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Insecta (Insects)

EA Jarzembowski, Maidstone Museum and Bentlif Art Gallery, Maidstone, UK

Insects are the most diverse organisms on Earth with a long evolutionary history and are
one of the most successful groups of organisms to have existed.
Judging by their diversity, insects (superclass Hexapoda)
are perhaps the most successful organisms of all time, with
an estimated 230 million species. Most of these are yet to
be described scientically (only about a million have been
namedsofar). However, already well over 370 000dierent
kinds of beetle have beenrecognized compared with a mere
quarter of a million owering plants (angiosperms). The
size of the task of documenting insect species is possibly
one of the single biggest challenges facing entomologists in
the new millennium. Many insects have very specialized
lifestyles and are restricted to particular environments and
microclimates. With the present rate of global develop-
ment, it is perhaps inevitable that many habitats will shrink
or vanish during the twenty-rst century and many species
will become extinct.
Insects occupy an important role in terrestrial and
freshwater ecosystems and without themit is inconceivable
that the worlds food chains would survive in their present
form. For example, what else would pollinate many species
of owering plants, or provide food for many small
vertebrates? Some insects are injurious to human develop-
ment, particularly agriculture (e.g. locusts) and health (e.g.
malaria-spreading mosquitoes). Such species have been
studied intensively by applied entomologists, but for every
pest there are thousands of harmless or benecial species
whichusually receive less attention. Those that doare often
colourful, such as butteries, or large and active, like
dragonies. Even small unobtrusive insects, however, may
have a fascinating biology which can provide insight into
the complexity of life. For example, the study of ants,
wasps and bees has revealed much about the evolution of
social behaviour in which rearing the young is a communal
responsibility. For many years, the Drosophila fruity has
been a laboratory animal for important genetic work
including how genes control body development. Insects
can thus help us to understand the workings of life itself.
Why are insects so successful as a group? There is no
simple answer, but it may be that they are about the right
size, most insects ranging from a few millimetres to a few
centimetres long. The Earth can support only a few
thousand species of large animals such as birds and
mammals. At the other end of the scale, the planet can
support a lot of microorganisms that are smaller than
insects, but because their populations spread so easily,
their biodiversity is correspondingly low despite there
being a very large number of individuals. The upper size
limit of insects (e.g. the Hercules beetle 16 cm long) is
usually attributed to the constraints of their special
(tracheate) respiratory system (see below). Apart from
size, insects also have some special adaptations of their
arthropod inheritance:
. Their bodies are segmented and the segments have
become fused and specialized (tagmosis) producing
three mainbody divisions a head, thorax andabdomen
(Figure 1).
. Their appendages (limbs) are composed of several
segments and have become specialized to produce
walking legs and other useful parts including several
pairs of mouthparts and the sexual apparatus.
. They have a tough external skeleton (exoskeleton) to
protect them and support the internal soft parts (see
below). The exoskeleton has the added function of
reducing desiccation which has enabled insects to
successfully colonize the terrestrial environment.
Insects dier from other arthropods in that:
. They have basically three pairs of legs, one pair on each
thoracic segment, and walk in a tripodal manner.
. They have evolved powered ight, unlike any other
invertebrates. Over 99% of insects are winged or belong
to basically winged insect groups (collectively called
Pterygota or pterygotes, see below). Unlike in other
animals, insect wings evolved independently of limbs
Article Contents
Introductory article
. Introduction
. Key Stages in Insect Evolution
. Insect Groups
. General Biology
Abdomen Abdomen
Compound eye
Antenna Mouthparts
Figure 1 Giant ant Formicium showing insect body organization (after
Jarzembowski). Wingspan 144 mm.
1 ENCYCLOPEDIA OF LIFE SCIENCES 2001, John Wiley & Sons, Ltd.
and therefore without loss of function in other
Key Stages in Insect Evolution
Insects are thus distinguished by a number of important
adaptations. Insects also have a long geological history.
The four mainstages inthe evolution of insects hadalready
been accomplished by the close of the Palaeozoic era (245
million years ago). These were:
1. The rise of pterygote insects from some primitively
wingless insects (apterygotes, see below). This helped
dispersal and escape from ground enemies (e.g.
spiders) prior to the evolution of any other ying
2. The ability of many insects (Neoptera, see below) to
fold their wings over their bodies when at rest. This
gave adult winged insects the advantage of entering
conned spaces such as nooks and crannies in rocks
and plants.
3. The appearance of metamorphosis, especially com-
plete metamorphosis (holometaboly). Insects that
reach the latter evolutionary level undergo a major
structural reorganization during a resting (pupal)
stage between the juvenile and adult stages (Figure 2,
4. A near-modern ecological spectrum of plant-feeding
activity (herbivory) arising over one hundred million
years before the spread of owering plants (angios-
Holometaboly enabled the adult (imago) and juvenile
(larva) to evolve marked dierences in form and lifestyle
andthus exploit dierent natural resources without mutual
competition. It is perhaps inevitable that insects possessing
a combination of neoptery and holometaboly (2 and 3)
include the most diverse organisms on Earth. Complex
adaptations such as parasitismand sociality appeared later
in the Mesozoic era (the age of the dinosaurs). Insects were
modern in many respects by the dawn of the Cenozoic era
(65 million years ago) in which we still live.
The origin and relationships of insects are, however, still
a subject of debate and research (Figure 3). The earliest
known hexapod is a springtail, Rhyniella praecursor, from
the Lower Devonian Rhynie Chert of Scotland dated at
396 million years (Ma) (Figure 4). Pterygote insects appear
much later in the mid Carboniferous, c. 330 Ma. Insects
share features with myriapods (millipedes and centipedes),
including the presence of tracheae andMalpighiantubules.
They also share features with crustaceans (crabs and
shrimps), such as similarities in the nervous systemand the
possession of compound eyes. A relationship with Crus-
tacea has been supported recently by the discovery of
shared genetic characters at the molecular level. There are,
however, important dierences in the body plans of insects
and crustaceans, notably the presence of a second pair of
antennae (antennules) and a nauplius larva in the latter. If
Hexapoda is most closely related to Crustacea, then
myriapods are related to apterygotes.
Insect Groups
There is broad agreement about the composition of the
major divisions (orders) of living Hexapoda. There is a
variable number of extinct orders some of which are
currently being reinterpreted in the light of newstudies and
palaeoentomological discoveries. Some entomologists
Figure 2 Life cycles of hemimetabolous (left) andholometabolous (right)
insects compared (nymph = larva, pupa = chrysalis). (Adapted from
Chinery, 1993.)
Insecta (Insects)
restrict the term insect to pterygotes and their closest
apterygote relatives and regard the remaining apterygotes
as other groups of hexapods. The insect orders, with their
main attributes and included groups, are summarized
Apterygotes are primitively wingless insects. They range in
length from less than 1 to 50 mm. Under 1% of all known
hexapod species are apterygotes: they include Collembola,
Diplura, Protura, Thysanura and Monura.
. Order Collembola (springtails): Found from the Lower
Devonian onwards, these small or minute, white or
pigmented entognathous hexapods are called springtails
because of the ability of many species to leap consider-
able distances when disturbed. The leap is accomplished
using a special appendage called the spring (furca) on
abdominal segment 4. There is usually a glue tube
(collophore) on segment 1 which is used for imbibing
water. Collembola are among the most important
consumers in many soil ecosystems.
. Order Diplura (two-pronged bristletails): Dating from
the Upper Carboniferous, diplurans are small or large,
narrow-bodied and mostly unpigmented, blind entog-
nathous hexapods. There is a pair of cerci (tails) on
abdominal segment 10 (hence the common name) which
may resemble antennae or forceps. Diplura are mainly
found in damp soil under logs or stones.
. Order Protura: There is no fossil record. proturans are
very small, cryptic, mostly white entognathous hexa-
pods with no eyes or antennae. The front legs point
forward. They have no common name but Protura
means single tail, referring to the simple pointed
Figure 3 Provisional relationships among the major hexapod groups (after Jarzembowski).
Insecta (Insects)
abdomen. Proturans are found in soil but are uncom-
. Order Thysanura (three-tailed bristletails): These
ectognathous hexapods have a carrot-shaped body with
shiny scales and two long, thread-like antennae plus
three long, segmented tails, hence the common name.
The order is often split into two separate orders
Archaeognatha and Zygentoma, the former including
Petrobius found on the sea shore and the latter the
domestic silversh and rebrat. The extinct order
Monura (with only one tail) from the Upper Carbo-
niferous and Permian is related to Archaeognatha.
Zygentoma are considered to be the closest relatives of
winged insects (pterygotes) because of similar jaw
articulation and together form the Dicondylia.
Primitively winged insects comprising 99% of all known
insect species. sizes range from very small to very large
(under 1 mm long to about 700 mm wingspan (fossils)).
Two pairs of wings are present, one each on the middle and
hind thoracic segments, although the hind pair may be
reduced or lost, e.g. in certain parasites and underground
dwellers. Some fossil pterygotes had an additional pair of
winglets on the front thoracic segment. Pterygotes have
two main divisions: exopterygotes and endopterygotes.
The wings develop externally in the juvenile insect (larva or
nymph). Metamorphosis is incomplete (pauro- or hemi-
metabolous; Figure 2, left) and there is no pupal stage.
Exopterygotes comprise three groups: Palaeoptera, Poly-
neoptera and Paraneoptera; Polyneoptera and Paraneop-
tera are the Neoptera.
These insects cannot normally fold their wings at over the
body when at rest (due to lack of a structure at the wing
base known as the third axillary sclerite). Some Palaeop-
tera can, however, bend their wings back. Palaeopterous
insects comprise two living orders and several extinct ones.
. Order Ephemeroptera (mayies): Upper Carboniferous
onwards. Delicate insects with two or three tails.
Larvae with abdominal gills living in freshwater. Adult
with reduced hindwings, non-functional (vestigial)
mouthparts, and a subimago stage. Mayies are not
conned to the month of May, but many adults live for
less than a day, hence their scientic name (derived from
the Greek ephemeros, short lived).
. Order Odonata (dragonies and damselies, sometimes
called dragonies collectively): Upper Carboniferous
onwards. Adults and larvae (nymphs) are active
predators. Larvae usually live in freshwater with
tracheal gills and extendable mouthparts for seizing
prey. Adults with two pairs of well-developed wings and
toothed jaws: they take insect prey on the wing, hunting
by sight. The adults may be distinctly patterned,
associated with sexual dimorphism and well-developed
vision. Dragonies include the largest insects of all time
in the extinct order Protodonata (Upper Carboniferous
to Triassic) some of which attained wingspans of 70 cm.
The Palaeoptera includes several extinct late Palaeozoic
orders collectively called the Palaeodictyopteroidea (or
Archeoptera) which had beak-like mouthparts and are
thought to have fed on plants. Some Palaeodictyopter-
oidea (Diaphanopterodea) developed a unique wing-
folding mechanism.
These are the more primitive Neoptera with nine living
orders and several extinctions. Both adults and larvae
(nymphs) are terrestrial except in the order Plecoptera
where the larvae are aquatic. The larvae are usually
miniature adults as in Paraneoptera. Mouthparts are
adapted for biting (cutting and chewing) a variety of food.
. Order Blattodea (cockroaches): Upper Carboniferous
onwards. Body attened with large head shield (prono-
tum) and leathery forewings (tegmina).
. Order Dermaptera (earwigs): Jurassic onwards. Abdo-
men often with forceps. Forewings reduced to small
tegmina (like rove beetles). Hindwings, when present,
large andfan-shaped. Some earwig females are noted for
their maternal care. The common name stems from the
fact that Dermaptera have been found occasionally in
peoples ears (e.g. when camping or sleeping on straw
mattresses) associated with the insects cryptic beha-
Figure 4 Rhyniella praecursor reconstructed leaping(after Jarzembowski).
Length 1.5 mm.
Insecta (Insects)
. Order Embioptera (web-spinners): Permian onwards.
Forelegs with distinct swellings packed with glands
which are used to line tunnels with silk in which the
insects live.
. Order Isoptera (termites): Lower Cretaceous onwards.
Social insects with reproductive forms (queens and
kings) and sterile soldiers and workers. They live in
family units (nests) in wood or earthen constructions,
sometimes in buildings. The reproductives can y to
colonize new sites and shed their wings on landing.
. Order Mantodea (praying mantises): Cretaceous on-
wards. Predators with distinctive spiny front legs used
for catching insect prey. The name mantis is derived
from the Greek for holy person, and the common name
also reects the resemblance to the act of praying when
awaiting prey.
. Order Orthoptera (crickets, grasshoppers, katydids and
locusts): Upper Carboniferous onwards. Insects with
hind legs usually adapted for jumping and pronotum
extended down the side of the body. Males of many
species produce distinctive calls (songs) by rasping the
forewings or forewings/hind legs together.
. Order Phasmatodea (stick insects): Permian onwards.
Plant-feeding insects with a remarkable resemblance to
sticks and leaves. Reproduction without males (parthe-
nogenesis) occurs in some species.
. Order Plecoptera (stoneies): Permian onwards. Larvae
(nymphs) usually live in freshwater and have no gills and
only two long tails. The adults often have distinctive
veins resembling a pair of ladders intheir wings, andmay
be found among waterside stones, from which they get
their common name.
The polyneopterous insects also include the following
orders: Grylloblattodea, Protelytroptera, Caloneurodea,
Miomoptera and Protorthoptera of which the rst is still
living but rare and the others are all extinct.
These are characterized mainly by reductions in internal
organs and foot segmentation. There are ve living orders
and some extinct groups.
. Order Hemiptera (true bugs): Permian onwards. The
dominant order of exopterygote insects with piercing
mouthparts adapted for sucking. In the suborder
Homoptera (including leafhoppers, cicadas, aphids,
scale insects and whitey) the forewings are usually of
even consistency whereas in Heteroptera (true bugs)
they may have a soft outer part, e.g. shield bugs.
Homopterans feed only on plant juices, whereas some
heteropterans alsosuckbloodandinclude aquatic forms
(e.g. pond skaters, water boatmen). Bugs can transmit
plant and occasionally human diseases (Chagas dis-
. Order Phthiraptera (lice): Eocene onwards. Wingless
insects, parasitic on birds and mammals. Body attened
when viewed from above with biting or sucking
mouthparts in the divisions Mallophaga and Ano-
plura, respectively.
. Order Psocoptera (book and bark lice): Permian
onwards. Small insects with a round head, long
antennae, biting mouthparts and wings often held
roofwise (pitched at an angle) over the abdomen. They
are mainly scavengers.
. Order Thysanoptera (thrips): Permian onwards. Small,
slender insects with asymmetric mouthparts adapted
for sucking plants. Wings, when present, narrow with
reduced venation and a hairy fringe resembling tiny
feathers. (The latinized name of the order means fringe-
Paraneopterous insects also include the tiny living
order Zoraptera, and some extinct Palaeozoic proto-
Endopterygotes (Oligoneoptera, Endopterygota,
These are insects in which the wings develop internally
(hence Endopterygota) and metamorphosis is complete
(holometabolous, Figure 2, right). They comprise 80%
of known insect species with one extinct and eleven living
orders, although most of the biodiversity is in four
orders (Coleoptera, Diptera, Hymenoptera, Lepidoptera):
. Order Coleoptera (beetles; Permian onwards): These
minute tolarge insects normally have biting mouthparts.
The front pair of wings are modied into rigid wing
cases called elytra. The elytra usually meet edge to edge
in a straight line when at rest and partly or wholly
cover the hindwings and abdomen. 40% of all insects
and 30% of all animals are beetles; the number of
beetle species is six (or more) times greater than that of
vertebrates. The single most important factor contribut-
ing to the success of the Coleoptera is the evolution of
the elytra, protecting the body and facilitating life in
enclosed spaces and cryptic habitats. Beetles are mainly
terrestrial, but some are aquatic.
. Order Diptera (true or two-winged ies, including gnats,
midges, mosquitoes and bluebottles; Triassic onwards):
These minute to moderately large insects have only the
front pair of wings usually present, the hindwings
having become altered to club-like balance organs
(halteres). Wings are translucent with few cells, absent
in parasitic forms. Mouthparts are adapted for sucking
or mopping up liquids, e.g. houseies and for biting
(piercing and sucking), e.g. mosquitoes. Some species
transmit virulent diseases and many mimic bees and
wasps for protection (see below). Larvae lack true
legs and are usually maggot-like. Pupae sometimes
Insecta (Insects)
develop in a tough skin (puparium). Diptera are
often associated with moist situations and decaying
. Order Hymenoptera (sawies, wasps, hornets,
ichneumons, ants and bees; Triassic onwards): These
minute to large insects usually have two pairs of
translucent wings of which the front pair is larger.
The wings are coupled by a row of minute hooks on the
front edge of the hindwing. The wing venation consists
of large cells, or may be greatly reduced. Biting
mouthparts may be accompanied by a tongue for
lapping up liquids, as in bees, to collect nectar for
making honey as larval food. A well-developed egg-
laying organ (ovipositor) is usually found in females,
sometimes used as a sting. Larvae are terrestrial, with
or without legs. Wasps, hornets, ichneumons and bees
have a very narrow waist which is absent in sawies.
The ovipositor in sawies is used for cutting slits in
plants, hence the common name. Ichneumons and
many wasps are parasitic on other insects and important
in biological control. Some bees are also parasitic, e.g.
cuckoo bees. Wasps, ants and bees have evolved
eusociality independently of the termites and have
reproductives and workers inhabiting earth or paper
nests. The colonies are dominated by queens. Ant
reproductives can shed their wings, like termites,
although workers lack wings, unlike wasp and bee
workers which are noted insect predators and pollina-
tors of owers, respectively. There are many solitary
wasps and bees, unlike ants.
. Order Lepidoptera (moths and butteries, Jurassic
onwards): These insects, which may be small or large,
usually have two pairs of wings which, together with
the body, are more or less covered with tiny scales
(attened hairs). Most adult Lepidoptera have tubular
sucking mouthparts (proboscises) for feeding on juices
like nectar (although some have primitive biting
mouthparts) with the proboscis coiled like a watch
spring when not in use. Larvae (caterpillars) have biting
mouthparts mostly for chewing plants and include
pests of crops and garden plants. Exceptions include
the clothes moth, which feeds on animal material such
as wool. Some caterpillars spin a protective cocoon
before pupating, as in the silk moth, the threads of which
are used to weave a quality fabric. Day-ying Lepidop-
tera include some of the most colourful insects, the wing
patterns being formed by natural pigment and irides-
cence, the latter provided by the microscopic structure
of the scales.
. Order Mecoptera (scorpionies; Permian onwards):
Scorpionies are small or medium-size insects in which
the head is typically extended downwards to form a
beak with the mouthparts at the end. The tip of the male
abdomen is often curved up and swollen, resembling a
scorpions sting. Some species hang from plants by
the forelegs. Larvae are terrestrial.
. Order Neuroptera (lacewings and antlions; Permian
onwards): These insects, which may be small or large,
usually have two similar pairs of wings which are
large and covered with a complex network of veins
and held roofwise when not ying. Neuroptera are
predators and larvae have distinctive sucking jaws used
to catch other insects, e.g. ants and aphids. Larvae are
terrestrial or aquatic. Adult antlions resemble dragon-
ies and mantisies resemble praying mantises.
. Order Megaloptera (alder ies): These medium-size
or large insects are sometimes included in Neuroptera.
Larvae have biting mouthparts and are aquatic.
. Order Raphidioptera (snake ies): These medium-
size insects are sometimes included in Neuroptera.
Larvae have biting mouthparts and are terrestrial.
. Order Siphonaptera (eas; Cretaceous onwards):
Fleas are small, wingless insects compressed sideways
(unlike lice) with blood-sucking mouthparts. They
are parasites of birds and mammals.
. Order Strepsiptera (stylopids; Eocene onwards): Small
parasites with extreme sexual dimorphism. Males are
free-living, and have functional hindwings with the
forewings modied as halteres (unlike in Diptera).
The females are grub-like and generally stay in the host.
. Order Trichoptera (caddisies, sedge ies or rails of
shermen; Permian onwards): These are small or
medium-size, moth-like insects with long antennae,
large compound eyes and wings held roofwise
when not ying. Adults are readily distinguished
from moths by being covered in small hairs and few (if
any) scales, having reduced mouthparts, and being
found near water where the larvae live. The larvae
have biting mouthparts and often live in xed shelters
or portable cases which they construct of various
There is one extinct order of Oligoneoptera: the Glossely-
trodea (Permian to Jurassic).
General Biology
Exoskeleton and growth
The insect exoskeleton (cuticle) is made of chitin and
sclerotin, the latter forming the hard outer layer, as well as
protein and a waxy external coating. Insect larvae grow by
shedding their exoskeleton periodically, e.g. nymphs of a
hemimetabolous grasshopper moult ve to eight times
(Figure 2, left). Ametabolous insects (without metamor-
phosis) continue to moult into adult life, altogether as
much as 50 times in the silversh. The growth period of
insects tends to be faster in hot climates. A tropical
buttery may develop from egg to adult in 3 weeks,
Insecta (Insects)
whereas in temperate regions there are usually only one to
three generations per annum.
Internal organs
The gut is a tube (with pouches) extending fromthe mouth
to the anus. Excretion is done mainly via the Malpighian
tubules, which are blind, narrow tubes attached to the gut
and producing uric acid. Blood bathes the organs in large
cavities (the haemocoel) and there are few blood vessels.
Air is supplied to the living tissues (and not the blood) by a
network of chitinous tubes (tracheae and tracheoles) from
about 10 pairs of openings (spiracles) on the sides of the
body. Many aquatic larvae get their oxygen by simple
The compound eyes are made up of a number of similar
light-sensitive units (ommatidia). Each unit has a lens
giving the eye a faceted appearance. Dragonies have good
visionwithupto30 000 ommatidia. Insects alsohave single
eyes (ocelli).
This sense is located in the antennae. The branched
antennae of some male moths can detect female scent over
several miles, e.g. in the emperor moth.
Sound and hearing
Crickets, cicadas and other insects produce songs which
they hear with ears (tympanal organs) on various parts of
the body. Night-ying moths can hear bat calls and take
evasive action.
Insects can taste with their feet (tarsi) as well as their
mouths, e.g. the red admiral buttery can detect even very
weak sugar solution with its forefeet.
Insects also have touch sensors, heat and temperature
Many small insects depend on sheer fecundity to outstrip
predation. Larger insects with smaller populations may
hide or display protective camouage, e.g. leaf insects. The
black and yellow or red stripes of wasps are warning
colours that the insects can sting and birds soon learn to
avoid them. To discourage predators, many insects have a
nasty taste, e.g. burnet moths, or irritant hairs, e.g. the
brown-tail moth caterpillar. Some defenceless species
mimic wasp coloration for protection, e.g. the wasp beetle.
Palatable species can resemble unpalatable ones, e.g. in
butteries the viceroy resembles the monarch. Some insect
larvae are protected by being surrounded in food, e.g. gall-
forming and leaf-mining insects and dung-ies.
Further Reading
Carpenter FM (1992) Superclass Hexapoda. Treatise on Invertebrate
Paleontology, Part R, Arthropoda 4, 3 & 4. Lawrence: Kansas
University Press.
Chinery M (1993) Insects of Britain and Northern Europe, 3rd edn.
London: HarperCollins.
Commonwealth Scientic and Industrial Research Organisation (1991)
The Insects of Australia, 2nd edn. vols 1 and 2. Carlton: Melbourne
University Press.
Daly HV, Doyen JT and Purcell AH III (1998) Introduction to Insect
Biology and Diversity, 2nd edn. Oxford: Oxford University Press.
Jarzembowski EA and Ross AJ (1996) Insect origination and extinction
in the Phanerozoic. Biotic Recovery from Mass Extinction Events,
Geological Society, Special Publication, no. 102, pp. 6578. London.
Jolivet P (1998) Interrelationships Between Insects and Plants. Boca
Raton, FL: CRC Press.
Parker SP (ed.) (1982) Synopsis and Classication of Living Organisms,
vol. 2. New York: McGraw-Hill.
Wilson EO (1971) The Insect Societies. Cambridge, MA: Belknap/
Harvard University Press.
Insecta (Insects)