Table Of ContentAbbreviations 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 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). Sub-
seript numerals are oeeasionally used to designate the particular abdominal seg-
ment on which a sclerite is loeated.
a, anal vein aos, anterior oblique sulcus be, bursa copulatrix elm, calamistrum
A, anal vein, anal cell on mesepisternum bcv, bridge cross vein elp, elypeus
ab, abdomen, opisthosoma ap, appendix bg, book gills elpl, elypellus
ac, accessory, lanceolate, or Ap, apical cell bk, beak, proboscis, cls, claval suture
subanal vein apc, apical cross vein rostrum, or snout elt, claw tuft, clypeal
Ac, anal crossing (A apd, apodeme bl, blastoderm tubercle
branching posteriorly apo, apophysis bln, banksian line elv, claval vein
from Cu, often called the ar, arista bm, basal medial cell, cm, gastric caeca or caecum
cubito-anal cross vein) are, arculus basement membrane en, colon
acc, accessory cell are, areolet bm-cu, basal mediocubital ena,cornea
acg, accessory gland aro, arolium cross vein enge, corneagenous cells
acg, accessory gland an, point of aniculation bms, basalar muscle cnu, eleavage nuclei
ael, antennal elub as, antennal sulcus, anterior bp, brood pouch colm, collum, tergite
aelp, anteclypeus spiracle br, brain or basal radial cell of the first body
acr, acrostichal bristles ase, antennal sclerite brv, bridge vein segment
aes, acrosternite ask, antennal socket bt, breathing tube com, commissural trachea
act, acrotergite asp, apical spur buc, buccula or bucca como, tritocerebral
acv, anterior cross vein aspr, anterior spiracle bv, basal vein commissure
adf, adfrontal area at, alimentary tract bvn, brace vein cor, corim
aed, aedeagus ata, anterior tentorial arm covd, common oviduct
af, antennal fossa atb, anal tube e, costal vein cp, crop
agr, scrobe, groove in beak atp, anterior tentorial pit C, costal vein, costal cell cph, prosoma or
for reception of antenna au, auricle ca, corpus allatum cephalothorax
al, anallobe av, auxiliary vein cal, calypter or squama cpl, cortical cytoplasm
alp, anal loop aw, anterior wart cb, corbicula cr, cercus, lateral caudal
alu, alula awp, anterior no tal wing cbr, costal break filament, superior
am, axillary muscle process cc, crystalline cone appendage
an, antenodal cross vein ax, axilla cd, cardo crb, cribellum
AN, alinotUm AX, axillary cell cee, circumesophageal cre, cremaster
anc, anal cleft axc, axillary cell connective crl, crystalline lens
anes, antecostal suture axcr, axillary cord cen, cenchri cm, cornicle
anp, anal plate axs, axillary sclerite cg, cerebral ganglion cro, crochets
anr, anal ring axv, axillary vein ch, chelicera crp, carapace
ans, anus cho,chorion es, coronal suture
ant, antenna B, basal cell chp, cheliped esp, cusp of mandible,
ante, antecosta ba, basalare el, clypeus, clavus caudal spiracle
antl, antennule BA, basal anal cell, basal cla, clasper cu, cubital vein
ao, dorsal aorta areole clc, movable spines or calcaria Cu, cubital vein
cual>anterior cubital cell exl, exite lobe hc, humeral callus mdu, microduct
cuf,cubitalfork (fork of CuA) exm, extensor muscle hcl, hypostigmatic or truss cell mdv, median vein
cun, cuneus exo, exocuticle hd, head mem, membrane
cup, posterior cubital cell exp, exopodite hg, anterior portíon of the met, metasomatic segment
Cup,posterior cubital vein hindgut mf, medial fork (fork of
cut, cuticle f, frenulum ho,hom MP2)
cva, clava fa, face hp, humeral plate mg, midgut or mesenteron
cvs, cervical sclerite lb, frontal bristles hr, heart mh, movable hook or palp
CVXc,ervix fc, food channel hst, haustellum mi, median lobe
ex, coxa fch, lilter chamber hv, humeral or recurrent vein mm, marginal macroduet
exc, coxal cavity fen, frontal ganglion hyb, hypopleural bristles mn, mentum
exg,groove in coxa connectíve hyp, hypopharynx, mo, mouth
exp, eoxopodite of re, facial fovea intermediate stylet mp, mouthparts
abdominal appendages fg, frontal ganglion hypl, hypopleuron Mp, posterior media
lib, libula mpb, mesopleural bristles
d, discoidal or intercostal fl, flagellum iab, intra-alar bristles mpo, marginal 8-shaped
vein flb, flabellum
iap, interior appendage pore
D, diseal cell, or discoidal flm, flexor muscle (paraproct) ms, mesoderm
eell fm, femur
iar, interantennal ridge msd, mesoderm
de, dorsocentral bristles fmb, femoral bristles ias, interantennal suture msl, mesostemallobe
dcv,discal cross vein fn, fang of chelicera iep, infraepistemum mspl, medial supplement
dlm, dorsallongitudinal fob, fronto-orbital bristles il, ileum mst, mental seta
muscle fon, fontanelle
ism, intersegmental mt, Malpighian tubule
dm, domelike layer of for, foramen magnum membrane mts, metatarsus or lirst
euticle over nerve ending, fr, frons it, intercalated triangle tarsal segment
or discal medial cell frl, frontallunule ivb, inner vertical bristles mu, mucro
dm-cu, discal medio-cubital fs, frontal suture mv, marginal or radial
crossvein vein
fu, sternal apophysis, furca j, jugum
do, dorsal ostiole fun, funiculus or funicle jl, jugallobe mx, maxilla, dorsal stylet
dp, distal process of fv, frontal vitta mxa, maxillary
sensory cell I,leg articulatíon
DS],disjugal furrow g, galea L, length, lanceolate mxl, maxillary lobe
dta, dorsal tentorial arm gap, gonapophysis Iba, labial articulation mxn, maxillary nerve
dtra, dorsal trachea ge, genal comb Ibl, labellum mxp, maxillary palp
gcl, germ cell Ibm, labium, ventral stylet mxt, maxillary tentacle
e, eye,compound eye gex, gonocoxa Ibn, labial nerve
ee, eyeeap ge, gena Ibr, labrum, rostrum n, notum
eet, eetoderm gen, male copulatory lbrn, labral nerve nb, notopleural bristles
ef, epigastrie furrow apparatus le, lacinia ne, ventral nerve cord
eg, egg gf, genital forceps let, layer of the cutícle nod,nodus
ejd, ejaeulatory duct gh, gland hair 19, ligula, median lobe nll, pronotallobe
el, elytron gi, gills 11,lamina lingualis npl, notopleuron
emb, embolium gl, glossa lo, lorum npls, notopleural suture
emp, empodium gle, gland cell lp, labial palp nt, notaulus
en, endophallus gld, duct of gland cell Lp, lateral plate nu, nucleus
end, endocuticle, endoderm gis, gland spines 15, labial suture nv, neuron
endr, endodermal rudiments glt, gland tubercle 1st, lateral setae
enl, endite lobe gn, ganglion of ventral Itra, main longitudinal o,opening
enp, endopodite nerve cord tracheal trunk ob, oeellar bristles
ep, epidermis gna, gnathochilarium obv, oblique vein
eper, epicranium gon, gonangulum m, medial cross vein, mouth, oe, ocellus
epg, epigynum gpl, gonoplacs or recurrent nerve oep, oeciput
eph, epipharynx gr, gill remnants M, medial vein, medial eell oepd, ocellar pedicel
epi, epicuticle gs, guiar suture ma, mandibular articulation oes, ocular suleus
epm, epimeron gst, gonostylus MA, anterior media oeg, oecipital ganglion
epp, epipleurite gt, genal tooth MC, marginal eell og, optic ganglion
epr, epistomal ridge gu, gula mcf, median eaudallilament op,operculum
eps, epistemum gvp, genovertical or orbital mep, mieropyle opl, optie lobe
ept, epiproet, median caudal plate md, mandible opt, ocular point
filament, or inferior MD, median eell orp, orbital plate
appendage h, humeral eross vein mdn, mandibular nerve os, oecipital suleus
es, epistomal su\cus hal, haltere mdp, median plate (of osm, osmeterium (seent
eso, esophagus hb, humeral bristles wing), middle plate (of gland)
ex, exuvium hbr, hypostomal bridge embryo) ot, oeellar triangle
Continued on back endpaper
---
Borrorand DeLong's
Introduction to the Studyof Insects
Seventh Edition
UNIVERSIDADDECALDAS
B;.8lI0TECA
-
BC
CG
595.7
1385
Borrorand DeLong's
2005EJ.2
Introduction
to the Studyof Insects
Seventh Edition
CharlesA.Triplehorn
The Ohio State University
Norman F.Johnson
The Ohio State University
UNIVERSIDADDECALDAS
CENTRODEBIBLIOTECA
Fecha:MQr~," tr:ooo.l9~2:¡
Facultad:í11otora
VoIúmen: E¡empB: 2.- ~ LIBRO:595.73737 BO
e ~!OR ANO OELONG'S 1
Convré};>t Donaci6n: CmIje:
1- \.~ OOUCTION TO THE
! ~~ ~1Iimll~Mm~llrn~i; 21
iii 84.178
THOMSON
.
BROOKS/COLE
Australia. Canada . Mexico . Singapore · Spain · United Kingdom . United Statf;s'
"
-
THOMSON
.
BROOKS/COLE
Publisher: Peter Marshall Production Service: Graphic World
Development Editor: Elizabeth Howe Text Designer: Tani Hasegawa
Assistant Editor: Elesha Feldman Photo Researcher: Sarah Evertson
Editorial Assistant: Lisa Michel Copy Editor: Linda Purrington
Technology Project Manager: Travis Metz Cover Dtsigner: lrene Morris
Marketing Manager: Ann Caven Cover Image: Walter Hodges/CORBlS
Project Manager, Editorial Production: Belinda Krohmer Cover Printer: Phoenix Color Corp
Art Director: Rob Hugel Compositor: Graphic World
PrintIMedia Buyer: Barbara Britton Printer: R.R. Donnelley/Willard
Permissions Editor: Sarah Harkrader
COPYRIGHT @ 2005 BrookslCole, a division of Thomson Thomson Brooks/CoIe
Learning, Inc. Thomson Learning™ is a trademark used 10 Davis Drive
herein under license. Belmont, CA 94002
USA
ALL RIGHTS RESERVED. No part of this work covered by
the copyright hereon may be reproduced or used in any form Asia
or by any means-graphic, electronic, or mechanical, includ- Thomson Learning
ing but not limited to photocopying, recording, taping, Web 5 Shenton Way #01-01
distribution, information networks, or information storage UIC Building
and retrieval systems-without the written permission of the Singapore 068808
publisher.
AustraliaINew ZeaIand
Printed in the United States of America Thomson Learning
3 4 5 6 7 08 07 06 102 Dodds Street
Southbank, Victoria 3006
Aus tralia
For more information about our products, contact us at:
Thomson Learn
1in-8g00A-c4a2d3e-m05i6c3Resource Center Canada
Nelson
For permission to use material from this text or product, 1120 Birchmount Road
submit a request online at http://www.thomsonrights.com.
Any additional questions about permissions can be TCoarnoandtao, Ontario MIK 5G4
submitted by email
---
Preface
n Introduction to the Study oIInsects: this is the seventh edition of a textbook
Athat has been widely used in entomology classes in North America over
more than 50 years. Its value has been 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 imme-
diately apparent to knowledgeable readers even though the formal authorship
has now passed on to subsequent generations. We have prepared this new edi-
tion 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 Cran-
berry 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 subordi-
nation of the Homoptera into an enlarged concept of the order Hemiptera. Be-
yond that, though, the classification of nearly every order has been 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 classifi-
cations, 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 v
-~
vi Preface
(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 de-
velopment 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 evo-
lution, 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 be-
come 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 litera-
ture, 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
e are indebted to many individuals who contributed to this revision in
Wmany ways, from criticisms and suggestions to the complete rewriting of
some of the chapters. Some are cited in individual chapters, but we take this op-
ponunity 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, Be-
havior 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 refer-
ences. 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.
vii
-
Jable of Contents
1 InsectsandTheirWays 1
2 TheAnatomy,Physiologya, ndDevelopmenot f Insects 5
3 SystematicsC, lassification,Nomenclature,
and Identification 52
4 BehaviorandEcology 62
5 PhylumArthropoda 99
6 Hexapoda 152
7 TheEntognathousHexapodsP: rotura,
Collembola,Diplura 169
8 TheApterygoteInsects:MicrocoryphiaandThysanura 177
9 OrderEphemeropteraM: ayflies 181
10 OrderOdonata:Dragonfliesand Damselflies 193
11 OrderOrthoptera:GrasshoppersC,rickets,
and Katydids 209
12 OrderPhasmatodeaW: alkingsticksand LeafInsects 227
13 OrderGrylloblattodea:RockCrawlers 230
14 OrderMantophasmatodea 232
15 OrderDermaptera:Earwigs 234
16 OrderPlecopteraS: toneflies 239
17 OrderEmbiidina:Web-Spinners 247 ix
