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    Shieldbugs
    Shieldbugs
    Shieldbugs
    Ebook973 pages8 hours

    Shieldbugs

    By Richard Jones

    ()

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    About this ebook

    An eagerly anticipated addition to the New Naturalist series.

    The shieldbug is an amazing and beautiful species, rich with diversity in shape, form, size, life history, ecology, physiology and behaviour. But they are not commonly known, outside of specialist circles.

    Richard Jones’ groundbreaking New Naturalist volume on shieldbugs encourages those enthusiasts who would otherwise be put off by the, to date, rather technical literature that has dominated the field, providing a comprehensive natural history of this fascinating and beautiful group of insects.

    LanguageEnglish
    PublisherHarperCollins UK
    Release dateJul 6, 2023
    ISBN9780008334901
    Shieldbugs
    Author

    Richard Jones

    Richard Jones has more than fifteen years of experience in the creative arts and worked in a children's library for over a decade. Combining his two passions, he began illustrating children's books in 2016. Winter Dance is his first picture book in the US. He lives in Devon, England. www.paintedmouse.com

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      Book preview

      Shieldbugs - Richard Jones

      Copyright

      William Collins

      An imprint of HarperCollinsPublishers

      1 London Bridge Street

      London SE1 9GF

      WilliamCollinsBooks.com

      HarperCollinsPublishers

      Macken House, 39/40 Mayor Street Upper

      Dublin 1, D01 C9W8, Ireland

      This eBook edition published by William Collins in 2023

      Copyright © Richard Jones 2023

      Photographs © individual photographers unless otherwise credited

      Cover art by Robert Greenhalf

      Richard Jones asserts his moral right to be identified as the author of this work

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

      All rights reserved under International and Pan-American Copyright Conventions. By payment of the required fees, you have been granted the non-exclusive, non-transferable right to access and read the text of this eBook on-screen. No part of this text may be reproduced, transmitted, downloaded, decompiled, reverse engineered, or stored in or introduced into any information storage and retrieval system, in any form or by any means, whether electronic or mechanical, now known or hereinafter invented, without the express written permission of HarperCollins Publishers.

      Source ISBN: 9780008334895

      Ebook Edition © 2023 ISBN: 9780008334901

      Version: 2023-06-20

      EDITORS

      SARAH A. CORBET, SCD

      DAVID STREETER, MBE, FRSB

      JIM FLEGG, OBE, FIHORT

      PROF. JONATHAN SILVERTOWN

      PROF. BRIAN SHORT

      *

      The aim of this series is to interest the general

      reader in the wildlife of Britain by recapturing

      the enquiring spirit of the old naturalists.

      The editors believe that the natural pride of

      the British public in the native flora and fauna,

      to which must be added concern for their

      conservation, is best fostered by maintaining

      a high standard of accuracy combined with

      clarity of exposition in presenting the results

      of modern scientific research.

      Contents

      Cover

      Title Page

      Copyright

      About the Editors

      Editors’ Preface

      Authors’ Acknowledgements

      1 What is a Shieldbug?

      2 Shieldbug Structure

      3 Life Histories: Breeding and Feeding

      4 Dangers and Defences

      5 Evolution of Shieldbugs and a History of British Species

      6 History of Shieldbug Study

      7 Key to British Shieldbug Species

      8 British Shieldbug Species

      9 How to Study Shieldbugs

      Appendix 1: Selected British and Irish Shieldbug Foodplants and Prey

      Appendix 2: Shieldbug-related Websites and Apps

      Glossary

      References

      Picture Credits

      Species Index

      General Index

      About the Publisher

      Editors’ Preface

      Among other aspects of natural history, the New Naturalist Library has a distinguished history of publishing titles on individual insect groups. The very first volume in the series, published in 1945, was E. B. Ford’s Butterflies , and over the years this was followed by books on honeybees, moths, bumblebees, dragonflies, ants, ladybirds, grasshoppers and crickets, beetles, and most recently solitary bees. Whereas most of these groups were already popular when the books were written, shieldbugs currently have a smaller fan club and are less well known to non-specialists.

      Following his excellent book on Beetles, we are delighted that Richard Jones has been willing to introduce us to this new group. We expect that this book, with its lively and informative text and its colourful illustrations, will make shieldbugs attractive and accessible to many more naturalists. The keys and illustrations, supported by notes on habitats and food plants, will be of great assistance in species identification, and the inclusion of recent additions to the British shieldbug fauna, and others likely to follow, challenges us to watch out for new arrivals from the continent.

      This book will bring great pleasure and satisfaction to those who are inspired to study shieldbugs seriously, but even readers who have no intention of grubbing around the base of food plants or sniffing the bugs on their broad beans will enjoy the vivid colours and curious antics presented in this book. Perhaps in the future we will look back at its publication as the trigger for a fresh surge of interest in British shieldbugs.

      Author’s Foreword and Acknowledgements

      Shieldbugs are lovely. Why else would we give them such an elegant heroic heraldic name? Sure enough, they are glossy, chunky, often large and brightly coloured, and suitably shield-shaped. Shieldbugs have a regal and aristocratic air about them. My stylish pentatomid-design enamel lapel badge often draws comments because its obvious shield shape looks as if it could be the emblem of some noble family. It is, of course, but not necessarily in the way most people imagine.

      Shieldbugs walk with a friendly clockwork gait and take to the wing with a solid model-aircraft rattle. Each year common Green Shieldbugs (Palomena prasina) emerge from hibernation in my garden and disport themselves on the garden fence in the early spring sunshine. At that time, March or April, they are a rich brownish purple, seemingly well camouflaged against the brown wooden panels, but soon they will physiologically ferment themselves back to their original emerald. My family are inured to my childish enthusiasm, and politely look on whenever I eagerly rush inside with one cupped in my hands to show them.

      Having said that, in North America especially they are called, less appealingly perhaps, stink bugs. True, they smell a bit, but I’ve always thought their aroma rather pleasant – a kind of oily marzipan scent. After I release that inevitable first show-and-tell Palomena, I can’t resist sniffing my fingers. It’s funny; as a child I couldn’t stand marzipan, and always peeled that layer off my slice of Christmas cake, and yet the shieldbug ‘stink’ is terribly evocative of my childhood. Maybe my psychotherapist could make something of that. On a more positive note, some very brightly coloured tropical species of shieldbug (especially those in the family Scutelleridae) are called jewel bugs, for their mesmerising colours and bright patterns.

      One of the most appealing things about shieldbugs is that they are seen as non-threatening. Even the biggest do not bite, and their short broad feet do not scratch as they walk up the outstretched finger. Sharp spines on the pronotum are not regarded as a danger to life or limb. The cats aren’t scared of them – well, the cats aren’t interested in them, but that’s cats. Importantly they are not seen as significant garden, farm or forest pests. They feed, reassuringly (and usually discreetly), on a variety of wild plants. Mind you, there was that time I got into trouble for blithely dismissing them as ‘harmless’ in a wildlife article in a gardening magazine. A reader wrote an irate letter to the editor complaining of my indifference, and pointing out (with photos, if memory serves) that they were decimating her broad beans. A suitable apology was issued. On the whole, though, they rarely get out of hand in the garden. And mostly they are, indeed, harmless. Mostly.

      In the autumn of 1992 my partner and I took a last-minute adventurous holiday to Costa Rica. A few days in, and we had driven our small hire car to the tiny town of Quepos on the Pacific coast with a view to visiting the Parque Nacional Manuel Antonio the next day in search of tree sloths, agouti, white-faced monkeys and hummingbirds. We saw them all, but being an entomologist my lingering memory of the place (shortly after the sloth sighting) is of a loud rattling buzz as a large insect flew over my head. Without a thought, I jumped up and caught it in my bare hands. It was a huge orange shieldbug, probably one of the Edessa species so diverse there in the Neotropics. I held it gently, as I had done for many large insects, but it did not recognise my friendly behaviour and immediately exuded a copious amount of defensive chemical from its thoracic glands. Ordinarily I would have savoured the delicate rancid marzipan smell I knew from shieldbugs back home in England, but I was surprised to see my thumb completely stained a rich ochre brown. There was none of the accompanying pungent smell that I had come to associate with shieldbugs, but my skin was marked, immovably, for the next five days. It was quite unnerving to think of the chemical power from this insect, particularly as I’d always thought of plant bugs as being at the mild end of the insect danger spectrum. I’d like to be able to say that I was chastened by this episode, more circumspect in my future dealings with large tropical insects, but this was not the case; I continue with my cavalier pick-it-up-and-see-what-happens attitude.

      A few years later, demonstrating insects to small children at a local environmental event at the annual Nunhead Cemetery Open Day, I took great delight in getting my visitors to smell the mating pairs of the small bronze shieldbug Eysarcoris venustissimus that many of them were bringing to the bug-identification stall. Holding them in the palm of my left hand I gave the still-conjoined shieldbug couples a rude poke with my right index finger and held them out to the noses of my audience. Sometimes I was given a screwed-up face of disgust; sometimes an amazed eyes-wide-open recognition from those who knew what a cocktail of almonds and diesel smelled like. But mostly what I got, by the end of the day, was a memory of that Costa Rican encounter when I noticed that following the repeated prod-and-sniff performances my left palm was marked with streaks of vague brown stain that would not be washed away. The home-grown shieldbugs, smaller, less loaded with bodily secretions, nevertheless had the same ability to tan my hide, albeit in a more subdued and gentle fashion. Despite these assaults on my epidermis, I remain unafraid of shieldbugs and continue to handle them as gently as I can. Plenty of the photos in this book are of shieldbugs crawling over my hand, or held gently between finger and thumb. Thankfully my fingers have never been tested so again.

      The North American suspicion of shieldbugs may be down to the possibility that North American pentatomids stink more than European species, though it is more likely to be a general distrust of insects there. We can be rather blasé about potential insect nuisances in the gentle temperate and oceanic climate we enjoy here in north-west Europe, but North Americans have had more than their fair share of significant invasive alien insect pests – from Gypsy Moths (Lymantria dispar) and Tiger Mosquitoes (Aedes albopictus) to Emerald Ash-borer beetles (Agrilus planipennis). One of the latest is the Brown Marmorated Shieldbug (Halyomorpha halys), a pentatomid originally from East Asia that is now a widespread orchard and soft-fruit pest in the USA. It is not just farmers and market-gardeners who notice this large and striking insect; it has a propensity to enter homes to hibernate in clusters, and the appearance of large numbers indoors is alarming and olfactorily offensive. Reports of many thousands invading houses, each buzzing about ‘like an angry overweight wasp’, are commonplace. A prominent article on them in the New Yorker clearly demonstrates that tabloid spluttering has now been replaced by mainstream angst (Schulz 2018). Incidentally, this is the Marmorated Shieldbug, not the Marmite-rated Shieldbug as suggested by the autocorrect on my phone.

      Halyomorpha halys arrived in Europe (in Liechtenstein) in 2004 and now occurs sporadically in Switzerland, Italy, Germany and France. Its arrival into the British Isles has been widely anticipated; indeed, it has been intercepted at British ports in imported goods and passenger luggage, and live specimens have recently turned up in pheromone monitoring traps. If Halyomorpha does become well established here maybe its agricultural afflictions and household invasions will cause people to shy away from the benign aura that presently surrounds shieldbugs.

      Before shieldbugs become blighted with bad press, however, I’d like to re-emphasise their usual harmlessness, their striking and attractive forms, their interesting behaviours and physiology, and their amenable accessibility to the field entomologist. I therefore humbly offer up this book in the furtherance of shieldbug appreciation.

      Acknowledgements

      During 2020 and 2021, it became a slightly facetious joke that everyone should be using the downtime – lockdown, curfew, furlough, hibernation, whatever you care to call it – to write a book. From the privileged armchair of the middle-class, middle-aged, white man with a university education I consider myself very fortunate that this is exactly what I was able to do. Although initiated in 2018 through reading and research, the writing began in earnest in May 2020, and has followed the on/off time I have had available. There have been stoppages and slow-downs, but by early December of that year I was back on track again. As ever, my partner Catrina Ure helped me find a safe place – physically, mentally, spiritually, and emotionally – to get on with the task. She can only guess at the importance I place on my entomology and what sociologist Harriet Martineau so eloquently described as the ‘need of utterance’ to write down and get out what is on my mind. It has been both cathartic and calming to sit writing about shieldbugs, amidst the raging tempests of shambolic domestic and international politics and a tragic medical calamity unfolding on a world scale. I hope that my prose has not been too distorted to either rant or whimsy, and that I have been able to steer a steady and neutral course through what, for many, has been a time of confusion, chaos, and personal heartache.

      The text is only half this book, and the illustrations also needed considerable work. Many shieldbugs are common and striking and relatively easily identified from photographs, so they feature heavily in the several stock photo agencies through which I have been able to trawl for the usual portraits and even some behavioural shots. However, I have not always agreed with the identifications offered. I’ve done my best, but if there are errors in the names then I accept full responsibility. Thankfully I have also been able to call on several specialist entomologist photographers to fill the many gaps and provide spectacular photographs of some of the more difficult and obscure species, and also their unusual behaviours – Tristan Bantock, Yvonne Couch, Maria Justamond, Penny Metal and Simon Robson, you are all stars. My daughter Verity Ure-Jones skilfully produced all the line illustrations for the identification key, coping admirably with my sometimes garbled instructions and ill-formed ideas.

      Roger Hawkins kindly read through some of the text, particularly the long list of British shieldbug species in Chapter 8; he made many helpful suggestions and contributed some interesting personal observations. Hugh Brazier then patiently edited my original text using subtle variations on a theme of ‘did you really mean this?’ when he had spotted a complete boo-boo. And thanks to designers Namrita and David Price-Goodfellow for arranging the sometimes complex page layouts that resulted when all the disparate pictures, verbose fact boxes and my obsessive footnotes needed to be slotted in together sensibly.

      Many other people have helped with contributions, suggestions, photographs and more: Sybil Baldwin, Max Barclay, Ian Beavis, Nathaniel Blair, Steve Covey, Alan Watson Featherstone, Graham Fisher, Jim Flanagan, Will George, Ian Harding, Joanne Hatton, Graham Hopkins, Finley Hutchinson, Richard Lewington, Jerzy Lis, Graeme Lyons, Emma Nicholls, Matthew Oates, Simon Oliver, Colin Purrington, Stuart Reed, Jony Russell, Matt Shardlow, Arabella Sock, Stras Strekopytov, Lillian Ure-Jones, Leon van der Noll, Sue Vincent, Simon Warry, David Williams, Gary Williamson, Ron Woollacott, Aidan Wren, Antony Wren, Elaine Wright. To all of you go my grateful thanks.

      CHAPTER 1

      What is a Shieldbug?

      First – what is a bug?

      ‘Bug’ is a rather vague term. Today it can mean a disease-causing germ from virus to bacterium, any invertebrate from tardigrade to giant squid, a computer coding problem, a mechanical glitch or an electronic failure. In natural history, though, it has mainly become a flexible and broad term for creepy-crawlies in general. I became Bugman Jones when I started working with local schoolchildren who knew that I studied the small insects, spiders, woodlice, centipedes and snails that they just lumped all together as ‘bugs’. This trend is sometimes lamented as unfortunate, and for some people ‘minibeast’ is an even more unspeakable heresy. Yet these are inclusive, friendly and non-threatening terms in a science often mired in jargon, and they all have their place when used properly. Meanwhile, for the more academic entomologist, a ‘true bug’ is a member of the insect order Hemiptera. This is a large and diverse collection of creatures including aphids, whitefly, scales, jumping plant lice, leafhoppers, froghoppers, cicadas, water boatmen, back-swimmers, skaters, capsids and shieldbugs.

      This was not always the case. Although ‘bug’ had sometimes been used disparagingly to mean a small nondescript insect before about 1622 (according to the Oxford English Dictionary),¹ it had previously mostly meant something completely different – a bugbear, boggard, bogey, hobgoblin or some other general night fear – and variants include bugabo and buglarde. The etymological origins are unclear, but it may have some connection with the Welsh bwg or bwgan – ghost (again, the OED is my source here). This ill-defined night dread is actually the meaning used in the oft-misquoted (by entomologists) Psalm 91, verse 5, from the Matthew Bible (1537), ‘So that thou shalt not nede to be afrayed of eny bugges by nyght.’ For the 1611 King James edition, though, this was retranslated: ‘Thou shalt not be afraid for the terror by night.’ In other words, creepy, but not crawly.

      FIG 1. The glory of shieldbugs, shown in examples of some of the groups covered in this book. (a) Brassica Bug (Eurydema oleracea, Pentatomidae). (b) Box Bug (Gonocerus acuteangulatus, Coreidae). (c) Firebug (Pyrrhocoris apterus, Pyrrhocoridae). (d) Cinnamon Bug (Corizus hyoscyami, Rhopalidae). (e) Common Tortoise Bug (Eurygaster testudinaria, Scutelleridae). (f) Bordered Shieldbug (Legnotus limbosus, Cydnidae). (Photos by Penny Metal)

      However, in 1730 Londoner John Southall firmly fixed the name to the blood-sucking Bedbug (Cimex lectularius) when he wrote his Treatise of Buggs, mainly to promote his nonpareil liquor (2 shillings a bottle) to get rid of the damn things (Fig. 2). He was also appointed ‘buggman’ to Sir Thomas Coke, Lord Lovell, later first Earl of Leicester, to control the vermin at a salary of a guinea a year (Strekopytov 2021). Whether he was a genuine naturalist or just a copywriter and pamphleteer is not clear; his booklet has some scientific information, but also much myth and misunderstanding, and Weiss (1931) concluded that it was just a clever advertisement for pest control. Up until that point Cimex bedbugs had mostly been called wall-lice, for their habit of secreting themselves away in cracks in the walls, ready to emerge to suck the blood of their unfortunate victims during the night. The modern German word for almost any hemipteran true bug, Wanze, is a corruption of Wandlaus, meaning wall-louse, while the French punaise is a word also used for drawing pin or thumb tack – an item you might use to pin something to the wall. Like ‘bug’, ‘louse’ was another one of those vague old words used to mean almost any small crawling critter from head louse to woodlouse. Cimex lectularius is now classed as a genuine ‘true bug’ in the order Hemiptera, family Cimicidae.

      FIG 2. Frontispiece from Southall’s Treatise of Buggs (1730), depicting the well-known Bedbug (Cimex lectularius), although he was of the opinion that European and American bedbugs were different species.

      Uptake of the word ‘bug’ was slow to start; nevertheless, it soon crawled decisively out onto the entomological world stage. Cimex the wall-louse (also wall-lowse, or punie) is included in one of the most important early English books on insects, physician Thomas Mouffet’s Theater of Insects (1658), along with three obvious shieldbugs, but he calls these ‘wood wall-lice of the sheath-winged kinde’, not bugs – a word he uses only once in his entire 400-page tract as an alternative name (along with ‘klock’) for beetle. Later, John Ray (1710) uses the Latin term cimices sylvestres (wood cimexes, i.e. wood wall-lice) for all of his shieldbugs too. After Southall’s 1730 booklet, though, ‘bug’ increasingly meant not just Cimex, the bedbug/wall-louse, but gradually, by association, other hemipteran relatives. Just over 30 years later ‘bug’ is already the default word used by Brookes (1763) for all manner of hemipterans, at least 40 species, from the original Cimex (‘only a common bug’) to Notonecta the ‘water bug or boat-flie’ and several probable shieldbugs, although he also includes May-bug (and May Bug) for the cockchafer.

      what’s in a name?

      So – shield bug, shield-bug or shieldbug? As is clear from the title of this book, I have already decided which one I will be using. But this is not without some trepidation, and I realise that I have exposed myself to potential argument. My reasoning for going down this route is as follows.

      There was once supposed to be a tradition amongst naturalists that composite insect names were two words if they were scientifically logical, but one word if they were not. Thus: hover flies, house flies, bot flies and dung flies are all actual flies so two words each, but butterflies, dragonflies, mayflies and greenflies are patently not. By this same reckoning we should have lace bugs, spittle bugs, leaf bugs and shield bugs, which are all bugs, but Maybugs, which are beetles. But, as is all too common with English common names, they wash with the times and with the fashions of both popular and scientific publishing. Supposedly authoritative dictionaries and encyclopaedias are just as fickle, and the several I consulted from my extensive etymological library, published between 1895 and 1987, offer between them all three variants of shield bug, shield-bug and shieldbug, often as alternatives, and it is clear that no clear consensus has ever really existed.

      Fly/flye/flie and bug/bugg/bugge are ancient words anyway, coined long before any real understanding of insect phylogeny or taxonomy was either necessary or available. There have long been transgressors and intermediates to the ‘rule’. Maybugs, sometimes May-bugs but frequently May bugs, are beetles (Melolontha species) that just happen to fly about in May. They are also called cockchafers or cock-chafers (no hen-chafers known though) and are obviously chafers, from the Old English word ceafor, meaning beetle. However, the Black Beetle isn’t a beetle, but a cockroach, Blatta orientalis; and cockroaches are nothing to do with cockchafers. Nor are they types of roach, which is a very modern contraction. And there are no hen-roaches. Fire-flies (family Lampyridae) and the Spanish Fly (Lytta vesicatoria) are not flies – they are beetles. Neither woodworms nor glow-worms are worms. Bumblebees and honeybees are certainly bees, and though they are sometimes two words, occasionally hyphenated, they too are frequently and inconsistently one. It turns out there isn’t really a rule at all.

      To conflate this muddle, there has been a general (by which I also mean rather haphazard) pattern in the entomological literature of the last 100 years or so, to first hyphenate some of these double word names, then unify them. My old A-level chemistry teacher Mr Warren maintained that this reflected the assimilation of large numbers of German scientists (particularly chemists) into British and American research programmes after the Second World War, bringing with them a long-standing German-language tradition of combining short words wholesale to make new longer ones. As hover flies have become more popular subjects for study, they first became hover-fly (Coe 1953), and now hoverfly (Stubbs & Falk 1983, 2002). Likewise we also regularly have housefly, and botfly, but strangely dung fly remains two words – for now. Bee-fly also has had to remain in limbo, hyphenated (Stubbs et al. 2001), since beefly looks like a brand of bovine stock cube.

      Consequently I have partly followed this supposedly modern trend from the shield bugs of the nineteenth century, through a vague transition into shield-bugs in the mid-twentieth, to give us the shieldbugs we have today. This form was already well established, and although the more technical literature still often hangs on to two separate words, every popular British treatment I could find during the last 25 years now gives shieldbugs on a unified front. There are also leatherbugs, lacebugs, flatbugs and bedbugs, although contrariwise there remain stink bugs, ground bugs, squash bugs, tortoise bugs, Dock Bug and Parent Bug. No, this is not completely satisfactory, but it’s what has evolved through the piecemeal jumble that is the English language, and it does just go to show how unhelpful, confusing and illogical English names can be.

      In truth any logic in these common English names has long been broken. This is why scientific names were invented. Throughout this book the scientific names are the ones that hold authority, and any English names, if they are available, are given only subsidiary status. Nevertheless, certain groups of insects are currently enjoying increased popularity, particularly the relatively large and photogenic bug groups covered here. And many have acquired or have been newly christened with English names. The current style of the New Naturalist Library is to capitalise English common names. This works well enough for the well-established name pedigrees that support plants, fungi, birds, mammals, butterflies and moths, but can be counterproductive for the more obscure groups of insects. When I wrote the Beetles volume in 2018 I argued that not enough of the over 4,000 British beetle species had good enough common names, and the names that did exist (violet ground beetle, stag beetle, black oil beetle, red soldier beetle) were still too vague and nebulous to be conferred with capital letter importance. The editors of the series agreed, so in that case scientific clarity was upheld at the lesser expense of literary consistency.

      Shieldbugs are on the cusp. They are now part of a burgeoning array of stepping-stone insect groups like dragonflies, grasshoppers, bees and hoverflies – slightly more difficult than butterflies and moths, but not as tricky as fleas or flea beetles. Here, English names can be a genuine help in understanding and do not get in the way of identification. Some are long-standing, some are inventive new coinings. In this case, with the scientific names to lead the way, I am happy to wash along with the publishing fashions of the day, and although I normally try to avoid too many extraneous capitals in my text, I do here meekly, if slightly grumpily, follow the prescribed style of the series.

      Today the ‘true bugs’, the Hemiptera, are thought to comprise in excess of 100,000 species worldwide (that’s an impressive 10% of all known insect species), and about 1,850 Hemiptera species occur in Britain and Ireland. Like most insects, many of these are tiny and never acquired useful common names, though very general terms like aphid, plant-louse, scale insect or leafhopper cover vast swathes of them. Shieldbugs represent a small group of some of the largest, brightest, most obvious and best-known amongst this broad and varied classification.

      What makes it a shieldbug?

      It’s not simply a case of claiming they are bugs which are shaped a bit like shields. What something looks like at first glance is not often a sound judgement on which to base any classification. There are now many hundreds of years of studious observation by which the organisms of this planet can be counted, accounted and arranged. In order to define shieldbugs properly, the other bugs in the insect order Hemiptera have to be identified and cut loose.

      The Hemiptera in general are characterised by having four wings, sucking tubular mouthparts, prominent compound eyes, often also ocelli (simple eyes), and long to very short antennae, and they go through hemimetabolous development from egg to adult.² This ‘incomplete metamorphosis’ means that the tiny nymphs which hatch from the eggs are merely miniature wingless versions of the adults, already showing the correct body segmentation, full leg articulation, properly developed eyes and antennae. As they grow they moult their skins several times, and at each point the wing buds increase until the final winged adult form emerges. This is a very different process from the holometabolous development (‘complete metamorphosis’) of flies, beetles, bees and butterflies, which pass through a feeding and growing maggot, grub or caterpillar stage before the entire body plan is reshuffled (liquidised might be more accurate) inside the chrysalis to produce a winged adult insect wholly different in form and structure from the larva. The importance of the sucking mouthparts of the Hemiptera has long been appreciated, and indeed they are what characterise all true bugs. The order has also variously been designated as Rhyngota, Rhynchota or Rhyngotorum over the years, all derived from the Greek ρινο rhino, meaning snout or nose. The fine structure of the rostrum, as the beak-like tube mouthparts are called, is unique, and defines the Hemiptera. More details are given in Chapter 2.

      Today the order Hemiptera (Greek ήμι hemi = half, and plural of πτερόν pteron = wing) is divided into 2–5 suborders, depending upon whose scheme you follow (Fig. 3). The suborder Homoptera (Greek όμο homo = same, and plural of πτερόν pteron = wing) have front and back wings very similar in shape, size and texture; at rest they are folded tent-like along the abdomen, and they are most often transparent and clear, but a few are coloured or patterned. This is a large group including leafhoppers, cicadas, aphids and psyllids, though it is often considered defunct because its evolutionary origins are unclear. Instead this slightly artificial grouping is usually subdivided along evolutionary lines into the suborders Coleorrhyncha (moss bugs, only 30 world species in a single family, Peloridiidae), Auchenorrhyncha Cicadamorpha (cicadas, leafhoppers, spittle bugs etc.), Auchenorrhyncha Fulgoromorpha (planthoppers) and Sternorrhyncha (aphids, scales, whitefly etc.). Little else needs to be said about these insects here except that they all share that hemimetabolous upbringing and tubular piercing sucking mouthparts.

      The shieldbugs belong to the remaining suborder, Heteroptera (Greek έτερο hetero = different, and plural of πτερόν pteron = wing), which has very different front and back wings, usually resting more or less flat over the abdomen. The front wings are normally long and slim, partially coloured and sclerotised (hardened) in the basal section, or at least coriaceous (leathery) and opaque, whilst the hind wings are broadly triangular, membranous, clearly transparent and partially folded concertina- or origami-style. At rest the more delicate hind wings are furled away and covered by the tougher protective layer of the partly overlapping front wings. Together with the large toughened upper front plate of the thorax (the pronotum), this gives the ‘het’ bugs (as they are sometimes known) a more solid, robust form than many of the more flimsy Homoptera. Their under-bodies also tend to be tougher and harder.

      This hard toughness comes from the chemical structure of the insect body. Insect cuticle is mostly made up of chitin, a long-chain polymer of N-acetylglucosamine; it is a polysaccharide analogous to the cellulose found in plants. Chitin is laid down in a series of parallel fibres to form thin flat sheets; a series of sheets, layered one on top of the other, with the grain of the fibres of each sheet at slightly different angles, gives a superior strength and rigidity much as do the laminated layers of plywood. The chitin fibres and layers are impregnated, interlinked and reinforced with proteins, and it is this combination of chitin sheets and protein glue that gives insects their tough yet flexible bodies. All insects use chitin, but in some hard and tough insects (het bugs are a very good example) the chitin layers are laid down so thickly that they achieve real armour-plating strength.

      FIG 3. Examples of the Hemiptera. (a) Moss bug (Xenophyes rhachilophus, suborder Coleorrhyncha). (b) Rhododendron Leafhopper (Graphocephala fennahi, suborder Auchenorrhyncha Cicadamorpha). (c) Spotted Lanternfly (Lycorma delicatula, suborder Auchenorrhyncha Fulgoromorpha). (d) Cottony Scale (Icerya purchasi, suborder Sternorrhyncha). (e) Back-swimmer (Notonecta species, Hemiptera Heteroptera Nepomorpha). (f) Green Shieldbug (Palomena prasina, Hemiptera Heteroptera Pentatomomorpha), perhaps the most shieldish of shieldbugs.

      With their hardened wing structure, stout head and pronotum, and toughened underbellies, the Heteroptera slightly resemble beetles (Coleoptera), another heavily sclerotised group of insects. Beetles are one of the most diverse and species-rich groups of insects on the planet, a status they are thought to have achieved by the development of their hard shell-like bodies and tough protective wing cases (called elytra, singular elytron), allowing them the joint advantages of flying and of also pressing hard into tight spaces to hide without damaging their fragile flight wings. The convergent evolution that also toughened the bodies and hardened the front wings of the Heteroptera has conveyed a similar protective advantage to these bugs, and this large grouping has many species that regularly push down into flowers, the general foliage, plant root thatch or leaf litter, burrow into loose soil, live under rocks and stones, push between the scales of pine cones, under rotten bark or into fungi, occur as parasites on birds and mammals, live in or on water, including inside rocks on the seashore, or ride on the open ocean. Arguably the Heteroptera occupy a more diverse array of habitat types than any other insect group. No Hemiptera burrow through wood, or are internal parasites of other animals, but this is more a reflection of their non-chewing mouthparts, as discussed later.

      The hemi ‘half’ in the name Hemiptera refers to the fact that most large bugs have the basal half of the front wings hardened and coloured, whilst the terminal half (called rather unimaginatively the ‘membrane’) remains transparent and membranous. This reiterates that comparison with beetle elytra; thus the hemipteran forewing is usually referred to as a hemelytron (occasionally hemielytron) – half a wing case.

      A key body part that helps define a shieldbug is the large flat triangular plate, behind the pronotum, right in the middle of the back. This is the scutellum – quite literally Latin for ‘little shield’, being the diminutive form of scutum, the large, rectangular, slightly curved shield carried by Roman legionary soldiers. All hemipteran bugs have a scutellum, but it is particularly large and obvious in many of the shieldbugs, and this shield-plate is often given as the reason behind the common name, rather than these insects simply being generally shield-shaped. John Ray (1710) used the term interscapulum (Latin, ‘between the shoulders’), but this word has almost completely fallen out of use except in medical textbooks referring to the upper back, or some rather technical bird descriptions to denote the area between the wings just behind the head.

      Traditionally, true shieldbugs comprised the family Pentatomidae, so named for the five (penta-) rather than four segments that made up the antennae of most other bugs. But as I bemoaned above, English names for insects can be rather vague and amorphous, and this narrow classification rather unfairly excluded many groups with four-segmented antennae which had very obviously similar body form and which were also regularly called shieldbugs. It is at this point that we have to admit that ‘shieldbug’ is a somewhat arbitrary conglomeration of groups, of which most (but not all) are shield-shaped, most (but not all) have a prominent scutellum, and some (but not all) have five-segmented antennae.

      FIG 4. Keeping good company, Pentatoma rufipes (number 5) represents the Hemiptera Heteroptera, along with various ‘other orders’ (i.e. not the big four groups Coleoptera, Diptera, Hymenoptera, Lepidoptera) in this illustration from Kirby & Spence’s Introduction to Entomology (1815–26). Also shown are: (1) Xenos peckii (Strepsiptera), (2) Gryllotalpa gryllotalpa (Orthoptera), (3) Blatella germanica (Blattodea) and (4) Ledra aurita (Hemiptera Homoptera).

      Conveniently, modern scientific treatments of the suborder Heteroptera recognise an intermediate subdivision – the infraorder Pentatomomorpha, one of seven infraorders into which all het bugs are grouped. This will form the outer limits of what might reasonably be considered shield-type bugs. It is defined by numerous very abstruse technical characters including antennae with 3–5 segments, labium mostly four-segmented, large scutellum, costal fracture of forewings never present, pretarsus with claws equally developed, and pulvilli well developed, divided into basipulvillus and lamellate distipulvillus, and absence of a true egg operculum. This doesn’t really help with fieldwork, where these awkward characters are often difficult to understand or impossible to see. And covering all of these groups would make this book unwieldy, and it would be overwhelmed by the very numerous much smaller and less shield-shaped members. Instead, I have chosen a personal pragmatic selection. For the basis of this book, I am including all the British families of the Pentatomomorpha, except the superfamilies Aradoidea and Lygaeoidea. This was, I admit, a bit of a wrench, but there is barely enough time and space so something had to give. But I can’t quite bring myself to get rid of them entirely and have sneaked in, under the editorial radar, a paragraph on each (see here). This finally gives a workable 79 British ‘shieldbug’ species across 11 families, and although some are not particularly shieldish, there is a genuine biological cohesiveness in that they share a close evolutionary descent, have many similar body structures and forms, are likely to be found by the field naturalist on or under plants in the general landscape, without recourse to specialist searching techniques; and they are mostly large enough to appreciate and identify using a hand lens, often with just the naked eye (Fig. 5).

      FIG 5. Two extremes of shieldbug form: (left) typical shield-shaped Palomena prasina (Pentatomidae) and (right) lacy and spined southern European Phyllomorpha lacerata (Coreidae).

      These, then, are shieldbugs, or at least my broad interpretation, and this is a permutation on the same group of families that have been covered in many other recent British and European publications including those by Hawkins (2003), Evans & Edmondson (2005), Boardman (2014), and Nielsen & Skipper (2015). I hope specialist entomologists will not consider my choices too jarring, or less specialist naturalists find it too confusing. I can see a logic here, and I hope I can justify it to you, dear reader. A summary of world and British shieldbug families, genera and species is given in Table 1, at the end of this chapter.

      Some non-British shieldbug groups

      As in most insect groups, shieldbugs are seriously under-represented in Britain when compared to the rest of the world. They are especially diverse in the tropics and subtropics. A limited number of genera and families occur here, but a much broader diversity occurs elsewhere. Before going into more detail on our native species, it is worth a brief diversion across the globe to consider some non-British shieldbug groups.

      excluded groups – british pentatomomorpha deliberately left out

      flat, and not really shield-shaped – aradoidea

      Commonly called flatbugs, or bark bugs, there are only seven British species anyway, in the family Aradidae; they are all small (less than 7 mm) and secretive, strongly flattened bugs that only occur under fungoid tree bark (Fig. 6). The commonest are Aneuris laevis and Aradus depressus. Worldwide, though, this is a huge group (2,050 species at least) and many are large (up to 20 mm) and attractive – not necessarily brightly coloured, but interestingly granulate or sculptured, and variously shaped oval, rectangular or triangular. Because of their secretive life under the bark, many tropical species no longer have winged forms. The mouthparts are much longer than those of other bugs, with the piercing stylets coiled around inside the head. These are used, under the bark, to penetrate deep into the hyphae of the fungal strands, and this mode of feeding may be ancestral for all of the Pentatomomorpha. Some exotic species live solely in termite nests, as do the eight known species of the related Termitaphididae. All British aradids are adequately keyed out by Southwood & Leston (1959).

      FIG 6. Flatbugs occur only under fungoid bark. Aneuris laevis, adults (left) and pale nymphs (top left); and Aradus depressus mating pairs (right).

      fast, and not really shield-shaped – lygaeoidea

      Usually called ground bugs for their habit of running about on the ground, or seed bugs because many feed on seeds, lygaeids are attractive under the microscope, but most are small or very small, and rather ‘other’ compared to shieldbugs (Fig. 7). And with over 100 British species, they would swamp this book if they were included. Most British species belong to the family Lygaeidae (although this has been subdivided by some authors), and typical species include the common Heterogaster urticae, which often occurs in large aggregations on Stinging Nettles (Urtica dioica), Kleidocerys resedae, a small but very smelly bug infesting birch trees, and Rhyparochromus vulgaris, a handsome well-marked and long-legged denizen of rough grassland where it rushes about on the soil at top speed if disturbed. The related Berytinidae are slim and very long-legged and despite their fragile forms some are thought to be at least partially predatory. Most British species can be identified using Southwood & Leston (1959), but the most up-to-date European treatment of the Lygaeidae is provided in the three volumes by Péricart (1998), PDFs of which are now freely available to download from the faunedefrance.org website.

      FIG 7. Rhyparochromus vulgaris (left) is a typical ground bug, with its mottled earthy palette and large shield-like scutellum. Lygaeus equestris (right) could easily be mistaken for Pyrrhocoris or Corizus; it is widespread in mainland Europe.

      Despite the fact that the Australian family Henicocoridae has just a single species and the Australian/South American Idiostolidae just five species, these families are interesting because they have several characters intermediate between true shieldbugs and ground bugs in the superfamily Lygaeoidea. These are mainly to do with the distribution of trichobothria, long sensory bristles that detect air movements or vibrations, on the underside of the abdomen. Tiny and insignificant as these hairs might seem, they are remarkably persistent across shieldbug groups, even in fossils, and they confirm the close relatedness of the various families. This deep southern-hemisphere distribution is typical of truly ancient lineages (like marsupials, kauri conifer trees, southern beech Nothofagus forests), and were this a 1950s science fiction book I might be describing them as missing links in the classification. They superficially resemble and were originally described as Lygaeidae, but are now given their own separate family statuses. Little is known of their life histories, but they inhabit moss and leaf litter in humid temperate forests and are thought to be plant-feeding.

      There are only nine species in a single genus in the family Canopidae – all from Central and South America. These strange small hemispherical shining black, greenish or purplish bugs are nearly all scutellum, and both front and back wings have particular folding mechanisms to furl them away. They feed on the fruiting bodies of bracket fungi in tropical forests, and fungal spores have been confirmed in their guts. They are often found in conjunction with similarly globose shining metallic fungus beetles in the family Erotylidae. No British Hemiptera are known to feed on fungal fruiting bodies.

      FIG 8. Unnamed dinidorid shieldbug, its broad stout form showing its clear relatedness to the Pentatomidae.

      The Dinidoridae are large, heavily built insects mainly from Asia and Africa (Fig. 8). They are obviously shieldbugs, but separated into their own family on minor details of head structure and sensory hairs under the abdomen. Some have two tarsal segments like the Acanthosomatidae, some three like the Pentatomidae.

      Two Australian species make up the family Lestoniidae. Though hemispherical and subglobular, with a huge scutellum and massive rounded pronotum, these lentil-shaped insects share many evolutionary features with the Acanthosomatidae, including two-segmented tarsi together with the presence and position of various abdominal glands. They feed on the growing tips of Australian cypress trees in the genus Callitris.

      The Phloeidae are large broadly flattened bugs, up to 30 mm long, with wide leafy flanged body edges adapted to clamping down to hide on South American or Bornean tree trunks, where they vanish in the rough bark and lichen layers (Fig. 9). They are renowned for their maternal habit of protecting the young nymphs by carrying them about tucked in on the underside of the body, concealed by the skirt-like flange of their outline. Their eyes are separated into one pair of lobes on the top edge of the head looking upwards, and another pair just under the flange looking down at the tree bark. They also have the ability to squirt a jet of liquid a considerable distance from their bodies, though this is probably excretory rather than defensive.

      The Tessaratomidae are a large group of large (to 45 mm long), mostly tropical shieldbugs which all have remarkably small heads. The nymphs

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