Borror & Delong 2005. Study of Insects PDF

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Abbreviations Used in the Figures The following list includes all the abbreviations used in the figures in Chap- ters 3 through 35. The abbreviations used on wing drawings for the veins and eells (using the Comstoek-Needham terminology) are not listed in the figure legends, but are included in the ...

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Abbreviations Used in the Figures The following list includes all the abbreviations used in the figures in Chapters 3 through 35. The abbreviations used on wing drawings for the veins and eells (using the Comstoek-Needham terminology) are not listed in the figure legends, but are included in the following listo Subseript numerals are used to designate branehes of the longitudinal veins. Sueh numerals are often used to designa te the particular thorade segment on which a strueture is loeated O, designating the prothorax; 2, the mesothorax; and 3, the metathorax). Subseript numerals are oeeasionally used to designate the particular abdominal segment on which a sclerite is loeated.

a, anal vein A, anal vein, anal cell ab, abdomen, opisthosoma ac, accessory, lanceolate, or subanal vein Ac, anal crossing (A branching posteriorly from Cu, often called the cubito-anal cross vein) acc, accessory cell acg, accessory gland acg, accessory gland ael, antennal elub aelp, anteclypeus acr, acrostichal bristles aes, acrosternite act, acrotergite acv, anterior cross vein adf, adfrontal area aed, aedeagus af, antennal fossa agr, scrobe, groove in beak for reception of antenna al, anallobe alp, anal loop alu, alula am, axillary muscle an, antenodal cross vein AN, alinotUm anc, anal cleft anes, antecostal suture anp, anal plate anr, anal ring ans, anus ant, antenna ante, antecosta antl, antennule ao, dorsal aorta

aos, anterior oblique sulcus on mesepisternum ap, appendix Ap, apical cell apc, apical cross vein apd, apodeme apo, apophysis ar, arista are, arculus are, areolet aro, arolium an, point of aniculation as, antennal sulcus, anterior spiracle ase, antennal sclerite ask, antennal socket asp, apical spur aspr, anterior spiracle at, alimentary tract ata, anterior tentorial arm atb, anal tube atp, anterior tentorial pit au, auricle av, auxiliary vein aw, anterior wart awp, anterior no tal wing process ax, axilla AX, axillary cell axc, axillary cell axcr, axillary cord axs, axillary sclerite axv, axillary vein B, basal cell ba, basalare BA, basal anal cell, basal areole

be, bursa copulatrix bcv, bridge cross vein bg, book gills bk, beak, proboscis, rostrum, or snout bl, blastoderm bln, banksian line bm, basal medial cell, basement membrane bm-cu, basal mediocubital cross vein bms, basalar muscle bp, brood pouch br, brain or basal radial cell brv, bridge vein bt, breathing tube buc, buccula or bucca bv, basal vein bvn, brace vein e, costal vein C, costal vein, costal cell ca, corpus allatum cal, calypter or squama cb, corbicula cbr, costal break cc, crystalline cone cd, cardo cee, circumesophageal connective cen, cenchri cg, cerebral ganglion ch, chelicera cho,chorion chp, cheliped el, clypeus, clavus cla, clasper clc, movable spines or calcaria

elm, calamistrum elp, elypeus elpl, elypellus cls, claval suture elt, claw tuft, clypeal tubercle elv, claval vein cm, gastric caeca or caecum en, colon ena,cornea enge, corneagenous cells cnu, eleavage nuclei colm, collum, tergite of the first body segment com, commissural trachea como, tritocerebral commissure cor, corim covd, common oviduct cp, crop cph, prosoma or cephalothorax cpl, cortical cytoplasm cr, cercus, lateral caudal filament, superior appendage crb, cribellum cre, cremaster crl, crystalline lens cm, cornicle cro, crochets crp, carapace es, coronal suture esp, cusp of mandible, caudal spiracle cu, cubital vein Cu, cubital vein

cual>anterior cubital cell cuf,cubitalfork (fork of CuA) cun, cuneus cup, posterior cubital cell Cup,posterior cubital vein cut, cuticle cva, clava cvs, cervical sclerite CVX, cervix ex, coxa exc, coxal cavity

exg, groove in coxa exp, eoxopodite of abdominal appendages d, discoidal or intercostal vein D, diseal cell, or discoidal eell de, dorsocentral bristles dcv,discal cross vein dlm, dorsallongitudinal muscle dm, domelike layer of euticle over nerve ending, or discal medial cell dm-cu, discal medio-cubital cross vein do, dorsal ostiole dp, distal process of sensory cell DS],disjugal furrow dta, dorsal tentorial arm dtra, dorsal trachea e, eye, compound eye ee, eye eap eet, eetoderm ef, epigastrie furrow eg, egg ejd, ejaeulatory duct el, elytron emb, embolium emp, empodium en, endophallus end, endocuticle, endoderm endr, endodermal rudiments enl, endite lobe enp, endopodite ep, epidermis eper, epicranium epg, epigynum eph, epipharynx epi, epicuticle epm, epimeron epp, epipleurite epr, epistomal ridge eps, epistemum ept, epiproet, median caudal filament, or inferior appendage es, epistomal su\cus eso, esophagus ex, exuvium

exl, exite lobe exm, extensor muscle exo, exocuticle exp, exopodite f, frenulum fa, face lb, frontal bristles fc, food channel fch, lilter chamber fen, frontal ganglion connectíve re, facial fovea fg, frontal ganglion lib, libula fl, flagellum flb, flabellum flm, flexor muscle fm, femur fmb, femoral bristles fn, fang of chelicera fob, fronto-orbital bristles fon, fontanelle for, foramen magnum fr, frons frl, frontallunule fs, frontal suture fu, sternal apophysis, furca fun, funiculus or funicle fv, frontal vitta g, galea gap, gonapophysis ge, genal comb gcl, germ cell gex, gonocoxa ge, gena gen, male copulatory apparatus gf, genital forceps gh, gland hair gi, gills gl, glossa gle, gland cell gld, duct of gland cell gis, gland spines glt, gland tubercle gn, ganglion of ventral nerve cord gna, gnathochilarium gon, gonangulum gpl, gonoplacs gr, gill remnants gs, guiar suture gst, gonostylus gt, genal tooth gu, gula gvp, genovertical or orbital plate h, humeral eross vein hal, haltere hb, humeral bristles hbr, hypostomal bridge

hc, humeral callus hcl, hypostigmatic or truss cell hd, head hg, anterior portíon of the hindgut ho,hom hp, humeral plate hr, heart hst, haustellum hv, humeral or recurrent vein hyb, hypopleural bristles hyp, hypopharynx, intermediate stylet hypl, hypopleuron iab, intra-alar bristles iap, interior appendage (paraproct) iar, interantennal ridge ias, interantennal suture iep, infraepistemum il, ileum ism, intersegmental membrane it, intercalated triangle ivb, inner vertical bristles j, jugum jl, jugallobe I,leg L, length, lanceolate Iba, labial articulation Ibl, labellum Ibm, labium, ventral stylet Ibn, labial nerve Ibr, labrum, rostrum lbrn, labral nerve le, lacinia let, layer of the cutícle 19, ligula, median lobe 11,lamina lingualis lo, lorum lp, labial palp Lp, lateral plate 15, labial suture 1st, lateral setae Itra, main longitudinal tracheal trunk m, medial cross vein, mouth, or recurrent nerve M, medial vein, medial eell ma, mandibular articulation MA, anterior media MC, marginal eell mcf, median eaudallilament mep, mieropyle md, mandible MD, median eell mdn, mandibular nerve mdp, median plate (of wing), middle plate (of embryo)

mdu, microduct mdv, median vein mem, membrane met, metasomatic segment mf, medial fork (fork of MP2) mg, midgut or mesenteron mh, movable hook or palp mi, median lobe mm, marginal macroduet mn, mentum mo, mouth mp, mouthparts Mp, posterior media mpb, mesopleural bristles mpo, marginal 8-shaped pore ms, mesoderm msd, mesoderm msl, mesostemallobe mspl, medial supplement mst, mental seta mt, Malpighian tubule mts, metatarsus or lirst tarsal segment mu, mucro mv, marginal or radial vein mx, maxilla, dorsal stylet mxa, maxillary articulatíon mxl, maxillary lobe mxn, maxillary nerve mxp, maxillary palp mxt, maxillary tentacle n, notum nb, notopleural bristles ne, ventral nerve cord nod,nodus nll, pronotallobe npl, notopleuron npls, notopleural suture nt, notaulus nu, nucleus nv, neuron o,opening ob, oeellar bristles obv, oblique vein oe, ocellus oep, oeciput oepd, ocellar pedicel oes, ocular suleus oeg, oecipital ganglion og, optic ganglion op,operculum opl, optie lobe opt, ocular point orp, orbital plate os, oecipital suleus osm, osmeterium (seent gland) ot, oeellar triangle Continued on back endpaper

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Borror and DeLong's Introduction to the Study of Insects

Seventh Edition

UNIVERSIDAD DECALDAS

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Borrorand DeLong's Introduction to the Studyof Insects

Seventh Edition

CharlesA. Triplehorn The Ohio State University

Norman F.Johnson The Ohio State University

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Preface

to the Study oIInsects: this is the seventh edition of a textbook AnthatIntroduction has been widely used in entomology classes in North America over more than 50 years. Its value has be en demonstrated by the fact that it retains a prominent place on the bookshelves of professional entomologists, long after their first exposure to insects in class. Because the book has been widely known by the names of its first two authors, we are adding their names to the title. The contributions of these two men, in both style and substance, will still be immediately apparent to knowledgeable readers even though the formal authorship has now passed on to subsequent generations. We have prepared this new edition in recognition of the important role the text has played in the education of biologists of all specializations and in the hope that it can continue to play that role in the future. NF] clearly recalls the nights and weekends spent at Cranberry Lake Biological Station in the Adirondack Mountains of New York, poring over this book in theexcitement of new discoveries and with an ever-growing appreciation for the diversity of insects. CAT, too, was greatly influenced by Borror and DeLong, but in a more direct way. He took undergraduate courses from both of them and quickly abandoned his original goals in herpetology when exposed to "the wonderful world of insects" in a beginning entomology course taught by Borror. In this new edition we have concentrated our attention on the subject of insect systematics. The most obvious changes in content are the addition of a chapter for a newly described order, the Mantophasmatodea, and the subordination of the Homoptera into an enlarged concept of the order Hemiptera. Beyond that, though, the classification of nearly every order has be en modified, sometimes substantially, to reflect new discoveries and scientific hypotheses. The chapter on beetles has been updated considerably to reflect the changes in our understanding of the diversity and phylogeny of Coleoptera. Many new families have been added throughout the book, some reflecting revised classifications, but many the result of the discovery of new groups within the United States and Canada, particularly from the New World tropics. These include the families Platystictidae (üdonata), Mackenziellidae (Collembola), Mantoididae

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(Mantodea), and Fauriellidae (Thysanoptera), to name just a few. Changes in classifications also have been brought about by the widespread adoption of the methods of phylogenetic systematics and their application to a new source of information on insect relationships, molecular sequence data. Although these new data will not help the beginning student to identify specimens, the results of molecular analyses are beginning to substantively con tribute to the the development of a robust and predictive classification. Thus, our best hypotheses of the phylogeny of insects has changed rather drastically from the last edition, incorporating molecular data. The most conspicuous change is the recognition that the order Strepsiptera is most closely related to the true flies (Diptera), rather than to the Coleoptera. As we turn to focus our efforts on the issues of insect systematics and evolution, a better appreciation for the magnitude of the diversity of life and Earth as well as the immediate and long-term threats to that same diversity have become important societal issues. lt is our hope that this text will continue to have an important role to play in understanding and preserving this diversity for the benefit of all. Donald Joyce BOITorwas the senior author on the first six editions of this book. He died before the last edition was printed. He was unsurpassed in his ability to construct keys for the identification of insects and was constantly modifying them to make certain that the user would arrive at the COITecttaxon. His discussion of the various families, containing facts gleaned from the literature, is amazing, considering that it was done before computers were available. Furthermore, the entire manuscripts were typed by BOIToron an old manual typewriter. He was well-versed in Greek and Latin and also knew shorthand. His influence was missed in preparation of this edition, and we hope that it would have met with his approval. C. A. T. N. EJ.

Acknowledgments

W

e are indebted to many individuals who contributed to this revision in many ways, from criticisms and suggestions to the complete rewriting of some of the chapters. Some are cited in individual chapters, but we take this opponunity to list them here along with our sincere thanks: Roben Anderson, Richard W Baumann, Brian Brown, George W Byers, Kenneth Christiansen, Shawn M. Clark, Peter Cranston, Neal Evenhuis, Paul H. Freytag, Gary A. P. Gibson, Ronald Hellenthal, Ronald W Hodges, Michael A. Ivie, David Kistner, Michael Kosztarab, Kumar Krishna, Roben E. Lewis, Jeremy A. Miller, Edward L. Mockford, John Morse, Luciana Musetti, Steve Nakahara, David Nickle, Manuel Pescador, Norman D. Penny, Hans Pohl,Jerry Powell, Roger Price,John E. Rawlins, Edward S. Ross, David Ruiter,James Slater, Manya Stoetzel, Catherine A. Tauber, Maurice J. Tauber, Kenneth J. Tennessen, Darrell Ubick, Tatyana S. Vshivkov, ThomasJ. Walker,James B. Whitfield, MichaelJ. Whiting. We would like to thank Woodbridge A. Foster for his careful revision of Chapter 4, Behavior and Ecology. We also gratefully acknowledge the services of Kathy Royer, Sue Ward, and Bruce Leach for help with preparation of the manuscript and in locating references. We accept the responsibility for all errors and cases in which the keys fail to work, or taxa are omitted or misplaced. We hope that these are few and not serious.

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Jable of Contents

1 InsectsandTheirWays 1 2 TheAnatomy,Physiology,and Developmentof Insects 5 3 Systematics, Classification,Nomenclature, and Identification 52 4 Behaviorand Ecology 62 5 PhylumArthropoda 99 6 Hexapoda 152 7 TheEntognathousHexapods:Protura, Collembola,Diplura 169 8 TheApterygoteInsects:MicrocoryphiaandThysanura 177 9 OrderEphemeroptera: Mayflies 181 10 OrderOdonata:Dragonfliesand Damselflies 193 11 OrderOrthoptera:Grasshoppers, Crickets, and Katydids 209 12 OrderPhasmatodea: Walkingsticksand LeafInsects 227 13 OrderGrylloblattodea:RockCrawlers 230 14 OrderMantophasmatodea 232 15 OrderDermaptera:Earwigs 234 16 OrderPlecoptera:Stoneflies 239 17 OrderEmbiidina:Web-Spinners 247

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Contents

18 19 20 21

OrderZoraptera:Zorapterans,Angellnsects 250 OrderIsoptera:Termites 252 OrderMantodea:Mantids 260 OrderBlattodea:Cockroaches 263

22 OrderHemiptera:TrueBugs,Cicadas,Hoppers,Psyllids, Whiteflies,Aphids,and ScaleInsects 268 23 OrderThysanoptera: Thrips 333 24 OrderPsocoptera:Psocids 341 25 OrderPhthiraptera:Lice 356 26 OrderColeoptera:Beetles 365 27 OrderNeuroptera:Alderflies,Dobsonflies,Fishflies, Snakeflies,Lacewings,Antlions,and Owlflies 469 28 OrderHymenoptera:Sawflies,ParasiticWasps,Ants,Wasps, and Bees 481 29 OrderTrichoptera:Caddisflies 558 30 OrderLepidoptera:Butterfliesand Moths 571 31 OrderSiphonaptera:Fleas 648 32 OrderMecoptera:Scorpionfliesand Hangingflies 662 33 OrderStrepsiptera:Twisted-WingParasites 669 34 OrderDiptera:Flies 672 35 Collecting,Preserving,and StudyingInsects 745 Glossary 779 Credits 798 Index 805

1

Insects and Their Ways

he science of taxonomy takes as its arbitrary startthe publication of the 10th edition of Linnaeus's Systema Naturae in 1758. More than two centuries later, nearly one million species of insects have been described and named. Biology in the 21st century has changed in many fundamental ways, led primarily by the revolution in molecular biology. Yet the study of diversity of life on Earth has not faded into the past. Rather, it has become reinvigorated by advances in other sciences and in technology. We continue to discover new species at an increasing rate, even as habitat destruction by the growing human population brings the threat of extinction. In 2002, entomologists announced the discovery of a new order of insects, the Mantophasmatodea, illustrating that our understanding of even the major groups is imperfecto Our goal in writing this book is to provide an introduction to the diversity of insects and their relatives and a resource for identifying the fauna of temperate North America. We thus hope to encourage the study of these fascinating creatures so that we all may better understand the world in which we live. lnsects are the dominant group of animals on Earth today. They far outnumber all other terrestrial animals, and they occur practically everywhere. Several hundred thousand different kinds have been described-three times as many as there are in the rest of the animal kingdom-and some authorities believe the total number of different kinds may approach 30 million. More than a thousand kinds may occur in a fair-sized backyard, and their populations of...