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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 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
CVX, cervix 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
cross vein fu, sternal apophysis, furca j, jugum vein
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, eye eap 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

Borror and DeLong's

Introduction to the Study of Insects

Seventh Edition

Borrorand DeLong's
to the Studyof Insects

Seventh Edition

CharlesA. Triplehorn
The Ohio State University

Norman F.Johnson
The Ohio State University

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iii 84.178

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AnthatIntroduction to the Study oIInsects: this is the seventh edition of a textbook

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 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 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 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.

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 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


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 ix
x Contents

18 OrderZoraptera:Zorapterans,Angellnsects 250
19 OrderIsoptera:Termites 252
20 OrderMantodea:Mantids 260
21 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
Insects and Their Ways

he science of taxonomy takes as its arbitrary start- without them. By their pollinating activities, they make
Ting point the publication of the 10th edition of possible the production of many agricultural crops, in-
Linnaeus's Systema Naturae in 1758. More than two cluding many orchard fruits, nuts, clovers, vegetables,
centuries later, nearly one million species of insects and cotton; they provide us with honey, beeswax, silk,
have been described and named. Biology in the 21st and other products of commercial value; they serve as
century has changed in many fundamental ways, led food for many birds, fish, and other beneficial animals;
primarily by the revolution in molecular biology. Yet they perform valuable services as scavengers; they help
the study of diversity of life on Earth has not faded keep harmful animals and plants in check; they have
into the past. Rather, it has become reinvigorated by been useful in medicine and in scientific research; and
advances in other sciences and in technology. We people in all walks of life look on them as interesting
continue to discover new species at an increasing animals. A few insects are harmful and cause enormous
rate, even as habitat destruction by the growing hu- losses each year in agricultural crops and stored prod-
man population brings the threat of extinction. In ucts, and some insects transmit diseases that seriously
2002, entomologists announced the discovery of a affect the health of humans and other animals.
new order of insects, the Mantophasmatodea, illus- Insects have lived on Earth for about 350 million
trating that our understanding of even the major years, compared with less than 2 million for humans.
groups is imperfecto Our goal in writing this book is During this time, they have evolved in many directions
to provide an introduction to the diversity of insects to become adapted to life in almost every type of habi-
and their relatives and a resource for identifying the tat (with the notable and puzzling exception of the sea)
fauna of temperate North America. We thus hope to and have developed many unusual, picturesque, and
encourage the study of these fascinating creatures so even amazing features.
that we all may better understand the world in which Compared with humans, insects are peculiarly
we live. constructed animals. Humans might think of them as
lnsects are the dominant group of animals on inside out, because their skeleton is on the outside, or
Earth today. They far outnumber all other terrestrial upside down, because their nerve cord extends along
animals, and they occur practically everywhere. Sev- the lower side of the body and the heart lies above the
eral hundred thousand different kinds have been alimentary canal. They have no lungs, but breathe
described-three times as many as there are in the through a number of tiny holes in the body wall-all
rest of the animal kingdom-and some authorities be- behind the head-and the air entering these holes is
lieve the total number of different kinds may ap- distributed over the body and directly to the tissues
proach 30 million. More than a thousand kinds may through a multitude of tiny branching tubes. The heart
occur in a fair-sized backyard, and their populations and blood are unimportant in transporting oxygen to
often number many millions per acre. the tissues. lnsects smell with their antennae, some
A great many insects are extremely valuable to hu- taste with their feet, and some hear with special organs
mans, and society could not exist in its present form in the abdomen, front legs, or antennae. 1
.". -
2 Chapter1 Insects andTheirWays

In an animal whose skeleton is on the outside of ture also drops, and their physiological processes slow
the body, the mechanics of support and growth limit down. Many insects can withstand short periods of
the animal to a relatively small size. Most insects are freezing temperatures, and some can withstand long
relatively small: Probably three fourths or more are less periods of freezing or subfreezing temperatures. Some
than 6 mm in length. Their small size enables them insects survive these low temperatures by storing in
to live in places that would not be available to larger their tissues ethylene glycol, the same chemical that we
animals. pour into our car radiators to protect them from freez-
Insects range in size from about 0.25 to 330 mm in ing during the winter.
length and from about 0.5 to 300 mm in wingspread; Insect sense organs often seem peculiar compared
one fossil dragonfly had a wingspread of over 760 mm! with those of humans and other vertebrates. Many in-
Some of the longest insects are very slender (the 330- sects have two kinds of eye-two or three simple eyes
mm insect is a walking stick occurring in Borneo), but located on the upper part of the face and a pair of com-
some beetles have a body nearly as large as one's fist. pound eyes on the sides of the head. The compound
The largest insects in North America are some of the eyes are often very large, occupying most of the head,
moths, with wingspreads of about 150 mm, and the and may consist of thousands of individual "eyes."
walking stick, with a body length of about 150 mm. Some insects hear by means of eardrums, whereas oth-
The insects are the only invertebrates with wings, ers hear by means of very sensitive hairs on the anten-
and these wings have had an evolutionary origin dif- nae or elsewhere on the body. An insect that has
ferent from that of the vertebrates. The wings of flying eardrums may have them on the sides of the body at
vertebrates (birds, bats, and others) are modifications the base of the abdomen (grasshoppers) or on the front
of the forelimbs; those of insects are structures in addi- legs below the "knee" (katydids and crickets).
tion to the paired "limbs," and might be likened to the The reproductive powers of insects are often
wings of the mythical horse Pegasus. With wings, in- tremendous; most people do not realize just how great
sects can leave a habitat when it be comes unsuitable; they are. The capacity of any animal to build up its
adult aquatic insects, for example, have wings when numbers through reproduction depends on three char-
adult, and if their habitat dries up they can fly to an- acteristics: the number of fertile eggs laid by each fe-
other habitat. Under similar adverse conditions, fish male (which in insects may vary from one to many
and other aquatic forms usually perish. thousands), the length of a generation (which may
Insects range in color from very drab to brilliant; vary from a few days to several years), and the propor-
no other animals on Earth are more brilliantly colored tion of each generation that is female and will produce
than some of the insects. Some insects, such as the the next generation (in some insects there are no
japanese beetle and the morpho butterfly are glittering males).
and iridescent, like living jewels. Their colors and An example that might be cited to illustrate insects'
shapes have inspired artists for millennia. reproductive powers is Drosophila, the fruit fly that has
Some insects have structures that are amazing been studied by so many geneticists. These flies develop
when we compare them to vertebrates. The bees and rapidly and under ideal conditions may produce
wasps and some of the ants have their ovipositor, or 25 generations in ayear. Each female will lay up to
egg-Iaying organ, developed into a poison dagger 100 eggs, of which about half will develop into males
(sting) that serves as an excellent means of offense and and half into females. Suppose we start with a pair of
'j defense. Some ichneumonids have a hairlike ovipositor these flies and allow them to increase under ideal con-
100 mm long that can penetra te solid wood. Some ditions, with no checks on increase, for a single year-
snout beetles have the front of the head drawn out into with the original and each female laying 100 eggs before
a slender structure longer than the rest of the body, she dies and each egg hatching, growing to maturity,
with tiny jaws at the end. Some flies have their eyes sit- and reproducing again, at a 50:50 sex ratio. With 2 flies
uated at the ends of long, slender stalks, which in one in the first generation, there would be 100 in the sec-
South American species are as long as the wings. Some ond, 5000 in the third, and so on, with the 25th gener-
of the stag beetles have jaws half as long as their bod- ation consisting of about 1.192 X 1041flies. lf this many
ies and branched like the antlers of a stag. Certain in- flies were packed tightly together, 1000 to a cubic
dividuals in some of the honey ants become so en- inch, they would form a ball of flies 96,372,988 miles in
gorged with food that their abdomens become greatly diameter-or a ball extending approximately from Earth
distended. These serve as living storehouses of food, to the sun!
which they regurgitate "on demand" to other ants in Throughout the animal kingdom, an egg usually
the colony. develops into a single individual. In humans and some
Insects are cold-blooded creatures. When the en- other animals one egg occasionally develops into two
vironmental temperature drops, their body tempera- individuals (that is, identical twins) or, on rare occa-
InsectsandTheirWays 3

sions, three or four. Some insects carry this phenome- with that of vertebrates); and many have "invented"
non of polyembryony (more than one young from a things we may think of as strictly human accomplish-
singleegg) much further; some platygastrid wasps have ments. Some groups of insects have developed complex
as many as 18, some dryinid wasps have as many as 60, and fascinating social behavior.
and some encyrtid wasps have more than 1000 young Insects feed on an almost endless variety of foods,
developing from a single egg. A few insects have an- and they feed in many different ways. Thousands of
other unusual method of reproduction, paedogenesis species feed on plants, and practically every kind of
(reproduction by larvae). This occurs in the gall gnat plant (on land or in fresh water) is fed on by some kind
genus Miastor and the beetle genera Micromalthus, of insecto The plant feeders may feed on almost any
Phengodes,and Thylodrias. parl of the plant; caterpillars, leaf beetles, and leafhop-
In the nature of their development and life cycle, pers feed on the leaves, aphids feed on the stems, white
insectsron the gamut from very simple to complex and grubs feed on the roots, certain weevil and moth larvae
evenamazing. Many insects undergo very little change feed on the fruits, and so on. These insects may feed on
as they develop, with the young and adults having sim- the outside of the plant, or they may burrow into it.
ilar habits and differing principally in size. Most in- Thousands of insects are carnivorous, feeding on other
sects,in contrast, undergo in their development rather animals; some are predators, and some are parasites.
remarkable changes, both in appearance and in habits. Many insects that feed on vertebrates are blood suck-
Mostpeople are familiar with the metamorphosis of in- ing; some of these, such as mosquitoes, lice, fleas, and
sects and possibly think of it as commonplace, which, certain bugs, not only are annoying pests because of
as a matter of fact, it is. Consider the development of a their bites, but may serve as disease vectors. Some in-
butterfly: An egg hatches into a wormlike caterpillar; sects feed on dead wood; others feed on stored foods of
this caterpillar eats ravenously and every week or two all types; some feed on various fabrics; and many feed
sheds its exoskeleton; after a time it becomes a pupa, on decaying materials.
hanging from a leaf or branch; and finally a beautiful, The digger wasps have an interesting method of
wingedbutterfly emerges. Most insects have a life cycle preserving food collected and stored for their young.
like that of a butterfly; the eggs hatch into wormlike These wasps dig burrows in the ground, provision
larvae,which grow by periodically shedding their outer them with a certain type of prey (usually other insects
exaskeleton (together with the linings of the foregut, or spiders), and then lay their eggs (usually on the
hindgut, and breathing tubes), finally transforming body of the prey animal). If the prey animals were
into an inactive pupal stage from which the winged killed before being put into the burrows, they would
adult emerges. A fly grows from a maggot; a beetle dry up and be of little value as food by the time the
growsfram a grub; and a bee, wasp, or ant grows from wasp eggs hatched. These prey animals are not killed;
a maggotlike larval stage. When these insects become they are stung and paralyzed, and thus "preserved" in
adult, they stop growing; a little fly (in the winged good condition for the young wasps when they hatch.
stage)does not grow into a bigger one. Insects often have interesting and effective means
An insect with this sort of development (complete of defense against intruders and enemies. Many "play
metamorphosis) may live as a larva in a very different dead," either by dropping to the ground and remaining
type of place from that in which it lives as an adult. motionless or by "freezing" in a characteristic position.
One common household fly spends its larval life in Others are masters of the art of camouflage, being so
garbage or some other filth; another very similar fly colored that they blend with the background and are
mayspend its larvallife eating the insides out of a grub very inconspicuous; some closely resemble objects in
ar caterpillar. The june beetle that flies against the their environment-dead leaves, twigs, thorns, or even
screensat night spends its larvallife in the ground, and bird droppings. Some insects become concealed by cov-
the long-horned beetle seen on flowers spends its lar- ering themselves with debris. Others that do not have
vallife in the wood of a tree or log. any special means of defense very closely resemble an-
Many insects have unusual features of structure, other that does, and presumably are afforded some
physiology,or life cycle, but probably the most in ter- protection because of this resemblance. Many moths
esting things about insects are what they do. In many have the hind wings (which at rest are generally con-
instances the behavior of an insect seems to surpass in cealed beneath the front wings) brightly or strikingly
intelligence the behavior of humans. Some insects give colored-sometimes with spots resembling the eyes of
the appearance of an amazing foresight, especially as a larger animal (for example, giant silkworm moths; see
regards laying eggs with a view to the future needs of Figure 30-76)-and when disturbed display these hind
the young. Insects have very varied food habits; they wings; the effect may sometimes be enough to scare off
have some interesting means of defense; many have a potential intruder. Some of the sound-producing in-
what might be considered fantastic strength (compared sects (for example, cicadas, some beetles, and others)

4 Chapter 1 InsectsandTheirWays

produce a characteristic sound when attacked, and this lift some 60 tons, and an elephant could lift a fair-sized
sound often scares off the attacker. building! When it comes to jumping, many insects put
Many insects use a "chemical warfare" type of de- our best Olympic athletes to shame. Many grasshop-
fense. Some secrete foul-smelling substances when dis- pers can easily jump a distance of 1 meter, which is
turbed; stink bugs, broad-headed bugs, lacewings, and comparable to ahuman long-jumping the length of a
some beetles might well be called the skunks of the in- football field, and a flea jumping several inches up in
sect world, because they have a very unpleasant odor. the air is comparable to ahuman jumping over a 30-
A few of the insects using such defensive mechanisrns story building.
can eject the substance as a spray, in some cases even Many insects do things that we might consider
aiming it at an intruder. Some insects, such as the milk- strictly an activity of civilized humans or a product of
weed butterflies, ladybird beetles, and net-winged bee- our modern technology. Caddisfly larvae were probably
tles, apparently have distasteful or mildly toxic body the first organisms to use nets to capture aquatic or-
fluids, and predators avoid them. ganisrns. Dragonfly nymphs, in their intake and expul-
Many insects inflict a painful bite when handled. sion of water to aerate the gills in the rectum, were
The bite may be simply asevere pinch by powerful among the first to use jet propulsion. Honey bees were
jaws, but the bites of mosquitoes, fleas, black-flies, as- air-conditioning their hives long before humans even
sassin bugs, and many others are much like hypoder- appeared on Earth. Hornets were the first animals to
mic injections; the irritation is caused by the saliva in- make paper from wood pulpo Long before people began
jected at the time of the bite. making crude shelters, many insects were constructing
Other means of defense indude the stinging hairs shelters of day, stone, or "logs" (Figure 29-8), and
some caterpillars have (for example, the saddleback some even induce plants to make shelters (galls) for
caterpillar and the larva of the io moth), body fluids them. Long before the appearance of humans on Eanh,
that are irritating (for example, blister beetles), death the insects had "invented" cold light and chemical war-
feigning (many beetles and some insects in other or- fare and had solved many complex problems of aero-
ders), and warning displays, such as eyespots on the dynamics and celestial navigation. Many insects have
wings (many moths and mantids) or other bizarre or elaborate communication systems, involving chemicals
grotesque structures or patterns. (sex, alarm, trail-following, and other pheromones),
One of the most effective means of defense insects sound (cicadas, many Orthoptera, and others), behav-
possess is a sting, which is developed in the wasps, ior (for example, honey bee dance "language"), light
bees, and some ants. The sting is a modified egg-Iaying (fireflies), and possibly other mechanisms.
organ; hence only females sting. lt is located at the pos- These are only a few of the ways in which insects
terior end of the body, so the "business" end of a sting- have become adapted to life in the world about uso
ing insect is the rear. Some of the detailed stories about these animals are
Insects often perform feats of strength that seem fantastic and almost incredible. In the following chap-
nearly impossible compared with those of human be- ters, we point out many of the interesting and often
ings. lt is not unusual for an insect to be able to lift 50 unique features of insect biology-methods of repro-
or more times its own weight, and researchers have duction, ways of obtaining food, techniques for de-
found that some beetles, when rigged with a special positing eggs, methods of rearing the young, and fea-
harness, can lift more than 800 times their own weight. tures of life history-as well as the more technical
If they were as strong as such beetles, ahuman could phases that deal with morphology and taxonomy.

2 The Anatomy, Physiology,

and Development of Insects

standing of insect anatomy necessary to use the rest of

Aknowledge of anatomy and physiology is essential
to an understanding of insects. It is also necessary the book.
to havenames for structures in order to be able to talk Insects are more or less elongate and cylindrical in
about them. The nomenclature of insect anatomy form and are bilaterally symmetric; that is, the right
shouldbe viewed as a language, a tool, that makes pre- and left sides of the body are essentially alike. The body
cise discussions about insects possible, and not as a is divided into a series of segments, the metameres, and
barrier to understanding. In fact, many of the terms these are grouped into three distinct regions or tagmata
(for example, femur, trochanter, mandible) have analo- (singular, tagma): the head, thorax, and abdomen (Fig-
gousmeaningsin the context of vertebrate anatomy. ure 2-1). The primary functions of the head are sen-
Theterms that have special meanings in individual or- sory perception, neural integration, and food gather-
dersare discussed in the appropriate chapters. In ad- ing. The thorax is a locomotory tagma and bears the
dition,all terms used are defined in the glossary at the legs and wings. The abdomen houses most of the vis-
backof this book. Our primary purpose in this chap- ceral organs, including components of the digestive,
ter is to provide the student with the basic under- excretory, and reproductive systems.

hd th A
~~/ , ept
/,," __ cr

I f
\1/ ovp

Figure 2-1 General structure of an insectoab, abdomen; ant, antenna; cr, cercus; e,
compound eye; epm, epimeron; eps, epistemum; ept, epiproct; hd, head; Ibm, labium; md,
mandible; mp, mouthparts; mx, maxilla; n, nota of thorax; Ovp,ovipositor; pIs, pleural su-
ture; ppt, paraproct; spr, spiracles; t,_IO,terga; th, thorax; th¡, prothorax; thz, mesothorax;
th3,metathorax. (Modified from Snodgrass, 1935, Principies of Insect Morphology,
Comell University Press.) 5
:::z:: -

6 Chapter 2 TheAnatomy,Physiology,and Developmentof Insects

even differing in its characteristics from one part of an

The Body Wall insect to another. It is made up of chains of a polysac-
charide, chitin, embedded in a protein matrix. Chitin
The skeleton of an animal supports and protects the primarily consists of monomers of the sugar N-acetyl-
body, and transfers the forces generated by the contrac- glucosamine (Figure 2-3). Individual chitin chains are
tion of muscles. One of the fundamental features of intertwined to form microfibrils, and these microfibrils
arthropods is the development of hardened plates, or are often laid down in parallel in a layer called a lamina.
sclerites, and their incorporation into the skeletal sys- Chitin itseU is a very resistant substance, but it
tem of the animal. This is usually called an exoskele- does not make the cuticle hard. The hardness is de-
ton, because the sclerites are part of the outer body rived from modifications of the protein matrix in
wall of the arthropod. In fact, however, arthropods also which the microfibrils are embedded. The cuticle ini-
have an extensive endoskeleton of supports, braces, tially secreted by the epidermis, called procuticle, is
and ridges for the attachment of muscles. The charac- soft, pliant, pale in color, and somewhat expandable.
teristics of the body wall also influence the way in The formation of sclerites in this cuticle is the process
which substances such as water and oxygen move into of hardening and darkening, or sclerotization. This re-
and out of the animal. sults from the formation of cross bonds between pro-
The integument of an insect consists of three prin- tein chains in the outer portions of the procuticle. Such
cipal layers (Figure 2-2): a cellular layer, the epider- sclerotized cuticle is called exocuticle (Figure 2-2, exo).
mis; a thin acellular layer below the epidermis (that is, Below the exocuticle may be unsclerotized cuticle
toward the inside of the animal), the basement mem- called endocuticle (Figure 2-2, end). This pliant endo-
brane (or basal lamina); and another acellular layer, cuticle forms the "membranes" that connect sclerites
outside of and secreted by the cells of the epidermis, and can be resorbed into the body before molting.
the cuticle. Atop the endo- and exocuticle is a very thin, acel-
The cuticle is a chemically complex layer, not only lular layer, the epicuticle (Figure 2-2, epi). This itseU
differing in structure from one species to another but consists of layers: those generally present are cuticulin,






tmg trg gle

Figure2-2 Structure of the body wall (diagrammatic). bm, basement membrane; cut,
cuticle; end, endocuticle; ep, epidermis; epi, epicuticle; exo, exocuticle; glc, gland cell; gld,
duct of glandcell;Ict,layerof the cuticle;pcn,pore canal;se, seta;ss,setalsocket;tmg,
tormogen cell (which forms the setal socket); trg, trichogen cell (which forms the seta).
Abdomen 7




Figure 2-3 Chemical structure of chitin and its primary monomeric component,
N-acetylglucosamine. (From Arms and Camp 1987.)

su .....
a waxlayer, and a cement layer. The epicuticle contains ---'" "....
no chitin. The wax layer is very important to terrestrial ",
insects,because it serves as the primary mechanism to
limit the 1055of water across the body wall (both exo- cut -_
cuticle and endocuticle are permeable to water). As a ep E3
soliddecreases in size (as measured by volume, surface bm///
area,or some linear dimension) the ratio of its surface-
areato volume, that is, the relative amount of surface area,
increases.Therefore, the loss of water across the body
surface is relatively much more important to a small
creaturethan to a large one. Many small terrestrial an-
imals,such as snails and isopods, do not have such a
Figure 2-4 Diagram of external and internal features of
protective wax layer, but these creatures usually live
the body wall. apo, apophysis; bm, basement membrane;
only in regions of high relative humidity, which de-
creasesthe rate at which water is lost from their body. COS,costa; cut, cuticle; ep, epidermis; su, sulcus or suture.
The outermost cement layer is thought to protect the
waxbeneath from abrasion.
The sclerites of the body wall are often subdivided
bygroovesand crests, or may project into the body as in-
temalstruts. In general, an external groove marking an Abdomen
infoldingof cuticle of the outer body wall is called a sul-
cus (plural, sulcO (Figure 2-4, su). The term suture, We begin our discussion of the three tagmata of insects
alsovery widely used, refers to a line of fusion between with the abdomen beca use in contrast to the head and
two formerly separated sclerites. The distinction is a thorax, it is relatively simple in structure. Arthropods,
subtleone and often difficult or impossible to make sim- like vertebrates, are built on a basic ground plan of re-
ply by looking at the external structure of a specimen. peated body segments, or metameres. These are most
Therefore,in this book we generally use these terms clearly visible in the abdomen. In general, the abdomen
moreor less synonyrnously. The lines of inflection seen of an insect is made up of a maximum of 11 metameres
extemally usually correspond to internal ridges, or (Figure 2-1, ab). Each metamere typically has a dorsal
costae (Figure 2-4, cos). The internal costae may serve sclerite, the tergum (plural, terga; Figure 2-1, t¡-tlO;
as strengthening braces or as the sites of muscle attach- Figure 2-5A, t); a ventral sclerite, the sternum (plural,
ment.An external crest may be referred to as a costa or stema; Figure 2-1, stn; Figure 2-5A, stn); and a mem-
carina (or any number of common English names such branous lateral region, the pleuron (plural, pleura; fig-
as heel).Internal projections of cuticle are also referred ure 2-5A, plm). The openings to the respiratory sys-
to as apodemes or apophyses (Figure 2-4, apo). tem, the spiracles (Figure 2-1, spr) , typically are
8 Chapter2 TheAnatomy,Physiology,and Development of Insects


>-- ante
acs....... /
..... I

I \
I I \ ,
I I anes stn
stn ism


Figure 2-5 Structure of a typical abdominal segment (diagrammatic). A, Cross section;

B, Sagittal section. aes, acrosternite; aet, acrotergite; anes, antecostal suture; ante, ante-
costa; ism, intersegmental membrane; plm, pleural membrane; stn, sternum; t, tergum.

located in the pleuron. Terga and sterna may be subdi- these segments have no appendages. In the primitively
vided; these parts are referred to as tergites and ster- wingless insects,l the orders Microcoryphia and Thysa-
nites. Sclerites in the pleural wall are called pleurites. nura, the ventral portion of a pregenital segment gen-
This segmentation of the abdomen differs from erally consists of a small medial sternum and two large
that found in other nonarthropod protostomes such as plates laterad of the sternum, the coxopodites (see Fig-
the segmented worms (Annelida). In these, the exter- ure 8-1B, cxp and stn). The coxopodites are remnants
nally visible grooves in the body wall delimiting the of the bases of the abdominal legs, and apically they
metameres serve as the points of attachment for the bear a musculated stylus (Figure 8-1A,B, sty). The styli
dorsal and ventrallongitudinal muscles. In arthropods, probably represent the apical portions of these legs (the
these muscles attach to internal costae, the antecostae, telopods), but they are not segmented as are the tho-
which are located near, but not at, the anterior margins racic legs. The styli generally function as sensory or-
of the terga and sterna (Figure 2-5B, ante). Externally, gans and also support the abdomen, much like the run-
the position of the antecosta is indicated by a groove, ners of a sled. Mesal to the styli are one or sometimes
the antecostal suture (Figure 2-5B, ancs). The region two pairs of eversible vesicles, which function in water
of the tergum anterior to the antecostal sulcus is the absorption. They are everted from the body by hydro-
acrotergite (Figure 2-5B, act). The corresponding re- static pressure and retracted by muscles. In many cases
gion of the sternum is the acrosternite (Figure 2-5B, the coxopodites and the sternum are fused into a sin-
acs). The main dorsallongitudinal muscles extend be- gle composite sclerite, the coxosternum.
tween the antecostae of successive segments. Contrac- Pregenital abdominal appendages are present in
tion of these muscles results in a telescoping, or retrac- winged insects only in immature stages (with the ex-
tion, of the abdominal segments. This body plan, in ception of male Odonata). In embryos, the appendages
which the externally visible segmentation does not of the first abdominal segment, known as pleuropodia,
conform to the attachment of the longitudinal muscles, are present. These are glandular structures and are lost
is known as secondary segmentation. before the insect hatches from the egg. The larvae of
The genitalia of insects are generally located on or some Neuroptera (Figure 27-6A-C) and Coleoptera
about abdominal segments 8 and 9. These segments (Figures 26-19A,B, 26-21A) bear lateral styluslike
have a number of specializations associated with copu- structures that have variously been interpreted as rep-
lation and oviposition; our discussion of them is there-
fore included in the section later on the reproductive 1 As we describe in Chapter 6, we are distinguishing between the
systems. Segments 1-7, anterior to the genitalia, are the terms Hexapoda and Insecta, restricting the latter to refer to the
pregenital segments. In most adult winged insects, Pterygota, Thysanura, and Microcoryphia.
Thorax 9

resentingleg rudiments, styli, or secondarily developed are highly reduced or normally retracted within the
gills. The nymphs of Ephemeroptera bear a series of body.
platelike gills along the upper lateral portions of the
body (Figure 9-2). Again, from just what structures
thesegills were derived and what their serial homologs
maybe on the thorax have been considerably debated. Thorax
The immature stages of a number of orders have
prolegson the pregenital segments. These are typically The thorax is the locomotory tagma of the body, and it
Oeshy,short appendages that are important in walking bears the legs and wings. It is made up of three seg-
orcrawling (see, for example, Figures 30-3, prl; 30-74, ments, the anterior prothorax, mesothorax, and poste-
ventral,plg; and 28-37). Hinton (1955) concluded that rior metathorax (Figure 2-1, th¡-th3). Among insects,
prolegshad evolved independently a number of times; a maximum of two pairs of spiracles open on the tho-
others,such as Kukalová-Peck (1983), interpret these rax, one associated with the mesothorax, one with the
structures as modified abdominal legs, both homolo- metathorax. The mesothoracic spiracle serves not only
gousbetween orders and serially homologous with the that segment but also the prothorax and head. The
segmentedthoracic legs. terga of the thorax are typically called nota (singular,
The postgenital segments are typically reduced in notum). Among present-day insects, wings are borne at
insects. Among hexapods, the Protura are unique in most on the mesothoracic and metathoracic segments;
that theyhave 12 well-developedsegments in the ab- these two segments are collectively called the
domen (representing 11 metameres and an apical non- pterothorax to reflect this (pteron, Greek for "wing").
metamerictelson). In general, the only indications of These segments have several modifications associated
an Ilth segment among insects are a dorsal sclerite, the with flight that are not shared with the prothorax.
epiproct, and two lateral sclerites, the paraprocts (Fig- The prothorax is connected to the head by a mem-
ure 2-1, ept, ppt). Between them are inserted the ap- branous necklike region, the cervix (Figure 2-6, cvx).
pendages of the apical abdominal segment, the cerci Dorsallongitudinal muscles extend from the mesotho-
(singular, eereus). Typically the cerci are sensory or- rax through the prothorax and insert on the head; the
gans,but in some cases they are modified as organs of pronotum has no antecosta. Movements of the head co-
defense(as in the forceps of Dermaptera, Figures 15-1 ordinate with the rest of the body by one or two pairs
and 15-2) or may be specialized as accessory copula- of cervical sclerites (Figure 2-6, cvs) that articulate
toryorgans. Very often the apical abdominal segments with the prothorax posteriorly and the head anteriorly.

, epp
hd \
\ \

Figure 2-6 Thoraxof Panorpa,

lateral view. AN, alinotum; evs, cer-
vical sclerite; evx, cervix; ex, coxa;
epm, epimeron; epp, epipleurite;
eps, episternum; hd, head; nI>
pronotum; pIs, pleural suture;
PN, postnotum; pwp, pleural wing
process; se!, scutellum; set, scutum;
spr, spiracle; stn, abdominal
sternum; t, abdominal tergum;
wb, base of wing. (Redrawn from
Ferris and Rees 1939.)
-- ..

10 Chapter 2 The Anatomy, Physiology, and Development of Insects

The system of seeondary segmentation just de- others have suggested that, in essenee, both are eorrect
seribed in referenee to the abdomen is modified in the in that the thorade pleurites are eomposite in origino In
pterothorax lo aecommodate the flight musculature. any case, the selerotized portion of the pleuron is di-
The dorsal longitudinal museles of the meso- and vided by a suture that extends from the base of the leg
metathorax are greatly enlarged and are involved in to the base of the wing; this is the pleural suture (Fig-
depression (downward movement) of the wings (Fig- ure 2-6, pis). This suture divides the pleuron into an
ure 2-7B, dlm). As a eorollary, the sites of insertion of anterior episternum (Figure 2-6, eps¡-eps3) and a pos-
these museles-that is, the anteeostae of the mesotho- terior epimeron (Figure 2-6, epm¡-epm3). Aeeording to
rax, metathorax, and first abdominal segments-are the subeoxal theory of the origin of pleural selerites, the
also greatly enlarged and projeet downward within the pleurites originally eonsisted of a pair of ineomplete
thorax and base of the abdomen. These enlarged ante- rings above the base of the leg: an upper anapleurite
eostae are called the first, second, and third phragmata and a lower eatepleurite (the latter also ealled katepleu-
respeetively (singular, phragma; Figure 2-7B, ph¡-ph3). rite, eatapleurite, or eoxopleurite). Sueh selerites are vis-
Extemally, the mesonotum and metanotum are typi- ible in the primitively wingless hexapods and in a few
eally divided transversely by a sulcus that gives added pterygotes. The eatepleurite articulates with the leg.
flexibility. The sulcus divides eaeh notum into an ante- Thus the combination of the pleural suture and the two
rior seutum (Figure 2-6, set2, set3) and a posterior rings of the subeoxa ean theoretically define four re-
seutellum (Figure 2-6, sel2, sel3). In addition, the por- gions of the pleuron: anepimeron, anepistemum,
tions of the notum bearing the seeond and third eatepimeron, and eatepistemum (for an example, see
phragma are often separated from the following seu- terminology of McAlpine et al. 1981 for the dipteran
tum from whieh they are derived, and moved forward, thorax in Chapter 33). In addition to its dorsal articula-
sometimes even entirely fused with the selerites ante- tion with the eatepleurite, the leg articulates anteroven-
rior to them. These selerites bearing the seeond and trally with a narrow selerite (often entirely fused with
third phragmata are ealled postnota (Figures 2-6 and the epistemum), the troehantin. The wing rests on the
2-7, PN2, PN3). pleural wing proeess (Figure 2-6, pwp), whieh forros
The lateral portion of the thorax in winged inseets the dorsal apex of the pleural suture. Anterior lo the
is very different from the abdomen in that typieally it is pleural wing proeess is a small selerite, the basalare
strongly selerotized and relatively rigid. The origin of (Figure 2-I2A,B, ba); posterior to the pleural wing pro-
these pleural selerites has been eonsiderably debated. eess is another selerite, the subalare (Figure 2-I2A, sb;
Some researehers have argued that these pleural seler- oeeasionally two small selerites are found here instead
ites evolved de novo and have no serial homologs in of one). These selerites (sometimes eolleetively ealled
other parts of the body. Many others postulate that the epipleurites) are attaehed lo the base of the wing and
pleural selerites represent the ineorporation of a basal serve as means of eontrolling the attitude of the wings
leg segment, the subeoxa, into the body wall. Finally, or may be direetly involved in wing movement.

n2 PN2ancs
, , n3
, PN3ancs
i I I I I I
,' /t,

II dlm I dl'm I
I , I
ph, ph2 ph3


Figure 2-7 Endoskeleton of the thorax (diagrammatic). A, Cross section of a thoracic

segment; B, Longitudinal section of the thoradc dorsum. anes, antecostal suture; ex, coxa;
dlm, dorsallongitudinal muscles;fu, stemal apophyses or furca; n, notum; nI>pronotum;
n2>mesonotum; nJ, metanotum; ph, phragmata; pl, pleuron; plap, pleural apophyses;
PN2,mesopostnotum; PNJ, metapostnotum; tI>first abdominal tergum. (Redrawn from
Snodgrass 1935, PrincipIesoflnseet Morphology,1993, Comell University Press.)

Thorax 11

The external pleural suture corresponds to an in- second long segment; the tarsus (ts), usually a series of
ternal costa, the pleural ridge. This ridge extends in- small subdivisions beyond the tibia; and the pretarsus
ternallyon either side as a pair of pleural apophyses (ptar), consisting of the claws and various padlike or se-
(or pleural arms; Figure 2-7 A, plap). These pleural talike structures at the apex of the tarsus. A true seg-
apophyses are connected with a corresponding pair of ment of an appendage (including the six just described)
apophysesarising from the sternum (Figure 2-7 A, fu). is a subdivision with musculature inserted at its base.
The two may be connected by muscle or tendon, or in The subdivisions of the tarsus, though commonly called
somecasesthey are fused together. The bases of the tarsal segments, are not true segments in this sense and
sternalapophyses are often fused together, particularly are more properly called subsegments or tarsomeres. The
in species in which the legs are contiguous ventrally. pretarsus usually includes one or more padlike struc-
The apophyses then have a Y-shaped appearance, and tures between or at the base of the claws. A pad or lobe
the structure is called the furca. between the claws is usually called an arolium (Fig-
ure 2-8A,B, aro), and pads located at the base of the
claws are usually called pulvilli (Figure 2-8C, pul).
Legs The movements of a leg depend on its musculature
The thoracic legs of insects are sclerotized and subdi- and the nature of the joints between its segments. These
videdinto a number of segments. There are typically six leg joints may be dicondylic, with two points of articu-
segments (Figure 2-8): the coxa (cx), the basal seg- lation, or monocondylic, with a single point of articula-
ment;the trochanter (tr), a small segment (occasionally tion (Figure 2-9). The movement at a dicondylic joint is
twosegments) following the coxa; the femur (fm), usu- largely limited to the plane perpendicular to a line con-
allythe first long segment of the leg; the tibia (tb), the necting the two points of articulation, whereas that at a



ptar -- D

Figure2-8 Leg structure in insects. A, Middle leg of a grasshopper (Melanoplus);

B, Last tarsal segment and pretarsus of Melanoplus;e, Last tarsal segment and pretarsus
of a robber fly; D, Front leg of a katydid (Scudderia). aro, arolium; ex, coxa; emp,
empodium;fm, femur; ptar, pretarsus; pul, pulvillus; tb, tibia; tel, tarsal claw; tr,
trochanter; ts, tarsus; tym, tympanum.
12 Chapter2 TheAnatomy,Physiology,
andDevelopmentof Insects

, ,,- - ~~
I ''
I .~..::'::.::,.
.,: ...... ' ,art2

/ :;":':: ,,/
/ I
/ I
art, flm

A B e

Figure2-9 Articular mechanisms in insect legs. A, A monocondylic joint; B, C, End

view and side view of a dicondylic joint. art, points of articulation; exm, extensor muscle;
Jlm, Ilexor muscle. (Redrawn from Snodgrass 1935, PrincipIes oIInsect Morphology, 1993,
Comell University Press.)

monocondylic joint (which is like a ball-and-socket wing venation is very useful as a means of identifica-
joint) can be more varied. The joints between the coxa tion. Several venational terminologies have been devel-
and body may be monocondylic. If it is dicondylic, the oped, and the most widely used has been the Comstock-
axis of rotation is usually more or less vertical, and the Needham system (Cornstock and Needham 1898, 1899)
leg moves forward and backward (promotion and remo- (see Figure 2-10). This system basically recognizes a
tion). The coxo-trochanteral, trochantero-femoral, and series of six major longitudinal wing veins (with their
femoro-tibial joints are usually dicondylic. The move- abbreviations in parentheses): costa (C) at the leading
ment between the coxa and trochanter and the femur edge of the wing, followed by the subcosta (Sc), radius
and tibia is dorsal and ventral (elevation and depression (R), media (M), cubitus (Cu), and anal veins (A).
of the leg). The tibio-tarsal joint is usually mono- Each of these veins, with the exception of the costa,
condylic, thus permitting more varied movements. may be branched. The subcosta may branch once. The
branches of the longitudinal veins are numbered from
anterior to posterior around the wing by means of sub-
Wings script numerals: the two branches of the subcosta are
The wings of insects are outgrowths of the body wall designated Sc¡ and SC2'The radius first gives off a pos-
located dorsolaterally between the nota and pleura. terior branch, the radial sector (Rs), usually near the
They arise as saclike outgrowths, but when fully devel- base of the wing; the anterior branch of the radius is R¡;
oped are flattened and flaplike, and are strengthened by the radial sector may fork twice, with four branches
a series of sclerotized veins. Among living insects, fully reaching the wing margino The media may fork twice,
developed and functional wings are usually present with four branches reaching the wing margino The cu-
only in the adult stage. The one exception is the pres- bitus, according to the Comstock-Needham system,
ence of functional wings in the penultimate instar in forks once, the two branches being Cu¡ and CU2;ac-
Ephemeroptera (the subimago). At most, two pairs of cording to some other authorities, Cu. forks again dis-
wings are found in living insects, 10cated on the tal1y,with the two branches being CUla and CU¡b' The
mesothoracic and metathoracic segments. Most of the anal veins are typical1y unbranched and are usualIy
muscles that move the wings are attached to sclerites in designated from anterior to posterior as the first anal
ilie iliorade wa\\ railiet ilian to tne wings directly, and vein (11\), second anal vein (21\), and so on.
the wing movements are ptoduced indirecuy by CTossveins connect the major longitudinal veins
changes in the shape of the thorax. and are usually named accordingly (for example, th!
The wing veins are hollow structures that may medio-cubital crossvein, m-cu). Some crossveins havl
contain nerves' tracheae, and hemolymph (blood). The special names: two common examples are the humera
paccem ol venation varies considerably among differenr crossvein (h) and the sectorial crossvein (s).
groups of insects. Urde is known abour rhe funcrional The spaces in rhe wing membrane berween (he
significance of these differences, but the pattem of veins are calIed celIs. CelIs may be open (extending to
Thorax 13

Figure2-10 Generalized wing venation, according to Comstock; for a key to the letter-
ing, see accompanying textoIn some orders the vein here labeled Cu¡ is called Cu in the
Comstock-Needham system (and its branches Cu¡ and Cu2), and the remaining veins anal

thewing margin) or closed (completely surrounded by spedal names, for example, the triangles of the drag-
veins).The cells are named according to the longitudi- onfly wing and the discal cell of the Lepidoptera.
nal vein on the anterior side of the cell; for example, The wings of insects are attached to the thorax at
the open cell between R2 and R3 is called the R2 cell. three points (see Figures 2-11 and 2-12): with the no-
Wheretwo cells separated by a crossvein would ordi- tum at the anterior and posterior notal wing processes
narilyhave the same name, they are individually desig- (Figure 2-11, awp, pnwp), and ventrally at the pleural
nated by number; for example, the medial crossvein wing process (Figure 2-12A, pwp). In addition, small
connectsM2 and M3 and divides the M2 cell into two sclerites, the axillary sclerites (or pteralia) at the base
cells,the basal one is designated the first M2 cell and of the wing are important in translating the movements
the distal one the second M2 cell. Where a cell is bor- of the thoracic sclerites into wing movements. Most
deredanteriorly by a fused vein (for example, R2+3),it living insects (the Neoptera) have three axillary scler-
is named after the posterior component of that fused ites (Figure 2-11, axsCaxs3). Anteriorly the first axil-
vein(cell R3). In some insects, certain cells may have lary articulates with the anterior no tal wing process,

=_ __R
- --M
- - -Cu Figure2-11 Diagram showing the articula-
tion of the wing with the thoradc notum.
=---1A am, axillary musc1es; awp, anterior notal wing
process; axcr, axillary cord; axs, axillary
- - -2A sc1erites 1-3; hp, humeral plate;jl, jugallobe;
mdp, median plates; n, notum; pnwp, posterior
- - -3A notal wing process; tg, tegula. The letters at
the right side of the figure indicate the veins.
\ (Redrawn from Snodgrass 1935, Principies of
\ InsectMorphology,1993,CornellUniversity
jl Press.)
-'ro - _'1111111ii181

14 Chapter2 TheAnatomy,Physiology,
of Insects

the subcostal vein, and the second axillary. The second Odonata and in most Orthoptera, the phase difference
axillary then artieulates with the first, the radial vein, is less pronounced, with the front wings moving a lit-
the pleural wing process, and the third axillary. The tle ahead of the hind wings.
third axillary articulates with the second, the anal The forces needed to fly-lift, thrust, and attitude
veins, and the posterior notal wing process. In the control-are generated by the movement of the wings
Neoptera, a muscle (am) inserted on the third axillary through the air. These movements, in turn, are gener-
causes it to pivot about the posterior no tal wing pro- ated by the thoracic muscles pulling either directly on
cess and thereby to fold the wing over the back of the the base of the wing (direct flight mechanism) or caus-
insect. (Some groups of Neoptera, such as butterflies, ing changes in the shape of the thorax, whieh in turn
have lost this ability to flex the wings over the back.) are translated by the axillary sclerites into wing move-
Two groups of winged insects, the Ephemeroptera and ments (indirect flight mechanism). In most insects,
Odonata, have not evolved this wing-flexing mecha- the primary flight muscles are indirect: the dorsal lon-
nism, and their axillary sclerites are arranged in a pat- gitudinal muscles (Figure 2-12A,B, dlm) cause the no-
tern different from that of the Neoptera. Some special- tum to bow, thereby raising the no tal wing processes in
ists classify these two orders (with a number of extinct relation to the pleural wing process, depressing the
orders) together as the Paleoptera. wing. The antagonistie movement is generated by con-
The Comstock-Needham system made great tracting the tergosternal (dorsoventral or tergopleural)
strides in recognizing the homology in wing veins (Figure 2-12B, tsm); these pull down on the notum,
among the orders and in reducing the number of drawing the no tal wing processes down in relation
names associated with them. Kukalová-Peck (1978, to the pleural wing process, thereby elevating the wing.
1983, 1985) and Riek and Kukalová-Peck (1984) have In addition, muscles inserted on the basalare (Fig-
proposed a reinterpretation of the origin and basic ure 2-12, bms) and subalare (Figure 2-12A, sbm) can
structure of insect wings. The veins are interpreted to be involved in direct depression of the wing (by means
consist of a series of paired blood channels that loop of their connection to the wing margin at XI and x2) or
from the base of the wing to the apex and back again. may be important in controlling the angle at whieh the
The anterior vein in the loop protrudes from the dorsal wing moves through the air.
surface (a convex vein) and the posterior vein in the Flight, however, is not a simple matter of flapping
loop protrudes from the ventral surface (a concave the wings up and down. In addition, the wings are
vein). The fundamental venation in this interpretation brought forward (promotion) and backward (remo-
consists of eight major longitudinal vein systems: the tion), and twisted; that is, the leading edge is turned
precosta, costa, subcosta, radius, media, cubitus, anal downward (pronation) or the trailing edge is turned
vein, and jugal vein. The "vein" in the costal margin of downward (supination). The manner in whieh these
living insects is thus formed from the fusion of the pre- wing movements are produced involves a complex in-
costa, costa, and sometimes portions of the subcosta. tegration of the anatomical details of the attachment of
This interpretation has been applied to the orders the wing to the thorax and the contraction of muscles.
Ephemeroptera and Odonata, cases in whieh peculiar- The details are not completely known for any species,
ities in venation and axillary structure had led some to and only in a few can we begin to say we understand
postulate that wings had evolved in insects more than them at al!. In fact, it is clear that insects of different
once (see Chapters 9 and 10). sizes and shapes fly in different ways. A minute para-
sitie wasp of about 1 mm in length moves its wings dif-
Flight ferently and has wings of a different shape from a house
fly's, for example, and the aerodynamies of its flight are
Many insects have powers of flight that exceed those of probably also quite different.
all other flying animals; they can steer accurately and
quiekly, hover, and go sideways or backward. Only the
hummingbirds rival insects in their ability to maneuver
on the wing. Head
Most insects have two pairs of wings, and the two
wings on each side may be overlapped at the base or The head of insects consists of a series of metameric
hooked together in some way so that they move to- body segments, together specialized for food gathering
gether as one, or they may be capable of independent and manipulation, sensory perception, and neural inte-
movement. In many Odonata, the front and hind wings gration. Exactly how many segments are in the head
move independently, and there is a phase difference in has long been a matter of contention among morphol-
the movements of the two pairs; that is, when one pair ogists, with the postulated number ranging from 3 to 7,
is moving up, the other pair is moving down. In other The head bears the eyes, antennae, and mouthparts. Its
Head 15



Figure2-12 Diagram of the wing muscles of an insectoA, Lateral view; B, Cross section
of a wing-bearing segmento am, axillary muscles; AN, alinotum; axs2and axs3,second and
third axillary sclerites; ba, basalare; bms, basalar muscles; ex, coxa; dlm, dorsallongitudi-
nal muscles; ph, phragma; pl, pleuron; pIs, pleural suture; PN, postnotum; pwp, pleural
wing process; sb, subalare; sbm, subalar muscles; tsm, tergostemal muscles; w, wing;
Xl and X2,connections between basalare and subalare and wing base. (Redrawn from
Snodgrass 1935, PrincipIesoflnsectMorphology,1993, Comell University Press.)

shapediffersquite widely between groups of insects, magnum; for); through it ron the ventral nerve cord,
but somelandmarks are consistently visible to enable traeheae, the digestive system, muscles, so me times the
identificationof its component parts. dorsal blood vessel, and so on. The most posterior line
The head is divided by grooves into a number of of infleetion on the head eapsule outside of the occipi-
moreor less distinct sclerites (Figure 2-13). Typically tal foramen is generally the postoccipital suture (pos).
thereis a transverse sulcus extending across the lower This suture defines the limits of the posterior segment
partof the face just above the base of the mouthparts; of the head, the labial segment, named because it bears
the medial or anterior part of this sulcus is the episto- ventrally the most posterior set of mouthparts, the la-
mal sulcus (es), and the lateral portions above the bium. The area behind the postoecipital suture is the
mandiblesand maxillae are the subgenal sulci (sgs). postocciput (po); the area on the side of the head an-
The anterior portion of the head capsule, above the terior to this suture is the postgena (pg); and the dor-
epistomal sulcus and between the large compound sal portion of the head anterior to the suture is the oc-
eyes,is the frons (fr). The anterior area below the epi- ciput (oep). In so me cases an oecipital sulcus (os) is
stomalsulcus is the clypeus (clp). The area below the present that defines the anterior limits of the oeciput
eye,on the side of the head, and above the subgenal and postgenae (separating them fram the vertex and
su\cusis the gena (ge). The top of the head, between genae), but this is far fram universally presento
the eyes, is the vertex (ver). In many, if not most in- The points on the head where the arms of the ten-
sects,the frons, vertex, and genae are general areas of torium (a set of internal braces, see later) meet the
the head, and their edges are not clearly defined by head wall are usually marked by pits or slits visible ex-
sulci. ternally. The anterior tentorial pits (atp) are at the lat-
The head is connected to the thorax by the mem- eral ends of the epistomal sulcus; the posterior tentor-
branous cervix (cvx). The opening on the posterior ial pits (ptp) are at the lower ends of the postoecipital
sideof the head is the occipital foramen (or foramen suture.

16 Chapter 2 The Anatomy, Physiology, and Development of Insects


___ fr--l
_ - sas
505----- ---

Figure2-13 General structure of

B an insect head. A, Anterior view;
B, Lateral view; C, Posterior view.
ant, antenna; as, antennal sulcus;
atp, anterior tentorial pit; e/p, clypeus;
es, coronal suture; evx, cervix; e, com-
pound eye; es, epistomal sulcus;for,
foramen magnum;fr, frons;fs, frontal
suture; ge, gena; Ibm, labium; Ibr,
labrum; md, mandible; mx, maxilla;
oe, ocelli; oep, occiput; oes, ocular
sulcus; os, occipital sulcus; p, palps;
pg, postgena; po, postocciput; pos,
postoccipital suture; ptp, posterior ten-
lOrial pit; sas, subantennal sulcus; sgs,
subgenal sulcus; sos, subocular sulcus;
ver, vertex. (Modified from Snodgrass
1935, PrincipIes of lnseet Morphology,
1993, Comell University Press.)

In different insect groups the sulci and sclerites are located at the lower ends of the postoccipital su-
just described may be absent, or they may be supple- tures. These arms unite from side to side to form a ten-
mented by others. The nomenclature usually makes torial bridge (Figure 2-14, ttb). Some insects have dor-
use of the landmarks mentioned; for example, the fron- sal arms on the tentorium (dta) that extend to the
togenal sulci are lines of inflection separating the genae upper pan of the face near the antennal bases. The ten-
from the frons. In many groups, however, the naming torium serves to brace the head capsule against the pull
of these parts follows the traditions taxonomists devel- of the powerful mandibular muscles, as the point of at-
oped over the last century or more, and may not be tachment for muscles moving the head appendages,
standardized. and as protection for the subesophageal ganglion and
The head is braced intemally by a group of pharynx.
apophyses forming the tentorium (Figure 2-14). This The head appendages of hexapods, starting poste-
structure is usually H-shaped, X-shaped, or shaped like riorly and moving forward, are (1) the labium (fig-
the Greek letter 'ir (pO with the principal arms in a ure 2-13, lbm); (2) the maxillae (mx); (3) the mandibles
more or less horizontal plane and extending from the (md); (4) the labrum (lbr); and (5) the antennae (ant).
lower pan of the rear of the head to the face. The points These are described in more detail later. These repre-
where the anterior arms of the tentorium (Figure 2-14, sent modified appendages, serially homologous to the
ata) meet the face are marked extemally by the anterior thoracic walking legs. In the ancestral condition, the
tentorial pits (atp). which are located at eitner end of moutnparts are directed downward~ sucn a nead is
the epistomal sulcus between the frons and the called hypognathous. In many predalOry and burrow-
clypeus. The posterior arms of the tentorium meet the ing species, the mouthparts are directed anteriorly, the
head wall at the posterior tentorial pits (ptp) , which prognathous condition. FinallY' in some groups, espe-
Head 17


_-ptp Figure2-14 Head of an insect with a section

of the head wall cut away to show the tentorium
(diagrammatic). ata, anterior tentorial arms;
atp, anterior tentorial pits; e/p, clypeus; dta,
dorsal tentorial arms; epr, epistomal ridge;
es, epistomal sulcus; ge, gena; Iba, labial articu-
lation; Ibr, labrum; ma, mandibular articulation;
mxa, maxillary articulation; ocp, occiput; po,
postocciput; por, postoccipital ridge; pos, post-
\ occipital suture; ptp, posterior tentorial pit; sgr,
\ subgenal ridge; sgs, subgenal sulcus; ttb, tentor-
\ ial bridge. (Redrawn from Snodgrass 1935, Prin-
\ cipIes oIInsect Morphology, 1993, Comell
University Press.)

ciallythe Hemiptera, the mouthparts are directed pos- thoracic legs) of three postoral (behind the mouth)
teriorly; this is the opisthognathous condition (or, segments. The area anterior to the mouth bears the
when speaking of the beak of Hemiptera, the compound eyes, ocelli, antennae, and labrum; the in-
opisthorhynchous condition). terpretation of this region is a matter of contention,
The posterior surface of the head, between the and some of the hypotheses and the evidence in sup-
foramenand the labium, is membranous in most in- port of them are succinctly summarized by Rempel
sects,but in a few this region is sclerotized. This scler- (1975). Recent developmental and molecular studies
otizationmay be the result of the hypostomal areas support the idea that both the labrum and antennae are
(areasbelow the subgenal sulci posterior to the mandi- modified appendages associated with independent
bles) extending ventrally and toward the midline to head segments.
formwhat is called a hypostomal bridge, or (particu-
larlyin prognathous insects) the result of the postoc- Antennae
cipitalsutures extending forward onto the ventral side
of the head, with a sclerite developing between these The antennae are paired segmented appendages 10-
suturesand the foramen. In the latter case the sclerite cated on the head, usually between or below the com-
iscalledthe gula (see Figure 26-4, gu) and the anterior pound eyes. The basal segment is called the scape
extensionsof the postoccipital sutures are called guIar (Figure 2-15N, scp), the second segment the pedicel
sutures. In some groups, the hypostomal bridge may (ped), and the remainder the flagellum (O). In insects
be "overgrown" by extensions of the postgenae, thus (the Pterygota and the apterous orders Thysanura and
creatinga postgenal bridge. Microcoryphia), the "segments" of the flagellum lack
The number of segments making up the head is intrinsic musculature and therefore are thought to rep-
notapparenrin the adult insect, as the head sulci rarely resent subsegments of the apical, third true antennal
coinicidewith the sutures between the original seg- segmento These are often called flagellomeres to distin-
ments.Entomologists do not agree on the number of guish them from true musculated segments (although
segmentsin the insect head; the one area of consensus this anatomical characteristic is widely recognized,
is that the posterior three sets of mouthparts corre- these subsegments are still often called segments). This
spond to appendages (serially homologous with the type of antenna is called an annulated antenna, refer-
18 Chapter2 TheAnatomy,Physiology,and Developmentof Insects




~ K


Figure 2-15 Types of antennae. A, Setaceous (dragonfly); B, Filiform (ground beetle);

C, Moniliform (wrinkled bark beetle); D, Clavate (darkling beetle); E, Clavate (ladybird
beetle); F, Capitate (sap beetle); G, Serrate (click beetle); H, Pectinate (fire-colored bee-
tle); 1, Plumose (male mosquito);], Aristate (syrphid fly); K, Stylate (snipe fly); l, Flabe1-
late (cedar beetle); M, lamellate Oune beetle); N, Geniculate (chalcid). Antennae such as
those in D-F, l, and M are also called clubbed. ar, arista; as, antennal sulcus; ase, anten-
nal sclerite; ask, antennal socket;jl, flagellum; ped, pedicel; sep, scape; sty, style.
Head 19

ring to the subsegmentation of the flagellum. In the Mouthparts

orders Diplura and Collembola, more than the basal
Insect mouthparts typically consist of a labrum, a pair
three antennal segments are musculated; these are
each of mandibles and maxillae, a labium, and a hy-
calledsegmented antennae. An antenna arises from an
antennal socket that is membranous but is surrounded popharynx. These structures are modified, some times
significantly, in different insect groups and are often
by a ringlike antennal sclerite that often bears a small
used in classification and identification. The type of
process, the antennifer, on which the scape pivots.
mouthparts an insect has determines how it feeds and
Theantennae are primarily sensory in function and act
(in the case of most injurious species) what sort of
astactile organs, organs of smell, and in some cases or-
damage it does. We describe next the basic structure of
gansof hearing.
the mouthparts, followed by a few of the significant
Insect antennae vary greatly in size and form and modifications. Further information on the variations in
areimportant in identification. The following terms are
mouthpart structure can be found in the discussion of
usedto describe their shapes: individual insect orders.

Setaceous-bristlelike, the segments becoming MandibulateMouthparts

more slender distally; for example, dragonfly The most generalized condition of the mouthparts is
(Figure 2-I5A), damselfly, leafhopper. found in chewing insects, such as a cricket. These are
Filiform-threadlike, the segments nearly uniform in said to be "chewing" or "mandibulate" mouthparts be-
size and usually cylindrical; for example, ground cause of the heavily sclerotized mandibles that move
beetle (Figure 2-15B), tiger beetle. transversely and are able to bite off and chew particles
Moniliform-like a string of beads, the segments simi- of food. You can most easily see and study the mouth-
lar in size and more or less spherical in shape; for parts by removing them from a preserved specimen one
example, wrinkled bark beetle (Figure 2-15C). at a time and examining them under a microscope.
Serrate~sawlike, the segments, particularly those The labrum, or upper lip (Figure 2-13, lbr; fig-
in the distal half or two thirds of the antenna, ure 2-16E), is a broad, flaplike lobe located below the
more or less triangular; for example, click beetle clypeus on the anterior side of the head, in front of the
(Figure 2-15G). other mouthparts. Gn the posterior or ventral side of
Pectinate-comblike, most segments with long, slen- the labrum may be a swollen area, the epipharynx.
der, lateral processes; for example, fire-colored The mandibles (Figure 2-13, md; Figure 2-16D)
beetle (Figure 2-15H). are the paired, heavily sclerotized, unsegmented jaws
Clubbed-the segments increasing in diameter distally lying immediately behind the labrum. In the winged
(Figure2-15D-F,L,M). If the increase is gradual, the insects and the order Thysanura, they articula te with
condition may be termed clavate (Figure 2-15D,E). the head capsule at two points, one anterior and one
Thisname is also used more or less synonyrnously posterior, and move transversely (and therefore these
with the term clubbed. If the terminal segments are two taxa are classified together as the Dicondylia). The
rather suddenly enlarged, the condition is termed mandibles of chewing insects may vary somewhat in
capitate (Figure 2-15F). If the terminal segments structure; in so me insects (including the cricket), they
areexpanded laterally to form rounded or oval bear both cutting and grinding ridges, whereas in oth-
platelikelobes, the condition is termed lamellate ers (such as certain predaceous beetles) they are long
(Figure2-15M). Where the terminal segments have and sicklelike.
long,parallel-sided, sheetlike, or tonguelike lobes The maxillae (Figure 2-13, mx; Figure 2-16A) are
extending laterally, the condition is termed flabel- paired structures lying behind the mandibles; they are
late (Figure 2-15L). segmented, and each maxilla bears a feelerlike organ,
Geniculate-elbowed, with the first segment long and the maxillary palp (mxp). The basal segment of the
the following segments small and arising at an an- maxilla is the cardo (cd, plural cardines); the second
gle to the first; for example, stag beetle, ant, chal- segment is the stipes (stp, plural stipites). The palp is
cid (Figure 2-15N). borne on a lobe of the stipes called the palpifer (plC).
Plumose-feathery, most segments with whorls of long The stipes bears at its apex two processes: the lacinia
hair; for example, male mosquito (Figure 2-151). (lc), an elongate jawlike structure; and the galea (g), a
Aristate-the last segment usually enlarged and bear- lobelike structure.
ing a conspicuous dorsal bristle, the arista; for ex- The labium, or lower lip (Figure 2-13, lbm; Fig-
ample, housefly, syrphid fly (Figure 2-15J). ure 2-16C), is a single median structure (although it
Stylate-the last segment bearing an elongate terminal is derived from two maxilla-like mouthparts fusing
stylelike or fingerlike process, the style; for exam- along the midline) lying behind the maxillae. It is di-
pIe, robber fly,snipe fly (Figure 2-15K). vided by a transverse sulcus into two portions, a basal
20 Chapter2 TheAnatomy,Physiology,and Developmentof Insects

Figure 2-16 Mouthparts of a

cricket (Gryllus). A, Maxilla;
B, Median vertical section of the
head, showing relation of hy-
popharynx (hyp) to the other
parts (somewhat diagrammatic);
e, Labium; D, Mandible, showing
muscle attachments and points of
articulation; E, Labrum. art,
points of articulation of mandible;
ed, cardo; clp, clypeus;fr, frons;
g, galea; gl, glossa; hyp, hypophar-
ynx; Ibm, labium; Ibr, labrum;
le, ¡acinia; Ig, ligula; Ip, labial
palp; Is, labial suture; m, mouth;
md, mandible; mn, mentum;
mx, maxilla; mxp, maxillary palp;
art::: pgl, paraglossa; phx, pharynx;
plf, palpifer; plg, palpiger; pmt,
postmentum; prmt, prementum;
A smt, submentum; stp, stipes;
ver, vertex.

postmentum (pmt) and a distal prementum (prmt). haustellate mouthparts either are elongate and stylet-
The postmentum may be divided into a basal submen- like or are lacking.
tum (smt) and a distal mentum (mn). The prementum
bears a pair of labial palps (lp) and a group of apical The Mouthparts of Hemiptera. The beak in this order
lobes that constitute the ligula (lg). The labial palps (Figure 2-17) is elongate, usually segmented, and
are borne on lateral lobes of the prementum, called arises from the front (Heteroptera) or rear (Auchenor-
palpigers (plg). The ligula consists of a pair of mesal rhyncha, Sternorrhyncha) of the head. The external
lobes, the glossae (gl), and a pair of laterallobes, the segmented structure of the beak is the labium which is
paraglossae (pgl). sheathlike and eneloses four piercing stylets: the two
mandibles and the two maxillae. The labrum is a short
If the mandible and maxilla on one side of a spec-
imen are removed, the hypopharynx (Figure 2-16B, lobe at the base of the beak on the anterior side, and
hyp) becomes visible; this is a short, tonguelike struc- the hypopharynx is a short lobe within the base of the
ture located immediately in front of or above the la- beak. The labium does no piereing, but folds up as the
bium and between the maxillae. In most insects, the stylets enter the tissue fed on. The inner stylets in
ducts from the salivary glands open on or near the hy- the beak, the maxillae, fit together in such a way as to
popharynx. Between the hypopharynx, mandibles, and form two channels, a food channel and a salivary chan-
labrum lies the preoral food cavity, the cibarium, which nel. The palps are lacking.
leads dorsally to the mouth.
The Mouthparts of the Diptera. The biting Diptera in
Variations in Insect Mouthparts the suborder Nematocera and the Tabanomorpha in-
Insect mouJhparts can be elassified into two general elude the mosquito es (Figm:e l=l8~sand flies,
types, mandibulate (chewing) and haustellate (suck- punkies, black flies, horse flies, and snipe flies. Females
ing). In mandibulate mouthparts, the mandibles move of these insects have six piereing stylets: the labrum,
transversely, that is, from side to side, and the insect is the mandibles, the maxillae, and the hypopharynx; the
usually able to bite off and chew its food. Insects with labium usually serves as a sheath for the stylets. The
haustellate mouthparts do not have mandibles of this stylets may be very slender and needlelike (mosqui-
type and cannot chew food. Their mouthparts are in toes) or broader and knifelike (the other groups). The
the form of a somewhat elongated proboseis or beak maxillary palps are well developed, but labial palps are
through which liquid food is sucked. The mandibles in lacking (some dipterists regard the labellar lobes as
Head 21


Figure2-18 Mouthparts of a mosquito. A, Head of

Aedes,lateralview;B, Crosssectionof proboseisof
2-17 Mouthparts of the large milkweed bug, Anopheles. ant, antenna; bk, proboseis; clp, clypeus; e,
fasciatus(Dalias).A, Lateralviewof the head
Oncopeltus compound eye;fe, food channel; hyp, hypopharynx; lbm,
showingbeak, with the labrum detched from front of labium; lbr, labrum; md, mandible; mx, maxilla; mxp,
beak; B, Cross section of stylets (somewhat diagram- maxillary palp; se, salivary channel. (B, Redrawn from
matic).ant, antenna; bk, beak; bue, buccula; e, compound Snodgrass, after Vogel1921.)
eye;fe,food channel;j,jugum; lbm, labium; lbr, labrum;
lo,lorum;md, mandible; mx, maxilla; oe, ocellus; se, sali-
varychannel;sty, stylets; ty, tylus.

labialpalps). The salívary channel is in the hypophar-

ynx, and the food channel is located between the
groovedlabrum and the hypopharynx (for example,
the mosquitoes) or between the labrum and the man-
dibles(for example, punkies and horse flies). The la- Ibr fe
\ I
bium does no piereing and folds up or back as the \ I
styletsenter the tissue pierced.
The Muscomorpha lack mandibles, and the maxil-
lae are represented by the palps (maxillary stylets are
usuallylacking). The proboseis consists of the labrum,
hypopharynx, and labium. There are two modifications
ofthe mouthparts in these flies: (a) a piereing type and
(b) a sponging or lapping type. A B
The Muscomorpha with piereing mouthparts in-
eludethe stable fly (Figure 2-19), tsetse fly, horn fly,
and louse flies. The prineipal piereing structure in Figure2-19 Mouthparts of the stable fly,Stomoxys eal-
theseflies is the labium; the labrum and hypopharynx citrans (L.). A, Anterior view of head; B, Cross section
areslender and styletlike and líe in a dorsal groove of through haustellum. bk, rostrum; clp, clypeus;fe, food
the labium. The labium terminates in a pair of small, channel; hst, haustellum; hyp, hypopharynx; lbl, label-
hard plates, the labella, which are armed with teeth. lum; lbm, labium; lbr, labrum; mxp, maxillary palp; se,
The salivary channel is in the hypopharynx, and the salivary channel. (Redrawn from various sources; some-
foodchannel is between the labrum and hypopharynx. what diagrammatic.)
- "
22 Chapter2 TheAnatomy,Physiology,and Developmentof Insects

The proboscis in the louse flies (Hippoboscidae) is bella, a pair of large, 50ft, ovallobes. The lower surface
somewhat retracted into a pouch on the ventral side of of these lobes bears numerous transverse grooves that
the head when not in use. serve as food channels. The proboscis can usually be
The remaining Diptera with sponging or lapping folded up against the lower side of the head or into a
mouthparts include the nonbiting species such as the cavity there. These flies lap up liquids; the food may be
house fly (Figure 2-20), blow flies, and fruit flies. The already in liquid form, or it may first be liquefied by
mouthpan structures are suspended from a conical salivary secretions of the fly.
membranous projection of the lower pan of the head
called the rostrum. The maxillary palps arise at the dis- The Mouthparts of Lepidoptera. The proboscis of adult
tal end of the rostrum, and the pan of the proboscis be- Lepidoptera (Figure 2-21) is usually long and coiled
yond the palps is termed the haustellum. The labrum and is formed of the two galeae of the maxillae; the food
and hypopharynx are slender and lie in an anterior channel is between the galeae. The labrum is a narrow
groove of the labium, which forms the bulk of the transverse band across the lower margin of the face, and
haustellum. The salivary channel is in the hypophar- there are no mandibles and hypopharynx (except in the
ynx, and the food channellies between the labrum and Micropterigidae). The maxillary palps are usually small
the hypopharynx. At the apex of the labium are the la- or absent, but the labial palps are usually well devel-

Figure2-20 Mouthpans of the house Oy,

Musea domestica L. A, Anterior view of head;
B, Cross section through haustellum. bk, ros-
trum; clp, clypeus;fe, food channel; hst, haustel-
B lum; hyp, hypopharynx; Ibl, labellum; Ibm,
labium; Ibr, labrum; mxp, maxillary palp; se,
salivary channel. (Redrawn from Snodgrass
1935, Principies of Insect Morphology, 1993,
A Comell University Press.)


Figure2-21 Mouthparts of a moth. A, Lateral view of head; B, Anterior view of head;

C, Cross section through proboscis. ant, antenna; atp, anterior tentorial pit; e, compound
eye;fe, food channel;fr, frons; Ibr, labrum; Ip, labial palp; m.x,maxilla (galea); oe, ocellus;
pf, pilifer; prb, proboscis. (Redrawn from Snodgrass 1935, Principies of Inseet Morphology,
1993, Comell University Press.)
Digestive System 23

oped.Thereis no speeialsalivarychannel. This type of The power of a muscle varies with the size of its cross-
mouthpart structure is sometimes called siphoning- sectional area, or with the square of its width; what the
sucking,because there is usually no piereing and the in- muscle moves (¡he mass of the body) varies with the
sect merely sucks or siphons liquids up through the cube of the linear dimension. Thus as the body be-
proboseis.Some moths in Southeast Asia and northem comes smaller, the muscles become relatively more
Australia,however, use the proboseis to pierce the skins powerful.
of 50ftfruits and then siphon liquids from the tissues Insect muscles are often capable of extremely
undemeath. When used, the proboscis is uncoiled by rapid contraction. Wing stroke rates of a few hundred
bloodpressure; it recoils by its own elastieity: per second are fairly common in insects, and rates up
to 1000 or more per second are known. In insects with
relatively slow wing-stroke rates and in most other
skeletal muscles, each muscle contraction is initiated
Insed Muscles by a nerve impulse. Such muscles are called synchro-
nous or neurogenic muscles because of this one-to-one
Ihe muscular system of an insect is comprised of from correspondence between action potentials and muscle
severalhundred lO a few thousand individual muscles. contractions. In insects with higher wing-beat fre-
Allconsist of striated muscle cells, even those around quencies, the muscles contract much more often than
the alimentary canal and the heart. The skeletal mus- the rate at which neural impulses reach them. The
des, which attach lO the body wall, move the various rates of contraction in such asynchronous muscles
parts of the body, including the appendages. The cell (found prineipally in flight muscles but sometimes in
membranesof the muscle and epidermis are interdigi- other oscillating systems) depend on the characteris-
tatedand interconnected by desmosomes; from the tics of the muscles themselves and the assoeiated
desmosomes,microtubules run to the outer epidermal sclerites. Nerve impulses are necessary to initiate con-
cellmembrane, and from there attachment fibers run tractions, but thereafter serve to generally maintain
through the cuticle to the epicuticle. The attachment the rate of contractions rather than to stimulate each
fibersare not broken down between the times when the one individually:
epidermisis separated from the old cuticle (apolysis) That insect muscles may have such an extremely
andthe shedding of that cuticle (ecdysis; see section on rapid contraction frequency, which is sometimes main-
moltinglater). Thus the muscles remain attached to tained for a prolonged period, attests to the effieiency
the body wall, and the insect continues to be able to of their metabolismo The tracheal system provides the
moveduring the period when a new cuticle is being large volumes of oxygen needed for such metabolic
formed.The locations of the points of attachment of rates. In most insects, the tracheoles (across whose
theskeletal muscles are sometimes useful in determin- walls gas exchange takes place) indent the cell mem-
ingthe homologies of various body parts. The visceral branes of the muscles, thus minimizing the distance
musdes, which surround the heart, the alimentary across which diffusion of gases must take place. Insects
canal,and the ducts of the reproductive system, pro- use a variety of fuels for flight. Carbohydrates are im-
duce the peristaltic movements that move materials portant for many speeies; in others, lipids are the pri-
alongthese tracts. They usually consist of longitudinal mary fuels; in some flies (such as the tsetse), amino
andcircular muscle fibers. acids form the substrate for generating the energy nec-
Ihe muscles moving the appendages are arranged essary for flight.
segmentally,generally in antagonistic pairs. Some ap-
pendageparts (for example, the galea and laeinia of the
maxillaeand the pretarsus) have only flexor muscles.
Thesestructures are usually extended by a combina- Digestive System
tionof hemolymph pressure and the elastieity of the
cutide.Eachsegment of an appendagenormally has its Insects feed on almost every organic substance found
ownmuscles. The tarsal and flagellar "segments" do in nature, and their digestive systems exhibit consider-
nothave their own muscles and thus are not true seg- able variation. The alimentary canal is a tube, usually
ments. somewhat coiled, which extends from the mouth to the
lnsect muscles seem to us to be very strong: Many anus (Figure 2-22). It is differentiated into three main
insectscan lift 20 or more times their body weight, and regions: the foregut, or stomodaeum; the midgut, or
jumping insects can often jump distances equal to mesenteron; and the hindgut, or proctodaeum. Both
manytimes their own length. These feats appear very the foregut and hindgut are derived from ectodermal
remarkablewhen compared to what humans can do; tissue and are lined intemally by a thin layer of cuticle,
theyare possible not because the muscles of insects are the intima. This cuticle is shed at each molt along with
inherently stronger, but because insects are smaller. the outer exoskeleton.
;8: - ~

24 Chapter2 TheAnatomy,Physiology,and Developmentof Insects


gn mg

Figure 2-22 Internal organs of a grasshopper, shown in longitudinal section (somewhat

diagrammatic). ans, anus; ao, dorsal aorta; be, bursa copulatrix; ea, corpus allatum; eee,
circumesophageal connective; eg, cerebral ganglion (part of the brain); em, gastric caeca;
en, colon; ep, crop; eg, eggs; eso, esophagus; gn, ganglia of ventral nerve cord; hr, heart;
hyp, hypopharynx; iI, ileum; lbm, labium; lbr, labrum; mg, midgut or mesenteron; mt,
Malpighian tubules; ne, ventral nerve cord; og, optic ganglion (part of the brain); ovd,
oviduct; ovp, ovipositor; ovt, ovarian tubules; phx, pharynx; ree, rectum; segn, subesopha-
geal ganglion; slg, salivary gland; sld, salivary duct; spth, spermatheca; vag, vagina.
(Redrawn from Robert Matheson: Entomologyfor Introductory Courses,Seeond Edition.
Comstock Publishing Company, Inc.)

Most insects have a pair of glands lying below the tima. The anterior part of the foregut is provided with
anterior part of the alimentary canal (Figure 2-22, dilator muscles, which have their origins on the walls
slg). The ducts from these glands extend forward and and apodemes of the head and thorax and their inser-
unite into a common duct that opens near the base of tions on the stomodaeal muscle layers, the epithelium,
the labium or hypopharynx. These labial glands (so or intima. These are best developed in the pharyngeal
named because they open at the base of the labium) region in sucking insects, where they make the phar-
generally function as salivary glands. There is often an ynx into a sucking pump. The crop is specialized for
enlargement of the duct from each gland that serves the temporary storage of food. It may be a simple en-
as a reservoir for the salivary secretion. The labial largement of the foregut, or, as in mosquitoes and Lep-
glands in the larvae of Lepidoptera, Trichoptera, and idoptera, it may be a lateral diverticulum off the diges-
Hymenoptera secrete silk, which is used in making tive tract. Uttle or no digestion of food takes place in
cocoons and shelters and in food gathering by net- the foregut.
spinning caddisflies. The midgut (mg) is usually an elongate tube of
The foregut is usually differentiated into a phar- rather uniform diameter, sometimes differentiated into
ynx (phx, immediately beyond the mouth), esophagus two or more parts. It often bears diverticula, the gastric
(eso, a slender tube extending posteriorly from the caeca (cm), near its anterior end. The midgut is not
pharynx), crop (cp, an enlargement of the posterior lined by cuticle. The epitheliallayer of the midgut is in-
portion of the foregut), and proventriculus. At its pos- volved both with the secretion of digestive enzymes
terior end is the stomodaeal valve, which regulates the into the lumen and in the absorption of the products of
passage of food between the foregut and midgut. In digestion into the body of the insect. Individual midgut
some groups, such as cockroaches and termites, the epithelial cells are generally rather short-lived and are
proventriculus may bear an armature of teeth inter- constantly being replaced. These dividing cells may be
nally; these are used to further crush the food before it scattered throughout the midgut, or may be concen-
enters the midgut. The intima is secreted by the foregut trated as pockets of growth. Such areas are sometimes
epithelium and is relatively impermeable. The intima visible from the lumen of the gut as invaginated crypts
and epithelium are often longitudinally folded. Outside and from the outer side as bulges (called nidi). The
of the epithelium is an inner layer of longitudinal mus- midgut is the primary site of digestion and absorption
cles and an outer layer of circular muscles. The longi- in the alimentary canal. In many species, the midgut
tudinal muscles sometimes have insertions on the in- epithelium and the food are separated by a peritrophic
Digestive System 25

membrane-a nonliving, permeable network of chitin has large, thick rectal pads that are important in re-
and protein that is secreted by the epithelium. The moving water fram the feces.
function of the peritraphic membrane is unclear. It The filter chamber is a modification of the ali-
mayserve to limit abrasion of the epithelium, to inhibit mentary canal in which two normally distant parts are
the movement of pathogens fram the food to the in- held close together by connective tissue; it occurs in
sect'stissues, or as a means of separating endo- and ec- many of the Hemiptera and varies somewhat in form
toperitrophic spaces within which digestive specializa- in different members of the order. The midgut in these
tioncan occur. insects is differentiated into three regions: the first,
The hindgut extends fram the pyloric valve, second, and third ventriculi. The first and second ven-
which lies between the midgut and hindgut, to the triculi are saclike structures immediately posterior to
anus. Posteriorly it is supported by muscles extending the esophagus, and the third ventriculus is a slender
to the abdominal wall. The hindgut is generally differ- tube. Typically, the third ventriculus turns forward and
entiatedinto at least two regions, the anterior intestine comes to lie close to the first ventriculus, often coiling
and the posterior rectum (rec). The anterior intestine about it, where it is held in place by connective tissue.
maybe a simple tube, or it may be subdivided into an This complex-the first ventriculus, the coiled third
anterior ileum (il) and a posterior colon (en). The ventriculus, and the connective tissue-forms the fil-
Malpighian tubules (mt), which are excretory organs ter chamber (Figure 2-23). Beyond the filter chamber,
(seelater), arise at the anterior end of the hindgut, and the alimentary canal continues backward, usually as a
their contents empty into it. The hindgut is the final slender tube, to enter the rectum. The Malpighian
site for resorption of water, salts, and any nutrients tubules emerge either fram the filter chamber or just
framthe feces and urine. The rectum in several species beyond it.


eso / eso

/ vnt


rec rec

Figure2-23 The filter chamber of Hemiptera (diagrammatic). A, A simple type of filter

chamber, in which the two ends of the midgut (the first and third ventriculO are bound
together; B, The filter chamber of a scale insect (Leeanium); e, A filter chamber in which
the posterior pan of the midgut (the third ventriculus) coils about the anterior pan (the
first ventriculus), with the hindgut emerging from the anterior end. In A and B the junc-
tion of the midgut and hindgut (where the Malpighian tubules, which are not shown, en-
ter the alimentary tract) is in the filter chamber. eso, esophagus; feh, filter chamber; hg,
anterior portion of the hindgut; mt, Malpighian tubules; prct, proctodeum; ree, rectum;
vnt, ventriculus (1, first; 2, second; 3, third).
26 Chapter2 TheAnatomy,Physiology,
andDevelopmentof Insects

Many Hemiptera live on plant juices, which they their food, and aphids inject amylase into the plant tis-
usually ingest in large quantities. Entomologists think su es and thus digest starch in the food plant. Such ex-
the filter chamber is a device that allows water from in- traintestinal digestion is also found in the predaceous
gested sap to pass directly from the anterior portion of larvae of antlions and predaceous diving beetles, and
the midgut to the hindgut, concentrating sap before di- bugs that feed on dry seeds.
gestion in the posterior part of the midgut. This excess Most chemical digestion of the food takes place
fluid passes from the anus as honeydew. However, be- within the midgut. Some of the midgut epithelial cells
cause honeydew is often rich in nutrients such as car- produce enzymes, and others absorb digested food.
bohydrates and amino acids, there is some doubt about Sometimes the same cells carry out secretion and ab-
the exact function of the filter chamber. sorption. Enzymes may be released into the lumen of
Digestion is the process of changing food chemi- the midgut by the disintegration of the secretory cells
cally and physically so that it can be absorbed and (holocrine secretion) or by the release of small
nourish various parts of the body. This process may be- amounts of enzymes across the cell membrane (mero-
gin even before the food is ingested but usually occurs crine secretion).
as the ingested materials pass through the digestive Only a few species of insects produce enzymes
tract. Solid foods are broken down by various mechan- that digest cellulose, but some can use cellulose as food
ical means (chiefly the mouthparts and proventricular as a result of symbiotic microorganisms present in their
teeth), and all foods are subjected to a battery of en- digestive tracts. These microorganisms, usually bacte-
zymes as they pass through the digestive tract. ria or flagellated protists, can digest the cellulose, and
Insects feed on a great variety of living, dead, and the insects absorb the products of this digestion. Such
decomposing animals, plants, and fungi and on their microorganisms are present in termites and many
products. In some cases, liquids such as blood or plant wood-boring beetles and are often housed in special or-
juices may constitute their entire food supply. The di- gans connected to the gut.
gestive system varies considerably with the different The fat body is a large, often somewhat amor-
kinds of foods consumed. The food habits may even phous organ housed in the abdomen and thorax. In
vary greatly in a single species. Larvae and adults many ways its function is analogous to that of the liver
usually have entirely different food habits and differ- among vertebrates. It serves as a food reservoir and is
ent types of digestive systems. Some adults do not feed an important site of intermediate metabolismo In some
at all. species it is also important in storage excretion (see
Most insects take food into the body through the page 27). The fat body is usually best developed in the
mouth. Some larvae that live endoparasitically in a host late nymphal or larval instars. By the end of meta mor-
animal can absorb food through the surface of their phosis, it is often depleted. Some adult insects that do
bodies from host tissues. Many insects have chewing not feed retain their fat body in adult life.
mandibles and maxillae that cut, crush, or macerate
food materials and force them into the pharynx. In
sucking insects, the pharynx functions as a pump that
brings liquid food through the beak into the esopha- Excretory System
gus. Peristaltic action moves food is along the alimen-
tary canal. The primary excretory system of an insect consists of a
Saliva is usually added to the food, either as it en- group of hollow tubes, the Malpighian tubules, which
ters the alimentary canal or before, as in the case of arise as evaginations at the anterior end of the hindgut
many sucking insects that inject it into the fluids they (Figure 2-22, mt). These tubules vary in number from
siphon up as foods. Saliva is generally produced by the one to more than several hundred, and their distal, free
labial glands, and the labial glands of many insects also ends are closed. These tubules function in removing ni-
produce amylase. In certain bees, these glands secrete trogenous wastes and in regulating, together with the
invertase, which is later taken into the body with nec- hindgut, the balance of water and various salts in the
tar. In bloodsucking insects such as mosquitoes, the hemolymph. Ions apparently are actively transponed
saliva generally contains no digestive enzymes but con- across the outer membrane of the tubule, generating an
tains a substance that prevents coagulation of the blood osmotic flow of water into the lumen. Along with this
and the consequent mechanical plugging of the food water a number of small solutes-amino acids, sugars,
channel. This saliva causes the irritation produced by and nitrogenous wastes-enter the tubule passively.
the bite of a bloodsucking insect. This primary urine is therefore an isosmotic solution
Many insects eject digestive enzymes on food, and containing the small molecules present in the he-
partial digestion may occur before the food is ingested. molymph. Some of these solutes and the water may be
Flesh fly larvae discharge proteolytic enzymes onto actively resorbed into the hemolymph in the basal por-
Circulatory System 27

molt, and in the pratrusion of eversible structures such

as eversible vesicles and genitalia. It may also function
in the animal's internal defenses, in the phagocytic ac-
tion of hemocytes against invading micraorganisms, in
plugging wounds, and in walling off certain foreign
bodies such as endoparasites. Finally, the hemolymph
is also a storage tissue, serving as a reservoir for water
2-24 Stucture of uric acid. and such food materials as fat and carbohydrates.
The circulatory system of an insect is open. The
main (and often only) blood vessel is located dorsal to
the alimentary tract and extends thraugh the thorax
tionsof the Malpighian tubules or in the hindgut. The and abdomen (Figure 2-22). Elsewhere the hemo-
principal nitragenous waste is usually uric acid (Fig- lymph flows unrestricted thraugh the body cavity Cthe
ure 2-24), a chemical that is relatively nontoxic (and hemocoel). The posterior part of the dorsal blood ves-
can therefore be tolerated in higher concentrations) sel, which is divided by valves into a series of cham-
and insoluble in water (again, recall the water balance bers, is the heart (hr), and the slender anterior part is
problemsinherent to a small terrestrial organism). the aorta (ao). Extending fram the lower surface of the
In some insects, most notabIy many beetle and heart to the lateral portions of the terga are pairs of
moth larvae, the Malpighian tubules are bound very sheetlike muscle bands. These constitute a dorsal di-
dosely to the hindgut; these are called cryptone- aphragm more or less separating the region around the
phridia. In species such as the mealworm, Tenebrio heart (the pericardial sinus) from the main body cavity
molitorL., that live in conditions of high draught (or perivisceral sinus, sometimes further divided into a
stress,this arrangement of the tubules is apparently in- perivisceral sinus and a perineural sinus). The heart is
volvedin extracting water fram the fecal pellets. The pravided with paired lateral openings called ostia, one
mealworm,in fact, can extract water vapor fram the air pair per heart chamber, thraugh which hemolymph en-
whenthe relative humidity exceeds 90%. ters the heart. The number of ostia varies in different
In addition to the Malpighian tubules, various in- insects. Some have as few as two pairs.
seetshave a range of methods of removing wastes or The hemolymph is usually a more or less clear
toxicsubstances fram the hemolymph. One method is fluid in which are suspended a number of cells (the he-
to store chemicals, such as uric acid, more or less per- mocytes). lt may be yellowish or greenish but is only
manendywithin individual cells or tissues. This pra- rarely red (as in some aquatic midge larvae and some
eessis known as storage excretion. Cockraaches store aquatic Hemiptera, owing to the exceptional presence
urieacid in their fat body, and the white pigment in the of hemoglobin). lt makes up fram 5% to 40% of the
sealesof pierid butterflies derives from uric acid stored body weight (usually about 25% or less).
within them. At the anterior end of the dorsal blood The liquid part of the hemolymph (the plasma)
vesselmay be a graup of cells, the pericardial cells, that contains a great many dissolved substances (such
are important in absorbing and breaking down col- as salts, sugars, prateins, and hormones). These vary
loidalparticles in the hemolymph. In other cases, sim- considerably-in different insects and in the same in-
ilareeUsmay be widely distributed thraughout the he- sect at different times. The plasma contains very little
moeoel. oxygen; the transport of oxygen is the function of the
respiratory system and is decoupled fram the circula-
tory system.
The hemocytes vary considerably in number-
CirculatorySystem fram about 1000 to 100,000 per mm3-but average
about 50,000 per mm3. These cells vary greatly in
Theprincipal function of the blood, or hemolymph, is shape and function. Some circulate with the he-
the transport of materials-nutrients, hormones, molymph, and some adhere to the surface of tissues.
wastes,and so forth. In most cases it plays a relatively The functions of the various types of hemocytes are not
minorrole in the transport of oxygen and carbon dio x- well known, but many are capable of phagocytosis.
ideoThe hemolymph is also involved in osmoregula- They may ingest bacteria, and they play a rale in re-
tion, the balance of salts and water in the body; this moving dead cells and tissues during metamorphosis.
function also involves other organs, particularly the The hemolymph of different insects differs in clotting
Malpighiantubules and the rectum. The hemolymph ability; the hemocytes may migrate to wounds and
has an important skeletal function-for example, in form a plug. Hemocytes often congrega te around for-
molting,in the expansion of the wings after the last eign bodies such as parasitoids and parasites, forming
-. ..

28 Chapter2 TheAnatomy,Physiology,and Developmentof Insects

a sheath around them and walling them off from the

body tissues. Other than the action of these hemocytes,
insects have no immune system comparable to the an-
tibodies of vertebrates (thus facilitating tranplantation
experiments) .
Hemolymph is moved about by pulsations of the
heart and in other parts of the body, such as the base
of the legs and wings, by accessory pulsatile organs.
The heartbeat is a peristaltic wave that starts at the
posterior end of the dorsal blood vessel and moves for-
ward. Hemolymph enters the heart through the ostia,
which are closed during the systolic phase of the
heartbeat, and is pumped anteriorly. The rate of the
heartbeat varies greatly: Observed rates in different in-
sects range from 14 to about 160 beats per minute.
This rate increases during periods of increased activity.
The pulsations of the heart may be initiated within the
heart muscle (myogenic), or they may be under ner-
vous control (neurogenic). A reversal of the direction
of the peristaltic wave of contractions, thus moving
the hemolymph backward instead of forward, is not
Very little pressure develops in the general flow of
hemolymph through the body. The hemolymph pres-
sure may sometimes be less than atmospheric pressure.
It can be increased by muscular contraction and com-
pression of the body wall or by dilation of the alimen-
tary canal (produced by swallowing air). This is how
pressure is developed to break out of the remainder of
the old exoskeleton at molting time and to inflate the
wings. Figure2-25 Diagram of a horizontal section of an in-
sect showing the arrangement of the principal tracheae.
ant, antenna; com, commissural trachea; dtra, dorsal tra-
chea; e, compound eye; 1,legs; Itra, main longitudinal tra-
Respiratory System cheal trunk; spr, spiracles; stra, spiracular trachea; vtra,
ventral tracheae.
Gas transport in insects is the function of the tracheal
system. The circulatory system of insects, unlike that
of vertebrates, usually plays only a minor role in this The spiracles are located laterally in the pleural
process. wall and vary in number from 1 to 10 pairs (some
The tracheal system (Figure 2-25) is a system of species have no functional spiracles). There is typically
cuticular tubes (the tracheae) that extemally open at a pair on the anterior margin of both the meso- and
the spiracles (spr) and intemally branch and extend metathorax, and a pair on each of the first eight (or
throughout the body. They terminate in very fine fewer) abdominal segments. They vary in size and
closed branches, called tracheoles, that permeate and shape and usually have some sort of valvelike closing
actually penetra te the living tissues (indenting but not device. These valves therefore play an important role in
actuaUy breaking through the cell membranes). The retaining body water.
tracheae are lined with a layer of cuticle, and in the In insects with an open tracheal system (that is,
larger branches this is thickened to form helical rings, with functional spiracles), air enters the body through
called taenidia, that simultaneously give the tracheae the spiracles, then passes through the tracheae to the
strength (against collapse) and flexibility (lo bend and tracheoles, and oxygen ultimately enters body cells by
twist). The tracheoles (also lined with cuticle) are diffusion. Carbon dioxide leaves the body in similar
minute intracellular tubes with thin walls, and they of- fashion. The spiracles may be partly or completely
ten contain fluido It is across the walls of the tracheoles closed for extended periods in some insects. Water los5
through the spiracles may be minimized in this way.
~~ts.~~hange.ac.tually takes. ~~!ce.
Respiratory System 29

Insects generally have longitudinal tracheal trunks Insects that live in water and get their oxygen from
(connectives, ltra) connecting the tracheae from adja- atmospheric air do this in one of three general ways:
centspiracles on the same side of the body and trans- from air spaces in submerged parts of certain aquatic
versecommissures (com) connecting the tracheae on plants, through spiracles placed at the water surface
oppositesides of the body, so that the entire system is (with the body of the insect submerged), or from a film
interconnected. The movement of air through the tra- of air held somewhere on the surface of the body while
chealsystem is by simple diffusion in many small in- the insect is submerged. A few larvae (for example,
sects,but in most larger insects this movement is aug- those of the beetle genus Donada and the mosquito
mentedby active ventilation, chiefly by the abdominal genus Mansonia) have their spiracles in spines at the
muscles;the movements of the internal organs, or of posterior end of the body, and these spines are inserted
the legs and wings, may also aid ventilation. Where into the air spaces of submerged aquatic plants. Many
ventilation occurs, air may move in and out of each aquatic insects (for example, waterscorpions, rattailed
spiracle,but generally enters through the anterior spi- maggots, and the larvae of culicine mosquitoes) have a
raclesand leaves by the posterior ones. This flow of air breathing tube at the posterior end of the body, which
through the tracheal system is effected by controlling is extended to the surface. Hydrophobic hairs around
which spiracles are open and when. Sections of the the end of this tube enable the insect to hang from the
maintracheal trunks are often dilated to form air sacs, surface film, and they prevent water from entering the
whichhelp in ventilation. breathing tube. Other aquatic insects (for example,
Closed tracheal systems have the spiracles perma- backswimmers and the larvae of anopheline mosqui-
nentlyclosed but have a network of tracheae just un- toes) get air through posterior spiracles placed at the
der the integument, distributed either widely over the water surface. These insects do not have an extended
bodyor particularly below certain surfaces (the gills). breathing tube.
Someaquatic and parasitic insects have closed systems. The insects that get their oxygen from atmo-
In these species, gases enter and leave the body by dif- spheric air at the water surface do not spend all their
fusionacross the body wall between the tracheae and time at the surface. They can submerge and remain un-
theexternal environment, and gases move through the derwater for a considerable period, getting oxygen
trachealsystem by diffusion. from an air store either inside or outside the body. The
A great many insects live in water; these get their air stores in the tracheae of a mosquito larva, for ex-
oxygenfrom one (rarely both) of two sources: the oxy- ample, enable the larva to remain underwater for a long
gendissolved in the water or atmospheric oxygen. Gas time.
exchangein many small, soft-bodied aquatic nymphs Many aquatic bugs and beetles carry a thin film of
andlarvae (and possibly some adults) occurs by diffu- air somewhere on the body surface when they sub-
sionthrough the body wall, usually into and out of a merge. This film is usually under the wings or on the
trachealsystem. The body wall in some cases is un- ventral side of the body. The air film acts like a physi-
modifiedexcept perhaps for having a fairly rich tra- cal gill, with dissolved oxygen in the water diffusing
cheal network just under the integument. In other into the bubble when the partial pressure of oxygen in
cases,there are special thin extensions of the body wall the film falls below that of the water. The insect may
thathave a rich tracheal supply and through which gas get several times as much oxygen from this temporary
exchangeoccurs. These structures, called tracheal gills, structure as was originally in it, because of gas ex-
comein a variety of shapes and may be located on dif- changes between the air film and the surrounding wa-
ferentparts of the body. The gills in mayfly nymphs are ter. A few insects (for example, elmid beetles) have a
inthe form of leaflike structures on the sides of the first permanent layer of air around the body surface, held
sevenabdominal segments (Figure 9-2). In dragonfly there by a body covering of thick, fine, hydrophobic
nymphs, they are folds in the rectum, and water is hairs; such a layer is called a plastron. The air reser-
movedinto and out of the rectum and over these folds. voirs of aquatic insects not only playa role in gas ex-
In damselfly nymphs, the gills are three leaflike struc- change but also may have a hydrostatic function (like
turesat the end of the abdomen as well as folds in the the swim bladder of fish). Two crescent-shaped air sacs in
reClUm(Figure lO-l). In stonefly nymphs, the gills are Chaoborus larvae (Diptera: Chaoboridae, Figure 34-30A)
fingerlike or branched structures located around the are apparently used to regulate this insect's specific
basesof the legs or on the basal abdominal segments gravity: to hold it perfectly motionless or to enable it to
(Figurel6-2B). Gas exchange may occur through the migrate up or down in the water column.
generalbody surface of these insects, and in some cases Parasitic insects that live inside the body of their
(suchas damselfly nymphs), the exchange through the host get oxygen from the body fluids of the host by dif-
bodysurface may be more important than that through fusion through their integument, or (for example, in
the tracheal gills.

30 Chapter2 TheAnatomy,Physiology,
andDevelopmentof Insects

tended to the body surface of the host or attach to one

of the host's tracheal trunks. ocpd
" ao

Body Temperature
Insects are generally considered cold-blooded or poiki-
lothermic; that is, their body temperature rises and falls
with the environmental temperature. This is the case
with most insects, particularly if they are not very ac-
tive, but the action of the thoracic muscles in flight
usually raises the insect's temperature above that of the
environment. The cooling of a small object is fairly
rapid, and the body temperature of a small insect in
flight is very close to that of the environment. In in-
sects such as butterflies and grasshoppers, the body
temperature in flight may be 5°C or 10°C above the en-
vironmental temperature, and in insects such as moths
and bumble bees (which are insulated with scales or
hair), the metabolism during flight may raise the tem-
perature of the flight muscles 20°C or 30°C above the
environmental temperature.
With most flying insects, the temperature of the
flight muscles must be maintained above a certain
point to produce the power necessary for flight. Many
larger insects may actively increase the temperature of Figure2-26 Anterior part of the nervous system of a
their flight muscles before flight by a "shivering" or a grasshopper. ao, dorsal aorta; br¡, protocerebrurn; br2'
vibration of the wing muscles. deutocerebrum; br3,tritocerebrurn; ea, corpus allatUrn;
Honey bees remain in the hive during the winter, cee, circurnesophageal connective; comn, tritocerebral
but do not go into a state of dormancy at the onset of commissure; ep, crop;fen, frontal ganglion connective;fg,
cold weather (as most other insects do). When the frontal ganglion; Ibn, labial nerve; Ibm, labral nerve; mdn,
temperature gets down to about 14°C, they form a clus- rnandibular nerve; mxn, rnaxillary nerve; oeg, oCcipital
ter in the hive and, by the activity of their thoracic ganglion; oepd,ocellar pedicel; opl, optic lobe; phx, phar-
muscles, maintain the temperature of the cluster well ynx; m, recurrent nerve; segn, subesophageal ganglion; sld,
over 14°C (as high as 34°C to 36°C when they are rear- salivary duct; tnt, tentoriurn. (Redrawn frorn Snodgrass
ing brood). 1935, PrincipIesoflnsect Morphology, 1993, Comell
University Press.)

Nervous System by a commissure that passes under the esophagus

(comn). The ventral nerve cord (Figure 2-22, nc) is typi-
The central nervous system of an insect consists of a cally double and has segmental ganglia (Figure 2-22, gn).
brain in the head above the esophagus, a subesopha- Frequently some of these ganglia are fused, particularly
geal ganglion (Figure 2-26, segn) connected to the toward the end of the abdomen, resulting in fewer visi-
brain by two nerves (the circumesophageal connectives, ble ganglia than segments.
cec) that extend around each side of the esophagus, and The ganglia of the central nervous system (brain,
a ventral nerve cord extending posteriorly from the sub- subesophageal ganglion, and segmental ganglia of the
esophageal ganglion. The brain consists of three pairs of central nerve cord) serve as the coordinating centers.
lobes, the protocerebrum (br.), deutocerebrum (br2), Each of these has a certain amount of autonomy; tha!
and tritocerebrum (br3). The protocerebrum innervates is, each may coordinate the impulses involved in activ-
the compound eyes and ocelli, the deutocerebrum in- ities in particular regions of the body. Activities involv-
nervates the antennae, and the tritocerebrum innervates ing the en tire body may be coordinated by impulses
the labrum and foregut. The two lobes of the tritocere- from the brain, but many of these can occur with the
brum are separated by the esophagus and are connected brain absent.
Nervous System 31

SenseOrgans many (for example, butterflies, moths, and flies) have

taste organs on the tarsi.
Aninsect receives information about its environment
The exact mechanism by which a particular sub-
(including its own internal environment) through its stance initiates a nerve impulse in the sensory cells of
senseorgans. These organs are located mainly in the a chemoreceptor is not completely understood. The
bodywall, and most are microscopic in size. Each is substance may penetrate to the sensory cells and stim-
usuallyexcited only by a specific stimulus. Insects have ulate them direct1y, or it may react with something in
senseorgans receptive to chemical, mechanical, audi- the receptor to produce one or more other substances
tory,and visual stimuli, and possibly such stimuli as that stimulate the sensory cells. In any event, an in-
relativehumidity and temperature. sect's sensitivity to different substances varies; two very
similar chemicals (such as the dextro and levo forms of
Chemical Senses
a particular sugar) may be quite different in their stim-
Chemoreceptors-those involved in the senses of taste
ulating effect. Some scents (for example, the sex at-
(gustation)and smell (olfaction)-are important parts
tractant produced by a female) can be detected by one
ofan insect's sensory system and are involved in many
sex (in this case, the male) but not by the other. The
Iypesof behavior. Feeding, mating, habitat selection,
sensitivity of chemoreceptors to some substances is
and parasite-host relationships, for example, are often
very high. Many insects can detect certain odors at very
directedby the insect's chemical senses:
low concentrations up to a few miles from their source.
Generally each sensillum consists of a group of
sensorycells whose distal processes form a bundle ex- Mechanical Senses
lendingto the body surface (Figure 2-27C). The end- Insect sense organs sensitive to mechanical stimuli re-
ings of the sensory processes are usually in a thin- act to touch, pressure, or vibration, and provide the
walled,peglike structure (scn). The peglike process insect with information that may guide orientation,
maybe sunk in a pit, or the sensory processes may end general movements, feeding, flight from enemies, re-
in a thin cuticular plate set over a cavity in the cutic1e. production, and other activities. These sense organs are
Insomecases the endings of the sensory processes may of three principal types: hair sensilla, campaniform
liein a pit in the body wall and may not be covered by sensilla, and scolopophorous organs.
cuticle. The simplest type of tactile receptor is a hair sen-
Ihe organs of taste are located principally on the sillum (Figures 2-27A and 2-28). A process from the
mouthparts, but some insects (for example, ants, bees, sensory neuron extends to the base of the seta, and
andwasps) also have taste organs on the antennae, and movements of the seta initiate impulses in the neuron.

I dp

- --cut- --

/ I \ / I
trg snc nv tro I \
nv bm
A B e

Figure2-27 Insect sensilla. A, Hair sensillum; B, Campaniform sensillum; C, Chemo-

receptor. bm, basement membrane; cut, cuticle; dm, domelike layer of cuticle over nerve
ending; dp, distal process of sensory cell; ep, epidermis; nv, neuron; scn, sense cone; se,
seta; snc, sensory cell; trg, trichogen cell. (A, C, Redrawn from Snodgrass 1935, PrincipIes
of Insect MorphoIogy,1993, Comell University Press.)

32 Chapter2 The Anatomy.Physiology.and Developmentof Insects


- B
~~~.". - e
\'~ "- \\;l~'" ~
\\\, 1. .

Figure2-28 Scanning electron microscope photographs of some insect sensilla. A, Sen-

silla on the antenna of an aphid, 1750x; B, Hair sensilla on the antenna of a clearwing
moth, 95 X; e, Hair sensilla on the antenna of a braconid, 500 x.

In a campaniforrn sensillum, the neuron ending lies just ble to the statocysts of crustaceans, although air bub-
under a domelike area of the cuticle (Figure 2-27B), bles carried on the body surfaces by certain aquatic in-
and distortion of this dome elicits a neuronal response. sects when they submerge may act in a similar manner.
Scolopophorous organs (also known as chordotonal or- The forces of gravity and pressure generally are de-
gans) are more complex sensilla that consist of a bun- tected by other means.
dle of sensory neurons whose dendrites are auached to Many joints in insects have tactile setae that regis-
the body wall; they are sensitive to movements of the ter any movements, providing the insect with informa-
body (including pressure and vibration). These organs, tion on the position of the joint (lhis is known as pro-
which are widely distributed over the body, include the prioception). Pressure on the body wall, whether
subgenual organs (usually located within the proximal produced by gravity or some other force, is usually de-
end of the tibiae), ]ohnston's organ (in the second an- tected by campaniforrn sensilla. Pressure on the legs
tennal segment, sensitive to movements of the anten- may be detected by subgenual organs or by sensitive
nal flagellum), and the tyrnpanal organs (involved in setae on the tarsi.
hearing). Mechanical stimuli act by displacement. The An insect detects movements of the surrounding
stimuli may come from outside the insect (for example, medium (air or water currents) chiefly by tactile setae.
touch and hearing) or from inside it (stimuli resulting lt receives information on its own movements both by
from position or movement). The mechanical stimuli mechanoreceptors and by visual cues. Movements of
initiate a series of nerve impulses, the character of air or water past an insect (whether the insect is sta-
which is deterrnined by the stimulus. In some cases, tionary and the medium is moving, or the insect is
the nerve impulses may be transmitted at frequencies moving) are detected largely by the antennae or sen-
as high as several hundred per second. sory setae on the body. In the Diptera and Hy-
The sense of touch in insects operates mainly menoptera, the antennae seem the most important de-
through hair sensilla (trichoid sensilla). The character tectors of such movements. In other insects, sensory
of the nerve impulses initiated is deterrnined by the setae on the head or neck may be the most important
rate and direction of the hair deflection. This sense is receptors. The halteres of the Diptera play an impor-
generally quite acute: Very liule hair deflection may be tant role in maintaining equilibrium in flight. They
necessary to initiate a series of neuronal impulses. move through an arc of nearly 180 degrees at rates of
Many insects show a response to gravity, for ex- up to several hundred times per second. Any change in
ample, in the surfacing of aquatic insects and in the the insect's direction strains the cuticle beca use of the
vertical constructions (burrows in the ground, combs gyroscopic property of the rapidly beating halteres and
in a beehive, and the like) some insects make. Insects is detected by the campaniforrn sensilla distributed on
generally do not have organs of equilibrium compara- the base of the haltere.

NervousSystem 33

Hearing ences (or changes) in the pitch of a sound, at least at

Theability to detect sound (vibrations in the substrate the higher frequencies. In contrast, tympanal organs
or in the surrounding medium) is developed in many are very sensitive to amplitude modulation, that is, the
insects,and sound plays a role in many types of behav- rhythmic features of the sound. These are the most im-
ior. Insects detect airborne sounds by means of two portant features of an insect's "song."
types of sense organs, hair sensilla and tympanal or-
gansoThey detect vibrations in the substrate subgenual Vision
organs. The primary visual organs of insects generally are of
Manyinsects apparently can detect sound, but en- two :ypes, the frontal ocelli (singular, ocellus) and the
tomologistsdo not always know which sensilla are in- many-faceted compound eyes.
volved.In some of the Diptera (for example, mosqui- Ocelli have a single corneal lens that is some-
toes), however, entomologists know that the setae of what elevated or domelike; beneath this lens are two
the antennae are involved in hearing (the sensilla being cell layers, the corneagenous cells and the retina
]ohnston's organ in the second antennal segment). (Figure 2-29A). The corneagenous cells, which se-
Tympanal organs are scolopophorous organs in crete the cornea, are transparent. The light-sensitive
which the sensory cells are attached to (or very near to) portion of insect photoreceptors consists of closely
tympanic membranes. The number of sensory cells in- packed microvilli on one side of the retinal cells
volvedranges from one or two (for example, in certain called the rhabdom. In the ocelli, the rhabdoms are in
moths) up to several hundred. The tympanic mem- the outer part of the retina. The basal portions of the
brane (or tympanum) is a very thin membrane with air retinal cells are often pigmented. The ocelli appar-
on both sides of it. Tympanal organs are present in cer- ently do not form focused images (the light is fo-
tain Orthoptera, Hemiptera, and Lepidoptera. The cused below the retina); they seem to be organs sen-
tympana of short-horned grasshoppers (Acrididae) are sitive mainly to differences in light intensity.
located on the sides of the first abdominal segment. The most complex light receptors in insects are
Those of katydids (Tettigoniidae) and crickets (GryIli- the compound or faceted eyes, which consist of many
dae), when present, are located at the proximal end of (up to several thousand) individual units called om-
the front tibiae (Figure 2-8D, tym). The tympana of ci- matidia (Figure 2-29C,D). Each ommatidium is an
cadas are located on the first abdominal segment (Fig- elongate group of cells capped externally by a hexago-
ure 22-45, tym). Moths may have tympana on the nal corneallens. The corneallenses are usually convex
metathorax or the base of the abdomen. externally, forming the facets of the eye. Beneath this
Vibrations in the substrate may be initiated in the corneal lens is usually a crystalline cone of four
substrate directly or may be induced (through reso- Semper cells (Figure 2-29D, cc) surrounded by two
nance) by airborne sound vibration. The detection of pigmented corneagenous cells (pgc), and beneath the
substrate vibration is mainly by subgenual organs. The crystalline cane is a group of elongate sensory cells,
frequency range to which these organs are sensitive usually eight in number, surrounded by a sheath of epi-
varies in different insects, but is mainly between about derrnal pigment cells (pgc). The striated portions of the
200 and 3000 Hz. Some insects (for example, bees) sensory cells forrn a central or axial rhabdom (rh) in
may be largely insensitive to airborne sound but can the ommatidium.
detect sound vibrations reaching them through the The pigment surrounding an ommatidium (Fig-
substrate. ure 2-29D, pgc) generally extends far enough inward
The sensory setae that detect airborne sound are so that the light reaching a rhabdom comes through
generally sensitive only to relatively low frequencies (a just the one ommatidium; the image the insects gets is
few hundred Hertz or less; rarely, a few thousand). thus a mosaic, and such an eye is spoken of as an ap-
Probably the most efficient auditory organs in insects position eye. If the pigment is located more distally in
are the tympanal organs. These are often sensitive to relation to the rhabdom, light from adjacent ommatidia
frequencies extending well into the ultrasonic range may reach a given rhabdom; this is a superposition eye.
(up to 100,000 Hz or more),2 but their discriminatory In some insects that fIy both day and night, such as
ability is targeted to amplitude modulation rather than moths, the migration of the pigment around an omma-
to frequency modulation. An insect's response (its be- tidium operates somewhat like the iris of the human
havior and the nerve impulses initiated in the auditory eye: In bright light, the pigment migrates inward, sur-
nerves) is not affected by differences in the frequencies rounding the rhabdom so that the only light reaching
of the sound as long as these frequencies are within the the rhabdom is that coming through that ommatidium
detectable range; an insect thus does not detect differ- (an apposition eye); in the dark, the pigment moves
outward, so that light from adjacent ommatidia can
2 The upper limit of hearing in humans is generally about 15,000 Hz. also reach the rhabdom (a superposition eye). The time
-- --

34 andDevelopmentof Insects
Chapter2 TheAnatomy,Physiology,

cna cna
1 I
I !

--- cc ---
pgc ¿ --


A B e D

Figure 2-29 Eye structure in insects (diagrammatic). A, Dorsal ocellus of an ant;

B, Lateral stemma of a caterpillar; C, Vertical section of part of a compound eye;
D, Ommatidium of a compound eye. bm, basement membrane; ee, crystalline cone; ena,
comea; enge, comeagenous cells; al, crystalline lens; cut, cuticle; ep, epidermis; pge,
pigment cells; ret, retina; rh, rhabdom; sne, sensory cells of retina. (B, C, Redrawn
from Snodgrass 1935, PrincipIesof Inseet Morphology,1993, Comell University Press;
D, Redrawn from Matheson 1951, Entomologyfor Introduetory Courses, 1947.)

required for this movement of the pigment varies in range is shifted to shorter wavelengths in comparison
different species. For the codling moth, it is fram 30 to to that of vertebrates. Many insects appear to be color
60 minutes. blind, but some can distinguish colors, including ul-
Many immature insects (and some adults) lack traviolet. The honey bee, for example, can distinguish
compound eyes, and in their place may be small groups blue and yellow but cannot see red. Just how an insect
of visual organs similar to ocelli in external appearance distinguishes different colors is not clear. There is ev-
(Figure 2-29B). These are called stemmata (or some- idence it may result fram different retinal cells being
times lateral oeellí). These structures are quite varied in sensitive to light of different wavelengths. Some in-
their structure, but all appparently represent highly sects (for example, the honey bee) are able to analyze
modified compound eyes (see Paulus 1979). In most polarized light. Fram the pattern of polarization in a
larvae of Holometabola, the larval eyes degenerate dur- small patch of sky, they can determine the position of
ing metamorphosis and are replaced by new adult com- the sun. The head capsule of some weevils contains a
pound eyes. region that transmits only far-red and near-infrared
The flicker-fusion frequency in insects (the rate of light. In the alfalfa weevil, Hypera postica, this ex-
flicker at which the light appears continuous) is much traocular cutoff filter apparently works in conjunc-
higher than in humans: 45 to 53 per second in humans tion with the compound eyes, enabling the insect to
and up to 250 or more in insects. This higher rate use visual cues in locating and recognizing its host
means that insects can perceive fonn even when in planto
rapid flight and that they are very sensitive to motion.
In some insects (for example, dragonflies), the omma- Other Sense Organs
tidia .of ope eye are oriented so that their axes intersect lnsects usually have a well-developed temperature
with .thbse of the other eye,allowing steroscopic vi- sense. The sense organs involved are distributed over
sion. If a dragonfly nymph is blinded in one eye, the the body but are more numerous on the antennae and
ny¡rtpli cannot judge the position of its prey very accu- legs. Some insects also have a well-developed humidity
rately. sense. The sensilla involved in these senses are quite
The range in wavelength to which insect eyes are diverse in structure, and in many cases the relationship
sensitive is fram about 2540 to 6000 Á, compared between the observed structure and function is not
with about 4500 to 7000 Á in humans. lnsects' visual well understood (see Altner and Loftus 1985).
Reproductive Systems 35

vitelIogenesis, accessory reproductive gland activity,

EndocrineSystem pheromone production, and sexual behavior (see
Raabe 1986).
Severalorgans in an insect are known to produce hor- Substances chemicalIy related to ecdysone and ju-
mones,the principal functions of which are the control venile hormone occur in certain plants and may pro-
of the reproductive processes, molting, and metamor- tect the plants from feeding by insects. Chemical
phosis. Chemicals similar to hormones of vertebrates, analogs of ecdysone and juvenile hormone are being
includingandrogens, estrogens, and insulin, have been studied to see whether they can function as new kinds
detectedin insects, but their function is yet unknown. of insecticides.
The neurosecretory celIs in the brain are neurons
thatproduce one or more hormones that playa role in
growth, metamorphosis, and reproductive activities.
One of these, commonly calIed the brain honnone or
Reproductive Systems
prothoracicotropichonnone (PTTH), plays an important
role in molting by stimulating a pair of glands in the
Internal Reproductive Systems
prothorax to produce the hormone ecdysone that
causes apolysis. Other hormones produced by the The internal reproductive system of the female (Fig-
brain may have other functions. For example, ento- ure 2-31A) consists of a pair of ovaries (ovy), a system
mologistsbelieve that a brain hormone plays a role in of ducts through which the eggs pass to the outside,
caste determination in termites and in breaking dia- and associated glands. Each ovary generalIy consists of
pausein some insects. a group of ovario les (ov1). These lead into the lateral
Ecdysone (Figure 2-30A) initiates growth and de- oviduct posteriorly (ovd) and unite anteriorly in a sus-
velopmentand causes apolysis. This hormone occurs pensory ligament (51)that usualIy attaches to the body
in all insects groups that have been studied, in crus- walI or to the dorsal diaphragm. The number of ovari-
taceans,and in arachnids, and it is probably the molt- oles per ovary varies from 1 to 200 or more, but it is
inghormone of alI arthropods. It also plays a role in the usualIy in the range of 4 to 8. Oogonia (the primary
differentiationof the ovarioles and accessory reproduc- germ celIs) are located in the anterior apical portion of
tiveglands in females and in several steps in the pro- the ovariole, the germarium. The oogonia undergo mi-
cessof egg production (oogenesis). Ecdysone, in fact, tosis, giving rise to the oocytes and trophocytes (or
is alsoproduced within the ovaries of insects. nurse celIs; entomologists do not know what mecha-
Thecorpora alIata (Figure 2-26, ca) produce a hor- nism determines which daughter celIs become oocytes
monecalled the juvenile honnone üH) (Figure 2-30B), and which become trophocytes). Ovarioles in which
theeffectof which is the inhibition of metamorphosis. trophocytes are produced are calIed meroistic ovarioles;
Various substances, particularly terpenes such as no trophocytes are produced in panoistic ovario les.
famesol,show considerable juvenile hormonelike ac- The oocytes pass down through the ovario les, matur-
tivity.JH also has effects on other processes besides ing as they go. Thus the spatial sequence in the ovari-
the inhibition of metamorphosis. It is involved in ole reflects the temporal sequence of oocyte matura-



o CH2-CH3 CH3
/ \ I I



CH3 2-30
COOCH3 Figure Structure of two insect hor-
B mones. A, Ecdysone; B,Juvenile hormone.
l. --

36 Chapter2 TheAnatomy,Physiology,and Developmentof Insects

tion. The trophocytes may be connected to the oocyte haploid number of chromosomes. In the lower portion
by cytoplasmic filaments and may remain in the ger- of the ovariole, a vitelline membrane forms around the
marium (telotrophic ovario les) or pass down the ovar- oocyte, and the follicular epithelium secretes the
iole with each oocyte (in polytrophic ovario les). The chorion (or eggshell) around the mature oocyte.
trophocytes are important in passing ribosomes and In many insects, all or most of the oocytes mature
RNA to the oocyte. An oocyte, the surrounding epithe- before any are laid, and the egg-swollen ovaries may
lium, and trophocytes (in polytrophic ovario les) to- occupy a large part of the body cavity and may even
gether form a folliele. Yolk proteins (vitellogenins) are distend the abdomen. The two lateral oviducts usually
synthesized outside the ovariole and transported into unite posteriorly to form a single common (or median)
the oocyte by the follicular epithelium. In this region of oviduct, which enlarges posteriorly iOlo a genital
the ovariole Cthe vitellarium), the oocytes greatly in- chamber or vagina. The vagina extends to the outside,
crease in size owing to the deposition of yolk Cthe pro- the opening being called either the ovipore (in reference
cess of vitellogenesis). Yolk consists of protein bodies to the opening through which the eggs are laid) or
(largely derived from hemolymph proteins), lipid vulva (the copulatory opening). Because the vagina
droplets, and glycogen. Many insects harbor microor- usually also receives the male genitalia during copula-
ganisms in their bodies, and in some cases these may tion, it is sometimes known as the bursa copulatrix. As-
get into the egg during its developmeOl, usually sociated with the vagina are usually a saelike structure
through the folliele cells. The maturation divisions of called the spermatheca, in which spermatozoa are
the oocyte may occur at about the end of vitellogenesis stored, and often various accessory glands, which may
or even after insemination, resulting in eggs with the secrete adhesive material to fasten the eggs to some ob-



Figure 2-31 Reproductive systems of insects. A, Female reproductive system; B, Male

reproductive system. acg, accessory gland; aed, aedeagus; covd, common oviduct; ejd,
ejaculatory duct; ovd, oviduct; ovl, ovariole; ovy, ovary; sI, suspensory ligament; smv, sem-
inal vesicle; spt, sperm tube; spth, spermatheca; spthg, spermathecal gland; tst, testis; vag,
genital chamber or vagina; vd, vas deferens; ve, vas efferens. (Redrawn from Snodgrass
1935, PrincipIesoflnsect Morphology,1993, Comell University Press.)
Reproductive Systems 37

ject orprovidematerialthat covers the eggmass with a duct (ejd), which opens to the outside on a penis or an
protectivecoating. aedeagus (aed). Some insects have an enlargement of
In many Lepidoptera (the Ditrysia), there are two or a lateral diverticulum fram each vas deferens in
openings to the reproductive tract of the female. The which spermatozoa are stored. These are called semi-
commonoviduct leads to the vagina and the ovipore to nal vesicles(smv). The accessory glands (acg) secrete
the outside on segment 9. During copulation, however, fluids that serve as a carrier for the spermatozoa or
the male places his genitalia and deposits the sper- that harden about them to form a sperm-containing
matophore in a separate opening, the vulva, on seg- capsule, the spermatophore.
ment8. The vulva leads to the bursa copulatrix, which The sperm begin their development in the distal
is connected to the vagina and thence to the sperma- (anterior) ends of the sperm follieles of the testes and
thecabya spermduct, through which the spermatozoa continue development as they pass toward the vas ef-
mustmove. ferens. The processes of spermatogenesis (production
Egg development is usually complete about the of haploid germ cells fram diploid spermatogonia) is
timethe adult stage is reached, but in some cases it is usually completed by the time the insect reaches the
completed later. In those aphids in which the female adult stage or very shortly thereafter.
gives birth to living young parthenogenetically, the The spermatozoa of insects come in an almost be-
eggsare matured and development begins before the wildering variety of shapes and sizes, often differing
adultstage is reached. In the beetle genus Micromalthus, quite strikingly fram the typical tadpole-shaped cells
the eggs are matured and begin their development one thinks of (see jamieson 1987). One notable char-
(withoutfertilization) in the ovaries of the larvae, and acteristic is that the flagellum, or axoneme, is made up
arepassed to the outside (as either eggs or larva e) be- of the typical 9 + 2 arrangement of microtubules char-
forethe mother becomes an adult. In the cecidomyiid acteristic of flagella and ciba, but in addition has an
genusMiastor, egg development is also completed in outer ring of 9 single microtubules that are derived
the larval stage, but in this case the young larvae from the ring of 9 doublets. This 9 + 9 + 2 arrange-
(whichhave developed parthenogenetically) break out ment is characteristic of hexapods as a whole. In addi-
of the ovaries inlo the body cavity and develop there. tion to the more familiar threadlike or tadpolelike cells,
Theyeventually rupture the cutiele of the mother larva some hexapods have spermatozoa with two flagella in-
and escape to the outside. Reproduction by a preadult stead of one, cells in which the axoneme is "encysted"
stageis called paedogenesis. so that they are immobile, and, in some Protura, even
Eggproduction appears to be controlled in many simple disk-shaped immobile cells. Among species of
insectsby one or more hormones from the corpora al- Drosophila, the spermatozoa range in length from
lata,incIuding juvenile hormone (Figure 2-30B), that about 55 ¡Lmto 15 mm (the total body length of the fa-
act by controlling the initial stages of oogenesis and miliar D. melanogaster is less than 5 mm)!
yolkdeposition. Removal of the corpora allata prevents
nonnal egg formation, and their reimplantation (from External Genitalia
either a male or a female) induces ovarian activity
again.The corpora allata have nerve connections with The external genitalia of most insects are generally
thebrain, and nerve impulses affect their activity. Re- thought to be derived from appendages of abdominal
searchersalso believe that (at least in some cases) neu- segments 8, 9, and possibly 10. The male genitalia are
rosecretorycells in the brain may produce a hormone primarily organs involved with copulation and the
thataffects the activity of the corpora allata. Many ex- transfer of sperm to the female. The female genitalia
ternalfactors (for example, photoperiod and tempera- are involved in depositing the eggs on or in a suitable
ture)affect egg production, and these factors probably substrate. These structures are called external genitalia
actthroughthe corpora allata. even though they may be retracted within the apical
Thereproductive system of the male (Figure 2-31B) abdominal segments when not in use and are often (es-
issimilarin general arrangement to that of the female. pecially in the male) not visible without dissection.
It consistsof a pair of gonads, the testes, ducts to the The appendicular ovipositor of pterygote insects is
outside,and accessory glands. Each testis (tst) con- believed to have evolved from a structure similar to
sistsof a group of sperm tubes (spt) or follieles sur- that now found in the female genitalia in the Thysa-
rounded by a peritoneal sheath. Each sperm folliele nura (Figure 2-32A). This consists of an ovipositor,
opensinto a short connecting tube, the vas efferens which is formed fram the appendages (gonopods) of
(ve;plural, vasa efferentia), and these connect lo a segments 8 and 9. The first gonocoxa (the first valvifer,
singlevas deferens (vd; plural, vasa deferentia) on from segment 8, gcx¡) articulates dorsally with tergum 8;
eachside of the animal. The two vasa deferentia usu- the second gonocoxa (second valvifer, from segment 9,
allyunite posteriorly to form a median ejaculatory gcx2) articulates with tergum 9. Laterally, the gono-
38 Chapter2 TheAnatomy.Physiology.and Developmentof Insects



I gap,
A gcx,

Figure2-32 Ovipositor of insects. A, Ovipositor of Thysanura, ventral view; B, Ovipos-

itor of a leafhopper, lateral view with parts spread out; C, Secondary ovipositor of
Mecoptera, lateral view.atb, anal tube; gcx" first gonocoxa; gcx2,second gonocoxa; gap"
first gonapophysis; gap2,second gonapophysis; gpl, gonoplac; gst¡, first gonostylus; gst2,
second gonostylus. (A, C, Redrawn from Snodgrass 1935, Principies oIInsect Morphology,
1993, Comell University Press.)

coxae bear styli, the gonostyli; these are presumably substrate in order to oviposit. In the Orthoptera, how-
serial homologs of the styli on the pregenital segments ever, the gonoplacs are cutting or digging structures,
and thus represent the derived telopods of the primi- taking over the function of the second gonapophyses,
tive abdominal appendages. Medially each gonocoxa which are smaller and function as egg guides.
bears an elongate process known as a gonapophysis There are a number of modifications of this basic
(also called a valvula); the second gonapophyses (gap2, appendicular ovipositor structure within the ptery-
segment 9) lie above the first gonapophyses (gapi, seg- gotes, but the most generalized condition is found in
ment 8), and together these form the shaft of the some Odonata, Hemiptera (Auchenorrhyncha), Or-
ovipositor (ovp). Coordination of the movement of thoptera, and Hymenoptera. In many Holometabola,
these four elongate structures is achieved by two mech- the appendicular components of the ovipositor are very
anisms. First, the two gonapophyses on each side are much smaller and are not involved in oviposition. In-
connected by a tongue-in-groove mechanism known as stead, the terminal abdominal segments form a tele-
an olistheter. In addition, a small scIerite on each side, scoping tube, called the pseudovipositor or oviscapt, which
the gonangulum, articulates with the second gonocoxa the female extends when ovipositing (Figure 2-32C). In
(from which it is derived), the first gonapophysis, and some cases, such as tephritid flies, this type of oviposi-
tergum 9. Again, this interconnects the movements of tor bears apical cutting plates, enabling the female to
the first and second gonapophyses on each side. place her eggs deep within a suitable substrate.
In the pterygote insects that retain an appendicular The external genitalia of male insects show such
ovipositor, the first gonostylus is lost, and the second incredible diversity that it has been difficult for ento-
gonocoxa is elongate (perhaps incorporating remnants mologists to infer the primitive structures from which
of the second gonostylus) to form a sheathlike outer they evolved and to homologize the parts in different
covering for the ovipositor shaft, the gonoplacs (also orders. The genitalia of the Thysanura and Microco-
known as the third valvulae, Figure 2-328, gpl). In most ryphia are generally similar to that of the females,
insects, the gonoplacs serve both a protective and a sen- but with an additional median penis derived from
sory function and are not involved in penetrating the segment 10 (Figure 2-33A). However, the mal e geni-
Development and Metamorphosis 39

Development and Metamorphosis

Sex Determination
The chromosomes of insects (as well as other animals)
usually occur in pairs, but in one sex the members of
one pair do not match or are represented by one chro-
mosome only. The chromosomes of this odd pair are
,/, called sex chromosomes; those of the other pairs, auto-
somes. In most insects, the males has just one X (sex)
chromosome (and is called heterogametic) and the fe-
A B male has two (homogametic). The male condition is gen-
erally referred to as XO (if only one chromosome in this
pair is present) or XY Cthe y chromosome being differ-
Figure2-33 A, Male genitalia of Machilidae, ventral ent in size or shape from the X chromosome), and the fe-
view;B, External genitalia of pterygotes (diagrammatic). male as XX CtwoX chromosomes). One major exception
ard,aedeagus; gap2' second gonapophysis; pmr, paramere; to this generalization is in the Lepidoptera; in most
phtr,phallotreme. (A, Redrawn from Snodgrass 1935, species in this order, it is the female that is heteroga-
PrincipIes01[nsectMorphology, 1993, Comell University metic (the WZJZZ system, females are WZ, males ZZ).
Press;B,Redrawnfrom Snodgrass 1951.) The autosomes appear to contain genes for
"maleness," whíle the X chromosomes contain genes
for "femaleness." More accurately, it seems the sex is
determined by the balance between these two groups
taliaof silverfish and bristletails are not involved with of genes. With two autosomes of each pair and only
eopulation.In these insects as well as the entognathous one X chromosome, the genes for maleness predomi-
hexapods,sperm transfer is indirect: The male places nate and the animal becomes a maleo With two auto-
hisspermatophore or a sperm droplet on the substrate, somes of each pair and two X chromosomes, the
andthe female actively places it in her gonopore. The genes for femaleness predominate and the animal be-
penisof lepismatids is used to spin a silken web on comes a female.
whichthespermatophore is placed. Sex is determined a httle differently in the Hy-
Thereis considerable debate conceming the origin menoptera and a few other insects. In these insects, the
. ofthemale genitalia in pterygotes. Some entomologists males are generally haploid (only very rarely diploid),
eontendthat they are derived from the appendages of and the females are diploid. The males develop from
segment9 alone, and some include both these ap- unfertilized eggs and the females develop from fertíl-
pendagesand the penis of segment 10 (as seen in ized eggs (a type of parthenogenesis called arrheno-
maehiloidsand lepismatoids). Snodgrass (1957) held toky). Just how a haploid condition produces a male
thatthe genitalia are derived from outgrowths of ster- and a diploid condition produces a female is less well
num10. In very general terms, the genitalia consist of understood, but geneticists beheve that sex in these in-
outerclasping organs and a median intromittant organ sects depends on a series of multiple alleles (Xa, Xb, Xc,
(Figure2-33B). The outer claspers, or parameres and so on): haploids and homozygous diploids (Xa/Xa,
(pmr),may arise from a common base, the gonobase or Xb/Xb, Xc/Xc, and so on) are males, whíle heterozygous
basal ring (gb). The median intromittant organ is the diploids (Xa/Xb, Xc/Xd, and so on) only are females.
aedeagus(aed). The opening of the aedeagus through Parthenogenetic development producing females
whichthe spermatophore or the semen passes is the occurs in many insects (this type is called thelytoky). In
phallotreme(phtr). In many species, the ejaculatory some of these species, males are relatively rare or are
duetisevertedthrough the phallotreme during copula- unknown. These insects usually have the XO or XY
lion;this eversible lining is called the endophallus. Such male and XX female sex-determining mechanism,
diversityexists in the structure and nomenclature of which means that either the eggs faíl to undergo meio-
themale genitalia that it is beyond the scope of this sis and are diploid or they do undergo meiosis and two
bookto describe them in more detail (see Tuxen 1970 cleavage nuclei fuse to restore the diploid condition.
foranorder-by-order account of genitalic structure and Some insects (for example, gall wasps and aphids) pro-
thenomenclature used). This diversity of structure is duce both males and females parthenogenetically (at
veryuseful in identifying many groups of insects at the certain seasons). The production of a male apparently
specieslevel; such identification usually requires dis- involves the 1055of an X chromosome, and the pro-
seetionand mounting of the genitalia for closer study. duction of a female involves either a fusion of two
40 Chapter 2 The Anatomy, Physiology, and Development of Insects

cleavage nuclei to restore the diploid condition or enters the egg and undergoes cleavage to produce hap-
diploid eggs arising from tetraploid germ tissue. loid (male) tissue in an otherwise female individual.
Recent research has revealed the widespread pres- lndividuals with a sexual condition intermediate be-
ence of the bacterium Wohlbachia in a great diversity of tween maleness and femaleness are called intersexes.
insects. In some cases, the bacterium kills embryos that These usually result from gene tic imbalance, particu-
would develop into males. lady in polyploids (for example, a triploid Drosophila
Individual insects sometimes develop with aber- with an XXY sex chromosome content is an intersex
rant sex characters. lndividuals having some male tis- and is sterile).
sues and some female tissues are called gynandro-
morphs; such individuals sometimes occur in the
Hymenoptera and Lepidoptera. In the Hymenoptera,
where the sex-determining mechanism is haploidy = The eggs of different insects vary greatly in appearance
male and diploidy = female, a gynandromorph may (Figures 2-34 through 2-36). Most eggs are spherical,
develop from a binucleate egg in which only one of the oval, or elongate (Figure 2-34B,C,G), but some are
nuclei is fertilized or may develop when an extra sperm barrel shaped (Figure 2-35), some are disk shaped, and

Figure 2-34 lnsect eggs. A, Fall army-
worm, Spodopterafrugiperda O. E. Smith);
B, Grape leaf-folder, Desmiafuneralis
(Hübner); C, Southern corn rootworm,
Diabroticaundecimpunctata howardi Barber;
D, Horse bot fly,Gasterophilusintestinalis
(De Geer); E, Snowy tree cricket, Oecanthus
fultoni Walker; F,Anopheles mosquito;
G, Seedcorn maggot, Hylemya platura
(Meigen); H, Culex mosquito, egg raft;
1, Lacewing, Chrysopa sp.;], Fall canker-
worm, Alsophila pometaria (Harris). (A-C,
1, courtesy of USDA;], courtesy of Ohio
Agricultural Research and Development
Development and Metamorphosis 41

.. $c::

- EQ)

~ ,.
,. -.. a;>
.( .J( '"
J( c::

Figure2-35 Eggs of a stink bug.

so on. The egg is covered with a shell that varies in

thickness,sculpturing, and color: Many eggs are pro-
videdwith characteristic ridges, spines, or other pro-
cesses,and some are brightly colored.
Most insect eggs are laid in a situation where they
areafforded some protection or where the young, on
hatching,will have suitable conditions for develop- Figure2-36 Egg of a cIearwing moth (Sesiidae). A, Egg
ment.Many insects enclose their eggs in some sort of of Podosesiasyringae (Harris), 50 X; B, Same, showing de-
protectivematerial. Cockroaches, mantids, and other tail of egg surface, 290 x. (Courtesy of the Ohio Agricul-
insectsenclose their eggs in an egg case or capsule. The tural Research and Development Center.)
tentcaterpillars cover their eggs with a shellaclike ma-
terial.The gypsy moth lays its eggs in a mass of its
bodyhairs. Grasshoppers, june beetles, and other in-
sectslay their eggs in the ground. Tree crickets insert
theireggsin plant tissues (Figure 2-34E). Most plant- the ground and pupates. Thus the only active feeding
feedinginsects lay their eggs on the food plant of the stage is the adult parasitic fly.
young.Insects whose immature stages are aquatic usu-
allylay their eggs in or near water, often attaching
Embryonic Development
themto objects in the water. Parasitic insects usually
laytheir eggs in or on the body of the host. Some The egg of an insect is a cell with two outer coverings,
insectsdeposit their eggs singly, whereas others lay a thin vitelline membrane surrounding the cytoplasm
their eggs in characteristic groups or masses (Fig- and an outer chorion. The chorion, which is the hard
ures2-34H,j, and 2-35). The number laid varies from outer shell of the egg, has a minute pore or set of pores
onein certain aphids to many thousands in some of the (the micropyle) at one end, through which sperm en-
socialinsects, but most insects lay from 50 to a few ter the egg (Figure 2-37A). just inside the vitelline
hundredeggs. membrane is a layer of cortical cytoplasm. The central
Most insects are oviparous; that is, the young portion of the egg, inside the cortical cytoplasm, is
hatchfram the eggs after they have been laido In a few largely yolk.
insects,the eggs develop within the body of the female, Most insect eggs undergo what is termed superfi-
andliving young are deposited. The extreme in this cial cleavage. The early cleavages involve only the nu-
caseis seen in the sheep ked, for example: The female cleus, giving rise to daughter nuclei scattered through
flyretainsthe egg and larva within her body for an ex- the cytoplasm (Figure 2-37B). Eventually these nuclei
tendedperiod of time. When parturition ("birth") 6- migrate to the periphery of the egg (to the layer of cor-
nallyoccurs, the larva almost immediately burrows in tical cytoplasm). After nuclear migration, the periph-

42 Chapter2 TheAnatomy.Physiology.and Developmentof Insects

cpl- -
nu, á1T!F.~~~
\ 0:r~:.~~~~~.
.'. :",
\-, .(...
o:.Y.. 0";:,('.'~:
. .'

?e: --vm bl
B .....

- -yc
.?¡t;;~ " cpl



Figure2-37 A, Diagram of a typical insect egg; 8, Early cleavage; C, Peripheral blasto-

derm layer formed. bl, blastoderm; cho, chorion; cnu, cleavage nuclei; cpl, cortical cyto-
plasm; gel, germ cells; mcp, micropyle; nu, nucleus; vm, vitelline membrane; yc, yolk cells;
yo, yolk. (Redrawn from Snodgrass 1935, PrincipIesoflnsect Morphology. 1993, Comell
University Press.)

eral cytoplasm becomes subdivided into cells, usually germ layers--ectoderm, mesoderm, and endoderm-
each with one nucleus, forming a celllayer, the blasto- the various organs and tissues of the insect develop: the
derm (Figure 2-37C, bl). This is the blastula stage. ectoderm gives rise to the body waIl, tracheal system,
Within the blastoderm, in the mass of yolk material, nervous system, Malpighian tubules, and anterior and
are a few cells that do not take pan in forming the blas- posterior ends of the alimentary tract; the mesoderm
toderm; these consist mainly of yolk cells. gives rise to the muscular system, heart, and gonads;
The blastoderm cells on the ventral side of the egg the endoderm develops into the midgut.3
enlarge and thicken, forming a germ band or ventral The alimentary tract is formed by invaginations
plate that will eventually form the embryo. The re- from each end of the embryo, which extend to and
maining cells of the blastoderm become the serosa and unite with the primitive midgut (Figure 2-39). The an-
(later) the amnion. The germ band differentiates into a terior invagination becomes the foregut, the posterior
median area or middle plate and two lateral areas, the invagination becomes the hindgut, and the central part
lateral plates (Figure 2-38A). The gastrula stage begins (lined with endoderm) becomes the midgut. The cells
when the mesoderm is formed from the middle in lining the foregut and hindgut are ectodermal in origin
one of three ways: by an invagination of this plate (fig- and secrete cuticle.
ure 2-38B,C), by the lateral plates growing over it (Fig- Body segmentation becomes evident fairly early in
ure 2-38D,E), or by a proliferation of ceIls from its in- embryonic development, appearing first in the anterior
ner surface (Figure 2-38F). CeIls proliferate from each pan of the body. It involves ectoderm and mesoderm,
end of the mesoderm and eventuaIly grow around the
yolk. These cells represent the beginnings of the endo-
derm, and they form the lining of what wiIl be the 3Note that these embryonic tissues, in particular, the so-called endo-
midgut of the insect (Figure 2-39). From the three derm, may not be homologous to that of deuterostomes.

Development and Metamorphosis 43

Figure2-38 Cross-section diagrams show-

ing mesoderm formation in insects. A, Germ
- , -. L. -....ect band differentiated into middle and lateral
A B e plates; B, C, Stages in mesoderm formation
by invagination of middle plate; D, E, Stages
in mesoderm formation by lateral plates
growing over middle plate; F, Mesoderm for-
mation by internal proliferation from middle
plate. bl, blastoderm; eet, ectoderm; mdp,
middle plate; LP, lateral plate; msd, meso-
, - ---., - -, "- derm. (Redrawn from Snodgrass 1935, Prin-
LP ciples of Inseet Morphology, 1993, Cornell
'mdp 'ect 'ect
O E F University Press.)

but not endoderm, and is reflected in the segmental I
arrangement of the struetures developing from these
germlayers (nervous system, heart, tracheal system,
and appendages). The appendages appear soon after
segmentationbecomes evident. Typically, each segment
beginsto develop a pair of appendages, but most of
theseare resorbed and do not develop further.
Ihe molecular mechanisms governing the pattern
.,.,. f'..,<.? o c~ Q C)rJ o 0::..~
ofdevelopment in embryos is a particulariy active area
ofresearch.The initial orientation of the embryo is me- f.<jl)}.;/:r.:\?(H~;'{2}
diatedby proteins derived from maternal RNA in the
~~.<;>.o~)~)(?'¡?S-:(~ . ' endr
egg.Subsequently, a hierarchy of genes are transcribed B
thatdefine the segmentation pattern in the embryo. Of
particular interest are the homeobox genes, or HOX
genes.In the insects that have been studied, there are
eightHOXgenes. In Drosophila, these are alllocated on
chromosome3 and oceur in the order lab, pb, dfd, ser,
antp,ubx, abd-A and abd-B. These abbreviations stand e
forlabial,proboscidea, deformed, sex combs reduced, an- "-
tennapedia,ultrabithorax, abdomen-A, and abdomen-B.
These genesare differentiallyexpressed in the embryo,
bothinspaceand time, and are cruciaI to defining seg-
mentsand in the subsequent differentiation of limbs.
Theymay well have further developmentaI roles that /
o \mg
havenot yet been cleariy defined. The interest, from an
evolutionarypoint of view, arises beca use very similar
genesare found in distantly related phyla, implying an Figure2-39 Diagrams showing the formation of the ali-
ancientevolutionary origino Further, the expression of mentary canal. A, Early stage in which the endoderm is
thesegenes and their enhaneers have provided further represented by rudiments; B, C, Development of endo-
testsof hypotheses concerning the homology of struc- derm around yoIk; D, CompIetion of alimentary canal. ans,
tures,for example, the mandibles in Crustacea, Myri- anus; eet, ectoderm; mdr, endodermal rudiments; mg,
apoda,and Hexapoda. midgut; mo, mouth; ms, mesoderm; prct, proctodaeum; std,
At some time eariy in its development, the embryo stomodaeum or foregut;yo, yoIk. (Redrawn from Snodgrass
becomessurrounded by two membranes, an inner am- 1935, Principies of Insect Morphology, 1993, Cornell Uni-
nionand an outer serosa. Later it aequires a cuticular versity Press.)

44 Chapter2 TheAnatomy,Physiology,and Developmentof Insects

membrane secreted by the epidermis. The formation of is rigid, it cannot expand very mucho Therefore, as the
the amnion and serosa sometimes involves a reversal of insect grows or increases in size, the exoskeleton must
position of the embryo in the egg; the embryo turns taíl be periodically shed and replaced with a larger one.
first into the yolk, away from the blastoderm. This The process of digesting portions of the old cuticle and
turning carries part of the extraembryonic blastoderm synthesizing the new cuticle is called molting (also
into the yolk, and when the turning is complete, the spelled moulting), which culminates in the shedding of
opening into the embryonic cavity is closed. The ex- the old cuticle (ecdysis).
traembryonic blastoderm thus forms a lining (the am- The molt involves not only the cuticle of the body
nion) around the embryonic cavity, and the outer part 'vall but also the cuticular linings of the tracheae,
of the blastoderm, which surrounds the egg, becomes foregut, and hindgut and the endoskeletal structures.
the serosa. The embryo later returns to its original po- The tracheal linings usually remain attached to the
sition on the ventral side of the egg. In other cases the body wall when it is shed. The linings of the foregut
amnion and serosa are formed by folds of the blasto- and hindgut usually break up, and the pieces are
derm, which grow out from the edge of the germ band passed out through the anus. The tentorium usually
and unite beneath it. These membranes usually disap- breaks into four pieces, which are withdrawn through
pear before the embryo is ready to leave the egg. Cu- the tentorial pits during the molt. The cast skins, called
ticular coverings of the embryo (sometimes called exuviae, often retain the shape of the insects from
pronymphal membranes) occur in insects with simple which they were shed.
metamorphosis and in a few with complete metamor- The initial stages in the molting cycle are triggered
phosis. These are shed, by a process akin to molting, by the release of PTTH (brain hormone) from neurose-
before or very shortly after hatching. cretory cells in the brain. This stimulates the protho-
A young insect may escape from the egg in various racic glands (also sometimes called the molting glands)
ways. Most insects with mandibulate mouthparts chew to release ecdysone into the hemolymph. Ecdysone, in
their way out of the egg. Many insects have what are turn, stimulates the separation of the old cuticle from
called egg-bursters: spinelike, knifelike, or sawlike pro- the under1ying epidermis, a process known as apolysis.
cesses on the dorsal side of the head-which are used The epidermis undergoes mitosis and grows in size; af-
in breaking through the eggshell. The eggshell is some- ter this, the new cuticle is produced. Molting fluid se-
times broken along weakened lines, either by the wrig- creted from the epidermal cells contains enzymes that
gling of the insect within or by the insect taking in air digest the old endocuticle (but do not affect the epicu-
and rupturing the shell by internal pressure. The ticle or exocuticle), and as a new cuticle is being de-
hatching from the egg is called eclosion. posited, the digestive products are resorbed into the
Polyembryony is the development of two or more body. Once this new exoskeleton is complete, the in-
embryos from a single egg. It occurs in some of the par- sect is ready to shed or break out of the old one. Ecdy-
asitic Hymenoptera and in the Strepsiptera. In the em- sis is triggered by a molting hormone, and begins with
bryonic development of such an insect, the dividing nu- a splitting of the old cuticle along lines of weakness,
cleus forms cell clusters, each of which develops into an usually in the midline of the dorsal side of the thorax.
embryo. The number of embryos that grow to maturity The rupturing force is pressure of the hemolymph (and
in a given host depends on the relative sizes of the par- sometimes air or water), forced into the thorax by con-
asite larvae and the host. In some cases, there are more traction of the abdominal muscles. This split in the
parasite larvae than the food supply (the body contents thorax grows, and the insect eventually wriggles its
of the host) can support, and some of them die and may way out of the old cuticle.
be eaten by the surviving larvae. The number of young When it first emerges from the old cuticle, the in-
from a single egg varies in Maaocentrus (Braconidae) sect is pale in color, and its cuticle is 50ft. Within an
from 16 to 24, but in M. ancylivorus Rohwer only one hour or two, the exocuticle begins to harden and
parasite larva leaves the host. In Platygaster (Platy- darken. During this brief period, the insect enlarges to
gastridae) from 2 to 18 larvae develop from a single egg, the size of that instar, usually by taking in air or water.
and in Aphelopus (Dryinidae) from 40 to 60 develop The wings (if present) are expanded by forcing he-
from a single egg; in some of the Encyrtidae, more than molymph into their veins. The alimentary tract often
1500 young develop from a single egg. serves as a reservoir of the air used in this expansion:
If the crop of a cockroach, for example, is punctured
with a needle, the insect does not expand but collapses;
Postembryonic Growth
if the wing tips of an emerging dragonfly are cut off,
Having an exoskeleton presents a problem as far as hemolymph escapes from the cut end and the wings
growth is concerned. To function as an exoskeleton, faíl to expand. In addition to allowing the cuticle to ex-
r." , the insect's body wall must be.relatively rigid, but if it pand, this period between ecdysis and hardening of the
..!\'. ,¡P..' .'"

Development and Metamorphosis 45

cuticleallows insects that pupa te in the soil, for exam- enced by a number of environmental conditions. In
pIe,to crawl to the surface, there to expand the cuticle. many insects, however, the increase generally follows a
Insomespecies,researchershave identified a proteina- geometric progression. The increase in the width of the
ceoushormone, bursicon, that controls the process of larval head capsule in Lepidoptera, for example, is of-
sclerotization. ten a factor of 1.2 to 1.4 at each molt (Dyar's rule). In
Ihe number of molts varies among most insects species where the individual molts are not actually ob-
fram4 to 8, but some of the Odonata undergo 10 or 12 served, Dyar's rule can sometimes be applied to head
molts,and some of the Ephemeroptera may undergo as capsule measurements of a series of different-sized lar-
manyas 28 molts. A few hexapods, such as the ento- vae to estimate the number of instars.
gnathousorders, silverfish, and bristletails, continue to
moltafter reaching the adult stage, but winged insects
neithermolt nor increase in size once the adult stage is
reached.(Mayfiies have a sexually immature winged Most insects change in form during postembryonic de-
instarpreceding the adult, the subimago, that molts.) velopment, and the different instars are not all alike.
Ihe stage of the insect between ecdyses is generally This change is called metamorphosis. Some insects un-
calledan instar.The first instar is between hatching and dergo very little change in form, and the young and
thefirstlarval or nymphal molt; the second instar is be- adults are very similar except for size (Figures 2-40 and
tweenthe first and second molts; and so on. However, 2-41). In other cases the young and adults are quite dif-
thefull process of molting is not instantaneous. There ferent, in habits as well as in form (Figure 2-42).
is a period of time, usually short, but sometimes very There is quite a bit of variation in the metamor-
long,between apolysis and ecdysis during which the phosis occurring in different insect groups, but these
nextinstar of the insect is "hidden" within the old cuti- variations can be roughly grouped into two general
eIe.Hinton (1971) suggested that the term instar be types: simple metamorphosis and complete metamor-
usedto refer to period of time from one apolysis to the phosis. In simple metamorphosis, the wings (if any)
next,and he proposed the term pharate instar to refer to develop externally during the immature stages, and
theinsectduring the time between apolysis and ecdysis. there is ordinarily no quiescent stage preceding the last
Inmanycases this time period is sufficiently short that molt (Figures 2-40 and 2-41). In complete metamor-
littleconfusion arises concerning which event signals phosis, the wings (if any) develop internally during the
the end of one instar and the beginning of the next. immature stages, and a quiescent or pupal stage nor-
However,in some, such as the cyclorrhaphous Diptera, mally precedes the last molt (Figure'2-42). The pupal
thedistinction is important. In these fiies, larval-pupal stage is quiescent in that the insect at this time ordi-
apolysisis not followed by an immediate ecdysis. In- narily does not move around, but a very considerable
stead,the last larval cuticle is hardened to form a sort of amount of change (lo the adult) is taking place in this
cocoon withinwhich lies the pharate pupa. Full devel- stage.
opmentof the pupa is followed by the pupal-adult The changes during metamorphosis are accom-
apolysis.The adult cuticle is then formed, and at that plished by two processes, histolysis and histogenesis.
pointecdysis occurs with the adult fiy shedding both Histolysis is a process whereby larval structures break
thelastlarval and pupal cuticle at the same time. down into material that can be used in developing
Ihe increase in size at each molt varies in different adult structures. Histogenesis is the process of devel-
speciesand in different body parts and can be infiu- oping the adult structures from the products of histol-

Figure 2-40 Stages in the

development of the straw-
berry aphid, Chaetosiphonfra-
gaefolii (CockereIl). A, First
instar; B, Second instar;
C, Third instar; D, Fourth
instar; E, Adult female.
(Courtesy of Baerg and
Arkansas AgriculturaI
Experiment Station.)

A B e D E

46 Chapter2 TheAnatomy,Physiology,and Developmentof Insects



Figure 2-41 Stages in the development of the grass bug, Arhyssus sidae (Fabricius).

A, Egg; B, First instar; C, Second instar; D, Third instar; E, Fourth instar; F, Fifth instar;
G, Adult female. (Courtesy of Readio 1928 and the Entomological Society of America.)

ysis. The chief source s of material for histogenesis are ders (Protura, Collembola, Diplura, Microcoryphia, and
the hemolymph, fat body, and histolyzed tissues such Thysanura) and in most wingless members of the other
as larval muscles. Ectodermal structures, such as wings orders with simple metamorphosis. In hemimetabolous
and legs, develop beneath the larval cuticle as epider- metamorphosis (with "incomplete" metamorphosis),
mal thickenings called imaginal discs. These tissues re- the nymphs are aquatic and gill-breathing and differ
spond quite differently from other larval tissues to the considerably from the adults in appearance. This type of
hormonal milieu of the insect. In the late larval instars, development occurs in the Ephemeroptera, Odonata,
these tissues are elaborated to form the adult struc- and Plecoptera, and the young of these insects are
tures, and when the insect pupates, they are everted sometimes called naiads. Paurometabolous insects
(hence one name for holometabolous insects, the En- (with "gradual" metamorphosis) include the remaining
dopterygota, referring to the development of the wings insects with simple metamorphosis. The adults are
inside the body of the larva). Other organs may be re- winged; the nymphs and adults live in the same habitat;
tained from the larva to the adult or may be completely and the principal changes during growth are in size,
rebuilt from regenerative cells. body proportions, the development of the ocelli, and
occasionally the form of other structures.
Simple Metamorphosis
The young of insects with this type of metamorphosis Complete Metamorphosis
are called nymphs' and are usually very similar to the The immature and adult stages of insects that undergo
adults. If compound eyes are present in the adult, they complete metamorphosis are usually quite different in
are present in the nymph. If the adults are winged, the form, often live in different habitats, and have very dif-
wings appear as budlike outgrowths in the eariy instars ferent habits. The early instars are often more or less
(Figure 2-41) and increase in size only slightly up to wormlike, and the young in this stage are called larvae
the last molt. After the last molt, the wings expand to (Figures 2-42 and 2-43). The different larval instars
their full adult size. Simple metamorphosis occurs in are usually similar in form but differ in size. The wings,
the hexapod orders 1 to 22 (see list, Chapter 6). when they are present in the adult, develop intemally
Differences in the kind and amount of change oc- during the larval stage and are not everted until the end
cur in the insects with simple metamorphosis, and of the last larval instar. Larvae generally have chewing
some entomologists recognize three types of metamor- mouthparts, even in those orders in which the adults
phosis in these insects: ametabolous, paurometabolous, have sucking mouthparts.
and hemimetabolous. Ametabolous insects (with "no" Following the last larval instar, the insect trans-
metamorphosis) are wingless as adults, and the only ob- forms into a stage called the pupa (Figure 2-44). The
vious difference between nymphs and adults is size. insect does not feed in this stage and is usually inac-
This type of development occurs in the apterygote or- tive. Pupae are often covered by a cocoon or some
other protective material, and many insects pass the
4 In the European literature of entomology, the immature stages of all winter in the pupal stage. The final molt occurs at the
insects are generallyreferred to as larvae. end of the pupal stage, and the last stage is the adulto

Development and Metamorphosis 47



Figure2-42 Stages in the development of the sugarbeet root maggot, Tetanopsmyopae-

fonnis (Roder). A, Adult female; B, Adult male; C, Egg; D, Larva; E, Puparium (pupa in-
side). (From Knowlton 1937, used courtesy of the Utah Agricultural Experiment Station.)

L = 30 mm
1 B

L = 45 mm

Figure2-43 Insect larvae. A, Maggot or vermiform larva of Hylemya platura (Meigen)

(Diptera, Anthomyiidae); B, Grub or scarabaeiform larva of Phyllophaga rugosa
(Melsheimer) (Coleoptera, Scarabaeidae); C, Elateriform larva of Cardiophorussp.
(Coleoptera, Elateridae); D, Elateriform larva of Alaus oculatus (L.) (Coleoptera,
Elateridae); E, Campodeiform larva of Attagenus megatoma (Fabricius) (Coleoptera,
Dermestidae); F,Vermiform larva of Cylasfonnicarius elegantulus (Summers) (Coleoptera,
Brentidae); G, Eruciform larva of Calima aethiops (Fabricius) (Hymenoptera, Tenthre-
dinidae). a, antenna; as, anterior spiracIe; L, length; ps, posterior spiracIe; sp, spiracIe.
(A, E-G, Courtesy of USDA;B-D, From Peterson, 1948, by permission.)

Ihe adult is usually pale in color when it first emerges holometabolous (see list, Chapter 6), and these orders
[ramthe pupa, and its wings are short, 50ft, and wrin- are cIassified together as the HoIometaboIa.
kled.In a short time, from a few minutes to several Hypermetamorphosis is a type of complete meta-
hoursor more, depending on the species, the wings ex- morphosis in which the different larval instars are not
pandand harden, the pigmentation develops, and the of the same type. The first instar is active and usually
insectis ready to go on its way. This type of metamor- campodeiform, and the subsequent larval instars are
phosis,which occurs in orders 23 to 31, is often called vermiform or scarabaeiform (see definitions of these

48 Chapter2 TheAnatomy.Physiology.and Developmentof Insects


Figure2-44 Insect pupae. A, Chrysalis of the sulphur butterfly, Co/ias eurytheme

Boisduval (Lepidoptera, Pieridae); B, Fall armyworm, Spodopterafrugiperda O. E. Smith)
(Lepidoptera, Noctuidae); C, Clover seed chalcid, Bruchophagusplatyptera (Walker)
(Hymenoptera, Eurytomidae); D, Sweetpotato weevil, Cylas fonnicarius degantulus
(Summers) (Coleoptera, Brentidae); E, Sawtoothed grain beetle, Oryzaephilus surinamen-
sis (L.) (Coleoptera, Silvanidae); F, Seedcorn maggot, Hylemya platura (Meigen) (Diptera,
Anthomyiidae). A and B are obtect pupae, C-E are exarate pupae, and F is a coarctate
pupa. (Courtesy ofUSDA.)

tenns later). Hypermetamorphosis occurs in parasitic three instars are inactive, sessile, and scalelike, with the
insects; the first instar seeks out the host and, once in wings developing internally. The fourth instar, calIed the
the host, molts into a less active type of larva. This type pupa, has external wings. The first three instars are usu-
of complete metamorphosis occurs in the Meloidae alIy called larvae. The molt from the last larval instar to
(Figure 26-68) and Ripiphoridae (Coleoptera), the the pupa takes place inside the last larval skin, which
Mantispidae (Neuroptera), the Strepsiptera, and a few forros the puparium. This metamorphosis is essentialIy
Diptera and Hymenoptera. complete, although most other members of this order
(Hemiptera) have simple metamorphosis.
Intermediate Typesof Metamorphosis
The males of scale insects have a type of meta-
Not all insects have a type of metamorphosis that can
morphosis that is very similar to that in whiteflies. The
be readily classified as simple or complete. Some have
first instar (Figure 22-63B), the "crawler," is active and
a metamorphosis that is somewhat intennediate be-
wingless, but the remaining preadult instars are sessile
tween these two types. Such metamorphosis is found
and inactive. The last preadult instar, which has exter-
in thrips (Chapter 23), whiteflies (Chapter 22), and
nal wings, is calIed the pupa. The development of
male scale insects (Chapter 22). In fact, these groups
wings is at least partly interna!.
have gone far toward evolving complete metamorpho-
sis independently of the orders just discussed. Control of Metamorphosis
The first two instars of thrips (Thysanoptera) are The metamorphosis of insects is controlIed by three
wingless and active and are usually called larvae. The honnones: PTTH (prothoracicotropic or brain hor-
next two instars (the next three in the suborder Tubu- mone), ecdysone, andJH (juvenile honnone). PTTH is
lifera) are inactive, with external wings. The first of produced by neurosecretory cells in the brain and stim-
these (the first two in Tubulifera) is calIed a prepupa, ulates the prothoracic glands (also known as the malt-
and the last a pupa. The final instar is the adulto Ap- ing glands) to produce ecdysone, which induces apoly-
parently at least some of the wing development is in- sis and promotes growth. JH is produced by celIs in the
ternal during the first two instars. This metamorphosis corpora alIata and inhibits metamorphosis, thereby
resembles complete metamorphosis in that at least promoting further larval or nymphal development. Re-
some of the wing development is internal, and an inac- moving JH from a larva or nymph (by removing the
tive, "pupal" stage precedes the adult. It is similar to corpora alIata) causes the larva to pupa te and the
simple metamorphosis in that the early instars have nymph to develop into an adult when ecdysone is pres-
compound eyes, and external wing buds are present in ent. Injection of juvenile honnone into a pupa (in the
more than one preadult instar. presence of ecdysone) will cause the pupa to develop
Whiteflies have five instars, the last of which is the into a second pupa. Injecting JH into a last-instar
adulto The first instar is active and wingless, and the next nymph or larva causes another nymphal or larval stage
Variations in LifeHistory 49

tobe produced at the next molt. The corpora allata are a silken cocoon formed by the larva before it molts
activeduring the early instars and usually cease secret- to the pupal stage.
ingJH in the last pre-adult instar. The absence of the Exarate-with the appendages free and not glued to
hormonein this instar results in metamorphosis. the body (Figure 2-44C-E). Such a pupa looks
The changes from instar to instar in insects with much like a pale, mummified adult and is usually
simplemetamorphosis are generally relatively slight and not covered by a cocoon. This type occurs in most
gradual,being most marked at the final molt to the insects with complete metamorphosis, except the
adult,but insects with complete metamorphosis show Diptera and Lepidoptera.
considerablereorganization within the insect in the pu- Coarctate-essentially like an exarate pupa, but re-
palstage.Some structures in the larva, such as the heart, maining covered by the hardened cuticle of the
nervoussystem, and tracheal system, change very little last larval instar, which is called a puparium (Fig-
at metamorphosis. Other adult structures are present in ure 2-44F). This type occurs in the Diptera (sub-
a rudimentary form in the larva and remain so during order Brachycera).
successivelarval instars. Then, more or less suddenly, Decticous-with the mandibles movably articulated
theydevelop to their adult form in the pupal stage. Still with the head. This type is also always exarate and
otheradult structures are not represented in the larva occurs in the Neuroptera, Trichoptera, and some
andmust be developed at the time of metamorphosis. Lepidoptera.
Adecticous-with the mandibles immovably attached
Typesof Larvae
to the head. This type of pupa is found in the re-
Thelarvae of insects that undergo complete metamor-
maining groups of holometabolous insects.
phosis differ considerably in form, and several types
havebeen recognized:
Eruciform-caterpillar-like (Figure 2-43G); body
cylindrical, the head well developed but with very Variations in Life History
short antennae, and with both thoracic legs and
abdominal prolegs. This type occurs in the Lepi- The length of a generation and the way it is fitted to the
doptera, Mecoptera, and some Hymenoptera (sub- different seasons vary quite a bit in different insects.
order Symphyta). Most insects in temperate regions have what entomolo-
Scarabaeiform-grublike (Figure 2-43B); usually gists call a "heterodynamic life cycle"; that is, the adults
curved, the head well developed, with thoracic appear for a limited time during a particular season, and
legsbut without abdominal prolegs, and relatively some life stage passes the winter in a state of dormancy.
inactive and sluggish. This type occurs in certain The overwintering stage may be the egg (for example,
Coleoptera (for example, Scarabaeidae). most Orthoptera and Hemiptera), nymph (for example,
Campodeiform-resembling diplurans in the genus most Odonata and many Orthoptera), larva (for exam-
Campodea (Figure 7-4A);body elongate and some- pIe, many Lepidoptera), or adult (for example, most
what flattened, the cerci and antennae usually well Hemiptera and many Coleoptera and Hymenoptera).
developed, the thoracic legs well developed, and Many insects, particularly those living in the tropics,
the larvae usually active. This type occurs in the have a homodynamic life cycle; that is, development is
Neuroptera, Trichoptera, and many Coleoptera. continuous and there is no regular period of dormancy.
Elateriform-wirewormlike (Figure 2-43C,D); body Most insects in the United States have a single gen-
elongate,cylindrical, and hard-shelled, the legs eration ayear. Some require 2 or more years to complete
short, and the body bristles reduced. This type oc- their life cycle, as is usually the case with large insects
cursin certain Coleoptera (for example, Elateridae). occurring in the northem part of the country. Some of
Vermiform-maggotlike (Figure 2-43A,F); body elon- the large beetles, dragonflies, and moths in the northem
gateand wormlike, legless, and with or without a states and Canada require 2 or 3 years to complete their
well-developedhead. This type occurs in the development. Probably the longest life cycle of any in-
Diptera,Siphonaptera, most Hymenoptera (suborder sect is that of some of the periodical cicadas (Magicicada
Apocrita),and some Coleoptera and Lepidoptera. spp.), which lasts 17 years (see Chapter 22).
Many insects have more than one generation a
Types of Pupae
year. In some cases, the number of generations in a
The pupae of insects with complete metamorphosis
year is constant throughout the range of the species. In
vary,and five types may be recognized:
other cases, the species may have more in the southem
Obtect-with the appendages more or less glued to part of its range. A few insects, usually rather small
the body (Figure 2-44A,B). This type occurs in the species that can complete their life cycle in a few
Lepidoptera and some Diptera (suborder Nemato- weeks, have many generations ayear. Such insects con-
cera). The pupa in many Lepidoptera is covered by tinue to reproduce through the season as long as

50 Chapter2 TheAnatomy,Physiology,and Development of Insects

weather conditions are favorable. Insects of tropical spring. This factor of day length apparently operates
origin, such as those of the household and those which similarly in the case of the codling moth. Individuals of
attack stored products, may continue breeding the first generation pupate and emerge as adults in the
throughout the entire 12 months. summer, but individuals of the second generation (in
In many insects, development is arrested during a autumn) do noto In some cases (for example, Antheraea,
specific stage of the annual cycle. This period of genet- family Satumiidae), day length may also control emer-
icaIly programmed (that is, predetermined) dormancy gence from diapause. The larvae of the second genera-
is known as diapause, in contrast to periods of quies- tion of the codling moth in Ohio remain as diapausing
cence in response to adverse environmental conditions. larvae in silk-lined cells under the bark of apple trees
A period of winter dormancy in temperate or arctic re- unless subjected to a short period (some three weeks) of
gions is often caIled hibemation, and a period of dor- low temperatures (O°C or lower). When then retumed
mancy during high temperatures is called aestivation. to normal developmental temperatures, they pupate and
Diapause in insects is geneticaIly controlled, and complete their development. The effect of day length is
both onset and termination may be induced by environ- usually direct, on the insect itself, but may occasionally
mental factors such as photoperiod or temperature. The be indirect, its effect being on the food eaten by the in-
chief factor initiating diapause seems to be photoperiod sect. Photoperiod may also be an important cue for the
(day length). Studies of homworm larvae (Sphingidae) initiation of diapause in tropical insects, even though the
have shown that aIl individuals that enter the soil for pu- changes in day length are much smaller than in temper-
pation before a certain date wiIl complete their develop- ate regions. Diapause in tropical insects may be associ-
ment, emerge as moths, and reproduce, but individuals ated with altemating wet and dry seasons, high temper-
entering the soil after this date go into diapause and do atures, or the availability of appropriate food, instead of
not complete their development until the following the avoidance of winter (Denlinger 1986).


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Dethier, V.G. 1976. The Hungry Fly: A Physiological SLUdyof field, IL: C. C Thomas, 255 pp.
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vard University Press, 489 pp. Insect Physiology, Biochemistry and Pharmacology. 13 vols.
Eberhard, W G. 1985. Sexual selection and animal genitalia. Vo\. 1: Embryogenesis and Reproduclion; 482 pp. Vo\. 2:
Cambridge, MA: Harvard University Press, 224 pp. Poslembryonic Development; 505 pp. Vo\. 3: Integu-
Ferris, G. E, and B. E. Rees. 1939. The morphology of menl, Respiralion and Circulalion; 625 pp. Vo\. 4: Regu-
Panorpa nuptialis Gerstaecker (Mecoptera: Panorpide). lalion: Digeslion, Nutrition, Excrelion; 639 pp. Vo\. 5:
Microentomology 4:79-108. Nervous Syslem: Structure and MOlor Funclion; 646 pp.
References 51

Vol. 6: Nervous Systern: Sensory; 710 pp. Vol. 7: En- Rernpel, J. G. 1975. The evolution of the insect head: The
docrinology 1; 564 pp. Vol. 8: Endocrinology Il; 595 pp. endless dispute. Quaest. Entornol. 11:7-25.
Vol. 9: Behaviour; 735 pp. Vol. 10: Biochernistry; 715 pp. Riek, E. E, andJ. Kukalová-Peck. 1984. A new interpretation
Vol. 11: Pharrnacology; 740 pp. Vol. 12: lnsect Control; of dragonfly wing venation based on Early Upper Car-
849 pp. Vol. 13: Curnulative lndexes; 314 pp. New York: boniferous fossils frorn Argentina (Insecta: Odonatoidea)
Pergamon Press. and basic character states in pterygote wings. Can. J.
Knowlton, G. E 1937. Biological control of the beet leafhopper Zool. 62:1150-1166.
in Utah. Proc. Utah Acad. Sci., Arts, Letters 14:111-139. Rockstein, M. (Ed.). 1973-1974. The Physiology of lnsecta.
Kukalová-Peck, J. 1978. Origin and evolution of insect wings 6 vols., illus. Vol. 1: Physiology of Ontogeny-Biology,
and their relation to rnetarnorphosis, as docurnented by Peveloprnent, and Aging, 512 pp. (1973). Vol. 2: The ln-
the fossil record. J. Morphol. 156:53-126. sect and the External Environrnent. Part A. I. Environrnen-
Kukalová-Peck, J. 1983. Origin of the insect wing and wing tal Aspects. Part B. Il. Reaction and lnteraction, 517 pp.
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61:1618-1669. Part A. Il. Reaction and lnteraction. Part B. IlI. Locorno-
Kukalová-Peck,J. 1985. Epherneroid wing venation based on tion, 517 pp. (1974). Vol. 4: The Insect and the External
new gigantic Carboniferous rnayflies and basic rnorphol- Environrnent. Hornoeostasis I. 488 pp. (1974). Vol. 5: The
ogy, phylogeny, and rnetarnorphosis of pterygote insects. lnsect and the External Environrnent. Hornoeostasis Il.
Can. J. Zool. 63:933-955. 648 pp. (1974). Vol. 6: The lnsect and the External Envi-
Manton, S. M. 1977. The Arthropoda, Habits, Functional ronrnent. Hornoeostasis IlI. 548 pp. (1974). New York:
Morphology, and Evolution. Oxford, UK: Clarendon Acadernic Press.
Press, 527 pp. Rodriguez, J. G. (Ed.). 1972. lnsect and Mite Nutrition. New
Matheson, R. 1951. Entornology for introductory courses. York: American EIsevier, 701 pp.
lthaca, NY: Cornstock Publishing Cornpany, lnc., 600 pp. Roeder, K. D. 1967 (rev. ed.). Nerve cells and insect behavior.
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Matsuda, R. 1976. Morphology and Evolution of the lnsect Macrnillan, 499 pp.
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299-383. New York: Van Nostrand Reinhold, 762 pp. in lnsects. Copenhagen: Munksgaard, 359 pp.
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Corizus (Coreidae: Herniptera). Ann. Entornol. Soco Arner.

3 Systematics, Classification,
Nomenclature, and Identification

theme of diversity inevitably dominates any dis-
of insects and their relatives. Not only is
expression of that gene pool-to look for the smallest
set of phenotypic homogeneity while acknowledging
there a large number ofindividual insects (lhe biomass and incorporating known aspects of variability, such as
of ants in the tropics probably exceeds that of all verte- sexual dimorphism, developmental stages, seasonal
brates combined), but there is an almost overwhelming changes, geographic variation, and individual variabil-
number of different kinds of insects. The study of the ity. Hence researchers often must reIy on morphologi-
diversity of organisms and of the reIationships between calor other characters to determine specific limits;
them is the scientific fieId of systematics. This disci- caution is needed, because there are "good" species
pline also includes the study of classification, the field (groups reproductively isolated) that cannot be distin-
of taxonomy. Systematics forms the foundation on guished by morphological characters (known as sibling
which all other biological disciplines rest. The names species), and converseIy, within a single species there
of organisms provide a key to the published literature may be a number of different forms.
and enable us to communicate with one another. Clas- The subspecies category is sometimes used to refer
sifications serve both as information retrieval systems to recognizable and geographically restricted popula-
and, as they reflect the relationships among organisms, tions of a species. Because different subspecies of a
provide predictions of the distribution of characteris- given species are capable of interbreeding, the differ-
tics among organisms. ences between them are usually not clear-cut but inter-
The fundamental unit of systematics is the grading, particularly where adjacent subspecies come
species. Concepts of what a species is and how they in contact. This category has been misused, especially
origina te are based largely on studies of vertebrates. in the early part of the 20th century, to refer to any dis-
Extrapolation beyond them to other animals, especially tinguishable variant, often a color variant.
to fundamentally different organisms in other king-
doms, must be done with some careo Basically, most re-
searchers dealing with living animals define the species
to be a group of individuals or populations in nature Systematics
that (l) are capable of interbreeding and producing fer-
tile offspring, and (2) under natural conditions are re- Classifications of insects and their relatives form the
productiveIy isolated from (lhat is, ordinarily not in- heart of this book. Classifications are tools that pro-
terbreeding with) other such groups. This is known as vide names for groups of species and serve as a short-
the biologicalspeciesconceptoBecausethese criteria in- hand for communicating information about those
volve characteristics of living organisms, they are diffi- species. These instruments are utilitarian products
cult, sometimes perhaps impossible, to address directly. that emerge from systematics, the study of the diver-
Therefore, the first steps in systematics usually involve sity and interrelationships of organisms. Although the
attempts to infer the limits of a species (the extent of a fieId is as old as any area of biological inquiry, it has
52 reproductive community) by observing the phenotypic undergone a remarkable resurgence over the past two

Systematics 53

This renaissance has been fueled by (1) theo- cerned. Thus the primary goal of phylogenetic system-
reticaladvances in the nature and analysis of system- atics is the discovery of monophyletic graups.
aticdata; (2) the development of new sources of data These ideas that a structure found in two different
fromnuc\eic acid sequences; (3) the development of species is the same structure and that they share it be-
powerfuland affordable computers with which to an- cause they inherited the structure fram their common
alyzethese data; and (4) the grawth of the size and ancestor constitute the definition of a homologous
scope of natural history collections. As a result, the character. As you can see, the definitions of homolo-
process that began approximately 50 years ago has gous characters and synapomorphies are equivalent.
now transformed the systematics fram a field do mi- Characters that are not homologous, but only seem so,
natedby arcane facts and argument by authority, into arise thraugh the evolutionary processes of conver-
a science driven by data, explicit hypotheses, and gence, parallelism, and reversals. Convergence and
quantitativeanalysis. Nevertheless, systematics retains parallelism are the development of similar structures
theaesthetic appeal that comes fram the continual dis- independently in two different species. A reversal is a
coveryof new, beautiful, sometimes bizarre, but al- character that "Iooks" like the ancestral condition,
waysfascinatingforms of life. usually the loss or reduction of so me feature. The ab-
The conceptual advances in systematics were first sence of a structure may be due either to the fact that
coherently articulated by an entomologist, Willi the species (and its ancestors) never had it in the first
Hennig,a specialist on Diptera. Hennig published his place, or to its subsequent loss. It is usually difficult, if
principiesin German (Hennig 1950), but after publi- not impossible, to distinguish between these two hy-
cationin English his ideas received much more atten- potheses simply by studying the character itself. That
tion(Hennig 1965,1966). He called this new appraach conclusion relies on considering all the evidence on the
phylogenetic systematics;today it is often referred to phylogeny of the species.
simplyas phylogenetics or as cladistics. The ideas are How, then, can you determine if a character is an-
fairly straightforward. The characters of a species cestral or derived? First, we must divorce ourselves
arepassed down fram ancestor to descendant. Species, fram the idea that in science we can actually determine
in general, do not interbreed; therefore characteristics the truth of any statement. Rather, we form hypotheses
in one species cannot be transferred to a different, that are based on evidence and subject to testing. The
parallellineage. The appearance of new lineages, the dominant means of hypothesizing character polarity,
processof speciation, involves splitting an ancestral whether it is ancestral or derived, is to look at the form
speciesinto two (or possibly more) descendant species. in which the character occurs in graups outside of our
Ifa character changes, that is, becomes modified fram group of interest. The species being studied constitute
itsancestralstate, then this new, derived character will the in-group; those species with which the characters
appearonly in the species in which it arase or in its are being compared inorder to hypothesize their po-
descendants.Therefore, the sequence of appearance of larity make up the out-group. The state in which the
lineagesthrough time can be reconstructed by docu- character is found in the out-group is hypothesized to
mentingthe distribution of derived characters. be the plesiomorphic condition. Thus the choice of
Asin any specialty, entomology has a fair amount species lObe included in the out-graup is critical to the
ofjargon that the student must understand. Derived analysis. Ultimately, out-group analysis is based on the
charactersare called apomorphies (even if the charac- principIe of parsimony (see later discussion).
teris chemical, behavioral, or so on, and not morpho- The hierarchical pattern of distribution of synapo-
logicalat all). The ancestral characters are referred to morphies is represented as a treelike branching dia-
asplesiomorphies. Branches on the phylogenetic tree gram known as a cladogram (Figure 3-1). The two
arehypothesized on the basis of shared derived char- groups of species that diverge at any node are sister
acters among the species, called synapomorphies. groups. A cladogram resembles many of the classical il-
Suchgroupsare considered monophyletic (sometimes lustrations of the "tree of life," but with an important
calledholophyletic) graups and consist of an ancestral exception. Gn a cladogram, the units that are being
speciesand all its descendants. Shared ancestral char- studied-often species, but the unit could be a group
actersare symplesiomorphies; graups defined on the of species as well-are shown only at the apex of the
basisof symplesiomorphies are paraphyletic. Unnat- diagram, at the ends of the branches. If a fossil species
uralgroups, such as a graup containing only insects is included, it is found at the end of a branch, just as
andmosses, are polyphyletic. This is an extreme ex- any living species would be. The internodes, then, do
ample,of course, but polyphyletic graups usually re- not represent ancestral species. The order in which the
sultfrom the misinterpretation of characters, such as graups are listed (for example, from top to bottom in
thinkingthat two structures are "the same" when in Figure 3-1) is of no significance; the cladogram can be
factthey evolved independently in the species con- ratated about any of its nodes. The meaning of a clado-
54 Chapter3 Systematics,Classification,Nomenclature,and Identification

E. zephyrus

H 5
6 I
O. canadensis
O. americanus

4 I
C. dorsalis
C. ventralis
B. niger
H B. flavens
H A. longipes
A H A. curtipes

67 O. canadensis
1 I O. americanus
C. dorsalis
C. ventralis

B. niger
B. flavens
64 A. longipes
B 1 I A. curtipes

C. dorsalis
2 C. ventralis
t--t---t---t1 : B. niger
1 1 2 2 I 1
1 12 15 A. longlpes
e 1 1 A. curtipes

Figure 3-1 Samples of drawing conventions for depicting cladograms. A, Cladogram

with a standard length used for each branch. The numbers above each branch are the
number of synapomorphies defining it, the numbers below are decay indices, indicating
the minimum number of additional steps in a cladogram that did not include that
branch. B, Cladogram illustrating a polytomy (at the base), an area of the cladogram for
which either no evidence is available to resolve it into a series of dichotomous branches,
or for which the available data conflict. In this case the number above a branch is a boot-
strap value (in %), indicating the frequency with which that branch appears when the
data set is randomly resampled for characters; the number below the branch is a decay
index. C, A cladogram in which the synapormophies are indicated on the branches in
which they appear. The number above the branch represents the character, the number
below indicates which state the character exhibits. The different format of the cross-
hatch (either solid black or gray) is used here to indicate if the synapormorphy is unique
or if it also occurs elsewhere in the cladogram. None of the conventions shown is stan-
dardized, and the caption should always explain such details to the reader.
Systematics 55

gramliesin the hierarchy illustrated by the branching veloped to find the most likely phylogenetic hypothe-
diagram. sis (cladogram) given the observed data and model of
Relationship, in the sense of phylogenetic system- base substitution.
atíes,is defined as the relative recency of common an- Often the internodes of a cladogram are labeled
cestry.In other words, given three species-A, B, and with information that indicates the support for the hy-
C-species A and B are more closely related if they pothesis that it represents a monophyletic group. This
sharea more recent common ancestor with each other may be a listing of the apomorphic characters that de-
thaneither does with C. If we assume that the animal fine that section of the cladogram; also commonly
kingdomis monophyletic, that all animals share a com- found are bootstrap support values or Bremer support
mon ancestor, then any pair of animals is related in values. A bootstrap value represents the percentage of
somesense. The critical issue is not if two species are times that grouping (or clade) appears when a large
related,but how closely related they are. Thus any number of subsets of the original data set are analyzed.
meaningful phylogenetichypothesis must deal with at Some researchers think a high number (say, 98%) indi-
Icastthree species. cates that the data strongly support the hypothesized
In the simplest case of three taxa (species or monophyletic group. Note, however, that this is not a
groupsof species), there are only three possible pat- statistical test in the usual sense of the termo The hy-
tems of relationship: either A+B are most closely re- pothesis, the proposed monophyletic group, is not re-
lated,MC, or B+C. With four terminal taxa (A to D) jected on the basis of this number. A Bremer support
the number of possible patterns increases to 15, with value indicates the number of further steps that would
fivetaxa there are 105 possible patterns; for ten taxa, be added to the entire cladogram if the clade under
over34 million possibilities; and the number continues consideration were not included and its species were
togrowexponentially. What criteria are used to distin- placed in a different pattern. The significance of the ab-
guishamong all these possibilities? solute value of such a number, then, depends on the to-
The most commonly used metric is the number of tallength of the cladogram, which, in turn, depends on
"steps"on each tree. Each individual step represents the number of taxa and characters.
thechange in a character from the ancestral to derived The student will often see published cladograms
condition.These changes include the possible hypoth- in which a node do es not give rise to two branches but
esisthat a character changes back to what appears to be instead gives rise to three or more. This polytomy in-
the ancestral sta te, that is, a character reversal. One dicates either that there are no data to support any one
dominantmethod of analysis then applies the principIe of the possible sets of dichotomous possibilities at that
of parsimony to select the optimal cladogram. The point, or that the data conflict so that no one possibil-
dadogram that has the fewest number of steps, the ity is more parsimonious (or likely) than another. Poly-
shortestcladogram, is held to be the best explanation tomies are expressions ofignorance or ambiguity in the
of the data at hand. All other, longer cladograms re- underlying data. Although it is biologically reasonable
quireunnecessary hypotheses of character change. The to believe that an ancestral species could give rise,
principIeof parsimony (Occam's razor) holds that the more or less simultaneously, to three or more daughter
bestexplanationfor a phenomenon is the simplest one species, the methodology of cladistics, and of science
thataccounts for the observed data. This should not be in general, compels us to search for a completely di-
interpreted to mean that evolution proceeds by the chotomously branching cladogram. To do otherwise
mastparsimonious path. We have clear evidence that would be to actively ignore the phylogenetic signal that
thatnotion would be false. Rather, parsimony simply do es exist in the data.
requiresus to have evidence to support hypotheses of It is probably becoming clear at this point that cal-
relationship. To do otherwise means that any clado- culating the most parsimonious cladogram for a group
gramthatappealed to a researcher could be claimed to of any size requires the use of a computer and cladistic
bethe "best" on grounds that had nothing to do with software. Only in cases with a relatively small number
of species or an unambiguous data set is it possible to
Phylogenetic analyses of sequence data often use a arrive at the best answer manually. A number of
differentoptimality criterio n by which to judge the software packages are now available for these analyses;
bestcladogram: maximum likelihood. This method re- two of the most popular are PAUP (Phylogenetic
lieson specifying a model of base transformation; for Analysis Using Parsimony) and the WinClada package.
example,what is the probability of an adenine residue MacClade is a popular tool for visualizing and manip-
changingto a thymine at a given position? The overall ulating data on cladograms. However, even with the
likelihoodof a set of cladograms is calculated, and the most powerful hardware and software some phylo-
onewith the highest likelihood is given preference. Re- genetic problems are inherently intractable. In many
cently,Bayesian analysis methods have begun to be de- cases, the analysis results in large numbers of equally
56 Chapter3 Systematics.Classification.Nomenclature.andIdentification

parsimonious trees. In such cases, authors often resort no harm, that is, to change the formal classification
to publication of consensus trees to depict at least the where necessary to preserve the criterion of mono-
components that all the solutions have in common. phyly, but to keep such changes to a minimum. There
is, obviously, a great deal of room here for individual
interpretation, and, as a result, several altemative clas-
sifications may be in use at any point in time, particu-
Classifications larly for popular groups (such as butterflies).
Many groupings of insects first recognized by early
In a formal biological classification, species are systematists such as Fabricius and Latreille are still rec-
grouped into monophyletic groups. Such groups are ognized today as valid monophyletic groups. Examples
called taxa (sing., taxon). These taxa are arranged in a include the Apocrita (Hymenoptera), Brachycera
hierarchical pattem. The most commonly used cate- (Diptera), Ditrysia (Lepidoptera), and Polyphaga
gories or levels in the system of zoological classifica- (Coleoptera). However, in traditional classifications
tion are the following (listed from most inclusive to these taxa were often paired off in contrast with an-
least) : other group, Symphyta versus Apocrita, Nematocera
versus Brachycera. These "sister" taxa often are not
Phylum true sisters in the phylogenetic sense, because they are
Class widely acknowledged to be paraphyletic groups. As
Subclass such, according to the principIes of phylogenetic sys-
Order tematics they should not be formally recognized. Ento-
Suborder mologists are, indeed, moving in that direction, but it
is a slow and gradual process. We have continued to in-
clude some of these groups in the classifications in
each chapter, partly beca use the names are so widely
Tribe used that it is important for the student to leam them.
Genus However, in the long run classifications are based on
scientific hypotheses, and these hypotheses may be
changed or rejected as understanding of insect diver-
sity increases. Thus expect to see changes in those clas-
sifications reflecting the underlying scientific research,
In this scheme, the animal kingdom is divided into and embrace the changes as indications of advances in
a number of phyla (sing., phylum). Each phylum is di- the understanding of insect diversity and relationships.
vided into classes, classes into orders, orders into fam-
ilies, families into genera (sing., genus), and genera
into species. Finer distinctions in the levels is afforded
by prefixes attached to these category names, such as Nomenclature
supeifamily or subgenus. This list provided is not com-
prehensive, but does illustrate the normal categories Animals have two types of names, scientific and com-
that are used. The species is probably the only level mono Scientific names are used throughout the world,
that can be assessed by objective criteria. A genus is and every known animal taxon has a scientific name
one or more species classified together in a mono- unique to it. Common names are vernacular names,
phyletic group; a family is one or more genera; and so and they are often Iess precise than scientific names.
on. Any scheme of classification that is developed for a (Some common names are used for more than one
group of animals will be affected by the particular char- taxon, and a given animal taxon may have several.)
acters used, the relative weight they are given, and how Many animals lack common names beca use they are
they are analyzed. If different people use different char- small or seldom encountered.
acters or a different "weighting" of a series of charac-
ters, they may arrive at different classifications. Scientific Nomenclature
A classification is derived from a phylogeny by
designating monophyletic groups for each category. Of The scientific naming of animals follows certain rules,
course, a dichotomous cladogram of n species will have outlined in the IntemationaI Code of ZooIogicaI Nomen-
n -1 monophyletic groups. For a cladogram of any clature (ICZN 1999). Scientific names are latinized, but
size, particularly in insects where one order may have they may be derived from any language or from the
more than 100,000 species, not all these groups are for- names of people or places. Most names are derived
mally named. The usual approach systematists take is from Latin or Greek words and usually refer to some
similar to the admonition in the Hippocratic oath to do characteristic of the animal or group named.
Nomenclature 57

The names of graups above genus are latinized and dark brawnish (not violet) coloration. These
nouns in the nominative (subject case) plural. The three subspecies are listed as Argia fumipennis vio-
namesof genera and subgenera are latinized nouns in laeea (Hagen), Argia fumipennis fumipennis
the nominative singular. Specific and subspecific (Burmeister), and Argiafumipennis atra Gloyd.
namesmay be adjectives (descriptors, or modifiers), Hagen's name in parentheses means violaeea was
present or past participles (forms) of verbs (action described in a genus other than Argia, but there is
words),or nouns (names of things or places). Adjec- no way of knowing fram this name whether vio-
tivesand participles must agree in gender (masculine, laeea was originally described as a species, as a
feminine,or neuter) with the genus name, and nouns subspecies of fumipennis, or as a subspecies of
are in either the nominative or the genitive (posses- some other species. Gloyd's name without paren-
sive)case. theses means atra was originally described in the
Thescientific name of a species is a binomen; that genus Argia, but there is no way of knowing fram
is,it consists of two words: the genus name and a spe- this name whether atra was originally described as
ciflcepithet. That of a subspecies is a trinomen: the a species of Argia, as a subspecies of fumipennis, or
genusname, the specific epithet, and a subspecific epi- as a subspecies of another species of Argia. In fact,
thet.These names should be printed in italies (if writ- Hagen originally described violacea as a species of
tenor typewritten, italics can be indicated by underlin- Agrion, and Gloyd originally described atra as a
ing).Names of species and subspecies are sometimes subspecies of Argia fumipennis.
followedby the name ofthe author, the person who de-
Some entomologists have used trinomials for what
scribedthe species or subspecies. Authors' names are
notitalicized.The appended author name is sometimes they have called "varieties," for example, A-us b-us, varo
e-uso Other ranks may also be found, including form,
importantin dealing with the systematics of a species or
stirps, race, and so forth. Such names, if published be-
in caseswhere the same name has been praposed for
fore 1961, are assumed to be names of subspecies, in
twodifferent species (a case of homonyrny). In most
which case the "var." in the name is drapped, or if they
othercases, it is simply superfluous and adds nothing
are shown to designa te an individual variant, they are
morethana falsesense of authority to the person citing
thename. The names of genera and higher categories considered "infrasubspecific" categories, which are not
covered by the Intemational Code of Zoological Nomen-
alwaysbegin with a capital letter; species and sub-
clature. Such names published after 1960 are considered
speciesnames never do and generally are not cited
to designate individual variants ("infrasubspecific" cat-
withoutthe name or abbreviation of the genus. If the
author'sname is in parentheses, it means that the egories), not covered by the mIes. The taxonomic cate-
species(or subspecies, in the case of a subspecies name) gories just listed apply to populations and not to indi-
vidual variants such as color forros, sexual forros, and
wasdescribed in some genus other than the one in seasonal forros.
whichit is now placed. For example,
A species referred to but not named is often desig-
PapilioglaucusLinnaeusl-the tiger swallowtail.The nated simply by "sp." For example, "Gomphus sp."
speciesglaucus was described by Linnaeus in the refers to a species of Gomphus. More than one species
genusPapilio. may be designated by "spp."; for example, "Gomphus
deeemlineata (Say)-the Colorado potato spp." refers to two or more species of Gomphus.
beetle.The species deeemlineata was described by The names of categories fram tribe through super-
ThomasSay in the genus Doryphora, and this family have standardized endings and hence can always
specieshas since been transferred to the genus be recognized as referring to a particular category. These
Leptinotarsa. can be illustrated by some taxa of bees, as follows:
fumipennis(Burmeister)- The speciesfumipen-
Superfamily names end in -oidea;for example,
niswas described by Hermann Burmeister in some
Apoidea: the bees. Note that the names of some
genusother than Argia and has subsequently been
genera and categories above the level of family can
transferred to the genus Argia. There are three
have the same ending.
subspeciesof this species in the eastern United
Family names end in -idae; for example, Apidae:
States:a northern subspecies (violaeea) with clear
wingsand considerable violet coloration, a south- cuckoo bees, digger bees, carpenter bees, or-
chid bees, bumble bees, and honey bees.
em subspecies (fumipennis) with smoky wings and
Subfamily names end in -inae; for example,
considerable violet coloration, and a subspecies in
Apinae: orchid bees, bumble bees, and
peninsular Florida (atra) with very dark wings
honey bees.
Tribe names end in -ini; for example, Apini,
lThroughoutthis book Linnaeus is abbreviated "L." honey bees.
58 Chapter3 Systematics.

Types. Whenever a new taxon (from subspecies to su- (homonyrns and synonyrns) are not easy to discover.
perfamily) is described, the describer is supposed to As they are discovered, it becomes necessary to change
designa te a type, which serves to anchor the name to a names, not only of genera and species, but also of fam-
taxonomic concepto The type of a species or subspecies ilies and even orders. The problems of priority in sci-
is a single specimen (rhe type, or holotype); the type of entific nomenclature are often very intricate, and it is
a genus or subgenus is a species (the type species); and sometimes difficult to determine just what name is the
the type of a taxon from tribe through superfamily is a correct one. Name changes may also result from in-
genus (the type genus). Names of taxa from tribe creased knowledge. This added knowledge may indi-
through superfamily (see the previous examples) are ~ate that groups should be split or combined, resulting
formed by adding the appropriate ending to the root of in name changes for some of the groups involved.
the name of the type genus. For the type genus Apis in In cases where two or more names for a group
the previous examples, the stem is Ap-. If a species is have been in fairly wide use, we have listed in this book
divided into subspecies, the particular subspecies that first what we believe to be the correct name and have
includes the holotype of the species has the same sub- listed other names in parentheses.
specific name as its specific name (for example, Argia
fumipennisfumipennis). Similarly, if a genus is divided Pronunciation
into subgenera, the subgenus that includes the type
species of the genus has the same subgenus name as The pronunciation of some of the technical names and
genus name-for example, Formica (Formica) ruja L. terrns used in entomology may be found in a good dic-
(The name in parentheses is the subgenus.) As con- tionary or glossary, but very few texts or references give
cepts of the limits of species or other taxa are revised, the pronunciation of the bulk of scientific names.
it is often necessary to refer to the available types (and There are roles for the pronunciation of these names,
ultimately the holotypes of the relevant species) to de- but few entomologists are familiar with them, and
termine to which concepts of taxa of the systematist many names are pronounced differently by different
the types belong, and, thus, to determine which name people and in different countries. We have therefore in-
or names are applicable to it. The significance of holo- cluded some of the general roles by which technical
types has to do with nomenclature; they do not repre- names and terms used in zoology are generally pro-
sent a "typical" member of a species. nounced in American English. We realize that not all
entomologists will agree with the pronunciation of
Priority. The staning point of modem binomial zoolog- some names. There are two reasons for such disagree-
ical nomenclature was the publication of 10th edition of ment: (1) a given pronunciation, whether it follows the
Unnaeus' Systema Naturae; the date is taken as 1January roles or not, may become established through usage as
1758. lt often happens that a panicular taxon is de- the "correct" pronunciation; and (2) the correct pro-
scribed independently by two or more people, and hence nunciation of many scientific names depends on the
may have more than one name. In such cases the first derivation of the name and the vowel sound in the
name used from 1758 on (if the describer has followed source language, and it is difficult or impossible to de-
certain roles) is the correct name, and any other names termine the derivation of some names. Hence there will
become synonyrns and should no longer be used. A par- always be a question as to the correct pronunciation of
ticular name is often used for a long time before someone some scientific names.
discovers that another name has priority over it. The principal roles for the pronunciation of scien-
Sometimes a person describing a new taxon gives tific names and terms are outlined here:
it a name that has previously been used for another
taxon; if those taxa involved are at the same taxonomic Vowels. All vowels in scientific names are pro-
nounced. Vowels are generally either long or short, and
level, the names are termed homonyms, and all but the
oldest must be discarded and the taxa renamed. There in the examples that follow a long vowel sound is indi-
cated by a grave accent O) and a short vowel sound by
cannot be two (or more) species or subspecies with the
an acute accent (O; for example, mate, mát, mete, mét,
same name in a given genus. There cannot be two (or
bite, bU, rope, rót, cute, cút, by, symmetry. A vowel at the
more) genera or subgenera in the animal kingdom with
the same name. Nor can there be two (or more) taxa in end of a word has the long sound, except when it is an
-a; a final -a has the uh sound, as in idea. The vowel in
the family group of categories (rribe through super-
the final syllable of a word has the short sound, except
family) with the same name (although the names of the
-es, which is pronounced ease.
typical subdivisions will be the same except for their
endings). The fundamental role is that every animal Diphthongs. A diphthong consists of two vowels writ-
taxon must have a unique name. ten together and pronounced as a single vowel. The
Because of the large number of animal taxa and the diphthongs are ae (pronounced e, rarely é), oe (usually
vast amount of zoological literature, errors in naming pronounced e, rarely é), oi (pronounced as in oi/), eu
Nomenclature 59

(pronouncedu), ei (pronounced 1), ai (pronounced a), Sphecinae) and tribe names (for example,
andau (pronounced as in August). The final -ae in fam- Sphecini); in tribe names, the final i is also long.
ily and subfamily names is generally pronounced e. 2. In other cases the antepenult is accented.
Thecombination ii is pronounced as two syllables and
is thereforenot a diphthong. Common Names of Inseds
Consonants. Ch has the k sound, except in words de- Because there are so many species of insects, and be-
rivedfram a language other than Greek. When e is fol- cause so many of them are very small or poorly known,
lowedby ae, e, oe, i, or y, it has the soft (s) sound; relatively few have common names. Those that do are
whenit is followed by a, o, oi, or u, it has the hard (k) generally particularly showy insects or insects of eco-
sound.When g is followed by ae, e, i, oe, or y, it has the nomic importance. American entomologists recognize
soft(j) sound; when it is followed by a, o, oi, or u, it as "official" the common names in a list published
hasthe hard sound (as in go). In words beginning with every few years by the Entomological Society of Amer-
ps,pt, et, en, gn, or mn, the initial letter is not pro- ica, but this list do es not include all species of insects
nounced,but when these letters appear together in the (and other arthropods) to which common names have
middleof a word, the first letter is pronounced (for ex- be en applied. The common names used in this book
ample,the p is not pronounced in the word pteromorph, for individual species have be en taken from this list or
but it is pronounced in the word Orthoptera). An x at have been obtained from other sources.
the beginning of a word is pronounced as z, but as ks Many common names of insects refer to groups such
whenit appears elsewhere in a word. When a double e as subfamilies, families, suborders, or orders, rather than
is followedby e, i, or y, it is pronounced as ks. to individual species. The name bark beetle, for example,
refers to all species in the subfamily Scolytinae of the
Accent. The accented syllable is either the penult or
family Curculionidae. The name leafbeetle applies gener-
theantepenult (very long words may have a secondary
ally to all species in the family Chrysomelidae. The name
accenton a syllable near the beginning of the word).
beetle applies to all species in the order Coleoptera. The
Theprincipal roles governing the syllable accented and
name damselfly applies to the entire suborder Zygoptera,
thevowelsound (whether long or short) are as follows:
of which there are hundreds of species.
1. The accent is on the penultimate syllable in the Most common names of insects that consist of a
following cases: single word refer to entire orders (for example, beetle,
a. When the name contains only two syllables; for bug, eaddis, coekroaeh, and termite). Some (such as bee,
example,Apis, Bómbus. damselfly, grasshopper, and laeewing) refer to suborders
b. When the penult contains a diphthong; for ex- or groups of families. Onlya few (such as ants) refer to
ample, Culicoides, Hermileuea, Lygaeus. families. Most common names applying to families
c. When the vowel in the penult is followed by x consist of two or more words, the last being the name
or z; for example, Coríxa, Prodóxus, Agromyza, of the larger group, and the others descriptive (for ex-
Trioza. ample, brown laeewings, dick beetle, soldier flies, and
d. When the vowel of the penult is long. Whether small winter stoneflies).
the penult vowel is long or short often depends The members of a group are often referred to by an
on the derivation of the word and the vowel anglicized form of the group name. For example, insects
sound in the source language. The vowel e in a in the order Hymenoptera may be called hymenopterans;
word derived from the Greek is long if the the wasps in the superfamily Chalcidoidea are called
vowel in the Greek word is eta (1J), but short if ehalcidoids; those in the Sphecidae are called sphecids;
it is epsilon (E); for example, in words derived and those in the subfamily Nyssoninae may be called
from the Greek fL1JPOc:J>, meaning thigh, the e is nyssonines. lt is a common practice to use such a form of
long (Diapheromera, epimeron), whereas in the family or subfamily name as a common name.
those derived from the Greek fLEpOIT, meaning The names "fly" and "bug" are used for insects in
part, the e is short (Heterómera). Similarly, the more than one order, and the way the names of these
vowel o in a word derived from the Greek is insects are written may indicate the order to which the
long if the vowel in the Greek word is omega insect belongs. For example, when a fly belongs to the
(w), but short if it is omicron (o); for example, order Diptera, the "fly" of the name is written in this
in words derived from the Greek ITWfLCX mean- book as a separate word (for example, blaek fly, horse
ing body, the o is long (Calosoma, Malaeosoma), fly, and blow fly); when it belongs to another order, the
whereas in those derived from the Greek "fly" of the name is written together with the descrip-
<TTOfLCX, meaning mouth, it is short (Melanó- tive word (for example, dragonfly, butterjly, and sawfly).
stoma, Belóstoma, epístoma). The penult vowel When a bug belongs to the order Hemiptera, the "bug"
is long in subfamily names (for example, of the name is written as a separate word (for example,
60 Chapter3 Systematics, Classitication,
and Identitication

damsel bug, stink bug, and lace bug); when it belongs to pictures is often unsafe, beca use in many instances one
another order, the "bug" of the name is wriuen to- type of insect looks a great deallike another.
gether with the descriptive word (for example, mealy- As a general rule, in this book identification is car-
bug, ladybug, junebug, and sowbug). Unfortunately, this ried only to family. To go much further usually requires
practice is not universally accepted. specialized knowledge and is beyond the scope of this
book. Identifying insects only to family reduces the num-
ber of names from many thousand to several hundred,
and of these probably only 200 or fewer are likely to be
The Identification of Insects encountered by the average student. We reduce the prob-
lem still further by being concemed largely with adults.
When someone encounters an insect, one of the first Thus insect identification becomes less formidable.
questions askedis "What kind of insect is it?" (Or per-
haps that is the second question after "Will it bite
me?") One of the principal aims of the beginning stu-
dent in any field of biology is to become able to iden- The Keys in This Book
tify the organisms he or she is studying. The identifica-
tion of insects differs from the identification of other Analytical keys are devices used to identify all sorts of
types of organisms only in that it is likely to be some- things, living as well as nonliving. Different keys may
what more difficult, because there are more kinds of in- be arranged somewhat differently, but all involve the
sects than anything else. same general principIes. You run an insect (or other or-
Four things complicate the problem of insect iden- ganism) through a key in steps. At each step you face
tification. First, there are so many different species of two (rarely more) altematives, one of which should
insects that the beginner may be discouraged about apply to the specimen at hand. In our keys either a
ever becoming proficient in insect identification. Sec- number or a name follows the altemative that best fits
ond, most insects are small, and the identifying char- the specimen. If there is a number, the next step is the
acters are often difficult to see. Third, many insects are couplet with this number. Thus each step leads to an-
poorIy known, and when they are finally identified, other step and its altematives until a name is reached.
you may have only a technical name (which you may The couplets of altematives in our keys are num-
not understand) and liule biological information. bered 1 and 1', 2 and 2', and so on. In the first half of
Fourth, many insects go through very different stages each couplet after the first is a number in parentheses.
in their life history, and you may come to know insects This is the number of the couplet from which that cou-
in one stage of their life cycle and still know very liule plet is reached, and it enables the student to work
about those same insects in another stage. backward in the key if a mistake is discovered. This
There are about five ways in which a student may method of numbering also serves as a check on the ac-
identify an unknown insect: (1) by having it identified curacy of the organization of the key. In a few large
by an expert, (2) by comparing it with labeled speci- keys, certain couplets may be reached from more than
mens in a collection, (3) by comparing it with pictures, one previous couplet, and this fact is indicated by two
(4) by comparing it with descriptions, (5) by using an or more numbers in parentheses.
analytical key, or by a combination of two or more of The keys in this book have been prepared from
these procedures. Of these, obviously the first is the three principal sources: other published keys, descrip-
simplest, but this method is not always available. Sim- tions, and an examination of specimens of the groups
ilarIy, the second method may not always be available. concemed. Many are taken largely from previously pub-
In addition, the use of a collection is of limited value if lished keys (generally with some changes in wording or
you do not know the characteristics that distinguish organization), but some represent a new approach. Our
the species in a particular group. In the absence of an aim with each key has been to prepare something that
expert or a labeled collection, the next best method is is workable for practically every species (or specimen)
usually to use a key. In the case of particularIy striking in the groups covered. Most of these keys have been
or well-known insects, the identification can often be tested by student use over a number of years. In many
made by the third method mentioned, but in many of the insect orders (particularIy the larger ones), some
groups this method is unsatisfactory. No book can il- groups key out in more than one place. This is the case
lustrate all kinds of insects and still sell for a price a in two types of situations: (1) where there is significant
student can afford to pay. Where an unknown insect variation within the group, and (2) with borderline
cannot be definitely identified by means of illustra- cases where the specimen seems to fit both altematives
tions, the best procedure is to use an analytical key and of a couplet. In the latter situation, a specimen should
then to check the identificationby as many of the other key out correctly from either altemative. Although our
methods mentioned as possible. Identification from intent is that these keys will work for every specimen,

Geographic(overageof ThisBook 61

werealizethat species or specimens in many groups are Analytical keys are made for people who do not
erraticin their characters. We have sought to include as know the identity of a specimen they have. Once a
manyas possible of these atypical forms in our keys, specimen has be en identified with a key, subsequent
buta fewmay not key out correctly. We believe our keys identifications of this same insect may often be based
shouldwork for 95% or more of the insects of the con- on such characters as general appearance, size, shape,
tinentalUnited States and Canada. We believe the user and color, without reference to minute characters.
ofour keys is more likely to reach an impasse beca use lt will be apparent very early during your work in
ofan inability to see or interpret a character than be- identifying insects that you need a good binocular mi-
causeof a discrepancy in the key. croscope to see many characters of the insect. Most
When a determination is reached in the key, check insects, once you know what to look for, can be iden-
thespecimen against any illustrations or descriptions tified with a good hand lens (magnification about
available.If these do not fit the specimen, then either a lOX).
mistakehas been made somewhere or the specimen is The mere identification and naming of insects
onethat will not work out correctly in the key. In the should not be the student's final objective in insect
[atterevent, save the specimen until you can show it to study. There is much more of interest in insects than
anexpert.It may be something rare or unusual. just identifying them. Go further, and learn some-
Success in running an insect through a key de- thing of the habits, distribution, and importance of
pendslargely on understanding the characters used. In insects.
manycasesin this book, the key characters are illus-
trated.Often several characters are given in each alter-
native.In case one character cannot be seen or inter-
preted,use the other characters. If at any point in the Geographic Coverage of This Book
keyyou cannot decide which way to go, try following
upboth alternatives and then check further with illus- The taxonomic treatment in this book of the various
trationsand descriptions when you reach a name. insect orders and the other groups of arthropods ap-
A great many families of insects are very unlikely plies to the fauna of North America north of Mexico. A
tobeencountered by the general collector beca use they few insects in other parts of the world are mentioned,
containsmall or minute forms that may be overlooked, but the characters given for each group (and the keys)
becausethey are quite rare, or because they have a very apply to North American species and may not apply to
restrictedgeographic range. Such families are indicated all other species occurring outside of North America.
in most of our keys by an asterisk, and couplets con- The terms North America and North American refer to
tainingsuch groups can often be skipped by the begin- that portion of the continent north of Mexico. Where
ningstudent. the geographic range of a group in North America is
Measurements in this book are given in metric more or less limited, information on this range is given.
units.Tables for converting these to English units fol- Groups for which there is no information on range are
lowthe table of contents. widely distributed in North America.


Borror,D. J. 1960. Dictionary of Word Roots and Combining lnternational Commission on Zoological Nomenclature.
Forms. Palo Alto, CA: Mayfield, 134 pp. 2000. International Code of Zoological Nomenclature,
Brown,R. W 1978. Composition of Scientific Words. Wash- 4th ed. London: The International Trust for Zoological
ington, DC: Smithsonian lnstitution Press, 882 pp. Nomenclature 1999,306 pp.
Crowson, R. A. 1970. Classification and Biology. New York: Mayr, E., and P. D. Ashlock. 1991. PrincipIes of Systematic
Atherton Press, 350 pp. Zoology, 2nd ed. New York: McGraw-Hill, 475 pp.
Eldredge,N., andJ. Cracraft. 1980. Phylogenetic Patterns and Schuh, R. T. 2000. Biological Systematics: PrincipIes and Ap-
the Evolutionary Process: Method and Theory in Com- plícations. Ithaca, NY: Comstock Publíshing Associates,
parative Biology. New York: Columbia University Press, Cornell University Press, 236 pp.
349 pp. Stoetzel, M. B. 1989. Common Names of Insects &: Related
Hennig,W 1950. Grundzüge einer Theorie der phylogenetis- Organisms 1989. College Park, MD: Entomological Soci-
chen Systematik. Berlín: Deutscher Zentralverlag. ety of America, 199 pp.
Hennig,W 1965. Phylogenetic systematics. Annu. Rev. Ento- Wiley, E. O. 1981. Phylogenetics: The Theory and Practice of
mal. 10:97-116. Phylogenetic Systematics. New York: Wiley, 439 pp.
Hennig,W 1966. Phylogenetic Systematics. Urbana: Univer- Winston,j. E. 1999. Describing Species: Practical Taxonomic
sity of lllinois Press, 263 pp. Procedure for Biologists. New York: Columbia University
Press, 518 pp.
- ;;----

4 Behavior and Ecology1

stimulus-response rule engage large suites of reflexes,

Thewhatimportance of insects is detennined largely by
they do. The accounts of various insect groups involving dozens or hundreds of sensory receptors and
in this book saya good deal about their behaviors and muscle sets. These combined reflexes result in a coor-
natural histories. In this chapter we discuss and illus- dinated group of movements that perfonn some useful
trate general principIes of insect behavior and ecology element of behavior. An example is food chewing,
and draw together the kinds of activities common to which consists of coordinated movements among man-
insects in general. dibles, maxillae, hypopharynx, and other structures
that handle and shred food after it is bitten off but be-
fore it is swallowed. The stimulus that triggers a par-
ticular complex of movements is often called a re/easer.
Properties of Behavior The resulting behavior, once initiated, is exactly the
same each time it is perfonned; that is, it is stereotypic
A common misconception is that compared to verte- and is called a fixed-action pattem. Most motor pat-
brates, insects have simple and limited behavioral terns, however, vary according to stimulus input, both
repertoires, because insects are constrained by small when they are initiated and while they are being per-
nervous systems and short lives. Yet, like other ani- fonned. Thus, even though the end result is the same,
mals, insects are not predictabIe automatons. They do the insect's behavior is adjusted to meet the precise cir-
not remain inert until provoked by particular stimuli to cumstances of the situation. These modal-action pat-
perform specific acts, and when stimulated to respond tems are less stereotypic, because they constantly adapt
in a characteristic way, they are not limited to a rigid to changes in body position relative to external objects.
preprogrammed set of movements. Instead, much of Thus insects walk or fly according to a very specific se-
insect behavior is outwardly spontaneous and can ad- quence of leg or wing movements, operated by a cen-
just to particular circumstances. The jobs insects carry tral pattem generator, but they continuously make fine
out during their lives are the result of a complex in ter- adjustments in these movements to stay on course over
play between environmental stimuli, internal sta te, and rough terrain or in air turbulence. Even nest building,
experience, and the way their activities are organized which is a complicated process and may be interrupted
often is exceedingly complex. by the need to gather more nest material or to feed or
rest, is a modal-action pattern in the broad sense. Piece
Composition and Pattern by piece, the insect adds nest material where it is
The basic unit of insect behavior is the reflex: One kind needed most, adjusts for damage, and works around
natural obstacles. However, the insect adds material to
of receptor, when stimulated, causes a specific group of
the nest in a species-specific way, and completed nests
muscles to contract, outwardly visible as a body move-
of a species are nearly identical.
ment. Yet even the simplest behaviors following this
Orientation is the ability of an insect to be guided
by local external circumstances. Technically, it is the
62 'This chapter was contributed by Woodbridge A. Foster. modification of body position or movement with re-
I~h;'" ,~,,~, ..~..' ,
t:.Jl\\t~~ ;",.~",'-"J;~ ': (~:

Propertiesof Behavior 63

spectto somevariation in the distribution of environ- lus is lost, the insect ceases to fly upwind and makes er-
mentalstimuli. Even behavior patterns that start spon- ratic turns or reverts to a crosswind or downwind
taneously,such as the search for food or a mate, never- searching flight. In moths, males progressing upwind
theless typically are guided by the arrangement of in the presence of female sex pheromone automatically
variousvisual, chemical, or mechanical stimuli. In- zigzag across the odor plume continuously, making it
sects'oriented movements are categorized as either ki- more difficult to lose the scent altogether.
"esesor taxes. Kineses are simply changes in the speed Some kinds of orientation remain enigma tic. The
oflocomotionor rate of turning as an insect moves and migration of monarch butterflies is noteworthy because
encountersan increased or decreased stimulus in ten- they trave! thousands of kilometers between the north-
sity,withoutregardto the direction, source, or gradient ern reaches of their distribution and very localized
ofthe stimulus. Kineses have the effect, through ran- southern overwintering sites. The slow northward mi-
dommovements, ofbringing the insect into a favorable gration involves up 10 three successive generations of
zoneor away from an unfavorable one. For example, butterflies. Thus all adults that overwintered are dead
an insectexhibiting positive klinokinesis with respect by the time the population reaches its northern limit
to temperature makes more turns as it encounters in- during the summer, and those that fly back to the
creasinglyfavorable temperatures, so it tends to remain overwintering site in the south have never been there
inzonesof optimal temperature. Taxes, in contrast, are before. Research evidence indicates that they use a
positionsand movements with respect 10 the source, time-compensated sun compass, and perhaps also geo-
direction,or gradient of a stimulus (for example, magnetic cues, to guide them toward the correct longi-
chemotaxis,phototaxis). Different mechanisms may be tude and latitude. If this is correct, it suggests that the
involved.Insects can move up or down a stimulus gra- monarch butterflies have an innate sense of what those
dient(towardor away from its source) by balancing the coordinates should be, which vary considerably be-
stimulion the right and left, either by turning alter- tween eastern and western populations.
natelyright and left or by sampling both sides simulta-
neously(klinotaxis and tropotaxis, respectively), or by Behavior Modification
maintaininga "fix" on the source itse!f Ctelotaxis). They
alsocan use te!otaxis to maintain themse!ves at a fixed All behavioral abilities of an insect are inherited, but
anglefrom the stimulus source (menotaxis). This al- they are not invariably expressed, even under identical
lowsthemto remain right side up with respect to sky- environmental conditions. Insects respond to certain
lightor gravity and to walk or fly in a straight line stimuli or perform particular behaviors only when in-
throughvaried landscape, using the sun or moon as a ternal organs, hemolymph chemistry, and the nervous
fixedframe of reference-essentially as a compass. system itseU are in particular states. The state of re-
Thesebasic kinds of orientation mechanisms, applied sponsiveness, or motivational state, regulates the ex-
toproblemssuch as nest building in a difficult spot, al- pression of even the most rudimentary and stereotypic
lowan insect to behave effective!y in a complex and instincts. Even learning, the ability of an insect to mod-
changingworld, even when the guided behavior is en- ify its behavior adaptively, following experience with
tirelyindependent of experience. particular external stimuli and their consequences, is
When insects locate food and mates through vi- an innate capacity.
sion,sound, or chemicals, accurate orientation is es-
Motivation: A variety of internal factors influence an
sentia\.They typically use tropo taxis or telotaxis 10 ori- insect's motivational state.
ent toward attractive sounds and visual objects at a
distance.At close range they follow gradients of a Rhythms: Nearly all insects go through cycles of
chemical,temperature, or humidity by employing activity and inactivity, the most common type being the
klinotaxisor tropotaxis. However, the need for flying 24-hour die! (daily) cycle. During its active phase, the
insectsto locate a distant source of volatile chemicals insect may become active spontaneously and conduct
presentsa special problem. It is impossible for them to searches for one or more necessities: mates, food,
geta fix on the source by scanning the environment, oviposition sites, or shelter. Also, at this time it is re-
andstimulusgradients are lost beyond a few centime- sponsive to particular stimuli that it ignores when in-
tersfromthe source, as chemicals waft downwind in active. These die! rhythms are maintained by an inter-
dilute,meandering swirls and filaments. Instead, they nal clock, which is repeatedly set by the periods of dark
orientupwind (positive anemotaxis) in the presence of and light. When an insect is artificially deprived of the
pulsesofthe stimulating odor, guided in their forward natural time-setting light-dark transitions at regular
progressby the passage of objects on the ground be- intervals, the rhythm continues but drifts away from
neaththem.This behavior has the effect of bringing the 24-hour period, giving rise to the term circadian
themverycloseto the odor's source. If the odor stimu- rhythm (around a day).

64 Chapter4 Behaviorand Ecology

Inhibitory Feedback: An insect's state of respon- programmed and modified behaviors, so four cate-
siveness changes when particular behaviors cause a gories of behavior are useful (based on Alcock 1979):
temporary physiological change. After eating food, an (l) closed instincts, the fixed programs, such as
insect's threshold of response to food stimuli rises, so courtship sequences; (2) open instincts, in which feed-
that it requires stronger stimuli before it wiIl taste or back from experience alters the program, such as im-
ingest more. If it has ingested enough, it wiIl not re- proved nectar-collecting effieiency or ignoring sudden
spond to stimuli of any strength. A classic example is movements (habituation); (3) restricted learning, in
the tarsolabeIlar reflex of the blow fly.When its crop is which limited stimuli alter behavior in a preeise way,
empty, if it tastes sugar with its front tarsi the fly ex- such as attaching new meanings to stimuli (classical
tends its proboseis. When the crop contains a substan- conditioning); and (4) flexible learning, in which an ex-
tial sugar meal, nerves from foregut stretch receptors perience can lead to a wide range of changes in the
block the reflex. The time it takes for the fly to become behavior pattern, such as familiarity with unique land-
motivated again (that is, has a lower threshold of the scapes (exploration or latent learning) and reinforce-
tarsolabeIlar reflex) is simply a function of crop volume ment of modified behavior (operant conditioning). The
and the rate at which the crop empties, which in turn ability to achieve any of these acts, and the constraints
depends on how fast absorbed sugar is depleted from placed on them, have a genetic basis. Modification by
the hemolymph. Some changes are the result of cyclic learning is most valuable to species that live where in-
internal processes. For example, female insects do not formation is reliable for extended periods within the
exhibit oviposition behaviors until they have a batch of lifetime of the insect, but not reliable over many gen-
eggs ready to be laido erations or over the population's entire range. Informa-
tion that remains reliable need not be learned and can
Long- Term Physiological Changes: Other be-
be committed to rigid stimulus-response programming.
havioral changes are long-Iasting or permanent. These
Thus, with instinct, short-lived insects get it right the
often are the result of development and maturation. Af- first time.
ter the final molt, adult insects do not immediately ex-
Insects are capable of most types of learning
press sexual or other reproductive behaviors, but known to animals. The first three, here, are well estab-
hormone-induced nervous or other internal changes
lished: (1) Habituation is a diminished response to a
lead to their expression and may last a lifetime. Juve-
nile hormone-a misnomer in these cases--often is the stimulus after repeated exposures with no relevant
consequences or assoeiations. An example from nature
mediator. For example, juvenile hormone causes adult
is the ability of male wasps to learn to avoid being
female mosquito es lO become sexuaIly receptive, and
tricked into "mating" with orchid flowers that crudely
after mating and insemination they immediately be-
mimic female wasps in appearance and odor. In captiv-
come unreceptive to further sexual advances by males.
ity, insects usually habituate to startling noises, move-
First mechanical stimuli from copulation and insemi-
nation, then chemical stimuli from the male's semen, ments, and handling. (2) Associative learning takes two
major forms: (a) In classical conditioning, a previously
abolish her receptivity for a long period or a lifetime.
neutral stimulus, when paired with a favorable or un-
The tendency to migrate or to enter diapause (dis-
favorable stimulus that normaIly causes a speeific re-
cussed later in this chapter) may occur only early in
sponse, becomes a conditioned stimulus, elieiting the
adult life, only when the insect is crowded during de-
response by itself (the new stimulus becomes associ-
velopment, or only when photoperiod shortens and ated with the old stimulus because of its effect). For ex-
temperatures begin to drop. Complete metamorphosis
ample, a honey bee, instinctively attracted to a blue
is an extreme example of developmentally altered be- flower that has a neutral odor, obtains nectar and sub-
havior. Larval and adult forms of the same individual
sequently is attracted to that odor when the blue color
are often so different, both morphologically and eco-
is absent. A variation of this is preimaginal conditioning,
logically, that they act like different speeies with en- in which a neutral stimulus that an immature insect
tirely different behavioral repertoires.
encounters while feeding on a suitable food, such as an
Learning: Insect learning generally is the experience- unusual host plant, becomes the stimulus that, after
induced modification of instinctive behavior patterns, metamorphosis, the adult female chooses for oviposi-
rather than formation of new behavior patterns. Learn- tion. This is similar to imprinting, weIl known in birds
ing differs from instinct mainly in that it involves prop- that become attached to the appearance of their par-
erties of the nervous system that permit useful modifi- ents, but imprinting has a narrow sensitive period soon
cations in behavior, but the distinctions among after hatching and occurs without an assoeiated favor-
motivational change, sensory adaptation, and true able or unfavorable stimulus. (b) Operant conditioning
learning are not always obvious. Even in clear cases of is a particular act, followed by a positive or negative
learning, there exists a continuous range between pre- consequence that reinforces either performing or the
Properties of Behavior 65

refrainingfrom that act (the act becomes associated direct route home or to a feeding station, from places
withits consequence); a particular variant of this is they have never been before. This requires that a bee
sometimesknown as "trial-and-error learning." For ex- learn the position of distant landmarks relative to one
ample,ants can learn how to make the appropriate left another and then interpolate the correct direction to its
andright turns in a maze, without the help of a trail destination when it views the landmarks from a unique
pheromone,if the reward is to reach their nest at the perspective.
endof the maze. (3) In latent leaming, an insect be-
comesfamiliar with the relationships among various
Temporal Structure of Behavior
neutralstimuli, even though these stimuli have no im-
mediatepositive or negative consequences. It is also Insects and other animals generally can do only one
cal\edexploratory leaming. A classic example is the thing at a time, and their various behaviors are not per-
abilityof digger wasps to memorize the configuration formed in some haphazard sequence, although often
oflandmarks around their hidden nests so they can there is an element of randomness in the probabilistic
findthemmuch later (Figure 4-1). (4) In insight leam- ordering of different activities. Instead, they are guided
ing,the animal combines information from several by sets of rules that are assumed to maximize repro-
leamingexperiences and applies it to a situation never ductive success over the long run. From a study of
beforeencountered. Whether insects are capable of this these rules, a basic principIe emerges: Different behav-
veryadvanced form of learning is controversial. They iors compete with one another and are mutually in-
areclearlycapable of computations based on a multi- hibitory, and at any given time one of them is domi-
tudeof sense data, so there is no a priori reason why nant. Dominant or subordinate status is a property of
theycannot process learned information from several the behavior's causal system, which is based on two
sourcesand use it to meet new circumstances. The best things: the insect's internal causal state (or motiva-
evidencefor cognition of this sort comes from tests of tional state) and its external stimulus situation. Thus,
theability of honey bees to form mental maps. Some whatever an insect "decides" to do (with no implica-
experimentsindicate that bees can figure out the most tion that learning-based cognition is involved) reflects
the dominant status of one of the causal systems. A
change in behavior may occur in four instances: (1) In

competition, a formerIy inactive behavior system be-
comes active beca use of either internal changes or
changes in external stimuli, causing a rise in its domi-
nance status, that is, increased tendency to perform or
lowered threshold for expression. (2) In chaining, per-
formance of the dominant behavior causes feedback
that reduces its status, allowing dominance and ex-
pression of a behavior previously subdominant and
therefore inhibited. Fixed chains are deterministic, and
at the finest level they may be caused by (a) sequential
releasers (as in the case of appetitive sequences, see
later discussion in this chapter), in which each behav-
ior leads to an encounter with the next releaser; (b) in-
terlocking releasers, in which each behavior causes the
next releaser (for example, the queen butterfly
courtship, in which male and female interact in a chain
of alternating behaviors); and (c) inflexibly linked se-
quences, in which one behavior follows inexorably after
another, regardless of feedback (for example, the series
of elements in some male courtship displays). The al-
Figure 4-1 FemalediggerwaspAmmophila(Spheci- ternative to fixed chains are Markov chains, in which
clae)at the entrance to its underground nest. Initially, she individual behaviors follow one another in a proba-
becomesfamiliar with its position while digging. When bilis tic manner, according to what behaviors preceded
aboutto leave, she hides the nest by filling the opening it and how long ago each one occurred, a pattern that
withsand grains evenly with the ground's surface. As she occurs often in insect grooming movements. (3) In an-
fiiesaway,she updates her memory of its position rela- tagonistic induction, an insect prevented by circum-
tiveto surrounding landmarks, so that she can find it stances from performing a behavior is more likely to
when she returns. perform it when the opportunity occurs. For example,

66 Chapter4 Behaviorand Ecology

aphids that are prevented from l1ying because of wind bernetic models), which borrow heavily from engi-
willl1y sooner than aphids not exposed to wind. (4) In neering and use 110wdiagrams, electrical schematics,
temporary switching, insects express competing behav- and computer-like notations. These more closely rep-
iors randomly or according to a time-sharing method resent nervous systems, are more l1exible (they can eas-
that operates independently of dominance. ily depict both hierarchical and weblike arrangements
Conceptual models have been developed to help of behaviors), and minimize the number of assump-
people understand the organization of these underly- tions required. They are formed from basic conceptual
ing roles. The simplest one with broad application to patterns of control, such as chains, meshes, and posi-
insects was developed by European ethologists in the tive and negative feedback loops. For example, they
mid-I900s and assumes that behavioral control is have been used to explain the mechanism by which a
strictly hierarchical. At the top of the hierarchy are the praying mantis can strike with its fore legs in the cor-
most basic alternative causal systems (or "centers"), rect direction toward prey, even though its head moves
such as reproduction, feeding, and migration. Within independently of its thorax and can be held at different
each basic system are levels of competing subsystems angles. A broader approach employs optimality theory,
and sub-subsystems, each subordinate to the level the general evolutionary assumption that animals eval-
above it, sharing common causal factors and a common uate their causal-sta te variables and tend to make deci-
goal but controlling finer elements of the behavior. For sions that minimize costs and maxirnize gain according
example, in the reproductive mode an insect may per- to some criterion. This type of rnodeling can incorpo-
form mating, nesting, oviposition, or food gathering rate probabilistic roles and take a variety of mathemat-
for offspring, but only one task at a time. A male in the ical forrns. The dynamic stochastic modeling method,
mating mode may engage in territoriality or mate pur- widely used in behavioral ecology, relies heavily on
suit, but not both simultaneously. The mate-pursuit computer analysis to predict sequences of decisions.
system controls behaviors that lead progressively to- The simple ethological model can account for
ward the final sequence: courtship, mate acceptance, many aspects of an insect's behavioral organization,
mounting, copulation, and insemination. At each level such as spontaneous behavior, the shifting dominance
the competing behaviors are under the control of a par- of different behaviors, and sequences that lead to a
ticular system. The insect's readiness, tendency, or re- goal. A famous example of an appetitive sequence is
sponsiveness (states giving rise to the unfortunate con- predation by the bee-wolf Philanthus, a digger wasp
cept of "drive") for each behavior is an outcome of that specializes in catching honey bees to provision its
internal and external stimuli and central excitatory nests. lt starts when the wasp l1ies from 110wer to
state. According to this hierarchical hypothesis, the ex- 110wer,searching for a bee. When it sees a moving ob-
pression of the behavior controlled by each system at ject, the visual stirnulus releases the behavior of hover-
each level is blocked until its postulated innate release ing downwind of the object. If bee odor is perceived,
mechanism receives a particular stimulus, its releaser, that releases seizing the bee. And if tactile stimuli (ei-
which often is a simple "sign stimulus" or "token stim- ther mechanical or chemical) are beelike, then stinging
ulus." Between successive steps in this hierarchy, ap- behavior is released. Thus the goal is achieved through
petitive behavior is expressed, in which oriented move- a series of behaviors, each of which prepares the inseCl
ments bring the insect to the next releaser. When the for encounter with the next releaser. A similar series is
goal has been reached and the very stereotypic consum- performed by beetles locating host plants for oviposi-
matory act is performed, feedback to the system re- tion, mosquitoes locating vertebrate animals for blood
duces its motivational status, relative to other behavior feeding, and male moths locating pheromone-releasing
subcategories, and one of those others becomes domi- fernales in order to mate. In the Philanthus case, re-
nant. searchers can assume that stinging the prey is the con-
Hierarchical models have been greatly expanded, summatory act, which lowers the motivation to collecl
and therefore complicated, to account for feedback prey, so one of the competing behaviors connected
loops and the proba bilis tic nature of behavioral expres- with reproduction now becomes dominant, probably
sion. However, as models become progressively more oviposition, a sequence involving taking the prey to a
comprehensive, they lose their heuristic value, and hy- nest and laying an egg on it. This process also has been
potheses about particular insects become difficult to studied in detail. A case made famous by]. Henri Fabre
formula te. Alternatives to hierarchical models have is Sphex,a wasp that digs a nest in the ground, then
been used chiel1y to test and explain one specific ma- finds and stings a grasshopper and brings it to the nest
neuver, sequence, or choice between two alternatives, entrance. The wasp uncovers the entrance, makes an
rather than to offer guidelines for understanding the inspection visit of the tunnel that terminates in a cham-
organization of an insect's entire behavioral repertoire. ber, then returns to the surface. She grasps the prey by
One of these approaches is control system models (cy- its antennae, pulls it down into the chamber, lays an
Properties of Behavior 67

eggon it, then exits the tunnel, covers the entrance, an inspection. Lack of feedback from the previous in-
andleaves.Aside from raising the question of whether spection makes her incapable of improvising in this
preycapture and nest provisioning is one or two ap- rare circumstance.
petitivesequences,the Sphcxstudy is revealingbecause The complexity of the en tire repertoire of nesting
it tested some of the limits of flexibility in the se- behavior and its hierarchical organization is epito-
quence.lf the prey is moved a few centimeters away mized in some digger wasp species (Figure 4-2). Am-
fromthe nest entrance while the wasp is making her mophila adriaansei females can maintain several nests
inspectionvisit, when she comes to the surface she at the same time, in various stages of founding and pro-
drags thepreyback to the entrance and makes another gressive provisioning with caterpillars. Experiments

inspectionvisit. She will do this repeatedly, indicating have demonstrated that, for each nest serviced, there
a tightlink between bringing the prey and conducting are three levels of subsystems within the reproductive

t ~~ I
r I
M 111 ... I









Figure4-2 Hierarchical control of nest provisioning in the digger wasp Ammophila adri-
aansei (Sphecidae). Nesting behavior is pan of the reproductive behavioral system (Rp),
which competes with the maintenance system (Mt). Several nests (a, b, e, d, e) are in
memory (M) at one time, but during a phase of activity on a nest, attention is not diverted
to other nests. Phases 1, U, and III are characterized by the amount of food (caterpillars)
required, and they utilize different combinations of subsystems and sub-subsystems.
Founding occurs only in Phase 1and Final CIosing only in Phase IlI. Each phase begins
with lnspecting. An evaluation of the presence of a larva, its size, and the amount of food
remaining determines, by heterogeneous summation (HS), which of the three phases wiIl
occur. If no larva is present, or if a phase is completed, the wasp scans (S) its memory and
redirects attention to the oldest uncompleted nest. If none needs funher provisioning at
the time, a new one is founded and its position learned (L) during digging.
68 Chapter4 Behaviorand Ecology

"system." The first level consists of three phases, each assume that the way insects organize and distribute
with its own motivational state and each beginning their behaviors is roughly optimized for a particular in-
with an inspection visit prior to foraging for one or sect niche. Using this kind of thinking, we can divide
more larvae: Phase 1, one caterpillar stored, foIlowed insects into the time minimizers, which are favored by
by oviposition; Phase 2, one to two more caterpillars spending the least time gathering food but the most
provided to young larva; Phase 3, five to seven cater- time defending it (for example, territorial ants or bury-
pillars added and nest closed for final time. Within ing beetles), and the energy maximizers, which gain an
each phase are four subsystems (founding, provision- advantage by spending most of the time available gath-
ing, final closing, and defending), founding occurring ering food until it is nearly gone, then dispersing to
only in Phase 1, final closing only in Phase 3. And the find more (for example, blow flies that develop in car-
four subsystems, plus inspecting, are expressed in one casses). These extremes in a continuum of time budget
or more still finer subsystems: digging, closing, hunt- strategies fit various modes of insect life in different
ing, and transporting. Each of these consists of two to kinds of environments. In some ways they correspond
six distinct behaviors, such as the appetitive sequence to the oversimplified life history dichotomy based on
of events involved in hunting just described. During reproductive strategies: r-strategists produce huge
the inspection visit, the status of the larva or remaining numbers of cheaply made eggs, but eggs and immature
prey determines the total amount of provisioning that stages suffer massive mortality (for example, floodwa-
wiIl be done before the next phase commences. In be- ter mosquitoes); K-strategists invest large amounts of
tween two phases for one nest, a phase is performed for time and energy on relatively few eggs and offspring,
one or even two other nests. particularly remarkable is whose survival rates are high (for example, tsetse flies).
the conclusion that the motivational state of each sys- The r-strategists grow quickly and are good colonizers
tem persists independently for each of several nests be- of unexploited resources, whereas the K-strategists
ing cared for simultaneously, each with its own memo- grow slowly but are successful against competitors and
rized location. Furthermore, the wasp juggles these natural enemies.
reproductive activities with competing maintenance
Conditional Strategies: In the course of a life cycle, an
activities, such as feeding, grooming, sun basking, and
insect shifts between two broad phases, each with its
own set ofbehaviors and priorities: vegetative, in which
the immature insect is committed primarily to feeding,
Life History Strategies growth and storage of food, and reproductive, in which
the adult insect is devoted primarily to dispersal and
Time and Energy Budgets: The lives of insects, at the
reproduction. In addition, three tactical alternatives
broadest level, are organized to produce the greatest
may occur in either phase: active, migrating, and dor-
number of successful, mature offspring. This number
manto These are induced by particular environmental
defines an insect's fitness. There are many ways to max-
conditions. Active insects carry on their vegetative or
imize fitness, but they aIl involve economizing time
reproductive functions because conditions allow it.
and energy, the common currencies of life. Time and
Both the ability to migrate and the ability to be come
energy budgets are shaped, evolutionarily, according to
dormant allow insects to exploit temporary resources.
the principIes of stringency and aIlocation. This means
Dormancy occurs when the insect remains in place but
that the budgets are geared to fit the worst conditions
endures inimical conditions, such as winter, dry sea-
an insect might encounter and that time and energy are
son, or unpredictable drought, when growth and re-
aIlocated among survival and reproductive activities so
production are impossible. Insects may behave in par-
as to maximize fitness. Each sex of an insect species
ticular ways to prepare for dormancy but are otherwise
typicaIly aIlocates a certain portion of its time carrying
inactive. During dormancy the insect reduces its meta-
out several necessary activities, such as feeding, nest
bolic rate and exists in one of two distinctive physio-
making, mating, grooming, and quiescence. Quies-
logical states: quiescence or diapause (see Chapter 2).
cence serves to avoid natural enemies, inimical
weather, or stressful times of day, while important in- Migratory Behavior: Like dormancy, migration often
ternal activities continue, such as food digestion. The occurs because of inimical conditions, but in this case
energy obtained by feeding is likewise budgeted among the insect escapes in space instead of time, by moving
several activities that compete for energy use, such as to a better environment. These migrations may be ei-
metabolic activity, formation of sperm or eggs, locomo- ther round trips or one-way trips, and unlike migratory
tion, and behaviors for obtaining specific nu,trients birds, the round trip may be achieved by members of
finding mates, and defense. The best budgets are those two separa te generations. Well-known migrations ac-
that lead to the largest numbers of offspring and be- commodate either the seasonal deterioration of an in-
come propagated through natural selection. Thus we sect's resources or the shrinkage or overexploitation of

Properties of Behavior 69

resources.The famous monarch butterfly, Danaus plex- their local resources, mainly insect prey. During their
ippus(L.), migration is a seasonal round trip between statary phase, the bivouac remains in one place, and
northernNorth America, including southern Canada, daily foraging parties extend in all directions for many
andits more mild southern reaches. In this migration, days. Migration also occurs for reasons other than re-
dormancyplays an integral part. Ihe southward-flying source deterioration. Some insects make a round trip
butterfliesare in a state of reproductive diapause dur- between two different kinds of resources that are spa-
ingtheir approximately 3,300-km (2,000-mile) jour- tially separated, or simply make a one-way dispersal to
ney,although they continue to feed on nectar. In their colonize new areas. Saltmarsh mosquitoes, such as
mainoverwintering habitats in coastal California and Aedestaeniorhynchus (Wiedemann), which hatch and
centralMexico, they are usually torpid but will fly and develop in synchrony after an exceptionally high tide,
drinkwater on warm days. Only in the spring, when do both: They go on a spectacular postemergence exo-
theystart the northward journey, do they become re- dus that results in dispersal, and once they settle, the fe-
productivelyactive, breeding on milkweed plants. Up males translocate between two habitats during each re-
10 three generations are passed during this more productive cycle: Ihey often find their blood meals
leisurelyspring migration. Many other insects that mi- inland but must return to the saltmarsh to lay their
grateseasonally simply die out at the end of their eggs. Ihe swarming dispersal of newly emerged winged
northernmigration as winter approaches, rather than termites is an example of a purely colonizing migration.
retumingsouth. Quite different are the migrations of
thenotorious locusts, such as the desert locust, Schis- The Genetic Basis of Behavior
tacerca gregaria(Forskal), which are species of grass-
hoppers(Acrididae) capable of developmental dimor- Both instincts themselves and the ability to modify
phism.They perform a one-way trip with no diapause, them through learning have a genetic basis. A great
noparticulardestination, no periodicity, and no finish- many insect behavioral traits are demonstrably herita-
ingtime.The root of migration is a shrinking suitable ble, as shown by the methods of classic genetics, such
habitat,which causes crowding, and mutual stimula- as hybridization and backcrossing, mutation, and arti-
tionofgrasshoppers leads 10 their transformation from ficial selection. Entomologists have explored traits' 10-
a solitaryphaseto a gregarious phase. The complete calized expression in the insect body through the use
transfonnationtakes up to three generations, and the of sexual mosaics (gynandromorphs) and biochemical
gregariouslocusts are strikingly colored and behave analysis, and have demonstrated their molecular basis
quitedifIerentlyfrom solitary ones. The gregarious forms through genome manipulation. Identifying heritable
remaina cohesive unit, even when their swarms num- units that underlie behavior has been difficult. Ihis is
berin the tens of billions and cover up 10 1,000 km2. partly because behavior itself is not easily fractionated
Theymay travel across continents, eating plants and into single elements, each caused by the expression of
reproducingas they go. If unchecked, they eventually one or a few genes. Unlike color or so me morphologi-
arecarriedby wind to regions where rain is falling and cal features, which may be expressed through a unitary
forageis growing over wide areas, so they become dis- developmental process, behavior requires the partici-
persedand revert to the inconspicuous solitary phase. pation of sensory receptors, nervous circuitry, appro-
Locustmigrations, known throughout all history, are priate neurochemical states, and a system of muscles to
stilla serious problem in Africa and the Middle East. move body parts. A small difference in the genes that
Locustsalso occur in China, South America, and Aus- control any of these components can result in a differ-
tralia.The species that plagued the North American ence in behavior. Research indicates that even in cases
plainsin the 18005 is now extinct. Like locusts, when where a genetically based difference between two in-
aphidsbecome too crowded on a plant they begin de- sects can be traced to a specific difference in neural cir-
velopingwings and migrate to new plants, also a one- cuitry or neurochemistry, that difference usually is the
waytrip. The emigration of some aphids has a round- result of many genes having multiple effects; that is,
trip aspect, because they migrate from a spring the trait is polygenic and the genes have pleiotropic ef-
host-plantspecies to a summer one, the shift being fects. Among the traits studied in greatest detail are
causedby a physiological change in the spring plant. cricket calls, honey bee food hoarding, honey bee hy-
Theaphids return to the spring host species the fol- gienic behavior, and a variety of behaviors in the fly
lowingyear. Tropical army ant (for example, Eciton) Drosophila: geotaxis, larval foraging, odor learning, cir-
coloniesmake impressive daily raids from a central cadian activity rhythms, sex recognition, and courtship
bivouac but are nomads on a perpetual one-way trip. songs. The hygienic behavior of worker honey bees in
Duringtheir nomadic phase,the entire bivouac, queen colonies that are resistant to foulbrood bacteria was
andall,must be moved daily while the larvae are grow- found, by backcrossing experiments, to consist of two
ing,because these carnivores are continually depleting separa te acts, uncapping cells of infected larvae and re-
70 Chapter4 BehaviorandEcology

moving infected larvae from the hive. These behaviors tecture, a permanent record of behavior that can be
assort independently, and analysis indicates that prob- gathered and stored, even from fossils. Independently
ably at least three genes are involved. of their use in understanding relationships, behavioral
In some behaviors, researchers have discovered characters can be applied to phylogenetic reconstruc-
simple one-gene control. In the case of larval foraging, tions based on morphological and molecular data.
two behavioral tYpesof DrosophilamelanogasterMacquan From them, researchers can infer the evolution of be-
have been identified: rovers, which wander around on havior by its appearance and modification among vari-
their food, and sitters, which travellittle. These differ- ous groups in the evolutionary tree. Similarly, the ob-
ent types occur naturally, each one being favored by servation of polymorphic behaviors within one
different food conditions. Hybrids between rovers and recognized species has led to the discovery of distinct
sitters produce offspring in a 3:1 rover-sitter ratio, sug- but cryptic species within what then becomes recog-
gesting that foraging is controlled by a single gene nized as a species complex.
(dg2) and that the sitter trait is recessive. The two be-
havioral types have been mapped to a single locus on a
chromosome. Apparently differences in the gene (that
is, its nudeotide sequences in the DNA) result in dif- Behaviorallnteractions between
ferences in the way RNA splicing occurs, giving rise to Conspecifics
different kinds and amounts of the enzyme protein ki-
nase. Protein kinase affects the excitability of nerve
cell5, which probably causes food receptors or foraging-
movement neurons to Cire more rapidly. Experimen- SexualEncounter: The vast majority of insect species
tal transfer of dg2 DNA from rovers to sitters causes sit- are sexually dioecious, occurring as either female or
ters to behave like rovers. In the case of circadian-dock male, and most engage in mating, even those whose
genetics, researchers have found mutant Drosophila females have the option of parthenogenesis (see Chap-
strains that differ from the normal 24-hour rhythm of ter 2). Mating presents the problem of finding a recep-
activity by having 28-hour or 19-hour cydes or no cy- tive member of the opposite sex, and insects have two
des at all. Each mutant differs only at the per locus and solutions. (1) In sexual aggregation, individuals that
only by a single pair of nudeotides among a large need to mate convene in special sites, often at certain
group that codes for a particular protein that differs by times of the day or night, during the mating season.
a single amino acid. The per locus also is responsible These sites may be species-specific places of emer-
for very specific tonal features of the male Drosophila gence, feeding, or oviposition, or distinctive land-
courtship song that are important to the female for marks. The aggregations typically have an operational
species recognition and mating speed. These features sex ratio strongly skewed toward males, beca use much
have been traced to a very small part of the gene. The of a male's fitness depends on his ability to obtain
species-specific songs of D. melanogaster and D. simu- mates, whereas a female's fitness is distributed more
lans Sturtevant apparently derive from a difference of broadly among activities connected with feeding and
only four amino acids in the protein translated from the nesting, and she may have less to gain from multiple
relevant region of the gene. However, the entire male matings. Because of their intense competition, the
courtship ritual involves a series of female-oriented and males of many species establish territories that they de-
female-behavior-dependent movements: facing, tap- fend against other males. The territories can have ac-
ping, cirding, wing-extending, licking, and mounting. tual value to females for feeding or oviposition, as in
These must involve many other loci, all contributing to the case of dragonflies, or they may be leks, symbolic
properties of the nervous system that allow this se- territories that have no intrinsic value but are sought
quence to unfold. by females and therefore are prized by males and de-
Because behavioral traits, like morphological and fended by those with the best competitive ability-for
molecular ones, are heritable and shared among related example, Hawaiian Drosophila on plant leaves. High-
taxa, they are used as tool5 in insect systematics. De- density sexual aggregations over landmarks usually are
spite a reputation for evolutionary flexibility, behav- aerial, the males forming a seething mating swarm that
ioral characters show approximately the same range, makes establishment of a territory virtually impossible.
from stability to volatility, as morphological ones. Nest In that case, competition is based on a male's ability to
building and courtship displays have been particularly loca te a female quickly as she approaches or enters the
useful, beca use they are rich in complexity, providing swarm, to seize her, and to deny other males access to
many different traits that can be measured or catego- her before the couple departs. Male-biased mating
rized, independently of one another. Nesting offers the swarms are typical of many ants and small nematocer-
additional advantage of leaving behind, in nest archi- ous Diptera, such as gnats, midges, and mosquitoes.
Behaviorallnteractions between Conspecifics 71

(2)In sexualattraction, one sex of a speciesmay broad-

casta visual, chemical, or auditory signal indicating ~4 . ... ...
that the signalerwants to mate, and members of the 3
oppositesex use the signal to find its source. To pre- 2 Á
eludeuseless hybrid matings, these signals must be 1 .
speciesspecific for a given area and season. Visual sig-
nalsare transmitted in the flashing colors of butterflies'
~5 ~d~~IIII~
wingsin sunlight and in the blinking lights of fireflies' ~3
photicorgans. In common firefly species, both males "'02
andfemalesproduce light, the flying mal e giving a pe-
gu 1
riodic,species-specific calling flash, and the sedentary
female answeringwith a flash from the ground. In this O)a
case,it is the female that attracts the male, by answer- 0.5
ingeachof his calls after a prescribed interval, allow-
inghimto locate her. Malesof some Pteroptyx species,
bestknown in Southeast Asia, gather together in trees
~2 :C
1I 1I
alongthe banks of rivers and flash in unison. This
combinedeffort is thought to be effective in attracting .S: 8
flyingfemales over long distances; at close range, indi- ga
vidualattraction still is necessary.Males of some tree 0)5
cricketand katydid species seem to do the same thing, ~ 3
bysynchronizing their calling songs. The best known u-2
auditory attractants are produced by cicadas and 1
variousOrthoptera, such as crickets and katydids (Fig- 7
ure4-3). Cicada males buzz by rapidly vibrating two 6
membranes(tymbals) within chambers of the first ab- 5
dominalsegment. The much quieter leafhoppers and 3
planthoppersalso use tymbals. Most orthopteran males
relyon wing stridulation. A scraper on one wing rubs
21 -e
O.O 0.5' 1.0' 1.5' 2.0'
alonga ridgedfile on the other, causing the wings to vi- Time in Seconds
brate.Some grasshoppers stridulate by rubbing their
legsagainsttheir foldedwings. For typical chemicalat-
traction,as in moths, females release a volatile sex
Figure4-3 Audiospectrographsof someinsect sounds.
pheromone,which is distinct for each species (even the A, Four chirps of the caIling song of a fieIdcricket, Gryl-
isomermust be correct) and often is a mixture of two
lus sp.; B, A portion of the caIling song of the tree
ormorechemicals in a specific ratio. Sex pheromones cricket, Oecanthus nigricomis Walker; e, Two 2-puIse
canelicitreactions in males many kilometers from the
songs of the northem true katydid, Pterophylla camellifo-
female caller.As a general rule, males emit attractants lia (Fabricius); D, StriduIation sounds produced by the
thatarerisky or energetically costly, such as sound, and sawyer beetle, Monochamus notatus (Drury) Ctheinsect il-
females emitattractants that are safeand cheap, such as lustrated in Figure 26-75); E, A part of the caUingsong
pheromones.Risky attractants are those that predators of the cicadaTibicenchloromera(WaIker).Frequency
andparasites can use to locate the sendero is shown here in kilohertz (or kilocydes per second);
4 kHz is approximateIy the pitch of the top note of a
Courtship:Once males and females have encountered piano.
other,they may proceed directly to copulation or
mayfirst engage in various rituals, collectively known
ascourtship.Although both sexesmay play active roles,
involvingtwo-way communication, usually the male male's body (so-called aphrodisiac), large and edible
conducts most of the overt performances and appears extensions of spermatophores, or the body of the male
lobeseeking acceptance. Obversely, typically it is the himself. In the last instance, including the widely pub-
femalethat rejects a potential partner. These rituals in- licized cases of praying mantises and black widow spi-
elude leg and wing displays (Figure 4-4), dances and ders, the male usually attempts to escape with his liCe
songs, the use of silken traps and guides, and various after inseminating the female, and the frequency of
feedingdevices. Feeding includes items of food (nup- sexual cannibalism in nature is disputed. But at least in
tialgifts),such as prey, secretions from glands on the the Australian red-back spider, the male clearly offers

72 Chapter4 Behaviorand Ecology

cept or reject particular individuals, according to how

fit they seem. Most evidence comes from studies in
which females choose large males over small ones. Per-
haps body size is among the most important messages
being communicated during visual displays, but
Drosophila research demonstrates that females also se-
lect males according to their courtship sounds. Insect
body symmetry also is important. Any courtship fea-
tllres may be indicators of underlying fitness character-
/ istics, such as ability to obtain food and resist parasites.
/' Hangingfly femaIes (Mecoptera: Bittacidae) accept
more sperm from males that present them with larger
gifts of prey, which may indica te not only his size, per
~ se, but his ability to obtain prey. Often the mate-choice
.E pathway appears to have Ied to the sexual selection for
~ males with exceptionally eIaborate appearances and
performances that have little to do with a male's health
or vigor, but rather confer mainly his ability to sire sons
Figure 4-4 Courtship of the neo tropical mosquito Sa- that aIso are stimuIating to females. Furthermore, fe-
bethes cyaneus (Fabricius), which is iridescent blue and males that choose such stimulating males are more
silver. Mating consists of three phases: precoupling, su- likely to produce daughters that are stimulated by such
perficial genital coupling, and full copulation. Within males. The combined effect is "run-away sexual selec-
these phases, the male performs a series of courting tion." An altemative to courtship is male-male fight-
stages with overt movements that involve extensive use ing, which provides for sexual selection of a different
of the midlegs, which on both sexes have "paddles." The nature. If a female attracts more than one male at a
stages always occur in this order: free-Iegwaving, swing- time, she does not necessarily choose among them. In-
ing, waving, and waggling. If the female is receptive, she stead, a fight ensues, and the female accepts the win-
lowers her abdomen during the swinging stage (shown ner. Intrasexual selection for fighting ability has re-
here), making genital coupling possible. Full copulation sulted in conspicuous horns and large mandibles on
occurs right after waving, and it is during the waggling male beetles. They serve as weapons against each other,
stage that insemination occurs. rather than as devices to defend against enemies or to
impress females, and probably are detrimental to all as-
pects of life other than fighting for mates.
himself to be eaten. The male gains from self-sacrifice
by contributing to the development of eggs in her Copulation: Copulation and/or insemination are the
body, which may become fertilized by his sperm. end result of a successful mating. In some apterygote
Courtship seems to have a number of purposes. insects, including many Collembola, not only is spenn
Among arthropods with indirect insemination, such as transfer indirect, but the male may place sper-
scorpions, some springtails, and silverfish, a dance fa- matophores on the substrate and never encounter the
cilitates the transfer of the spermatophore from male to female that picks them up. Yet various forms of non-
substrate, then to female. Among predators, such as contact and contact courtship also occur in Collem-
jumping spiders, a male's signals are thought to inhibit bola. Even copulation, which involves direct transfer of
the female's inclination to kill. Courtship interchanges sperm from male to female, may not occur between
also may allow the female to communicate to the male genitalia but rather between genitalia and some other
whether she is receptive, which benefits both sexes by structures on the opposite partner. This extragenital in-
precluding time-wasting, strenuous, and potentially semination is the rule in spiders, in which the pedipalps
damaging attempts by the male to copulate. The two of males are modified into secondary intromittent or-
most common explanations for courtship are species ganso A male deposits sperm in these small appendages
isolation and individual mate choice. If courtship dis- prior to mating and during copulation introduces them
plays are species specific, a female that discriminates into the female's genital opening. A similar arrange-
among them can avoid mating with males of other ment occurs in Odonata. In this case, the male deposits
species, and thus can avoid laying nonfertile eggs or sperm in an organ at the base of his abdomen so that at
producing aberrant offspring. Furthermore, if the fe- the time of mating, he clasps the neck of the female
male can discrimina te among male performances with his primary genitalia at the tip of the abdomen,
within her species, she also has the opportunity to ac- and she curves her abdomen ventrally forward to clasp
Behaviorallnteractions between Conspecifics 73

hissecondary genitalia and receive sperm. This strange

configurationis the frequently observed "wheel posi-
!ion" of dragonflies and damselflies. In bed bugs
(Hemiptera:Cimicidae) and some relatives, it is the fe-
malethat has a secondary sexual structure: a cleft and
associatedorgan (spermalege) in the cuticle on the
ventralside of her abdomen, which the male pierces
withhis strong, hooked aedeagus. The injected sperm
mustmigrate posteriorly to reach her reproductive
tract.Ihe vast majority of insects mate by genital cou-
pling,in which female and male genitalia become
lockedtogether while a spermatophore or free semen is
transferredto the female. The emire process may last
onlya few seconds or may go on for hours, during
whichtime the male may continue with courtship
movements.Female genitalia tend to be uncompli-
cated,whereas those of males exhibit a diversity of
structuresthat vary widely among species. Some of Figure4-5 Postcopulatory mate-guarding during
theseare devices for clinging 10 the female (for exam- oviposition by the yellow dung fiy,Scathophaga sterco-
pIe,claspers) or for transferring sperm, but many oth- raria (L.). The polyandrous females oviposit only in
ershaveno known function. A long-held view is that fresh ungulate dung, and the larger males aggregate there
elaborate male genitalia fit perfectly only into females lO mate with them. A gravid female copulates with the
ofthesame species, so that species isolation is assured, first male quick enough to catch her, usually in vegeta-
knownas the lock-and-keyhypothesis.A more recent tion close to the dung, then they uncouple genitalia and
explanationis that in most insects, these structures, she proceeds to oviposit on the dung while the male
likecourtship itseU, stimulate females in specific ways stands over her, fending off sexually aggressive males. If
sothat she may choose to accept or use more sperm there is a takeover, the female is inseminated by the new
framsomemales than others. These phenomena, re- maleoAbout 80% of her eggs are fertilized by the male
ferredto as copulatorycourtship or internal courtship, currently guarding her. Thus, the male assures his pater-
areused in cryptic female choice. This hypothesis nity by protecting the female, and guarding serves both
wouldexplain why species with polygamous females sexes by making oviposition possible.
alsohave males with more elaborate genitalia, the re-
sultofsexual selection.
or their resources are not concentrated, males typically
Mating Systems: The multitude of insect mating sys- either establish leks (buuerflies) or form massive mat-
temsamong insects appears related to the wide species ing assemblages (mosquitoes) at landmarks. In the lat-
differencesin the scarcity and distribution of their re- ter, no courtship or female choice is evident. From the
sourcesand thus to the likelihood of encountering the female point of view, monandry (having one mate) is
oppositesex, but the evolutionary origins of these sys- advantageous, especially if life is short and mating in-
temsis not always clear. Monogyny (having one female volves time, energy, or risk. But polyandry provides
mate)tends to be high when females are thinly distrib- sperm replenishment, allows access to a male's re-
utedor males are excessively abundant. Some monogy- sources and services (nuptial gifts, access to defended
nousmales guard their mates to prevent cuckoldry, food or oviposition sites, protection from aggressive
suchaslovebugs(Diptera: Bibionidae),which prevent males), and allows the female to diversify the patemity
heraccessto other males by remaining in copula for a of her offspring or possibly to select sperm from among
longperiodoOthers assist females with parental care of several matings. The conflict between the best interests
offspring,such as Nicrophorus burying beetles (Silphi- of males and females derives from the fact that sperm
dae) andLethrusdung beetles (Scarabaeidae).Polygyny are numerous and cheap, eggs fewer and expensive.
iscommonwhen there is a low male operational sex ra- This reasoning leads to the explanation for females'
tio.When females are concentrated in patches, males more limited sexual activity, higher mate selectivity, and
attemptto monopolize them by defending them
either extensive investment in parental careo Males lose liule
againstother males (for example, yellow dung flies, by making mating mistakes and usually can gain greater
Scathophaga) (Figure 4-5) or by defending territories fitness by inseminating more mates rather than by help-
thatcontain resources a female needs (dragonflies). ing to rear the offspring of one mate, especially if his pa-
Whenfemale monopoly is unlikely, because the females temity is not assured. By extension, the asymmetry be-

74 Chapter4 BehaviorandEcology

tween the sex roles also explains why males perfonn

Social Systems
most courtship displays, take the risks, do the fighting,
defend territories, and protect the females they mate Insects that fonn cooperative units have two large ad-
With. There are a few exceptions to this general sex-role vantages over insects that live alone: (1) They are quick
rule. Among giant water bugs (Belostomatidae), females to discover and monopolize resources, by communi-
of some species lay their eggs on the males' backs, and cating the location of food, mounting mass attacks on
. the males give the eggs special care until after hatching. invaders, and maintaining territories to exclude com-
A male wiIl not let a female lay eggs on his back unless petitors. (2) They can build large nests rapidly, which
she first lets him inseminate her, assuring his patemity protects them and their offspring against natural ene-
of the offspring. Females of other giant water bug mies, harsh weather, and stressful environments, and
species lay their eggs on emergeOl vegetation, and the their close association aIlows easy transmission of mu-
male guards them there. Another female may attempt to tualistic organisms.
kill the eggs and become his mate, so that he wiIl guard The rudiments of advanced sociality among
her eggs instead. Monnon cricket females court males, groups of arthropods are evident in paren tal careo In a
whose mating services are exceptionally valuable be- convenient classification, presocial insects are divided
cause the males produce large, edible spennatophores. iOlo those that are subsocial, rearing their offspring
A male weighs courting females (by assessing their heCt) alone, and those that are parasocial, sharing their rear-
and mates with the heaviest one. Heavier females pro- ing with other parents of the same generation. These
duce more eggs, so his choice results in a greater num- presocial arthropods include members of many insect
ber of offspring sired by him. orders and also crustaceans and spiders. Each subcate-
gory has gradations that end in eusociality. Eusocial in-
sects traditionally meet three criteria: Their groups
Nesting and Parental Care
contain overlapping generations, perfonn cooperative
Insect parents exhibit a broad spectrum of care for their brood care, and have a caste of nonreproductives, the
offspring. Some show none at aIl, such as stick insects "sterile caste." Within the subsocial category of preso-
that simply drop eggs to the ground beneath the vege- ciality are primitive forros that lack complete overlap of
tation they are feeding on. But the majority of solitary generations, intennediate fonns whose generations
insects do things that improve an offspring's chance of overlap, and advanced fonns in which the offspring re-
survival. UsuaIly the female alone does this, but in main with the group and share in caring for further off-
some cases the male-female mated pair coopera te. The spring, including their own. Within the parasocial
most elementary form of care is oviposition-site selec- group, which consists primarily of bees, are primitive
tion. Females of plant- and animal-feeding species with species in which females nest together, either gregari-
narrow host ranges seek hosts on which their larvae ously (separate nest entrances) or communally (a
wiIl thrive. So do insects that develop in very specific shared nest entrance). Tent caterpillars and webwonns
aquatic habitats. Beyond this, some females remain (both Lepidoptera) sometimes are considered commu-
with the eggs, guarding them from natural enemies or nal, because they build, expand, and repair their pro-
physical damage until the eggs hatch, or until some- tective nest together and cooperate on foraging expedi-
time later. A further step is taken by insects that create tions. Nevertheless, the group consists only of
a nest to protect the eggs and immatures, a nest that immatures and thus does not involve parental careo A
may be as simple as a hole in the ground or a curled more advanced parasocial fonn is quasisociality, in
leaf. Others construct elabora te structures and carry which communal nesters coopera te in providing brood
out necessary maiOlenance. Still others provide the im- careo Still more advanced is semisociality, in which the
matures with food, either just once or repeatedly as the cooperatively brooding adults include some nonrepro-
offspring develop (progressive provisioning). For ex- ductives. Yet this group abandons the nest before the
ample, burying beetles work as mated pairs to lower a emergence of the next generation, and those offspring
smaIl, dead bird or mammal iOlo a cavity in the ground start new nests elsewhere. Primitively eusocial insects
and to prepare it as a cuplike nest in which the larvae meet aIl three criteria but there is only one relatively
develop, first feeding them mouth-to-mouth and later undifferentiated sterile caste, the workers. Progres-
aIlowing them to devour the cup itself. Among solitary sively more advanced eusocials show more differentia-
wasps that huOl prey for their offspring, a range of tion among colony members, including large differ-
species perform various numbers of steps in nest build- ences in fonn and behavior between reproductives and
ing and prey preparation and provisioning. The most workers. Colonies of advanced termites may contain
advanced fonns stay with their offspring, feeding and several kinds of worker and soldier castes, and ad-
defending them and cleaning the nest until they are vanced ant societies have multiple worker subcastes,
mature and can fend for themselves. including the large majors, the soldiers, medias that do
Behaviorallnteractions between Conspecifics 75

of the foraging, and tiny minors that never leave the flightless soldiers are not sterile. And cottonwood
thenest. gall aphids do have a sterile female soldier caste to de-
Insectsknown to meet the three conventional cri- fend the all-female family in the gall, but there is no co-
teriaofeusociality are found only in the orders Isoptera operative brood careo Brood care is a necessity chiefly
(termites),Hymenoptera (bees, wasps, and ants), and of holometabolous insects that have larva e unable to
perhapsColeoptera (beetles). All termites and all ants feed themselves. Furthermore, members of a sterile
(Formicidae)are eusocial. Among the wasps, eusocials caste of many eusocial insects are not permanently
occurin some Vespidae (all Vespinae, some Polistinae) sterile but may assume reproductive duties if needed
andonespecies of Sphecidae. Among the bees, they oc- (as discussed later in this chapter). Delayed reproduc-
curin some Halictidae (some Augochlorini and some tion of adults, overlapping generations, and coopera-
Halictiniwithin the Halictinae), and some Apidae tive brood care are not uncommon in groups of other
(someCeratini within the Xylocopinae, all Apinae). kinds of animals, including birds and mammals, so the
Amongthe beetles, one species of ambrosia beetle demarcation between presociality and eusociality can
(Curculionidae:Platypodinae) seems to meet the euso- be blurred. Yet these other animals lack fixed morpho-
cial criteria. One noninsect arthropod is eusocial: logical caste distinctions, so they can be considered
Synalpheus pistol shrimp (Decapoda: Alpheidae), primitively eusocial at best. Despite its imperfections,
whose colonieslivein sponges. Amongvertebrates, the . the social classification provides a useful tool for un-
onlyclearlyeusocial animals are the naked mole rats derstanding how advanced sociality in insects may
(Rodentia:Bathyergidae), whose underground soci- have arisen. Living examples of degrees of subsociality
etiesroughly parallel those of termites, except that the and parasociality demonstrate alternate routes to euso-
ratsmayhave several reproductive males ("kings"). ciality from different directions. A third, the polygynous
The preceding scheme of presocial and eusocial family route, proposed for some Hymenoptera and per-
insectsis not entirely neat. Aphids, which usually seem haps applicable to spiders as well, combines features of
obliviousto one another on a plant stem, in some cases each, progressing from intermedia te or advanced sub-
releasean alarm pheromone when attacked, causing sociality to quasisociality as females from different gen-
othersin the group to save themselves by dropping off erations cooperatively tend their brood.
the plant (Figure 4-6). Communal caterpillars live Entomologists have used three basic ideas to explain
muchlike a society as they develop, but they cannot the emergence of societies, particularly to address the
reproduce, and adults are solitary. Australian gall evolutionarily counterintuitive origin of self-sacrificing
thrips,which live as family groups of multiple genera- sterile castes by natural selection: (l) In mutualistíc
tionsinside plant galls, have an anatomically distinct aggregation, group living and reciprocal altruism bene-
soldiercaste, as do some advanced eusocial insects, but fits all members, especially under certain environmen-


Figure4-6 A, A damseI bug, Nabís americoferus Carayon, attacking the aphid

Acyrthosiphon pisum (Harris), and the aphid secreting cornicle droplets in response to the
attack. B, Secreted droplets on the tips of the cornicles of the aphid Acyrthosiphon solani
(Kaltenback) .

76 Chapter4 Behaviorand Ecology

tal conditions; only one or a few reproduce, yet all act as primers, most of them pheromones, either stim-
have a chance to become reproductive at some time. ulate or inhibit development into a particular caste or
(2) In parental manipulation, the reproductive individ- prevent particular behaviors from being expressed.
ual (the queen) controls the nutrition of her helpless Primer pheromones commonly are transmitted during
larvae, so by restricting their food, she diminishes or trophallaxis, the exchange of food and secretions, or
eliminates their reproductive capacity. (3) In kin selec- during grooming. Mouth-to-mouth trophallaxis is typ-
tion, members of the group are closely related, so non- ical of Hymenoptera, whereas termites transfer many
reproductive members gain inclusive fitness (genetic materials through their feces, and communication is
propagation) from the success of their reproductive rel- therefore anus-to-mouth trophallaxis. The best under-
atives. The third idea offers an attractive explanation stood recruitment systems are trail making in ants and
for the fact that eusocial, female-dominated societies dancing in honey bees (discussed later). Contact chem-
have evolved at least 12 times among Hymenoptera, icals are important in allowing a worker to identify the
which all have an uncommon method of sex determi- stages and castes of its nestmates and to recognize dead
nation: Males develop from unfertilized eggs and are nestmates. In addition, the ability to recognize nest-
haploid; the females develop from fertilized eggs and mates in general is universal among social insects. The
are diploid. Because of this arrangement, hymenopter- characteristic odor of each colony is carried on the in-
ans have pedigrees in which, on average, full sisters are sect and appears to involve both genetic and environ-
more closely related to each other (0.75) than to their mental factors. An individual entering another nest,
own female offspring (0.5) or their brothers (0.25). even if of the same species, generally is attacked.
This means that a female hymenopteran can indirectly Social insects often work together to accomplish
propaga te her genes quite efficiently by self-sacrifice extraordinary tasks that suggest planning and insight.
that directly or indirectly benefits her reproductive sis- Honey bees are recruited to the best sources of nectar,
terso However, this one type of selective pressure favor- shape the cells of their wax combs into structurally
ing altruism does not account for eusociality in insects ideal hexagons, and thermoregulate their hives; ants
in which both sexes are diploid, notably termites. Re- take the shortest routes to food, make flanking move-
searchers have suggested a similar inclusive fitness ments during raids, and build neat walls of uniform
mechanism for termites, the result of alternating peri- thickness to seal the nest; termites build underground
ods of inbreeding and outbreeding, as well as several arches that meet in the middle; weaver ants coopera-
ecological factors. Even in Hymenoptera, it is plausible tively pullleaves together so that others, holding silk-
that all three proposed mechanisms have been in- producing larvae, can bind them (discussed later). You
volved. One researcher has proposed that these three might infer that all these activities require either un-
mechanisms be integrated into an appealing scenario derstanding of the larger goals or an exceedingly com-
consisting of five stages: colony mutualism, gambling plex system of communication, entailing explicit in-
for reproduction, parental manipulation, kin recogni- structions from a leader or detailed information passed
tion and selection, and the superorganism. In the final back and forth among individual members. Yet a close
superorganism stage, the colony may be thought of as examination of individuals engaged in cooperative ac-
the unit of selection, a single organism whose sterile tivities often reveals afine level of disorder and appar-
castes are merely cellular extensions of the queen's ent chaos. Effectiveness emerges at a higher, statistical
body and whose new reproductives are her gametes. A level, through mass action. Studies indica te that even
major difference, however, is that the individuals the most clever and efficient activities are performed by
within an insect colony are not genetically identical, individual workers following very simple behavioral
which reveals itself occasionally in conflicts of interest. rules, through a process of self-organization. Commu-
For example, a honey bee queen mates multiple times, nication simply meshes the behaviors in particular
so her worker offspring come from different fathers ways to make this possible.
and are not all related by 0.75. Workers with the same
father (full sisters) cooperate with each other better
than with half-sisters. Comparison between Termites
and Eusocial Hymenoptera

Social Communication The eusocial societies of termites and hymenopterans

are fundamentally different because only the latter
Members of insect societies communicate by volatile have complete metamorphosis and are female domi-
pheromones, contact stimulation (both pheromonal nated. The developmental instars of termites are func-
and mechanical), and sound. The messages that act as tionally similar to the adults. After one or two molts,
releasers of behavior communicate alarm, attraction termites can fend for themselves, and in one form or
and assembly, recruitment, and recognition. Those that another both sexes of immatures do most of the work
~~.~" '4' "'\ ~."" i '.Í"
('..'''''\q/~~ =)n~3i:f. !
~.¡ ',:itf] ..
Behaviorallnteractions between Conspecifics 77

ofthe colony. Typically, termites have a single repro-

ductivepair, a king and a queen. Hymenopterans, in
contrast,have helpless larvae that usually contribute
liuleor nothing to the colony, so various forms of fe-
maleadults do all the work. Adult males also con-
tributenothing and are present only during mating pe-
riods.Asa general rule, there is a single reproductive,
Mosttermites are pale or white, whereas ants tend
tobeyellow,red, brown, or black, and have a narrow
constrictionat the base of the abdomen. Yet termites
and many ant species share superficial similarities,
suchas having colonies that last for years, royalty that
shedtheir wings, workers that never have wings, sol-
dierforms,and a readily visible reproductive exodus
swarm.Their nests often are constructed of "carton"
(madeof dead plant materials, feces, or soil) or are 10-
catedin protected cavities in tree trunks, logs, or un-
derground.These similarities give rise to the termite
misnomer"white ants."

Termite Societies

Termitecolonies may be completely underground, in

earthenmounds that rise above ground, on or inside
deadwood (human habitations included), or high in
livingtrees, depending on the species (Figure 4-7).
They function for years, sometimes decades. All
speciesneed dead plant material as food. Their ability
tousethis food effectively depends on a mutualistic re-
lationshipwith internal bacteria or protists, or with
culturedfungí. The nest consists of chambers and tun- Figure4-7 Visible portion of subterranean-based Cu-
nelsformedfrom mud, plant material, and fecal pellets. bitermes termite colony in equatorial Africa. The archi-
Thespecific chambers are dedicated to housing the tecture apparently is designed to shed water during the
kingandqueen (royal chamber), the brood, plant ma- heavy rains, yet allow exhaust of stale air and intake of
terial,or a fungus garden. The tunnels serve as con- fresh air, by convection.
duitsbetween chambers or as heat chimneys and air
circulators.Even on open surfaces, such as tree
branchesor the side of a house, termites typically
travelin covered passageways. The various body forms (mandibulate soldiers) or to squirt them with noxious
in a typical colony fall into two categories: nonfixed or sticky substances from a gland on the front of the
(astesandfixed castes. Nonfixed castes are develop- head (nasute soldiers). In some species, the soldiers
mentalstages,that is, nymphs (but called larvae, pseud- can do both. Lower (primitive) termites lack a fixed
crgatcs,presoldiers, and nymphs, depending on devel- worker caste, and all stages and castes are divided
opmentalstate), that serve as unfixed workers and equally between females and males. Some advanced
eventuallycan molt into one of the fixed castes (work- termites may have two or more kinds of fixed worker
ers,soldiers,primary reproductives, or supplementary and soldier castes, each consisting of males or females
reproductives).Members of fixed castes molt no fur- only. Colonies of some termite species often contain
ther.Workers and soldiers may be viewed either as de- more than one reproductive female, especially when
velopmentallyarrested immatures or as sterile adults. the colony is large and diffuse.
Theworkersare devoted to constructing and repairing New termite colonies are founded when bra-
thenest,nursing larvae, feeding the royal pair, and for- chypterous nymphs (that is, nymphs with externally vis-
agingforplant matter. Soldiers protect the colony by ible wing buds) in mature colonies molt by the thou-
attackinginvaders. Their modified heads are used ei- sands into alates (winged adults that are future primary
ther to grasp enemies in their large mandibles reproductives) and leave the nest at the same time as

78 Chapter4 BehaviorandEcology

those in other colonies in the area. These exodus

swarms occur typically during the onset of a warm or
wet season, triggered by a heavy rain. After a period of
weak flight, the alates land on the ground, males locate
females by a female pheromone, both shed their wings,
and the pair crawling in tandem quickly loca te a site
for a new nest. The newly established primary repro-
ductives (king and queen) begin to rear offspring, but
it may take several years before their own colony can
produce alates. In the meantime, larvae are developing
into pseudergates (unfixed workers that can remain
uncommitted to a fixed caste), presoldiers and soldiers,
brachypterous nymphs, or supplementary reproduc-
tives, depending on the needs of the colony. In well-
defined colonies of primitive termites and many
advanced ones, a supplementary reproductive with func-
tional gonads appears only if the queen or king dies,
the replacement having molted from an older larva,
pseudergate, or nymph. Otherwise, separa te primer
(development-controlling) pheromones from the king
and queen are passed via their feces to all members of
the colony by their anal-oral communication system,
preventing unfixed termites of each sex from taking
over reproductive duties. If substitution accidentally
occurs, or if several supplementary reproductives ap-
pear at once, a contact pheromone from the oldest re-
productive causes workers to kill the others. An in-
Figure4-8 Nest of the paper wasp Polistesexclamans
(Viereck), with nearly identical foundress (queen), sub-
hibilOry primer-pheromone system apparently also
ordinate foundresses, and workers. Centrally located nest
keeps soldiers at a small but constant proportion of the
cells with white caps contain pupae; open cells closer to
total number of colony members.
the periphery are uncapped and contain growing larvae.
In one cell, an unhatched egg is visible.
Paper Wasp Societies
Paper wasps build their nests from a mixture of wood
fibers and saliva, pressed by their mandibles into thin lack functional ovaries. They collect wood fibers, ex-
sheets of paper that make up both the general nest pand the nest, forage for nectar, meat (prey or carrion),
structure and the comb of hexagonal cells in which and water, and defend the nest against intruders, using
they rear their brood (Figure 4-8). These cells typically a venomous sting. There is some tendency among
are inverted, so the larva growing in each is oriented workers to specialize in certain tasks, but they are ca-
head-downward. In common Polistes wasps of North pable of doing any of them. The helpless larvae con-
America, a single comb is attached to the underside of tribute lO the colony only by supplying nutrients lO the
an overhanging substrate (commonly the eaves of adults by traphallaxis.
houses in residential areas) by one or more pedicels. In New colonies are founded by inseminated females
yellow jackets and hornets (for example, genera Vespa, destined to be queens. In temperate climates, these
Vespula, Dolichovespula) and also many tropical paper foundress wasps overwinter in sheltered sites, either in
wasps, many combs are attached one below the other, a cluster (Polistes) or in isolation (for example,
and all combs together are enclosed within severallay- Vespula). In the spring they start new nests (old nests
ers of paper, perforated by an opening for adult wasps are rarely used) by themselves. Vespula foundresses al-
to pass in and out. These enveloped, multitiered nests ways perform all foraging, construction, egg laying,
may be attached to tree limbs or overhangs or be situ- feeding, and defending duties until the first workers
ated in rock piles or underground with a tunnellead- emerge as adults. Polistes foundresses sometimes are
ing to the surface. During most of the season of nest joined by one or two females fram the same generation,
growth, the colony consists of a queen and numerous which assume the rank of subordinate foundress with
slightly smaller female workers. The queen spends nonfunctioning ovaries and perform the tasks of work-
most of her time at the nest, laying eggs. The workers ers. If at any time the foundress dies, one of the subor-
Behaviorallnteractions between Conspecifics 79

dinates will rejuvenate her own reproductive system predators, capturing mainly arthropods. Fungus gar-
andbecome the new queen. During colony growth, fer- deners (¡he leafcutter ants, tribe Attini) cut fresh leaves
tilized eggs are laid in worker cells and receive only and petals from shrubs and trees and place them in
moderate amounts of food. Near the end of the season, chambers where fungus is cultured (see later). Slave
the queen begins laying unfertilized eggs, which de- makers specialize in raiding or taking over the nests of
velop into males, and fertilized eggs in cells slightly other ant species and using the labor of the conquered
larger than those of workers (in the case of Vespula), workers. Nest parasites maintain their entire colonies
which develop into foundress daughters. The latter re- within the nests of other ant species, often lacking a
ceive more food than workers and become larger worker caste altogether and depending on the host for
adults. Males await foundress daughters near nests or defense and food. Most kinds of foraging ants use trail
potential overwintering sites, but all males are dead by pheromones to lead other foragers to a source of food.
the time foundress daughters are safely sequestered. In A forager that has found food leaves a trail on the sub-
subtropical and tropical regions, paper wasps may have strate as she returns directly to the nest (having used
many foundresses and maintain nests for much longer the sun compass to keep track of her position relative
periods of time, with periodic queen replacement and to the nest), then recruits other ants to follow the trail.
seasonalproduction of reproductives for dispersal. Each ant that finds the food adds to the trail on her re-
turn, but when all food has been removed the trail no
Ant Societies longer is reinforced and soon disappears.
Colonies of ants typically start like those of paper
Ant species form nests in a wide variety of sites: deep wasps: periodic production and dispersal of a new gen-
under ground, beneath rocks, inside old logs, inside eration of reproductives. At a particular season winged
hollowbranches, inside chambers provided by mutual- adults of both sexes leave the colony, the males form
istic plants (se e discussion of myrmecophytes later in aerial mating swarms that females enter to find a mate.
this chapter), and high in trees. Some construct their Inseminated females, destined to be come queens, shed
nests of carton, others from living materials, such as their wings and find a site to establish a colony on their
the leaf nests of weaver ants (Oecophylla), which are own. A queen begins by laying eggs and feeding the lar-
held together by silk, from larvae, that workers manip- vae from her metabolic stores to develop a generation
ulate in a stitching action, just prior to forming co- of workers. As the worker base expands, the colony be-
COOllS. The nomadic army ants (such as Eciton) and sa- comes self-maintaining by foraging for food, feeding
fari or driver ants (such as Dorylus) form a temporary the larvae, and leaving the queen to egg production.
nest from the interlocking bodies of living workers. After several years, the colony is sufficiently large that
Nests of solid construction may have tunnels and winged males and reproductive females are produced
chambers devoted to the queen, brood, food, or fungus and exodus flights occur. Another type of colony for-
gardens. A typical growing colony has one wingless mation, seen in army ants, is division of the parent
queen and numerous wingless female workers. Some colony into two groups, one with the old queen and
ant species, especially those with large and dispersed one with a few new, uninseminated female reproduc-
colonies, may have secondary reproductives (worker- tives. All but one are removed from the new colony and
queen intermediates that lay eggs) or multiple queens. allowed to die. The remaining new queen is insemi-
Workersmay be all of one type or may be divided mor- nated by males from her own or other colonies. Con-
phologically into several subcastes with different func- trol of female caste generally is pheromonal-nutritional.
tions: Minors are small and often remain within the The queen produces a pheromone that prevents work-
nest, cleaning and tending to the brood; medias are of ers from feeding female larva e a royal diet until the
intermediate size and do most of the foraging; majors time of winged exodus approaches, when she also lays
Coftenreferred to as soldiers) are large-bodied, with ex- unfertilized eggs that will be fed a male diet. Larval di-
ceptionally large heads and mandibles, and defend the ets that allow development of various female subcastes
nestor columns of foraging workers. In the case of no- are determined by the needs of the colony, also pre-
madic ants, majors also guard the procession of all sumed to be mediated by pheromones.
colonymembers (many carrying a larva) as they move
to a new bivouac. The foods of ants are extremely di- Honey Bee Society
verse.Harvester species collect seeds and store them in
chambers. Honeydew collectors are closely associated Honey bee (Apis mellifera L.) colonies have been as-
with honeydew-producing Hemiptera (discussed sisted or maintained by humans for thousands of years,
later), in some cases storing the fluid underground to obtain their stored honey or pro mote their pollinat-
within the bodies of designated workers (honeypot ing abilities, which are extremely important to agricul-
ants). Carnivores may be either generalist or specialist ture. Consequently they are the most intensively stud-
80 Chapter4 BehaviorandEcology

ied and best understood of the social insects. In nature, which remains in the intruder and attracts other work-
nests are established in rock and tree cavities that have ers to the site of the sting and induces further stinging.
a narrow opening. In cultivation, colonies are main- An assembly pheromone, released by a forager after re-
tained in baskets or wooden boxes (referred to as hives) turning from a successful expedition, causes other for-
with similar features. The nest consists of a series of agers to gather around to obtain information about the
parallel combs, differing from those of paper wasps in food source. Much of this information is transmitted
that the combs are made of wax and are vertical, so that during a dance, which has components suggesting
the cells are horizontal and occur on both sides of the symbolic language. The dance language differs slightly
combo Furthermore, the cells are used not only for among different races of honey bee. The following di-
rearing brood but also for storing honey (concentrated alect applies to the Carniolan race of northern Europe:
flower nectar) and pollen. In addition, the combs serve When food is less than 20 meters from nest, the forager
as platforms where workers exchange food and infor- performs a round dance, in which it walks in an almost
mation. The colony consists of one winged queen, a complete circle, alternating between clockwise and
few hundred winged males (drones), and many thou- counterclockwise directions. This pattern indicates
sands of smaller winged female workers. The queen is that the source is nearby, its richness is indicated by the
entirely dedicated to producing eggs and laying them duration and vigor (vibrations) of the dance, and its
in cells. Drones are present only during the reproduc- odor is imparted by the dancer. Between 20 and 80 me-
tive season, contribute nothing to the colony, and serve ters, a transitional sickle dance is performed, which
to mate with new queens from other colonies. Workers contains elements of the waggle dance. Foragers do a
generally follow a sequence of tasks and associated perfect waggle dance when food is more than 80 m
gland activity: (l) cleaning cells; (2) feeding larvae away. It consists of alternating left and right semicir-
salivary secretions (known as bee milk or royal jelly) cles, with a straight waggle run in between, in which
and pollen mixed with honey (bee bread); (3) building the bee vigorously wags its abdomen from side to side
cells, using wax from their abdominal wax glands; while walking forward. Richness and odor of the food
(4) storing pollen and honey, which has been taken source are transmitted as in the round dance. The ap-
from incoming foragers; (5) guarding the colony, by re- proximate distance to the food is indicated by the du-
maining near the entrance and attacking intruders with ration of the waggle run, and consequently the number
a suicidal sting; and finally (6) foraging, which in- of waggles in it and number of waggle runs per unit of
ocludes collecting nectar, pollen, resins, and water, up to time. The direction to the food is indicated by the an-
8 km from the nest (Figure 4-9). gle between the direction of the waggle run and the
Communication among workers includes the re- vertically upward direction. This angle represents the
lease of an alarm pheromone from the sting apparatus, angle between the food source and the sun's azimuth,
which is the sun's coordinate on the horizon. The
dance occurs within an almost completely dark nest on
a vertical surface, so recruited workers detect its geo-
metric features entirely by mechanical stimuli coming
from the dancing bee, which they follow during the
course of her dance, and from their sense of gravity.
The most abstract feature of this communication is the
representation of the real sun's azimuth on a horizontal
landscape by the direction "up" on a vertical dancing
New honey bee colonies are founded when a large
one divides into two parts. The process starts when
workers start building queen cells, large cells that hang
vertically off the face of ordinary honeycomb, and rear-
ing future queens in them. Typically, the old queen
leaves with about half the workers in a prime swarm, a
mass of bees that comes to rest on a tree branch or
other aerial site. By using the surface of this cluster of
bees, permanent nest-site foragers employ the waggle
dance to recruit each other to cavities that might be
suitable, and those dances reporting the best site grad-
Figure4-9 Foraging worker honey bee, Apis me/lifera, ually recruit more workers until a critical mass agrees.
gathering nectar and pollen froro a flower. Then they allleave together with the queen to relocate
CommunityAssociations 81

in the chosen cavity. Meanwhile, at the old nest, the intertidal zone. Green plants are the universal produc-
firstnew female reproductive to emerge destroys the ers of organic materials, using energy derived from the
otherqueen ce11s.If two emerge at about the same sun. Phytophagous (herbivorous) arthropods feed on
time,they fight to the death, using their venomous plants to obtain these materials and are thus primary
stings,so that only one remains as queen. This repro- consumers. Zoophagous anhropods are secondary, ter-
ductivefemale then goes on a mating flight (nuptial tiary, or quaternary consumers, feeding at so me more
flight)to a location high in the air where drones are ac- distant link in the food chain, either on animals that
tive.She attracts them with a pheromone and mates feed on plants or on animals that feed on other ani-
with10-15 in succession, a process that tears the gen- mals. Detritivores feed on dead matter at any of several
italiaoff the abdomen of each male and causes his trophic links, further using the organic materials there.
death.When fi11edwith a lifetime's supply of sperm, Some insect detritivores are omnivorous, feeding op-
sheretums to the nest to begin laying eggs lo replenish portunistica11y on living plant or animal tissue as we11.
thestockof workers. In temperate climates, swarming At each step in the transfer of food from consumed to
occursin late spring and may involve two or three af- consumer, roughly 90% of the energy contained in the
terswanns,fo11owingthe prime swarm. An afterswarm food is lost. Thus the total amount of biomass among
consistsof another large proportion of workers and the primary consumers is large, but at the top of this
newqueen, who wil\ be replaced by a subsequent trophic pyramid the biomass is sma11. This means the
queenstill developing. environment sustains a greater mass of herbivores
Female caste control is pheromonal-nutritional. (grasshoppers and buffalo) than carnivores (lice and
Asin other hymenopterans, drones are derived from eagles). Actual relations among members in a commu-
unfertilizedeggs, which the queen lays in moderately nity do not form a simple pyramid based on a few food
largecellsnear the corners of the combs. Workers de- chains. Rather, they usua11y form a food web, with each
velopfram fenilized eggs in sma11ce11s,identical lo species either eating or being eaten by several other
thoseused to store food, typica11yin the central pan of species (Figure 4-10). This is beca use in every feeding
thecomboThey are fed royal je11y (nurse bee saliva) category, species vary along a continuum of food pref-
duringthe first three days of development, then a erences, from extreme generalists lo extreme special-
steadydiet of bee bread. Future queens, in contrast, are ists. Furthermore, not a11 relationships involve just
fedlargeamounts of royal je11y throughout larvallife. eating and being eaten. The exact nature of the rela-
Iftheresident queen dies, workers hastily build queen tionship between one insect species and other organ-
cellsand transfer diploid eggs from worker brood ce11s isms, whether based on food and or another vital
tothequeen ce11s,so that new queens can be reared. resource, defines its ecological role¡ or niche. By con-
(Inthe capensis race, workers, lacking sperm, never- vention, in mutualism both organisms bepefit from the
thelesscan produce diploid eggs on their own.) The relationship. In commensalism, one benefits and the
nursebees decide which diet to give a larva by assess- other is unaffected. In parasitism, the sma11 organism
ingits development and the kind of ce11it is develop- (the parasite) benefits and the large one (the host) is
ingin.Ultimately, it is the queen who controls caste, by weakened. In predation, the large organism (the preda-
releasingan inhibitory mandibular-gland pheromone tor) benefits and the sma11one (the prey) is killed. Be-
("queensubstance") that is passed through the colony tween parasitism and predation, there are pathogens
andpreventsworkers from building queen ce11s.Ento- (bacteria, protozoa, nematodes, fungí) and parasitoids
rnologists think that during swarming season either the (insects), which are sma11parasites that seriously harm
colonyhas grown so large that the pheromone becomes or gradua11y kill the larger host. The term symbiosis is
toadilute to be effective or that the queen herself re- reserved for the physica11y close and long-lived rela-
duces itsproduction, so new female reproductives can tionships observed in mutualism, commensalism, and
beproduced. parasitismo

Microbial Relationships
Community Mutualists: A variety of microbes assist insects nu-
tritiona11y, living in close association with them or in-
Arthropodsin general are major players in a11biotic terna11y. In return, the insect provides a home, a
communities.In the marine environment, crustaceans source of food, and a mechanism for transmission to
havelargeroles, whereas on land and in freshwater, in- other insects. Many wood-eating termites, and also
sectsare dominant but are virtua11y absent in marine wood-eating cockroaches, harbor colonies of fiagel-
cornmunities.Exceptions are a few insect species on late protists and bacteria in a special chamber in the
theocean'ssurfaces and along marine beaches and the anterior hindgut. These microbes break down ce11u-
82 Chapter4 Behaviorand Ecology

TERTlARY Parasites




Figure 4-10 Major feeding links within a community in one English woodland, illus-
trating the general nature of food webs and the integral roles of insects in them. Tortrix is
a moth, Cyzenis is a tachinid fly,Philonthus is a staphylinid beetle, and Abax and Feronia
are carabid beetles.

lose, which is otherwise indigestible. They are passed mutualistie fungus from being destroyed by a parasitie
to newly hatched termites by the fecal-oral transmis- fungus. In both kinds of fungus-growing insects, new
sion route used also in communieation. Similarly, reproductives that leave the parent colony carry the
wax moths, whieh live in honey bee colonies and eat fungus with them, to seed a new garden. Some bark
the wax of their honey combs, contain bacteria that beetles (Scolytinae) and ambrosia beetles (Platypodi-
digest the wax. Other insects maintain colonies of nae) also cultiva te fungi (Ascomycetes and Fungi Im-
bacteria that provide vital nutrients, such as vita- perfecti) in their tunnels beneath tree bark and eat spe-
mins. These bacteria may be maintained in gut crypts cial growth forms.
or intracellularly in mycetocytes, which form a spe-
Pathogens and Parasites: Insects are attacked by a
cial structure, the mycetome. For example, insects
wide variety of mieroorganisms, induding viruses,
that feed on vertebrate blood during their entire life
rickettsiae, spirochetes, eubacteria, protists, and fungí.
cyde (kissing bugs, bed bugs, lice, tsetse flies) obtain
insufficient B-vitamins from the blood, but the bacte- Insects also are infected by flukes, tapeworms, round-
worms, hairworms, thorny-headed worms, and para-
ria in their guts or mycetomes provide these vita- sitie insects (se e discussion later). Most of these infec-
mins. The bacteria are passed to new generations via tions result in death of the host insect and form the
eggs or feces.
basis of several kinds of successful biologieal control of
Some termites and ants grow mutualistie fungi in
insect pests.
gardens as a source of food, and in return the insects
provide the fungi with a substrate for growth and Arthropod Vectorsof Pathogensand Parasites: Arthro-
with transmission mechanisms. Advanced termites pods often serve as vectors, transmitting pathogens and
(Macrotermitinae) grow Tennitomyces fungi in special internal parasites to vertebrates or to plants. The most
chambers on combs of fecal pellets containing plant important vectors of human and livestock diseases are
fibers. The termites eat both combs and fungus, whieh mosquitoes and ticks. Some pathogens are transmitted
contain cellulases allowing digestion. Leafcutter ants accidentally or incidentally, having no biologieal rela-
(Attini) grow various basidiomycete fungi on a mulch tionship with the arthropod (mechanical transmission),
of freshly cut leaves and flowers and eat the swollen for example, a variety of viruses and bacteria carried by
tips (gongylidia) of the hyphae. The ants also keep the house flies and cockroaches. In doser vector-pathogen
CommunityAssociations 83

relationships(biological transmission), which are obli- mor (leafhoppers), sugar beet curly top (leafhoppers),
galeandfairlyspecies specific, the parasite completes its aster yellows (leafhoppers), corn stunt (leafhoppers),
tXtrinsic cycle in the vector, by multiplying (propaga- rice stunt (planthoppers), tomato spotted wilt (lhrips),
tive),multiplyingand transforming (cyclopropagative), cotton leaf curl (whitefIies), cucurbit wilt (cucumber
or only Iransforming and growing (cyclodevelopmen- beetles), and Dutch elm disease (bark beetles).
tal).Propagativepathogens include viruses, rickettsiae,
spirocheles,and eubacteria. Cyclopropagative patho-
Plant Relationships
gensinelude sporozoan and fIagellate protists. Cyclo-
developmentalparasites include fIukes, tapeworms, Pollination: The best known of the mutualistic
roundworms,and thorny-headed worms. Many of plant-insect relationships is cross-pollination of an-
¡hesecause serious diseases in humans and livestock, giosperms by insects that visit fIowers. This service is
affectinghundreds of millions of people. On a global much more efficient than windborne pollination and is
basis,among the most noteworthy livestock and pet thought to have brought the angiosperms to plant
diseases(and their vectors) are babesiosis (ticks), thei- dominance, as well as to diversify the insects associated
leriasis(ticks), nagana (tsetse flies), equine encephali- with them. The most speciose orders of insects-
lis (mosquitoes), and dog heartworm (mosquitoes). Coleoptera, Hymenoptera, Diptera, and Lepidoptera-
Amongthe most infamous human diseases are malaria also are those that do most of the pollinating. Basically,
(mosquitoes),river blindness (black fIies), elephantia- the insects transfer pollen between plants by picking
sis(mosquitoes), encephalitis (mosquitoes), Lyme dis- up pollen from anthers of one fIower and depositing it
ease(ticks), spotted fever (ticks), dengue (mosqui- on stigmas of another. In most cases, this activity is
toes),epidemic typhus (lice), relapsing fever (lice and purely incidental on the part of the insect. The primary
ticks),Chagas disease (kissing bugs), yellow fever incentive for insects is food provided in the fIower:
(mosquitoes),sleeping sickness (tsetse fIies), leishma- nectar, a secreted solution of sugars with a variable
niasis(sand flies), and plague (fIeas). Most arthropod amino acid component, and excess pollen, a valuable
veclorsare blood feeders (discussed later), offering the source of protein and lipid. This reciprocity has led to
palhogenor parasite an easy way to pass between ver- a variety of special features of both pollinators and
¡ebraleand invertebrate hosts without exposure to the plants. Insects have body modifications for handling
outsideenvironment, but there are many variations in the pollen and storing the nectar, and behavioral re-
¡heexact manner of host inoculation. These include finements that help them quickly loca te and exploit
venebrateautoinoculation with the crushed vector or sources of nectar and pollen: ability to learn and dis-
itsfecesor inadvertent ingestion of the infected arthro- criminate among colors and scents, an accurate time
podo Within the vertebrate, the pathogen completes its sense, fIower constancy on each foraging expedition,
intrinsiccycleand then is infectious to the arthropod and a tendency to specialize on just one kind of plant
again.Ihis alternation between hosts is called horizon- (monolexy). These insect behavioral characteristics are
talIransmission.By contrast, some propagative patho- promoted by, and benefit, the plants. Plants, in con-
gensalsoare transmitted from generation to generation trast, have evolved features that assist the insect in 10-
ofarthropodby transovarian transmission, that is, ver- cating the nectar and that at the same time facilitate
ticaltransmission. This helps maintain the chain of in- pollen transfer: fragrance, colorful petals, nectar guides
fectionin the face of vertebrate immunity and condi- (petal arrangements and markings), landing platforrns,
tionswhen horizontal transmission is impossible, as reliable and synchronous fIowering times and nectar
during winter. fIows, mechanical pollen-transfer devices, temporary
Whenreferring to arthropods that act as vectors of insect traps, and defenses against nectar robbers. Blue
plan!pathogens, plant pathologists speak of nonpersis- and yellow fIowers tend to be visited by bees, orange
tentandpersistent pathogens, instead of mechanical and red fIowers by butterfIies, and white fIowers by
andbiologicaltransmission, but the meanings are sim- moths or generalist short-tongued pollinators, such as
ilar.Viruses,phytoplasmas, spiroplasmas, and bacteria flies and beetles. Flowers often promote monolexy by
arecommonand economically important pathogens of secreting nectar into a corolla that can be reached only
plantsthat multiply in their insect vectors. Fungi are by an insect with an exceptionally long proboscis.
typicallycarried mechanically.Most vectors are thrips, Some plant-pollinator relationships are particulairIy
aphids,leafhoppers, planthoppers, and similar insects close, involving insect development within the fIower.
thalpierce plant tissue with haustellate mouthparts, In the case of the yucca moth (Prodoxidae: Tegitecula),
bUIseveral kinds ofbeetles also transmit important dis- the female gathers a pollen ball in her tentacles (pre-
easeorganisms. Among the well-known vector-borne hensile palps) on one fIower, then fIies to another, lays
diseasesof crops (and their vectors) are sugarcane mo- her eggs in its ovary, then places the pollen ball on the
saic(aphids), cucumber mosaic (aphids), wound tu- fIower's stigma so that seeds will develop, some to be
84 Chapter4 Behaviorand Ecology

eaten by the moth's larvae. Fig wasps (Agaonidae) are

even more complex. Both sexes of wasp develop in the
ovaries of modified female (neuter) flowers of fig trees
(Ficus) within the fig's "fmit" (the syconium, a large re-
ceptacle lined with flowers). The wingless males come
out first and mate with females while they are inside
their cocoons. A female leaves the fmit while it is in the
male flowering phase, so that she picks up pollen. She
then finds a fmit that is in the female phase and polli-
nates normal female flowers while ovipositing on the
neuter ones. The normal, fertilized flowers develop
seeds. In some species of figs, predominantly male, fe-
male, or neuter types of fmit occur at different times of
year. Commercial Smyma figs lack wasps and must be
pollinated by wasps from "goat figs" that growers place
in the orchards, in order for the fruit to develop its
thick, edible flesh.
Not all flower visiting and pollination is mutualis-
tic. Nectar-robbing bumble bees that cannot reach the
nectar of specialized flowers with their mouthparts cut Figure 4-11 Two ants drinking nectar from an ex-
into the side of the corolla to take the nectar without trafloral nectary on an Inga tree. These ants attack in-
contacting anthers or stigmas. Nectar thieves merely sects that land on nearby leaves.
take accessible nectar but lack the appropriate equip-
ment or behavior to transfer pollen. In the reverse sit-
uation, plants employ deception to achieve pollination mosquito and fly larvae fall into this category. Still oth-
without providing the insect with anything in retum. ers create tiny shelters (domatia) for predaceous mites,
Flowers that release odors similar to rotting flesh or ex- or have glands (extrafloral nectaries) that provide
crement attract saprophagous flies and beetles, which sugar for parasitoid wasps and ants that patrol the
pick up or deposit polI en while wandering around on plant and disturb or attack plant-feeding insects (Fig-
the flower. Blow flies that are attracted to the star ure 4-11). Myrmecophytes, or ant plants, provide shel-
flower (Stapelia) even lay their eggs at the center of the ter, food, or both, to the ants, in retum for protection,
flower, where the hatched larvae die. Many of these and have a fairly species-specific and interdependent
flowers are brown or purple, and the dead-animal effect relationship with them. The best known of these is the
may be enhanced by simulated hairs. Ground orchids bull's hom acacia, a small tree in the Neotropics that
in Europe (Ophrys) and Australia (Drakonorchis) provides both food and shelter for one Pseudomyrmex
loosely resemble female Hymenoptera visually and also ant colony. The ants, including queen and larvae, are
produce pheromone-mimicking volatile chemicals and housed in large, hollow, twin thoms that are scattered
tactile stimulation that target particular bee and wasp over the branches and twigs. Sugar is provided from
species. This mimicry is sufficient to cause males to extrafloral nectaries at the base of leaf petioles, and
make repeated attempts to copulate with a succession protein and fat are derived from Beltian bodies, pellet-
of flowers, in the process transferring sticky pollen like extensions of the leaflets that the ants pick and
packets. carry to the thoms for colony distribution. For their
part, the ants act as ferocious body guards, biting and
Myrmecophytes: Arthropods have a variety of mutual- stinging both insects and vertebrates that come into
istic relationships with plants that do not involve pol- contact with the tree. They also prevent competing
lination. Some are relatively simple, such as the plant plants from growing around the base of the tree. An en-
providing a fleshy attachment (elaiosome) to its seeds, tirely different kind of relationship occurs between
to make them more attractive to ants, which will dis- Philidris ants and bulbous epiphytes (Rubiaceae:
perse and bury them in the process of harvesting. This Myrmecodia and Hydnophytum) that grow on small
is called myrmecochory. Many plants collect rainwater trees in nutrient-poor sandy soils of Southeast Asia.
in their leafaxils or other stmctures, providing a spe- The pseudobulb has rootlets clinging to the host tree
cialized habitat for aquatic insects. In retum, the in- and has a single twig of leaves. It is riddled with cav-
sects process the organic matter that falls into these ems that house the ants and provide them with cavities
phytotelmata, making nutrients more quickly available for depositing refuse, which consists mainly of remains
to the host planto Insectivorous pitcher plants and their of the ant's insect prey and of dead ants. The linings of
CommunityAssociations 85

these cavities appear able to absorb the nitrogenous beetles, corn-ear worrns, gypsy moths), and the para-
nutrients from this waste. The ants forage widely and sites, especially the endoparasites, are the most host
are ineffective in defending either the host tree or the specific (such as leafminers, gall-makers). Yet, many
epiphyte against herbivores. species with leaf-chewing larvae have be come dedi-
cated to particular kinds of plants containing toxic sub-
Herbivores: Very few terrestrial or freshwater plants stances or other defenses that only a specialized insect
are not fed on by insects. Together, phytophagous in- can overcome (such as monarch butterflies, pipevine
sectsmake up almost half of all insect species. Both tis- swallowtails, zebra butterflies). Herbivores make host
sue feeders and sap suckers may be considered para- choices during the appetitive sequence that leads from
sites, grazers, or predators, depending on the duration a general search to successive and overlapping re-
and outcome of the relationship between plant and in- sponses to volatiles, visual appearance, taste, and ini-
sect. A single insect that attacks seeds (such as seed tial ingestion or oviposition. The behaviors often are
bugs, seed weevils) always kills the embryo, and some elicited by chemical "sign stimuli," kairomones and
kinds of stem borers do sufficient damage to the main phagostimulants that are associated only with the fam-
stalk that an entire plant dies. However, typical grazers ily, genus, or species of plant that the insect is adapted
(such as leaf, stem, and root eaters with chewing to feed on. Specialists have host-detecting equipment
mouthparts) and parasites (such as leaf raspers, and digestive and metabolic systerns dedicated to the
leafminers, leaf skeletonizers, and phloem or xylem particular host, giving them a competitive advantage
drinkers with piercing-sucking mouthparts) only over generalists or allowing them to eat plants from
weaken the plant or even stimulate new growth, unless which generalists are excluded altogether. A further ad-
theyoccur in tremendous numbers, as locusts do. Even vantage is that the host-plant poisons can be incorpo-
when gypsy moth caterpillars completely defoliate a rated into the insects' bodies, giving them protection
tree,new leaves are produced later the same year or fol- from their own predators. Generalists, however, have
lowing year. Tiny phloem-feeding aphids, ectoparasites more food options, making it relatively easy to locate a
equivalent to the blood-feeding lice of mammals, often host plant and providing a large resource base for their
havean imperceptible impact on the plant's health. Yet populations. Also, their evolutionary fates are not tied
heavyinfestations of aphids, scale insects, froghoppers, to the success of one or a few species of plant.
leafhoppers, and various piercing-sucking bugs can Plant defenses against phytophagous insects fall
cause significant wilting, spotting, browning, fruit into anatomical, chemical, developmental, behavioral,
drop, and leaf curling, and can precipita te death of the and mutualistic defense categories. Anatomical quali-
plant. Similarly, endoparasites such as leafminers, ties include (l) visual devices such as unattractive col-
stem-borers, and gall-makers can have either marginal ors and low reflectance, divergent leaf shapes within a
effects or serious ones. Stem-borers usually kill the genus (making it harder to learn and recognize suitable
stemin which they develop, thus retarding growth, and hosts), and visual mimicry of inedible plants; (2) me-
bark-boring and wood-boring beetles can weaken trees chanical and structural devices such as thick and fi-
sufficiently that they die from infection by microbial brous tissues, spines, hooks, and hairs (trichomes),
pathogens. Gall-makers seldom threaten a plant's life, and sticky sap as a wound response; (3) small seeds
but by applying their secretions they create conspicu- that make searching and feeding inefficient. Chemicals
ous and sometimes disfiguring swellings on the plant. include repellents, distasteful substances, natural in-
The secretions subvert the plant's own developmental secticides, digestive enzyme inhibitors, antimetabo-
program to produce a distinctive plant structure in lites, and growth regulators (hormone mimics). These
which one or more insects can feed and grow to matu- properties are produced by various alkaloids, ter-
rity.Most of these are made by gall wasps (most Cynip- penoids, phenolics, proteins, gIycosides, and cyanides.
idae) and gall midges (Diptera: most Cecidomyiidae). Developmental defenses include seed tactics that make
Gallsalso are made by other Hymenoptera (so me chal- seeds and seedlings an unreliable food source: wide
cidoids, braconids, and tenthredinids), other Diptera dispersal, erratic production, and unpredictable germi-
(some tephritids and agromyzids), Hemiptera (some nation or long dormancy. Another developmental de-
aphids, psyllids, and coccids), Coleoptera (some wee- fense connects the rate of growth to chemical and
vils,cerambycids, and buprestids), Lepidoptera (some structural defenses in one of two strategies. According
gelechiids), and Acari (some mites). to the plant apparency hypothesis, plants tend to be ei-
Plant feeders range widely in host specificity but ther (l) inapparent (fast growing, herbaceous, annual,
are conveniently divided into generalists (polypha- with edible parts available for short periods and with
gous) and specialists (oligophagous and monopha- highly toxic substances produced in low concentra-
gous).In general, the grazers that chew leaves, flowers, tions) and are fed on by specialist insects, or (2) ap-
and fruit have the broadest tastes (such as japanese parent (slowly growing, woody, perennial, with tough
86 Chapter4 BehaviorandEcology

but edible tissues available for long periods and with some associations are more involved. Aphids may re-
substances of low toxicity but high concentration, tain their honeydew until an ant solicits it or have spe-
making digestion difficult and slow) and are fed on by cial structures for holding the droplet rather than dis-
generalists. Behavioral defenses consist of diel rhythms charging it. Others have grasping devices to cling to the
of movement or responses to insect attack that make ants when disturbed, or they may reproduce vivipa-
further attack difficult. These include collapsing leaves rously throughout the year, so that clones of them are
and petioles and enhanced production of toxic sub- constantly producing large amounts of honeydew.
stances both at the site of injury and throughout Ants, for their part, solicit honeydew; carry aphid eggs
healthy tissues. Mutualistic defenses are provided by int.J the ant nest for the winter; select appropriate food
insects that are natural enemies of herbivores. Ex- plants and carry the aphids to them; apply substances
trafloral nectaries encourage ants and parasitoid wasps similar to juvenile hormone to the aphids, to prevent
to remain around the plants and attack or infect herbi- them from developing wings; respond to an aphid
vores. In the case of some myrmecophytes, this alarm pheromone by searching for possible enemies;
arrangement is a deeply involved interdependency (as build shelters on the plant to house scale insects; and
discussed earlier). Other plants, when injured by phy- carry scale insects on nuptial flights in order to estab-
tophagous insects, release volatile compounds that at- lish them in new colonies. Lycaenid caterpillars pro-
tract parasitoid wasps. These natural enemies inject vide a special substance from a dorsal gland on the ab-
eggs into the herbivores, and the parasitoids' larvae in- domen. Ants covet the secretion and provide
flict a slow death. protection in return. When disturbed, the caterpillars
emit a sound that attracts the ants, and within the ant
Insectivorous Plants: A few plants have turned the ta-
nest the caterpillars are nurtured and protected. Hu-
bles on insects by eating them. These insectivores typ-
mans and honey bees also have a long history of food-
ically live in nitrogen-poor soils, such as sandy or clay
for-care mutualism, although it is not interdependem
soils and acid bogs, and the prey are biodegraded to re-
and is cultural rather than genetic.
lease elementary nitrogenous compounds. The plants
derive energy from photosynthesis and are not carniv-
Commensals:A large assortment of insects gain by liv-
orous heterotrophs. Active traps, such as the Venus fly-
ing in close association with other animals, doing little
trap (Dionaea), have specialized leaf devices that
or no harm by their presence. Typically they live in the
quickly snap shut to trap small arthropods when trig-
animal's nest and are known collectively as inquilines.
gered by sensitive hairs. In the case of the aquatic blad-
In addition to accessing shelter and protection, they eat
derwort (Vtrícularía), the prey is sucked into a cham-
the host's waste, other inquilines, or the host's food
ber. Semiactive traps, such as sundews (Drosera),
supplies, the latter causing a very slight drain on re-
flycatchers (Byblis), and butterwort (Pinguicula), have
sources and thus crossing the line into social para-
sticky tentacles or hairs on their leaves to trap the in-
sitism. A few species of domestic cockroaches and their
sect, then gradually enfold it to form a digestive cupo
human hosts fall into this category. Termites and ants
Passive traps, such as pitcher plants (Sarracenia, Ne-
provide an ancient habitat for many specialized in-
penthes, and so forth) and cobra lilies (Darlingtonia),
have leaves modified into vessels of water surrounded quilines, termed termitophiles and myrmecophiles, re-
spectively. These fall into three categories: (1) Synec-
by downward-pointing hairs and spines that make es-
thrans are attacked by the hosts but can escape
cape nearly impossible. Most of these plants have ei-
or protect themselves; they scavenge food or waste.
ther visually or chemically attractive features, and
(2) Synoeketes are ignored by the host beca use they
some bear nectaries that encourage insects to feed near
move quickly or do not appear foreign; they generally
the trap's entrance. Digestion is aided by enzymes, but
scavenge food or waste but also sometimes kill and ea!
at least in pitcher plants bacterial degradation of the
host immatures. (3) Symphiles are accepted by the hos!
drowned insect is equally important.
as members of the colony; they may be fed, carried, and
groomed by the host. Symphiles share various charac-
Animal Relationships
teristics that suit them to this role: They secrete ap-
Mutualists: Ants have two well-known trophobiotic peasement substances that the hosts find attractive and
relationships, one with various stenorrhynchans palatable; they have chemical, tactile, or visual resem-
(Hemiptera) that produce honeydew and one with but- blances to the host (Wasmannian mimicry), or they
terfly caterpillars (Lycaenidae) that produce a glandu- have a sheltering cara pace that prevents hosts from
lar secretion. In the case of aphids and scale insects, the contacting vulnerable parts; they exhibit morphologi-
basic arrangement is that these plant suckers defecate cal regression, including flightlessness and reduction
honeydew (excess phloem sap, rich in sugars and or loss of eyes and appendages; and they use the chem-
amino acids), and the ants protect them. However, ical and behavioral communication systems of the

CommunityAssociations 87

host,includingalarm, attraction, mutual grooming, 50- toes or small domestic flies. When these carriers land
of food, and traíl following. on the skin, the larvae immediately hatch and burrow
in. Endoparasites obtain oxygen by tapping into the
Parasites:Arthropodsparasitize a wide variety of ani- host's tracheal system (in the case of insect hosts) or
mals,including other arthropods. Typical ectoparasites maintaining a hole in the host's skin where the spira-
suckthehemolymph or blood of their hosts, either re- des can be exposed to air. Stomach bot-fly larvae of
mainingon the host continuously (that is, symbioti- horses circumvent this necessity by using hemoglobin
cally)(such as lice, host fleas, and some mites) or vis- cells to store oxygen that comes as occasional air bub-
itingit only periodically (such as kissing bugs, nest bles with the horse's food. As with plant feeders, animal
Oeas,mosquitoes, and horse flies) (Figure 4-12). The parasites find their hosts in a series of oriented maneu-
closelyadapted ones have a number of characteristics vers, starting with a general search and ending in land-
suitedto \ife on another animal, induding the reduc- ing, probing, and feeding. Periodic blood feeders use
tionor elimination of wings, legs, and eyes; special chemical stimuli (kairomones such as fatty acids from
clinging and attachment devices; and body flattening. skin bacteria and carbon dioxide from breath) to locate
Periodic blood feeders share behavioral and physiolog- mammals at a distance, to which are added visual
icalfeatures,induding special receptors for locating stimuli when the host is doser, heat and humidity at
hosts,sneakiness, painless biting, antihemostatic and very dose range, and skin chemicals on contact. Host-
anticoagulant properties of the saliva, and presence of specific sign stimuli, perceived by the parasite during
mutualistic bacteria (discussed earlier). Endoparasites, these steps, are critical in the decision to continue the
suchas strepsipterans parasitic in insects and scabies attack or desisto Parasites tend to be relatively host spe-
mites orbot fliesin vertebrate animals, also have dis- cific, the periodic ones least so, the continuously ec-
tinctive traits that allow them to find, enter, and live toparasitic species much more so, and the endopara-
withinthehost. The so-called human bot fly,or tórsalo, sites most of all.
whose larvadevelops in the skin of mammals and large Important variations on the parasite theme are
birds,is a large, free-living adult. To avoid disturbing parasitoids (often referred to simply as parasites),
itshostduring oviposition, it lays its eggs on mosqui- whose larvae first are parasites on or inside an appar-

-- lB
.j'" a.
:s: Jj ...;

Figure4-12 Periodic blood feeders of vertebrate animals. A, Human bed bug feeding.
This species (Cimex lectularius) lives within human habitations. All members of the fam-
ily Cimicidae are wingless blood feeders throughout life and most live in the roosts and
nestsof bats and birds. B,Asiantiger mosquitofeeding.This species,Aedesalbopictus
(Skuse), often lives near human dwellings, but not in them, and occurs mainly in rural
or wooded areas. All adult members of the family Culicidae have wings, and the females
of most species feed on blood. Their larvae are mostly free-living detritivores.
88 Chapter4 BehaviorandEcology

ently healthy insect host but gradually eat more and tmpoidea, Platygastroidea Chrysidoidea, and Vespoidea.
more of the host's internal organs until it becomes Rhipiphorid beetles, which undergo hypermetamor-
moribund and dies. Essentially the same thing happens phosis, are parasitoids during the second phase of their
when a solitary hunting wasp paralyzes its prey, uses it larval lives. The Strepsiptera also are hypermetamor-
to provision a nest, and lays an egg on it, although in phic but do not kill their hosts. Instead, they often ster-
that case the host immediately ceases to function nor- ilize them or turn them into intersexes, so in one sense
mally. Parasitoid insects typically are small and lay they are parasitoids, beca use they can kill the host's re-
their egg(s) in the host where they find it. They tend to productive potential.
be very host and stage specific, developing, for exam-
pIe, only in the eggs of a particular moth, or only in the Predators: Predatory arthropods inelude most orders
parasitoids that develop in those moth eggs (that is, of Arachnida, all üdonata and Mantodea, and nearly all
hyperparasitoids). The latter are exceedingly small. But Neuroptera. A great many Hemiptera, Coleoptera, and
as a group parasitoids attack a wide variety of insects, Diptera also are predators. Like the parasitoids, they
ineluding many pests, and therefore are use fuI in bio- prey on a variety of small animals, but mostly other
logical control. Females may lay many eggs in a single arthropods, so they are said to be entomophagous. Preda-
host, so that a large family develops in it. Among some tors with chewing mouthparts eat most or all of their
of the parasitic hymenopterans (e.g., Braconidae) are prey; those with piercing-sucking mouthparts pierce the
females that lay only a single egg in the host, but the prey's cutiele, inject a lytic saliva to liquefy the tissues,
egg undergoes polyembryony, resulting in numerous and suck out the resulting soup (Figure 4-14). Like
larvae. Parasitoids may pupate within the host if it plant feeders and animal parasites, there are generalists
dries out by the time they are mature. Typically they and specialists, but in this case the reasons for special-
exit the host and pupate on its cutiele or nearby (Fig- ization are less often physiological and have more to do
ure 4-13). Parasitoids are most common among with habitats, competition, methods of hunting, and
Diptera and Hymenoptera. Tachinids are the most im- special techniques of prey capture. Most species of
portant fly parasitoids; others inelude sarcophagids, dragonflies, assassin bugs, mantids, and many kinds of
pyrgotids, pipunculids, acrocerids, and bombyliids. army ants and driver ants eat diverse types of prey that
Hymenopteran parasitoids inelude many hundreds of occur within the habitats they frequent. By contrast, dif-
species in the Ichneumonoidea, Chalcidoidea, Procto- ferent genera of digger wasps (Sphecidae: Sphecinae)
specialize in spiders, cockroaches, grasshoppers, katy-
dids, tree crickets, tme crickets, plant-dwelling cater-
pillars, or soil-dwelling caterpillars. There are two basic
approaches to prey capture: (1) Hunting is most efficient
if the prey are sedentary, slow moving, or are most
abundant in locations where a predator cannot wait (for
example, in mid-air). The predator moves though like1y
habitats, increasing its chances of encountering the
prey. (2) Awaiting and stalking work well when the prey
are mobile and likely to pass by. The predator remains
in one place until prey is detected, then stalks it ancl/or
grasps it.
Equipment that assists predatory arthropods in
seizing prey inelude raptorial fore legs, often with
spines or sticky setae, and raptorial mouthparts, such
as the extensible labium of odonate naiads and the
Figure4-13 Tobacco homworm, Manduca sexta (L.) long, curved mandibles of diving beetle larvae and
(Sphingidae), in final stage of parasitization by the para- tiger beetles. Insects with a venomous bite or sting
sitoid Cotesia (Braconidae). The larvae have completed (spiders, scorpions, antlions, female hymenopterans)
deve10pment within the caterpillar, then exited to spin can paralyze the prey with a single, well-placed injec-
cocoons in which they pupate. Some have already tion, making them easier to handle. Beaded lacewing
emerged as adults, and some cocoons have dropped off, larvae (Berothidae) subdue their prey with a chemical
leaving black spots at the point where the larva had bur- ejected from the anus. Hunting wasps do their hunting
rowed out. This brood of parasitoids typically results primarily to provision nests for their larvae, and the
from a single egg that has been inserted into the caterpil- paralyzed prey forms a living meal when the larva is
lar and undergone polyembryony, creating a family of ready to eat. Some of the sit-and-wait predators have
identical offspring. devices that improve their catch: traps, lures, or a
CommunityAssociations 89

Figure4-15 White crab spider (Thomisidae) feeding

on a syrphid fly it has captured at the flower. Predators
with this ambush tactic often blend well with the flowers
on which they wait, either to avoid detection by poten-
tial prey or to avoid detection by their own predators.

look like flowers themselves. lt is not clear if this con-

cealment or disguise facilitates prey capture or pre-
vents predation by the predators' own enemies. Insect-
generated lures include bright lights (Waitomo
.:g worms, larvae of mycetophilid larva e that live in caves
~ in New Zealand and attract insects to their sticky-
<!:i stranded webs), flashing lights (female Photuris fire-
--, flies that attract Photinus males to their deaths by mim-
icking the answering signal of a receptive Photinus
Figure4-14 A, Robber fly (Asilidae) feeding on moth. female), and attractive chemicals (bolas spiders that
Robber!lies typically await prey fram perches, then fly mimic female moths by releasing moth sex
out to seize them in mid-air. The proboseis is inserted pheromones and snaring male moths that come within
into the prey, lytic saliva is injected, and then digested range of their sticky bolas). The last two examples ex-
tissuesare withdrawn as liquido The proboseis also can hibit the tactic known as aggressive mimicry.
be used as a defensive weapon. B, Flower fly larva (Syr- Arthropods are prey for many vertebrate animals.
phidae) feeding on an aphid. Because aphids tend to be They form much of the diet of freshwater fish, am-
sedentary,they are easily seized by predators, including phibians, reptiles (notably lizards and small snakes),
those that have smaller bodies. many groups of birds (such as flycatchers, bee eaters,
warblers, creepers, woodpeckers), and some mammals
(notably shrews, small bats, and anteaters). Even ver-
tebrates that do not specialize in arthropods make
combinationof the two. Traps include the sticky aer- them an important part of the diet during the breeding
ialwebsof spiders, the aquatic nets of some caddisfly season (for example, hummingbirds, squirrel mon-
larvae,and the sand pits of antlion (Myrmeleontidae) keys) or include them as a minor constituent of their
andwormlion (Vermileonidae) larvae. Lures include omnivorous diets (for example, field mice, chim-
fiowers,which attract insect prey to within striking panzees, humans). Predation pressure on arthropod
distance of predators that wait on or behind the populations has led to a spectacular array of protective
fiower.Crab spiders and ambush bugs apply this tech- measures directed primarily against vertebrate preda-
nique.Often they are the same color as the flower torso These may be classified as follows: (l) Defense and
wherethey wait (Figure 4-15), and some mantids offense discourage an attack by resisting, threatening,
90 Chapter4 BehaviorandEcology

Figure4-16 Eye spot on owl butterfly Caligo (Satyridae: JiI!'
.- ~
Brassolinae). Eye spots occur on many kinds of insects. .:1
Experiments demonstrate that vertebrate animals are ~o
frightened by them, probably because of their resemblance
B :¡:
to the eyes of large predators, such as owls and cats.

Figure4-17 Mimicry of a yellow jacket (B) by a elear-

wing moth (Parathene sp.)(A).
or injuring a predator. The devices may be mechanical,
chemical, or visual, may involve mutualistic defenders,
such as pugnacious ants (discussed earlier), and may
be used actively, passively, or both. Active mechanical
defenses inelude biting, spiking, stinging, and frighten- ing dark background. These patterns are easily learned
ing sounds; passive defenses inelude spines, hairiness, and easily seen. The insects enhance the patterns' ef-
and thick, hard cutieles. Chemically defended insects fectiveness by behaving ostentatiously (for example,
deploy allomones, ineluding noxious, painful, and poi- being exposed openly, walking and flying slowly, wav-
sonous substances that are actively ejected from the ing conspicuous body parts) and forming aggregations.
body or injected by sting into the victim's body, or are Familiar Müllerian mimics are the warning-colored
deployed passively within the insect or available in and aggressive vespid wasps, which share yellow-and-
needlelike spines that penetrate the predator on con- black markings. Tropical Müllerian "rings" inelude a
tact. Visual defenses inelude frightening or threatening host of slow-flying, long-winged butterflies and moths
appearance (sudden movements, startle displays, flash that share a common orange-yellow-black color pat-
colors, eyespot exposure, predator simulation = defen- tern and behavior. Some members of such rings are not
sive mimicry) (Figure 4-16), aposematism (warning distasteful and therefore are Batesian, rather than Mül-
colors, lights, and sounds that are combined with lerian, mimics. (2) Disguise involves appearing to be
chemical or mechanical defenses that the predator something else, so the attack is not initiated. This in-
learns), and Müllerian mimicry (warning signals eludes the following: (a) Batesian mimicry, in which an
shared among several distasteful species, which serve unprotected mimic resembles a distasteful species-
as both models and mimics of each other, providing the model-with which the predator has had an un-
mutual protection). Aposematic insects typically use pleasant experience. Batesian mimic-model systems are
bold colors, such as red, orange, yellow, or white, pre- ubiquitous, most commonly seen among various flies,
sented as stripes, spots, bars, and rings on a contrast- beetles, and moths (Figure 4-17) that mimic wasps
CommunityAssociations 91

and bees. This tactic includes insects whose body parts

resemble a separa te insecto A widely cited North Amer-
ican Batesian mimic is the viceroy (Figure 4-18), a
usually edible butterfIy that closely resembles the
monarch, a butterfly whose body contains poisonous
cardiac glycosides derived from milkweed on which
the larva feeds. A variation on Batesian mimicry is the
resemblance of an edible mimic to a model that the
A predator learns is difficult to catch. Thus, some weevils
closely resemble sarcophagid fIies. (b) Wasmannian
mimicry, in which symphiles resemble their carnivo-
;;; rous, defensive social hosts (discussed earlier) and
] thereby prevent host attack. Symphiles that travel out-
g side the nest with their hosts frequently function as
~ Batesian mimics, too. (c) In object resemblance, the in-
C> sect looks like an inanimate and inedible object, such
-g as a rock, stick, leaf (Figure 4-19) , or feces (Figure 4-20).
~ (d) In thanatosis, the insect feigns death. (3) In con-
~ cealment, the insect seems not to be there. This ap-
~ proach includes background resemblance, counter-
! shading (= obliterative shading: body appears fIat,
.§, rather than solid and three-dimensional), shadow elim-
~ ination (body appears joined to substrate), disruptive
¡§ coloration (body outline is obliterated), natural cam-
oufIage (body covered with natural materials, such as
tree bark, living lichens, the remains of an insect's prey,
4-18 An example of mimicry in butterflies.
or its own feces), and hiding in an unexposed place;
A,Theviceroy, Basilarchia archippus (Cramer); B, The
insects can hide under physical objects or in nests
monarch,Danaus plexippus (L.).
or cases of their own construction. Open conceal-
ment works best when the insect remains motionless.



Figure4-19 Katydids (Tettigoniidae) that resemble (A) diseased or dying leaf,

(B) completely dead leaL Oetails of this disguise include designs that suggest the
presence of a leaf midrib and methods for concealing the presence of legs.
92 Chapter4 BehaviorandEcology

evasive maneuvers, and detachable body parts. (b) In

disorientation various deceptive structures or behaviors
cause the predator to desist from attack or fail to com-
plete the attack. This includes dejlection, the use of
conspicuous marks, a false head, and autotomy (drop-
ping a body pan), to direct the predator's attention to-
ward expendable items while the prey escapes in an
A unanticipated direction; and confusion, the use of
strange shapes and movements (predator does not rec-
ognize it as prey) and swarming (predator has difficulty
concentrating on one individual).
There are two things to keep in mind about the
preceding antipredator-device categories: First, not all
cases fall neatIy into a single category, but rather com-
bine aspects of two or more tactics. Second, many spe-
cific instances of disguise, concealment, warning, and
so on have been inferred from their appearance to the
human eye and have yet to be tested experimentally.
Yet, so far, all carefully reasoned inferences have been
confirmed by experimentation.

A large proponion of anhropod species feed on dead
and decaying plants and animals, recycling them into
living biomass. These saprophagous anhropods oc-
B cur in many orders but are found chiefiy in the Acari,
Blattodea, Isoptera, Coleoptera, and Diptera. They all
have chewing, rasping, or chelate mouthparts. Most
specialize either in plant or animal materials, partIy
beca use ovipositing aduIts locate them by very differ-
ent means but also because different digestive mech-
anisms are employed. Technically, most detritivores
do not feed exclusively on dead material itseU but
feed instead on a mixture that includes bacteria,
fungi, nematodes, and other minute organisms that
occur with dead tissue and share in its breakdown.
The detritus community also includes predators and
Figure4-20 Bird-feces resemblance: Insects using this parasites that feed on the detritivores. What makes
disguise depend on a composite of brown and white col- saprophagous insects especially imponant, ecologi-
ors, typical of bird feces. A, Moth openly exposed on a cally, is that they are relatively large and mobile, dis-
leaf at rest, resembling bird feces. The elongate scales ex- persing large amounts of detritus by working it into
tend laterally, giving the impression of splatter. B, Swal- the soil, carrying it to scattered locations, or com-
lowtail butterfly larva. Its glossy cuticle gives the impres- pleting their life cycles in it so that the next genera-
sion of wet bird feces. Its second line of protection, when tion fiies away.
disturbed, is to extend the two-pronged glandular hom Plant detritivores live chiefiy in the immediate
(the osmeterium) behind its head, which releases a nox- vicinity of standing crops of plants, such as forests,
ious vapor. grasslands, and ponds, where the dead material accu-
mulates on the ground or in aquatic sediments. A large
part of the soil microfauna of the world consists of var-
(4) Escapeandevasioninvolvebeing hard to capture, so ious mites, millipedes, springtails, and beetIes that feed
the predator cannot complete the attack. This method on finely divided plant material and its associated mi-
occurs in two forms: (a) In jleeing the predator cannot croorganisms. More obvious on the forest floor are
maintain pursuit or achieve capture, such as fast escape fallen leaves, branches, and tree trunks, which also
(by running, jumping, swimming, flying, dropping), suppon a diversity of arthropods and hasten their con-
InsectImpactin Biotic Communitiesand Ecosystems 93

versioninto soil. Even recently killed trees that are still Animal carcasses and dung piles are "fugitive habi-
standing attract specialists whose larvae develop tats," environments with resources that quickly dissi-
withinthe wood. By processing the larger components pateo During the period of utility, they are invaded by
of plants, large arthropods make plant matter more successive but overlapping waves of insects that deal
readilyavailable to microscopic arthropods, to mi- effectively with certain stages in the degradation and
erobes,and ultimately to the root systems of living drying process and with each other. A cow pat in North
plantsas mineral nutrients. America often is invaded first by horn flies and face
Animaldetritivores form a much smaller part of flies, whose females oviposit on it immediately after the
mostcommunities, because much less animal material feces drop to the ground. Yellow dung flies, sepsids,
isavailablefor degradation. Some arthropods, such as and ulidiid flies come, then leave, at various stages as it
adultseorpionflies, specialize in the dead bodies of cools and a crust forms. These are followed by scarab
otherarthropods, but the most obvious ones process and hydrophilid beetles, and by staphylinid beetles that
the dead bodies of medium-sized and large verte- either feed on the eggs of other insects or parasitize de-
brates.In its early stages of decay, a carcass is so rich veloping fly larvae. Soldier flies show up still later. By
inorganicnutrients and is so easily digested, that sar- the time the cowpat is stiff and only moist in the mid-
eophagous(= flesh-eating or carrion-feeding) insects dIe, it is occupied primarily by the larvae and pupae of
engagein intense scramble competition. The most slow-growing fly and scarab-beetle larvae, each devel-
quiekand efficient of these are the blow flies and flesh oping in zones of the cow pat that differ in physical
fiies,which in warm, humid weather can reduce a characteristics (crust, center, base). Arthropod succes-
largecorpse to a skeleton in a few days. Gravid females sion in carcasses follows a similar pattern, typically
startlayingeggs in an animal's orifices within an hour with blow flies appearing first, beetles later. In late
ofits death, and the eggs hatch in less than a day (or stages of decay, the carcass community may consist
aredeposited as ready-to-feed larvae, in the case of mainly of dermestid beetles and moths that eat dried
fieshflies).When the hundreds or thousands of fly lar- skin and ligaments. The exact species composition and
vae(maggots)have matured, they crawl away and pu- rate of change depend not only on the geographic 10-
pateunderground nearby. Other carrion feeders spe- cality but also on the exact habitat (carcasses transform
cializein small birds and rodents. A mated pair of very differently in sun and shade), weather, and season.
bUlyingbeetles lowers the carcass into the ground and Knowledge of the pattern and speed of faunal succes-
eonvertsinto an edible nest for its young (as discussed sion in a carcass can be critical to forensic investiga-
earlier). torso From data on the arthropod species and their
Dung-feeding (coprophagous) arthropods are, stages in a corpse when it is discovered, it is possible to
eeologicalIy,either plant or animal detritivores, de- estimate the postmortem interval and thus the time of
pendingon the diet of the animal producing the feces. death.
lnsectstend to specialize in the dung of kinds of dif-
ferentkinds of animals, beca use the size and consis-
teneyof the dung pile determines its shape and vol-
umeand therefore how quickly it will dry out. Dung
beetles(Scarabaeidae), as a group, are able to use the Insect Impact in Biotic Communities
greatest variation in dung-pile sizes, beca use some
and Ecosystems
speciesdevelop within the dung in situ, but others
quiekly gather the dung into packets or balls and
lowerit into tunnels beneath the pile or roll it away,lO
Insed Diversity
provisiontheir underground nests. Yet different dung Insects are vitally important members of biotic com-
beetlespecies focus on dung from herbivorous mam- munities for two reasons: There are so many of them,
malsof different sizes. larval development of flies oc- and there are so many different kinds. The average
eurs within the dung pile itself, so relatively large abundance of any one species is simply a function of
dungpiles, such as the "cow pats" produced by large how small it is, how large its resource base is, and how
ungulates, form the best breeding medium. One im- many enemies exploit it. But insects are unusual
portantconsequence of. this situation was that intro- among organisrns in their abundance of species. Ac-
ducing cattle to Australia also introduced a huge cording to conventional ecological theory, each species
soureeof dung, which the native kangaroo-dung bee- occupies a unique niche, a unique way of living that it
des could not handle, but which was ideal for bush performs better than any other species and that is de-
fiiesand buffalo flies. Importation of African dung fined by where it lives and what it consumes. In some
beetlesthat disintegrate cow pats have helped bring communities insect species are so tightly packed that
pestfly populations under control. they have extensively overlapping niches, resulting in

94 Chapter4 BehaviorandEcology



~ T'"S
~h :)




.~/ ~:~~//

'~D ~




Figure4-21 Food web on a single plant, illustrating the presence of three guilds of
plant-feeding insects: 18 species of pit feeders, 17 species of strip feeders, and 59 species
of sap feeders. The parasitoids and predators of these insects, not numbered, also form

competition, but partitioning and environmental insta- This is the strongest and most efficient skeleton Cor
bility prevent the exclusion of one competitor by an- small-bodied animals. In addition to giving body sup-
other. Thus we find many insect guilds, groups of in- port, external protection, and opportunities for evolv-
sect species that use the same food source but do so in ing a wide variety of hard tools for handling food and
slightly different ways (Figure 4-21). The fact that over other materials, it provides leverage for long ap-
half of all described species of organisms, and about pendages, allowing elevated body support and quick
three quarters of all animal species, are insects suggests movements, and it offers resistance to desiccation.
they are exceptional in their ability to assume a myriad These features allowed arthropods to become the firsl
of unique lifestyles. animals to invade land, along with vascular plants, so
There appear to be four interconnected reasons for a vast variety of potential niches were available 10
this diversity, all directly or indirectly resulting in a them, and they could diversify along with the plants.
high speciation-to-extinction ratio: (1) Exoskeleton: (2) Small size: More niches are available to small-bodied
InsectImpactin Biotic Communitiesand Ecosystems 95

animals, because many kinds of food and space are numbers. The latter are responsible for wide and
available only in small quantities and limited sizes. erra tic population fluctuations. The endogenous fac-
Smallanimals also have shorter generation times, al- tors are all density dependent, including changes in
lowing rapid evolution and adaptation to new condi- disease-inducing stress levels, social interactions, emi-
tions.And small animals have a large surface-to-volume gration rates, and gene pool composition. The density-
ratio,which has beneficial consequences: (a) proximity dependent/independent distinction is a useful concept,
of organs to each other, which allows use of simple but insect populations often are controlled by a com-
circulatoryand respiratory systems; (b) greater muscle plex interplay of weather-related biotic factors, so that
strengthin relation to size, which allows thinner limbs; equilibrium densities are superimposed on erratic or
and (c) greater air resistance, which allows greater seasonal changes caused by changing physical condi-
wingand body 10ft, facilitating airborne dispersal and tions. In other words, an insect may have several equi-
evolution of powered flight. However, a large surface librium densities, depending on the weather, and if the
areais disadvantageous beca use desiccation and heat effects of weather change often enough, population
1055are more rapid. (3) Wings: The greater mobility regulation is almost completely obscured. Some insect
provided by flight allows animals to exploit widely populations may never reach the point where reduced
scattered resources, which form distinct niches. Fur- resources and increasing predation prevent further
thermore, it allows them to colonize new areas rapidly, population growth. In others, fluctuations are caused
reducing the likelihood of extinction and providing by density-dependent factors that have effects extend-
opportunities for genetic isolation and species forma- ing long beyond the immediate causes (for example,
tion. (4) Complete metamorphosis: Transformation of desert locusts, as mentioned earlier) or are the result of
the body from one kind of animal to another has the environmental feedback that works over time scales of
followingconsequences: (a) a temporary resource that many years (many forest Lepidoptera). Therefore, an
isonlypart of a unique niche can be exploited without intimate knowledge of natural population control may
having to sustain all active stages; (b) more unique be necessary before insect outbreaks can be predicted
nichesare available, because the "compound niche" of with any reliability.
the two different active forms (larva and adult) is, as a
whole,another niche, reducing competition with over- ArtificialControl: The principies that determine popu-
lapping species and avoiding competitive exclusion lation density in nature can be applied by humans to the
even by species that occupy only one niche or the judicious control of insect pests. Pest control is not ap-
other; (c) individuals escape from natural enemies by propriate, economically, unless pests are dense enough
disappearingfrom one of the microhabitats before ene- to reduce the value of some product or resource more
miesbecome too numerous; (d) specialization in either than it costs to prevent that reduction. The breakeven
growthand storage (the larva) or dispersal and repro- point, the pest density at which the gain in value from
duction (the adult) results in increased efficiency of control is equal to the cost of that control, is the eco-
eachstage. The fact that most insect species occur in nomic injury leve!, or EIL. For insects that are largely
orderswith complete metamorphosis suggests that this density dependent, the relationship between the EIL
featureis exceptionally important. and the equilibrium density determines whether mea-
sures should ever be taken against them. Some pests
cause relatively minor damage to crops, livestock, or
Size and Its Control
human health, rarely exceeding their EILs, because
Thenumbers of individuals in an insect population is their equilibrium densities are low, thanks to natural
determinedfirst by the size of the entire community in enemies or restricted breeding sites. Others may cause
whichthey can exist, their biotope. Beyond that, their extensive damage but still rarely exceed their EILs. This
densityis controlled by an array of exogenous and en- occurs if control is very expensive, the product has a
dogenous factors affecting reproduction, death, and low value, or it is tolerant to high damage so the eco-
migration rates. Exogenous factors are principally nomic 1055is small. However, when pests surpass a cer-
[ood, space, natural enemies (predators, parasites, tain lower level Cthe economic threshold) indicating that
pathogens),and weather. The first three vary in their they soon will exceed the EIL, control measures should
intensityaccording to how dense the population is, be implemented immediately in order to take effect by
and are termed density-dependent factors, responsible the time the EIL is reached. Aside from environmental
for regulating the density within somewhat narrow concerns, the decision to control pests in agricultural
bounds around the species' equilibrium density. systems has an exclusively economic basis. The deci-
Weather-relatedfactors, in contrast, usually are density- sion to control insects of medical importance is compli-
independent,having an effect proportional to insect cated by the need to consider not only losses in human
96 Chapter4 BehaviorandEcology



Figure4-22 Simple freshwater aquatic food web, showing the pivotal roles of insects
at the primary- and secondary-consumer trophic leve\s.

productivity and costs of treatment, but also the ing how weather-related factors and biotic factors com-
amount of suffering caused. A knowledge of what de- bine to determine population growth and depression
termines the natural equilibrium density may be crucial help entomologists anticipa te conditions when the EIL
to effective time management. For example, the appli- wiIl be exceeded. The ideal goal is to find low-impact
cation of pesticides may temporarily elimina te natural ways to alter a pest's environment so that the EIL is
enemies of the pest, not just the pest, causing its rapid never reached and insecticides are unnecessary.
rebound and biotic re/ease. In this reaction, the pest den-
sity soars until it finds a new, higher equilibrium posi- Insed Roles in Food Chains and Webs
tion, causing more damage and requiring more frequent
application of insecticides. This situation, which is Insects and other arthropods are major constituents of
costly and can lead to insecticide resistance more trophic pyramids because of the sheer mass of num-
quickly, is known as the "pesticide treadmill.» For both bers in populations of each species at aIl consumer
density-independent and density-dependent pests, know- leve\s. In every case studied, insects eat and incorpo-

References 97

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same communities. In fields in North America, more which generally are perceived as being occupied by
than three quarters of the energy flow from plants such carnivores as eagles and lions, the ectoparasitic
passes through populations of a few species of Or- lice, fleas, biting midges, and skin mites feed on these
thoptera, dwarfing the importance of birds and mice. so-called top predators. In addition, arthropods are
In terms of living biomass, just the ants of the world, pivotal in a myriad of intersections within food webs
a single family consisting of about 11,000 species, (Figure 4-22) because of their diversity, as reflected in
weigh as much as the 6 billion humans of the world. the number of species in every terrestrial and freshwa-
In the Amazon rain forest, ants and termites make up ter community. Without arthropods in the picture,
about one third of the biomass of all animals, from communities most likely would consist of fewer
mites to tapirs, and the dry weight of ants there is four trophic levels, with drastically reduced trophic links
times as great as the dry weight of allland vertebrates among the remaining organisms.


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5 PhylumArthropoda1

e are concerned in this book principally with in- 11. Respiration by means of gills, or tracheae and spi-
Wsects,but it is appropriate to point out the place of racles
theinsectsin the animal kingdom and to include at 12. No cilia or nephridia
leasta briefaccount of the animals most closely related 13. The sexes nearly always separate
toandsometimes confused with the insects.
The position of the Arthropoda within the animal
Theinsects belong to the phylum Arthropoda, the
kingdom and, indeed, even the reality of the arthro-
principalcharacters of which are as follows:
pods as a monophyletic group are subjects of active re-
1.Thebody segmented, the segments usually search and controversy. The most serious challenge to
groupedin two or three rather distinct regions the hypothesis of arthropod monophyly began in the
2. Paired,segmented appendages (from which the 1940s and 1950s (see, for example, Tiegs and Manton
phylumgets its name) 1958) and reached its pinnacle with the publication of
3. Bilateralsymmetry The Arthropoda by Sidnie Manton in 1977. The essence
4. Achitinous exoskeleton, which is periodically of this position is that the detailed study of arthropod
shedand renewed as the animal grows functional morphology reveals a tremendous amount
5. Atubular alimentary canal, with mouth and anus of diversity, not only in structure, but in the way in
6. Anopen circulatory system, the only blood vessel which those structures actually work. Manton's own
usuallybeing a tubular structure dorsal to the ali- studies dealt largely with mandibular and locomotory
mentarycanal with lateral openings in the abdom- mechanisms, and this work was supplemented by
inalregion other research in embryonic development (Anderson
7. Thebody cavity a blood cavity or hemocoel, the 1973). The structures of modern arthropods are closely
coe\omreduced integrated and coordinated to forrn a complex, func-
8. Thenervous system consisting of an anterior gan- tional whole, and they were interpreted to be so dis-
glionor brain located above the alimentary canal, tinctive between the major groups that no interrnediate
a pairof connectives extending from the brain between them could be conceived that, in turn, could
aroundthe alimentary canal, and paired ganglion- have been functional in the common ancestor. As a re-
atednerve cords located below the alimentary sult, these features must have evolved independently in
canal the groups that we call arthropods. Manton proposed
9. Striatedskeletal muscles and strongly defended the idea that those groups tradi-
10.Excretionusually by means of tubes (rhe tionally placed within Arthropoda had, in fact, inde-
Malpighiantubules) that empty into the alimen- pendently evolved those characteristics listed at the be-
tarycanal, the excreted materials passing to the ginning of this section and, therefore, that the
outsideby way of the anus Arthropoda are polyphyletic. This position has been
defended recently by Fryer (1997).
The weakness in this argument, of course, is that
IAnhropoda:arthro,joint or segment; poda, [oot or appendage. it addresses the weaknesses in evidence for monophyly 99

100 Chapter5 PhylumArthropoda

but doesnot propose a testable alternative hypothesis.

To adequatelydemonstrate that a hypothesis of mono- Classification of the Arthropoda
phyly is false (that the Arthropoda are not mono-
phyletic), it is necessaryto provide evidence showing There are differences of opinion regarding the relation-
that one or more "arthropod" taxa are more closely re- ships of the various arthropod groups and the taxo-
lated to a nonarthropod group. No such evidence has nomic level at which they should be recognized. A
been offered. Today,most workers accept that the evi- number of different taxonomic arrangements of these
dence available is most parsimoniously explained by groups have been proposed. We follow here the classifi-
the hypothesis that the Arthropoda are, indeed, mono- cation of Barnes (1987) and recognize four major
phyletic. groups within the arthropods as subphyla; this arrange-
Traditionally, researchershave held arthropods to ment is as follows (with synonyrns in parentheses):
be most closely related to the Annelida and the Ony-
Phylum Arthropoda-arthropods
chophora. The Annelida, which include the segmented Subphylum Trilobita-trilobites (fossils only)
worms (earthworms, marine worms, and leeches) dif-
Subphylum Chelicerata
fer from the arthropods in lacking segmented ap- Class Merostomata-horseshoe crabs
pendages,a chitinous exoskeleton, and a tracheal sys-
(Xiphosura) and the fossil eurypterids
temoThey havea closedcirculatory system;the skeletal (Eurypterida)
muscles are not striated; and excretion is by means of Class Arachnida-arachnids
ciliated tubes called nephridia. Someinsect larvae lack
Class Pycnogonida-sea spiders
appendages and superficially resemble annelids. They
Subphylum Crustacea-crustaceans
can be recognized as insects by their internal organiza-
Class Cephalocarida
tion (different types of circulatory and excretory sys- Class Branchiopoda
tems and the presence oEtracheae). The Onychophora Class Ostracoda
resemble the arthropods in having (1) segmentation Class Copepoda
(the body is indistinctly segmented, the segmentation Class Mystacocarida
being indicated by the legs, which are unsegmented Class Remipedia
but bear claws at their apex), (2) a chitinous exoskele- Class Tantulocarida
ton that is periodically shed and renewed, (3) a tra- Class Branchiura
cheal system, and (4) an open circulatory system. They Class Cirripedia
resemble the annelids in having nephridia and unstri- Class Malacostraca
ated skeletal muscles. The Onychophora are wormlike Subphylum Atelocerata
or sluglike animals, varying in length up to several cen- Class Diplopoda-millipedes
timeters. They occur only in the southern hemisphere, Class Chilopoda-centipedes
where they live in moist situations. Class Pauropoda-pauropods
Recent molecular studies, however, have com-
Class Symphyla-symphylans
pletely changed the perspective on animal relationships Class Hexapoda (Insecta)-hexapods
(Aguinaldo et al. 1997 and many following works). In
the resulting classification, the Arthropoda are grouped Three phyla sometimes included in the Arthro-
together with the Nematoda and a few other small poda are the Onychophora, Tardigrada, and Pentasto-
phyla into the Ecdysozoa. This huge group of animals mida (= Linguatulida). The Onychophora are some-
was recognized by similarities in the sequences of ribo- what intermediate between the Annelida and the
somal DNA, but apparently are all characterized by a Arthropoda (as noted previously), but the other two
cuticle that is periodically molted (hence the name: ec- groups are somewhat degenera te morphologically (for
dysis, escape from; zoon, animal). Several subsequent example, they lack circulatory and excretory organs).
studies, incorporating data from other sources, have The preceding arrangement of the arthropod
confirmed this resulto The segmented worms are now groups is based primarily on the character of the ap-
placed together in another huge group of protostome pendages (partially the jaws and legs) and the nature of
animals, the Lophotrochozoa, that includes not only the body regions. The trilobites, crustaceans, and Ate-
the Annelida, but the Mollusca, Platyhelminthes, and locerata have a pair of antennae (usually two pairs in
Rotifera. There is, as yet, no consensus on the position the crustaceans), whereas the chelicerates lack anten-
of the Onychophora, although most researchers agree nae. The endite lobes at the base of certain appendages
they should not be classified within the Arthropoda. (the gnathobase) function as jaws in the trilobites, che-
This classification is not unequivocally accepted, and licerates, and crustaceans, whereas (according to
the entire subject is a focus of current research. Manton 1964, 1977) the Myriapoda and Hexapoda bite
Classification of the Arthropoda 101


A B e

Figure5-1 Some major variations in arthropod appendages. A, Generalized; B, Trilobite;

C, Crustacean. enl, endite lobe; enp, endopodite; exl, exite lobe; exp, exopodite.

withthe tips of the mandibles. However, recent studies tudinal divisions of the body. (The lateral divisions
ofthe expression of the gene distal-less in arthropod were extensions over the bases of the legs.) Trilobites
mouthpartsindicate that the mandible of hexapods is had a pair of antennae, with the remaining appendages
alsognathobasicin nature. similar and leglike (Figure 5-IB). The anterior part of
Arthropod appendages are subject to a great deal the body (the preoral region and the first three postoral
of variation but are basically seven-segmented (Fig- segments) was covered with a carapace. The legs had
ure5-l). The basal one or two segments sometimes an exite lobe or epipodite of the basal segment; this
bearmesal(endite) or lateral (exite) lobes or processes. bore a series of lamellae and apparently functioned as a
Ihese lobes or processes frequently have important gill (lhese animals were marine). The endite lobes of
functionsand sometimes have special names. The en- the anterior legs apparently functioned as jaws.
ditelobes of certain appendages usually have a chew-
ingfunction in the trilobites, chelicerates, and crus- Subphylum Chelicerata3
taceans,and an exite lobe of the basal segment often
Animals belonging to the subphylum Chelicerata lack
functionsas a gill in the trilobites and crustaceans. In
antennae and typically have six pairs of appendages.
the crustaceans, the second segment usually bears a
The first pair are the chelicerae, and the rest are leglike.
well-developed exite lobe, which is generally seg-
Endite lobes of the pedipalps (lhe second pair of ap-
mentedand is sometimes as large as or larger than the
pendages in the arachnids), or the leglike appendages
restof the appendage, giving the appendage a forked
in the horseshoe crabs, function as jaws. The body of a
appearance.Such an appendage is spoken of as bira-
chelicerate usually has two distinct divisions, an ante-
mous.The exite lobe of the second segment is called
rior region called the prosoma (or cephalothorax) and a
theexopodite,and the rest of the appendage the en-
posterior region called the opisthosoma (or abdomen).
dopodite (Figure5-IC).
The prosoma bears the chelicerae and the leglike ap-
pendages. The genital ducts open to the outside near
SubphylumTrilobita2 the anterior end of the opisthosoma. Most chelicerates
Ihe trilobites lived during the Paleozoic era (see Table
8-2), but were most abundant during the Cambrian 'Trilobita: tri, three; lobita, lobed (referring to the three longitudinal
andOrdovician periods. These animals were somewhat divisions of the body).
elongateand flattened, with three rather distinct longi- 3Chelicerata: with chelicerae.
102 Chapter5 PhylumArthropoda

have an extra leg segment, the patella, between the fe-

mur and the tibia. The legs are generally uniramous;
that is, there is no exite or exopodite.

Class Merostomata4

Subclass Eurypteridas: The Eurypterida lived during

the Paleozoic era, from the Cambrian to the Carbonif-
erous periods. They were aquatic and somewhat simi-
lar to the present-day Xiphosura, and some reached a A
length of more than 2 meters. The prosoma bore a pair
of chelicerae, five pairs of leglike appendages, and a
pair each of simple and compound eyes; the opistho-
soma was 12-segmented, with platelike appendages
concealing gills on the first 5 segments, and the telson
was either spinelike or lobelike.
Subclass Xiphosura6-horseshoe crabs or king crabs:
The horseshoe crabs are marine forms and are quite
common along the Atlantic Coast from Maine to the
Gulf of Mexico. They live in the shallow water and
along sandy or muddy shores where they spawn. They
feed chiefly on marine worms. Horseshoe crabs are eas-
ily recognized by their characteristic oval shell and
long, spinelike tail (Figure 5-2).

Class Arachnida-Arachnids7
The Arachnida constitute by far the largest and most
important class of the Chelicerata (about 65,000 de-
scribed species, with about 8,000 in North America),
and a person studying insects will probably encounter
more of them than of any other noninsect group of
arthropods. Its members occur almost everywhere, of- Figure 5-2 A horseshoe crab, Limulus sp. (subclass
ten in considerable numbers.
Xiphosura). A, Dorsolateral view; B, Ventral view. bg,
Most authorities recognize 11 major groups of book gills; ch, chelicera.
arachnids (all represented in North America), but not
all agree on the names to use for these groups or the
taxonomic level they represent. We call these groups
ordcrs (some would call them subclasscs) and arrange Schizomida (Tartarides, Schizopeltida, Colopyga,
Pedipalpida in part)-short-tailed whipscorpions
them as follows (with other names and arrangements
in parentheses): Amblypygi (Amblypygida, Phrynides, Phryneida,
Scorpiones (Scorpionides, Scorpionida)-scorpions Phrynichida, Pedipalpida in part)-tailless
whipscorpions, whipspiders
Palpigradi (Palpigrada, Palpigradida, Microthely-
phonida)-micro whipscorpions Araneae (Araneida)-spiders

Uropygi (Thelyphonida, Holopeltidia, Pedipalpida in Ricinulei (Ricinuleida, Meridogastra, Podogonata,

part)-whipscorpions Rhinogastra)-ricinuleids
Opiliones (Phalangida, including Cyphophthalmi)-
Acari (Acarina, Acarida)-mites and ticks
4Merostomata: mero, part; stomata, mouth.
'Eurypterida: eury, broad; pterida, wing or fin. Pseudoscorpiones (Pseudoscorpionida, Chernetes,
.Xiphosura: xipho, sword; ura, tail. Chelonethida)-pseudoscorpions
7 Arachnida: from the Greek, meaning a spider. Solifugae (Solpugida)-windscorpions
Keyto the Ordersof Arachnida 103

Keyto the Orders of Arachnida

l. Opisthosoma (abdomen) unsegmented or, if segmented, with spinnerets

posteriorly on ventral side (Figures 5-5 and 5-8, ALS,PLS,PMS) 2
1', Opisthosoma distinctly segmented, without spinnerets 3
2(1). Opisthosoma petiolate (Figures 5-5 and 5-9 through 5-13) Araneae p.105
2'. Opisthosoma not petiolate, but broadly joined to prosoma (Figures 5-15
through 5-22) Acari p.129
3(1'). Opisthosoma with a tail-like prolongation that is either thick and
terminating in a sting (Figure 5-3) or slender and more or less whiplike
(Figure 5-4A,B); mostly tropical 4
3'. Opisthosoma without a tail-like prolongation or with a very short
leaflike appendage 7
4(3). Opisthosoma ending in a sting (Figure 5-3); first pair of legs not greatly
elongated; second ventral segment of opisthosoma with a pair of comblike
organs Scorpiones p. 104
4', Opisthosoma not ending in a sting; first pair of legs longer than other
pairs (Figure 5-4A,B); second ventral segment of opisthosoma without
comblike organs 5
5(4'). Pedipalps slender, similar to legs (Figure 5-4A); minute forms, 5 mm
or less in length Palpigradi p.104
5'. Pedipalps usually much stouter than any of the legs (Figure 5-4B);
mostly larger forms 6
6(5'). With two median eyes on a tUbercle anteriorly and a group of three eyes
on each lateral margin; taillong, filiform, and many-segmented; pedipalps
nearly straight or curved mesad, extending forward, and moving laterally
(Figure 5-4B); body blackish, more than 50 mm in length Uropygi p.104
6'. Eyeslacking;tail short, one- to four-segmented;pedipalpsarchingupward,
forward, and downward and moving vertically; body yellowish or
brownish, less than 8 mm in length Schizomida p.105
7(3'), Pedipalps chelate (pincerlike) (Figure 5-23); body more or less oval,
fiattened, usually less than 5 mm in length Pseudoscorpiones p.135
7'. Pedipalps raptorial or leglike, but not chelate; body not particularly
fiattened; size variable 8
8(7'). Chelicerae very large, usually about as long as prosoma, and extending
forward, and body slightly narrowed in middle (Figure 5-4C); color pale
yellow to brownish; length 20 to 30 mm; mostly nocturnal desert forms
occurring in western United States Solifugae p.135
8'. Without the preceding combination of characters 9
9(8'). First pair of legs very long, with long tarsi; opisthosoma constricted at
base; mainly tropical 10
9'. First legs similar to the others and (except in the Opiliones) not usually
long; opisthosoma not particularly constricted at base 11
10(9). Prosoma longer than wide, lateral margins nearly parallel, and with a
transverse membranous suture in caudal third; opisthosoma with a very
short terminalappendage;length 5-8 mm (see also couplet 6') Schizomida p.105
lO'. Prosoma wider than long, lateral margins rounded or arched, and without
a transverse suture; opisthosoma without a terminal appendage; length
variable, up lOabout 50 mm Amblypygi p.105
104 Chapter5 PhylumArthropoda

11(9'). Orange-red to brown arachnids, about 3 mm in length, with broad flap

at anterior end of prosoma concealing chelicerae; legs not unusually
long, animal somewhat mitelike or ticklike in appearance; recorded
from Rio Grande Valley of Texas Ricinulei p.128
11'. Size and color variable, but without broad flap at anterior end of
prosoma covering chelicerae; legs variable in length, often very long
and slender (Figure 5-14); widely distributed Opiliones p.129

ORDERScorpionesB-Scorpions:The scorpions are sting. The young are born alive and, for a time after
well-known animals that occur in the southern and birth, ride about on the mother's back. Scorpions grow
western parts of the United States. They are fair-sized slowly, and some species require several years to reach
arachnids, varying in length up to about 125 mm. The maturity. The function of the pectines is not known,
opisthosoma is broadly joined to the prosoma and is dif- but they are believed to be tactile organs.
ferentiated into two portions, a broad seven-segmented The effect of a scorpion sting depends primarily
mesosoma and a much narrower five-segmented poste- on the species of scorpion involved. The sting of most
rior metasoma that terminates in a sting (Figure 5-3). species is painful and usually accompanied by local
The prosoma bears a pair of eyes near the midline and swelling and discoloration, but it is not dangerous. Of
two to five along the lateral margin on each side. The the 40-odd species of scorpions in the United States,
pedipalps are long and chelate. On the ventral side of the only one-Centruroides sculpturatus Ewing-is very
second opisthosomal segment is a pair of comblike venomous, and its sting may be fatal. This species is
structures, the pectines. slender and rarely exceeds 65 mm in length. It varies in
Scorpions are largely nocturnal and during the day color from almost entirely yellowish to yellowish-
remain concealed in protected places. When a scorpion brown with two irregular black stripes down the back.
runs, the pedipalps are held outstretched and forward, There is a slight dorsal protuberance at the base of the
and the posterior end of the opisthosoma is usually sting. As far as is known, this species occurs only in
curved upward. Scorpions feed on insects and spiders, Arizona.
which they catch with their pedipalps and sometimes Scorpions do not ordinarily attack people but will
sting quickly if disturbed. In areas where scorpions oc-
8Scorpiones: from the Latin. meaning a scorpion. cur, be careful in picking up boards, stones, and simi-
lar objects, and brush off, rather than swat, a scorpion
found crawling on your body.
ORDER Palpigradi9-Micro Whipscorpions: These
are tiny arachnids, 5 mm or less in length, somewhat
spider-like in appearance but with a long segmented
tail (Figure 5-4A). The pedipalps are leglike, and the
first pair of legs is the longest. These animals usually
live under stones or in the soil. This group is repre-
sented in the United States by three species in Texas
and California.
ORDER UropygPO-Whipscorpions: The whipscor-
pions are mainly tropical and, in the United States, oc-
cur only in the South. They are elongate and slightly
flattened, with a slender, segmented tail about as long
as the body, and powerful pedipalps (Figure 5-4B); the
maximum body length is about 80 mm, and the total
length including the tail may be 150 mm or more. They

.Palpigradi: palpi, palp or feder; gradi, walk (referring to the leglike

character of the pedipalps).
Figure5-3 A scorpion, 2X. IOUropygi: uro, tail; pygi, rump (referring to the whiplike tail).

Keyto the Ordersof Arachnida 105

A B e

Figure5-4 Arachnids. A, A micro whipscorpion (Order Palpigradi); B, A whipscor-

pion (Order Uropygi); C, A windscorpion (Order Solifugae). (Courtesy of the Institute of

look somewhat like scorpions but have a very slender in California, is yellowish- to reddish-brown in color
"tail" and no sting. The first pair of legs is slender and and 4.5 to 7.5 mm in length; it lives under rocks and in
used as feelers; only the three hind pairs of legs are leaf litter in the desert regions of southern California.
used in walking. When disturbed, these animals emit A Florida species, Schizomus floridanus Muma, is 3.0 to
(or spray, up to 0.5 meter) a substance with a vinegar- 3.3 mm in length and pale yellowish in color; it lives in
like odor from glands at the base of the tail, and hence crevices in bark and in organic debris, from the Miami
they are often called vinegaroons (or vinegarones) . area south into the Keys.
Whipscorpions are nocturnal and predaceous. They are aRDER Amblypygj12-Tailless Whipscorpions or
generally found under logs or burrowing in the sand. Whipspiders: These arachnids are somewhat spiderlike
The eggs are carried in a membranous sac under the in appearance, but the opisthosoma is distinctly seg-
opisthosoma, and the young ride around on the back of mented and, although narrowed at the base, is not peti-
the female for a time after they hatch. The whipscor- olate. There are no spinnerets; the pedipalps are large,
pion most likely to be encountered in the southern powerful, and spiny (and are used in capturing prey);
states is Mastigoproctus giganteus (Lucas), which oc- and the first legs are very long and whiplike. The pro-
curs in Florida and the Southwest; it has a body length soma is wider than long and has rounded sides. There
of 40 to 80 mm and is the largest species in the order. are no venom glands. The few North American species
aRDERSchizomida"-Short-TailedWhipscorpions: vary in length from about 10 to 55 mm and occur
These animals are somewhat similar to the Uropygi but chiefly in the southern states. They are found under
much smaller and more slender. In addition, the termi- bark or stones (usually scurrying sideways when dis-
nal appendage is not long and whiplike; the pedipalps turbed) and sometimes enter houses.
arch upward and forward and move vertically; and aRDER Araneae13-Spiders: Spiders are a large, dis-
there is a transverse suture on the prosoma. The first tinct, and widespread group with more than 3,700 de-
legs are slender and are not used for walking. The scribed species in North America and more than
fourth legs are modified for jumping, and in the field 38,000 worldwide. The earliest evidence of spiders
these animals superficially resemble small crickets. comes from a 380-million-year-old Devonian fossil
There are no venom glands or eyes. The eight species (Shear et al. 1989). Spiders occur in many types of
ofSchizomida in the United States occur in Florida and habitats and are often very abundant. Typical nondesert
California. Trithyreus pentapeltis (Cook), which occurs
12Amblypygi: ambly, blunt; pygi, rump.
schizo, split (referring to the transverse suture on the 13Araneae: from the Latin, meaning a spider. This section was writ-
prosoma). ten by Jeremy A. Miller and Oarrell Ubick.

106 Chapter5 PhylumArthropoda

habitats may support up to 800 spiders per square me- Two other species in North America, Cheiracanthium
ter. Point estimates of spider diversity range from 20 inclusum (Hertz) (family Miturgidae) and Argiope au-
species per hectare in the temperate zone to more than rantia Lucas (family Araneidae) are reported to have a
600 species per hectare in tropical forests (Coddington mild neurotoxic venom. Despite their imposing ap-
and Colwell 2001). pearance, North American tarantulas (family Thera-
Unique derived characters that define spiders in- phosidae) have relatively innocuous venoms. However,
elude cheliceral venom glands (tost in the family Ulo- they are capable of releasing a eloud of urticating hairs
boridae), abdominal spinnerets, and the modification that can cause discomfort in the mucous membranes of
of the male pedipalps into sperm transfer organs. mammals.

Silk. The ability to produce silk has evolved indepen-

MedicallyImportantSpiders. Although spider bites are
dently in several arthropod lineages. However, spiders
widely feared, few species are dangerous to humans.
are the only group to use silk throughout their lives
Brown reeluse spiders and their relatives (13 North
(Coddington and Colwe1l2001). In addition to its con-
American species ineluding 2 exotics, genus Loxosceles,
spicuous use as a snare to trap prey, silk is used to line
family Sicariidae) and black widow spiders (5 North
burrows, construct retreats and molting chambers,
American species, genus Latrodectus, family Theridi-
make sperm webs, protect developing eggs, and serve
idae) are the most medically important spiders in the
United States. as a dragline. Complex snares, ineluding orb webs, in-
corporate several distinct types of silk. Some spiders
Loxosceles (Figures 5-91, 5-10) lives mostly in the
periodically eat their web and are capable of rapidly re-
Midwest and Southwest. A species introduced from Eu-
cyeling most of the protein into fresh silk (Peakall
rope has been found on the eastern seaboard and else-
1971). Some spiders, especially small species and im-
where in the United States, but the venom of this
mature individuals, use silk for a form of airborne
species is relatively mild (Gertsch and Ennick 1983).
travel called ballooning. To balloon, the spider elimbs to
Brown reeluse venom causes a small, dry, irregular
a high point and releases silk into the air. When the
necrotic lesion that heals very slowly. Contrary to pop-
drag on the silk exceeds the spider's mass, the spider
ular belief, brown reeluse bites cannot be conelusively releases itself into the air.
diagnosed from the wound. Serious medical condi-
Silk is a protein fiber produced in glands that
tions, ineluding Lyme disease, chemical burn, and an-
terminate in spigots on the abdominal spinnerets (Fig-
thrax infection, have been misdiagnosed as brown
ures 5-6, 5-8). In the gland, silk is a water-soluble liq-
reeluse bites, delaying proper treatment (Osterhoudt et
al. 2002, Sandlin 2002). It is not uncommon for such uid protein soup. As the silk is spun, it passes through
an acid bath (Vollrath and Knight 2001). The acid
misdiagnoses to occur outside the range of the brown
hardens the silk by causing the molecules to reorient.
reeluse spider (Vetter and Barger 2002, Vetter and Bush
2002). Complementary regions of the silk molecule align and
bond together in multilayered stacks, forming protein
Latrodectus (Figure 5-HA) is widespread in the
United States. Black widow venom is a neurotoxin. crystals. These crystals are interspersed in a matrix of
loosely arranged amino acids. The protein crystals give
Typical symptoms of envenomation inelude swelling of
the silk its strength, and the loose matrix provides elas-
the lymphatic nodes, profuse sweating, rigidity of the
ticity (Gosline et al. 1986).
abdominal museles, facial contortions, and hyperten-
The physical properties of silk are remarkable.
sion. Antivenin is readily available to counteract the ef-
fects of Latrodectus envenomation. No deaths have Tensile strength is the greatest stress a material will tol-
been attributed to widow bites in the United States erate before failure. Silk is stronger than most natural
since the 1940s. materials and is about half as strong as steel. However,
silk is extremely extensible: It can tolera te substantial
Neither Loxosceles nor Latrodectus species known
distortion (strain) before failure. The product of stress
from North America are aggressive. Unlike groups such
to strain is expressed as toughness and is the total
as fieas, ticks, biting fiies, and bed bugs, spiders do not
amount of energy a material will absorb before failure.
take blood meals from humans and do not normally
Silk has extremely high toughness; steel tolerates very
bite humans except in self-defense.
little distortion in shape, and so is not a very tough ma-
Two other spiders have been implicated in causing
minor necrotic lesions. Both are introduced from Eu- terial (Figure 5-5).
rope. Tegenaria agrestis (Walckenaer) (family Age- Anatomy. The body of a spider is divided into two re-
lenidae) is found in the Pacific Northwest; Cheiracan- gions, the cephalothorax and the abdomen (Figure 5-6).
thium mildei Koch (family Miturgidae) is now The abdomen, usually soft and unselerotized, is at-
widespread in human structures in North America. tached to the cephalothorax by a narrow pedicel. The
r Keyto the Ordersof Arachnida 107

2 cephalothorax bears the eyes, mouthparts, legs, pedi-

Steel palps, and stomach. The abdomen contains the pri-
N mary reproductive structures, respiratory system, in-
E testine, anus, silk glands, and spinnerets.
z The cephalothorax is covered dorsally by the cara-
'"o Frame silk
pace and ventrally by the sternum. Anterior to the ster-
X num is a small sclerite called the labium, which may be
fused to the sternum. Spiders primitively have eight
simple eyes, but some taxa have fewer eyes. Eye num-
ber and arrangement is important in identifying spi-
ders. The area between the anterior eye and the edge of
0.1 0.2 0.3 the carapace is the clypeus.
The first pair of appendages is called the che-
Strain= A length licerae. Chelicerae have two segmems, a base and a
initiallength fango The opening for the poison gland is located near
the tip of the fango The base may be quite robusto The
base may receive the fang in a furrow, which is usually
Figure5-5 Structural properties of silk compared to lined with rows of teeth. In some families, the che-
othermaterials(FromGoslineet al. 1986). liceral bases may be fused by a membranous lamella

Figure 5-6 Structural characteristics of spiders. A, Dorsal view (generalized); B, Ven-

tral view, araneomorph; e, Ventral view, mygalomorph.
108 Chapter5 PhylumArthropoda




--- --- . -,::::-"--
~,- , -<~..'''. ~--,,:::. :::. -,--
- i\"""m_~"'I.\\\U\\"_'"
--':~ -CT
\ H

Figure5-7 Legs, chelicerae, and male palp characters of spiders. A, First leg of Mime-
tus puritanus Chamberlin (Mimetidae); B, Fourth metatarsus of Achaearanea tepidariorum
(c. L. Koch) (Theridiidae); C, Fourth tarsus and metatarsus of Callobius deces
(Chamberline and Ivie) (Amaurobiidae); D, Tarsus of Araneus diadematus Clerck
(Araneidae); E, Tarsus of Tibellus oblongus (Keyserling) (Philodromidae); F, Trilobe
membrane between metatarsus and tarsus of Heteropodavenatoria (L.) (Sparassidae);
G, Stridulatory striae on lateral face of chelicera, Mermessus dentiger; H, Chelicera of
Araneus diadematus showing condyle on proximolateral parto 1, Male palp of a haplogyne
spider, Latrodectus rec/usaGertsch and Mulaik (Sicariidae); J, Palp of an entelegyne male,
Mermessus dentiger O. Pickard-Cambridge (Linyphiidae). CA, calamistrum; CB, ventral
comb of setae; Cl, claws; CN, condyle; CT, claw tufts; CY, cymbium; MT, metatarsus;
P, paracymbium, SC, scopula; SS, stridulatory striae; TA, tarsus; TM, trilobe membrane,
TR, trichobothrium.
Keyto the Ordersof Arachnida 109

(Figure 5-9F,I). The basal segment of the chelicera just anterior to this furrow. She fertilizes her eggs just
sometimes bears a small rounded lateral prominence before exuding them through the epigastric furrow.
(¡hecondyle) at its base (Figure 5-7H, CN). Some spi- On the anterolateral portion of the epigastric fur-
dershave a series of filelike ridges on the lateral surface row are the anterior respiratory organs, the book lungs
of the chelicerae (Figure 5-7G, SS). Spiders may use a (rarely lost or modified in some families). The book
plectrum on the pedipalp to stridulate against these lungs consist of alternating layers of sheetlike air pock-
ridges. ets and hemolymph-filled (blood-filled) lamellae. The
Ihe second pair of appendages is the pedipalps. air- and hemolymph-filled regions are incredibly thin,
The pedipalps are somewhat leglike, except that they which facilitates gas exchange. Primitively, spiders had
lacka metatarsus and have only one terminal claw (lost a second pair of book lungs immediately posterior to
in some spiders). The basal segment, the endite, is en- the anterior pair. In most spiders, the posterior respira-
largedand forms part of the oral cavity. The anterior tory organs are modified into a tracheal system. The
marginof the endite usually has a serrula for cutting tracheal system opens to the environment via the tra-
into prey. The labium lies between the two endites. cheal spiracle, which may be a pair of ventrolateral slits
Adult male spiders have their pedipalps (or simply or a single ventral slit. The single tracheal spiracle usu-
palpus)modified into sperm transfer organs. The form ally lies just anterior to the spinnerets, but in so me spi-
of the male palpal organ is extremely variable and is der taxa is closer to the epigastric furrow.
criticalin spider taxonomy (Figures 5-7I,J). Minimally, The spinnerets are usually on the posterior part of
the palpal organ consists of a bulb containing a sperm the abdomen (Figure 5-8). Extant spiders have no
duct and a terminal embolus. The bulb is attached to more than six functional spinnerets: the anterior lat-
the palpal tarsus, which is called the cymbium (Fig- eral, posterior median, and posterior lateral pairs. The
ure 5-7J, CY). The cymbium may be ventrally exca- anterior median spinnerets are not functional in any
vated, partially enclosing the palpal bulbo A para- living spider. However, a platelike spinning organ
cymbium is an appendage arising from the cymbium called the cribellum (Figure 5-8A, CR) is believed to
(Figure5-7J, P). The testes are located in the abdomen. be derived from the anterior median spinnerets. The
Thereis no connection between the testes and the pedi- cribellum is lost in many araneomorph spiders, but a
palps. Instead, sperm is extruded from the epigastric vestige called the colulus may remain (Figure 5-8B-E).
furrow.A sperm web is typically spun to receive the Posterior to the spinnerets is the anal tubercle (Fig-
spenn. 10 transfer the sperm, the spider inserts his em- ure 5-6A,C). This structure is usually quite small but
bolus into the sperm droplet and draws the fluid into is enlarged and modified in oecobiids.
the palpal organ, probably by capillary action. During
copulation, the male transfers his sperm to the female Reproduction andDevelopment. The two sexes of spi-
byinserting the embolus into the female genitalia. ders often differ considerably in size, the female being
Ihe four pairs of ambulatory legs, numbered I-IV larger and heavier, and the male being smaller but with
from front to back, have seven segments (coxa, relatively long legs. Mating is often preceded by elabo-
trochanter, femur, patella, tibia, metatarsus, and tar- rate courtship, which may involve a variety of body or
sus), and each leg bears two or three terminal claws. pedipalp movements or stridulation. In some cases, the
Twoclaws are paired; the third, if present, is smaller male may present the female with a dead insect or
and set between them. The tip of the tarsus may have other nuptial gift before mating. Males usually do not
numerous setae, the claw tuft, which may obscure the live long after mating.
claws(Figure 5-7E, CT). Usually claw tufts are present Fertilized eggs are deposited in a silk saco These
only in two-clawed spiders, but there are exceptions sacs vary in their construction. They may be suspended
(noted later). In some spiders, the ventral part of the in a web, deposited in some sheltered location, or car-
tarsus(and sometimes the metatarsus) is covered with ried about by the female. The number of eggs in a sac
a verydense field of modified hairs, the scopula (Fig- varies between 1 and over 2,000. The eggs usually
ure S-7E, SC). This structure may allow spiders to hatch soon after they are laid, but if the eggs are laid in
walkon smooth vertical surfaces. Various leg segments the autumn the young spiders may remain in the sac
mayhave trichobothria (Figure 5-7C, TR). These long, until the following spring.
straighthairs are sensitive to air currents. The position Spiders undergo very little metamorphosis during
andnumber of trichobothria can be a useful taxonomic their development. When hatched, they usually look
character. like miniature adults but without developed genitalia.
Near the anterior end of the abdomen on the ven- If legs are lost during development, they can usually be
tralsideis a transverse groove called the epigastric fur- regenerated (Vollrath 1991). The point at which legs
row,which at its midpoint bears the gonopore. The fe- usually break off can be a useful taxonomic character.
malestores sperm received from males in spermathecae Spiders generally molt from 3 to 15 times during their

110 Chapter5 PhylumArthropoda

Figure 5-8 Abdominal characters of spiders. A, Spinnerets of Callobius deces

(Chamberlin and Ivie) (Amaurobiidae); B, Spinnerets of Agelenopsisoregonensis
Chamberline and Ivie (Agelenidae); C, Spinnerets of Cheiracanthium meldei L. Koch
(Miturgidae); D, Spinnerets of Clubiona obesa Hentz (Clubionidae); E, Spinnerets of Sco-
tophaeus blackwalli (Thorell) (Gnaphosidae). ALS, anterior lateral spinnerets; ca, colu-
lus; CR, cribellum; PLS, posterior lateral spinneret; PMS, posterior median spinneret.

growth to maturity; males typically undergo fewer other spiders and stealing prey from the host's web. A
molts than females. Female mygalomorphs and filista- small number of spider species have some degree of so-
tids continue to molt once or twice a year as long as cial organization. Social spiders may cooperate to build
they live. Most spiders live 1 or 2 years, but mygalo- webs of several cubic meters and feed communally on
morphs often take several years to mature, and some the large prey items they subdue.
females may live as long as 20 years. Spiders play an important role in nearly all terres-
trial ecosysterns. They are generally quite numerous,
Ecology. AIIspiders are predaceous and feed mainly on and their predation impact keeps in check the numbers
insects. Some spiders may occasionally feed on small of many other animals, particularly insects. They are, in
vertebrates. The prey is usually killed or disabled by the tum, preyed on by various other animals, particularly
poison injected into it by the spider bite. Different spi- wasps. There is growing evidence that spiders can keep
ders capture their prey in different ways. The wolf spi- populations of pest insects in check in agroecosysterns
ders and jumping spiders actively forage and pounce on (Wise 1993, Riechert 1999). Conventional integrated
their prey. Many crab spiders are sit-and-wait predators, pest management prograrns tend to rely on predators
waiting for their prey on flowers and feeding on visiting and parasitoids that attack specific pest species, but
pollinators. Many spiders capture their prey in webs. A dense populations of generalist predators, including spi-
few spiders are kleptoparasites, living in the webs of ders, can effectively limit some pest populations. How-
Classificationof Spiders 111

ever,broad-spectrum insecticides can be more effective out from the cribellum using tq.e calamistrum, a group
in killing spiders than they are in killing the pests being of specialized, curved setae on the metatarsus of the
targeted.Agricultural practices that encourage increases founh leg (Figure 5-7C, CA), which may be a single
inspider density can greatly decrease the need for chem- row of setae, two rows, or an oblong field. Mature male
ical insecticides without 1055 in crop yield (Riechert cribellate spiders lose the ability to make cribellate silk
1999). Fear of spiders (arachnophobia) is one of the and typically retain only a vestige of the cribellum and
rnostcommon phobias is Western culture. However, spi- calamistrum. The cribellum and calamistrum have
dersperform a service by destroying noxious insects and be en lost numerous times in spider evolution. A well-
do not damage household items. As pointed out earlier, known example of ecribellate sticky silk comes from
the venomous nature of spiders is generally greatly ex- araneoid spiders, the ecribellate orb-weavers and their
aggerated.Most spiders do not bite if handled carefully. descendents. Araneoid sticky silk consists of a pair of
core lines dotted with sticky droplets. Adult male spi-
ders rarely spin prey-capture webs and sometimes lose
the ability to produce sticky silk.
of Spiders Spider taxonomy is based heavily on characters of
the genitalia. For this reason, juvenile spiders are usually
Spidersare divided into two suborders: Mesothelae and impossible to identify to species unless they are found
Opisthothelae. The Mesothelae are currently restricted associated with adults or unless the local fauna is very
to southeastern Asia andJapan. These relict taxa retain well known. Determining whether a spider is an adult
rnany features of primitive spiders, such as abdominal can be difficult for the beginner. Adult female spiders
segmentation and ventral spinnerets. The vast majority usually have a sclerotized plate called an epigynum (Fig-
of spiders belongs to the suborder Opisthothelae, ure 5-6B) located near the epigastric furrow between
which itself is divided into two infraorders, Mygalo- the anterior respiratory organs (the genital region). In
rnorphae and Araneomorphae. Both occur in North some spiders (for example, lycosids), a rudimentary,
America. A few basal mygalomorphs have vestiges of nonfunctional plate may be visible during late-stage ju-
abdominal segmentation in the form of abbreviated ter- venile development. Female spiders sometimes lack an
gites. A sclerotized scutum covering pan of the ab- external sclerotized plate and can be easily confused
domen is found sporadically among araneomorph spi- with juveniles. In such spiders, adult females differ by
der taxa and is not derived from primitive abdominal the presence of dense hairs in the genital region; hairs in
segmentation. Mesothele and mygalomorph spiders the genital region are undifferentiated in juveniles. All
have chelicerae oriented so that the fangs are more or adult female spiders have a set of spermathacae, which
lessparallel (Figure 5-6C); araneomorph spiders usu- can be examined by carefully dissecting the integument
allyhave opposing fangs (Figure 5-6B) , although some anterior to the epigastric furrow. Few student collections
taxahave secondarily modified chelicerae. Mesotheles, will contain many species where the female lacks and
rnygalomorphs, and the most basal groups of araneo- epigynum. Adult male spiders always have the distal
rnorph spiders have two pairs of book lungs. In North segments of the pedipalp modified. Late-stage juvenile
America, only hypochilids among the araneomorphs males may have a swollen palpal tarsus, indicating that
retain the posterior pair of book lungs. In most arane- the palpal organ is developing inside.
omorph spiders, the posterior pair of book lungs is
rnodified into a tracheal system (Figure 5-6B). Sec- Infraorder Mygalomorphae
ondary modifications of the respiratory system occur in Antrodiaetidae-folding-door spiders
some derived groups of araneomorph spiders. Atypidae-purse-web spiders
Individual species can produce up to seven dis-
tinct types of silk, each with a specialized function, Dipluridae
such as dragline production, egg sac construction, and
both the adhesive and nonadhesive parts of a web. Cyrtaucheniidae
Basalaraneomorph spiders produce adhesive silk from
a platelike cribellum just anterior to the spinnerets Theraphosidae-tarantulas
(Figure 5-8A, CR). The cribellum is covered in tiny Infraorder Araneomorphae
spigots. Cribellate silk consists of hundreds of very Hypochilidae-Iampshade spiders
fine, dry silk fibers around a few thicker core fibers. Haplogynae
The physical basis of stickiness in cribellate silk is not Filistatidae
wellunderstood, but the adhesive force is proponional Caponiidae
to the surface area contact between the silk and the ob- Dysderidae
ject being held (Opell 1994). Cribellate silk is combed Segestriidae
112 Chapter5 PhylumArthropoda

Oonopidae Hahniidae
Pholcidae-cellar spiders Agelenidae
Diguetidae Cybaeidae
Plectreuridae Desidae
Ochyroceratidae Amphinectidae
Leptonetidae Amaurobiidae
Telemidae Tengellidae
Sicariidae Zorocratidae
Scytodidae-spitting spiders Superfamily Lycosoidea
Entelegynae Ctenidae-wandering spiders
Eresoidea Zoropsidae
Oecobiidae Miturgidae
Hersiliidae Oxyopidae-Iynx spiders
Orbiculariae Pisauridae-nursery-web and fishing
Uloboridae-hackled-band orb-weavers spiders
Deinopidae-ogre-faced spiders Trechaleidae
Araneidae Lycosidae-wolf spiders
Tetragnathidae Clade Dionycha
Theridiosomatidae Clubionidae
Symphytognathidae Anyphaenidae
Anapidae Corinnidae
Mysmenidae Liocranidae
Mimetidae-pirate spiders Zoridae
Theridiidae-cobweb spiders Gnaphosidae
Nesticidae Prodidomidae
Linyphiidae Homalonychidae
Pimoidae Selenopidae
Titanoecidae Sparassidae-giant crab spiders
RTA Clade Philodromidae
Dictynidae Thomisidae
Zodariidae Salticidae-jumping spiders

Keyto SpiderFamiliesof NorthAmerica

This key is adapted from D. Ubick, in preparation, Key to Nearctic Spider Families. In D. Ubick, P. Paquin, P. E.
Cushing, and V. Roth (Eds.), Spiders of North America: A Guide to Genera (available in ]anuary 2005 from the
American Arachnological Society). The following abbreviations are used in the key. Eyes: AER = anterior eye row;
PER = posterior eye row; AME = anterior median eyes; ALE = anterior lateral eyes; PME = posterior median eyes;
PLE = posterior lateral eyes; LE = lateral eyes. Spinnerets: AMS = anterior median spinnerets; ALS = anterior lat-
eral spinnerets; PMS = posterior median spinnerets; PLS = posterior lateral spinnerets. Measurements: L = length;
W = width; PT/C is the ratio of the patella+tibia length divided by carapace length. The term procurved refers to
a line in which the ends are anterior to the center of the line; a recurved line has the ends posterior to the middle.

1. Chelicerae paraxial, fangs parallel (Figure S-6C); with 2 pairs of book

lungs (Figure S-6C); never with cribellum or colulus; with 8 eyes; legs
stout (PT/C < 2) (Mygalomorphae) 2
1'. Chelicerae diaxial, fangs opposing each other or oblique (Figure S-6B);
usually with at most I pair of book lungs (Figure S-6B), if with 2 pairs of
book lungs then with cribellum (Figure S-8A) and slender legs
(PT/C = 3 to S); cribellum or colulus may be present; with 8 or fewer
eyes; leg thickness variable (Araneomorphae) 11
2(1). Abdomen with 1-3 tergites; anal tubercle separated from spinnerets 3

to SpiderFamiliesof NorthAmerica 113

2/. Abdomen without tergites; anal tubercle adjaeent to spinnerets 5

3(2). Endites long, % width of sternum; labium and sternum fused; thorade
furrow (Figure 5-6A) quadrangular or suboval; eastern United States Atypidae p.120
3'. Endites short, at most 1(2width of sternum; labium and sternum
separated by a suture, groove, or at least a depression; thorade furrow
longitudinal or rounded and pidike; widespread 4
4(3'). Distal segment of PLS slender, at least 5 X as long as basal width,
tapering to a point, flexible, pseudosegmented; Washington lO
California, Arizona Mecicobothriidae p.120
4/. Distal segment of PLS slOuter, about 3X as long as wide, neither flexible
nor pseudosegmented; widespread in distribution Antrodiaetidae p.120
5(2/). Tarsi with 2 claws and claw tufts (Figure 5-7E). Florida to southwestern
United States Theraphosidae p.122
5/. Tarsi with 3 claws, laeking claw tufts (Figure 5-7D). widespread in
distribution 6
6(5/). PLS long, at least 1(2earapaee length, and slender, length of distal
segment > 2X width 7
6/. PLS short, at most 1/2earapaee length, and stout, length of distal
segment < 2X width 8
7(6). Endites with euspules (Figure 5-6C); ehelieeral inner surfaee with row
of 4-7 short, blaek, rodlike setae; thorade furrow transverse; PLS 1(2to
%X earapaee length; size 16-23 mm; living in open burrows;
California Nemesiidae(Calisoga) p.122
7/. Endites laeking euspules; ehelieeral inner surfaee without row of blaek,
rodlike setae; thorade furrow pitlike or longitudinal; PLS at least
V.X earapaee length; size 2.5-17 mm; living in silken tubes or on
sheet webs; British Columbia to Oregon, Arizona to Texas,
North Carolina, Tennessee Dipluridae p.120
8(6'). Abdomen posteriorly truneated, sclerotized, with longitudinal grooves; Ctenizidae
southeastern United States (Cyclocosmia) p. 120
8'. Abdomen unmodified, posteriorly rounded, not sclerotized, and
without grooves 9
9(8/). Females with a dense seopula on tarsus 1 (Figure 5-7E); ehelieeral
promargin toothed, retromargin of ehelieera with row of short, stout, Cyrtaucheniidae
rounded tubercles; Texas (Eucteniza) p.122
9'. Males or females with tarsus llaeking seopula, but with lateral spines;
if tarsus 1with a weak seopula and laeking spines, then retromargin of
chelicera without teeth (but may have a few denticles); widespread in
distribution 10
10(9/). Both promargin and retromargin of ehelieera toothed; thorade furrow
strongly proeurved; anterior tarsi and metatarsi of female with lateral
rows of short, thornlike spines Ctenizidae(inpart) p.120
10'. Only promargin of ehelieera toothed; thoraeic furrow varying from
strongly proeurved to straight; anterior tarsi and metatarsi of female Cyrtaucheniidae
with few spines, these usually long and slender (in part) p.122
ll(l'). Cribellum (Figure 5-8B-E) and ealamistrum absent 12
ll'. Cribellum (Figure 5-8A) and ealamistrum (Figure 5-7C) present 14
12(1l). With 8 eyes 13
12/. With fewer than 8 eyes 27
114 Chapter5 PhylumArthropoda

13(12). Tarsi 3-clawed, scopulae and claw tufts absent (Figure S-7D); mostly
web builders; if legs are slender and relatively delicate, assume tarsi are
3-clawed 51
13'. Tarsi 2-clawed, usually with scopulae and claw tufts (Figure S-7E);
if claw tufts are present, assume tarsi are 2-clawed 81
14(11'). With 2 pairs of book lungs (Figure S-6C) Hypochilidae
(Hypochilus) p.122
14'. With 1 pair of book lungs (Figure S-6B) 15
15(14'). With fewer than 8 eyes; cribellum entire 16
15'. With 8 eyes; cribellum entire or divided 17
16(15). With 4 eyes; leg I greatly enlarged; south Texas Uloboridae
(Miagrammopes) p.124
16'. With 6 eyes; leg I not enlarged; widespread in distribution Dictynidae(Lathys) p.126
17(15'). PME huge, several times the diameter of remaining eyes (Figure S-9G) Deinopidae(Deinopis) p.124
17'. PME not so enlarged 18
18(17'). Eyes clustered on central mound, occupying < 1(2width of cephalon 19
18'. Eyes spread across carapace, occupying > 1/2width of cephalon 20
19(18). Anal tubercle enlarged and fringed with long setae; chelicerae free;
entelegyne (epigynum present) Oecobiidae(Oecobius) p.124
19'. Anal tubercle not so modified; chelicerae fused; haplogyne (epigynum
absent) Filistatidae p.122
20(18'). Anterior tibia with at least 4 pairs of ventral spines 21
20'. Anterior tibia with fewer ventral spines 22
21(20). Anterior tibia with 4-5 pairs of ventral spines; PER straight to
weakly procurved; tarsi U-IV with 2 claws; body unicolorous; Zorocratidae
Arizona to Texas (Zorocrates) p.127
21'. Anterior tibia with 6-7 pairs of ventral spines; PER recurved; all tarsi
with 2 claws; body patterned; California (introduced) Zoropsidae(Zoropsis) p.127
22(20'). Calamistrum extends over more than half the length of metatarsus IV 23
22'. Calamistrum extends over no more than half the length of metatarsus IV 25
23(22). Femora U-IV with rows of long trichobothria; metatarsus IV dorsally
concave and with ventral row of short spines extending to tip of tarsus;
cribellum entire Uloboridae(in part) p.124
23'. Femora U-IV without rows of long trichobothria; metatarsus IV not so
modified and without ventral row of short spines; cribellum divided or
entire 24
24(23'). Endites converging apically; cribellum usually entire; legs usually without
spines Dictynidae(in part) p.126
24'. Endites parallel; cribellum divided; legs with spines Titanoecidae
(Titanoeca) p.126
25(22'). Cheliceral margins with 5 to 7 stout teeth; male palpus with embolus
threadlike, enclosed in membranous conductor; California, Texas Amphinectidae
to Florida (Metaltella) p.126
25'. Cheliceralmarginsusuallywith no more than 4 teeth, if more, then
teeth small and slender; male palpus with embolus variable; widespread
in distribution 26
26(25'). AME 1.4X largerthan ALE;malepalpuswith emboluslong and
sigmoid, enclosed in membranous conductor Desidae(Badumna) p.126
Keyto SpiderFamiliesof NorthAmerica 115

26'. AME at most 1.2X larger than ALE; male palpus with embolus usually
short and stout, iflong, then arcuate, never enc10sed in conductor Amaurobiidae(in part) p.126
27(12'). Eyes complete1y lacking (but may have small eye spots) 28
27'. At least 2 eyes present 34
28(27). Anterior tibia with 2-3 pairs of ventral spines 29
28'. Anterior tibia with few scattered ventral spines or none 30
29(28). ALScontiguous, longer than PLS Cybaeidae
(in part) p.126
29'. ALSslightly separated, shorter than PLS Dictynidae(in part) p.126
30(28'). Male palp with exposed bulb (Figure 5-71), female lacking epigynum
(haplogynes) 31
30'. Male palp with bulb enc10sed by cymbium (Figure 5-7J), female with
epigynum (ente1egynes) 32
31(30). Abdomen with anterodorsal sc1erotized ridge; with a pair of tracheal
spirac1es between epigastric furrow and spinnerets; colulus pentagonal,
wider than ALS;California Telemidae
(Usofila) p.123
31'. Abdomen without anterodorsal sc1erotized ridge; with one tracheal
spirac1e near spinnerets; colulus not so modified; Texas, Georgia leptonetidae(in part) p.123
32(30'). Tarsus IV lacking ventral comb; retromargin of che1icera toothed;
chelicera usually with stridulatory file on outer face (Figure 5-7G);
leg break at patella-tibia joint Linyphiidae(in part) p.126
32'. Tarsus IV with ventral comb of serrated setae (Figure 5-7B); retromargin
of chelicera edentate or with small dentic1es; che1icera without
stridulatory file; leg break at coxa-femur joint 33
33(32'). Retromargin of chelicera edentate, lacking both teeth and dentic1es; Theridiidae
male palp with inconscpicuous paracymbium; southern Arizona (Thymoites) p.125
33'. Retromargin of chelicera with small dentic1es; male palp with large
retrolateral paracymbium; California, Texas, Appalachian Mountains Nesticidae(in part) p.126
34(27'). Two eyes present Caponiidae(in part) p.122
34'. More than 2 eyes present 35
35(34'). Four eyes present 36
35'. Six eyes present 37
36(35). Two eyes pigmented; size 0.6 mm; litter spiders; Florida Symphytognathidae
(Anapistula) p.125
36'. All eyes unpigmented; size 3.6 mm; cave spiders; Tennessee Nesticidae(Nesticus) p.126
37(35'). Male palp with exposed bulb, not enc10sed by cymbium (Figure 5-71);
female lacking epigynum, but may have some sc1erotization at epigastric
area (haplogynes) 38
37'. Male palp with bulb partially enc10sed by cymbium (Figure 5-7J) ,
female with epigynum (ente1egynes) 48
38(37). Chelicerae fused at base (Figure 5-9F) 39
38'. Che1iceraenot fused at base, can be moved apart (Figure 5-9E) 42
39(38). Eyes in 2 triads Pholcidae(in part) p.123
39'. Eyes in 3 dyads (Figure 5-9G) 40
40(39'). Carapace strongly convex (Scytodes) p.123
40'. Carapace fiat 41
41(40'). PER strongly recurved; carapace pear-shaped (Figure 5-91) Sicariidae(Loxosceles)p.123
41'. PER slightly recurved; carapace oval Diguetidae(Oiguetia) p.123
r ..

116 Chapter5 PhylumArthropoda

42(38'), Size >5 mm; tracheal spiracles paired, conspicuous, located near book
lung openings (Figure 5-9H) 43
42', Size < 5 mm; tracheal spiracles inconspicuous, if paired, then not near
book lungs 44
43(42), Tarsi with 2 claws; leg 1IInot anteriorly directed Dysderidae(Dysdera) p.122
43'. Tarsi with 3 claws; leg 1IIanteriorly directed Segestriidae p.123
44(42'). Abdomen with sclerotized ridge on anterior dorsal surface Telemidae(Usofí/a) p.123
44'. Abdomen without sclerotized ridge 45
45(44'). Abdomen with 1 or 2 scuta Oonopidae(in part) p.123
45'. Abdomen without scuta 46
46(45'). Legs relatively long (PT/C > 1.5); PME usually posteriorly displaced
from LE, if eyes contiguous then occupying less than '/2 cephalon width Leptonetidae
(in part) p.123
46'. Legs shorter (PT/C "" 1); eyes in transverse arrangement, if contiguous
then occupying more than 1/2cephalon width 47
47(46'). Eyes in compact group; if in transverse row, then femur IV enlarged;
colulus absent; widespread in distribution Oonopidae(in part) p.123
47'. Eyes in transverse row; femur IV not enlarged; colulus large, broad; Ochyroceratidae p.123
southem Florida (Theotíma)
48(37'). Tibia I with 2 to 3 pairs of ventral spines 49
48'. Tibia I with fewer ventral spines 50
49(48). ALS contiguous, longer than PLS; eyes small, widely separated Cybaeidae p. 126
49', ALSslightly separated, shorter than PLS; eyes larger, in two triads of
contiguous eyes Dictynidae(in part) p.126
50(48'). Tarsus IV with ventral comb of serrated bristles (Figure 5-7B); male
palpus with large, rigid basal paracymbium Nesticidae(in part) p.126
. 50'. Tarsus IV without ventral comb; male palpus with smaller basal
paracymbium flexibly attached by membrane (Figure 5-7]) Linyphiidae(in part) p.126
51(13). Male palpus with bulb exposed, not enveloped by cymbium
(Figure 5-71); female without epigynum (haplogynes) 52
51'. Male palpus with bulb partially enveloped by cymbium (Figure 5-7]);
female with epigynum (entelegynes) 55
52(51). Eyes contiguous with AME surrounded by the others Caponiidae(Ca/ponía) p.122
52'. Eyes not so arranged 53
53(52'). Eyes in 3 groups, with AME forming a dyad and the others 2 triads
(Figure 5-9F); chelicerae fused at base (Figure 5-9F) Pholcidae(in part) p.123
53'. Eyes in 2 transverse rows; chelicerae fused or not 54
54(53'). Chelicerae fused at base; endites converging apically Plectreuridae p.123
54'. Chelicerae not fused; endites parallel (in part) p.125
55(51'). Chelicerae fused at base (Figure 5-9F); eyes in 3 groups with AME
forming a dyad and the others 2 triads (Figure 5-9F); legs long and
slender, PT/C > 1.6, with flexible tarsi (Note: Although pholcids are
haplogynes, some have complex genitalia and may be mistaken for
entelegynes) Pholcidae(in part) p.123
55'. Chelicerae not fused; eyes not so arranged; legs usually shorter with
rigid tarsi 56
56(55'). Tarsi with at most a single trichobothrium 57
56'. Tarsi with 2 or more trichobothria (Figure 5-7C) 67
Keyto SpiderFamiliesof NorthAmerica 117

57(56). PLS with long apical segment, about as long as abdomen; eyes clustered
on a mound at center of cephalon; Texas, Florida Hersiliidae(Tama) p.124
57/, PLS shorter; eye arrangement different; widespread in distribution 58
58(57). Anterior tibiae and metatarsi with prolateral row of curved spines in
serrated series (Figure 5-7 A) Mimetidae p.125
58/. Anterior legs without such spines 59
59(58'). Tarsus IV with ventral comb of serrated bristles (sometimes not distinct,
lacking in Argyrodes) (Figure 5-7B); legs usually lacking spines;
chelicerae usually with basal extension; epigynum without scape 60
59/, Tarsus IV without ventral comb; legs with spines; chelicerae lacking
basal extension; epigynum sometimes with scape 61
60(59). Labium with anterior margin thickened; endites parallel; male palpus
with large basal paracymbium Nesticidae(in part) p.126
60', Labium with anterior margin unmodified; endites converging apically;
male palpus with paracymbium inconspicuous or represented by an
apical or ectal (outer) notch Theridiidae(in part) p.125
61(59/). Tarsi as long as or longer than metatarsi; tiny spiders, < 2 mm 62
61'. Tarsi shorter than metatarsi; size usually > 2 mm 63
62(61). Pedicel originating from opening on posterior slope of carapace;
male abdomen with scutum; male metatarsus 1without clasping spur; Anapidae
female without palpi (Gertschanapis) p.125
62'. Pedicel origin below edge of carapace; male abdomen without scutum;
male metatarsus 1 usually with clasping spur; female with palpi Mysmenidae p.125
63(61'). Sternum with a pair of pits at labial margin; tibia IV with long Theridiosomatidae
trichobothria; legs stout; size < 2.5 mm ( Theridiosoma) p.125
63'. Sternum without such pits; tibia IV without long trichobothria;
legs variable; size usually larger 64
64(63'). Clypeus higher than 4 diameters of AME; chelicerae usually with
stridulatory file on lateral surface, lacking condyle (Figure 5-7G);
legs weakly spined; tarsi cylindrical; leg break at patella-tibia joint;
sheet web builders 65
64/. Clypeus lower than 3 diameters of AME; chelicerae without stridulatory
file, usually with condyle (Figure 5-7H); legs with strong spines;
tarsi tapering distally; no leg break at patella-tibia joint; orb web builders 66
65(64). Male cymbium with integral retrolateral paracymbium; cymbium usuaUy
with a retromedian process armed with spines or cuspules; female
epigynum forming stout, tonguelike projection with apical openings;
size > 5 mm Pimoidae(Pimoa) p.126
65/. Male cymbium with retrolateral paracymbium attached by membrane
(Figure 5-7)); cymbium usually lacking retromedian process, if present
lacks spines or cuspules; female epigynum variable; size usually < 5 mm Linyphiidae(in part) p.126
66(64'). Endites square or rectangular; epigynum usually 3-dimensional,
usually with scape, a central fingerlike projection, fIat in Hypsosinga,
Mastophora, and Zygiella; palpal tibia cup-shaped with irregular distal
margin (except in Zygidla); builders of vertical orb webs Araneidae p.125
66'. Endites elongate, widest at distal edge; epigynum absent or a fIat plate
with at most a swelling (Meta) or pointed (Metleucauge);palpal tibia Tetragnathidae
conical; builders of horizontal orb webs (except Nephila) (in part) p.125

118 Chapter5 PhylumArthropoda

67(56'). Tarsal and metatarsal trichobothria in dorsal row, increasing in length

distally (Figure 5-7C); at least anterior trochanters not notched 68
67'. Tarsal and metatarsal trichobothria in 2 irregular rows; all trochanters
with shallow to deep notches 78
68(67). ALS enlarged, others reduced; clypeus high; endites strongly converging,
lacking serrula Zodariidae p.126
68'. ALSat most only slightly larger than the others; endites not strongly
converging; serrula present 69
69(68'). Colulus large and broad, occupying at least half the width of the
spinning area 70
69'. Colulus width less than 1/2of the spinning area 71
70(69). Distribution: southern Florida Desidae(Paratheuma) p.126
70'. Distribution: southern California Dictynidae(Saltonia) p.126
71(69'). Spinnerets arranged in transverse row; tracheal spiracle positioned
well anterior of spinnerets Hahniidae(in part) p.126
71'. Spinneret arrangement not so modified; tracheal spiracle near spinnerets 72
72(71'). Legs with plumose hairs; both eye rows strongly procurved, straight in
Tegenaria,which has distinctive sternum markings Agelenidae p.126
72'. Legs lacking plumose hairs; eye rows straight; sternum typically 1 color 73
73(72'). ALScontiguous or nearly so, thicker and usually longer than PLS Cybaeidae
(in part) p.126
73'. ALSdistinctly separated, shorter than PLS 74
74(73'). Anterior tibia with 4 or more pairs of ventral spines Hahniidae(in part) p.126
74'. Anterior tibia with 3 or fewer pairs of ventral spines 75
75(74'). Retromargin of chelicera with 2-5 equal-sized teeth, no denticles 76
75'. Retromargin of chelicera with both teeth and denticles or with more
than 5 teeth 77
76(75). Size > 5 mm Amaurobiidae(in part) p.126
76'. Size < 5 mm Hahniidae(in part) p.126
77(75'). Legs long and slender, PT/C > 1.25; male palp with apically produced Hahniidae
cymbium (Calymmaria) p.126
77'. Legs shorter and stouter, PT/C < 1; male palp with unmodified cymbium Dictynidae(in part) p.126
78(67'). Tarsi long and flexible Trechaleidae
(Trechalea) p.127
78'. Tarsi not flexible 79
79(78'). Clypeus high; AME small, others larger, forming hexagon (PER
procurved) (Fígure 5-9A); cheliceral margins with at most a single tooth;
trochanters with shallow notches Oxyopidae p.127
79'. Clypeus low; PER recurved; chelicerae strongly toothed; trochanters
deeply notched 80
80(79'). PER strongly recurved with PLE posterior to PME so that eyes appear
as 3 rows (Fígure 5-9C); male palp lacking retrolateral tibial apophysís Lycosidae p.127
80'. PER not as strongly recurved, PLE lateral to PME (Figure 5-9D); male
palp with retrolateral tibial apophysis Pisauridae p.127
81(13'). Legs laterigrade: extending laterally from the body and twisted so that
the morphologically pro lateral surface of the anterior legs ís functionally
dorsal (Figure5-11 C) 82
81'. Legs prograde: anterior pair directed forward, posterior pair backward
(Figure 5-lID) 85

Keyto SpiderFamiliesof NorthAmerica 119

82(81). Anterior legs thicker and longer than posterior ones (Figure 5-11C);
tarsi lacking scopulae; chelicerae lacking teeth (except Isaloides) Thomisidae p.128
82'. Anterior legs not so enlarged, although leg II may be significantly longer
than the others; tarsi with scopulae; chelicerae with teeth or denticles 83
83(82'). Smaller spiders (size < 10 mm); cheliceral retromargin lacking teeth (in part) p.128
83'. Larger spiders (size usually > 10 mm); cheliceral retromargin with teeth
or denticles 84
84(83'). Metatarsi with dorsoapical trilobed membrane permitting hyperextension
of tarsi (Figure 5-7F); tarsi with thick claw tufts; cheliceral retromargin
with teeth; trochanters with deep notches, if not notched, then cheliceral
retromargin with denticles, not teeth (Pseudosparianthis);southwestern
United States and Florida Sparasiidae
(in part) p.128
84'. Metatarsi lacking dorsoapical trilobed membrane; tarsi lacking claw
tufts, but tarsal scopulae apically produced and may give the impression
of tufts; cheliceral retromargin with teeth; trochanters with shallow
notches; Arizona, New Mexico (Lauricius) p.126
85(81'). AME greatly enlarged, PER on lateral edge of carapace; carapace
anteriorlytruncate,facenearlyvertical(Figure 5-9B) Salticidae p.128
85'. Eyes and carapace not so modified 86
86(85'). PER strongly recurved, eyes appear as 3 rows with anterior edge of PLE
at or behind the posterior edge of PME 87
86'. PER variable, but not strongly recurved 90
87(86). Anterior tibia with at least 5 pairs of strong ventral spines 88
87'. Anterior tibia with fewer ventral spines 89
88(87). AER strongly recurved with ALE small and contiguous with PME and
PLE; size > 6 mm Ctenidae p.128
88'. AER straight to slightly recurved; size < 6 mm Zoridae(Zara) p.128
89(87'). Carapace narrow, longer than wide; grass spiders; widespread in Philodromidae
distribution (Tibellus) p.128
89'. Carapace broad, as long as wide; ground spiders; southern California to Homalonychidae
Arizona (Homalonychus) p.128
90(86'). Tracheal spiracle at middle of abdomen; claw tufts of broad,
lamelliform setae Anyphaenidae p.128
90'. Tracheal spiracle near spinnerets; claw tufts of normal setae 91
91(90'). ALScylindrical, noticeably separated (may appear contiguous in Micaria
and some Orodrassus) (Figure 5-8B); PME usually modified, either
elliptical, oval or triangular; cheliceral margins usually weakly toothed,
with only denticles, or lacking armature, sometimes keeled or with
only 2-3 teeth; endites usually with distinct oblique median depression
(not conspicuous in Callilepis); claw tufts always present 92
91', ALSconical, contiguous at base (Figure 5-8A, C); PME usually round,
if oval then anterior tibiae with 5-6 pairs of ventral spines; cheliceral
margins strongly toothed; endites nearly parallel, often widened distally,
usually lacking transverse median depression (if so, then anterior tibiae
with 5-6 pairs of ventral spines); claw tufts present or absent 93
92(91). PER straight or recurved, rarely procurved (Scopoides);ALSwith
few moderately elongated spigots; tarsal claws and at least cheliceral
promargin toothed Gnaphosidae
(in part) p.128
92'. PERstronglyprocurved;ALSwith numerousconspicuouslyelongated
spigots; tarsal claws and cheliceral margins lacking teeth (in part) p.128
120 Chapter5 PhylumArthropoda

93(91'). PLSdistinctly 2-segmented, distal segment conical (Figure 5-8A) Miturgidae p.127
93'. PLS l-segmented or, if 2-segmented, distal segment rounded 94

94(93'). Legs lacking spines, but anterior tibiae with ventral cusps Corinnidae (Trachelinae) p. 128
94'. Legs with spines 95
95(94'). Anterior tibiae with more than 4 pairs of ventral spines 96
95'. Anterior tibiae with fewer than 4 pairs of ventral spines 98
96(95). Margin of stemum with triangular processes pointing toward ,:oxae Corinnidae(Corinninae,
(precoxal triangles); trochanters with at most shallow notches Phrurolithinae) p. 128
96'. Precoxal triangles absent; trochanters deeply notched 97
97(96'). Size > 5 mm; tarsi not pseudosegmented Tengellidae(in part) p. 126
97'. Size < 4 mm; tarsi pseudosegmented, posterior ones bent Liocranidae(Apostenus)p. 128
98(95'). Abdomen of adults with scutum, covering entire abdomen of male and Corinnidae
only anterior portion of female (Castianeirinae) p. 128
98'. Abdomen of adult without scutum 99
99(98'). Endites concave ectally; precoxal triangles present (see couplet 96) Clubionidae p. 127
99'. Endites straight or convex ectally; precoxal triangles usually absent,
present in Hesperocranum,which has a distinctive dense brush of paired
ventral bristles on anterior tibiae and metatarsi Liocranidae (in part) p. 128

Infraorder Mygalomorphae
debris. If an insect comes in contact with the tube, the
These are the tarantulas and their allies. These spiders spider may bite through, grab the insect, and pull it
have large and powerful chelicerae with the fangs through the tube. Eight species occur in the eastern
parallel to each other and two pairs of book lungs (Fig- United States, extending as far north as Wisconsin and
ure 5-6C). Females are long-lived and molt after sex- New England.
ual maturity. Most species are large (5-34 mm), heavy- Family Mecicobothriidae:Members of this group
bodied, and stout-Iegged. Many species are burrowers; have one or two dorsal abdominal tergites; long, flexi-
some dose their burrows with a trapdoor or silk collar. ble, posterior spinnerets; a longitudinal thoracic fur-
This group is largely tropical, but 138 species occur in row; and separation between the sternum and labium.
North America. Most of these occur in the South and They build webs consisting of sheets and tubes under
Southwest, but a few occur as far north as Massachu- debris. They are small to medium-sized mygalomorphs
setts and southeastern Alaska. (4-18 mm) with six species known from western
Family Antrodiaetidae-Folding-Door Spiders: North America.
Members of this group have one to three dorsal ab- Family Dipluridae: Members of this group lack ab-
dominal tergites, some separation between the labium dominal tergites and have very long posterior spin-
and the sternum, and a longitudinal or pitlike thoracic nerets. They may build a sheet web with a funnel, or a
furrow. The burrows of these spiders are dosed with a silk tube. This family indudes the Appalachian species
pair of flexible, silken doors (Antrodiaetus) or with a MicrohexuramontivagaCrosby and Bishop, which is on
single thin door (Aliatypus), or remain open on a tur- the endangered species list because of habitat destruc-
ret (Atypoides). They are medium-sized mygalomorphs tion associated with the invasive balsam woolly adelgid
(6-16 mm) with 25 species widely distributed in North (Hemiptera: Adelgidae). At 4 mm, Microhexura monti-
America. vaga is the smallest mygalomorph in North America.
Family Atypidae-Purse-Web Spiders: These mod- Five species are known from North America.
erately large spiders (8-30 mm) have one dorsal ab- Family Ctenizidae:Spiders in the families Ctenizidae
dominal tergite, sometimes enlarged into a scutum in and Cytraucheniidae are trapdoor spiders, construct-
males. The sternum and labium are fused, and the tho- ing tunnels in the ground that are dosed by a door
racic furrow is transverse. These spiders build silken hinged with silk. The door fits snugly and is usually
~'.tI'! . ¡ubesthat may líe hO~W1tá\ly on the ground or extend well camouflaged. The tunnels may be simple or
ft{" . up the base of a tree. The tUbesare camouflagedwith branched, and may even contain side chambers that are
Keyto SpiderFamiliesof NorthAmerica 121

Figure5-9 A-G, 1,Eyearrangementsof spiders;H, Ventralview.A, Peucetiaviridans

(Hentz) (Oxyopidae); B, Salticus scenicus (Clerck) (Salticidae); C, Rabidosa rabida
(Walckenaer) (Lycosidae); D, Pisaurina sp. (Pisauridae); E, Ozyptila pacifica Banks
(Thomisidae); F, Pho/cusphalangioides (Fuesslin) (Pholcidae); G, Deinopis:&inosaMarx
(Deinopi~ae); ,H, ?ysdera crocataC. L. ~och (Dysderidae); 1, Loxosce/esrevNJVfRSIDAD DE CA' ...".
and Mulalk (SlCarudae). TR, tracheal splracle.
B IB ,l .r . ,-""",A
, 122 Chapter5 PhylumArthropoda

closed off from the main tunnel by hinged doors. These fication of the posterior pair of book lungs into a sys-
spiders can often be observed at night positioned at the tem of tracheae.
entrance of a slightly opened door. When they detect Family Hypochilidae-LampshadeSpiders: These
prey passing close by, they rush out, capture the prey, cribellate, long-legged spiders are the only araneo-
and take it down into the tunnel. Ctenizids are distin- morphs that retain the second pair of book lungs. The
gUished by rows of short, stout spines on the first and cribellum is undivided. North American species sit at
second tarsi and metatarsi, and by the presence of a the center of a circular mesh web, usually built on rock
strongly procurved thoracic furrow. There are 15 walls or the underside of overhanging ledges, often
species in the southern and western United States, in- along streams. They are moderately large (8-12 mm),
cluding two members of the genus Cyelocosmia, which spindly spiders. There are 10 North American species
has the posterior part of the abdomen strongly trun- in the Appalachians and Southwest.
cated and covered in a heavily sclerotized disco When
the spider is inverted, the abdomen fits tightly against
Clade Haplogynae
the walls of the burrow, forming a false bottom.
Family Cyrtaucheniidae: These trapdoor spiders The term haplogyne condition refers to the female re-
(see preceding entry) have a transverse or slightly productive system in which sperm is deposited and ex-
procurved thoracic furrow. Only the promargin of the pelled via the same passages (contrast with Entelegy-
chelicerae is toothed, although the retromargin may nae). This condition is primitive for spiders, being
have a row of low, rounded tubercles. There are 17 found in mesotheles, mygalomorphs, and hypochilids.
species in the southern and western United States. However, the grouping of the Haplogynae is supported
Family Nemesiidae:These spiders construct an as monophyletic by other lines of evidence, including
open burrow that lacks a trapdoor. They are distin- the fusion of parts of the male palp to form a pyriform
guished by a row of 4-7 stout setae on the upper inner bulb (Figure 5-71) and details of the spinnerets
surface of the chelicerae. There are five species in (Platnick et al. 1991). The Haplogynae include the
California. next 13 families listed.
Family Theraphosidae- Tarantulas:These distinc- Family Filistatidae:These small to medium-sized
tive, conspicuously hairy, two-clawed spiders are spiders (3-15 mm) have the eight eyes tightly grouped
among the largest u.s. species (8-34 mm). They build on a rounded hump, a nearly horizontal clypeus, weak
open burrows, although these may be closed during chelicerae fused medially, spinnerets and anal tubercle
the day by a thin sheet web or plugged with soil in the advanced from the posterior part of the abdomen to a
winter. Despite their reputation, the venom of North ventral position, and a divided cribellum. They are the
American tarantulas is not dangerous. However, they only araneomorph spiders in which the female molts
can produce clouds of urticating hairs by rubbing their after sexual maturity. They build sheet webs with radi-
abdomen with their hind legs. These hairs can irritate ating lines and a central funnel-shaped retreat. There
the mucous membranes of mammals, such as the eyes are seven species in the southern and western United
and respiratory passages. There are 57 species in the States.
southwestern United States. Family Caponiidae: This family includes our only
two-eyed spiders, although the number of eyes is vari-
Infraorder Araneomorphae able. The lateral face of the chelicera has a stridulatory
file. These small spiders (2-6 mm) lack book lungs,
This group contains the vast bulk of spider diversity. having the anterior and posterior respiratory organs
They differ from the Mygalomorphae in having modified into tracheae. Eight species are described
the chelicerae hinged at the base to move in and out, from the southwestern United States. They live in litter
rather than up and down as in mygalomorphs. and under stones.
They also have the fangs articulated so that they op- Family Dysderidae: This family is represented in the
pose each other, rather than moving in parallel (Fig- United States by one introduced species, Dysdera crocata
ure 5-6B). The Araneomorphae are also united by the C. L. Koch. This two-clawed, ecribellate species has six
origin of the cribellum (Figure 5-8A), a specialized eyes arranged in a tight cluster and very large, distally
spinning organ apparently modified from the anterior projecting chelicerae. The coxae of the two anterior
median spinnerets. Most spiders have three functional pairs of legs are longer and thinner than those of the two
pairs of spinnerets, although some mesotheles have posterior pairs (Figure 5-9H). They can be further dis-
nonfunctional remnants of this fourth pair. The tinguished from other spider families except Segestriidae
cribellum has been lost multiple times in spider evo- and Oonopidae by having paired tracheal openings 10-
lution. The Araneomorphae exclusive of Hypochili- cated just posterior to the book lungs (Figure 5-9H).
dae (and so me of its non-North American relatives) These spiders live under bark or stones, where they con-
forms the Araneoclada, which is united by the modi- struct a silken retreat and prey on isopods.
Key to Spider Families of North America 123

FamilySegestriidae:These three-clawed,ecribellate in the southem United States. Some live in caves; oth-
spidershave six eyes arranged in three dyads. They are ers are associated with rocks and leaf litter.
distinguished by having the third pair of legs directed FamilyTelemidae: Tiny (1-2 mm) three-clawed spi-
anteriorly. The tracheal system is as in dysderids. ders that may be six-eyed or blindo Legs are long and
Segistriids are widespread in the United States, with thin. The book lungs are modified into an anterior set
seven described species. of tracheae. The abdomen has a distinctive sclerotized
Family Oonopidae: Tiny 0.5-3 mm) ecribellate, "zigzag" above the pedicel. There are four species in
two-clawed spiders with six eyes arranged in a tight westem North America, usually associated either with
group. Many species have large sclerotized plates in caves or damp leaf litter. .
various configurations covering parts of the abdomen. Family Sicariidae:Small to medium-sized (5-12 mni.~
The tracheal system is as in dysderids. These spiders two-clawed spiders with six eyes arranged in three
can be found in leaf litter or buildings; some species are widely separated dyads (Figure 5-91). The chelicerae are
saltatory.There are 24 North American species, mostly fused basally; the lateral face has coarse stridulatory
in the southem United States. striae. The legs are relatively long and slender. This fam-
Family Pholcidae-Cellar Spiders: Small to medium- ily is represented in North America by the genus Lox-
sized (1.5-9 mm) ecribellate, three-clawed spiders oseeles, which includes the brown recluse (Figure 5-10).
with fused chelicerae, six or eight eyes (Figure 5-9F) , Loxoseeles contains some of the most poisonous species
and very long, thin legs with tarsi subdivided into found in North America. Nevertheless, as mentioned,
many pseudosegments. Unlike most haplogynes, the the frequency of envenomation has been exaggerated.
female genital regio n may be covered by a sclerotized Thirteen species of Loxoseeles can be found in the Mid-
"epigynum," as in most entelegyne spiders. However, west, Southwest, and Atlantic seaboard.
the ducts of pholcid genitalia are haplogyne. Pholcids Family Scytodidae-Spitting Spiders: Small (3.5-
build irregular webs and may vibrate their webs when 5.5 mm), three-clawed spiders with six eyes arranged in
disturbed. Eggs are carried in the chelicerae loosely three widely separated dyads, carapace domed in pro-
bound with silk. Pholcids are common in basements, file, highest posteriorly. The chelicerae are weak and
garages,and other such unimproved structures. Thirty- fused basally. The legs are long and slender. These slow-
four species are represented in North America. moving spiders do not construct snares, but capture
Family Diguetidae: Small to medium-sized (4- their prey by spitting out a mucilaginous substance that
10 mm) ecribellate, three-clawed spiders with six engulfs the prey and fastens it to the substrate. There
eyes arranged in three dyads, long thin legs, and first are five North American species.
tarsus of the male subdivided into many pseudoseg-
ments. The chelicerae are fused medially with a flexi- Clade Entelegynae
ble membrane; the outer surface has a stridulatory
The term entelegyne condition refers to the female gen-
file. A pair of tracheal spiracles are located anterior to
italia, which has separate ducts for the reception and
the spinnerets about one third the distance to the epi-
expulsion of sperrn. Entelegyne spiders usually have
gastric furrow. These spiders live in silken tubes at-
tached to a series of overlapping silken disks that are
arranged like shingles on a roof and attached to vege-
tation. There are seven species in the southwestern
United States.
Family Plectreuridae: Medium-sized (5.5-12 mm)
ecribellate, three-clawed spiders with eight eyes. The
chelicerae are as in diguetids. These spiders construct
silk tubes with silk radiating out at the entrance and
are often found in hot, arid habitats. There are 16
species in the southwestem United States.
Family Ochyroceratidae:Tiny (1-2 mm), ecribellate,
six-eyed, three-clawed spiders with long, thin legs.
These spiders resemble pholcids but lack pseudoseg-
mented tarsi. The chelicerae are free, although a me-
dian lamella is presento These spiders have a distinctive
purplish coloration. One species is found in Florida.
Family Leptonetidae: Tiny (1-3 mm), three-clawed
spiders that may be six-eyed, four-eyed, or blindo Most Figure5-10 Brown recluse spider, Loxosee/es recluse
six-eyed species have a posterior dyad set apart from an Gertsch and Mulaik. (Courtesy of U.5. Public Health
anterior row of four eyes. There are 34 species, mostly Service).
124 Chapter5 PhylumArthropoda

the female genitalia covered by a sclerotized plate, the Family Hersiliidae:These flat, ecribellate three-
epigynum. The origin of the entelegyne condition is clawed spiders are easily distinguished by their ex-
synapomorphic for a large group of spiders, but rever- tremely long, tapered posterior spinnerets, with the dis-
sal to the "haplogyne" condition has occurred several tal segment about as long as the abdomen. Two species
times. Entelegyne spiders usually have a very compli- are, rarely, collected in southern Texas and Florida.
cated male pedipalp, featuring inflatable sacs and nu-
merous sclerites. Prior to copulation, the sacs may fill Clade Orbiculariae
with hemolyrnph, radically altering the configuration
This large group of three-clawed spiders includes the
of the palpal bulbo The following spiders all belong to
orb-weavers (Figure 5-llB) and their descendents.
the Entelegynae.
The Uloboridae and Deinopidae have an undivided
Clade Eresoidea cribellum and comprise the Deinopoidea; the 11 re-
maining families are ecribellate and belong to the Ara-
This small group of spiders is mostly tropical and con- neoidea. Many araneoids have a paracyrnbium attached
tains some social species. lt includes cribellate and to the retrolateral part of the cymbium (Figure 5-7J);
ecribellate spiders in three families, two of which are this structure is of taxonomic importance.
represented in North America. Family Uloboridae-Hackled-BandOrb-Weavers: This
Family Oecobiidae: This familyof tiny 0-4.5 mm) four- or eight-eyed family lacks poison glands and builds
three-clawed spiders is distinguished by its enlarged, a complete or reduced orb web. The family of small to
jointed anal tubercle fringed with long curved hairs. medium-sized spiders (2-6 mm) is widespread in North
Four eyes are well developed, four are degenera te. America, with 14 species.
North American species have a divided cribellum, al- Family Deinopidae-Ogre-Faced Spiders: These
though other family members are ecribellate. This fam- night-active spiders are easily recognized by their ex-
ily is widely distributed and is often found in houses. tremely large posterior median eyes (Figure 5-9G).
There are eight North American species. They have an elongate body (12-17 mm) and long,

A e

Figure5-11 A, Black
widow spider, Latrodectus
sp. (Theridiidae), female
hanging from its web;
B, A garden spider, Argiope
aurantia Lucas; C, A crab
spider, Misumenops sp.
(Thomisidae); D, Ajump-
ing spider, Phidippus audax
(Hentz) (Salticidae).
(A, Courtesy of Utah
Agricultural Experiment
D Station.)
Keyto SpiderFamiliesof NorthAmerica 125

slender legs. They build a modified orb web, which licerae. The U.s. anapid species is a tiny (1-1.3 mm)
theyhold in their first two pairs of legs and cast like a eight-eyed spider found in Oregon and California. The
netto capture prey. There is one species in the south- book lungs are modified into tracheae, and the poste-
eastemUnited States. rior tracheal system is lost (variable within the family).
FamilyAraneidae:This is a diverse family,nearly all The abdomen of the male has a dorsal scutum; the fe-
ofwhichconstruct an orb web. Included in this group male has the dorsal surface of the abdomen coriaceous
arethe distinctive day-active spiny orb-weavers of the and lacks a pedipalp.
generaGasteracantha and Micrathena, the garden spi- Family Mysmenidae:These are minute (0.5-2 mm)
dersofthe genus Argiope, and the Bolas spiders (genus eight-eyed spiders with 50ft abdomens. Females have a
Mastophora).Bolas spiders have abandoned the orb pedlpalp. In most of our species, the male has a clasp-
web;theyuseaggressivechemicalmimicry to lure male ing spur on metatarsus I. The natural history of this
moths,which they disable by striking them with a group is diverse. Some build horizontal webs similar to
globuleof sticky silk swung on the end of a line. There those of anapids; others build a "three-dimensional"
are 164 araneid species widely distributed in North orb web, with radii not restricted to a single plane
Ameriea. (Eberhard 1982, 1986; Coddington 1986); still others
FamilyTetragnathidae:This family of orb-weavers have abandoned web building to live as kleptoparasites
includesNephila clavipes (L.), which builds a golden in the webs of other spiders. Five species are wide-
orb web that may be a meter in diameter. Female spread in North America.
Nephila are very large (22-26 mm), but the males are Family Mimetidae-Pirate Spiders: Mimetids are
muehsmaller (6-8 mm) and may be overlooked by characterized by a distinctive pattern of prolateral
colleetors.Nephila can be common in the southern macrasetae on the first and second pair of legs (Fig-
states.Nephila and most araneids build vertical orb ure 5-7 A). These setae are arranged in a repeating
webs;the remaining tetragnathids typically build hori- pattern with each series containing several sequen-
zontalorb webs. Members of the genus Tetragnatha tially longer setae. Mimetids are araneophagous, ag-
oftenbuild webs over water. These spiders are long- gressive mimics that typically invade the webs of
bodiedwith very long and protruding chelicerae, espe- other spiders. In the web, mimetid locomotion is typ-
ciallyin the males. There are 43 tetragnathid species ically very slow and characterized by frequent pauses.
widelydistributed in North America. The mimetid then jerks or plucks the web, apparently
FamilyTheridiosomatidae: These tiny (1-2.5 mm) imitating either a struggling insect or a courting maleo
orb-weaversare recognized by the presence of a pair of When the victim moves close enough, the mimetid
pitson the anterior part of the sternum adjacent to the delivers a lethal bite of fast-acting venom (Bristowe
!abium,by their truncated sternum, and long tri- 1958). The higher-Ievel classification of mimetids is
chobothriaon tibiae III and IV.In our genus, the spider controversial.
sitsat the center of a vertical orb pulled into a cone by Family Theridiidae-Cobweb Spiders: The follow-
athreademanating from the hubo There are two species ing four families have abandoned the orb web in favor
ineastemNorth America. of a mesh sheet or three-dimensional cobweb. Theridi-
FamilySymphytognathidae: Symphytognathiclsand ids typically build cobwebs, although some have
anapidsboth build small horizontal orb webs. These adopted a kleptoparasitic lifestyle. They are usually
webs aredistinctivebecause the spider builds a second eight-eyed (rarely six-eyed or blind) with a high
seriesof radii after the sticky spiral is laid down. As a clypeus; they may have a stridulatory organ above the
result,the spiral threads to not change direction at pedicel between the carapace and abdomen. Theridiids
everyradius, as they do in all other orb webs. Symphy- usually have a comb of serrated setae on the fourth tar-
tognathidwebs differ from anapid webs in having sus (Figure 5-7B). The tarsal comb is used to wrap
manymore of these secondary radii, and by the prey in silk before envenomation. They rarely have
absenceof any radii outside the plane of the web teeth on the retrolateral margin of the chelicerae; teeth
(Eberhard1982, 1986; Coddington 1986). Symphyto- on the promargin occur but are also rare. The para-
gnathidsare distinguished by the basal fusion of their cymbium, if present, is fused to the retrodistal part of
chelicerae,the absence of the female pedipalp (shared the cymbium; the palpal tibia never has apophyses.
withAnapidae), the modification of the book lungs The colulus is reduced or absent in some theridiids.
intotraeheae,and the loss of the posterior tracheal sys- This family includes the widow spiders (Latrodectus;
temoThere is one tiny (0.5 mm) four-eyed species in Figure S-HA), known for their potent venom (see ear-
Florida. lier section on medically important spiders). AIso rep-
FamilyAnapidae:Anapids are distinguished fram resented among the 250 or so North American
otherspiders by the presence of a spur arising from just theridiid species are social spiders (Anelosimus) and
abovethe oral cavity behind and between the che- kleptoparasites (Argyrodes).
126 Chapter5 PhylumArthropoda

Family Nesticidae: Nesticids may have eight, six, or Family Dictynidae:Dictynids are eight-eyed (occa-
four eyes or lack them altogether. They are typically sionally six-eyed or blind), three-clawed spiders. The
pale in color and many species are associated with family includes both cribellate and ecribellate mem-
caves. Nesticids share several characteristics with bers; when present, the cribellum is usually entire, but
theridiids, including a comb on tarsus IV; nesticids occasionally is divided. Some dictynids have lost the
have a well-developed colulus. Males are readily distin- tarsal trichobothria. There are over 280 North Ameri-
guished from theridiids by presence of a conspicuous can species.
paracymbium fused to the retrobasal part of the cym- Family Zodariidae:These three-clawed, ecribellate
bium. There are 37 species in North America, mostly spiders are distinguished by the absence of serrula on
endemic to the Appalachian region. the distal margin of the endites, and by the reduction
Familylinyphiidae: Linyphiidsare the most species- or absence of the posterior lateral and posterior median
rich spider family in North America (>800 species). spinnerets. There are five North American species.
Whereas some species build conspicuous sheet or Family Hahniidae:Hahniids are ecribellate, three-
dome-shaped webs, most species are tiny (1-3 mm) and clawed, sheet web builders. Many North American
live in leaf litter. The males of some species have bizarre hahniids are easily distinguished by the configuration
head modifications that are associated with mating. of the spinnerets in a transverse row and the advanced
Linyphiids are typically eight-eyed spiders (rarely six- position of the tracheal spiracle. Other hahniids lack
eyed or blind) with a high clypeus, stridulatory striae these characters, but are distinguished by rows of ven-
on the lateral face of the chelicerae (Figure 5-7G), and tral spines on metatarsus and tibia 1, by the presence of
a tendency for legs to break at the patella-tibia joint. an undivided colulus, and the absence of plumose
The male palp features a small, often hooklike para- hairs.
cymbium attached to the retrobasal part of the cym- Family Agelenidae: These three-clawed spiders are
bium by a membrane (Figure 5-7J). The palpal tibia characterized by the presence of a divided colulus (Fig-
may or may not have one or more apophyses. ure 5-8B). They typically have feathery hairs on the ab-
Family Pimoidae: These spiders resemblelinyphiids domen, elongated posterior spinnerets, and build a
but are larger than most of those species (4.5-10.5 mm). funnel web. Their webs can be quite numerous and
They share with linyphiids a high clypeus, stridulatory conspicuous. There are 89 North American species.
striae, and leg breakage at the patella-tibia joint; theyare Family Cybaeidae: These three-clawed spiders have
best separated from them by details of the genitalia. contiguous anterior spinnerets that are longer than the
The pimoid paracymbium is fused to the retrobasal posterior spinnerets and a more or less transverse, en-
part of the cymbium; the palpal tibia never has an tire colulus. There are 44 North American species.
apophysis. Thirteen species are represented in western Family Desidae:These three-clawed spiders are
North America. characterized by their greatly enlarged, distally project-
The remaining spiders all belong to a large clade of ing chelicerae. We have only two species in North
spiders that, along with some non-North American America. Paratheuma is a three-clawed, ecribellate spi-
groups, is the sister group to the Orbiculariae. This group der with widely separated anterior spinnerets. It is as-
does not have a formal taxonomic name at this time. Re- sociated with intertidal habitats in coastal Florida. Bad-
lationships among the taxa within this clade are not all umna has a divided cribellum and is introduced to
well defined, but some distinctive groups are noted. California.
FamilyTitanoecidae: Smallto medium-sizedspiders Family Amphinectidae: These three-clawed spiders
(3.5-8 mm) with a well-developed, divided cribellum. have a divided cribellum and 5-7 strong teeth on both
These spiders sometimes have short, inconspicuous cheliceral margins. The male palpus has a filiform em-
tarsal trichobothria. Males have ventral rows of short bolus enclosed in a membranous conductor. One
macrosetae on metatarsi and tibiae I and Il, reduced in species is introduced to California and the Gulf coastal
females. Four species are widespread in North America. region.
Family Amaurobiidae:Amaurobiids are three-
RTA (retrolateral tibial apophysis) clade clawed, cribellate or ecribellate spiders. They may
build large sheet webs, or live in silk tubes with only a
This large group of spiders all have a retrolateral few lines radiating from the opening. There are 124
apophysis on the male palpal tibia and have tri- North American species.
chobothria on the tarsi, usually increasing in length Family Tengellidae:North American species are
distally (Figure 5-7 C). Some families have two rows of ecribellate spiders with two or three claws, claw tufts,
trichobothria or an irregular cluster; others have the and two rows of tarsal trichobothria. There are 23
trichobothria secondarily reduced or lost. Lycosids North American species, including several associated
have lost the retrolateral tibial apophysis. with caves.
Keyto SpiderFamiliesof NorthAmerica 127

Family Zorocratidae: These large (5-13 mm) spi- Family Pisauridae-Nursery-Web and Fishing Spi-
ders have a large, divided cribellum, scopulate tarsi, ders: These medium to large (6.5-31 mm) spiders re-
and two rows of tarsal trichobothria. Tarsus 1has three semble wolf spiders, but differ in the eye arrangement:
claws; other tarsi have third claw absent. The che- The anterior eye row is variable, usually smaller than
licerae have lateral stridulatory striae. There are five the posteriors; the posterior row is strongly recurved,
speciesin the southwestern United States. with the eyes subequal in size (Figure 5-9D). The egg
sac is carried by the female under her cephalothorax,
held there by the chelicerae and pedipalps. Before the
Supeñamily Lycosoidea eggs hatch, the female attaches the sac to a plant,
The Lycosoidea includes the seven families listed builds a web around it, and stands guard nearby.
next. The primitive condition for this group is that of Pisaurids forage for their prey and build webs only for
a cribellate web builder. With the exception of one their young. Some spiders in this group, particularly
Iycosid in the southeastern United States and an in- the fishing spiders of the genus Dolomedes, are quite
troduced species of zoropsid, all North American large. Dolomedes live near water; they may walk over
members of this group are ecribellate wandering the surface of the water or dive beneath it to feed on
hunters. This group is united by specializations of aquatic insects and sometimes small fish. There are 13
the eyes and characteristics of the male genitalia. species in North America.
Most members have three tarsal claws, but there are Family Trechaleidae: Large (14-16 mm), flattened,
exceptions. fast-moving, three-clawed spiders with long, flexible
FamilyCtenidae-Wandering Spiders: These medium tarsi. There is one species in the Arizona, found near
to large (6.5-30) spiders are recognized by their eye permanent streams in xeric regions. Females may carry
arrangement in which tiny anterior lateral eyes are al- a disc-shaped egg case attached to the spinnerets.
most contiguous with both the posterior median and Family Lycosidae-Wolf Spiders: Lycosids are dis-
posterior lateral eyes. They have two claws and claw tinctive three-clawed spiders recognized by their eye
tufts; scopulae vary from absent to dense and extend- pattern: anterior eyes small in a more or less straight
ing onto the metatarsi. Tarsal trichobothria are scat- row; posterior median eyes very large, posterior lateral
tered, not in a defined row. This group is chiefly trop- eyes smaller, positioned well behind the posterior me-
ical; seven species occur in the southern United dians (Figure 5-9C). The male palp lacks a tibial
States. apophysis. Most North American species are foragers,
Family Zoropsidae: These two-clawed spiders have except Sosippus,which constructs a sheet web with a
6-7 pairs of ventral spines on the anterior tibiae; a funnel-shaped retreat. The egg sac is carried by the fe-
calamistrum consisting of an oval patch, rather than male, attached to her spinnerets. When the young
one or two rows of setae; and a divided cribellum. A hatch, they are carried for a time on the back of the fe-
single species has be en introduced to California. maleo Lycosids are widely distributed, with over 250
Family Miturgidae: These ecribellate, two-clawed species in North America. They occur in many differ-
spiders have dense scopulae and claw tufts. The poste- ent habitats and are often quite common.
rior spinnerets are distinctly two-segmented and the
distalsegment is conical (Figure 5-8A). The absence of Clade Dionycha
a thoracic furrow is indicative of Cheiracanthium, a mi-
turgid genus of minor medical importance. There are The Dionycha include the 13 families listed next. This
10species in North America. is a large group of two-clawed, eight-eyed (North Amer-
Family Oxyopidae-Lynx Spiders: These three- ican fauna), ecribellate spiders; it is probably not mono-
clawed spiders can be recognized by the eye arrange- phyletic. Unless otherwise noted, they have scopulae
ment, with the anterior lateral and posterior pairs of (at least on tarsi 1 and ll, often on the metatarsi also)
eyesforming a hexagon, and the anterior median eyes and claw tufts. All North American representatives for-
below the anterior laterals (Figure 5-9A). The ab- age without a web, although this does not hold true
domen often tapers to a point posteriorly, the legs have worldwide. Sparassidae, Selenopidae, Philodromidae,
many long macrosetae, and the tarsi have two rows of and Thomisidae are crab spiders with laterigrade legs
trichobothria; some are a distinctive bright green. Most (Figure 5-100; the remaining families have the typical
Iynxspiders forage over foliage and ambush their prey; prograde orientation. Laterigrade legs are rotated so
others sit and wait on limbs or twigs. Members of the that the morphologically pro lateral surface is dorsal.
genus Oxyopes are strong jumpers. North American Family Clubionidae: These small to medium
species do not construct a web or retreat, although (2.5-10.5 mm) spiders have dense claw tufts and thin
web-building species can be found in the tropics. Eigh- scopulae. They are usually pale yellow or light gray.
teenspecies occur in North America. Clubionids typically spend the day in silk tube retreats

128 Chapter5 PhylumArthropoda

and forage for prey at night. There are 61 species in are flattened, nocturnal, fast-moving, two-clawed spi-
North America. ders that lack a colulus. There are six species in the
Family Anyphaenidae:These spiders resemble clu- southern United States.
bionids, but have the tracheal spiracle advanced at least Family Sparassida~iant Crab Spiders:These large
half way between the spinnerets and the epigastric fur- spiders 00-25 mm) are distinguished by a unique
row, and the hairs of the claw tufts are somewhat flat- trilobed membrane on the dorsoapical part of the
tened. Tarsus and metatarsus I and Il have sparse scop- metatarsi (Figure 5-7F). Eleven species are generally con-
ulae. Their behavior is similar to clubionids, retreating fined to the southern United States, but are sometimes
to silk tubes by day and foraging at night. There are 39 found in the North associated with shipments ofbananas.
species in North America. Family Philodromidae:These somewhat flattened
Family Corinnidae:The legs usually have numerous spiders have dense scopulae and claw tufts. The eyes
ventral macrosetae (absent in Trachelas), dense claw are usually subequal in size and not on tubercles. Teeth
tufts, and sparse scopulae. The abdomen often has scler- are present on the promargin of the chelicerae, absent
otized plates. Some species are ant mimics. North Amer- from the retromargin. The legs are either subequal in
ica has 122 species.. length, or leg Il is much longer than the others. There
Family Liocranidae:These spiders typically lack are nearly 100 species in North America.
paired ventral macrosetae on tibiae and metatarsi I and FamilyThomisidae: Thesesomewhatflattenedspiders
Il; scopulae and claw tufts are absent or very thin. The lack scopulae and claw tufts. The lateral eyes are often
abdomen often has iridescent setae or one or more larger than medians and positioned on tubercles (Fig-
scuta. There are eight species in North America. ure 5-9E) The chelicerae are almost always without
Family Zoridae:These small to medium-size (2.5- teeth, or have both margins toothed. Legs I and Il are
7 mm) spiders have the anterior eye row nearly straight longer and thicker than legs III and IV (Figure 5-11C).
and the posterior eye row strongly recurved. The tibiae One common species in this group is the goldenrod spi-
have 6-8 pairs of overlapping macrosetae. There are der, Misumena vatia (Clerk), which is white or yellow
two species in North America. with a red band on either side of the abdomen. This spi-
Family Gnaphosidae: These spiders have the poste- der is a sit-and-wait predator, often occupying a flower
rior median eyes flattened and irregularly shaped. The and attacking pollinators. This spider can change color,
anterior spinnerets are cylindrical, sclerotized, and sep- over a period of a few days, depending on the color of the
arated by at least their width (Figure 5-8B). The en- flower. There are over 140 species of thomisids in North
dites usually have an oblique depression on the ventral America.
surface. These spiders are usually dark, but some have Family Salticidae-Jumping Spiders: These stout-
light markings on the abdomen. The males sometimes bodied, short-Iegged spiders have a distinctive eye pat-
have a dorsal scutum. Unlike prodidomids, gnaphosids tern, with the anterior median eyes by far the largest
have teeth on the tarsal claws and at least the promar- (Figures 5-9B, 5-11D). The body is rather hairy and is
gin of the chelicerae toothed. These spiders are typi- often brightly colored or iridescent. Some species are
cally active at night. Over 250 species are in North antlike in appearance. Jumping spiders forage for prey
America. in the daytime. They approach their prey slowly, and
Family Prodidomidae:These spiders are closely re- then suddenly leap onto it. They can jump many times
lated to gnaphodids, but are distinguished in having their own body length. Before jumping, they attach a
the posterior eye row strongly procurved, both che- silk thread dragline, which they can use to climb back
liceral and tarsal claw teeth are absent, and by details to their starting position. Salticids are the world's most
of the spinnerets. Two species live in the southern diverse spider family, with over 330 representatives in
United States. North America.
Family Homalonychidae:Moderately large (6.5-
9 mm) spiders with strongly converging endites; they Order Ricinulei14-Ricinuleids
lack serrula on the anterior margin of the endites, teeth
on the tarsal claws, and a colulus. Juveniles and adult This is a small group of rare tropical arachnids. They
females are usually covered with fine soil, which ad- are somewhat ticklike in appearance, and one of their
heres to modified setae, providing effective camouflage. distinctive features is a movable flap at the anterior end
There are two species in the southwestern United of the prosoma that extends over the chelicerae. The
States. tarsi of the third pair of legs in the male are modified as
Family Selenopidae:These spiders have the one copulatory organs. One species, Cryptocellus dorotheae
pair of posterior eyes advanced so that the anterior eye
row appears to include six eyes; the posterior eye row 14Ricinulei: Rícin, a kind of mite or tick; uleí, small (a diminutive
includes two relatively large, widely spaced eyes. They suflix).
Keyto SpiderFamiliesof NorthAmerica 129

Gertsch and Mulaik, has been reported from the Rio

GrandeValley of Texas. This arachnid is about 3 mm in
length,is orange-red to brown in color, and occurs un-
der objects on the ground. Some tropical species are
larger(up to about 15 mm in length), and the majority
havebeen taken in caves.

Order Opiliones15-Harvestmen,
Ihese arachnids have the body rounded or oval, with the
prosomaand opisthosoma broadly joined. There are usu-
alIytwo eyes, generally located on each side of a median
elevation.Scent glands are present, their ducts opening
to the outside above the first or second coxae. These
glandssecret a peculiar-smelling fluid when the animal is
disturbed. Most species are predaceous or feed on dead
animalsor plant juices. The eggs are laid on the ground
in the fall and hatch in the spring. Most species live ayear
or two. This order is divided into three suborders, the
Cyphophthalmi, the Laniatores, and the Palpatores.
SuborderCyphophthalmi-Mite Harvestmen
Ihese are small, short-legged, mitelike forms, 3 mm in
lengthor less, which differ from the other two suborders
in having the scent glands opening on short conical pro-
cesses.The eyes, if present, are far apart and indistinct.
Ihis group is represented in the United States by four Figure 5-12 A harvestman or daddy-longlegs (Order
species,which occur in the Southwest and the Far West. Opiliones). A, Lateral view; B, Anterior view. eh, che-
SuborderLaniatores licera; 1,front leg; De,eyes or ocelli; pdp, pedipalp. The
Ihis group is mainly tropical, but more than 60 species harvestmen typically have two eyes located on a tubercle,
and the chelicerae are clawlike, as shown in B.
occur in the southern and western states, many of them
in caves. These differ from the Palpatores in having the
tarsi on the third and fourth legs with two or three million more are still undescribed. The body is usually
claws (or one claw with three teeth). The pedipalps are oval, with little (Figures 5-13,5-14) or no (Figures 5-15
largeand robust, and their tarsi are armed with a strong through 5-20) differentiation of the two body regions.
claw.The legs are not unusually long. Newly hatched young, called larvae, have only three
SuborderPalpatores-Daddy-Longlegs pairs of legs (Figures 5-17B, 5-20A) and acquire the
Ihis group includes the harvestmen commonly called fourth pair after the first molt. A few mites have fewer
daddy-longlegs, which have very long and slender legs than three pairs of legs (Figure 5-18). The instars be-
(Figure5-12A). About 150 species occur in North Amer- tween larva and adult are called nymphs.
ica.These forms have the second pair of legs the longest, The Acari occur in practically all habitats in which
and the tarsi of alllegs have just one claw. The pedipalps any animal is found and rival the insects in their varia-
are smaller, their tarsi having a weak claw or none. Four tions in habits and life histories. This group includes
families of this suborder occur in North America, but both aquatic and terrestrial forms, and the aquatic
most U.5. species belong to the Phalangidae. forms occur in both fresh and salt water. icari are abun-
dant in soil and organic debris, where they usually out-
Order Acari16-Mites and Ticks number other arthropods. Many are parasitic, at least
during part of their life cycle, and both vertebrates and
Ihe Acari constitute a very large group of small to invertebrates (including insects) serve as hosts. Most
minute animals. More than 30,000 have been de-
of the parasitic forms are external parasites of their
scribed, and it has be en estimated that perhaps half a hosts. Many of the free-living forms are predaceous,
and some of these prey on undesirable arthropods.
"Opiliones: from the Latin, meaning a shepherd. Many are scavengers and help break down forest liuer.
"Acari: from the Greek, meaning a mite. Many are plant feeders, and some of these harm crops.
130 Chapter5 PhylumArthropoda

Some of the parasitic forms are pests of humans and Group IlI. Acariformes
other animals, causing damage by their feeding and Suborder Prostigmata (Trombidiformes,
sometimes serving as vectors of disease. This group is Actinedida)
of considerable biological and economic importance. Suborder Astigmata (Sarcoptiformes,
The groups of Acari have been variously arranged Acaridida)
by different authorities; we follow here the arrange- Suborder Oribatida (Oribatei, Cryptostigmata)
ment ofBarnes (1987), but call the three major groups
of Acari "groups" rather than orders. This arrangement Groupl. Opilioacariformes
(with other spellings, names, or arrangements in The members of this group (Figure 5-13) are elongate
parentheses) is as follows: and somewhat leathery, and have the abdomen seg-
Order Acari-mites and ticks mented. They are brightly colored and are superficially
Group I. Opilioacariformes (Opilioacarida, Noto- similar to some of the harvestmen (Opiliones). They
are usually found under stones or in debris, and are
stigmata; Parasitiformes in part)
Group 11.Parasitiformes omnivorous or predatory. The U.s. species occur in the
Suborder Holothyrina (Holothyrida, Tetrastig-
mata) Group11.Parasitiformes
Suborder Mesostigmata (Gamasida) These are medium-sized to large mites that have an un-
Suborder Ixodida (Ixodides, Metastigmata)- segmented abdomen, and the tracheal system has ven-
ticks trolateral spiracles.


Figure5-13 A female opilioacariform mite. A, Dorsal view; B, Ventral view. (From

Johnston 1968, redrawn by van der Hammen.)
Keyto SpiderFamiliesof NorthAmerica 131

Figure5-14 The American dog tick, Dennacentor variabilis (5ay) (Ixodidae). A, Larva;
B, Nymph; e, Adult (unengorged). (Courtesy of U5DA.)

SUBORDER Holothyrina:The members of this group

are fair-sized (2-7 mm in length), rounded or oval
mites that occur in Australia, New Guinea, New
Zealand, some islands in the Indian Ocean, and in the
American tropies. They are found under stones or in
decaying vegetation and are predaceous.
SUBORDER Mesostigmata:This is the largest subor-
der of the Parasitiformes and includes predaceous,
scavenging, and parasitie forms. The majority are free-
living and predaceous and are usually the dominant
mites in leaf litter, humus, and soil. The parasitic mites
in this group attack birds, bats, small mammals,
snakes, insec15, and rarely humans. One parasitic
species, the chicken mite, Dennanyssus gallinae (De Figure5-15 The fowl tick, Argas persicus (Oken),
Geer), is a serious pest of poultry. lt hides during the adult female (Argasidae). A, Dorsal view; B, Ventral view.
day and attacks poultry and sucks their blood at night. (Courtesy of U5DA.)
Ihis species also causes a dermatitis in humans.
SUBORDER Ixodida-Ticks:Twofamiliesof ticks oc-
cur in North America, the Ixodidae or hard ticks and the upon one host, drops off, and mol15; the nymph on a
Argasidaeor 50ft ticks. Ticks are larger than most other second; and the adult on a third. The hard ticks take
Acariand are parasitic, feeding on the blood of mammals, only one blood meal in each of their three instars. They
birds,and reptiles. Those attacking humans are annoying remain on the host for several days while feeding, but
pests,and some species serve as disease vectors. Ticks are usually drop off to molt. The 50ft ticks generally hide
the most important vectors of disease to domestic ani- in crevices during the day and feed on their hos15 at
malsand second only to mosquitoes as vectors of disease night; each instar may feed several times. The hard
to humankind. Certain ticks, especially engorging fe- ticks ordinarily have two or three hos15 during their de-
males feeding on the neck or near the base of the skull of velopment, whereas the 50ft ticks may have many
their host, inject a venom that produces a paralysis; the hos15. The cattle tick that transmi15 Texas cattle fever
paralysismay be fatal if the tick is not removed. The most feeds on the same host individual during all three in-
important tick-bome diseases are Rocky Mountain spot- stars, and the protist that causes the disease is trans-
ted fever, relapsing fever, Lyme disease, tularemia, Texas mitted transovarially, that is, through the eggs to the
cattle fever, and Colorado tick fever. tiek's offspring. The hard ticks (Figure 5-14) possess a
Ticks lay their eggs in various places, but not on hard dorsal plate called the scutum, and they have the
the host; the young seek out a host after hatching. Most mouthpar15 protruding anteriorly and visible from
species have a three-host life cycle: the larva feeds above. The 50ft ticks (Figure 5-15) lack a scutum and
132 Chapter5 PhylumArthropoda

are soft-bodied, and the mouthparts are ventrally 10- are widely distributed, multivoltine, and sometimes oc-
cated and are not visible from above. cur in tremendous numbers. The eggs are laid on the
plant and, during the warm days of summer, hatch in 4
Group 11I.Acariformes or 5 days. There are fOUTinstars (Figure 5-17), and
These are small mites that have the abdomen unseg- growth from egg to adult usually requires about three
mented, and the spiracles are near the mouth parts or weeks. Most species overwinter in the egg stage. The
absent (Figure 5-16). immature instars are usually yellowish or pale in color,
SUBORDERProstigmata:This is a large group and the adults are yellowish or greenish. (These ani-
whose members vary considerably in habits. Some are mals are sometimes called red mites, but they are sel-
free-living (in litter, moss, or water) and vary in food dom red.) Sex in these mites is determined by the fer-
habits; some are parasitic; and some are parasitic as lar- tilization of the egg. Males develop from unfertilized
vae but predaceous as adults. This group includes the eggs, and females from fertilized eggs.
spider mites, gall mites, water mites, harvest mites, The gall mites (Eriophyoidea) are elongate and
feather mites, and others. wormlike and have only two pairs oflegs (Figure 5-18).
The spider mites (Tetranychidae) are plant feed- A few species form small, pouchlike galls on leaves, but
ers, and some species do serious damage to orchard the majority feeds on leaves without forming galls and
trees, field crops, and greenhouse plants. They feed on produces a rusting of the leaves. Some attack buds, and
the foliage or fruits and attack a variety of plants. They one forms the conspicuous witches' -broom twig gall on


r - - - - - - - - - _1_ - - - - - - - - - - -,
I Figure 5-16 A generalized
acariform mite. A, Anterior
view; B, Lateral view.
ch, socket of chelicera;
L__,__~L r ~ ex, coxa; DS], disjugal fur-
GNATHOSOMA PODOSOMA row; Jm, femur; p, socket
L ,
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __J
of palp; ptar, pretarsus;
ptl, genu or patella; tb, tibia;
L_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __~ tr, trochanter; ts, tarsus.
B (Courtesy of Johnston.)
Keyto SpiderFamiliesof NorthAmerica 133

larvae are parasitic on aquatic insects, and most

nymphs and adults are predaceous. Water mites are

small, round-bodied, usually brightly colored (red or
green), and often quite common in ponds. They crawl
on the bottom and on aquatic vegetation and lay their
eggs on the undersides of leaves or on aquatic animals
(mainly freshwater mussels). Water mi te larvae

ti e D E
(mostly in the genus Arrenurus) are often abundant on
the bodies of dragonflies and damselflies. They crawl
from the nymph to the adult when the latter emerges
and may remain there for a couple of weeks, feeding on
the body fluids of the insect and eventually dropping
off and developing into adults (if they happen to get
into a suitable habitat).
Figure5-17 The fourspotted spider mite, Tetranychus The harvest mites (also called chiggersor redbugs)
canadensis(McGregor). A, Egg; B, First instar or larva; (Trombiculidae) are ectoparasites of vertebrates in the
e, Secondinstar, or protonymph; D, Third instar, or larval stage, whereas the nymphs and adults are free-
deutonymph; E, Fourth instar, adult female. (Courtesy of living and predaceous on small arthropods and arthro-
USDA,after McGregor and McDunough.) pod eggs. Harvest mites lay their eggs among vegeta-
tion. Gn hatching, the larvae crawl over the vegetation
and attach to a passing host. They insert their mouth-
hackberry. Many gall mites are serious pests of orchard parts into the outer layer of the skin, and their saliva
trees or other cultivated plants. partly digests the tissues beneath. The larvae remain on
The water mites (Hydrachnidia, with 45 families the host for a few days, feeding on tissue fluid and di-
in nine superfamilies) include a number of common gested cellular material, and then drop off. These mites
and widely distributed freshwater species and a few are small (Figure 5-19A) and are seldom noticed. Their
marine forms. A few species occur in hot springs. The bites, however, are quite irritating, and the itching per-
sists for some time after the mites have left. Gn people,
these mites seem to prefer areas where the clothing is
tight. A person going into an area infested with chig-
gers can avoid being attacked by using a good repel-
lent, such as dimethyl phthallate or diethyl toluamide.
This material can be put on the clothing, or the cloth-
ing can be impregnated with it. A good material to re-
duce the itching caused by chiggers is tincture of ben-
zyl benzoate. In Asia, the Southwest Pacific, and
Australia, certain harvest mites serve as vectors of
scrub typhus, or tsutsugamushi disease. This disease
caused more than 7,000 casualties in the V.s. armed
forces during World War n.
The feather mites 09 families, in the superfamilies
Analgoidea, Pterolichoidea, and Freyanoidea) consti-
tute a large group whose members occur on the feath-
ers or skin or (rarely) in the respiratory system ofbirds.
Many are found on particular feathers or feather areas
of their hosts. They are seldom of economic impor-
tance, although they often occur in considerable num-
bers on poultry or avian pets. They appear to be scav-
engers, feeding on feather fragments and oily
secretions on the feathers. Some of those occurring on
aquatic birds feed on diatoms.
The Tarsonemidae is a large family of mites that
Figure5-18 A gall mite, PhyllocoptesvariabilisHodgkiss, includes species associated with insects or plants, and
a species attacking sugar maple; 460 X . A, Dorsal view; some cause human dermatitis. Some species live in the
B, Ventral view. (Redrawn fram Hodgkiss 1930.) galleries of bark beetles, where they feed on the bark

134 Chapter5 PhylumArthropoda


Figure5-19 A, A chigger, the larva of Eutrombiculaalfreddugesi(Oudemans), 215x;

B, An oribatid mite (family Euphthiracaridae). (A, Redrawn from a US Public Health Ser-
vice release; B, Courtesy of the lnstitute of Acarology).

beetle eggs. They are carried from gallery to gallery on

the bodies of the beetles. Species in the genus Acarapis
occur on the bodies of honey bees. Acarapis woodi
(Rennie) causes what is known as lsle of Wight disease
in honey bees. A few species (Tarsonemus and other
genera) have been implicated in cases of human der-
SUBORDER Astigmata:The mites in this group are
mostly terrestrial and nonpredatory. Some are parasitic,
and a few of the parasitic forms are important pests of
people and animals. The most important mites in this
group are probably those that infest stored foods and
those that cause dermatitis in humans and animals.
The Acaroidea are principally scavengers, occur-
ring in animal nests, stored foods, and plant tissues.
Figure5-20 The sheep scab mite, Psoroptesovis
Those occurring in stored foods (cereals, dried meats,
(Hering), female. A, Dorsal view; B, Ventral view.
cheese, and the like) not only damage or contaminate (Courtesy of USDA.)
these materials, but they may get on people and cause
a dermatitis called grocers itch or millers itch.
The most important mites in this suborder that SUBORDEROribatida: This is a large group of
cause dermatitis in humans and animals are in the fam- small mites (0.2-1.3 mm) that vary considerably in
ilies Psoroptidae and Sarcoptidae. The Psoroptidae in- formo Some superficially resemble small beetles (Fig-
elude the mange mites, which attack various animals ure 5-20B) and are called beetle mites. Some species
(Figure 5-20), and the Sarcoptidae inelude the itch or have winglike lateral extensions of the notum. In a few
scab mites. These mites burrow into the skin and cause cases these extensions, called pteromorphs, are hinged,
severe irritation, and the resulting scratching often contain "veins," and are provided with museles. Orib-
causes additional injury or leads to secondary infec- atid mites are found in leaf litter, under bark and
tion. One of the best treatments for scabies (infection stones, and in the soil. They are mainly scavengers.
by these mites) is the application of a solution of ben- They make up a large percentage of the soil fauna and
zyl benzoate. Species of Dermatophagoides (Pyroglyphi- are important in breaking down organic matter and
dae) are common inhabitants of houses and have been promoting soil fertility. Some species of Oribatuloidea
have been found to serve as the intermedia te hosts of
_J<f:~d jn housedust allergi:e~~:
Keyto SpiderFamiliesof NorthAmerica 135

A B e

Figure5-21 Pseudoscorpions. A, Dactylochelifer copiosusHoff; B, Larca granulata

(Banks); C, Pselaphochernesparvus Hoff. (Courtesy of Hoff and Illinois Natural History

certain tapeworms that infest sheep, cattle, and other Florida). They are called by a variety of names: wind-
ruminants. scorpions (they run "like the wind"), sunscorpions, sun-
spiders, and camelspiders. They are 20-30 mm in
Order Pseudoscorpiones17-Pseudoscorpions length, are usually pale colored and somewhat hairy,
and the body is slightly constricted in the middle (Fig-
The pseudoscorpions are small arachnids, seldom ure 5-4C). One of their most distinctive features is
more than 5 mm in length. They resemble true scorpi-
their very large chelicerae (often as long as the pro-
ons in having large chelate pedipalps, but the opistho-
soma), giving them a very feroeious appearance. They
somais short and oval, there is no sting (Figure 5-21),
may bite, but they do not have venom glands. The
and the body is quite flato The pseudoscorpions differ
males have a flagellum on the chelicera. The fourth legs
fram most other arachnids in lacking a patellar seg-
bear, on the ventral side of the coxae and trochanters,
mentin the legs. Eyes may be present or absent; if pres-
ent, there are two or four, located at the anterior end of five short, broad, T-shaped structures (attached by the
base of the T) called racket organs or malleoli, which are
the prosoma.
probably sensory in function.
The Pseudoscorpiones are a fair-sized group,
Windscorpions are largely nocturnal, hiding dur-
with about 200 speeies in North America, and its
ing the day under objects or in burrows. They are fast-
members are common animals. They live under bark
and stones, in leaf litter and moss, between the boards running and predaceous, sometimes even capturing
small lizards. The pedipalps and first legs are used as
of buildings, and in similar situations. They some-
feelers, and these animals run on the last three pairs of
times cling to and are carried about by large insects.
legs. Two families occur in the United States, the Am-
They feed chiefly on small insects, which they catch
motrechidae (first legs without claws, and anterior
with their pedipalps. Most speeies have venom glands
edge of prosoma rounded or pointed) and the Eremo-
that open on the pedipalps. These animals have silk
batidae (first legs with one or two claws, and anterior
glands, the ducts from which open on the che-
edge of prosoma straight).
licerae.18The silk is used in making a cocoon in
which the animal overwinters.
Class Pycnogonida20-Sea Spiders
ORDERSolifugaeI9-Windscorpions: This is a fair-
sized group of arachnids (about 120 speeies in North The pycnogonids are marine, spiderlike forms with
America) whose members occur chiefly in the arid or long legs. They are occasionally found under stones
desert regions of the West (one speeies occurs in near the low-tide mark, but usually occur in deep wa-
ter. They are predaceous and have a sucking proboseis.
The body consists prineipally of prosoma; the opistho-
17Pseudoscorpiones: pseudo, false; scorpiones, scorpion. soma is very small. The sea spiders vary in length from
¡'Another name given 10 this order, Chelonethida, refers to this fea-
ture: chelo, claw; neth, spin.
¡'Solifugae: sol, sun; fugae, flee (referring to the nocturnal habits of 20Pycnogonida: pycno, thick or dense; gonida, offspring (referring to
these animals) the eggs).

136 Chapter5 PhylumArthropoda

one to several centimeters. Uttle is known of their for preserving gall mites. Such mites are best collected
habits, because they are uncommon. by wrapping infested plant parts in 50ft tissue paper
and allowing them to dry. This dried material can be
kept indefinitely, and the mites may be recovered for
study by warming the dried material in Kiefer's solu-
Collecting and Preserving Chelicerates tion (50 grams of resorcinal, 20 grams of diglycolic
acid, 25 milliliters of glycerol, enough iodine to pro-
To obtain a large collection of chelicerates, one needs duce the desired color, and about 10 milliliters of wa-
primarily to collect in as many different types of habi- ter). Specialislf on mites prefer specimens in fluid,
tats as possible. Chelicerates are frequently very abun- rather than mounted on permanent microscopic slides,
dant, often as abundant as insects or more so. The gen- so that all aspects and structure can be studied.
eral collector of insects is likely to encounter more Chelicerates can be collected by means of a net,
spiders and mites than any other types of chelicerates; forceps, vial, or small brush, or by hand. For biting or
therefore, the following suggestions are concerned pri- stinging forms, it is safer to use some method other
marily with these groups. than collecting them with the fingers. Specimens col-
Chelicerates occur in a great variety of situations lected with a net can be transferred directly to a vial of
and can often be collected with the same techniques alcohol, or collected in an empty vial and later trans-
and equipment used in collecting insects. Many can be ferred to alcohol. Because some species are quite active,
taken by sweeping vegetation with an insect net. Many it is some times preferable to put them first into a
can be obtained with beating equipment, that is, using cyanide bottle and transfer them to alcohol after they
a sheet or beating umbrella beneath a tree or bush and have been stunned and are quiet. Specimens collected
beating the bush to knock off the specimens. The from the ground or debris may be picked up with for-
ground forms can be found running on the ground or ceps or coaxed into a bottle; the smaller specimens
under stones, boards, bark, or other objects. Many can (found in any situation) may be picked up with a small
be found in the angles of buildings and similar pro- brush moistened with alcohol.
tected places. Many of the smaller forms can be found Spider webs that are flat and not too large can be
in debris, soillitter, or moss and are best collected by collected and preserved between two pieces of glass.
means of sifting equipment such as a Berlese funnel One piece of glass is pressed against the web (which
(Figure 35-5) or a screen sieve or by means of pitfall will usually stick to the glass beca use of the viscous
traps. Many are aquatic or semiaquatic and can be col- material on some of the silk strands), and then the
lected in marshy areas with aerial collecting equipment other piece of glass is applied to the first. It is often de-
or in water with aquatic equipment. The parasitic sirable to have the two pieces of glass separated by thin
species (various mites and ticks) usually must be strips of paper araund the edge of the glass. Once the
looked for on their hosts. web is between the two pieces of glass, bind the glass
Many chelicerates are nocturnal, and collecting at edges together with binding tape. Spider webs are best
night may prove more successful than collecting dur- photographed when covered with moisture (dew or
ing the day. Very few are attracted to lights, but they fog) or dust. They can often be photographed dry if il-
can be spotted at night with a flashlight or a headlamp. luminated fram the side and photographed against a
The eyes of many spiders reflect light, and with a little dark background.
experience and a light, you can locate many spiders at
night. Scorpions are fluorescent and can be collected at Subphylum CrustaceaZ1-Crustaceans
night with a portable ultraviolet light to make them
visible. The crustaceans are a large and varied group of arthro-
Chelicerates should be preserved in fluids rather pods, with more than 44,000 known species. Most of
than on pins or points. Many forms, such as the spi- them are marine, but many occur in fresh water, and a
few are terrestrial. In addition to the larger and more
ders, are very soft-bodied and shrivel when dry. They
are usually preserved in 70 to 80% alcohol. There familiar types, such as lobsters, crayfish, (Figure 5-22)
should be plenty of alcohol in the bottle in relation to crabs, and shrimp, a multitude of small to minute
the specimen, and it is often desirable to change the al- aquatic forms are very important in aquatic food webs.
cohol after the first few days. Many workers preserve The appendages and body regions vary greatly in this
mites in Oudeman's fluid, which consists of 87 parts of group, but typically there are two pairs of antennae, the
70% alcohol, 5 parts of glycerine, and 8 parts of glacial functional jaws consist of endite lobes of the gnathal (jaw)
acetic acid. The chief advantage of this fluid is that the appendages, and many of the appendages are biramous. A
mites die with their appendages extended so that sub- 2lCrustacea: from the Latin, referring to the crustlike exoskeleton
sequent examination is easier. Alcohol is not suitable possessed by many of these animals.
Collecting and PreservingChelicerates 137


I Figure5-22 A crayfish (Cambarus sp.),

natural size. ab, abdomen; ant, antenna;
,, I ¡
I 1
1 antl, antennule; chp, cheliped; crp, cara-
__ ' ',1/11/ 11 pace; e, eye; 1,legs (including cheliped);
sw, swimmerets; ur, uropod.

biramousappendage bears a process at its base (arising There are differences of opinion regarding the clas-
fromthe second segment of the appendage) that is more sification of these crustaceans, but four fairly distinct
or lessleglike, giving the appendage a two-branched ap- groups are usually recognized, the Anostraca, Noto-
pearance(Figure 5-1 C). There may be an exite lobe on straca, Conchostraca, and Cladocera. The first two of
thebasalsegment, as in trilobites, and in some cases this these are sometimes placed in a group called the Phyl-
functionsas a gil!. There are differences in this group in lopoda, and the last two are sometimes called the
thenature of the body regions. Sometimes there are two Diplostraca.
fairlydistinct (and nearly equal-sized) body regions, the The Anostraca,23 or fairy shrimps (Figure 5-23A),
cephalothoraxand the abdomen, with the cephalothorax have an elongated and distinctly segmented body, with-
bearing the antennae, gnathal appendages, and legs. out a carapace and with 11 pairs of swimming legs, and
Sometimesthe abdominal appendages occupy only a the eyes are on stalks. The fairy shrimps are often abun-
smallportion of the total body length. There is typically a dant in temporary pools. The Notostraca,24 or tadpole
terminaltelson. The cephalothorax in many crustaceans shrimps, have an oval convex carapace covering the an-
iscoveredby a shieldlike portion on the body wall called terior part of the body, 35 to 71 pairs of thoracic ap-
the carapace.The abdomen lacks paired appendages ex- pendages, and two long, filamentous caudal ap-
ceptin the Malacostraca. pendages. These animals range in size from about 10 to
The smaller crustaceans, particularly those in the 50 mm and live only in the westem states. The Con-
Branchiopoda, Copepoda, and Ostracoda, are abundant chostraca,25 or clam shrimps, have the body somewhat
in both salt and fresh water. The chief importance of flattened laterally and entirely enclosed in a bivalved
mostspecies is that they are food for larger animals and carapace and have 10 to 32 pairs of legs. Most species
thus are an important pan in the food webs leading to are 10 mm in length or less. The Cladocera,26 or water
fishand other larger aquatic animals. A few species are fleas (Figure 5-24A), have a bivalved carapace, but the
parasitic on fish and other animals, and the bamacles head is not enclosed in the carapace. There are four to
are often a nuisance when they encrust pilings, boat six pairs of thoracic legs. The water fleas are 0.2 to 3.0 mm
bottoms, and other surfaces. Many of the small crus- in length and are very common in freshwater pools.
taceanscan be easily maintained in indoor aquaria and Three groups of small crustaceans living in fresh wa-
arefrequently reared as food for other aquatic animals. ter have a bivalved carapace, and observers are likely to
confuse these groups. The Ostracoda (Figure 5-24B,C)
ClassBranchiopoda22 and Conchostraca have the body completely enclosed in
the carapace, whereas in the Cladocera (Figure 5-24A)
Most members of class Branchiopoda occur in fresh
the head is outside the carapace. The Ostracoda have
water. Males are uncommon in many species, and
only three pairs of thoracic legs; the Conchostraca have
parthenogensis is a common mode of reproduction.
10 to 32 pairs.
Both unisexual (parthenogenic) and bisexual repro-
duction occur in many species, and the factors control- 23 Anostraca:an, without; ostraca,shell.
lingthe production of males are not well understood. HNotostraca; not, back; ostraca, shell.
"Conchostraca: conch, shell or shellfish; ostraca, shell.
"Branchiopoda: branchio, gill; poda, foot or appendage '6Cladocera: dado, branch; cera, hom (referring to the antennae).
138 Chapter5 PhylumArthropoda

Figure5-23 Crustaceans. A, A fairy shrimp, Eubranchipus (subclass Branchiopoda, or-

der Anostraca), 6X; B, A female copepod, Cyclops sp., 50X, with two egg sacs at poste-
rior end of body.

Figure5-24 Crustaceans. A, A
water flea or cladoceran, Daphnia
sp., 25x; B, An ostracod, Cypri-
dopsis sp., lateral view; C, Same,
but with left valve of carapace re-
moved. ans, anus; ant, antenna;
antl, antennule; at, alimentary
tract; bp, brood pouch; cm, cae-
cum; e, compound eye;fu, furca;
hr, heart; 1, first and second tho-
racic legs; Ibr, labrum; md, mandi-
ble; mx, maxilla; pa, postabdomen;
sh, bivalved shell; thap, thoradc
appendages; y, developing young.
B e (C, Modified from Kesling.)
Collectingand PreservingChelicerates 139

ClassCopepoda27 bers of this group are hermaphroditic; that is, each in-
dividual has both male and female organs. Some bar-
Someof the copepods are free-swimming, and others are
nacles, such as the goose barnacle (Figure 5-25A), at-
parasiticon fish. The parasitic forms are often peculiar
tach the shell to some object by a stalk. Others, such as
in body fono and quite unlike the free-swimming forms the rock barnacle (Figure 5-25B), are sessile and do
in generalappearance. This group includes both marine not have stalks.
andfreshwater forms. The female of most copepods car-
riesher eggs in two egg sacs located laterally near the The Smaller Crustacean Classes
endof the abdomen (Figure 5-23B). The parasitic cope-
pods,often called fish lice, live on the gills or the skin or lncluded in the smaller crustaceans are the classes
burrow ioto the flesh of their host. When numerous, Cephalocarida,30 Mystacocarida,31 Branchiura,32 Tantu-
theymayseriously injure the host. Some species serve as locarida,33 and Remipedia.34 The nine known species of
intermediatehosts of certain human parasites for exam- Cephalocarida are marine bottom-dwelling forms that
pie, the fish tapeworm, Diphyllobothnum latum (L.). often occur in very deep water. The Mystacocarida are
minute (mostly about 0.5 mm in length) marine forms
ClassOstracoda28 living in the intertidal zone. Nine species of these have
been described. The Branchiura are ectoparasites on
Theostracodshave a bivalved carapace that can be closed the skin or in the gill cavities of fish (both marine and
bya muscle, and when the valves are closed the animal freshwater fish). About 130 species are known. The
lookslike a miniatureclam (Figure5-24B,C).When the Tantulocarida (four known species) are ectoparasites
valvesof the carapaceare open, the appendagesare pro- of deep-water crustaceans. The Remipedia (eight
trudedand propel the animal through the water. Many known species) have a long and wormlike body, and
speciesare parthenogenic. Most of the ostracods are ma- live in island caves connected to the sea.
rine,but there are also many common freshwater species.
Class Malacostraca35
The Malacostraca, the largest of the crustacean classes,
Thebest-known members of the class Cirripedia29 are includes the larger and better known forms, such as the
the bamacles, the adults of which live attached to
lobsters, crayfish, crabs, and shrimps. They differ from
rocks,pilings, seaweeds,boats, or marine animals, and
whichare enclosed in a calcareous shell. A few species
areparasitic, usually on crabs or mollusks. Most mem- 30Cephalocarida: cephalo, head; carida, a shrimp.
3lMystacocarida: mystaco, mustache; carida, a shrimp.
17Copepoda:cope, oar; poda, foot or appendage. 32Branchiura: branchí, gill; ura, tail.
"Ostracoda: from the Greek, meaning shel\-like (referring to the 33Tantulocarida: tantula, little; carida, a shrimp.
clarnlikecharacter of the carapace). 34Remipedia: remí, an oar; pedía, foot or appendage.
"Cirripedia: cirri, a cur\ of hair; pedía, foot or appendage. 35Malacostraca: malac, 50ft; ostraca, shell.

Figure5-25 Bamades. A, A goosebamade,

Lepassp., 3X; B, A rock bamade, Balanussp., 2X.
The basalstalk or pedunde of the goosebamacleis
B at the animal's posterior end; the biramous ap-
pendages protruding from the shell at the top of
the figure are the posterior thoracic legs; a second
small individual is shown attached to the first.

140 Chapter 5 PhylumArthropoda

the preceding classes (sometimes called the Ento- forms generally live under stones or among seaweed,
mostraca) in having appendages (swimmerets or where they are scavengers or omnivores, but a few are
pleopods) on the abdomen. There are typically 19 pairs wood-boring (apparendy feeding chiefly on the fungi
of appendages, the first 13 being cephalothoracic and in the wood), and some are parasitic on fish or other
the last 6 abdominal. The leglike appendages on the crustaceans. The most common isopods away from the
cephalothorax are often chelate. This class contains ocean are the sowbugs or woodlice-blackish, gray, or
some 13 orders. Only the more common ones can be brownish animals usually found under stones, boards,
mentioned here. or bark. Some sowbugs (often called pillbugs) can roll
ORDER Amphipoda:36The body of an amphipod is into a ball. In some areas sowbugs are important pests
elongate and more or less compressed; there is no cara- of cultivated plants.
pace; and seven (rarely six) of the thoracic segments ORDER Stomatopoda38-Mantis Shrimps: These are
are distinct and bear leglike appendages. The abdomi- predaceous marine forms, mosdy 5 to 36 cm in
nal segments are often more or less fused, and hence length, with the body dorsoventrally flattened. There
the six or seven thoracic segments make up most of the are three pairs of legs, in front of which are five pairs
body length (Figure 5-26). This group contains both of maxillipeds, the second of which is very large and
marine and freshwater forms. Many of them, such as chelate. A small cara pace covers the body in front of
the beach fleas (Figure 5-26B), live on the beach under the legs, and the abdomen is a litde wider than the
stones or in decaying vegetation. Most amphipods are carapace. The mantis shrimps are often brighdy co\-
scavengers. Some tropical species are common on the ored: green, blue, red, or patterned. This group is
floor of moist forests. principally tropical. U.5. species occur chiefly along
ORDERIsopoda:37The isopods are similar to the the southern coasts.
amphipods in lacking a carapace, but are dorsoven- ORDER Decapoda:39This order contains the largest
trally flattened. The last seven thoracic segments are and probably the best known of the crustaceans, the
distinct and bear leglike appendages. The abdominal lobsters, crayfish (Figure 5-22), crabs (Figure 5-28),
segments are more or less fused, and hence the thoracic and shrimps. The cara pace of a decapod covers the en-
segments (with their seven pairs of legs) make up most tire thorax. Five pairs of the cephalothoracic ap-
of the body length (Figure 5-27). The anterior abdom- pendages are leglike, and the first pair of these usually
inal appendages of the aquatic forms usually bear gills. bears a large claw. The abdomen may be well developed
The terminal abdominal appendages are often enlarged Clobsters and crayfish) or much reduced (crabs). This
and feelerlike. The isopods are small (most are less is a very important group, beca use many of its mem-
than 20 mm in length), and most are marine, but some bers are used as food, and their collection and distri-
live in fresh water and some are terrestrial. The marine bution underpin a large coastal industry.

3<>Amphipoda:amphi, on both sides, double; poda, foot or appendage. 3.Stomatopoda: stomato, mouth; poda, foot or appendage.
"Isopoda: iso, equal; poda, foot or appendage. 39Decapoda: deca, ten; poda, foot or appendage.

Figure 5-26 Amphipods.

A, A common freshwater scud,
Dikerogammarusfasciatus (Say),
10-15 mm in length; B, A sand
flea or beach flea, Orchestia agilis
Smith, abundant underneath sea-
weed along the coast near the high
tide mark; C, A common freshwa-
ter scud, Hyalella knickerbockeri
(Bate), about 7 mm in length;
D, A sea scud, Gammarus annula-
tus Smith, a common coastal form,
about 15 mm in length. (Courtesy
of Kunkel and the Connecticut
State Geology and Natural History
e Survey; C, After Smith.)
Collectingand PreservingCrustacea 141

A B e

Figure 5-27 Isopods. A, Oniscus asellus L., a common sowbug; B, Asellus communis
Say,a common freshwater isopod; C, Cylisticus convexus (DeGeer), a pillbug capable of
rolling itself into a ball. (Courtesy of the Connecticut State Geology and Natural History
Survey. A, C, Courtesy of Kunkel 1918, after Paulmier; B, Courtesy of Kunkell918,
after Smith.)

Figure5-28 A green crab, Carcinides

sp., P/2X.

Collectingand Preserving Crustacea towed by a boato Many larger forms are collected by traps.
Such traps (or "pots") are the standard means of collect-
Theaquatic crustaceans must be collected with various ing lobsters and crabs. The shore-dwelling and terrestrial
typesof aquatic collecting equipmenl. Most can be col- forms can be collected by hand or forceps or possibly (for
lected by a dip nel. A white enamel dipper is the best example, beach fleas) with an aerial insect neto Handle
meansof collecting many of the smaller forms. The dip- the larger forms with well-developed claws with care, be-
per is simply dipped into the water, and any small ani- cause the claw can inflict serious injury. The safest way to
malsin the dipper can be easily seen. Forms so collected pick up a large crayfish or lobster is from above, grasping
canbe removed by means of an eye dropper or (if fairly the animal at the back of the carapace.
large)by forceps. The smaller forms in ponds, lakes, and To obtain a variety of Crustacea, collect in a vllri-
the ocean are often collected in afine-mesh plankton net, ety of places. When collecting in water, investiga te
142 Chapter5 PhylumArthropoda

every possible aquatic niche. Some crustaceans are ies have done little to alleviate the considerable uncer-
free-swimming; some burrow in bottom mud; some tainty concerning the interrelationships among Crus-
live under stones; and many are found on aquatic veg- tacea, Myriapoda, and Hexapoda. Some researchers
etation. The shore-dwelling forms are usually under suggest that the myriapods are a polyphyletic or para-
stones, debris, or decaying vegetation along the shore. phyletic group; others that hexapods are most closely
Preserve crustaceans in fIuids (for example, 70 to related to a subset of crustaceans (and, therefore, that
95% alcohol). Most smaller forms must be mounted on Crustacea is paraphyletic).
microscope slides for detailed study. Some of the
Class Diplopoda41-Millipedes. The millipedes are
smaller Malacostraca can be preserved dry (for exam-
elongatt:, wormlike animals with many legs (Fig-
pIe, pinned), but specimens preserved in fIuid are more ure 5-29). Most millipedes have 30 or more pairs of
satisfactory for study.
legs, and most body segments bear 2 pairs. The body
is cylindrical or slightly fIattened, and the antennae
Subphylum Atelocerata40
are short and usually seven-segmented. The external
The members of this subphylum have a single pair of openings of the reproductive system are located at
antennae and uniramous (unbranched) appendages. the anterior end of the body, between the second and
According to Manton (1977), the mandibles of these third pairs of legs. One or both pairs of legs on the
species differ from those of Crustacea in that the entire seventh segment of the male are usually modified
appendage makes up the functional portion of the into gonopods, which function in copulation. Com-
mandible (and not only the basal portion). She placed pound eyes are usually present. The first tergum be-
the groups included here (following) together with the hind the head is usually large and is called the collum
Onychophora as the phylum Uniramia (concluding (Figure 5-30A).
that the Arthropoda is a polyphyletic taxon). We be- The head in most millipedes is convex above,
lieve this position has not been adequately supported. with a large epistomal area, and fIat beneath. The
In terms of her conclusion regarding the structure of bases of the mandibles form a part of the side of the
the mandible, both Crustacea and Hexapoda have sim- head. Beneath the mandibles, and forming the fIat
ilar expression patterns of the gene distal-less, falsifying ventral surface of the head, is a characteristic liplike
the hypothesis that one (the crustacean mandible) is structure called the gnathochilarium (Figure 5-30B,
gnathobasic, whereas the other represents the entire gna). The gnathochilarium is usually divided by su-
appendage of the mandibular segment. Further, Manton's tures into several areas: a median more or less trian-
conclusion requires evidence that one or more taxa gular plate, the mentum (mn); two laterallobes, the
now classified as arthropods are more closely related to stipites (stp); two median distal plates, the laminae
a nonarthropod group. Until such evidence is available, linguales (1l); and usually a median transverse basal
we continue to treat the Arthropoda as a monophyletic sclerite, the prebasalare (pbs), and two smalllatero-
unit and do not consider the onychophorans to be basal sclerites, the cardines (cd). The size and shape
arthropods. of these areas differ among groups of millipedes, and
Other than Onychophora, the Atelocerata includes the gnathochilarium often provides characters by
the Myriapoda and Hexapoda. Recent molecular stud- which the groups are recognized.

"'Atelocerata: ate/os, defective; keras, horn; referring to the fact that the ilDiplopoda: diplo, two; poda, foot or appendage (referring to the
second antennae are present in these taxa only as embryonic rudiments. fact that most body segments bear two pairs of legs).

Figure5-29 A common millipede, Nar-

ceus sp. (order Spirobolida), P/2x.
Keyto the Ordersof Diplopoda 143

material, but some attack living plants and sometimes

do serious damage in greenhouses and gardens, and a
few are predaceous. These animals overwinter as adults
in protected situations and lay their eggs during the
summer. Some construct nestlike cavities in the soil in
which they deposit their eggs; others lay their eggs in
damp places without constructing any sort of nest. The
eggs are usually white and hatch within a few weeks.
Newly hatched millipedes have only three pairs of legs.
The remaining legs are added in subsequent molts.
B There are a number of arrangements of orders and
families in this group. We follow the arrangement of
Chamberlin and Hoffman (1958), which is outlined
here (with alterna te names or arrangements in paren-
Subclass pselaphognatha (Pencillata)
5-30 Head structure in a millipede (Nareeus, Order Polyxenida
orderSpirobolida). A, Lateral view of head; B, Gnatho- Subclass Chilognatha
chilarium.ant, antenna; ed, cardo; eolm, collum, tergite Superorder Pentazonia (Opisthandria)
ofthe first body segment; e, eye; gna, gnathochilarium; Order Glomerida (Oniscomorpha)
1,firstleg; lbr, labrum; 11,lamina lingualis; md, mandible; Superorder Helminthomorpha (Olognatha,
mn,mentum; pbs, prebasilare; stn, sternum of first body Eugnatha)
segment;stp, stipes. Order Polydesmida (Proterospermophora)
Order Chordeumida (Chordeumatida,
Order Julida (Opisthospermophora in part)
Millipedes are usually found in damp places: un- Order Spirobolida (Opisthospermophora
der leaves, in moss, under stones or boards, in rotting in part)
wood, or in the soil. Many species can give off an ill- Order Spirostreptida (Opisthospermophora
smellingfluid through openings along the sides of the in part)
body.This fluid is some times strong enough to kill in- Order Cambalida (Opisthospermophora
sectsthat are placed in ajar with the millipede, and it in part)
hasbeen shown (in some cases at least) to contain hy- Superorder Colobognatha
drogen cyanide. Millipedes do not bite people. Most Order Polyzoniida
millipedes are scavengers and feed on decaying plant Order Platydesmida

Keyto the Ordersof Diplopoda

1. Adults with 13 pairs of legs; integument soft; body hairs forming long
lateral tufts; 2-4 mm in length Polyxenida p.144
l' Adults with 28 or more pairs of legs; integument strongly sclerotized;
body hairs not forming long tufts; larger millipedes 2
2(1'). Body with 14-16 segments and with 11-13 tergites; male gonopods at
caudal end of body, modified from last 2 pairs of legs; southern and
western United States Glomerida p. 144
2' Body with 18 or more segments; male gonopods modified from legs on
seventh segment 3
3(2'). Head small, often concealed, the mandibles much reduced; 8 pairs of
legs anterior to the male gonopods 4
3'. Head and mandibles of normal size; 7 pairs of