippiger, Sympetrum fonscolombii and S. meridionale. Trithemis annulata reached the French mainland in 1994 and is now fairly widespread in the south and south-west. The related T. kirbyi is expected to

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cold and torrential to harbour dragonflies, but moderately flowing rivers are suitable. The numerous peatbogs and swamps are full of promise for dragonfly-watchers. Common species such as Lestes sponsa, Coenagrion puella, Enallagma cyathigerum, Pyrrhosoma nymphula, Aeshna grandis, Cordulia aenea, Somatochlora metallica and Libellula quadrimaculata can easily be Lake in the Upper Drugeon Valley, Jura, France. found. Somatochlora arctica is widespread in peatbogs and to the many waterbirds in this scenic area, visitors may be encountered at any may encounter Leucorrhinia caudalis, L. pectoralis suitable location and at any altitude, except in the or Epitheca bimaculata. Pyrenees, where only a few localities are known. Standing waters constitute the most attractive Somatochlora alpestris is much more restricted in dragonfly habitats in north-east France. Ponds altitude and reproduces with certainty only in the with flourishing aquatic plants and reedy fringes Vosges above 800m, and in the Alps within and in more or less forested landscapes in the above the upper tree line, above 1,800m. In the Lorraine and Champagne-Ardenne regions hold southern Alps, an area with a fairly sunny climate, populations of Epitheca bimaculata. In some it can easily be observed some kilometres north localities, more than 500 exuviae have been of the Salèse pass in the Mercantour National collected in one season on a single pond. The Park. While Aeshna juncea, Leucorrhinia dubia mid-May emergence period is the best time to and Sympetrum danae are abundant species in see the species, as adults become very elusive most boggy habitats, in France Aeshna subarctica later. The best spots are in the forested parts reproduces only in the Vosges and Jura above of the Parc Naturel Régional de Lorraine, an 700m, where it is scarce. Coenagrion hastulatum area devoted to balance both human activities is another speciality confined to small populations and nature conservation, especially the Pays in true peatbogs and acidic pools. In addition to des Étangs, in central Moselle department, peaty habitats, spring areas are very promising and the Forêt de la Reine, west of Nancy. and support good populations of Cordulegaster These areas are also noted for Leucorrhinia boltonii. In travertine springs (calcareous caudalis, and more widespread species such as seepages), Cordulegaster bidentata is the main Aeshna mixta, A. isoceles, Brachytron pratense, speciality, also occurring at low levels in the northCrocothemis erythraea, Erythromma viridulum east quarter of France. and Lestes virens. In the small Territoire-de-Belfort J-P Boudot and G Jacquemin department, numerous artificial fish ponds in the Parc Naturel Régional des Ballons des Vosges Switzerland Switzerland is a small, mountainous country, with support Sympetrum pedemontanum. the Alps in the south and the Jura Mountains in Both the north-east Lorraine and north-west the north-west together comprising almost threeAlsace regions lie on red sandstone, particularly quarters of its area. Between these mountain in the so-called Vosges du Nord. The very ranges is the densely populated and intensely sandy rivers and brooks here typically have farmed Swiss Plateau, with its rolling hills, plains good populations of Ophiogomphus cecilia, and some large lakes. together with Cordulegaster boltonii, Gomphus With 77 species, the dragonfly fauna is vulgatissimus and G. pulchellus. comparatively rich, and includes Mediterranean as The mountainous areas have a distinct fauna, which includes boreal species. The most attractive well as alpine elements. The presence of Boyeria irene, at the Vierwaldstättersee (Lake Lucerne), habitats here are acid bogs, but spring areas can Zugersee and Ägerisee, and of Oxygastra curtisii also be promising. Many running waters are too

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at Lago di Lugano and Lago di Origlio, is exceptional for central Europe. The Alpine region, with its fantastic scenery and small lakes, ponds and mires between 1,200m and 2,000m, is most rewarding for dragonfly-watching. However, diversity is reduced to some specialised species found around and above the tree line. Many sites are remote, small and hard to find, and can be reached only on foot, while others are easily accessible by car or public transport. The best season is July and August, but note that dragonfly collecting is not allowed. One of the characteristic habitats of the Jura Mountains are peatbogs, although most have been damaged by former peat exploitation. Lac de Gruère, near Seignelégier, is a small bog lake at 1,000m, set in beautiful surroundings, with 22 species, including Coenagrion hastulatum and Aeshna juncea, typical for the region. The fish ponds of Bonfol, partly situated in woodland, are the best place in Switzerland to see Epitheca bimaculata. The ponds hold more than 40 dragonfly species, including Libellula fulva and Anax parthenope. At the northern foot of the Jura Mountains, on the Swiss Plateau, lies the Lac de Neuchâtel with ‘La Grande Cariçaie’ on its southern border, comprising 800ha of managed reedbeds and sedge swamps, with 45 dragonfly species. Important populations of Sympecma fusca, Nehalennia speciosa, Brachytron pratense and Somatochlora flavomaculata occur here, the latter representing the main food for nestlings of Savi’s Warbler (Locustella luscinioides) during the emergence period. The best site for dragonflywatching is Champ Pittet, near Yverdon. In the German-speaking part of the Swiss Plateau, the semi-urban Greifensee region, near Uster, is another area with a rich dragonfly fauna, thanks to several large ponds that were recently created in the fenland, especially at the southern end of the lake. They have been colonised by more than 35 species. As well as common lowland fauna, rare and Mediterranean species such as Orthetrum albistylum, Sympetrum fonscolombii and Crocothemis erythraea may be seen. Tributary streams support Calopteryx virgo and C. splendens, and from the outlet, the Glatt river, Gomphus vulgatissimus and Onychogomphus forcipatus emerge in good numbers every year. Within the Alps, the Flumserberg region above the Walensee, in the east, is a rewarding

place to find some of the typical alpine species. Alp Panüöl and Alp Fursch, with their vegetated ponds of different sizes, just above the tree line, are within walking distance of the Maschgakamm cableway station. Aeshna juncea, A. caerulea, Somatochlora alpestris and Leucorrhinia dubia are most characteristic here. At the three small lakes of Seebenalp, 11 species have been recorded, with strong populations of Somatochlora metallica and Enallagma cyathigerum, the latter being the only damselfly at this altitude. The Upper Engadine, a wide, high-altitude valley in the Grisons, is another excellent place for watching alpine dragonflies. The small bog lake of Lej da Statz, near the tourist resort of St Moritz, and its woodland surroundings interspersed with boggy clearings support three Somatochlora species, together with Aeshna caerulea, A. subarctica, Leucorrhinia dubia and Coenagrion hastulatum. A similar fauna may be found further up in the valley, e.g. in the bogs around Maloja and Lago Cavloggio. In the Lower Engadine, near Scuol/Tarasp, at the small lake named Lai Nair because of its black water, 14 species may be encountered. Quite close by, just beyond the Swiss border, above Nauders in Tyrol, another black-water lake known as Schwarzer See is worth a visit, as is the Grüner See and a number of nearby marshy forest clearings. Somatochlora arctica, S. alpestris, S. metallica, Aeshna grandis, Erythromma najas and Coenagrion hastulatum are typical of the Tiefwald region. In the western part of the Swiss Alps, the Aletsch region, a UNESCO World Heritage Site, warrants an excursion. At Breite Bode, above Riederalp, with fantastic views over the longest glacier in the Alps, some small ponds supporting alpine species can be found; Somatochlora alpestris and Aeshna juncea can even be encountered in the botanical garden of the Riederfuka nature conservancy visitor centre. In contrast are two sites situated in the warmer Rhône valley floor in the Valais: the Pfinwald, a pine forest on a prehistoric landslide near Sierre, with a number of large ponds; and the oxbows of Leukerfeld, near Leuk. These sites each harbour at least 30 species, including Erythromma lindenii, E. viridulum, Aeshna isoceles and Leucorrhinia albifrons. Alpine species may also be found in the southern parts of the Alps. Several good locations are situated at higher altitudes of the Ticino valley, such as Piora, above Ambri Piotta. H Wildermuth

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also occur at many places in Tyrol, e.g. at Hartkaseralp near Ellmau, at Wildschönau/ Schatzberg near Auffach, at the Filzenscharte/ Trattenbach pass between Westendorf and Wald (Pinzgau, Salzburg), and in the Swiss Stone Pine (Pinus cembra) forest near Obergurgl/Ötztal. In the lower Inn Valley, the Thierberg region near Kufstein and Kramsach (with the Reintaler lakes) harbour 42 and 37 species, respectively, including Sympecma paedisca and Libellula fulva. The Lech valley between Stanzach and Weissenbach is one of the very last wild rivers of the Alps. It harbours an interesting flora and fauna, and is also rewarding for dragonfly-watching. Among many other species, Coenagrion hylas may be found along the edge of the river plain, in ponds and small lakes with cold, slow-flowing water. The whole region is proposed as a national park, and the LIFE-Infostelle in Weissenbach is worth a visit. In the east of Austria, one of the most interesting areas for dragonflies is the region of Neusiedler See-Seewinkel, which is the lowestlying part of the country. The Neusiedler See is the most westerly shallow steppe lake in Europe. Extensive reedbeds, up to 5km wide, fringe the lake. The adjacent Seewinkel is characterised by about 45 small, shallow saline lakes, with seasonally fluctuating water levels. Both are included in the Nationalpark Neusiedler See-Seewinkel, with an excellent visitor centre at Illmitz. The mosaic of large and diverse wetland habitats makes it one of the most important sites in the region for many species, particularly Lestes macrostigma, but also L. barbarus, L. virens, Chalcolestes parvidens, Ischnura pumilio, Aeshna affinis, A. isoceles, Sympetrum fonscolombii, S. meridionale and Leucorrhinia pectoralis. Altogether, 49 species have been found in the area. Another fantastic area to visit Marginal reedbeds of the Neusiedler See, Austria. is the floodplain of the Donau (Danube), east of Vienna, with currently 50 species. Downstream from Vienna, these are the only remaining large floodplains and riverine forests on a free-flowing stretch of the Danube in Austria. The whole area is protected in the Nationalpark Donauauen. The large bodies of standing fresh water in the Untere (Lower) Lobau, such as the Mittelwasser and Kühwörter Wasser, only a

Austria is dominated by high mountains in the west and south. The gently sloping northern and north-eastern parts form the Danube plain, which passes into the shallow basin of Vienna, with the large lake of Neusiedler See in the extreme east. With 78 species, Austria’s dragonfly fauna is diverse and encompasses Mediterranean as well as alpine elements, including specialities such as Lestes macrostigma, Coenagrion hylas, Stylurus flavipes, Cordulegaster heros and Somatochlora meridionalis, as well as Chalcolestes parvidens, the latest addition to the Austrian list. Aside from S. flavipes, none of these species is present in adjacent Switzerland. On the other hand, the typical alpine species are the same in both countries. Access to the rewarding alpine sites in Austria is rather similar to those in Switzerland. In contrast to the alpine habitats, where the best season is July and August, the lowland sites such as the Neusiedler See or the Danube floodplains are rewarding places to visit for dragonfly fauna from April to September. All dragonflies and damselflies are strictly protected in Austria and may not be collected. In Vorarlberg, the westernmost part of the country, alpine species such as Aeshna juncea, A. caerulea, Somatochlora alpestris and Leucorrhinia dubia may be encountered in the Montafon region, on a walk from Tafamuntalpe cableway station near Partenen to the Wiegensee (1,900m), and at the Kops reservoir on the Zeinisjoch. The same fauna is found at small ponds among alpine pastures near the Sonnenkopf cableway station, between Dalaas and Silbertal. Alpine species, including Coenagrion hastulatum and Aeshna subarctica,

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few kilometres from central Vienna, support Austrian Cordulegaster species occur together: important populations of Coenagrion pulchellum, bidentata inhabits the areas around the springs Brachytron pratense, Aeshna isoceles and Epitheca of small tributaries; heros is found in the central, bimaculata. Further west, and even closer to the mostly forested, reaches of the stream; and city centre in the Obere (Upper) Lobau, Anax boltonii occurs mainly along the lower reaches parthenope and Libellula fulva are interesting where meadows border the stream. The ditches species to look out for. The most important site and streams of south-eastern Styria are home for Stylurus flavipes is situated where the river to an interesting variety of dragonflies, including March meets the Danube, on the Austrian– Somatochlora meridionalis. This species prefers Slovakian border. The March and its floodplains half-shaded banks of flowing waterbodies, e.g. are home to significant populations of Epitheca the Mühlgang, south-west of Poklitsch. bimaculata and Orthetrum albistylum. Where the R Raab March annually inundates large parts of lowland Czech Republic and Slovakia areas, Aeshna affinis and Sympetrum flaveolum Lying in the heart of central Europe, these republics are likely to be found. A good place to start a are largely hilly, sloping to higher mountains on dragonfly tour of the floodplain is Marchegg, the western and northern borders, and to the from where you can walk north to Baumgarten. Pannonian lowlands in the south. Mediterranean The Wienerwald is a forest covering the northeastern foothills of the Alps and extends to within species creep up into the lowlands, which mark the northern edge of their range, while isolated the south-western city boundaries of Vienna. populations of boreal species are found in the Here, Cordulegaster heros occurs along the small mountains. Seventy-four species have been streams and may even be seen inside the city in reported from the Czech Republic, including the Lainzer Tiergarten, at the Rotwasserbach and recent records of Erythromma lindenii, Gomphus the Gütenbach. pulchellus, Cordulegaster heros and Somatochlora The Waldviertel, in northern Austria, lies at meridionalis. Sixty-nine species are currently known the southern end of the granite Bohemian Mass, from Slovakia, with Anax ephippiger being the which separates the river systems of the Elbe and latest addition. Danube, and is home to 58 species of dragonfly. The Šumava, Krušné hory, Jizerské hory, There are about 1,400 fish ponds here, with a Krkonoše, Králický Snežník and Hrubý Jeseník total surface area of around 1,700ha, many of which are not used intensively. Larger ponds often mountains on the Czech border provide various habitats to explore. Small lakes hold Aeshna have extensive stands of reeds and other aquatic juncea and Somatochlora alpestris, while small vegetation. Good places to watch the fascinating peatbogs harbour the rarer Somatochlora arctica variety of species found here are the ponds and those at higher altitudes support numerous around Pürbach south of Schrems, or those populations of Aeshna caerulea. Lakes with south and west of Heidenreichstein, such as floating Sphagnum are often home to Coenagrion Brünauteich. Apart from the fish ponds, the bogs and streams of the Waldviertel Lake in Šumava National Park, Czech Republic. hold important populations of dragonflies, e.g. Lestes dryas, L. sponsa, L. virens, Coenagrion hastulatum, Erythromma najas, Cordulegaster boltonii, Sympetrum flaveolum and Leucorrhinia dubia. The only Austrian sites where Leucorrhinia rubicunda is currently found are three bogs in the north-western Waldviertel. Interestingly, there is a small stream in the southern Waldviertel, the Felbringbach, where all three

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hastulatum, Aeshna subarctica, Cordulia aenea and Leucorrhinia dubia. In Slovakia’s Karpaty (Carpathian) mountains, forest springs and small brooks support Cordulegaster bidentata, while rivers with shingle banks are ideal for Onychogomphus forcipatus and Gomphus vulgatissimus. Hundreds of artificial ponds characterise the hills of Stredoceská pahorkatina and Ceskomoravská vrchovina, between Praha (Prague) and Brno. Coenagrion puella, Aeshna mixta, A. grandis, Libellula quadrimaculata and Lestes and Sympetrum species are common here. Where peat moss fringes small ponds, Coenagrion hastulatum, Leucorrhinia pectoralis, Sympetrum danae and, rarely, Coenagrion lunulatum and Leucorrhinia rubicunda appear. Calopteryx splendens, C. virgo, Cordulegaster boltonii and Somatochlora metallica grace the streams in these hills. At lower altitudes, Erythromma viridulum, Aeshna isoceles and Crocothemis erythraea can be found at numerous ponds and sand pits around Trebon, east of Ceské Budejovice, and Ophiogomphus cecilia and Cordulegaster boltonii occur along the pristine streams. Rarities such as Nehalennia speciosa, Somatochlora arctica, S. flavomaculata, Leucorrhinia dubia, L. pectoralis and L. rubicunda are found in peatbogs, some of which contain lakes created by peat extraction. Sympecma paedisca may still be found in the lowlands near Most, in the north-west of the Czech Republic, while Libellula fulva and Sympetrum depressiusculum can be found in old mines around Ostrava in the northeast. Natural stretches of the nearby Odra river support Stylurus flavipes, Gomphus vulgatissimus, Ophiogomphus cecilia and Onychogomphus forcipatus. The lowlands along the Dunaj (Danube) and its tributaries Dyje, Morava, Ipel and Tisa have permanent populations of southern species such as Coenagrion scitulum, Crocothemis erythraea, Orthetrum brunneum and Sympetrum meridionale, while Aeshna affinis is regular. Epitheca bimaculata inhabits gravel pits and where the rivers are still natural, populations of Stylurus flavipes, Ophiogomphus cecilia and sometimes Onychogomphus forcipatus are present. Clear streams in forests are the habitat of Cordulegaster heros, while muddy ones in open land harbour Coenagrion ornatum, Orthetrum coerulescens and Sympetrum pedemontanum. In 1995,

Somatochlora meridionalis was first discovered here, and in 2002 Leucorrhinia caudalis. O Holuša

Hungary Hungary is generally a very flat country, its vast Pannonian lowlands watered by many meandering streams and rivers, which provide a broad variety of habitats along them. The puszta landscape of the Great Plain, which dominates huge lowland areas of central Hungary, conceals many shallow alkaline steppe lakes, which often dry out in summer and harbour a characteristic dragonfly fauna. The discoveries of a population of Erythromma lindenii in the south-east (2014) and a single male of Trithemis annulata (2016) bring the number of known species to 66. The shallow alkaline lakes of the Great Plain are the most important stillwater habitats for dragonflies. Their temporary character does not suit all dragonflies, but are a paradise for Lestes macrostigma, Sympetrum depressiusculum, S. meridionale and a plethora of other Sympetrum and Lestes species, which are well adapted to such ecological conditions. Lakes with a more generous summer water level allow Sympetrum fonscolombii and Anax ephippiger to reproduce within a few months, and locally Coenagrion scitulum may also be encountered. Kelemen-szék in Kiskunság National Park (about 30km south of Budapest) is one of the best sites. Swampy areas in the Kiskunság area and around KisBalaton are characterised by strong populations of Coenagrion pulchellum, Aeshna isoceles, Somatochlora flavomaculata and Leucorrhinia pectoralis. One of the most robust populations of the latter in central Europe is at Lake Kolon, in Kiskunság National Park. These and other waterbodies in the park can best be visited with local guides. Similar habitats can be found at the crossborder national park of Neusiedler See and Ferto-Hanság, shared with Austria. It consists of a shallow steppe lake, with a huge reed belt, adjacent small soda lakes and traditional pastures. It is very popular among birdwatchers and other naturalists, including those seeking dragonflies. In June, Lestes macrostigma and Leucorrhinia pectoralis can be seen. The extensive canal network of the peatland of NorthHanság is worth a visit to see good populations of Coenagrion ornatum and Libellula fulva, accompanied by Orthetrum coerulescens and

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Szentgyörgyvölgy and Mecsek, in northern and western Hungary, are the best places to look for Cordulegaster heros. A Ambrús and M Bedjaniˇc

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Ditch in Hansági National Park, Hungary. O. brunneum. In the summer, Aeshna grandis and Sympetrum pedemontanum are also frequently seen here. Some Hungarian rivers, such as the upper Tisza in the north-east (e.g. near Tiszabecs, Tarpa, Vásárosnamény and Tokaj) and Rába in the west (Szentgotthárd, Magyarlak, Körmend, Rábahídvég), are among the best localities to find all four central European gomphids together: Stylurus flavipes, Onychogomphus forcipatus, Ophiogomphus cecilia and, most widespread, Gomphus vulgatissimus. The Tisza is famous for the ‘river flowering’ in the middle of June, caused by the mass emergence of the mayfly Palingenia longicauda. Oxbows, flooded areas and gravel pits along the river are important habitats for the extremely local Leucorrhinia caudalis and L. pectoralis, and more abundant Epitheca bimaculata. Sites near Tiszadob, Tiszafüred and Tiszacsege-Egyek can be visited with guides from Hortobágy National Park. Stands of Water-soldier (Stratiotes aloides) in north-east Hungary may harbour Aeshna viridis, with accessible sites along the Csaronda at Tiszaszalka and Lónya on the Ukrainian border. Streams in the hills of Sopron, Orség,

Forming the south-west corner of the Iberian peninsula, Portugal boasts a wide variety of landscapes. The altitude drops from the Spanish border to the Atlantic coast, and from north to south. The Tejol (Tagus) river divides the mountainous northern region from an area of plateaux and plains in the south. The dragonfly fauna of continental Portugal, the westernmost country of Europe, comprises 65 species. In the north-west, Parque Nacional da Peneda-Gerês contains a network of streams where Boyeria irene, Cordulegaster boltonii, Onychogomphus uncatus, Calopteryx virgo and Pyrrhosoma nymphula are ubiquitous. In addition, Macromia splendens is known from Lindoso reservoir, at the Spanish border. In the north-east, one of the most interesting places to visit is the valley of Rio Sabor and its tributaries, where more than 20 species occur, including the protected Coenagrion mercuriale, Gomphus graslinii, Macromia splendens and Oxygastra curtisii. Also worth visiting is Salreu, a large marshy area near Aveiro, very close to the Atlantic coast, where dragonflies may be observed almost throughout the year, sometimes in huge numbers. The most interesting species recorded from the area is Brachytron pratense, already on the wing by the beginning of March, but many others are present, including Aeshna mixta, Crocothemis erythraea and Chalcolestes viridis. A little to the south, and closer to the sea, is Lagoa da Vela, a small lake where the scarce Diplacodes lefebvrii and Brachythemis impartita are most notable. Aeshna juncea is known from only one area in the country, Serra da Estrela, the highest mountain and the largest natural park in mainland Portugal. The various lakes on its high plateau also harbour Lestes dryas, Enallagma cyathigerum, Anax imperator, Libellula quadrimaculata and Sympetrum flaveolum, while the clean streams and rivers are home to Calopteryx virgo, Platycnemis latipes, P. acutipennis, Boyeria irene, Cordulegaster boltonii, Onychogomphus forcipatus and O. uncatus. The Rio Mira valley, near Santa Clara-a-Velha, is a good site to look for Lestes virens and Ceriagrion tenellum, and a diligent searcher may find Coenagrion scitulum.

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The natural valleys of numerous small, clean rivers in the Rio Guadiana catchment have a most attractive fauna, including Calopteryx xanthostoma, Ischnura pumilio, Gomphus graslinii, G. simillimus, Onychogomphus costae, O. forcipatus, Paragomphus genei, Macromia splendens, Oxygastra curtisii, Orthetrum chrysostigma, O. nitidinerve, O. trinacria and Trithemis annulata. A beautiful example is Ribeira do Vascão, on the border of Beja and Faro districts. In the Algarve, the marshes of Alvor harbour, among other species, Diplacodes lefebvrii, Anax parthenope and Libellula quadrimaculata. Along the southern coast, mass migrations of Anax ephippiger and Sympetrum fonscolombii are sometimes witnessed, especially in autumn. Selysiothemis nigra was recently discovered near Faro, the westernmost point of its distribution range. Trithemis kirbyi bred at Ribeira de Asseca in 2017 is expected elsewhere in the south-east. F Weihrauch and S Ferreira

Spain There are 80 species in Spain (excluding the Canaries), with Trithemis kirbyi added last. After its arrival in Andalucía in 2007, the species rapidly colonised much of the east and interior, and recently reached the foothills of the Pyrenees. The climate of northern Spain, with humid summers, contrasts with the desert-like climate of the south, and this is reflected in the composition of the fauna. Species such as Macromia splendens, Oxygastra curtisii, Lestes dryas, L. sponsa, Calopteryx virgo, Aeshna juncea, Gomphus vulgatissimus and Cordulegaster bidentata are Lagoon at El Rocío, Coto Doñana, Spain.

found only in the north, or become progressively rarer to the south, while Lestes macrostigma, Anax parthenope, A. ephippiger, Paragomphus genei, Orthetrum trinacria, O. chrysostigma, Brachythemis impartita, Diplacodes lefebvrii, Trithemis annulata, T. kirbyi, Selysiothemis nigra and Zygonyx torridus are typical southern species, many of which are also common in North Africa. Rich dragonfly communities are found in rivers in Galicia (e.g. Cabe, Sil, Lérez, Tea, Tambre, Arnoia, Limia) in the north-west, where Oxygastra curtisii is very common and is usually accompanied by Macromia splendens and sometimes by Gomphus graslinii. Lagoons along the coast of Galicia, such as Doniños, Valdoviño, Louro, Xuño and Vixán, have large populations of Ischnura elegans (a rare species in most of Spain), Lestes virens, L. barbarus, Aeshna mixta and Sympetrum fonscolombii. At some of these there are mixed communities of I. elegans and I. graellsii, and hybridisation between the two is common. In the mountainous region of Asturias the rivers are short and swift. The commonest species here are Calopteryx virgo, Boyeria irene and Cordulegaster boltonii. Calopteryx haemorrhoidalis is common and the males have a metallic violet body: they are sometimes treated as the distinct subspecies asturica. The ibones (mountain lakes) of Aragón harbour large populations of Aeshna juncea, with A. cyanea, Libellula depressa, L. quadrimaculata, Sympetrum flaveolum, Lestes dryas, L. sponsa, Pyrrhosoma nymphula and Enallagma cyathigerum as accompanying species. These northern species are generally scarce and confined to high altitudes in the Iberian peninsula. Some rare species in Spain, such as Sympetrum vulgatum (the pale subspecies ibericum), S. flaveolum, Lestes sponsa, Aeshna juncea and Gomphus vulgatissimus, are found in the mountains of the León region, between Asturias and the plains of central Spain. Other northern species are found only in the Pyrenees: Coenagrion hastulatum, Somatochlora metallica, Leucorrhinia dubia and L. pectoralis. An area with a rich fauna is Madrid, probably owing to its geographical situation in the

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Tyrrhenian Islands in particular harbour a fauna with an African flavour, and the south is even home to an endemic species, Cordulegaster trinacriae. Several species were added recently, with Aeshna subarctica in the Alps (2009) and Sympetrum sinaiticum (2010), Pantala flavescens (2012), Diplacodes lefebvrii (2013) and Tramea basilaris (2016) on small Mediterranean islands; the latter even represented the first record north of the Sahara and in Europe. The southern Alps are a Mountain lake, Lago de Erina, in the Covadonga National Park, good place to start looking for Picos de Europa, Spain. dragonflies in the north of the middle of the peninsula, where southern and country. Just below the mountain hut at Baita northern species can be found together. The Segantini, at Passo Rolle in Trentino, is a bog Mediterranean coast has many African species, with Aeshna caerulea and Somatochlora alpestris. such as Orthetrum nitidinerve, O. chrysostigma, In the upper Valtellina, at Trivigno in Lombardy, Brachythemis impartita, Trithemis annulata, T. the peatbog Torbiera di Pian Gembro supports kirbyi, Selysiothemis nigra and Zygonyx torridus; plenty of Coenagrion hastulatum, Somatochlora S. nigra is also regularly recorded on the Balearic arctica and Leucorrhinia dubia. The small Lago Islands. The Valencia area and the river Ebro, di Lod near the village Antey-Saint-André in especially the delta, are two of the richer parts of Valtournenche (Valle d’Aosta) has dozens of this region. Some Andalucían rivers, for instance Aeshna grandis. the Yeguas and Hozgarganta, have good Moving south, the large wetland Torbiere populations of western European endemics, such d’Iseo, between Iseo and Provaglio, has more as Platycnemis acutipennis, P. latipes, Gomphus than half the dragonfly species known in Italy. graslinii, G. pulchellus and Macromia splendens. The rare Oxygastra curtisii occurs here, along Onychogomphus costae is locally abundant in with Brachytron pratense, the only viable Italian the catchments of the Guadalquivir, Júcar and population of Leucorrhinia pectoralis and, until Ebro. Sympetrum sinaiticum breeds in all types a few decades ago, Nehalennia speciosa. Here, of water, and is fairly common along the entire and in the other basins along the edge of the coast from Andalucía to Cataluña (Catalonia). Lombardy Pre-Alps, Erythromma najas and An area of particular interest is the Doñana Cordulia aenea are found alongside Coenagrion National Park, with rarer species such as pulchellum, Erythromma viridulum, E. lindenii, Lestes macrostigma, Brachytron pratense, Anax Ceriagrion tenellum, Aeshna isoceles and ephippiger, Paragomphus genei and Diplacodes Somatochlora flavomaculata. lefebvrii, although B. pratense has not been found The western part of Pianura Padana (Po there recently. valley) is an agricultural landscape with large A Cordero Rivera channels and streams where, among the numerous Orthetrum albistylum and Sympetrum Italy pedemontanum, Ophiogomphus cecilia may Stretching from the high Alps to the subtropical be seen along the edges of the cornfields, or climes of the Mediterranean, Italy has an even Stylurus flavipes patrolling over the water. extraordinary richness of landscapes, with a Huge swarms of Sympetrum depressiusculum consequently diverse fauna of 95 dragonfly occurred in the ricefields until the 1970s, but the species. Although many western and eastern population has now declined dramatically. Any Mediterranean specialities are absent, the small stream may have Calopteryx splendens,

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Pyrrhosoma nymphula, Gomphus vulgatissimus, Libellula fulva and, occasionally, Calopteryx virgo, Boyeria irene and Cordulegaster boltonii. Try the area of Tromello (Lombardy, Pavia province), close to the channels Naviglio Langosco and Canale Cavour and the stream Terdoppio. Between Lombardy and Emilia, the relatively undisturbed sections of the river Po are good for S. flavipes, some of them supporting large populations. Several streams in Liguria, e.g. the Erro near The Ampola valley in the southern Alps, near Lake Garda, Italy. Pontinvrea, hold Boyeria irene, Onychogomphus uncatus and Oxygastra curtisii. Around the Lago di usually found on streams and rivers, while the Massaciuccoli in Toscana (Tuscany), you may be former mostly inhabits still waters, for instance in lucky enough to encounter the elusive Lindenia the north-west at the Lago Baratz and the small tetraphylla. The mountain streams of southern lake Stagno di Platamona. Clear streams with a Italy, such as the upper course of the rivers sandy bottom and shore, such as Rio Terramala Bussento (Campania) and Agri (Basilicata), near Monti or the streams near Posada, may may reveal the Italian endemic Cordulegaster yield large numbers of Paragomphus genei. You trinacriae. have to be really lucky to see Lindenia tetraphylla, The Tyrrhenian Islands are the best place to see but try at the Stagno di Platamona or around the African species, preferably during the summer, mouth of Rio Padrogianu near Olbia. Around the and the only place for Ischnura genei, which is small saline ponds at Su Palosu near Oristano common here. Sicily offers many opportunities. in late May and early June, the clumps of rushes Visit the coastal lake Biviere di Gela, near Gela on are covered with hundreds of Lestes macrostigma, the southern coast, to see numbers of Orthetrum looking like pale blue flowers. This is almost trinacria, Trithemis annulata and Selysiothemis the only dragonfly able to breed here. Similarly, nigra, with possibly also Paragomphus genei. small temporary ponds that are reduced to a dry Even before you reach its shores, dozens of reedbed in summer, such as those scattered on Brachythemis impartita will fly all around your the Monte Limbara near Tempio Pausania, are feet as you walk on the bare ground. In the unsuitable for most species, apart from Lestes mountains near Palermo, head for the southern barbarus and L. virens, which may occur in their slopes of the rocky-topped Rocca Busambra, hundreds. at Ficuzza. The trickles here may harbour the M Pavesi brightly yellow-marked subspecies Cordulegaster bidentata sicilica, together with the rare Orthetrum Slovenia nitidinerve. Cordulegaster trinacriae occurs around Slovenia is small green country south of the Alps, characterised by beautiful landscapes and a rich Castelbuono and Mezzojuso. fauna and flora. Its position at the confluence of Should you have a sun-and-sea holiday on the small island of Pantelleria, then visit the volcanic the Mediterranean, Pannonian, Alpine and Dinaric biogeographic regions, and its wealth of different lake Bagno dell’Acqua or Specchio di Venere, freshwater habitats, are the main reasons for its which have the only European populations of diverse dragonfly fauna of 72 species. Slovenia Ischnura fountaineae known to date – practically is the best place in Europe to see the enigmatic the only dragonfly to be found there. Cordulegaster heros and Somatochlora meridionalis. The African Orthetrum trinacria and Trithemis Just a few kilometres south of the capital, annulata also occur on Sardinia. The latter is

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Ljubljana, is one of the most interesting localities in the country. Once the most famous marshland in southern Europe, the Ljubljansko barje moor today is characterised by an extensive but beautiful agricultural landscape, criss-crossed by a dense and ecologically diverse network of drainage channels, ditches and streams. Among the 40-plus species recorded here and in the near surroundings, the largest Slovene population of Somatochlora flavomaculata and Coenagrion ornatum are of special interest. Early summer is the best time to find the largest European dragonfly, Cordulegaster heros. It favours small, slow- to moderate-flowing forest streams, typical of the hilly country leading down to the flatlands. North of Ljubljana, try the forest streams west of the Kranj–Škofja road and north of the Cerklje–Preddvor road. C. heros is also very common in central and eastern Slovenia, and for dragonfly-watchers with a little experience it is almost impossible not to encounter this species, which is often accompanied by Calopteryx virgo, and sometimes also by Cordulegaster bidentata and Somatochlora meridionalis. The latter is also very common, but it prefers slightly larger, shaded, meandering streams in the lowlands. It is most frequently accompanied by Calopteryx virgo, C. splendens, Platycnemis pennipes and Onychogomphus forcipatus. Lying on the sunny side of the Alps, northern Slovenia has some of the southernmost peatbogs in Europe: the Pohorje mountains and the Pokljuka and Jelovica plateaux in the Julian Alps are good for Aeshna juncea, Somatochlora arctica and Leucorrhinia dubia. Within an hour’s drive is one of the greatest karst curiosities, the intermittent Lake Cerknica, with its constantly changing appearance and 36 dragonfly species. Despite the summer crowds, it is worth visiting the Slovene coast in June: the old saltpans in Secovlje, the brackish lagoon Škocjanski zatok, near Koper, and two claypit lakes in Fiesa, near Piran, harbour more than 40 species, including a strong population of Ceriagrion tenellum in Fiesa. In the north-east of Slovenia, extensive fish ponds around the villages of Podvinci and Zgornji Velovlek near Ptuj, Komarnik reservoir near Lenart, and Medvedce reservoir south of Pragersko, each harbour more than 35 species. May is the best time to see Epitheca bimaculata, which has been recorded at more than 70 localities in this part of the country. In June, more

than 30 species can be easily encountered during a weekend, including Leucorrhinia pectoralis, Aeshna isoceles and Erythromma najas. Further to the east, the Mura floodplain, with its oxbows and numerous gravel pits, represents another dragonfly hotspot, with more than 50 species. Some of the older oxbows along the border section of the Mura, near Petišovci, are inhabited by both Chalcolestes viridis and C. parvidens, and some even support the endangered Aeshna viridis, Leucorrhinia caudalis and L. pectoralis, while Aeshna isoceles and Coenagrion pullchelum are common. M Bedjaniˇc

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Croatia With the exception of peatbogs and mires, Croatia has it all as far as dragonfly habitats are concerned: beautiful lowland rivers with preserved floodplains and oxbows, extensive fish ponds, scenic limestone rivers and, of course, the picturesque Adriatic coast and its islands, with their innumerable small pools and cattle ponds. However, care is needed in certain parts of East Slavonia, Banovina and Dalmatia as a result of minefields placed during the war. At present, the dragonfly fauna of Croatia totals 68 species. In northern Croatia, on the border with Hungary, lower sections of the Drava river are very interesting, offering the possibility of seeing Ophiogomphus cecilia, Stylurus flavipes and Gomphus vulgatissimus together. In the far east is the Kopacki rit nature park, one of the largest natural marshlands and floodplains in Europe, where the Drava flows into the Dunav (Danube). More than 40 dragonfly species have been recorded here. In central Croatia, similar habitats with a rich dragonfly fauna can be also found in Lonjsko polje national park. The outstanding cultural landscape around the scenic village of Cigoc is famous for its breeding storks and free-ranging hairy pigs, which often unintentionally create small dragonfly habitats in flooded pastures. These small temporary ponds provide suitable habitat for Lestes barbarus, L. dryas, Sympecma fusca, Ischnura pumilio, Aeshna affinis and Sympetrum meridionale. Heading out from Zagreb, through the Kapela mountains towards the Adriatic coast, the famous Plitvice lakes are certainly worth visiting. Although somewhat crowded on weekends and holidays, Plitvice is one of the most stunning examples

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of karst scenery, and is protected as a national park and UNESCO World Heritage Site. The series of numerous scenic lakes on the Korana river, separated by travertine barriers and lovely waterfalls, is further enhanced by its rich dragonfly life. More than 30 species have been recorded here. The typical dragonfly assemblage of limestone rivers is characterised by numerous Calopteryx splendens and C. virgo, Platycnemis pennipes, Gomphus vulgatissimus and Orthetrum coerulescens. The Adriatic coast is magnificent in its beauty and diversity, and offers great possibilities to combine family holidays with dragonfly-watching. Some of the bigger Kvarner islands, such as Pag and Krk, host more than 35 dragonfly species and represent the northern extent of the rare Selysiothemis nigra and Lindenia tetraphylla. In late June and July, lake Velo Blato in southern Pag is among the best places in Europe to observe the two. Pantala flavescens was recorded for the first time in Croatia on Krk in 2010. The scarcity of bigger streams and lakes on the Croatian islands is compensated for by hundreds, or even thousands, of small cattle ponds, called lokve. Their dragonfly communities are characterised by Anax imperator, Libellula depressa, Crocothemis erythaea, Sympetrum striolatum and S. meridionale, but may include Chalcolestes parvidens, Coenagrion scitulum, Ceriagrion tenellum and also, as in the case of the national park of Brijuni, Lestes macrostigma. The southern coast of Dalmacija (Dalmatia) is among the richest areas in the Mediterranean for dragonflies. In the Neretva delta, at least 36 species have been recorded, including Sympecma fusca, Ceriagrion tenellum, Coenagrion ornatum, Erythromma lindenii, Lindenia tetraphylla and Selysiothemis nigra. The streams in the Konavli area are frequently inhabited by Caliaeschna microstigma, Cordulegaster bidentata and the subspecies Calopteryx splendens balcanica. T Bogdanovi´c and M Bedjaniˇc

Other states of former Yugoslavia Serbia, Montenegro, Bosnia and Herzegovina, Kosovo and North Macedonia occupy the largest portion of the north-western Balkans. In the northeast, dragonfly habitats are largely influenced by the Dunav (Danube) and its tributaries. The central part of the region is mountainous with karst scenery, which is relatively unfavourable to dragonflies. Further south, towards

Montenegro and North Macedonia, the increasing Mediterranean influence is reflected in the climate, dragonfly habitats and species encountered. The dragonfly fauna of the whole region is poorly known. Sixty-three dragonfly species have been recorded from Serbia, 64 from Bosnia and Herzegovina, 67 from Montenegro, 62 from North Macedonia and 42 from Kosovo. Minefields were laid across what is now Bosnia and Herzegovina during the Bosnian War, and as a result care is needed with fieldwork here. The lowland Pannonian part of Serbia has many suitable rivers, ponds and marshes, supporting a fauna similar to that of the adjacent regions of Croatia and Hungary. From a zoogeographical point of view, the widely disjunct relict populations of Coenagrion hastulatum, Leucorrhinia dubia and Aeshna juncea occurring in the mountain lakes of the Durmitor National Park, in Montenegro, and on Mt Golija, in southern Serbia, are interesting. Streams in the hilly and mountainous areas of Serbia and North Macedonia are frequently inhabited by Cordulegaster heros and C. insignis. The two regions of Bosnia and Herzegovina are very diverse in terms of dragonfly habitats. Plains along the Sava and Bosna rivers are suitable for many common species such as Calopteryx splendens, C. virgo and Libellula depressa, but populations of Leucorrhinia pectoralis and L. caudalis are also found locally. Streams in the high mountains, especially in central Bosnia, host species such as Cordulegaster bidentata and Gomphus vulgatissimus. The unique mix of Mediterranean and continental influences on the dragonfly fauna of the region is seen in Calopteryx splendens populations, with characters of the subspecies balcanica and ancilla expressed to a varying degree. Although generally dry and rocky, Herzegovina is rich in small rivers. Interesting combinations of species, such as Lindenia tetraphylla, Cordulia aenea, Somatochlora meridionalis and Coenagrion ornatum can be found here, for instance in Hutovo Blato National Park, near Capljina. While the mountainous karst interior of Montenegro is not very suitable for dragonflies, the lowland surrounding lake Skadar, together with the coastal Adriatic region around Ulcinj, may offer the chance of seeing interesting southern species such as Gomphus schneiderii, Trithemis annulata, Selysiothemis nigra and the subspecies Platycnemis pennipes nitidula. The lake harbours one of the

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largest populations of Lindenia tetraphylla known, as well as a very isolated (and only recently discovered) population of the otherwise western Mediterranean Gomphus pulchellus. In North Macedonia, the dragonflies of lakes Doiran and Prespa and their surroundings are well known. The latter lies in the hills on the border with Greece and Albania. More than 30 species have been recorded, the most interesting being Leucorrhinia pectoralis, Cordulia aenea and Sympetrum River Veleka in southern Bulgaria, habitat of Somatochlora borisi. vulgatum, which become very rare towards the south of Europe. The lowland lake Doiran lies on the southernmost occurrence. During glaciation, border with Greece and hosts more than 35 these and other northern species penetrated dragonfly species, including a large population southwards along the Carpathian and Balkan of Lindenia tetraphylla, while Epallage fatime is mountains. Peatbogs in the higher mountains found in the hills north of the lake. Caliaeschna are now home to isolated communities of microstigma can be easily observed on warm Lestes sponsa, Pyrrhosoma nymphula, Aeshna summer afternoons along some streams on the juncea, Cordulia aenea, Libellula quadrimaculata slopes of Mt Belasica. and Sympetrum flaveolum. Erythromma najas, M Bedjaniˇc and T Bogdanovi´c Coenagrion hastulatum, C. lunulatum, Aeshna grandis, Epitheca bimaculata, Somatochlora Bulgaria and Romania metallica, S. alpestris and Leucorrhinia pectoralis The region has one of the most diverse landscapes are very local, but may be found more widely in Europe. Numerous tributaries in vast lowlands once Romanian Moldavia and Transylvania are drain to the majestic Danube (Dunare in Romanian, explored. In the Carpathians of Romania’s far Dunav in Bulgarian), with countless lakes along its north, Coenagrion armatum and Nehalennia course leading to Europe’s largest delta. Coastal speciosa may still occur. lagoons are bordered by flooded forests with an In complete contrast, the lowlands and broad almost tropical appearance. Undulating plateaux valleys of the rivers flowing to the Aegean and Black with karst lakes lead up to large mountain chains seas are favourite haunts of Mediterranean with some of Europe’s southernmost peatbogs and species. Burgas is a good place to start exploring montane lakes. In the Balkans, continental European the southern Black Sea coast. Check the and Mediterranean species mix. Recent records of numerous small rivers for Epallage fatime, Pantala flavescens (2012) and Epitheca bimaculata Caliaeschna microstigma, Cordulegaster picta and (2017) bring the Bulgarian list to 71 species. The Somatochlora meridionalis. Lestes macrostigma dragonflies of Romania are still rather poorly can be seen around vegetated pools in the dunes understood; currently, 71 species are also known south of the town. Small tributaries of the Danube in the country, with Aeshna subarctica (2009) and are good for Cordulegaster insignis, e.g. near Selysiothemis nigra (2013) added most recently. Vetren. Somatochlora borisi has been found near The Balkans are an attractive destination, with the Ropotamo and Veleka rivers, but it was first a rich but still insufficiently known biodiversity discovered as new to science (in 1999) in the valley that offers much potential for new discoveries. of the Byala reka in the heart of the eastern Somatochlora arctica and Leucorrhinia dubia were Rhodops, close to the villages of Byalgradets only recently discovered in the Rila mountains and Gugutka. At least S. meridionalis and L. of south-west Bulgaria, representing their macrostigma also reach into central Romania.

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The characteristic fauna of small rivers in the plains includes Calopteryx splendens, Coenagrion ornatum, Orthetrum brunneum and O. coerulescens. Libellula fulva often occurs with them, as may Erythromma lindenii. The best place to look for the nomadic Selysiothemis nigra is in brackish habitats near the Black Sea in the far south, where Lindenia tetraphylla was found reproducing in 2017 and Trithemis annulata may also await discovery. The Danube forms a dynamic artery through the region, supporting a vast range of habitats. Stylurus flavipes inhabits its entire stretch. The attractive delta covers much of Romania’s northern Dobrogea and is rich in lakes, channels and flooded areas. Known as ‘the living delta’, its dense vegetation forms small floating islands that move with the wind and tides. Aeshna isoceles and Anax imperator are typical over open areas with reeds, while Erythromma viridulum prefers to perch on floating leaves. More than 30 species are reported from the delta, but several recent records, such as of Selysiothemis nigra, suggest that the number present is more likely to be around 40. Similar diversity is found in Srebarna Nature Reserve, a wetland along the Danube in the Bulgarian Dobrudja. M Marinov

Greece and Albania

Almost four-fifths of the Greek mainland consists of mountains, where some northern species reach their southern limit. The coastal areas have a Mediterranean climate and fauna. The 2,000plus Greek islands make up almost one-fifth of Gorge in the Pindos Mountains, north Greece.

the country. Only 150 of them are inhabited and many are poor in dragonflies, although some of the larger islands are more favourable. Currently, 79 species are known from Greece. The dragonfly fauna of Albania is similar to that of northern Greece, but requires further study. The discoveries of Gomphus pulchellus (2014) and Somatochlora metallica (2015) brought the list up to 59 species. In the number of (near) endemic dragonfly species, Greece is unrivalled by any other European country. These species are scattered across the isles and mainland, making it difficult to see them all in a single visit. Although many southern species are found in standing waters, running waters harbour most of the (near) endemics. The recently described Somatochlora borisi is known from a small number of localities in the eastern Rhodopes and the Istranca Mountains, and also occurs in adjacent Bulgaria and Turkey. Try the river Diavolorema near Dadia, 10km southwest of Souflion near the Turkish border. Another near-endemic of the Greek mainland is Pyrrhosoma elisabethae, which is known only from a number of brooks in the northern Pelopónnisos (Peloponnese) (try the surroundings of Kalávrita and Dáfni), one location in southern Albania and from the isle of Kérkira (Corfu). On Kérkira (Corfu), Ceriagrion georgifreyi, another red damselfly with a small distribution, is also found. In addition, this species occurs on the isles of Thásos, Zákinthos and Lesbos, and locally in the Pelopónnisos. Strangely enough, only the more widely distributed Ceriagrion tenellum is present on Crete and the nearby island of Ios, although this species is absent from the rest of Greece. Crete is home to two endemic species: Coenagrion intermedium is a rather unimpressive C. puella lookalike and is fairly common around streams across the island; Boyeria cretensis is common only locally, being known from about 15 streams, including the Kabanos south-east of Koufi and the Petres north-west of Kaloniktis. It is endangered owing to its small distribution and the ongoing deterioration of its stream habitat. With five Cordulegaster species, Greece is heaven

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for anyone who enjoys the delicate differences between the many members of this genus. On the mainland, C. bidentata, C. heros, C. picta and C. helladica occur. The latter is endemic to Greece and is represented by three subspecies, of which buchholzi is present only on the Cyclades. Impressive numbers of the subspecies helladica can be found, together with C. heros, at the large well 8km west of Kalávrita. The springs at the ruins of Delphi are a classic site for the subspecies kastalia. The brooks just south-west of Neos Pandeleimónas, 30km south of Katerini, are good for C. bidentata and C. heros. The fifth species, C. insignis, can be found on the Greek islands of Thassos, Lesbos and Samos, where it occurs together with C. picta. The south Asian Anax immaculifrons, which recalls an oversized Cordulegaster, has its westernmost populations on the islands of Rhodes, Kárpathos and Ikaria. Another enigmatic Greek species, Lindenia tetraphylla, is found at large lakes such as Volvi and Iliki. The island of Lesbos is a popular birdwatching destination, and with well over 40 Odonata is equally good for dragonfly-watching. Among the species found here are goodies such as Epallage fatime, Lestes macrostigma, Ceriagrion georgifreyi, Caliaeschna microstigma, Gomphus schneiderii, Cordulegaster picta, C. insignis and Selysiothemis nigra. V J Kalkman

Turkey and Cyprus

microstigma and Trithemis festiva grace most streams. The best place to start is around Lake Köycegiz, a region with more than 50 recorded species. The lake itself can be explored by boat, and has remarkably high densities of Erythromma lindenii, Aeshna isoceles, Lindenia tetraphylla and Libellula fulva. The nearby Esen river harbours Onychogomphus flexuosus. Streams in Oriental Sweetgum (Liquidambar orientalis) forest are sites for Gomphus schneiderii, Cordulegaster insignis and C. picta. Nearby rocky mountain streams hold the impressive Anax immaculifrons. Runnels near the village of Köycegiz are inhabited by Ceriagrion georgifreyi and an unusual dark variety of C. pulchellum. The delta of the Göksu, especially the dune slacks and reedy ditches around Akgöl lagoon, is a good place to look for Lestes macrostigma, Brachythemis fuscopalliata, Diplacodes lefebvrii, Pantala flavescens, Selysiothemis nigra and Trithemis arteriosa. Crocothemis servilia flies with C. erythraea here. Most of these species may also be found around Adana. Canals and pools in the delta here (e.g. near Kadirli) also harbour Coenagrion syriacum, Platycnemis dealbata, P. kervillei, Gomphus davidi, Paragomphus lineatus and Libellula pontica. Four Onychogomphus species (O. forcipatus, O. flexuosus, O. lefebvrii and O. macrodon) live in the delta’s Ceyhan river, as does Stylurus ubadschii, a species that occurs throughout Turkey but is seldom observed. Between the Göksu and Adana lies the stunning limestone gorge of Limonlu, where Gomphus schneiderii, Onychogomphus assimilis and Anax immaculifrons occur. The most westerly

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Turkey is a large, rugged country with varied habitats ranging from snowy peaks and temperate forests, through cold steppes and hot deserts to subtropical coasts. The dragonfly fauna is accordingly diverse, and Carputz Cay in Turkey, habitat of Brachythemis fuscopalliata. with at least 101 species is the richest of the countries covered in this guide. Many tropical African and Asian species reach the extreme limits of their range on the Mediterranean shores of Turkey and Cyprus. With its pleasant climate, beautiful scenery and interesting dragonflies, southern Turkey is a popular destination. Orthetrum chrysostigma, O. sabina, O. taeniolatum and Trithemis annulata are common and widespread. Epallage fatime, Caliaeschna

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populations of the Central Asian Sympetrum haritonovi lie high in the mountains above Alanya. The west of Turkey, with its many archaeological sites, is less often visited for dragonflies and much remains to be discovered. A large population of Libellula pontica occurs at a brook 5km south of Taraklı (40km south of Sakarya), an extension of its known range by more than 500km. In north-west Turkey, lakes Kus (Manyas) and Ulubat (Apolyont), and the delta of the Koca, all west of Bursa, look promising. The numerous lagoons and rivers on the west coast should be searched for Lestes macrostigma and the poorly known Stylurus ubadschii. Thousands of Erythromma lindenii skim above the large lakes near Isparta, and the surrounding hills hold good habitat for Leucorrhinia pectoralis and Sympetrum vulgatum decoloratum. The ecology of the latter is poorly known and its exuviae remain undiscovered: freshly emerged individuals were found at a springfed lake just north of Çayıryazi, 35km south of Çay. Turkey’s north has a very different landscape and fauna, reminiscent of the mountains of central Europe. In Turkey, species such as Pyrrhosoma nymphula, Aeshna cyanea, Cordulia aenea and Sympetrum danae occur only here. The rainy far east of the Black Sea coast (not covered in this guide) is home to deviant relatives of Calopteryx splendens, Coenagrion puella and Cordulegaster insignis, each often considered distinct species. The dragonfly fauna of Cyprus is similar to that of the opposite mainland, but with 37 species it is relatively impoverished; glaring absentees are the genera Coenagrion, Gomphus, Cordulegaster and Libellula. Nonetheless, Anatolia’s charismatic stream species – Epallage fatime, Caliaeschna microstigma and Anax immaculifrons – are present throughout the island, although the latter is uncommon. Streams in the Diarizos river basin are best to find the island’s speciality, Ischnura intermedia, which was added to the European list in 2013. Lestes macrostigma is found at the brackish lagoons at Larnaca and Akrotiri. V J Kalkman and K-D B Dijkstra

Canaries, Madeira and the Azores These volcanic Atlantic archipelagos are noted for their extremely high and rugged mountains, whose slopes descend steeply to the coast; the record-breaking Pico del Teide on Tenerife reaches 3,715m. Trade winds from the north-east unload

their humidity on the steep mountain slopes on most islands of the Canaries and determine the islands’ climate, making the northern parts fertile while the southern parts are arid and hot. Natural freshwater habitats are scarce, except in the Azores, and the dragonfly fauna is accordingly poor: there are just ten regularly recorded species in the Canaries, seven in Madeira and five in the Azores. The dragonfly fauna is strongly influenced by the proximity to Africa, as demonstrated by the only two damselfly species in the Canaries, Ischnura saharensis and I. senegalensis. An American species of Ischnura makes a visit to the Azores worthwhile: I. hastata occurs on all major islands, but only females are found. This is the only parthenogenetic species known among Odonata. Ischnura pumilio has also colonised the Azores, as well as Madeira. The Canarian landscape is shaped by barrancos, steep gorges descending from the mountains, which are dry for most of the year or offer only ephemeral freshwater habitats. The lack of running water in the barrancos is compensated for by open irrigation channels, which may serve as a larval habitat for the African species Orthetrum chrysostigma, Trithemis arteriosa and Zygonyx torridus. Good sites are Barranco del Infierno and Barranco de Masca on Tenerife. Mass influxes from Africa of Anax ephippiger and the ubiquitous Sympetrum fonscolombii can be recorded from time to time, while Pantala flavescens probably followed in their wake: two individuals on Gran Canaria in 2013 represent the first record of this species for the Canaries. The first European record of Orthetrum ransonnetii was at Barranco de Rio Cabras on Fuerteventura in 2018, while Ischnura senegalensis has become established at a few sites on La Palma and southern Tenerife. Sympetrum nigrifemur is known only from Madeira, La Gomera, Gran Canaria and Tenerife, although some regard this species merely as a large, dark form of Sympetrum striolatum. It is fairly common and on the wing almost throughout the year. F Weihrauch

Morocco For a European traveller, Morocco is a perfect link with Africa. Separated from Spain by only a 15km strait, the landscape and habitats in the north appear very similar to those of Andalucía. In contrast, the south and east are more arid,

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the Sahara forming the border between the Palearctic and tropical African biogeographic realms. This ‘European’ and ‘African’ blend is shared with Algeria and Tunisia (together, the three countries form the Maghreb), but Morocco is special in two ways. First, the Atlantic Ocean provides a relatively moist climate with distinctive landscapes and habitats all along its coast. And second, high mountains dissect the country, providing remarkable upland habitats. In the north is the rather wet Valley in the High Atlas, Morocco. Rif (2,448m). In the centre are the Middle and High Atlas (3,340m and 4,165m), only a handful of species are visible at one place. a formidable backbone that divides the country Standing waterbodies are rare, apart from large into two strongly contrasting sides: the northirrigation dams and daya (temporary pools) after west, which is relatively moist; and the south-east, good rains. It is important, therefore, not to which is almost a desert. In the south are the neglect oued (rivers and streams), which are by Anti-Atlas (2,531m), relatively well watered due far the dominant habitats. They often have many to the proximity of the Atlantic, and the arid Jbel slower and still sections, which harbour most Saghro (2,712m). Moroccan species. The Moroccan dragonfly fauna of 64 species The oued in the western Rif are an excellent reflects this diversity, with a majority of starting point when exploring the north and the Mediterranean species, especially western ones Atlantic coast of Morocco. Trithemis annulata, such as Calopteryx haemorrhoidalis, Ischnura Orthetrum nitidinerve and O. chrysostigma graellsii and Boyeria irene, but also some boreal are easily found, with Paragomphus genei, species with isolated highland populations, such Onychogomphus costae or Trithemis kirbyi as Lestes dryas and Libellula quadrimaculata, and often also present. Oued Loukkos, between species found only in the Maghreb and the Iberian Chefchaouen and Ouezzane, is a good example; peninsula. The latter includes Onychogomphus Oued Laou, near Chefchaouen, offers a more costae and Orthetrum nitidinerve, three montane feel. The coastal marshes of Larache, Maghreb endemics (Calopteryx exul, Platycnemis at the mouth of Oued Loukkos, provide an subdilatata and Enallagma deserti), one Sahara opportunity to observe many species. The African endemic (Ischnura saharensis) and even two Diplacodes lefebvrii is difficult to see elsewhere. Moroccan endemics (Cordulegaster princeps and Further south, at the brackish lagoon of Sidi Bou Onychogomphus boudoti). About 15 species Ghaba near Qenitra, the ‘African’ Orthetrum are typically African. Most of these also occur in trinacria and Brachythemis impartita cohabit Europe’s extreme south (Paragomphus genei, with the ‘European’ O. cancellatum and Aeshna Trithemis annulata, T. kirbyi, Brachythemis impartita mixta. Visit the Oued Massa, and the Youssef and Zygonyx torridus), but Trithemis arteriosa Ben Tachfine dam at the edge of the Anti-Atlas and Pseudagrion sublacteum do not. Ischnura between Agadir and Tiznit, to see southern fountaineae extends deep into the drylands of Asia. species such as Trithemis arteriosa, Ischnura Watching dragonflies in Morocco can be saharensis and, possibly, Pantala flavescens. difficult, and patience and tenacity are needed. The inland and Atlas mountains also offer Suitable habitats are not always accessible by car, interesting opportunities. The aguelmam (lakes) and many can be completely dry in bad years. in the Middle Atlas harbour Palearctic species The flight period is very long and individuals can such as Lestes dryas, Enallagma cyathigerum thus be rather scarce, with an obvious decrease and Libellula quadrimaculata, together with in numbers during the hottest months. Generally, specialities such as Enallagma deserti and the

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subspecies Gomphus simillimus maroccanus. Separating the two Enallagma species here can be very challenging. Recommended aguelmam are Sidi Ali and Azigza, both very scenic mountain lakes, and dayet (sometimes dry ponds) such as Ifrah, Iffer or Aaoua. On these still waterbodies, typical Mediterranean species such as Coenagrion scitulum and Aeshna affinis can be found. The Oued Tizguid near Ifrane (e.g. at a place called ‘source Vittel’), is a wonderful river, where one is almost guaranteed to see such specialities as Calopteryx exul and Cordulegaster princeps. Two African species, Pseudagrion sublacteum and Zygonyx torridus, are among the most attractive in Morocco and are likely to be seen on small or medium-sized oued in the semiarid inner regions of the country, where they generally occur with Onychogomphus costae. Look for them (in rapid sections only) on the Oued Sebou above Fès, or on its tributaries (e.g. Oued El-Ammar at Matmata), and also on the tributaries of the Oued Oum Er-Rbia as they descend from the Atlas (such as Oued El-Abiod, at Imezgane bridge). Onychogomphus boudoti may yet be discovered at such tributaries, having been described as new to science from Khénifra province only in 2014. In the east, you need to cross the Middle Atlas, towards a region called ‘l’Oriental’ by the Moroccans, to see the eastern Ischnura fountaineae. The best site is probably the Oued Za and its tributaries, near Aïn Bni Matar. Here, it is abundant, together with I. saharensis and Enallagma deserti (which here occurs without E. cyathigerum). These three species can also be seen in the Guir and Ziz basins. In the south, the palm groves in the valleys of the Ziz, Todgha, Dades or Draa are magnificent places. It is a delight to stroll along the rivers and saqiya (irrigation ditches) and be rewarded with sightings of Platycnemis subdilatata, Coenagrion caerulescens, Ischnura saharensis, Anax parthenope, Orthetrum chrysostigma, O. nitidinerve, Sympetrum fonscolombii and various Trithemis species. Look for Selysiothemis nigra at a small reservoir near Oued Tata south-west of Tata. Orthetrum ransonnetii was found for the first time in Morocco at this river in 2003. It can also be seen at Oued Tarkal near Zawyat Sidi Blal, together with Sympetrum sinaiticum, another species added to the Moroccan list only recently. Finally, Orthetrum sabina was first discovered in Morocco at Oued Ez-Zahar near Akhfennir in 2007. G Jacquemin

Algeria Algeria is the second-largest country in Africa and has a very varied climate and topography. It can be divided into three regions: a wet coastal strip, with evergreen oak forest and maquis surrounding numerous marshes and garaet (shallow lakes); an area of flat, semi-arid high plateaux, with immense salt lakes known as chott and sabkhas; and the Sahara, sparsely dotted with oases, and where ergs (sand-dunes) alternate with regs (rocky plateaux). A total of 63 species has been recorded; Lestes numidicus was described from Algeria in 2003. Algeria shares many species with Morroco and Tunisia, but less is known about them and the many gaps in the distribution maps in the guide are simply a result of this lack of information. Aside from the prospect of new discoveries, Algeria hosts specialities such as Gomphus lucasii, Urothemis edwardsii and Acisoma inflatum. A visit also offers an opportunity to explore the splendid canyons of the Tassili N’Ajjer and the Ahaggar mountains in the deep south (outside the scope of this book), where African species such as Pseudagrion hamoni and desert specialists like Orthetrum ransonnetii are found. Owing to its subtropical climate, Numidia in the north-east, which has also long been known for its importance for waterbirds, is home to many dragonfly species. Sites such as Lac Tonga, Lac des Oiseaux and Lac Oubeïra have benefited from conservation measures. An exploration of the mosaic of wetlands within the El Kala National Park and environs is likely to be rewarded with swarms of Ischnura graellsii, Erythromma viridulum, Aeshna isoceles and the African species Diplacodes lefebvrii and Brachythemis impartita. The latter follows large mammals, including humans, in the manner of a Cattle Egret (Bubulcus ibis). The fastflowing oued and wadi (rivers) here are likely to yield Calopteryx haemorrhoidalis, Coenagrion mercuriale, Gomphus lucasii and the crepuscular Boyeria irene, while in slower rivers Platycnemis subdilatata and Paragomphus genei are likely to be seen. The coastal dunes house small ponds with a rich fauna and flora. More than 30 species of dragonfly have been recorded in Lac Bleu, including such rarities as Urothemis edwardsii and Acisoma inflatum. There are large seasonal shifts in species composition and behaviour. The beauty of the short North African spring is enhanced by the busy flight of Sympecma

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fusca, Anax ephippiger and Sympetrum fonscolombii. The dark wet alder forest of Lac Tonga is frequented in spring by Aeshna isoceles. Later in the season, seeking relief from the desiccating summer, species such as Lestes barbarus and Chalcolestes viridis fill the shady, moist alder scrub. Shortly before the first autumn rains cast a veil of relief over parched habitats, one can witness the spectacular return flight of millions of Oued Mellègue, Tunisia. Aeshna mixta, Sympetrum meridionale and S. striolatum from their montane extensive stands of bulrushes (Typha). It is worth summer refuges, bustling over the Mekhada investigating any river that crosses the road. marsh and adjacent wetlands. El Kala National Boyeria irene and Onychogomphus uncatus are Park is known for Lindenia tetraphylla and abundant at the rivers Lebgâa and Bransia near Rhyothemis semihyalina. While the former was Aïn Draham, while Pyrrhosoma nymphula and rediscovered at Lac Noir in 2018 after 170 years, Aeshna cyanea, the latest additions to the Tunisian the latter is still considered extinct in Algeria. list, were found in 2014. The Oued Zarga in the The scarcer freshwater wetlands of the high Beja region, the Oued Maden near Nefza and plateaux, such as the lake of Timerganine near the Oued Ghezala near Fernana, for instance, all Oum El Bouaghi, hold abundant Enallagma host the North African endemics Calopteryx exul, deserti, Coenagrion coerulescens, Anax Platycnemis subdilatata and Gomphus lucasii. parthenope and Orthetrum cancellatum. Some The huge central steppe is situated between the permanent streams support Orthetrum nitidinerve southern slopes of the Tell Atlas and the northern or the elusive North African endemic Calopteryx fringe of the Sahara. In this dry region, reservoirs exul. The northern fringe of the Sahara is a may be excellent sites. The best example is a small good place to see Ischnura saharensis and I. lake at the south-east outflow of the Barrage el fountaineae, as well as Sympetrum sinaiticum, Habib, north of the Kairouan–Sbeitla road. Here, Trithemis arteriosa and T. kirbyi. 25 species have been recorded so far, including B Samraoui Lindenia tetraphylla, Onychogomphus costae and Orthetrum nitidinerve. Brachythemis impartita Tunisia is typical of big reservoirs, whereas Enallagma Tunisia is a typical Maghreb country, with a deserti prefers small ones. checklist of 58 dragonfly species. European Most southern sites have the scenic visitors will find a fascinating species composition, combination of dragonflies and palm trees. The containing several Maghreb endemics and, in oases in the northern Sahara provide many addition to well-known Palearctic representatives, a suitable habitats: irrigation channels with pumped good portion of tropical African species. With a car, water, and brackish lakes or swamps where the road map and at least a week to explore, you can water seeps away. The best sites can be found visit a selection of representative sites throughout around Douz and Tozeur, which typically host the country. As the summers are hot, the best times thousands of Ischnura fountaineae, I. saharensis, to visit are mid-May to June and in September. In Sympetrum sinaiticum, Diplacodes lefebvrii and the southern oases, the most species can be seen Selysiothemis nigra. The road north of Gabès between late March and late October. Collecting is crosses the Oued Akarit, one of several fine allowed, except in the few nature reserves. coastal streams hosting Orthetrum trinacria, O. In the Mediterranean north, several small but sabina and Paragomphus genei. Zygonyx torridus perennial rivers have their source in the Tell was found in 2002 at one of the three falls in the Atlas. Their beds are stony and flanked with mountain river that rises at the Tamerza oasis. Oleanders (Nerium oleander), or muddy with R Jödicke

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Identification

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Genus and species order The order of the families is taxonomic (see: Appendix 3). Such an order reflects the evolutionary relationships and richness of the families by always listing the branch of the family tree with the fewest species first. Recent molecular research has confirmed most inferences about odonate phylogeny made previously from morphology. However, the species in Lestidae and a few related families (not found in Europe) have proven to be more closely related to each other than to all other damselflies. Therefore they, rather than Calopterygidae and Euphaeidae, are now listed first. Similarly, Platycnemididae and Coenagrionidae are the nearest relatives to one another, but the former (smaller) family is listed first. Finally, molecular research has shown that the genera Oxygastra and Macromia do not to belong within Corduliidae and they have consequently been moved forward. As a result, the family order in this edition differs somewhat from the 2006 version, but is consistent with contemporary (and future) publications. The order of the genera and species within the families is intended to place more familiar taxa towards the front and similar taxa close to each other. Where possible, we have placed pairs of similar species on opposite pages to facilitate direct comparisons. Genus and species texts The scientific and vernacular names used in this section are discussed on p. 15 of the Introduction and also in the Appendices (pp. 323–330). The genus texts discuss the separation of each genus from other genera, and of the species within the genus. Because similar species tend to have similar behaviour, most behavioural details are also provided here. The Introduction also provides a section on identifying dragonflies, from the level of suborders through families to genera (pp. 21–35). Each species text is divided into an Identification section, including a General opening statement, Field characters, Hand characters, Variation and Behaviour, followed by an Occurrence section, including Range, Habitat and Flight season. For genera with only one species, the genus and species texts have been merged. More background to the arrangement and content of the descriptions is provided in the Introduction. For details, see: Dragonfly identification (pp. 15–21); Dragonfly behaviour (p. 8–9); Dragonfly occurrence (pp. 9–12); Habitat (p. 12–14); Flight season (p. 14); and Map key (p. 12).

Scale of artworks The main colour illustrations are drawn to scale. The damselflies are drawn at 1.8 × life-size, the true dragonflies at 1.4 × life-size. Annotations accompanying the artworks are intended to highlight diagnostic features (or peculiarities of the individual) and should be used in conjunction with the main description text. In each artwork, the head of the animal is to the left or above.

Calyopteryx splendens ?

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Lestidae

Lestes Leach, 1815, Spreadwings & & Chalcolestes Willow Spreadwings Kennedy, 1920 Emerald Damselflies Identification Diagnosis Most damselflies that have metallic colouring, often with a partly pruinose body, and large rectangular Pt (bordering two or three underlying cells), and that perch with wings spread, belong to Lestes or Chalcolestes. Separation from other genera Teneral Lestes and Chalcolestes may perch with wings shut, and damselflies of other families may also occasionally spread them, especially when agitated. The venation (with many pentagonal rather than rectangular cells), rectangular Pt and male appendages recall Sympecma. However, these have a pale brown body marked with dark bronze, and perch with closed wings, in which the Pt of Fw and Hw overlap (see Sympecma for more details). Calopteryx also have metallic bodies and Epallage even combines pruinosity, a long Pt and open wings at rest, but both have numerous (not just two) cross-veins between the wing base and node. Some small damselflies, such as Erythromma, Ceriagrion and especially Nehalennia, can have metallic bodies, but they have small lozenge-shaped Pt, venation like squared mesh and differently configured markings. In case of doubt, non-lestids can always be recognised because no longitudinal veins branch off the vein connecting the arculus and subnode. Separation of the species Lestes is a cosmopolitan genus of about 80 species. Our species have a similar ecology, and therefore

up to five frequently occur together. The two Chalcolestes species known are larger and sleeker than Lestes and never develop pruinosity, and their habit of laying eggs in living wood is unique among European odonates. Besides rather subtle differences in adult and larval morphology, Chalcolestes is genetically closer to Sympecma than Lestes. Species recognition in these genera is instant by reference to the male appendages and, with some experience, by the female ovipositor (plus sheath). Mature males can usually be identified on sight. Separation of C. parvidens and C. viridis in the south-east and L. barbarus and L. virens in the south-west requires additional care. The similar L. dryas and L. sponsa often coexist, so either may be overlooked. Behaviour Hardy species, with cold- and droughtresistant eggs, fast-growing larvae and highly mobile adults, allowing survival in seasonally dry habitats. These features are most strongly developed in L. barbarus and least in both Chalcolestes species, which instead have specialised in ovipositing into the bark of living twigs. Males are very forward, often forming tandems with other species or other males, but hybrids are known only in the closely related C. parvidens and C. viridis. The male escorts the egg-laying female in tandem, and eggs are typically placed in living plant tissue, often well above or away from water. K-D B Dijkstra

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Comparison of Lestes and Chalcolestes species  If the statement agrees, compare the given species. If it disagrees, go to the next statement. Lower border of metallic green area on thorax with prominent spur. Never pruinose

C. parvidens C. viridis

Underside of back of head yellow, sharply contrasting with dark upperside. Pt pale brown and/or yellow, never blackish. At most S9–10 pruinose; S1–2 and S8 never pruinose

L. barbarus L. virens

Thorax without metallic green or bronze areas, wholly pruinose. Pt large, bordering about three underlying cells. Lower appendages less than half as long as uppers. Ovipositor sheath rounded, all dark

L. macrostigma

At least upperside of thorax is bright metallic green or bronze. Pt small, bordering about two underlying cells. Lower appendages more than half as long as uppers. Ovipositor sheath pointed, partly pale

L. dryas L. sponsa

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Lestes sponsa (Hansemann, 1823)

Common Spreadwing



Common Emerald Damselfly dark pterostigma

/ thorax

spots on S1 triangular

70 /

pruinosity covers all of S2

no metallic area on lateral lobes of prothorax

mature ?

/ abdomen tip

pointed bicoloured sheath

ovipositor only reaches end of S10

? abdomen tip

mature ? lower appendages straight with narrow tips

Identification General The most widespread and numerous Lestes in many areas, probably because it is less partial to ephemeral habitats. Field characters Tot 35–39mm, Ab 25–33mm, Hw 17–24mm. Average size and build for a spreadwing. Separated from L. barbarus and L. virens by the dark underside of the head, dark Pt (when mature) and more extensive pruinosity. C. parvidens and C. viridis lack pruinosity, have whitish appendages, a

larger and paler Pt, and a diagnostic thorax marking. L. macrostigma has a larger Pt and a darker, more pruinose body with almost no hint of green. See the similar L. dryas for a full comparison. Hand characters Male’s lower appendages are unique: long and straight, with narrow tips. Female shares the strongly pointed bicoloured ovipositor sheath with L. dryas, but differs by its weaker and shorter ovipositor and details of metallic markings (see L. dryas). Variation Pale areas are distinctly orangish after emergence, and metallic areas are brighter and bluer before pruinosity develops. Females may develop pruinosity as in males, especially on S1–2.

Occurrence Range and status One of the commonest damselflies in most of northern Europe across to Japan, but (largely) absent from most of the south. Habitat Almost any standing water with ample reed-like vegetation. May be more numerous at recent shallow or acidic sites, but not specific to pioneer, ephemeral or bog-like conditions. Flight season Generally from mid-May to midOctober, peaking in August. Most emergence tends to be a few weeks later than L. dryas.

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Lestes dryas Kirby, 1890

Robust Spreadwing

Scarce Emerald Damselfly, Emerald Spreadwing (NA) bright blue eyes

spots on S1 rectangular

/

/ thorax

71 apical third of S2 not pruinose

metallic area extends to lateral lobes of prothorax

mature ?

/ abdomen tip robust build

ovipositor reaches beyond S10

? abdomen tip

mature ?

lower appendages with broad incurved tips

Identification General Often occurs with L. sponsa, with which it is easily confused. Typically occurs in lower numbers, but can be more abundant in sites that are only seasonally wet. With some experience may easily be picked out by jizz, but examination of details with a hand lens is preferred for definite identification. Field characters Tot 35–40mm, Ab 26–33mm, Hw 20–25mm. Size and general appearance close to L. sponsa, but more heavily built (thicker abdomen) and more brightly coloured. Females are especially robust and dark. Males may be separated from L. sponsa by several not entirely reliable features: (1) pruinosity covers only two-thirds of S2, leaving a metallic square on apical half; (2) Pt is less elongate, with sides fairly contrastingly pale; (3) eyes deeper blue (usually duller or darker in L. sponsa, especially dorsally).

Hand characters Tips of male’s long lower appendages are expanded and bent towards each other. Female’s ovipositor is heavier than that of L. sponsa, typically surpassing S10. Two small details of the metallic marking corroborate identification of female: (1) metallic area extends onto lower, lateral lobes of prothorax; (2) the two metallic marks on upperside of S2 are squarish, not triangular. These characters apply to males too, but become covered by pruinosity with age. Variation Similar to L. sponsa, not pruinose and tinged rufous when teneral.

Occurrence Range and status Range similar to L. sponsa, but relatively more common southwards and typically more localised and less numerous than that species in most of its northern range. Our only Lestes that also occurs in North America.

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 Lestes dryas male. Note that pruinosity covers only two-thirds of S2.

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Habitat A wide variety of still waters, which typically dry out in the course of summer or have shallow borders providing warm microhabitats for the larvae. Sites usually have dense growths of rushes or sedges, e.g. dune lakes, reedy shallows, small meadow ponds or edges of bogs; turloughs in Ireland. Flight season The earliest Lestes in most areas, emerging from late May in northern Europe and April in Spain; most abundant in July and August, with the last records in October.

G Lestes macrostigma pair ovipositing, showing the distinct blue pruinosity and large, dark pterostigmas.

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Lestes macrostigma (Eversmann, 1836) Dark Spreadwing extensive bluish pruinosity covering all of head, thorax, S1–2 and S8–10

large dark pterostigmas

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mature ?

/

/ abdomen tip

? abdomen tip

short and straight lower appendages dark ovipositor with rounded sheath

Identification General The dark body, with extensive pruinosity, and the species’ preference for brackish habitats make this spreadwing unmistakable. Field characters Tot 39–48mm, Ab 31–38mm, Hw 24–27mm. Larger and more robust than most other Lestes, never displaying the extensive bright green or bronze colours of those species. At most, S3–7 are bronzy green, otherwise is dark overall and heavily pruinose on head, entire thorax, S1–2 and S8–10. This pruinosity has a distinctive purple to bluish hue, and is extensive in both sexes and already present shortly after emergence. Pt is notably large and dark, bordering about three adjacent underlying cells, and blackish when mature (pale brown in tenerals). Other Lestes have at least the upperside of the thorax bright metallic, and smaller (usually bordering two cells) and/or paler Pt (e.g. L. viridis has large creamy Pt). Hand characters Male’s rather short and simple lower appendages are distinctive. Female’s ovipositor is moderately heavy, and entirely dark and pruinose, with rounded sheath. Variation Body is more metallic when teneral, giving overlying pruinosity a peculiar lustre.

and largely coastal owing to habitat preferences. The Hungarian plains are the best-known inland stronghold. Isolated records suggest this species may wander widely, but low numbers inhibit detection. Habitat Almost exclusively in shallow water with dense rushes, particularly Sea Club-rush (Bolboschoenus maritimus), of coastal and saline inland wetlands, e.g. abandoned saltpans, dune ponds, saltmarsh fringes and steppe lakes. Flight season Emerges as early as late February and March in the Spanish and Turkish Mediterranean, but in central Europe probably mainly present in June, July and August.

Occurrence Range and status Extends from the steppes of Central Asia to Europe, where it is highly localised

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Lestes barbarus (Fabricius, 1798)

Migrant Spreadwing



Southern Emerald Damselfly yellow ‘jowls’

/ thorax pale bicoloured pterostigma

wide antehumeral stripe

74 mature ?

/

? abdomen tip

/ abdomen tip

lower appendages diverge at the tips

pale rounded sheath

pale abdomen tip with no (or limited whitish) pruinosity

Identification General This mobile species is typical of habitats prone to desiccation. It is recognised in most areas by its pale coloration and bicoloured Pt. Field characters Tot 40–45mm, Ab 26–35mm, Hw 20–27mm. Paler and slightly larger than most other Lestes. Shares yellow underside of back of head only with L. virens. This can be viewed from below (the entire area between eyes and labium appears yellow) or from behind, so the clear contrast with the dark metallic upperside is apparent. Tenerals of dark-jowled species may appear pale below, but lack the sharp contrast. Differs from L. virens and others by: (1) pale brown Pt, with outer half pale yellow; (2) pale yellow markings more extensive, most notably in the wide antehumeral stripes, pale-sided S9–10 and whitish appendages; (3)

male is barely pruinose, at most lightly and notably white on S10. In parts of the Iberian peninsula, L. virens also has a two-tone Pt, but its pale portion occupies only about one-third, and bluish pruinosity is typically present on S9–10 of male. Hand characters Male’s lower appendages are uniquely slender and divergent at the tips. Female’s ovipositor and its sheath are wholly pale. The latter is rounded (unlike in L. dryas, L. sponsa and L. virens). Variation Varies considerably in size. Pale areas of tenerals are a bright lemon-yellow. Behaviour Often oviposits in wholly dry areas.

Occurrence Range and status A strong wanderer found east to Mongolia; may be absent from areas for many years, suddenly establishing large colonies that may persist for some time. A southern species that has increased dramatically in northern Europe since the mid-1990s, first reaching Great Britain in 2002. Habitat Favours ephemeral conditions even more than other Lestes species, being typical of sites that dry out early in summer, such as dune slacks, meadow pools and shallow depressions. Flight season From March to October in the southern Mediterranean, but in the north most adults probably emerge in June and July, reaching the greatest densities in August.

Lestes Spreadwings

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Lestes virens (Charpentier, 1825)

Small Spreadwing

includes: Lestes numidicus Samraoui, Weekers & Dumont, 2003

Late Spreadwing

Small Emerald Damselfly / thorax ssp. virens

bicoloured rostigma

back of head yellow pale brown pterostigma

75 /

/ thorax ssp. vestalis

mature ?

no pruinosity at abdomen base

green more extensive on prothorax

? abdomen tip

green extends to metapleural suture

lower appendages short and straight

/ abdomen tip pruinosity confined to S9–10

more pointed sheath than L. barbarus

Identification General Our most delicate Lestes, which is normally easily separated by its stature and coloration, although some populations on the Iberian peninsula recall L. barbarus. These are part of a confusing and unresolved taxonomic situation, which also includes the recently named cryptic species L. numidicus from Algeria. Field characters Tot 30–39mm, Ab 25–32mm, Hw 19–23mm. Slightly smaller and sleeker than L. sponsa. Separated from all species except L. barbarus by the yellow underside of the back of the head. The male’s abdominal pattern is unique, the blue pruinosity being neatly confined to S9–10 but absent from S1–2. Pt is usually uniformly pale brown with whitish sides, but some populations on the Iberian peninsula have the outer third pale yellow and are deceptively close to L. barbarus (see that species), and also easily confused with that species because they are rather pale (see below). Hand characters Male’s lower appendages are noticeably short and straight. Female’s ovipositor is fairly weak; its pale sheath is more pointed than in

L. barbarus but less than L. dryas and L. sponsa. Variation Specimens from Africa, the Iberian peninsula, southern France and the Tyrrhenean Islands have reduced dark markings, most clearly on the thorax: (1) green area on prothorax is small; (2) pale antehumeral stripes are longer and wider; (3) green area does not reach metapleural suture; (4) this suture lacks a narrow dark line. These populations have been known as ssp. virens. Populations to the north and east pertain to ssp. vestalis, which becomes gradually paler towards the Balkans and Turkey. Specimens from the wetter west and south of the Iberian peninsula, including those from which virens was described, have bicoloured Pt (see above). So-called virens in Algeria consists of two populations, separated by reproductive period rather than distribution and morphology. Recently, the autumnal group was split off on genetic grounds as L. numidicus. Specimens are described as tinged reddish rather than green, but in all L. virens forms the metallic colour tends to become more coppery with age. Analysis throughout north-west Africa and south-

Lestes Spreadwings

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 Lestes virens male. Note the pruinose S9–10, but S1–2 not pruinose.

76

F Lestes barbarus male. Note the wide antehumeral stripe and pale abdomen sides.

west Europe, where L. virens also has a long flight season and could thus have distinct reproductive seasons, may resolve this complex situation.

Occurrence Range and status Widespread, although seldom the dominant Lestes species. Distribution recalls that of L. barbarus and also tends to wander like that species, though rarely in similarly great numbers. Habitat A wide variety of seasonally dry shallow and reedy waters in the south (similar to L. barbarus), becoming more critical in the north-west, where it is most abundant in heath and bog lakes with peat moss (Sphagnum) and rushes (Juncus). Flight season Northern populations mostly emerge in July, flying into November. Flies as late in Europe’s far south, but may emerge as early as April here. Emergence is in the second half of April and in May in Algeria; probably includes both individuals that are reproductively active from June to August and those active from late August to early November, the latter named L. numidicus.

Lestes Spreadwings

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Chalcolestes viridis (Vander Linden, 1825) Identification General This and its sister species C. parvidens differ from Lestes species by their appearance and habits, and both species are easily noted for their vivid green colour, pale appendages, large size and habit of hiding in trees and bushes. They are the only European odonates to lay eggs in live wood. Field characters Tot 39–48mm, Ab 29–39mm, Hw 23–28mm. Larger and longer-bodied than other Lestes. Typically relatively bright metallic green, without blue accents (e.g. eyes, pruinosity). Both C. viridis and C. parvidens are characterised by a prominent spur-like marking on the lower border of the metallic green area of the thorax. Neither is ever pruinose; the abdomen tip is entirely dark metallic green, contrasting with the whitish appendages. Pt is large and paler than other species when mature, typically pale brown with a paler cream centre. Note that teneral adults of Lestes species are often mistaken for C. viridis

Western Willow Spreadwing Willow Emerald Damselfly because their Pt is still white and pruinosity has yet to develop. However, the spur marking is absent or reduced in these individuals. Hand characters Male has very short lower appendages and a prominent tooth close to the tip of the upper appendages. Female’s ovipositor is typically partly dark with a pale border (the reverse, or uniformly pale or dark, in Lestes), with a limited number of large and discrete teeth on the lower border. Lestes species have more numerous, but weaker, serrations. Variation Limited, compared to Lestes species, as Pt barely darkens and pruinosity does not develop. Behaviour Adults are usually found hanging in trees and bushes, sometimes far from water. Unlike Lestes, usually oviposits in woody and other compact tissues (less so in the north-east of its range), preferably thin twigs overhanging water. Signs of oviposition remain as track-like scars in the bark.

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 Chalcolestes viridis pair ovipositing into a branch.

Lestes Spreadwings

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body metallic green without pruinosity

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large pale pterostigmas

mature ?

/

? abdomen tip

short lower appendages pale appendages contrasting with dark S10

upper appendages with black on tip extending along border

/ abdomen tip

/ thorax

prominent spur-like marking dark ovipositor with pale border

10–14 large teeth on lower edge

Occurrence Range and status Widespread and common in most of our area; inexplicably absent until recently from Great Britain, but now locally common in south-eastern England. Benefits from urbanisation, favouring garden and park ponds, and possibly expanding northwards. South-eastern range limits are still insufficiently known owing to confusion with C. parvidens, but range extends at least to Sicily and central Greece. Habitat Almost any type of standing or slowflowing water with bordering trees or bushes. Unlike Lestes species, does not favour ephemeral conditions (e.g. shallows, drying-out pans). Flight season In central Europe seldom emerges before mid-July, but already in early May on the Iberian peninsula. Abundant in August and September, persisting into November.

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Chalcolestes parvidens (Artobolevsky, 1929)

Eastern Willow Spreadwing

/ abdomen tip uniform brown pterostigmas

6–8 large teeth

79

Identification General First described as a subspecies of C. viridis from the Crimea, but only in the 1980s and 1990s found to overlap widely with that species in Italy and the Balkans. Can be separated reliably only under magnification, preferably by examination of the male appendages. Field characters Tot 44–50mm, Ab 34–39mm, Hw 22–26mm. Size and general appearance as C. viridis. Two differences may be observed in the field, but cannot be used for reliable identification: (1) Pt darker, uniformly brown (like L. virens) rather than cream-coloured (but these are white in tenerals); (2) upper appendages black only at tips, never on exterior borders, although C. viridis may also have largely white upper appendages, especially in the south-west of its range. Hand characters Male’s appendages bear the most reliable characters (examine with hand lens): (1) inner border of upper appendages with smaller tooth, which is placed slightly more dorsal than in C. viridis (view from above); (2) lower appendages with more slender and strongly up-curved fine tips (view from side). Females have 6–8 teeth on the border of the ovipositor, whereas C. viridis has 10–14; occasional females with nine teeth are inseparable. Variation Hybrids with C. viridis, intermediate in male’s appendage features, occur in areas of overlap, but are quite rare. Behaviour As with C. viridis, but may oviposit more frequently in non-woody materials. In mixed populations in Italy, most activity of C. parvidens was observed in the morning, of C. viridis in the afternoon.

mature ?

? abdomen tip

relatively small tooth upper appendages with black tip

lower appendages (side view) up-curved fine tip

L. parvidens

L. viridis

Habitat As C. viridis; possible ecological differences are poorly known. Flight season Late April to late November. Emergence occurs in late May in Italy and the north-west Balkans; after a maturation period of several weeks, adults return to their breeding sites late in July. Emergence is possibly earlier on average than in co-occurring C. viridis.

Occurrence Range and status Overlaps with C. viridis across a huge area south of the Alps and Tatra. Outnumbered by C. viridis in Italy and the northern Balkans, but much the commoner species in the southern Balkans (e.g. Bulgaria and Greece). Extends to the Caucasus region, northern Iran and the Levant.

Lestes Spreadwings

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Sympecma Burmeister, 1839

Winter Damsels Winter Damselflies

80

 Sympecma fusca male.

Identification Diagnosis Body brownish, with dark bronze dorsal markings, characteristically torpedo-shaped on the upperside of S3–6. The pale brown Pt is an elongate rectangle, standing in front of two cells. In the Fw the Pt is distinctly nearer to the wing apex than in the Hw; this feature is easily seen when the wings are closed in rest and the Pt of Fw and Hw hardly overlap. Separation from other genera Venation (with many pentagonal rather than rectangular cells), long Pt and male appendages recall Lestes and Chalcolestes but, although the metallic markings are greenish in teneral Sympecma, there is never the bright green coloration of Chalcolestes and

Lestes. In Sympecma the wings are narrower and more pointed; moreover Chalcolestes and Lestes typically rest with wings half-spread, and if their wings are closed, Fw and Pt of the Hw broadly overlap. The hind rim of the pronotum in Chalcolestes and Lestes is simply rounded, but in Sympecma it is trilobed: two incisions produce a pronounced central lobe and two lateral lobes. The Sympecma ovipositor is weak and short, its tip extending only halfway along S10; the female appendages are notably large and pale. Brown females of Enallagma have a similar abdominal pattern, but have a small lozenge-shaped Pt, short dark appendages, a spine at the ovipositor base, and differently configured and blacker

Sympecma Winter Damsels

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markings on the head and thorax, including postocular spots. Separation of the species This Palearctic genus contains three species, one confined to Central Asia and Iran, the other two ranging into Europe. Their separation requires close examination (preferably in the hand) of the thorax markings and the male appendages. Behaviour Cryptic, due to colour and behaviour. At rest, the wings are closed and carried along one side of the abdomen. Individuals often press their body against the perch for camouflage. Adults emerge in late summer and reproduce in the

following spring, after hibernation, unlike other European odonates that winter as eggs or larvae. After emergence they appear in open, structured landscapes, such as rough meadows and heaths with shrubs or forest borders. Here, they forage during autumn and also spend the winter, surviving the cold on an exposed perch, pressed against a stem, or concealed under stones and bark. Few adults are seen from November to March, but they may appear on sunny days. Oviposition takes place in tandem, most often in floating dead plant material. R Jödicke

G Hibernating Sympecma fusca female covered by frost.

G Pair of Sympecma paedisca copulating. Note the long brown pterostigmas that do not overlap.

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Sympecma Winter Damsels

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Sympecma fusca

Common Winter Damsel

(Vander Linden, 1820)

Winter Damselfly straight outer margin to stripe on upper thorax

long brown pterostigmas

82 mature ? mature /

/ abdomen tip ? abdomen tip

tips of lower appendages reach to or beyond tips of basal teeth of uppers

Identification General A common but easily overlooked damselfly, dainty and drab. Males and tandems may suddenly appear on the first sunny days of spring, perching and ovipositing on the reeds and rushes of the previous summer. In the north and east, lookalike S. paedisca may be concealed among populations of S. fusca, or vice versa. Field characters Tot 34–39mm, Ab 25–30mm, Hw 18–23mm. Slightly longer than Enallagma cyathigerum. The body is pale brown, with glossy

dark markings on the upperside of the head, thorax and abdomen. Can be mistaken only for S. paedisca (see that species), but has straight outer margins of the dark band on the upperside of the thorax. Hand characters The male’s lower appendages are slim, and their tips reach as far as the tips of the basal teeth of the upper appendages. Variation Dark markings tend to be less extensive in the south. The brown ground colour darkens until autumn and may become blackish in very old individuals. In spring, mature individuals have a blue spot on the upperside of the eyes.

Occurrence Range and status Common in a large part of our region, extending to Central Asia. Scarcer and fluctuating at the northern fringe of its range, where currently rapidly expanding. Habitat All kinds of well-vegetated standing waters, especially where there are floating dead reeds or rushes. Flight season Adults may be seen throughout the year, but are most reproductively active in April and May, while late summer activity peaks in August and September. Especially southwards, succeeding generations may overlap in summer.

Sympecma Winter Damsels

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Sympecma paedisca  (Brauer, 1877)

Siberian Winter Damsel bulge on outer margin of stripe on upper thorax

oblique line on side of thorax narrower than in S. fusca and sometimes broken

mature ?

83

? abdomen tip / abdomen tip tips of lower appendages fall short of tips of basal teeth of uppers

Identification

Occurrence

General More northern and eastern sister species of S. fusca, closely resembling it in coloration, behaviour and biology. Suitable habitats in central Europe often harbour both species, requiring careful comparison. Field characters Tot 36–39mm, Ab 25–29mm, Hw 18–22mm. Same size as S. fusca. Separation in the field is subtle: the outer margins of the dark area on the upperside of the thorax (just behind the middle) bear an abrupt, roughly rectangular bulge (followed by a slight indentation). The oblique dark streak on the thorax sides is normally thinner than in S. fusca. Hand characters The only reliable structural character is the shape of the male appendages: the lowers are significantly shorter than in S. fusca, reaching only about halfway to the tips of the basal teeth of the upper appendages. Those teeth also lie slightly further from the base in S. paedisca. Variation Asian individuals may lack the distinctive bulge, but this variation is not known to occur in our area. Darkens with age like S. fusca, which may make thorax markings hard to discern. The dorsal half of the eyes in mature males (in spring) is blue, whereas females develop a blue eye spot like S. fusca. Behaviour Like S. fusca, eggs are laid in dead plants, but also in living ones.

Range and status Huge northern Eurasian range extends to Japan. Common in eastern Poland and further eastwards, but very local and often endangered to the west, where limited to two fronts: one through the northern German and Dutch lowlands, the other along the foothills of the Alps. Habitat Similar to S. fusca, breeding in almost any type of standing water in the east. The more localised occurrence in western Europe is difficult to associate with a particular habitat. Flight season Emergence begins in late July, oviposition takes place mostly in April and May.

Sympecma Winter Damsels

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Calopterygidae

Calopteryx Leach, 1815

Demoiselles Jewelwings (NA)

84

 Calopteryx splendens male. The colourful wings of demoiselle males are displayed in aerial courtship dances

Identification Diagnosis Large, broad-winged damselflies with metallic bodies and veins. Males are colourful (most often blue) and often have extensively coloured wings. Females are a metallic green or brown, with clear to brown wings; rarely they develop male colours. The legs are noticeably long and spidery, with especially numerous and long bristles. The wings are unstalked (i.e. the bases narrow gradually and have numerous cross-veins) and exceptionally densely veined (e.g. with 18 or more antenodal veins). Males lack Pt and females have pale pseudopterostigmas, which, unlike true Pt, are weakly defined and are crossed by veins.

Separation from other genera The wing coloration, metallic bodies, absence of (true) Pt and dense venation are diagnostic. Lestes and Chalcolestes species are metallic green, but are smaller, with narrow, stalked and unmarked wings. The large Epallage has similar wings but lacks a metallic sheen, and has short leg spines and long Pt. Separation of the species Structural differences between species are insignificant (e.g. male appendages are almost identical in all), but the wing shape and coloration, metallic colour and the colour of the ‘tail-light’ (underside of abdomen tip) are sufficient to separate males. Distribution is

Calopteryx Demoiselles

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also helpful: in most of our area only C. splendens and C. virgo occur. Females can be difficult to identify; it is often easier to look for nearby males. Numerous, generally weakly defined subspecies have been named for the wide ranging species C. haemorrhoidalis, C. splendens and C. virgo. The two others, C. exul and C. xanthostoma, are sometimes considered subspecies of C. splendens. Hybridisation between (alleged) subspecies is common, producing broad transition zones between them. Hybrids between overlapping species also occur in nature, even between the widespread C. splendens and C. virgo, although

these are not closely related. Male hybrids show intermediate characters with C. splendens-like wing spots and a C. virgo-like reddish ‘tail-light’. Behaviour The resting posture is distinctive, with raised abdomen and closed wings. Both sexes are found along flowing water, often in great numbers. Here, males avidly defend territories near suitable oviposition sites (e.g. submerged aquatic vegetation). They attract females with elaborate aerial courtship dances, demonstrating their colourful wings and ‘tail-lights’. Often gather in large bankside roosts in the evening. H J Dumont

85

Calopteryx splendens (Harris, 1780) Banded Demoiselle Identification General This ‘streamside butterfly’ is our most familiar riverine odonate. It is a variable species and numerous forms have been named. C. xanthostoma and C. exul are most distinct in this complexity and are here treated separately as good species. Field characters Tot 45–48mm, Ab 33–41mm, Hw 27–36mm. Largest damselfly, except for Epallage and other Calopteryx. Males are metallic blue with a dark blue band across each wing (actually brown, crossed by metallic blue veins). In extreme cases, wing markings may be absent, although the metallic veins still make the wings appear blue. Male ‘tail-light’ (underside S8–10) is bright yellow to greyish white, with a black stripe along the middle of S8 and usually S9. S10 is often whitish pruinose in older males. Typical females are less conspicuous, with a metallic green body and veins, clear greenish wings (not tinted brown) and usually a white pseudopterostigma. In andromorph females, which have male wing colours, the wing band contrasts with the white pseudopterostigma (lacking in males). See other species for their separation. Variation Wing markings are present at emergence, but not yet fully coloured. The extent of the male wing colour varies widely individually and geographically. Many subspecies have been proposed based on this variation, but there is much variation within these and also broad zones of introgression between them. Most of this diversity results from extensive hybridisation following

the expansion of populations that were isolated during the last Ice Age. The mixed populations are thus not genetically or geographically distinct and therefore do not warrant subspecies status. Beyond our area, entirely unbanded C. splendens populations (recalling C. exul) occur.

Occurrence Range and status Often numerous, extending to Lake Baikal and north-west China. Being conspicuous, wandering males are often noted far from suitable habitat. Habitat Most (partially) open running waters, avoiding cold torrents, high mountains and deep shade, and scarce on large rivers. Flight season Late April to October in the south; May to August in the north.

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yellowish

? abdomen apical segments (view from below)

86

pale pseudopterostigma

? ssp. splendens

wings greenish

/

? ssp. balcanica forms with even more blue (including wing tips and more of the base) occur in Turkey

/ ssp. balcanica (androchrome)

Calopteryx Demoiselles

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Calopteryx xanthostoma 

Western Demoiselle Yellow-tailed Demoiselle

(Charpentier, 1825) Identification General Replaces C. splendens on the Iberian peninsula, southern France and the Ligurian coast of Italy. Sometimes treated as a subspecies of that species, but genetics indicate that it is distinct. In the area of overlap, C. xanthostoma is best recognised by the male’s completely dark wing tips. Southeastern subspecies of C. splendens are inseparable by wing pattern, but do not overlap in range. Field characters Tot 45–48mm, Ab 35–37mm, Hw 28–31mm. Size and habits as that of C. splendens, but the male’s wing colour always reaches the wing tip and rarely extends far beyond the node. The male’s yellow ‘tail-light’ is often given as a difference with C. splendens, where it is supposedly whitish, but this is unreliable. The underside of the last three abdominal segments is much more broadly black: a wide stripe on S8, a stripe and often additional spots on S9. In old males, most of S8 darkens, sometimes even becoming metallic. Females are green metallic and hard to separate from C. splendens, although typically the dorsal side of

the last abdominal segments is not metallic green, but dull brown. Variation Unlike C. splendens, the wing colour is absent at emergence, developing during the first week of adult life. There is some variation in the extent of the wing colour between the base and node. No andromorph females are known, unlike the similarly marked south-eastern subspecies of C. splendens, where such females are frequent.

87

Occurrence Range and status Abundant in southern France, becoming scarce towards the south of the Iberian peninsula. Frequently hybridises with C. splendens, e.g. in Liguria and between the Loire and Garonne in France. North African records are probably erroneous. Habitat Like C. splendens, but inhabits larger watercourses where that species also occurs, e.g. lowland rivers and canals. Flight season April to September.

pigmented area always extends to wing tip

?

Calopteryx Demoiselles

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Calopteryx virgo (Linnaeus, 1758) Identification

88

General The largest and darkest Calopteryx, with (almost) entirely metallic cobalt wings in males and transparently greenish to deep ebony wings in females. The flight is butterfly-like, but heavier than in C. splendens, and the species is more often seen in deep forest shade. Field characters Tot 45–49mm, Ab 31–42mm, Hw 24–36mm. Relatively large and robust. Wings, especially the male Hw, are broader than in other Calopteryx, and almost paddle-shaped. Male wings appear completely purplish blue (but see Variation). Male ‘tail-light’ is brown to reddish, extensively marked with black (paler or unmarked in other species). Only large, dark male forms of C. splendens (e.g. Turkish ssp. intermedia) could be mistaken for C. virgo (especially the paler ssp. meridionalis of south-west Europe), but potentially confusing forms are widely separated geographically. Moreover, the C. splendens ‘taillight’ is yellowish. Females are metallic (brownish) green and are hard to separate from C. splendens, but the wings tend to be broader and browner (sometimes very darkly so). Variation Three main forms can be distinguished by wing colour, and are normally treated as subspecies. In most areas (roughly north of the Loire, Alps and Danube), virgo is found, characterised by males with only the extreme tips (especially Fw) and base paler (looking ‘washed out’) than the rest of the wing. The rather distinctive meridionalis occurs roughly

Beautiful Demoiselle

from Normandy and Geneva southwards. It has distinctly clear wing bases but entirely dark tips; the delimitation of the clear area is rather sharp (not ‘washed out’). In Turkey and the southern Balkans (along the coast at least as far as Montenegro and south-east Romania) festiva is found. This is the largest European Calopteryx, with entirely purpleblue wings in males. Females with dark brown wings may emulate the male’s subspecific wing pattern. The extent and distribution of intergrades between the three forms are poorly known. For instance, in Italy, meridionalis, festiva and intermediates occur. Occasional females around the Black Sea (festiva) and western Mediterranean (meridionalis) have darker Hw tips, recalling C. haemorrhoidalis.

Occurrence Range and status Locally common, but due to the poor differentiation of its eastern sister species, C. japonica, it remains unclear how far C. virgo extends beyond the Urals into Siberia. Absent from large areas in the south, e.g. many major islands, and in Anatolia mainly limited to coastal areas; rare on the plateau. Habitat Prefers cooler running waters than other Calopteryx, typically smaller, more shaded, at higher latitudes or karstic: small forest streams are the classic habitat. Where streams broaden or open up, C. virgo gives way to C. splendens. There may be broad overlap and occasional hybridisation. Flight season May to late September.

? ssp. festiva

wings entirely pigmented

Calopteryx Demoiselles

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reddish

wing pigment extensive

? abdomen apical segments (view from below)

89 hindwing broad

? ssp. virgo with relatively restricted wing markings /

wings brownish

/

? ssp. meridionalis

dark tips

well-defined clear basal area

Calopteryx Demoiselles

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Calopteryx haemorrhoidalis (Vander Linden, 1825) Identification

90

General A stunning demoiselle, easily identified by the carmine ‘tail-light’ of the male and dark apical Hw spot of the female. The copper-red body of the male is also distinctive in most areas, but not present in all populations of this western Mediterranean endemic. Field characters Tot 45–48mm, Ab 30–43mm, Hw 23–37mm. Typical males are unmistakable by their reddish-bronze body and largely dark wings, which appear dark brown (rather than blue) when closed and have a

clear forewing tips

? ssp. occasi

Copper Demoiselle

conspicuous clear area at the base. The demarcation of the clear base is notably oblique because the coloured area runs almost to the wing base on the fore edge. Locally, males appear brown, purplish, dark blue, olive or almost black. This colour is always darker and duller than the bright green to blue of C. splendens and C. virgo, while the wings appear brown to black, not blue. Males have the underside of the abdomen tip uniformly bright pink to vivid red. The metallic green to bronze females are easily recognised by their contrasting dark Hw tips and, unlike other females, have narrow, pale humeral lines. In both sexes, the tibiae are frequently rusty brown (black in most other Calopteryx). Variation Male body colour may vary (see above). Variation in male wing patterns intergrades widely, and does not seem to warrant the recognition of subspecies. In typical haemorrhoidalis, two-thirds of the male wing is dark; at most, narrow apical fringes are clear. North-western populations have paler wings: males with broadly clear Fw tips (known as occasi) occur along the Ligurian coast of northern Italy, in France away from the Mediterranean, and on the southern slopes of

copper-red colour

 Calopteryx haemorrhoidalis female, showing the dark hindwing tips with contrasting pale pseudopterostigma, and brown tibiae.

Calopteryx Demoiselles

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? abdomen apical segments (view from below)

91 oblique border to pigmented area of wing

narrow, pale humeral lines

? ssp. haemorrhoidalis (very dark individual; typical colour more copper-red as male on previous page)

dark hindwing tip

/

the Pyrenees. On the Spanish north coast, the clear apical area extends basally along the Fw hind margin, leaving a distinct squarish dark spot. Some individuals from the Iberian peninsula are notably small. Smallish, pale males from coastal Asturias have been named asturica. Records of C. haemorrhoidalis from outside the western Mediterranean refer to exceptionally dark-tipped females of subspecies C. virgo festiva.

Occurrence Range and status Occurs only around the western Mediterranean, including the major islands. Generally common; the most widespread Calopteryx in north-west Africa. Habitat Clear streams and rivers, where it occurs with other Calopteryx; these are often smaller and more shady. Also found in larger waters in full sun. Flight season May to September.

Calopteryx Demoiselles

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Calopteryx exul Selys, 1853

Glittering Demoiselle

clear, narrower wings

92

 Calopteryx exul male.

?

Identification General Replaces C. splendens in north-west Africa, where it may be noted by its narrow, unbanded

EN

wings, which flash conspicuously in its rather rapid flight, often skimming the water surface. Field characters Tot 45–50mm, Ab 34–36mm, Hw 27–29mm. In both sexes the wings are narrower than in all other members of this genus, and remain completely clear. The male’s venation is metallic, producing a light blue flash on each wingbeat. Legs, especially femora, tend to be paler than in other species, and in both sexes are chocolate brown rather than black, or black with yellow internal sides.

Occurrence Range and status Localised endemic of the mountains from Morocco to Tunisia. Currently rare in Algeria, where most of its habitat has been polluted. Typically only a few individuals are seen at a time. Habitat Streams and rivers between 200m and 2,000m altitude. Flight season May to August.

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Euphaeidae

Epallage Charpentier, 1840 Odalisques Epallage fatime (Charpentier, 1840)  Identification General A large and heavy damselfly immediately recognised by its ‘anisopteran’ build and posture. The abdominal pattern of females and fresh males and the non-metallic blue abdomen of mature males are unmistakable in combination with the black wing tips. The single-species genus Epallage is the only representative of the Euphaeidae in our area. The family otherwise consists of more than 70 south-east Asian species, mostly satinwings of the genus Euphaea. Field characters Tot 40–50mm, Ab 28–37mm, Hw 30–34mm. Similar in size to Calopteryx splendens but very differently proportioned, with a much more robust body, narrower wings and short leg spines. Unmistakable, although flying males could be confused with pruinose libellulids at first sight. Females have a whitish-yellow or pale bluish body, with a black pattern that becomes more extensive with age. Fresh males are largely black, with pale stripes on the sides of the thorax and abdomen. They become entirely blue with pruinosity, within days of emergence. The very long Pt is unique; it

Odalisque

is dark greyish blue in males and conspicuously white to grey in females. The dark wing tips usually extend to halfway along the Pt (but see Variation). Female wings are often suffused with pale yellow and have an amber patch at their base. The presence of 12–14 antenodal veins distinguishes E. fatime from all other regional species (Calopteryx, also with unstalked wings, has 18 or more). All other damselflies have stalked wings (i.e. narrowed bases) with just two antenodals. Variation Both the extent and intensity of the dark wing tip can vary considerably within and between populations. Some specimens have wholly smoky wings, while others lack markings altogether. This variation led to the description of several invalid subspecies. Behaviour Both sexes are most often seen perched on twigs or stones above or near the water, sometimes in high densities, showing little interaction. It is the only damselfly in our area to sit with its wings spread and the abdomen held horizontally, or even raised. The short, fast flight also appears ‘anisopteran’. Oviposits in tandem, in

93

F Epallage fatime pair in tandem, showing the robust ‘anisopteran’ body, dark wing tips and pruinose body in male.

Epallage Odalisques

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organic material such as roots, or on twigs and pine needles jammed between boulders or vegetation.

Occurrence

94

Range and status Not uncommon in Turkey, Cyprus and Greece, including several of the latter’s larger islands. Scarce in the east of Bulgaria. Likely to be present in the south of Albania. Extends east to Afghanistan and Pakistan. Habitat Running waters with mostly rocky beds; highest densities are along streams, but also found on rivers. Flight season In Turkey, from the end of April to mid-August. V J Kalkman

dark wing tip and very long pterostigma

/

mature ? pruinose body

G Epallage fatime female. This sex may have only dark wing tips, but can have entirely pigmented wings.

fresh ?

short leg spines

Epallage Odalisques

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Platycnemididae

PlatycnemisFeatherlegs Burmeister, 1839 White-legged Damselflies Identification Diagnosis Pale, non-metallic damselflies with a noticeably wide head. The legs have numerous, relatively long bristles, often widened tibiae and much white (especially males), making them feather-like in appearance in both sexes. The quadrilateral cell in all wings is almost rectangular; usually separated from subnode (the vein running down from the node) by two cells. Head, legs and venation are unique among our damselflies. Details of markings are also unequalled. Separation from other genera Coenagrionid damselflies with similar coloration have stouter heads, and unexpanded, less bristly and darker legs. Also, the anterior border of the discoidal cell is distinctly shorter than the posterior one, with most often three cells to subnode. Only P. pennipes becomes blue, but even so the black markings are very differently configured (see species text).

Separation of the species Identification of our six species is simplified when ranges are regarded. Males are separated by their colour at maturity and details of tibiae and appendages. Females are coloured similarly between species and differ less in tibial features. They should be separated by careful study of the prothorax. See table below. Behaviour Males perch on the banks or in nearby vegetation. They search for mates along the shoreline or borders of vegetation with a slow, characteristic zigzagging flight. Males grasp the female by landing on her thorax, often pouncing directly onto her in flight. After copulation, the pair flies off in tandem to oviposit. Sites with egg-laying pairs attract other pairs, resulting in large groups laying together. In both sexes, the legs are not used in courtship, but for threatening behaviour. They are presented to, and waved at, individuals of the same species in flight. A Martens

95

Comparison of species. Tibiae are white with a black line of variable length. The upper appendages are usually notched at the tip; the relative length of the tips above and below this notch is distinctive. The / pronotal hindlobe may have a paired set of teeth, whose size and position differ. Compare species in:

Range

NW Africa

Turkey

SW Europe

France, Spain and Portugal

Widespread; not on Iberian peninsula or in Africa

SE Turkey

Abdomen mature ?

Width hind tibiae; black markings ?

Tips of upper appendages ?

Pronotal hindlobe teeth /

Species

White to green-blue

Narrow; complete line

About equal

Absent

subdilatata

White

Very broad; at most small at base

Lower longer

Small, sublateral

latipes

Orange to reddish

Very narrow; complete line

Upper longer

Large, lateral

acutipennis

Pale blue

Broad; at least small at base, often a line

About equal

Absent

pennipes

White

Broad; unmarked

Tip unnotched

Small, lateral

dealbata

Grey-blue pruinose

Very narrow; complete line

About equal

Absent

kervillei

Platycnemis Featherlegs

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Platycnemis pennipes (Pallas, 1771)

Blue Featherleg White-legged Damselfly

hind tibia of ?

96 immature

/

/

? black distributed more distally than in other blue damselflies

/ pronotum (from behind) ‘double’ antehumerals

mature

?

? appendages (side view)

Identification General The pale blue males, showing off their white legs to each other, are a familiar sight along rivers and calm streams in large parts of Europe. Here, it is often the only Platycnemis and is easily recognised as being the only small damselfly with a wide head and feather-like legs. Field characters Tot 35–37mm, Ab 27–31mm, Hw 19–23mm. Somewhat larger than Coenagrion puella or Enallagma cyathigerum. Males are the only Platycnemis that become all blue, but appear paler than other blue damsels (e.g. Coenagrion) due to less black and a lighter blue colour. In details, the black markings in this and other Platycnemis differ markedly: (1) face more extensively pale and band across head runs between ocelli, not in front of them; (2) it appears as if there are two pairs of antehumeral stripes rather than one; (3) black markings on S7–10 are not interrupted by a pale ‘tail-light’ but are separated by a fine pale central line, thus appearing ‘paired’. Overlaps with other Platycnemis

species only in south-western and south-eastern extremes of range; confusion is most likely with P. latipes in France, and P. dealbata in Turkey. Both these species have porcelain-white mature males, with wide (almost) unmarked tibiae, but remember that P. pennipes males are white before they turn blue (see Variation). Hand characters Male upper appendages are as long above apical notch as below it. Hind margin of female pronotum lacks lateral teeth. Variation Highly variable in the extent of black markings, depending on ambient humidity at emergence: individuals that emerge with dry weather have smaller markings, e.g. black markings on tibiae and S2–6 may not be apparent. Thus the patterns may be regionally and seasonally different. Colour changes from transparent pink immediately after emergence, through white, in both sexes, to blue in males and yellowish brown, greenish or sometimes blue in females. Males on the Adriatic coast of Montenegro, Albania and Greece have, on average, broader tibiae with

Platycnemis Featherlegs

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fewer markings. They are treated as the subspecies nitidula (also known as hyalinata) and may recall the (non-overlapping) P. latipes and P. dealbata. Behaviour Egg-laying tandems aggregate on flower stems of Yellow Water-lily (Nuphar lutea), as well as on water-milfoils (Myriophyllum) and pondweeds (Potamogeton), driftwood and roots.

Flight season From the start of May to the end of September in central Europe, longer in Turkey and shorter in England and Fennoscandia; the main months are June and July.

Occurrence Range and status Occurs from the Atlantic to the Jenisei in Siberia; often abundant. Not threatened, but the range-restricted ssp. nitidula may require closer monitoring and protection. Habitat Characteristic of floodplains, and dominant on oxbows, rivers and open stretches of streams. Also lakes and a wide range of man-made habitats such as canals, gravel pits and fish ponds; the larvae are well adapted to occur with fish. In the north-west (e.g. England, the Netherlands, Fennoscandia), almost confined to flowing waters.

97

F Platycnemis dealbata male. Note the largely white abdomen and legs.

F Platycnemis pennipes male. Note the distal distribution of the black on the abdomen, and the ‘double’ antehumeral stripes typical of all Platycnemis.

Platycnemis Featherlegs

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Platycnemis subdilatata Selys, 1849 Barbary Featherleg

98

? appendages (side view)

/

?

hind tibia of ?

/ pronotum (from behind)

Identification General Endemic to north-west Africa, where it is unique by genus characters. Field characters Tot 33–36mm, Ab 22–28mm, Hw 17–21mm. The only small, pale damselfly in its range with a wide head, almost rectangular

discoidal cell and feather-like legs. Most similar to P. pennipes in extent of black on abdomen and legs, but mature male is white to pale greenish blue instead of blue; females become white. Tibiae are narrower than in P. pennipes. Hand characters Male appendages and female pronotum are very similar to those of P. pennipes. Variation Like congeners, is variable in extent of black markings. Head, legs and thorax of young females are conspicuously reddish.

Occurrence Range and status Endemic to Morocco, Algeria and Tunisia, where it is common; no records south of the Atlas Mountains except from some Algerian oases. Probably occurs throughout northern Algeria. The record of a single museum specimen labelled as from Tenerife (Canary Islands) has not been confirmed in the field. Locally threatened by overexploitation of water in more arid regions. Habitat All permanent stretches of running waters. Flight season Mid-April to the end of September.

Platycnemis Featherlegs

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Platycnemis acutipennis Selys, 1841 Orange Featherleg

Orange White-legged Damselfly

upper part of tip longer than lower

99

? appendages (side view) hind tibia of ?

/

?

lateral tooth on hind edge of pronotum

combination of blue eyes and reddish body is unique

/ pronotum (from behind)

Identification

Occurrence

General Reddish colour and narrow tibiae may not immediately suggest a Platycnemis species. The blue-eyed orange-bodied male is one of our most attractive damselflies. Field characters Tot 34–37mm, Ab 24–28mm, Hw 18–19mm. In markings and shape of the head, a typical Platycnemis, but mid- and hind tibiae not conspicuously enlarged and mature adults have an orange-red abdomen. P. pennipes and P. latipes, which overlap in France, have significantly expanded tibiae and only some females may have a reddish wash. The mature colour is less intense than that of Ceriagrion tenellum or Pyrrhosoma nymphula, which are differently marked. Moreover, P. acutipennis has bluish eyes, like other mature Platycnemis males – a unique colour combination. Hand characters Upper tip of male’s upper appendages longer than lower tip. Female hind margin of pronotum with pair of distinct lateral teeth. Variation Abdomen whitish in immatures, more closely recalling congeners. Behaviour Deposits eggs in fresh, floating aquatic plants, as well as roots or driftwood.

Range and status Endemic to Iberian peninsula and France, where generally common. Habitat Running waters, mostly with fast or medium current. Rarely at standing waters. Flight season End of May to mid-August in most of range; earlier in southern Spain, from late March to late June.

Platycnemis Featherlegs

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Platycnemis latipes Rambur, 1842

White Featherleg

upper part of tip shorter than lower

100 ? appendages /

(side view)

? hind tibia of ?

largely white abdomen and legs

sublateral tooth on hind edge of pronotum

/ pronotum (from behind)

Identification General Within the species’ range, mature males are unique by their porcelain-white body and expanded legs with limited black markings. These ghost-like damsels are often abundant on rivers in Europe’s warm south-west. Field characters Tot 33–37mm, Ab 25–30mm, Hw 18–22mm. Males are paler, with broader tibiae, than overlapping species; the abdomen is creamy

white, with usually at least S2–5 unmarked. In both sexes, the mid- and hind tibiae are without or with only a short black basal marking. P. subdilatata (North Africa) and P. dealbata (Turkey) are similar, but do not overlap in range. Beware of teneral P. pennipes males, which are whitish (not yet blue) and may have less intense dark markings. Hand characters Male upper appendages with upper tip shorter than lower. Pronotum of female with pair of short lateral teeth on the hind margin, which are larger and placed more outward in P. acutipennis, but mostly absent in P. pennipes. Variation The extent of the black markings on S6–10 varies greatly.

Occurrence Range and status Endemic to Iberian peninsula and the south-western half of France; more common than P. pennipes where they overlap. Habitat Running waters, mostly with medium or slow current, such as larger rivers; rare in mountain areas. Flight season Mid-June until the end of September. In southern Spain, from early May to late August.

Platycnemis Featherlegs

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Platycnemis dealbata 

Ivory Featherleg

Selys in Selys & Hagen, 1850

tip of upper appendage unnotched

101

? appendages (side view)

/

?

both sexes may have more black on abdomen than shown

largely white abdomen and legs

hind tibia of ?

weak tooth on hind edge of pronotum

/ pronotum (from behind)

Identification General Within their range, mature males are unique in having an ivory body and enlarged white legs. Almost identical in this regard to P. latipes, but separated by structural details and range. Field characters Tot 28–38mm, Ab 21–30mm, Hw 16–19mm. Both sexes differ from overlapping P. pennipes and P. kervillei by (almost) unmarked white mid- and hind tibiae. Moreover, male abdomen is almost all white, seldom more than S7–9 marked with black; usually not blue (P. pennipes) and never pruinose (P. kervillei). Hand characters Unlike other Platycnemis, upper appendages of male lack a notch at tip. Hind margin of pronotum in female with a small but prominent tooth laterally. Variation As in P. latipes, the extent of the black markings on S6–10 varies. In old males the white of thorax, abdomen and tibiae may show a bluish, almost iridescent, tinge.

coast, Central Asia and northernmost India and Pakistan; limited in the south by the Sinai, Syrian Desert and Persian Gulf. Often abundant. Habitat All types of running waters. Flight season In Turkey, from late April to the end of September; from March to October in the Near East.

Occurrence Range and status Just occurs in the far south-east of our area, in Turkey. Extends to the Caspian

Platycnemis Featherlegs

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Platycnemis kervillei (Martin, 1909) Powdered Featherleg tip of upper appendage notched

? appendages (side view)

102

/

? hind tibia of ?

entirely pruinose when mature

/ pronotum (from behind)

Identification General Mature male does not instantly recall Platycnemis, with its ‘normal’ (not widened) tibiae and whitish-blue pruinose body, but head shape, bristly legs, venation and appendages are typical of the genus. Field characters Tot 30–38mm, Ab 24–28mm, Hw 18–22mm. Females and teneral males appear as typical Platycnemis by markings, but tibiae in both sexes not clearly widened. Males become almost uniformly pruinose on head, thorax and abdomen. The Syrian Sprite Pseudagrion syriacum occurs in the Levant, with Turkish records just outside the scope of the guide. Mature P. syriacum males are

also pruinose, but more patchily, e.g. forming distinctive antehumeral stripes. They also often retain small greenish postocular spots and lack typical Platycnemis features (see genus text p. 95). Hand characters Upper appendages of males with notched tip. Hind margin of female pronotum without prominent lateral teeth. In male Pseudagrion syriacum the upper appendages are longer than the lowers, instead of the reverse. Variation Immatures have a white abdomen bearing two black spots at the tip of each segment. Development of pruinosity progresses from the head to the thorax, legs and finally abdomen tip. Females are less pruinose. Behaviour Similar to other Platycnemis species, but comparatively poorly known. Males perch in the shade of stream banks.

Occurrence Range and status Endemic to the Near East, from south-eastern Turkey to Israel and northern Iraq and south-western Iran; status needs closer examination because of the restricted range and the vulnerability of aquatic habitats in this arid region. Range in our area is almost identical to that of P. dealbata, but seems less common. Habitat Mainly running waters with lush shady banks, such as streams, rivers, canals and ditches. Flight season Earlier and shorter than overlapping congeners: early April to late July in Turkey; most records in May.

Platycnemis Featherlegs

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Coenagrionidae

Ischnura Charpentier, 1840

Bluetails

Blue-tailed Damselflies, Forktails (NA)

Identification Diagnosis Delicate damselflies, the males (and females with ‘male colours’) are easily recognised by their ‘tail-light’, a striking blue marking at the end of the abdomen. This stands out because the abdomen is otherwise largely bronze-black, with the underside of roughly S3–6 yellow. Males have a bicoloured Pt in the Fw; the inner part is dark grey, the outer whitish. Females usually have a small vulvar spine on the underside of S8, at the base of the ovipositor. Separation from other genera The dark abdomen with blue ‘tail-light’ may recall dark-bodied Coenagrion or Erythromma species. However, both these have all-dark Pt, Coenagrion has S2 mainly blue, and dark Erythromma have red eyes. The vulvar spine is shared only with Enallagma females, but these differ in their abdominal pattern. Separation of the species Nine species of this cosmopolitan genus occur in the region, although three largely extralimital ones occur only on islands at its periphery. In most of our area only two species occur together: I. elegans (or one of its relatives) and I. pumilio. The elegans-group species, I. fountaineae

and I. senegalensis, overlap locally; up to three of them may occur together on the edge of the Sahara. These species are essentially similar in markings and must be separated by the male’s appendages and hind border of the pronotum. Only the latter structure may be used in females, but it is highly variable. Viewed from behind, the male’s upper appendages appear as an inverted ‘U’; the relative shape and position of its branches is species-specific. Identification is complicated by the existence of various female colour forms. Three basic types exist, which we name A, B and C here; they are defined in the table below. Coloration depends on form, but also varies with age. Females that ultimately develop bright male-like colours are called andromorphs, otherwise they are called gynomorphs. Behaviour Unlike other small damselflies, females oviposit alone, rather than in tandem with the male, following a prolonged copulation. After emergence, many adults remain close to the waterside. For these reasons, the proportions of mating pairs and tenerals at a site are often high. R Jödicke

103

Basic definition of female colour forms. * Applies to elegans-group females only; the ‘tail-light’ is absent in females of I. fountaineae, I. hastata and I. pumilio. Type

Andro-/ Gynomorph

Black humeral stripe when teneral

Thorax and abdomen base when teneral

Thorax and abdomen base when mature

‘Tail-light’ colour when mature*

A

Andromorph

Present

Lilac or orange

Bright green to blue

Blue

B

Gynomorph

Present

Lilac

Olive green or brown

Brown

C

Gynomorph

Absent

Pink or orange

Brown to green

Brown

Simple key to (groups of) species (males only). If statement disagrees, then go to next line. On Azores, beware of I. hastata females (not in table).

1

Pt in Fw larger than in Hw. Blue ‘tail-light’ covers S9 and part of S8, rather than being concentrated on S8

pumilio intermedia

2

Lower appendages about as long as uppers, not distinctly longer. N Sahara, SE Turkey

fountaineae

3

Black on S2 bulges downward at front, forming a saddle. Canary Islands and tropical greenhouses.

senegalensis

4

Black on S2 separated from coloured lower section by straight margin. Widespread, also Canary Islands.

elegans-group

Ischnura Bluetails

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Comparison of species of elegans-group:

104

Range

Overlap

Median lobe pronotum

Inner branches of upper appendages (from behind)

Tips of lower appendages (from above)

Species

Corsica, Sardinia, Sicily, etc.

Elba and Giglio

Weakly bilobed; low

Long and crossed

Parallel

genei

Long; upright

Diverging

elegans

Widely on the Iberian peninsula

Short and parallel, but crossed in SE

Weakly arched

Long and diverging

Converging

graellsii

Rarely in NW fringe of Sahara

Weakly arched

Long and crossed

Parallel

saharensis

Widespread

Iberian peninsula and NW Africa Sahara

Male Ischnura appendages viewed from behind.

I. elegans

I. genei

I. graellsii

I. saharensis

I. senegalensis

I. fountaineae

I. pumilio

I. intermedia

Male Ischnura pronotums.

I. elegans

I. senegalensis

I. genei

I. fountaineae

I. graellsii

I. pumilio

I. saharensis

I. intermedia

Ischnura Bluetails

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Ischnura elegans (Vander Linden, 1820) Common Bluetail Identification General One of the most familiar, widespread and abundant odonates in Europe, well known from eutrophic habitats and garden ponds. Replaced by very similar species in (parts of) the western Mediterranean and Africa. Field characters Tot 30–34mm, Ab 22–29mm, Hw 14–21mm. Similar size to Enallagma cyathigerum. Males are bronze-black, with head, thorax and abdomen base and tip marked sky blue at maturity; I. genei and I. saharensis remain relatively green on the head, thorax and abdomen base, but can be separated only in the hand, as can I. graellsii. Hand characters The tips of the male’s lower appendages diverge (view from above). The outer branches of the upper appendages (view from behind) are relatively thick and long; the

Blue-tailed Damselfly inner branches are parallel to each other (but see Variation). A narrow, upright projection on the hind border of the pronotum separates this species from all others, but is absent in some females and regionally in males. Variation Young males are green. Females appear in three colour forms: mature A-females are very like the male, but have a lilac thorax when immature. B-females are identical when immature, but become brownish or dull greenish at maturity. C-females have a pink thorax at first, but become rather like B-females later. Old B- and C-females may become very dark. The taxonomy of this and related species is complex. South-western relatives have been split off as distinct species, but variation towards the south-east is poorly understood, with several doubtful subspecies named. Males from southern Italy, Turkey, Crete, Cyprus and the Middle East usually have crossed inner branches of the upper appendages, but so do some individuals from central Europe and elsewhere. The upright projection on the pronotum is usually absent or reduced in males from the Hungarian Great Plain and northern Balkans to Central Asia.

105

Occurrence

G Pair of Ischnura elegans mating (andromorph female).

Range and status Extends from Ireland to Japan. Common in the region and typically the most abundant odonate at very eutrophic sites, but can be local and scarce in more nutrient-poor areas of northern and eastern Europe. Habitat Abundant at running and especially standing waters; tolerant of some salinity but avoids acidic habitats such as Sphagnum bogs. Flight season Late April to late September in central and northern Europe, normally with only one generation a year, but longer flight season with more generations in the south.

Ischnura Bluetails

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‘A-type’ mature /

teneral ?

immature ?

mature ?

106 bicoloured pterostigma on ? forewing

characteristic upright projection viewed from the side (note its absence in the photographs of similar species)

? pronotum

mature ? long median projection

/ pronotum

‘A-type’ immature /

? abdomen tip

S8 blue distinct black humeral stripe

‘B-type’ mature /

lower appendages diverge ‘C-type’ immature /

? appendages

no black humeral stripe

inner branches of upper appendages parallel

‘C-type’ mature /

rear view

Ischnura Bluetails

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Ischnura genei (Rambur, 1842)

? pronotum

Island Bluetail

small, weakly bilobed lobe

107 G Ischnura genei male. Note the green face and thorax, and absence of upright projection on pronotum. / pronotum

inner branches crossed

? appendages (rear view)

G Ischnura genei C-type immature female. Note black markings on S8.

Identification General Closely related to I. elegans, which it replaces on the Tyrrhenian Islands west of the Italian mainland, and very much like I. saharensis structurally. Occurs with I. elegans on two small islands close to the Italian mainland. Field characters Tot 29–32mm, Ab 18–26mm, Hw 12–18mm. Looks like I. elegans, but the colour of the face, thorax, pronotum and abdomen base of males and andromorph A-females is usually

green (although a turquoise tinge develops in full maturity), and females of all colour forms may have black markings on S8. Mature females without humeral black stripes may be confused with the coexisting I. pumilio, but the relative size of the Pt is discriminatory; that in the Fw is larger in I. pumilio. Hand characters The inner and outer branches of the male’s upper appendages are of fairly similar length and thickness; the tips of the former are crossed (view from behind). I. elegans has differently proportioned upper appendages. Hind border of pronotum with a central lobe, but without an upright projection. Variation Female colour forms are as in I. elegans, but form A does not become as blue and form C is rather salmon pink when teneral.

Occurrence Range and status Endemic to Corsica, Sardinia, Sicily, Malta, Capraia and Linosa; occurs with I. elegans on Elba and Giglio. Widespread and locally abundant within this restricted range. Habitat Inhabits standing and running waters with rich vegetation. Flight season Recorded from May to September.

Ischnura Bluetails

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Ischnura graellsii (Rambur, 1842)

Iberian Bluetail

108

G Mature male Ischnura graellsii. As seen here, the pale antehumeral stripes and postocular spots are often (but not always) reduced or lacking in this species.

hind border weakly arched

? pronotum

Identification General A member of the elegans-group, confined to the westernmost parts of Europe and North Africa. Overlaps and hybridises with I. elegans locally on the Iberian peninsula, especially in the north and west, while only I. elegans occurs in the Balearics. Meets I. saharensis very locally on the edge of the Sahara, but hybrids are unknown. Some mature females in particular may not be reliably identifiable in areas of overlap. Field characters Tot 26–31mm, Ab 20–25mm, Hw 13–19mm. Smaller on average than I. elegans. Unlike I. elegans, postocular spots and antehumeral stripes are often reduced or totally lacking in both sexes, while the majority of gynomorph females (B and C) have variable black markings on S8. On the Iberian peninsula, larger, well-marked (postoculars, antehumerals) males should be carefully checked as

inner branches diverge tips of lower appendages directed inwards

? appendages (rear view)

/ pronotum they may be I. elegans (but see Variation). Females without postocular spots may be confused with Erythromma viridulum, but have a vulvar spine. Hand characters Unlike I. elegans, the tips of the male’s lower appendages are directed towards each other (view from above). The inner branches of the upper appendages diverge (view from behind). Hind border of the pronotum is weakly arched, without an upright projection as in I. elegans. Variation The spring generation is generally larger-bodied and darker, with the antehumeral stripes and postocular spots reduced or absent. As in I. elegans, female forms A, B and C all occur, but the teneral thorax colours are distinctive: (greenish) yellow in A; whitish lilac in B; orange in C. The latter may recall I. pumilio; black on S2 is restricted to the distal quarter. Pink individuals, as often seen in I. elegans, and burgundy ones are rare.

Occurrence Range and status Widespread and abundant within its western Mediterranean range. Locally hybridises with I. elegans. Habitat All types of running and standing waters. Flight season Mid-April to mid-October in Catalonia; March to November in Andalucia and North Africa. Two generations a year in the north and up to four in the south.

Ischnura Bluetails

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Ischnura saharensis Aguesse, 1958

Sahara Bluetail

? appendages inner branches crossed

rear view

G Mature male Ischnura saharensis. Note green thorax and rather vivid orange of abdomen.

side view

? pronotum

Identification General This close relative of I. elegans is endemic to the western Sahara and the Canary Islands. It may be picked out among I. graellsii by the green thorax, and among I. fountaineae also by its smaller size, but can be distinguished with certainty only by the male’s appendages. Females may be inseparable in the field. Field characters Tot 26–31mm, Ab 19–25mm, Hw 12–17mm. One of our smallest Ischnura. Similar to I. elegans but in mature males the head, thorax and abdomen base are green. Blue individuals are likely to pertain to the blue colour form of the female or the locally coexisting I. graellsii or I. fountaineae. The underside of S3–7 in males is more vividly orange than in other species except I. fountaineae. Unlike I. graellsii and I. fountaineae, the upperside of S8 in females is never marked with black. In both sexes, the postocular spots tend to be small and the antehumeral stripes are narrow, but this is similar to I. graellsii and I. fountaineae. Hand characters As in I. genei, the inner branches of the male’s upper appendages are crossed; the proportion of the inner and outer branches resembles I. genei. The tips of the lower appendages are longer, more slender and less denticulate than in I. elegans. The posterior lobe of the pronotum is very similar to I. graellsii in both sexes. Variation The spring generation is particularly dark, with reduced postocular spots and antehumeral stripes. Female variation and development is not fully understood. Form C is most frequent, and has a pink thorax at first and

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/ pronotum

large postocular spots, becoming greenish and finally brown. Form B is similar when mature, but the immature colour is unknown. Andromorph A-females may have either a bright green or a bright blue thorax.

Occurrence Range and status Widespread and locally abundant along the northern fringe of the Sahara, ranging east to Libya and south to northern Niger. Together with I. senegalensis, the only damselfly breeding on the Canary Islands, where it is rare. Habitat Running and standing waters, preferably with dense vegetation; avoids higher salinity. Flight season Several generations throughout the year; scarce from December to February.

C

Ischnura Bluetails

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Ischnura fountaineae Morton, 1905

Oasis Bluetail

 Ischnura fountaineae pair copulating. Note blueness of male and absence of ‘tail-light’ in female.

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? pronotum

/ pronotum upper and lower appendages of similar length

rear view

? appendages

side view

Identification General A desert species that just reaches the region on the fringe of the Sahara, where it occurs with the smaller I. saharensis and (rarely) I. graellsii. In mixed populations, is picked out by its slightly larger size and the absence of a ‘tail-light’ in most females. Field characters Tot 27–34mm, Ab 21–25mm, Hw 19–24mm. One of the larger Ischnura species. In mature males, the thorax and abdomen base have a sky-blue ground colour, without a hint of green, and shining black markings; the underside of S3–6 is noticeably orange (only I. saharensis and I. senegalensis are equally bright). In both sexes, the postocular spots are small and the antehumeral stripes in males are narrow, but this is similar to I. graellsii and I. saharensis. The great majority of

females are gynomorph (form C) and lack humeral stripes and a ‘tail-light’, the upperside of their abdomen being black on S2–9. Their positive identification may be difficult under field conditions. The combination of their larger size, small postoculars and black on S2 and S8 tends to be diagnostic. Hand characters Males are reliably identified by the appendages, which are all of similar length. Variation Unlike other species, areas that become blue in males are whitish, not greenish, at immaturity. Postocular spots and antehumeral stripes are often reduced or absent. Immature females are orange with black markings. Their legs are unmarked at first, but develop brownish stripes. With maturity females turn olive, and later to brown. The rare andromorph females (A) are much like males; their teneral colours are unknown. B-females are unknown.

Occurrence Range and status Extends from Central Asia through the Middle East to Tunisia, Algeria, Morocco and Pantelleria (an island between Sicily and Tunisia). Abundant in suitable habitat. Occurs in eastern Turkey outside the region covered by this guide. Habitat Mainly in desert oases and brackish coastal sites. Springs and rivers with little vegetation, shallow lakes, glasswort (Salicornia) marsh; tolerant of high salinity. Flight season All year in the oases of Tunisia, with very low numbers in winter, otherwise March to November; several generations a year.

Ischnura Bluetails

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Ischnura senegalensis (Rambur, 1842)

? pronotum

Tropical Bluetail

 pronotum

inner branches not crossed

111 black patch on S2 bulges downwards

?appendages (rear view)

mature ?

immature 

mature 

?abdomen tip

upperside of S8 black

Identification General One of the most widespread damselflies in the Old World tropics, which just reaches the Canary Islands. Long mistaken for the co-occurring I. saharensis, its presence was confirmed there only in 2014. As the species is often imported with aquatic plants, can reproduce in greenhouses, and benefits from higher temperatures and human impacts, it is worth keeping an eye out more widely for the male’s subtle but distinctive abdominal markings. Field characters Tot 28–31mm, Ab 19–25mm, Hw 13–16mm. Recalls I. saharensis and other elegansgroup species in size and overall appearance, and best distinguished in the hand. However, in males: (1) the front of the glossy black patch on S2 clearly bulges downward, appearing like a black saddle, rather than being separated by a fairly straight margin from the green lower section; (2) the blue

on S8 appears square-cut at the front, not or barely extending onto S7, while in the other species it extends below the black upperside for about half the length of that segment; (3) the black line on the interpleural suture of the thorax tends to be shorter and thinner, at least in individuals in the Canary Islands. Only gynomorph females (form C) have been found there, which are orange when immature and black on the upperside of S8. In contrast, all forms of I. saharensis are blue or brown on S8, like I. elegans. Both sexes are similar to I. fountaineae, which could show up on the Canary Islands. The male is usually larger, with a blue or whitish (when immature) rather than green thorax. The somewhat similar saddle on S2 is less pronounced. Definitive identification requires examination of the very different appendages. Most males of I. genei are also green, but that species

Ischnura Bluetails

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is confined to Mediterranean islands. Hand characters The inner branches of the male’s upper appendages are in close contact but not crossed as in I. saharensis. Unlike the latter and other elegans-group species, the tips of the lower appendages are pointed inwards. The tubercle on the tip of S10 (above the appendages) is also smaller and thus rather inconspicuous. Variation The green markings on mature males infrequently develop a blue tinge. Andromorph females are known in other parts of the range of I. senegalensis and may have a black saddle on S2 as in the male, no saddle, or only part of it.

Occurrence Range and status Widespread and often common from Africa to Japan, extending through the Nile Valley to the Levant. Localised and rare on Tenerife and La Palma, but likely to be found elsewhere on the Canary Islands or in southern Morocco. This is the most frequent species to be imported to

G Ischnura senegalensis male. Europe with aquatic plants and can even maintain populations in greenhouses. Habitat A wide range of stagnant and slowflowing waters, often with dense vegetation. Tolerant to organic pollution and salinity, thus often more common at brackish, eutrophic, urban and agricultural sites than other species in its range. On the Canaries, found at man-made waterbodies such as water tanks and reservoirs. Flight season Recorded from late February to November on the Canary Islands, but as it can have multiple generations each year, may be on the wing in all months.

Ischnura pumilio (Charpentier, 1825) Identification General A diminutive but effective disperser, which swiftly colonises pioneer habitats. Young females are remarkably orange; males have the typical Ischnura ‘tail-light’, but it is shifted to S9. Field characters Tot 26–31mm, Ab 22–25mm, Hw 14–18mm. One of the smallest European species. Easily separated from I. elegans and its relatives by: (1) Pt in Fw distinctly bigger than in Hw, especially in males; (2) in males, the blue ‘tail-light’ is on S9 and the adjacent half of S8, rather than being concentrated on S8; (3) females always lack the ‘tail-light’, and when immature are bright orange with limited black on the head, thorax and abdomen (e.g. S1–2 and base S3 unmarked), and when mature are very different and inconspicuous (see Variation). Immature I. graellsii and I. fountaineae females can also be orange but have more black on the head (enclosing postocular spots) and on the abdomen base (at least upperside S2 partly black). Mature females

Small Bluetail Scarce Blue-tailed Damselfly

are distinguished by the bigger Pt of the Fw, and the completely black upperside of the abdomen and pronotum (see below). Because of their size, greenish females may be mistaken for Nehalennia speciosa, which is metallic and much more delicate. Hand characters The male’s upper appendages lack the inner branches of I. elegans and its relatives (view from behind). The hind border of the pronotum forms a simple arch, without any projecting or upright stuctures. Variation The male’s colour changes from whitish, through yellowish and green, to blue. The ‘taillight’ is variably (sometimes extensively) covered with black markings. Distal part of Pt frequently becomes blue in mature males. All females typically lack the black humeral stripes, having at most a thin black line along the suture, and are orange when teneral. They have big postocular patches, which narrow to round spots during maturation, while the upperside of the abdomen becomes completely black. They fit the definition of the

Ischnura Bluetails

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immature /

mature /

? pronotum

orange extensive on head, thorax and abdomen base

mature ?

hind border weakly arched

pterostigma distinctly larger in forewing than in hindwing

bicoloured pterostigma on ? forewing; distal part often bluish

S9 and apical part of S8 blue

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female never with ‘tail-light’

? abdomen tip

inner branch absent

developing black humeral stripe is initially absent

? appendages (rear view)

/ pronotum

transitional phase /

F Mature male Ischnura pumilio. Note larger bicoloured pterostigma on forewings.

F Immature female Ischnura pumilio. This bright orange developmental phase is sometimes referred to as the form ‘aurantiaca’.

Ischnura Bluetails

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gynomorph form C. However, occasional females do not become green when mature, but blue, and could effectively be considered andromorph. Rare, largely blackish females and green females without solid black markings are also known.

A M

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Behaviour In rare cases, tandem oviposition has been reported.

Occurrence Range and status This effective coloniser ranges from the Azores and Madeira to north-east China, and is locally abundant. Populations are often short-lived, disappearing with vegetation succession. Habitat Recently created habitats with sparse vegetation (for instance, in quarries) can harbour huge populations, but the species can also be found on a wide variety of sites, such as small or temporary ponds (also nutrient-poor and acidic ones), springs and ditches. Flight season March to October in the south, with at least two generations a year; from the end of May until mid-September in the north, where a second generation seems to be becoming more regular.

Ischnura intermedia Dumont, 1974 Identification General An inconspicuous Ischnura from southwest Asia, which just reaches Cyprus and southeast Turkey. Unlike its congeners, the male’s blue ‘tail-light’ typically covers the two penultimate abdominal segments entirely. This scarce and localised species is probably overlooked due to its resemblance to I. pumilio, and was first discovered on Cyprus only in 2013. Field characters Tot 26–29mm, Ab 19–21mm, Hw 11–13mm. Tiny, even smaller on average than I. pumilio. Males are readily separated from that and other Ischnura species by the blue ‘taillight’ extending over the full length of both S8 and S9, but see Variation below. Both sexes are distinguished from other species except I. pumilio by the notably small Pt in the Hw; in males, those in the Fw are even greater than twice the size and have a larger dark portion than in I. pumilio. Females are similar to I. pumilio (see Variation) and best identified in the hand. Hand characters The shape and relative length of the male appendages are unique among our Ischnura species. Seen from the side, the upper appendages end in a small hook and are slightly (but distinctly) longer than the lower appendages. From above, the uppers appear like diverging

Persian Bluetail

pliers and (almost) completely eclipse the lowers. As in I. pumilio, the upper appendages lack the inner branch found in most other Ischnura species. Viewed from above, the hind border of the female pronotum is slightly notched, not arched as in I. pumilio. The ovipositor is shorter and more pointed than in I. pumilio, projecting obliquely from the abdomen rather than being pressed against and parallel to it. Variation The male’s blue ‘tail-light’ can variably be covered with black markings, as in I. pumilio. Their extent varies seasonally, with more black in spring. Such early males may recall I. pumilio, but at least three-quarters of S8 remains blue while at most half of the segment is blue in the bluest I. pumilio males. Mature males can have a greenish or blueish thorax. Observed females are gynomorph (form C) and probably display the same variability as I. pumilio, the bright orange immatures with big postocular patches and becoming dark olive at maturity. Blue andromorph females have not been recorded but seem likely to occur.

Occurrence Range and status Seems scarce and localised, but also easily overlooked due to its small size and

Ischnura Bluetails

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? pronotum

mature ?

immature 

mature 

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?appendages (rear view)

females lack ‘tail-light’

S9 and at least 3 ⁄4 of S8 blue

?abdomen tip

 pronotum

upper appendages longer than the lower

hind border slightly notched

immature 

unobtrusive behaviour. Since its discovery along the Turkish Euphrates (just outside the area covered by this book), only two dozen sites scattered across northern Syria, Iraq, Iran and southern Turkmenistan have become known. In our area it is known only from single records near Birecik and Adana in Turkey, and five valleys in south-western Cyprus, where it is moderately common. Habitat Small, shallow channels of springs, streams and small rivers, bordered by dense emergent vegetation and with little flow. Occurs with I. elegans and I. pumilio, although no extant populations of the latter are known from Cyprus. Flight season On Cyprus from late March to late November: numbers peak from the end of April to the middle of June, then drop, and rise again in August, suggesting there are at least two generations a year.

Ischnura Bluetails

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Ischnura hastata (Say, 1839)

Citrine Forktail

S1–4 largely unmarked

immature /

pterostigma on ? forewing

mature ?

/ pronotum this sex has not yet been found in our region

116 reduced hind border

? S8-10 side view

Identification General Unique in many ways, this is the only American species reproducing in our area. The only population here, in the Azores, was misidentified until 1990, although records suggest it may have been present at least a century before. It is the only parthenogenic Odonata population known in the world: all generations lack males, unfertilised eggs producing females only. Surveys in the archipelago and experiments aimed at interrupting parthenogenesis have failed to produce males. These males are the only odonates in the world with the Pt inside the wing, rather than on its leading edge. Field characters Tot 20–27mm, Ab 17–21mm, Hw 11–15mm. Distinctly smaller than I. pumilio. The male, unknown from Europe, is unique by its largely yellow abdomen. The deep orange Pt of the Fw is fully enclosed by the wing (females have normal Pt). Fresh females have the basal half of the abdomen orange, S1–4 being (largely) unmarked; in I. pumilio only the very base (S1–2) is unmarked. Mature I. hastata females turn pale brown to dull olive, with a wholly black abdominal dorsum. Eventually, they become largely covered with greyish pruinosity (see photograph). I. pumilio turns green or occasionally blue, but is never pruinose. Olive females of both may be easily confused, but can be separated by the pronotal hindlobe.

pterostigma on ? hindwing

Hand characters The male has a unique rod-like projection on S10. The female’s pronotal hindlobe is reduced in comparison to I. pumilio. Behaviour The Azorean females lay eggs without needing to mate, producing a new generation of females. In other invertebrates a bacterial infection is usually responsible for parthenogenesis, but experiments with antibiotics have not yet proved this for I. hastata. Males of I. pumilio (the only other damselfly on the Azores) have been seen holding I. hastata females in tandem.

Occurrence Range and status Inhabits all islands of the Azores, where sometimes hundreds of females have been seen at a pond, making it the commonest species there (always outnumbering I. pumilio). Widespread in North and South America (also the Galápagos Islands), where parthenogenesis is unknown. A strong disperser, which could appear on eastern Atlantic shores. Habitat Almost every well-vegetated Azorean pond, typically with pondweeds (Potamogeton) and spike-rushes (Eleocharis). Flight season Recorded from the Azores from late May to early August. Almost year-round in the southern United States; April to November further north.

F Ischnura hastata female in the Azores.

Ischnura Bluetails

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Enallagma 

‘American’ Bluets

Charpentier, 1840 Identification Diagnosis Typical bluets, the males being blue, distinctively marked with black (see E. cyathigerum). Females are usually either brown or blue, with torpedo-like markings on S3–7, and have a small but prominent vulvar spine on the underside of S8, at the base of the ovipositor. Separation from other genera The blue-andblack pattern of males recalls most Coenagrion species and Erythromma lindenii (see E. cyathigerum for a detailed comparison). Only Ischnura females share the vulvar spine, although it is weaker. Brown females may be confused with Sympecma, which has similar markings on S3–6, but differs in its long Pt and details of markings and venation.

Separation of the species The genus is almost entirely confined to Eurasia and North America, the vast majority of the almost 50 species restricted to the latter. Only two very similar species occur in our region, one being among our most widespread species, the other restricted to north-west Africa. Their separation requires close examination of the male appendages (see E. deserti). Behaviour Strongly recalls Coenagrion species, but unlike them males prefer open water, where they fly and perch on floating plants close to the water’s surface. R Jödicke

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 Enallagma cyathigerum male. Note the broad antehumeral stripes and ‘plain’ side of the thorax.

Enallagma cyathigerum (Charpentier, 1840) Common Bluet Identification General Probably the most widespread species in the guide. One of many bluets whose identity is best confirmed in the hand. Easily identified by the black markings, e.g. the ‘mushroom’ on the male’s S2, the ‘torpedoes’ on the female’s S3–7 and the ‘single stripe’ on the thorax sides of both. Its behaviour and habitat also differ from other bluets, for example, myriads of bluets skimming over an open lake will rarely be another species. Field characters Tot 29–36mm, Ab 22–28mm, Hw 15–21mm. Size and shape close to other bluets, but appears slightly more robust and purer blue. Combines the following markings: (1) broad

Common Blue Damselfly antehumeral stripes and large postocular spots in both sexes, the antehumeral normally at least as wide as the black humeral stripe beneath it; (2) the sides of the thorax appear plain because there is not usually a short black interpleural stripe between the humeral stripe and the small black stripe on the lower suture; (3) the male has a small mushroom-like marking on S2, and S8–9 are completely blue, creating a conspicuous ‘tail-light’; (4) the female has black torpedo-like markings on the abdominal dorsum. Erythromma lindenii males share the skimming habit and broad antehumerals of Enallagma cyathigerum, but the postocular spots are reduced, there are two short black stripes

Enallagma ‘American’ Bluets

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Coenagrion species, where there are often prominently projecting lobes, especially in females. Unlike Coenagrion, at least one cell below the wing’s leading edge between the Pt and wing tip in the Hw is usually doubled. Variation The male’s S2 ‘mushroom’ may be round or pointed, its stem may be missing or lateral winglike extensions can be present. Very dark males occur occasionally, especially in the north and east, defying the familiar diagnosis. They may have extensive black on abdomen, narrowed or even broken antehumerals, and well-developed black interpleural and metapleural stripes, recalling Coenagrion hylas and C. johanssoni. Andromorph females are blue and not uncommon, but are often outnumbered by greenish and brownish gynomorphs.

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on the thorax sides, the entire upperside of S2 is black and the blue is shifted to S9–10 (S8 is black). Coenagrion males are hardly distinguishable at a distance, but prefer to remain among bankside vegetation. They have antehumerals narrower than the humeral, two short thorax stripes, and a different marking on S2 (but see Variation). The aforementioned features of the head and thorax also apply to Coenagrion and Erythromma females. Hand characters A quick look for the female’s vulvar spine with a hand lens rules out confusion with all Coenagrion, Erythromma and Sympecma species. In both sexes, the hind margin of the pronotum is only weakly curved, unlike most

Occurrence Range and status Common in most parts of the region, its range extending north of the Arctic Circle and to the Moroccan Atlas Mountains and Kamchatka. North American populations have been found to pertain to a separate species. Habitat All types of ponds, lakes and also slow rivers. Huge densities occur in acidic waters without fish. Flight season Late April to October. Larval development needs more than one year in the north, while in the south up to two generations emerge each year.

Enallagma deserti (Selys, 1871) Identification General A typical bluet, patrolling and perching close to the water’s surface. General appearance

Desert Bluet

and ecology are very similar to E. cyathigerum, of which some workers consider it a subspecies, but the two overlap in northern Morocco. Identification is possible only under magnification. Field characters Tot 32–37mm, Ab 24–29mm, Hw 19–23mm. Slightly larger than E. cyathigerum, but otherwise seems identical. See that species and genus text for separation from other bluets. Hand characters The tips of the male’s upper appendages have a distinctive shape, which can be recognised through a hand lens with some experience. The protruding tip of E. cyathigerum is much reduced in E. deserti (view from side), and the tooth lying above it appears like an inwardpointing finger in E. cyathigerum, but is much broader and shorter in E. deserti (view from above and behind). Males with intermediate features have been found. Slight structural differences of the female mesostigmal plate (the part of the thorax

Enallagma ‘American’ Bluets

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immature / (dull form)

mature / (dull form)

mature / (blue form)

immature ?

mature ?

? S1–2 showing ‘mushroom’ mark

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blue antehumeral stripe at least as broad as black humeral stripe below it

/ abdomen tip

mature ? ‘plain’ side of thorax, with black interpleural stripe not or hardly developed

prominent spine below S8

/ pronotum

mature / (blue form)

? appendages

upper appendage (rear view) side view

directly behind the pronotum) are too subtle and microscopic for reliable identification. Variation Gynomorph females are prevalent and are all greenish.

Occurrence Range and status Endemic to the Maghreb. Generally rather scarce, but may occur in huge populations locally; the species has benefited from the increase in reservoirs. Habitat Calm river sections or lakes, dams and ponds, with floating leaves of submerged plants. Flight season Mid-April to late September; probably with several generations a year.

Enallagma deserti / pronotum pronotum does not differ from E. cyathigerum; markings vary according to paleness of individual

? appendages

upper appendage (rear view) side view

Enallagma ‘American’ Bluets

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Coenagrion 

‘Eurasian’ Bluets

Kirby, 1890 Identification

120

Diagnosis Males are small blue damselflies marked with black. The blue areas may be (partly) greenish or whitish. Females are blacker, and more often green or brownish. Because this is a large and diverse group, the genus is best identified by the exclusion of the smaller genera (see below). A damselfly with plain Pt, roundish postocular spots, the coloured antehumeral stripes narrower than the black lines below (humeral stripes), and two short but well-defined black lines on the thorax sides (interpleural and metapleural stripes) will in most cases belong to Coenagrion. Separation from other genera Enallagma normally have the antehumeral stripes wider than the black humeral stripes below them, and they lack the black interpleural stripes on the thorax sides below that. Ischnura males have two-toned Pt. Enallagma and Ischnura females possess a vulvar spine. Erythromma species are darker, with reduced or no postocular spots, and males have red or all-blue eyes, whereas these are blue with dark caps in Coenagrion. The blue Erythromma lindenii is superficially very similar, but differs in numerous details of markings, including wider antehumerals,

an all-black upperside to S2 and a shorter ‘tail-light’ shifted to the extreme tip of the abdomen. Separation of the species The richest genus of damselflies in our area, with 13 species. Additional species occur east to Japan, with a few in North America. Males can normally be separated by their markings, aided by their appendages. The pattern on the male’s S2 is a popular identification feature, but as all markings are highly variable this may serve only as an indication, and it may be wise to check the appendages and pronotum in individual cases. A simple key to males is provided below. Females may be difficult to identify, as markings differ less between species and vary (even) more than in males. Study of the hind border of their pronotum is required. The species typically occur in fairly high densities and do not wander far from their habitat, so a conspecific male is usually close at hand. Behaviour Males typically fly low among or along riparian vegetation, seldom venturing far from it. Tandems often oviposit in concentrations on floating vegetation, the male adopting an upright posture. K-D B Dijkstra

Simple key to groups of similar species (males only). If the statement agrees, compare the given species. If it disagrees, then go to the next line. See also photographs opposite.

1

Abdomen is all black, except for bluish areas at base and near tip. Lower appendages are massive, dwarfing uppers.

armatum

2

Abdomen sides black along lower border. Black markings are present at base of hind legs. Exclusively boreal or alpine species.

johanssoni hylas

3

Black on middle segments of abdomen is drawn out into two long spikes, one on each side of segment.

puella pulchellum syriacum intermedium

4

Undersides of eyes, face and often thorax are green, not blue. Black marking on S1 always restricted to base, not covering segment’s full length. Black marking on S2 usually broken up into three marks, not a single one. Upper appendages never hooked at tip. Northern rather than southern species.

hastulatum lunulatum

5

S6 is largely black; terminal half of abdomen therefore appears black, apart from blue ‘tail-light’. Pt are elongate and pale.

scitulum caerulescens

6

At least basal third of S6 is blue; whole abdomen appears to have alternating pattern of black and blue. Pt are lozenge-shaped and dark.

mercuriale ornatum

Coenagrion ‘Eurasian’ Bluets

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 Coenagrion armatum male.

H Below left C. caerulescens male. H Below right C. puella male.

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 C. hastulatum male.

 C. lunulatum male

 C. hylas male.

Coenagrion ‘Eurasian’ Bluets

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Coenagrion pulchellum  (Vander Linden, 1825)

Variable Bluet Variable Damselfly

antehumeral stripe broken

? pronotum (similar shape in /)

122 mature ?

/ andromorph form

/ gynomorph form

? S1–2 showing ‘Y’ mark

uppers (nearly) touching at base

? abdomen tip

? appendages (side view)

Identification General A slender blue damselfly that is often confused with the paler C. puella. Males typically have blue exclamation marks on the ‘shoulders’ and a black ‘Y’ at the abdomen base, rather than blue lines and a black ‘U’. Although the two species widely overlap in range, they often occupy different habitats. The vernacular name refers to the variability of this species, particularly

in females, although through a hand lens the distinctive W-bordered pronotum of the female is characteristic. Field characters Tot 34–38mm, Ab 23–32mm, Hw 16–23mm. Similar build to C. puella, with which it will most often be confused. Good pointers to recognise males in the field are: (1) darker, with a deeper blue colour and more black markings; (2) antehumeral stripes usually broken, appearing like exclamation marks; (3) black figure on S2 connected by a stalk to the segment border, usually appearing like a ‘Y’ rather than a ‘U’; (4) S3–6 substantially blacker, e.g. S4–5 at least half black (not less). Females normally have complete antehumeral stripes and are more variable than those of any other Coenagrion (see below). Hand characters Male appendages somewhat like those of C. puella, but uppers as long as lowers (view from side) and (nearly) touching each other at their base (view from above). Hind margin of female pronotum deeply incised at two points, resulting in three distinct lobes. Variation Eastern and southern males (e.g in the Mediterranean and Turkey) can be exceptionally dark. Males with more blue that recall C. puella

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may be found among typical ones, showing a black ‘U’ on S2, but S6 is typically more than half black and the antehumeral stripes are usually broken. Andromorph females with broad blue bases to the abdominal segments are common and not unlike males (i.e. with diagnostic long lateral spikes), but with more black on S2 and S8–9. Green gynomorph females with an all-black abdominal dorsum, not dissimilar to the common type of C. puella female, are also frequent. Other varieties are scarce or local, such as a black-backed variety with S1–2 and S8 noticeably pale, reminiscent of C. armatum. Like the latter, C. pulchellum is the only species in which the black marking on S2 is

frequently free from the basal segment border in females.

Occurrence Range and status Widespread and numerous locally, but overall much scarcer than C. puella, especially in south. Ranges to the West Siberian Plain and Central Asia. Habitat Wide range of still, occasionally slow-flowing, waters. These are typically lushly vegetated, non-acidic and meso- to eutrophic, such as oxbows and fenlands. Flight season From April to mid-September; most abundant in late May and June.

Coenagrion puella (Linnaeus, 1758)

123

Azure Bluet Azure Damselfly

? pronotum

immature ?

? abdomen tip

mature ?

/ pronotum

upper appendages widely separated

? S1–2 showing ‘U’ mark blue antehumeral stripe complete

? appendages

/ S1–2

mature ?

/ gynomorph / andromorph form

(side view)

form

black spikes extend forwards

short black interpleural stripe present in all Coenagrion but very rarely in Enallagma

Identification General This sleek, pale blue damselfly is one of the commonest odonates in large parts of Europe. Males are usually recognised by the black ‘U’ on S2 and two-spiked markings on the abdomen, but

mature /

may be confused with the equally widespread but often darker and more local C. pulchellum. Two lookalikes replace the species in Crete and southeast Turkey. Field characters Tot 33–35mm, Ab 22–31mm,

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124

Hw 15–24mm. Longer and thinner than most similar species, such as Enallagma cyathigerum. The typical blue damselfly in many areas. The male is light blue, typically with narrow but complete antehumeral stripes, black ‘U’ on S2 and apical fifths of S3–5 with black rings. A thin black spike extends from these rings on each side (absent on S3 in pale specimens), almost reaching segment bases. These spikes are shared with the near-identical C. syriacum of southern Turkey and C. intermedium of Crete, and the darker but widespread C. pulchellum (see that species). Gynomorph females usually predominate; these are yellowish green with the entire abdominal upperside black and are best identified by the pronotal hind border. Hand characters Male upper appendages are distinctly shorter than lowers, widely separated from one another and with inward-directed hooks at tips (view from above). The wide gap between the upper appendages is unique among related species. Hind margin of female pronotum is wavy, forming three very shallow lobes. Variation Males vary greatly in the extent of the black markings, and care is required to use these characters. In north-west Africa, males have two black spots on S8, which are absent or smaller elsewhere. These specimens are sometimes considered as the subspecies kocheri, but similar males from Spain suggest the variation is gradual. An andromorph variety of the female has the basal fifths of S3–6 blue, the black markings on these segments being three-toothed. They are

C. intermedium (VU)

reminiscent of typical andromorph C. pulchellum and C. ornatum females, but these normally have more blue on S7–9.

Occurrence Range and status The most abundant species of the genus; common in a very large part of our area, with its range extending to the West Siberian Plain and eastern Kazakhstan. Habitat A wide variety of waters, especially still but also flowing, although generally scarce on peaty or clay soils. Favours the presence of aquatic vegetation. Flight season From April (even March in Algeria) to September, but most common from mid-May to the end of July.

Coenagrion intermedium Lohmann, 1990

Cretan Bluet

Identification General Endemic bluet of Crete, where it replaces C. puella. It differs mainly by the shape of the male appendages and can therefore most easily be recognised by its range. Field characters Tot 35–36mm, Ab 26–32mm, Hw 19–22mm. Larger and slightly darker on average than C. puella from central and northern Europe, but otherwise appears identical to it and C. syriacum; C. puella from the adjacent mainland are similarly dark. Hand characters Male appendages are similar to those of C. puella, but the upper appendages more or less touch each other near their base (view from above), as in C. syriacum and C. pulchellum. Hind margin of female pronotum intermediate between those of C. puella and C. pulchellum.

? pronotum

/ pronotum

? appendages

? S1–2

(side view)

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Variation An andromorph form of the female (see C. puella) may occur but is unrecorded to date.

Occurrence Range and status Confined to Crete (shown in blue on map p. 124), where it is localised because

of habitat; classified as Vulnerable on the IUCN Red List. Habitat Unlike its mainland counterparts, appears to be confined to running water, inhabiting permanent streams. Flight season From mid-May to early August.

125

G Coenagrion intermedium male.

Coenagrion syriacum (Morton, 1924)

Syrian Bluet

Identification General Replaces the deceptively similar C. puella in the Near East, the two species meeting in southern Turkey. Can be separated reliably only by the male appendages. Field characters Tot 30–37mm, Ab 24–29mm, Hw 16–20mm. Male appears identical to C. puella. It is blacker than typical males of that species from central and northern Europe (e.g. S6 is largely black instead of about half) and may recall C. pulchellum, but Turkish C. puella are similarly dark. Pale females are bluer, more like those of C. pulchellum. Hand characters Male upper appendages more or less touch each other near their base (view from above). Lower appendages are longer and more horizontal than in related species, about twice as long as the upper appendages and pointing straight backwards, not half upwards. Hind margin of female pronotum more strongly three-lobed than in C. puella (recalling C. pulchellum), with rather narrow central lobe. Variation Has the same female colour forms as C. puella.

? pronotum

? appendages

/ pronotum

? abdomen tip

? S1–2

(side view)

NT

Occurrence Range and status Replaces C. puella in a band along the Mediterranean Sea from Antalya to northern Israel. Habitat Still and slow-flowing waters, with ample vegetation such as pondweeds (Potamogeton). Flight season Turkish records are from mid-April to the end of May.

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Coenagrion mercuriale (Charpentier, 1840) Mercury Bluet

Southern Damselfly

/ pronotum ?

lozenge-shaped pterostigma with dark centre

/

gynomorph form (side view)

126 internal tooth near base

? appendages

? S1–2 showing ‘Mercury’ mark NT HD II+IV

Identification General Small, deep-blue damselfly, typical of small streams in south-western Europe. Male’s S2 is marked with a black double-horned triangle, recalling the ancient alchemist symbol for the element mercury, to which the species owes its name. Details of markings and appendages further identify it. Field characters Tot 27–31mm, Ab 19–27mm, Hw 12–21mm. Stouter than C. puella and among the smallest members of the genus. Male S2 marking is variable, not unlike that of other Coenagrion species, but typically resembles a head bearing a horned helmet. S3–6 seldom more than half black, S7 always with some blue at the base. Males are perhaps most easily identified by the exclusion of congeners likely in the same habitat: C. puella bears long lateral black spikes on the black end rings of S3–6; C. ornatum has more lanceolate markings on S3–4 and frayed postocular spots; C. caerulescens and C. scitulum have elongated Pt and largely black S6–7. Typical females are gynomorph, marked yellowish green but rather dark and nondescript, again best recognised by the exclusion of other species. Hand characters Male upper appendages are strongly hooked at the tip, as in C. scitulum and C. caerulescens, but are about as long as the lowers and also with distinct internal teeth near their base. Hind margin of female pronotum with tiny lobe in the middle, much smaller than that of C. scitulum. Variation Andromorph females with the pale areas on the abdomen larger and bluer are rather rare. Females with distinctly pinkish thorax and

abdomen might already be reproductively active but probably darken with age.

Occurrence Range and status Locally common in France and the north of the Iberian peninsula; populations are very scattered in the north of range. Not known to occur east of Italy, Austria and Germany; numerous old records from eastern Europe are presumed to be erroneous. Habitat Sunny streams and springs with rich aquatic and riparian vegetation, such as submerged banks of water-starworts (Callitriche) or stands of Lesser Water-parsnip (Berula erecta). These are often calcareous, but can occur on heathy areas flushed with calcareous water in England. Flowing drainage ditches may also offer suitable habitat. Flight season Mainly June and July in north of range (from mid-May to mid-August), but longer further south, e.g. from early March to late September in Morocco, suggesting two generations a year.

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Coenagrion ornatum (Selys, 1850)

Ornate Bluet

lozenge-shaped dark pterostigma

/ pronotum / andromorph form

? lanceolate markings on S3–9

127

much blue at segment bases

? appendages (side view)

? S1–2 showing

toothed rear margin to postocular spots

‘winged’ mark

Identification General The eastern ecological counterpart of C. mercuriale, inhabiting similar streams. The two species overlap widely in central Europe, where both species are very local, but often occur together in suitable streams. The black marking on the male’s S2 has been likened to a bird, bat, butterfly or other winged creature, but the ragged border of the postocular spots is perhaps the best distinguishing feature. Details of markings and appendages confirm identification. Field characters Tot 30–31mm, Ab 20–30mm, Hw 17–24mm. Smaller and stouter than C. puella. Both sexes typically have the hind border of the postocular spots toothed. Male S2 is usually marked with a trident or stalked ‘W’. Male has a turnip-shaped black marking at the ends of S3–4, characteristically drawn out into very long spikes in the centre. Andromorph females predominate, with more blue at segment bases than most species, reminiscent of andromorph females of C. pulchellum. In the latter, the black markings basally have longer lateral spikes flanking the central spike, while in C. ornatum the central spike is longest, especially on S3–4. Hand characters Male upper appendages shorter than lowers (view from side) and without strongly hooked tip, therefore more like C. pulchellum than other southern Coenagrion. Hind margin of female pronotum with slight lobe projecting from the

middle, weakly notched in its centre. Variation Gynomorph females without bluebased segments exist, but should be identifiable by their postocular spots and pronotum. As in C. mercuriale, females are occasionally pinkish on the thorax and abdomen.

Occurrence Range and status Very local in the north and west of range, to fairly common in the south-east. Habitat Similar to C. mercuriale, being found by sunny streams and flowing ditches, often calcareous and with structured vegetation. Flight season From early May to mid-August.

HD II

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Coenagrion scitulum (Rambur, 1842)

Dainty Bluet Dainty Damselfly

long pale pterostigma

/ pronotum

?

/

128

much blue on abdomen, but may be blacker

? S1–2 showing ‘cat’s head’ mark

strongly incurved hooks

less than half the length of S10 side view

? appendages

/ andromorph form

Identification General Another small bluet of the south, preferring calmer waters than its relatives, these often with rich aquatic vegetation. The black-andblue pattern of the males appears as if the basal half of the abdomen is of a Coenagrion and the terminal half of the abdomen is of an Ischnura. Care is required to differentiate from C. caerulescens along running waters in the western Mediterranean. Field characters Tot 30–33mm, Ab 20–27mm, Hw 14–20mm. Shorter-bodied than C. puella and one of the smallest species of Coenagrion. Males differ from their usual neighbours, C. mercuriale, C. ornatum and C. puella, by having S3–5 about half black, while S6–7 are all black. Male S2 is marked with a black ‘tuning fork’ or ‘cat’s head’, but this pattern is highly variable. Males of C. caerulescens appear almost identical (see that species and Hand characters for distinguishing features). Gynomorph females are greenish or brown, but are usually outnumbered by andromorphs, in which the blue areas creep up rather broadly at the bases of the abdomen segments and the black markings appear torpedo-shaped. Best identified by the long, pale Pt (see below). Hand characters Shares the following diagnostic features with C. caerulescens: (1) Pt in both sexes more elongate and often paler (brownish, not black) than other Coenagrion, the anterior side being clearly longer than the basal side, while in other Coenagrion all sides are about equal; (2) male upper appendages distinctly longer than lowers (view from side), with a distinct tooth at the tip

of each but not at base (view from above). Unlike C. caerulescens, male upper appendages have strongly incurved hooks (view from above) and are less than half as long as S10. The hind margin of the female pronotum has a small lobe in the middle, while the male pronotal margin is broadly triangular and pointed in the middle. Variation Females are variable and can be darker than described.

Occurrence Range and status Widespread in the Mediterranean and fairly common in France and parts of the Iberian peninsula, becoming more localised towards the Caspian Sea and rare in Central Asia. Has expanded considerably northwards in Europe since the 1990s. Habitat Sunny, still, sometimes slow-flowing, waters with rich aquatic vegetation, such as water-milfoils (Myriophyllum) and hornworts (Ceratophyllum). Flight season From April to September; most abundant from mid-May to the end of July.

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Coenagrion caerulescens 

Mediterranean Bluet

(Fonscolombe, 1838) ? pterostigma appears triangular

/ pronotum over half the length of S10

/ andromorph form

?

129 side view

straightish hooks

? appendages

? S1–2 blue tip

Identification General Doppelganger of C. scitulum, occurring in the south-west of the latter’s range and favouring flowing rather than still waters. It is relatively more common than its sibling on the Iberian peninsula and northern Africa. The male appendages and female pronotum are the most reliable means of separating the two. Field characters Tot 30–33mm, Ab 18–27mm, Hw 14–21mm. Similar in size and shape to C. scitulum. The markings differ subtly from this species, although there is overlap and these characters cannot entirely be relied upon: (1) both sexes tend to have large, pale paired spots on the pronotum (not small or absent); (2) males usually have more than half (not less) of S3–4 black, often with relatively more on S3 than S4 (not the reverse); (3) male S9 usually has some black (seldom unmarked blue). Females are not unlike C. scitulum (also with long Pt), but are generally very pale with broad antehumeral stripes (usually wider than the black humeral stripes), larger postocular spots and narrower black markings on abdominal upperside than congeners. In most of its range (but not, or less so, in Italy and the Tyrrhenian Islands), they have the abdomen tip (S9–10 and appendages) largely blue, not black on top as in overlapping Coenagrion species, including C. scitulum. Hand characters See C. scitulum for shared features that differ from other Coenagrion. Outer corner of the Pt is more acute than in C. scitulum (i.e. Pt appears more triangular than quadrangular).

Male upper appendages are longer and straighter than in C. scitulum, only weakly hooked at tips and over half as long as S10. Female is recognisable by the pale projecting ‘V’ in the hind margin of the pronotum. Male pronotal margin has a small knob in the middle. Variation The extent of the markings varies strongly in both sexes but appears to be no basis for the distinction of subspecies, such as the dark Italian ssp. caesarum.

Occurrence Range and status Endemic to the western Mediterranean. Local and seldom numerous in Europe, but commoner towards the south and the most common Coenagrion in the Maghreb. Habitat Sunny seepages, streams and rivers with ample vegetation. Flight season From mid-April to mid-October in parts of north-west Africa (possible proof of two generations a year), but elsewhere mainly from May to August.

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Coenagrion hastulatum  (Charpentier, 1825)

130

Spearhead Bluet Northern Damselfly

usually more green visible at base than in C. puella

mature ?

? S1–2 showing ‘spearhead’ mark

/ pronotum

/ gynomorph form

? appendages (side view)

/ abdomen tip /

Identification General The commonest blue damselfly in much of boreal Europe, but very local relict populations further south. Males are recognised by their greenish hue and the black spearhead markings on the basal abdominal segments. Field characters Tot 31–33mm, Ab 22–26mm, Hw 16–22mm. Shorter-bodied than C. puella. Male is pale blue, with a turquoise tinge, and can be quite green on the underside (as in C. lunulatum), especially of the eyes. Marking on S2 is variable, usually consisting of a black spearhead- or mushroom-shaped mark and two black dashes. Usually less than half of S3–4 is black, the marking on S3 being characteristically spearhead-shaped. Typical female is a green gynomorph, with the entire abdominal dorsum black, making it similar to C. puella, but in C. hastulatum the black gradually narrows towards the base: from above, more lateral green is visible on S3–4 than on S6–9. See C. lunulatum for a comparison with this species. Hand characters Male upper appendages shorter than lowers. Hind margin of female pronotum with two straight oblique borders that meet in a small point, like a shallow ‘V’. Variation Marking on male’s S2 may be reduced, without a stem, recalling that of C. lunulatum.

Occurrence Range and status Abundant in many parts of northern Eurasia east to Kamchatka, but isolated in elevated or bog-like sites towards the west and south; in our region has undergone a sharp decline in recent decades in many countries south of its Fennoscandian core range. Habitat Wide variety of pools and lakes in the north, but restricted to somewhat acidic and oligotrophic or mesotrophic habitats in much of its range. Favours well-vegetated borders, e.g. with sedges. Flight season Up to early September in the north and at higher altitudes, but most numerous in the second half of May and June in the lowlands of central Europe, and seldom seen after July.

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Coenagrion lunulatum (Charpentier, 1840) Crescent Bluet

Irish Damselfly

/ pronotum much black on middle of abdomen

/ gynomorph form

mature ?

/ abdomen tip

131

conspicuous blue base of S8

? S1–2 showing ‘sad face’ pattern

? appendages (side view)

? showing green underside

Identification General A fairly robust bluet, with a noticeably greenish colour, rather black abdomen, and diagnostic pattern on S2 consisting of a black crescent and two dashes. Rather local in much of its range, often in acidic habitats. Field characters Tot 30–33mm, Ab 22–26mm, Hw 16–22mm. Stouter than C. puella. Darker and more robust than C. hastulatum, male fairly deep greenish blue, green on underside, with extensive black markings differing as follows: (1) postocular spots not ‘connected’ by blue line across back of head; (2) S2 typically with a ‘sad face’ (crescent and two dashes forming mouth and eyes), but this pattern is variable and not always reliable; (3) S3–4 more than half black; (4) S6 almost all black, thus terminal half of abdomen appears black, apart from blue ‘tail-light’. Both green to brownish gynomorph and bluer andromorph females occur frequently, both types resembling females of E. cyathigerum (see that species), with the entire abdominal dorsum black, but S8 broadly interrupted by pale area at base, which is characteristic. Hand characters Unlike C. hastulatum, male upper appendages are at least as long as lowers, and hind margin of female pronotum has large protruding lobe. Variation Andromorph females vary widely in the extent and intensity of blue coloration, e.g.

abdominal markings and postocular spots can be bright blue, contrasting with brown-grey thorax.

Occurrence Range and status Widespread in the temperate regions of Eurasia, as far east as Kamchatka, with an isolated population in the mountain steppes where Turkey, Georgia and Armenia meet. Generally rather local, but may occur at high densities. Habitat Well-vegetated pools and lakes, often acidic and nutrient-poor, such as heathy lakes. Less critical towards the east, where also found in gravel pits or eutrophic ponds. Flight season End of April to late July, especially May and June, perhaps later occasionally.

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Coenagrion hylas (Trybom, 1889)

Siberian Bluet

? ? pronotum

/ pronotum

extensive areas of black on abdomen sides and thorax

132

? black at base of hindlegs

? S1–2

? appendages / showing characteristic ‘three-coloured’ pattern

(side view)

Identification

Occurrence

General Large, dark bluet that is widespread in Siberia, but one of the rarest species in Europe. It is known only as a postglacial relict from a few sites on the northern slopes of the Alps. At present, only about a dozen Austrian populations survive. Within its European range, males are easily identified by the black-sided abdomen. Field characters Tot 33–38mm, Ab 25–32mm, Hw 19–28mm. Our largest Coenagrion species. Both sexes differ from all other small blue damsels (except the tiny C. johanssoni, which does not overlap in our region) by the black sides and underside of the abdomen. Black markings on the thorax at the base of the hindlegs are also rarely seen in species other than these two. Differs from C. johanssoni by its range and size. Moreover, males of C. hylas have well-developed black metapleural stripes extending from Hw bases to the hindlegs (also in some females of C. johanssoni). Females are rather massive, with blue postocular and abdominal spots, a green thorax and heavy black markings, rendering them unmistakable. Hand characters Not important. Variation In females, length of the black interpleural stripes varies, sometimes joining the metapleural stripes at their broadened lower ends. Behaviour Adults commute from foraging habitats up to 500m from the water in the late morning. Their numbers peak around noon but most unpaired males leave almost simultaneously an hour later, egg-laying pairs remaining at most two hours longer.

Range and status Ranges from the Urals to Japan, with an isolated population in the northern Alps and another in the far north of European Russia, ouside the scope of this book. There, C. hylas occurs on peaty taiga lakes with its congeners C. johanssoni, C. hastulatum and C. glaciale (the latter not included in this book). Extinct due to habitat destruction in Bavaria, and in our area now present in only about 14 sites in western Austria. Habitat In Austria, small lakes and peaty marshes in mountain valleys between 800m and 1,600m altitude, densely bordered with sedges and fed by cold, nutrient-poor, neutral to calcareous spring water. Flight season May to August; most abundant in June and July.

HD II+IV

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Coenagrion johanssoni (Wallengren, 1894)

Arctic Bluet

/ pronotum /

?

133

? S1–2 lateral view

? S1–2 blue tip

/

black on abdomen sides and at base of hindlegs

? appendages (side view)

Identification General Small, dark bluet with the most northern range of any European damselfly. Here they are easily identified by their size and black-sided abdomen. Field characters Tot 27–30mm, Ab 20–24mm, Hw 15–19mm. Shorter-bodied than C. puella, this is one of the smallest species of the genus and the most diminutive in our area. Because of the black-sided abdomen, both sexes are most similar to the much larger Alpine C. hylas, but they are not known to occur together in the region covered by this book. Unlike that species, C. johanssoni males usually have the U-shaped black marking on S2 connected to the segment base by a thin stem. Overlaps most widely with C. hastulatum and C. lunulatum, but those males are greenish on the underside, lack black flanks on the abdomen and usually have the black marking on S2 broken up into three parts. Female C. johanssoni has tip of abdomen (S10 and adjacent part of S9) blue, a feature otherwise restricted to the Mediterranean C. caerulescens. Hand characters The pear-shaped postocular spots are not a reliable character. Variation Extent of black on the thorax sides of females varies, with metapleural and interpleural stripes sometimes converging. A gynomorph form

with beige to yellowish thorax and abdomen is infrequent.

Occurrence Range and status Widespread and moderately common in Fennoscandia; rather local in the Baltic States. Ranges to Kamchatka and the Bering Strait. Habitat Mainly aapa mires (the predominant wetland type of the boreal taiga), mire lakes, raised bogs, and marshy bays or peaty edges of lakes. Flight season June to August.

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Coenagrion armatum (Charpentier, 1840)

Dark Bluet Norfolk Damselfly

antehumeral stripes reduced

/ pronotum ?

134

/ distinctive black abdomen in both sexes, with blue tip and base

? S1–2 large lower appendages

? appendages

Identification General A dark, inconspicuous damselfly that lives concealed in marshy vegetation. At first glance, more reminiscent of an Ischnura than a Coenagrion. Both sexes can be identified by the black abdomen marked diagnostically with blue at the base and the tip, and are easily recognised by their different but equally distinctive markings. The male is armed (hence its scientific name) with peculiar lower appendages, so large that they stand out even when seen with the naked eye. Field characters Tot 31–34mm, Ab 22–26mm, Hw 16–21mm. Shorter-bodied than C. puella. Male largely black with greenish-blue markings, antehumeral stripes are absent or reduced to one

(side view)

or two spots, and abdomen has diagnostic pattern on S1–3 and S8–9. Female is unique in having a largely black abdomen with bases S2 and S8 broadly pale. Like other Coenagrion and unlike the male, it has complete antehumeral stripes. Hand characters Male lower appendages are huge, twice as long as S10, covered with thin pruinosity. Hind margin of female pronotum with narrow rectangular lobe. Variation Female colour ranges from olive green to grey-blue. Behaviour Males wind low down through the vegetation, and can be hard to follow.

Occurrence Range and status A north-eastern species that is generally very local in our area and has disappeared from large parts of its former range, such as England, although locally (e.g. northern Germany) it seems to have recovered recently. Range extends to the Russian Far East and Kamchatka, with a relict population in the mountain steppes in Georgia and Armenia. Habitat Fields of sedges (Carex), Water Horsetail (Equisetum fluviatile) or similarly reedy plants, usually standing in shallow mesotrophic water. Here, the vegetation is quite uniform, with a critical plant density (not too open, but also not too dense), offering adults both shelter and room to move. Flight season Short and early in the southern fringe of its range, peaking in May, but as late as August in the north and at higher altitudes.

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Erythromma 

Charpentier, 1840 Identification Diagnosis This genus includes two types of species, both with bright-eyed males, sharing the following diagnostic features: (1) male’s eyes are bright red or blue, not capped with black; thus, the top of the eyes contrasts with the black upperside of the head; (2) the postocular spots are reduced to narrow stripes, or are absent; (3) the black marking on S2 on males extends along the segment, from base to tip; (4) males have no blue on the upperside of S8, while S9–10 are all or largely blue, and the ‘blue tail’ is therefore shorter and closer to the abdomen’s tip; (5) upper appendages of males are much longer than lowers, about as long as S10; (6) the wing tips, especially in the Hw, are usually densely veined, because numerous cells have been subdivided, and the Pt are rather long. Separation from other genera and of the species Females have no vulvar spine, unlike Enallagma and Ischnura. The species are compared with other blue-and-black damselflies in the species texts. The red-eyed Erythromma males are the only damselflies combining red eyes with blue markings

Brighteyes Red-eyed Damselflies on the body and an all-dark back of head. Their females have dark eyes and no postocular spots, and the entire abdomen upperside is dark. Males of other blue-tailed damselflies (Coenagrion, Enallagma, Ischnura) do not have red eyes and usually have postocular spots. Most damselflies with red eyes (Ceriagrion, Pyrrhosoma) also have red bodies. The red-faced male of Pseudagrion sublacteum is likely to be found only in Africa and the Near East. The combination of characters in Erythromma females may also be found in all-dark forms of Ceriagrion and Pyrrhosoma. However, these will often have traces of red and have differently configured black markings on the thorax; the legs of Ceriagrion have no black. If seen well, males of the two red-eyed species can be identified in the field by details of the blue pattern on the abdomen. Behaviour Unlike other damselflies, males tend to stay away from the water’s edge, preferring floating and emergent perches. Their flight is not slow and undulating, but direct and linear, quickly skimming low over the water’s surface. K-D B Dijkstra

135

F Male Erythromma viridulum showing the bronzy-black abdomen and blue coloration extending to the sides of S2–3 and S8.

F Male Erythromma lindenii. Note the contrasting blue eyes and wide antehumeral stripes.

Erythromma Brighteyes

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Erythromma najas (Hansemann, 1823)

Large Redeye



Red-eyed Damselfly

short antehumeral stripes

no antehumeral stripes

? abdomen tip upper appendages straight

136

?

/

many subdivided cells in hindwing tip

mature ?

? abdomen appearing matt and greyish

blue coloration on S1 and S9–10 square-cut

immature ? (females are similarly coloured)

Identification General Still the only red-eyed blue damselfly in much of northern Europe, although E. viridulum is expanding into its range. The classic image of this species is of a male perched flat on a lily-pad, appearing dark grey with three patches of colour: red eyes, blue thorax and blue tail-end. Field characters Tot 30–36mm, Ab 25–30mm, Hw 19–24mm. Somewhat larger and thicker than Ischnura elegans. Darker, duller and heavier than E. viridulum. Legs are all black, and the antehumeral stripes are absent in the male and

reduced in the female. Male has S2–8 largely black without blue sides, and S9–10 all blue, the blue areas therefore appearing isolated and square-cut. The dark areas on the abdomen become pruinose, appearing matt and greyish. Female is blacker than most damselflies, otherwise mainly green; brown eyes often reddish. Teneral females are remarkably bronzy, and can be mistaken for Lestes and Chalcolestes species. Hand characters Male upper appendages are straight (from above). Female hind border of pronotum with projecting narrow lobe (view from above). Behaviour Males often perch flat on floating vegetation, making quick sorties over the water.

Occurrence Range and status Common in many areas east to Japan, but can be local (especially in south) and seems to wander relatively little, not readily colonising new habitats. Habitat Richly vegetated calm or standing waters, such as ponds, ditches and canals, usually with floating leaves of water-lilies (Nuphar, Nymphaea, Nymphoides) or pondweeds (Potamogeton). Flight season From April to August, with a peak in June, therefore about a month earlier than E. viridulum on average.

Erythromma Brighteyes

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Erythromma viridulum (Charpentier, 1840) Small Redeye complete antehumeral stripes

complete or occasionally partial antehumeral stripes

Small Red-eyed Damselfly

? abdomen tip ‘X’ mark on S10 and upper appendages curved inwards

?

/

137

mature ?

? abdomen glossy black

blue coloration extends to sides of S2–3 and S8

Identification General Brighter, sleeker and daintier than E. najas, with a more southerly range and later flight season. However, the overlap in range and season is broad, and E. viridulum is rapidly expanding its range northwards into areas once inhabited only by E. najas. Like that species, males often perch far from banks, low on aquatic vegetation, although they are less partial to floating leaves. Field characters Tot 26–32mm, Ab 22–25mm, Hw 16–20mm. About as large as Ischnura elegans. Paler, smaller and more slender than E. najas. Underside of the legs is usually pale. Antehumeral stripes complete. Male has upperside S2–8 black, more narrowly so on S2–3 and S8, leaving the sides of these segments blue. The blue of the thorax and tail-end thus extends onto the sides of adjacent segments (and legs), giving a brighter but less clean-cut appearance. S9–10 largely blue, but upperside S10 marked with a black ‘X’. The abdomen is never pruinose, being glossy bronzeblack. Female is best separated from E. najas by the complete antehumeral stripes. Unlike E. lindenii, female is often marked bluish on the thorax sides and abdomen base and tip. In Morocco, Pseudagrion sublacteum males are also red-faced, but the ranges do not overlap. Hand characters Male superior appendages incurved at tips (view from above). Female hind border of pronotum not lobed (view from above). Variation Tends to become paler in south of range; the Near Eastern subspecies orientale merely

represents such a pale variety and does not warrant taxonomic status. Behaviour Typically perches on emergent aquatic vegetation.

Occurrence Range and status Expanding northwards; once absent in parts of north-west Europe, it became one of the commoner species in the 1980–90s. First reached Great Britain in 1999, now occurring widely in the south-east. Colonised Lithuania, Latvia, Belarus, Denmark and Sweden in the 21st century. Habitat Favours eutrophic standing water, clogged with aquatic vegetation. Typical emergents include algal mats, hornworts (Ceratophyllum), watermilfoils (Myriophyllum) and other plants. May also appear on other emergent vegetation, including floating peat moss (Sphagnum) in bogs. Flight season From May to September in the south, but mainly in July and August in the north.

Erythromma Brighteyes

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Erythromma lindenii (Selys, 1840) Blue-eye

Goblet-marked Damselfly

bright blue eyes contrast with black head

postocular spots reduced to slits or even absent long, pale pterostigma

broad antehumeral stripes

round knob

138 ?

/

/ pronotum and front of thorax

lanceolated S3–6

? S1–2 showing goblet-shaped mark

dull with blue middle of abdomen

? abdomen tip long, curved upper appendages

short ‘tail-light’ at extreme abdomen tip

Identification General The slender all-blue males skim swiftly over water and rest on floating perches, far from the banks. With binoculars, the broad antehumerals and terminally shifted ‘blue tail’ can be seen. In the hand, the unique configuration of markings on the head, thorax and abdomen are apparent. Field characters Tot 30–36mm, Ab 24–28mm, Hw 19–21mm. In size and general appearance, close to Enallagma cyathigerum and Coenagrion puella. Postocular spots are linear, often connected

pale appendages

across the back of the head, sometimes absent. Pale antehumerals wider than the black humerals (as in E. cyathigerum), but short black line below humeral is also present (as in Coenagrion). Male abdomen is blue, upperside S2 with a complete black goblet-shaped band, S3–6 with elongate black spearheads, S7–8 largely black and S9–10 largely blue. Coenagrion and Enallagma males have rounder postocular spots, often less blue on the thorax and S3–6 and S9–10, but more on S2 and S8, resulting in a different distribution of black and blue. At a distance, the blue males of Platycnemis pennipes appear to have a similar distribution of black on their bodies, but the details are quite different, and their broad white legs readily exclude confusion. The female is peculiarly three-coloured: the head, thorax and abdomen base are pale yellowish brown to greenish, the middle of the abdomen (especially S4–6) is blue, and the tip is pale brown. The upperside of the abdomen is almost continuously black from the base to tip. The female’s three colour zones and markings (which are similar to those of the male) are diagnostic. Hand characters Male upper appendages longer than in most bluets. Female with round knobs on sides of thorax behind pronotum. Female appendages are pale (but black in most similar species). Unlike E. cyathigerum, there is no vulvar spine.

Erythromma Brighteyes

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Variation Subspecies zernyi and lacustris have been separated in the Near East and the German–Polish border region, but differences are insubstantial and in the case of the former reflect seasonal variation. Behaviour Males fly low, and often fast, over water away from banks, perching horizontally on emergent stakes and aquatic vegetation. They are easily searched for by scanning prominent isolated perches in open water.

Occurrence Range and status A southern species that can be abundant in the Mediterranean region. Slowly expanding its range northwards. Habitat Larger, well-oxygenated waters, including lakes, gravel pits, slow-flowing rivers and wide canals, with rich aquatic vegetation such as watermilfoil (Myriophyllum). Flight season From March to October in the south, but mainly in July and August in the north.

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Pyrrhosoma 

Large Red Damsels

Charpentier, 1840 Identification Diagnosis Rather large damselflies with a mainly red abdomen, no postocular spots, black legs and dark Pt. Some females lack any red, being black marked with yellow. The thorax markings are unique: the antehumeral stripes are (often red) exclamation marks that cross the humeral sutures (as their name indicates, these stripes are normally entirely anterior to these sutures). Separation from other genera Red specimens can be confused only with Ceriagrion species, but differ by leg and Pt colour, and the distinctive

antehumeral stripes. Black females can be confused with females of Erythromma and the black form of Ceriagrion, but differ by details of the thorax markings. Separation of the species Two species are found in the region, but one occurs only locally in the southern Balkans. They can only be distinguished in the hand by the male appendages and the female pronotum. Behaviour Males behave aggressively, but do not have clear territories. Oviposits in tandem. V J Kalkman

 Pyrrhosoma nymphula male. Note the prominent red antehumeral stripes and the black legs.

Pyrrhosoma Large Red Damsels

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Pyrrhosoma nymphula (Sulzer, 1776) Large Red Damsel

140

Large Red Damselfly

/

/

/

/

blackish pterostigma

?

? appendages (side view) f. melanota without red

f. melanota

f. typica

f. fulvipes

?

note black legs and broad antehumeral stripe

Identification General The Large Red Damsel is the first sign of spring in much of our area, being the only Pyrrhosoma present in north-west Africa, Turkey and most of Europe. Although it is easy to identify to genus level, there are no characters that separate P. nymphula from its southern Balkans counterpart, P. elisabethae, in flight. Field characters Tot 33–36mm, Ab 25–30mm, Hw 19–24mm. Somewhat larger and more robust than Enallagma cyathigerum. Both sexes have a red abdomen, with only the last segments (especially S7–9) largely black. The thorax is black with yellow sides and antehumeral stripes in young specimens, the latter becoming red with age. Hand characters See P. elisabethae for structural characters that distinguish it from P. nymphula. Variation Females vary in the extent of black and red coloration. This variation probably differs between regions and populations. Detailed information is lacking, although females are generally blacker in the south. Various female

/ pronotum

forms have been named, but a wide range of intermediates exists. The three main forms are: (1) f. typica (f. intermedia is included here) – S2–6 are red, with a median black line that expands into a spot at the end of each segment and narrow yellow rings at their base; (2) f. fulvipes – this

Pyrrhosoma Large Red Damsels

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form is similar to the male, although yellow rings are present, the black line and spots on S2–6 are reduced compared with the previous form, S7–9 are almost totally black, and the red rim of the pronotum is interrupted by the black hind margin; (3) f. melanota – abdomen is largely bronze-black, the antehumeral stripes do not become red but remain yellow, unlike the other two forms, and it recalls Erythromma females. Some individuals appear to lack red on the abdomen.

Occurrence Range and status Common in large areas west of the Urals, becoming more local to the south. Habitat Wide range of waters, with the highest abundance in well-vegetated standing water. Mainly in running waters in parts of eastern Europe. Flight season One of the earliest species in northern Europe. From April to August, most abundant in May and June. Probably a few weeks earlier in the south.

141

Pyrrhosoma elisabethae Schmidt, 1948 Greek Red Damsel ventral hook is only onethird length of appendage lower appendages are slightly longer than uppers

deep folds in hind margin

/ pronotum

? appendages (side view)

Identification

Occurrence

General Replaces P. nymphula in the southern Balkans. All individuals of Pyrrhosoma here should be carefully identified (under slight magnification), because the limits of both Pyrrhosoma species are insufficiently known and their ranges may overlap. Field characters Tot 36–38mm, Ab 28–30mm, Hw 20–24mm. Coloration and markings as P. nymphula. Hand characters Differs from P. nymphula in three subtle characters that are visible with a hand lens: (1) the ventral hook of the male’s upper appendages is about one-third the length of the appendage, and about two-thirds as long as uppers in P. nymphula; (2) the male’s lower appendages reach further than the uppers (in P. nymphula, the lowers are about the same length or even shorter than the uppers); (3) in females, the hind margin of the pronotum has a deep and raised fold on each side, these folds appearing prominent when viewed both from the side and above (P. nymphula females have only slight pleats in the same position on the hind margin, and these are not raised above the margin in side view). Variation Probably the same colour forms occur as in P. nymphula. The black f. melanota seems to be relatively common in P. elisabethae, while f. fulvipes has not yet been found.

Range and status A rare and vulnerable species, currently known from a small number of localities on the Peloponnese, Kérkira (Corfu) and southern Albania. Probably present in a larger part of Albania and north-west Greece. Habitat Poorly known: found in richly vegetated streams. Flight season Records range from the last third of April to the middle of June.

CR

Pyrrhosoma Large Red Damsels

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Ceriagrion 

Small Red Damsels

Selys, 1876 Identification

142

Diagnosis The only damselflies with a combination of no postocular spots, reddish legs and Pt, and usually a (partly) red abdomen. Some females have a totally black abdomen. The yellow antehumeral stripes are very thin or even absent. Across the face (in front of the antennae) is a diagnostic sharp transverse ridge. Separation from other genera Other damselflies with red and/or without postoculars have black on the legs. This includes the more robust Pyrrhosoma species, which in addition have black Pt and differently configured thorax and abdomen markings. Black Ceriagrion females are most likely

to be confused with other small damselflies, but they never have black-marked legs. Separation of the species Only two superficially identical species are found in the area. These cannot be distinguished on sight, but in most areas only one species occurs. The two may overlap in Greece, and should be separated in the hand. Behaviour Weak fliers that stay low in the vegetation, most often in close proximity to the water’s edge. Males interact aggressively, but do not have clear territories. Oviposition takes place in tandem. V J Kalkman

Ceriagrion tenellum (de Villers, 1789)

Small Red Damsel

no or reduced antehumeral stripes

Small Red Damselfly

reddish pterostigma

/

/

/

? ? S10 and appendages

crown of black spines on tip of S10

more slender lower appendages C. tenellum

C. georgifreyi

/ pronotum and front of thorax upright lobes at front of thorax f. melanogastra

f. typica

?

f. erythrogastra

note reddish legs and all-red abdomen

Identification General This small, weak-flying damselfly is the only Ceriagrion found in north-west Africa and most of Europe. Field characters Tot 25–35mm, Ab 22–30mm, Hw 15–21mm. The only Ceriagrion in most of our

C. tenellum

C. georgifreyi

area, and therefore easily recognised by the genus characters: entirely red male abdomen, reddish legs and Pt, and absence of postocular spots. Hand characters See C. georgifreyi, which appears identical in the field, for distinguishing characters.

Ceriagrion Small Red Damsels

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Variation Four colour forms of females are recognised. The relative abundance of these varies between regions and populations. Their ranges are unclear, although f. melanogastra is more common in the south and seems to be absent in the north. The forms differ by the extent of black on the abdomen and face: (1) f. typica – abdomen black with most of S1–3 and S9–10 red, extent of black may vary slightly, face cream-coloured, with postclypeus and base of labrum black; (2) f. erythrogastra – abdomen and face all red, as in male; (3) f. melanogastra – entire upperside of abdomen and most of face black; (4) f. intermedia – includes all largely red but black-marked varieties between the first two forms, thus is quite variable, and face is marked as f. typica.

G Ceriagrion tenellum male.

143

Occurrence Range and status Widespread in the western Mediterranean, extending into north-western Europe, where locally common. Rare in a narrow strip along the Adriatic Sea (Slovenia, Croatia and Albania) and also present on Crete and Ios. Habitat Small streams and seepages (often calcareous), but in the north-west mainly bogs and heathy lakes with peat moss (Sphagnum) and often

C. georgifreyi (VU)

Marsh St John’s-wort (Hypericum elodes). Flight season From the end of May to early September in the north, peaking in July and August, but from the start of April to October in the south.

Ceriagrion georgifreyi Schmidt, 1953 Turkish Red Damsel Identification General Replaces C. tenellum in the Near East, Turkey, and on the Greek mainland and some Greek islands. While the two species are not known to occur together, the range of C. georgifreyi is still poorly understood and careful separation is required in the hand. Field characters Tot 35–40mm, Ab 28–33mm, Hw 17–20mm. Somewhat larger than the superficially identical C. tenellum. Hand characters Differs from C. tenellum in three subtle characters that are visible with a hand lens: (1) the male’s lower appendages are more slender; (2) in males, the slightly raised tip of S10 bears a crown of black spines. C. tenellum males have no such crown, but those from Crete (and possibly Ios) sometimes bear a few small spines, when the shape of their appendages is diagnostic; (3) females have two conspicuous, upright lobes on the thorax, just behind the pronotum, which are higher than the hind rim

of the pronotum (in C. tenellum these lobes are inconspicuous and never exceed the pronotal rim). Variation Probably has the same female colour forms as C. tenellum. So far, f. typica and f. erythrogastra have been found.

Occurrence Range and status Found in a narrow fringe along the Mediterranean (shown in blue on map), from Israel to south-west Turkey, and also on the Greek islands of Thásos, Zákinthos, Kérkira (Corfu), and Lésbos. Recent records near Athens and from the north-west Peloponnese suggest that older records of C. tenellum from the north-east Greek mainland may prove to be this species too. Uncommon within its small range, and possibly threatened by habitat destruction. Habitat Seepages and small streams. Flight season From the beginning of May to the end of September.

Ceriagrion Small Red Damsels

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Nehalennia Selys, 1850

Sedglings



Sprites (NA)

Nehalennia speciosa (Charpentier, 1840) Sedgling

Pygmy Damselfly pale blue line across back of head

144

?

? mature /

upper appendages large and blunt blue tip to metallic green body

pale pterostigma

Identification General Called ‘Déesse Précieuse’ in French, meaning ‘precious goddess’. Nehalennia was a goddess worshipped in the Rhine area in Roman times. This tiny odonate is perhaps the most fragile in our area, because of its weak flight, delicate size and build, and vulnerability to landscape change. These petite, metallic green damselflies mostly inhabit stands of sleek sedges in bogs, where they live concealed among the leaves. They are very inconspicuous and their presence may not be

NT

? S10 and appendages apparent until the adults are flushed from their retreats. Field characters Tot 24–26mm, Ab 19–25mm, Hw 11–16mm. Smaller than any other damselfly in our area. Not only tiny but also very slim-bodied and short-winged. Mature specimens of both sexes are bright metallic green, marked diagnostically with pale blue: a bow-shaped line across the back of the head, the lower half of the thorax, and blotches on S8–10 forming a pale blue tip to the abdomen. Unlike many coenagrionids, round postocular spots, antehumeral stripes and black markings on S10 are absent. The metallic body colours are reminiscent of Lestes and Chalcolestes species, an illusion strengthened when these are marked with pale blue pruinosity. However, all Lestes and Chalcolestes are considerably larger (over 30mm total length), have long Pt and typically perch with half-spread wings. Because of their colour and size, mature orangish females have been mistaken for Ischnura pumilio. The latter always have a vulvar spine. Hand characters Frons with sharp transverse ridge in front of the antennae, seen otherwise only in Ceriagrion. Male upper appendages are large and blunt, lower appendages are virtually absent. Variation Pale blue is replaced by (reddish) brown in older females.

Nehalennia Sedglings

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Behaviour Lives hidden among vegetation, making only short, slow, feeble, low flights between leaf blades.

Occurrence Range and status The species’ specific habitat has been fragmented or lost throughout its range, making it one of Europe’s most endangered species. While it is extinct in large parts of its western range, it seems to have recovered in some areas and has been rediscovered in several countries, such as in France in 2009 and the Netherlands in 2015. Becomes somewhat less rare towards the east (e.g. widespread and regionally not uncommon in the Baltic States), reaching northern Japan. Easily overlooked because of its

size, behaviour and poorly accessible habitat. Habitat Confined to specifically structured vegetation, usually in wet depressions of bogs or shallow borders of acidic, nutrient-poor lakes, with uniform growths of narrow-leaved grass-like plants spaced densely enough to provide protection, but loosely enough to allow free movement. Slender Sedge (Carex lasiocarpa) and Bog-sedge (C. limosa) are most characteristic, but the exceptional association with Purple Moor-grass (Molinia caerulea) and Water Horsetail (Equisetum fluviatile) has also been documented. Flight season From mid-May to mid-September; most abundant in June and July. K-D B Dijkstra

145

 Nehalennia speciosa male. Note the short wings and pale legs, as well as the blue bowshaped stripe along the back of the head.

Pseudagrion Selys, 1876

Sprites

Pseudagrion sublacteum (Karsch, 1893) Cherry-eye Sprite Identification General Our only member of Pseudagrion, one of the largest damselfly genera, with over 150 species in the warm regions of the Old World. Males are brownish grey, with a bright rufous face. Field characters Tot 32–39mm, Ab 25–33mm, Hw 17–23mm. Slightly larger than Enallagma

cyathigerum. The front half of the head, including the eyes, is brick-red or orange in mature males. Black markings on the head and thorax are limited, outlining large pale brown to dull orange postocular spots and antehumeral stripes. The sides of the thorax are bluish in males and the abdomen has almost continuous black dorsal markings,

Pseudagrion Sprites

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border of S10 toothed

/

? ? S10 and appendages front half of head red

146 /

which are distinctly torpedo-shaped on S3–6. S8–9 are blue (before they become pruinose). Apart from the face, males become covered with whitish pruinosity; this is most dense on S8–9. Females are pale brown, with vestigial black markings. The extent and shape of these markings are unlike any other coenagrionid females in our area. Erythromma species also have red-eyed males but do not co-occur, and have bright blue thorax sides and tail-end. The distribution of red on the head also differs. Hand characters Apical border of male S10 has several small black teeth, a feature found in no other non-red damselfly in our area. Male upper

appendages long and blunt, clearly surpassing lower appendages. Latter with a prominent dorsal knob (seen from side) and rounded tips (seen from side and above). The Swarthy Sprite P. hamoni may be found in north-west Africa and shares the red face and toothed S10. The mature male is very dark, with the lower appendages marginally longer than the upper ones. The lower appendages are not or only weakly knobbed and pointed. The Syrian Sprite P. syriacum is known from southeastern Turkey but lacks the red face and is more likely to be mistaken there for another pruinose damselfly, Platycnemis kervillei (see that species). Variation Extent of black markings and intensity of pruinosity varies individually, regionally and with age. Combined with the underlying colours, this may produce an array of grey, brown and violet hues. Behaviour Males perch on stones and sticks.

Occurrence Range and status Common in much of tropical Africa. Relict outposts in Morocco and the Near East. Habitat Sunny ponds, ditches, streams and rivers in most of its range, but in Morocco confined to fast-flowing lowland rivers in semi-arid areas. Flight season Recorded from the end of May to early August in Morocco. K-D B Dijkstra

Pseudagrion Sprites

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Aeshnidae

Aeshna Fabricius, 1775 Mosaic Hawkers

Mosaic Darners (NA)

Identification Diagnosis Best identified by the exclusion of other aeshnid genera; specialists do not regard all species as closely related. It is quite certain that A. isoceles will be placed in a separate genus in the future. Typical Aeshna species have dark bodies inlaid with coloured bands on the thorax and ‘mosaics’ of spots on the abdomen. The wings have a median space (large central space at base) without cross-veins. Fields before medial supplement and radial supplement have three or more irregular rows of cells. Male has auricles and an anal triangle of two to four cells (except A. isoceles). Separation from other genera Closest to Brachytron, which differs by numerous (although somewhat relative) features, including early flight season, small size, hairy body, thin Pt and abdomen not waisted (unlike Aeshna males). In the hand, unique details of venation and markings are apparent. Anax differs in shape and wing venation, but is ruled out in the field by the plain thorax and

pale abdomen with a black band on the upperside (except the huge A. immaculifrons). Boyeria and Caliaeschna have cross-veins in the median space. Separation of the species A large and diverse group, the majority of the world’s species occurring in North America. Aside from a few pairs of similar species, each species has several unique features. The simple key below leads to the main groups. Females may develop male coloration, although the intensity and extent of blue markings vary strongly. Behaviour Often seen hawking in open but sheltered places, such as glades and gardens, as well as over predominately still waters. Hunting individuals often concentrate at good sites, especially towards dusk. Males make aggressive and swift patrols over water, usually following a fairly fixed and often extensive route, frequently interrupting direct flight with hovering pauses. Females of all species, except A. affinis, usually oviposit alone.  K-D B Dijkstra

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Simple key to (pairs of similar) species. If the statement agrees, compare the given species. If it disagrees, then go to the next line.

1

Prominent green antehumeral stripes that are wider than the dark area between them. Sides of thorax are always green with black lines.

cyanea viridis

2

Body is mostly brown, the abdomen without a clear ‘inlaid’ pattern of spots on upperside. Marking on upperside of frons is vague or a single dark cross-bar, but not a distinct black ‘T’.

grandis isoceles

3

The distance along which the eyes touch each other is less than twice the length of the occipital triangle between them.

caerulea

4

Total length is less than 65mm. Facial suture with no or only thin dark line, not a thick black one. Anal loop encloses two columns of cells, not three or more. Anal triangle is of three cells, not two.

mixta affinis

5

Male has a group of small teeth on the upperside of upper appendages (view from side). In female, hind border of ovipositor plate at base is rather straight (view from below).

crenata serrata

6

Male has a smooth upperside of upper appendages. In female, hind border of ovipositor plate is deeply incised (view from below).

juncea subarctica

Aeshna Mosaic Hawkers

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Aeshna mixta Latreille, 1805

Migrant Hawker  Aeshna mixta male.

148

Identification General The commonest small hawker, but beware confusion with the similar-sized Brachytron pratense, especially earlier in the season. Numerous in much of our area, and although it can be on the wing during most months in the Mediterranean, further north it is especially associated with late summer and autumn, when it may appear in massive migrations. It is usually identified by its size, relatively dull colours and the diagnostic yellow ‘nail’ mark at the abdomen base. Field characters Tot 56–64mm, Ab 43–54mm, Hw 37–42mm. Distinctly smaller than most Aeshna, it resembles a small A. juncea. The upperside S2, with its central T- or nail-shaped yellow marking, is most notable in mature males, where it contrasts with the blue abdominal spotting; females and

fresh males have yellowish spots. The antehumeral stripes are reduced to a short yellow stripe in both sexes, but are prominent in other species, especially males (e.g. A. juncea). See A. affinis for a comparison. Hand characters Unlike A. juncea and other large mosaic-marked Aeshna, the facial suture (between frons and clypeus) is not marked with a distinct black line. Together with A. affinis, differs from most Aeshna by having two (not three) columns of cells in the anal loop, three (not two) cells in the anal triangle, and seven to nine (not more) cross-veins between the node and Pt. Unlike A. affinis, the male upper appendages lack a tooth on the underside near the base, and the female appendages are longer than S9–10 combined. Variation In cool weather the blue markings of the male change to lilac. Males from the Urals and Western Siberia have entirely blue abdomen markings, even lacking the yellow nail, and are easily confused with A. affinis. Considering the species’ migratory behaviour, such individuals could show up anywhere in Europe. Behaviour Prefers to hawk swiftly quite high up along trees, often above 2m, especially in late afternoon. Male patrols at breeding sites are lower down, often along reedy banksides, and include frequent hovering.

Occurrence Range and status Widespread and common in large parts of Europe, east to Japan. Influxes in late summer can often lead to sudden surges in the

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three cells in anal triangle

149 two columns of cells in anal loop

short antehumeral stripes

mature ?

distinctive yellow ‘nail’ mark on S2

immature ?

/

facial suture at most darkened somewhat

mature ?

number of individuals present in an area. Currently expanding its range, having colonised parts of northern Britain, as well as Ireland (2000) and Finland (2003). Habitat Breeds in a wide range of still and slowflowing waters with some riparian vegetation, such

as reeds; avoids acidic water but tolerates brackish water. May be found anywhere when migrating or hunting. Flight season From May to December in the south, but mainly August and September in most of its range.

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Aeshna affinis Vander Linden, 1820

Blue-eyed Hawker Southern Migrant Hawker

Identification

150

G Aeshna affinis male. Freshly emerged, its colours are still as in the female. Note the distinctive fine-lined green side and small antehumerals.

General In flight, often confused with the related and similarly small A. mixta. Ranges less far north, but also migratory and may be invasive in good summers. Males are often observed when making low patrols over drying wetlands, showing their noticeably bright colours. The male’s vivid blue eyes and abdomen and largely green thorax sides are especially distinctive, reminiscent of a miniature Anax imperator. Field characters Tot 57–66mm, Ab 39–49mm, Hw 37–42mm. Size and build as A. mixta, but male lacks yellow ‘T’ mark on S2. Sides of thorax green, with fine black lines along sutures, but with only small green antehumeral stripes (unlike other greensided Aeshna such as A. cyanea and A. viridis). Blue markings on the male abdomen are brighter and more extensive than in A. mixta, most noticeable being the larger middle spot-pairs on S3–7 and the all-blue S2 with its characteristic black mask-like marking. Beware of Russian A. mixta with wholly blue abdomens that may wander, especially into eastern Europe. The green- or yellow-marked females (and young males) generally also appear brighter and paler than A. mixta. In size and colour may resemble Brachytron pratense. Hand characters See A. mixta for venational peculiarities of both these species. Pt of A. affinis is longer than in A. mixta. Male upper appendages have a blunt tooth on underside near base, unlike A. mixta (view from side). Female appendages are shorter than S9–10 combined. Variation Mature males develop a blue hue on the green thorax. Like other Aeshna, females may develop male colours, an especially startling phenomenon in this bright species. Behaviour Males patrol low, at about chest height, often beating to and fro over a small area; frequently perch and not easily disturbed. This is the only Aeshna that regularly lays its eggs in tandem.

Occurrence Range and status Seldom abundant, but may migrate in huge aggregations. Most frequent in areas with a continental climate but also permanently present around the Mediterranean, although scarce in much of the Iberian peninsula and north Africa. Hot summer weather may lead to influxes further north with a vagrant male in

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relatively long pterostigma

? S2 blue with black ’mask’ marking

151 venation as A. mixta

/

mature ?

large blue spots

bright blue eyes

rather short appendages mature ? showing strong blue wash to green sides

H Aeshna affinis male. Note bright colours and vivid blue eyes.

Iceland in 2018 being only the third odonate species recorded there, all of them aeshnids. Habitat Prefers standing waterbodies that dry up over the course of summer, often overgrown with

low rushes, bulrushes or reeds. Flight season On average, emerges earlier than A. mixta. Seen mainly from May to August, especially in the later months.

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Aeshna isoceles (Müller, 1767)

Green-eyed Hawker Norfolk Hawker

152

G Aeshna isoceles male. Note the contrasting green eyes.

Identification General A brown and rather plain hawker, with largely clear wings and conspicuous green eyes. The yellow triangle on S2 is diagnostic, as are the colour and shape of the Hw base. Males patrol marshy ditches, reedy lakesides and other lush, calm waters. Field characters Tot 62–66mm, Ab 47–54mm, Hw 39–45mm. Between A. mixta and A. juncea in size. Entire body is brown and virtually unmarked,

apart from a slightly dark crest on the frons, two yellow bands on the thorax sides, dark lines on the abdomen and a conspicuous yellow triangle on upperside S2. When mature, the green eyes contrast with the brown body. Wings clear, with dark veins and a diagnostic deep amber patch at the Hw base. The all-brown Aeshna grandis is larger and has entirely amber wings, and blue or yellow spots on the sides of the abdomen. The brownish Anax ephippiger and A. parthenope can have greenish eyes, but have an unbanded thorax and blue ‘saddle’ at the abdomen base. Hand characters Male membranule and anal triangle elongated, the latter with three to seven cells (often two and seldom more than four in Aeshna). Male upper appendages more slender than those of other Aeshna, with tooth near base on underside. Variation Individuals from the southern Balkans eastwards tend to have extended yellow thorax markings with prominent nail-shaped antehumeral stripes. These have been treated as ssp. antehumeralis but are probably just paler due to the warmer climate and therefore do not warrant taxonomic status.

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green eyes

yellow triangle on S2

153

amber patch on wing base covering drawn-out and many-celled anal triangle, and bordered by extended membranule

/

mature ?

never with blue markings

mature ? no pale spots on S3-10

Behaviour Unlike most hawkers, males frequently perch during patrols.

Occurrence Range and status Widespread but very local in much of its range, especially in south-west; often numerous where present. Extends to Central Asia in the east.

Habitat Ditches, marshes, ponds and lakes with rich vegetation. Favours swamps of Water-soldier (Stratiotes aloides) in most of its northern range. Sometimes on running water in the south. Flight season May to August in most of its range, most abundant in June; earlier than most Aeshna.

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Aeshna grandis (Linnaeus, 1758)

Brown Hawker

154

G Aeshna grandis male. Note the blue spots on the side of the abdomen.

Identification General Heavy hawker with a tobacco-coloured body and amber wings, often seen gliding majestically down forest glades or high over woodland lakes. A mere glimpse is usually sufficient for a positive identification. Field characters Tot 70–77mm, Ab 49–60mm, Hw 41–49mm. Size similar to A. juncea, but appears larger because of its deeply tinted wings. Body, including eyes, entirely reddish brown; thorax with two lemon-yellow bands on each side. Mature male with blue highlights in eyes, six blue spots on upperside (one at each wing base and a pair on S2) and a series of spots on sides of S3–8. Latter

markings yellow in most females and immatures, the four spots at the wing bases being the only blue on females. Wings deep yellow to golden brown, including veins. ‘T’ mark on frons indistinct (more like a crescent), with no or very thin stem. See A. isoceles for comparison. Will seldom be seen with the similarly large, brown Anax ephippiger and A. parthenope. Hand characters Not really needed. Seen from above, the male upper appendages have rounder tips than most species. Variation Individuals from northern Fennoscandia, known as the subspecies linnaei, are smaller on average and have narrower thorax bands. Behaviour Hawks with a characteristic slowseeming gliding flight, interspersed with bursts of shallow wingbeats and graceful turns and loops. Often hunts deep into dusk. Appears to mate predominantly around sunrise, when males enter dense vegetation to find roosting females. Mating wheels formed while the female clings to her perch can be awkwardly twisted, with the male facing down and his abdomen lying along the female’s face.

Occurrence Range and status One of the commoner hawkers of central and eastern Europe, especially in forested areas; very common over much of the boreal forest belt of northern Europe, being the most abundant

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155

golden brown wings

/

mature ?

mature ?

blue markings on side of abdomen

dragonfly species in Finland, for example. A female in Reykjavík in 2011 was only the second odonate species recorded on Iceland after Anax ephippiger. Becomes scarce and restricted towards highlands in south of range. East to Lake Baikal.

Habitat Breeds in a wide range of calm waters, usually with rich bankside or submerged vegetation, such as abandoned canals, oxbows and fenlands. Adults are often seen in forested areas. Flight season Late May to early October; most abundant at the end of July and in August.

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Aeshna caerulea (Ström, 1783)

Azure Hawker

156

Identification General Small, very blue hawker of the far north and high mountains, known for its habit of basking on light surfaces. This behaviour warms the body, enabling the species to survive in cold regions, but also enables detailed examination by observers. Through binoculars, the diagnostic short area of contact between the eyes can be seen. Field characters Tot 54–64mm, Ab 42–48mm, Hw 37–41mm. A similar size to Aeshna mixta,

G Aeshna caerulea male characteristically perched flat on a light surface. making it distinctly smaller than the other Aeshna species with which it is usually found. Lateral stripes of the thorax are narrow and S-shaped, but the abdominal spots are relatively large, leaving little black between them. Antehumeral stripes are reduced in males and absent in females. All pale areas are blue in mature males, yellow in females and immatures. Hand characters The distance along which the eyes touch each other is less than twice the length of the occipital triangle between them. The fork in vein IR3 is not well defined, unlike in other Aeshna. Variation Although colour change resulting from temperature fluctuation is common in dragonflies, it is especially notable in this species. Cold males have dark grey spots that become pale blue as the insects heat up. Females can have either yellow or blue markings, like males. Behaviour Dependent on sunshine to be fully active; frequently perches horizontally on bright, reflective surfaces, such as rocks and birch trunks, allowing the sun to heat its body. Males patrol low along pool edges.

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short antehumeral stripes

small area of contact between the eyes

IR3 without welldefined forking near base of pterostigma

157

/

mature ?

blue markings comparatively extensive

narrow sinuous bands on side of thorax

mature ?

Occurrence Range and status A boreal species, most common above or north of the tree line. One of the few species numerous in Eurasia’s polar regions eastwards to Kamchatka, and indeed the dominant dragonfly of the shrub tundra of subarctic Europe. Occurs as a postglacial relict in Scotland and central European mountains, where it is scarce and local. Habitat Alpine and arctic moors, heaths and

tundras; seldom below 1,000m in the Alps (mainly 1,400–2,200m in Switzerland). Breeds in bog pools and sedge swamps, and in northern Fennoscandia in almost any type of standing and slow-flowing water. Flight season Mid-July to mid-September in most of Europe, but relatively early in Scotland, where it can be on the wing as early as late May, although typically from mid-June.

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Aeshna cyanea (Müller, 1764)

Blue Hawker



Southern Hawker

Identification

158

General Large and gaudy hawker, its dark body inlaid with bright nuggets of apple-green and sky blue. Typically hunts low down along borders, often in the half-dark, like a phantom. Markings, morphology and behaviour render it unmistakable. It is not closely related to other European Aeshna. Field characters Tot 67–76mm, Ab 51–61mm, Hw 43–53mm. One of the larger Aeshna species, close in size to A. juncea. Mature male has pairs of green spots on upperside of S1–7, in marked contrast to the blue spots on sides and on S8–10. Upperside of S9–10 each with a single blue band, rather than paired spots, forming a conspicuous ‘tail-light’. Markings on female and immature are (yellow-) green. Thorax with two broad oval green antehumeral stripes and green sides interrupted by two thick black lines, one of which is complete and one only half complete. Hand characters Hind border of eyes strongly curved, revealing pale spot behind eyes even when viewed from side. In male, the anal triangle has three cells, sometimes up to six, but rarely two as in other large Aeshna species. Pt is notably short (only about 3mm), and twice as long as wide. In male, upperside S10 is flat and smooth (unlike the tooth-like ridge found in other Aeshna, seen from side). Male upper appendages expand towards the end, then abruptly narrow to form a down-turned spine (seen from the side, tip appears like a bird’s head).

G Aeshna cyanea male, showing the ‘bird’s head’ profile of the upper appendages.

Behaviour Has a diagnostic hawking flight, in which it winds about erratically, usually alone, low to the ground in shade or dusk. The abdomen is held in a slight downward curve. Curious, inspecting every nook and cranny, and diving for midges around the observer. Frequently caught by cats. Males aggressively guard a site, therefore usually only a single male is seen hovering over a small pond from which hundreds of adults may have emerged.

Occurrence Range and status Breeding habitat is favoured by few other dragonflies, and larvae can live in huge densities therein. Often common in urban or heavily forested areas. One of the commoner Aeshna species in central Europe, becoming scarcer towards the north-east (to Urals) and south. Habitat Breeds in wide range of waterbodies, preferring those that are stagnant, small and shaded, and often murky with no substrate other than leaf litter, such as garden ponds or forest pools. Feeds along woodland rides and clearings. Flight season Most typical of the late summer (July and August) and seldom seen before June, but flies well into autumn, with occasional records in November.

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broad antehumeral stripes pterostigma short

159

mature /

mature ?

immature ? blue spots on S9–10 coalesce to form ‘tail-light’

upperside S10 untoothed

mature ?

contrasting blue spots on sides of abdomen green thorax sides with thick black stripes

‘bird’s head’ profile of

? upper appendages

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Aeshna viridis Eversmann, 1836

Green Hawker

160

Identification General A light blue male hawker patrolling the length of a mass of Water-soldier (Stratiotes aloides), or a very green female egg-laying in the heart of a rosette of this plant, are the classic images of this large aeshnid. The rustling of the female’s wings as she oviposits is often the first sign of the species’ presence. This behaviour demonstrates the species’ dependence on this spiny water plant. But beware, as a large blue-andgreen dragonfly above Water-soldier could also be Anax imperator.

G Aeshna viridis pair mating. The twisted position results from the male grabbing the female in the early morning while she is perched among dense vegetation.

Field characters Tot 65–75, Ab 43–54mm, Hw 38–45mm. Slightly smaller than A. juncea. Mark on the frons has an extremely thin stalk, appearing as a single black cross-bar rather than a ‘T’. Thorax with two broad green antehumeral stripes, as in A. cyanea, but all-green sides have scarcely discernible black lines. Eyes and abdomen spots pale blue in the mature male, green in females and fresh males. Leading edges of rather yellowishtinged wings are bright yellow, as opposed to dark brown in A. cyanea. Superficially similar to Anax imperator, which differs in many details of markings, most notably by the all-green thorax. Compare with other green-sided Aeshna, such as A. affinis and A. cyanea, which have very different habitat preferences. Hand characters No black line on facial suture. Has a small, sharp yellow dot behind the eyes, which is not as clearly visible in side view as in A. cyanea, while A. affinis lacks such dots. Behaviour Normally found close to Water-soldier swamps but may travel further afield to hunt, and may be seen swarming in meadows, clearings,

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‘T’ mark on frons almost reduced to a crescent prominent antehumeral stripes

161

yellowish tinged wings mature ?

/

eyes and abdomen rather pale blue

very green overall

side of thorax almost unmarked

mature ?

along roadsides or over mats of Water-soldier before sunrise and after sunset. Seeks the shelter of rough herbage, reeds and coppices in open landscapes, often found resting low down in such vegetation. Like A. grandis, males seek roosting females to mate with at sunrise, leading to similarly irregular mating wheels.

Occurrence Range and status Holland to western Siberia. Habitat specialisation makes this species scarce and under threat in much of its range. Habitat Marshlands, ditches and lakes with sizeable masses of Water-soldier. Flight season From late June onwards, perhaps occasionally to early October; most abundant in August.

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Aeshna juncea (Linnaeus, 1758) Identification

162

Moorland Hawker Common Hawker, Sedge Hawker (NA)

General Could be considered the archetypal hawker. Large and dark, marked with yellow to bluish spots and bands, best identified by the exclusion of other species. It is the most widespread and numerous of a complex of similar species, which also includes A. subarctica, A. serrata and A. crenata. These species must always be identified in careful comparison with A. juncea. Field characters Tot 65–80mm, Ab 50–59mm, Hw 40–48mm. One of the larger Aeshna species. Most large hawkers with the following characters will belong to this species: (1) frons with thick black ‘T’ mark; (2) thorax dark brown with two broad yellow bands on each side (upper ends sometimes bluish) and two narrow antehumeral stripes (smaller in female); (3) leading edge of wings yellow; (4) abdomen brown-black, with a pattern of small ‘inlaid’ blue and yellow spots in mature males. Females (like young males) have predominantly yellowish spots, turning greenish (rarely bluish) with age. Males of A. crenata and A. subarctica

(see for comparison) are darker and more uniform, with a brown leading edge on wings, and have generally blue markings. In the field, A. serrata differs most notably by the narrow-stemmed ‘T’ mark on the frons. Hand characters Has a small, sharp yellow dot behind the eyes, where similar species are all black. Female appendages bluntly pointed, tilted relative to each other, together forming a ‘V’ in cross section (view from behind). Variation Markings of thorax are very variable, sometimes showing considerably broadened yellow bands or even almost entirely yellow sides. The yellow spots behind the eyes may be absent in southern mountain populations, but are consistently present in the north where A. juncea overlaps with A. subarctica. Behaviour Strong flier, males patrolling fast and tirelessly along and over water. They are aggressive and inquisitive, chasing off other dragonflies and searching actively for females egg-laying amid vegetation.

Comparison of species similar to A. juncea. Note: Dark = leading edge of wings brown, mature male abdominal spots rather evenly coloured; Pale = leading edge of wings yellow, mature male with more distinct yellow and blue spots on abdomen. / A. serrata

/ A. juncea

hind border of ovipositor plate rather straight (view from below)

hind border of ovipositor plate deeply incised (view from below)

? A. crenata appendages (side view) showing toothed upper appendage

Diagnostic features

General

Incised ovipositor plate

Toothed upper appendages

Species

Hw often over 53mm. / appendages acute.

Dark

no

yes

crenata

Pale spot behind each eye.

Pale

yes

no

juncea

Narrow stem of ‘T’ mark on frons. Membranule evenly pale, not two-toned. ? upper appendages strongly upcurved, at least twice as long as lower.

Pale

no

yes

serrata

None of above.

Dark

yes

no

subarctica

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163 /

mature ?

/ ovipositor (from below)

abdomen spots may be yellow, green or blue in /

hind border of basal plate is deeply incised

yellow spot behind eye

black line on facial suture narrowed at base

two broad yellow bands on side of thorax

mature

?

Occurrence Range and status One of the commonest hawkers in most northern and montane areas, becoming more local and restricted to higher altitudes to the south. Has the largest range of any Aeshna species, stretching over much of Eurasia and North America. Habitat Largely restricted to acidic heathy lakes and bogs in the European lowlands, but regularly found in smaller numbers in many other types of standing water, such as gravel pits, richly vegetated ponds or ditches with Water-soldier (Stratiotes aloides); habitat range widens at higher altitudes and latitudes.

Flight season Occasionally emerges in late May and June and may persist to November, but most numerous from July to September.

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Aeshna subarctica Walker, 1908

Bog Hawker Subarctic Hawker

Identification

164

General Doppelgänger of A. juncea, found exclusively within that species’ range and often with it, occurring in similar habitats, but largely confined to bogs with floating mats of peat moss (Sphagnum). Appears darker and more uniform than A. juncea, but careful study of markings is needed for positive identification. Field characters Tot 70–76, Ab 47–57mm, Hw 39–46mm. Size and build like A. juncea. Generally darker than the latter, the body appearing blacker and leading edge of wings more brown than yellow. Colour pattern of thorax and abdomen more uniform, quite evenly bluish, unlike more clearly differentiated yellows and blues of A. juncea. Flight identification is unreliable. Hand characters The two most reliable differences with A. juncea are on the head: (1) the black line along the facial suture (between frons and clypeus) is of constant thickness or even (often) widens at the eyes, while in A. juncea it narrows at the sides; (2) all black behind the eyes, without the yellowish dot of A. juncea. Other distinguishing features are: (3) underside of thorax often has two dropshaped yellow spots, but is usually uniformly dark in A. juncea; (4) male upper appendages are wider than in A. juncea; (5) female appendages are also wider, more symmetrical and with rounder tips. They lie in a single horizontal plane (view from behind). Variation Tends to be significantly paler in central European lowland bogs than in boreal and alpine regions. Such pale specimens (not illustrated) possess the following supplementary differences to A. juncea: (6) thorax sides with a thin, pale line

G Aeshna subarctica pair mating. anterior to the first lateral band, and a rather large, pale marking between the two lateral bands (pale thorax may be noticeable in flight); (7) middle pair of spots on abdominal segments almost as large as terminal pairs. Behaviour Males patrol above floating peat moss. Females egg-laying there are often sufficiently still and out in the open to allow identification through binoculars.

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leading edge of wing dark brown

165

/

? dark mature ?

black line on facial suture wide at base all black behind eyes

/ ovipositor (from below)

hind border of basal plate is deeply incised

Occurrence Range and status Rather localised because of its habitat; has a scatter of relict populations in much of our area. Ranges to Japan and also occurs in North America, from where the species was first described. European populations are known as the subspecies elisabethae. Habitat Moors and bogs with floating peat moss. These very wet communities appear like a thick soup, and recognising them is often the best way to find the species. Flight season Emerges from late May, but seldom before mid-July in most areas; most abundant in August and September.

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Aeshna serrata Hagen, 1856

Baltic Hawker Identification

166

G Aeshna serrata male.

General A large hawker, occurring only in and around the Baltic Sea in our area. Often known as A. osiliensis, described from the Estonian island, Saaremaa (German name: Ösel). Morphological differences are insubstantial, while scattered populations across northern Russia suggest that the Baltic population is connected with the species’ largely Central Asian range, weakening the justification for a distinct species even further. Field characters Tot 75–81mm, Ab 50–62mm, Hw 48–53mm. Larger than A. juncea, but very close in general appearance, notably the yellow leading edge of the wings and clear differentiation of yellow and blue markings. The abdominal spots are larger and deeper in colour, making the species appear bluer. Unlike A. juncea, females bear large, pale antehumeral stripes. The most notable difference from similar species is the narrow-stemmed black ‘T’ mark on the frons. See table for a summary of the differences with similar species (p. 162). Hand characters Membranule evenly whitish to pale grey, unlike the largely dark, white-based membranule of related species. Lower appendage just under half the length of upper appendages, the latter with a curved profile and cluster of four to seven small teeth near the tip, on the upperside, all visible in side view. Lower appendage of similar species is over half the length of upper appendages, the latter being straight and only toothed in A. crenata. Variation Andromorph females are frequent and may outnumber the yellowish gynomorphs locally. Behaviour Appears heavier with a slightly slower flight than A. juncea. Individuals of all sexes and ages are often seen patrolling open areas such as fields and meadows. Unlike other large aeshnids, A. serrata stays notably low, frequently roosting in vegetation on the shore and even perching flat on bare ground such as gravel roads, boardwalks, jetties or boats, especially in the early morning and during cool windy weather.

Occurrence Range and status Locally abundant on Baltic coast; discovered in Denmark in 2006. Known from eastern Turkey and mountain steppes of the Transcaucasus. Not confirmed, but may be found on reedy lakes and in brackish marshlands of central and northern Turkey. Main range is in the

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‘T’ mark on frons with narrow stem prominent antehumeral stripes in both sexes

leading edge of wing yellow

167

/

?

abdomen spots comparatively large

/ ovipositor (from below) hind border of basal plate straight

? appendages (side view) upper appendages curved with small teeth near tip

lower appendage under half length of uppers

(forest) steppes from the southern Ural eastwards; also found in the taiga of northern Russia, east to Kamchatka. Habitat Brackish (i.e. coastal) water with reedbeds, including shallow bays of the Baltic Sea and

adjacent lagoons. In inland Sweden also inhabits shallow eutrophic lakes, ponds and sluggish rivers dominated by reeds and bulrushes, in an agricultural landscape. Flight season July to September.

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Aeshna crenata Hagen, 1856

Siberian Hawker

168

G Aeshna crenata male.

Identification General An impressive hawker, noted for its large size and vivid colour. When mature, males are clear blue and females have large, dark brown patches on their wings. It is an inhabitant of the Siberian taiga, just reaching southern Finland and northeast Lithuania. Field characters Tot 71–86mm, Ab 53–67mm, Hw 44–60mm. Our largest Aeshna species, larger even than Anax imperator. Black ‘T’ mark on frons is very thick-stemmed. The leading edge of the wings are brown, unlike the yellow of A. juncea and A. serrata. Males of these species also have the

thorax markings and the smaller abdominal spots mainly yellow (contrasting with blue spots), while in mature A. crenata all markings are blue. Most females are marked yellow to green. Older females often have dark patches between the node and Pt, noticeable in flight. See table for a summary of the differences with similar species (p. 162). Hand characters Male upper appendages with two or three rather robust teeth placed on a ridge on the upperside, just before the tip. A. serrata also has such teeth, but about four to seven of them (A. crenata may have up to six teeth, but the additional ones tend to be rather small). Female appendages are pointed, unlike those of similar species, which are rounded. Variation Antehumeral stripes are well developed and often bluish in mature andromorph females but absent or reduced to a small yellowish-green patch in gynomorphs. Occasionally both sexes can have strikingly enlarged and washed-out antehumerals that may cover almost the entire front of the thorax. Behaviour Males persistently patrol over open water along the vegetation’s edge, only rarely pausing to hover. They are often very aggressive towards other Aeshna males, forcing them to adjust their behaviour, such as by flying less conspicuously or when A. crenata males are absent.

Aeshna Mosaic Hawkers

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‘T’ mark on frons with thick stem

leading edge of wing brown

169

? all blue markings on abdomen

? appendages

/

(side view) upper appendages straight, with large teeth near tip

lower appendage more than half length of uppers

Occurrence Range and status Ranges from the Baltic shore across Siberia to Japan. Seems to be very local in our area, but may be under-recorded in its European range. Habitat Westermost populations are in small or medium-sized nutrient-poor, acidic forest lakes bordered by bog-moss (Sphagnum) and sedges (Carex). More of a generalist further east, e.g. in large lakes, oxbows and even overgrown watering holes for lifestock. Flight season Late June to mid-September, not flying well into autumn, unlike other Aeshna species; most records in the second half of July and August.

/ ovipositor (from below) pointed appendages

hind border of basal plate not deeply incised

Aeshna Mosaic Hawkers

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Anax Leach, 1815 Identification

170

Diagnosis Among our largest dragonflies. With the exception of one aberrant east Mediterranean species, our species have a uniformly brown or green thorax, and a green, blue or brown abdomen with a broad black mid-dorsal stripe. The aberrant species, A. immaculifrons, has broad black thorax bands and a striking black-and-yellowringed abdomen. Males, like all aeshnid females, lack auricles on S2 and the anal triangles. IR3 is not forked proximal of the Pt, and R3 is abruptly arched forward near the distal end of the Pt. The Rspl curves forward strongly, its distal end pointing towards a point between the Pt and the wing tip. Separation from other genera Other aeshnids have a banded thorax and a largely dark abdomen with a series of paired coloured spots. The males of these genera possess auricles and anal triangles, their IR3 is usually forked, R3 gradually curves backward and the Rspl points to the wing tip. A. immaculifrons may suggest a Cordulegaster in flight, but can be recognised by its thicker

Emperors Green Darners (NA) abdomen. In the hand, it is easily separated by its broadly confluent eyes. Separation of the species With the exception of A. immaculifrons, flight identification can be difficult because most characters (e.g. colour of abdomen, thorax and eyes) vary and have some overlap. In young individuals, for instance, the body colour is often green instead of brown or blue. In the hand, important characters are the frons markings, male appendages and the presence of occipital tubercles in females. The table below compares the three most widespread species. On the Atlantic coast, careful observations may produce new records of the American vagrant A. junius. Behaviour All species are fast, powerful fliers. Males of most species patrol effortlessly over the centre of larger waterbodies. Will often rest low among waterside vegetation, for instance during poor weather, hanging vertically. Oviposition mode varies between the species and can help species diagnosis.  V J Kalkman

H Anax imperator male in flight. Note the slight downward curve of the abdomen.

Anax Emperors

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Field characters for adult Anax imperator, A. parthenope and A. ephippiger. imperator

parthenope

ephippiger

Size and shape

Large, abdomen thick and longer than Hw

Large, abdomen thick and longer than Hw

Smaller, abdomen slender and as long as Hw

Frons

Bluish bar behind crest, black pentagon at base

Bluish bar behind crest, black triangle at base

Black bar on crest and at base, never with blue

Eyes

171

Green to blue above, with yellow to green underside

Uniformly green (a shade darker than in imperator), sometimes tinged yellow, blue and/or brown

Brown above, with a (greenish-) yellow underside, never with blue

Thorax in front of Fw

Green, normally blue in males

Normally brown

Normally brown

Thorax

Green, sometimes brownish in older specimens

Normally brown, sometimes tinged grey or green

Normally brown, often lower sides contrastingly green-yellow

S1

Green, sometimes blue

Normally brown

Normally brown

Basal ring S2

Green, only yellow when young

Normally yellow

Normally brown, sometimes yellow

‘Blue saddle’ S2

Not distinct because S3–10 are also blue (or S2–10 all green in most females and tenerals)

Normally distinct, wrapped around segment and extending onto base S3

Normally distinct but sometimes difficult to see (often absent or inconspicuous in females), usually restricted to top of segment (sides of segment are dull yellow)

Ground colour S3–7

Green, turning blue in all mature males and some mature females

Brown, sometimes bluish

Brown or straw-coloured

S8–10

Colour and markings similar to preceding segments, sometimes brownish

Colour and markings similar to preceding segments, sometimes blacker

Blacker than preceding segments, leaving only clearly defined pale spots at hind corners of segments (these often paler than brown of S3–7)

Wings

Sometimes tinted between triangle and tip

Often conspicuously tinted brown between node and Pt

Often inconspicuously tinted in central part of the wing

Anax Emperors

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Anax imperator Leach, 1815

Blue Emperor



Emperor Dragonfly

Identification

172

General A common and conspicuous dragonfly of African origin, which only recently has colonised large parts of northern Europe. Patrolling males are easily recognised by their size, unmarked green thorax and blue abdomen with a black mid-dorsal stripe. Field characters Tot 66–84mm, Ab 50–61mm, Hw 45–52mm. The largest aeshnid in most of our area. Most Anax individuals with an all-blue (mature males and some older females) or green (tenerals, most females) abdomen belong to A. imperator, but a minority of A. parthenope and A. ephippiger may appear similar. A. imperator has: (1) eyes green to blue, at most tinged brown; (2) the thorax and S1 green, not (largely) brown, greyish or green-brown; (3) a thin yellow ring on S2 only in fresh individuals, which are still largely green. Moreover, males are typically blue on the thorax just in front of the Fw bases. The yellow S2 ring is typical of mature A. parthenope, which are largely brown. The frons marking differs in details from A. ephippiger, A. parthenope and A. junius. See the species text of the latter for a comparison, and the table of field characters for separating the first two (p. 171). May be confused with some Aeshna species, especially those with conspicuous blue eyes and abdomen, and a green thorax (A. affinis, A. viridis). These are smaller and have black thorax markings and a mosaic-like abdominal pattern. Hand characters Male lower appendage at least one-quarter as long as uppers, square (about as long as wide), only distal margin with dorsal denticles. Male uppers shorter than S9–10, with broadly rounded tips. Back of female occiput smooth, without tubercles. Variation The female’s abdominal colour may become obscured by dirt after oviposition, thus they may resemble A. parthenope or A. ephippiger. Behaviour Patrolling males have a slight downward curve in the abdomen. Our only Anax in which females oviposit without the male. This is similar to many other aeshnids, but they often choose floating vegetation and are therefore very exposed and more visible than most aeshnids.

/ / occiput

smooth, with no tubercles

A C M

Anax Emperors

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largely green thorax in both sexes

eyes green to blue

black pentagon at base of frons blue in front of wings in mature ?

173

immature ?

mature ?

blue abdomen with black dorsal line

square lower appendage

uniform green sides to thorax

Occurrence Range and status Common in large parts of Africa and western Eurasia. Has expanded northwards considerably in the last decades.

mature ?

Habitat Standing waters, often large and well vegetated. Flight season From March to December in north Africa, but much shorter in the north; most abundant from June to August.

Anax Emperors

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Anax junius (Drury, 1773)

Common Green Darner

174

G Anax junius male. Note the bull’s-eye marking on the frons and how the abdomen is partly brownish.

Identification General An American vagrant that superficially is most like A. imperator, as males are large dragonflies with a green thorax and blue abdomen, but is more like A. parthenope in details. Suspect Anax on the Atlantic coast should be scrutinised with extra care. Field characters Tot 68–80mm, Ab 51–57mm, Hw 45–56mm. Similar in size and general appearance to A. imperator. Distinguished from it as follows: (1) male’s mid-dorsal black stripe more tapered, largely absent on S2 but widened towards the end of the abdomen; (2) frons with unique ‘bull’seye’ marking, and not black along eyes laterally of the vertex; (3) in males the thorax in front of the Fw bases is often green (blue in A. imperator males). The very blue S2, combined with the darker abdomen end, may suggest A. parthenope. Moreover, in A. junius the abdomen is often partly brown (especially terminally and in females) and becomes dark purplish in cooler conditions; the blue S2 then stands out strongly. A. parthenope has a brownish (not green) thorax and S1. Hand characters Male lower appendage very short, less than one-fifth as long as uppers. The

uppers are long (as long as S9–10 combined) and truncated, with a large spine on their outer corner. Back of female occiput bears two tubercles, like A. parthenope but unlike A. imperator. Behaviour Oviposits in tandem, like A. parthenope but unlike A. imperator.

Occurrence Range and status Common and widespread in North and Central America. Vagrant in Europe. At least six individuals were recorded on the Isles of Scilly and in Cornwall in September 1998, and one male was caught on the French coast, near Nantes, in September 2003. The British records coincided with an Atlantic depression with strong westerly winds. Habitat Standing, marshy waters, including temporary or slightly brackish ones. Flight season All year in the south of its American range, but from April to October in the north, which is colonised by migrants from the south each year. These migrants are often the earliest anisopterans seen there. Migration southwards starts in late summer.

Anax Emperors

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bull’s-eye marking on frons

175

?

blue abdomen with black dorsal line reduced on S2 but relatively broad towards tip

very short lower appendage

long, spined upper appendages

/

Anax Emperors

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Anax parthenope (Selys, 1839)

Lesser Emperor

Identification

176

General A large and rather dull aeshnid. Its drab colouring makes the greenish eyes and blue abdominal saddle particularly striking. Typical individuals can be confused only with A. ephippiger, but occasional ones with a largely blue abdomen can closely resemble A. imperator. Field characters Tot 62–75mm, Ab 46–53mm, Hw 44–51mm. Slightly smaller than A. imperator. Typical specimens are most likely to be confused with the smaller and more slender A. ephippiger, while those with a bluish, instead of brown, abdomen may be confused with A. imperator (see table p. 171 and those species’ texts for distinguishing features). In A. imperator females the wings are clear, or with a brownish tint from the triangles to the tips, while the wings of older A. parthenope females often are tinged brown only between the Pt and the node. Note that the pattern on the frons is different from all other Anax species. Hand characters Male lower appendage is less than one-fifth the length of the uppers, and shorter than wide (view from above), with numerous denticles on upperside; uppers have truncated tips bearing a small spine on the outer corner. Unlike A. ephippiger and A. imperator, there are two tubercles behind the female occiput.

C M

Variation In fresh specimens, the blue parts of the abdomen are grass green. Behaviour The male’s abdomen is straighter in flight than in A. imperator. Unlike that species, the male usually accompanies the female in tandem during oviposition.

Occurrence Range and status Common in the Mediterranean countries; scarce further north, although locally abundant. Has expanded its range since the 1990s and, although it is not expanding as quickly as A. imperator, northwards vagrancy is more often observed than in A. ephippiger. Range extends to Japan and China, and into Sahara. Habitat Standing, often larger, waters. Flight season In the Mediterranean from March to November, but northern records mostly from June to August.

 Anax parthenope mature male.

Anax Emperors

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177

wings often tinted between node and pterostigma in mature individuals

? abdomen mostly brown, making blue saddle stand out

very short lower appendage

small spine at tip of upper appendages

basal ring on S2 typically yellow

/ rather blue individual

mature ?

distinct blue ‘saddle’ wrapped around waist

/ occiput

two tubercles on occiput

Anax Emperors

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Anax ephippiger (Burmeister, 1839)

Vagrant Emperor F Anax ephippiger pair mating.

178

Identification General These sandy-coloured emperors are strongly migratory, occurring mostly in the dry regions of Africa and Asia and only irregularly straying beyond the Mediterranean. Their scientific name refers to the characteristic blue saddle at the abdomen base on the male. Often placed in the genus Hemianax. Field characters Tot 61–70mm, Ab 43–56mm, Hw 43–48mm. Slightly smaller than other Anax species, with a broad globular head and a shorter, more slender abdomen. Most males have a largely dark abdomen with a blue ‘saddle’ on S2 (and sometimes part of S3). The saddle is restricted to the upper side of the abdomen, making it harder

C M

to see in flight than in A. parthenope, where it is ‘wrapped around’ the abdomen. S3–10 are brown with a black dorsal band; this band is wider on S8–10, enclosing a pair of large pale brown spots on each segment. The blue saddle is less intense in young males and also in females, where it is crossed by the dorsal bands. The eyes and thorax are brown, often with their lower halves characteristically yellow-green. Neither the frons nor eyes show any trace of blue, whereas in A. parthenope and A. imperator there is a blue bar on the frons and the eyes are typically quite uniformly green, often tinged blue. The two bold black bars on the frons of A. ephippiger are distinctive. Hand characters The only Anax with one ridge only (not two) on each side of the abdomen. The only Anax with a broad cubital field in Hw with three cell rows, i.e. with several cells enclosed between the field’s two cell rows. Male upper appendages with sharply pointed, tapered tips. Lower at least one-quarter the length of the uppers, triangular (view from above), upper surface covered with denticles. Back of female occiput is smooth, without tubercles. Female appendages broad, similar to male’s uppers. Variation Blue ‘saddle’ may be absent in some females. Behaviour Usually oviposits in tandem. Mainly diurnal, but more often active at dusk than other Anax species; sometimes attracted to light.

Anax Emperors

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brown eyes

two black bars on frons

179

this field differs in shape from all other Anax, with 3 irregular rows of cells, rather than 2 regular rows

? slender brown abdomen

pale brown paired spots on S8–10 triangular lower appendage with numerous denticles

blue ‘saddle’ less distinct on immatures and females

pointed upper appendages

/

basal ring to S1 brown

lower part of eyes and thorax greenish yellow

mature ?

blue ‘saddle’ mainly restricted to upperside

Occurrence Range and status Occurs mainly in strongly seasonal parts of Africa and south-west Asia, wandering to and fro to breed after rains. Appears to breed annually in the Mediterranean basin, but present further north only in some years, when invasions can even reach Iceland. Subsequent reproduction (sometimes in great numbers) has recently occurred as far north as the Netherlands.

broad, pointed appendages

Wanderers have also reached Brazil, Japan and Thailand; has recently even bred on some Caribbean islands. Habitat Shallow, warm (often temporary) pools and lakes. Flight season Throughout the year around the Mediterranean. Mostly from July to September in the north, but can be recorded in all months.

Anax Emperors

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Anax immaculifrons Rambur, 1842 Magnificent Emperor Identification

180

General The largest and one of the most exotic European dragonflies, with its bold black-andyellow abdominal rings and the male’s peculiar blue-washed head, thorax and abdomen base. A predominantly southern Asian species, with its western outposts in streams flowing into the eastern Mediterranean Sea. Field characters Tot 80–86mm, Ab 55–60mm, Hw 54–60mm. Our largest dragonfly, and unmistakable due to its size and black abdomen with pale anterior halves of S2–8, creating seven rings. Thorax pale, with two broad black bands on each side. Frons without dark markings. Eyes, frons and pale markings are (greenish) yellow in females and young males. Males become pale bluish from frons to abdomen base, with bright blue eyes. Males often have strongly brown-stained wings; females’ wings are at most faintly tinted. Overall appearance suggests a Cordulegaster, some of which also have blue eyes.

Hand characters Male’s lower appendage long (about half of uppers) and triangular in shape (view from above). Variation In eastern Asia, males lack the blue tinge and develop a brick-red abdomen: possibly a separate (sub-) species is involved. Behaviour Males patrol over long stretches of streams at great speed. Uniquely for an aeshnid, the male guards the female during oviposition by hovering over her.

Occurrence Range and status Widespread across southern Asia to China, reaching its western limit on the Aegean coast. Found in south Turkey, Cyprus and the Greek islands of Rhodes, Kárpathos and Ikaria. Habitat Unlike other Anax species, avoids still water, inhabiting larger (often partly shaded) rocky streams. Flight season In Greece and Turkey, seen from May to September.

 Anax immaculifrons pair mating.

Anax Emperors

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unmarked frons blue eyes, frons and thorax in mature males

181

?

black abdomen with pale rings

long, triangular lower appendage

banded thorax

/

/ like large Cordulegaster

Anax Emperors

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Brachytron Evans, 1845 Hairy Hawkers Brachytron pratense (Müller, 1764) Identification

182

General A small hawker that is often confused with the smaller Aeshna species. It has an earlier flight season, and its hairy body and stout abdomen create the impression of a more compact insect. Males are often seen hawking low down, closely following marshy margins, in late spring and early summer. In the hand, numerous details of the venation, body shape and markings reveal that Brachytron pratense – the only species of its genus – is very distinct from other aeshnids. Field characters Tot 54–63mm, Ab 37–46mm, Hw 34–37mm. Shorter and more stubby than Aeshna mixta. Thorax noticeably downy, with unique markings: sides green, interrupted by two complete black lines (not just one). Unlike Aeshna, in males the abdomen is not waisted at the base, but is cylindrical, as in the females. The abdomen is black, with ‘inlaid’ pale blue spots in the male and a diagnostic central yellow (or green) dot on S1. The terminal paired spots on the segments are more elongated than in Aeshna. The markings are greenish yellow in females and immatures. Compare Aeshna cyanea, and the similar-sized A. mixta and A. affinis. Hand characters Details of the wings make it distinct from Aeshna: Pt are longer and thinner; largely just a single row of cells before Rspl; usually has only two cross-veins between base

Hairy Hawker Hairy Dragonfly

and triangles; anal triangle is short and anal angle rounded. The male lower appendage is about a third the length of the upper appendages, and distinctly notched at the tip (about half as long as uppers and more pointed in Aeshna). Behaviour Males make low and very inquisitive patrols, flying in and out of clearings among the vegetation.

Occurrence Range and status Widespread west of the Urals, but generally localised, although often common where found.

G Brachytron pratense pair mating.

Brachytron Hairy Hawkers

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long thin pterostigma

single row of cells before Rspl

central dot on S1

183 abdomen not waisted in ?

rounded edge to ? hindwing

mature ?

noticeably hairy body

pairs of elongated blue spots on abdomen central dot on S1 relatively short lower appendage with notched tip

/

pairs of elongated yellow spots on abdomen

green side to thorax with two complete black lines

mature ?

Habitat Standing or slow-flowing waters with rich riparian and aquatic vegetation, such as reedy canals, marshes, oxbows and coastal grazing marshes.

Flight season Late March to early August, with the emphasis on May and June in most of its range, well ahead of most Aeshna, which peak in late summer.  K-D B Dijkstra

Brachytron Hairy Hawkers

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Boyeria McLachlan, 1896 Identification

184

Diagnosis Coloration rather dull and blotchy, like military camouflage. Most males and some females have dark-tipped wings, especially the Hw. Fields before Mspl and Rspl with two rows of cells, median space (between arculus and base) of all wings with two to four cross-veins, and vein IR3 not forked. Pt moderately long, above four to seven cells. Separation from other genera In the field, habitat, behaviour and colour are important, and in the hand venation rules out any confusion. Caliaeschna has similar habits, but does not co-occur and is much smaller, with more contrasting markings and a short Pt. Most Aeshna species are more strongly marked, all lack median cross-veins and Mspl and Rspl subtend three or more irregular rows. Both these genera have IR3 forked and clear wing tips.

Spectres Spotted Darners (NA)

Separation of the species A genus with isolated relict populations, with two species in eastern North America and three in eastern Asia. Two species are endemic to our area. Behaviour Males make long and furtive patrols, slowly flying low over the water and closely trailing its edge. They keep to shade and carefully inspect dark corners, e.g. under tree roots, often with hovering pauses. They avoid sunlight, becoming more active towards the late afternoon, and aggregate at dusk, hunting in fast, zigzagging flight over clearings. They remain on the wing until nightfall and may be attracted to light thereafter. Unlike B. irene, B. cretensis males do not appear to establish and defend territories.  K-D B Dijkstra

Boyeria irene (Fonscolombe, 1838)

Western Spectre Dusk Hawker

Identification

Occurrence

General Known as the ‘peaceful hawker’ in French and the ‘ghost dragonfly’ and ‘twilight dragonfly’ in German and Dutch, this aeshnid is characterised by its slow, inconspicuous, shade-hugging streamside patrols. Nonetheless, at dusk it may be seen more in the open, whizzing wildly to and fro in pursuit of prey. Aside from behaviour, it can be identified by generic characters in most of its range and is unlikely to be mistaken (see above). Field characters Tot 63–71mm, Ab 44–48mm, Hw 39–45mm. Size between Aeshna mixta and A. cyanea, near A. isoceles. Greyish brown, marked with rather weakly contrasting greyish or bluish pale green blotches. Pattern is characteristic, including broadly green bases of S2–9 and allgreen upperside of S9–10, forming five to seven rings and a ‘tail-light’. Frons is marked above with an indistinct dark blotch. Eyes are green. Hand characters See genus text. Variation Extent of wing markings varies. The female has two forms: the typical form (f. typica) has very long appendages (about 6mm, three times the length of S10), while they are only about 2mm (as long as S10) in f. brachycerca. Despite its name, the typical form is scarcer, constituting at most half of the females at a site.

Range and status Endemic to the western Mediterranean, where common on most streams and rivers. A highly isolated but strong population occurs at the Örtze river in northern Germany. Habitat Streams and rivers with shaded borders. Locally inhabits shores of large lakes, which is the only habitat in Switzerland. Flight season From the end of June to late September; most abundant in July and August.

B. cretensis (EN)

Boyeria Spectres

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B. cre

B. irene

green eyes

IR3 not forked

B. cretensis dark wing tips

two rows of cells before Mspl and Rspl

cross-veins in median space

?

immature /

markings darker and more extensive

185

?

camouflagelike markings

B. irene

/ frons more pointed B. cretensis green ‘tail-light’

f. brachycerca

f. typica

Boyeria cretensis Peters, 1991 Identification General This endemic of Crete is very similar to its western counterpart, B. irene. It appears to be an ancient relict: Caliaeschna microstigma replaces Boyeria on the surrounding mainland. First reported in 1850, its status as distinct from B. irene was realised only 141 years later. Field characters Tot 69–71mm, Ab 45–49mm, Hw 44–47mm. On average, slightly larger than B. irene. Mature male has green eyes, contrasting with its dark, dull body. The dark markings are more extensive than in B. irene, leaving only small greyish-yellow (rather than green) spots. S2–8 in particular are much darker basally, the species thus lacking the ringed appearance of B. irene. Teneral specimens are considerably paler, with broad, pale bases to S2–8. Confusion is unlikely within its restricted range, but Caliaeschna microstigma could potentially occur as a vagrant from the mainland. The nearest sites for this species from Crete are about 100km north-west and 150km north-east. Hand characters Differs from B. irene in these

Cretan Spectre

minor characters: (1) Pt of Hw 4–5mm, rather than less than 4mm; (2) wings more densely veined on average; (3) profile of frons (view from side) more pointed; (4) dorsal ridge of male’s upper appendages not raised before end (view from side, not illustrated). Variation The female form with long appendages, known in B. irene, has never been observed in this species.

Occurrence Range and status Confined to Crete (blue on map opposite), where scarce and difficult to observe; only a fraction of the streams on the island are permanent and thus suitable. The species, whose entire population is restricted to 15 river systems, is seriously threatened by impoundment, piping and pollution of streams. Habitat Rocky and partly shaded streams. Flight season Most adults have been recorded in July and August, but probably on the wing from late May into September.

Boyeria Spectres

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Caliaeschna 

Eastern Spectres

Selys, 1883 Caliaeschna microstigma (Schneider, 1845) Eastern Spectre F Caliaeschna microstigma male.

186

Identification General A small, sleek aeshnid, named for its unusually short Pt. Shares its cryptic, crepuscular habits and stream habitat with Boyeria, but is not known to overlap, is considerably smaller and is more contrastingly marked. Field characters Tot 50–60mm, Ab 39–47mm, Hw 35–41mm. The smallest aeshnid in our area. The Pt is very dark and short (at most 2mm, and subtended by only about two cells), rather more like that of some libellulids than that of an aeshnid. Mature males are brown-black, with crisp, pale blue markings, usually with ‘7’-shaped antehumeral stripes. The dorsal blue spots on S1–7 are narrow, but on S8–10 are broader and even (partly) fused, creating a distinct ‘tail-light’. Females are browner, with more yellowish markings. The frons is whitish, with a contrasting black bar on the crest. Unlikely to be confused within its range. Other small aeshnids (e.g. Aeshna mixta, Brachytron pratense) differ by habitat and behaviour.

Hand characters Fields before Mspl and Rspl with one row of cells, like most Brachytron but unlike Boyeria (usually two rows) and Aeshna (three or more irregular rows). Median space (between arculus and base) of all wings with cross-veins, a character shared only with Boyeria. Vein IR3 is forked, as in Aeshna and Brachytron (not forked in Anax and Boyeria). The female appendages are very short and stubby, and the sheath enclosing the ovipositor is longer and straighter than in other genera, appearing as a spike reaching beyond S10. Variation Eastern populations can be considerably darker, with the antehumeral stripes reduced to a tiny dot. Behaviour Males behave similarly to Boyeria. They have a low, slow patrolling flight, closely following the streamside. They avoid sunlight and inspect every nook and cranny for the presence of females. Most activity takes place in the latter part of the day, especially late afternoon and dusk. Groups hunt over streams and along trees at twilight with fast, erratic flight.

Caliaeschna Eastern Spectres

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short dark pterostigma usually hooked antehumeral stripes

single row of cells before Rspl IR3 forked

187 cross-veins in median space

single row of cells before Mspl

?

antehumeral stripes may be reduced and dot-like

blue marking partially fused on S9–10 to form a ‘tail-light’



/

black bar on whitish frons

? / abdomen tip appendages minute

ovipositor sheath long and pointed

Occurrence Range and status Ranges to the Caucasus region, Afghanistan and northern Israel. Notably absent from Crete, where it is replaced by Boyeria cretensis. Common in most of its range. Habitat Strictly in streams, which are often fastflowing and shaded. Larvae live among submerged moss, leaf litter or roots, or under stones when other substrates are absent. Flight season From mid-May to August, occasionally to October.  K-D B Dijkstra

Caliaeschna Eastern Spectres

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Gomphidae

Gomphus Leach, 1815 and Stylurus Needham, 1897 Identification

188

Diagnosis Medium-sized dragonflies with a yellow to greenish body, marked with black. The almost uninterrupted yellow line running from S1 to S7, or even S10, is distinctive. The Hw lacks an anal loop, thus an uninterrupted perpendicular vein connects the subtriangle and the hind margin. The abdomen is thickened in most species to a club shape, but lacks leaf-like lateral flaps. The male’s upper appendages are rather short (about equal to S10) and clearly diverge. The lower is only slightly shorter, and its branches are largely eclipsed by the uppers when viewed from above. The short, splayed, eclipsing appendages are unique in our area. Separation from other genera All our gomphids, characterised by clearly separated eyes, are rather similar in general appearance and size, except for the large Lindenia and small Paragomphus. Males of both these genera bear abdominal flaps and have very long, parallel upper appendages. Ophiogomphus and Onychogomphus have an anal loop, and incurved upper appendages that do not eclipse the parallel branches of the lower appendage. Moreover, the Onychogomphus male appendages are long and pincer-like, and the abdomen appears ringed or spotted, rather than striped. In the field, confusion with Ophiogomphus is most likely, but mature adults are unmistakable because of their bright green head and thorax, as well as the yellow upper appendages. When seen poorly, particularly in flight, females and immatures of some libellulids, particularly Orthetrum cancellatum, resemble Gomphus and Stylurus. The eyes in these touch each other. Separation of the species Few of our genera seem as uniform as Gomphus and Stylurus

Clubtails

combined, and all nine species may be confused. Stylurus is a mainly North American genus with only a few Eurasian species. Until recently the latter were usually placed in Gomphus, but their longsuspected relationship has now been confirmed by genetic evidence. They also differ subtly in markings, the more slender and acute posterior hamules, and the distinctly elongated larvae, which have reduced hooks on the legs, an adaptation for burrowing in soft sediments. The separation of three species (G. schneiderii, G. lucasii, S. ubadschii) from their more widespread counterparts (G. vulgatissimus, G. simillimus, S. flavipes) is still somewhat questionable, although the species in each pair are geographically (largely) segregated. A simple table, based on markings of both sexes and ranges, is provided, but for positive identification careful comparison is required, preferably in the hand. The male’s appendages and female’s vulvar scales may then be useful, but are fairly uniform. The shape of the male’s posterior hamules is underappreciated as a character, but the hamules are almost as easily examined with a hand lens as the appendages, and may be more informative! Behaviour Relatively shy; the large numbers of exuviae found on riverbanks may compare poorly to the occasional adult seen at the same site. After emergence, adults mature in surrounding rough terrain, such as overgrown fields, borders and fallow land, and these are often better places to search than the waterside. Breeding males perch on twigs or rocks at the water’s edge, waiting for females, but at larger rivers may also fly for long periods in midstream (particularly S. flavipes but also G. vulgatissimus), or search the edges (G. pulchellus): it may be useful search for individuals in the middle of a river with binoculars. Females are especially shy, dashing out to open water, often in the centre of a stream, and depositing many eggs in only one or several dips.  F Suhling and O Müller

 Gomphus pulchellus pair mating. Note the black-and-yellow body typical of all Gomphus and Stylurus species.

Gomphus Clubtails

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Comparison of Gomphus and Stylurus species Thorax stripes: Antehumeral stripe: yellow band between second and third black line counted from head Interpleural stripe: fourth black line, just before metastigma Metapleural stripe: fifth black line, just behind metastigma S8–9

Usually black dorsally

Yellow lines on legs

Antehumeral stripe/second black line

Usually none

Narrower to equal

Femora

Narrower

Diagnostic Approximate features of range thorax markings (indicated on artwork below)

Interpleural stripes not deadended just above metastigmas, but complete from base of Fw to middle leg

G. vulgatissimus

G. pulchellus

G. graslinii

189

Turkey, S Balkans G. schneiderii

Equal

Wider

G. vulgatissimus

Europe

None

With yellow central Femora stripes and tibiae

Species

France, Iberian peninsula,

G. graslinii

Turkey

G. davidi

SW Europe, Morocco

G. simillimus

Algeria, Tunisia

G. lucasii

W Europe

G. pulchellus

S. flavipes

Europe Metapleural stripes not forked at level of metastigmas; antehumeral Turkey stripes not limited ventrally by black and thus extend to middle leg

S. flavipes

G. davidi

S. ubadschii

S. ubadschii

G. lucasii

G. simillimus

Gomphus Clubtails

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Gomphus vulgatissimus (Linnaeus, 1758) Common Clubtail

Club-tailed Dragonfly

Identification

190

General The commonest and most widespread Gomphus in much of our area. It is darker than all other species, with a greener ground colour and more extensive black markings. Field characters Tot 45–50mm, Ab 33–37mm, Hw 28–33mm. More robust than congeners, with a thicker clubbed abdomen. Darker than other Gomphus species, relevant characters are: (1) legs are usually completely black; (2) S8–10 are usually dorsally black, without a central yellow stripe; (3) the yellow ground colour becomes distinctly green in older males, whereas other species remain (greenish) yellow; (4) eyes (blue-)green when mature. Another much-used, but not always reliable, feature is that the pale antehumeral stripes are narrower than both black stripes bordering them; these black stripes are often (almost) in contact. Males of G. graslinii, G. pulchellus, G. simillimus and S. flavipes, all of which can occur with this species, have blue eyes when mature, and possess yellow stripes on femurs and upperside S8–10. G. vulgatissimus is most similar to the generally blue-eyed G. schneiderii, which overlaps in range in the southern Balkans (see that species for details).

G Gomphus vulgatissimus male. Note that the yellow thorax is tinged green, as is typical for older males. Hand characters Male’s anal triangles typically contain four or more cells, but usually only three in other Gomphus and Stylurus species. Variation Hybridises with G. schneiderii in a broad contact zone in the southern Balkans, often making identification there impossible. Behaviour Males perch on bankside plants and rocks. Both sexes usually stay near the water, but may retreat to the canopy of nearby trees, making them hard to find.

Occurrence Range and status The most frequent Gomphus in most of northern and central Europe, ranging east to the the West Siberian Plain. Habitat Middle and lower sections of rivers and streams. Favours a calm flow and sandy bottom (e.g. lowland streams and small rivers), avoiding small, fast-flowing, rocky-bottomed waters. Locally abundant in reservoirs, gravel pits and large lakes. The larvae burrow in fine sand, preferably covered with detritus. Flight season From April to June in the south of range, from June to August in the north. Typically emerges in great numbers in a short period in spring, followed by a short flight season, e.g. in central Europe emergence peaks in early May.

Gomphus Clubtails

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greenish eyes

yellow coloration turns green in older individuals

191

anal triangle 4 or more cells (5 here)

mature ?

no central yellow stripes on upperside of S8–10

immature ?

entirely black legs

sharply pointed posterior hamule

? secondary genitalia

? appendages (side view)

/ vulvar scale /

Gomphus Clubtails

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Gomphus schneiderii Selys, 1850

Turkish Clubtail

blue eyes black legs

192

? appendages (side view) anal triangle usually with 4 or more cells, as in G. vulgatissimus

/

mature ?

at most narrow central spots on S8–9

/ vulvar scale

Identification General Replaces G. vulgatissimus in Turkey and the southern Balkans. It is yellower and has bluer eyes, therefore appearing more like other Gomphus species, but both it and G. vulgatissimus are variable and may be distinguished only by the male appendages and female’s vulvar scale, although these too barely differ. This and the extensive hybridisation in a zone from Montenegro to Thrace suggest that, if no evidence of their

genetic isolation is forthcoming, the two must be considered synonymous. Field characters Tot 40–48mm, Ab 30–34mm, Hw 29–31mm. Tends to be smaller and more slender than G. vulgatissimus, with eyes of mature males blue rather than greenish. Otherwise, very similar to that species. While the black lines on the thorax may be thinner, with the yellow antehumeral stripes about as wide as the black humeral stripes behind them, this is not a reliable character. Yellow markings in females in particular can be more extensive, e.g. with yellow-streaked legs, rather like other Gomphus species, e.g. G. davidi and S. ubadschii, with which it occurs in Turkey (see these species). Hand characters All structures are very similar to those of G. vulgatissimus. Male appendages are shaped almost like those of G. simillimus.

Occurrence Range and status Ranges from the southern Balkans to Iran, Lebanon and the Caucasus region. Fairly widespread in Turkey and southern Greece. Habitat Probably resembles that of G. vulgatissimus and G. simillimus, breeding in rivers and streams but also in tiny runnels and large lakes. Flight season From April to July; probably most abundant in May and June.

Gomphus Clubtails

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Gomphus graslinii Rambur, 1842

Pronged Clubtail legs black with yellow stripes, tibiae often all black

large lateral tooth

? appendages (side view)

193 mature ?, rather green individual

/ blunt tip to posterior hamule

pale antehumeral stripe narrower than adjoining black stripes

? secondary genitalia

‘forked’ male appendages

yellow posterior border to S9 forms foot of goblet-shaped marking

/ vulvar scale

Identification General A bright yellow clubtail with such prominently toothed male appendages that it can be identified by this feature without capture. The species occurs only in France, Spain and Portugal. Field characters Tot 47–50mm, Ab 33–38mm, Hw 27–30mm. Fairly weakly club-tailed; S8–9 are only slightly widened. A richly coloured Gomphus with a bright yellow body, sharp black markings and bright blue eyes. Blacker than the co-occurring G. simillimus and G. pulchellus, but yellower than G. vulgatissimus (see these species). Note the following characters: (1) yellow antehumeral stripes are much thinner than the black stripes flanking them, the anterior black stripes especially broad, and their dorsal ends often connecting with the black markings anterior and/or posterior of them (in G. simillimus the antehumerals and their flanking stripes are all about equally thick, and the anterior ones are dorsally unconnected); (2) S9 has a yellow posterior border, appearing like the foot of a goblet-shaped yellow central marking (G. vulgatissimus is usually all black dorsally, and in G. simillimus the posterior border of S9 is black, although sometimes the yellow central marking reaches it); (3) the legs are black, typically with yellow lines only on the femora (G. simillimus

always also has lines on the tibiae; G. vulgatissimus typically has all-black legs). Hand characters The male’s upper appendages each have a large lateral tooth, making them appear forked. This unique character is often visible through binoculars. The posterior hamule is also distinctive, with a prominent, but blunt, point. This point is very sharp in G. vulgatissimus and G. simillimus, but is small and concealed in G. pulchellus.

NT HD II+IV

Gomphus Clubtails

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Occurrence Range and status Endemic to south-western France (where locally common) and the Iberian peninsula (widespread but rather scarce). Due to its restricted range and the frequent alteration of river habitats there, G. graslinii is ranked as Near Threatened globally. Habitat Larger rivers in hilly terrain, favouring slow-flowing sections; also large impoundments.

194

The larvae burrow in fine sand, usually at sites with leaf detritus. On the Iberian peninsula, larvae were found in abundance in drying-up river sections. Flight season In France, from mid-June to the end of August; perhaps slightly earlier in the Iberian peninsula. All co-occurring Gomphus species have an earlier flight season (although S. flavipes is also later) and thus G. graslinii is rarely seen with them in summer.

Gomphus simillimus Selys, 1840

Yellow Clubtail

Identification General A bright yellow Gomphus endemic to north-west Africa and south-west Europe, where it overlaps with up to three congeners and S. flavipes. Rather nondescript, and therefore most easily identified by the exclusion of others, although the female’s large vulvar scale is unique. Field characters Tot 45–50mm, Ab 33–36mm, Hw 29–33mm. A very average Gomphus in size, shape and colour. Co-occurring Gomphus and Stylurus species are excluded as follows (see each

G. lucasii (VU)

 Gomphus simillimus mature male. Note the central line on S8–9, the clubbed tail and the antehumeral stripe which is equal to the black lines each side of it.

Gomphus Clubtails

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legs black with yellow lines

195 /

mature ?

metapleural stripe forked (compare G. flavipes)

long, bluntly tipped vulvar scale

/ vulvar scale

sharply pointed posterior hamule

? appendages

? secondary

(side view)

species for details): (1) G. vulgatissimus lacks yellow lines on legs and central line on S8–9, and mature males are greenish with green (not blue) eyes; (2) G. pulchellus is paler and duller, scarcely clubtailed, and the black line in front of the metastigma is not dead-ended but extends towards Fw base; (3) S. flavipes has a pair of pale black-encircled stripes on the front of the thorax and the black metapleural stripes are not forked; (4) G. graslinii has forked male appendages and thicker black markings on the thorax; (5) G. lucasii may overlap in Morocco or Algeria, and is almost identical to G. simillimus and can be distinguished with certainty only by hand characters (see that species). Hand characters Male’s appendages and posterior hamule are almost identical to those of G. vulgatissimus, but female’s vulvar scale is unique within the genus, being longer (about half of S9) and with a less acute apical notch. Behaviour Males usually perch on twigs and plants, less often on rocks. Variation As in all gomphids, black markings are

genitalia

reduced in hot, arid areas. The paler Moroccan populations have been considered as the subspecies maroccanus, but this distinction is probably not very useful.

Occurrence Range and status Rather common within a range that is largely restricted to France, Spain, Portugal and Morocco. Mainly a vagrant in adjacent countries, but a large isolated population occurs on the upper Rhine. Shows a general decline and is classified as Near Threatened globally. Habitat Almost all types of running water, from large, slow rivers and their side channels to faster-flowing mountain streams, where the larvae inhabit sections with a reduced current (margins, deeper reaches). The larvae burrow in fine sand, often at sites with detritus. Flight season From early May to late July in France, but earlier in the south, e.g. emergence starts in early April in Morocco. The Rhine population, in contrast, emerges as late as July.

Gomphus Clubtails

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Gomphus lucasii Selys, 1849

Algerian Clubtail narrow black stripes on thorax

yellow margin to vertex also present in pale Moroccan G. simillimus

? appendages (side view)

196

mature ?

/ vulvar scale

vulvar scale shorter than in G. simillimus

uppers less diverging and more tapering than in G. simillimus, thus eclipsing lower appendage less perfectly

? secondary genitalia

Identification General Replaces G. simillimus in Tunisia and most of Algeria, where it is the only species of its genus. Field characters Tot 44–48mm, Ab 31–40mm, Hw 28–37mm. Strongly resembles G. simillimus but may be even paler than individuals of that species from Morocco. S8–9 often distinctly brighter yellow than basal half of abdomen and thorax, giving the impression of a yellow ‘tail-light’. Black lines on the thorax are especially thin; the yellow antehumeral stripes are bordered by two much finer black lines, of which the anterior one is usually not connected to the black flanking the mid-dorsal crest. There are mid-dorsal yellow dots on S8 and S9, which may just reach the posterior segment margins. Legs are black, with yellow lines along the segments. Because both species are variable, identification must be based on hand characters. Hand characters Male’s upper appendages are less diverging and with more tapering tips than G. simillimus, and they eclipse the lower appendage less perfectly (view from above). The female’s

G Gomphus lucasii male. vulvar scale is shorter. Unlike G. simillimus, back of female’s vertex is raised and produced over ocelli.

Occurrence Range and status Restricted to waters flowing off the Atlas to the Mediterranean Sea in Tunisia and Algeria. Not reported with certainty from Morocco, where only G. simillimus occurs. The two may meet in the border region with Algeria (see map p. 194). Habitat Low-lying rivers and streams. Flight season Emerges at least from late March to May; on the wing until the end of June.

Gomphus Clubtails

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Gomphus davidi Selys, 1887

Levant Clubtail

legs extensively marked yellow

197 / mature ?

? appendages (side view)

/ vulvar scale broad stripe on S8–9

swollen tip to posterior hamule

? secondary genitalia

Identification General Another ordinary-looking Gomphus, which resembles G. simillimus but occurs far from that species on the eastern Mediterranean seaboard. A rather indistinct species, but can be identified instantly by inspecting the male’s ‘roundheaded’ posterior hamule.

Field characters Tot 46–55mm, Ab 35–40mm, Hw 31–35mm. Slightly larger than the co-occurring G. schneiderii. Yellower than that species: at least all femora have much yellow and S8–9 have a broad and almost uninterrupted yellow central stripe, which often leaves the posterior border of S9 largely yellow. S. ubadschii, which also occurs in the same area, also has a rather yellow S8–9, but the segments (including S7) are wider and the thorax is marked very differently (see under S. flavipes). Hand characters The shape of the male appendages and female vulvar scale may be useful characters, although they are rather average for the genus. The male’s posterior hamule is unique, with a swollen rounded head.

Occurrence Range and status Endemic to the Near East, from around Adana in southern Turkey (where it is common) to the Jordan Valley. Habitat Slow-flowing rivers and associated waters; also nearly stagnant ditches. Flight season Recorded in Turkey in May and June, and as early as March elsewhere.

Gomphus Clubtails

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Gomphus pulchellus Selys, 1840

Western Clubtail

198 /

mature ? posterior hamule appears to lack distinct point

lines on thorax very thin

? secondary genitalia

interpleural stripe complete and wavy

S8–9 not strongly club-shaped

/ vulvar scale

? appendages (side view)

Identification General A sleek, rather faded clubtail that usually inhabits calmer (frequently even still) waters than its congeners. Field characters Tot 47–50mm, Ab 34–38mm, Hw 27–31mm. Not as club-tailed as other Gomphus species; S8–9 scarcely expanded. Ground colour pale yellow, more sallow than other species, and may be tinged greenish. The eyes are rather pale blue. The black thorax lines are thinner than in co-occurring Gomphus species and have a unique

feature: the interpleural stripes form complete curvy black lines between the Fw bases and midlegs. G. graslinii, G. simillimus, G. vulgatissimus or S. flavipes may occur with G. pulchellus, but are brighter in colour, with thicker thorax markings and a more club-shaped abdomen. The interpleural stripes always stop just above the metastigma. The thin thorax lines resemble Ophiogomphus cecilia, which has a bright green thorax. Hand characters Male’s upper appendages are somewhat angled laterally (view from above). Unlike overlapping gomphid species, the point of the male’s posterior hamule is small and bent inwards, and therefore not conspicuous from the side. Behaviour Males search along the shoreline for mates, flying in a wave-like flight; constantly varying their height but staying just a few centimetres above the water.

Occurrence Range and status Near endemic to south-western Europe and generally quite common. Expanded north and east of the Rhine, approximately to the Elbe, during the 20th century, probably in reaction to the creation of artificial habitats. A southeastern outpost was recently discovered in Lake Skadar in Montenegro and Albania.

Gomphus Clubtails

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Habitat All kinds of slow-flowing and standing waters; avoids strong currents. Most common in slow lowland rivers and their stagnant side channels, on residual floodplain pools and impoundments. In the north, usually in gravel and sand pits, recreational lakes, oxbow lakes, fish ponds and canals, but also sluggish rivers. The larvae occupy sandy patches, which may be covered with coarse detritus.

Flight season Rather short and early: emerges from early April or even late March in southern Iberian peninsula, but at the northern end of its range normally from May. Here, individuals may be seen as late as August, but most active in late May and June.

Stylurus flavipes (Charpentier, 1825)

River Clubtail

199

Yellow-legged Clubtail completely yellow collar and middorsal crest enclosed by black to form a distinct ‘T’

yellow antehumeral stripe wider than adjoining black lines and extends to legs

black lines on front of thorax enclose two yellow ovals

legs noticeably yellow

metapleural stripe not forked

G. simillimus (for comparison)

/

mature ?

black at base of legs interrupts yellow antehumeral

short vulvar scale

metapleural stripe forked

/ vulvar scale

? appendages (side view)

Identification General Prefers very large rivers and may easily be overlooked. Populations can best be found by searching for exuviae on riverbanks. Replaced in Turkey by the very similar S. ubadschii, which is often treated as a subspecies of S. flavipes. Field characters Tot 50–55mm, Ab 37–42mm, Hw 30–35mm. Slightly larger than most Gomphus species, with more slender abdomen and strongly widened S8–9. Legs are relatively yellow, often appearing yellow with black lines rather than

short, pointed posterior hamule

? secondary genitalia

the reverse. Most easily identified by the unique configuration of the thorax markings, from front to back: (1) the mid-dorsal crest is not marked with black, but forms a completely yellow line that is connected anteriorly to the transverse yellow ‘collar’, together forming a yellow ‘T’; (2) the flanking black lines lie close to the next set of black lines and typically connect with these ventrally and dorsally, thus enclosing two long yellow ovals and appearing like a pair of eyes (view from above); (3) the antehumeral stripes (the next pair of yellow

Gomphus Clubtails

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200

G Stylurus flavipes male. Note the two long yellow ovals enclosed by black on the front of the thorax. stripes) are very broad, wider than the black lines behind them, and continue uninterrupted to the middle legs; (4) the black metapleural stripes are not forked ventrally, just behind the metastigmas. G. simillimus is similar in shape and yellowness, but overlaps in range only locally in France and southwestern Germany; while males have blue eyes, the female’s eyes are greenish. Hand characters The male’s upper appendages have relatively slender tapering tips, although they are slightly angled laterally (view from above). The female’s vulvar scale is shorter than in congeners, about one-quarter of S9. The male’s posterior hamule is small, tapering and acute in comparison. Behaviour Adults are most easily found among bushes and over rough meadows near rivers or at emergence on the riverbanks. Mature males are seldom found at the water’s edge but patrol in the middle of the river, very low over the water. It may be useful to scan for them with binoculars.

in eastern Europe. Deemed extinct in most of western and central Europe, but staged a strong comeback in the 1990s. The reasons are largely unknown; changes in water quality and climate are favoured explanations. Now abundant in rivers such as the Rhine, Rhône, Elbe and Danube, right down to their estuaries. Whether this increase marks an expansion from the east or the recovery of (overlooked) relict populations is uncertain. The

Occurrence Range and status A north Asian species, with western outposts in large rivers such as the Rhine, Loire and Po. Common in large lowland rivers

S. ubadschii

Gomphus Clubtails

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species is increasingly being discovered in smaller tributaries and therefore its distribution is probably more diffuse than suggested by the map. Habitat Slow-flowing lower sections of large rivers with sandy beds. Larvae burrow shallowly in fine substrates with relatively high concentrations of organic matter.

Flight season Late and protracted compared with other gomphid species: from early June to early October, with maximum emergence in June and July.

Stylurus ubadschii (Schmidt, 1953)

Syrian Clubtail 201

thorax markings and secondary genitalia differ from Gomphus in the same way as S. flavipes

mature ?

S7–9 widened to form strongly clubbed tail

? appendages upper appendages have smooth, unangled outer borders

short, pointed posterior hamule

? secondary

(side view)

Identification General Replaces S. flavipes in Turkey, the Caucasus region and Central Asia. Often treated as a subspecies of S. flavipes, but has a noticeably larger and more yellow abdominal club. Male appendages and larvae also differ in some characters. Formerly known as Gomphus lineatus, but that name was originally used for Paragomphus lineatus and was therefore replaced. Field characters Tot 44–51mm, Ab 31–40mm, Hw 25–31mm. Identical to S. flavipes, except smaller and with an even more strongly clubbed tail. S7, in addition to S8–9, is also distinctly widened, to almost as wide as S9, whereas it is clearly narrower in S. flavipes. Moreover, S9–10 are largely yellow, instead of extensively black. Occurs with G. davidi and G. schneiderii, but differs from these by features of the thorax markings (see S. flavipes).

genitalia

Hand characters The male’s upper appendages differ from those of S. flavipes by having smooth, unangled outer borders. Variation The extent of the abdominal yellow varies. Central Asian populations are darker than Near Eastern ones.

Occurrence Range and status Occurs south of S. flavipes, from Georgia, Turkey and Syria to Central Asia. The westernmost reports from Thrace and the Aegean island of Lemnos are unconfirmed. The two taxa may meet in north-western Turkey (see blue on map p. 200). Habitat Found in large rivers with sandy beds, as is S. flavipes. Flight season From May to early August.

Gomphus Clubtails

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Ophiogomphus Selys, 1854 Snaketails Ophiogomphus cecilia (Fourcroy, 1785)

Green Clubtail

Identification

202

Green Snaketail

General The only Ophiogomphus species in our area. It has a similar appearance to Gomphus and Stylurus species, but is heavier and is characteristically three-coloured, with an applegreen head and thorax, and a black-and-yellow abdomen. It is the typical summer gomphid of many sandy lowland rivers. Field characters Tot 50–60mm, Ab 37–42mm, Hw 30–36mm. Larger and more robust than Gomphus and Stylurus species; our largest gomphid besides Lindenia tetraphylla. The male is clearly club-tailed, with S8–9 thickened. The head, eyes, thorax and S1–2 are distinctly pale green, with thin black markings (excepting the green-eyed greenish G. vulgatissimus, Gomphus and Stylurus species have a yellowish head and thorax with contrasting blue eyes when mature). The thorax pattern resembles Gomphus pulchellus and especially G. simillimus. S3– 10 have pale yellow dorsal markings. These are less linear and more triangular than those in Gomphus and Stylurus species, but not as short and broad as in co-occurring Onychogomphus species. Unlike Gomphus and Stylurus, the upper appendages are yellow, not black. The legs are yellow, with only thin black stripes. The green colour will not have

HD II+IV

developed in tenerals and these are best identified by hand characters. Hand characters Hw has an anal loop, usually of three cells; therefore there is not a perpendicular vein running directly to the wing’s hind margin from the subtriangle, as in Gomphus and Stylurus. The male’s modest appendages are very unlike those of Gomphus, Onychogomphus and Stylurus, being distinctly short (as long as S10), straight and parallel. The female has two unique features: the occiput is crowned with two irregularly toothed crests, and the vulvar scale is drawn out into two very fine diverging points. Ophiogomphus is a mainly North American genus but several species occur in Central Asia, of which only O. cecilia reaches us. Variation The thorax lines are always thin, but may be even more reduced. Southern individuals are possibly smaller. Behaviour Males perch by smaller waters waiting for females, but at larger rivers (20m wide or more) they often fly in the middle of the watercourse in search of passing females.

Occurrence Range and status Widespread over most of eastern Europe up to Germany, but found only in isolated populations in France and Italy; records from the Iberian peninsula remain unconfirmed. The species has a history of decline in many areas, but in the last decades populations in central Europe have recovered so well that it is now regionally common. Habitat Small highland rivers to large lowland rivers; most abundant in lower sections of large rivers with a sandy bottom. The larvae prefer to burrow in gravel and sand, avoiding muddy areas. Flight season Emergence varies locally, usually starting at the end of May, but can be in early May in large rivers in warm regions, and not before July in very cold, shady streams. Most abundant in July and August, but may fly until October.  F Suhling and O Müller

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eyes green, like frons and thorax

distinctive threecoloured pattern in both sexes

203

triangular yellow central markings on abdomen

anal loop (here 2-celled) interrupting course of perpendicular vein between subtriangle and wing border

mature ?

/ immature

short, pale appendages

green thorax with thin black markings

vulvar scale with fine points

/ occiput with two

/ vulvar scale

crests

? appendages (side view)

H Ophiogomphus cecilia male, showing the characteristic green colour of the head and thorax.

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Onychogomphus Selys, 1854 Pincertails Hooktails

204

G Onychogomphus assimilis male on a streamside rock in typical posture with its abdomen raised.

Identification Diagnosis Best known for the male’s large appendages: the upper and lower appendages are both distinctly longer than S10 and are curved strongly towards one point, like pincers. With the exception of the faintly marked brown O. costae, all species have a yellow abdomen ringed with black, and a yellow thorax with black stripes. The Hw has a small anal loop of one to three cells, thus there is not a straight perpendicular vein running directly to the wing’s hind margin from the last thick lengthwise vein in the wing base. Separation from other genera The males can hardly be misidentified because of their appendages, but females may be mistaken for Gomphus, Ophiogomphus, Paragomphus and Stylurus. Only Ophiogomphus has an anal loop, but in this genus the thorax is largely green and the female has two crests on top of the occiput. Paragomphus species are smaller; the male has broad flaps on S8–9 and the female has a row of small black denticles on the rear of the occiput. Gomphus and Stylurus are marked differently, the abdomen appearing blacker and longitudinally striped, rather than ringed. Separation of the species In a large part of Europe the only species of this genus that occurs is O. forcipatus. In the Mediterranean there are seven

G Onychogomphus flexuosus pair mating.

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additional species, four of which are found only in Turkey and one of which is endemic to Morocco. These can quite easily be recognised by the pattern on the abdomen and thorax, although separation on sight between O. forcipatus and O. uncatus (in the south-west) is difficult, and between O. lefebvrii and the various subspecies of O. forcipatus

is not possible. Male appendages and the presence of ear-like tubercles behind the eyes in females are important characters in the hand. Behaviour Males often perch on gravel banks or boulders lying near or in the water, raising their abdomen and accentuating their appendages.  V J Kalkman and K-D B Dijkstra

Comparison of Onychogomphus species (recently added O. cazuma not included) Range

SW Europe, NW Africa

S3–6

Brown

? upper/ lower apps

Unique features of ? lower appendage

Longer

Smooth, branches diverge, not parallel

One black ring

Turkey

Equal

Longer Two black rings

Species

205 costae

Absent

uncatus

With subbasal knobs only

Widespread Morocco

/ postocular tubercles

With subbasal and subterminal knobs

Strong

Lacks knobs, but has weak hump roughly halfway

Weak

forcipatus boudoti

Almost smoothly crescent-shaped

lefebvrii

Straight, slightly thickened at midlength

assimilis Absent

Slender and wavy

flexuosus macrodon

Large triangular teeth at midlength

Appendages of male Onychogomphus (side view)

O. costae

O. uncatus

O. assimilis

O. forcipatus

O. flexuosus

O. lefebvrii

O. macrodon

O. boudoti

Postocular tubercles of female Onychogomphus (view from above)

O. uncatus

O. forcipatus

O. lefebvrii

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Onychogomphus uncatus  (Charpentier,

Large Pincertail

1840)

Blue-eyed Hooktail yellow ‘collar’ interrupted in middle by black

yellow stripes connected dorsally

206

/ occiput /

?

? appendages (view from above)

extensive confluent black lines on side of thorax

narrow, finger-like lobes

/ vulvar scale

no dorsal lobes

anal triangle normally 4-celled

? appendages

no subterminal knobs

(side view)

Identification General An impressive black-and-yellow gomphid, found within the south-western range of O. forcipatus. Must be examined closely while perched, to distinguish it from its compatriot. Field characters Tot 50–53mm, Ab 34–42mm, Hw 29–33mm. General appearance is close to O. forcipatus, but slightly larger and the yellow abdominal markings are more extensive on average, although this varies greatly in O. forcipatus. Best distinguished as follows: (1) the vertex is all black, and there is no extra yellow bar

between the yellow of frons and occiput (although this is sometimes also unclear in O. forcipatus); (2) the ‘collar’ (the transverse yellow area on the anterior ridge of the thorax) is severed by the black area along the mid-dorsal keel; (3) the yellow antehumeral stripe dorsally connects with the broader yellow stripe before it, thus the black stripe separating them is not connected with the black along the mid-dorsal keel; (4) the black on the side of the thorax is more extensive, and the stripes are not broken but often partly confluent (especially in southern O. forcipatus – i.e. where it overlaps with O. uncatus – these stripes are reduced). Occurs with the similar O. boudoti in a tiny area in the Middle Atlas of Morocco (see that species). Hand characters Male anal triangle is normally four-celled, but is three-celled in most O. forcipatus. The male appendages are like those of O. forcipatus, including subbasal knobs of the lower, but lack the subterminal knobs of that appendage in O. forcipatus. Uppers curved towards each other (view from above), without the thumb-like dorsal lobe in the bend; the tips meet, but do not overlap. Female lacks tubercles behind the eyes. The vulvar scale is reduced to a pair of finger-like processes pointing towards each other. In O. forcipatus, the vulvar scale appears as two closely apposed, roundly triangular lobes.

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Occurrence Range and status Fairly common in Morocco, but rare elsewhere in northern Africa. Fairly common in Spain, southern France and parts of Italy, but rare in the rest of range.

Habitat Streams, less often rivers. Prefers fasterflowing, more shaded and smaller waters than O. forcipatus. Flight season From May to September.

Onychogomphus forcipatus 

Small Pincertail

(Linnaeus, 1758)

Green-eyed Hooktail 207

/

?

?

normally 3-celled anal triangle

variable thorax and abdomen markings cannot be used to separate subspecies reliably

ssp. forcipatus (dark individual)

ssp. unguiculatus

?

subbasal knobs

ssp. unguiculatus

ssp. forcipatus

/ two tubercles

rounded lobes

behind eyes

/ vulvar scale

Identification General The most common and widespread Onychogomphus species and the only one occurring in large parts of Europe. Males perched on a streamside rock, with their claspers raised, are a typical sight in summer, especially in the south.

? appendages (view from above) ssp. forcipatus

dorsal lobe

Field characters Tot 46–50mm, Ab 31–37mm, Hw 25–30mm. A medium-sized anisopteran that is about as large as Orthetrum brunneum. The only Onychogomphus that can have dark (not yellow) upper appendages and green eyes (without a hint of blue). Unfortunately, the species

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208

is variable and these characters usually do not apply in Mediterranean countries where O. forcipatus overlaps with its congeners. Here, it is best identified by exclusion. In Turkey, O. flexuosus and O. macrodon are much paler, with a ‘double-ringed’ abdomen and pale Pt, while O. assimilis has a clearly different thorax pattern. O. lefebvrii can best be separated in the hand, but hardly overlaps. In south-west Europe and north-west Africa, O. costae is almost devoid of black, while O. uncatus differs mainly in details of the thorax (see that species); O. boudoti is clearly darker than the regional O. forcipatus unguiculatus and differs in markings and male appendages (see those species). Hand characters Upper and lower appendages of similar length. Lower has two small dorsal tooth-like knobs at the base, as well as a dorsal knob near the tip. The subbasal knobs are shared only with O. uncatus. The subterminal knobs are unique, their shape defining the subspecies. The uppers typically overlap at their tips and each bears a thumb-like dorsal lobe, lying just past the appendage’s bending point (visible from above or behind). The female has a small but distinct yellow(-tipped) tubercle behind each eye. Other than the less developed tubercles in O. lefebvrii, these are unique. Variation The three subspecies were long separated by coloration, but this is unreliable. Compared with ssp. forcipatus, the southern ssp. unguiculatus and albotibialis have less black (e.g. usually more yellow

on the thorax and upper appendages) and the eyes are more blue-green than green. It is necessary to examine the appendages of several males within a population to identify the subspecies to which they belong (see table).

Occurrence Range and status Common in the south, but rather local in the north, extending to northeastern Kazakhstan. Habitat Largely unshaded (usually rocky) rivers and streams. Occasionally occurs at large lakes. Flight season Typical gomphid of summer, from May to September.

? appendages (side view) see table below

dorsal lobe

subterminal knob ssp. forcipatus

ssp. unguiculatus

subbasal knob

ssp. albotibialis

Subspecies of O. forcipatus Subterminal knobs of lower appendage (view from side) Range

Length/ width

Angle between it and rest of the appendage (median and range)

Subspecies

S France, N two-thirds of Italy, Iberia, NW Africa

1–2.5×

90° (25–120°)

Remainder of Europe

1–2.5×

>140° (120–160°)

forcipatus

Cyprus, Turkey and adjacent Greek islands

0.5–1×

90° (25–120°)

albotibialis

unguiculatus

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Onychogomphus lefebvrii (Rambur, 1842)

Pale Pincertail

pale pterostigma

209

?

black markings on thorax reduced

paler than O. forcipatus, but adjoining populations of that species also relatively pale

no knobs on lower appendage

/ two blunt tubercles behind eyes ? appendages (side view)

Identification

Occurrence

General Replaces O. forcipatus in the arid regions of south-western and Central Asia. Has been considered a subspecies of O. forcipatus, which it resembles closely. Must be identified in the hand. Field characters Tot 45–52mm, Ab 31–40mm, Hw 26–30mm. Similar in shape and pattern to O. forcipatus, although paler and perhaps sleeker on average. Indications in the field are the paler Pt (yellow-brown, not blackish brown) and greener eyes, but these cannot be relied on for identification. May occur with O. flexuosus and O. macrodon, with which it shares a three-coloured appearance: the pale yellow thorax contrasts with the ivory abdomen base and bright orange-yellow club; these species have two separate black rings on each of S3–6, rather than just one. Hand characters Male appendages are like those of O. forcipatus, but the lower is completely devoid of knobs or teeth near the base or tips. Careful comparison is needed for female identification: a yellow tubercle is present behind each eye, but these are less prominent than in O. forcipatus.

Range and status Ranges from Adana to Afghanistan, Israel and possibly Egypt. Common in south-east Turkey, where O. forcipatus is largely absent; both species show little overlap. Habitat Rocky and largely unshaded streams and rivers. Flight season From mid-May to early August.

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Onychogomphus boudoti  Ferreira, 2014

Boudot’s Pincertail

210 /

no subterminal knobs

? / vulvar scale

normally 4–5 celled anal triangle

Identification General This species was discovered only in 2011 and is still known from just two small populations in the Middle Atlas of Morocco. At first sight appears identical to the two other medium-sized black-and-yellow Onychogomphus species found there, but is easily separated by the distinctive male appendages and female vulvar scale. Field characters Tot 49–50mm, Ab 39–40mm, Hw 31–33mm. Due to its extensively black thorax, clearly appears darker than co-occuring O. forcipatus and at least slightly darker than O. uncatus. Markings combine features of both these species: like O. uncatus has (1) confluent black lines on the thorax sides and (2) yellow ‘collar’ anteriorly on the thorax interrupted by black; but like O. forcipatus has (3) yellow antehumeral stripe not connected dorsally to the broader yellow stripe before it, and (4) yellow patch on the vertex, rather than a wholly black vertex. Especially when seen from the side or above, the male appendages appear bulkier in O. boudoti: the robust lower appendage has neither subbasal nor subterminal knobs, but does have a (variably prominent) tubercle at about mid-length. This unique character is often visible with binoculars or the naked eye, so O. boudoti males may picked out by jizz with some experience. O. assimilis is similarly dark, with a rather blunt lower male appendage, but is excluded by range. Hand characters The shape of the appendages is the most reliable character in males. Anal

? appendages (side view)

blunt thickening at midlength

triangles of Hw in males usually contains four or five cells (as in O. uncatus), while most of O. forcipatus have only three. Females have two small tubercles behind the eyes, as in O. forcipatus but not O. uncatus. The lobes of the vulvar scale are rounded (as in O. forcipatus) instead of being narrow and finger-like (as in O. uncatus), but the cleft separating them is less than half as deep as the vulvar scale is long, rather than dividing the lobes almost completely. Behaviour Males, females and mating wheels perch mainly on stems, sticks or plants, but may settle on the bare ground under cloudy skies. Both sexes appear not to wander far from the small streams where they mate and breed.

Occurrence Range and status Extremely local Moroccan endemic, confined to just a few streams in Khénifra province. Due to tiny population size, limited genetic diversity, and unregulated land use, it is classed as Critically Endangered. Habitat Diffuse shallow trickles that form tiny pools with aquatic and emergent vegetation on calcareous gravel and cobbles. These are part of grassy swamps that may almost dry up in summer. The water runs through pasture and agricultural land to a river nearby with O. forcipatus and O. uncatus. Flight season Emergence occurs in late June and early July, but the species is likely to be on the wing until August.

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Onychogomphus cazuma

Cazuma Pincertail

Barona, Cardo & Díaz, 2020

/ vulvar scale

211 ? appendages (side view)

? appendages (view from above)

G Onychogomphus cazuma male.

Identification General This book was almost in print when news seeped through that in 2017 citizen scientists in Spain had found a new species to science. This is the first from Europe in two decades, and the first from south-western Europe since the 19th century! While it appears like a mix of the cooccurring O. forcipatus and O. uncatus, the male appendages and female vulvar scale are unique. Genetically the species is closest to O. boudoti from Morocco, discovered only six years earlier. As both seem highly localized due to their specific habitat preferences, more new species may yet be found. Field characters Tot 43–47 mm, Ab 29–35 mm, Hw 25–30 mm. Appears paler and smaller than O. uncatus, with the lower male appendage never being darker than the uppers. May thus also be taken for O. forcipatus, but with a confusing mix of features recalling both species, such as (1) yellow antehumeral stripes connected dorsally to the broader yellow stripes before them like O. uncatus; (2) yellow ‘collar’ anteriorly on the thorax not (or very narrowly) interrupted by black like O. forcipatus; and (3) yellow patch on the vertex that is small and circular, but typically a wider bar in O. forcipatus and absent in O. uncatus. Hand characters Identification requires close examination of male and female sexual characters. The shape of the male appendages is somewhat in-between O. uncatus and O. boudoti. The lower appendage not only lacks the subterminal knobs of O. forcipatus, but also the subbasal knobs found in that species and O. uncatus. Instead, there is a

subbasal ridge with a peak that is directed straight upward, which thus does not project outwards when viewed from above. The upper appendages are not lobed dorsally like O. forcipatus, but have a spoon-shaped tip similar to O. uncatus. Anal triangles of Hw in males usually contain 3 cells (as in O. forcipatus), while most O. uncatus have 4 or 5. Females lack the two small tubercles behind the eyes of O. forcipatus (thus like O. uncatus). However, the lobes of the vulvar scale are not narrow and finger-like as in O. uncatus, but broad (more like O. forcipatus), although more pointed and the cleft separating them is about half as deep as the vulvar scale is long, rather than dividing the lobes almost completely (thus most like O. boudoti). Behaviour Similar to other Onychogomphus.

Occurrence Range and status So far found on well-preserved tributaries of two river systems in Valencia province of eastern Spain. As these habitats have very low flow and are vulnerable to development, pollution and drought, the species may well be threatened under IUCN Red List criteria. Habitat Springs, streams and upper courses of small rivers in low mountains. These are calcareous with very clear, nutrient-poor water. Males perch at more vegetated and slower-flowing sections (often alternating between shallow parts and small pools) than those of O. uncatus. Flight season So far observed from early June to early September.

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Onychogomphus assimilis (Schneider, 1845) Dark Pincertail vertex all black

black stripes broad and partly confluent

212

pale central area on S2 smaller than on S3

/

?

single broad black stripe uppers longer than lower

/ vulvar scale

? appendages (side view)

Identification General The largest, blackest and most contrasting Onychogomphus species in our area, where it is confined to mountain streams along the Turkish Mediterranean and the Black Sea coasts. Field characters Tot 50–55mm, Ab 35–42mm, Hw 31–34mm. Somewhat larger than O. forcipatus. The black abdominal pattern is largely similar, albeit usually more extensive, to that of O. forcipatus, which also occurs in Turkey. Instantly identified by pattern on the head and thorax: (1) the vertex is all black and there is no extra yellow bar between the yellow of frons and occiput; (2) the humeral

VU

lower straightish, thickened at midlength

black stripe and the black stripe before it are broad and partly confluent; (3) the largely pale sides are interrupted only by a single broad black stripe. In females, the yellow dorsal marking on S2 is smaller than that on S3; in O. forcipatus it is about as broad as that on S3. Coloration may recall the co-occurring Gomphus schneiderii, but that species has a yellow-striped (rather than ringed) abdomen. Hand characters Male upper appendages slender and much longer than the lower appendage. Lower appendage has a straight underside and blunt dorsal thickening at mid-length, but lacks the subbasal knobs of O. forcipatus. Variation The abdomen may vary from bright yellow to almost white. Behaviour Males regularly perch on boulders at or in the water, but unlike congeners also often rest on branches a few metres above the water.

Occurrence Range and status Occurs from Turkmenistan west to Turkey, where it is not uncommon along the southern coast. Recent records near the Black Sea suggest the species has been overlooked in northwestern Turkey. Habitat Partly shaded and often cold rivers, and at the foot of mountainous areas. Flight season From late May to the end of August.

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Onychogomphus costae Selys, 1885

Faded Pincertail

pale pterostigma

virtually devoid of black

/

213

?

lower appendage relatively short and without knobs

/ vulvar scale

? appendages (side view)

lower appendage with splayed branches partly concealed by longer uppers

Identification

otherwise often seen resting on bare ground.

General Small, odd-looking western Mediterranean Onychogomphus, whose body has different shades of ivory, ochre and rust but is almost devoid of black. Easily overlooked owing to its small size and camouflaging coloration. Field characters Tot 43–46mm, Ab 30–34mm, Hw 22–27mm. Smallest and palest Onychogomphus. Easily told from congeners by size, pallid but warm brown ground colour and the absence of large, distinct black markings. It is the only Onychogomphus within its range with a beige rather than dark brown to black Pt. Resembles Paragomphus genei in size, posture, pale Pt and indistinct markings, but latter has a green face and thorax, more abdominal black and broad flaps on S8–9 in the male (appendages also differ). Hand characters Male appendages are slender and very distinct from other Onychogomphus. The lower appendage is much shorter than the uppers, is elongate and lacks knobs or teeth. It is deeply incised (view from below), the two branches widely diverging. In other Onychogomphus the branches are close and parallel; in O. forcipatus and O. uncatus they bear subbasal knobs. Males have long whitish hairs at the base of the underside of S7, which are three to four times as long as those on S6. Behaviour Males wait for females near the water; mating wheels usually perch on bushes and trees,

Occurrence Range and status Endemic to arid parts of the Iberian peninsula and north-west Africa. Rare in most of its range, but not uncommon locally in Morocco and the Spanish Guadalquivir and Ebro valleys. Habitat In Spain, strongly connected to large lowland rivers with pebbly stretches. Regionally also in stagnant pools of intermittent streams (oueds, wadis). In Morocco, also found at higher altitudes in arid uplands. Flight season From May to August.

NT

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Onychogomphus flexuosus 

Waved Pincertail

(Schneider, 1845)

brownish pterostigma bordered in black

thin lines on side of thorax

214 /

S3–6 ‘double-ringed’

?

S7–10 orange-brown

/ vulvar scale

? appendages

wavy lower appendage

(side view)

Identification General An attractive, sleek Onychogomphus species with a bright straw-yellow appearance. Uncommon in southern Turkey and unlikely to be found elsewhere in our area. Field characters Tot 41–46mm, Ab 30–35mm, Hw 25–30mm. Somewhat smaller and more slender than O. forcipatus. Note the following characters: (1) black on S3–6 separated into one black ring at apex and one near base, giving the abdomen a double-ringed impression, subbasal ring thinnest, sometimes reduced to spots; (2) S7–10 orange-yellow with dark patches of variable size, distinctly brighter than the whitish basal half of the abdomen and pale yellow thorax, and

VU

appearing as a yellow ‘tail-light’; (3) Pt brownyellow, bordered by thick black veins; (4) thorax with thin, straight black lines, anterior to the metastigma often complete from the leg to wing base. O. macrodon, O. lefebvrii and the regional O. forcipatus albotibialis have a similar three-coloured appearance, with yellow thorax, pale abdomen base and bright club, but only O. macrodon shares the double-ringed abdomen. It must be separated in the hand (see species text). Paragomphus lineatus is superficially similar, but does not have doubled black abdominal rings. Hand characters Male has slender upper appendages, longer than lower; the tips meet but do not overlap. Latter is not thickened or toothed, but gently wavy, like a shallow ‘W’ in side view. Variation The extent of the black markings on the thorax and abdomen varies. Behaviour Wanders widely. Mainly found perching in dry scrubland away from water; copulates mainly in the afternoon in open landscape at some distance from the river. Likes to perch on sand or pebbles, where it is perfectly camouflaged; difficult to follow when flying swiftly low over the ground.

Occurrence Range and status Distributed from Afghanistan to Turkey, where it is scarce. Habitat Large, unshaded sandy rivers with gravel banks. Flight season From mid-May to the end of July.

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Onychogomphus macrodon Selys, 1887 Levant Pincertail

215 /

‘double-ringed’ abdominal pattern

? / vulvar scale

more extensive black markings on S7–9

Identification General A largely orange to brown-yellow Onychogomphus species with limited black markings, confined to the Near East. Superficially most similar to O. flexuosus. Field characters Tot 50–52mm, Ab 35–37mm, Hw 26–28mm. Larger and more robust than O. flexuosus, thus nearer O. forcipatus. With the exception of the former, distinguished from other Turkish congeners by its reduced black markings, which on S3–6 form a double set of black rings. Distinguished from O. flexuosus by less contrasting black thorax pattern, with the two black stripes next to the mid-dorsal keel on front of the thorax (one pair on each side) usually rather blurred and not connected across the keel (usually well developed and connected with the keel in O. flexuosus). Beware confusion with Paragomphus lineatus (see O. flexuosus text). Hand characters Male has slender upper appendages, longer than lower. The latter bears a pair of large triangular dorsal teeth at about mid-length. The underside of female S9, behind the vulvar scale, bears strong longitudinal and transverse ridges, which are short and hardly developed in O. flexuosus. Behaviour Males frequently perch on gravel banks in the riverbed some distance from the water’s edge,

well-developed ridges on underside of S9

? appendages (side view)

large tooth on each branch of lower appendage

but apparently not on rocks in the water, bushes or twigs; both sexes may hunt among shrubs on river banks, where they are perfectly camouflaged.

Occurrence Range and status Ranges only from northern Israel to the region of Adana and Antakya in Turkey. Uncommon within its restricted range; vulnerable as a result of habitat destruction. Habitat Large, mainly lowland rivers with gravel banks. Flight season From late May to mid-July.

VU

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Paragomphus Cowley, 1934 Hooktails Identification

216

Diagnosis Small gomphids. Males have broad flaps on S8–9 and diagnostic appendages: uppers are long, parallel and down-curved, like hooks or walking canes, lower is at most half their length. Separation from other genera Lindenia tetraphylla is the only other species with broad flaps, but these are on S7–8 and the species is 70–80mm long. Small and pale Onychogomphus species differ in details of markings, and males lack

broad flaps and have different appendages. Lacks a small anal loop in the Hw: there is a straight perpendicular vein running directly to the wing’s hind margin from the last thick lengthwise vein in the wing base. Separation of the species Numerous species occur in tropical Africa and Asia, but only two reach the Mediterranean. These differ in coloration. Behaviour Often perches with abdomen raised in V J Kalkman obelisk position. 

Paragomphus genei (Selys, 1841)

Green Hooktail

darkish pterostigma with pale centre

green thorax with indistinct markings

/

?

perpendicular vein not interruped by an anal loop

/ vulvar scale

anvil-shaped posterior hamule

? secondary genitalia

abdomen marked with mottled brown and black

long, slender down-curved upper appendages

flaps on S8–9

semicircular ridge on underside of S9

Identification General The only Paragomphus species in the western Mediterranean. It is quite easily

? appendages (side view)

distinguished by its blotched abdomen, and green face and thorax. Field characters Tot 37–50mm, Ab 30–36mm, Hw 21–26mm. Smaller than Onychogomphus forcipatus. Not yet found overlapping with its congener P. lineatus, but size, shape and coloration are similar to its compatriot O. costae. Unlike both, P. genei has: (1) pale green face and thorax with indistinct dark markings; (2) sandy- to strawcoloured abdomen, mottled with brown and black, giving a somewhat indistinct pattern. Both O. costae and P. lineatus lack the greenish coloration: O. costae is a rather warm, sandy brown overall, with virtually no black, and males lack the flaps on S8–9. P. lineatus is quite uniformly straw-coloured, marked less extensively but more sharply with black. See P. lineatus for a fuller comparison. Hand characters Male’s upper appendages are long and slender, strongly curved downwards and

Paragomphus Hooktails

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usually ending in two teeth. The lower is only half as long and is curved upwards, almost touching the uppers. O. costae has the lower appendage well over half as long as the uppers. Male P. genei is the only gomphid with an anvil-shaped posterior hamule. Females have a semicircular area on the underside of S9, surrounding the vulvar scale. This area is encircled by a low ridge of even thickness. Variation Extent of the dark markings, especially on the abdomen, varies greatly.

Occurrence Range and status Scarce to fairly common in northwest Africa. Rare on Corsica and Sicily; commoner and increasing in south-western Iberian peninsula and on Sardinia, benefitting from construction of reservoirs and ponds. Not uncommon in the Levant and therefore likely to be found in southern Turkey. Numerous in tropical Africa. Habitat Still or slow-flowing, often seasonal, waters, such as pools in dry streambeds. Flight season Mainly from April to October, but throughout the year in Saharan oases.

Paragomphus lineatus (Selys, 1850)

217

Lined Hooktail

sandy coloured thorax with distinct linear markings

/

? dark individual

/ vulvar scale

thicker upper appendages

? secondary genitalia

distinct swellings beside semicircular area on underside of S9

club may be largely yellow

Identification General A southern Asian species that reaches its western limits in south-east Turkey. There it may occur with P. genei, although the latter has not yet been recorded from here. It is a small, pale gomphid, with a distinct pattern of black lines. Field characters Tot 45–50mm, Ab 32–35mm, Hw 23–28mm. Similar in size to P. genei, with which it is most likely to be confused. Has a sandy-coloured abdomen, thorax and face, with well-defined black markings: fine lines on the thorax and thicker bars on the abdomen. P. genei has a green face and thorax with fainter markings, and the dark abdominal pattern is less well defined. The flaps on S8–10 are largely yellow, but brown and black-bordered in P. genei. Colour and build may recall some Onychogomphus species, O. flexuosus

long posterior hamule ending in small hook

? appendages (side view)

appearing most similar and best separated by sexual characters. Hand characters The male’s upper appendages are thicker and blunter, and the lower shorter, than those of P. genei. Unlike that species, the male’s posterior hamule is long, ending in a small hook. In females, the semicircular area surrounding the vulvar scale is bordered on each side by a distinct swelling.

Occurrence Range and status Not uncommon near Adana and Antakya in Turkey, from where its range extends to northern India and Nepal. Not mapped. Habitat Poorly known; includes open and sluggish, often seasonal, streams. Flight season Recorded from May to September, but probably active from April to October.

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Lindenia de Haan, 1826 Bladetails Lindenia tetraphylla (Vander Linden, 1825) Identification

218

Diagnosis This gomphid’s combination of great length (about 70–80mm), large leaf-like flaps on S7–8 and triangles in wings crossed by veins is unique in our area. Only a single species of this genus is known worldwide. This slender dragonfly exhibits remarkable colour variation, from largely sandy to almost evenly black individuals. Field characters Tot 69–80mm, Ab 49–57mm, Hw 36–40mm. Our largest gomphid, being 10–20mm longer than Onychogomphus forcipatus. The abdomen is long and slender, appearing abruptly thickened near the end. Its length and the flaps on S7 rule out all other species. Paragomphus and Onychogomphus have flaps on S8–9 instead of S7–8, but only in Paragomphus are they similarly large. Both these genera are not longer than 60mm. The body is typically pale yellow, marked with dark brown or black, but the markings vary considerably (see below). The male has numerous dark microscopic spines on the abdomen, especially from the middle of S3 to S7, giving it a darker appearance. The yellow to pale brown Pt is relatively large and long. At a glance, may be mistaken for the long-bodied skimmers Orthetrum sabina and O. trinacriae, but these lack wide flaps and separated eyes.

Bladetail

Hand characters Triangles in the wings are divided by cross-veins into three to four cells, but are single-celled in our other gomphids. The upper appendages are finger-like and almost straight, about twice as long as S10, dwarfing the lower. Paragomphus and Onychogomphus have strongly (down- and/or in-) curved upper appendages. Variation The thorax and abdomen may be largely dark, with obscured pale areas. Although darkening could progress with age, there are records of largely black tenerals, suggesting the darkness is determined by climatic conditions. Old individuals develop some bluish pruinosity on the thorax. Dark, pruinose individuals have a totally different appearance to the pallid fresh ones. Both the darkening and the pruinosity are unusual in gomphids. Behaviour Often perches on the ground, with the abdomen slightly raised above horizontal. Has migratory tendencies, which is very unusual for a gomphid.

Occurrence Range and status Patchily distributed from Central Asia, through the Middle East into the H Lindenia tetraphylla mature male.

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many-celled triangle

large pale pterostigma

219 dark speckling on upper surface of abdomen

?

largest gomphid in region

flaps on S7–8

long straight upper appendages

/

Mediterranean basin. In our area, only a handful of permanent populations are known, mostly in Greece, Montenegro and Turkey, although some of these are very large. Occurrence is very sporadic in the western Mediterranean. Habitat Lakes and slow-flowing rivers, both often large. Frequently associated with extensive beds of Common Reed (Phragmites australis), but also found at sparsely vegetated lakes; can colonise new barrage lakes within a few years. Flight season Most records from May to August, emerging in Turkey from the end of May.  V J Kalkman

HD II+IV

Lindenia Bladetails

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Cordulegastridae

Cordulegaster Leach, 1815 Goldenrings 

Spiketails (NA)  Cordulegaster insignis female, laying eggs by rhythmically piercing the soil at the bottom of a shallow stream.

220

Identification Diagnosis Easily recognised by the very large size (70–100mm), the black-and-yellow body pattern and the eyes just meeting in a point on top of the head. Males have an anal triangle in the Hw and auricles on the sides of S2. Females have a unique vulvar scale that serves as an ovipositor, projecting well beyond the tip of the abdomen. It is used to deposit the eggs into the bottom substrate of springs and small brooks, where goldenrings are often the only species of dragonfly present. Separation from other genera Aeshnids are often large and have a similar appearance and venation, but the eyes are broadly confluent. Many gomphids also have a black-and-yellow coloration, but are smaller and the eyes are widely separated above. The local Macromia splendens in southern France and Iberian peninsula, and Anax immaculifrons in southern Turkey, are similar in appearance but differ in various markings. Separation of the species The genus consists of two species groups. Each is a complex of similar species, the distributions of which hardly overlap. If two Cordulegaster species co-occur, these mostly belong to different species groups. In areas where both C. boltonii and C. bidentata occur (western

and central Europe), the former inhabits generally larger brooks than the latter. As a result, C. boltonii is more a lowland species, whereas the range of C. bidentata reflects the presence of lower mountain ranges. This partial segregation of the two species groups is less clear in the Balkans and Turkey After assigning an individual to a species group (see the first table below), the locality is often the easiest way to identify the species (second and third tables). Identification on the basis of the markings on the various body parts is hampered by the fact that it is still unclear which characters are reliable. Moreover, individuals within a population may vary in a number of characters, e.g. the size of the black marking on the frons, the extension of black on other parts of the head, the shape and size of the yellow marking on the thorax between the two large lateral bands, the shape and size of the yellow marking on S1 and the presence of apical spots on S2–8. These characteristics are thus unsuitable for identification purposes, unless stated otherwise in the species descriptions. In members of the Cordulegaster boltonii group, and especially in C. heros and C. picta, a small spot may be present near the upper lateral corner of the antehumeral stripe, and it may vary in size. Note

Cordulegaster Goldenrings

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Comparative table of Cordulegaster species groups (See pp. 223 and 229 for illustrations) Yellow marking on side of S1

Upper appendages (view from above)

Group

Along lower hind margin, normally shaped as reversed ‘C’, but sometimes reduced to a rectangle in the lower half

Diverging, with curved outer borders and close (almost touching) at base

boltonii

Oblique spot or broad triangle near middle of segment; if descending towards lower hind margin, then linear and not continuing along the lower edge of the segment, thus not forming a reversed ‘C’

Parallel, with straight outer borders and separated at base

bidentata

221

Geographic key to the species of the boltonii group Moroccan High and Middle Atlas

princeps

S Italy and Sicily

trinacriae *

N and W Turkey, islands in N and E Aegean Sea, SE Balkans

picta

W and S Balkans

heros **

Other parts of Europe or NW Africa

boltonii

* C. trinacriae and C. boltonii overlap and hybridise in Lazio and Marche regions of central Italy ** C. heros and C. boltonii overlap in eastern Austria and potentially western Ukraine Geographic key to the species of the bidentata group S Greece (including Peloponnese, Euboea and Cyclades)

helladica

Turkey, islands in N and E Aegean Sea (including Ikaria), SE Balkans

insignis ***

Other parts of Europe

bidentata

*** C. insignis and C. bidentata overlap from eastern Serbia to Bulgaria and Greek Macedonia that females are larger than males, with larger abdominal markings. The measurements for sexes are therefore provided separately. Within various species a number of subspecies are recognised, with narrow or wide geographical zones in which intermediate specimens occur. Some subspecies have been described on the basis of such intermediate specimens, resulting in taxonomical confusion. Others were described on material of unknown or doubtful origin, complicating the determination of the exact distribution of the subspecies. Behaviour All species breed in permanent and intermittently flowing springs and smaller streams, where the larvae live for three to seven years. Males patrol over the water surface, often

slowly, in search of a female. Not easily disturbed, patrolling males can be observed at close quarters: if you stand astride a small brook and remain motionless, a male may fly through your legs! Females visit the water only for copulation and oviposition. The oviposition mode is highly characteristic, with the female hovering over shallow water in a vertical position, repeatedly pushing her ovipositor into the substrate. An ovipositing female may do this more than 500 times, remaining at a particular site for 15 minutes, and therefore can be easily observed. Females tend to avoid males during oviposition, and often hide under overhanging vegetation.  G J van Pelt

Cordulegaster Goldenrings

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Cordulegaster boltonii  (Donovan, 1807)

Common Goldenring Golden-ringed Dragonfly

Identification

222

General The only Cordulegaster species in a large part of western and northern Europe, e.g. the British Isles, Fennoscandia around the North and Baltic seas, lower France and south of the Pyrenees. In higher parts of western and central Europe, found together with C. bidentata, from which it can be distinguished by the more yellow appearance, the largely yellow costa, the wholly yellow occipital triangle, and the characters listed in the first table on p. 221. Field characters Male: Tot 74–80mm, Ab 52–64mm, Hw 40–47mm; female: Tot 80–85mm, Ab 57–69mm, Hw 45–51mm. Approximately as large as Anax imperator. In most of range typical individuals occur (ssp. boltonii) with small abdominal markings that run down and reach onto the underside of S4–8, but are interrupted above on all segments except S2(–3). Pairs of spots are present on the apical margin of S2–7(8), especially in females; S9–10 are (largely) black. The foreside of the occipital triangle and the costa are yellow. A small black bar is present across the frons. In south-west Europe and north-west Africa, C. boltonii is much yellower (see Variation). The species overlaps broadly with C. bidentata, overlaps less so with C. heros in eastern Austria, hybridises with C. trinacriae in central Italy, and almost meets C. princeps in the Moroccan Atlas. See these species for their separation. Hand characters Upper appendages relatively short; as long as S10. Variation Exceptionally has blue eyes, mostly in Spain. In a small part of north-west Italy individuals occur with a (partly) black occipital triangle. The following subspecies lack the black marking on the frons, and have larger dorsally confluent abdominal markings than ssp. boltonii, with additional marking on S9–10. In north-west Italy, southern France and in Spain, ssp. immaculifrons is found, with a broad zone (especially in central Spain) in which intermediates with ssp. boltonii occur. The marking on S3 is sometimes longer than broad (view from above), and the hind margin of S3–4 may show a ‘W’ shape. In south-east Spain, ssp. iberica has the hind margin of S2 indented by black, the rings on S3–4 longer than wide, and

/ ssp. boltonii

pairs of spots on apical margins of S2–8

spike-like vulvar scale diagnostic of genus

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? ssp. boltonii eyes touch in a point

foreside of costa yellow

? ssp. immaculifrons

? ssp. algirica

223

longer marking on S3 yellow occipital triangle

longer marking small black bar on frons

upper appendages diverging and almost touching at base

lower appendage broad, parallelsided and at most shallowly notched

additional yellow markings on S8–10

well-developed stripe between the broad yellow stripes

? appendages

? ssp. boltonii reversed ‘C’ on side of S1

the hind margin of those on S3–4 and S7 show a ‘W’ shape. Ssp. algirica, from southern Spain and north-west Africa, is similar, also with the ring on S5 sometimes longer than wide, and the hind margin of those on S3–7 show a ‘W’ shape. All subspecies are weakly defined and, except for the African populations of C. b. algirica, none are clearly separated genetically.

Occurrence Range and status Locally common. Reaches the Arctic Circle in Finland and adjacent Russia; extends to southern Urals. Habitat Streams and sometimes small rivers, often in forests but also in open moors and heaths. Flight season Beginning of May to late September, especially July.

Cordulegaster Goldenrings

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Cordulegaster princeps Morton, 1916 Identification

?

224

Atlas Goldenring

markings end in a point on side of abdomen

General Replaces C. boltonii in most of the Moroccan Atlas. It is identifiable by its large size, the shape of the abdominal markings, the largely black S9–10, and the largely pale area behind the eyes. Field characters Male: Tot 75–86mm, Ab 56–65mm, Hw 45–49mm; female: Tot 79–87mm, Ab 60–66mm, Hw 47–53mm. The abdominal markings on S3–6 do not descend onto the underside of the segments, as in C. boltonii, but end in a point halfway down. They are broadly connected dorsally on S2–8, and on S3–8 are indented at the hind and often the fore margin (view from above). S9 has a basal yellow spot, and S10 is usually all black, unlike C. b. algirica. Foreside of frons and occipital triangle are yellow, as C. b. algirica, but the back of the head (behind the eyes) is largely yellow. Hand characters Upper appendages as C. boltonii, as long as S10, outer margins relatively straight.

Occurrence ? appendages

Range and status Endemic to the High and Middle Atlas. Not known to overlap with C. b. algirica, but the two may meet on Jbel Tazzeka in the north of the Middle Atlas range. Habitat Streams (but not yet found in very small ones, as is C. b. algirica), roughly between 1,000m and 2,500m. Flight season Late May to mid-September.

Cordulegaster Goldenrings

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Cordulegaster trinacriae 

Italian Goldenring

Waterston, 1976 ?

Identification General Replaces C. boltonii in Italy roughly south of Rome, where they overlap in the Lazio region and are best separated in the hand by the shape of the appendages. Field characters Male: Tot 73–79mm, Ab 55–63mm, Hw 45–49mm; female: Tot 83–93mm, Ab 64–72mm, Hw 51–53mm. Resembles C. b. boltonii but with smaller abdominal markings that are usually dorsally connected on S2–4(5), and those on S4–7 do not completely descend to the underside. S2–6 have apical spots, and S9 small basal markings. Occipital triangle is yellow, and foreside of the frons is yellow with sometimes a trace of black in males and always with a small black bar in females (never extending onto the peaks), whereas C. b. boltonii has a small but distinct bar in both sexes. Hand characters Upper appendages are relatively long, slender and sinuous, somewhat separated at their base and slightly longer than S10 (view from above). Hind margin of lower appendage is deeply notched, unlike any other species. Variation A broad hybrid zone with C. boltonii extends from central Marche to southern Lazio. The male appendages of hybrids are smaller and more like those of C. boltonii, with the hind margin of the lower appendage less deeply notched. Genetic studies suggest that these differences limit boltonii-like males to only mating successfully with C. trinacriae females, which would allow C. trinacriae to intrude into the range of C. boltonii, extending the hybridisation zone further northwards.

reduced abdominal markings

225

hind margin of lower appendage notched

? appendages

NT HD II+IV

Occurrence Range and status Endemic to southern Italy and Sicily. Habitat As C. boltonii. Flight season June to August.

Cordulegaster Goldenrings

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Cordulegaster heros Theischinger, 1979 Balkan Goldenring Identification

?

226

abdominal rings connected above

apical spots on S5–8 absent or reduced

? appendages

NT HD II+IV

General In the Balkans replaces C. boltonii, from which it is distinguished by its larger size and larger abdominal markings. Field characters Male: Tot 77–84mm, Ab 56–64mm, Hw 45–50mm; female: Tot 88–96mm, Ab 68–76mm, Hw 53–58mm. The largest Cordulegaster species, and one of our largest dragonflies. Differs from C. b. boltonii as follows: (1) occipital triangle black, not yellow, but may have two small yellow spots as in C. picta, especially in females; (2) the narrow yellow stripe between the two broad lateral thorax bands has the hind margin usually bent at the middle, so its lower half is placed before the upper half; (3) median abdominal yellow rings larger (usually dorsally connected on S2–7 and reaching well onto the underside of S3–8), but apical spots reduced, absent on S7–8 and often on S5–6, especially in males (see artwork of rather similar C. picta). Another popular character, the angular upper outer corners of the antehumeral stripes, is not deemed reliable. See C. picta for differences with that species. Black bar on face of frons is always very extensive in females. This, the black occipital triangle, and its similar appearance may cause confusion with the co-occurring C. bidentata, which is smaller and differs by bidentata-group characters (see first table on p. 221). Hand characters Upper appendages somewhat longer, but similar in shape, compared to C. boltonii. Variation Two subspecies are recognised, differing only by the size of the black bar on the frons in males, which is at most vaguely indicated in ssp. heros from Austria, Slovenia, Hungary and Serbia, but is a distinct rectangular bar in ssp. pelionensis from Greece, Albania and Bulgaria. As in the other species, these taxa may have little standing.

Occurrence Range and status Endemic to central and southeast Europe. Overlaps and locally coexists with C. boltonii in eastern Austria; both species occur and may overlap on the north-western slopes of the Ukrainian Carpathians. Habitat As C. boltonii. Flight season June to August.

Cordulegaster Goldenrings

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Cordulegaster picta Selys, 1854

Turkish Goldenring

Identification General Replaces C. heros in the south-eastern Balkans and Turkey; any overlap with the distribution of the latter has not yet been established. As C. picta is highly variable, a positive identification should be made based on the male appendages. Field characters Male: Tot 72–80mm, Ab 53–60mm, Hw 43–46mm; female: Tot 80–89mm, Ab 61–68mm, Hw 48–53mm. A rather large Cordulegaster, but smaller on average than C. heros. Differences include: (1) occipital triangle never all black but with two yellow patches; (2) the narrow yellow stripe between the two broad lateral thorax bands has a rather straight hind margin; (3) the black bar across the frons is variable, may be absent or even extending onto peaks in males, and never covers peaks fully in females as in C. heros; (4) unlike C. heros, females often have the yellow band on S2 extended centrally towards the base (view from above). Abdominal markings similar to C. heros, median rings typically connected dorsally on S2–6(7) and running onto underside of S3–9, but may also be connected even on S8 or only on S2–3. Apical spots usually on S2–7, but may be limited to S2–3. Overlaps widely with C. insignis, but in a large part of its range that species is blue-eyed and has a yellow occipital triangle (not in northern Anatolia), aside from species-group differences (see first table on p. 221). Hand characters Upper appendages long, slender and diverging; longer than S10. Variation Abdomen pattern is highly variable and reliable identification is possible only by the male appendages. Very yellow populations are known from the eastern Rhodopes, western and southern Turkey, and some eastern Aegean islands like Samos and Lesbos; these individuals may be strikingly similar to C. heros, C. insignis or C. helladica. By contrast, populations from the western Rhodopes and northern Turkey (including the European part) are darker, resembling C. bidentata.

?

227

abdominal rings connected above

apical spots on S4–7 absent or reduced

? appendages

Occurrence Range and status The zone separating C. picta and C. heros probably runs from Kalkhidikhi to the Danube Delta, but is imperfectly known. Ranges to the Transcaucasus and north-western Iran. Habitat As C. boltonii and C. heros. Flight season Late May to mid-August.

Cordulegaster Goldenrings

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Cordulegaster bidentata Selys, 1843 Sombre Goldenring

Two-toothed Goldenring

228

G Cordulegaster bidentata male. Note the dark occipital triangle and pointed ends to the markings on the side of the abdomen.

Identification General The smallest and darkest Cordulegaster species outside Turkey. Field characters Male: Tot 69–78mm, Ab 52–60mm, Hw 41–46mm; female: Tot 74–83mm, Ab 55–63mm, Hw 45–50mm. Smaller on average than C. boltonii. Overlaps widely with the usually more common and more widespread C. b. boltonii. Separation from it is possible by species-group characters (see first table on p. 221), of which the lateral S1 marking is easiest, as well as: (1) markings small, median rings do not reach abdomen underside and appear rather triangular from side, apical spots present on S2–3(4) only, absent on S5–8; (2) costa predominantly black, not yellow (view from front); (3) occipital triangle black; (4) black bar on foreside of frons very extensive, normally extending onto

peaks of frons. Note that characters 1–4 are not group specific, e.g. they do not separate C. bidentata from C. heros, nor can they be used to distinguish C. insignis and C. picta. See C. insignis for differences with it and C. helladica. Hand characters Male appendages differ from those of most other Cordulegaster species by: (1) each upper appendage bears two teeth, which are about as close to each other as to S10 (view from side) – these give the species its scientific name, but note that both are present in other Cordulegaster species too, although the basal tooth lies closer to S10 and is often (partly) concealed; (2) lower appendage tapers towards rather narrow hind margin, not broad and parallel-sided (view from below). Variation On Sicily the more yellow ssp. sicilica occurs, having almost confluent abdominal

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oblique mark on side of S1

?

black occipital triangle

large black bar on frons

229

abdominal rings appear triangular from the side

apical spots on S5–8 absent

upper appendages rather parallel and well separated at base

lower appendage has narrow hind margin

basal tooth rather clearly visible, although present in all species

? appendages

rings on S5–8 that descend more towards the underside, and yellow markings on S9–10 present. Intermediates with ssp. bidentata have been found in Calabria, but as there is no genetic support, the validity of these subspecies is questionable. Regionally in Albania and southern Greece, but also further north in Slovenia, there are individuals with larger abdominal markings and sometimes a yellow foreside of the occipital triangle.

Occurrence Range and status Endemic to Europe. Generally more local than overlapping species of the boltoniigroup, but may be regionally common. Habitat Generally smaller streams than C. boltonii, including dripping rockfaces, near-vertical gulleys and calcareous springs. Flight season Mid-May until the end of August.

Cordulegaster Goldenrings

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Cordulegaster insignis 

Blue-eyed Goldenring

Schneider, 1845 blue eyes characteristic, but not present in all populations

yellow occipital triangle

230

? ssp. mzymtae ? ssp. charpentieri ? appendages lower appendage not tapering

basal tooth near S10

? ssp. insignis

C. helladica (EN)

Identification General The only Cordulegaster species with bluish eyes, although in parts of Turkey and on the Greek island of Ikaria green-eyed forms occur. Field characters (Ssp. insignis only) Male: Tot 71–78mm, Ab 52–58mm, Hw 40–46mm; female: Tot 71–83mm, Ab 51–63mm, Hw 41–49mm. Similar in size to C. bidentata but generally much paler. Differs by: (1) eyes blue; (2) frons anteriorly completely yellow, but may sometimes bear a small black marking; (3) occipital triangle is yellow, not black, and its backside is swollen, not flat; (4) abdominal markings are more extensive – median rings typically connected above on S2–8, apical spots present on S2–6 (occasionally also on S7 or only on S2) and S9 with a small basal marking. Regional forms with green eyes and often a black occipital triangle exist that can only be separated from C. helladica and C. bidentata by range and appendages. Only C. picta overlaps widely, but it has green eyes and boltonii-group characters (see first table on p. 221), with long, diverging appendages. Hand characters Male upper appendages resemble those of C. bidentata (parallel and widely separated at base), but the basal tooth is closer to S10 (view from side). The lower appendage is wider and more rectangular, not tapering distally (as in C. bidentata and C. helladica). Variation In the south-east Balkans and western and southern Turkey, the blue-eyed typical ssp. insignis occurs (see above). From south-east Romania ssp. montandoni has been described, but its only discriminating character is not useful, as a small black marking on the frons may also be present in typical ssp. insignis. In central Turkey (south of the mountains along the Black Sea), ssp. charpentieri is similar but rather small (male: Tot 64–67mm), with very large abdominal markings, those on S9–10 often connected to form ‘7’-shaped markings. Other, more local, forms are green-eyed: in north-east Turkey, possibly as far west as Samsun, the rather small (male: Tot 64–67mm) ssp. mzymtae has very small abdominal markings and a flat-backed black occipital triangle. In north-west Turkey, roughly between Bursa and Samsun, a similar form occurs with larger, but still rather small, abdominal markings. Individuals from the mountains north of Adana have a mixed set of characters, e.g. green eyes and larger markings

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than in typical insignis. Ongoing molecular work may reveal that certain forms are, in fact, species, whereas others are genetically indistinguishable and not recognisable even as subspecies. A large, green-eyed Cordulegaster from the Greek island of Ikaria, formerly considered nearest C. insignis, may be a hybrid between that species and C. helladica from the Cyclades (see that species).

Occurrence Range and status The zone separating C. bidentata and C. insignis probably runs from Kalkhidikhi to the Danube Delta. It is the only Cordulegaster species in much of central and southern Turkey. Habitat Smaller streams than C. picta, but occasionally in larger and deeper waters. Flight season Late May to mid-August.

Cordulegaster helladica 

Greek Goldenring

231

(Lohmann, 1993) Identification General Endemic to southern Greece. Resembles C. insignis, but is always green-eyed and is separated from it by the Aegean Sea and a large area of northern Greece, where C. bidentata occurs. Without information on locality, single individuals may be impossble to separate from C. insignis. Field characters Male: Tot 68–78mm, Ab 50–60mm, Hw 41–46mm; female: Tot 78–83mm, Ab 59–63mm, Hw 46–49mm. Differs from C. bidentata in much the same way as C. insignis (see that species): large abdominal markings (median rings dorsally connected on S2–6(7) and apical spots on S2–5), predominantly yellow foreside of costa, and the black bar across frons is small or absent. Like C. bidentata, it has green eyes and normally a black occipital triangle. Hand characters Upper appendages as in C. insignis; lower appendage not quadrate but tapering towards hind margin. Variation On the Peloponnese, Euboea (Evia) and areas between these, the ssp. helladica occurs, with the foreside of the occipital triangle normally black. Around Mt Parnassos (and perhaps a small part of north-eastern Peloponnese), ssp. kastalia has the stripe between the two large lateral bands on the thorax very large and almost rectangular (normally it is restricted to a small, often triangular, upper spot). The foreside of the occipital triangle is usually yellow and the yellow abdominal markings are more extensive. Both forms may coexist, although ongoing molecular work suggests that the two subspecies cannot be distinguished. In the Cyclades (Andros, Tenos, Naxos), ssp. buchholzi has smaller abdominal markings and a black or yellow occipital triangle. Formerly considered a possible subspecies of C. insignis (see that species), the green-eyed population on the nearby island of Ikaria may consist of hybrids between that

? ssp. helladica large stripe

ssp. kastalia

? appendages

lower appendage tapering

species and ssp. buchholzi. Individuals there are large (male: Tot 75–83mm) and have a black stripe across the frons, the foreside of the occipital triangle is black, the backside is flat, and abdominal markings are moderately extensive.

Occurrence Range and status This Greek endemic meets C. bidentata near Mt Parnassos (see blue on map, p. 230). Habitat As C. insignis. Flight season Mid-May to mid-August.

Cordulegaster Goldenrings

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Family affiliation uncertain

Oxygastra Orange-spotted Emeralds Selys, 1870 Oxygastra curtisii (Dale, 1834) Orange-spotted Emerald 232

Identification General A dark, slender emerald easily identified by the combination of the brilliant green eyes and the deep yellow streaks on the upperside of the dark, slender, clubbed abdomen. Males patrol short stretches of calm, tree-lined rivers. The only species of its genus worldwide, with venation and appendages that are unique among European dragonflies. Field characters Tot 47–54mm, Ab 33–39mm, Hw 33–36mm. Darker (the bright green eyes stand out more) and more slender than Cordulia aenea and Somatochlora metallica; the abdominal club lies nearer the tip (S8 widest). Face all dark and metallic (marked yellow in Somatochlora species). Thorax is metallic green, loosely covered with short

hairs (densely covered with long hairs in C. aenea). Wing bases with rather extensive areas of saffron; in females, this extends along the leading edges of the wings. Abdomen is dark metallic green, with a chain of orange-yellow spots on the upperside of S1–7 and S10. C. aenea males have an unmarked abdomen, with a thicker, less distal club (S7–8 widest). Somatochlora species are usually not clubtailed, and at most have paired yellow spots on the abdomen sides. Hand characters Male S10 bears a yellowish membranous crest on the upperside. Male’s upper appendages are blunt and somewhat divergent (view from above), each with an elongate ventral spine near the base (view from side). Lower appendage with a broad notched tip (view from below).

H Oxygastra curtisii male. Note the slender abdomen with the club-shaped tip and the pale ridge on S10.

Oxygastra Orange-spotted Emeralds

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triangles without cross-veins

233 membranule all-white

/

chain of yellow spots along upperside of abdomen

arculus and base of triangle in hindwing distinctly not aligned mature ?

dorsal crest

abdomen widest at S8

yellowish crest on S10

side view

? S10 and appendages

/ face entirely dark

Female’s appendages are short; vulvar scale tiny, almost invisible. Venation differs from corduliids: (1) anal loop consists of two long arched rows of cells, not foot-shaped; (2) all triangles without cross-veins; (3) Hw triangle is distinctly separated from arculus and shifted towards the wing tip; (4) membranule is all whitish, not dark-tipped; (5) anal angle of male Hw is rounded, less angular. Variation Intensity and extent of saffron tint on wing base varies, being more distinct in immatures. Behaviour Males leisurely patrol short stretches (up to 20m long) in sun or shade, in a straight or zigzag flight, up to 1m from the bank and rather low above the water. Eggs are deposited near the waterline, often under overhanging trees and in crevices, but also on roots, soaked logs, mossy rocks and floating mats of algae.

side of abdomen unmarked

/ appendages and vulvar scale inconspicuous

Habitat Slow-flowing tree-lined rivers and streams, rarely lake shores. Flight season From late May to the end of August; most abundant in July, maybe a month or more earlier in the south.  H Wildermuth

NT HD II+IV

Occurrence Range and status Endemic to south-western Europe and Morocco. Locally common; occurrence north of France is sporadic. Extinct in Great Britain.

Oxygastra Orange-spotted Emeralds

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Macromiidae

Macromia Rambur, 1842

River Cruisers (NA)

Macromia splendens (Pictet, 1843) Identification 234

Cruisers

General An enigmatic large dragonfly that cruises rapidly along the banks of calm river sections and reservoirs. Endemic to south-western Europe, but only rarely encountered. Its nearest relatives occur in North America and Asia. Unique in appearance, although its large size and yellow-marked black body is not unlike Cordulegaster at first sight. Field characters Tot 70–75mm, Ab 48–55mm, Hw 42–49mm. Similar in size to Cordulegaster but the abdomen is more slender and less clubbed. Large eyes are bright green. Face is dark with broad yellow markings, dorsally strongly inflated with a wide central furrow flanked by two large yellow spots. Thorax is dark metallic green, engirdled by a yellow band running between the wing pairs and through the metastigma, anteriorly with a pair of yellow streaks and a yellow semicircle in front of the Fw. Legs are long and spidery; hindlegs reach S5 when stretched. Abdomen black, upperside with yellow bar on S2, yellow spots S3–4(–6), becoming smaller rearwards, conspicuously larger spot on S7 and (in male only) a small one on S8. May be confused with Cordulegaster species in flight. Latter has eyes barely touching each other, no green lustre, shorter legs and protruding

VU HD II+IV

Splendid Cruiser

spike-like ovipositor, and differently configured yellow markings and venation. The genera regularly co-occur, but Cordulegaster makes slower and more concealed patrols and more frequently perches near water. Hand characters Rather unique in venation and appendages: 12–15 antenodal cross-veins on Fw; usually three cross-veins between triangle and wing base; triangles without cross-veins; Hw triangle far distal of arculus; anal loop almost circular. Appendages only about as long as S10: male’s uppers shorter than the triangular lower, each with a large external tooth. Female’s vulvar scale flat and broad, with a rather round border. Variation Yellow abdominal spots vary greatly in size; sometimes (almost) absent on S5–6. Behaviour A strong, seemingly tireless flier; seldom seen perched, hanging vertically from twigs. Forages along forest edges, near trees and over fallow land. Seen mainly over the water in the late morning and afternoon, preferring sunny and calm weather. Males patrol sections several hundred metres long, 1m or more from the bank and about 0.5m above the surface. May also be seen flying over paths or between trees near water, a few metres above ground. Ovipositing females skim over the water, tapping the abdomen tip onto the surface three to ten times at different sites close to each other.

Occurrence Range and status Found only in southern France and in the Iberian peninsula but very local and rare; always in small numbers. Habitat Warm watercourses with calm sections or dammed rivers in deep valleys, margins often with rocky and overhanging trees and bushes. May also colonise reservoirs with strongly oscillating water levels and no bankside vegetation. Flight season From the end of May to midAugust; mainly mid-June to mid-July, perhaps earlier in the southern Iberian peninsula. H Wildermuth

Macromia Cruisers

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distinctive facial markings yellow crescent in front of wings

235

venation unique (see p. 30)

/

?

large yellow spot on S7

metallic green lustre on thorax

large yellow band on thorax sides

?

lower appendage longer than uppers

very long legs

Macromia Cruisers

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Corduliidae

Cordulia Leach, 1815

Downy Emeralds

Cordulia aenea (Linnaeus, 1758) Identification 236

General Males of this compact, medium-sized, dark species restlessly patrol the edges of ponds and small lakes, with their clubbed abdomen slightly raised. Their flight is fast and close to the water’s surface, interrupted by bouts of hovering, when their brilliant green eyes flash up. A small genus; two virtually identical species occur in North America and eastern Asia. Field characters Tot 47–55mm, Ab 30–39mm, Hw 29–35mm. Somewhat smaller than Somatochlora metallica and similar in appearance, with a wholly dark metallic body of variable tone, but often less brilliant green and more bronzy. The thorax appears more fuzzy (more densely coated with long pale hairs) than Somatochlora species, and the wings tend to be tinged saffron at the base, especially in Hw. However, C. aenea is best separated by: (1) frons all dark, lacking yellow spots; (2) male abdomen clubbed, clearly thickest at S7–8 (except for the local S. borisi of the south-eastern Balkans, Somatochlora males have the abdomen

Downy Emerald

more evenly thickened at S5–7), and with large yellow (also whitish in female) markings beneath, especially near the base, but none on upperside. In Somatochlora, the abdominal markings are more fragmented and dispersed, e.g. spots may be visible from above on S2–3. Oxygastra has a dark frons and clubbed abdomen, but the abdomen bears a dorsal chain of yellow spots and the anal loop is not foot-shaped. Hand characters Hw with only one cross-vein between triangle and base, versus usually two in Somatochlora. Male appendages are unmistakable, the upper ones cylindrical and blunt, the lower deeply forked, both branches split into two strong hooks. Vulvar scale of female is short, flat and deeply notched. Behaviour Territorial males closely patrol the water’s edge, preferring small coves and avoiding deep shade and areas of dense vegetation. They seldom rest at the water, often feeding in nearby wooded areas. Relatively cold-tolerant, on the wing early in the year, often earlier and later in the day than

F Cordulia aenea male patrolling. Note the clubbed abdomen, bronzy tone, hairy body and absence of pale markings.

Cordulia Downy Emeralds

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downy thorax

saffron patches at wing bases

237

one cross-vein between triangle and hindwing base

/

?

no yellow areas on frons

noticeably clubbed abdomen, thickest at S7–8

/

large pale spots on underside of abdomen

vulvar scale not protruding

two hooks

blunt upper appendages and forked lower

? appendages

other dragonflies; also frequently active in cloudy weather. Females are secretive, ovipositing alone.

Occurrence Range and status Common and locally abundant in northern Eurasia. Strong populations mainly restricted to lowland, but in the south local and mostly restricted to mountain lakes. Habitat Standing waters, such as large ponds and small lakes in woodlands, bogs and heaths, but also oxbows, gravel pits, fish ponds, sluggish rivers and canals. Flight season Earlier than other emeralds: from late April to late August, mainly in May and June but not before mid-June in the far north.  H Wildermuth

Cordulia Downy Emeralds

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Somatochlora 

Striped Emeralds

Selys, 1871 Identification

238

Diagnosis Rather elusive medium-sized dragonflies, with largely dark bodies that have a metallic green lustre. Eyes reddish brown at emergence, becoming brilliant green. Frons dark metallic green with yellow spots on both sides. With the exception of the ‘Cordulia-shaped’ S. borisi, abdomens of males have the diagnostic ‘Somatochlora shape’: S1–2 bulbous, S3 waisted, S4–10 gradually widened up to about halfway (S6–7), and then gradually narrowed; not clubbed near tip. Abdomen is dark green to almost black, with restricted yellowish markings on the sides. Male appendages are rather long, often with up-curled tips and several irregular ventral teeth. Lower appendage is triangular, with a narrow, upcurved tip. Appendages of females are very long, their vulvar scale large, visible when viewed from the side, often shaped like a spout or trough, and distinctly projecting in most species. Separation from other genera Both Cordulia and Oxygastra have a uniformly dark frons, the

Cordulia aenea

S. metallica

male has a conspicuously clubbed abdomen (S7 and/or S8 widest) and a deeply notched lower appendage, and the female has an inconspicuous vulvar scale. Moreover, Cordulia has only one crossvein between the Hw triangle and base (not two) and Oxygastra has deep yellow streaks down the middle of the abdomen. Separation of the species This is a large genus, especially in North America (from where the misleading name ‘striped emeralds’ originates), and our only corduliid genus with more than a single species. Our seven species may be most easily determined in the hand by the shape of the anal appendages in males and by the vulvar scale in females. The patterns of yellow spots on the face, thorax and abdomen are also distinctive features. Behaviour Male patrols are swift and often stealthy or erratic. Females are especially shy, but may be detected by the rustling of wings while ovipositing alone under the cover of dense vegetation. Both sexes rarely perch near the waterside, but rather up in trees. H Wildermuth

S. arctica

S. borisi

Simple key to (pairs of similar) species. If the statement agrees, compare the given species. If it disagrees, then go to the next line.

1 2 3 4 5

Yellow spots (almost) connected on centre of frons. Abdomen strongly metallic green, never glossy black. / vulvar scale is a long, perpendicularly projecting spike.

metallica meridionalis

Most abdominal segments, as well as thorax, with lateral yellow spots.

flavomaculata

Yellow spots on frons continue onto postclypeus. ? abdomen club-shaped (S7–8 widest). South-east Balkans only.

borisi

? appendages incurved, like an earwig’s claspers seen from above. / S3 dorsally with two round yellow spots.

arctica

? appendages incurved at base and then up-curved and slightly outcurved. / S3 dorsally black.

alpestris sahlbergi

Somatochlora Striped Emeralds

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Somatochlora metallica 

Brilliant Emerald

(Vander Linden, 1825)

entirely metallic green

239

/

?

yellow spots on frons forming a bar

yellow spots at base of abdomen

triangular lower appendage

? appendages

/ abdomen tip

long vulvar scale perpendicular to abdomen

Identification General The largest and greenest Somatochlora species, along with its south-eastern counterpart S. meridionalis. The commonest Somatochlora species in much of Europe. Often flies in deep shade over lowland waters, but in full sunshine at mountain lakes. Field characters Tot 50–55mm, Ab 37–44mm, Hw 34–38mm. The largest emerald dragonfly (Somatochlora, Cordulia), together with the very similar S. meridionalis. Both are more brilliant green

than other Somatochlora species, being most notable for the shiny emerald abdomen, although this is not always reliable. Both also differ from other species on account of the female’s impressive vulvar scale (see below) and the yellow facial bar, in which the yellow sides of the frons are connected. This connecting bar is sometimes narrowly severed down its middle. The abdomen in both sexes has yellow spots at the base, smaller in males than in females. S. metallica and S. meridionalis hardly differ, the presence of yellow spots on the

Somatochlora Striped Emeralds

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240

thorax sides best separating S. meridionalis (see that species for more details). Often occurs with Cordulia aenea, which is also entirely metallic green, but Cordulia males are clearly club-tailed and their colour is less pure green, more bronze. However, both features may be hard to judge in flight. Hand characters Male’s upper appendages appear lean and jagged from above, divergent at base, allowing a clear view of the triangular lower appendage, then converging. Underside with two teeth. S. alpestris appendages are similar, but straighter and more parallel at base, while more bent towards their tips. Female’s vulvar scale is very large and spout-like, jutting out perpendicularly

from the abdomen, like a pick. Variation The extent and intensity of the abdominal spotting and amber at the Hw base varies, as does the colour of the Pt and the position of the teeth on the male’s upper appendages. The labrum has a vertical yellow bar in some individuals. Considering local variation, subspecies based on Pt and Hw colour (e.g. the Siberian ssp. abocanica, with a clear Hw base and black Pt) are doubtful, as is their occurrence in Europe. Behaviour Males secretively patrol over water close to the edge, and, in contrast to C. aenea, often fly under overhanging trees. Females oviposit alone by tapping the vulvar scale into water, mud, moss or other moist and soft substrates, preferring shady banks above the waterline.

Occurrence Range and status Common over a large part of Europe, especially throughout Fennoscandia; east to central Siberia. Restricted to mountain lakes in the south. Habitat Various standing and slow-flowing waters such as ponds, oxbows, rocky lake shores, moorland lakes, canals and sluggish rivers, but also in open tundra. Often favours some shading by trees and steep banks that suit oviposition, conditions often found in woodland. Flight season From late May to late September; most abundant from June to August. From the end of June in the far north.

F Somatochlora metallica male hovering while patrolling its territory. Compare colour and shape with Cordulia aenea on p. 236.

Somatochlora Striped Emeralds

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Somatochlora meridionalis Nielsen, 1935 Balkan Emerald single spot on side of thorax

/

241

dark pterostigma

? appendages

G Somatochlora meridionalis male. Note the pale spot on the side of the thorax.

Identification

Occurrence

General Replaces the very similar S. metallica in south-eastern Europe, where it is found mainly at shaded brooks. Can be distinguished only by close inspection of perched individuals or, better still, in the hand. The species’ status is much debated, because differences are very slight and individuals with mixed features occur. Field characters Tot 50–55mm, Ab 35–44mm, Hw 34–38mm. Similar to S. metallica in size and appearance. The presence of one, rarely two, yellow spots on the thorax sides is really the only usable field character. The Pt are a darker black, rather than orange-brown, although black Pt are also frequent in some S. metallica populations. The pale markings on S2–3 tend to be larger. Hand characters Male’s upper appendages are somewhat longer and thicker than in S. metallica, with the ventral teeth slightly stronger, but differences are often subtle, even in direct comparison. Vulvar scale is identical. Behaviour Males patrol long stretches of streams, following precisely its course at low height, often in deep shade, avoiding sunny spots and clearings. The fast flight is interspersed with hovering stops. May also be active in overcast weather.

Range and status Rather common in some parts of its range. Extends north to Austria, the Czech Republic, Slovakia and south-western Ukraine. S. metallica tends to inhabit higher altitudes in areas where the two species overlap. Endemic to our area. Habitat Unlike S. metallica, almost exclusively breeds in running water, usually heavily shaded lowland streams and rivers, but also sunny stretches at greater altitudes. Flight season Mainly from June to August.

Somatochlora Striped Emeralds

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Somatochlora flavomaculata 

Yellow-spotted Emerald

(Vander Linden, 1825)  Somatochlora flavomaculata pair mating.

242

Identification General Readily recognised by the string of yellow spots on each side of the abdomen. Often found patrolling woodland edges far from water. Field characters Tot 45–54mm, Ab 34–43mm, Hw 32–39mm. Medium-sized Somatochlora species; smaller than S. metallica, and nearer Cordulia aenea. Easily separated from other metallic green dragonflies by yellow spotting of the thorax and abdomen: thorax with two long spots on each side, S2–3 with large lateral spots, and S4–8 each with a pair of triangular paired spots at the segment bases. The spots are larger in females and brighter in young individuals. Oxygastra curtisii has dorsal yellow streaks on the abdomen but has no yellow spots on both sides of the frons. Hand characters Upper appendages of male are straight, smooth and almost parallel (view from

above), slightly convergent towards the up-curled tips, with a rather fine but prominent ventral tooth near the base (view from side). Vulvar scale of female is shorter than S9, a perpendicularly projecting trough with a rounded, notched margin. Variation Abdominal spots darken, and may become almost invisible, in mature individuals. Behaviour Males often defend their territories over dry vegetation or paths near bushes and trees, or in glades in reedbeds, but also patrol over small waterbodies, especially towards the end of the breeding season. Mating wheels may circle low over reedbeds for minutes at a time.

Occurrence Range and status Rather rare, but locally common in extensive marshy areas and mires in southern Fennoscandia and north-eastern Europe.

Somatochlora Striped Emeralds

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/

243

? lateral abdominal spots distinctive, but may be indistinct in older males

upper appendages straight and smooth sided

? appendages /

yellow spots on side of thorax and abdomen

vulvar scale shorter than S9

/ abdomen tip

Declining in much of its range, with only a very few highly isolated populations in the south. Range extends to north-eastern Kazakhstan. First recorded in Great Britain in 2018. Habitat Unlike most other Somatochlora species, has a relatively southern range, being rather typical of temperate valleys and lowlands. Inhabits marshes, wet meadows, mire lakes, bog edges and reedbeds (sometimes sluggish rivers or lake shores), where there are small, richly vegetated waters, such as ponds and ditches. Flight season From the end of May to midAugust; most abundant in June and July (not before July in the north).

Somatochlora Striped Emeralds

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Somatochlora arctica (Zetterstedt, 1840) Northern Emerald F Somatochlora arctica male. Note the sharp contrast between the green eyes and the glossy black body. Note also the distinctive pincer-like appendages and narrow-waisted abdomen.

244

Identification General A rather small, dark, elusive Somatochlora of bogs. Often occurs with the similar S. alpestris in boreal and alpine areas, but additionally survives in lowland bogs inbetween. It seems less critical of climate but more of habitat, the reverse of S. alpestris, having a larger but more fragmented range. Indeed, it is seldom common and is often under threat where it occurs. Field characters Tot 45–51mm, Ab 30–37mm, Hw 28–35mm. One of the smaller corduliids. Slightly more slender than S. alpestris, the male’s ‘waist’ (S3) is narrower, and lacks that species’ conspicuous white rings at the base of the abdomen (see S. alpestris for more comparative features). S. arctica can be separated from it and other Somatochlora species by the male’s unique appendages and, in the female, by the vulvar scale and markings: abdomen all dark from above, with

a distinct pair of orange-yellow spots at the base of S3, preceded by a yellow ring. Hand characters In contrast to S. alpestris, has only one cross-vein between the Fw triangle and base. The upper appendages of the male are calliper-shaped (view from above): parallel at the base, then arched and converging apically, recalling an earwig’s ‘claspers’. No teeth are visible from above, but it is irregularly toothed beneath. The vulvar scale of the female is a large trough with an obtusely pointed margin, hardly projecting from the abdomen and often extending beyond S9. Behaviour Forages between trees, often at treetop height. Males fly low over boggy or marshy vegetation with concealed puddles, often zigzagging or dashing to and fro. They also patrol small peat pools. Unlike S. alpestris, they avoid more open water, such as large ponds and lake shores.

Somatochlora Striped Emeralds

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two large spots on S3 with a yellow ring

245

relatively narrow waist

/

?

upper appendages shaped like callipers

/

? appendages

vulvar scale hardly protrudes

/ abdomen tip

Occurrence Range and status Usually rather local, mostly in small numbers and declining in many areas; more frequent and regionally common in Fennoscandia and north-eastern European regions with large bogs. Ranges to Japan. Habitat Breeds in tiny wet depressions, which can hardly be classified as waterbodies, in raised bogs and mesotrophic lowland moors. Habitats are often surrounded by coniferous forests, but may also be situated in open tundra. Flight season From mid-June to mid-September; from July to mid-August in the far north.

Somatochlora Striped Emeralds

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Somatochlora alpestris (Selys, 1840)

Alpine Emerald

usually 2 cross-veins between base and triangle

upper appendages doubleangled

246 pale ring between dorsally unmarked S2 and S3

/

? appendages

? projecting triangular vulvar scale

Identification General A rather small, robust, seemingly black Somatochlora with brilliant green eyes, seen over small waterbodies in central European mountains and in the far north. Recognised when perched by the bright ring near the abdomen base and the male’s appendages. Field characters Tot 45–50mm, Ab 31–36mm, Hw 30–34mm. Similar in size to S. arctica. In comparison with that species, the eyes tend to be blue-green rather than olive green, the thorax is more hairy and male’s S3 is constricted but less strongly so. The abdomen appears all dark from above (also in females) but the pale ring between

/ abdomen tip

S2 and S3 is whiter. Being brighter and set in a dark background, this ring is more conspicuous than in other Somatochlora species, except the very similar S. sahlbergi. Both species should be identified by the male’s appendages and female’s vulvar scale. Hand characters In contrast to other corduliids, usually has two, rather than one, cross-veins between the Fw triangle and base. The male’s upper appendages have a strongly double-angled outer border, convergent tips and two ventral teeth in the basal half. The female’s vulvar scale is large and triangular, reaching about halfway along S9 and projecting at a right angle to the abdomen’s axis. Behaviour Males hover or erratically zigzag low over pools for a few seconds or minutes, chasing conspecifics.

Occurrence Range and status Not rare in most suitable areas, but usually encountered as single individuals. Has a classic boreo-alpine range: found almost exclusively above 1,200m in central Europe, and above 60°N in the north, east to Japan. These thresholds may shift with climate change. Habitat Various stagnant waters (puddles, pools, tarns and small vegetated lakes) in bogs, moorlands, subalpine pastures, taiga and tundra. Flight season From mid-June to August or September, depending on altitude and latitude, e.g. from the end of June to mid-August in the far north.

Somatochlora Striped Emeralds

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Somatochlora sahlbergi Trybom, 1889 Treeline Emerald ? appendages

usually one cross-vein between base and triangle

pterostigmas rather pale

upper appendages sharply angled with hooked tip

pale ring between S2 and S3

247

/ abdomen tip

? vulvar scale not always projecting as shown here

tip of vulvar scale notched

Identification

Occurrence

General This species has the northernmost distribution of any dragonfly, found only in remote regions north of the Arctic Circle. Appears robust and black, very similar to the more common S. alpestris, and should be identified by the male’s ‘exaggerated’ appendage shape. Field characters Tot 48–50mm, Ab 32–35mm, Hw 30–33mm. Same size, with the same rather stout build and relatively hairy abdomen, as S. alpestris. Has a similarly dark, unmarked abdomen with a whitish or yellowish ring between S2 and S3. The Pt tends to be paler brown. See S. alpestris for separation of other northern corduliids. Hand characters Only one cross-vein between the Fw triangle and base, while S. alpestris mostly has two. The upper appendages of the male are rather massive and hairy, running almost parallel from the base, then abruptly bent inwards and downwards, with a sharply upcurled tip. The outer borders are smooth when viewed from above. The vulvar scale of the female is shorter and less protruding than in other northern Somatochlora species, its margin being notched rather than rounded or pointed. Behaviour Males are strong fliers, patrolling 30–50m sections of lake edges or flying criss-cross over the water 0.5–1m from the shore. Active only with sunshine and 20°C or more, suddenly vanishing with overcast weather. Unusually for a corduliid, may perch flat on the ground in cold wind rather like a gomphid.

Range and status In Europe very local in the far north of Fennoscandia and Russia, being a climate specialist restricted to the driest and coldest but sunniest parts of the subarctic. World range largely coincides with so-called palsa mires, characterised by raised peat hummocks with a frozen core, which develop only under these distinct climatic conditions in Alaska, Canada and Eurasia. Habitat Rather deep (50cm or more) ponds and lakes with boggy margins of sedges (Carex), cottongrasses (Eriophorum) or peatmoss (Sphagnum) in the transition zone between taiga and tundra, and further north in open tundra with low woody vegetation. Flight season July and August; most adults recorded in the last ten days of July.

Somatochlora Striped Emeralds

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Somatochlora borisi Marinov, 2001

Bulgarian Emerald  Somatochlora borisi male. Note the yellow spots on the abdomen, the unmarked thorax side and the yellow sides to the postclypeus.

248

Identification General First discovered in 1999, and so far found only in a very restricted area of the south-east Balkans, where it flies mostly in June on shaded, calm rivers. An aberrant Somatochlora, recalling Cordulia in some respects, from which it can be separated by the large yellow spots on the head and abdomen base. Although the genus Corduliochlora was created for this species, that name has not been generally accepted because more in-depth analysis of related genera is still wanting. Field characters Tot 45–50mm, Ab 34–37mm, Hw 31–34mm. Somewhat smaller than the cooccurring S. meridionalis. Similar in size and shape to Cordulia aenea. Unlike other Somatochlora species, the male abdomen has a thick clubtail, widest at S7–8. The frons is metallic green, with large yellow spots on the sides extending towards each other, but not touching as in S. meridionalis. The yellow extends onto the sides of the shining

black postclypeus, which is all dark in C. aenea and other Somatochlora species. The abdomen is unmarked except for conspicuous yellow spots at the base: the male has three on each side of S2 and two large basal spots on S3, and the female has four spots on the upperside of S2–3 and a chain of large spots on the underside of S2–4 and sometimes S5. Uniquely, the ventral spots on S3–5 are bordered with black below. In contrast with this extensive spotting, the thorax is entirely metallic green, lacking spots on the sides (unlike S. meridionalis). Hand characters Unlike C. aenea and like typical Somatochlora, it has two cross-veins between the Hw triangle and base. The male’s upper appendages are unique: bent inwards and then outwards (view from above), with thick and blunt down-curved tips (view from side) and two small ventral teeth near the base. The lower appendage is rather like that of other Somatochlora species,

Somatochlora Striped Emeralds

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spots on S2 and S3

yellow patches on sides of frons extend to but do not meet at middle

? thorax sides unmarked

yellow sides of postclypeus diagnostic

249

/

abdomen thickest at S7–8 yellow spots on underside bordered with black ventrally

upper appendages with thick, blunt down- and outcurved tips

? appendages

short vulvar scale split almost to base lower appendage with broad, notched tip

/ abdomen tip (dorsal view)

but with a broader and slightly notched tip (not forked as in C. aenea). Genital lobe with a projecting point, unlike any other Somatochlora species (not illustrated). Vulvar scale of female is

VU

less than half as long as S9, semicircular, with a very narrow slit from the apex almost to the base (view from below). Variation Abdominal spots may vary in shape and size. Behaviour Males seem to defend territories over water.

Occurrence Range and status Endemic to Thrace, the southeastern corner of the Balkans between the Aegean and Black seas, where Bulgaria, Greece and Turkey meet. Found only in small numbers. Habitat Shaded sections of slow and often intermittently flowing rivers and streams, in wooded areas below 300m altitude with a hot and moist climate. Flight season Early, like C. aenea but unlike other Somatochlora; emerges from early May onwards and is mainly seen until the beginning of July.

Somatochlora Striped Emeralds

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Epitheca Burmeister, 1839 Baskettails Epitheca bimaculata (Charpentier,

1825)

Identification

250

General Baskettails are large, strong fliers, most often seen cruising over open water, rarely perched. The single European species (more occur in Japan and especially North America) is a shy brownish corduliid, with boldly marked wings and without metallic green colours. Although its sustained flight, vertical resting posture, lobed eye-margin, long legs, and male’s anal triangle and tibial keels make it a typical corduliid, confusion with Libellula species is more likely. With its larger size and habit of roving about over water far from the edge, the ‘twinspot’ can best be observed with binoculars. Field characters Tot 55–65mm, Ab 37–43mm, Hw 36–44mm. Larger than any co-occurring corduliid or libellulid. Unlike other corduliids, the body is brown, black and yellow, without a metallic lustre, and when mature the eyes are greyish blue-green

Eurasian Baskettail Two-spotted Dragonfly rather than vivid green. Face is yellow, with a contrasting black labrum and upper frons. Thorax is brown-yellow, with bold black bands. Wings are lightly smoky, with a large dark brown area on the Hw base. Abdomen is awl-shaped in the male, broad-based but terminally very slender; black with a yellowish base, S3–8 (also S9 and sometimes S10 in female) largely black with yellow to orangebrown lateral spots. Appearance recalls Libellula quadrimaculata and (unpruinose) L. fulva, but both of these are distinctly smaller and stay more to the water’s edge, fly low and frequently perch (in a less penduline position). L. quadrimaculata has black spots on nodes, L. fulva often has black on the wing tips. Hand characters Venation similar to Cordulia and Somatochlora. Triangle usually three-celled in Fw and two-celled in Hw, rather than two- (sometimes three-) and one- to two-celled, respectively. Legs

H Epitheca bimaculata female. Note the long legs and the smoky tint on the wings of this freshly emerged individual.

Epitheca Baskettails

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251

black patches on hindwing bases

immature /

mature ?

tapering abdomen

V-shaped appendages

much longer than in Libellula, the hind pair reaching S6 when stretched. Male’s upper appendages are long and divergent, like a ‘V’ or fishtail when viewed from above. Note that L. quadrimaculata also has splayed appendages. Lower appendage with a broad, deeply notched tip. Hamules massive, enveloped in huge, sheath-like genital lobes (view from side). Female’s equally unique vulvar scale is huge and V-shaped (view from below), its two lobes almost reaching S10’s end (not illustrated). Variation The intensity of the yellowish tint on the wings varies with age and sex. Behaviour Territorial males fly over open water steadily and persistently, fast and rectilinear, seldom closer than 3m to the shore, 30–50cm above the water. Co-occurring corduliids also have sustained patrols but stay closer to the water’s edge and surface. Females carry an egg mass under their upcurved abdomen tip while searching for a suitable oviposition site. The ball of eggs unravels into a gelatinous strand when it is deposited in flight on floating plants.

and elusive behaviour. More easily recorded by searching for its large, spidery, spiny exuviae in May. Occurs east to Japan. Habitat Small and rather deep lakes, meso- to eutrophic, with much submerged and floating vegetation, containing fish, and partly or completely surrounded by trees or bushes, e.g. oxbows, forest lakes, gravel pits and fish ponds. Flight season From mid-May to the end of June. Mid-June to August in the north.  H Wildermuth

Occurrence Range and status Generally very local, but may be abundant in regions with many suitable sites. Often overlooked because of its short flight season

Epitheca Baskettails

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Libellulidae

Libellula Linnaeus, 1758

Chasers F Libellula pontica male. Note the biting midge Forcipomyia paludis near the base of the right forewing.

252

Identification Diagnosis Bulky, medium-sized dragonflies, with diagnostic large triangular blackish patches at the Hw base, normally crossed by pale veins. With 12–20 antenodal cross-veins, and the only libellulid genus in our area with always more than one cross-vein in each so-called bridge space (long triangular space below the subnode). Separation from other genera Leucorrhinia is the only genus with dark Hw patches whose distributions overlap widely, but in these the patches are smaller and not pale-veined. The species are also much smaller, have seven to eight antenodal cross-veins and bright white snouts. Mature males of Diplacodes lefebvrii and Trithemis festiva have dark Hw patches but are much

smaller, sleeker and darker. Orthetrum species can be similarly robust and pruinose, but never have more than a slight yellow colouring at the wing base. Separation of the species Identification is straightforward: by abdomen shape, wing patterns and body colours. The three widespread species also have rather different habitat preferences. Behaviour The aggressive males chase off rivals with fast, defensive flights undertaken from prominent perches, such as stakes. Egg-laying females are usually guarded by the male hovering overhead, although L. fulva and L. pontica males more frequently leave the female after some time to oviposit by herself.  K-D B Dijkstra

Simple key to (pairs of similar) species. If the statement agrees, compare the given species. If it disagrees, then go to the next line.

1 2 3

All wings with blackish spots at node (on leading edge, midway between base and tip). Fw base amber but without blackish streaks or patch. Abdomen never pruinose.

quadrimaculata

Fw and Hw with equally large, dark basal dark patches, covering median spaces and Hw triangle. Abdomen very broad, often with yellow sides.

depressa

Forewing at most with small basal dark streaks; median spaces and Hw triangle clear. Abdomen not so broad, never with yellow sides.

fulva pontica

Libellula Chasers

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Libellula quadrimaculata  Linnaeus, 1758

Four-spotted Chaser Four-spotted Skimmer (NA)

dark spots on node of each wing

253 black base on hindwings only

immature /

mature ? abdomen tapered, sides narrowly yellow

f. praenubila

S6-10 black

Identification

Occurrence

General Common by still waters in most of Europe. Easily recognised by its large size, brown body and the black wing spots for which it is named, one at the node of each wing. Field characters Tot 40–48mm, Ab 27–32mm, Hw 32–40mm. As long as L. depressa, but more slender and appearing smaller. Body largely and uniformly translucent brown in both sexes. Abdomen tapered, terminal three-fifths (S6–10) black, sides S4–8 narrowly yellow. Male abdomen does not become bluish-grey pruinose like L. depressa and L. fulva. All wings deeply amber at base, with unique black spot at node. Hw base with large blackish triangles but, unlike other Libellula species, Fw is amber at base rather than black. Hand characters Upper appendages rather long in both sexes and splayed in male. Variation Extent of wing markings is variable, especially at the node. Dark specimens with large nodal spots as well as brown bands behind the Pt are known as f. praenubila. Behaviour Males have a powerful, aggressive flight and frequently perch on stakes overlooking the water, unlike the similarly coloured Epitheca bimaculata, for which it may be mistaken at a distance.

Range and status Widespread and abundant in much of range, across temperate Eurasia and North America. In the past was known to have massive migrations in Eurasia. Habitat Most still waters, preferably with welldeveloped vegetation, and can be very numerous on acidic lakes. Flight season From late April to mid-September, but most abundant in early summer.

Libellula Chasers

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Libellula depressa Linnaeus, 1758 Broad-bodied Chaser F Libellula depressa male. Note the pale antehumeral stripes, wing markings and very broad abdomen.

254

Identification

Occurrence

General A large, pale blue male of this species, aggressively defending a garden pond or small natural pool, is a familiar sight in much of our area. The abdomen shape and four large black areas at the wing bases instantly distinguish this species. Field characters Tot 39–48mm, Ab 22–31mm, Hw 32–38mm. Appearing very robust because of its remarkably broad, flattened abdomen. Fw with black bars at base; Hw base with triangular black patches. Markings reach about equally far in Fw and Hw, to the outer corner of the triangles. Female and immature male are quite evenly yellowish brown, the front of the thorax with two broad whitish antehumeral stripes and the sides of S3–8 narrowly yellow. Both the latter markings can disappear with age, especially in the male, where S3–9 become blue-grey pruinose. The wing bases are more extensively marked and the abdomen is broader than Orthetrum or other Libellula species. Hand characters The underside of S1 in the male (anterior to secondary genitalia) bears two robust spines, a structure not found in any other European dragonfly. Behaviour Males make fast, direct dashes from a conspicuous perch, often controlling an entire pool by fiercely chasing off competition.

Range and status Among the commonest species in much of Europe, extending to Central Asia. A strong flier that is often the first species to claim newly created or cleared habitats. Habitat A wide range of mostly stagnant waters, especially favouring those that are small, shallow, sunny and bare, such as cattle-drinking pools or quarry lakes. Flight season From late April to mid-September; most abundant in May and June.

Libellula Chasers

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broad antehumeral stripes

extensive dark wing bases on both wings

255 mature ?

mature /

old /

very broad abdomen becomes pruinose blue in male, concealing characteristic yellow sides

mature ?

two spines in front of secondary genitalia

immature ?

 Libellula depressa female. In flight it can look like a giant wasp!

Libellula Chasers

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Libellula fulva Müller, 1764

Blue Chaser Scarce Chaser F Libellula fulva female. Note the orange colour and dark wing tips.

256

 Libellula fulva male. The hint of orange on the abdomen and pristine pruinosity (no scraping marks following mating) indicate it is immature.

Identification General A local species throughout our area that prefers slow-moving waters, bordered with tall emergent vegetation. The appearance of the male changes dramatically with maturation: vivid orange tenerals turn into black and grey-blue territorial males. Nonetheless, both extremes are easily recognised by the coloration of the body and wings. Field characters Tot 42–45mm, Ab 25–29mm, Hw 32–38mm. Shorter-bodied but more bulky than Orthetrum cancellatum. Both sexes are entirely bright tawny orange when immature, with a black stripe down the length of the abdomen. Combined with the dark wing bases (including dark streaks at the Fw base), dark wing tips (especially in females) and the glaring yellow central veins of the wings, this renders immatures unmistakable. Females darken to a dull brown, males blacken entirely and develop greyish eyes

and grey-blue pruinose S3–7. The mature male’s larger black abdomen tip and glassy grey eyes separate it from other Libellula species, the black wing markings from Orthetrum, and the black snout from both. Hand characters Not important. Variation Although a feature seen in all-pruinose male libellulids, the dark scraping marks on S5–6

Libellula Chasers

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blue-grey eyes and black face dark streak

dark hindwing bases

immature ?

blue pruinose abdomen

bright orange abdomen with black dorsal stripe

males may have dark wing tips

257

mature ?

S8–10 black

old /

dark wing tips

mature /

inflicted by the female’s legs during copulation are especially clear in L. fulva. Pruinosity extends onto S9 in Turkish males. Behaviour Males frequently perch on waterside plants, making darting flights between perches.

Occurrence Range and status Widespread but habitatspecific, therefore only locally common. Range extends to Caspian Sea. Habitat Appears to require a certain combination of water quality and habitat structure, the latter usually borders of reeds or other rich riparian growth. Habitats include slow-flowing rivers

and streams, abandoned canals, reedy lakes and ditches, oxbows and fish ponds. Flight season From late April to early August, but activity is concentrated in May and June in most of range.

Libellula Chasers

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Libellula pontica Selys, 1887

Red Chaser black pterostigma

red frons

no dark streak

258

white membranule

? secondary

mature ?

genitalia

red abdomen with narrow black dorsal stripe

Identification

Occurrence

General Small counterpart of L. fulva, which it replaces to the south-east, although these twin species overlap widely in Turkey. Both are very similar at emergence, but L. pontica males become brick red on the face and abdomen when mature, not pruinose. The first impression of the species is thus more of a Sympetrum or Crocothemis than a Libellula. Field characters Tot 39–42mm, Ab 22–26mm, Hw 29–32mm. Noticeably smaller and somewhat more slender than L. fulva. Immature males and females differ from L. fulva in their size, narrower black abdominal stripe, the absence of dark wing tips and dark streaks at the Fw base, and the presence of a whitish membranule. The latter is grey in L. fulva and this does not stand out, while it contrasts with the dark Hw base and brown to red abdomen in L. pontica. Mature males with a brown thorax and red face and abdomen (retaining black stripe over its length). Similar to other red libellulids, especially the broad-bodied Crocothemis erythraea. Diagnostic features are the black Pt, dark triangular patches at Hw base, unmarked brown thorax and the black line along the full length of the abdomen. Hand characters Structurally, essentially identical to L. fulva.

Range and status Local in our area; range extends to the Transcaucasus, Iraq, Iran and northern Israel. Habitat Mainly slow-flowing waters with dense bankside vegetation, such as reedy streams, channels and ditches; also outflows of wellvegetated gravel pits and ponds. Flight season From early May to mid-July.

NT

Libellula Chasers

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Orthetrum Newman, 1833

Skimmers

Identification Diagnosis Almost every blue to grey pruinose dragonfly with clear wings will be an Orthetrum male. The combination of a clear Hw base (at most yellow-tinted) and 10–14 Fw antenodal crossveins (thus last one is complete) is unique for a libellulid. Separation from other genera Libellula species are similar in venation, stature and pruinosity, but have large black patches at the Hw base. Pruinose Leucorrhinia species are dark, with a contrasting G Orthetrum brunneum pair mating. The female and back of white face and also dark Hw the male’s head show the contrasting markings of the Sardinian patches. The rather plain brown subspecies cycnos. females and young males of most Orthetrum species are often mistaken for This is especially true for males, whose markings Sympetrum and Crocothemis. The species in both have become obscured by pruinosity, thus these genera normally have the last antenodal appearing very unlike females and younger males. cross-vein incomplete; Sympetrum has only six For convenience, the species can be divided into to seven complete Fw antenodal cross-veins, three groups (see key below). In most of Europe, and Crocothemis has large yellow patches at the separating O. albistylum from O. cancellatum Hw base. Especially fresh yellow O. cancellatum (group 1), and O. brunneum from O. coerulescens and the club-tailed black-and-pale O. sabina (group 3), will cause the greatest difficulty. In the superficially resemble gomphids, but their eyes Mediterranean, two diagnostic species in group touch each other. 2 and four additional species in group 3 occur. Separation of the species Some 60 Orthetrum Because up to five species of group 3 may occur species occur in Australia, Eurasia and particularly together in the Iberian peninsula, North Africa Africa. The species are often difficult to identify. or Turkey, and identification is difficult, the table

259

Simple key to similar species. If the statement agrees, compare the given species. If it disagrees, then go to the next statement. Table applies to both sexes.

1

Pt blackish, not yellow to brown. When not pruinose, abdomen with two thick longitudinal lines. When abdomen pruinose, terminal third is black.

albistylum cancellatum

2

Abdomen thin and at least as long as Hw, with S1–3 swollen and three times as thick as other segments. When not pruinose, abdomen black with pale streaks or rings. When pruinose, abdomen very dark blue-grey. Total length never under 43mm.

trinacria sabina

Abdomen thick and at most as long as Hw; not strongly swollen at base. When not pruinose, abdomen brown with black lines. When pruinose, abdomen pale blue to grey. Total length never over 52mm.

brunneum chrysostigma coerulescens nitidinerve ransonnetii taeniolatum

3

Orthetrum Skimmers

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below summarises the main characters of these species. Examination of the male’s secondary genitalia with a hand lens will be most decisive. Behaviour Active species, often perching on the

260

ground or stones. Male guards the female during oviposition, usually by hovering above her and chasing off rival males.  V J Kalkman

Main characters of group 3 Orthetrum species. See species texts for details. Explanation of characters: (1) Only two species are widespread (Ws), the other three are confined to the south (S), with one only in western (SW) and eastern Mediterranean (SE) respectively. (2) Middle size has Hw and total length around 30mm and 40mm. Small and large are usually at least 2mm above or below that. (3) M  iddle-sized Pt is around 3mm, and just over a tenth of Hw length, but Pt can range from 2–5mm and 7–13%. (4) R  spl (check all wings) subtends 1 or 2 rows of cells (seldom more than three or less than four cells doubled respectively). (5) Membranule pale (whitish) to dark brown. Range (1)

Diagnostic characters

Size (2)

Pt (3)

Rspl (4)

Memb (5)

Species

Ws

Face never pale blue

middle

large

1

interm.

coerulescens

Ws

None

large

small

2

pale

brunneum

S

Thorax side with one pale, dark-bordered stripe. Abdomen waisted, not parallel-sided

middle –large

middle

1 (2)

dark

chrysostigma

SW

Radius between base and node contrastingly yellow, not dark. Lower border S8 of / straight, not curved

large

large

2

pale

nitidinerve

SW

Black subcostal cross-veins between wing basis and node

large

small

1

pale

ransonnetii

SE

Thorax side with two pale, dark-bordered stripes. Upperside of eyes deep brown, not grey.

small

small

1 (2)

interm.

taeniolatum

Orthetrum cancellatum 

Black-tailed Skimmer

(Linnaeus, 1758) Identification General Known as ‘blue arrows’, the males of this active species, sheering fast and low above water or perching on open ground beside it, are one of the most familiar dragonfly sights in our area. Field characters Tot 44–50mm, Ab 29–35mm, Hw 35–41mm. Slightly longer than Libellula depressa and L. quadrimaculata. The only other libellulid that combines clear wings, black Pt, and an abdomen with two thick lengthwise black stripes that becomes grey-blue pruinose with a black tip

(S(6–)8–10) in males is O. albistylum; on Sardinia and Corsica, females and immature males of O. brunneum may look similar (see these species). O. cancellatum is altogether darker and more uniform, without pale bands or appendages. With their yellow bodies marked with a black latticework, tenerals are often mistaken for Gomphus species. These, however, are much less robust and the abdominal black stripes are much thicker than the yellow stripe in between, instead of vice versa. Hand characters Male’s hamule has a very large

Orthetrum Skimmers

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black pterostigma

yellow abdomen with longitudinal black bands

261 mature ?

/

black markings and eventually yellow sides disappear under bluish pruinosity in older males pointed anterior lamina immature ?

black tip is retained even in very pruinose males

rounded hamule

? secondary genitalia

/ old /

and rounded head, the anterior lamina consists of two sharp points (view from behind). Variation The abdomen of older females becomes brown and often thinly pruinose. Behaviour Male makes fast, skimming patrols and often rests on open patches on the bank.

Occurrence Range and status Found throughout our area, with the exception of northern Fennoscandia, extending to northern China and Mongolia. One of the most common species. Habitat Larger standing or slow-flowing waters, generally open and often with unvegetated margins, such as lakes, sandpits, rivers and canals; in Finland and adjacent region, mainly in bays of the Baltic Sea.

Flight season From the end of April to the beginning of September; most abundant from June to August.

Orthetrum Skimmers

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Orthetrum albistylum (Selys, 1848) White-tailed Skimmer Identification

262

General Very similar to O. cancellatum, with which it is found especially in the south-east, and as far west as France. However, it is sleeker, paler and more contrasting. Named for the contrasting white appendages of both sexes, but beware occasional males with black appendages! Field characters Tot 45–50mm, Ab 30–37mm, Hw 33–38mm. As large as O. cancellatum but less robust, with a narrower abdomen. As with O. cancellatum, has black Pt and two distinct lengthwise black stripes on the abdomen that

disappear under pruinosity as the male ages, S(6–)8–10 becoming black. It can be difficult to distinguish it in flight, but O. albistylum has the following: (1) whitish appendages in both sexes, a character that can be indiscernible in the male, while the female’s S10 is also white; (2) two contrasting milky bands on the sides of the thorax and a pale stripe running dorsally over the thorax between the wings; (3) ground colour of abdomen paler and the black lines on each segment slightly more curved; (4) mature male’s pruinosity paler, almost white; (5) relatively larger Pt.

F Orthetrum albistylum female. Note the white appendages, curved bands on the abdomen and black pterostigmas. Hand characters Secondary genitalia much like those of O. cancellatum, but hamule more pointed and anterior lamina less deeply bifid. Behaviour Like O. cancellatum, male often sits on open ground near the water, making very fast, low flights over the water.

Occurrence Range and status Distribution is patchy, but the species is generally not uncommon, stretching to China and Japan. May be expanding gradually, e.g. first found in Belgium in 2016. Habitat Open ponds and lakes. Flight season From the end of May to mid-September.

Orthetrum Skimmers

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black pterostigma

263

mature ?

/

longitudinal bands on abdomen noticeably curved

black tip to abdomen

S10 and appendages white

abdomen narrower than O. cancellatum and pruinosity whiter

white appendages

? secondary

relatively pointed hamule

genitalia whitish bands on sides of thorax

/

old /

 Orthetrum albistylum male, showing the contrasting white appendages and the pale bands on the thorax.

Orthetrum Skimmers

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Orthetrum coerulescens (Fabricius, 1798) Keeled Skimmer F Orthetrum coerulescens female. Note the thin dorsal line with ‘crossbars’ and the large yellow pterostigmas.

264

Identification General The common small Orthetrum of flowing water throughout much of our area. Often found with the similar O. brunneum and several other lookalikes in the far south. Differences are summarised in the table on p. 260. Field characters Tot 36–45mm, Ab 23–38mm, Hw 28–33mm. About the size of Sympetrum striolatum. In most areas, the mature male is separated from all libellulids, except O. brunneum, by the all-pruinose abdomen and yellow Pt, but young males and females are often confused

with Sympetrum (see description of O. brunneum for a full comparison, and descriptions of O. chrysostigma, O. nitidinerve, O. ransonnetii and O. taeniolatum for their distinctive characters). O. coerulescens is characterised by its rather small size, fairly slender, tapering abdomen and large Pt (around 4mm long). Unlike similar species, the mature male develops little pruinosity on the thorax, and has a brown face. It thus appears twocoloured rather than all blue (but see Variation). Usually, two pale stripes on the front of the thorax are visible, but the sides are a uniform colour. Hand characters Almost invariably has Rspl subtending a single row of cells. At most, five cells are doubled, whereas similar species often or always have more doublings. Anterior lamina of male dwarfs hamule by comparison with similar species (but see Variation). Variation Wing bases are often suffused with amber. Two subspecies are recognised. The southern ssp. anceps has been considered a species, formerly often called O. ramburii, but this is unjustified as intermediate individuals occur in large areas. Seen from the side, typical ssp. coerulescens males have the anterior lamina standing nearly perpendicular to the abdominal axis. Its apex is inflated and rounded. Typical ssp. anceps has a more pointed and triangular

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pale antehumeral stripes

large yellow pterostigma

brown face

Rspl usually subtends single row of cells (only one doubling here)

thin dorsal line with ‘cross-bars’, sometimes separated into pairs of dots

265

mature ?

/

no black tip to abdomen

thorax and abdomen may be all-pruinose in south of range immature ? (notably dark individual)

mature ?

ssp. coerulescens

old / anterior lamina with rounded tip

/

ssp. anceps

anterior lamina triangular

? secondary genitalia

anterior lamina, which is directed more backwards. Southern males (thus including ssp. anceps) tend to be more densely and extensively pruinose and may seem totally blue. Behaviour Normally sits on vegetation, seldom on the ground.

Occurrence Range and status Common around the Mediterranean, but generally local in central and northern Europe. Typical ssp. anceps is found in

north Africa, Asia (to northern India), Sardinia and the southern Balkans. Intermediate forms predominate in the Iberian peninsula, Sicily and large parts of south-eastern Europe. Elsewhere, typical ssp. coerulescens is found. Habitat Running waters, such as streams and ditches. In north of range, mainly runnels in boggy areas. Flight season From April to November; most abundant from June to August.

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Orthetrum brunneum  Southern Skimmer (Fonscolombe, 1837) relatively small brown pale blue face

pterostigma

Rspl subtends double row of cells (6–7 cells doubled here)

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/

mature ?

? secondary genitalia thin dorsal line flanked by pairs of spots on each segment

all-blue abdomen and thorax

comparatively low anterior lamina

large hook on hamule

/ ssp. cycnos

/ abdomen tip

S8 expanded ventrally

Identification General A rather plain, medium-sized skimmer that is common on running waters in the south. Males become totally blue, females are brown overall. Being somewhat featureless, it is most easily separated from similar species around the Mediterranean by exclusion.

Field characters Tot 41–49mm, Ab 25–32mm, Hw 33–37mm. Size and shape between O. cancellatum and O. coerulescens. Distinguished from the former (and O. albistylum) by its reddishbrown Pt and almost unmarked body, which becomes largely blue in mature males. Most likely to be confused with O. coerulescens in most of its range, but visual distinguishing features are: (1) somewhat larger and heavier, with a broader abdomen; (2) adult male has a whitish face with a blue hue, rather than a dirty pale brown; (3) thorax is plain brown, becoming entirely blue pruinose in mature male (in O. coerulescens the thorax seldom becomes densely pruinose and two paler antehumeral stripes are often visible); (4) when not pruinose, the abdomen is uniformly brown with a thin lengthwise black line down the centre (except for Sardinia and Corsica; see Variation), which is flanked by two spots near the end of each segment (these spots are usually fused to a cross-bar in O. coerulescens, but seldom so in O. brunneum); (5) when mature, the Pt is reddish brown, rather than yellow as in O. coerulescens. These characters

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should be used as an indication of the identity of individuals (4 and 5 especially are not fully reliable), and identification in the hand is recommended. See O. nitidinerve, O. chrysostigma and O. taeniolatum for their separation. Hand characters Additional distinguishing features are: (6) between IR3 and Rspl, four to nine (rarely less) cells are doubled; there are no (rarely one to five) doublings in O. coerulescens; (7) Pt is relatively short (2.5–3mm); (8) the male’s hamule has a very large hook, and unlike in other Orthetrum species it juts out well beyond the anterior lamina. Variation On Sardinia and Corsica, ssp. cycnos has distinct brown lengthwise stripes on the abdomen, resembling O. cancellatum or dark O. coerulescens;

Orthetrum nitidinerve 

in males this pattern disappears under pruinosity with maturity. However, both sexes of all ages show bold black and white bars behind the eyes, unlike the yellowish and brown pattern in ssp. brunneus. Behaviour Often perches on the ground or on stakes beside running water.

Occurrence Range and status Common around the Mediterranean; range extends to eastern China. Has expanded northwards since the 1990s. Habitat Mostly small streams, running ditches and seepages, preferring scantily vegetated sites more than O. coerulescens. Favours bare runnels in the north, e.g. in chalk or marl quarries. Flight season From April to September.

267

Yellow-veined Skimmer

(Selys, 1841) large yellow pterostigma radius yellow from base to node costa yellow from node to pterostigma

/

mature ?

? secondary / abdomen tip

genitalia

S8 not expanded ventrally, its sides as straight as S7

Identification General Fairly large Orthetrum with large golden Pt, endemic to the western Mediterranean. Named for, and best identified by, its thick, bright yellow veins. Field characters Tot 46–50mm, Ab 28–33mm, Hw 31–38mm. Similar in size, shape and general appearance to O. brunneum. Distinguished from that species, O. coerulescens and O. chrysostigma by: (1) radius (third longitudinal vein counted from leading edge of wing) is yellow from base to node (in other species this is brown to black, as are the

other longitudinal veins), and the costa between the node and Pt is similarly bright, standing out more than in other Orthetrum species; (2) the yellow Pt are notably large (about 4–5mm) (those of O. brunneum are clearly smaller and darker in the field); (3) body becomes wholly whitish-blue pruinose in males, a shade paler than other species. Females and young males are very plain, typically without obvious pale or dark markings, other than a thin black central line on the abdomen. Hand characters Unlike the similar species, S8 is

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not expanded in the female, its lower border being straight rather than convex. Behaviour Females oviposit while perched on a stone, dipping their abdomen into the water and attaching gelatinous strands of white eggs to the stone, with the guarding male perching nearby. This behaviour is exceptional among Libellulidae.

Occurrence

268

Range and status Endemic to our area. Most common in north-west Africa, more local in Iberian peninsula, Sardinia, Sicily and near Naples. Habitat Mainly small running waters (springs, streamlets, seepages) with marshy vegetation in rather arid areas, but sometimes larger waterbodies, like pools of intermittent streams and rivers. Flight season From April to October.

Orthetrum chrysostigma 

Epaulet Skimmer

(Burmeister, 1839) Identification General One of tropical Africa’s commonest dragonflies, extending to Rhodes and the Iberian peninsula. Usually easily distinguished by its ‘epaulette’: broad white badges bordered with black on each ‘shoulder’. Field characters Tot 39–48mm, Ab 26–33mm, Hw 27–32mm. Size near O. brunneum and especially O. coerulescens. One of several similar species

with a largely pruinose body in the mature male and a yellowish Pt. Diagnostic features are: (1) abdomen rather slender and waisted near base, and constricted between S3 and S4 when seen from above; (2) thorax with a conspicuous whitish stripe beneath each Fw base, accentuated by black lines on both sides (note that this is obscured in old males); (3) membranules are sooty and usually bordered by a small yellow patch. In other species,

dark membranule with small yellow patch in adjacent hindwing

? secondary genitalia

waisted abdomen broad hook on hamule

/

mature ?

distinctive white ‘epaulet’ bordered with black

lobe small but distinct

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are less white. In the even smaller O. taeniolatum, there are three stripes on each side, and these are accentuated by a black line on only one side. Both smaller species are unwaisted and have grey-brown membranules. Hand characters Hamule has a broad hook that dwarfs a small, but separate, lobe beside it. Behaviour In Turkey, said to perch rarely in the obelisk position, in contrast to O. taeniolatum.

C

Occurrence

the Hw base is generally clear. The larger, broaderbodied and unwaisted O. brunneum, O. nitidinerve and O. ransonnetii lack pale stripes on the thorax, and have whitish membranules. In the smaller O. coerulescens, the stripes lie more anteriorly and

Range and status Ubiquitous outside forests throughout Africa. Not uncommon in the southwestern Iberian peninsula, the Canaries, coastal Turkey and adjacent Greek islands. Habitat Larger streams and shallow still waters in open landscapes. Flight season In Turkey, recorded from April to the end of August. In north-west Africa, up until November and suspected to be on the wing throughout the year.

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Orthetrum taeniolatum (Schneider, 1845) Small Skimmer Identification General A diminutive Orthetrum, with a pretty pattern of milky stripes on the thorax that, like the whole body, becomes pale blue pruinose in old males. Inhabits the dry lands of Asia, just reaching the Aegean.

Field characters Tot 33–38mm, Ab 21–26mm, Hw 25–28mm. The smallest Orthetrum species. This is another species with yellowish Pt and wholly blue mature males, but distinct by: (1) total length normally less than 40mm (O. coerulescens is at least 5mm longer on average); (2) the thorax has

? secondary genitalia

/

abdomen straight sided and quite narrow

mature ?

thick black dorsal line two stripes on thorax side with a black stripe only along the front edge

lobe enveloped by base of broad hook

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only about 2–2.5mm long; (6) the upperside of the eyes are brown in mature males, contrasting with the blue pruinosity. Females and young males are easily identified by their striking pattern, but the markings become totally obscured by pruinosity in older males. Mature males of other species are larger, with broader or waisted abdomens and (green-) bluish eyes. Hand characters Hamule has a broad hook, the base of which envelops a small lobe. Behaviour Prefers to perch on rocks or sandy ground. Unlike other Orthetrum species, often sits in the obelisk position.

270

Occurrence two pale stripes on the front and two on each side, and the lateral stripes are accentuated by a black line on their anterior side; (3) the abdomen is slender, straight-sided and gradually tapering; (4) the sandy-yellow abdomen has a relatively thick black central line, accentuated by paler areas along it; (5) the Pt is short, even relative to its small size,

Range and status Widespread Asian species (Middle East to India) that extends along the Mediterranean shore in southern Turkey and some Greek islands, where it is not uncommon. Habitat Standing or slow-flowing waters, often sparsely vegetated, such as in rocky stream beds. Flight season From early April to the beginning of October.

Orthetrum ransonnetii (Brauer, 1865) Identification General One of world’s few true desert dragonflies, this scarce species was recently discovered on the northern edge of the Sahara in Morocco and on Fuerteventura in the Canary Islands. May first be noted by the female’s rather broad tan abdomen with dark rings. Males are similar initially but become completely pruinose, and may be noted among the generally smaller and more common O. chrysostigma by lacking the

C

Desert Skimmer

abdomen waist and Hw yellow base, as well as by the shorter Pt and brownish hue of the eyes. Field characters Tot 45–59mm, Ab 28–34mm, Hw 27–32mm. Larger on average than O. chrysostigma. While males become entirely pruinose with age, the pale brown females and young males are distinctive by: (1) darker chestnut apical rings of the abdominal segments; (2) discontinuous or even absent thin black dorsal line on the abdomen; (3) absence of dark-bordered whitish stripes on the plain thorax. Wholly pruinose males differ from O. chrysostigma by: (4) almost parallel-sided abdomen without a constriction (‘waist’) between S3 and S4 when viewed from above; (5) entirely clear Hw bases and whitish membranules; (6) notably small (about 2.5mm) rufous Pt. Uniquely among Orthetrum species in the Sahara, (7) all cross-veins and most longitudinal veins (except parts of the costa) are black and ultimately become pruinose in males; at least the subcostal cross-veins (the second row between the node and wing base) are yellowish in other species. Hand characters Usually no (rarely one to two) cells are doubled between IR3 and Rspl. The hook of the male’s hamule has a broad triangular base and pointed, out-turned tip, while the lobe is just a

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Rspl usually subtends single row of cells

small rufous pterostigma

271

whitish membranule and entirely clear hindwing base immature /

mature /

mature ?

? secondary genitalia

darker abdominal rings in females and young males anterior lamina and hook of hamule pointed

basal swelling; the anterior lamina is also pointed. Variation The abdominal rings are especially clear and triangular in females but may narrow, fade and even vanish with age. Older females can develop pale pruinosity on the underside of the body. Behaviour Perches on rocks and bare ground, regularly sheltering from the sun on vertical surfaces or under overhanging rocks.

Occurrence Range and status Highly localised from Morocco and Mauritania to Iran and Afghanistan, but not uncommon where found, especially in the mountains of the central Sahara, for example in southern Algeria and Libya just outside the mapped region. An old record from eastern Turkey is unconfirmed. A young male photographed on

Fuerteventura in 2018 represents the first record for the Canary Islands and Europe. This was near a ravine with an intermittent stream and stagnant pools. As such barrancos are typical of the islands, more records may be expected. Habitat Contrary to most desert species that breed at ephemeral sites and wander widely, this is a resident of permanent pools fed by intermittent and perennial streams and springs. These are usually rocky, can be brackish and artificial, and often persist in the deep wadis of desert mountains, such as guelta. Flight season Recorded mainly between March and October, but most abundant in spring at lower elevations: may fly throughout the year in the central Sahara.

Orthetrum trinacria (Selys, 1841) Identification General Large, elongate and powerful species that may recall an aeshnid in flight. Its thin cylindrical abdomen with a bulbous base, crisply marked females and young males, and slate-coloured mature males should render it unmistakable. Field characters Tot 51–67mm, Ab 38–44mm, Hw 34–38mm. Our longest and most slender Orthetrum. Nearest to O. sabina by the swollen

Long Skimmer

S1–2 (more than three times as thick as remaining segments), olive-yellow ground colour, rather black abdomen and large, pale Pt. Unique for: (1) total length always more than 50mm (O. cancellatum about 10mm shorter, on average); (2) uniformly slender S3–10; (3) black abdomen with pale streaks at least up to S7, rather than rings up to S6, and the longest part of the pale spots on S4–6 lies against the dorsal keel, not along the lateral

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large pale pterostigma

? secondary genitalia

272 / pale longitudinal streaks

mature ?

/

long, thin abdomen

down-turned hook halfway between anterior and posterior ridges bulbous abdomen base

dark colour of mature male

long appendages

keels as in O. sabina; (4) males and old females darkening, with relatively thin bluish pruinosity, appearing darker than other pruinose Orthetrum males, and the abdomen may appear largely black. The abdomen shape may recall the much smaller O. chrysostigma, which is marked differently. O. sabina is smaller and never pruinose, has different markings and never has grey-blue eyes. Hand characters Appendages are longer than in other species, almost three times S10 in female, and upper appendages almost twice as long as lower in males. Hamule has a down-turned hook between two ridges, unlike any other species except O. sabina. Behaviour Aggressive; frequently takes other dragonflies as prey.

of the Iberian peninsula. Fairly common in Sicily and Sardinia; has recently colonised the Canaries (Fuerteventura) and Malta, but still very rare and local in Greece and Turkey. Habitat Standing waters, these often large, such as reedy lakes. Flight season From April to October.

C

Occurrence Range and status Common in tropical Africa and parts of south-west Asia. Widespread but not common in north-west Africa. First recorded in Spain and Portugal in 1980 and 1991, now regionally common in the south and south-west

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Orthetrum sabina (Drury, 1773)

/

distinctive black-andwhite marked abdomen, never pruinose

Slender Skimmer

273 ? secondary mature ?

markedness of characteristic pale stripes depends on darkness of thorax

diagnostic clubshaped abdomen (S7–10) conspicuous from side

/

General A swift, nervous species, lacking pruinosity, with strongly contrasting markings and a peculiar wasp-waisted, clubbed abdomen. Often taken for a gomphid rather than an Orthetrum at first sight. One of the commonest tropical Asian dragonflies extending into our area along the Mediterranean coast. Field characters Tot 43–50mm, Ab 31–36mm, Hw 28–33mm. Size is similar to O. cancellatum, but shape is totally different. Unmistakable, with these unique characters: (1) abdomen club-shaped, S1–2 swollen (more than three times thicker than the very slender S4–6) and S7–10 expanded (view from side); (2) abdomen predominantly black and never pruinose, wih a diagnostic pattern of three whitish rings on thin ‘waist’ (S4–6) and conspicuous (partly) white appendages; (3) thorax is pale greenish yellow with numerous bold black stripes, becoming dark with age and with a single bold whitish band on each side; (4) eyes never grey-bluish, as in mature males of other species, but greenish or brownish. Confusion is most likely with females and young males of the similarly shaped O. trinacria, but that species is larger and lacks a clubbed tail-end. Hand characters Unlike any other Orthetrum, anterior lamina bears a conspicuous tuft of dark orange bristly hairs. Variation Colour of thorax and extent of abdominal black varies considerably.

tuft of orange hairs on anterior lamina

bulbous base

white appendages

Identification

genitalia

Behaviour Active, and often very wary, perching on the ground or twigs for just a few seconds before speeding off again.

Occurrence Range and status Ubiquitous species in southern Asia, reaching Australia. Common on the Turkish Mediterranean coast and some Greek islands. Less common in Tunisia and Algeria. Recently discovered in Morocco; may be found on the Greek mainland in future. Habitat All kinds of standing or slow-flowing waters. Flight season From April to October.

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Leucorrhinia Brittinger, 1850 Whitefaces

White-faced Darters

Identification

274

Diagnosis Easily distinguished, even from afar, by the bright white face, contrasting with a largely black body. This feature is combined with unique wings that have dark spots at the Hw bases, only seven to eight (rarely nine) Fw antenodal cross-veins, and notably short, rectangular Pt. The abdomen is predominantly black, with a single series of pale dorsal spots that turn deep red in mature males in three species. In two others they disappear and the abdomen becomes grey pruinose at its base. Separation from other genera Other libellulids have coloured faces, although this may be poorly developed in young individuals, and in most of those genera pale colours predominate on the abdomen. Males developing pruinosity (e.g. Orthetrum, Libellula) tend to have this on more than half the abdomen, but the distinction may be difficult in old and worn pruinose Leucorrhinia males. The only largely black libellulid occurring widely with Leucorrhinia species is Sympetrum danae. The smallest species, L. dubia, in particular may be confused with it when viewed from a distance; S. danae has a yellow to black face, and at most a yellow Hw base. Finally, the white-faced North American vagrant Pachydiplax longipennis may recall a Leucorrhinia species, but it is only likely to appear on the Atlantic seaboard. Separation of the species Five species occur in our area. All increase in abundance towards the northeast, ranging deeply into Siberia. Another seven inhabit North America. Our species can easily be

G Leucorrhinia caudalis male. separated into two groups. Two species have white appendages and males that become dark, with pruinosity at the abdomen base. Three others have dark appendages and develop deep red markings but no pruinosity. Note that the species in these groups often occur together. The white appendages are easily seen from a distance, making separation possible without capture. To identify species, especially the red-spotted ones, examination in the hand is preferable; the male’s hamule and female’s vulvar scale rule out all confusion. Young individuals of all species are black with yellowish spots, and can be separated safely only by close examination, and by also referring to the pattern of spots. Behaviour All species have an erratic flight, especially the smaller red-marked species, which can often be seen ‘dancing’ over bogs and fens. The larger, pruinose species fly more slowly, often over open water or even among the trees surrounding a breeding site. Females of all species may oviposit alone, or are guarded by the male flying or perching nearby. In hot weather all species may perch with abdomen raised in obelisk position.  G Sahlén

Simple key to (pairs of similar) species. If the statement agrees, compare the given species. If it disagrees then go to the next line.

1 2 3

Appendages white, not black. Abdomen grey-blue pruinose at base; if not so, S7 at most with minute spot, never with red markings on thorax and abdomen.

albifrons caudalis

Spot on S7 yellower than the spots on S4–6, which are reddish brown in males and yellow-brown in females.

pectoralis

Spot on S7 at most more reddish or orange than those on S4–6.

dubia, rubicunda

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Leucorrhinia dubia (Vander Linden, 1825) Small Whiteface

White-faced Darter approx. basal half of costa dark (view from front)

pterostigma usually blackish patches semicircular with rounded outher edge

/

275

immature ?

mature ? relatively small spots up to S7 distinctive white face of the genus

?

? secondary

/ vulvar scale

genitalia

straighter hook of hamule, at right angle to abdomen

Identification General The dominant whiteface in many areas, often the only one at the edge of its range, but also widely overlapping with the closely similar L. rubicunda. The two are difficult to distinguish with certainty without capture, although relative differences in size, shape, and abdominal and wing spots aid recognition in the field. Field characters Tot 31–36mm, Ab 21–27mm, Hw 23–28mm. The smallest Leucorrhinia; almost as small as Sympetrum danae. Easily confused with L. rubicunda, unless caught or closely examined (see that species for comparison). Both are black with (yellowish-) brown spots on the thorax and S2–7. The spots are smaller and darker in males, especially mature ones, turning orange and eventually deep red between the wings and on the abdomen. In L. dubia, all spots are usually smaller than in other red-spotted species; those on S4–6 are usually darker than that on S7 and may even disappear with age. Patches at Hw bases are almost semicircular, with a rounded outer edge that is blunt where it meets the wing border; in L. rubicunda these are more triangular with rather straight and sharply angled edge. Sympetrum danae has similar range, habitat and stature. It is also sleek and black, but the face and markings are at most yellowish, not white and red. Females and

two triangular processes

young males may be mistaken for L. albifrons and L. caudalis, but these species both lack the large spot on S7 and have whitish appendages. Hand characters In red-spotted species in particular, the blackish basal spots in the Hw are also present in the Fw. This is most extensive in L. dubia, broadly covering the Fw’s hind corner. Male’s hamule has a straighter hook than other Leucorrhinia species, pointing perpendicularly away from the abdomen. Female’s vulvar scale has two triangular processes. Variation Andromorph females occur regularly.

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Behaviour Like L. rubicunda, often displays a rather erratic flight across the water and the surrounding area. Frequently perches on the ground, among dense vegetation.

Occurrence Range and status Often abundant, but like other Leucorrhinia species its occurrence is relictual in the south-west of its range. Here it is relatively frequent, being the only Leucorrhinia species in

276

Great Britain and the higher Alps and Pyrenees. Habitat Bogs, ponds, tarns and lakes, usually acidic and often in forests. The larvae are sensitive to fish predation, and so the species does better where fish are not present, such as small pools in raised bogs. Flight season From mid-April to early September. In temperate lowlands, most abundant from late May to mid-July.

Leucorrhinia rubicunda (Linnaeus, 1758) Ruby Whiteface Northern White-faced Darter



entire costa yellow (view from front)

pterostigma red-brown patches triangular with rather straight outer edge

/

relatively large spots up to S7

mature ?

? secondary / vulvar scale

genitalia

hamule with strongly curved hook

large round lobe

without large processes

Identification General Often occurs with the smaller, sleeker and smaller-spotted L. dubia. May be easily confused without close examination, but with binoculars the shape of Hw patch and entirely yellow leading edge of Fw can be made out. Field characters Tot 31–38mm, Ab 23–28mm, Hw 27–31mm. Medium-sized Leucorrhinia; near the average Sympetrum. See L. dubia for similarities. Differentiating field marks are: (1) somewhat larger and more robust; (2) abdominal spots (red in males, yellow in immatures and females) usually larger, approaching those of L. pectoralis, especially in females, and the large dorsal spots on S3–4 extend downwards (normally strictly dorsal and separated from some lateral spots in L. dubia); (3) patches at Hw bases triangular with rather straight outer edge meeting the wing border at a sharp angle (more

rounded and blunt in L. dubia); (4) costa (leading vein) of Fw yellow from base to Pt (view from front) (in L. dubia it is dark brown to black basally (almost) to node); (5) may be picked out by the redbrown rather than blackish Pt, but this character is not consistent as some L. dubia have reddish Pt. Features 1–4 recall L. pectoralis, but that generally has the S7 spot yellower than those on S2–6. Hand characters Basal dark spot in Fw usually smaller than in L. dubia, barely covering the wing’s hind corner. Male’s hamule appears like a towing hook: a relatively small, curved hook with a large round lobe. Female’s vulvar scale lacks double triangles of L. dubia and L. pectoralis. Variation Very old individuals can be very dull and dark, with only the S7 spot distinct. Andromorph females occur regularly. Behaviour Similar to L. dubia.

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Occurrence Range and status May be abundant in good habitat, but is the least widespread Leucorrhinia in the warm temperate parts of Europe. The populations on the south-western edge of its range, such as along the northern Alps, are isolated and vulnerable. Habitat Like L. dubia prefers acidic, oligotrophic lakes, tarns and bogs, often in forest. Also occasionally breeds in more nutrient-rich, richly vegetated habitats (similar to those of L. pectoralis). Better adapted to coexist with higher fish densities than L. dubia. Flight season Probably the earliest whiteface; from mid-April to early August. In temperate lowlands, most abundant from mid-May to mid-June.

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Leucorrhinia pectoralis  (Charpentier, 1825)

Yellow-spotted Whiteface Large White-faced Darter

dark pterostigma

mature ?

/

large yellow markings

spot on S7 does not always stand out as pale

? secondary genitalia

/ vulvar scale pale spot on S7 triangular processes

Identification General The largest whiteface and the prevailing one in many lowland and more eutrophic areas, although almost always very local. The conspicuous yellow spot on S7 allows males and most females to be identified through binoculars. Field characters Tot 32–39mm, Ab 23–27mm, Hw 30–33mm. Largest Leucorrhinia species, somewhat larger and plumper than Sympetrum species. Nearest is L. rubicunda in all features, but L. pectoralis is even larger, with a thicker abdomen and larger spots. The markings on S1–6 are fairly dull brown, yellower when fresh and with a strong orange or red tinge in males. In contrast,

long plume on tip of anterior lamina

large hook of hamule

the spot on S7 is bright lemon yellow, becoming darker and duller with age. The Pt is darker than that of L. rubicunda, a feature that may aid field identification. May easily be confused with L. dubia and L. rubicunda, especially very old and very young individuals that lack bright colours. However, these species never have the spot on S7 lemon yellow; if its colour differs from S1–6 (brighter, less brown), the tone is redder rather than yellower. If all spots are yellowish, identification must be made by hand characters. Hand characters Male’s hamule is like that of L. rubicunda but the hook is much larger. Although all Leucorrhinia males have very hairy secondary

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genitalia, those of L. pectoralis have an especially conspicuous plume of long hairs on the tip of the anterior lamina. Female’s vulvar scale with two triangular processes, like those of L. dubia. These processes are absent in L. rubicunda, this being the only reliable character to distinguish the similar females of these species. Variation Old males are duller; the contrast of the S7 spot may be less apparent. Behaviour Males often patrol among tall emergent plants, seldom perching on low, flat surfaces.

278

L. rubicunda, preferring marshy borders of bogs, forest lakes, fenlands, marshy ditches, oxbows, and even sluggish rivers or canals. Vegetation is typically lush and varied, with both emergent and submerged species. The water is frequently mesotrophic and often coloured brown due to peat. Flight season From early May to early August, with most records in late May and June.

HD II+IV

Occurrence Range and status Fairly common and widespread in the Baltic region and southern Sweden, but generally uncommon elsewhere; populations are usually local and small but may disperse in good numbers. First recorded in Great Britain in 2012. The most thermophilous Leucorrhinia, with a relatively southern range. The only Leucorrhinia species in Turkey and much of the Balkans, where it is very local. Habitat Generally occurs in habitats that are not as acidic and nutrient poor as those of L. dubia and

G Leucorrhinia rubicunda andromorph female. H Leucorrhinia caudalis female, which has become somewhat pruinose with age.

G Leucorrhinia pectoralis young female. H Leucorrhinia dubia andromorph female.

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Leucorrhinia albifrons (Burmeister, 1839) Dark Whiteface Eastern White-faced Darter



? secondary genitalia

pterostigma dark

hook almost parallel to abdomen

279 larger spots confined to S2–6

/

abdomen base pruinose, especially S3–4

appendages white

mature ?

/ vulvar scale

virtually indiscernible processes

Identification General The darkest and dullest of the Leucorrhinia species, males appearing rather grey. In the hand, easily separated from congeners by the white spots on the labium. Field characters Tot 33–39mm, Ab 22–27mm, Hw 23–28mm. A medium-sized Leucorrhinia. Differs from all Leucorrhinia species except L. caudalis as follows: (1) largely dark without reddish spots, and yellowish markings restricted and visible only in females and young males; (2) abdominal spots small and confined to S2–6, at most a trace on S7; (3) appendages white in both sexes; (4) mature males are all dark with bluish-grey pruinosity, especially at the abdomen base and wing joints. The thickest pruinosity is confined to S3–4, but thin pruinosity may also cover the thorax (e.g. between wings) and abdomen up to S5. The pale ring at the abdomen base thus appears shorter and more diffuse than in L. caudalis. Appears smaller and sleeker than that species (abdomen scarcely clubbed). The Pt are blackish brown in both sexes. Hand characters Labium laterally white, but all black in other Leucorrhinia species. Hook of male’s hamule is small and straight, pressed inwards, almost parallel to abdomen. Female’s vulvar scale with two very short, almost invisible processes. Females are most easily identified by the labium. Variation Old individuals can appear very grey. Only spots on S2–3 may be apparent in dark females. Dark basal Hw marking in females is enclosed by diffuse yellow, which may cover wings quite extensively. Teneral females can appear distinctly club-tailed and must be separated from L. caudalis

white patches on labium

HD IV

by careful comparison of the yellow markings on S3: in L. albifrons there is a basal ring that reaches to the segment sides and a central patch that seems separated into two spots lying beside each other, while in L. caudalis there is a larger central patch that consists of two parts following each other. Behaviour Often perches in bankside bushes or trees. Unlike L. caudalis, seldom perches on lilypads.

Occurrence Range and status Locally abundant in Fennoscandia and the Baltic region, but generally scarce in central Europe, becoming very local towards the south and west, where it is usually under threat. Habitat Wide selection of lakes and bogs with abundant floating and emergent vegetation. Mainly confined to mesotrophic lakes in periphery of its range. Elsewhere also in rather acidic, oligotrophic or eutrophic waters. Flight season From late May (or end of April in the south) to mid-August; most abundant in June.

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Leucorrhinia caudalis (Charpentier, 1840) Lilypad Whiteface

Dainty White-faced Darter ? secondary genitalia

pterostigma white

280

/ vulvar scale

S3–5 pruinose pattern of dorsal spots different to L. albifrons

/

S7–8 expanded

narrow gap between large hook and lobe

mature ?

2nd cross-vein between hindwing triangle and base

finger-like lobes

appendages white

Identification General Large, broad club-tailed species. Males often perch on lily-pads. They are dark, marked with white highlights on the face, waist, and at the tips of the wings and abdomen. This makes identification through binoculars easy. Field characters Tot 33–37mm, Ab 23–25mm, Hw 29–32mm. Almost as large as L. pectoralis, appearing short, plump and wide-winged. Differs from other Leucorrhinia species by features shared with L. albifrons (see that species) and differs from that species in that: (1) abdomen is distinctly club-shaped (but see Variation in L. albifrons) and S7–8 are very broad; (2) abdominal spots (visible in females and young males) are more extensive and differently configured – the dorsal spots on S3–4 are often fused with each other, but are separate from small lateral spots (in L. albifrons, these segments typically have a pale ring at their base, plus a separate dorsal spot); (3) males have whitish uppersides of Pt (dark brown in females, young males and other species); (4) male’s pruinosity is thicker, whiter and sharper, distinct on S3–5. Hand characters Unlike its congeners, Hw usually has two (not one) cross-veins between the wing base and triangle. Unlike L. albifrons, the labium is all dark. Hook and lobe of male’s hamule is rather large and pressed together. Female’s vulvar scale ends in two finger-like, almost parallel processes. These extend over about one-third of S9. Variation Female wing bases are often diffusely yellow. Some females also have brown patches under the Pt of each wing. Behaviour Both sexes fly actively over open water,

HD IV

frequently resting on floating vegetation (Nuphar, Nymphaea, Potamogeton), where mating may also take place. Male often raises clubbed tail.

Occurrence Range and status Occurrence is scattered and populations are normally rather small, although may be abundant locally. Fairly common in southern Fennoscandia and the Baltic States. Was local and under threat in south and west of its range, but showing recent signs of recovery with westward expansion in the Netherlands, Belgium, France and Germany. Extends to central Siberia. Habitat Pools and lakes, often in forests (but not shaded), with moderate nutrient levels and rich (especially aquatic) vegetation. The larvae are very spiny, unlike its congeners, surviving high fish densities. In the periphery of its range, often found in oxbows and fish ponds. Flight season Mid-May to early August; most abundant in June.

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Sympetrum Newman, 1833

Darters Meadowhawks (NA)

281

G Sympetrum sinaiticum male.

Identification Diagnosis Rather small libellulids. Mature males – with the exception of a black species – have an abdomen that is a shade of red. Black markings are variable, but usually at least traces on the thorax sutures, legs and abdomen. Hw base clear or marked with yellow or amber. Fw with distinctive venation: 6½–8½ antenodal cross-veins; discoidal field narrowing towards wing border and of three rows at base; triangle and subtriangle with cross-veins, thus normally with two and three cells, respectively; Rspl subtends a single row of cells. Separation from other genera The only smaller libellulids without a dark Hw patch and often a red abdomen in much of our range. Crocothemis has Fw with 8½–10½ antenodals and widening discoidal field; moreover, the legs lack any black and the abdomen is broader (only some Sympetrum occurring regionally in Turkey, southern France and Spain have almost no black on their legs). Red Trithemis are similar in stature but have 9½–12½ antenodals, two rows before Rspl, and mature males have a much brighter appearance.

Brachythemis has similar venation, but has a plump, cylindrical abdomen and the Fw triangle and subtriangle lack cross-veins. Separation of the species A large and often common genus. Many species are readily identified, especially those in the first half of the key. The species in the latter half can cause identification problems, especially when observing with binoculars only (see table overleaf). The greatest challenge is to pick out the increasingly widespread but deceptively variable S. meridionale among the common S. striolatum and S. vulgatum. Behaviour All darters behave similarly. They spend most of their time on a prominent perch, from which short sorties are undertaken to chase prey or rivals. Males often attain high densities along the water’s edge, defending a small ‘private space’ around themselves rather than a territory. Oviposition always commences in tandem, but species differ somewhat in their mode of flight and choice of oviposition site.  K-D B Dijkstra

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Simple key to (groups of similar) species. If the statement agrees, compare the given species. If it disagrees, then go to the next line.

282

1

Wings with brown bands between Pt and node.

pedemontanum

2

Tibiae and usually femora completely black, without yellow streaks.

danae, depressiusculum sanguineum

3

Hw with prominent yellow patch at base. Vulvar scale deeply incised in middle (view from below).

flaveolum fonscolombii

4

Back of head all-pale, without dark bars. In Turkey only.

haritonovi vulgatum (Turkey)

5

Black on sutures of thorax almost absent, not as distinct lines. Legs with only some black on inner surface, rather than black with pale stripes.

meridionale (typical) sinaiticum vulgatum (Spain)

6

Black at base of frons extending downwards along eye-margins, like drooping moustache. Vulvar scale projecting at right angle to abdomen.

vulgatum (typical)

7

Black at base of frons not extending downwards. Vulvar scale pressed more against abdomen.

striolatum (including nigrescens & nigrifemur)

Comparison of similar (sub)species near S. meridionale, S. striolatum (s.) and S. vulgatum (v.). Pale: black on sutures of thorax almost absent; legs with only some black on inner surface. Very pale: pale, with even back of head all-pale, without dark bars; legs almost unmarked. Dark: distinct black thorax lines present; black legs with pale stripes. Very dark: dark, with black thorax stripes partly confluent; femora largely black. Hhk: hamule hook long (+) or short (–). Vsc: vulvar scale (strongly) protruding ((+)+) or appressed (–). Diagnostic features

Range Turkey

Tiny (total length