The Insects

Structure and Function

4th edition

R. F. Chapman


The Pitt Building, Trumpington Street, Cambridge, United Kingdom

The Edinburgh Building, Cambridge CB2 2RU, UK
40 West 20th Street, New York, NY 10011-4211, USA
477 Williamstown Road, Port Melbourne, VIC 3207, Australia
Ruiz de Alarcon
13,28014 Madrid, Spain
Dock House, The Waterfront, Cape Town 8001, South Africa
http://www.cambridge.org
Cambridge University Press 1998
This book is in copyright. Subject to statutory exception
and to the provisions of relevant collective licensing agreements,
no reproduction of any part may take place without
the written permission of Cambridge University Press.
First published by Edward Arnold 1969
Second edition 1971, reprinted 1972, 1974, 1975, 1978, 1980
Third edition 1982, reprinted 1983, 1985, 1988, 1991
Fourth edition published by Cambridge University Press 1998
Reprinted 2003
Printed in the United Kingdom at the University Press, Cambridge
Typeset in MT Ehrhardt 9/12pt []
A catalogue record for this book is available from the British Library
Library of Congress Cataloguing in Publication data
Chapman, R.F. (Reginald Frederick)
The insects : structure and function / R.F. Chapman. 4th ed.
p. cm.
Includes indexes.
ISBN 0 521 57048 4 (hb). ISBN 0 521 57890 6 (pb)
1. Insects. I. Title.
QL463.C48 1998
595.7 dc21 97-35219 CIP
ISBN 0 521 57048 4 hardback
ISBN 0 521 57890 6 paperback

Contents

Preface xi
Acknowledgments xiii

PART I

The Head, Ingestion, Utilization and Distribution of Food

1 Head 3
1.1 Head 3
1.2 Neck 6
1.3 Antennae 8
References 11

2 Mouthparts and feeding 12

2.1 Ectognathous mouthparts 12
2.2 Mechanics and control of feeding 18
2.3 Regulation of feeding 26
2.4 The consequences of feeding 30
2.5 Head glands 30
References 36

3 Alimentary canal, digestion and absorption 38

3.1 Alimentary canal 38
3.2 Passage of food through the gut 51
3.3 Digestion 51
3.4 Absorption 60
3.5 Efficiency of digestion and absorption 65
References 66
4 Nutrition 69
4.1 Nutritional requirements 69
4.2 Balance of nutrients 78
4.3 Feeding on nutritionally poor substrates 82
4.4 Nutritional effects on growth and development
References 91

89

5 Circulatory system, blood and immune systems 94

5.1 Circulatory system 94
5.2 Hemolymph 106
5.3 Immunity 121
5.4 Connective tissue 126
References 127
[v]

vi

CONTENTS

6 Fat body 132

6.1 Structure 132
6.2 Functions 135
References 141
PART II

The Thorax and Locomotion

7 Thorax 145
7.1 Segmentation 145
7.2 Thorax 146
References 150

8 Legs and locomotion 151

8.1 Basic structure of the legs 151
8.2 Modifications of the basic leg structure
8.3 Maintenance of stance 160
8.4 Locomotion 160
8.5 Locomotion in aquatic insects
References 182

157

174

9 Wings and flight 185

9.1
9.2
9.3
9.4
9.5
9.6
9.7
9.8
9.9
9.10
9.11
9.12

Occurrence and structure of wings 185

Modifications of the wings 190
Wing coupling 192
Articulation of the wings with the thorax 194
Sensilla on the wings and the halteres 195
Muscles associated with the wings 196
Mechanisms of wing movement 198
Movements of the wings 202
Aerodynamics 206
Control of wingbeat 210
Stability in flight 214
Power for flight 218

References

225

10 Muscles 229
10.1 Structure 229
10.2 Changes during development 239
10.3 Muscle contraction 244
10.4 Energetics of muscle contraction 249
10.5 Muscular control in the intact insect 251
References 253

PART III

The Abdomen, Reproduction and Development

11 Abdomen 259
11.1 Segmentation 259
11.2 Abdominal appendages and outgrowths
References 267

261

vii

CONTENTS

12 Reproductive system: male 268

12.1 Anatomy of the internal reproductive organs
12.2 Spermatozoa 270
12.3 Transfer of sperm to the female 276
12.4 Other effects of mating 288
12.5 Sperm capacitation 291
References 292

268

13 Reproductive system: female 295

13.1 Anatomy of the internal reproductive organs
13.2 Oogenesis 298
13.3 Ovulation 312
13.4 Fertilization of the egg 313
13.5 Oviposition 313
References 321

295

14 The egg and embryology 325

14.1 The egg 325
14.2 Embryology 332
14.3 Viviparity 351
14.4 Polyembryony 355
14.5 Parthenogenesis 356
14.6 Pedogenesis 358
References 359
15 Postembryonic development 363
15.1 Hatching 363
15.2 Larval development 365
15.3 Metamorphosis 378
15.4 Control of postembryonic development
15.5 Polyphenism 400
15.6 Diapause 403
References 408

PART IV

394

The Integument, Gas Exchange and Homeostasis

16 Integument 415
16.1 Epidermis 415
16.2 Basic structure of cuticle 417
16.3 Different types of cuticle 422
16.4 Molting 427
16.5 Cuticle formation 432
16.6 Expansion of the new cuticle 434
16.7 Changes in the intermoult cuticle 435
16.8 Functions of the cuticle 438
References 438

17 Gaseous exchange 441

17.1 Tracheal system 441
17.2 Spiracles 448

viii

CONTENTS

17.3 Cutaneous respiration 452

17.4 Gaseous exchange in terrestrial insects 452
17.5 Gaseous exchange in aquatic insects 461
17.6 Insects subject to occasional submersion 469
17.7 Gas exchange in endo-parasitic insects 472
17.8 Other functions of the tracheal system 473
17.9 Respiratory pigments 473
References 475

18 Excretion and salt and water regulation 478

18.1 Excretory system 478
18.2 Nitrogenous excretion 481
18.3 Urine production 484
18.4 Water regulation 491
18.5 Non-excretory functions of the Malpighian tubules
18.6 Nephrocytes 501
18.7 Detoxification 502
References 505

501

19 Thermal relations 509

19.1 Body temperature 509
19.2 Thermoregulation 514
19.3 Behavior and survival at low temperatures 517
19.4 Activity and survival at high temperatures 522
19.5 Acclimation 523
16.6 Cryptobiosis 523
19.7 Temperature and humidity receptors 524
19.8 Temperature-related changes in the nervous system
References 528
PART V

A.

527

Communication

Physiological Co-ordination Within the Insect

20 Nervous system 533

20.1 Basic components 533
20.2 Basic functioning 537
20.3 Anatomy of the nervous system
20.4 Brain 550
20.5 Controlling behavior 560
References 566

546

21 Endocrine system 570

21.1 Chemical structure of hormones
21.2 Endocrine organs 571
21.3 Transport of hormones 578
21.4 Regulation of hormone titer 578
21.5 Mode of action of hormones 580
References 582

570

ix

CONTENTS

B.

Perception of the Environment

22 Vision 587
22.1 Compound eyes 587
22.2 Functioning of the eye 592
22.3 Vision 600
22.4 Dorsal ocelli 603
22.5 Stemmata 605
22.6 Other visual receptors 606
References 607

23 Mechanoreception 610
23.1 Cuticular mechanoreceptors

610

23.2 Chordotonal organs 617

23.3 Stretch receptors 629
References 633

24 Chemoreception 636
24.1 Olfaction 636
24.2 Contact chemoreception
References 652

C.

645

Communication with Other Organisms

25 Visual signals: color and light production 657

25.1 The nature of color 657
25.2 Physical colors 657
25.3 Pigmentary colors 660
25.4 The colors of insects 665
25.5 Color change 665
25.6 Significance of color 671
25.7 Light production 674
References 678
26 Mechanical communication: producing sound and substrate vibrations 680
26.1 Mechanisms producing vibrations 680
26.2 Signal transmission 692
26.3 Patterns of vibrational signals 692
26.4 Neural regulation of sound production 696
26.5 Significance of vibrational signals 698
References 701
27 Chemical communication: pheromones and chemicals with interspecific significance 704
27.1 Pheromones 704
27.2 Secretions with interspecific significance 725
References 736
Taxonomic index 741
Subject index 749

Head

Insects and other arthropods are built up on a segmental

plan and their characteristic feature is a hard, jointed
exoskeleton. The cuticle, which forms the exoskeleton, is
continuous over the whole of the outside of the body and
consists of a series of hard plates, the sclerites, joined to
each other by flexible membranes, which are also cuticular.
Sometimes the sclerites are articulated together so as to
give precise movement of one on the next. Each segment
of the body primitively has a dorsal sclerite, the tergum,
joined to a ventral sclerite, the sternum, by lateral membranous areas, the pleura. Arising from the sternopleural
region on each side is a jointed appendage.
In insects, the segments are grouped into three units,
the head, thorax and abdomen, in which the various basic
parts of the segments may be lost or greatly modified.
Typical walking legs are only retained on the three thoracic
segments. In the head, the appendages are modified for
sensory and feeding purposes and in the abdomen they are
lost, except that some may be modified as the genitalia and
in Apterygota some pregenital appendages are retained.

1.1 HEAD

The insect head is a strongly sclerotized capsule joined to

the thorax by a flexible membranous neck. It bears the
mouthparts, comprising the labrum, mandibles, maxillae
and labium, and also important sense organs, the antennae,
compound eyes and ocelli. On the outside it is marked by
grooves most of which indicate ridges on the inside, and
some of these inflexions extend deep into the head, fusing
with each other to form an internal skeleton. These structures serve to strengthen the head and provide attachments for muscles as well as supporting and protecting the
brain and foregut.
The head is derived from the primitive pre-oral
archecerebrum and a number of primitively post-oral segments. Molecular studies of Drosophila suggest that there
are seven postoral segments: labral, ocular, antennal, intercalary, mandibular, maxillary and labial (Schmidt-Ott et
al., 1994). The last three segments are often called the

gnathal segments because their appendages form the

mouthparts of the insect.
Reviews: Bitsch, 1973; Denis & Bitsch, 1973; DuPorte, 1957;
Matsuda, 1965; Snodgrass, 1935, 1960
1.1.1 Orientation
The orientation of the head with respect to the rest of the
body varies (Fig. 1.1). The hypognathous condition, with
the mouthparts in a continuous series with the legs, is
probably primitive. This orientation occurs most commonly in phytophagous species living in open habitats. In
the prognathous condition the mouthparts point forwards
and this is found in predaceous species that actively pursue
their prey, and in larvae, particularly of Coleoptera, which
use their mandibles in burrowing. In Hemiptera, the elongate proboscis slopes backwards between the forelegs.
This is the opisthorhynchous condition.
The mouthparts enclose a cavity, the pre-oral cavity,
with the mouth at its inner end (Fig. 1.2). The part of the
pre-oral cavity enclosed by the proximal part of the hypopharynx and the clypeus is known as the cibarium. Between
the hypopharynx and the labium is a smaller cavity known
as the salivarium, into which the salivary duct opens.
1.1.2 Rigidity
The head is a continuously sclerotized capsule with no
outward appearance of segmentation, but it is marked by a
number of grooves. Most of these grooves are sulci (singular: sulcus), marking lines along which the cuticle is
inflected to give increased rigidity. The term suture should
be retained for grooves marking the line of fusion of two
formerly distinct plates. The groove which ends between
the points of attachment of maxillae and labium at the back
of the head is generally believed to represent the line of
fusion of the maxillary and labial segments and it is therefore known as the postoccipital suture.
Since the sulci are functional mechanical developments
to resist the various strains imposed on the head capsule,
they are variable in position in different species and any
one of them may be completely absent. However, the needs

[3]

HEAD

a) hypognathous
brain
frons
pharynx

antenna

salivary duct

mandible
maxillary
palp

mesothoracic
leg

labial
palp

prothoracic
leg

pharyngeal
dilator
muscles

subesophageal
ganglion

frontal ganglion

suspensory
sclerite

mouth
salivarium
cibarium

b) prognathous

lingual
sclerite

clypeus

antenna

epipharynx

maxillary
palp

labial
palp
prothoracic
leg

c) opisthorhynchous
antenna

prothoracic
leg

labium

labrum

mandible

proboscis

mesothoracic
leg

Fig. 1.1. Orientation of the head. (a) Hypognathous

mouthparts ventral, in a continuous series with the legs
(grasshopper). (b) Prognathous mouthparts in an anterior
position (beetle larva). (c) Opisthorhynchous sucking
mouthparts with the proboscis extending back between the front
legs (aphid).

for strengthening the head wall are similar in the majority

of insects, so some of the sulci are fairly constant in occurrence and position (Fig. 1.3). The most constant is the
epistomal (frontoclypeal) sulcus, which acts as a brace

hypopharynx

Fig. 1.2. Pre-oral cavity and some musculature. Diagrammatic

vertical section through the head of an insect with biting and
chewing mouthparts. Sclerites associated with the hypopharynx
are black with white spots. Muscles attached to these sclerites
move the hypopharynx (after Snodgrass, 1947).

between the anterior mandibular articulations. At each end

of this sulcus is a pit, the anterior tentorial pit, which
marks the position of a deep invagination to form the anterior arm of the tentorium. The lateral margins of the head
above the mandibular articulations are strengthened by a
horizontal inflexion indicated externally by the subgenal
sulcus. This sulcus is generally a continuation of the epistomal sulcus to the postoccipital suture. The part of the
subgenal sulcus above the mandible is called the pleurostomal sulcus, the part behind the mandible is the hypostomal sulcus. Another commonly occurring groove is the
circumocular sulcus, which strengthens the rim of the eye
and may develop into a deep flange protecting the inner
side of the eye. Sometimes this sulcus is connected to the
subgenal sulcus by a vertical subocular sulcus; the
inflexions associated with these sulci act as a brace against
the pull of the muscles associated with feeding. The circumantennal ridge, marked by a sulcus externally,
strengthens the head at the point of insertion of the
antenna, while running across the back of the head, behind
the compound eyes, is the occipital sulcus.
The areas of the head defined by the sulci are given
names for descriptive purposes, but they do not represent
primitive sclerites. Since the sulci are variable in position, so

HEAD

a) anterior

b) lateral
occipital sulcus

ecdysial line
vertex

ecdysial line

occiput
postoccipital
suture
postocciput
neck
membrane

circumocular sulcus
gena

postgena

circumantennal sulcus
frons

subocular sulcus

frons

gena

anterior tentorial pit

clypeus
anterior
articulation of
mandible

labrum

subgenal
sulcus
posterior
tentorial pit
subgena

epistomal sulcus

labium

mandible
maxilla

Fig. 1.3. Common lines or grooves on the insect head and the areas which they define (italicized) (modified after Snodgrass,
1960).

too are the areas which they delimit. The front of the head,
the frontoclypeal area, is divided by the epistomal sulcus into
the frons above and the clypeus below (Fig. 1.3). It is
common to regard the arms of the ecdysial cleavage line as
delimiting the frons dorsally, but this is not necessarily so
(Snodgrass, 1960). From the frons, muscles run to the
pharynx, the labrum and the hypopharynx; from the clypeus
arise the dilators of the cibarium. The two groups of muscles
are always separated by the frontal ganglion and its connectives to the brain (Fig. 1.2). Dorsally the frons continues into
the vertex and posteriorly this is separated from the occiput
by the occipital sulcus. The occiput is divided from the postocciput behind it by the postoccipital suture, while at the
back of the head, where it joins the neck, is an opening, the
occipital foramen, through which the alimentary canal,
nerve cord and some muscles pass into the thorax.
The lateral area of the head beneath the eyes is called
the gena, from which the subgena is cut off below by the
subgenal sulcus, and the postgena behind by the occipital
sulcus. The region of the subgena above the mandible is
called the pleurostoma and that part behind the mandible
is the hypostoma.
In hypognathous insects with a thick neck, the posterior ventral part of the head capsule is membranous. The
postmentum of the labium is contiguous with this

membrane, articulating with the subgena on either side.

The hypostomal sulci bend upwards posteriorly and are
continuous with the postoccipital suture (Fig. 1.4a). In
insects with a narrow neck, permitting greater mobility of
the head, and in prognathous insects, the cuticle of the
head below the occipital foramen is sclerotized. This
region has different origins. In Diptera, the hypostomata
of the two sides meet in the midline below the occipital
foramen to form a hypostomal bridge which is continuous
with the postocciput (Fig. 1.4b). In other cases,
Hymenoptera and the water bugs Notonecta and Naucoris,
a similar bridge is formed by the postgenae, but the bridge
is separated from the postocciput by the postoccipital
suture (Fig. 1.4c). Where the head is held in the prognathous position, the lower ends of the postocciput fuse
and extend forwards to form a median ventral plate, the
gula (Fig. 1.4d), which may be a continuous sclerotization
with the labium. Often the gula is reduced to a narrow strip
by enlargement of the postgenae and sometimes the postgenae meet in the midline, so that the gula is obliterated.
The median ventral suture which is thus formed at the
point of contact of the postgenae is called the gular suture.
In all insects, the rigidity of the head is increased by
four deep cuticular invaginations, known as apodemes,
which usually meet internally to form a brace for the head

HEAD

Fig. 1.4. Sclerotization at the back of

the head. Notice the position of the
bridge below the occipital foramen
with reference to the posterior
tentorial pit. Membranous areas
stippled, compound eyes crosshatched. The names of areas defined
by sulci are italicized (after Snodgrass,
1960). (a) Generalized condition, no
ventral sclerotization; (b) hypostomal
bridge (Deromyia, Diptera); (c)
postgenal bridge (Vespula,
Hymenoptera); (d) gular bridge
formed from the postoccipital sclerites
(Epicauta, Coleoptera).

a) generalized condition

b) hypostomal bridge

postoccipital
suture
postocciput
posterior
tentorial pit

occipital
foramen

postgena
hypostomal
sulcus

postmentum

hypostomal
bridge

subgena

prementum

maxilla

postmentum

mandible

d) gula
c) postgenal bridge

postoccipital
suture
postocciput
posterior
tentorial pit
postgena
hypostomal
sulcus

postgenal
bridge

occipital
foramen

gula
submentum

subgena
labium

mentum

maxilla
mandible

and for the attachment of muscles. The structure formed

by these invaginations is called the tentorium (Fig. 1.5).
Its two anterior arms arise from the anterior tentorial pits,
which in Apterygota and Ephemeroptera are ventral and
medial to the mandibles. In Odonata, Plecoptera and
Dermaptera the pits are lateral to the mandibles, while in
most higher insects they are facial at either end of the epistomal sulcus. The posterior arms arise from pits at the
ventral ends of the postoccipital suture and they unite to
form a bridge running across the head from one side to the
other. In Pterygota the anterior arms also join up with the
bridge, but the development of the tentorium as a whole is
very variable. Sometimes a pair of dorsal arms arise from
the anterior arms and they may be attached to the dorsal
wall of the head by short muscles. In Machilidae
(Archaeognatha) the posterior bridge is present, but the
anterior arms do not reach it, while in Lepismatidae
(Thysanura) the anterior arms unite to form a central
plate near the bridge and are joined to it by very short
muscles.

1.1.3 Molting

Immature insects nearly always have a line along the dorsal

midline of the head dividing into two lines on the face so as to
form an inverted Y (Fig. 1.3). There is no groove or ridge
along this line, and it is simply a line of weakness, continuous
with that on the thorax, along which the cuticle splits when
the insect molts (see Fig. 16.11). It is therefore called the
ecdysial cleavage line, but has commonly been termed the
epicranial suture. The anterior arms of this line are very variable in their development and position and, in Apterygota,
they are reduced or absent. The ecdysial cleavage line may
persist in the adult insect and sometimes the cranium is
inflected along this line to form a true sulcus. Other ecdysial
lines may be present on the ventral surface of the head of
larval insects (Hinton, 1963).
1.2 NECK

The neck or cervix is a membranous region which gives

freedom of movement to the head. It extends from the postocciput at the back of the head to the prothorax, and possibly

NECK

occiput
postoccipital ridge
postocciput

dorsal arm of
tentorium

Fig. 1.5. Tentorium. Cutaway of the head capsule to show

the tentorium and its relationship with the grooves and
ridges of the head (after Snodgrass, 1935).

postoccipital
suture
tentorial
bridge

subgenal
ridge
anterior arm of
tentorium

posterior
tentorial pit

epistomal
sulcus

subgenal
sulcus
anterior
tentorial pit

clypeus
labrum

epistomal ridge

a) musculature
head

prothorax

neck
membrane

postocciput

occipital
condyle

second cervical
sclerite

third cervical
sclerite

b) head movement
head retracted

head

head protracted

prothorax

head

neck membrane
folded
cervical
sclerites

cervical
sclerites

prothorax
neck membrane
extended

Fig. 1.6. Neck and cervical sclerites of a grasshopper.

(a) Seen from the inside to show the muscles (after
Imms, 1957). (b) Diagrams showing how a change in
the angle between the second and third cervical
sclerites retracts or protracts the head. (The first
cervical sclerite is small and is not shown). Arrows
indicate points of articulation.

HEAD

a) annulated
flagellum

b) segmented
pedicel

flagellum

scape
extensor
muscle

pedicel

annuli
flexor
muscles

levator
muscle

depressor
muscle

scape
intrinsic muscles
of flagellar
segments
flexor
muscles

extensor
muscles

levator
muscle
Fig. 1.7. Antenna. Proximal region showing the musculature. (a) Typical insect annulated antenna. There are no muscles in
the flagellum (Locusta, Orthoptera). (b) Segmented antenna of a non-insect hexapod (Japyx, Diplura) (after Imms, 1940).

represents the posterior part of the labial segment together

with the anterior part of the prothoracic segment. Laterally
in the neck membrane are the cervical sclerites. Sometimes
there is only one, as in Ephemeroptera, but there may be two
or three. In Schistocerca (Orthoptera) the first lateral cervical
sclerite, which articulates with the occipital condyle at the
back of the head, is very small. The second sclerite articulates
with it by a ball and socket joint allowing movement in all
planes. Posteriorly it meets the third (posterior) cervical sclerite and movement at this joint is restricted to the vertical
plane. The third cervical sclerite connects with the prothoracic episternum, relative to which it can move in all planes.
Muscles arising from the postocciput and the pronotum are inserted on the cervical sclerites (Fig. 1.6a) and
their contraction increases the angle between the sclerites
so that the head is pushed forwards (Fig. 1.6b). A muscle
arising ventrally and inserted on to the second cervical
sclerite may aid in retraction or lateral movements of the
head. Running through the neck are longitudinal muscles,
dorsal muscles from the antecostal ridge of the mesothorax
to the postoccipital ridge, and ventral muscles from the
sternal apophyses of the prothorax to the postoccipital
ridge or the tentorium. These muscles serve to retract the
head on to the prothorax, while their differential contraction will cause lateral movements of the head. Schistocerca
has 16 muscles on each side of the neck, each of which is
innervated by several axons, often including an inhibitory

fiber. This polyneuronal innervation, together with the

versatility of the cervical articulations and the complexity
of the musculature, permits movement of the head in a
highly versatile and accurately controlled manner.

1.3 ANTENNAE

All insects possess a pair of antennae, but they may be

greatly reduced, especially in larval forms. Amongst the
non-insectan Hexapoda, Collembola and Diplura have
antennae, but Protura do not.
1.3.1 Antennal structure

The antenna consists of a basal scape, a pedicel and a

flagellum. The scape is inserted into a membranous region
of the head wall and pivoted on a single marginal point, the
antennifer (Fig. 1.8a), so it is free to move in all directions.
Frequently the flagellum is divided into a number of similar
annuli joined to each other by membranes so that the
flagellum as a whole is flexible. The term segmented should
be avoided with reference to the flagellum of insects since
the annuli are not regarded as equivalent to leg segments.
The antennae of insects are moved by levator and
depressor muscles arising on the anterior tentorial arms and
inserted into the scape, and by flexor and extensor muscles
arising in the scape and inserted into the pedicel (Fig. 1.7a).
There are no muscles in the flagellum, and the nerve which

ANTENNAE

a) filiform

b) capitate

pedicel

c) lamellate

scape

Fig. 1.8. Antennae. Different forms

occurring in different insects. Not all
to same scale.

flagellum

antennifer

flagellum
part of
head
capsule

pedicel
pedicel
scape

d) cyclorraphous
Diptera

scape

scape

e) Lepidopteran
larva

pedicel
arista
basal
annulus

pedicel
scape

traverses the flagellum is purely sensory. This is the annulated type of antenna. In Collembola and Diplura the musculature at the base of the antenna is similar to that in
insects, but, in addition, there is an intrinsic musculature in
each unit of the flagellum (Fig. 1.7b), and, consequently,
these units are regarded as true segments.
The number of annuli is very variable between species.
Adult Odonata, for example, have five or fewer annuli
while adult Periplaneta have over 150, increasing from
about 48 in the first stage larva.
The form of the antenna varies considerably depending on its precise function (Fig. 1.8). Sometimes the
modification produces an increase in surface area allowing
a large number of sensilla to be accommodated on the
antenna (Fig. 1.9) and, in the case of the plumose antennae
of some male moths, enabling them to sample a large
volume of air. Sexual dimorphism in the antennae is
common, those of the male often being more complex than
those of the female. This often occurs where the male is
attracted to or recognizes the female by her scent.
Conversely, in chalcids scent plays an important part in
host-finding by the female and in this case the females
antennae are more specialized than the males.
The antennae of larval hemimetabolous insects are
similar to those of the adult, but with fewer annuli. The

number increases at each molt (see Fig. 15.10). In

Periplaneta, for example, there are only 48 annuli in the
first stage larva compared with over 150 in the adult. The
antennae of larval holometabolous insects are usually
considerably different from those of the adult. The larval
antennae of Neuroptera and Megaloptera have a number
of annuli, but in larval Coleoptera and Lepidoptera the
antennae are reduced to three simple segments. In some
larval Diptera and Hymenoptera the antennae are very
small and may be no more than swellings of the head wall.
1.3.2 Sensilla on the antennae

The antennae are primarily sensory structures and they are

richly endowed with sensilla in most insects. It is characteristic of insects that the pedicel contains a chordotonal organ,
Johnstons organ, which responds to movement of the
flagellum with respect to the pedicel (see section 23.2.3.2).
In addition, the scape and pedicel often have hair plates and
groups of campaniform sensilla that provide information on
the positions of the basal segments with respect to the head
and to each other. Scattered mechanosensory hairs are also
often present on these segments.
The principal sensilla on the flagellum of most insects are
olfactory, and these have a variety of forms (see section 24.1.1).
It is common for contact chemoreceptors, mechanoreceptors

10

HEAD

a)

Fig. 1.9. Antenna. Plumose form providing

space for large numbers of sensilla (male of
the moth Telea polyphemus) (after Boeckh,
Kaissling & Schneider, 1960). (a) The whole
antenna seen from above. Two slender
branches arise on opposite sides of each
annulus. (b) Detail of two annuli from the
side showing the bases of the branches and
arrangement of long trichoid olfactory
sensilla along the branches.

b)
annular
branches

annulus
21

flagellum
pedicel
basal part
of branch

and thermohygroreceptors also to be present. Where the

flagellum is made up of a series of similar annuli, successive
annuli often have a similar arrangement of sensilla, but the sensilla are often concentrated in particular regions. In Melanoplus
(Orthoptera), for instance, there are no basiconic or coeloconic
pegs on the proximal annuli; most of these sensilla are found on
the annuli in the middle of the flagellum (Fig. 1.10). In Pieridae
(Lepidoptera), most of the antennal sensilla are aggregated on
the terminal club. The terminal annulus often has a group of
contact chemoreceptors at its tip. The total numbers of sensilla
on an antenna are often very large. Adult male Periplaneta, for
instance, have about 250 000 sensilla on each antenna and male
corn borer moths, Ostrinia, about 8000. When the antennae are
sexually dimorphic, as in many Lepidoptera, the more complex
antenna bears a much larger number of sensilla. For example,
male Telea have over 65 000 sensilla on one antenna, while the
female has only about 13 000.
Review: Zacharuk, 1985
1.3.3 Functions of antennae
The antennae function primarily as sense organs and they
are the primary olfactory receptors of all insects (see
section 24.1.1). They also have a tactile function by virtue
of the large number of mechanosensitive sensilla that are
often present. Very long antennae, such as occur in the
cockroach, are possibly associated with their use as feelers.
Johnstons organ is important in the regulation of airspeed
in flying insects (see Fig. 9.36) and in some insects, male

mosquitoes, female Drosophila and worker honeybees, for

example, it is concerned in the perception of near-field
sounds (see Fig. 23.11).
Sometimes the antennae have other functions. The adult
water beetle Hydrophilus submerges with a film of air over
its ventral surface which it renews at intervals when it comes
to the surface. At the water surface the body is inclined to
one side and a funnel of air, connecting the ventral air
bubble to the outside air, appears between the head, the prothorax and the distal annuli of the antenna, which is held
along the side of the head. The four terminal annuli of the
antenna are enlarged and are clothed with hydrofuge hairs

200
number of sensilla

scape
annulus
20

trichoid
sensilla

basiconic
sensilla

150
100

coeloconic
sensilla

50
0
5

10
15
20
annulus number

25

Fig. 1.10. Distribution of sensilla along the flagellum of a male

grasshopper. Only olfactory sensilla are shown. 1 most
proximal annulus, adjacent to the pedicel; 25 most distal
annulus (Melanoplus) (data from Slifer et al., 1959).

11

REFERENCES

which facilitate the formation of the air funnel. In the newly

hatched larva of Hydrophilus the antennae assist the
mandibles in masticating the prey. This is facilitated by a
number of sharp spines on the inside of the antennae.
In fleas and Collembola the antennae are used in
mating. Male fleas use the antennae to clasp the female
from below and the inner surfaces bear large numbers of
adhesive discs. These discs, about 5 m in diameter, are set

on stalks above the general surface of the cuticle and

within each one there is a gland, presumably secreting an
adhesive material. Species with sessile or semi-sessile
females lack these organs (Rothschild and Hinton, 1968).
In many Collembola the males have prehensile antennae
with which they hold on to the antennae of the female and,
in Sminthurides aquaticus, the male may be carried about by
the female, holding on to her antennae, for several days.

REFERENCES

Bitsch, J. (1973). Morphologie de la tte des

insectes. A. Partie Gnrale. In Trait de
Zoologie, vol. 8, part 1, ed. P.-P.Grass,
pp. 3100. Paris: Masson et Cie.
Boeckh, J., Kaissling, K.-E. & Schneider,
D. (1960). Sensillen und Bau der
Antennengeissel von Telea polyphemus.
Zoologische Jahrbcher (Anatomie), 78,
55984.
Denis, J.R. & Bitsch, J. (1973).
Morphologie de la tte des insectes. B.
Structure cphalique dans les ordres
dinsectes. In Trait de Zoologie, vol. 8,
part 1, ed. P.-P.Grass, pp. 101593.
Paris: Masson et Cie.
DuPorte, E.M. (1957). The comparative
morphology of the insect head. Annual
Review of Entomology, 2, 5570.
Hinton, H.E. (1963). The ventral ecdysial
lines on the head of endopterygote
larvae. Transactions of the Royal
Entomological Society of London, 115,
3961.

Imms, A.D. (1940). On the antennal

structure in insects and other arthropods. Quarterly Journal of Microscopical
Science, 81, 273320.
Imms, A.D. (1957). A General Textbook of
Entomology. London: Methuen.
Matsuda, R. (1965). Morphology and
evolution of the insect head. Memoirs of
the American Entomological Institute, no.
4, 334 pp.
Rothschild, M. & Hinton, H.E. (1968).
Holding organs on the antennae of
male fleas. Proceedings of the Royal
Entomological Society of London, (A) 43,
1057.
Schmidt-Ott, U., Gonzlez-Gaitn, M.,
Jckle, H. & Technau, G.M. (1994).
Number, identity, and sequence of the
Drosophila head segments as revealed
by neural elements and their deletion
patterns in mutants. Proceedings of the
National Academy of Sciences of the
United States of America, 91, 83637.

Slifer, E.H., Prestage, J.J. & Beams, H.W.

(1959). The chemoreceptors and other
sense organs on the antennal flagellum
of the grasshopper, (Orthoptera:
Acrididae). Journal of Morphology, 105,
14591.
Snodgrass, R.E. (1935). Principles of Insect
Morphology. New York: McGraw-Hill.
Snodgrass, R.E. (1947). The insect
cranium and the epicranial suture.
Smithsonian Miscellaneous Collections,
104, no. 7, 113 pp.
Snodgrass, R.E. (1960). Facts and theories
concerning the insect head.
Smithsonian Miscellaneous Collections,
142, 161.
Zacharuk, R.Y. (1985). Antennae and sensilla. In Comprehensive Insect
Physiology, Biochemistry and
Pharmacology, vol. 6, ed. G.A. Kerkut
& L.I. Gilbert, pp. 169. Oxford:
Pergamon Press.