Insect

Insects (from Latin insectum) are hexapod invertebrates of the

class Insecta. They are the largest group within the arthropod Insect
phylum. Insects have a chitinous exoskeleton, a three-part Temporal range:
body (head, thorax and abdomen), three pairs of jointed legs,
compound eyes, and a pair of antennae. Insects are the most
diverse group of animals, with more than a million described
species; they represent more than half of all animal species.

The insect nervous system consists of a brain and a ventral

nerve cord. Most insects reproduce by laying eggs. Insects
breathe air through a system of paired openings along their
sides, connected to small tubes that take air directly to the
tissues. The blood therefore does not carry oxygen; it is only
partly contained in vessels, and some circulates in an open
hemocoel. Insect vision is mainly through their compound
eyes, with additional small ocelli. Many insects can hear, using
tympanal organs, which may be on the legs or other parts of
the body. Their sense of smell is via receptors, usually on the
antennae and the mouthparts.
Insects have a three-part body: head
Nearly all insects hatch from eggs. Insect growth is constrained
with large compound eyes and
by the inelastic exoskeleton, so development involves a series
of molts. The immature stages often differ from the adults in antennae, a thorax with three pairs of
structure, habit and habitat. Groups that undergo four-stage legs, and a segmented abdomen.
metamorphosis often have a nearly immobile pupa. Insects that Many groups also have two pairs of
undergo three-stage metamorphosis lack a pupa, developing wings.
through a series of increasingly adult-like nymphal stages. The
Scientific classification
higher level relationship of the insects is unclear. Fossilized
insects of enormous size have been found from the Paleozoic Domain: Eukaryota
Era, including giant dragonfly-like insects with wingspans of Kingdom: Animalia
55 to 70 cm (22 to 28 in). The most diverse insect groups
Phylum: Arthropoda
appear to have coevolved with flowering plants.
Clade: Pancrustacea
Adult insects typically move about by walking and flying;
Subphylum: Hexapoda
some can swim. Insects are the only invertebrates that can
achieve sustained powered flight; insect flight evolved just Class: Insecta
once. Many insects are at least partly aquatic, and have larvae Linnaeus, 1758
with gills; in some species, the adults too are aquatic. Some Subgroups
species, such as water striders, can walk on the surface of
water. Insects are mostly solitary, but some, such as bees, ants Archaeognatha
and termites, are social and live in large, well-organized
Dicondylia
colonies. Others, such as earwigs, provide maternal care, Zygentoma
guarding their eggs and young. Insects can communicate with
Pterygota
each other in a variety of ways. Male moths can sense the
pheromones of female moths over great distances. Other Synonyms
species communicate with sounds: crickets stridulate, or rub
their wings together, to attract a mate and repel other males. Ectognatha
Lampyrid beetles communicate with light.
Entomida
Humans regard many insects as pests, especially those that
damage crops, and attempt to control them using insecticides and
other techniques. Others are parasitic, and may act as vectors of
diseases. Insect pollinators are essential to the reproduction of
many flowering plants and so to their ecosystems. Many insects
are ecologically beneficial as predators of pest insects, while a few
provide direct economic benefit. Two species in particular are
economically important and were domesticated many centuries Insects live in a world of motion.
ago: silkworms for silk and honey bees for honey. Insects are This leaf-footed bug climbs wind
blown grass and flies off.
consumed as food in 80% of the world's nations, by people in
roughly 3,000 ethnic groups. Human activities are having serious
effects on insect biodiversity.

Etymology
The word insect comes from the Latin word insectum from in, "cut up",[1] as insects appear to be cut into
three parts. The Latin word was introduced by Pliny the Elder who calqued the Ancient Greek word
ἔντομον éntomon "insect" (as in entomology) from ἔντομος éntomos "cut in pieces";[2] this was
Aristotle's term for this class of life in his biology, also in reference to their notched bodies. The English
word insect first appears in 1601 in Philemon Holland's translation of Pliny.[3][4]

Insects and other bugs

Distinguishing features
In common speech, insects and other terrestrial arthropods are often called bugs.[a] Entomologists to
some extent reserve the name "bugs" for a narrow category of "true bugs", insects of the order Hemiptera,
such as cicadas and shield bugs.[6] Other terrestrial arthropods, such as centipedes, millipedes, woodlice,
spiders, mites and scorpions, are sometimes confused with insects, since they have a jointed
exoskeleton.[7] Adult insects are the only arthropods that ever have wings, with up to two pairs on the
thorax. Whether winged or not, adult insects can be distinguished by their three-part body plan, with
head, thorax, and abdomen; they have three pairs of legs on the thorax.[8]

Insects and other bugs that could be confused with them

 Woodlouse: seven pairs
of legs, seven body
Spider: eight legs, segments (plus head and
Insect: Six legs, three- two-part body tail)
part body
(head, thorax, abdomen),
up to two pairs of wings

Centipede: many legs,

one pair per segment

Millipede: many legs,

two pairs per segment

Diversity
Estimates of the total number of insect
species vary considerably, suggesting
that there are perhaps some 5.5 million
insect species in existence, of which
about one million have been described
and named.[9] These constitute around
half of all eukaryote species, including
animals, plants, and fungi.[10] The most
diverse insect orders are the Hemiptera
(true bugs), Lepidoptera (butterflies and
moths), Diptera (true flies),
About half of all eukaryotes are insects (left side of diagram).
Hymenoptera (wasps, ants, and bees),
and Coleoptera (beetles), each with
more than 100,000 described species.[9]

Insects are extremely diverse. Five groups each have over 100,000 described species.
 True bugs Butterflies and moths Flies
(Hemiptera) (Lepidoptera) (Diptera)

Wasps Beetles
(Hymenoptera) (Coleoptera)

Distribution and habitats

Insects occur in habitats as varied as snow, freshwater, the tropics, desert, and even
the sea.
 The snow scorpionfly The great diving beetle The green orchid bee
Boreus hyemalis on Dytiscus marginalis larva Euglossa dilemma of
snow in a pond Central America

The desert locust Sea skater Halobates on a

Schistocerca gregaria Hawaii beach
laying eggs in sand

Insects are distributed over every continent and almost every terrestrial habitat. There are many more
species in the tropics, especially in rainforests, than in temperate zones.[11] The world's regions have
received widely differing amounts of attention from entomologists. The British Isles have been
thoroughly surveyed, so that Gullan and Cranston 2014 state that the total of around 22,500 species is
probably within 5% of the actual number there; they comment that Canada's list of 30,000 described
species is surely over half of the actual total. They add that the 3,000 species of the American Arctic must
be broadly accurate. In contrast, a large majority of the insect species of the tropics and the southern
hemisphere are probably undescribed.[11] Some 30–40,000 species inhabit freshwater; very few insects,
perhaps a hundred species, are marine.[12] Insects such as snow scorpionflies flourish in cold habitats
including the Arctic and at high altitude.[13] Insects such as desert locusts, ants, beetles, and termites are
adapted to some of the hottest and driest environments on earth, such as the Sonoran Desert.[14]

Phylogeny and evolution

External phylogeny
Insects form a clade, a natural group with a common ancestor, among the arthropods.[15] A phylogenetic
analysis by Kjer et al. (2016) places the insects among the Hexapoda, six-legged animals with segmented
bodies; their closest relatives are the Diplura (bristletails).[16]

Hexapoda ⁠
⁠
 ⁠
Collembola (springtails)
⁠
⁠ Protura (coneheads)
⁠

⁠ ⁠ Diplura (two-pronged bristletails)

⁠
⁠
⁠
⁠ Insecta (=Ectognatha)
⁠

Internal phylogeny
The internal phylogeny is based on the works of Wipfler et al. 2019 for the Polyneoptera,[17] Johnson et
al. 2018 for the Paraneoptera,[18] and Kjer et al. 2016 for the Holometabola.[19] The numbers of described
extant species (boldface for groups with over 100,000 species) are from Stork 2018.[9]
⁠Insecta
⁠Monocondylia Archaeognatha (hump-backed/jumping bristletails, 513 spp)
⁠
⁠Dicondylia
⁠ Zygentoma (silverfish, firebrats, fishmoths, 560 spp)
⁠
⁠Pterygota
⁠ Odonata (dragonflies and damselflies, 5,89
⁠
⁠Palaeoptera
⁠
⁠ Ephemeroptera (mayflies, 3,240 spp)
⁠

⁠Neoptera
⁠ Zoraptera (angel insects, 37
⁠
⁠
⁠
⁠ Dermaptera (earwigs, 1,978
⁠
⁠
⁠ Plecoptera (stoneflies, 3,743
⁠
⁠
⁠ Orthoptera (grasshoppers,
⁠
⁠
Gryllo
⁠
⁠

⁠Notoptera
⁠ Manto
⁠
⁠Polyneoptera ⁠
⁠
⁠
 ⁠ ⁠
Phasm
⁠
⁠
⁠
⁠ ⁠
⁠ Embio
⁠
⁠

⁠ Mantodea (ma
⁠
⁠Dictyoptera
⁠ Blattodea (coc
⁠
⁠

Psocodea (book
⁠
⁠ spp)

⁠Paraneoptera
⁠ ⁠ Hemiptera (true
⁠
⁠
⁠
⁠ Thysanoptera (t
⁠
⁠ ⁠Holometabola
Hymenoptera (sa
⁠
⁠
⁠wings ⁠

⁠
⁠wings flex over abdomen

⁠Co

⁠Neuropteroidea
⁠

⁠Ne

⁠Eumetabola
⁠
 ⁠larvae, pupae

⁠Am

⁠Panorpida
⁠

Taxonomy

Early
Diagram of Linnaeus's key to his seven orders of insect, 1758[20]
⁠Aptera
⁠wingless

⁠Diptera
⁠2‑winged

⁠Coleoptera
⁠forewings fully hardened
⁠
⁠dissimilar pairs
⁠ emiptera
H
⁠ ⁠forewings partly hardened
⁠Insecta
⁠
⁠winged ⁠ epidoptera
L
⁠ ⁠wings scaly
⁠4‑winged

⁠ ⁠Neuroptera
⁠similar pairs ⁠no sting
⁠
 ⁠wings membranous
⁠Hymenoptera
⁠sting

Aristotle was the first to describe the insects as a distinct group. He placed them as the second-lowest
level of animals on his scala naturae, above the spontaneously generating sponges and worms, but below
the hard-shelled marine snails. His classification remained in use for many centuries.[21]

In 1758, in his Systema Naturae,[22] Carl Linnaeus divided the animal kingdom into six classes including
Insecta. He created seven orders of insect according to the structure of their wings. These were the
wingless Aptera, the two-winged Diptera, and five four-winged orders: the Coleoptera with fully-
hardened forewings; the Hemiptera with partly-hardened forewings; the Lepidoptera with scaly wings;
the Neuroptera with membranous wings but no sting; and the Hymenoptera, with membranous wings and
a sting.[20]

Jean-Baptiste de Lamarck, in his 1809 Philosophie Zoologique, treated the insects as one of nine
invertebrate phyla.[23] In his 1817 Le Règne Animal, Georges Cuvier grouped all animals into four
embranchements ("branches" with different body plans), one of which was the articulated animals,
containing arthropods and annelids.[24] This arrangement was followed by the embryologist Karl Ernst
von Baer in 1828, the zoologist Louis Agassiz in 1857, and the comparative anatomist Richard Owen in
1860.[25] In 1874, Ernst Haeckel divided the animal kingdom into two subkingdoms, one of which was
Metazoa for the multicellular animals. It had five phyla, including the articulates.[26][25]

Modern
Traditional morphology-based systematics have usually given the Hexapoda the rank of superclass,[27]
and identified four groups within it: insects (Ectognatha), Collembola, Protura, and Diplura, the latter
three being grouped together as the Entognatha on the basis of internalized mouth parts.[28]

The use of phylogenetic data has brought about numerous changes in relationships above the level of
orders.[28] Insects can be divided into two groups historically treated as subclasses: wingless insects or
Apterygota, and winged insects or Pterygota. The Apterygota traditionally consisted of the primitively
wingless orders Archaeognatha (jumping bristletails) and Zygentoma (silverfish). However, Apterygota is
not monophyletic, as Archaeognatha are sister to all other insects, based on the arrangement of their
mandibles, while the Pterygota, the winged insects, emerged from within the Dicondylia, alongside the
Zygentoma.[29]

The Pterygota (Palaeoptera and Neoptera) are winged and have hardened plates on the outside of their
body segments; the Neoptera have muscles that allow their wings to fold flat over the abdomen. Neoptera
can be divided into groups with incomplete metamorphosis (Polyneoptera and Paraneoptera) and those
with complete metamorphosis (Holometabola). The molecular finding that the traditional louse orders
Mallophaga and Anoplura are within Psocoptera has led to the new taxon Psocodea.[30] Phasmatodea and
Embiidina have been suggested to form the Eukinolabia.[31] Mantodea, Blattodea, and Isoptera form a
monophyletic group, Dictyoptera.[32] Fleas are now thought to be closely related to boreid
mecopterans.[33]

Evolutionary history
The oldest fossil that may be a primitive wingless insect is Leverhulmia from the Early Devonian
Windyfield chert.[34] The oldest known flying insects are from the mid-Carboniferous, around 328–324
million years ago. The group subsequently underwent a rapid explosive diversification. Claims that they
originated substantially earlier, during the Silurian or Devonian (some 400 million years ago) based on
molecular clock estimates, are unlikely to be correct, given the fossil record.[35]

Four large-scale radiations of insects have occurred: beetles (from about 300 million years ago), flies
(from about 250 million years ago), moths and wasps (both from about 150 million years ago).[36]

The remarkably successful Hymenoptera (wasps, bees, and ants) appeared some 200 million years ago in
the Triassic period, but achieved their wide diversity more recently in the Cenozoic era, which began 66
million years ago. Some highly successful insect groups evolved in conjunction with flowering plants, a
powerful illustration of coevolution. Insects were among the earliest terrestrial herbivores and acted as
major selection agents on plants.[37] Plants evolved chemical defenses against this herbivory and the
insects, in turn, evolved mechanisms to deal with plant toxins. Many insects make use of these toxins to
protect themselves from their predators. Such insects often advertise their toxicity using warning
colors.[38]
 The giant dragonfly-like insect Meganeura Beetle Moravocoleus
monyi grew to wingspans of 75 cm (2 ft permianus, fossil and
6 in) in the late Carboniferous, around reconstruction, from the Early
300 million years ago.[39] Permian

Hymenoptera such as this

Iberomaimetsha from the Early
Cretaceous, around 100 million
years ago.

Morphology and physiology

External

Three-part body
Insects have a segmented body supported by an exoskeleton, the hard outer covering made mostly of
chitin. The body is organized into three interconnected units: the head, thorax and abdomen. The head
supports a pair of sensory antennae, a pair of compound eyes, zero to three simple eyes (or ocelli) and
three sets of variously modified appendages that form the mouthparts. The thorax carries the three pairs
of legs and up to two pairs of wings. The abdomen contains most of the digestive, respiratory, excretory
and reproductive structures.[8]

Segmentation
The head is enclosed in a hard, heavily sclerotized, unsegmented head capsule, which contains most of
the sensing organs, including the antennae, compound eyes, ocelli, and mouthparts.[40] The thorax is
composed of three sections named (from front to back) the prothorax, mesothorax and metathorax. The
prothorax carries the first pair of legs. The
mesothorax carries the second pair of legs and the
front wings. The metathorax carries the third pair of
legs and the hind wings.[8][40] The abdomen is the
largest part of the insect, typically with 11–12
segments, and is less strongly sclerotized than the
head or thorax. Each segment of the abdomen has
sclerotized upper and lower plates (the tergum and
sternum), connected to adjacent sclerotized parts by
membranes. Each segment carries a pair of
spiracles.[40] Insect morphology
A- Head B- Thorax C- Abdomen
Exoskeleton 1. antenna 17. anus
2. ocellus (lower) 18. oviduct
The outer skeleton, the cuticle, is made up of two
layers: the epicuticle, a thin and waxy water-resistant 3. ocellus (upper) 19. nerve cord
4. compound eye (abdominal
outer layer without chitin, and a lower layer, the thick
5. brain (cerebral ganglia)
chitinous procuticle. The procuticle has two layers: an
ganglia) 20. Malpighian
outer exocuticle and an inner endocuticle. The tough tubules
and flexible endocuticle is built from numerous layers 6. prothorax
7. dorsal blood 21. tarsal pads
of fibrous chitin and proteins, criss-crossing each
vessel 22. claws
other in a sandwich pattern, while the exocuticle is
8. tracheal tubes 23. tarsus
rigid and sclerotized.[41][42] As an adaptation to life
(trunk with 24. tibia
on land, insects have an enzyme that uses atmospheric
spiracle) 25. femur
oxygen to harden their cuticle, unlike crustaceans
9. mesothorax 26. trochanter
which use heavy calcium compounds for the same
10. metathorax 27. foregut (crop,
purpose. This makes the insect exoskeleton a
lightweight material.[43] 11. forewing gizzard)
12. hindwing 28. thoracic ganglion
13. midgut (stomach) 29. coxa
Internal systems 14. dorsal tube 30. salivary gland
(heart) 31. subesophageal
Nervous 15. ovary ganglion
16. hindgut 32. mouthparts
The nervous system of an insect consists of a brain (intestine,
and a ventral nerve cord. The head capsule is made up rectum, anus)
of six fused segments, each with either a pair of
ganglia, or a cluster of nerve cells outside of the brain.
The first three pairs of ganglia are fused into the brain, while the three following pairs are fused into a
structure of three pairs of ganglia under the insect's esophagus, called the subesophageal ganglion.[44] The
thoracic segments have one ganglion on each side, connected into a pair per segment. This arrangement is
also seen in the first eight segments of the abdomen. Many insects have fewer ganglia than this.[45]
Insects are capable of learning.[46]

Digestive
An insect uses its digestive system to extract nutrients and other substances from the food it
consumes.[47] There is extensive variation among different orders, life stages, and even castes in the
digestive system of insects.[48] The gut runs lengthwise through the body. It has three sections, with
paired salivary glands and salivary reservoirs.[49] By moving its mouthparts the insect mixes its food with
saliva.[50][51] Some insects, like flies, expel digestive enzymes onto their food to break it down, but most
insects digest their food in the gut.[52] The foregut is lined with cuticule as protection from tough food. It
includes the mouth, pharynx, and crop which stores food.[53] Digestion starts in the mouth with enzymes
in the saliva. Strong muscles in the pharynx pump fluid into the mouth, lubricating the food, and enabling
certain insects to feed on blood or from the xylem and phloem transport vessels of plants.[54] Once food
leaves the crop, it passes to the midgut, where the majority of digestion takes place. Microscopic
projections, microvilli, increase the surface area of the wall to absorb nutrients.[55] In the hindgut,
undigested food particles are joined by uric acid to form fecal pellets; most of the water is absorbed,
leaving a dry pellet to be eliminated. Insects may have one to hundreds of Malpighian tubules. These
remove nitrogenous wastes from the hemolymph of the insect and regulate osmotic balance. Wastes and
solutes are emptied directly into the alimentary canal, at the junction between the midgut and hindgut.[56]

Reproductive
The reproductive system of female insects consist of a pair of ovaries, accessory glands, one or more
spermathecae to store sperm, and ducts connecting these parts. The ovaries are made up of a variable
number of egg tubes, ovarioles. Female insects make eggs, receive and store sperm, manipulate sperm
from different males, and lay eggs. Accessory glands produce substances to maintain sperm and to protect
the eggs. They can produce glue and protective substances for coating eggs, or tough coverings for a
batch of eggs called oothecae.[57]

For males, the reproductive system consists of one or two testes, suspended in the body cavity by
tracheae. The testes contain sperm tubes or follicles in a membranous sac. These connect to a duct that
leads to the outside. The terminal portion of the duct may be sclerotized to form the intromittent organ,
the aedeagus.[58]

Respiratory
Insect respiration is accomplished without lungs. Instead, insects have a system of internal tubes and sacs
through which gases either diffuse or are actively pumped, delivering oxygen directly to tissues that need
it via their tracheae and tracheoles. In most insects, air is taken in through paired spiracles, openings on
the sides of the abdomen and thorax. The respiratory system limits the size of insects. As insects get
larger, gas exchange via spiracles becomes less efficient, and thus the heaviest insect currently weighs
less than 100 g. However, with increased atmospheric oxygen levels, as were present in the late
Paleozoic, larger insects were possible, such as dragonflies with wingspans of more than two feet
(60 cm).[59] Gas exchange patterns in insects range from continuous and diffusive ventilation, to
discontinuous.[60][61][62][63]

Circulatory
Because oxygen is delivered directly to tissues via tracheoles, the circulatory system is not used to carry
oxygen, and is therefore greatly reduced. The insect circulatory system is open; it has no veins or arteries,
and instead consists of little more than a single, perforated dorsal tube that pulses peristaltically. This
dorsal blood vessel is divided into two sections: the heart and aorta. The dorsal blood vessel circulates the
hemolymph, arthropods' fluid analog of blood, from the rear of the
body cavity forward.[64][65] Hemolymph is composed of plasma in
which hemocytes are suspended. Nutrients, hormones, wastes, and
other substances are transported throughout the insect body in the
hemolymph. Hemocytes include many types of cells that are
important for immune responses, wound healing, and other
functions. Hemolymph pressure may be increased by muscle
contractions or by swallowing air into the digestive system to aid
in molting.[66]

Sensory
The tube-like heart (green) of the
Many insects possess numerous specialized sensory organs able to mosquito Anopheles gambiae
detect stimuli including limb position (proprioception) by extends horizontally across the
campaniform sensilla, light, water, chemicals (senses of taste and body, interlinked with the diamond-
smell), sound, and heat. [67] Some insects such as bees can shaped wing muscles (also green)
and surrounded by pericardial cells
perceive ultraviolet wavelengths, or detect polarized light, while
(red). Blue depicts cell nuclei.
the antennae of male moths can detect the pheromones of female
moths over distances of over a kilometer.[68] There is a trade-off
between visual acuity and chemical or tactile acuity, such that most insects
with well-developed eyes have reduced or simple antennae, and vice
versa. Insects perceive sound by different mechanisms, such as thin
vibrating membranes (tympana).[69] Insects were the earliest organisms to
produce and sense sounds. Hearing has evolved independently at least 19
times in different insect groups.[70]

Most insects, except some cave crickets, are able to perceive light and
dark. Many have acute vision capable of detecting small and rapid
movements. The eyes may include simple eyes or ocelli as well as larger
compound eyes. Many species can detect light in the infrared, ultraviolet
and visible light wavelengths, with color vision. Phylogenetic analysis
suggests that UV-green-blue trichromacy existed from at least the
Most insects have a pair of
Devonian period, some 400 million years ago.[71] large compound eyes and
other sensory organs such
The individual lenses in compound eyes are immobile, but fruit flies have as antennae able to detect
photoreceptor cells underneath each lens which move rapidly in and out of movements and chemical
focus, in a series of movements called photoreceptor microsaccades. This stimuli on their heads.
gives them, and possibly many other insects, a much clearer image of the
world than previously assumed.[72]

An insect's sense of smell is via chemical receptors, usually on the antennae and the mouthparts. These
detect both airborne volatile compounds and odorants on surfaces, including pheromones from other
insects and compounds released by food plants. Insects use olfaction to locate mating partners, food, and
places to lay eggs, and to avoid predators. It is thus an extremely important sense, enabling insects to
discriminate between thousands of volatile compounds.[73]
Some insects are capable of magnetoreception; ants and bees navigate using it both locally (near their
nests) and when migrating.[74] The Brazilian stingless bee detects magnetic fields using the hair-like
sensilla on its antennae.[75][76]

Reproduction and development

Life-cycles
The majority of insects hatch from eggs. The fertilization and
development takes place inside the egg, enclosed by a shell
(chorion) that consists of maternal tissue. In contrast to eggs of
other arthropods, most insect eggs are drought resistant. This is
because inside the chorion two additional membranes develop
from embryonic tissue, the amnion and the serosa. This serosa
secretes a cuticle rich in chitin that protects the embryo against
desiccation.[77] Some species of insects, like aphids and tsetse
Butterflies mating flies, are ovoviviparous: their eggs develop entirely inside the
female, and then hatch immediately upon being laid.[78] Some
other species, such as in the cockroach genus Diploptera, are
viviparous, gestating inside the mother and born alive.[79] Some insects, like parasitoid wasps, are
polyembryonic, meaning that a single fertilized egg divides into many separate embryos.[80] Insects may
be univoltine, bivoltine or multivoltine, having one, two or many broods in a year.[81]

Other developmental and reproductive variations include

haplodiploidy, polymorphism, paedomorphosis or peramorphosis,
sexual dimorphism, parthenogenesis, and more rarely
hermaphroditism.[82][83] In haplodiploidy, which is a type of sex-
determination system, the offspring's sex is determined by the
number of sets of chromosomes an individual receives. This
system is typical in bees and wasps.[84]
Aphid giving birth to live female
Some insects are parthenogenetic, meaning that the female can young by parthenogenesis from
reproduce and give birth without having the eggs fertilized by a unfertilized eggs
male. Many aphids undergo a cyclical form of parthenogenesis in
which they alternate between one or many generations of asexual
and sexual reproduction.[85][86] In summer, aphids are generally
female and parthenogenetic; in the autumn, males may be
produced for sexual reproduction. Other insects produced by
parthenogenesis are bees, wasps and ants; in their haplodiploid
system, diploid females spawn many females and a few haploid
males.[78]
A female leaf-footed bug deposits
an egg before flying off.
Metamorphosis
Metamorphosis in insects is the process of development that converts young to adults. There are two
forms of metamorphosis: incomplete and complete.

Incomplete
Hemimetabolous insects, those
with incomplete
metamorphosis, change
gradually after hatching from
the egg by undergoing a series
of molts through stages called
instars, until the final, adult,
Incomplete metamorphosis in a locust with multiple instars. Egg is not
stage is reached. An insect
shown. The largest specimen is adult.
molts when it outgrows its
exoskeleton, which does not
stretch and would otherwise restrict the insect's growth. The molting process begins as the insect's
epidermis secretes a new epicuticle inside the old one. After this new epicuticle is secreted, the epidermis
releases a mixture of enzymes that digests the endocuticle and thus detaches the old cuticle. When this
stage is complete, the insect makes its body swell by taking in a large quantity of water or air; this makes
the old cuticle split along predefined weaknesses where it was thinnest.[87][88]

Complete
Holometabolism, or complete
metamorphosis, is where the
insect changes in four stages, an
egg or embryo, a larva, a pupa
and the adult or imago. In these
species, an egg hatches to
produce a larva, which is
generally worm-like in form.
Life-cycle of butterfly, undergoing complete metamorphosis from egg
This can be eruciform through caterpillar larvae to pupa and adult
(caterpillar-like), scarabaeiform
(grub-like), campodeiform
(elongated, flattened and active), elateriform (wireworm-like) or vermiform (maggot-like). The larva
grows and eventually becomes a pupa, a stage marked by reduced movement. There are three types of
pupae: obtect, exarate or coarctate. Obtect pupae are compact, with the legs and other appendages
enclosed. Exarate pupae have their legs and other appendages free and extended. Coarctate pupae develop
inside the larval skin.[89] Insects undergo considerable change in form during the pupal stage, and emerge
as adults. Butterflies are well-known for undergoing complete metamorphosis; most insects use this life
cycle. Some insects have evolved this system to hypermetamorphosis. Complete metamorphosis is a trait
of the most diverse insect group, the Endopterygota.[82]

Communication
Insects that produce sound can generally hear it. Most insects can hear only a narrow range of frequencies
related to the frequency of the sounds they can produce. Mosquitoes can hear up to 2 kilohertz.[90]
Certain predatory and parasitic insects can detect the characteristic sounds made by their prey or hosts,
respectively. Likewise, some nocturnal moths can perceive the ultrasonic emissions of bats, which helps
them avoid predation.[91]

Light production
A few insects, such as Mycetophilidae (Diptera) and the beetle families Lampyridae, Phengodidae,
Elateridae and Staphylinidae are bioluminescent. The most familiar group are the fireflies, beetles of the
family Lampyridae. Some species are able to control this light generation to produce flashes. The
function varies with some species using them to attract mates, while others use them to lure prey. Cave
dwelling larvae of Arachnocampa (Mycetophilidae, fungus gnats) glow to lure small flying insects into
sticky strands of silk.[92] Some fireflies of the genus Photuris mimic the flashing of female Photinus
species to attract males of that species, which are then captured and devoured.[93] The colors of emitted
light vary from dull blue (Orfelia fultoni, Mycetophilidae) to the familiar greens and the rare reds
(Phrixothrix tiemanni, Phengodidae).[94]

Sound production
Insects make sounds mostly by mechanical action of appendages. In grasshoppers and crickets, this is
achieved by stridulation. Cicadas make the loudest sounds among the insects by producing and
amplifying sounds with special modifications to their body to form tymbals and associated musculature.
The African cicada Brevisana brevis has been measured at 106.7 decibels at a distance of 50 cm
(20 in).[95] Some insects, such as the Helicoverpa zea moths, hawk moths and Hedylid butterflies, can
hear ultrasound and take evasive action when they sense that they have been detected by bats.[96][97]
Some moths produce ultrasonic clicks that warn predatory bats of their unpalatability (acoustic
aposematism),[98] while some palatable moths have evolved to mimic these calls (acoustic Batesian
mimicry).[99] The claim that some moths can jam bat sonar has been revisited. Ultrasonic recording and
high-speed infrared videography of bat-moth interactions suggest the palatable tiger moth really does
defend against attacking big brown bats using ultrasonic clicks that jam bat sonar.[100]

Grasshopper stridulation
0:00 / 0:00
Several unidentified grasshoppers stridulating

Problems playing this file? See media help.

Very low sounds are produced in various species of Coleoptera, Hymenoptera, Lepidoptera, Mantodea
and Neuroptera. These low sounds are produced by the insect's movement, amplified by stridulatory
structures on the insect's muscles and joints; these sounds can be used to warn or communicate with other
insects. Most sound-making insects also have tympanal organs that can perceive airborne sounds. Some
hemipterans, such as the water boatmen, communicate via underwater sounds.[101]

Communication using surface-borne vibrational signals is more widespread among insects because of
size constraints in producing air-borne sounds.[102] Insects cannot effectively produce low-frequency
sounds, and high-frequency sounds tend to disperse more in a dense environment (such as foliage), so
insects living in such environments communicate primarily using
substrate-borne vibrations.[103] 0:00 / 0:00
Cricket in garage with familiar call
Some species use vibrations for communicating, such as to attract
mates as in the songs of the shield bug Nezara viridula.[104]
Vibrations can also be used to communicate between species; lycaenid caterpillars, which form a
mutualistic association with ants communicate with ants in this way.[105] The Madagascar hissing
cockroach has the ability to press air through its spiracles to make a hissing noise as a sign of
aggression;[106] the death's-head hawkmoth makes a squeaking noise by forcing air out of their pharynx
when agitated, which may also reduce aggressive worker honey bee behavior when the two are close.[107]

Chemical communication
Many insects have evolved chemical means for communication.
These semiochemicals are often derived from plant metabolites
including those meant to attract, repel and provide other kinds of
information. Pheromones are used for attracting mates of the
opposite sex, for aggregating conspecific individuals of both
sexes, for deterring other individuals from approaching, to mark a
trail, and to trigger aggression in nearby individuals. Allomones
benefit their producer by the effect they have upon the receiver. Social insects such as ants have
Kairomones benefit their receiver instead of their producer. multiple types of pheromonal
Synomones benefit the producer and the receiver. While some glands, producing different
semiochemicals for communication
chemicals are targeted at individuals of the same species, others
with other insects.[108]
are used for communication across species. The use of scents is
especially well-developed in social insects.[108] Cuticular
hydrocarbons are nonstructural materials produced and secreted to the cuticle surface to fight desiccation
and pathogens. They are important, too, as pheromones, especially in social insects.[109]

Social behavior
Social insects, such as termites, ants and many bees and wasps, are eusocial.[110] They live together in
such large well-organized colonies of genetically similar individuals that they are sometimes considered
superorganisms. In particular, reproduction is largely limited to a queen caste; other females are workers,
prevented from reproducing by worker policing. Honey bees have evolved a system of abstract symbolic
communication where a behavior is used to represent and convey specific information about the
environment. In this communication system, called dance language, the angle at which a bee dances
represents a direction relative to the sun, and the length of the dance represents the distance to be
flown.[111] Bumblebees too have some social communication behaviors. Bombus terrestris, for example,
more rapidly learns about visiting unfamiliar, yet rewarding flowers, when they can see a conspecific
foraging on the same species.[112]

Only insects that live in nests or colonies possess fine-scale spatial orientation. Some can navigate
unerringly to a single hole a few millimeters in diameter among thousands of similar holes, after a trip of
several kilometers. In philopatry, insects that hibernate are able to recall a specific location up to a year
after last viewing the area of interest.[113] A few insects
seasonally migrate large distances between different
geographic regions, as in the continent-wide monarch
butterfly migration.[114]

Care of young
Eusocial insects build nests, guard eggs, and provide
food for offspring full-time. Most insects, however, A cathedral mound Honey bee's figure-eight
lead short lives as adults, and rarely interact with one created by eusocial waggle dance. An
another except to mate or compete for mates. A small mound-building orientation 45° to the right
number provide parental care, where they at least guard termites. of ‘up' on the comb
their eggs, and sometimes guard their offspring until indicates food 45° to the
right of the sun. The
adulthood, possibly even feeding them. Many wasps
dancer's rapid waggling
and bees construct a nest or burrow, store provisions in blurs her abdomen.
it, and lay an egg upon those provisions, providing no
further care.[115]

Locomotion

Flight
Insects are the only group of invertebrates to have developed
flight. The ancient groups of insects in the Palaeoptera, the
dragonflies, damselflies and mayflies, operate their wings directly
by paired muscles attached to points on each wing base that raise
and lower them. This can only be done at a relatively slow rate.
All other insects, the Neoptera, have indirect flight, in which the
flight muscles cause rapid oscillation of the thorax: there can be
more wingbeats than nerve impulses commanding the muscles.
One pair of flight muscles is aligned vertically, contracting to pull
Insects such as hoverflies are
the top of the thorax down, and the wings up. The other pair runs capable of rapid and agile flight.
longitudinally, contracting to force the top of the thorax up and the
wings down.[116][117] Most insects gain aerodynamic lift by
creating a spiralling vortex at the leading edge of the wings.[118] Small insects like thrips with tiny
feathery wings gain lift using the clap and fling mechanism; the wings are clapped together and pulled
apart, flinging vortices into the air at the leading edges and at the wingtips.[119][120]

The evolution of insect wings has been a subject of debate; it has been suggested they came from
modified gills, flaps on the spiracles, or an appendage, the epicoxa, at the base of the legs.[121] More
recently, entomologists have favored evolution of wings from lobes of the notum, of the pleuron, or more
likely both.[122] In the Carboniferous age, the dragonfly-like Meganeura had as much as a 50 cm (20 in)
wide wingspan. The appearance of gigantic insects is consistent with high atmospheric oxygen at that
time, as the respiratory system of insects constrains their size.[123] The largest flying insects today are
much smaller, with the largest wingspan belonging to the white witch moth (Thysania agrippina), at
approximately 28 cm (11 in).[124]

Unlike birds, small insects are swept along by the prevailing winds[125] although many larger insects
migrate. Aphids are transported long distances by low-level jet streams.[126]

Walking
Many adult insects use six legs for walking, with an
alternating tripod gait. This allows for rapid walking with a
stable stance; it has been studied extensively in cockroaches
and ants. For the first step, the middle right leg and the front
and rear left legs are in contact with the ground and move the
insect forward, while the front and rear right leg and the
middle left leg are lifted and moved forward to a new 0:00
position. When they touch the ground to form a new stable
triangle, the other legs can be lifted and brought forward in
turn.[127] The purest form of the tripedal gait is seen in insects Spatial and temporal stepping pattern of
moving at high speeds. However, this type of locomotion is walking desert ants performing an
not rigid and insects can adapt a variety of gaits. For example, alternating tripod gait. Recording rate:
500 fps, Playback rate: 10 fps.
when moving slowly, turning, avoiding obstacles, climbing or
slippery surfaces, four (tetrapodal) or more feet (wave-gait)
may be touching the ground.[128] Cockroaches are among the fastest insect runners and, at full speed,
adopt a bipedal run. More sedate locomotion is seen in the well-camouflaged stick insects (Phasmatodea).
A small number of species such as Water striders can move on the surface of water; their claws are
recessed in a special groove, preventing the claws from piercing the water's surface film.[62] The ocean-
skaters in the genus Halobates even live on the surface of open oceans, a habitat that has few insect
species.[129]

Swimming
A large number of insects live either part or the whole of their
lives underwater. In many of the more primitive orders of insect,
the immature stages are aquatic. In some groups, such as water
beetles, the adults too are aquatic.[62]

Many of these species are adapted for under-water locomotion.

Water beetles and water bugs have legs adapted into paddle-like
structures. Dragonfly naiads use jet propulsion, forcibly expelling
water out of their rectal chamber.[130] Other insects such as the The backswimmer Notonecta glauca
underwater, showing its paddle-like
hindleg adaptation
rove beetle Stenus emit pygidial gland surfactant secretions that reduce surface tension; this enables them
to move on the surface of water by Marangoni propulsion.[131][132]

Ecology
Insects play many critical roles in ecosystems, including soil turning and aeration, dung burial, pest
control, pollination and wildlife nutrition.[133] For instance, termites modify the environment around their
nests, encouraging grass growth;[134] many beetles are scavengers; dung beetles recycle biological
materials into forms useful to other organisms.[135][136] Insects are responsible for much of the process by
which topsoil is created.[137]

Defense
Insects are mostly small, soft bodied, and fragile compared to
larger lifeforms. The immature stages are small, move slowly or
are immobile, and so all stages are exposed to predation and
parasitism. Insects accordingly employ multiple defensive
strategies, including camouflage, mimicry, toxicity and active
defense.[138] Many insects rely on camouflage to avoid being
noticed by their predators or prey.[139] It is common among leaf
beetles and weevils that feed on wood or vegetation.[138] Stick
Reduvius personatus, the masked
insects mimic the forms of sticks and leaves.[140] Many insects use
hunter bug nymph, camouflages
mimicry to deceive predators into avoiding them. In Batesian itself with sand grains to avoid
mimicry, edible species, such as of hoverflies (the mimics), gain a predators.
survival advantage by resembling inedible species (the
models).[138][141] In Müllerian mimicry, inedible species, such as
of wasps and bees, resemble each other so as to reduce the sampling rate by predators who need to learn
that those insects are inedible. Heliconius butterflies, many of which are toxic, form Müllerian
complexes, advertising their inedibility.[142] Chemical defense is common among Coleoptera and
Lepidoptera, usually being advertised by bright warning colors (aposematism), as in the monarch
butterfly. As larvae, they obtain their toxicity by sequestering chemicals from the plants they eat into their
own tissues. Some manufacture their own toxins. Predators that eat poisonous butterflies and moths may
vomit violently, learning not to eat insects with similar markings; this is the basis of Müllerian
mimicry.[143] Some ground beetles of the family Carabidae actively defend themselves, spraying
chemicals from their abdomen with great accuracy, to repel predators.[138]

Pollination
Pollination is the process by which pollen is transferred in the reproduction of plants, thereby enabling
fertilisation and sexual reproduction.[144] Most flowering plants require an animal to do the
transportation. The majority of pollination is by insects.[145] Because insects usually receive benefit for
the pollination in the form of energy rich nectar it is a mutualism.
The various flower traits, such as bright colors and pheromones
that coevolved with their pollinators, have been called pollination
syndromes, though around one third of flowers cannot be assigned
to a single syndrome.[146]

Parasitism
European honey bee carrying pollen
Many insects are parasitic. The largest group, with over 100,000
in a pollen basket back to the hive
species[147] and perhaps over a million,[148] consists of a single
clade of parasitoid wasps among the Hymenoptera.[149] These are
parasites of other insects, eventually killing their hosts.[147] Some are hyper-parasites, as their hosts are
other parasitoid wasps.[147][150] Several groups of insects can be considered as either micropredators or
external parasites;[151][152] for example, many hemipteran bugs have piercing and sucking mouthparts,
adapted for feeding on plant sap,[153][154] while species in groups such as fleas, lice, and mosquitoes are
hematophagous, feeding on the blood of animals.[152]

A parasitoid wasp Human head-lice are

ovipositing into an directly transmitted
aphid[155] obligate ectoparasites.
Plant parasite or
micropredator: a coreid
bug sucking plant sap

Relationship to humans

As pests
Many insects are considered pests by humans. These include parasites of people and livestock, such as
lice and bed bugs; mosquitoes act as vectors of several diseases. Other pests include insects like termites
that damage wooden structures; herbivorous insects such as locusts, aphids, and thrips that destroy
agricultural crops, or like wheat weevils damage stored agricultural produce. Farmers have often
attempted to control insects with chemical insecticides, but increasingly rely on biological pest control.
This uses one organism to reduce the population density of a pest organism; it is a key element of
integrated pest management.[156][157] Biological control is favored because insecticides can cause harm to
ecosystems far beyond the intended pest targets.[158][159]

In beneficial roles
Pollination of flowering plants by insects including bees, butterflies,
flies, and beetles, is economically important.[162] The value of insect
pollination of crops and fruit trees was estimated in 2021 to be about
$34 billion in the US alone.[163]

Insects produce useful substances such as honey,[164] wax,[165][166]

lacquer[167] and silk.[168] Honey bees have been cultured by humans
for thousands of years for honey.[169] Beekeeping in pottery vessels
began about 9,000 years ago in North Africa.[170] The silkworm has Aedes aegypti, the yellow fever
greatly affected human history, as silk-driven trade established mosquito, is a vector of several
relationships between China and the rest of the world.[171][172] diseases.

Insects that feed on or parasitise other insects are beneficial to

humans if they thereby reduce damage to agriculture and human
structures. For example, aphids feed on crops, causing economic loss, but
ladybugs feed on aphids, and can be used to control them. Insects account
for the vast majority of insect consumption.[173][174][175]

Fly larvae (maggots) were formerly used to treat wounds to prevent or

stop gangrene, as they would only consume dead flesh. This treatment is
finding modern usage in some hospitals. Insects have gained attention as
potential sources of drugs and other medicinal substances.[176] Adult
insects, such as crickets and insect larvae of various kinds, are commonly
used as fishing bait.[177]
Silkworms were
domesticated for silk over
Population declines 5,000 years ago.[160][161]
Here, silk cocoons are
At least 66 insect species extinctions have been recorded since 1500, being unrolled.
many of them on oceanic islands.[178] Declines in insect abundance have
been attributed to human activity in the form of artificial lighting,[179]
land use changes such as urbanization or farming,[180][181] pesticide use,[182] and invasive
species.[183][184] A 2019 research review suggested that a large proportion of insect species is threatened
with extinction in the 21st century,[185] though the details have been disputed.[186] A larger 2020 meta-
study, analyzing data from 166 long-term surveys, suggested that populations of terrestrial insects are
indeed decreasing rapidly, by about 9% per decade.[187][188]

In research
Insects play important roles in biological research. For example, because of its small size, short
generation time and high fecundity, the common fruit fly Drosophila melanogaster is a model organism
for studies in the genetics of eukaryotes, including genetic linkage, interactions between genes,
chromosomal genetics, development, behavior and evolution. Because genetic systems are well
conserved among eukaryotes, understanding basic cellular processes like DNA replication or transcription
in fruit flies can help to understand those processes in other eukaryotes, including humans.[189] The
genome of D. melanogaster was sequenced in 2000, reflecting the
organism's important role in biological research. It was found that
70% of the fly genome is similar to the human genome, supporting
the theory of evolution.[190]

As food
Insects are consumed as food in 80% of the world's nations, by
people in roughly 3,000 ethnic groups.[192][193] In Africa, locally The fruit fly Drosophila
abundant species of locusts and termites are a common traditional melanogaster is a widely used
model organism.
human food source.[194] Some, especially deep-fried cicadas, are
considered to be delicacies. Insects have a high protein content for
their mass, and some authors suggest their potential as a major source
of protein in human nutrition.[195] In most first-world countries,
however, entomophagy (the eating of insects), is taboo.[196] They are
also recommended by armed forces as a survival food for troops in
adversity.[194] Because of the abundance of insects and a worldwide
concern of food shortages, the Food and Agriculture Organization of
the United Nations considers that people throughout the world may Witchetty grubs are prized as
have to eat insects as a food staple. Insects are noted for their high-protein foods by Aboriginal
nutrients, having a high content of protein, minerals and fats and are Australians.[191]
already regularly eaten by one-third of the world's population.[197]

In other products
Black soldier fly larvae can provide protein and fats for use in cosmetics.[198] Insect cooking oil, insect
butter and fatty alcohols can be made from such insects as the superworm (Zophobas morio).[199] Insect
species including the black soldier fly or the housefly in their maggot forms, and beetle larvae such as
mealworms, can be processed and used as feed for farmed animals including chicken, fish and pigs.[200]
Many species of insects are sold and kept as pets.[201]

In religion and folklore

Scarab beetles held religious and cultural symbolism in ancient
Egypt, Greece and some shamanistic Old World cultures. The
ancient Chinese regarded cicadas as symbols of rebirth or
immortality. In Mesopotamian literature, the epic poem of
Gilgamesh has allusions to Odonata that signify the impossibility
of immortality. Among the Aborigines of Australia of the Arrernte
language groups, honey ants and witchetty grubs served as Ancient Egyptian scarab with
separate wings, c. 712-342 BC
personal clan totems. In the case of the 'San' bush-men of the Kalahari, it is the praying mantis that holds
much cultural significance including creation and zen-like patience in waiting.[202]

See also
Entomology
Ethnoentomology
Flying and gliding animals
Insect-borne diseases

Notes
a. The Museum of New Zealand notes that "in everyday conversation", bug "refers to land
arthropods with at least six legs, such as insects, spiders, and centipedes".[5] In a chapter
on "Bugs That Are Not Insects", entomologist Gilbert Walbauer specifies centipedes,
millipedes, arachnids (spiders, daddy longlegs, scorpions, mites, chiggers and ticks) as well
as the few terrestrial crustaceans (sowbugs and pillbugs).[6]

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External links
Insect species and observations on iNaturalist (https://www.inaturalist.org/observations?plac
e_id=any&subview=map&taxon_id=47158&view=species)
Overview of Orders of Insects (http://bugguide.net/node/view/222292)
"Insect" (https://web.archive.org/web/20230000000000/http://www.eol.org/pages/344). The
Encyclopedia of Life.
A Safrinet Manual for Entomology and Arachnology (https://web.archive.org/web/200911220
74740/http://www.spc.int/PPS/SAFRINET/inse-scr.pdf) SPC
Tree of Life Project (http://tolweb.org/Insecta/8205) – Insecta, Insecta Movies (http://tolweb.
org/movies/Insecta/8205)
Fossil Insect Database (http://edna.palass-hosting.org/): Holotypes at the International
Palaeoentological Society
UF Book of Insect Records (http://entomology.ifas.ufl.edu/walker/ufbir/)
InsectImages.org (http://www.insectimages.org/) 24,000 high resolution insect photographs

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