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The Royal Entomological Society Book of British I suppose you are an entomologist? – I said with a note of interrogation.

Not quite so ambitious as that, sir. I should like to put my eyes on the individual entitled to that name! A society may call itself an Entomological Society, but the man who arrogates such a broad title as that to himself, in the present state of science, is a pretender, sir, a dilettante, an imposter! No man can be truly called an entomologist, sir; the subject is too vast for any single human intelligence to grasp.

Oliver Wendell Holmes, 1882, The Poet at the Breakfast Table The Royal Entomological Society Book of British Insects

Peter C. Barnard

A John Wiley & Sons, Ltd., Publication This edition fi rst published 2011 © 2011 by Royal Entomological Society

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Library of Congress Cataloging-in-Publication Data Barnard, Peter C. (Peter Charles), 1949– The Royal Entomological Society book of British insects / Peter C. Barnard. p. cm. Includes bibliographical references and index. ISBN 978-1-4443-3256-8 (hardcover : alk. paper) 1. Insects—British Isles—Identifi cation. I. Royal Entomological Society of London. II. Title. III. Title: Book of British insects. QL482.G8B37 2011 595.70941—dc23 2011019098

A catalogue record for this book is available from the British Library.

This book is published in the following electronic formats: ePDF 9781444344950; Wiley Online Library 9781444344981; ePub 9781444344967; Mobi 9781444344974

Set in 9.5/12 pt Palatino by Toppan Best-set Premedia Limited

1 2011 Contents

Preface vii Foreword by Quentin D. Wheeler ix Acknowledgements xi

1 General introduction 1

PART 1 ENTOGNATHA 17 2 Order Collembola: the springtails 19 3 Order Diplura: the two-tailed or two-pronged bristletails 25 4 Order Protura: the proturans 27

PART 2 INSECTA – ‘’ 29 5 Order or Microcoryphia: the bristletails 31 6 Order Zygentoma: the silverfi sh and fi rebrats 33

PART 3 35 7 Order Ephemeroptera: the mayfl ies or upwing fl ies 37 8 Order Odonata: the dragonfl ies and damselfl ies 42

PART 4 49 9 Order Dermaptera: the 51 10 Order : the cockroaches, termites and mantids 54 11 Order Orthoptera: the grasshoppers, crickets and bush-crickets 57 12 Order Phasmida: the stick-insects 64 13 Order Plecoptera: the stonefl ies 67

PART 5 71 14 Order : the true bugs 73 15 Order Phthiraptera: the sucking and biting lice 106 16 Order : the booklice and barklice 112 17 Order Thysanoptera: the thrips 118

PART 6 123 18 Order Coleoptera: the beetles 125 19 Order Diptera: the true fl ies 177

v CONTENTS

20 Order Hymenoptera: the ants, bees and wasps 226 21 Order Lepidoptera: the butterfl ies and moths 268 22 Order Mecoptera: the scorpionfl ies 308 23 Order Megaloptera: the alderfl ies 311 24 Order : the lacewings 313 25 Order Raphidioptera: the snakefl ies 317 26 Order Siphonaptera: the fl eas 319 27 Order Strepsiptera: the stylops 323 28 Order Trichoptera: the caddisfl ies or sedge fl ies 326

Index to taxa 337

vi Preface

In 2007 the Royal Entomological Society (RES) standing series Handbooks for the Identifi cation of took the momentous decision to move from British Insects . central London to a more rural environment near St Inevitably such a book highlights the contrast Albans in Hertfordshire, the fi rst time that the between popular groups such as Lepidoptera and Society had left London since its foundation in 1833. Coleoptera, relatively well - served by identifi cation The move was seen as controversial by some, literature, and the lesser studied groups such as although the new headquarters are close to the many of the Hymenoptera. However, the aim is not London Orbital M25 motorway and only just to bemoan the lack of information on these less outside the Greater London conurbation. Apart popular orders of insects, but to encourage poten- from the fi nancial relief of no longer having to tial authors to fi ll some gaps. This call to arms is as maintain an impressive but aged building in South much aimed at the keen amateur entomologist as it Kensington, the sale of 41 Queen ’ s Gate realized a is at the professional, since few of the latter are substantial capital sum that enabled the RES to encouraged or even allowed to prepare taxonomic press ahead with several new projects, such as the works as part of their duties. In these sadly unen- appointment of the fi rst full - time entomologist - in - lightened times the perpetual need for basic taxo- residence. nomic information is overshadowed by the political This book is one of the tangible products funded insistence that good science can be defi ned by its by the move out of London and it represents part ability to attract external funding, which may refl ect of the re - positioning of the RES as the central hub the inability of some infl uential managers to make of information on British insects, as well as main- informed and independent decisions about the real taining its role as the premier society for profes- value of science. With luck, the overwhelming need sional entomologists. The book is designed as a key for better taxonomic information, coupled with the reference work for professional entomologists as development of systems to coordinate it, will cause well as being a readable and attractively illustrated this short- sighted approach to be overturned. It will account for the interested student of insects. It helps be gratifying if this book helps to restore the rightful to bridge the gap between the popular but incom- place of systematic entomology at the centre of plete guides to the more conspicuous groups of studies in this country. British insects and the highly specialized works that currently can cover only a fraction of the Peter C. Barnard entire fauna. Of these detailed works the most pres- Royal Entomological Society tigious are, of course, the Society’ s own long- May 2011

vii

Foreword

One could proclaim the study of insects to be the insect studies. And fi nally, they would have author- study of biodiversity and be at least half right. The itative and comprehensive books that provide an more than 1 million insect species described to date entré e into insect biodiversity for the public and account for fully half of all kinds of plants and scientists alike. No place on earth meets these , but no more than 25% of the estimated requirements as fully as the British Isles. total number of living insect species (Foottit & The concentration of expertise and great institu- Adler, 2009 ; Chapman, 2009 ; Grimaldi & Engel, tions, from natural history museums to the Royal 2005 ). No full understanding of either evolutionary Entomological Society (RES), combined with a history or the function of terrestrial or freshwater diverse yet fi nite assemblage of species make the ecosystems is possible without detailed knowledge British insect fauna accessible and approachable on of insects, yet our ignorance of insect diversity, and a level unmatched elsewhere. The growth of our inadequate description and classifi cation of knowledge of insects is so fast - paced that general their kinds, prohibit rapid progress. So how are we summaries of the fauna are soon outdated and to proceed? become impediments themselves; the number of In the long term, we need to accelerate funda- species recorded for Britain has increased threefold mental taxonomic and natural history investiga- since the time of Stephens (1846) . While popular tions and make relentless progress toward a accounts of a few charismatic taxa such as dragon- complete exploration of insect diversity. This will fl ies and butterfl ies exist as well as superb treat- involve investments in museums, a cyberinfrastruc- ments of particular groups in journals and the ture specifi cally engineered to meet the unique outstanding RES Handbook series, there is a need needs of taxonomists, and the inspiration and edu- for an up - to - date introduction to the insect fauna cation of the next generation of insect specialists. of the UK as a whole. This volume fi lls that gap We need to ensure also that insects are integral to exceedingly well. research into all aspects of basic and applied biology, The need for such comprehensive overviews is from agriculture to behaviour, developmental not new. In 1828, Kirby and Spence were convinced biology and ecosystem science. Our immediate that it was the absence of such an adequately com- need is for natural laboratories where insects are plete and affordable introductory volume on the more approachable than they are on the global insect fauna of Britain that was the primary impedi- stage. What might such natural laboratories look ment to the advancement of entomology. Their like? First, they would have insect faunas with infl uential book was an answer to this obstruction enough species and diversity to be broadly repre- and was the result of their resolve ‘ to do what was sentative, but not so many species as to be unman- in their power to remove it, and to introduce their ageably large. Second, their fauna would be countrymen to a mine of pleasure, new, boundless, relatively well known, their species reasonably well and inexhaustible’ (Kirby & Spence, 1828 : ix). described and mirrored in comprehensive and well- Thanks to the vision of the RES and Dr Peter curated museum collections. Third, they would Barnard, this need is once again met for a new gen- have a vibrant community of amateur and profes- eration of entomologists. The knowledge and expe- sional entomologists. Fourth, they would have rience required to adequately summarize what we visionary organizations to encourage and enable know of 24,000 species representing 558 families are

ix FOREWORD great, and few would be up to the challenge. I can play its unique role as a natural laboratory of bio- imagine no better choice than Dr Barnard whose diversity exploration. research and curatorial work at the Natural History Museum and dedicated service to the RES span Quentin D. Wheeler, PhD, FRES, FLS more than 30 years and have prepared him for this Vice President and Dean singularly important challenge. Arizona State University Insects remain a largely untapped mine of pleas- ures for those who take the time to identify, observe REFERENCES and enjoy them. The discoveries yet to be made through a more thorough study of entomology Chapman , A. 2009 . Numbers of living species in Australia remain similarly new, boundless and inexhaustible. and the World ( 2nd edn .). Australian Biological Resources Circumstances threatening biodiversity and the Study , Canberra . environment make the importance and timeliness Foottit , R.G. & Adler , P.H. (eds.). 2009 . Insect biodiver- sity: science and society . Blackwell Publishing , Oxford . of this volume and increased attention to research Grimaldi , D.A. & Engel , M. 2004 . Evolution of the insects . on insects obvious. Of equal importance is increas- Cambridge University Press , Cambridge . ing understanding and appreciation of insects Kirby , W. & Spence , W. 1828 . An introduction to entomology among the general public. There is no surer or more or elements of the natural history of insects . Vol. I . Longman, effective means to that end than making the insects Rees, Orme, Brown, and Green, London . around us more easily accessible. Thanks to Dr Stephens , J.F. 1846 . Illustrations of British entomology . Barnard and the RES, the British fauna is poised to Supplement. Henry G. Bohn , London .

x Acknowledgements

My initial thanks must go to the Council of fi shing competitions; he has a keen interest in fresh- the Royal Entomological Society and the Registrar, water insects and has written many articles on Bill Blakemore, for entrusting me with the task fl y - fi shing, entomology and river management, all of writing this book on their behalf. Val McAtear, illustrated by his own photos. the RES librarian, has helped me on numerous I am very grateful to Peter Chandler for sending occasions with bibliographic queries, and Mike me a copy of his Dipterist ’ s Handbook before it was Claridge has provided a generous supply of useful published so that I could make use of the invaluable advice. information therein. Much of the value of this book lies in the range Quentin Wheeler ’ s tireless championing of insect of insect photos, and I take great pleasure in thank- has always been a source of inspiration, ing the four photographers for their unique and I thank him for his stirring Foreword; his clear contributions: understanding of the needs and potential of British Roger Key is a well- known entomologist, for- entomology should ideally help to shape the merly of English Nature/Natural England, who research priorities in this country. now lives in North Yorkshire. As a freelance lec- My wife Jane has been very patient with my ento- turer, ecologist and fi eld entomologist he specializes mological obsession, though if she hadn’ t kept the in inspiring young people to appreciate wildlife; he kitchen so clean it might have been easier to fi nd is also an experienced broadcaster and his photos some domestic psocids to photograph. I should also of all groups of insects have been published widely thank my greyhound Reene for providing a supply in many books and magazines. of fl eas (now safely exterminated). Robin Williams, from Wedmore in Somerset, is Several staff at Wiley - Blackwell have made the an authority on the Hymenoptera, but his long planning and production of this book a much experience as a naturalist and photographer is smoother process than it might otherwise have evident in the wide range of insect groups and been, and I particularly thank Ward Cooper, Kelvin behaviour portrayed in his pictures. He is one of the Matthews, Kathy Palmer and Delia Sandford for few people who can recognize the numerous species their advice and support. In addition, freelance associated with oak galls, as well as other families copy- editor Lewis Derrick and freelance project of the ‘ parasitic ’ wasps. manager Nik Prowse have made invaluable Colin Rew lives near Huddersfi eld in West improvements with their editing of the manuscript, Yorkshire; he qualifi ed in Electrical Engineering and saving me from some potentially embarrassing then worked in industrial insurance, where his pho- slips. I take responsibility for any remaining errors tographic skills were in frequent use. Following that the assiduous reader will undoubtedly fi nd. retirement in 2008 he took a course in macro- Many modern authors owe a debt of gratitude to photography and became hooked on photograph- the automatic spell- checkers in their word - process- ing the insects in his garden; the stunning results of ing software, but not in a book with so many scien- his new - found interest are here for all to see, includ- tifi c names. I quickly ran out of patience when my ing on the front cover. computer kept trying to change Adelidae to Stuart Crofts is a lifelong fi sherman from South Adelaide, to Gertrude, to por- Yorkshire who has represented England in fl y - ridge and Sesiidae to seaside.

xi

1 General introduction

THE SYSTEMATIC APPROACH The terms classifi cation, taxonomy and systemat- ics are often used as if synonymous, but there are Systematic studies form the basis of all scientifi c some important differences. A classifi cation is basi- work, and this is no coincidence or artifi cial contriv- cally a way of groupings organisms in a logical way: ance. Classifying natural objects is an essential although it claims to be based on similarities it often instinct in human beings that enables us to make ends up emphasizing differences in order to defi ne sense of the world, and the origins of this ability are the boundaries of groups, and no assumptions not hard to trace. For an off - beat and thought - about processes or underlying causes of any per- provoking look at this subject see Yoon (2009) ; as ceived patterns are made. Such systems are often Quentin Wheeler says in a review of that book, tax- phenetic in nature, based on morphological differ- onomy is the ‘ oldest profession practiced by people ences to divide a large taxon into manageable with their clothes on’ , a reference to the belief that pieces. Systematics uses the principle that similari- Adam was given the task of naming all the crea- ties are based on shared, derived characters; it tures (Genesis 2: 19 – 20). therefore assumes an evolutionary, phylogenetic Superfi cial characters like colour, shape and even cause, and importantly it makes predictions about some behaviours are often quickly seen to be of the characteristics of groups, whether morphologi- little use in predicting the unknown, which is the cal, behavioural, ecological or physiological. value of systematics, even at a domestic level. From Taxonomy has traditionally been regarded as all careful observations about a few animals or plants about naming things, though it is linked to a clas- we can make generalizations about other species sifi cation or a systematic study, because any system that we have not yet encountered, and this ability of naming has to be based on principles of relation- to predict the unknown is what makes taxonomy a ships, whether real or artifi cial. This narrow defi ni- science rather than simply a technical procedure. tion has led to the perception that taxonomy is a Recognizing that plants related to nightshades are somewhat mechanical process, which simply pro- likely to be poisonous is clearly valuable, whereas duces names and groupings on behalf of the real wrongly classifying bats as a kind of bird might scientists who are carrying out systematic analysis. entail a long and fruitless wait for them to lay eggs! It is more useful to see taxonomy as the overarching Classifying organisms can, of course, be done for activity of studying phylogenetic relationships, pos- different purposes: there is a well- known cartoon tulating testable hypotheses about distributions of by Charles Keene in Punch magazine (1869, vol. 56, characters, and hence producing useful classifi ca- p. 96) in which a railway porter is informing a lady tions with meaningful systems of nomenclature that of the charge for carrying a tortoise on a train; he refl ect the underlying relationships. tells her that, ‘ Cats is “ dogs ” and rabbits is “ dogs ” Thus, the need to recognize true relationships and so ’ s parrots, but this ’ ere “ Tortis ” is an between organisms is essential to our perception of insect . . . ’ . the world and to our ability to exist in some kind of

The Royal Entomological Society Book of British Insects, First Edition. Peter C. Barnard. © 2011 Royal Entomological Society. Published 2011 by Blackwell Publishing Ltd.

1 GENERAL INTRODUCTION harmony with our environment. The importance of (608); Hymenoptera: Pteromalidae (600), Eulophidae taxonomic research is frequently cited, which makes (500); Coleoptera: Curculionidae (475); Diptera: it all the more surprising that support for taxonomy (471). Most of these fi gures are, of is increasingly hard to fi nd. Even national museums, course, approximate and it is unlikely that family the custodians of the most important natural history limits are interpreted in the same way in the differ- collections in the world, seem to be constantly ent insect orders. downgrading the emphasis on taxonomy in their The decision to cover the British insects at family organizational research priorities (Wheeler, 2008 ), level is not just for the convenience of dealing with though this is likely to be rooted in the fundamental 558 families rather than over 6000 genera! In many misunderstanding of the value and true aims of groups, families have a reality outside scientifi c tax- taxonomy, as stated above. This issue is discussed onomy; grasshoppers, pondskaters, ladybirds, further by Secord (1996) . hawk - moths, mosquitoes and ants are just a few examples of well -known groups of insects that cor- THE BRITISH INSECTS respond to family level taxa. There are, of course, many little- known families that have no common The terms ‘ British ’ and ‘ Britain ’ are deliberately names. used rather loosely to include Ireland as part of the Inevitably the coverage of each group varies British Isles, because many published ‘ British ’ lists because some are much better known than others, also include Irish species, even if they contain but every family is included, not just the common species found in Ireland but not in the UK. However, or conspicuous ones. This at least highlights those there are many special Irish lists (see, for example, groups – often quite large and diverse families – http://www.habitas.org.uk/invertebrateireland/ that still need far more work done on them. index.html ). With well over a million species of All the genera occurring in the British Isles are insects known in the world, and many more to be listed under each family, alphabetically within sub- discovered, our fauna of around 24,000 species families where these are commonly used in the looks rather insignifi cant. However, insect diversity larger groups. Tribes are not mentioned because is generally much greater in the tropics than in tem- they are used inconsistently in different insect perate regions (Foottit & Adler, 2009 ) and the British orders, and are often the subject of considerable insect fauna is in the range to be expected at our disagreement between authors. All generic names latitude. are listed in the index, along with every higher Table 1.1 shows the approximate numbers of taxon of insects mentioned in the text, as well as species, families and orders of insects known in the widely accepted vernacular names of groups at world, Europe and the British isles. World fi gures family level and above. are based on Foottit and Adler (2009) and Resh and The numbers of species listed here are in many Card é (2009) , European fi gures are from the Fauna cases an approximation; this is because classifi ca- Europaea website, and British fi gures from the tions change, species are added or synonymized, current book. Of course, these numbers can only and any checklist is out of date almost as soon as it refl ect the species described to date, and are not is published. Where the number is preceded by ‘ c . ’ necessarily indicative of the numbers of species that this indicates even less certainty about the total, actually occur. On a world scale, the fi ve largest particularly noticeable in groups like the parasitic groups of insects are: (i) Coleoptera; (ii) Lepidoptera; Hymenoptera. (iii) Diptera; (iv) Hymenoptera; and (v) Hemiptera. Totals normally include UK and Ireland, but not In the British Isles, the order of largest groups is the Channel Islands, though species found only in rather different, being: (i) Hymenoptera and Diptera the Channel Islands are occasionally mentioned (roughly equal); (iii) Coleoptera; (iv) Lepidoptera; where they are of particular interest or importance. and (v) Hemiptera. The differences may well refl ect The corresponding numbers of taxa in Europe and the fact that the diversity of groups like Coleoptera the rest of the world are given, so that the signifi - and Lepidoptera is higher in the tropics than in cance of the fauna of the British Isles can be placed temperate regions. For those who like impressive in context. For typical groups such as Lepidoptera statistics, the ten largest families of insects in the our fauna can seem insignifi cant in terms of world British Isles are Hymenoptera: Ichneumonidae species numbers, but for more cosmopolitan groups (2100 spp.), Braconidae (1045); Coleoptera: like the apparent proportion of British Staphylinidae (1000); Diptera: (652); species is larger in comparison with other parts of Hemiptera: (630); Diptera: the world. The reasons for these apparent differ-

2 CHAPTER 1

Table 1.1 Numbers of described insect species and families in the world, Europe and British Isles (see main text for sources).

Group World World European European British British species families species families species families

Collembola 8,000 29 2,000 23 250 19 Diplura 1,000 10 300 5 11 1 Protura 750 10 175 4 15 3 Archaeognatha 500 2 200 2 7 1 Zygentoma 400 5 60 2 2 1 Ephemeroptera 3,100 40 330 18 51 10 Odonata 5,600 33 130 11 49 9 Dermaptera 2,000 8 80 5 7 4 Dictyoptera 8,500 20 200 10 11 5 Embioptera 450 8 12 2 0 0 Grylloblattaria 30 1 0 0 0 0 Mantophasmatodea 20 1 0 0 0 0 Orthoptera 24,000 29 1,000 15 33 10 Phasmida 3,000 2 15 3 4 2 Plecoptera 3,000 16 400 7 34 7 Zoraptera 35 1 0 0 0 0 Hemiptera 100,000 104 8,000 94 1,830 63 Phthiraptera 5,000 24 800 17 540 17 Psocoptera 5,000 17 200 25 100 19 Thysanoptera 6,000 9 570 6 179 3 Coleoptera 350,000 175 12,500 144 4,000 112 Diptera 152,000 160 19,000 132 7,000 103 Hymenoptera 150,000 90 16,000 66 7,000 57 Lepidoptera 160,000 120 8,500 87 2,570 72 Mecoptera 600 9 23 3 4 2 Megaloptera 330 2 10 1 3 1 Neuroptera 6,000 17 290 12 69 6 Raphidioptera 225 2 75 2 4 1 Siphonaptera 2,600 15 260 8 62 7 Strepsiptera 600 10 30 7 10 4 Trichoptera 13,000 46 1,100 24 198 19

Totals 1,011,740 1,015 72,260 735 24,043 558

ences are discussed in each chapter. Subspecies are are created. Thus a traditional list of Trichoptera not included in the species totals, though they are families might begin: mentioned in the text where they are signifi cant. 1 ; All taxa are in alphabetical order at every level, 2 ; which may look different to some conventional lists 3 ; where ‘ similar ’ groups are put together, even 4 ; though not formally linked. The principle adopted 5 . here is that classifi cations are consulted mainly by To the specialist this may represent some kind of non - specialists, and they cannot know where to vague evolutionary sequence but it is not clear draw the boundaries unless formal higher groups whether families 1 and 2 are more closely related to

3 GENERAL INTRODUCTION

3 than family 3 is to 4, and so on. Any well - established recommended websites. This decision to omit the associations between taxa that are not represented specialist scientifi c papers will appear biased by formal groupings are described in the text; in towards certain insect groups: it will scarcely affect some cases these are the result of higher groups the Lepidoptera references, for example, whereas being split or combined by successive taxonomists. the hymenopterists will feel neglected because very Since all family names and higher taxa are listed in little of their literature is available in easily acces- the index they can be traced easily within the book. sible books or websites. This is a refl ection of how There can be differences in interpreting a ‘ British ’ groups like Hymenoptera, especially the ‘ Parasitica ’ species: some recording schemes include any are seen as the domain of professionals; perhaps species found, even as an accidental migrant or this will stimulate the wider dissemination of this import, whereas other schemes relegate these to an information in order to encourage more enthusias- appendix as ‘ unestablished ’ or even omit them alto- tic amateurs. Where identifi cation depends on spe- gether. Therefore totals cannot always be compared cialized literature this is clearly noted in the text. directly. Access to the specialist journals is often available In each group of insects outlines of the main bio- through the libraries of societies such as the British logical and morphological features are given. For Entomological & Natural History Society and Royal detailed identifi cation there are references to more Entomological Society. Even with societies that do specialized literature at the appropriate level. In the not maintain libraries such as the Amateur larger insect orders this will be at family level, but Entomologists’ Society, the contact with other spe- for the smaller orders one work may cover the cialists is immeasurably valuable to the beginner, so entire group. Much of the specialized identifi cation the message is clear: join a society! literature was covered in an earlier publication Works that can be obtained individually, such as (Barnard, 1999 ) and these lists have been updated complete issues or supplements of journals are with newer books and websites, but not with papers included, as they are frequently advertised by spe- in scientifi c journals because the latter are not easily cialist book dealers or are available directly from the available to the non- specialist. It is very noticeable publishers. Some of the older and rarer books are of in the last few years how much more information course diffi cult to obtain, but increasingly they are is available on the internet, though this ease of being digitized and made available online. access brings as many problems as advantages. In Species currently having legal protection in the general websites are not moderated, so the informa- UK are listed, though the status of these will vary tion they present can be incorrect or inappropriate over time; many other species are cited on various for certain purposes and the risks of misidentifying lists of conservation status. The exact details of the specimens from such sites are very real. Inexperienced protection are not listed here, and the latest infor- users are often unaware of the potential for gross mation should always be checked on the Joint errors from using accurate yet inappropriate sites; Nature Conservation Committee (JNCC) website numerous enquiries sent to the Royal Entomological ( http://www.jncc.gov.uk ). Inevitably it tends to be Society read something like, ‘ I found this insect in the larger and more conspicuous species that have my garden and have discovered a picture that looks gained any legal protection, and under current UK like it on an Australian website, so has it been intro- and European legislation there are just 55 British duced to the UK?’ For this reason, websites are species in this category. These comprise ten species mentioned only when they are known to be accu- of Coleoptera, 38 Lepidoptera (of which 30 are but- rate and useful, though the absence of a site does terfl ies), three Odonata, three Orthoptera and one not necessarily indicate the opposite. There is also Hemiptera. There are a great many more species the problem of URLs that change over time; some whose status causes concern; some of these are the organizations helpfully provide long - term links to subjects of species action plans and more informa- their new websites, but not always. Not all sites tion on these will be found on the appropriate web- contain primary data, but they may include valua- sites listed throughout the book. Perhaps more ble metadata, acting as useful portals to other informative is the List of UK Priority Species, result- sources of information. ing from the UK Biodiversity Action Plan ( http:// Articles published in scientifi c journals are www.ukbap.org.uk ). This UKBAP list includes 349 becoming more diffi cult to access as fewer libraries species of insects, comprised as follows: subscribe to the journals. In some cases taxonomic • Lepidoptera 176 species (of which 152 are papers are available online, though often needing a moths); paid subscription, but they may also be listed on the • Coleoptera 78 species;

4 CHAPTER 1

• Diptera 35 species; Developments continue in this fi eld and the true • Hymenoptera 35 species; phylogenetic relationships between the groups of • Hemiptera 10 species; insects are now becoming much clearer; however • Orthoptera 4 species; the story is far from complete. In some cases mor- • Trichoptera 4 species; phological and molecular data are in confl ict, and • Plecoptera 2 species; there is no agreement on an unambiguous higher • Ephemeroptera 2 species; classifi cation of all the insect orders at the present • Odonata 2 species; time. The relationships between the orders of • Neuroptera 1 species. insects are discussed in the appropriate chapters, Although this gives better coverage to some of the but any attempt to produce a formal classifi cation less conspicuous groups, it will be noted that only at this level generates an awkward hierarchy of eleven of the 24 orders of Insecta are represented. names, with some groups of uncertain affi nities; All species on this UKBAP list are noted in the therefore a more informal system seems most appropriate chapters in the book, under their cor- appropriate at present (Grimaldi & Engel, 2005 ). responding family. The authors of scientifi c names These issues are discussed by Cranston and Gullan are not given, as these are easily obtained from (2009) and a broad account of the relationships standard checklists. Species on these various lists of between and other groups is given by conservation concern have been given common Thorp (2009) . names if they did not already possess one; some of The family classifi cation used in this book is these names have changed over time, and the latest given at the end of this chapter. Several names are ones are not always correctly quoted on websites. deliberately not given any particular rank in order Clearly the scientifi c name is the defi nitive one, and to highlight their informal nature. In some texts, should always be used in preference. groups such as orthopteroids and hemipteroids are given formal names as the superorders Orthop- teroidea and Hemipteroidea, but these should be CLASSIFICATION OF INSECTS AND avoided as they resemble superfamily names, THEIR RELATIVES which always take the ‘ oidea ’ suffi x. Although the International Code of Zoological Nomenclature The exact relationships of the more primitive does not cover the formation of names above the insects, especially the ‘ apterygotes ’ has challenged family group level it seems sensible to avoid any authors for many decades. This can be seen in the possible confusion. various editions of the well - known Imms ’ General Any rigorous analysis of relationships based on Textbook of Entomology : in the fi rst edition (Imms, monophyly tends to create new groupings that have 1925 ) the Thysanura (including Diplura), Protura no equivalents in the everyday world. At the same and Collembola are placed unequivocally in the time many older convenience groups are lost, a subclass Apterygota on the grounds that they good example being the ‘ invertebrates ’ . It is clear shared the primitive wingless condition. This was that the Entognatha are distinct from the Insecta, repeated in other early editions, but by the time of but it seems reasonable to retain the common name the ninth edition (Richards & Davies, 1957 ) things of ‘ insects ’ for all the . Otherwise we have had changed; the subclass Apterygota was retained the awkward terminology of ‘ true insects ’ for the but with the admission that it was a ‘ diverse, Class Insecta, and no common name at all for the perhaps polyphyletic, assemblage ’ and acknowl- Class Entognatha or for the Hexapoda, except for edging that some authors argued that only the the somewhat artifi cial ‘ hexapods ’ . Thysanura seemed closely related to the true insects. In the tenth edition (Richards & Davies, ORDERS OF INSECTS NOT FOUND 1977 ) the discrepancies could no longer be ignored IN THE BRITISH ISLES and the authors admited that the apterygote orders should no longer be grouped together as a single There are just four insect orders not found in Britain, subclass ‘ as was done in older classifi cations ’ . By though one of them occurs in Europe; all are this time the subdivision of the Thysanura into two quite small and all are in the Polyneoptera (the distinct monophyletic groups, the Archaeognatha Orthopteroid group). (or Microcoryphia) and Zygentoma, was becoming Embioptera (also known as Embiidina): the web- established though Richards and Davies conserva- spinners are a fairly small group with around 450 tively retained them as suborders. known species, found mainly in the tropical regions

5 GENERAL INTRODUCTION of all continents; they also extend into some tem- the RES really beginning in 1806 (Gardiner, 2002 ), perate parts of the USA and southern Europe. which would make it the oldest entomological Around a dozen species are found in southern society in the world. At present that record is con- Europe in the families Embiidae and Oligotomidae. sidered to be held by the Soci é t é Entomologique de The Embioptera, together with their sister group France, so any further clarifi cation of the early Zoraptera, seem distantly related to Plecoptera, but origins of the RES might reverse this position! It has also share some characteristics with the Phasmida been suggested that the notorious lawsuit brought and Orthoptera. Their most distinctive feature is a by J.F. Stephens against J. Rennie in 1832/3 for large silk - producing gland in each fore tarsus, alleged plagiarism, helped to unite entomologists in found in both sexes and in the nymphal stages; the a common cause, which led in turn to the formation silk is used to build extensive gallery systems in the of the Entomological Society itself (Clark, 2009 : 10). soil, leaf litter or on tree bark. Through the early years it is impossible to say Grylloblattaria (also known as Grylloblattodea): whether the Society collectively infl uenced the a small and obscure order with fewer than 30 species direction of entomological research any more than found only in the western USA and Canada, and the individual members did; the membership was parts of China, Korea and Japan. Sometimes known relatively small (growing from 127 members in as rock crawlers or ice crawlers, they are usually 1834 to 178 in 1848) and their personal interests found at high altitudes, and share some characters and efforts effectively set the agenda of the Society’ s with the Dictyoptera and Phasmida. activities. Through the fi rst half of the 19th century Mantophasmatodea: these small predatory the importance of specialists in insect taxonomy insects are the most recently discovered insect order, was becoming clear; the Society had of course con- recognized only in 2001. Fewer than 20 species are tributed greatly to this advance by publishing known, all in southern and east Africa. Because they papers in its Transactions from its founding date but hold the distal ends of their tarsi off the ground it was decided to compile an authoritative cata- when walking they have been dubbed heelwalkers; logue of British insects. There had been several their phylogenetic relationships are not yet clear such attempts made by individual entomologists in though they have strong links with the Gryllobla- previous years, beginning with Forster (1770) , who ttaria, and some authors place the two groups listed just under 1000 species, plus a few related together in the Notoptera. groups of invertebrates. Samouelle’ s (1819) book Zoraptera: another obscure order with only also included spiders, mites, crustaceans and myr- around 35 known species principally in tropical iapods, but his total was around 4000 species. Two regions, living in rotting wood and feeding mainly important works appeared ten years later; Curtis on fungal hyphae. They have no common name, (1829) mentioned around 15,000 species, but many and live in small colonies, superfi cially resembling of these were undescribed or were synonyms. termites and Psocoptera, but they are probably the Stephens’ (1829) total of around 10,000 species may sister group of the Embioptera. be a more accurate refl ection of the knowledge at the time though he later revised the fi gure down to THE ROLE OF THE ROYAL nearer 9000 (Stephens, 1846 ). The British Museum ENTOMOLOGICAL SOCIETY IN BRITISH had begun its Lists of the Specimens of British Animals INSECT TAXONOMY in the Collection of the British Museum in 1848, but these were based on the museum ’ s collections only, When the current RES was instituted in 1833, its and the series was never fi nished. So, in 1851 the stated intention was ‘ the improvement and diffu- Society resolved to produce a work entitled Insecta sion of Entomological Science’ and at the Jubilee Britannica, using different authors for each major Address in 1883 the then President, J.W. Dunning, group of insects. In the end the series was pub- asserted that this aim was being successfully exe- lished by Lovell Reeve, but only two sections cuted, a view that is even more true today. The early ever appeared, three volumes on the Diptera by origins of the Society have been well described Walker (1851 – 6) and the Lepidoptera Tineina by before (Neave, 1933 ). The fi rst Entomological Stainton (1854) ; it seems that the Society’ s strict Society of London was actually founded in 1806, rules about the arrangement and contents of the with Transactions published between 1807 and 1812; works deterred any more authors from contribut- although there was little further activity until 1833 ing (Neave, 1933 ). there is no evidence that this earlier group was actu- A new catalogue appeared by the Rev. Morris ally disbanded. One can certainly make a case for (1865 – 7) , listing 12,454 species and based largely on

6 CHAPTER 1

existing lists, but the complete version of this is a publication of short papers and descriptions of new rare book. In 1867 the Entomological Society began species. a similar project to the one abandoned some years The stalled Generic Nomenclature project was before, but this time with more determination. The effectively saved by the appearance of the fi rst Catalogue of British Insects was fi rst proposed by the modern checklist of British insects (Kloet & Hincks, Neuropterist Robert McLachlan. A committee was 1945 ). This was a list begun in the 1930s by the formed to oversee its production, but again there amateur entomologist and Manchester business- were arguments about the nomenclature and groups man George Sidney Kloet (1904 –81) and later aided such as Coleoptera were never published; however by Walter Douglas Hincks (1906 – 61), an authority fi ve volumes did appear (McLachlan & Eaton, 1870 ; on Dermaptera with a keen interest in other groups Smith, 1871 ; Marshall, 1872, 1873 ; Douglas & Scott, such as Coleoptera, and who eventually became 1876 ). By the end of this series only around 3400 Keeper of Entomology at Manchester Museum. species had been catalogued, and it was to be many They were well aware of the shortcomings of their years before a similar project was attempted. In the work, which listed just over 20,000 species, but its meantime separate lists of many groups of insects publication stimulated others to bring the critical were published by individual authors and inevita- sections up to scratch. As O.W. Richards perspica- bly they vary in quality and comprehensiveness, the ciously put it, ‘ Naturally, almost as soon as the list very diffi culties that a coordinated project was was published, it was criticized by the various designed to avoid. experts who had always been too busy to produce In 1933, the year that the Society received its lists themselves’ (Kloet & Hincks, 1964 ). Although ‘ Royal ’ status, the Committee on Generic Nomen- both authors were Fellows of the RES, they had clature was set up, with the object ‘ to prepare lists their book published privately, perhaps demon- of specifi c names to be fi xed as genotypes of genera strating that keen individuals can often be more of British insects with a view to the suspension effi cient at fi nishing projects than committees, where necessary of the law of priority in respect of however well- meaning the latter may be. those generic names’ which sounded an ambitious During this same decade, the RES launched its project. Eleven subcommittees were formed to most successful and long - standing series, the cover all groups of insects, but it was not long Handbooks for the Identifi cation of British Insects . The before the enormity and complexity of the task were fi rst parts were published in July 1949 and they apparent, partly because fi xing the type species of have appeared at irregular intervals up to the genera is clearly a task that requires study of the present day, with some of the more popular parts world fauna, not just the British species. The main being reprinted or revised. The ambitious intention committee had to admit in its fi rst report (Committee was to cover all the British orders, and altogether on Generic Nomenclature, 1933 ) that their optimism ten volumes were planned, with vol. 1 including the for completing the task in a reasonable time was smaller insect orders, vol. 2 the Hemiptera, vol. 3 unrealistic, stating that for some subcommittees the Lepidoptera, vols 4 and 5 the Coleoptera, vols ‘ several years must necessarily elapse before their 6, 7 and 8 the Hymenoptera, and vols 9 and 10 the work is complete’ . In the event, eight parts of the Diptera. A projected vol. 11 was set aside for updates proposed lists were published and formed Volume to the checklists. Inevitably the numbering system 1, together with the Committee’ s report, but no became rather complicated because individual more appeared (Hemming, 1934 ; Cowley, 1935 ; authors varied in the coverage of each handbook. Killington, 1937 ; Richards, 1937 ; Andrewes, 1939 ; The original intention was no doubt to have simple Balfour - Browne, 1940 ; China, 1943 ; Tottenham, subsections for each family or Superfamily, so that 1949 ). One suspects that many readers found the parts might be numbered vol. 5, part 3, or if neces- included checklists of insects more useful than sary vol 5, part 3a. However, diffi culties arose when the often lengthy discussions on nomenclatural some authors could only cover a subfamily at a problems. time, leading to complicated numbers like vol. 8, At the same time as these lists were being pre- part 3(dii); recently this system has been simplifi ed, pared, the RES was recognizing the increasing partly in recognition of the fact that some volumes importance of specialized journals to cover taxo- will never be complete. Because lepidopterists have nomic entomology. Longer revisionary papers had always published their own books in their preferred always been published in the Transactions, but in ways, vol. 3 has never been used. 1932 a new journal Stylops was begun, named after The early handbooks all carried lists of books the Society’ s logo; it was designed for the rapid already published by the RES, whether handbooks

7 GENERAL INTRODUCTION or separate publications like the Centenary History Entomology (Series B), and the latest journal for taxo- (Neave, 1933 ); in fact the Generic Names of British nomic publications is Systematic Entomology , begun Insects project was still being publicized until the in 1976. early 1960s, with the optimistic statement ‘ Parts so For a more general view of the history of ento- far published ’ though no more appeared after 1949. mology in Europe there are several recent books, However, the Handbooks had not supplanted that including d ’ Aguilar (2006) , Clark (2009) , Gilbert series because the early sections did not contain (1977, 2005, 2007), Salmon (2000) and Salmon and checklists of the groups they covered, an omission Edwards (2005) . The classic work on the earliest that was not put right until the 1970s. entomological history on a world scale is Boden- In the early 1960s the decision was made to heimer (1928 – 9) . update the Kloet and Hincks checklist. Hincks had died in 1961, and Kloet was not inclined to carry on GENERAL ENTOMOLOGICAL WORKS with the project alone so all their lists were handed over to the RES, along with the rights to publish all For the identifi cation of British insects there are future editions. The task was given to small groups many useful series, such as the Naturalists ’ of experts and the work eventually appeared in fi ve Handbooks , Handbooks for the Identifi cation of British volumes spread over 12 years. In recognition of the Insects (published by the Royal Entomological pioneering work of the original authors the series Society) and the AIDGAP keys (published by the was still entitled the Kloet and Hincks lists. Field Studies Council); these are listed in their In date order these new lists were: appropriate chapters throughout this book. For 1964, Kloet & Hincks, A check list of British insects, identifying insects in particular habitats or on par- second edition (revised). Part 1: small orders and ticular plants there are several useful titles in the Hemiptera. Handbooks for the Identifi cation of Naturalists’ Handbooks series, though these will not British Insects 11(1): 119 pp. always take identifi cations to species level. They 1972, Kloet & Hincks, A check list of British insects, cover topics such as insects on nettles (Davis, 1991 ), second edition (revised). Part 2: Lepidoptera. on thistles (Redfern, 1995 ), on dock plants (Salt & Handbooks for the Identifi cation of British Insects Whittaker, 1998 ), on cherry trees (Leather & Bland, 11(2): 153 pp. 1999 ), on cabbages and oilseed rape (Kirk, 1992 ), 1976, Kloet & Hincks, A check list of British insects, under logs and stones (Wheater & Reed, 1996 ), and second edition (revised). Part 5: Diptera and predators (Rotheray, 1989 ). There are similar Siphonaptera. Handbooks for the Identifi cation of titles in the ever- growing AIDGAP series. Insects British Insects 11(5): 139 pp. that cause plant galls are particularly well covered 1977, Kloet & Hincks, A check list of British insects, by several useful guides at different levels (Redfern second edition (revised). Part 3: Coleoptera and & Askew, 1998 ; Redfern & Shirley, 2002, 2004 ). Strepsiptera. Handbooks for the Identifi cation of At a more basic level, for those with little previ- British Insects 11(3): 105 pp. ous knowledge of insects, the books by Chinery 1978, Kloet & Hincks, A check list of British insects, (1986, 1993, 2009) cannot be bettered; and for a second edition (revised). Part 4: Hymenoptera. general entomological textbook Gullan and Handbooks for the Identifi cation of British Insects Cranston (2005) is the most comprehensive. 11(4): 159 pp. A completely new edition of checklists was planned REFERENCES in the 1990s; to date only the Diptera list has been published (Chandler, 1998 ) and it may be that all D ’ Aguilar , J. 2006 . Histoire de l ’ entomologie . Delachaux & future lists in this series will be published online to Niestlé , Paris . allow easier updating. Already there are many Andrewes , H.E. 1939 . The generic names of the British checklists being published on various websites, and Carabidae, with a check list of British species. Part 6 . the more reliable of these are listed in the appropri- The generic names of British insects . Vol. 1 : 151 – 92 . Royal ate chapters of this book. Entomological Society , London . Balfour - Browne , W.A.F. 1940 . The generic names of the Meanwhile the RES continues to support insect British Hydradephaga, with a check list of British taxonomy in many other ways. The journal Stylops species. Part 7 . The generic names of British insects . Vol. 1 : became Proceedings (Series B: Taxonomy) in 1936 as 193 – 209 . Royal Entomological Society , London . part of a general re- naming of the Society’ s journals. Barnard , P.C. (ed.) 1999 . Identifying British insects and In 1971 this was merged with Proceedings (Series A: arachnids: an annotated bibliography of key works . Cam- General Entomology) to become the Journal of bridge University Press , Cambridge .

8 CHAPTER 1

Bodenheimer , F.S. 1928 – 9 . Materialen zur Geschichte der Imms , A.D. 1925 . A general textbook of entomology . Methuen , Entomologie bis Linné . W. Junk , Berlin . 2 vols. London . Chandler , P.J. (ed.) 1998 . Checklists of insects of the Killington , F.J. 1937 . The generic names of the British British Isles (new series). Part 1: Diptera . Handbooks for Neuroptera, with a check list of British species. Part 4 . the identifi cation of British insects 12 ( 1 ): 234 pp. The generic names of British insects . Vol. 1 : 63 – 80 . Royal China , W.E. 1943 . The generic names of the British Entomological Society , London . Hemiptera– , with a check list of British Kirk , W.D.J. 1992 . Insects on cabbages and oilseed rape . species. Part 8 . The generic names of British insects . Vol. 1 : Naturalists’ Handbooks no. 18, Richmond , Publishing, 209 – 342 . Royal Entomological Society , London . Slough. Chinery, M. 1986 . Collins guide to the insects of Britain and wes- Kloet , G.S. & Hincks , W.D. 1945 . A check list of British tern Europe . Collins Pocket Guide, HarperCollins , London . insects . Privately published , Stockport . Chinery , M. 1993 . Insects of Britain and northern Europe Kloet , G.S. & Hincks , W.D. 1964 . A check list of British ( 3rd edn. ). Collins Field Guide, HarperCollins , London . insects (2nd edn., revised). Part 1: small orders and Chinery , M. 2009 . Collins complete guide to British insects . Hemiptera. Handbooks for the identifi cation of British HarperCollins , London . insects 11 ( 1 ): 119 pp. Clark , J.F.M. 2009 . Bugs and the Victorians . Yale University Leather , S.R. & Bland , K.P. 1999 . Insects on cherry trees . Press , New Haven & London . Naturalists’ Handbooks no. 27, Richmond , Publishing, Committee on Generic Nomenclature . 1933 . Recommen- Slough. dations relating to the publication of the Committee ’ s Marshall , T.A. 1872 . A catalogue of British Hymenoptera; reports. Part 1. The generic names of British insects . Vol. 1 : Chrysididae, Ichneumonidae, Braconidae and Evaniidae . 1 – 6 . Royal Entomological Society , London . Entomological Society of London . Cowley , J. 1935 . The generic names of the British Odonata, Marshall , T.A. 1873 . A catalogue of British Hymenoptera; with a check list of British species. Part 3 . The generic Oxyura . Entomological Society of London . names of British insects . Vol. 1 : 43 – 60 . Royal Entomological McLachlan , R. & Eaton , A.E. 1870 . A catalogue of British Society , London . Neuroptera . Entomological Society of London . Cranston , P.S. & Gullan , P.J. 2009 . Phylogeny of insects . Morris , F.O. 1865 – 7 . A catalogue of British insects, in all the In: Resh , V.H. & Card é , R.T. (eds.) Encyclopedia of insects orders . Longmans, Green, Reader & Dyer , London ( 2nd edn. ). Academic Press/Elsevier , San Diego & [issued in parts]. London, pp 780 – 93 . Neave , S.A. 1933 . The history of the Entomological Society of Curtis , J. 1829 . A guide to an arrangement of British insects . London, 1833 – 1933 . Entomological Society , London . London . Redfern , M. 1995 . Insects and thistles ( 2nd edn. ). Naturalists ’ Davis , B.N.K. 1991 . Insects on nettles ( 2nd edn. ). Naturalists ’ Handbooks no. 4, Richmond , Publishing, Slough . Handbooks no. 1, Richmond Publishing , Slough . Redfern , M. & Askew , R.R. 1998 . Plant galls ( 2nd edn. ). Douglas , J.W. & Scott , J. 1876 . A catalogue of British Naturalists’ Handbooks no. 17, Richmond , Publishing, Hemiptera; Heteroptera and Homoptera . Entomological Slough. Society of London . Redfern , M. & Shirley , P. 2002 . British plant galls: identi- Foottit , R.G. & Adler , P.H. (eds.) 2009 . Insect biodiversity: fi cation of galls on plants and fungi . Field Studies Council , science and society . Blackwell Publishing , Chichester . Preston Montford , OP270. Forster , J.R. 1770 . A catalogue of British insects . Warrington . Redfern , M. & Shirley , P. 2004 . A guide to plant galls Gardiner , B.O.C. 2002 . A short account of the Royal in Britain. Field Studies Council, Preston Montford, Entomological Society and of the progress of entomol- OP91. ogy in Great Britain (1833 – 1999 ). In: Pedersen , B. (ed.) Resh, V.H. & Cardé , R.T. (eds.) 2009 . Encyclopedia of insects A guide to the archives of the Royal Entomological Society . (2nd edn.). Academic Press/Elsevier, San Diego & London. Ashgate , Aldershot , pp. 1 – 30 . Richards , O.W. 1937 . The generic names of the British Gilbert , P. 1977 . A compendium of the biographical literature Hymenoptera, Aculeata, with a check list of British on deceased entomologists. British Museum (Natural species. Part 5 . The generic names of British insects . Vol. 1 : History), London . 79 – 149 . Royal Entomological Society , London . Gilbert , P. 2005 . The Entomological Club and Verrall Supper: Richards , O.W & Davies , R.G. 1957 . Imms ’ general text- a history (1826 – 2004) . The Entomological Club , London . book of entomology ( 9th edn. ). Methuen , London . Gilbert , P. 2007 . A source book for biographical literature on Richards , O.W & Davies , R.G. 1977 . Imms ’ general text- entomologists . Backhuys Publishers , Leiden . book of entomology ( 10th edn. ). Chapman & Hall , London . Grimaldi , D. & Engel , M.S. 2005 . Evolution of the insects . Rotheray , G.E. 1989 . Aphid predators . Naturalists ’ Hand- Cambridge University Press , Cambridge . books no. 11, Richmond , Publishing, Slough [reprinted Gullan , P.J. & Cranston , P.S. 2005 . The insects: an outline 2003]. of entomology ( 3rd edn. ). Blackwell , Oxford . Salmon , M.A. 2000 . The Aurelian legacy: British butterfl ies Hemming , F. 1934 . The generic names of the British and their collectors . Harley Books , Great Horkesley . Rhopalocera, with a check list of British species. Part 2 . Salmon , M.A. & Edwards , P.J. 2005 . The Aurelian ’ s fi reside The generic names of British insects . Vol. 1 : 6 – 40 . Royal companion: an entomological anthology . Paphia Books , Entomological Society , London . Lymington .

9 GENERAL INTRODUCTION

Salt , D.T. & Whittaker , J.B. 1998 . Insects on dock plants . Thorp, J.H. 2009 . Arthropoda and related groups. In: Resh , Naturalists’ Handbooks no. 26, Richmond , Publishing, V.H. & Cardé , R.T. (eds.) Encyclopedia of insects ( 2nd edn. ). Slough. Academic Press/Elsevier, San Diego & London, pp 50 – 6 . Samouelle , G. 1819 . A nomenclature of British entomology . Tottenham , C.E. 1949 . The generic names of the British London . Staphylinidae, with a check list of British species. Part Secord , J.A. 1996 . The crisis of nature . In: Jardine , N. , 9 . The generic names of British insects . Vol. 1 : 343 – 466 . Secord , J.A. & Spary , E.C. (eds.) Cultures of natural Royal Entomological Society , London . history . Cambridge University Press , Cambridge , pp. Walker , F. 1851 – 6 . Insecta Britannica. Diptera . Reeve , 447 – 59 . London , 3 vols. Smith , F. 1871 . A catalogue of British Hymenoptera; Aculeata . Wheater , C.P. & Read , H.J. 1996 . Animals under logs and Entomological Society of London . stones. Naturalists’ Handbooks no. 22, Richmond , Stainton , H.T. 1854 . Insect Britannica. Lepidoptera: Tineina . Publishing, Slough . Reeve , London . Wheeler , Q.D. 2008 . Taxonomic shock and awe . In: Stephens , J.F. 1829 . A systematic catalogue of British insects . Wheeler , Q.D. (ed.) The new taxonomy . Systematics London , 2 vols. Association/CRC Press , London/New York , pp 211 – 26 . Stephens , J.F. 1846 . Illustrations of British entomology Yoon , C.K. 2009 . Naming nature: the clash between instinct (Suppl.) . Henry G. Bohn , London . and science . Norton , New York .

FAMILY CLASSIFICATION OF THE BRITISH INSECTS Each group of higher taxa is arranged in alphabeti- Order Zygentoma (silverfi sh and fi rebrats) cal order Family Lepismatidae Phylum Arthropoda: Subphylum Hexapoda Insecta Palaeoptera Class Entognatha Order Ephemeroptera (mayfl ies) Order Collembola (springtails) Superfamily Baetoidea Suborder Entomobryomorpha Families Ameletidae, Baetidae, Siphlonuridae Superfamily Entomobryoidea Superfamily Caenoidea Families Cyphoderidae, Entomobryidae, Isotomidae Family Caenidae Superfamily Tomoceroidea Superfamily Ephemerelloidea Familes Oncopoduridae, Tomoceridae Family Ephemerellidae Suborder Neelipleona Superfamily Ephemeroidea Family Neelidae Families Ephemeridae, Potamanthidae Suborder Poduromorpha Superfamily Heptagenioidea Superfamily Hypogastruroidea Families Arthropleidae, Heptageniidae Family Hypogastruridae Superfamily Leptophlebioidea Superfamily Neanuroidea Family Leptophlebiidae Families Brachystomellidae, Neanuridae, Odonte- Order Odonata (damselfl ies and dragonfl ies) llidae Suborder Anisoptera Superfamily Onychiuroidea Superfamily Aeshnoidea Families Onychiuridae, Tullbergiidae, Poduridae Families Aeshnidae, Gomphidae Suborder Symphypleona Superfamily Cordulegastroidea Families Arrhopalitidae, Bourletiellidae, Dicyrtomi- Family Cordulegastridae dae, Katiannidae, Sminthuridae, Sminthuri didae Superfamily Libelluloidea Order Diplura (two -tailed bristletails) Families Corduliidae, Libellulidae Family Campodeidae Suborder Zygoptera Order Protura (proturans) Superfamily Calopterygoidea Suborder Eosentomata Family Calopterygidae Family Eosentomidae Superfamily Coenagrionoidea Suborder Acerentomata Families Coenagrionidae, Platycnemididae Families Acerentomidae, Protentomidae Superfamily Lestoidea Family Lestidae Class Insecta ‘ Apterygota ’ [ = Thysanura] Insecta Pterygota Polyneoptera Order Archaeognatha (bristletails) Order Dermaptera (earwigs) Family Machilidae Suborder Forfi culina

10 CHAPTER 1

Families , Forfi culidae, , Family Spongiphoridae Superfamily Order Dictyoptera (cockroaches, termites and Families Gerridae, mantids) Superfamily Hebroidea Suborder Blattodea Family Hebridae Families Blaberidae, Blattellidae, Blattidae Superfamily Hydrometroidea Suborder Isoptera Family Family Rhinotermitidae Superfamily Mesovelioidea Suborder Mantodea Family Family Mantidae Superfamily Naucoroidea Order Orthoptera (grasshoppers, crickets and Families , bush - crickets) Superfamily Nepoidea Suborder Caelifera Family Superfamily Acridoidea Superfamily Notonectoidea Family Acrididae Family Superfamily Tetrigoidea Superfamily Pleoidea Family Tetrigidae Family Suborder Ensifera Infraorder Superfamily Grylloidea Superfamily Families Gryllidae, Gryllotalpidae, Mogoplistidae Family Superfamily Rhaphidophoroidea Superfamily Family Rhaphidophoridae Families , , , Stenoce- Superfamily Tettigonioidea phalidae Families Conocephalidae, Meconematidae, Phane- Superfamily ropteridae, Tettigoniidae Families , , Order Phasmida (stick insects) Superfamily Families Bacillidae, Phasmatidae Families , , Pentato- Order Plecoptera (stonefl ies) midae, , Superfamily Nemouroidea Superfamily Piesmatoidea Families Capniidae, Leuctridae, Nemouridae, Family Taeniopterygidae Suborder Superfamily Perloidea Superfamily Cercopoidea Families Chloroperlidae, Perlidae, Perlodidae Families , Superfamily Cicadoidea Insecta Pterygota Neoptera Paraneoptera Family Tibicinidae Order Hemiptera (true bugs) Superfamily Suborder Heteroptera Families Cicadellidae, Membracidae, Ulopidae Infraorder Suborder Fulgoromorpha Superfamily Families , , , Tettigome- Families , , tridae Superfamily Miroidea Suborder Families , Superfamily Aleyrodoidea Superfamily Reduvioidea Family Aleyrodidae Family Superfamily Aphidoidea Superfamily Tingoidea Family Aphididae Family Superfamily Coccoidea Infraorder Families , , , Superfamily Dipsocoroidea , , , Margaro- Families , didae, , Pseudococcidae Infraorder Superfamily Superfamily Saldoidea Families , Families Aepophilidae, Superfamily Infraorder Families , , , Superfamily Corixoidea

11 GENERAL INTRODUCTION

Order Phthiraptera (lice) Insecta Pterygota Neoptera Endopterygota = Suborder Holometabola Superfamily Gyropoidea Order Coleoptera (beetles) Familiesy , Gliricolidae Suborder Adephaga Superfamily Laemobothrioidea Superfamily Caraboidea Family Families Carabidae, Dytiscidae, Gyrinidae, Superfamily Menopodoidea Haliplidae, Hygrobiidae, Noteridae Family Menopodidae Suborder Myxophaga Superfamily Ricinoidea Superfamily Sphaeriusoidea Family Family Sphaeriusidae Superfamily Trimenoponoidea Suborder Polyphaga Family Infraorder Bostrichiformia Suborder Anoplura Superfamily Bostrichoidea Superfamily Echinophthirioidea Families Anobiidae, Bostrichidae, Dermestidae, Family Echinophthiriidae Lyctidae Superfamily Linognathoidea Superfamily Derodontoidea Families , Hoplopleuridae, Linog- Family Derodontidae nathidae, Polyplacidae, Phthiridae Infraorder Cucujiformia Superfamily Pediculoidea Superfamily Chrysomeloidea Families Haematopinidae, Pediculidae Families Cerambycidae, Chrysomelidae, Megalo- Suborder podidae, Orsodacnidae Superfamily Goniodoidea Superfamily Cleroidea Family Goniodidae Families Cleridae, Dasytidae, Malachiidae, Superfamily Philopteroidea Phloiophilidae, Trogossitidae Family Superfamily Cucujoidea Superfamily Trichodectoidea Families Alexiidae, Biphyllidae, Bothrideridae, Byt- Family uridae, Cerylonidae, Coccinellidae, Corylophidae, Order Psocoptera (booklice and barklice) Cryptophagidae, Cucijidae, Endomychidae, Ero- Suborder tylidae, Kateretidae, Laemophloeidae, Lang- Infraorder Caeciliusetae uriidae, Latridiidae, Monotomidae, Nitidulidae, Families , , Stenop- Phalacridae, Silvanidae, Sphindidae socidae Superfamily Curculionoidea Infraorder Families Anthribidae, Apionidae, Attelabidae, Family Curculionidae, Dryophthoridae, Erirhinidae, Infraorder Nanophyidae, Nemonychidae, Platypodidae, Families , , , Raymondionymidae, Rhynchitidae , , , Superfamily Lymexyloidea Family Lymexylidae Infraorder Superfamily Tenebrionoidea Family Families Aderidae, Anthicidae, Ciidae, Melan- Suborder dryidae, Meloidae, Mordellidae, Mycetophagidae, Infraorder Mycteridae, Oedemeridae, Pyrochroidae, Pyth- Families , , Sphaerop- idae, Ripiphoridae, Salpingidae, Scraptiidae, Ten- socidae ebrionidae, Tetratomidae, Zopheridae Suborder Infraorder Elateriformia Infraorder Atropetae Superfamily Buprestoidea Families , , Family Buprestidae Infraorder Psocathropetae Superfamily Byrrhoidea Family Families Byrrhidae, Dryopidae, Elmidae, Hetero- Order Thysanoptera (thrips) ceridae, Limnichidae, Psephenidae, Ptiloda- Suborder Terebrantia ctylidae Families Aeolothripidae, Thripidae Superfamily Dascilloidea Suborder Tubulifera Family Dascillidae Family Phlaeothripidae Superfamily Elateroidea

12 CHAPTER 1

Families Cantharidae, Drilidae, Elateridae, Eucne- , , , midae, Lampyridae, Lycidae, Throscidae Stenomicridae Superfamily Scirtoidea Superfamily Families Clambidae, Eucinetidae, Scirtidae Families , , Phaeomyiidae, Infraorder Scarabaeiformia , Superfamily Scarabaeoidea Superfamily Families Aegialiidae, Aphodiidae, Bolboceratidae, Families , , Sphaero- Cetoniidae, Geotrupidae, Lucanidae, Melolon- ceridae thidae, Rutelidae, Scarabaeidae, Trogidae Superfamily Infraorder Staphyliniformia Families , , , Superfamily Hydrophiloidea , , Families Georissidae, Helophoridae, Histeridae, Infraorder Hydrochidae, Hydrophilidae, Spercheidae, Superfamily Sphaeritidae Families , Superfamily Staphylinoidea Superfamily Families Hydraenidae, Leiodidae, Ptiliidae, Scyd- Families , , , maenidae, Silphidae, Staphylinidae Order Diptera (true fl ies) Superfamily Suborder Families , Oestridae, , Infraorder Sarcophagidae, Superfamily Infraorder Families , , , Superfamily Families , Superfamily Superfamily Families , , , Families , , Spaniidae, , Microphoridae Tabanidae Superfamily Infraorder Xylophagomorpha Family Superfamily Xylophagoidea Infraorder Muscomorpha Family Superfamily Lonchopteroidea Suborder Family Infraorder Superfamily Superfamily Bibionoidea Families , Phoridae, Family Superfamily Superfamily Families , Syrphidae Families Bolitophilidae, Cecidomyiidae, Diado- Infraorder Muscomorpha Schizophora cidiidae, , , Myceto- Superfamily philidae, Families , , , Chloro- Infraorder pidae, , Tethinidae Superfamily Superfamily Conopoidea Families , Chironomidae, Simu- Family liidae, Superfamily Superfamily Families , , Strongyloph- Families , Culicidae, thalmyiidae, Infraorder Superfamily Superfamily Anisopodoidea Families , Campichoetidae, , Families , Mycetobiidae , Superfamily Psychodoidea Superfamily Family Families , Superfamily Scatopsoidea Superfamily Family Families , Pseudopomyzidae Superfamily Trichoceroidea Superfamily Family Families Acartophthalmidae, , Antho- Infraorder myzidae, , , , Superfamily Ptychopteroidea

13 GENERAL INTRODUCTION

Family Order Lepidoptera (moths and butterfl ies) Infraorder Suborder Glossata Superfamily Superfamily Alucitoidea Families , Limoniidae, , Family Alucitidae Tipulidae Superfamily Bombycoidea Order Hymenoptera (ants, bees and wasps) Families Endromidae, Saturniidae, Sphingidae Suborder Apocrita Aculeata Superfamily Choreutoidea Superfamily Apoidea Family Choreutidae Families Apidae, Crabronidae, Sphecidae Superfamily Cossoidea Superfamily Chrysidoidea Family Cossidae Families Bethylidae, Chrysididae, Dryinidae, Superfamily Drepanoidea Embolemidae Families Drepanidae, Thyatiridae Superfamily Vespoidea Superfamily Epermenioidea Families Formicidae, Mutillidae, Pompilidae, Family Epermeniidae Sapygidae, Scoliidae, Tiphiidae, Vespidae Superfamily Eriocranioidea Suborder Apocrita Parasitica Family Eriocraniidae Superfamily Ceraphronoidea Superfamily Gelechioidea Families Ceraphronidae, Megaspilidae Families Agonoxenidae, Amphisbatidae, Autosti- Superfamily Chalcidoidea chidae, Batrachedridae, Blastobasidae, Chimaba- Families Aphelinidae, Chalcididae, Elasmidae, chidae, Coleophoridae, Cosmopterigidae, Encyrtidae, Eucharitidae, Eulophidae, Eupel- Depressariidae, Elachistidae, Ethmiidae, Gele- midae, Eurytomidae, Mymaridae, Ormyridae, chiidae, Momphidae, Oecophoridae, Scythrididae, Perilampidae, Pteromalidae, Signiphoridae, Stathmopodidae Tetracampidae, Torymidae, Trichogrammatidae Superfamily Geometroidea Superfamily Cynipoidea Family Geometridae Families Cynipidae, Figitidae, Ibaliidae Superfamily Gracillarioidea Superfamily Evanioidea Family Bucculatricidae, Douglasiidae, Gracillariidae, Families Aulacidae, Evaniidae, Gasteruptiidae Roeslerstammiidae Superfamily Ichneumonoidea Superfamily Hepialoidea Families Braconidae, Ichneumonidae Family Hepialidae Superfamily Mymarommatoidea Superfamily Hesperioidea Family Mymarommatidae Family Hesperiidae Superfamily Platygastroidea Superfamily Incurvarioidea Families Platygastridae, Scelionidae Families Adelidae, Heliozelidae, Incurvariidae, Superfamily Proctotrupoidea Prodoxidae Families Diapriidae, Heloridae, Proctotrupidae Superfamily Lasiocampoidea Superfamily Trigonaloidea Family Lasiocampidae Family Trigonalidae Superfamily Nepticuloidea Suborder Symphyta Family Nepticulidae, Opostegidae Superfamily Cephoidea Superfamily Noctuoidea Family Cephidae Families Arctiidae, Ctenuchidae, Lymantriidae, Noc- Superfamily Orussoidea tuidae, Nolidae, Notodontidae, Thaumetopoeidae Family Orussidae Superfamily Papilionoidea Superfamily Pamphilioidea Families Lycaenidae, Nymphalidae, Papilionidae, Family Pamphilidae Pieridae, Riodinidae Superfamily Siricoidea Superfamily Pterophoroidea Family Siricidae Family Pterophoridae Superfamily Tenthredinoidea Superfamily Pyraloidea Families Argidae, Blasticotomidae, Cimbicidae, Families Crambidae, Pyralidae Diprionidae, Tenthredinidae Superfamily Schreckensteinioidea Superfamily Xiphrydioidea Family Schreckensteiniidae Family Xiphydriidae Superfamily Sesiodea Superfamily Xyeloidea Family Sesiidae Family Xyelidae Superfamily Tineoidea

14 CHAPTER 1

Families Psychidae, Tineidae Family Pulicidae Superfamily Tischerioidea Superfamily Vermipsylloidea Family Tischeriidae Family Vermipsyllidae Superfamily Tortricoidea Order Strepsiptera (stylops) Family Tortricidae Families Elenchidae, Halictophagidae, Stylopidae, Superfamily Yponomeutoidea Xenidae Families Acrolepiidae, Bedelliidae, Glyphiptery- Order Trichoptera (caddisfl ies) gidae, Heliodinidae, Lyonetiidae, Plutellidae, Suborder Yponomeutidae, Ypsolophidae Superfamily Superfamily Zygaenoidea Families Ecnomidae, , Polycentro- Families Limacodidae, Zygaenidae podidae, Suborder Zeugloptera Superfamily Philopotamoidea Superfamily Micropterigoidea Family Philopotamidae Family Micropterigidae Suborder Order Mecoptera (scorpionfl ies) Superfamily Leptoceroidea Familie Boreidae, Panorpidae Families , , Order Megaloptera (alderfl ies) Superfamily Family Sialidae Families , , , Order Neuroptera (lacewings and ) , Suborder Hemerobiiformia Superfamily Phryganeoidea Families , , Hemero- Family biidae, , Sisyridae Superfamily Suborder Families , Family Myrmeleontidae Suborder Order Raphidioptera (snakefl ies) Superfamily Glossosomatoidea Family Raphidiidae Family Glossosomatidae Order Siphonaptera (fl eas) Superfamily Hydroptiloidea Superfamily Ceratophylloidea Family Hydroptilidae Families Ceratophyllidae, Ischnopsyllidae, Lepto- Superfamily psyllidae Family Rhyacophilidae Superfamily Hystrichopsylloidea Families Ctenophthalmidae, Hystrichopsyllidae Superfamily Pulicoidea

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