Insect
Encyclopedia
Insects are a class
of living creatures within the arthropod
s that have a chitin
ous exoskeleton
, a three-part body (head
, thorax
, and abdomen
), three pairs of jointed leg
s, compound eyes, and two antennae
. They are among the most diverse groups of animal
s on the planet, including more than a million described species
and represent more than half of all known living organisms. The number of extant species is estimated at between six and ten million, and potentially represent over 90% of the differing metazoan life forms on Earth. Insects may be found in nearly all environments
, although only a small number of species occur in the oceans, a habitat dominated by another arthropod group, the crustacean
s.
The life cycles of insects vary but most hatch from eggs
. Insect growth is constrained by the inelastic exoskeleton
and development involves a series of molts. The immature stages can differ from the adults in structure, habit and habitat and can include a passive pupa
l stage in those groups that undergo complete metamorphosis. Insects that undergo incomplete metamorphosis
lack a pupal stage and adults develop through a series of nymph
al stages. The higher level relationship of the hexapoda
is unclear. Fossilized insects of enormous size have been found from the Paleozoic Era
, including giant dragonflies with wingspans of 55 to 70 cm (22–28 in). The most diverse insect groups appear to have coevolved with flowering plant
s.
Insects typically move about by walking, flying or occasionally swimming. As it allows for rapid yet stable movement, many insects adopt a tripedal gait in which they walk with their legs touching the ground in alternating triangles. Insects are the only invertebrates to have evolved flight. Many insects spend at least part of their life underwater, with larval adaptations that include gill
s and some adult insects are aquatic and have adaptations for swimming. Some species, like water striders, are capable of walking on the surface of water. Insects are mostly solitary, but some insects, such as certain bee
s, ant
s, and termite
s are social and live in large, well-organized colonies. Some insects, like earwig
s, show maternal care, guarding their eggs and young. Insects can communicate with each other in a variety of ways. Male moth
s can sense the pheromone
s of female moths over distances of many kilometers. Other species communicate with sounds: crickets
stridulate
, or rub their wings together, to attract a mate and repel other males. Lampyridae
in the beetle order Coleoptera
communicate with light.
Humans regard certain insects as pests and attempt to control them using insecticide
s and a host of other techniques. Some insects damage crops by feeding on sap, leaves or fruit
s, a few bite humans and livestock, alive and dead, to feed on blood and some are capable of transmitting disease
s to humans, pets and livestock. Nevertheless, without insects to pollinate flowers, the human race would soon run out of food because many of the crop plants that we rely on would not be able to reproduce. Many other insects are considered ecologically beneficial as predators and a few provide direct economic benefit. Silkworms and bees have been used extensively by humans for the production of silk
and honey
, respectively.
's loan-translation of the Greek
word or "insect" (as in entomology
), which was Aristotle's term for this class of life, also in reference to their "notched" bodies, first documented in English in 1601 in Holland's translation of Pliny. Translations of Aristotle's term also form the usual word for "insect" in Welsh
, Serbo-Croatian
(zareznik, from rezati "cut"), Russian
( nasekomoe, from sekat "cut"), etc.
, Hymenoptera
, and Lepidoptera
. There are almost as many named species of beetle as there are of all other insects added together, or all other non-insects (plant
s and animal
s).
ed bodies supported by an exoskeleton
, a hard outer covering made mostly of chitin
. The segments of the body are organized into three distinctive but interconnected units, or tagmata
: a head, a thorax
, and an abdomen
. The head supports a pair of sensory antennae
, a pair of compound eyes, and, if present, one to three simple eyes (or ocelli) and three sets of variously modified appendages that form the mouthpart
s. The thorax has six segmented leg
s—one pair each for the prothorax, mesothorax and the metathorax segments making up the thorax—and, if present in the species, two or four wing
s. The abdomen consists of eleven segments, though in a few species of insects these segments may be fused together or reduced in size. The abdomen also contains most of the digestive
, respiratory
, excretory
and reproductive internal structures. There is considerable variation and many adaptations in the body parts of insects especially wings, legs, antenna, mouth-parts etc.
: is enclosed in a hard, heavily sclerotized, unsegmented, exoskeletal head capsule, or epicranium
, which contains most of the sensing organs, including the antennae, ocellus or eyes, and the mouthparts. Out of all the insect orders, Orthoptera displays the most features found in other insects, including the sutures and sclerite
s. here, the vertex
, or the apex (dorsal region), is situated between the compound eyes for insects with a hypognathous and opisthognathous head. In prognathous insects, the vertex is not found between the compound eyes, but rather, where the ocelli
are normally. This is because the primary axis of the head is rotated 90 degrees to become parallel to the primary axis of the body. In some species this region is modified and assumes a different name.
The thorax
: is a segment composed of three sections, the prothorax
, mesothorax
, and the metathorax
. The anterior segment, closest to the head, is the prothorax, with the major features being the first pair of legs and the pronotum. The middle segment is the mesothorax with the major features being the second pair of legs and the anterior wings. The third and most posterior segment, abutting the abdomen, is the metathorax, which features the third pair of legs and the posterior wings. Each segment is dilineated by an intersegmental suture. Each segment has four basic regions. The dorsal surface is called the tergum (or notum) to distinguish them from the abdominal terga. The two lateral regions are called the pleura (singular: pleuron) and the ventral aspect is called the sternum. In turn, the notum of the prothorax is called the pronotum, the notum for the mesothorax is called the mesonotum and the notum for the metathorax is called the metanotum. Continuing with this logic, there is also the mesopleura and metapleura as well as the mesosternum and metasternum.
The abdomen
: is a the last segment of the insect, which typically consists of 11–12 segments and is less strongly sclerotized than the head or thorax. Each segment of the abdomen is represented by a sclerotized tergum, sternum, and perhaps a pleurite. Terga are separated from each other and from the adjacent sterna or pleura by a membrane. Spiracles are located in the pleural area. Variation of this ground plan includes the fusion of terga or terga and sterna to form continuous dorsal or ventral shields or a conical tube. Some insects bear a sclerite in the pleural area called a laterotergite. Ventral sclerites are sometimes called laterosternites. During the embryonic stage of many insects and the postembryonic stage of primitive insects, 11 abdominal segments are present. In modern insects there is a tendency toward reduction in the number of the abdominal segments, but the primitive number of 11 is maintained during embryogenesis.Variation in abdominal segment number is considerable. If the Apterygota are considered to be indicative of the ground plan for pterygotes, confusion reigns: adult Protura have 12 segments, Collembola have 6. The orthopteran family Acrididae has 11 segments, and a fossil specimen of Zoraptera has a 10-segmented abdomen.
, and a lower layer called the procuticle. The procuticle is chitinous and much thicker than the epicuticle and has two layers: an outer layer known as the exocuticle and an inner layer known as the endocuticle. The tough and flexible endocuticle is built from numerous layers of fibrous chitin and proteins, criss-crossing each others in a sandwich pattern, while the exocuticle is rigid and hardened. The exocuticle is greatly reduced in many soft-bodied insects (e.g., caterpillar
s), especially during their larva
l stages.
Insects are the only invertebrate
s to have developed active flight capability, and this has played an important role in their success. Their muscles are able to contract multiple times for each single nerve impulse, allowing the wings to beat faster than would ordinarily be possible. Having their muscles attached to their exoskeletons is more efficient and allows more muscle connections; crustacean
s also use the same method, though all spider
s use hydraulic pressure to extend their legs, a system inherited from their pre-arthropod ancestors. Unlike insects, though, most aquatic crustaceans are biomineralized with calcium carbonate
extracted from the water.
of an insect can be divided into a brain and a ventral nerve cord
. The head capsule is made up of six fused segments, each with 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
.
The thoracic segments have one ganglion on each side, which are connected into a pair, one pair per segment. This arrangement is also seen in the abdomen but only in the first eight segments. Many species of insects have reduced numbers of ganglia due to fusion or reduction. Some cockroaches have just six ganglia in the abdomen, whereas the wasp Vespa crabro has only two in the thorax and three in the abdomen. Some insects, like the house fly Musca domestica
, have all the body ganglia fused into a single large thoracic ganglion.
At least a few insects have nociceptor
s, cells that detect and transmit sensations of pain
. This was discovered in 2003 by studying the variation in reactions of larvae
of the common fruitfly Drosophila
to the touch of a heated probe and an unheated one. The larvae reacted to the touch of the heated probe with a stereotypical rolling behavior that was not exhibited when the larvae were touched by the unheated probe. Although nociception has been demonstrated in insects, there is not a consensus that insects feel pain consciously but see Pain in invertebrates
.
s and other complex substances like protein
s, polysaccharide
s, fat
s, and nucleic acid
s. These macromolecules must be broken down by catabolic reaction
s into smaller molecules like amino acid
s and simple sugars before being used by cells of the body for energy, growth, or reproduction. This break-down process is known as digestion.
The main structure of an insect's digestive system is a long enclosed tube called the alimentary canal, which runs lengthwise through the body. The alimentary canal directs food unidirectionally from the mouth
to the anus
. It has three sections, each of which performs a different process of digestion. In addition to the alimentary canal, insects also have paired salivary glands and salivary reservoirs. These structures usually reside in the thorax, adjacent to the foregut.
The salivary gland
s (element 30 in numbered diagram) in an insect's mouth produce saliva. The salivary ducts lead from the glands to the reservoirs and then forward through the head to an opening called the salivarium, located behind the hypopharynx. By moving its mouthparts (element 32 in numbered diagram) the insect can mix its food with saliva. The mixture of saliva and food then travels through the salivary tubes into the mouth, where it begins to break down. Some insects, like flies
, have extra-oral digestion
. Insects using extra-oral digestion expel digestive enzymes onto their food to break it down. This strategy allows insects to extract a significant proportion of the available nutrients from the food source. The gut is where almost all of insects' digestion takes place. It can be divided into the foregut
, midgut
and hindgut
.
The first section of the alimentary canal is the foregut
(element 27 in numbered diagram), or stomodaeum. The foregut is lined with a cuticular lining made of chitin
and protein
s as protection from tough food. The foregut includes the buccal cavity (mouth), pharynx
, esophagus
, and Crop
and proventriculus
(any part may be highly modified) which both store food and signify when to continue passing onward to the midgut.
Digestion starts in buccal cavity (mouth) as partially chewed food is broken down by saliva from the salivary glands. As the salivary glands produce fluid and carbohydrate-digesting enzyme
s (mostly amylase
s), strong muscles in the pharynx pump fluid into the buccal cavity, lubricating the food like the salivarium does, and helping blood feeders, and xylem and phloem feeders.
From there, the pharynx passes food to the esophagus, which could be just a simple tube passing it on to the crop and proventriculus, and then onward to the midgut, as in most insects. Alternately, the foregut may expand into a very enlarged crop and proventriculus, or the crop could just be a diverticulum
, or fluid filled structure, as in some Diptera species.
Once food leaves the crop, it passes to the midgut
(element 13 in numbered diagram), also known as the mesenteron, where the majority of digestion takes place. Microscopic projections from the midgut wall, called microvilli
, increase the surface area of the wall and allow more nutrients to be absorbed; they tend to be close to the origin of the midgut. In some insects, the role of the microvilli and where they are located may vary. For example, specialized microvilli producing digestive enzymes may more likely be near the end of the midgut, and absorption near the origin or beginning of the midgut.
In the hindgut
(element 16 in numbered diagram), or proctodaeum, undigested food particles are joined by uric acid
to form fecal pellets. The rectum absorbs 90% of the water in these fecal pellets, and the dry pellet is then eliminated through the anus (element 17), completing the process of digestion. The uric acid is formed using hemolymph waste products diffused from the Malpighian tubule
s (element 20). It is then emptied directly into the alimentary canal, at the junction between the midgut and hindgut. The number of Malpighian tubules possessed by a given insect varies between species, ranging from only two tubules in some insects to over 100 tubules in others.
s (element 30 in numbered diagram) in an insect's mouth produce saliva. The salivary ducts lead from the glands to the reservoirs and then forward through the head to an opening called the salivarium, located behind the hypopharynx. By moving its mouthparts (element 32 in numbered diagram) the insect can mix its food with saliva. The mixture of saliva and food then travels through the salivary tubes into the mouth, where it begins to break down. Some insects, like flies
, have extra-oral digestion
. Insects using extra-oral digestion expel digestive enzymes onto their food to break it down. This strategy allows insects to extract a significant proportion of the available nutrients from the food source.
e, and ducts connecting these parts. The ovaries are made up of a number of egg tubes, called ovarioles
, which vary in size and number by species. The number of eggs that the insect is able to make vary by the number of ovarioles with the rate that eggs can be develop being also influenced by ovariole design. Female insects are able make eggs, receive and store sperm, manipulate sperm from different males, and lay eggs. Accessory glands or glandular parts of the oviducts produce a variety of substances for sperm maintenance, transport, and fertilization, as well as for protection of eggs. They can produce glue and protective substances for coating eggs or tough coverings for a batch of eggs called ootheca
e. Spermathecae are tubes or sacs in which sperm can be stored between the time of mating and the time an egg is fertilized.
For males, the reproductive system is the testis, suspended in the body cavity by trachea
e and the fat body. Most male insects have a pair of testes, inside of which are sperm tubes or follicles that are enclosed within a membranous sac. The follicles connect to the vas deferens by the vas efferens, and the two tubular vasa deferentia connect to a median ejaculatory duct that leads to the outside. A portion of the vas deferens is often enlarged to form the seminal vesicle, which stores the sperm before they are discharged into the female. The seminal vesicles have glandular linings that secrete nutrients for nourishment and maintenance of the sperm. The ejaculatory duct is derived from an invagination of the epidermal cells during development and, as a result, has a cuticular lining. The terminal portion of the ejaculatory duct may be sclerotized to form the intromittent organ, the aedeagus. The remainder of the male reproductive system is derived from embryonic mesoderm, except for the germ cells, or spermatogonia, which descend from the primordial pole cells very early during embryogenesis.
s. Instead, the insect respiratory system
uses 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 trachea
(element 8 in numbered diagram). Since oxygen is delivered directly, the circulatory system is not used to carry oxygen, and is therefore greatly reduced. The insect circulatory system has no vein
s or arterie
s, and instead consists of little more than a single, perforated dorsal tube which pulses peristaltically
. Toward the thorax, the dorsal tube (element 14) divides into chambers and acts like the insect's heart. The opposite end of the dorsal tube is like the aorta of the insect circulating the hemolymph
, arthropods' fluid analog of blood
, inside the body cavity. Air is taken in through openings on the sides of the abdomen called spiracle
s.
There are many different patterns of gas exchange
demonstrated by different groups of insects. Gas exchange patterns in insects can range from continuous and diffusive
ventilation, to discontinuous gas exchange
. During continuous gas exchange, oxygen
is taken in and carbon dioxide
is released in a continuous cycle. In discontinuous gas exchange, however, the insect takes in oxygen while it is active and small amounts of carbon dioxide are released when the insect is at rest. Diffusive ventilation is simply a form of continuous gas exchange that occurs by diffusion
rather than physically taking in the oxygen. Some species of insect that are submerged also have adaptations to aid in respiration. As larvae, many insects have gills that can extract oxygen dissolved in water, while others need to rise to the water surface to replenish air supplies which may be held or trapped in special structures.
The majority of insects hatch from egg
s. The fertilization and development takes place inside the egg, enclosed by a shell (chorion
). Some species of insects, like the cockroach Blaptica dubia
, as well as juvenile aphids and tsetse flies, are ovoviviparous
. The eggs of ovoviviparous animals develop entirely inside the female, and then hatch immediately upon being laid. Some other species, such as those in the genus of cockroaches known as Diploptera
, are viviparous, and thus gestate inside the mother and are born alive
. Some insects, like parasitic wasps, show polyembryony
, where a single fertilized egg divides into many and in some cases thousands of separate embryos.
Other developmental and reproductive variations include haplodiploidy, polymorphism
, paedomorphosis or peramorphosis
, sexual dimorphism
, parthenogenesis
and more rarely hermaphroditism. In haplodiploidy, which is a type of sex-determination system
, the offspring's sex is determined by the number of sets of chromosome
s an individual receives. This system is typical in bees and wasps. Polymorphism is the where a species may have different morphs or forms, as in the oblong winged katydid
, which has four different varieties: green, pink, and yellow or tan. Some insects may retain phenotype
s that are normally only seen in juveniles; this is called paedomorphosis. In peramorphosis, an opposite sort of phenomenon, insects take on previously unseen traits after they have matured into adults. Many insects display sexual dimorphism, in which males and females have notably different appearances, such as the moth Orgyia recens
as an exemplar of sexual dimorphism in insects.
Some insects use parthenogenesis
, a process in which the female can reproduce and give birth without having the eggs fertilized by a male
. Many aphids undergo a form of parthenogenesis, called cyclical parthenogenesis, in which they alternate between one or many generations of asexual and sexual reproduction. 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 which they spawn males. However, overall, most individuals are female, which are produced by fertilization. The males are haploid and the females are diploid. More rarely, some insects display hermaphroditism, in which a given individual has both male and female reproductive organs.
Insect life-histories show adaptations to withstand cold and dry conditions. Some temperate region insects are capable of activity during winter, while some others migrate to a warmer climate or go into a state of torpor
. Still other insects have evolved mechanisms of diapause
that allow eggs or pupae to survive these conditions.
in insects is the biological process of development all insects must undergo. There are two forms of metamorphosis: incomplete metamorphosis and complete metamorphosis.
, or incomplete metamorphosis, change gradually by undergoing a series of molt
s. An insect 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. 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, which makes the old cuticle split along predefined weaknesses where the old exocuticle was thinnest. Other arthropods have a much different process and only molt; though must accommodate for the difference in exoskeleton structure and make up with other enzymes.
Immature insects that go through incomplete metamorphosis are called nymph
s or in the case of dragonflies and damselflies as naiads. Nymphs are similar in form to the adult except for the presence of wings, which are not developed until adulthood. With each molt, nymphs grow larger and become more similar in appearance to adult insects.
Holometabolism
, or complete metamorphosis, is where the insect changes all in four stages, an egg
or embryo
, a larva
, a pupa
, and the adult
or imago
. In these species, egg hatches to produce a larva
, which is generally worm-like in form. This worm-like form can be one of several varieties: eruciform (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 and often sealed within a cocoon
. 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. Insects undergo considerable change in form during the pupal stage, and emerge as adults. Butterflies are a well known example of an insects that undergo complete metamorphosis, although most insects use this life cycle. Some insects have evolved this system to hypermetamorphosis
.
Some of the oldest and most successful insect groups, such Endopterygota
, use a system of complete metamorphosis. Strangely though, complete metamorphosis is unique to certain insect orders, like Diptera
, Lepidoptera
, and Hymenoptera
, and no other arthropods undergo it, but incomplete metamorphosis.
. Some insects such as bees can perceive ultraviolet
wavelengths, or detect polarized light, while the antennae
of male moths can detect the pheromone
s of female moths over distances of many kilometers. There is a pronounced tendency for there to be 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. There are a variety of different mechanisms by which insects perceive sound, while the patterns are not universal, insects can generally hear sound if they can produce it. Different insect species can have varying hearing, though most insects can hear only a narrow range of frequencies related to the frequency of the sounds they can produce. Mosquitoes have been found to hear up to 2 MHz., and some grasshoppers can hear up to 50 MHz. Certain predatory and parasitic insects can detect the characteristic sounds made by their prey or hosts, respectively. For instance, some nocturnal moths can perceive the ultrasonic
emissions of bat
s, which helps them avoid predation. Insects that feed on blood have special sensory structures that can detect infrared
emissions, and use them to home in on their hosts.
Some insects display a rudimentary sense of numbers
, such as the solitary wasps that prey upon a single species. The mother wasp lays her eggs in individual cells and provides each egg with a number of live caterpillars on which the young feed when hatched. Some species of wasp always provide five, others twelve, and others as high as twenty-four caterpillars per cell. The number of caterpillars is different among species, but always the same for each sex of larva. The male solitary wasp in the genus Eumenes
is smaller than the female, so the mother of one species supplies him with only five caterpillars; the larger female receives ten caterpillars in her cell.
A few insects, such as members of the families Poduridae and Onychiuridae (Collembola), 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.
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. The colors of emitted light vary from dull blue (Orfelia fultoni, Mycetophilidae) to the familiar greens and the rare reds (Phrixothrix tiemanni, Phengodidae).
Most insects, except some species of cave crickets
, are able to perceive light and dark. Many species have acute vision capable of detecting minute movements. The eyes include simple eyes or ocelli
as well as compound eyes of varying sizes. Many species are able to detect light in the infrared, ultraviolet and the visible light wavelengths. Color vision has been demonstrated in many species and phylogenetic analysis suggests that UV-green-blue trichromacy existed from at least the Devonian
period between 416 and 359 million years ago.
s and crickets, this is achieved by stridulation
. Cicada
s make the loudest sounds among the insects by producing and amplifying sounds with special modifications to their body and musculature. The African cicada
Brevisana brevis
has been measured at 106.7 decibel
s at a distance of 50 cm (19.7 in). Some insects, such as the hawk moths and Hedylid
butterflies, can hear ultrasound
and take evasive action when they sense that they have been detected by bats. Some moths produce ultrasonic clicks that were once thought to have a role in jamming bat echolocation
. The ultrasonic clicks were subsequently found to be produced mostly by unpalatable moths to warn bats, just as warning coloration
s are used against predators that hunt by sight. Some otherwise palatable moths have evolved to mimic
these calls.
Very low sounds are also produced in various species of Coleoptera, Hymenoptera
, Lepidoptera
, Mantodea, and Neuroptera
. These low sounds are simply the sounds made by the insect's movement. Through microscopic stridulatory structures located on the insect's muscles and joints, the normal sounds of the insect moving are amplified and can be used to warn or communicate with other insects. Most sound-making insects also have tympanal organ
s that can perceive airborne sounds. Some species in Hemiptera
, such as the corixids (water boatmen), are known to communicate via underwater sounds. Most insects are also able to sense vibration
s transmitted through surfaces. For example, an insect is caught in a spider web
and struggles to escape. The vibrations it produces are sensed by the spider
, who is alerted to its presence. Through these vibrations, the spider can tell where on the web the insect is located, as well as how big it is.
Communication using surface-borne vibrational signals is more widespread among insects because of size constraints in producing air-borne sounds. 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. The mechanisms of production of vibrational signals are just as diverse as those for producing sound in insects.
Some species use vibrations for communicating within members of the same species, such as to attract mates as in the songs of the shield bug
Nezara viridula
. Vibrations can also be used to communicate between entirely different species; lycaenid (gossamer-winged butterfly) caterpillars which are myrmecophilous (living in a mutualistic association with ants) communicate with ants in this way. The Madagascar hissing cockroach
has the ability to press air through its spiracles to make a hissing noise as a sign of aggression; 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 in close proximity.
s, are often derived from plant metabolites include those meant to attract, repel and provide other kinds of information. Pheromone
s, a type of semiochemical, 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. Allomone
a benefit their producer by the effect they have upon the receiver. Kairomone
s benefit their receiver instead of their producer. Synomones benefit the producer and the receiver. While some chemicals are targeted at individuals of the same species, others are used for communication across species. The use of scents is especially well known to have developed in social insects.
Social insects, such as termite
s, ant
s and many bee
s and wasp
s, are the most familiar species of eusocial
animal. They live together in large well-organized colonies that may be so tightly integrated and genetically similar that the colonies of some species are sometimes considered superorganism
s. It is sometimes argued that the various species of honey bee
are the only invertebrates (and indeed one of the few non-human groups) to have evolved a system of abstract symbolic communication where a behavior is used to represent and convey specific information about something in 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.
Only insects which live in nests or colonies demonstrate any true capacity for fine-scale spatial orientation or homing. This can allow an insect to return unerringly to a single hole a few millimeters in diameter among thousands of apparently identical holes clustered together, after a trip of up to several kilometers' distance. In a phenomenon known as philopatry
, insects that hibernate
have shown the ability to recall a specific location up to a year after last viewing the area of interest. A few insects seasonally migrate large distances between different geographic regions (e.g., the overwintering areas of the Monarch butterfly
).
).
Most insects, however, lead short lives as adults, and rarely interact with one another except to mate or compete for mates. A small number exhibit some form of parental care, where they will at least guard their eggs, and sometimes continue guarding their offspring until adulthood, and possibly even feeding them. Another simple form of parental care is to construct a nest (a burrow or an actual construction, either of which may be simple or complex), store provisions in it, and lay an egg upon those provisions. The adult does not contact the growing offspring, but it nonetheless does provide food. This sort of care is typical for most species of bees and various types of wasps.
s suggest that the wings are from paranotal lobes, or extensions from the insect's exoskeleton called the nota
, called the paranotal theory. Other theories are based on a pleural origin. These theories include suggestions that wings originated from modified gills, spiracular flaps or as from an appendage of the epicoxa. The epicoxal theory suggests the insect wings are modified epicoxal exites, a modified appendage at the base of the legs or coxa
. In the Carboniferous
age, some of the Meganeura
dragonflies had as much as a 50 cm (19.7 in) wide wingspan. The appearance of gigantic insects has been found to be consistent with high atmospheric oxygen. The respiratory system of insects constrains their size, however the high oxygen in the atmosphere allowed larger sizes. The largest flying insects today are much smaller and include several moth species such as the Atlas moth and the White Witch (Thysania agrippina). Insect flight has been a topic of great interest in aerodynamics
due partly to the inability of steady-state theories to explain the lift generated by the tiny wings of insects.
Unlike bird
s, many small insects are swept along by the prevailing winds although many of the larger insects are known to make migrations. Aphid
s are known to be transported long distances by low-level jet stream
s. As such, fine line patterns associated with converging winds within weather radar
imagery, like the WSR-88D radar network, often represent large groups of insects.
gait
. The tripedal gait allows for rapid walking while always having a stable stance and has been studied extensively in cockroach
es. The legs are used in alternate triangles touching the ground. 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 position. When they touch the ground to form a new stable triangle the other legs can be lifted and brought forward in turn and so on. The purest form of the tripedal gait is seen in insects moving at high speeds. However, this type of locomotion is not rigid and insects can adapt a variety of gaits. For example, when moving slowly, turning, or avoiding obstacles, four or more feet may be touching the ground. Insects can also adapt their gait to cope with the loss of one or more limbs.
Cockroaches are among the fastest insect runners and, at full speed, adopt a bipedal run to reach a high velocity in proportion to their body size. As cockroaches move very quickly, they need to be video recorded at several hundred frames per second to reveal their gait. More sedate locomotion is seen in the stick insects or walking sticks (Phasmatodea
). A few insects have evolved to walk on the surface of the water, especially the bugs of the Gerridae
family, commonly known as water striders. A few species of ocean-skaters in the genus Halobates
even live on the surface of open oceans, a habitat that has few insect species.
s. The study of insects and biped
s has a significant impact on possible robotic methods of transport. This may allow new robots to be designed that can traverse terrain
that robots with wheel
s may be unable to handle.
A large number of insects live either parts or the whole of their lives underwater. In many of the more primitive orders of insect, the immature stages are spent in an aquatic environment. Some groups of insects, like certain water beetle
s, have aquatic adults as well.
Many of these species have adaptations to help in under-water locomotion. Water beetles and water bugs have legs adapted into paddle-like structures. Dragonfly naiad
s use jet propulsion, forcibly expelling water out of their rectal chamber. Some species like the water strider
s are capable of walking on the surface of water. They can do this because their claws are not at the tips of the legs as in most insects, but recessed in a special groove further up the leg; this prevents the claws from piercing the water's surface film. Other insects such as the Rove beetle Stenus
are known to emit pygidial gland secretions that reduce surface tension making it possible for them to move on the surface of water by Marangoni propulsion
(also known by the German
term Entspannungsschwimmen).
ary relationships of insects to other animal groups remain unclear. Although more traditionally grouped with millipede
s and centipede
s, evidence has emerged favoring closer evolutionary ties
with crustacean
s. In the Pancrustacea
theory, insects, together with Remipedia
and Malacostraca
, make up a natural clade
. Other terrestrial arthropods, such as centipede
s, millipede
s, scorpion
s and spider
s, are sometimes confused with insects since their body plans can appear similar, sharing (as do all arthropods) a jointed exoskeleton. However, upon closer examination their features differ significantly; most noticeably they do not have the six legs characteristic of adult insects.
A phylogenetic tree of the arthropods and related groups
The higher-level phylogeny of the arthropods continues to be a matter of debate and research. In 2008, researchers at Tufts University
uncovered what they believe is the world's oldest known full-body impression of a primitive flying insect, a 300 million-year-old specimen from the Carboniferous Period. The oldest definitive insect fossil is the Devonian
Rhyniognatha hirsti
, from the 396 million year old Rhynie chert
. It may have superficially resembled a modern-day silverfish
insect. This species already possessed dicondylic mandibles (two articulations in the mandible), a feature associated with winged insects, suggesting that wings may already have evolved at this time. Thus, the first insects probably appeared earlier, in the Silurian
period.
There have been four super radiations of insects: beetle
s (evolved ~300 million years ago), flies
(evolved ~250 million years ago), moth
s and wasp
s (evolved ~150 million years ago). These four groups account for the majority of described species. The flies and moths along with the flea
s evolved from the Mecoptera
.
The origins of insect flight
remain obscure, since the earliest winged insects currently known appear to have been capable fliers. Some extinct insects had an additional pair of winglets attaching to the first segment of the thorax, for a total of three pairs. As of 2009, there is no evidence that suggests that the insects were a particularly successful group of animals before they evolved to have wings.
Late Carboniferous
and Early Permian
insect orders include both extant groups and a number of Paleozoic species, now extinct. During this era, some giant dragonfly-like forms reached wingspans of 55 to 70 cm (21.7 to 27.6 in) making them far larger than any living insect. This gigantism may have been due to higher atmospheric oxygen levels that allowed increased respiratory efficiency relative to today. The lack of flying vertebrates could have been another factor. Most extinct orders of insects developed during the Permian period that began around 270 million years ago. Many of the early groups became extinct during the Permian-Triassic extinction event
, the largest mass extinction in the history of the Earth, around 252 million years ago.
The remarkably successful Hymenoptera
ns appeared as long as 146 million years ago in the Cretaceous
period, but achieved their wide diversity more recently in the Cenozoic
era, which began 66 million years ago. A number of highly successful insect groups evolved in conjunction with flowering plants, a powerful illustration of coevolution.
Many modern insect genera
developed during the Cenozoic. Insects from this period on are often found preserved in amber
, often in perfect condition. The body plan, or morphology
, of such specimens is thus easily compared with modern species. The study of fossilized insects is called paleoentomology
.
, through gradual evolutionary change, insectivory was the next diet type to evolve.
Insects were among the earliest terrestrial herbivore
s and acted as major selection agents on plants. 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. This successful evolutionary pattern has also been utilized by mimic
s. Over time, this has led to complex groups of coevolved species. Conversely, some interactions between plants and insects, like pollination
, are beneficial to both organisms. Coevolution has led to the development of very specific mutualisms in such systems.
! colspan="4" style="background:#E6D09D"| Classification
|-
|
Insects (from Latin
, a calque
of Greek
[], "cut into sections") are a class
of living creatures within the arthropod
s that have a chitin
ous exoskeleton
, a three-part body (head
, thorax
, and abdomen
), three pairs of jointed leg
s, compound eyes, and two antennae
. They are among the most diverse groups of animal
s on the planet, including more than a million described species
and represent more than half of all known living organisms. The number of extant species is estimated at between six and ten million, and potentially represent over 90% of the differing metazoan life forms on Earth. Insects may be found in nearly all environments
, although only a small number of species occur in the oceans, a habitat dominated by another arthropod group, the crustacean
s.
The life cycles of insects vary but most hatch from eggs
. Insect growth is constrained by the inelastic exoskeleton
and development involves a series of molts. The immature stages can differ from the adults in structure, habit and habitat and can include a passive pupa
l stage in those groups that undergo complete metamorphosis. Insects that undergo incomplete metamorphosis
lack a pupal stage and adults develop through a series of nymph
al stages. The higher level relationship of the hexapoda
is unclear. Fossilized insects of enormous size have been found from the Paleozoic Era
, including giant dragonflies with wingspans of 55 to 70 cm (22–28 in). The most diverse insect groups appear to have coevolved with flowering plant
s.
Insects typically move about by walking, flying or occasionally swimming. As it allows for rapid yet stable movement, many insects adopt a tripedal gait in which they walk with their legs touching the ground in alternating triangles. Insects are the only invertebrates to have evolved flight. Many insects spend at least part of their life underwater, with larval adaptations that include gill
s and some adult insects are aquatic and have adaptations for swimming. Some species, like water striders, are capable of walking on the surface of water. Insects are mostly solitary, but some insects, such as certain bee
s, ant
s, and termite
s are social and live in large, well-organized colonies. Some insects, like earwig
s, show maternal care, guarding their eggs and young. Insects can communicate with each other in a variety of ways. Male moth
s can sense the pheromone
s of female moths over distances of many kilometers. Other species communicate with sounds: crickets
stridulate
, or rub their wings together, to attract a mate and repel other males. Lampyridae
in the beetle order Coleoptera
communicate with light.
Humans regard certain insects as pests and attempt to control them using insecticide
s and a host of other techniques. Some insects damage crops by feeding on sap, leaves or fruit
s, a few bite humans and livestock, alive and dead, to feed on blood and some are capable of transmitting disease
s to humans, pets and livestock. Nevertheless, without insects to pollinate flowers, the human race would soon run out of food because many of the crop plants that we rely on would not be able to reproduce. Many other insects are considered ecologically beneficial as predators and a few provide direct economic benefit. Silkworms and bees have been used extensively by humans for the production of silk
and honey
, respectively.
's loan-translation of the Greek
word or "insect" (as in entomology
), which was Aristotle's term for this class of life, also in reference to their "notched" bodies, first documented in English in 1601 in Holland's translation of Pliny. Translations of Aristotle's term also form the usual word for "insect" in Welsh
, Serbo-Croatian
(zareznik, from rezati "cut"), Russian
( nasekomoe, from sekat "cut"), etc.
, Hymenoptera
, and Lepidoptera
. There are almost as many named species of beetle as there are of all other insects added together, or all other non-insects (plant
s and animal
s).
{| class="wikitable" style="text-align: center; width:80%;"
|+ Comparison of the Estimated Number of Species for Vertebrates and the Four Most Species Order
|-
|
! scope="col" | Described Species
! scope="col" | Average description rate (species per year)
! scope="col" | Publication effort
|-
! scope="row" | Coleoptera
| 300,000–400,000
| 2308
| 0.01
|-
! scope="row" | Lepidoptera
| 110,000–120,000
| 642
| 0.03
|-
! scope="row" | Diptera
| 90,000–150,000
| 1048
| 0.04
|-
! scope="row" | Hymenoptera
| 100,000–125,000
| 1196
| 0.02
|-
|}
ed bodies supported by an exoskeleton
, a hard outer covering made mostly of chitin
. The segments of the body are organized into three distinctive but interconnected units, or tagmata
: a head, a thorax
, and an abdomen
. The head supports a pair of sensory antennae
, a pair of compound eyes, and, if present, one to three simple eyes (or ocelli) and three sets of variously modified appendages that form the mouthpart
s. The thorax has six segmented leg
s—one pair each for the prothorax, mesothorax and the metathorax segments making up the thorax—and, if present in the species, two or four wing
s. The abdomen consists of eleven segments, though in a few species of insects these segments may be fused together or reduced in size. The abdomen also contains most of the digestive
, respiratory
, excretory
and reproductive internal structures. There is considerable variation and many adaptations in the body parts of insects especially wings, legs, antenna, mouth-parts etc.
: is enclosed in a hard, heavily sclerotized, unsegmented, exoskeletal head capsule, or epicranium
, which contains most of the sensing organs, including the antennae, ocellus or eyes, and the mouthparts. Out of all the insect orders, Orthoptera displays the most features found in other insects, including the sutures and sclerite
s. here, the vertex
, or the apex (dorsal region), is situated between the compound eyes for insects with a hypognathous and opisthognathous head. In prognathous insects, the vertex is not found between the compound eyes, but rather, where the ocelli
are normally. This is because the primary axis of the head is rotated 90 degrees to become parallel to the primary axis of the body. In some species this region is modified and assumes a different name.
The thorax
: is a segment composed of three sections, the prothorax
, mesothorax
, and the metathorax
. The anterior segment, closest to the head, is the prothorax, with the major features being the first pair of legs and the pronotum. The middle segment is the mesothorax with the major features being the second pair of legs and the anterior wings. The third and most posterior segment, abutting the abdomen, is the metathorax, which features the third pair of legs and the posterior wings. Each segment is dilineated by an intersegmental suture. Each segment has four basic regions. The dorsal surface is called the tergum (or notum) to distinguish them from the abdominal terga. The two lateral regions are called the pleura (singular: pleuron) and the ventral aspect is called the sternum. In turn, the notum of the prothorax is called the pronotum, the notum for the mesothorax is called the mesonotum and the notum for the metathorax is called the metanotum. Continuing with this logic, there is also the mesopleura and metapleura as well as the mesosternum and metasternum.
The abdomen
: is a the last segment of the insect, which typically consists of 11–12 segments and is less strongly sclerotized than the head or thorax. Each segment of the abdomen is represented by a sclerotized tergum, sternum, and perhaps a pleurite. Terga are separated from each other and from the adjacent sterna or pleura by a membrane. Spiracles are located in the pleural area. Variation of this ground plan includes the fusion of terga or terga and sterna to form continuous dorsal or ventral shields or a conical tube. Some insects bear a sclerite in the pleural area called a laterotergite. Ventral sclerites are sometimes called laterosternites. During the embryonic stage of many insects and the postembryonic stage of primitive insects, 11 abdominal segments are present. In modern insects there is a tendency toward reduction in the number of the abdominal segments, but the primitive number of 11 is maintained during embryogenesis.Variation in abdominal segment number is considerable. If the Apterygota are considered to be indicative of the ground plan for pterygotes, confusion reigns: adult Protura have 12 segments, Collembola have 6. The orthopteran family Acrididae has 11 segments, and a fossil specimen of Zoraptera has a 10-segmented abdomen.
, and a lower layer called the procuticle. The procuticle is chitinous and much thicker than the epicuticle and has two layers: an outer layer known as the exocuticle and an inner layer known as the endocuticle. The tough and flexible endocuticle is built from numerous layers of fibrous chitin and proteins, criss-crossing each others in a sandwich pattern, while the exocuticle is rigid and hardened. The exocuticle is greatly reduced in many soft-bodied insects (e.g., caterpillar
s), especially during their larva
l stages.
Insects are the only invertebrate
s to have developed active flight capability, and this has played an important role in their success. Their muscles are able to contract multiple times for each single nerve impulse, allowing the wings to beat faster than would ordinarily be possible. Having their muscles attached to their exoskeletons is more efficient and allows more muscle connections; crustacean
s also use the same method, though all spider
s use hydraulic pressure to extend their legs, a system inherited from their pre-arthropod ancestors. Unlike insects, though, most aquatic crustaceans are biomineralized with calcium carbonate
extracted from the water.
of an insect can be divided into a brain and a ventral nerve cord
. The head capsule is made up of six fused segments, each with 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
.
The thoracic segments have one ganglion on each side, which are connected into a pair, one pair per segment. This arrangement is also seen in the abdomen but only in the first eight segments. Many species of insects have reduced numbers of ganglia due to fusion or reduction. Some cockroaches have just six ganglia in the abdomen, whereas the wasp Vespa crabro has only two in the thorax and three in the abdomen. Some insects, like the house fly Musca domestica
, have all the body ganglia fused into a single large thoracic ganglion.
At least a few insects have nociceptor
s, cells that detect and transmit sensations of pain
. This was discovered in 2003 by studying the variation in reactions of larvae
of the common fruitfly Drosophila
to the touch of a heated probe and an unheated one. The larvae reacted to the touch of the heated probe with a stereotypical rolling behavior that was not exhibited when the larvae were touched by the unheated probe. Although nociception has been demonstrated in insects, there is not a consensus that insects feel pain consciously but see Pain in invertebrates
.
s and other complex substances like protein
s, polysaccharide
s, fat
s, and nucleic acid
s. These macromolecules must be broken down by catabolic reaction
s into smaller molecules like amino acid
s and simple sugars before being used by cells of the body for energy, growth, or reproduction. This break-down process is known as digestion.
The main structure of an insect's digestive system is a long enclosed tube called the alimentary canal, which runs lengthwise through the body. The alimentary canal directs food unidirectionally from the mouth
to the anus
. It has three sections, each of which performs a different process of digestion. In addition to the alimentary canal, insects also have paired salivary glands and salivary reservoirs. These structures usually reside in the thorax, adjacent to the foregut.
The salivary gland
s (element 30 in numbered diagram) in an insect's mouth produce saliva. The salivary ducts lead from the glands to the reservoirs and then forward through the head to an opening called the salivarium, located behind the hypopharynx. By moving its mouthparts (element 32 in numbered diagram) the insect can mix its food with saliva. The mixture of saliva and food then travels through the salivary tubes into the mouth, where it begins to break down. Some insects, like flies
, have extra-oral digestion
. Insects using extra-oral digestion expel digestive enzymes onto their food to break it down. This strategy allows insects to extract a significant proportion of the available nutrients from the food source. The gut is where almost all of insects' digestion takes place. It can be divided into the foregut
, midgut
and hindgut
.
The first section of the alimentary canal is the foregut
(element 27 in numbered diagram), or stomodaeum. The foregut is lined with a cuticular lining made of chitin
and protein
s as protection from tough food. The foregut includes the buccal cavity (mouth), pharynx
, esophagus
, and Crop
and proventriculus
(any part may be highly modified) which both store food and signify when to continue passing onward to the midgut.
Digestion starts in buccal cavity (mouth) as partially chewed food is broken down by saliva from the salivary glands. As the salivary glands produce fluid and carbohydrate-digesting enzyme
s (mostly amylase
s), strong muscles in the pharynx pump fluid into the buccal cavity, lubricating the food like the salivarium does, and helping blood feeders, and xylem and phloem feeders.
From there, the pharynx passes food to the esophagus, which could be just a simple tube passing it on to the crop and proventriculus, and then onward to the midgut, as in most insects. Alternately, the foregut may expand into a very enlarged crop and proventriculus, or the crop could just be a diverticulum
, or fluid filled structure, as in some Diptera species.
Once food leaves the crop, it passes to the midgut
(element 13 in numbered diagram), also known as the mesenteron, where the majority of digestion takes place. Microscopic projections from the midgut wall, called microvilli
, increase the surface area of the wall and allow more nutrients to be absorbed; they tend to be close to the origin of the midgut. In some insects, the role of the microvilli and where they are located may vary. For example, specialized microvilli producing digestive enzymes may more likely be near the end of the midgut, and absorption near the origin or beginning of the midgut.
In the hindgut
(element 16 in numbered diagram), or proctodaeum, undigested food particles are joined by uric acid
to form fecal pellets. The rectum absorbs 90% of the water in these fecal pellets, and the dry pellet is then eliminated through the anus (element 17), completing the process of digestion. The uric acid is formed using hemolymph waste products diffused from the Malpighian tubule
s (element 20). It is then emptied directly into the alimentary canal, at the junction between the midgut and hindgut. The number of Malpighian tubules possessed by a given insect varies between species, ranging from only two tubules in some insects to over 100 tubules in others.
s (element 30 in numbered diagram) in an insect's mouth produce saliva. The salivary ducts lead from the glands to the reservoirs and then forward through the head to an opening called the salivarium, located behind the hypopharynx. By moving its mouthparts (element 32 in numbered diagram) the insect can mix its food with saliva. The mixture of saliva and food then travels through the salivary tubes into the mouth, where it begins to break down. Some insects, like flies
, have extra-oral digestion
. Insects using extra-oral digestion expel digestive enzymes onto their food to break it down. This strategy allows insects to extract a significant proportion of the available nutrients from the food source.
e, and ducts connecting these parts. The ovaries are made up of a number of egg tubes, called ovarioles
, which vary in size and number by species. The number of eggs that the insect is able to make vary by the number of ovarioles with the rate that eggs can be develop being also influenced by ovariole design. Female insects are able make eggs, receive and store sperm, manipulate sperm from different males, and lay eggs. Accessory glands or glandular parts of the oviducts produce a variety of substances for sperm maintenance, transport, and fertilization, as well as for protection of eggs. They can produce glue and protective substances for coating eggs or tough coverings for a batch of eggs called ootheca
e. Spermathecae are tubes or sacs in which sperm can be stored between the time of mating and the time an egg is fertilized.
For males, the reproductive system is the testis, suspended in the body cavity by trachea
e and the fat body. Most male insects have a pair of testes, inside of which are sperm tubes or follicles that are enclosed within a membranous sac. The follicles connect to the vas deferens by the vas efferens, and the two tubular vasa deferentia connect to a median ejaculatory duct that leads to the outside. A portion of the vas deferens is often enlarged to form the seminal vesicle, which stores the sperm before they are discharged into the female. The seminal vesicles have glandular linings that secrete nutrients for nourishment and maintenance of the sperm. The ejaculatory duct is derived from an invagination of the epidermal cells during development and, as a result, has a cuticular lining. The terminal portion of the ejaculatory duct may be sclerotized to form the intromittent organ, the aedeagus. The remainder of the male reproductive system is derived from embryonic mesoderm, except for the germ cells, or spermatogonia, which descend from the primordial pole cells very early during embryogenesis.
s. Instead, the insect respiratory system
uses 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 trachea
(element 8 in numbered diagram). Since oxygen is delivered directly, the circulatory system is not used to carry oxygen, and is therefore greatly reduced. The insect circulatory system has no vein
s or arterie
s, and instead consists of little more than a single, perforated dorsal tube which pulses peristaltically
. Toward the thorax, the dorsal tube (element 14) divides into chambers and acts like the insect's heart. The opposite end of the dorsal tube is like the aorta of the insect circulating the hemolymph
, arthropods' fluid analog of blood
, inside the body cavity. Air is taken in through openings on the sides of the abdomen called spiracle
s.
There are many different patterns of gas exchange
demonstrated by different groups of insects. Gas exchange patterns in insects can range from continuous and diffusive
ventilation, to discontinuous gas exchange
. During continuous gas exchange, oxygen
is taken in and carbon dioxide
is released in a continuous cycle. In discontinuous gas exchange, however, the insect takes in oxygen while it is active and small amounts of carbon dioxide are released when the insect is at rest. Diffusive ventilation is simply a form of continuous gas exchange that occurs by diffusion
rather than physically taking in the oxygen. Some species of insect that are submerged also have adaptations to aid in respiration. As larvae, many insects have gills that can extract oxygen dissolved in water, while others need to rise to the water surface to replenish air supplies which may be held or trapped in special structures.
The majority of insects hatch from egg
s. The fertilization and development takes place inside the egg, enclosed by a shell (chorion
). Some species of insects, like the cockroach Blaptica dubia
, as well as juvenile aphids and tsetse flies, are ovoviviparous
. The eggs of ovoviviparous animals develop entirely inside the female, and then hatch immediately upon being laid. Some other species, such as those in the genus of cockroaches known as Diploptera
, are viviparous, and thus gestate inside the mother and are born alive
. Some insects, like parasitic wasps, show polyembryony
, where a single fertilized egg divides into many and in some cases thousands of separate embryos.
Other developmental and reproductive variations include haplodiploidy, polymorphism
, paedomorphosis or peramorphosis
, sexual dimorphism
, parthenogenesis
and more rarely hermaphroditism. In haplodiploidy, which is a type of sex-determination system
, the offspring's sex is determined by the number of sets of chromosome
s an individual receives. This system is typical in bees and wasps. Polymorphism is the where a species may have different morphs or forms, as in the oblong winged katydid
, which has four different varieties: green, pink, and yellow or tan. Some insects may retain phenotype
s that are normally only seen in juveniles; this is called paedomorphosis. In peramorphosis, an opposite sort of phenomenon, insects take on previously unseen traits after they have matured into adults. Many insects display sexual dimorphism, in which males and females have notably different appearances, such as the moth Orgyia recens
as an exemplar of sexual dimorphism in insects.
Some insects use parthenogenesis
, a process in which the female can reproduce and give birth without having the eggs fertilized by a male
. Many aphids undergo a form of parthenogenesis, called cyclical parthenogenesis, in which they alternate between one or many generations of asexual and sexual reproduction. 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 which they spawn males. However, overall, most individuals are female, which are produced by fertilization. The males are haploid and the females are diploid. More rarely, some insects display hermaphroditism, in which a given individual has both male and female reproductive organs.
Insect life-histories show adaptations to withstand cold and dry conditions. Some temperate region insects are capable of activity during winter, while some others migrate to a warmer climate or go into a state of torpor
. Still other insects have evolved mechanisms of diapause
that allow eggs or pupae to survive these conditions.
in insects is the biological process of development all insects must undergo. There are two forms of metamorphosis: incomplete metamorphosis and complete metamorphosis.
, or incomplete metamorphosis, change gradually by undergoing a series of molt
s. An insect 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. 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, which makes the old cuticle split along predefined weaknesses where the old exocuticle was thinnest. Other arthropods have a much different process and only molt; though must accommodate for the difference in exoskeleton structure and make up with other enzymes.
Immature insects that go through incomplete metamorphosis are called nymph
s or in the case of dragonflies and damselflies as naiads. Nymphs are similar in form to the adult except for the presence of wings, which are not developed until adulthood. With each molt, nymphs grow larger and become more similar in appearance to adult insects.
Holometabolism
, or complete metamorphosis, is where the insect changes all in four stages, an egg
or embryo
, a larva
, a pupa
, and the adult
or imago
. In these species, egg hatches to produce a larva
, which is generally worm-like in form. This worm-like form can be one of several varieties: eruciform (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 and often sealed within a cocoon
. 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. Insects undergo considerable change in form during the pupal stage, and emerge as adults. Butterflies are a well known example of an insects that undergo complete metamorphosis, although most insects use this life cycle. Some insects have evolved this system to hypermetamorphosis
.
Some of the oldest and most successful insect groups, such Endopterygota
, use a system of complete metamorphosis. Strangely though, complete metamorphosis is unique to certain insect orders, like Diptera
, Lepidoptera
, and Hymenoptera
, and no other arthropods undergo it, but incomplete metamorphosis.
. Some insects such as bees can perceive ultraviolet
wavelengths, or detect polarized light, while the antennae
of male moths can detect the pheromone
s of female moths over distances of many kilometers. There is a pronounced tendency for there to be 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. There are a variety of different mechanisms by which insects perceive sound, while the patterns are not universal, insects can generally hear sound if they can produce it. Different insect species can have varying hearing, though most insects can hear only a narrow range of frequencies related to the frequency of the sounds they can produce. Mosquitoes have been found to hear up to 2 MHz., and some grasshoppers can hear up to 50 MHz. Certain predatory and parasitic insects can detect the characteristic sounds made by their prey or hosts, respectively. For instance, some nocturnal moths can perceive the ultrasonic
emissions of bat
s, which helps them avoid predation. Insects that feed on blood have special sensory structures that can detect infrared
emissions, and use them to home in on their hosts.
Some insects display a rudimentary sense of numbers
, such as the solitary wasps that prey upon a single species. The mother wasp lays her eggs in individual cells and provides each egg with a number of live caterpillars on which the young feed when hatched. Some species of wasp always provide five, others twelve, and others as high as twenty-four caterpillars per cell. The number of caterpillars is different among species, but always the same for each sex of larva. The male solitary wasp in the genus Eumenes
is smaller than the female, so the mother of one species supplies him with only five caterpillars; the larger female receives ten caterpillars in her cell.
A few insects, such as members of the families Poduridae and Onychiuridae (Collembola), 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.
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. The colors of emitted light vary from dull blue (Orfelia fultoni, Mycetophilidae) to the familiar greens and the rare reds (Phrixothrix tiemanni, Phengodidae).
Most insects, except some species of cave crickets
, are able to perceive light and dark. Many species have acute vision capable of detecting minute movements. The eyes include simple eyes or ocelli
as well as compound eyes of varying sizes. Many species are able to detect light in the infrared, ultraviolet and the visible light wavelengths. Color vision has been demonstrated in many species and phylogenetic analysis suggests that UV-green-blue trichromacy existed from at least the Devonian
period between 416 and 359 million years ago.
s and crickets, this is achieved by stridulation
. Cicada
s make the loudest sounds among the insects by producing and amplifying sounds with special modifications to their body and musculature. The African cicada
Brevisana brevis
has been measured at 106.7 decibel
s at a distance of 50 cm (19.7 in). Some insects, such as the hawk moths and Hedylid
butterflies, can hear ultrasound
and take evasive action when they sense that they have been detected by bats. Some moths produce ultrasonic clicks that were once thought to have a role in jamming bat echolocation
. The ultrasonic clicks were subsequently found to be produced mostly by unpalatable moths to warn bats, just as warning coloration
s are used against predators that hunt by sight. Some otherwise palatable moths have evolved to mimic
these calls.
Very low sounds are also produced in various species of Coleoptera, Hymenoptera
, Lepidoptera
, Mantodea, and Neuroptera
. These low sounds are simply the sounds made by the insect's movement. Through microscopic stridulatory structures located on the insect's muscles and joints, the normal sounds of the insect moving are amplified and can be used to warn or communicate with other insects. Most sound-making insects also have tympanal organ
s that can perceive airborne sounds. Some species in Hemiptera
, such as the corixids (water boatmen), are known to communicate via underwater sounds. Most insects are also able to sense vibration
s transmitted through surfaces. For example, an insect is caught in a spider web
and struggles to escape. The vibrations it produces are sensed by the spider
, who is alerted to its presence. Through these vibrations, the spider can tell where on the web the insect is located, as well as how big it is.
Communication using surface-borne vibrational signals is more widespread among insects because of size constraints in producing air-borne sounds. 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. The mechanisms of production of vibrational signals are just as diverse as those for producing sound in insects.
Some species use vibrations for communicating within members of the same species, such as to attract mates as in the songs of the shield bug
Nezara viridula
. Vibrations can also be used to communicate between entirely different species; lycaenid (gossamer-winged butterfly) caterpillars which are myrmecophilous (living in a mutualistic association with ants) communicate with ants in this way. The Madagascar hissing cockroach
has the ability to press air through its spiracles to make a hissing noise as a sign of aggression; 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 in close proximity.
s, are often derived from plant metabolites include those meant to attract, repel and provide other kinds of information. Pheromone
s, a type of semiochemical, 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. Allomone
a benefit their producer by the effect they have upon the receiver. Kairomone
s benefit their receiver instead of their producer. Synomones benefit the producer and the receiver. While some chemicals are targeted at individuals of the same species, others are used for communication across species. The use of scents is especially well known to have developed in social insects.
Social insects, such as termite
s, ant
s and many bee
s and wasp
s, are the most familiar species of eusocial
animal. They live together in large well-organized colonies that may be so tightly integrated and genetically similar that the colonies of some species are sometimes considered superorganism
s. It is sometimes argued that the various species of honey bee
are the only invertebrates (and indeed one of the few non-human groups) to have evolved a system of abstract symbolic communication where a behavior is used to represent and convey specific information about something in 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.
Only insects which live in nests or colonies demonstrate any true capacity for fine-scale spatial orientation or homing. This can allow an insect to return unerringly to a single hole a few millimeters in diameter among thousands of apparently identical holes clustered together, after a trip of up to several kilometers' distance. In a phenomenon known as philopatry
, insects that hibernate
have shown the ability to recall a specific location up to a year after last viewing the area of interest. A few insects seasonally migrate large distances between different geographic regions (e.g., the overwintering areas of the Monarch butterfly
).
).
Most insects, however, lead short lives as adults, and rarely interact with one another except to mate or compete for mates. A small number exhibit some form of parental care, where they will at least guard their eggs, and sometimes continue guarding their offspring until adulthood, and possibly even feeding them. Another simple form of parental care is to construct a nest (a burrow or an actual construction, either of which may be simple or complex), store provisions in it, and lay an egg upon those provisions. The adult does not contact the growing offspring, but it nonetheless does provide food. This sort of care is typical for most species of bees and various types of wasps.
s suggest that the wings are from paranotal lobes, or extensions from the insect's exoskeleton called the nota
, called the paranotal theory. Other theories are based on a pleural origin. These theories include suggestions that wings originated from modified gills, spiracular flaps or as from an appendage of the epicoxa. The epicoxal theory suggests the insect wings are modified epicoxal exites, a modified appendage at the base of the legs or coxa
. In the Carboniferous
age, some of the Meganeura
dragonflies had as much as a 50 cm (19.7 in) wide wingspan. The appearance of gigantic insects has been found to be consistent with high atmospheric oxygen. The respiratory system of insects constrains their size, however the high oxygen in the atmosphere allowed larger sizes. The largest flying insects today are much smaller and include several moth species such as the Atlas moth and the White Witch (Thysania agrippina). Insect flight has been a topic of great interest in aerodynamics
due partly to the inability of steady-state theories to explain the lift generated by the tiny wings of insects.
Unlike bird
s, many small insects are swept along by the prevailing winds although many of the larger insects are known to make migrations. Aphid
s are known to be transported long distances by low-level jet stream
s. As such, fine line patterns associated with converging winds within weather radar
imagery, like the WSR-88D radar network, often represent large groups of insects.
gait
. The tripedal gait allows for rapid walking while always having a stable stance and has been studied extensively in cockroach
es. The legs are used in alternate triangles touching the ground. 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 position. When they touch the ground to form a new stable triangle the other legs can be lifted and brought forward in turn and so on. The purest form of the tripedal gait is seen in insects moving at high speeds. However, this type of locomotion is not rigid and insects can adapt a variety of gaits. For example, when moving slowly, turning, or avoiding obstacles, four or more feet may be touching the ground. Insects can also adapt their gait to cope with the loss of one or more limbs.
Cockroaches are among the fastest insect runners and, at full speed, adopt a bipedal run to reach a high velocity in proportion to their body size. As cockroaches move very quickly, they need to be video recorded at several hundred frames per second to reveal their gait. More sedate locomotion is seen in the stick insects or walking sticks (Phasmatodea
). A few insects have evolved to walk on the surface of the water, especially the bugs of the Gerridae
family, commonly known as water striders. A few species of ocean-skaters in the genus Halobates
even live on the surface of open oceans, a habitat that has few insect species.
s. The study of insects and biped
s has a significant impact on possible robotic methods of transport. This may allow new robots to be designed that can traverse terrain
that robots with wheel
s may be unable to handle.
A large number of insects live either parts or the whole of their lives underwater. In many of the more primitive orders of insect, the immature stages are spent in an aquatic environment. Some groups of insects, like certain water beetle
s, have aquatic adults as well.
Many of these species have adaptations to help in under-water locomotion. Water beetles and water bugs have legs adapted into paddle-like structures. Dragonfly naiad
s use jet propulsion, forcibly expelling water out of their rectal chamber. Some species like the water strider
s are capable of walking on the surface of water. They can do this because their claws are not at the tips of the legs as in most insects, but recessed in a special groove further up the leg; this prevents the claws from piercing the water's surface film. Other insects such as the Rove beetle Stenus
are known to emit pygidial gland secretions that reduce surface tension making it possible for them to move on the surface of water by Marangoni propulsion
(also known by the German
term Entspannungsschwimmen).
ary relationships of insects to other animal groups remain unclear. Although more traditionally grouped with millipede
s and centipede
s, evidence has emerged favoring closer evolutionary ties
with crustacean
s. In the Pancrustacea
theory, insects, together with Remipedia
and Malacostraca
, make up a natural clade
. Other terrestrial arthropods, such as centipede
s, millipede
s, scorpion
s and spider
s, are sometimes confused with insects since their body plans can appear similar, sharing (as do all arthropods) a jointed exoskeleton. However, upon closer examination their features differ significantly; most noticeably they do not have the six legs characteristic of adult insects.
A phylogenetic tree of the arthropods and related groups
The higher-level phylogeny of the arthropods continues to be a matter of debate and research. In 2008, researchers at Tufts University
uncovered what they believe is the world's oldest known full-body impression of a primitive flying insect, a 300 million-year-old specimen from the Carboniferous Period. The oldest definitive insect fossil is the Devonian
Rhyniognatha hirsti
, from the 396 million year old Rhynie chert
. It may have superficially resembled a modern-day silverfish
insect. This species already possessed dicondylic mandibles (two articulations in the mandible), a feature associated with winged insects, suggesting that wings may already have evolved at this time. Thus, the first insects probably appeared earlier, in the Silurian
period.
There have been four super radiations of insects: beetle
s (evolved ~300 million years ago), flies
(evolved ~250 million years ago), moth
s and wasp
s (evolved ~150 million years ago). These four groups account for the majority of described species. The flies and moths along with the flea
s evolved from the Mecoptera
.
The origins of insect flight
remain obscure, since the earliest winged insects currently known appear to have been capable fliers. Some extinct insects had an additional pair of winglets attaching to the first segment of the thorax, for a total of three pairs. As of 2009, there is no evidence that suggests that the insects were a particularly successful group of animals before they evolved to have wings.
Late Carboniferous
and Early Permian
insect orders include both extant groups and a number of Paleozoic species, now extinct. During this era, some giant dragonfly-like forms reached wingspans of 55 to 70 cm (21.7 to 27.6 in) making them far larger than any living insect. This gigantism may have been due to higher atmospheric oxygen levels that allowed increased respiratory efficiency relative to today. The lack of flying vertebrates could have been another factor. Most extinct orders of insects developed during the Permian period that began around 270 million years ago. Many of the early groups became extinct during the Permian-Triassic extinction event
, the largest mass extinction in the history of the Earth, around 252 million years ago.
The remarkably successful Hymenoptera
ns appeared as long as 146 million years ago in the Cretaceous
period, but achieved their wide diversity more recently in the Cenozoic
era, which began 66 million years ago. A number of highly successful insect groups evolved in conjunction with flowering plants, a powerful illustration of coevolution.
Many modern insect genera
developed during the Cenozoic. Insects from this period on are often found preserved in amber
, often in perfect condition. The body plan, or morphology
, of such specimens is thus easily compared with modern species. The study of fossilized insects is called paleoentomology
.
, through gradual evolutionary change, insectivory was the next diet type to evolve.
Insects were among the earliest terrestrial herbivore
s and acted as major selection agents on plants. 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. This successful evolutionary pattern has also been utilized by mimic
s. Over time, this has led to complex groups of coevolved species. Conversely, some interactions between plants and insects, like pollination
, are beneficial to both organisms. Coevolution has led to the development of very specific mutualisms in such systems.
! colspan="4" style="background:#E6D09D"| Classification
|-
|
Insects (from Latin
, a calque
of Greek
[], "cut into sections") are a class
of living creatures within the arthropod
s that have a chitin
ous exoskeleton
, a three-part body (head
, thorax
, and abdomen
), three pairs of jointed leg
s, compound eyes, and two antennae
. They are among the most diverse groups of animal
s on the planet, including more than a million described species
and represent more than half of all known living organisms. The number of extant species is estimated at between six and ten million, and potentially represent over 90% of the differing metazoan life forms on Earth. Insects may be found in nearly all environments
, although only a small number of species occur in the oceans, a habitat dominated by another arthropod group, the crustacean
s.
The life cycles of insects vary but most hatch from eggs
. Insect growth is constrained by the inelastic exoskeleton
and development involves a series of molts. The immature stages can differ from the adults in structure, habit and habitat and can include a passive pupa
l stage in those groups that undergo complete metamorphosis. Insects that undergo incomplete metamorphosis
lack a pupal stage and adults develop through a series of nymph
al stages. The higher level relationship of the hexapoda
is unclear. Fossilized insects of enormous size have been found from the Paleozoic Era
, including giant dragonflies with wingspans of 55 to 70 cm (22–28 in). The most diverse insect groups appear to have coevolved with flowering plant
s.
Insects typically move about by walking, flying or occasionally swimming. As it allows for rapid yet stable movement, many insects adopt a tripedal gait in which they walk with their legs touching the ground in alternating triangles. Insects are the only invertebrates to have evolved flight. Many insects spend at least part of their life underwater, with larval adaptations that include gill
s and some adult insects are aquatic and have adaptations for swimming. Some species, like water striders, are capable of walking on the surface of water. Insects are mostly solitary, but some insects, such as certain bee
s, ant
s, and termite
s are social and live in large, well-organized colonies. Some insects, like earwig
s, show maternal care, guarding their eggs and young. Insects can communicate with each other in a variety of ways. Male moth
s can sense the pheromone
s of female moths over distances of many kilometers. Other species communicate with sounds: crickets
stridulate
, or rub their wings together, to attract a mate and repel other males. Lampyridae
in the beetle order Coleoptera
communicate with light.
Humans regard certain insects as pests and attempt to control them using insecticide
s and a host of other techniques. Some insects damage crops by feeding on sap, leaves or fruit
s, a few bite humans and livestock, alive and dead, to feed on blood and some are capable of transmitting disease
s to humans, pets and livestock. Nevertheless, without insects to pollinate flowers, the human race would soon run out of food because many of the crop plants that we rely on would not be able to reproduce. Many other insects are considered ecologically beneficial as predators and a few provide direct economic benefit. Silkworms and bees have been used extensively by humans for the production of silk
and honey
, respectively.
's loan-translation of the Greek
word or "insect" (as in entomology
), which was Aristotle's term for this class of life, also in reference to their "notched" bodies, first documented in English in 1601 in Holland's translation of Pliny. Translations of Aristotle's term also form the usual word for "insect" in Welsh
, Serbo-Croatian
(zareznik, from rezati "cut"), Russian
( nasekomoe, from sekat "cut"), etc.
, Hymenoptera
, and Lepidoptera
. There are almost as many named species of beetle as there are of all other insects added together, or all other non-insects (plant
s and animal
s).
{| class="wikitable" style="text-align: center; width:80%;"
|+ Comparison of the Estimated Number of Species for Vertebrates and the Four Most Species Order
|-
|
! scope="col" | Described Species
! scope="col" | Average description rate (species per year)
! scope="col" | Publication effort
|-
! scope="row" | Coleoptera
| 300,000–400,000
| 2308
| 0.01
|-
! scope="row" | Lepidoptera
| 110,000–120,000
| 642
| 0.03
|-
! scope="row" | Diptera
| 90,000–150,000
| 1048
| 0.04
|-
! scope="row" | Hymenoptera
| 100,000–125,000
| 1196
| 0.02
|-
|}
ed bodies supported by an exoskeleton
, a hard outer covering made mostly of chitin
. The segments of the body are organized into three distinctive but interconnected units, or tagmata
: a head, a thorax
, and an abdomen
. The head supports a pair of sensory antennae
, a pair of compound eyes, and, if present, one to three simple eyes (or ocelli) and three sets of variously modified appendages that form the mouthpart
s. The thorax has six segmented leg
s—one pair each for the prothorax, mesothorax and the metathorax segments making up the thorax—and, if present in the species, two or four wing
s. The abdomen consists of eleven segments, though in a few species of insects these segments may be fused together or reduced in size. The abdomen also contains most of the digestive
, respiratory
, excretory
and reproductive internal structures. There is considerable variation and many adaptations in the body parts of insects especially wings, legs, antenna, mouth-parts etc.
: is enclosed in a hard, heavily sclerotized, unsegmented, exoskeletal head capsule, or epicranium
, which contains most of the sensing organs, including the antennae, ocellus or eyes, and the mouthparts. Out of all the insect orders, Orthoptera displays the most features found in other insects, including the sutures and sclerite
s. here, the vertex
, or the apex (dorsal region), is situated between the compound eyes for insects with a hypognathous and opisthognathous head. In prognathous insects, the vertex is not found between the compound eyes, but rather, where the ocelli
are normally. This is because the primary axis of the head is rotated 90 degrees to become parallel to the primary axis of the body. In some species this region is modified and assumes a different name.
The thorax
: is a segment composed of three sections, the prothorax
, mesothorax
, and the metathorax
. The anterior segment, closest to the head, is the prothorax, with the major features being the first pair of legs and the pronotum. The middle segment is the mesothorax with the major features being the second pair of legs and the anterior wings. The third and most posterior segment, abutting the abdomen, is the metathorax, which features the third pair of legs and the posterior wings. Each segment is dilineated by an intersegmental suture. Each segment has four basic regions. The dorsal surface is called the tergum (or notum) to distinguish them from the abdominal terga. The two lateral regions are called the pleura (singular: pleuron) and the ventral aspect is called the sternum. In turn, the notum of the prothorax is called the pronotum, the notum for the mesothorax is called the mesonotum and the notum for the metathorax is called the metanotum. Continuing with this logic, there is also the mesopleura and metapleura as well as the mesosternum and metasternum.
The abdomen
: is a the last segment of the insect, which typically consists of 11–12 segments and is less strongly sclerotized than the head or thorax. Each segment of the abdomen is represented by a sclerotized tergum, sternum, and perhaps a pleurite. Terga are separated from each other and from the adjacent sterna or pleura by a membrane. Spiracles are located in the pleural area. Variation of this ground plan includes the fusion of terga or terga and sterna to form continuous dorsal or ventral shields or a conical tube. Some insects bear a sclerite in the pleural area called a laterotergite. Ventral sclerites are sometimes called laterosternites. During the embryonic stage of many insects and the postembryonic stage of primitive insects, 11 abdominal segments are present. In modern insects there is a tendency toward reduction in the number of the abdominal segments, but the primitive number of 11 is maintained during embryogenesis.Variation in abdominal segment number is considerable. If the Apterygota are considered to be indicative of the ground plan for pterygotes, confusion reigns: adult Protura have 12 segments, Collembola have 6. The orthopteran family Acrididae has 11 segments, and a fossil specimen of Zoraptera has a 10-segmented abdomen.
, and a lower layer called the procuticle. The procuticle is chitinous and much thicker than the epicuticle and has two layers: an outer layer known as the exocuticle and an inner layer known as the endocuticle. The tough and flexible endocuticle is built from numerous layers of fibrous chitin and proteins, criss-crossing each others in a sandwich pattern, while the exocuticle is rigid and hardened. The exocuticle is greatly reduced in many soft-bodied insects (e.g., caterpillar
s), especially during their larva
l stages.
Insects are the only invertebrate
s to have developed active flight capability, and this has played an important role in their success. Their muscles are able to contract multiple times for each single nerve impulse, allowing the wings to beat faster than would ordinarily be possible. Having their muscles attached to their exoskeletons is more efficient and allows more muscle connections; crustacean
s also use the same method, though all spider
s use hydraulic pressure to extend their legs, a system inherited from their pre-arthropod ancestors. Unlike insects, though, most aquatic crustaceans are biomineralized with calcium carbonate
extracted from the water.
of an insect can be divided into a brain and a ventral nerve cord
. The head capsule is made up of six fused segments, each with 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
.
The thoracic segments have one ganglion on each side, which are connected into a pair, one pair per segment. This arrangement is also seen in the abdomen but only in the first eight segments. Many species of insects have reduced numbers of ganglia due to fusion or reduction. Some cockroaches have just six ganglia in the abdomen, whereas the wasp Vespa crabro has only two in the thorax and three in the abdomen. Some insects, like the house fly Musca domestica
, have all the body ganglia fused into a single large thoracic ganglion.
At least a few insects have nociceptor
s, cells that detect and transmit sensations of pain
. This was discovered in 2003 by studying the variation in reactions of larvae
of the common fruitfly Drosophila
to the touch of a heated probe and an unheated one. The larvae reacted to the touch of the heated probe with a stereotypical rolling behavior that was not exhibited when the larvae were touched by the unheated probe. Although nociception has been demonstrated in insects, there is not a consensus that insects feel pain consciously but see Pain in invertebrates
.
s and other complex substances like protein
s, polysaccharide
s, fat
s, and nucleic acid
s. These macromolecules must be broken down by catabolic reaction
s into smaller molecules like amino acid
s and simple sugars before being used by cells of the body for energy, growth, or reproduction. This break-down process is known as digestion.
The main structure of an insect's digestive system is a long enclosed tube called the alimentary canal, which runs lengthwise through the body. The alimentary canal directs food unidirectionally from the mouth
to the anus
. It has three sections, each of which performs a different process of digestion. In addition to the alimentary canal, insects also have paired salivary glands and salivary reservoirs. These structures usually reside in the thorax, adjacent to the foregut.
The salivary gland
s (element 30 in numbered diagram) in an insect's mouth produce saliva. The salivary ducts lead from the glands to the reservoirs and then forward through the head to an opening called the salivarium, located behind the hypopharynx. By moving its mouthparts (element 32 in numbered diagram) the insect can mix its food with saliva. The mixture of saliva and food then travels through the salivary tubes into the mouth, where it begins to break down. Some insects, like flies
, have extra-oral digestion
. Insects using extra-oral digestion expel digestive enzymes onto their food to break it down. This strategy allows insects to extract a significant proportion of the available nutrients from the food source. The gut is where almost all of insects' digestion takes place. It can be divided into the foregut
, midgut
and hindgut
.
The first section of the alimentary canal is the foregut
(element 27 in numbered diagram), or stomodaeum. The foregut is lined with a cuticular lining made of chitin
and protein
s as protection from tough food. The foregut includes the buccal cavity (mouth), pharynx
, esophagus
, and Crop
and proventriculus
(any part may be highly modified) which both store food and signify when to continue passing onward to the midgut.
Digestion starts in buccal cavity (mouth) as partially chewed food is broken down by saliva from the salivary glands. As the salivary glands produce fluid and carbohydrate-digesting enzyme
s (mostly amylase
s), strong muscles in the pharynx pump fluid into the buccal cavity, lubricating the food like the salivarium does, and helping blood feeders, and xylem and phloem feeders.
From there, the pharynx passes food to the esophagus, which could be just a simple tube passing it on to the crop and proventriculus, and then onward to the midgut, as in most insects. Alternately, the foregut may expand into a very enlarged crop and proventriculus, or the crop could just be a diverticulum
, or fluid filled structure, as in some Diptera species.
Once food leaves the crop, it passes to the midgut
(element 13 in numbered diagram), also known as the mesenteron, where the majority of digestion takes place. Microscopic projections from the midgut wall, called microvilli
, increase the surface area of the wall and allow more nutrients to be absorbed; they tend to be close to the origin of the midgut. In some insects, the role of the microvilli and where they are located may vary. For example, specialized microvilli producing digestive enzymes may more likely be near the end of the midgut, and absorption near the origin or beginning of the midgut.
In the hindgut
(element 16 in numbered diagram), or proctodaeum, undigested food particles are joined by uric acid
to form fecal pellets. The rectum absorbs 90% of the water in these fecal pellets, and the dry pellet is then eliminated through the anus (element 17), completing the process of digestion. The uric acid is formed using hemolymph waste products diffused from the Malpighian tubule
s (element 20). It is then emptied directly into the alimentary canal, at the junction between the midgut and hindgut. The number of Malpighian tubules possessed by a given insect varies between species, ranging from only two tubules in some insects to over 100 tubules in others.
s (element 30 in numbered diagram) in an insect's mouth produce saliva. The salivary ducts lead from the glands to the reservoirs and then forward through the head to an opening called the salivarium, located behind the hypopharynx. By moving its mouthparts (element 32 in numbered diagram) the insect can mix its food with saliva. The mixture of saliva and food then travels through the salivary tubes into the mouth, where it begins to break down. Some insects, like flies
, have extra-oral digestion
. Insects using extra-oral digestion expel digestive enzymes onto their food to break it down. This strategy allows insects to extract a significant proportion of the available nutrients from the food source.
e, and ducts connecting these parts. The ovaries are made up of a number of egg tubes, called ovarioles
, which vary in size and number by species. The number of eggs that the insect is able to make vary by the number of ovarioles with the rate that eggs can be develop being also influenced by ovariole design. Female insects are able make eggs, receive and store sperm, manipulate sperm from different males, and lay eggs. Accessory glands or glandular parts of the oviducts produce a variety of substances for sperm maintenance, transport, and fertilization, as well as for protection of eggs. They can produce glue and protective substances for coating eggs or tough coverings for a batch of eggs called ootheca
e. Spermathecae are tubes or sacs in which sperm can be stored between the time of mating and the time an egg is fertilized.
For males, the reproductive system is the testis, suspended in the body cavity by trachea
e and the fat body. Most male insects have a pair of testes, inside of which are sperm tubes or follicles that are enclosed within a membranous sac. The follicles connect to the vas deferens by the vas efferens, and the two tubular vasa deferentia connect to a median ejaculatory duct that leads to the outside. A portion of the vas deferens is often enlarged to form the seminal vesicle, which stores the sperm before they are discharged into the female. The seminal vesicles have glandular linings that secrete nutrients for nourishment and maintenance of the sperm. The ejaculatory duct is derived from an invagination of the epidermal cells during development and, as a result, has a cuticular lining. The terminal portion of the ejaculatory duct may be sclerotized to form the intromittent organ, the aedeagus. The remainder of the male reproductive system is derived from embryonic mesoderm, except for the germ cells, or spermatogonia, which descend from the primordial pole cells very early during embryogenesis.
s. Instead, the insect respiratory system
uses 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 trachea
(element 8 in numbered diagram). Since oxygen is delivered directly, the circulatory system is not used to carry oxygen, and is therefore greatly reduced. The insect circulatory system has no vein
s or arterie
s, and instead consists of little more than a single, perforated dorsal tube which pulses peristaltically
. Toward the thorax, the dorsal tube (element 14) divides into chambers and acts like the insect's heart. The opposite end of the dorsal tube is like the aorta of the insect circulating the hemolymph
, arthropods' fluid analog of blood
, inside the body cavity. Air is taken in through openings on the sides of the abdomen called spiracle
s.
There are many different patterns of gas exchange
demonstrated by different groups of insects. Gas exchange patterns in insects can range from continuous and diffusive
ventilation, to discontinuous gas exchange
. During continuous gas exchange, oxygen
is taken in and carbon dioxide
is released in a continuous cycle. In discontinuous gas exchange, however, the insect takes in oxygen while it is active and small amounts of carbon dioxide are released when the insect is at rest. Diffusive ventilation is simply a form of continuous gas exchange that occurs by diffusion
rather than physically taking in the oxygen. Some species of insect that are submerged also have adaptations to aid in respiration. As larvae, many insects have gills that can extract oxygen dissolved in water, while others need to rise to the water surface to replenish air supplies which may be held or trapped in special structures.
The majority of insects hatch from egg
s. The fertilization and development takes place inside the egg, enclosed by a shell (chorion
). Some species of insects, like the cockroach Blaptica dubia
, as well as juvenile aphids and tsetse flies, are ovoviviparous
. The eggs of ovoviviparous animals develop entirely inside the female, and then hatch immediately upon being laid. Some other species, such as those in the genus of cockroaches known as Diploptera
, are viviparous, and thus gestate inside the mother and are born alive
. Some insects, like parasitic wasps, show polyembryony
, where a single fertilized egg divides into many and in some cases thousands of separate embryos.
Other developmental and reproductive variations include haplodiploidy, polymorphism
, paedomorphosis or peramorphosis
, sexual dimorphism
, parthenogenesis
and more rarely hermaphroditism. In haplodiploidy, which is a type of sex-determination system
, the offspring's sex is determined by the number of sets of chromosome
s an individual receives. This system is typical in bees and wasps. Polymorphism is the where a species may have different morphs or forms, as in the oblong winged katydid
, which has four different varieties: green, pink, and yellow or tan. Some insects may retain phenotype
s that are normally only seen in juveniles; this is called paedomorphosis. In peramorphosis, an opposite sort of phenomenon, insects take on previously unseen traits after they have matured into adults. Many insects display sexual dimorphism, in which males and females have notably different appearances, such as the moth Orgyia recens
as an exemplar of sexual dimorphism in insects.
Some insects use parthenogenesis
, a process in which the female can reproduce and give birth without having the eggs fertilized by a male
. Many aphids undergo a form of parthenogenesis, called cyclical parthenogenesis, in which they alternate between one or many generations of asexual and sexual reproduction. 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 which they spawn males. However, overall, most individuals are female, which are produced by fertilization. The males are haploid and the females are diploid. More rarely, some insects display hermaphroditism, in which a given individual has both male and female reproductive organs.
Insect life-histories show adaptations to withstand cold and dry conditions. Some temperate region insects are capable of activity during winter, while some others migrate to a warmer climate or go into a state of torpor
. Still other insects have evolved mechanisms of diapause
that allow eggs or pupae to survive these conditions.
in insects is the biological process of development all insects must undergo. There are two forms of metamorphosis: incomplete metamorphosis and complete metamorphosis.
, or incomplete metamorphosis, change gradually by undergoing a series of molt
s. An insect 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. 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, which makes the old cuticle split along predefined weaknesses where the old exocuticle was thinnest. Other arthropods have a much different process and only molt; though must accommodate for the difference in exoskeleton structure and make up with other enzymes.
Immature insects that go through incomplete metamorphosis are called nymph
s or in the case of dragonflies and damselflies as naiads. Nymphs are similar in form to the adult except for the presence of wings, which are not developed until adulthood. With each molt, nymphs grow larger and become more similar in appearance to adult insects.
Holometabolism
, or complete metamorphosis, is where the insect changes all in four stages, an egg
or embryo
, a larva
, a pupa
, and the adult
or imago
. In these species, egg hatches to produce a larva
, which is generally worm-like in form. This worm-like form can be one of several varieties: eruciform (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 and often sealed within a cocoon
. 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. Insects undergo considerable change in form during the pupal stage, and emerge as adults. Butterflies are a well known example of an insects that undergo complete metamorphosis, although most insects use this life cycle. Some insects have evolved this system to hypermetamorphosis
.
Some of the oldest and most successful insect groups, such Endopterygota
, use a system of complete metamorphosis. Strangely though, complete metamorphosis is unique to certain insect orders, like Diptera
, Lepidoptera
, and Hymenoptera
, and no other arthropods undergo it, but incomplete metamorphosis.
. Some insects such as bees can perceive ultraviolet
wavelengths, or detect polarized light, while the antennae
of male moths can detect the pheromone
s of female moths over distances of many kilometers. There is a pronounced tendency for there to be 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. There are a variety of different mechanisms by which insects perceive sound, while the patterns are not universal, insects can generally hear sound if they can produce it. Different insect species can have varying hearing, though most insects can hear only a narrow range of frequencies related to the frequency of the sounds they can produce. Mosquitoes have been found to hear up to 2 MHz., and some grasshoppers can hear up to 50 MHz. Certain predatory and parasitic insects can detect the characteristic sounds made by their prey or hosts, respectively. For instance, some nocturnal moths can perceive the ultrasonic
emissions of bat
s, which helps them avoid predation. Insects that feed on blood have special sensory structures that can detect infrared
emissions, and use them to home in on their hosts.
Some insects display a rudimentary sense of numbers
, such as the solitary wasps that prey upon a single species. The mother wasp lays her eggs in individual cells and provides each egg with a number of live caterpillars on which the young feed when hatched. Some species of wasp always provide five, others twelve, and others as high as twenty-four caterpillars per cell. The number of caterpillars is different among species, but always the same for each sex of larva. The male solitary wasp in the genus Eumenes
is smaller than the female, so the mother of one species supplies him with only five caterpillars; the larger female receives ten caterpillars in her cell.
A few insects, such as members of the families Poduridae and Onychiuridae (Collembola), 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.
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. The colors of emitted light vary from dull blue (Orfelia fultoni, Mycetophilidae) to the familiar greens and the rare reds (Phrixothrix tiemanni, Phengodidae).
Most insects, except some species of cave crickets
, are able to perceive light and dark. Many species have acute vision capable of detecting minute movements. The eyes include simple eyes or ocelli
as well as compound eyes of varying sizes. Many species are able to detect light in the infrared, ultraviolet and the visible light wavelengths. Color vision has been demonstrated in many species and phylogenetic analysis suggests that UV-green-blue trichromacy existed from at least the Devonian
period between 416 and 359 million years ago.
s and crickets, this is achieved by stridulation
. Cicada
s make the loudest sounds among the insects by producing and amplifying sounds with special modifications to their body and musculature. The African cicada
Brevisana brevis
has been measured at 106.7 decibel
s at a distance of 50 cm (19.7 in). Some insects, such as the hawk moths and Hedylid
butterflies, can hear ultrasound
and take evasive action when they sense that they have been detected by bats. Some moths produce ultrasonic clicks that were once thought to have a role in jamming bat echolocation
. The ultrasonic clicks were subsequently found to be produced mostly by unpalatable moths to warn bats, just as warning coloration
s are used against predators that hunt by sight. Some otherwise palatable moths have evolved to mimic
these calls.
Very low sounds are also produced in various species of Coleoptera, Hymenoptera
, Lepidoptera
, Mantodea, and Neuroptera
. These low sounds are simply the sounds made by the insect's movement. Through microscopic stridulatory structures located on the insect's muscles and joints, the normal sounds of the insect moving are amplified and can be used to warn or communicate with other insects. Most sound-making insects also have tympanal organ
s that can perceive airborne sounds. Some species in Hemiptera
, such as the corixids (water boatmen), are known to communicate via underwater sounds. Most insects are also able to sense vibration
s transmitted through surfaces. For example, an insect is caught in a spider web
and struggles to escape. The vibrations it produces are sensed by the spider
, who is alerted to its presence. Through these vibrations, the spider can tell where on the web the insect is located, as well as how big it is.
Communication using surface-borne vibrational signals is more widespread among insects because of size constraints in producing air-borne sounds. 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. The mechanisms of production of vibrational signals are just as diverse as those for producing sound in insects.
Some species use vibrations for communicating within members of the same species, such as to attract mates as in the songs of the shield bug
Nezara viridula
. Vibrations can also be used to communicate between entirely different species; lycaenid (gossamer-winged butterfly) caterpillars which are myrmecophilous (living in a mutualistic association with ants) communicate with ants in this way. The Madagascar hissing cockroach
has the ability to press air through its spiracles to make a hissing noise as a sign of aggression; 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 in close proximity.
s, are often derived from plant metabolites include those meant to attract, repel and provide other kinds of information. Pheromone
s, a type of semiochemical, 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. Allomone
a benefit their producer by the effect they have upon the receiver. Kairomone
s benefit their receiver instead of their producer. Synomones benefit the producer and the receiver. While some chemicals are targeted at individuals of the same species, others are used for communication across species. The use of scents is especially well known to have developed in social insects.
Social insects, such as termite
s, ant
s and many bee
s and wasp
s, are the most familiar species of eusocial
animal. They live together in large well-organized colonies that may be so tightly integrated and genetically similar that the colonies of some species are sometimes considered superorganism
s. It is sometimes argued that the various species of honey bee
are the only invertebrates (and indeed one of the few non-human groups) to have evolved a system of abstract symbolic communication where a behavior is used to represent and convey specific information about something in 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.
Only insects which live in nests or colonies demonstrate any true capacity for fine-scale spatial orientation or homing. This can allow an insect to return unerringly to a single hole a few millimeters in diameter among thousands of apparently identical holes clustered together, after a trip of up to several kilometers' distance. In a phenomenon known as philopatry
, insects that hibernate
have shown the ability to recall a specific location up to a year after last viewing the area of interest. A few insects seasonally migrate large distances between different geographic regions (e.g., the overwintering areas of the Monarch butterfly
).
).
Most insects, however, lead short lives as adults, and rarely interact with one another except to mate or compete for mates. A small number exhibit some form of parental care, where they will at least guard their eggs, and sometimes continue guarding their offspring until adulthood, and possibly even feeding them. Another simple form of parental care is to construct a nest (a burrow or an actual construction, either of which may be simple or complex), store provisions in it, and lay an egg upon those provisions. The adult does not contact the growing offspring, but it nonetheless does provide food. This sort of care is typical for most species of bees and various types of wasps.
s suggest that the wings are from paranotal lobes, or extensions from the insect's exoskeleton called the nota
, called the paranotal theory. Other theories are based on a pleural origin. These theories include suggestions that wings originated from modified gills, spiracular flaps or as from an appendage of the epicoxa. The epicoxal theory suggests the insect wings are modified epicoxal exites, a modified appendage at the base of the legs or coxa
. In the Carboniferous
age, some of the Meganeura
dragonflies had as much as a 50 cm (19.7 in) wide wingspan. The appearance of gigantic insects has been found to be consistent with high atmospheric oxygen. The respiratory system of insects constrains their size, however the high oxygen in the atmosphere allowed larger sizes. The largest flying insects today are much smaller and include several moth species such as the Atlas moth and the White Witch (Thysania agrippina). Insect flight has been a topic of great interest in aerodynamics
due partly to the inability of steady-state theories to explain the lift generated by the tiny wings of insects.
Unlike bird
s, many small insects are swept along by the prevailing winds although many of the larger insects are known to make migrations. Aphid
s are known to be transported long distances by low-level jet stream
s. As such, fine line patterns associated with converging winds within weather radar
imagery, like the WSR-88D radar network, often represent large groups of insects.
gait
. The tripedal gait allows for rapid walking while always having a stable stance and has been studied extensively in cockroach
es. The legs are used in alternate triangles touching the ground. 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 position. When they touch the ground to form a new stable triangle the other legs can be lifted and brought forward in turn and so on. The purest form of the tripedal gait is seen in insects moving at high speeds. However, this type of locomotion is not rigid and insects can adapt a variety of gaits. For example, when moving slowly, turning, or avoiding obstacles, four or more feet may be touching the ground. Insects can also adapt their gait to cope with the loss of one or more limbs.
Cockroaches are among the fastest insect runners and, at full speed, adopt a bipedal run to reach a high velocity in proportion to their body size. As cockroaches move very quickly, they need to be video recorded at several hundred frames per second to reveal their gait. More sedate locomotion is seen in the stick insects or walking sticks (Phasmatodea
). A few insects have evolved to walk on the surface of the water, especially the bugs of the Gerridae
family, commonly known as water striders. A few species of ocean-skaters in the genus Halobates
even live on the surface of open oceans, a habitat that has few insect species.
s. The study of insects and biped
s has a significant impact on possible robotic methods of transport. This may allow new robots to be designed that can traverse terrain
that robots with wheel
s may be unable to handle.
A large number of insects live either parts or the whole of their lives underwater. In many of the more primitive orders of insect, the immature stages are spent in an aquatic environment. Some groups of insects, like certain water beetle
s, have aquatic adults as well.
Many of these species have adaptations to help in under-water locomotion. Water beetles and water bugs have legs adapted into paddle-like structures. Dragonfly naiad
s use jet propulsion, forcibly expelling water out of their rectal chamber. Some species like the water strider
s are capable of walking on the surface of water. They can do this because their claws are not at the tips of the legs as in most insects, but recessed in a special groove further up the leg; this prevents the claws from piercing the water's surface film. Other insects such as the Rove beetle Stenus
are known to emit pygidial gland secretions that reduce surface tension making it possible for them to move on the surface of water by Marangoni propulsion
(also known by the German
term Entspannungsschwimmen).
ary relationships of insects to other animal groups remain unclear. Although more traditionally grouped with millipede
s and centipede
s, evidence has emerged favoring closer evolutionary ties
with crustacean
s. In the Pancrustacea
theory, insects, together with Remipedia
and Malacostraca
, make up a natural clade
. Other terrestrial arthropods, such as centipede
s, millipede
s, scorpion
s and spider
s, are sometimes confused with insects since their body plans can appear similar, sharing (as do all arthropods) a jointed exoskeleton. However, upon closer examination their features differ significantly; most noticeably they do not have the six legs characteristic of adult insects.
A phylogenetic tree of the arthropods and related groups
The higher-level phylogeny of the arthropods continues to be a matter of debate and research. In 2008, researchers at Tufts University
uncovered what they believe is the world's oldest known full-body impression of a primitive flying insect, a 300 million-year-old specimen from the Carboniferous Period. The oldest definitive insect fossil is the Devonian
Rhyniognatha hirsti
, from the 396 million year old Rhynie chert
. It may have superficially resembled a modern-day silverfish
insect. This species already possessed dicondylic mandibles (two articulations in the mandible), a feature associated with winged insects, suggesting that wings may already have evolved at this time. Thus, the first insects probably appeared earlier, in the Silurian
period.
There have been four super radiations of insects: beetle
s (evolved ~300 million years ago), flies
(evolved ~250 million years ago), moth
s and wasp
s (evolved ~150 million years ago). These four groups account for the majority of described species. The flies and moths along with the flea
s evolved from the Mecoptera
.
The origins of insect flight
remain obscure, since the earliest winged insects currently known appear to have been capable fliers. Some extinct insects had an additional pair of winglets attaching to the first segment of the thorax, for a total of three pairs. As of 2009, there is no evidence that suggests that the insects were a particularly successful group of animals before they evolved to have wings.
Late Carboniferous
and Early Permian
insect orders include both extant groups and a number of Paleozoic species, now extinct. During this era, some giant dragonfly-like forms reached wingspans of 55 to 70 cm (21.7 to 27.6 in) making them far larger than any living insect. This gigantism may have been due to higher atmospheric oxygen levels that allowed increased respiratory efficiency relative to today. The lack of flying vertebrates could have been another factor. Most extinct orders of insects developed during the Permian period that began around 270 million years ago. Many of the early groups became extinct during the Permian-Triassic extinction event
, the largest mass extinction in the history of the Earth, around 252 million years ago.
The remarkably successful Hymenoptera
ns appeared as long as 146 million years ago in the Cretaceous
period, but achieved their wide diversity more recently in the Cenozoic
era, which began 66 million years ago. A number of highly successful insect groups evolved in conjunction with flowering plants, a powerful illustration of coevolution.
Many modern insect genera
developed during the Cenozoic. Insects from this period on are often found preserved in amber
, often in perfect condition. The body plan, or morphology
, of such specimens is thus easily compared with modern species. The study of fossilized insects is called paleoentomology
.
, through gradual evolutionary change, insectivory was the next diet type to evolve.
Insects were among the earliest terrestrial herbivore
s and acted as major selection agents on plants. 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. This successful evolutionary pattern has also been utilized by mimic
s. Over time, this has led to complex groups of coevolved species. Conversely, some interactions between plants and insects, like pollination
, are beneficial to both organisms. Coevolution has led to the development of very specific mutualisms in such systems.
! colspan="4" style="background:#E6D09D"| Classification
|-
|
|-
{|
| rowspan="4" style="background:#ECF4ED"| Insecta
| colspan="3" style="background:#ECF4ED"|
{| align="right" class="collapsible collapsed" style="background:Transparent"
! Monocondylia
|-
| style="border-top: 1px solid black;"| -Archaeognatha
- 470
|}
|-
| rowspan="3" style="background:#ECF4ED"| Dicondylia
| colspan="2" style="background:#ECF4ED"|
{| align="right" class="collapsible collapsed" style="background:Transparent;"
! Apterygota
|-
| style="border-top: 1px solid black;"| -Thysanura
<200
|-
| -Monura
|}
|-
| rowspan="2" style="background:#ECF4ED"| Pterygota
| style="background:#ECF4ED"|
{| align="right" class="collapsible collapsed" style="background:Transparent"
! Paleoptera
|-
| style="border-top: 1px solid black;"| -Ephemeroptera- 2,500–<3,000
|-
| -Odonata
- 6,500
|}
|-
| style="background:#ECF4ED"|
{| align="right" class="collapsible collapsed" style="background:Transparent"
! Neoptera
|-
| style="border-top: 1px solid black;"| -Blattodea - 3,684–4,000
|-
| -Coleoptera - 360,000–400,000
|-
| -Dermaptera - 1,816
|-
| -Diptera
- 152,956
|-
| -Embioptera
- 200–300
|-
| -Grylloblattaria - 24
|-
| -Hymenoptera
- 115,000
|-
| -Lepidoptera
- 174,250
|-
| -Mantodea - 2,200
|-
| -Mecoptera
- 481
|-
| -Megaloptera
- 250–300
|-
| -Neuroptera
- 5,000
|-
| -Orthoptera
- 24,380
|-
| -Phasmatodea
- 2,500–3,300
|-
| -Phthiraptera - 3,000–3,200
|-
| -Plecoptera
- 2,274
|-
| -Siphonaptera - 2,525
|-
| -Strepsiptera
- 596
|-
| -Trichoptera
- 12,627
|-
| -Zoraptera
- 28
|-
| -Zygentoma - 370
|-
|}
|}
{| align="right" style="background:Transparent; border:solid 0px #503DF9;"
! colspan="4" style="background:Transparent"|Cladogram
of living insect groups, with numbers of species in each group. Note that Apterygota
, Palaeoptera and Exopterygota
are possibly paraphyletic groups.
|}
Traditional morphology-based or appearance-based systematics
has usually given Hexapoda
the rank of superclass, and identified four groups within it: insects (Ectognatha), springtails (Collembola
), Protura and Diplura, the latter three being grouped together as Entognatha
on the basis of internalized mouth parts. Supraordinal relationships have undergone numerous changes with the advent of methods based on evolutionary history and genetic data. A recent theory is that Hexapoda is polyphyletic (where the last common ancestor was not a member of the group), with the entognath classes having separate evolutionary histories from Insecta. Many of the traditional appearance-based taxa have been shown to be paraphyletic, so rather than using ranks like subclass
, superorder
and infraorder
, it has proved better to use monophyletic groupings (in which the last common ancestor is a member of the group). The following represents the best supported monophyletic groupings for the Insecta.
Insects can be divided into two groups historically treated as subclasses: wingless insects, known as Apterygota, and winged insects, known as Pterygota. The Apterygota consist of the primitively wingless order of the silverfish (Thysanura). Archaeognatha make up the Monocondylia based on the shape of their mandibles, while Thysanura and Pterygota are grouped together as Dicondylia. It is possible that the Thysanura themselves are not monophyletic, with the family Lepidotrichidae
being a sister group to the Dicondylia (Pterygota and the remaining Thysanura).
Paleoptera and Neoptera are the winged orders of insects differentiated by the presence of hardened body parts called sclerite
s; also, in Neoptera, muscles that allow their wings to fold flatly over the abdomen. Neoptera can further be divided into incomplete metamorphosis-based (Polyneoptera and Paraneoptera
) and complete metamorphosis-based groups. It has proved difficult to clarify the relationships between the orders in Polyneoptera because of constant new findings calling for revision of the taxa. For example, Paraneoptera has turned out to be more closely related to Endopterygota than to the rest of the Exopterygota. The recent molecular finding that the traditional louse orders Mallophaga and Anoplura are derived from within Psocoptera
has led to the new taxon Psocodea. Phasmatodea
and Embiidina have been suggested to form Eukinolabia. Mantodea, Blattodea and Isoptera are thought to form a monophyletic group termed Dictyoptera
.
It is likely that Exopterygota is paraphyletic in regard to Endopterygota. Matters that have had a lot of controversy include Strepsiptera and Diptera grouped together as Halteria based on a reduction of one of the wing pairs – a position not well-supported in the entomological community. The Neuropterida are often lumped or split on the whims of the taxonomist. Fleas are now thought to be closely related to boreid mecopterans. Many questions remain to be answered when it comes to basal relationships amongst endopterygote orders, particularly Hymenoptera.
The study of the classification or taxonomy of any insect is called systematic entomology
. If one works with a more specific order or even a family, the term may also be made specific to that order or family, for example systematic dipterology.
s, individually or as a community, interact with the surrounding environment
or ecosystem
. Insects play one of the most important roles in their ecosystems, which includes many roles, such as dung burial, pest control, pollination, and wildlife nutrition. An example is beetle
s, which are scavenger
s that feed on dead animals and fallen trees and thereby recycle
biological materials into forms found useful by other organism
s. These insects, and others, are responsible for much of the process by which topsoil
is created.
and parasitism
. So insects have a variety of strategies to avoid being attacked by predators or parasitoid
s. These include camouflage
, mimicry, toxicity, and active defense.
Camouflage
is also important defense strategies, which involves the use of coloration or shape to blend into the surrounding environment. This sort of protective coloration is common and widespread among beetle families, especially those that feed on wood or vegetation, such as many of the leaf beetle
s (family Chrysomelidae) or weevil
s. In some of these species, sculpturing or various colored scales or hairs cause the beetle to resemble bird dung or other inedible objects. Many of those that live in sandy environments blend in with the coloration of the substrate. Most phasmids are known for effectively replicating the forms of sticks and leaves, and the bodies of some species (such as O. macklotti and Palophus centaurus) are covered in mossy or lichenous
outgrowths that supplement their disguise. Some species have the ability to change color as their surroundings shift (B. scabrinota, T. californica). In a further behavioral adaptation to supplement crypsis
, a number of species have been noted to perform a rocking motion where the body is swayed from side to side that is thought to reflect the movement of leaves or twigs swaying in the breeze. Another method by which stick insects avoid predation and resemble twigs is by feigning death (catalepsy
), where the insect enters a motionless state that can be maintained for a long period. The nocturnal feeding habits of adults also aids Phasmatodea in remaining concealed from predators.
Another defense that often uses color or shape to deceive potential enemies is mimicry. A number of longhorn beetle
s (family Cerambycidae) bear a striking resemblance to wasp
s, which helps them avoid predation even though the beetles are in fact harmless. Batesian
and Müllerian
mimicry complexes are commonly found in Lepidoptera. Genetic polymorphism and natural selection give rise to otherwise edible species (the mimic) gaining a survival advantage by resembling inedible species (the model). Such a mimicry complex is referred to as Batesian and is most commonly known by the mimicry by the limenitidine
Viceroy butterfly
of the inedible danaine Monarch. Later research has discovered that the Viceroy is, in fact more toxic than the Monarch and this resemblance should be considered as a case of Müllerian mimicry. In Müllerian mimicry, inedible species, usually within a taxonomic order, find it advantageous to resemble each other so as to reduce the sampling rate by predators who need to learn about the insects' inedibility. Taxa from the toxic genus Heliconius
form one of the most well known Müllerian complexes.
Chemical defense is another important defense found amongst species of Coleoptera, usually being advertised by bright colors, such as the Monarch butterfly. They obtain their toxicity by sequestering the chemicals from the plants they eat into their own tissues. Some Lepidoptera manufacture their own toxins. Predators that eat poisonous butterflies and moths may become sick and vomit violently, learning not to eat those types of species; this is actually the basis of Müllerian mimicry. A predator who has previously eaten a poisonous lepidopteran may avoid other species with similar markings in the future, thus saving many other species as well. Chemical defense is another important defense found amongst species of Coleoptera, with some species releasing chemicals in the form of a spray with surprising accuracy, such as ground beetle
s (Carabidae), may spray chemicals from their abdomen to repel predators.
Many insects are considered pests
by humans. Insects commonly regarded as pests include those that are parasitic (mosquito
es, lice, bed bugs), transmit diseases (mosquitoes, flie
s), damage structures (termite
s), or destroy agricultural goods (locust
s, weevil
s). Many entomologists are involved in various forms of pest control, as in research for companies to produce insecticides, but increasingly relying on methods of biological pest control
, or biocontrol. Biocontrol uses one organism to reduce the population density of another organism — the pest — and is considered a key element of integrated pest management
.
Despite the large amount of effort focused at controlling insects, human attempts to kill pests with insecticides can backfire. If used carelessly the poison can kill all kinds of organisms in the area, including insects' natural predators such as birds, mice, and other insectivores. The effects of DDT
's use exemplifies how some insecticides can threaten wildlife beyond intended populations of pest insects.
Although pest insects attract the most attention, many insects are beneficial to the environment
and to human
s. Some insects, like wasp
s, bee
s, butterflie
s, and ant
s, pollinate
flowering plant
s. Pollination is a mutualistic relationship between plants and insects. As insects gather nectar from different plants of the same species, they also spread pollen
from plants on which they have previously fed. This greatly increases plants' ability to cross-pollinate, which maintains and possibly even improves their evolutionary fitness
. This ultimately affects humans since ensuring healthy crops is critical to agriculture
. A serious environmental problem is the decline of populations of pollinator
insects, and a number of species of insects are now cultured primarily for pollination management
in order to have sufficient pollinators in the field, orchard
or greenhouse
at bloom
time.
Insects also produce useful substances such as honey
, wax
, lacquer
and silk
. Honey bee
s have been cultured by humans for thousands of years for honey, although contracting for crop pollination is becoming more significant for beekeeper
s. The silkworm has greatly affected human history, as silk-driven trade
established relationships between China and the rest of the world.
Insectivorous
insects, or insects which feed on other insects, are beneficial to humans because they eat insects that could cause damage to agriculture and human structures. For example, aphid
s feed on crops and cause problems for farmers, but ladybugs feed on aphids, and can be used as a means to get significantly reduce pest aphid populations. While bird
s are perhaps more visible predators of insects, insects themselves account for the vast majority of insect consumption. Without predators to keep them in check, insects can undergo almost unstoppable population explosions.
Insects are also used in medicine, for example fly larvae (maggot
s) 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. Recently insects have also gained attention as potential sources of drugs and other medicinal substances. Also adult insects, such as crickets, and insect larvae of various kinds are also commonly used as fishing bait.
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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 higher eukaryote
s. D. melanogaster has been an essential part of studies into principles like 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. The genome
of D. melanogaster was sequenced
in 2000, reflecting the organism's important role in biological research.
, while being a taboo
in other places. In some cultures, insects, especially deep-fried cicada
s, are considered to be delicacie
s, while in other places they form part of the normal diet as they have a high protein content for their mass. In most first-world countries, however, entomophagy
, or the consumption of insects, is taboo
.
There are proponents of developing this use to provide a major source of protein
in human nutrition
. Since it is impossible to entirely eliminate pest insects from the human food chain, insects are present in many foods, especially grains. Food safety
laws in many countries do not prohibit insect parts in food, but rather limit the quantity. According to cultural materialist
anthropologist Marvin Harris
, the eating of insects is taboo in cultures that have other protein sources such as fish or livestock.
held religious and cultural symbolism in Old Egypt, Greece
and some shamanistic Old World cultures. The ancient Chinese
regarded cicada
s as symbols of rebirth or immortality. In Mesopotamia
n literature, the epic poem of Gilgamesh
has allusions to Odonata
which signify the impossibility of immortality.
Amongst the Aborigines
of Australia
of the Arrernte language groups, honey ants and witchety grubs served as personal clan totems. In the case of the 'San' bush-men of the Kalahari, it is the praying mantis which holds much cultural significance including creation and zen-like patience in waiting.
Class (biology)
In biological classification, class is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, order, family, genus, and species, with class fitting between phylum and order...
of living creatures within the arthropod
Arthropod
An arthropod is an invertebrate animal having an exoskeleton , a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda , and include the insects, arachnids, crustaceans, and others...
s that have a chitin
Chitin
Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world...
ous exoskeleton
Exoskeleton
An exoskeleton is the external skeleton that supports and protects an animal's body, in contrast to the internal skeleton of, for example, a human. In popular usage, some of the larger kinds of exoskeletons are known as "shells". Examples of exoskeleton animals include insects such as grasshoppers...
, a three-part body (head
Head
In anatomy, the head of an animal is the rostral part that usually comprises the brain, eyes, ears, nose and mouth . Some very simple animals may not have a head, but many bilaterally symmetric forms do....
, thorax
Thorax (insect anatomy)
The thorax is the mid section of the insect body. It holds the head, legs, wings and abdomen. It is also called mesosoma in other arthropods....
, and abdomen
Abdomen
In vertebrates such as mammals the abdomen constitutes the part of the body between the thorax and pelvis. The region enclosed by the abdomen is termed the abdominal cavity...
), three pairs of jointed leg
Arthropod leg
The arthropod leg is a form of jointed appendage of arthropods, usually used for walking. Many of the terms used for arthropod leg segments are of Latin origin, and may be confused with terms for bones: coxa , trochanter , femur, tibia, tarsus, ischium, metatarsus, carpus, dactylus ,...
s, compound eyes, and two antennae
Antenna (biology)
Antennae in biology have historically been paired appendages used for sensing in arthropods. More recently, the term has also been applied to cilium structures present in most cell types of eukaryotes....
. They are among the most diverse groups of animal
Animal
Animals are a major group of multicellular, eukaryotic organisms of the kingdom Animalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their life. Most animals are motile, meaning they can move spontaneously and...
s on the planet, including more than a million described species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...
and represent more than half of all known living organisms. The number of extant species is estimated at between six and ten million, and potentially represent over 90% of the differing metazoan life forms on Earth. Insects may be found in nearly all environments
Natural environment
The natural environment encompasses all living and non-living things occurring naturally on Earth or some region thereof. It is an environment that encompasses the interaction of all living species....
, although only a small number of species occur in the oceans, a habitat dominated by another arthropod group, the crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...
s.
The life cycles of insects vary but most hatch from eggs
Egg (biology)
An egg is an organic vessel in which an embryo first begins to develop. In most birds, reptiles, insects, molluscs, fish, and monotremes, an egg is the zygote, resulting from fertilization of the ovum, which is expelled from the body and permitted to develop outside the body until the developing...
. Insect growth is constrained by the inelastic exoskeleton
Exoskeleton
An exoskeleton is the external skeleton that supports and protects an animal's body, in contrast to the internal skeleton of, for example, a human. In popular usage, some of the larger kinds of exoskeletons are known as "shells". Examples of exoskeleton animals include insects such as grasshoppers...
and development involves a series of molts. The immature stages can differ from the adults in structure, habit and habitat and can include a passive pupa
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...
l stage in those groups that undergo complete metamorphosis. Insects that undergo incomplete metamorphosis
Hemimetabolism
Hemimetabolism or hemimetaboly, also called incomplete metamorphosis, is a term used to describe the mode of development of certain insects that includes three distinct stages: the egg, nymph, and the adult stage, or imago. These groups go through gradual changes; there is no pupal stage...
lack a pupal stage and adults develop through a series of nymph
Nymph (biology)
In biology, a nymph is the immature form of some invertebrates, particularly insects, which undergoes gradual metamorphosis before reaching its adult stage. Unlike a typical larva, a nymph's overall form already resembles that of the adult. In addition, while a nymph moults it never enters a...
al stages. The higher level relationship of the hexapoda
Hexapoda
The subphylum Hexapoda constitutes the largest grouping of arthropods and includes the insects as well as three much smaller groups of wingless arthropods: Collembola, Protura, and Diplura . The Collembola are very abundant in terrestrial environments...
is unclear. Fossilized insects of enormous size have been found from the Paleozoic Era
Paleozoic
The Paleozoic era is the earliest of three geologic eras of the Phanerozoic eon, spanning from roughly...
, including giant dragonflies with wingspans of 55 to 70 cm (22–28 in). The most diverse insect groups appear to have coevolved with flowering plant
Plant
Plants are living organisms belonging to the kingdom Plantae. Precise definitions of the kingdom vary, but as the term is used here, plants include familiar organisms such as trees, flowers, herbs, bushes, grasses, vines, ferns, mosses, and green algae. The group is also called green plants or...
s.
Insects typically move about by walking, flying or occasionally swimming. As it allows for rapid yet stable movement, many insects adopt a tripedal gait in which they walk with their legs touching the ground in alternating triangles. Insects are the only invertebrates to have evolved flight. Many insects spend at least part of their life underwater, with larval adaptations that include gill
Gill
A gill is a respiratory organ found in many aquatic organisms that extracts dissolved oxygen from water, afterward excreting carbon dioxide. The gills of some species such as hermit crabs have adapted to allow respiration on land provided they are kept moist...
s and some adult insects are aquatic and have adaptations for swimming. Some species, like water striders, are capable of walking on the surface of water. Insects are mostly solitary, but some insects, such as certain bee
Bee
Bees are flying insects closely related to wasps and ants, and are known for their role in pollination and for producing honey and beeswax. Bees are a monophyletic lineage within the superfamily Apoidea, presently classified by the unranked taxon name Anthophila...
s, ant
Ant
Ants are social insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than...
s, and termite
Termite
Termites are a group of eusocial insects that, until recently, were classified at the taxonomic rank of order Isoptera , but are now accepted as the epifamily Termitoidae, of the cockroach order Blattodea...
s are social and live in large, well-organized colonies. Some insects, like earwig
Earwig
Earwigs make up the insect order Dermaptera, found throughout the Americas, Africa, Eurasia, Australia and New Zealand. With 1,800 species in 12 families, they are one of the smaller insect orders...
s, show maternal care, guarding their eggs and young. Insects can communicate with each other in a variety of ways. Male moth
Moth
A moth is an insect closely related to the butterfly, both being of the order Lepidoptera. Moths form the majority of this order; there are thought to be 150,000 to 250,000 different species of moth , with thousands of species yet to be described...
s can sense the pheromone
Pheromone
A pheromone is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting outside the body of the secreting individual to impact the behavior of the receiving individual...
s of female moths over distances of many kilometers. Other species communicate with sounds: crickets
Cricket (insect)
Crickets, family Gryllidae , are insects somewhat related to grasshoppers, and more closely related to katydids or bush crickets . They have somewhat flattened bodies and long antennae. There are about 900 species of crickets...
stridulate
Stridulation
Stridulation is the act of producing sound by rubbing together certain body parts. This behavior is mostly associated with insects, but other animals are known to do this as well, such as a number of species of fishes, snakes and spiders...
, or rub their wings together, to attract a mate and repel other males. Lampyridae
Firefly
Lampyridae is a family of insects in the beetle order Coleoptera. They are winged beetles, and commonly called fireflies or lightning bugs for their conspicuous crepuscular use of bioluminescence to attract mates or prey. Fireflies produce a "cold light", with no infrared or ultraviolet frequencies...
in the beetle order Coleoptera
Beetle
Coleoptera is an order of insects commonly called beetles. The word "coleoptera" is from the Greek , koleos, "sheath"; and , pteron, "wing", thus "sheathed wing". Coleoptera contains more species than any other order, constituting almost 25% of all known life-forms...
communicate with light.
Humans regard certain insects as pests and attempt to control them using insecticide
Insecticide
An insecticide is a pesticide used against insects. They include ovicides and larvicides used against the eggs and larvae of insects respectively. Insecticides are used in agriculture, medicine, industry and the household. The use of insecticides is believed to be one of the major factors behind...
s and a host of other techniques. Some insects damage crops by feeding on sap, leaves or fruit
Fruit
In broad terms, a fruit is a structure of a plant that contains its seeds.The term has different meanings dependent on context. In non-technical usage, such as food preparation, fruit normally means the fleshy seed-associated structures of certain plants that are sweet and edible in the raw state,...
s, a few bite humans and livestock, alive and dead, to feed on blood and some are capable of transmitting disease
Disease
A disease is an abnormal condition affecting the body of an organism. It is often construed to be a medical condition associated with specific symptoms and signs. It may be caused by external factors, such as infectious disease, or it may be caused by internal dysfunctions, such as autoimmune...
s to humans, pets and livestock. Nevertheless, without insects to pollinate flowers, the human race would soon run out of food because many of the crop plants that we rely on would not be able to reproduce. Many other insects are considered ecologically beneficial as predators and a few provide direct economic benefit. Silkworms and bees have been used extensively by humans for the production of silk
Silk
Silk is a natural protein fiber, some forms of which can be woven into textiles. The best-known type of silk is obtained from the cocoons of the larvae of the mulberry silkworm Bombyx mori reared in captivity...
and honey
Honey
Honey is a sweet food made by bees using nectar from flowers. The variety produced by honey bees is the one most commonly referred to and is the type of honey collected by beekeepers and consumed by humans...
, respectively.
Etymology
The word insect dates back to 1600, from the Latin word meaning "with a notched or divided body," literally "cut into," from neuter plural of "to cut into, to cut up," from in- "into" + secare "to cut"; from the fact that insects are "cut into" three sections. The word created by Pliny the ElderPliny the Elder
Gaius Plinius Secundus , better known as Pliny the Elder, was a Roman author, naturalist, and natural philosopher, as well as naval and army commander of the early Roman Empire, and personal friend of the emperor Vespasian...
's loan-translation of the Greek
Greek language
Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...
word or "insect" (as in entomology
Entomology
Entomology is the scientific study of insects, a branch of arthropodology...
), which was Aristotle's term for this class of life, also in reference to their "notched" bodies, first documented in English in 1601 in Holland's translation of Pliny. Translations of Aristotle's term also form the usual word for "insect" in Welsh
Welsh language
Welsh is a member of the Brythonic branch of the Celtic languages spoken natively in Wales, by some along the Welsh border in England, and in Y Wladfa...
, Serbo-Croatian
Serbo-Croatian
Serbo-Croatian or Serbo-Croat, less commonly Bosnian/Croatian/Serbian , is a South Slavic language with multiple standards and the primary language of Serbia, Croatia, Bosnia and Herzegovina, and Montenegro...
(zareznik, from rezati "cut"), Russian
Russian language
Russian is a Slavic language used primarily in Russia, Belarus, Uzbekistan, Kazakhstan, Tajikistan and Kyrgyzstan. It is an unofficial but widely spoken language in Ukraine, Moldova, Latvia, Turkmenistan and Estonia and, to a lesser extent, the other countries that were once constituent republics...
( nasekomoe, from sekat "cut"), etc.
Distribution and diversity
Even though the true dimensions of species diversity remain uncertain, estimates are ranging from 1.4 to 1.8 million species. This probably represents less than 20% of all species on Earth, and with only about 20,000 new species of all organisms being described each year, it seems that most species will not remain described for many years unless there is a rapid increase in species descriptions. About 850,000–1,000,000 of all described species are insects. Of the 30 or so orders of insects, four dominate in terms of numbers of described species, with an estimated 600,000–795,000 species include Coleoptera, DipteraDiptera
Diptera , or true flies, is the order of insects possessing only a single pair of wings on the mesothorax; the metathorax bears a pair of drumstick like structures called the halteres, the remnants of the hind wings. It is a large order, containing an estimated 240,000 species, although under half...
, Hymenoptera
Hymenoptera
Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...
, and Lepidoptera
Lepidoptera
Lepidoptera is a large order of insects that includes moths and butterflies . It is one of the most widespread and widely recognizable insect orders in the world, encompassing moths and the three superfamilies of butterflies, skipper butterflies, and moth-butterflies...
. There are almost as many named species of beetle as there are of all other insects added together, or all other non-insects (plant
Plant
Plants are living organisms belonging to the kingdom Plantae. Precise definitions of the kingdom vary, but as the term is used here, plants include familiar organisms such as trees, flowers, herbs, bushes, grasses, vines, ferns, mosses, and green algae. The group is also called green plants or...
s and animal
Animal
Animals are a major group of multicellular, eukaryotic organisms of the kingdom Animalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their life. Most animals are motile, meaning they can move spontaneously and...
s).
| Described Species | Average description rate (species per year) | Publication effort | |
|---|---|---|---|
| Coleoptera | 300,000–400,000 | 2308 | 0.01 |
| Lepidoptera | 110,000–120,000 | 642 | 0.03 |
| Diptera | 90,000–150,000 | 1048 | 0.04 |
| Hymenoptera | 100,000–125,000 | 1196 | 0.02 |
External
Insects have segmentSegmentation (biology)
Segmentation in biology refers to either a type of gastrointestinal motility or the division of some animal and plant body plans into a series of repetitive segments. This article will focus on the segmentation of animal body plans, specifically using the examples of the phyla Arthropoda,...
ed bodies supported by an exoskeleton
Exoskeleton
An exoskeleton is the external skeleton that supports and protects an animal's body, in contrast to the internal skeleton of, for example, a human. In popular usage, some of the larger kinds of exoskeletons are known as "shells". Examples of exoskeleton animals include insects such as grasshoppers...
, a hard outer covering made mostly of chitin
Chitin
Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world...
. The segments of the body are organized into three distinctive but interconnected units, or tagmata
Tagma (biology)
In invertebrate biology, a tagma is a specialized grouping of arthropod segments, such as the head, the thorax, and the abdomen with a common function. The segments of a tagma may be either fused or moveable.-Tagmata:...
: a head, a thorax
Thorax (insect anatomy)
The thorax is the mid section of the insect body. It holds the head, legs, wings and abdomen. It is also called mesosoma in other arthropods....
, and an abdomen
Abdomen
In vertebrates such as mammals the abdomen constitutes the part of the body between the thorax and pelvis. The region enclosed by the abdomen is termed the abdominal cavity...
. The head supports a pair of sensory antennae
Antenna (biology)
Antennae in biology have historically been paired appendages used for sensing in arthropods. More recently, the term has also been applied to cilium structures present in most cell types of eukaryotes....
, a pair of compound eyes, and, if present, one to three simple eyes (or ocelli) and three sets of variously modified appendages that form the mouthpart
Insect mouthparts
Insects exhibit a range of mouthparts, adapted to particular modes of feeding. The earliest insects had chewing mouthparts...
s. The thorax has six segmented leg
Arthropod leg
The arthropod leg is a form of jointed appendage of arthropods, usually used for walking. Many of the terms used for arthropod leg segments are of Latin origin, and may be confused with terms for bones: coxa , trochanter , femur, tibia, tarsus, ischium, metatarsus, carpus, dactylus ,...
s—one pair each for the prothorax, mesothorax and the metathorax segments making up the thorax—and, if present in the species, two or four wing
Insect wing
Insects are the only group of invertebrates known to have evolved flight. Insects possess some remarkable flight characteristics and abilities, still far superior to attempts by humans to replicate their capabilities. Even our understanding of the aerodynamics of flexible, flapping wings and how...
s. The abdomen consists of eleven segments, though in a few species of insects these segments may be fused together or reduced in size. The abdomen also contains most of the digestive
Digestion
Digestion is the mechanical and chemical breakdown of food into smaller components that are more easily absorbed into a blood stream, for instance. Digestion is a form of catabolism: a breakdown of large food molecules to smaller ones....
, respiratory
Respiration (physiology)
'In physiology, respiration is defined as the transport of oxygen from the outside air to the cells within tissues, and the transport of carbon dioxide in the opposite direction...
, excretory
Excretion
Excretion is the process by which waste products of metabolism and other non-useful materials are eliminated from an organism. This is primarily carried out by the lungs, kidneys and skin. This is in contrast with secretion, where the substance may have specific tasks after leaving the cell...
and reproductive internal structures. There is considerable variation and many adaptations in the body parts of insects especially wings, legs, antenna, mouth-parts etc.
Segmentation
The headHead
In anatomy, the head of an animal is the rostral part that usually comprises the brain, eyes, ears, nose and mouth . Some very simple animals may not have a head, but many bilaterally symmetric forms do....
: is enclosed in a hard, heavily sclerotized, unsegmented, exoskeletal head capsule, or epicranium
Epicranium
The Epicranium is the medical term for the collection of structures covering the cranium. It consists of the muscles, aponeurosis, and skin....
, which contains most of the sensing organs, including the antennae, ocellus or eyes, and the mouthparts. Out of all the insect orders, Orthoptera displays the most features found in other insects, including the sutures and sclerite
Sclerite
A sclerite is a hardened body part. The term is used in various branches of biology for various structures including hardened portions of sponges, but it is most commonly used for the hardened portions of arthropod exoskeletons....
s. here, the vertex
Vertex (anatomy)
In arthropod and vertebrate anatomy, the vertex refers to the upper surface of the head.In humans, the vertex is formed by four bones of the skull: the frontal bone, the two parietal bones, and the occipital bone...
, or the apex (dorsal region), is situated between the compound eyes for insects with a hypognathous and opisthognathous head. In prognathous insects, the vertex is not found between the compound eyes, but rather, where the ocelli
Ocellus
A simple eye refers to a type of eye design or optical arrangement that contains a single lens which detect light. A "simple eye" is so-called in distinction from a multi-lensed "compound eye", and is not necessarily at all simple in the usual sense of the word...
are normally. This is because the primary axis of the head is rotated 90 degrees to become parallel to the primary axis of the body. In some species this region is modified and assumes a different name.
The thorax
Thorax
The thorax is a division of an animal's body that lies between the head and the abdomen.-In tetrapods:...
: is a segment composed of three sections, the prothorax
Prothorax
The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites are the pronotum , the prosternum , and the propleuron on each side. The prothorax never bears wings in extant insects, though some fossil groups possessed...
, mesothorax
Mesothorax
The mesothorax is the middle of the three segments in the thorax of an insect, and bears the second pair of legs. Its principal sclerites are the mesonotum , the mesosternum , and the mesopleuron on each side...
, and the metathorax
Metathorax
The metathorax is the posterior of the three segments in the thorax of an insect, and bears the third pair of legs. Its principal sclerites are the metanotum , the metasternum , and the metapleuron on each side...
. The anterior segment, closest to the head, is the prothorax, with the major features being the first pair of legs and the pronotum. The middle segment is the mesothorax with the major features being the second pair of legs and the anterior wings. The third and most posterior segment, abutting the abdomen, is the metathorax, which features the third pair of legs and the posterior wings. Each segment is dilineated by an intersegmental suture. Each segment has four basic regions. The dorsal surface is called the tergum (or notum) to distinguish them from the abdominal terga. The two lateral regions are called the pleura (singular: pleuron) and the ventral aspect is called the sternum. In turn, the notum of the prothorax is called the pronotum, the notum for the mesothorax is called the mesonotum and the notum for the metathorax is called the metanotum. Continuing with this logic, there is also the mesopleura and metapleura as well as the mesosternum and metasternum.
The abdomen
Abdomen
In vertebrates such as mammals the abdomen constitutes the part of the body between the thorax and pelvis. The region enclosed by the abdomen is termed the abdominal cavity...
: is a the last segment of the insect, which typically consists of 11–12 segments and is less strongly sclerotized than the head or thorax. Each segment of the abdomen is represented by a sclerotized tergum, sternum, and perhaps a pleurite. Terga are separated from each other and from the adjacent sterna or pleura by a membrane. Spiracles are located in the pleural area. Variation of this ground plan includes the fusion of terga or terga and sterna to form continuous dorsal or ventral shields or a conical tube. Some insects bear a sclerite in the pleural area called a laterotergite. Ventral sclerites are sometimes called laterosternites. During the embryonic stage of many insects and the postembryonic stage of primitive insects, 11 abdominal segments are present. In modern insects there is a tendency toward reduction in the number of the abdominal segments, but the primitive number of 11 is maintained during embryogenesis.Variation in abdominal segment number is considerable. If the Apterygota are considered to be indicative of the ground plan for pterygotes, confusion reigns: adult Protura have 12 segments, Collembola have 6. The orthopteran family Acrididae has 11 segments, and a fossil specimen of Zoraptera has a 10-segmented abdomen.
Exoskeleton
Insect outer skeleton, the cuticle, is made up of two layers: the epicuticle, which is a thin and waxy water resistant outer layer and contains no chitinChitin
Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world...
, and a lower layer called the procuticle. The procuticle is chitinous and much thicker than the epicuticle and has two layers: an outer layer known as the exocuticle and an inner layer known as the endocuticle. The tough and flexible endocuticle is built from numerous layers of fibrous chitin and proteins, criss-crossing each others in a sandwich pattern, while the exocuticle is rigid and hardened. The exocuticle is greatly reduced in many soft-bodied insects (e.g., caterpillar
Caterpillar
Caterpillars are the larval form of members of the order Lepidoptera . They are mostly herbivorous in food habit, although some species are insectivorous. Caterpillars are voracious feeders and many of them are considered to be pests in agriculture...
s), especially during their larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...
l stages.
Insects are the only invertebrate
Invertebrate
An invertebrate is an animal without a backbone. The group includes 97% of all animal species – all animals except those in the chordate subphylum Vertebrata .Invertebrates form a paraphyletic group...
s to have developed active flight capability, and this has played an important role in their success. Their muscles are able to contract multiple times for each single nerve impulse, allowing the wings to beat faster than would ordinarily be possible. Having their muscles attached to their exoskeletons is more efficient and allows more muscle connections; crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...
s also use the same method, though all spider
Spider
Spiders are air-breathing arthropods that have eight legs, and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other groups of organisms...
s use hydraulic pressure to extend their legs, a system inherited from their pre-arthropod ancestors. Unlike insects, though, most aquatic crustaceans are biomineralized with calcium carbonate
Calcium carbonate
Calcium carbonate is a chemical compound with the formula CaCO3. It is a common substance found in rocks in all parts of the world, and is the main component of shells of marine organisms, snails, coal balls, pearls, and eggshells. Calcium carbonate is the active ingredient in agricultural lime,...
extracted from the water.
Nervous system
The nervous systemNervous system
The nervous system is an organ system containing a network of specialized cells called neurons that coordinate the actions of an animal and transmit signals between different parts of its body. In most animals the nervous system consists of two parts, central and peripheral. The central nervous...
of an insect can be divided into a brain and a ventral nerve cord
Ventral nerve cord
The ventral nerve cord makes up the nervous system of some phyla of the invertebrates, particularly within the nematodes, annelids and the arthropods. It usually consists of cerebral ganglia anteriorly with the nerve cords running down the ventral plane of the organism...
. The head capsule is made up of six fused segments, each with a pair of ganglia
Ganglion
In anatomy, a ganglion is a biological tissue mass, most commonly a mass of nerve cell bodies. Cells found in a ganglion are called ganglion cells, though this term is also sometimes used to refer specifically to retinal ganglion cells....
, 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
Esophagus
The esophagus is an organ in vertebrates which consists of a muscular tube through which food passes from the pharynx to the stomach. During swallowing, food passes from the mouth through the pharynx into the esophagus and travels via peristalsis to the stomach...
, called the subesophageal ganglion
Subesophageal ganglion
The subesophageal ganglion of insects is composed of three pairs of fused ganglia. It controls the mouthparts, the salivary glands and certain muscles....
.
The thoracic segments have one ganglion on each side, which are connected into a pair, one pair per segment. This arrangement is also seen in the abdomen but only in the first eight segments. Many species of insects have reduced numbers of ganglia due to fusion or reduction. Some cockroaches have just six ganglia in the abdomen, whereas the wasp Vespa crabro has only two in the thorax and three in the abdomen. Some insects, like the house fly Musca domestica
Housefly
The housefly , Musca domestica, is a fly of the suborder Cyclorrhapha...
, have all the body ganglia fused into a single large thoracic ganglion.
At least a few insects have nociceptor
Nociceptor
A nociceptor is a sensory receptor that responds to potentially damaging stimuli by sending nerve signals to the spinal cord and brain. This process, called nociception, usually causes the perception of pain.-History:...
s, cells that detect and transmit sensations of pain
Pain in animals
Pain is a sensory and emotional experience often caused by intense or damaging stimuli. The International Association for the Study of Pain says pain is a conscious experience involving unpleasantness, i.e. suffering...
. This was discovered in 2003 by studying the variation in reactions of larvae
Larvae
In Roman mythology, lemures were shades or spirits of the restless or malignant dead, and are probably cognate with an extended sense of larvae as disturbing or frightening...
of the common fruitfly Drosophila
Drosophila
Drosophila is a genus of small flies, belonging to the family Drosophilidae, whose members are often called "fruit flies" or more appropriately pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many species to linger around overripe or rotting fruit...
to the touch of a heated probe and an unheated one. The larvae reacted to the touch of the heated probe with a stereotypical rolling behavior that was not exhibited when the larvae were touched by the unheated probe. Although nociception has been demonstrated in insects, there is not a consensus that insects feel pain consciously but see Pain in invertebrates
Pain in invertebrates
Pain in invertebrates is a problematic issue. Although there are numerous definitions of pain, almost all involve two key components. First, nociception is required. This is the ability to detect noxious stimuli which evokes a reflex response that moves the entire animal, or the affected part of...
.
Digestive system
An insect uses its digestive system to extract nutrients and other substances from the food it consumes. Most of this food is ingested in the form of macromoleculeMacromolecule
A macromolecule is a very large molecule commonly created by some form of polymerization. In biochemistry, the term is applied to the four conventional biopolymers , as well as non-polymeric molecules with large molecular mass such as macrocycles...
s and other complex substances like protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...
s, polysaccharide
Polysaccharide
Polysaccharides are long carbohydrate molecules, of repeated monomer units joined together by glycosidic bonds. They range in structure from linear to highly branched. Polysaccharides are often quite heterogeneous, containing slight modifications of the repeating unit. Depending on the structure,...
s, fat
Fat
Fats consist of a wide group of compounds that are generally soluble in organic solvents and generally insoluble in water. Chemically, fats are triglycerides, triesters of glycerol and any of several fatty acids. Fats may be either solid or liquid at room temperature, depending on their structure...
s, and nucleic acid
Nucleic acid
Nucleic acids are biological molecules essential for life, and include DNA and RNA . Together with proteins, nucleic acids make up the most important macromolecules; each is found in abundance in all living things, where they function in encoding, transmitting and expressing genetic information...
s. These macromolecules must be broken down by catabolic reaction
Catabolism
Catabolism is the set of metabolic pathways that break down molecules into smaller units and release energy. In catabolism, large molecules such as polysaccharides, lipids, nucleic acids and proteins are broken down into smaller units such as monosaccharides, fatty acids, nucleotides, and amino...
s into smaller molecules like amino acid
Amino acid
Amino acids are molecules containing an amine group, a carboxylic acid group and a side-chain that varies between different amino acids. The key elements of an amino acid are carbon, hydrogen, oxygen, and nitrogen...
s and simple sugars before being used by cells of the body for energy, growth, or reproduction. This break-down process is known as digestion.
The main structure of an insect's digestive system is a long enclosed tube called the alimentary canal, which runs lengthwise through the body. The alimentary canal directs food unidirectionally from the mouth
Mouth
The mouth is the first portion of the alimentary canal that receives food andsaliva. The oral mucosa is the mucous membrane epithelium lining the inside of the mouth....
to the anus
Anus
The anus is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, unwanted semi-solid matter produced during digestion, which, depending on the type of animal, may be one or more of: matter which the animal cannot digest,...
. It has three sections, each of which performs a different process of digestion. In addition to the alimentary canal, insects also have paired salivary glands and salivary reservoirs. These structures usually reside in the thorax, adjacent to the foregut.
The salivary gland
Salivary gland
The salivary glands in mammals are exocrine glands, glands with ducts, that produce saliva. They also secrete amylase, an enzyme that breaks down starch into maltose...
s (element 30 in numbered diagram) in an insect's mouth produce saliva. The salivary ducts lead from the glands to the reservoirs and then forward through the head to an opening called the salivarium, located behind the hypopharynx. By moving its mouthparts (element 32 in numbered diagram) the insect can mix its food with saliva. The mixture of saliva and food then travels through the salivary tubes into the mouth, where it begins to break down. Some insects, like flies
Diptera
Diptera , or true flies, is the order of insects possessing only a single pair of wings on the mesothorax; the metathorax bears a pair of drumstick like structures called the halteres, the remnants of the hind wings. It is a large order, containing an estimated 240,000 species, although under half...
, have extra-oral digestion
Mastication
Mastication or chewing is the process by which food is crushed and ground by teeth. It is the first step of digestion and it increases the surface area of foods to allow more efficient break down by enzymes. During the mastication process, the food is positioned between the teeth for grinding by...
. Insects using extra-oral digestion expel digestive enzymes onto their food to break it down. This strategy allows insects to extract a significant proportion of the available nutrients from the food source. The gut is where almost all of insects' digestion takes place. It can be divided into the foregut
Foregut
The foregut is the anterior part of the alimentary canal, from the mouth to the duodenum at the entrance of the bile duct. At this point it is continuous with the midgut...
, midgut
Stomach
The stomach is a muscular, hollow, dilated part of the alimentary canal which functions as an important organ of the digestive tract in some animals, including vertebrates, echinoderms, insects , and molluscs. It is involved in the second phase of digestion, following mastication .The stomach is...
and hindgut
Hindgut
The hindgut is the posterior part of the alimentary canal. In mammals, it includes the distal third of the transverse colon and the splenic flexure, the descending colon, sigmoid colon and rectum.-Blood flow:...
.
Foregut
The first section of the alimentary canal is the foregut
Foregut
The foregut is the anterior part of the alimentary canal, from the mouth to the duodenum at the entrance of the bile duct. At this point it is continuous with the midgut...
(element 27 in numbered diagram), or stomodaeum. The foregut is lined with a cuticular lining made of chitin
Chitin
Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world...
and protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...
s as protection from tough food. The foregut includes the buccal cavity (mouth), pharynx
Pharynx
The human pharynx is the part of the throat situated immediately posterior to the mouth and nasal cavity, and anterior to the esophagus and larynx. The human pharynx is conventionally divided into three sections: the nasopharynx , the oropharynx , and the laryngopharynx...
, esophagus
Esophagus
The esophagus is an organ in vertebrates which consists of a muscular tube through which food passes from the pharynx to the stomach. During swallowing, food passes from the mouth through the pharynx into the esophagus and travels via peristalsis to the stomach...
, and Crop
Crop (anatomy)
A crop is a thin-walled expanded portion of the alimentary tract used for the storage of food prior to digestion that is found in many animals, including gastropods, earthworms, leeches, insects, birds, and even some dinosaurs.- Bees :Cropping is used by bees to temporarily store nectar of flowers...
and proventriculus
Proventriculus
The proventriculus is part of the digestive system of birds, invertebrates and insects.-Birds:The proventriculus is a standard part of avian anatomy...
(any part may be highly modified) which both store food and signify when to continue passing onward to the midgut.
Digestion starts in buccal cavity (mouth) as partially chewed food is broken down by saliva from the salivary glands. As the salivary glands produce fluid and carbohydrate-digesting enzyme
Enzyme
Enzymes are proteins that catalyze chemical reactions. In enzymatic reactions, the molecules at the beginning of the process, called substrates, are converted into different molecules, called products. Almost all chemical reactions in a biological cell need enzymes in order to occur at rates...
s (mostly amylase
Amylase
Amylase is an enzyme that catalyses the breakdown of starch into sugars. Amylase is present in human saliva, where it begins the chemical process of digestion. Food that contains much starch but little sugar, such as rice and potato, taste slightly sweet as they are chewed because amylase turns...
s), strong muscles in the pharynx pump fluid into the buccal cavity, lubricating the food like the salivarium does, and helping blood feeders, and xylem and phloem feeders.
From there, the pharynx passes food to the esophagus, which could be just a simple tube passing it on to the crop and proventriculus, and then onward to the midgut, as in most insects. Alternately, the foregut may expand into a very enlarged crop and proventriculus, or the crop could just be a diverticulum
Diverticulum
A diverticulum is medical or biological term for an outpouching of a hollow structure in the body. Depending upon which layers of the structure are involved, they are described as being either true or false....
, or fluid filled structure, as in some Diptera species.
Midgut
Once food leaves the crop, it passes to the midgut
Stomach
The stomach is a muscular, hollow, dilated part of the alimentary canal which functions as an important organ of the digestive tract in some animals, including vertebrates, echinoderms, insects , and molluscs. It is involved in the second phase of digestion, following mastication .The stomach is...
(element 13 in numbered diagram), also known as the mesenteron, where the majority of digestion takes place. Microscopic projections from the midgut wall, called microvilli
Microvillus
Microvilli are microscopic cellular membrane protrusions that increase the surface area of cells, and are involved in a wide variety of functions, including absorption, secretion, cellular adhesion, and mechanotransduction....
, increase the surface area of the wall and allow more nutrients to be absorbed; they tend to be close to the origin of the midgut. In some insects, the role of the microvilli and where they are located may vary. For example, specialized microvilli producing digestive enzymes may more likely be near the end of the midgut, and absorption near the origin or beginning of the midgut.
Hindgut
In the hindgut
Hindgut
The hindgut is the posterior part of the alimentary canal. In mammals, it includes the distal third of the transverse colon and the splenic flexure, the descending colon, sigmoid colon and rectum.-Blood flow:...
(element 16 in numbered diagram), or proctodaeum, undigested food particles are joined by uric acid
Uric acid
Uric acid is a heterocyclic compound of carbon, nitrogen, oxygen, and hydrogen with the formula C5H4N4O3. It forms ions and salts known as urates and acid urates such as ammonium acid urate. Uric acid is created when the body breaks down purine nucleotides. High blood concentrations of uric acid...
to form fecal pellets. The rectum absorbs 90% of the water in these fecal pellets, and the dry pellet is then eliminated through the anus (element 17), completing the process of digestion. The uric acid is formed using hemolymph waste products diffused from the Malpighian tubule
Malpighian tubule system
The Malpighian tubule system is a type of excretory and osmoregulatory system found in some Uniramia , arachnids and tardigrades....
s (element 20). It is then emptied directly into the alimentary canal, at the junction between the midgut and hindgut. The number of Malpighian tubules possessed by a given insect varies between species, ranging from only two tubules in some insects to over 100 tubules in others.
Endocrine system
The salivary glandSalivary gland
The salivary glands in mammals are exocrine glands, glands with ducts, that produce saliva. They also secrete amylase, an enzyme that breaks down starch into maltose...
s (element 30 in numbered diagram) in an insect's mouth produce saliva. The salivary ducts lead from the glands to the reservoirs and then forward through the head to an opening called the salivarium, located behind the hypopharynx. By moving its mouthparts (element 32 in numbered diagram) the insect can mix its food with saliva. The mixture of saliva and food then travels through the salivary tubes into the mouth, where it begins to break down. Some insects, like flies
Diptera
Diptera , or true flies, is the order of insects possessing only a single pair of wings on the mesothorax; the metathorax bears a pair of drumstick like structures called the halteres, the remnants of the hind wings. It is a large order, containing an estimated 240,000 species, although under half...
, have extra-oral digestion
Mastication
Mastication or chewing is the process by which food is crushed and ground by teeth. It is the first step of digestion and it increases the surface area of foods to allow more efficient break down by enzymes. During the mastication process, the food is positioned between the teeth for grinding by...
. Insects using extra-oral digestion expel digestive enzymes onto their food to break it down. This strategy allows insects to extract a significant proportion of the available nutrients from the food source.
Reproductive system
The reproductive system of female insects consist of a pair of ovaries, accessory glands, one or more spermathecaSpermatheca
The spermatheca , also called receptaculum seminis , is an organ of the female reproductive tract in insects, some molluscs, oligochaeta worms and certain other invertebrates and vertebrates...
e, and ducts connecting these parts. The ovaries are made up of a number of egg tubes, called ovarioles
Ovarioles
An ovariole is one of the tubes of which the ovaries of most insects are composed. Typically an insect will have two ovaries. The constituent ovarioles lead to two oviducts, which converge into a single oviduct. The ovarioles are composed of a germarium and a set of ovarial follicles.-In D...
, which vary in size and number by species. The number of eggs that the insect is able to make vary by the number of ovarioles with the rate that eggs can be develop being also influenced by ovariole design. Female insects are able make eggs, receive and store sperm, manipulate sperm from different males, and lay eggs. Accessory glands or glandular parts of the oviducts produce a variety of substances for sperm maintenance, transport, and fertilization, as well as for protection of eggs. They can produce glue and protective substances for coating eggs or tough coverings for a batch of eggs called ootheca
Ootheca
An ootheca is a type of egg mass made by any member of a variety of species .The word is a latinized combination of oo-, meaning "egg", from the Greek word ōon , and theca, meaning a "cover" or "container", from the Greek theke...
e. Spermathecae are tubes or sacs in which sperm can be stored between the time of mating and the time an egg is fertilized.
For males, the reproductive system is the testis, suspended in the body cavity by trachea
Invertebrate trachea
The invertebrate trachea refers to the open respiratory system composed of spiracles, tracheae, and tracheoles that terrestrial arthropods have to transport metabolic gases to and from tissues....
e and the fat body. Most male insects have a pair of testes, inside of which are sperm tubes or follicles that are enclosed within a membranous sac. The follicles connect to the vas deferens by the vas efferens, and the two tubular vasa deferentia connect to a median ejaculatory duct that leads to the outside. A portion of the vas deferens is often enlarged to form the seminal vesicle, which stores the sperm before they are discharged into the female. The seminal vesicles have glandular linings that secrete nutrients for nourishment and maintenance of the sperm. The ejaculatory duct is derived from an invagination of the epidermal cells during development and, as a result, has a cuticular lining. The terminal portion of the ejaculatory duct may be sclerotized to form the intromittent organ, the aedeagus. The remainder of the male reproductive system is derived from embryonic mesoderm, except for the germ cells, or spermatogonia, which descend from the primordial pole cells very early during embryogenesis.
Respiratory and circulatory systems
Insect respiration is accomplished without lungLung
The lung is the essential respiration organ in many air-breathing animals, including most tetrapods, a few fish and a few snails. In mammals and the more complex life forms, the two lungs are located near the backbone on either side of the heart...
s. Instead, the insect respiratory system
Respiratory system of insects
The respiratory system of insects is the means by which insects obtain oxygen from the air.-The Respiratory System:Air enters the respiratory systems of most insects through a series of external openings called spiracles. These external openings, which act as muscular valves in some insects, lead...
uses 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 trachea
Invertebrate trachea
The invertebrate trachea refers to the open respiratory system composed of spiracles, tracheae, and tracheoles that terrestrial arthropods have to transport metabolic gases to and from tissues....
(element 8 in numbered diagram). Since oxygen is delivered directly, the circulatory system is not used to carry oxygen, and is therefore greatly reduced. The insect circulatory system has no vein
Vein
In the circulatory system, veins are blood vessels that carry blood towards the heart. Most veins carry deoxygenated blood from the tissues back to the heart; exceptions are the pulmonary and umbilical veins, both of which carry oxygenated blood to the heart...
s or arterie
Artery
Arteries are blood vessels that carry blood away from the heart. This blood is normally oxygenated, exceptions made for the pulmonary and umbilical arteries....
s, and instead consists of little more than a single, perforated dorsal tube which pulses peristaltically
Peristalsis
Peristalsis is a radially symmetrical contraction and relaxation of muscles which propagates in a wave down the muscular tube, in an anterograde fashion. In humans, peristalsis is found in the contraction of smooth muscles to propel contents through the digestive tract. Earthworms use a similar...
. Toward the thorax, the dorsal tube (element 14) divides into chambers and acts like the insect's heart. The opposite end of the dorsal tube is like the aorta of the insect circulating the hemolymph
Hemolymph
Hemolymph, or haemolymph, is a fluid in the circulatory system of some arthropods and is analogous to the fluids and cells making up both blood and interstitial fluid in vertebrates such as birds and mammals...
, arthropods' fluid analog of blood
Blood
Blood is a specialized bodily fluid in animals that delivers necessary substances such as nutrients and oxygen to the cells and transports metabolic waste products away from those same cells....
, inside the body cavity. Air is taken in through openings on the sides of the abdomen called spiracle
Spiracle
Spiracles are openings on the surface of some animals that usually lead to respiratory systems.-Vertebrates:The spiracle is a small hole behind each eye that opens to the mouth in some fishes. In the primitive jawless fish the first gill opening immediately behind the mouth is essentially similar...
s.
There are many different patterns of gas exchange
Gas exchange
Gas exchange is a process in biology where gases contained in an organism and atmosphere transfer or exchange. In human gas-exchange, gases contained in the blood of human bodies exchange with gases contained in the atmosphere. Human gas-exchange occurs in the lungs...
demonstrated by different groups of insects. Gas exchange patterns in insects can range from continuous and diffusive
Diffusion
Molecular diffusion, often called simply diffusion, is the thermal motion of all particles at temperatures above absolute zero. The rate of this movement is a function of temperature, viscosity of the fluid and the size of the particles...
ventilation, to discontinuous gas exchange
Discontinuous gas exchange
Discontinuous gas-exchange cycles , also called discontinuous ventilation or discontinuous ventilatory cycles, follow one of several patterns of arthropod gas exchange that have been documented primarily in insects; they occur when the insect is at rest...
. During continuous gas exchange, oxygen
Oxygen
Oxygen is the element with atomic number 8 and represented by the symbol O. Its name derives from the Greek roots ὀξύς and -γενής , because at the time of naming, it was mistakenly thought that all acids required oxygen in their composition...
is taken in and carbon dioxide
Carbon dioxide
Carbon dioxide is a naturally occurring chemical compound composed of two oxygen atoms covalently bonded to a single carbon atom...
is released in a continuous cycle. In discontinuous gas exchange, however, the insect takes in oxygen while it is active and small amounts of carbon dioxide are released when the insect is at rest. Diffusive ventilation is simply a form of continuous gas exchange that occurs by diffusion
Diffusion
Molecular diffusion, often called simply diffusion, is the thermal motion of all particles at temperatures above absolute zero. The rate of this movement is a function of temperature, viscosity of the fluid and the size of the particles...
rather than physically taking in the oxygen. Some species of insect that are submerged also have adaptations to aid in respiration. As larvae, many insects have gills that can extract oxygen dissolved in water, while others need to rise to the water surface to replenish air supplies which may be held or trapped in special structures.
Reproduction and development
Egg (biology)
An egg is an organic vessel in which an embryo first begins to develop. In most birds, reptiles, insects, molluscs, fish, and monotremes, an egg is the zygote, resulting from fertilization of the ovum, which is expelled from the body and permitted to develop outside the body until the developing...
s. The fertilization and development takes place inside the egg, enclosed by a shell (chorion
Chorion
The chorion is one of the membranes that exist during pregnancy between the developing fetus and mother. It is formed by extraembryonic mesoderm and the two layers of trophoblast and surrounds the embryo and other membranes...
). Some species of insects, like the cockroach Blaptica dubia
Blaptica dubia
The Dubia Cockroach , also known as Orange-spotted Cockroach or Guyana Spotted Cockroach, is a medium/large species of cockroach, measuring about long. They are sexually dimorphic; the males have wings while the females have only tiny wing stubs...
, as well as juvenile aphids and tsetse flies, are ovoviviparous
Ovoviviparity
Ovoviviparity, ovovivipary, or ovivipary, is a mode of reproduction in animals in which embryos develop inside eggs that are retained within the mother's body until they are ready to hatch...
. The eggs of ovoviviparous animals develop entirely inside the female, and then hatch immediately upon being laid. Some other species, such as those in the genus of cockroaches known as Diploptera
Diploptera
Diploptera is a genus of cockroach, used particularly in molecular research into allatostatins. The species Diploptera punctata, the Pacific beetle cockroach, is the only known viviparous cockroach ....
, are viviparous, and thus gestate inside the mother and are born alive
Vivipary
Vivipary has two different meanings. In animals, it means development of the embryo inside the body of the mother, eventually leading to live birth, as opposed to laying eggs...
. Some insects, like parasitic wasps, show polyembryony
Polyembryony
Polyembryony is the phenomenon of two or more embryos developing from a single fertilized egg . Polyembryony occurs regularly in many plants and animals. The nine banded armadillo, for instance, usually gives birth to four identical young...
, where a single fertilized egg divides into many and in some cases thousands of separate embryos.
Polymorphism (biology)
Polymorphism in biology occurs when two or more clearly different phenotypes exist in the same population of a species — in other words, the occurrence of more than one form or morph...
, paedomorphosis or peramorphosis
Peramorphosis
In developmental biology, peramorphosis is a phylogenetic change in which individuals of a species mature past adulthood and take on hitherto unseen traits...
, sexual dimorphism
Sexual dimorphism
Sexual dimorphism is a phenotypic difference between males and females of the same species. Examples of such differences include differences in morphology, ornamentation, and behavior.-Examples:-Ornamentation / coloration:...
, parthenogenesis
Parthenogenesis
Parthenogenesis is a form of asexual reproduction found in females, where growth and development of embryos occur without fertilization by a male...
and more rarely hermaphroditism. In haplodiploidy, which is a type of sex-determination system
Sex-determination system
A sex-determination system is a biological system that determines the development of sexual characteristics in an organism. Most sexual organisms have two sexes. In many cases, sex determination is genetic: males and females have different alleles or even different genes that specify their sexual...
, the offspring's sex is determined by the number of sets of chromosome
Chromosome
A chromosome is an organized structure of DNA and protein found in cells. It is a single piece of coiled DNA containing many genes, regulatory elements and other nucleotide sequences. Chromosomes also contain DNA-bound proteins, which serve to package the DNA and control its functions.Chromosomes...
s an individual receives. This system is typical in bees and wasps. Polymorphism is the where a species may have different morphs or forms, as in the oblong winged katydid
Amblycorypha oblongifolia
The Oblong-Winged katydid is a species of katydid, or bush-cricket, of the family Tettigoniidae .-Appearance:...
, which has four different varieties: green, pink, and yellow or tan. Some insects may retain phenotype
Phenotype
A phenotype is an organism's observable characteristics or traits: such as its morphology, development, biochemical or physiological properties, behavior, and products of behavior...
s that are normally only seen in juveniles; this is called paedomorphosis. In peramorphosis, an opposite sort of phenomenon, insects take on previously unseen traits after they have matured into adults. Many insects display sexual dimorphism, in which males and females have notably different appearances, such as the moth Orgyia recens
Orgyia recens
The Scarce Vapourer is a species of moth of the Lymantriidae family. It is found in Europe.The wingspan is 35-40 mm for the males, the females are wingless. The moth flies from June to July depending on the location....
as an exemplar of sexual dimorphism in insects.
Some insects use parthenogenesis
Parthenogenesis
Parthenogenesis is a form of asexual reproduction found in females, where growth and development of embryos occur without fertilization by a male...
, a process in which the female can reproduce and give birth without having the eggs fertilized by a male
Male
Male refers to the biological sex of an organism, or part of an organism, which produces small mobile gametes, called spermatozoa. Each spermatozoon can fuse with a larger female gamete or ovum, in the process of fertilization...
. Many aphids undergo a form of parthenogenesis, called cyclical parthenogenesis, in which they alternate between one or many generations of asexual and sexual reproduction. 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 which they spawn males. However, overall, most individuals are female, which are produced by fertilization. The males are haploid and the females are diploid. More rarely, some insects display hermaphroditism, in which a given individual has both male and female reproductive organs.
Insect life-histories show adaptations to withstand cold and dry conditions. Some temperate region insects are capable of activity during winter, while some others migrate to a warmer climate or go into a state of torpor
Torpor
Torpor, sometimes called temporary hibernation is a state of decreased physiological activity in an animal, usually characterized by a reduced body temperature and rate of metabolism. Animals that go through torpor include birds and some mammals such as mice and bats...
. Still other insects have evolved mechanisms of diapause
Diapause
Diapause is the delay in development in response to regularly and recurring periods of adverse environmental conditions. It is considered to be a physiological state of dormancy with very specific initiating and inhibiting conditions...
that allow eggs or pupae to survive these conditions.
Metamorphosis
MetamorphosisMetamorphosis
Metamorphosis is a biological process by which an animal physically develops after birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation...
in insects is the biological process of development all insects must undergo. There are two forms of metamorphosis: incomplete metamorphosis and complete metamorphosis.
Incomplete metamorphosis
Insects that show hemimetabolismHemimetabolism
Hemimetabolism or hemimetaboly, also called incomplete metamorphosis, is a term used to describe the mode of development of certain insects that includes three distinct stages: the egg, nymph, and the adult stage, or imago. These groups go through gradual changes; there is no pupal stage...
, or incomplete metamorphosis, change gradually by undergoing a series of molt
Ecdysis
Ecdysis is the moulting of the cuticula in many invertebrates. This process of moulting is the defining feature of the clade Ecdysozoa, comprising the arthropods, nematodes, velvet worms, horsehair worms, rotifers, tardigrades and Cephalorhyncha...
s. An insect 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. 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, which makes the old cuticle split along predefined weaknesses where the old exocuticle was thinnest. Other arthropods have a much different process and only molt; though must accommodate for the difference in exoskeleton structure and make up with other enzymes.
Immature insects that go through incomplete metamorphosis are called nymph
Nymph (biology)
In biology, a nymph is the immature form of some invertebrates, particularly insects, which undergoes gradual metamorphosis before reaching its adult stage. Unlike a typical larva, a nymph's overall form already resembles that of the adult. In addition, while a nymph moults it never enters a...
s or in the case of dragonflies and damselflies as naiads. Nymphs are similar in form to the adult except for the presence of wings, which are not developed until adulthood. With each molt, nymphs grow larger and become more similar in appearance to adult insects.
Complete metamorphosis
Holometabolism
Holometabolism, also called complete metamorphism, is a term applied to insect groups to describe the specific kind of insect development which includes four life stages - as an embryo or egg, a larva, a pupa and an imago or adult. Holometabolism is a monophyletic trait that all insects in the...
, or complete metamorphosis, is where the insect changes all in four stages, an egg
Egg (biology)
An egg is an organic vessel in which an embryo first begins to develop. In most birds, reptiles, insects, molluscs, fish, and monotremes, an egg is the zygote, resulting from fertilization of the ovum, which is expelled from the body and permitted to develop outside the body until the developing...
or embryo
Embryo
An embryo is a multicellular diploid eukaryote in its earliest stage of development, from the time of first cell division until birth, hatching, or germination...
, a larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...
, a pupa
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...
, and the adult
Adult
An adult is a human being or living organism that is of relatively mature age, typically associated with sexual maturity and the attainment of reproductive age....
or imago
Imago
In biology, the imago is the last stage of development of an insect, after the last ecdysis of an incomplete metamorphosis, or after emergence from the pupa where the metamorphosis is complete...
. In these species, egg hatches to produce a larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...
, which is generally worm-like in form. This worm-like form can be one of several varieties: eruciform (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
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...
, a stage marked by reduced movement and often sealed within a cocoon
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...
. 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. Insects undergo considerable change in form during the pupal stage, and emerge as adults. Butterflies are a well known example of an insects that undergo complete metamorphosis, although most insects use this life cycle. Some insects have evolved this system to hypermetamorphosis
Hypermetamorphosis
Hypermetamorphosis is a kind of complete insect metamorphosis in which the different larval instars represent two or more different forms of larva. As the larva molts its morphology can change from that of a campodeiform to scarabaeiform or to vermiform...
.
Some of the oldest and most successful insect groups, such Endopterygota
Endopterygota
The Endopterygota, also known as Holometabola, are insects of the subclass Pterygota which go through distinctive larval, pupal, and adult stages. They undergo a radical metamorphosis, with the larval and adult stages differing considerably in their structure and behaviour...
, use a system of complete metamorphosis. Strangely though, complete metamorphosis is unique to certain insect orders, like Diptera
Diptera
Diptera , or true flies, is the order of insects possessing only a single pair of wings on the mesothorax; the metathorax bears a pair of drumstick like structures called the halteres, the remnants of the hind wings. It is a large order, containing an estimated 240,000 species, although under half...
, Lepidoptera
Lepidoptera
Lepidoptera is a large order of insects that includes moths and butterflies . It is one of the most widespread and widely recognizable insect orders in the world, encompassing moths and the three superfamilies of butterflies, skipper butterflies, and moth-butterflies...
, and Hymenoptera
Hymenoptera
Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...
, and no other arthropods undergo it, but incomplete metamorphosis.
Senses and communication
Many insects possess very sensitive and/or specialized organs of perceptionSense
Senses are physiological capacities of organisms that provide inputs for perception. The senses and their operation, classification, and theory are overlapping topics studied by a variety of fields, most notably neuroscience, cognitive psychology , and philosophy of perception...
. Some insects such as bees can perceive ultraviolet
Ultraviolet
Ultraviolet light is electromagnetic radiation with a wavelength shorter than that of visible light, but longer than X-rays, in the range 10 nm to 400 nm, and energies from 3 eV to 124 eV...
wavelengths, or detect polarized light, while the antennae
Antenna (biology)
Antennae in biology have historically been paired appendages used for sensing in arthropods. More recently, the term has also been applied to cilium structures present in most cell types of eukaryotes....
of male moths can detect the pheromone
Pheromone
A pheromone is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting outside the body of the secreting individual to impact the behavior of the receiving individual...
s of female moths over distances of many kilometers. There is a pronounced tendency for there to be 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. There are a variety of different mechanisms by which insects perceive sound, while the patterns are not universal, insects can generally hear sound if they can produce it. Different insect species can have varying hearing, though most insects can hear only a narrow range of frequencies related to the frequency of the sounds they can produce. Mosquitoes have been found to hear up to 2 MHz., and some grasshoppers can hear up to 50 MHz. Certain predatory and parasitic insects can detect the characteristic sounds made by their prey or hosts, respectively. For instance, some nocturnal moths can perceive the ultrasonic
Ultrasound
Ultrasound is cyclic sound pressure with a frequency greater than the upper limit of human hearing. Ultrasound is thus not separated from "normal" sound based on differences in physical properties, only the fact that humans cannot hear it. Although this limit varies from person to person, it is...
emissions of bat
Bat
Bats are mammals of the order Chiroptera "hand" and pteron "wing") whose forelimbs form webbed wings, making them the only mammals naturally capable of true and sustained flight. By contrast, other mammals said to fly, such as flying squirrels, gliding possums, and colugos, glide rather than fly,...
s, which helps them avoid predation. Insects that feed on blood have special sensory structures that can detect infrared
Infrared
Infrared light is electromagnetic radiation with a wavelength longer than that of visible light, measured from the nominal edge of visible red light at 0.74 micrometres , and extending conventionally to 300 µm...
emissions, and use them to home in on their hosts.
Some insects display a rudimentary sense of numbers
Numeracy
Numeracy is the ability to reason with numbers and other mathematical concepts. A numerically literate person can manage and respond to the mathematical demands of life...
, such as the solitary wasps that prey upon a single species. The mother wasp lays her eggs in individual cells and provides each egg with a number of live caterpillars on which the young feed when hatched. Some species of wasp always provide five, others twelve, and others as high as twenty-four caterpillars per cell. The number of caterpillars is different among species, but always the same for each sex of larva. The male solitary wasp in the genus Eumenes
Eumenes (genus)
Eumenes is the type genus of the subfamily Eumeninae of Vespidae. It is a large and widespread genus, with over 100 taxa , mostly occurring in the temperate portions of the Northern Hemisphere. Most species are black or brown, and commonly marked with strikingly contrasting patterns of yellow,...
is smaller than the female, so the mother of one species supplies him with only five caterpillars; the larger female receives ten caterpillars in her cell.
Light production and vision
Mycetophilidae
Mycetophilidae is a family of small flies, forming the bulk of those species known as fungus gnats. There are approximately 3000 described species in 150 genera but the true number of species is undoubtedly much higher...
(Diptera), and the beetle families Lampyridae, Phengodidae
Phengodidae
The beetle family Phengodidae is known also as glowworm beetles, whose larvae are known as glowworms. The females and larvae have bioluminescent organs. They occur throughout the New World from extreme southern Canada to Chile...
, Elateridae and Staphylinidae are bioluminescent. The most familiar group are the fireflies
Fireflies
Fireflies is a novel by Shiva Naipaul originally published in 1970. It was his first book, a comic novel set in Trinidad. In an essay in An Unfinished Journey, Naipaul described how in 1968 as a final year student at Oxford University studying Chinese, he had been moved to write down a sentence,...
, 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
Arachnocampa
Arachnocampa is a genus of four fungus gnat species which are, in their larval stage, glowworms. They are found mostly in New Zealand and Australia in caves and grottos, or sheltered places in forests....
(Mycetophilidae, Fungus gnats) glow to lure small flying insects into sticky strands of silk.
Some fireflies of the genus Photuris mimic the flashing of female Photinus
Photinus (beetle)
The rover fireflies are a genus of fireflies . They are the type genus of tribe Photinini in subfamily Lampyrinae. This genus contains, for example, the common eastern firefly , the state insect of Tennessee.Male Photinus beetles emit a flashing light pattern to signal for females...
species to attract males of that species, which are then captured and devoured. The colors of emitted light vary from dull blue (Orfelia fultoni, Mycetophilidae) to the familiar greens and the rare reds (Phrixothrix tiemanni, Phengodidae).
Most insects, except some species of cave crickets
Rhaphidophoridae
The orthopteran family Rhaphidophoridae includes the cave wetas, cave crickets, camelback crickets, camel crickets, spider crickets and sand treaders, of the suborder Ensifera; in some regions, such as Missouri and Virginia, these crickets are referred to as "Cricket Spiders"...
, are able to perceive light and dark. Many species have acute vision capable of detecting minute movements. The eyes include simple eyes or ocelli
Ocellus
A simple eye refers to a type of eye design or optical arrangement that contains a single lens which detect light. A "simple eye" is so-called in distinction from a multi-lensed "compound eye", and is not necessarily at all simple in the usual sense of the word...
as well as compound eyes of varying sizes. Many species are able to detect light in the infrared, ultraviolet and the visible light wavelengths. Color vision has been demonstrated in many species and phylogenetic analysis suggests that UV-green-blue trichromacy existed from at least the Devonian
Devonian
The Devonian is a geologic period and system of the Paleozoic Era spanning from the end of the Silurian Period, about 416.0 ± 2.8 Mya , to the beginning of the Carboniferous Period, about 359.2 ± 2.5 Mya...
period between 416 and 359 million years ago.
Sound production and hearing
Insects were the earliest organisms to produce and sense sounds. Insects make sounds mostly by mechanical action of appendages. In grasshopperGrasshopper
The grasshopper is an insect of the suborder Caelifera in the order Orthoptera. To distinguish it from bush crickets or katydids, it is sometimes referred to as the short-horned grasshopper...
s and crickets, this is achieved by stridulation
Stridulation
Stridulation is the act of producing sound by rubbing together certain body parts. This behavior is mostly associated with insects, but other animals are known to do this as well, such as a number of species of fishes, snakes and spiders...
. Cicada
Cicada
A cicada is an insect of the order Hemiptera, suborder Auchenorrhyncha , in the superfamily Cicadoidea, with large eyes wide apart on the head and usually transparent, well-veined wings. There are about 2,500 species of cicada around the world, and many of them remain unclassified...
s make the loudest sounds among the insects by producing and amplifying sounds with special modifications to their body and musculature. The African cicada
Cicada
A cicada is an insect of the order Hemiptera, suborder Auchenorrhyncha , in the superfamily Cicadoidea, with large eyes wide apart on the head and usually transparent, well-veined wings. There are about 2,500 species of cicada around the world, and many of them remain unclassified...
Brevisana brevis
Brevisana brevis
Brevisana brevis is a cicada found in Africa and is the loudest insect on record. It has been recorded producing sounds with pressure levels of 106.7 decibels at a distance of 50 cm....
has been measured at 106.7 decibel
Decibel
The decibel is a logarithmic unit that indicates the ratio of a physical quantity relative to a specified or implied reference level. A ratio in decibels is ten times the logarithm to base 10 of the ratio of two power quantities...
s at a distance of 50 cm (19.7 in). Some insects, such as the hawk moths and Hedylid
Hedylidae
Hedylidae, the "American moth-butterflies", is a family of insects in the lepidopteran order, representing the superfamily Hedyloidea. They are an extant sister group of the butterfly superfamilies Papilionoidea and Hesperioidea...
butterflies, can hear ultrasound
Ultrasound
Ultrasound is cyclic sound pressure with a frequency greater than the upper limit of human hearing. Ultrasound is thus not separated from "normal" sound based on differences in physical properties, only the fact that humans cannot hear it. Although this limit varies from person to person, it is...
and take evasive action when they sense that they have been detected by bats. Some moths produce ultrasonic clicks that were once thought to have a role in jamming bat echolocation
Animal echolocation
Echolocation, also called biosonar, is the biological sonar used by several kinds of animals.Echolocating animals emit calls out to the environment and listen to the echoes of those calls that return from various objects near them. They use these echoes to locate and identify the objects...
. The ultrasonic clicks were subsequently found to be produced mostly by unpalatable moths to warn bats, just as warning coloration
Aposematism
Aposematism , perhaps most commonly known in the context of warning colouration, describes a family of antipredator adaptations where a warning signal is associated with the unprofitability of a prey item to potential predators...
s are used against predators that hunt by sight. Some otherwise palatable moths have evolved to mimic
Batesian mimicry
Batesian mimicry is a form of mimicry typified by a situation where a harmless species has evolved to imitate the warning signals of a harmful species directed at a common predator...
these calls.
Very low sounds are also produced in various species of Coleoptera, Hymenoptera
Hymenoptera
Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...
, Lepidoptera
Lepidoptera
Lepidoptera is a large order of insects that includes moths and butterflies . It is one of the most widespread and widely recognizable insect orders in the world, encompassing moths and the three superfamilies of butterflies, skipper butterflies, and moth-butterflies...
, Mantodea, and Neuroptera
Neuroptera
The insect order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order contains some 6,010 species...
. These low sounds are simply the sounds made by the insect's movement. Through microscopic stridulatory structures located on the insect's muscles and joints, the normal sounds of the insect moving are amplified and can be used to warn or communicate with other insects. Most sound-making insects also have tympanal organ
Tympanal organ
A tympanal organ is a hearing organ in insects, consisting of a membrane stretched across a frame backed by an air sac and associated sensory neurons...
s that can perceive airborne sounds. Some species in Hemiptera
Hemiptera
Hemiptera is an order of insects most often known as the true bugs , comprising around 50,000–80,000 species of cicadas, aphids, planthoppers, leafhoppers, shield bugs, and others...
, such as the corixids (water boatmen), are known to communicate via underwater sounds. Most insects are also able to sense vibration
Vibration
Vibration refers to mechanical oscillations about an equilibrium point. The oscillations may be periodic such as the motion of a pendulum or random such as the movement of a tire on a gravel road.Vibration is occasionally "desirable"...
s transmitted through surfaces. For example, an insect is caught in a spider web
Spider web
A spider web, spiderweb, spider's web or cobweb is a device built by a spider out of proteinaceous spider silk extruded from its spinnerets....
and struggles to escape. The vibrations it produces are sensed by the spider
Spider
Spiders are air-breathing arthropods that have eight legs, and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other groups of organisms...
, who is alerted to its presence. Through these vibrations, the spider can tell where on the web the insect is located, as well as how big it is.
Communication using surface-borne vibrational signals is more widespread among insects because of size constraints in producing air-borne sounds. 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. The mechanisms of production of vibrational signals are just as diverse as those for producing sound in insects.
Some species use vibrations for communicating within members of the same species, such as to attract mates as in the songs of the shield bug
Shield bug
Pentatomoidea is a superfamily of insects in the Heteroptera suborder of the Hemiptera order and, as such, share a common arrangement of sucking mouthparts...
Nezara viridula
Nezara viridula
Nezara viridula, commonly known as the southern green stink bug or green vegetable bug , is a plant-feeding stink bug. Although believed to have originated in Ethiopia, it can now be found around the world...
. Vibrations can also be used to communicate between entirely different species; lycaenid (gossamer-winged butterfly) caterpillars which are myrmecophilous (living in a mutualistic association with ants) communicate with ants in this way. The Madagascar hissing cockroach
Madagascar hissing cockroach
The Madagascar hissing Cockroach , also known as the Hissing Cockroach or simply Hisser, is one of the largest species of cockroach, reaching 2–4 inches at maturity. They are from the island of Madagascar off the African coast, where they can be found in rotting logs.Unlike most cockroaches, they...
has the ability to press air through its spiracles to make a hissing noise as a sign of aggression; the Death's-head Hawkmoth
Death's-head Hawkmoth
The name Death's-head Hawkmoth refers to any one of the three species of moth in the genus Acherontia. The former species is primarily found in Europe, the latter two are Asian, and most uses of the common name refer to the European species...
makes a squeaking noise by forcing air out of their pharynx
Pharynx
The human pharynx is the part of the throat situated immediately posterior to the mouth and nasal cavity, and anterior to the esophagus and larynx. The human pharynx is conventionally divided into three sections: the nasopharynx , the oropharynx , and the laryngopharynx...
when agitated, which may also reduce aggressive worker honey bee behavior when the two are in close proximity.
Chemical communication
In addition to the use of sound for communication, a wide range of insects have evolved chemical means for communication. These chemicals, termed semiochemicalSemiochemical
A semiochemical is a generic term used for a chemical substance or mixture that carries a message. These chemicals acts as messengers within or between species...
s, are often derived from plant metabolites include those meant to attract, repel and provide other kinds of information. Pheromone
Pheromone
A pheromone is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting outside the body of the secreting individual to impact the behavior of the receiving individual...
s, a type of semiochemical, are used for attracting mates of the opposite sex, for aggregating conspecific
Conspecificity
Conspecificity is a concept in biology. Two or more individual organisms, populations, or taxa are conspecific if they belong to the same species....
individuals of both sexes, for deterring other individuals from approaching, to mark a trail, and to trigger aggression in nearby individuals. Allomone
Allomone
An allomone is any chemical substance produced and released by an individual of one species that affects the behaviour of a member of another species to the benefit of the originator but not the receiver...
a benefit their producer by the effect they have upon the receiver. Kairomone
Kairomone
A kairomone is a semiochemical, emitted by an organism, which mediates interspecific interactions in a way that benefits an individual of another species which receives it, without benefiting the emitter. This "eavesdropping" is often disadvantageous to the producer...
s benefit their receiver instead of their producer. Synomones benefit the producer and the receiver. While some chemicals are targeted at individuals of the same species, others are used for communication across species. The use of scents is especially well known to have developed in social insects.
Social behavior
Termite
Termites are a group of eusocial insects that, until recently, were classified at the taxonomic rank of order Isoptera , but are now accepted as the epifamily Termitoidae, of the cockroach order Blattodea...
s, ant
Ant
Ants are social insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than...
s and many bee
Bee
Bees are flying insects closely related to wasps and ants, and are known for their role in pollination and for producing honey and beeswax. Bees are a monophyletic lineage within the superfamily Apoidea, presently classified by the unranked taxon name Anthophila...
s and wasp
Wasp
The term wasp is typically defined as any insect of the order Hymenoptera and suborder Apocrita that is neither a bee nor an ant. Almost every pest insect species has at least one wasp species that preys upon it or parasitizes it, making wasps critically important in natural control of their...
s, are the most familiar species of eusocial
Eusociality
Eusociality is a term used for the highest level of social organization in a hierarchical classification....
animal. They live together in large well-organized colonies that may be so tightly integrated and genetically similar that the colonies of some species are sometimes considered superorganism
Superorganism
A superorganism is an organism consisting of many organisms. This is usually meant to be a social unit of eusocial animals, where division of labour is highly specialised and where individuals are not able to survive by themselves for extended periods of time. Ants are the best-known example of...
s. It is sometimes argued that the various species of honey bee
Honey bee
Honey bees are a subset of bees in the genus Apis, primarily distinguished by the production and storage of honey and the construction of perennial, colonial nests out of wax. Honey bees are the only extant members of the tribe Apini, all in the genus Apis...
are the only invertebrates (and indeed one of the few non-human groups) to have evolved a system of abstract symbolic communication where a behavior is used to represent and convey specific information about something in the environment. In this communication system, called dance language
Bee learning and communication
Honey bees learn and communicate in order to find food sources and for other means.-Learning:Learning is essential for efficient foraging. Honey bees are unlikely to make many repeat visits if a plant provides little in the way of reward...
, 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.
Only insects which live in nests or colonies demonstrate any true capacity for fine-scale spatial orientation or homing. This can allow an insect to return unerringly to a single hole a few millimeters in diameter among thousands of apparently identical holes clustered together, after a trip of up to several kilometers' distance. In a phenomenon known as philopatry
Philopatry
Broadly, philopatry is the behaviour of remaining in, or returning to, an individual's birthplace. More specifically, in ecology philopatry is the behaviour of elder offspring sharing the parental burden in the upbringing of their siblings, a classic example of kin selection...
, insects that hibernate
Hibernation
Hibernation is a state of inactivity and metabolic depression in animals, characterized by lower body temperature, slower breathing, and lower metabolic rate. Hibernating animals conserve food, especially during winter when food supplies are limited, tapping energy reserves, body fat, at a slow rate...
have shown the ability to recall a specific location up to a year after last viewing the area of interest. A few insects seasonally migrate large distances between different geographic regions (e.g., the overwintering areas of the Monarch butterfly
Monarch butterfly
The Monarch butterfly is a milkweed butterfly , in the family Nymphalidae. It is perhaps the best known of all North American butterflies. Since the 19th century, it has been found in New Zealand, and in Australia since 1871 where it is called the Wanderer...
).
Care of young
Some bees, ants, termites and some wasps are eusocial, build nest, guard eggs and provide food for offsprings fulltime (see EusocialityEusociality
Eusociality is a term used for the highest level of social organization in a hierarchical classification....
).
Most insects, however, lead short lives as adults, and rarely interact with one another except to mate or compete for mates. A small number exhibit some form of parental care, where they will at least guard their eggs, and sometimes continue guarding their offspring until adulthood, and possibly even feeding them. Another simple form of parental care is to construct a nest (a burrow or an actual construction, either of which may be simple or complex), store provisions in it, and lay an egg upon those provisions. The adult does not contact the growing offspring, but it nonetheless does provide food. This sort of care is typical for most species of bees and various types of wasps.
Flight
Insects are the only group of invertebrates to have developed flight. The evolution of insect wings has been a subject of debate. Some entomologistEntomology
Entomology is the scientific study of insects, a branch of arthropodology...
s suggest that the wings are from paranotal lobes, or extensions from the insect's exoskeleton called the nota
Notum
The notum is the dorsal portion of an insect's thoracic segment. It is typically a single large sclerite. It can be further divided into the alinotum which bears the wings and the postnotum which bears the phragma....
, called the paranotal theory. Other theories are based on a pleural origin. These theories include suggestions that wings originated from modified gills, spiracular flaps or as from an appendage of the epicoxa. The epicoxal theory suggests the insect wings are modified epicoxal exites, a modified appendage at the base of the legs or coxa
Arthropod leg
The arthropod leg is a form of jointed appendage of arthropods, usually used for walking. Many of the terms used for arthropod leg segments are of Latin origin, and may be confused with terms for bones: coxa , trochanter , femur, tibia, tarsus, ischium, metatarsus, carpus, dactylus ,...
. In the Carboniferous
Carboniferous
The Carboniferous is a geologic period and system that extends from the end of the Devonian Period, about 359.2 ± 2.5 Mya , to the beginning of the Permian Period, about 299.0 ± 0.8 Mya . The name is derived from the Latin word for coal, carbo. Carboniferous means "coal-bearing"...
age, some of the Meganeura
Meganeura
Meganeura is a genus of extinct insects from the Carboniferous period approximately 300 million years ago, which resembled and are related to the present-day dragonflies. With wingspans of more than 75 cm , M. monyi is one of the largest known flying insect species; the Permian Meganeuropsis...
dragonflies had as much as a 50 cm (19.7 in) wide wingspan. The appearance of gigantic insects has been found to be consistent with high atmospheric oxygen. The respiratory system of insects constrains their size, however the high oxygen in the atmosphere allowed larger sizes. The largest flying insects today are much smaller and include several moth species such as the Atlas moth and the White Witch (Thysania agrippina). Insect flight has been a topic of great interest in aerodynamics
Aerodynamics
Aerodynamics is a branch of dynamics concerned with studying the motion of air, particularly when it interacts with a moving object. Aerodynamics is a subfield of fluid dynamics and gas dynamics, with much theory shared between them. Aerodynamics is often used synonymously with gas dynamics, with...
due partly to the inability of steady-state theories to explain the lift generated by the tiny wings of insects.
Unlike bird
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...
s, many small insects are swept along by the prevailing winds although many of the larger insects are known to make migrations. Aphid
Aphid
Aphids, also known as plant lice and in Britain and the Commonwealth as greenflies, blackflies or whiteflies, are small sap sucking insects, and members of the superfamily Aphidoidea. Aphids are among the most destructive insect pests on cultivated plants in temperate regions...
s are known to be transported long distances by low-level jet stream
Jet stream
Jet streams are fast flowing, narrow air currents found in the atmospheres of some planets, including Earth. The main jet streams are located near the tropopause, the transition between the troposphere and the stratosphere . The major jet streams on Earth are westerly winds...
s. As such, fine line patterns associated with converging winds within weather radar
Weather radar
Weather radar, also called weather surveillance radar and Doppler weather radar, is a type of radar used to locate precipitation, calculate its motion, estimate its type . Modern weather radars are mostly pulse-Doppler radars, capable of detecting the motion of rain droplets in addition to the...
imagery, like the WSR-88D radar network, often represent large groups of insects.
Walking
Many adult insects use six legs for walking and have adopted a tripedalTripedal
Tripedal is the term used for any animal that stands on three legs. The terms bipedal and quadrupedal are used more commonly when referring to animals that either walk on two legs Tripedal (from the Latin tri = three + ped = foot) is the term used for (or would be used for) any animal that stands...
gait
Gait
Gait is the pattern of movement of the limbs of animals, including humans, during locomotion over a solid substrate. Most animals use a variety of gaits, selecting gait based on speed, terrain, the need to maneuver, and energetic efficiency...
. The tripedal gait allows for rapid walking while always having a stable stance and has been studied extensively in cockroach
Cockroach
Cockroaches are insects of the order Blattaria or Blattodea, of which about 30 species out of 4,500 total are associated with human habitations...
es. The legs are used in alternate triangles touching the ground. 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 position. When they touch the ground to form a new stable triangle the other legs can be lifted and brought forward in turn and so on. The purest form of the tripedal gait is seen in insects moving at high speeds. However, this type of locomotion is not rigid and insects can adapt a variety of gaits. For example, when moving slowly, turning, or avoiding obstacles, four or more feet may be touching the ground. Insects can also adapt their gait to cope with the loss of one or more limbs.
Cockroaches are among the fastest insect runners and, at full speed, adopt a bipedal run to reach a high velocity in proportion to their body size. As cockroaches move very quickly, they need to be video recorded at several hundred frames per second to reveal their gait. More sedate locomotion is seen in the stick insects or walking sticks (Phasmatodea
Phasmatodea
The Phasmatodea are an order of insects, whose members are variously known as stick insects , walking sticks or stick-bugs , phasmids, ghost insects and leaf insects...
). A few insects have evolved to walk on the surface of the water, especially the bugs of the Gerridae
Water strider
Gerridae is a family of true bugs in the order Hemiptera, commonly known as water striders, water bugs, magic bugs, pond skaters, skaters, skimmers, water scooters, water skaters, water skeeters, water skimmers, water skippers, water spiders, or Jesus bugs...
family, commonly known as water striders. A few species of ocean-skaters in the genus Halobates
Halobates
Halobates or sea skaters are a genus with over 40 species of water striders. While many are coastal, about five of these are able to survive and stand on the surface of the open ocean, a habitat containing very few insect species. They are predators, coastal species feeding mainly on fallen...
even live on the surface of open oceans, a habitat that has few insect species.
Use in robotics
Insect walking is of particular interest as an alternative form of locomotion in robotRobot
A robot is a mechanical or virtual intelligent agent that can perform tasks automatically or with guidance, typically by remote control. In practice a robot is usually an electro-mechanical machine that is guided by computer and electronic programming. Robots can be autonomous, semi-autonomous or...
s. The study of insects and biped
Biped
Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs, or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning "two feet"...
s has a significant impact on possible robotic methods of transport. This may allow new robots to be designed that can traverse terrain
Terrain
Terrain, or land relief, is the vertical and horizontal dimension of land surface. When relief is described underwater, the term bathymetry is used...
that robots with wheel
Wheel
A wheel is a device that allows heavy objects to be moved easily through rotating on an axle through its center, facilitating movement or transportation while supporting a load, or performing labor in machines. Common examples found in transport applications. A wheel, together with an axle,...
s may be unable to handle.
Swimming
Water beetle
A water beetle is a beetle adapted to living in water. Water beetles rise to the water surface and take atmospheric air into their tracheal systems. There are approximately 2000 species of water beetles. The rest marine species tend to live in the intertidal zone...
s, have aquatic adults as well.
Many of these species have adaptations to help in under-water locomotion. Water beetles and water bugs have legs adapted into paddle-like structures. Dragonfly naiad
Naiad
In Greek mythology, the Naiads or Naiades were a type of nymph who presided over fountains, wells, springs, streams, and brooks....
s use jet propulsion, forcibly expelling water out of their rectal chamber. Some species like the water strider
Water strider
Gerridae is a family of true bugs in the order Hemiptera, commonly known as water striders, water bugs, magic bugs, pond skaters, skaters, skimmers, water scooters, water skaters, water skeeters, water skimmers, water skippers, water spiders, or Jesus bugs...
s are capable of walking on the surface of water. They can do this because their claws are not at the tips of the legs as in most insects, but recessed in a special groove further up the leg; this prevents the claws from piercing the water's surface film. Other insects such as the Rove beetle Stenus
Stenus
Stenus is a genus of semiaquatic rove beetles in the subfamily Steninae. They are known for swimming by emitting pygidial gland secretions that reduce surface tension and allow them to move on the surface of water....
are known to emit pygidial gland secretions that reduce surface tension making it possible for them to move on the surface of water by Marangoni propulsion
Marangoni effect
The Marangoni effect is the mass transfer along an interface between two fluids due to surface tension gradient...
(also known by the German
German language
German is a West Germanic language, related to and classified alongside English and Dutch. With an estimated 90 – 98 million native speakers, German is one of the world's major languages and is the most widely-spoken first language in the European Union....
term Entspannungsschwimmen).
Phylogeny and systematics
The evolutionEvolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...
ary relationships of insects to other animal groups remain unclear. Although more traditionally grouped with millipede
Millipede
Millipedes are arthropods that have two pairs of legs per segment . Each segment that has two pairs of legs is a result of two single segments fused together as one...
s and centipede
Centipede
Centipedes are arthropods belonging to the class Chilopoda of the subphylum Myriapoda. They are elongated metameric animals with one pair of legs per body segment. Despite the name, centipedes can have a varying number of legs from under 20 to over 300. Centipedes have an odd number of pairs of...
s, evidence has emerged favoring closer evolutionary ties
Common descent
In evolutionary biology, a group of organisms share common descent if they have a common ancestor. There is strong quantitative support for the theory that all living organisms on Earth are descended from a common ancestor....
with crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...
s. In the Pancrustacea
Pancrustacea
Pancrustacea is a clade, comprising all crustaceans and hexapods. This grouping is contrary to the Atelocerata hypothesis, in which Myriapoda and Hexapoda are sister taxa, and Crustacea are only more distantly related. As of 2010, the Pancrustacea taxon is considered well accepted. The clade has...
theory, insects, together with Remipedia
Remipedia
Remipedia is a class of blind crustaceans found in coastal aquifers which contain saline groundwater, with populations identified in almost every ocean basin so far explored, including in Australia, the Caribbean Sea, the Atlantic Ocean...
and Malacostraca
Malacostraca
Malacostraca is the largest of the six classes of crustaceans, containing over 25,000 extant species, divided among 16 orders. Its members display a greater diversity of body forms than any other class of animals, and include crabs, lobsters, shrimp, krill, woodlice, scuds , mantis shrimp and many...
, make up a natural clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...
. Other terrestrial arthropods, such as centipede
Centipede
Centipedes are arthropods belonging to the class Chilopoda of the subphylum Myriapoda. They are elongated metameric animals with one pair of legs per body segment. Despite the name, centipedes can have a varying number of legs from under 20 to over 300. Centipedes have an odd number of pairs of...
s, millipede
Millipede
Millipedes are arthropods that have two pairs of legs per segment . Each segment that has two pairs of legs is a result of two single segments fused together as one...
s, scorpion
Scorpion
Scorpions are predatory arthropod animals of the order Scorpiones within the class Arachnida. They have eight legs and are easily recognized by the pair of grasping claws and the narrow, segmented tail, often carried in a characteristic forward curve over the back, ending with a venomous stinger...
s and spider
Spider
Spiders are air-breathing arthropods that have eight legs, and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other groups of organisms...
s, are sometimes confused with insects since their body plans can appear similar, sharing (as do all arthropods) a jointed exoskeleton. However, upon closer examination their features differ significantly; most noticeably they do not have the six legs characteristic of adult insects.
A phylogenetic tree of the arthropods and related groups
The higher-level phylogeny of the arthropods continues to be a matter of debate and research. In 2008, researchers at Tufts University
Tufts University
Tufts University is a private research university located in Medford/Somerville, near Boston, Massachusetts. It is organized into ten schools, including two undergraduate programs and eight graduate divisions, on four campuses in Massachusetts and on the eastern border of France...
uncovered what they believe is the world's oldest known full-body impression of a primitive flying insect, a 300 million-year-old specimen from the Carboniferous Period. The oldest definitive insect fossil is the Devonian
Devonian
The Devonian is a geologic period and system of the Paleozoic Era spanning from the end of the Silurian Period, about 416.0 ± 2.8 Mya , to the beginning of the Carboniferous Period, about 359.2 ± 2.5 Mya...
Rhyniognatha hirsti
Rhyniognatha
Rhyniognatha hirsti is the world’s oldest known insect. It emerged very early during the Early Devonian Period, around 400 million years ago, when earth’s first terrestrial ecosystems were being formed.-Evidence:...
, from the 396 million year old Rhynie chert
Rhynie chert
The Rhynie chert is an Early Devonian sedimentary deposit exhibiting extraordinary fossil detail or completeness . It is exposed near the village of Rhynie, Aberdeenshire, Scotland; a second unit, the Windyfield chert, is located some 700 m away...
. It may have superficially resembled a modern-day silverfish
Silverfish
Lepisma saccharina, frequently called silverfish, fishmoths, carpet sharks or paramites, are small, wingless insects in the order Thysanura...
insect. This species already possessed dicondylic mandibles (two articulations in the mandible), a feature associated with winged insects, suggesting that wings may already have evolved at this time. Thus, the first insects probably appeared earlier, in the Silurian
Silurian
The Silurian is a geologic period and system that extends from the end of the Ordovician Period, about 443.7 ± 1.5 Mya , to the beginning of the Devonian Period, about 416.0 ± 2.8 Mya . As with other geologic periods, the rock beds that define the period's start and end are well identified, but the...
period.
There have been four super radiations of insects: beetle
Beetle
Coleoptera is an order of insects commonly called beetles. The word "coleoptera" is from the Greek , koleos, "sheath"; and , pteron, "wing", thus "sheathed wing". Coleoptera contains more species than any other order, constituting almost 25% of all known life-forms...
s (evolved ~300 million years ago), flies
Fly
True flies are insects of the order Diptera . They possess a pair of wings on the mesothorax and a pair of halteres, derived from the hind wings, on the metathorax...
(evolved ~250 million years ago), moth
Moth
A moth is an insect closely related to the butterfly, both being of the order Lepidoptera. Moths form the majority of this order; there are thought to be 150,000 to 250,000 different species of moth , with thousands of species yet to be described...
s and wasp
Wasp
The term wasp is typically defined as any insect of the order Hymenoptera and suborder Apocrita that is neither a bee nor an ant. Almost every pest insect species has at least one wasp species that preys upon it or parasitizes it, making wasps critically important in natural control of their...
s (evolved ~150 million years ago). These four groups account for the majority of described species. The flies and moths along with the flea
Flea
Flea is the common name for insects of the order Siphonaptera which are wingless insects with mouthparts adapted for piercing skin and sucking blood...
s evolved from the Mecoptera
Mecoptera
Mecoptera are an order of insects with about 550 species in nine families worldwide. Mecoptera are sometimes called scorpionflies after their largest family, Panorpidae, in which the males have enlarged genitals that look similar to the stinger of a scorpion...
.
The origins of insect flight
Insect flight
Insects are the only group of invertebrates known to have evolved flight. Insects possess some remarkable flight characteristics and abilities, still far superior to attempts by humans to replicate their capabilities. Even our understanding of the aerodynamics of flexible, flapping wings and how...
remain obscure, since the earliest winged insects currently known appear to have been capable fliers. Some extinct insects had an additional pair of winglets attaching to the first segment of the thorax, for a total of three pairs. As of 2009, there is no evidence that suggests that the insects were a particularly successful group of animals before they evolved to have wings.
Late Carboniferous
Pennsylvanian
The Pennsylvanian is, in the ICS geologic timescale, the younger of two subperiods of the Carboniferous Period. It lasted from roughly . As with most other geochronologic units, the rock beds that define the Pennsylvanian are well identified, but the exact date of the start and end are uncertain...
and Early Permian
Permian
The PermianThe term "Permian" was introduced into geology in 1841 by Sir Sir R. I. Murchison, president of the Geological Society of London, who identified typical strata in extensive Russian explorations undertaken with Edouard de Verneuil; Murchison asserted in 1841 that he named his "Permian...
insect orders include both extant groups and a number of Paleozoic species, now extinct. During this era, some giant dragonfly-like forms reached wingspans of 55 to 70 cm (21.7 to 27.6 in) making them far larger than any living insect. This gigantism may have been due to higher atmospheric oxygen levels that allowed increased respiratory efficiency relative to today. The lack of flying vertebrates could have been another factor. Most extinct orders of insects developed during the Permian period that began around 270 million years ago. Many of the early groups became extinct during the Permian-Triassic extinction event
Permian-Triassic extinction event
The Permian–Triassic extinction event, informally known as the Great Dying, was an extinction event that occurred 252.28 Ma ago, forming the boundary between the Permian and Triassic geologic periods, as well as the Paleozoic and Mesozoic eras...
, the largest mass extinction in the history of the Earth, around 252 million years ago.
The remarkably successful Hymenoptera
Hymenoptera
Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...
ns appeared as long as 146 million years ago in the Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...
period, but achieved their wide diversity more recently in the Cenozoic
Cenozoic
The Cenozoic era is the current and most recent of the three Phanerozoic geological eras and covers the period from 65.5 mya to the present. The era began in the wake of the Cretaceous–Tertiary extinction event at the end of the Cretaceous that saw the demise of the last non-avian dinosaurs and...
era, which began 66 million years ago. A number of highly successful insect groups evolved in conjunction with flowering plants, a powerful illustration of coevolution.
Many modern insect genera
Genera
Genera is a commercial operating system and development environment for Lisp machines developed by Symbolics. It is essentially a fork of an earlier operating system originating on the MIT AI Lab's Lisp machines which Symbolics had used in common with LMI and Texas Instruments...
developed during the Cenozoic. Insects from this period on are often found preserved in amber
Amber
Amber is fossilized tree resin , which has been appreciated for its color and natural beauty since Neolithic times. Amber is used as an ingredient in perfumes, as a healing agent in folk medicine, and as jewelry. There are five classes of amber, defined on the basis of their chemical constituents...
, often in perfect condition. The body plan, or morphology
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....
, of such specimens is thus easily compared with modern species. The study of fossilized insects is called paleoentomology
Entomology
Entomology is the scientific study of insects, a branch of arthropodology...
.
Evolutionary relationships
Insects are prey for a variety of organisms, including terrestrial vertebrates. The earliest vertebrates on land existed 400 million years ago and were large amphibious piscivoresPiscivore
A piscivore is a carnivorous animal which eats primarily fish. Piscivory was the diet of early tetrapods , insectivory came next, then in time reptiles added herbivory....
, through gradual evolutionary change, insectivory was the next diet type to evolve.
Insects were among the earliest terrestrial herbivore
Herbivore
Herbivores are organisms that are anatomically and physiologically adapted to eat plant-based foods. Herbivory is a form of consumption in which an organism principally eats autotrophs such as plants, algae and photosynthesizing bacteria. More generally, organisms that feed on autotrophs in...
s and acted as major selection agents on plants. Plants evolved chemical defenses against this herbivory
Plant defense against herbivory
Plant defense against herbivory or host-plant resistance describes a range of adaptations evolved by plants which improve their survival and reproduction by reducing the impact of herbivores. Plants use several strategies to defend against damage caused by herbivores...
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. This successful evolutionary pattern has also been utilized by mimic
Mimic
In evolutionary biology, mimicry is the similarity of one species to another which protects one or both. This similarity can be in appearance, behaviour, sound, scent and even location, with the mimics found in similar places to their models....
s. Over time, this has led to complex groups of coevolved species. Conversely, some interactions between plants and insects, like pollination
Pollination
Pollination is the process by which pollen is transferred in plants, thereby enabling fertilisation and sexual reproduction. Pollen grains transport the male gametes to where the female gamete are contained within the carpel; in gymnosperms the pollen is directly applied to the ovule itself...
, are beneficial to both organisms. Coevolution has led to the development of very specific mutualisms in such systems.
Taxonomy
{| align="right" style="background:Transparent; border:solid 0px #503DF9;"! colspan="4" style="background:#E6D09D"| Classification
|-
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Insects (from Latin
Latin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...
, a calque
Calque
In linguistics, a calque or loan translation is a word or phrase borrowed from another language by literal, word-for-word or root-for-root translation.-Calque:...
of Greek
Ancient Greek
Ancient Greek is the stage of the Greek language in the periods spanning the times c. 9th–6th centuries BC, , c. 5th–4th centuries BC , and the c. 3rd century BC – 6th century AD of ancient Greece and the ancient world; being predated in the 2nd millennium BC by Mycenaean Greek...
[], "cut into sections") are a class
Class (biology)
In biological classification, class is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, order, family, genus, and species, with class fitting between phylum and order...
of living creatures within the arthropod
Arthropod
An arthropod is an invertebrate animal having an exoskeleton , a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda , and include the insects, arachnids, crustaceans, and others...
s that have a chitin
Chitin
Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world...
ous exoskeleton
Exoskeleton
An exoskeleton is the external skeleton that supports and protects an animal's body, in contrast to the internal skeleton of, for example, a human. In popular usage, some of the larger kinds of exoskeletons are known as "shells". Examples of exoskeleton animals include insects such as grasshoppers...
, a three-part body (head
Head
In anatomy, the head of an animal is the rostral part that usually comprises the brain, eyes, ears, nose and mouth . Some very simple animals may not have a head, but many bilaterally symmetric forms do....
, thorax
Thorax (insect anatomy)
The thorax is the mid section of the insect body. It holds the head, legs, wings and abdomen. It is also called mesosoma in other arthropods....
, and abdomen
Abdomen
In vertebrates such as mammals the abdomen constitutes the part of the body between the thorax and pelvis. The region enclosed by the abdomen is termed the abdominal cavity...
), three pairs of jointed leg
Arthropod leg
The arthropod leg is a form of jointed appendage of arthropods, usually used for walking. Many of the terms used for arthropod leg segments are of Latin origin, and may be confused with terms for bones: coxa , trochanter , femur, tibia, tarsus, ischium, metatarsus, carpus, dactylus ,...
s, compound eyes, and two antennae
Antenna (biology)
Antennae in biology have historically been paired appendages used for sensing in arthropods. More recently, the term has also been applied to cilium structures present in most cell types of eukaryotes....
. They are among the most diverse groups of animal
Animal
Animals are a major group of multicellular, eukaryotic organisms of the kingdom Animalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their life. Most animals are motile, meaning they can move spontaneously and...
s on the planet, including more than a million described species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...
and represent more than half of all known living organisms. The number of extant species is estimated at between six and ten million, and potentially represent over 90% of the differing metazoan life forms on Earth. Insects may be found in nearly all environments
Natural environment
The natural environment encompasses all living and non-living things occurring naturally on Earth or some region thereof. It is an environment that encompasses the interaction of all living species....
, although only a small number of species occur in the oceans, a habitat dominated by another arthropod group, the crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...
s.
The life cycles of insects vary but most hatch from eggs
Egg (biology)
An egg is an organic vessel in which an embryo first begins to develop. In most birds, reptiles, insects, molluscs, fish, and monotremes, an egg is the zygote, resulting from fertilization of the ovum, which is expelled from the body and permitted to develop outside the body until the developing...
. Insect growth is constrained by the inelastic exoskeleton
Exoskeleton
An exoskeleton is the external skeleton that supports and protects an animal's body, in contrast to the internal skeleton of, for example, a human. In popular usage, some of the larger kinds of exoskeletons are known as "shells". Examples of exoskeleton animals include insects such as grasshoppers...
and development involves a series of molts. The immature stages can differ from the adults in structure, habit and habitat and can include a passive pupa
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...
l stage in those groups that undergo complete metamorphosis. Insects that undergo incomplete metamorphosis
Hemimetabolism
Hemimetabolism or hemimetaboly, also called incomplete metamorphosis, is a term used to describe the mode of development of certain insects that includes three distinct stages: the egg, nymph, and the adult stage, or imago. These groups go through gradual changes; there is no pupal stage...
lack a pupal stage and adults develop through a series of nymph
Nymph (biology)
In biology, a nymph is the immature form of some invertebrates, particularly insects, which undergoes gradual metamorphosis before reaching its adult stage. Unlike a typical larva, a nymph's overall form already resembles that of the adult. In addition, while a nymph moults it never enters a...
al stages. The higher level relationship of the hexapoda
Hexapoda
The subphylum Hexapoda constitutes the largest grouping of arthropods and includes the insects as well as three much smaller groups of wingless arthropods: Collembola, Protura, and Diplura . The Collembola are very abundant in terrestrial environments...
is unclear. Fossilized insects of enormous size have been found from the Paleozoic Era
Paleozoic
The Paleozoic era is the earliest of three geologic eras of the Phanerozoic eon, spanning from roughly...
, including giant dragonflies with wingspans of 55 to 70 cm (22–28 in). The most diverse insect groups appear to have coevolved with flowering plant
Plant
Plants are living organisms belonging to the kingdom Plantae. Precise definitions of the kingdom vary, but as the term is used here, plants include familiar organisms such as trees, flowers, herbs, bushes, grasses, vines, ferns, mosses, and green algae. The group is also called green plants or...
s.
Insects typically move about by walking, flying or occasionally swimming. As it allows for rapid yet stable movement, many insects adopt a tripedal gait in which they walk with their legs touching the ground in alternating triangles. Insects are the only invertebrates to have evolved flight. Many insects spend at least part of their life underwater, with larval adaptations that include gill
Gill
A gill is a respiratory organ found in many aquatic organisms that extracts dissolved oxygen from water, afterward excreting carbon dioxide. The gills of some species such as hermit crabs have adapted to allow respiration on land provided they are kept moist...
s and some adult insects are aquatic and have adaptations for swimming. Some species, like water striders, are capable of walking on the surface of water. Insects are mostly solitary, but some insects, such as certain bee
Bee
Bees are flying insects closely related to wasps and ants, and are known for their role in pollination and for producing honey and beeswax. Bees are a monophyletic lineage within the superfamily Apoidea, presently classified by the unranked taxon name Anthophila...
s, ant
Ant
Ants are social insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than...
s, and termite
Termite
Termites are a group of eusocial insects that, until recently, were classified at the taxonomic rank of order Isoptera , but are now accepted as the epifamily Termitoidae, of the cockroach order Blattodea...
s are social and live in large, well-organized colonies. Some insects, like earwig
Earwig
Earwigs make up the insect order Dermaptera, found throughout the Americas, Africa, Eurasia, Australia and New Zealand. With 1,800 species in 12 families, they are one of the smaller insect orders...
s, show maternal care, guarding their eggs and young. Insects can communicate with each other in a variety of ways. Male moth
Moth
A moth is an insect closely related to the butterfly, both being of the order Lepidoptera. Moths form the majority of this order; there are thought to be 150,000 to 250,000 different species of moth , with thousands of species yet to be described...
s can sense the pheromone
Pheromone
A pheromone is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting outside the body of the secreting individual to impact the behavior of the receiving individual...
s of female moths over distances of many kilometers. Other species communicate with sounds: crickets
Cricket (insect)
Crickets, family Gryllidae , are insects somewhat related to grasshoppers, and more closely related to katydids or bush crickets . They have somewhat flattened bodies and long antennae. There are about 900 species of crickets...
stridulate
Stridulation
Stridulation is the act of producing sound by rubbing together certain body parts. This behavior is mostly associated with insects, but other animals are known to do this as well, such as a number of species of fishes, snakes and spiders...
, or rub their wings together, to attract a mate and repel other males. Lampyridae
Firefly
Lampyridae is a family of insects in the beetle order Coleoptera. They are winged beetles, and commonly called fireflies or lightning bugs for their conspicuous crepuscular use of bioluminescence to attract mates or prey. Fireflies produce a "cold light", with no infrared or ultraviolet frequencies...
in the beetle order Coleoptera
Beetle
Coleoptera is an order of insects commonly called beetles. The word "coleoptera" is from the Greek , koleos, "sheath"; and , pteron, "wing", thus "sheathed wing". Coleoptera contains more species than any other order, constituting almost 25% of all known life-forms...
communicate with light.
Humans regard certain insects as pests and attempt to control them using insecticide
Insecticide
An insecticide is a pesticide used against insects. They include ovicides and larvicides used against the eggs and larvae of insects respectively. Insecticides are used in agriculture, medicine, industry and the household. The use of insecticides is believed to be one of the major factors behind...
s and a host of other techniques. Some insects damage crops by feeding on sap, leaves or fruit
Fruit
In broad terms, a fruit is a structure of a plant that contains its seeds.The term has different meanings dependent on context. In non-technical usage, such as food preparation, fruit normally means the fleshy seed-associated structures of certain plants that are sweet and edible in the raw state,...
s, a few bite humans and livestock, alive and dead, to feed on blood and some are capable of transmitting disease
Disease
A disease is an abnormal condition affecting the body of an organism. It is often construed to be a medical condition associated with specific symptoms and signs. It may be caused by external factors, such as infectious disease, or it may be caused by internal dysfunctions, such as autoimmune...
s to humans, pets and livestock. Nevertheless, without insects to pollinate flowers, the human race would soon run out of food because many of the crop plants that we rely on would not be able to reproduce. Many other insects are considered ecologically beneficial as predators and a few provide direct economic benefit. Silkworms and bees have been used extensively by humans for the production of silk
Silk
Silk is a natural protein fiber, some forms of which can be woven into textiles. The best-known type of silk is obtained from the cocoons of the larvae of the mulberry silkworm Bombyx mori reared in captivity...
and honey
Honey
Honey is a sweet food made by bees using nectar from flowers. The variety produced by honey bees is the one most commonly referred to and is the type of honey collected by beekeepers and consumed by humans...
, respectively.
Etymology
The word insect dates back to 1600, from the Latin word meaning "with a notched or divided body," literally "cut into," from neuter plural of "to cut into, to cut up," from in- "into" + secare "to cut"; from the fact that insects are "cut into" three sections. The word created by Pliny the ElderPliny the Elder
Gaius Plinius Secundus , better known as Pliny the Elder, was a Roman author, naturalist, and natural philosopher, as well as naval and army commander of the early Roman Empire, and personal friend of the emperor Vespasian...
's loan-translation of the Greek
Greek language
Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...
word or "insect" (as in entomology
Entomology
Entomology is the scientific study of insects, a branch of arthropodology...
), which was Aristotle's term for this class of life, also in reference to their "notched" bodies, first documented in English in 1601 in Holland's translation of Pliny. Translations of Aristotle's term also form the usual word for "insect" in Welsh
Welsh language
Welsh is a member of the Brythonic branch of the Celtic languages spoken natively in Wales, by some along the Welsh border in England, and in Y Wladfa...
, Serbo-Croatian
Serbo-Croatian
Serbo-Croatian or Serbo-Croat, less commonly Bosnian/Croatian/Serbian , is a South Slavic language with multiple standards and the primary language of Serbia, Croatia, Bosnia and Herzegovina, and Montenegro...
(zareznik, from rezati "cut"), Russian
Russian language
Russian is a Slavic language used primarily in Russia, Belarus, Uzbekistan, Kazakhstan, Tajikistan and Kyrgyzstan. It is an unofficial but widely spoken language in Ukraine, Moldova, Latvia, Turkmenistan and Estonia and, to a lesser extent, the other countries that were once constituent republics...
( nasekomoe, from sekat "cut"), etc.
Distribution and diversity
Even though the true dimensions of species diversity remain uncertain, estimates are ranging from 1.4 to 1.8 million species. This probably represents less than 20% of all species on Earth, and with only about 20,000 new species of all organisms being described each year, it seems that most species will not remain described for many years unless there is a rapid increase in species descriptions. About 850,000–1,000,000 of all described species are insects. Of the 30 or so orders of insects, four dominate in terms of numbers of described species, with an estimated 600,000–795,000 species include Coleoptera, DipteraDiptera
Diptera , or true flies, is the order of insects possessing only a single pair of wings on the mesothorax; the metathorax bears a pair of drumstick like structures called the halteres, the remnants of the hind wings. It is a large order, containing an estimated 240,000 species, although under half...
, Hymenoptera
Hymenoptera
Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...
, and Lepidoptera
Lepidoptera
Lepidoptera is a large order of insects that includes moths and butterflies . It is one of the most widespread and widely recognizable insect orders in the world, encompassing moths and the three superfamilies of butterflies, skipper butterflies, and moth-butterflies...
. There are almost as many named species of beetle as there are of all other insects added together, or all other non-insects (plant
Plant
Plants are living organisms belonging to the kingdom Plantae. Precise definitions of the kingdom vary, but as the term is used here, plants include familiar organisms such as trees, flowers, herbs, bushes, grasses, vines, ferns, mosses, and green algae. The group is also called green plants or...
s and animal
Animal
Animals are a major group of multicellular, eukaryotic organisms of the kingdom Animalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their life. Most animals are motile, meaning they can move spontaneously and...
s).
{| class="wikitable" style="text-align: center; width:80%;"
|+ Comparison of the Estimated Number of Species for Vertebrates and the Four Most Species Order
|-
|
! scope="col" | Described Species
! scope="col" | Average description rate (species per year)
! scope="col" | Publication effort
|-
! scope="row" | Coleoptera
| 300,000–400,000
| 2308
| 0.01
|-
! scope="row" | Lepidoptera
| 110,000–120,000
| 642
| 0.03
|-
! scope="row" | Diptera
| 90,000–150,000
| 1048
| 0.04
|-
! scope="row" | Hymenoptera
| 100,000–125,000
| 1196
| 0.02
|-
|}
External
Insects have segmentSegmentation (biology)
Segmentation in biology refers to either a type of gastrointestinal motility or the division of some animal and plant body plans into a series of repetitive segments. This article will focus on the segmentation of animal body plans, specifically using the examples of the phyla Arthropoda,...
ed bodies supported by an exoskeleton
Exoskeleton
An exoskeleton is the external skeleton that supports and protects an animal's body, in contrast to the internal skeleton of, for example, a human. In popular usage, some of the larger kinds of exoskeletons are known as "shells". Examples of exoskeleton animals include insects such as grasshoppers...
, a hard outer covering made mostly of chitin
Chitin
Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world...
. The segments of the body are organized into three distinctive but interconnected units, or tagmata
Tagma (biology)
In invertebrate biology, a tagma is a specialized grouping of arthropod segments, such as the head, the thorax, and the abdomen with a common function. The segments of a tagma may be either fused or moveable.-Tagmata:...
: a head, a thorax
Thorax (insect anatomy)
The thorax is the mid section of the insect body. It holds the head, legs, wings and abdomen. It is also called mesosoma in other arthropods....
, and an abdomen
Abdomen
In vertebrates such as mammals the abdomen constitutes the part of the body between the thorax and pelvis. The region enclosed by the abdomen is termed the abdominal cavity...
. The head supports a pair of sensory antennae
Antenna (biology)
Antennae in biology have historically been paired appendages used for sensing in arthropods. More recently, the term has also been applied to cilium structures present in most cell types of eukaryotes....
, a pair of compound eyes, and, if present, one to three simple eyes (or ocelli) and three sets of variously modified appendages that form the mouthpart
Insect mouthparts
Insects exhibit a range of mouthparts, adapted to particular modes of feeding. The earliest insects had chewing mouthparts...
s. The thorax has six segmented leg
Arthropod leg
The arthropod leg is a form of jointed appendage of arthropods, usually used for walking. Many of the terms used for arthropod leg segments are of Latin origin, and may be confused with terms for bones: coxa , trochanter , femur, tibia, tarsus, ischium, metatarsus, carpus, dactylus ,...
s—one pair each for the prothorax, mesothorax and the metathorax segments making up the thorax—and, if present in the species, two or four wing
Insect wing
Insects are the only group of invertebrates known to have evolved flight. Insects possess some remarkable flight characteristics and abilities, still far superior to attempts by humans to replicate their capabilities. Even our understanding of the aerodynamics of flexible, flapping wings and how...
s. The abdomen consists of eleven segments, though in a few species of insects these segments may be fused together or reduced in size. The abdomen also contains most of the digestive
Digestion
Digestion is the mechanical and chemical breakdown of food into smaller components that are more easily absorbed into a blood stream, for instance. Digestion is a form of catabolism: a breakdown of large food molecules to smaller ones....
, respiratory
Respiration (physiology)
'In physiology, respiration is defined as the transport of oxygen from the outside air to the cells within tissues, and the transport of carbon dioxide in the opposite direction...
, excretory
Excretion
Excretion is the process by which waste products of metabolism and other non-useful materials are eliminated from an organism. This is primarily carried out by the lungs, kidneys and skin. This is in contrast with secretion, where the substance may have specific tasks after leaving the cell...
and reproductive internal structures. There is considerable variation and many adaptations in the body parts of insects especially wings, legs, antenna, mouth-parts etc.
Segmentation
The headHead
In anatomy, the head of an animal is the rostral part that usually comprises the brain, eyes, ears, nose and mouth . Some very simple animals may not have a head, but many bilaterally symmetric forms do....
: is enclosed in a hard, heavily sclerotized, unsegmented, exoskeletal head capsule, or epicranium
Epicranium
The Epicranium is the medical term for the collection of structures covering the cranium. It consists of the muscles, aponeurosis, and skin....
, which contains most of the sensing organs, including the antennae, ocellus or eyes, and the mouthparts. Out of all the insect orders, Orthoptera displays the most features found in other insects, including the sutures and sclerite
Sclerite
A sclerite is a hardened body part. The term is used in various branches of biology for various structures including hardened portions of sponges, but it is most commonly used for the hardened portions of arthropod exoskeletons....
s. here, the vertex
Vertex (anatomy)
In arthropod and vertebrate anatomy, the vertex refers to the upper surface of the head.In humans, the vertex is formed by four bones of the skull: the frontal bone, the two parietal bones, and the occipital bone...
, or the apex (dorsal region), is situated between the compound eyes for insects with a hypognathous and opisthognathous head. In prognathous insects, the vertex is not found between the compound eyes, but rather, where the ocelli
Ocellus
A simple eye refers to a type of eye design or optical arrangement that contains a single lens which detect light. A "simple eye" is so-called in distinction from a multi-lensed "compound eye", and is not necessarily at all simple in the usual sense of the word...
are normally. This is because the primary axis of the head is rotated 90 degrees to become parallel to the primary axis of the body. In some species this region is modified and assumes a different name.
The thorax
Thorax
The thorax is a division of an animal's body that lies between the head and the abdomen.-In tetrapods:...
: is a segment composed of three sections, the prothorax
Prothorax
The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites are the pronotum , the prosternum , and the propleuron on each side. The prothorax never bears wings in extant insects, though some fossil groups possessed...
, mesothorax
Mesothorax
The mesothorax is the middle of the three segments in the thorax of an insect, and bears the second pair of legs. Its principal sclerites are the mesonotum , the mesosternum , and the mesopleuron on each side...
, and the metathorax
Metathorax
The metathorax is the posterior of the three segments in the thorax of an insect, and bears the third pair of legs. Its principal sclerites are the metanotum , the metasternum , and the metapleuron on each side...
. The anterior segment, closest to the head, is the prothorax, with the major features being the first pair of legs and the pronotum. The middle segment is the mesothorax with the major features being the second pair of legs and the anterior wings. The third and most posterior segment, abutting the abdomen, is the metathorax, which features the third pair of legs and the posterior wings. Each segment is dilineated by an intersegmental suture. Each segment has four basic regions. The dorsal surface is called the tergum (or notum) to distinguish them from the abdominal terga. The two lateral regions are called the pleura (singular: pleuron) and the ventral aspect is called the sternum. In turn, the notum of the prothorax is called the pronotum, the notum for the mesothorax is called the mesonotum and the notum for the metathorax is called the metanotum. Continuing with this logic, there is also the mesopleura and metapleura as well as the mesosternum and metasternum.
The abdomen
Abdomen
In vertebrates such as mammals the abdomen constitutes the part of the body between the thorax and pelvis. The region enclosed by the abdomen is termed the abdominal cavity...
: is a the last segment of the insect, which typically consists of 11–12 segments and is less strongly sclerotized than the head or thorax. Each segment of the abdomen is represented by a sclerotized tergum, sternum, and perhaps a pleurite. Terga are separated from each other and from the adjacent sterna or pleura by a membrane. Spiracles are located in the pleural area. Variation of this ground plan includes the fusion of terga or terga and sterna to form continuous dorsal or ventral shields or a conical tube. Some insects bear a sclerite in the pleural area called a laterotergite. Ventral sclerites are sometimes called laterosternites. During the embryonic stage of many insects and the postembryonic stage of primitive insects, 11 abdominal segments are present. In modern insects there is a tendency toward reduction in the number of the abdominal segments, but the primitive number of 11 is maintained during embryogenesis.Variation in abdominal segment number is considerable. If the Apterygota are considered to be indicative of the ground plan for pterygotes, confusion reigns: adult Protura have 12 segments, Collembola have 6. The orthopteran family Acrididae has 11 segments, and a fossil specimen of Zoraptera has a 10-segmented abdomen.
Exoskeleton
Insect outer skeleton, the cuticle, is made up of two layers: the epicuticle, which is a thin and waxy water resistant outer layer and contains no chitinChitin
Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world...
, and a lower layer called the procuticle. The procuticle is chitinous and much thicker than the epicuticle and has two layers: an outer layer known as the exocuticle and an inner layer known as the endocuticle. The tough and flexible endocuticle is built from numerous layers of fibrous chitin and proteins, criss-crossing each others in a sandwich pattern, while the exocuticle is rigid and hardened. The exocuticle is greatly reduced in many soft-bodied insects (e.g., caterpillar
Caterpillar
Caterpillars are the larval form of members of the order Lepidoptera . They are mostly herbivorous in food habit, although some species are insectivorous. Caterpillars are voracious feeders and many of them are considered to be pests in agriculture...
s), especially during their larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...
l stages.
Insects are the only invertebrate
Invertebrate
An invertebrate is an animal without a backbone. The group includes 97% of all animal species – all animals except those in the chordate subphylum Vertebrata .Invertebrates form a paraphyletic group...
s to have developed active flight capability, and this has played an important role in their success. Their muscles are able to contract multiple times for each single nerve impulse, allowing the wings to beat faster than would ordinarily be possible. Having their muscles attached to their exoskeletons is more efficient and allows more muscle connections; crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...
s also use the same method, though all spider
Spider
Spiders are air-breathing arthropods that have eight legs, and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other groups of organisms...
s use hydraulic pressure to extend their legs, a system inherited from their pre-arthropod ancestors. Unlike insects, though, most aquatic crustaceans are biomineralized with calcium carbonate
Calcium carbonate
Calcium carbonate is a chemical compound with the formula CaCO3. It is a common substance found in rocks in all parts of the world, and is the main component of shells of marine organisms, snails, coal balls, pearls, and eggshells. Calcium carbonate is the active ingredient in agricultural lime,...
extracted from the water.
Nervous system
The nervous systemNervous system
The nervous system is an organ system containing a network of specialized cells called neurons that coordinate the actions of an animal and transmit signals between different parts of its body. In most animals the nervous system consists of two parts, central and peripheral. The central nervous...
of an insect can be divided into a brain and a ventral nerve cord
Ventral nerve cord
The ventral nerve cord makes up the nervous system of some phyla of the invertebrates, particularly within the nematodes, annelids and the arthropods. It usually consists of cerebral ganglia anteriorly with the nerve cords running down the ventral plane of the organism...
. The head capsule is made up of six fused segments, each with a pair of ganglia
Ganglion
In anatomy, a ganglion is a biological tissue mass, most commonly a mass of nerve cell bodies. Cells found in a ganglion are called ganglion cells, though this term is also sometimes used to refer specifically to retinal ganglion cells....
, 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
Esophagus
The esophagus is an organ in vertebrates which consists of a muscular tube through which food passes from the pharynx to the stomach. During swallowing, food passes from the mouth through the pharynx into the esophagus and travels via peristalsis to the stomach...
, called the subesophageal ganglion
Subesophageal ganglion
The subesophageal ganglion of insects is composed of three pairs of fused ganglia. It controls the mouthparts, the salivary glands and certain muscles....
.
The thoracic segments have one ganglion on each side, which are connected into a pair, one pair per segment. This arrangement is also seen in the abdomen but only in the first eight segments. Many species of insects have reduced numbers of ganglia due to fusion or reduction. Some cockroaches have just six ganglia in the abdomen, whereas the wasp Vespa crabro has only two in the thorax and three in the abdomen. Some insects, like the house fly Musca domestica
Housefly
The housefly , Musca domestica, is a fly of the suborder Cyclorrhapha...
, have all the body ganglia fused into a single large thoracic ganglion.
At least a few insects have nociceptor
Nociceptor
A nociceptor is a sensory receptor that responds to potentially damaging stimuli by sending nerve signals to the spinal cord and brain. This process, called nociception, usually causes the perception of pain.-History:...
s, cells that detect and transmit sensations of pain
Pain in animals
Pain is a sensory and emotional experience often caused by intense or damaging stimuli. The International Association for the Study of Pain says pain is a conscious experience involving unpleasantness, i.e. suffering...
. This was discovered in 2003 by studying the variation in reactions of larvae
Larvae
In Roman mythology, lemures were shades or spirits of the restless or malignant dead, and are probably cognate with an extended sense of larvae as disturbing or frightening...
of the common fruitfly Drosophila
Drosophila
Drosophila is a genus of small flies, belonging to the family Drosophilidae, whose members are often called "fruit flies" or more appropriately pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many species to linger around overripe or rotting fruit...
to the touch of a heated probe and an unheated one. The larvae reacted to the touch of the heated probe with a stereotypical rolling behavior that was not exhibited when the larvae were touched by the unheated probe. Although nociception has been demonstrated in insects, there is not a consensus that insects feel pain consciously but see Pain in invertebrates
Pain in invertebrates
Pain in invertebrates is a problematic issue. Although there are numerous definitions of pain, almost all involve two key components. First, nociception is required. This is the ability to detect noxious stimuli which evokes a reflex response that moves the entire animal, or the affected part of...
.
Digestive system
An insect uses its digestive system to extract nutrients and other substances from the food it consumes. Most of this food is ingested in the form of macromoleculeMacromolecule
A macromolecule is a very large molecule commonly created by some form of polymerization. In biochemistry, the term is applied to the four conventional biopolymers , as well as non-polymeric molecules with large molecular mass such as macrocycles...
s and other complex substances like protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...
s, polysaccharide
Polysaccharide
Polysaccharides are long carbohydrate molecules, of repeated monomer units joined together by glycosidic bonds. They range in structure from linear to highly branched. Polysaccharides are often quite heterogeneous, containing slight modifications of the repeating unit. Depending on the structure,...
s, fat
Fat
Fats consist of a wide group of compounds that are generally soluble in organic solvents and generally insoluble in water. Chemically, fats are triglycerides, triesters of glycerol and any of several fatty acids. Fats may be either solid or liquid at room temperature, depending on their structure...
s, and nucleic acid
Nucleic acid
Nucleic acids are biological molecules essential for life, and include DNA and RNA . Together with proteins, nucleic acids make up the most important macromolecules; each is found in abundance in all living things, where they function in encoding, transmitting and expressing genetic information...
s. These macromolecules must be broken down by catabolic reaction
Catabolism
Catabolism is the set of metabolic pathways that break down molecules into smaller units and release energy. In catabolism, large molecules such as polysaccharides, lipids, nucleic acids and proteins are broken down into smaller units such as monosaccharides, fatty acids, nucleotides, and amino...
s into smaller molecules like amino acid
Amino acid
Amino acids are molecules containing an amine group, a carboxylic acid group and a side-chain that varies between different amino acids. The key elements of an amino acid are carbon, hydrogen, oxygen, and nitrogen...
s and simple sugars before being used by cells of the body for energy, growth, or reproduction. This break-down process is known as digestion.
The main structure of an insect's digestive system is a long enclosed tube called the alimentary canal, which runs lengthwise through the body. The alimentary canal directs food unidirectionally from the mouth
Mouth
The mouth is the first portion of the alimentary canal that receives food andsaliva. The oral mucosa is the mucous membrane epithelium lining the inside of the mouth....
to the anus
Anus
The anus is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, unwanted semi-solid matter produced during digestion, which, depending on the type of animal, may be one or more of: matter which the animal cannot digest,...
. It has three sections, each of which performs a different process of digestion. In addition to the alimentary canal, insects also have paired salivary glands and salivary reservoirs. These structures usually reside in the thorax, adjacent to the foregut.
The salivary gland
Salivary gland
The salivary glands in mammals are exocrine glands, glands with ducts, that produce saliva. They also secrete amylase, an enzyme that breaks down starch into maltose...
s (element 30 in numbered diagram) in an insect's mouth produce saliva. The salivary ducts lead from the glands to the reservoirs and then forward through the head to an opening called the salivarium, located behind the hypopharynx. By moving its mouthparts (element 32 in numbered diagram) the insect can mix its food with saliva. The mixture of saliva and food then travels through the salivary tubes into the mouth, where it begins to break down. Some insects, like flies
Diptera
Diptera , or true flies, is the order of insects possessing only a single pair of wings on the mesothorax; the metathorax bears a pair of drumstick like structures called the halteres, the remnants of the hind wings. It is a large order, containing an estimated 240,000 species, although under half...
, have extra-oral digestion
Mastication
Mastication or chewing is the process by which food is crushed and ground by teeth. It is the first step of digestion and it increases the surface area of foods to allow more efficient break down by enzymes. During the mastication process, the food is positioned between the teeth for grinding by...
. Insects using extra-oral digestion expel digestive enzymes onto their food to break it down. This strategy allows insects to extract a significant proportion of the available nutrients from the food source. The gut is where almost all of insects' digestion takes place. It can be divided into the foregut
Foregut
The foregut is the anterior part of the alimentary canal, from the mouth to the duodenum at the entrance of the bile duct. At this point it is continuous with the midgut...
, midgut
Stomach
The stomach is a muscular, hollow, dilated part of the alimentary canal which functions as an important organ of the digestive tract in some animals, including vertebrates, echinoderms, insects , and molluscs. It is involved in the second phase of digestion, following mastication .The stomach is...
and hindgut
Hindgut
The hindgut is the posterior part of the alimentary canal. In mammals, it includes the distal third of the transverse colon and the splenic flexure, the descending colon, sigmoid colon and rectum.-Blood flow:...
.
Foregut
The first section of the alimentary canal is the foregut
Foregut
The foregut is the anterior part of the alimentary canal, from the mouth to the duodenum at the entrance of the bile duct. At this point it is continuous with the midgut...
(element 27 in numbered diagram), or stomodaeum. The foregut is lined with a cuticular lining made of chitin
Chitin
Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world...
and protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...
s as protection from tough food. The foregut includes the buccal cavity (mouth), pharynx
Pharynx
The human pharynx is the part of the throat situated immediately posterior to the mouth and nasal cavity, and anterior to the esophagus and larynx. The human pharynx is conventionally divided into three sections: the nasopharynx , the oropharynx , and the laryngopharynx...
, esophagus
Esophagus
The esophagus is an organ in vertebrates which consists of a muscular tube through which food passes from the pharynx to the stomach. During swallowing, food passes from the mouth through the pharynx into the esophagus and travels via peristalsis to the stomach...
, and Crop
Crop (anatomy)
A crop is a thin-walled expanded portion of the alimentary tract used for the storage of food prior to digestion that is found in many animals, including gastropods, earthworms, leeches, insects, birds, and even some dinosaurs.- Bees :Cropping is used by bees to temporarily store nectar of flowers...
and proventriculus
Proventriculus
The proventriculus is part of the digestive system of birds, invertebrates and insects.-Birds:The proventriculus is a standard part of avian anatomy...
(any part may be highly modified) which both store food and signify when to continue passing onward to the midgut.
Digestion starts in buccal cavity (mouth) as partially chewed food is broken down by saliva from the salivary glands. As the salivary glands produce fluid and carbohydrate-digesting enzyme
Enzyme
Enzymes are proteins that catalyze chemical reactions. In enzymatic reactions, the molecules at the beginning of the process, called substrates, are converted into different molecules, called products. Almost all chemical reactions in a biological cell need enzymes in order to occur at rates...
s (mostly amylase
Amylase
Amylase is an enzyme that catalyses the breakdown of starch into sugars. Amylase is present in human saliva, where it begins the chemical process of digestion. Food that contains much starch but little sugar, such as rice and potato, taste slightly sweet as they are chewed because amylase turns...
s), strong muscles in the pharynx pump fluid into the buccal cavity, lubricating the food like the salivarium does, and helping blood feeders, and xylem and phloem feeders.
From there, the pharynx passes food to the esophagus, which could be just a simple tube passing it on to the crop and proventriculus, and then onward to the midgut, as in most insects. Alternately, the foregut may expand into a very enlarged crop and proventriculus, or the crop could just be a diverticulum
Diverticulum
A diverticulum is medical or biological term for an outpouching of a hollow structure in the body. Depending upon which layers of the structure are involved, they are described as being either true or false....
, or fluid filled structure, as in some Diptera species.
Midgut
Once food leaves the crop, it passes to the midgut
Stomach
The stomach is a muscular, hollow, dilated part of the alimentary canal which functions as an important organ of the digestive tract in some animals, including vertebrates, echinoderms, insects , and molluscs. It is involved in the second phase of digestion, following mastication .The stomach is...
(element 13 in numbered diagram), also known as the mesenteron, where the majority of digestion takes place. Microscopic projections from the midgut wall, called microvilli
Microvillus
Microvilli are microscopic cellular membrane protrusions that increase the surface area of cells, and are involved in a wide variety of functions, including absorption, secretion, cellular adhesion, and mechanotransduction....
, increase the surface area of the wall and allow more nutrients to be absorbed; they tend to be close to the origin of the midgut. In some insects, the role of the microvilli and where they are located may vary. For example, specialized microvilli producing digestive enzymes may more likely be near the end of the midgut, and absorption near the origin or beginning of the midgut.
Hindgut
In the hindgut
Hindgut
The hindgut is the posterior part of the alimentary canal. In mammals, it includes the distal third of the transverse colon and the splenic flexure, the descending colon, sigmoid colon and rectum.-Blood flow:...
(element 16 in numbered diagram), or proctodaeum, undigested food particles are joined by uric acid
Uric acid
Uric acid is a heterocyclic compound of carbon, nitrogen, oxygen, and hydrogen with the formula C5H4N4O3. It forms ions and salts known as urates and acid urates such as ammonium acid urate. Uric acid is created when the body breaks down purine nucleotides. High blood concentrations of uric acid...
to form fecal pellets. The rectum absorbs 90% of the water in these fecal pellets, and the dry pellet is then eliminated through the anus (element 17), completing the process of digestion. The uric acid is formed using hemolymph waste products diffused from the Malpighian tubule
Malpighian tubule system
The Malpighian tubule system is a type of excretory and osmoregulatory system found in some Uniramia , arachnids and tardigrades....
s (element 20). It is then emptied directly into the alimentary canal, at the junction between the midgut and hindgut. The number of Malpighian tubules possessed by a given insect varies between species, ranging from only two tubules in some insects to over 100 tubules in others.
Endocrine system
The salivary glandSalivary gland
The salivary glands in mammals are exocrine glands, glands with ducts, that produce saliva. They also secrete amylase, an enzyme that breaks down starch into maltose...
s (element 30 in numbered diagram) in an insect's mouth produce saliva. The salivary ducts lead from the glands to the reservoirs and then forward through the head to an opening called the salivarium, located behind the hypopharynx. By moving its mouthparts (element 32 in numbered diagram) the insect can mix its food with saliva. The mixture of saliva and food then travels through the salivary tubes into the mouth, where it begins to break down. Some insects, like flies
Diptera
Diptera , or true flies, is the order of insects possessing only a single pair of wings on the mesothorax; the metathorax bears a pair of drumstick like structures called the halteres, the remnants of the hind wings. It is a large order, containing an estimated 240,000 species, although under half...
, have extra-oral digestion
Mastication
Mastication or chewing is the process by which food is crushed and ground by teeth. It is the first step of digestion and it increases the surface area of foods to allow more efficient break down by enzymes. During the mastication process, the food is positioned between the teeth for grinding by...
. Insects using extra-oral digestion expel digestive enzymes onto their food to break it down. This strategy allows insects to extract a significant proportion of the available nutrients from the food source.
Reproductive system
The reproductive system of female insects consist of a pair of ovaries, accessory glands, one or more spermathecaSpermatheca
The spermatheca , also called receptaculum seminis , is an organ of the female reproductive tract in insects, some molluscs, oligochaeta worms and certain other invertebrates and vertebrates...
e, and ducts connecting these parts. The ovaries are made up of a number of egg tubes, called ovarioles
Ovarioles
An ovariole is one of the tubes of which the ovaries of most insects are composed. Typically an insect will have two ovaries. The constituent ovarioles lead to two oviducts, which converge into a single oviduct. The ovarioles are composed of a germarium and a set of ovarial follicles.-In D...
, which vary in size and number by species. The number of eggs that the insect is able to make vary by the number of ovarioles with the rate that eggs can be develop being also influenced by ovariole design. Female insects are able make eggs, receive and store sperm, manipulate sperm from different males, and lay eggs. Accessory glands or glandular parts of the oviducts produce a variety of substances for sperm maintenance, transport, and fertilization, as well as for protection of eggs. They can produce glue and protective substances for coating eggs or tough coverings for a batch of eggs called ootheca
Ootheca
An ootheca is a type of egg mass made by any member of a variety of species .The word is a latinized combination of oo-, meaning "egg", from the Greek word ōon , and theca, meaning a "cover" or "container", from the Greek theke...
e. Spermathecae are tubes or sacs in which sperm can be stored between the time of mating and the time an egg is fertilized.
For males, the reproductive system is the testis, suspended in the body cavity by trachea
Invertebrate trachea
The invertebrate trachea refers to the open respiratory system composed of spiracles, tracheae, and tracheoles that terrestrial arthropods have to transport metabolic gases to and from tissues....
e and the fat body. Most male insects have a pair of testes, inside of which are sperm tubes or follicles that are enclosed within a membranous sac. The follicles connect to the vas deferens by the vas efferens, and the two tubular vasa deferentia connect to a median ejaculatory duct that leads to the outside. A portion of the vas deferens is often enlarged to form the seminal vesicle, which stores the sperm before they are discharged into the female. The seminal vesicles have glandular linings that secrete nutrients for nourishment and maintenance of the sperm. The ejaculatory duct is derived from an invagination of the epidermal cells during development and, as a result, has a cuticular lining. The terminal portion of the ejaculatory duct may be sclerotized to form the intromittent organ, the aedeagus. The remainder of the male reproductive system is derived from embryonic mesoderm, except for the germ cells, or spermatogonia, which descend from the primordial pole cells very early during embryogenesis.
Respiratory and circulatory systems
Insect respiration is accomplished without lungLung
The lung is the essential respiration organ in many air-breathing animals, including most tetrapods, a few fish and a few snails. In mammals and the more complex life forms, the two lungs are located near the backbone on either side of the heart...
s. Instead, the insect respiratory system
Respiratory system of insects
The respiratory system of insects is the means by which insects obtain oxygen from the air.-The Respiratory System:Air enters the respiratory systems of most insects through a series of external openings called spiracles. These external openings, which act as muscular valves in some insects, lead...
uses 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 trachea
Invertebrate trachea
The invertebrate trachea refers to the open respiratory system composed of spiracles, tracheae, and tracheoles that terrestrial arthropods have to transport metabolic gases to and from tissues....
(element 8 in numbered diagram). Since oxygen is delivered directly, the circulatory system is not used to carry oxygen, and is therefore greatly reduced. The insect circulatory system has no vein
Vein
In the circulatory system, veins are blood vessels that carry blood towards the heart. Most veins carry deoxygenated blood from the tissues back to the heart; exceptions are the pulmonary and umbilical veins, both of which carry oxygenated blood to the heart...
s or arterie
Artery
Arteries are blood vessels that carry blood away from the heart. This blood is normally oxygenated, exceptions made for the pulmonary and umbilical arteries....
s, and instead consists of little more than a single, perforated dorsal tube which pulses peristaltically
Peristalsis
Peristalsis is a radially symmetrical contraction and relaxation of muscles which propagates in a wave down the muscular tube, in an anterograde fashion. In humans, peristalsis is found in the contraction of smooth muscles to propel contents through the digestive tract. Earthworms use a similar...
. Toward the thorax, the dorsal tube (element 14) divides into chambers and acts like the insect's heart. The opposite end of the dorsal tube is like the aorta of the insect circulating the hemolymph
Hemolymph
Hemolymph, or haemolymph, is a fluid in the circulatory system of some arthropods and is analogous to the fluids and cells making up both blood and interstitial fluid in vertebrates such as birds and mammals...
, arthropods' fluid analog of blood
Blood
Blood is a specialized bodily fluid in animals that delivers necessary substances such as nutrients and oxygen to the cells and transports metabolic waste products away from those same cells....
, inside the body cavity. Air is taken in through openings on the sides of the abdomen called spiracle
Spiracle
Spiracles are openings on the surface of some animals that usually lead to respiratory systems.-Vertebrates:The spiracle is a small hole behind each eye that opens to the mouth in some fishes. In the primitive jawless fish the first gill opening immediately behind the mouth is essentially similar...
s.
There are many different patterns of gas exchange
Gas exchange
Gas exchange is a process in biology where gases contained in an organism and atmosphere transfer or exchange. In human gas-exchange, gases contained in the blood of human bodies exchange with gases contained in the atmosphere. Human gas-exchange occurs in the lungs...
demonstrated by different groups of insects. Gas exchange patterns in insects can range from continuous and diffusive
Diffusion
Molecular diffusion, often called simply diffusion, is the thermal motion of all particles at temperatures above absolute zero. The rate of this movement is a function of temperature, viscosity of the fluid and the size of the particles...
ventilation, to discontinuous gas exchange
Discontinuous gas exchange
Discontinuous gas-exchange cycles , also called discontinuous ventilation or discontinuous ventilatory cycles, follow one of several patterns of arthropod gas exchange that have been documented primarily in insects; they occur when the insect is at rest...
. During continuous gas exchange, oxygen
Oxygen
Oxygen is the element with atomic number 8 and represented by the symbol O. Its name derives from the Greek roots ὀξύς and -γενής , because at the time of naming, it was mistakenly thought that all acids required oxygen in their composition...
is taken in and carbon dioxide
Carbon dioxide
Carbon dioxide is a naturally occurring chemical compound composed of two oxygen atoms covalently bonded to a single carbon atom...
is released in a continuous cycle. In discontinuous gas exchange, however, the insect takes in oxygen while it is active and small amounts of carbon dioxide are released when the insect is at rest. Diffusive ventilation is simply a form of continuous gas exchange that occurs by diffusion
Diffusion
Molecular diffusion, often called simply diffusion, is the thermal motion of all particles at temperatures above absolute zero. The rate of this movement is a function of temperature, viscosity of the fluid and the size of the particles...
rather than physically taking in the oxygen. Some species of insect that are submerged also have adaptations to aid in respiration. As larvae, many insects have gills that can extract oxygen dissolved in water, while others need to rise to the water surface to replenish air supplies which may be held or trapped in special structures.
Reproduction and development
Egg (biology)
An egg is an organic vessel in which an embryo first begins to develop. In most birds, reptiles, insects, molluscs, fish, and monotremes, an egg is the zygote, resulting from fertilization of the ovum, which is expelled from the body and permitted to develop outside the body until the developing...
s. The fertilization and development takes place inside the egg, enclosed by a shell (chorion
Chorion
The chorion is one of the membranes that exist during pregnancy between the developing fetus and mother. It is formed by extraembryonic mesoderm and the two layers of trophoblast and surrounds the embryo and other membranes...
). Some species of insects, like the cockroach Blaptica dubia
Blaptica dubia
The Dubia Cockroach , also known as Orange-spotted Cockroach or Guyana Spotted Cockroach, is a medium/large species of cockroach, measuring about long. They are sexually dimorphic; the males have wings while the females have only tiny wing stubs...
, as well as juvenile aphids and tsetse flies, are ovoviviparous
Ovoviviparity
Ovoviviparity, ovovivipary, or ovivipary, is a mode of reproduction in animals in which embryos develop inside eggs that are retained within the mother's body until they are ready to hatch...
. The eggs of ovoviviparous animals develop entirely inside the female, and then hatch immediately upon being laid. Some other species, such as those in the genus of cockroaches known as Diploptera
Diploptera
Diploptera is a genus of cockroach, used particularly in molecular research into allatostatins. The species Diploptera punctata, the Pacific beetle cockroach, is the only known viviparous cockroach ....
, are viviparous, and thus gestate inside the mother and are born alive
Vivipary
Vivipary has two different meanings. In animals, it means development of the embryo inside the body of the mother, eventually leading to live birth, as opposed to laying eggs...
. Some insects, like parasitic wasps, show polyembryony
Polyembryony
Polyembryony is the phenomenon of two or more embryos developing from a single fertilized egg . Polyembryony occurs regularly in many plants and animals. The nine banded armadillo, for instance, usually gives birth to four identical young...
, where a single fertilized egg divides into many and in some cases thousands of separate embryos.
Polymorphism (biology)
Polymorphism in biology occurs when two or more clearly different phenotypes exist in the same population of a species — in other words, the occurrence of more than one form or morph...
, paedomorphosis or peramorphosis
Peramorphosis
In developmental biology, peramorphosis is a phylogenetic change in which individuals of a species mature past adulthood and take on hitherto unseen traits...
, sexual dimorphism
Sexual dimorphism
Sexual dimorphism is a phenotypic difference between males and females of the same species. Examples of such differences include differences in morphology, ornamentation, and behavior.-Examples:-Ornamentation / coloration:...
, parthenogenesis
Parthenogenesis
Parthenogenesis is a form of asexual reproduction found in females, where growth and development of embryos occur without fertilization by a male...
and more rarely hermaphroditism. In haplodiploidy, which is a type of sex-determination system
Sex-determination system
A sex-determination system is a biological system that determines the development of sexual characteristics in an organism. Most sexual organisms have two sexes. In many cases, sex determination is genetic: males and females have different alleles or even different genes that specify their sexual...
, the offspring's sex is determined by the number of sets of chromosome
Chromosome
A chromosome is an organized structure of DNA and protein found in cells. It is a single piece of coiled DNA containing many genes, regulatory elements and other nucleotide sequences. Chromosomes also contain DNA-bound proteins, which serve to package the DNA and control its functions.Chromosomes...
s an individual receives. This system is typical in bees and wasps. Polymorphism is the where a species may have different morphs or forms, as in the oblong winged katydid
Amblycorypha oblongifolia
The Oblong-Winged katydid is a species of katydid, or bush-cricket, of the family Tettigoniidae .-Appearance:...
, which has four different varieties: green, pink, and yellow or tan. Some insects may retain phenotype
Phenotype
A phenotype is an organism's observable characteristics or traits: such as its morphology, development, biochemical or physiological properties, behavior, and products of behavior...
s that are normally only seen in juveniles; this is called paedomorphosis. In peramorphosis, an opposite sort of phenomenon, insects take on previously unseen traits after they have matured into adults. Many insects display sexual dimorphism, in which males and females have notably different appearances, such as the moth Orgyia recens
Orgyia recens
The Scarce Vapourer is a species of moth of the Lymantriidae family. It is found in Europe.The wingspan is 35-40 mm for the males, the females are wingless. The moth flies from June to July depending on the location....
as an exemplar of sexual dimorphism in insects.
Some insects use parthenogenesis
Parthenogenesis
Parthenogenesis is a form of asexual reproduction found in females, where growth and development of embryos occur without fertilization by a male...
, a process in which the female can reproduce and give birth without having the eggs fertilized by a male
Male
Male refers to the biological sex of an organism, or part of an organism, which produces small mobile gametes, called spermatozoa. Each spermatozoon can fuse with a larger female gamete or ovum, in the process of fertilization...
. Many aphids undergo a form of parthenogenesis, called cyclical parthenogenesis, in which they alternate between one or many generations of asexual and sexual reproduction. 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 which they spawn males. However, overall, most individuals are female, which are produced by fertilization. The males are haploid and the females are diploid. More rarely, some insects display hermaphroditism, in which a given individual has both male and female reproductive organs.
Insect life-histories show adaptations to withstand cold and dry conditions. Some temperate region insects are capable of activity during winter, while some others migrate to a warmer climate or go into a state of torpor
Torpor
Torpor, sometimes called temporary hibernation is a state of decreased physiological activity in an animal, usually characterized by a reduced body temperature and rate of metabolism. Animals that go through torpor include birds and some mammals such as mice and bats...
. Still other insects have evolved mechanisms of diapause
Diapause
Diapause is the delay in development in response to regularly and recurring periods of adverse environmental conditions. It is considered to be a physiological state of dormancy with very specific initiating and inhibiting conditions...
that allow eggs or pupae to survive these conditions.
Metamorphosis
MetamorphosisMetamorphosis
Metamorphosis is a biological process by which an animal physically develops after birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation...
in insects is the biological process of development all insects must undergo. There are two forms of metamorphosis: incomplete metamorphosis and complete metamorphosis.
Incomplete metamorphosis
Insects that show hemimetabolismHemimetabolism
Hemimetabolism or hemimetaboly, also called incomplete metamorphosis, is a term used to describe the mode of development of certain insects that includes three distinct stages: the egg, nymph, and the adult stage, or imago. These groups go through gradual changes; there is no pupal stage...
, or incomplete metamorphosis, change gradually by undergoing a series of molt
Ecdysis
Ecdysis is the moulting of the cuticula in many invertebrates. This process of moulting is the defining feature of the clade Ecdysozoa, comprising the arthropods, nematodes, velvet worms, horsehair worms, rotifers, tardigrades and Cephalorhyncha...
s. An insect 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. 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, which makes the old cuticle split along predefined weaknesses where the old exocuticle was thinnest. Other arthropods have a much different process and only molt; though must accommodate for the difference in exoskeleton structure and make up with other enzymes.
Immature insects that go through incomplete metamorphosis are called nymph
Nymph (biology)
In biology, a nymph is the immature form of some invertebrates, particularly insects, which undergoes gradual metamorphosis before reaching its adult stage. Unlike a typical larva, a nymph's overall form already resembles that of the adult. In addition, while a nymph moults it never enters a...
s or in the case of dragonflies and damselflies as naiads. Nymphs are similar in form to the adult except for the presence of wings, which are not developed until adulthood. With each molt, nymphs grow larger and become more similar in appearance to adult insects.
Complete metamorphosis
Holometabolism
Holometabolism, also called complete metamorphism, is a term applied to insect groups to describe the specific kind of insect development which includes four life stages - as an embryo or egg, a larva, a pupa and an imago or adult. Holometabolism is a monophyletic trait that all insects in the...
, or complete metamorphosis, is where the insect changes all in four stages, an egg
Egg (biology)
An egg is an organic vessel in which an embryo first begins to develop. In most birds, reptiles, insects, molluscs, fish, and monotremes, an egg is the zygote, resulting from fertilization of the ovum, which is expelled from the body and permitted to develop outside the body until the developing...
or embryo
Embryo
An embryo is a multicellular diploid eukaryote in its earliest stage of development, from the time of first cell division until birth, hatching, or germination...
, a larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...
, a pupa
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...
, and the adult
Adult
An adult is a human being or living organism that is of relatively mature age, typically associated with sexual maturity and the attainment of reproductive age....
or imago
Imago
In biology, the imago is the last stage of development of an insect, after the last ecdysis of an incomplete metamorphosis, or after emergence from the pupa where the metamorphosis is complete...
. In these species, egg hatches to produce a larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...
, which is generally worm-like in form. This worm-like form can be one of several varieties: eruciform (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
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...
, a stage marked by reduced movement and often sealed within a cocoon
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...
. 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. Insects undergo considerable change in form during the pupal stage, and emerge as adults. Butterflies are a well known example of an insects that undergo complete metamorphosis, although most insects use this life cycle. Some insects have evolved this system to hypermetamorphosis
Hypermetamorphosis
Hypermetamorphosis is a kind of complete insect metamorphosis in which the different larval instars represent two or more different forms of larva. As the larva molts its morphology can change from that of a campodeiform to scarabaeiform or to vermiform...
.
Some of the oldest and most successful insect groups, such Endopterygota
Endopterygota
The Endopterygota, also known as Holometabola, are insects of the subclass Pterygota which go through distinctive larval, pupal, and adult stages. They undergo a radical metamorphosis, with the larval and adult stages differing considerably in their structure and behaviour...
, use a system of complete metamorphosis. Strangely though, complete metamorphosis is unique to certain insect orders, like Diptera
Diptera
Diptera , or true flies, is the order of insects possessing only a single pair of wings on the mesothorax; the metathorax bears a pair of drumstick like structures called the halteres, the remnants of the hind wings. It is a large order, containing an estimated 240,000 species, although under half...
, Lepidoptera
Lepidoptera
Lepidoptera is a large order of insects that includes moths and butterflies . It is one of the most widespread and widely recognizable insect orders in the world, encompassing moths and the three superfamilies of butterflies, skipper butterflies, and moth-butterflies...
, and Hymenoptera
Hymenoptera
Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...
, and no other arthropods undergo it, but incomplete metamorphosis.
Senses and communication
Many insects possess very sensitive and/or specialized organs of perceptionSense
Senses are physiological capacities of organisms that provide inputs for perception. The senses and their operation, classification, and theory are overlapping topics studied by a variety of fields, most notably neuroscience, cognitive psychology , and philosophy of perception...
. Some insects such as bees can perceive ultraviolet
Ultraviolet
Ultraviolet light is electromagnetic radiation with a wavelength shorter than that of visible light, but longer than X-rays, in the range 10 nm to 400 nm, and energies from 3 eV to 124 eV...
wavelengths, or detect polarized light, while the antennae
Antenna (biology)
Antennae in biology have historically been paired appendages used for sensing in arthropods. More recently, the term has also been applied to cilium structures present in most cell types of eukaryotes....
of male moths can detect the pheromone
Pheromone
A pheromone is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting outside the body of the secreting individual to impact the behavior of the receiving individual...
s of female moths over distances of many kilometers. There is a pronounced tendency for there to be 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. There are a variety of different mechanisms by which insects perceive sound, while the patterns are not universal, insects can generally hear sound if they can produce it. Different insect species can have varying hearing, though most insects can hear only a narrow range of frequencies related to the frequency of the sounds they can produce. Mosquitoes have been found to hear up to 2 MHz., and some grasshoppers can hear up to 50 MHz. Certain predatory and parasitic insects can detect the characteristic sounds made by their prey or hosts, respectively. For instance, some nocturnal moths can perceive the ultrasonic
Ultrasound
Ultrasound is cyclic sound pressure with a frequency greater than the upper limit of human hearing. Ultrasound is thus not separated from "normal" sound based on differences in physical properties, only the fact that humans cannot hear it. Although this limit varies from person to person, it is...
emissions of bat
Bat
Bats are mammals of the order Chiroptera "hand" and pteron "wing") whose forelimbs form webbed wings, making them the only mammals naturally capable of true and sustained flight. By contrast, other mammals said to fly, such as flying squirrels, gliding possums, and colugos, glide rather than fly,...
s, which helps them avoid predation. Insects that feed on blood have special sensory structures that can detect infrared
Infrared
Infrared light is electromagnetic radiation with a wavelength longer than that of visible light, measured from the nominal edge of visible red light at 0.74 micrometres , and extending conventionally to 300 µm...
emissions, and use them to home in on their hosts.
Some insects display a rudimentary sense of numbers
Numeracy
Numeracy is the ability to reason with numbers and other mathematical concepts. A numerically literate person can manage and respond to the mathematical demands of life...
, such as the solitary wasps that prey upon a single species. The mother wasp lays her eggs in individual cells and provides each egg with a number of live caterpillars on which the young feed when hatched. Some species of wasp always provide five, others twelve, and others as high as twenty-four caterpillars per cell. The number of caterpillars is different among species, but always the same for each sex of larva. The male solitary wasp in the genus Eumenes
Eumenes (genus)
Eumenes is the type genus of the subfamily Eumeninae of Vespidae. It is a large and widespread genus, with over 100 taxa , mostly occurring in the temperate portions of the Northern Hemisphere. Most species are black or brown, and commonly marked with strikingly contrasting patterns of yellow,...
is smaller than the female, so the mother of one species supplies him with only five caterpillars; the larger female receives ten caterpillars in her cell.
Light production and vision
Mycetophilidae
Mycetophilidae is a family of small flies, forming the bulk of those species known as fungus gnats. There are approximately 3000 described species in 150 genera but the true number of species is undoubtedly much higher...
(Diptera), and the beetle families Lampyridae, Phengodidae
Phengodidae
The beetle family Phengodidae is known also as glowworm beetles, whose larvae are known as glowworms. The females and larvae have bioluminescent organs. They occur throughout the New World from extreme southern Canada to Chile...
, Elateridae and Staphylinidae are bioluminescent. The most familiar group are the fireflies
Fireflies
Fireflies is a novel by Shiva Naipaul originally published in 1970. It was his first book, a comic novel set in Trinidad. In an essay in An Unfinished Journey, Naipaul described how in 1968 as a final year student at Oxford University studying Chinese, he had been moved to write down a sentence,...
, 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
Arachnocampa
Arachnocampa is a genus of four fungus gnat species which are, in their larval stage, glowworms. They are found mostly in New Zealand and Australia in caves and grottos, or sheltered places in forests....
(Mycetophilidae, Fungus gnats) glow to lure small flying insects into sticky strands of silk.
Some fireflies of the genus Photuris mimic the flashing of female Photinus
Photinus (beetle)
The rover fireflies are a genus of fireflies . They are the type genus of tribe Photinini in subfamily Lampyrinae. This genus contains, for example, the common eastern firefly , the state insect of Tennessee.Male Photinus beetles emit a flashing light pattern to signal for females...
species to attract males of that species, which are then captured and devoured. The colors of emitted light vary from dull blue (Orfelia fultoni, Mycetophilidae) to the familiar greens and the rare reds (Phrixothrix tiemanni, Phengodidae).
Most insects, except some species of cave crickets
Rhaphidophoridae
The orthopteran family Rhaphidophoridae includes the cave wetas, cave crickets, camelback crickets, camel crickets, spider crickets and sand treaders, of the suborder Ensifera; in some regions, such as Missouri and Virginia, these crickets are referred to as "Cricket Spiders"...
, are able to perceive light and dark. Many species have acute vision capable of detecting minute movements. The eyes include simple eyes or ocelli
Ocellus
A simple eye refers to a type of eye design or optical arrangement that contains a single lens which detect light. A "simple eye" is so-called in distinction from a multi-lensed "compound eye", and is not necessarily at all simple in the usual sense of the word...
as well as compound eyes of varying sizes. Many species are able to detect light in the infrared, ultraviolet and the visible light wavelengths. Color vision has been demonstrated in many species and phylogenetic analysis suggests that UV-green-blue trichromacy existed from at least the Devonian
Devonian
The Devonian is a geologic period and system of the Paleozoic Era spanning from the end of the Silurian Period, about 416.0 ± 2.8 Mya , to the beginning of the Carboniferous Period, about 359.2 ± 2.5 Mya...
period between 416 and 359 million years ago.
Sound production and hearing
Insects were the earliest organisms to produce and sense sounds. Insects make sounds mostly by mechanical action of appendages. In grasshopperGrasshopper
The grasshopper is an insect of the suborder Caelifera in the order Orthoptera. To distinguish it from bush crickets or katydids, it is sometimes referred to as the short-horned grasshopper...
s and crickets, this is achieved by stridulation
Stridulation
Stridulation is the act of producing sound by rubbing together certain body parts. This behavior is mostly associated with insects, but other animals are known to do this as well, such as a number of species of fishes, snakes and spiders...
. Cicada
Cicada
A cicada is an insect of the order Hemiptera, suborder Auchenorrhyncha , in the superfamily Cicadoidea, with large eyes wide apart on the head and usually transparent, well-veined wings. There are about 2,500 species of cicada around the world, and many of them remain unclassified...
s make the loudest sounds among the insects by producing and amplifying sounds with special modifications to their body and musculature. The African cicada
Cicada
A cicada is an insect of the order Hemiptera, suborder Auchenorrhyncha , in the superfamily Cicadoidea, with large eyes wide apart on the head and usually transparent, well-veined wings. There are about 2,500 species of cicada around the world, and many of them remain unclassified...
Brevisana brevis
Brevisana brevis
Brevisana brevis is a cicada found in Africa and is the loudest insect on record. It has been recorded producing sounds with pressure levels of 106.7 decibels at a distance of 50 cm....
has been measured at 106.7 decibel
Decibel
The decibel is a logarithmic unit that indicates the ratio of a physical quantity relative to a specified or implied reference level. A ratio in decibels is ten times the logarithm to base 10 of the ratio of two power quantities...
s at a distance of 50 cm (19.7 in). Some insects, such as the hawk moths and Hedylid
Hedylidae
Hedylidae, the "American moth-butterflies", is a family of insects in the lepidopteran order, representing the superfamily Hedyloidea. They are an extant sister group of the butterfly superfamilies Papilionoidea and Hesperioidea...
butterflies, can hear ultrasound
Ultrasound
Ultrasound is cyclic sound pressure with a frequency greater than the upper limit of human hearing. Ultrasound is thus not separated from "normal" sound based on differences in physical properties, only the fact that humans cannot hear it. Although this limit varies from person to person, it is...
and take evasive action when they sense that they have been detected by bats. Some moths produce ultrasonic clicks that were once thought to have a role in jamming bat echolocation
Animal echolocation
Echolocation, also called biosonar, is the biological sonar used by several kinds of animals.Echolocating animals emit calls out to the environment and listen to the echoes of those calls that return from various objects near them. They use these echoes to locate and identify the objects...
. The ultrasonic clicks were subsequently found to be produced mostly by unpalatable moths to warn bats, just as warning coloration
Aposematism
Aposematism , perhaps most commonly known in the context of warning colouration, describes a family of antipredator adaptations where a warning signal is associated with the unprofitability of a prey item to potential predators...
s are used against predators that hunt by sight. Some otherwise palatable moths have evolved to mimic
Batesian mimicry
Batesian mimicry is a form of mimicry typified by a situation where a harmless species has evolved to imitate the warning signals of a harmful species directed at a common predator...
these calls.
Very low sounds are also produced in various species of Coleoptera, Hymenoptera
Hymenoptera
Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...
, Lepidoptera
Lepidoptera
Lepidoptera is a large order of insects that includes moths and butterflies . It is one of the most widespread and widely recognizable insect orders in the world, encompassing moths and the three superfamilies of butterflies, skipper butterflies, and moth-butterflies...
, Mantodea, and Neuroptera
Neuroptera
The insect order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order contains some 6,010 species...
. These low sounds are simply the sounds made by the insect's movement. Through microscopic stridulatory structures located on the insect's muscles and joints, the normal sounds of the insect moving are amplified and can be used to warn or communicate with other insects. Most sound-making insects also have tympanal organ
Tympanal organ
A tympanal organ is a hearing organ in insects, consisting of a membrane stretched across a frame backed by an air sac and associated sensory neurons...
s that can perceive airborne sounds. Some species in Hemiptera
Hemiptera
Hemiptera is an order of insects most often known as the true bugs , comprising around 50,000–80,000 species of cicadas, aphids, planthoppers, leafhoppers, shield bugs, and others...
, such as the corixids (water boatmen), are known to communicate via underwater sounds. Most insects are also able to sense vibration
Vibration
Vibration refers to mechanical oscillations about an equilibrium point. The oscillations may be periodic such as the motion of a pendulum or random such as the movement of a tire on a gravel road.Vibration is occasionally "desirable"...
s transmitted through surfaces. For example, an insect is caught in a spider web
Spider web
A spider web, spiderweb, spider's web or cobweb is a device built by a spider out of proteinaceous spider silk extruded from its spinnerets....
and struggles to escape. The vibrations it produces are sensed by the spider
Spider
Spiders are air-breathing arthropods that have eight legs, and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other groups of organisms...
, who is alerted to its presence. Through these vibrations, the spider can tell where on the web the insect is located, as well as how big it is.
Communication using surface-borne vibrational signals is more widespread among insects because of size constraints in producing air-borne sounds. 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. The mechanisms of production of vibrational signals are just as diverse as those for producing sound in insects.
Some species use vibrations for communicating within members of the same species, such as to attract mates as in the songs of the shield bug
Shield bug
Pentatomoidea is a superfamily of insects in the Heteroptera suborder of the Hemiptera order and, as such, share a common arrangement of sucking mouthparts...
Nezara viridula
Nezara viridula
Nezara viridula, commonly known as the southern green stink bug or green vegetable bug , is a plant-feeding stink bug. Although believed to have originated in Ethiopia, it can now be found around the world...
. Vibrations can also be used to communicate between entirely different species; lycaenid (gossamer-winged butterfly) caterpillars which are myrmecophilous (living in a mutualistic association with ants) communicate with ants in this way. The Madagascar hissing cockroach
Madagascar hissing cockroach
The Madagascar hissing Cockroach , also known as the Hissing Cockroach or simply Hisser, is one of the largest species of cockroach, reaching 2–4 inches at maturity. They are from the island of Madagascar off the African coast, where they can be found in rotting logs.Unlike most cockroaches, they...
has the ability to press air through its spiracles to make a hissing noise as a sign of aggression; the Death's-head Hawkmoth
Death's-head Hawkmoth
The name Death's-head Hawkmoth refers to any one of the three species of moth in the genus Acherontia. The former species is primarily found in Europe, the latter two are Asian, and most uses of the common name refer to the European species...
makes a squeaking noise by forcing air out of their pharynx
Pharynx
The human pharynx is the part of the throat situated immediately posterior to the mouth and nasal cavity, and anterior to the esophagus and larynx. The human pharynx is conventionally divided into three sections: the nasopharynx , the oropharynx , and the laryngopharynx...
when agitated, which may also reduce aggressive worker honey bee behavior when the two are in close proximity.
Chemical communication
In addition to the use of sound for communication, a wide range of insects have evolved chemical means for communication. These chemicals, termed semiochemicalSemiochemical
A semiochemical is a generic term used for a chemical substance or mixture that carries a message. These chemicals acts as messengers within or between species...
s, are often derived from plant metabolites include those meant to attract, repel and provide other kinds of information. Pheromone
Pheromone
A pheromone is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting outside the body of the secreting individual to impact the behavior of the receiving individual...
s, a type of semiochemical, are used for attracting mates of the opposite sex, for aggregating conspecific
Conspecificity
Conspecificity is a concept in biology. Two or more individual organisms, populations, or taxa are conspecific if they belong to the same species....
individuals of both sexes, for deterring other individuals from approaching, to mark a trail, and to trigger aggression in nearby individuals. Allomone
Allomone
An allomone is any chemical substance produced and released by an individual of one species that affects the behaviour of a member of another species to the benefit of the originator but not the receiver...
a benefit their producer by the effect they have upon the receiver. Kairomone
Kairomone
A kairomone is a semiochemical, emitted by an organism, which mediates interspecific interactions in a way that benefits an individual of another species which receives it, without benefiting the emitter. This "eavesdropping" is often disadvantageous to the producer...
s benefit their receiver instead of their producer. Synomones benefit the producer and the receiver. While some chemicals are targeted at individuals of the same species, others are used for communication across species. The use of scents is especially well known to have developed in social insects.
Social behavior
Termite
Termites are a group of eusocial insects that, until recently, were classified at the taxonomic rank of order Isoptera , but are now accepted as the epifamily Termitoidae, of the cockroach order Blattodea...
s, ant
Ant
Ants are social insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than...
s and many bee
Bee
Bees are flying insects closely related to wasps and ants, and are known for their role in pollination and for producing honey and beeswax. Bees are a monophyletic lineage within the superfamily Apoidea, presently classified by the unranked taxon name Anthophila...
s and wasp
Wasp
The term wasp is typically defined as any insect of the order Hymenoptera and suborder Apocrita that is neither a bee nor an ant. Almost every pest insect species has at least one wasp species that preys upon it or parasitizes it, making wasps critically important in natural control of their...
s, are the most familiar species of eusocial
Eusociality
Eusociality is a term used for the highest level of social organization in a hierarchical classification....
animal. They live together in large well-organized colonies that may be so tightly integrated and genetically similar that the colonies of some species are sometimes considered superorganism
Superorganism
A superorganism is an organism consisting of many organisms. This is usually meant to be a social unit of eusocial animals, where division of labour is highly specialised and where individuals are not able to survive by themselves for extended periods of time. Ants are the best-known example of...
s. It is sometimes argued that the various species of honey bee
Honey bee
Honey bees are a subset of bees in the genus Apis, primarily distinguished by the production and storage of honey and the construction of perennial, colonial nests out of wax. Honey bees are the only extant members of the tribe Apini, all in the genus Apis...
are the only invertebrates (and indeed one of the few non-human groups) to have evolved a system of abstract symbolic communication where a behavior is used to represent and convey specific information about something in the environment. In this communication system, called dance language
Bee learning and communication
Honey bees learn and communicate in order to find food sources and for other means.-Learning:Learning is essential for efficient foraging. Honey bees are unlikely to make many repeat visits if a plant provides little in the way of reward...
, 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.
Only insects which live in nests or colonies demonstrate any true capacity for fine-scale spatial orientation or homing. This can allow an insect to return unerringly to a single hole a few millimeters in diameter among thousands of apparently identical holes clustered together, after a trip of up to several kilometers' distance. In a phenomenon known as philopatry
Philopatry
Broadly, philopatry is the behaviour of remaining in, or returning to, an individual's birthplace. More specifically, in ecology philopatry is the behaviour of elder offspring sharing the parental burden in the upbringing of their siblings, a classic example of kin selection...
, insects that hibernate
Hibernation
Hibernation is a state of inactivity and metabolic depression in animals, characterized by lower body temperature, slower breathing, and lower metabolic rate. Hibernating animals conserve food, especially during winter when food supplies are limited, tapping energy reserves, body fat, at a slow rate...
have shown the ability to recall a specific location up to a year after last viewing the area of interest. A few insects seasonally migrate large distances between different geographic regions (e.g., the overwintering areas of the Monarch butterfly
Monarch butterfly
The Monarch butterfly is a milkweed butterfly , in the family Nymphalidae. It is perhaps the best known of all North American butterflies. Since the 19th century, it has been found in New Zealand, and in Australia since 1871 where it is called the Wanderer...
).
Care of young
Some bees, ants, termites and some wasps are eusocial, build nest, guard eggs and provide food for offsprings fulltime (see EusocialityEusociality
Eusociality is a term used for the highest level of social organization in a hierarchical classification....
).
Most insects, however, lead short lives as adults, and rarely interact with one another except to mate or compete for mates. A small number exhibit some form of parental care, where they will at least guard their eggs, and sometimes continue guarding their offspring until adulthood, and possibly even feeding them. Another simple form of parental care is to construct a nest (a burrow or an actual construction, either of which may be simple or complex), store provisions in it, and lay an egg upon those provisions. The adult does not contact the growing offspring, but it nonetheless does provide food. This sort of care is typical for most species of bees and various types of wasps.
Flight
Insects are the only group of invertebrates to have developed flight. The evolution of insect wings has been a subject of debate. Some entomologistEntomology
Entomology is the scientific study of insects, a branch of arthropodology...
s suggest that the wings are from paranotal lobes, or extensions from the insect's exoskeleton called the nota
Notum
The notum is the dorsal portion of an insect's thoracic segment. It is typically a single large sclerite. It can be further divided into the alinotum which bears the wings and the postnotum which bears the phragma....
, called the paranotal theory. Other theories are based on a pleural origin. These theories include suggestions that wings originated from modified gills, spiracular flaps or as from an appendage of the epicoxa. The epicoxal theory suggests the insect wings are modified epicoxal exites, a modified appendage at the base of the legs or coxa
Arthropod leg
The arthropod leg is a form of jointed appendage of arthropods, usually used for walking. Many of the terms used for arthropod leg segments are of Latin origin, and may be confused with terms for bones: coxa , trochanter , femur, tibia, tarsus, ischium, metatarsus, carpus, dactylus ,...
. In the Carboniferous
Carboniferous
The Carboniferous is a geologic period and system that extends from the end of the Devonian Period, about 359.2 ± 2.5 Mya , to the beginning of the Permian Period, about 299.0 ± 0.8 Mya . The name is derived from the Latin word for coal, carbo. Carboniferous means "coal-bearing"...
age, some of the Meganeura
Meganeura
Meganeura is a genus of extinct insects from the Carboniferous period approximately 300 million years ago, which resembled and are related to the present-day dragonflies. With wingspans of more than 75 cm , M. monyi is one of the largest known flying insect species; the Permian Meganeuropsis...
dragonflies had as much as a 50 cm (19.7 in) wide wingspan. The appearance of gigantic insects has been found to be consistent with high atmospheric oxygen. The respiratory system of insects constrains their size, however the high oxygen in the atmosphere allowed larger sizes. The largest flying insects today are much smaller and include several moth species such as the Atlas moth and the White Witch (Thysania agrippina). Insect flight has been a topic of great interest in aerodynamics
Aerodynamics
Aerodynamics is a branch of dynamics concerned with studying the motion of air, particularly when it interacts with a moving object. Aerodynamics is a subfield of fluid dynamics and gas dynamics, with much theory shared between them. Aerodynamics is often used synonymously with gas dynamics, with...
due partly to the inability of steady-state theories to explain the lift generated by the tiny wings of insects.
Unlike bird
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...
s, many small insects are swept along by the prevailing winds although many of the larger insects are known to make migrations. Aphid
Aphid
Aphids, also known as plant lice and in Britain and the Commonwealth as greenflies, blackflies or whiteflies, are small sap sucking insects, and members of the superfamily Aphidoidea. Aphids are among the most destructive insect pests on cultivated plants in temperate regions...
s are known to be transported long distances by low-level jet stream
Jet stream
Jet streams are fast flowing, narrow air currents found in the atmospheres of some planets, including Earth. The main jet streams are located near the tropopause, the transition between the troposphere and the stratosphere . The major jet streams on Earth are westerly winds...
s. As such, fine line patterns associated with converging winds within weather radar
Weather radar
Weather radar, also called weather surveillance radar and Doppler weather radar, is a type of radar used to locate precipitation, calculate its motion, estimate its type . Modern weather radars are mostly pulse-Doppler radars, capable of detecting the motion of rain droplets in addition to the...
imagery, like the WSR-88D radar network, often represent large groups of insects.
Walking
Many adult insects use six legs for walking and have adopted a tripedalTripedal
Tripedal is the term used for any animal that stands on three legs. The terms bipedal and quadrupedal are used more commonly when referring to animals that either walk on two legs Tripedal (from the Latin tri = three + ped = foot) is the term used for (or would be used for) any animal that stands...
gait
Gait
Gait is the pattern of movement of the limbs of animals, including humans, during locomotion over a solid substrate. Most animals use a variety of gaits, selecting gait based on speed, terrain, the need to maneuver, and energetic efficiency...
. The tripedal gait allows for rapid walking while always having a stable stance and has been studied extensively in cockroach
Cockroach
Cockroaches are insects of the order Blattaria or Blattodea, of which about 30 species out of 4,500 total are associated with human habitations...
es. The legs are used in alternate triangles touching the ground. 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 position. When they touch the ground to form a new stable triangle the other legs can be lifted and brought forward in turn and so on. The purest form of the tripedal gait is seen in insects moving at high speeds. However, this type of locomotion is not rigid and insects can adapt a variety of gaits. For example, when moving slowly, turning, or avoiding obstacles, four or more feet may be touching the ground. Insects can also adapt their gait to cope with the loss of one or more limbs.
Cockroaches are among the fastest insect runners and, at full speed, adopt a bipedal run to reach a high velocity in proportion to their body size. As cockroaches move very quickly, they need to be video recorded at several hundred frames per second to reveal their gait. More sedate locomotion is seen in the stick insects or walking sticks (Phasmatodea
Phasmatodea
The Phasmatodea are an order of insects, whose members are variously known as stick insects , walking sticks or stick-bugs , phasmids, ghost insects and leaf insects...
). A few insects have evolved to walk on the surface of the water, especially the bugs of the Gerridae
Water strider
Gerridae is a family of true bugs in the order Hemiptera, commonly known as water striders, water bugs, magic bugs, pond skaters, skaters, skimmers, water scooters, water skaters, water skeeters, water skimmers, water skippers, water spiders, or Jesus bugs...
family, commonly known as water striders. A few species of ocean-skaters in the genus Halobates
Halobates
Halobates or sea skaters are a genus with over 40 species of water striders. While many are coastal, about five of these are able to survive and stand on the surface of the open ocean, a habitat containing very few insect species. They are predators, coastal species feeding mainly on fallen...
even live on the surface of open oceans, a habitat that has few insect species.
Use in robotics
Insect walking is of particular interest as an alternative form of locomotion in robotRobot
A robot is a mechanical or virtual intelligent agent that can perform tasks automatically or with guidance, typically by remote control. In practice a robot is usually an electro-mechanical machine that is guided by computer and electronic programming. Robots can be autonomous, semi-autonomous or...
s. The study of insects and biped
Biped
Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs, or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning "two feet"...
s has a significant impact on possible robotic methods of transport. This may allow new robots to be designed that can traverse terrain
Terrain
Terrain, or land relief, is the vertical and horizontal dimension of land surface. When relief is described underwater, the term bathymetry is used...
that robots with wheel
Wheel
A wheel is a device that allows heavy objects to be moved easily through rotating on an axle through its center, facilitating movement or transportation while supporting a load, or performing labor in machines. Common examples found in transport applications. A wheel, together with an axle,...
s may be unable to handle.
Swimming
Water beetle
A water beetle is a beetle adapted to living in water. Water beetles rise to the water surface and take atmospheric air into their tracheal systems. There are approximately 2000 species of water beetles. The rest marine species tend to live in the intertidal zone...
s, have aquatic adults as well.
Many of these species have adaptations to help in under-water locomotion. Water beetles and water bugs have legs adapted into paddle-like structures. Dragonfly naiad
Naiad
In Greek mythology, the Naiads or Naiades were a type of nymph who presided over fountains, wells, springs, streams, and brooks....
s use jet propulsion, forcibly expelling water out of their rectal chamber. Some species like the water strider
Water strider
Gerridae is a family of true bugs in the order Hemiptera, commonly known as water striders, water bugs, magic bugs, pond skaters, skaters, skimmers, water scooters, water skaters, water skeeters, water skimmers, water skippers, water spiders, or Jesus bugs...
s are capable of walking on the surface of water. They can do this because their claws are not at the tips of the legs as in most insects, but recessed in a special groove further up the leg; this prevents the claws from piercing the water's surface film. Other insects such as the Rove beetle Stenus
Stenus
Stenus is a genus of semiaquatic rove beetles in the subfamily Steninae. They are known for swimming by emitting pygidial gland secretions that reduce surface tension and allow them to move on the surface of water....
are known to emit pygidial gland secretions that reduce surface tension making it possible for them to move on the surface of water by Marangoni propulsion
Marangoni effect
The Marangoni effect is the mass transfer along an interface between two fluids due to surface tension gradient...
(also known by the German
German language
German is a West Germanic language, related to and classified alongside English and Dutch. With an estimated 90 – 98 million native speakers, German is one of the world's major languages and is the most widely-spoken first language in the European Union....
term Entspannungsschwimmen).
Phylogeny and systematics
The evolutionEvolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...
ary relationships of insects to other animal groups remain unclear. Although more traditionally grouped with millipede
Millipede
Millipedes are arthropods that have two pairs of legs per segment . Each segment that has two pairs of legs is a result of two single segments fused together as one...
s and centipede
Centipede
Centipedes are arthropods belonging to the class Chilopoda of the subphylum Myriapoda. They are elongated metameric animals with one pair of legs per body segment. Despite the name, centipedes can have a varying number of legs from under 20 to over 300. Centipedes have an odd number of pairs of...
s, evidence has emerged favoring closer evolutionary ties
Common descent
In evolutionary biology, a group of organisms share common descent if they have a common ancestor. There is strong quantitative support for the theory that all living organisms on Earth are descended from a common ancestor....
with crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...
s. In the Pancrustacea
Pancrustacea
Pancrustacea is a clade, comprising all crustaceans and hexapods. This grouping is contrary to the Atelocerata hypothesis, in which Myriapoda and Hexapoda are sister taxa, and Crustacea are only more distantly related. As of 2010, the Pancrustacea taxon is considered well accepted. The clade has...
theory, insects, together with Remipedia
Remipedia
Remipedia is a class of blind crustaceans found in coastal aquifers which contain saline groundwater, with populations identified in almost every ocean basin so far explored, including in Australia, the Caribbean Sea, the Atlantic Ocean...
and Malacostraca
Malacostraca
Malacostraca is the largest of the six classes of crustaceans, containing over 25,000 extant species, divided among 16 orders. Its members display a greater diversity of body forms than any other class of animals, and include crabs, lobsters, shrimp, krill, woodlice, scuds , mantis shrimp and many...
, make up a natural clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...
. Other terrestrial arthropods, such as centipede
Centipede
Centipedes are arthropods belonging to the class Chilopoda of the subphylum Myriapoda. They are elongated metameric animals with one pair of legs per body segment. Despite the name, centipedes can have a varying number of legs from under 20 to over 300. Centipedes have an odd number of pairs of...
s, millipede
Millipede
Millipedes are arthropods that have two pairs of legs per segment . Each segment that has two pairs of legs is a result of two single segments fused together as one...
s, scorpion
Scorpion
Scorpions are predatory arthropod animals of the order Scorpiones within the class Arachnida. They have eight legs and are easily recognized by the pair of grasping claws and the narrow, segmented tail, often carried in a characteristic forward curve over the back, ending with a venomous stinger...
s and spider
Spider
Spiders are air-breathing arthropods that have eight legs, and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other groups of organisms...
s, are sometimes confused with insects since their body plans can appear similar, sharing (as do all arthropods) a jointed exoskeleton. However, upon closer examination their features differ significantly; most noticeably they do not have the six legs characteristic of adult insects.
A phylogenetic tree of the arthropods and related groups
The higher-level phylogeny of the arthropods continues to be a matter of debate and research. In 2008, researchers at Tufts University
Tufts University
Tufts University is a private research university located in Medford/Somerville, near Boston, Massachusetts. It is organized into ten schools, including two undergraduate programs and eight graduate divisions, on four campuses in Massachusetts and on the eastern border of France...
uncovered what they believe is the world's oldest known full-body impression of a primitive flying insect, a 300 million-year-old specimen from the Carboniferous Period. The oldest definitive insect fossil is the Devonian
Devonian
The Devonian is a geologic period and system of the Paleozoic Era spanning from the end of the Silurian Period, about 416.0 ± 2.8 Mya , to the beginning of the Carboniferous Period, about 359.2 ± 2.5 Mya...
Rhyniognatha hirsti
Rhyniognatha
Rhyniognatha hirsti is the world’s oldest known insect. It emerged very early during the Early Devonian Period, around 400 million years ago, when earth’s first terrestrial ecosystems were being formed.-Evidence:...
, from the 396 million year old Rhynie chert
Rhynie chert
The Rhynie chert is an Early Devonian sedimentary deposit exhibiting extraordinary fossil detail or completeness . It is exposed near the village of Rhynie, Aberdeenshire, Scotland; a second unit, the Windyfield chert, is located some 700 m away...
. It may have superficially resembled a modern-day silverfish
Silverfish
Lepisma saccharina, frequently called silverfish, fishmoths, carpet sharks or paramites, are small, wingless insects in the order Thysanura...
insect. This species already possessed dicondylic mandibles (two articulations in the mandible), a feature associated with winged insects, suggesting that wings may already have evolved at this time. Thus, the first insects probably appeared earlier, in the Silurian
Silurian
The Silurian is a geologic period and system that extends from the end of the Ordovician Period, about 443.7 ± 1.5 Mya , to the beginning of the Devonian Period, about 416.0 ± 2.8 Mya . As with other geologic periods, the rock beds that define the period's start and end are well identified, but the...
period.
There have been four super radiations of insects: beetle
Beetle
Coleoptera is an order of insects commonly called beetles. The word "coleoptera" is from the Greek , koleos, "sheath"; and , pteron, "wing", thus "sheathed wing". Coleoptera contains more species than any other order, constituting almost 25% of all known life-forms...
s (evolved ~300 million years ago), flies
Fly
True flies are insects of the order Diptera . They possess a pair of wings on the mesothorax and a pair of halteres, derived from the hind wings, on the metathorax...
(evolved ~250 million years ago), moth
Moth
A moth is an insect closely related to the butterfly, both being of the order Lepidoptera. Moths form the majority of this order; there are thought to be 150,000 to 250,000 different species of moth , with thousands of species yet to be described...
s and wasp
Wasp
The term wasp is typically defined as any insect of the order Hymenoptera and suborder Apocrita that is neither a bee nor an ant. Almost every pest insect species has at least one wasp species that preys upon it or parasitizes it, making wasps critically important in natural control of their...
s (evolved ~150 million years ago). These four groups account for the majority of described species. The flies and moths along with the flea
Flea
Flea is the common name for insects of the order Siphonaptera which are wingless insects with mouthparts adapted for piercing skin and sucking blood...
s evolved from the Mecoptera
Mecoptera
Mecoptera are an order of insects with about 550 species in nine families worldwide. Mecoptera are sometimes called scorpionflies after their largest family, Panorpidae, in which the males have enlarged genitals that look similar to the stinger of a scorpion...
.
The origins of insect flight
Insect flight
Insects are the only group of invertebrates known to have evolved flight. Insects possess some remarkable flight characteristics and abilities, still far superior to attempts by humans to replicate their capabilities. Even our understanding of the aerodynamics of flexible, flapping wings and how...
remain obscure, since the earliest winged insects currently known appear to have been capable fliers. Some extinct insects had an additional pair of winglets attaching to the first segment of the thorax, for a total of three pairs. As of 2009, there is no evidence that suggests that the insects were a particularly successful group of animals before they evolved to have wings.
Late Carboniferous
Pennsylvanian
The Pennsylvanian is, in the ICS geologic timescale, the younger of two subperiods of the Carboniferous Period. It lasted from roughly . As with most other geochronologic units, the rock beds that define the Pennsylvanian are well identified, but the exact date of the start and end are uncertain...
and Early Permian
Permian
The PermianThe term "Permian" was introduced into geology in 1841 by Sir Sir R. I. Murchison, president of the Geological Society of London, who identified typical strata in extensive Russian explorations undertaken with Edouard de Verneuil; Murchison asserted in 1841 that he named his "Permian...
insect orders include both extant groups and a number of Paleozoic species, now extinct. During this era, some giant dragonfly-like forms reached wingspans of 55 to 70 cm (21.7 to 27.6 in) making them far larger than any living insect. This gigantism may have been due to higher atmospheric oxygen levels that allowed increased respiratory efficiency relative to today. The lack of flying vertebrates could have been another factor. Most extinct orders of insects developed during the Permian period that began around 270 million years ago. Many of the early groups became extinct during the Permian-Triassic extinction event
Permian-Triassic extinction event
The Permian–Triassic extinction event, informally known as the Great Dying, was an extinction event that occurred 252.28 Ma ago, forming the boundary between the Permian and Triassic geologic periods, as well as the Paleozoic and Mesozoic eras...
, the largest mass extinction in the history of the Earth, around 252 million years ago.
The remarkably successful Hymenoptera
Hymenoptera
Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...
ns appeared as long as 146 million years ago in the Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...
period, but achieved their wide diversity more recently in the Cenozoic
Cenozoic
The Cenozoic era is the current and most recent of the three Phanerozoic geological eras and covers the period from 65.5 mya to the present. The era began in the wake of the Cretaceous–Tertiary extinction event at the end of the Cretaceous that saw the demise of the last non-avian dinosaurs and...
era, which began 66 million years ago. A number of highly successful insect groups evolved in conjunction with flowering plants, a powerful illustration of coevolution.
Many modern insect genera
Genera
Genera is a commercial operating system and development environment for Lisp machines developed by Symbolics. It is essentially a fork of an earlier operating system originating on the MIT AI Lab's Lisp machines which Symbolics had used in common with LMI and Texas Instruments...
developed during the Cenozoic. Insects from this period on are often found preserved in amber
Amber
Amber is fossilized tree resin , which has been appreciated for its color and natural beauty since Neolithic times. Amber is used as an ingredient in perfumes, as a healing agent in folk medicine, and as jewelry. There are five classes of amber, defined on the basis of their chemical constituents...
, often in perfect condition. The body plan, or morphology
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....
, of such specimens is thus easily compared with modern species. The study of fossilized insects is called paleoentomology
Entomology
Entomology is the scientific study of insects, a branch of arthropodology...
.
Evolutionary relationships
Insects are prey for a variety of organisms, including terrestrial vertebrates. The earliest vertebrates on land existed 400 million years ago and were large amphibious piscivoresPiscivore
A piscivore is a carnivorous animal which eats primarily fish. Piscivory was the diet of early tetrapods , insectivory came next, then in time reptiles added herbivory....
, through gradual evolutionary change, insectivory was the next diet type to evolve.
Insects were among the earliest terrestrial herbivore
Herbivore
Herbivores are organisms that are anatomically and physiologically adapted to eat plant-based foods. Herbivory is a form of consumption in which an organism principally eats autotrophs such as plants, algae and photosynthesizing bacteria. More generally, organisms that feed on autotrophs in...
s and acted as major selection agents on plants. Plants evolved chemical defenses against this herbivory
Plant defense against herbivory
Plant defense against herbivory or host-plant resistance describes a range of adaptations evolved by plants which improve their survival and reproduction by reducing the impact of herbivores. Plants use several strategies to defend against damage caused by herbivores...
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. This successful evolutionary pattern has also been utilized by mimic
Mimic
In evolutionary biology, mimicry is the similarity of one species to another which protects one or both. This similarity can be in appearance, behaviour, sound, scent and even location, with the mimics found in similar places to their models....
s. Over time, this has led to complex groups of coevolved species. Conversely, some interactions between plants and insects, like pollination
Pollination
Pollination is the process by which pollen is transferred in plants, thereby enabling fertilisation and sexual reproduction. Pollen grains transport the male gametes to where the female gamete are contained within the carpel; in gymnosperms the pollen is directly applied to the ovule itself...
, are beneficial to both organisms. Coevolution has led to the development of very specific mutualisms in such systems.
Taxonomy
{| align="right" style="background:Transparent; border:solid 0px #503DF9;"! colspan="4" style="background:#E6D09D"| Classification
|-
|
Insects (from Latin
Latin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...
, a calque
Calque
In linguistics, a calque or loan translation is a word or phrase borrowed from another language by literal, word-for-word or root-for-root translation.-Calque:...
of Greek
Ancient Greek
Ancient Greek is the stage of the Greek language in the periods spanning the times c. 9th–6th centuries BC, , c. 5th–4th centuries BC , and the c. 3rd century BC – 6th century AD of ancient Greece and the ancient world; being predated in the 2nd millennium BC by Mycenaean Greek...
[], "cut into sections") are a class
Class (biology)
In biological classification, class is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, order, family, genus, and species, with class fitting between phylum and order...
of living creatures within the arthropod
Arthropod
An arthropod is an invertebrate animal having an exoskeleton , a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda , and include the insects, arachnids, crustaceans, and others...
s that have a chitin
Chitin
Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world...
ous exoskeleton
Exoskeleton
An exoskeleton is the external skeleton that supports and protects an animal's body, in contrast to the internal skeleton of, for example, a human. In popular usage, some of the larger kinds of exoskeletons are known as "shells". Examples of exoskeleton animals include insects such as grasshoppers...
, a three-part body (head
Head
In anatomy, the head of an animal is the rostral part that usually comprises the brain, eyes, ears, nose and mouth . Some very simple animals may not have a head, but many bilaterally symmetric forms do....
, thorax
Thorax (insect anatomy)
The thorax is the mid section of the insect body. It holds the head, legs, wings and abdomen. It is also called mesosoma in other arthropods....
, and abdomen
Abdomen
In vertebrates such as mammals the abdomen constitutes the part of the body between the thorax and pelvis. The region enclosed by the abdomen is termed the abdominal cavity...
), three pairs of jointed leg
Arthropod leg
The arthropod leg is a form of jointed appendage of arthropods, usually used for walking. Many of the terms used for arthropod leg segments are of Latin origin, and may be confused with terms for bones: coxa , trochanter , femur, tibia, tarsus, ischium, metatarsus, carpus, dactylus ,...
s, compound eyes, and two antennae
Antenna (biology)
Antennae in biology have historically been paired appendages used for sensing in arthropods. More recently, the term has also been applied to cilium structures present in most cell types of eukaryotes....
. They are among the most diverse groups of animal
Animal
Animals are a major group of multicellular, eukaryotic organisms of the kingdom Animalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their life. Most animals are motile, meaning they can move spontaneously and...
s on the planet, including more than a million described species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...
and represent more than half of all known living organisms. The number of extant species is estimated at between six and ten million, and potentially represent over 90% of the differing metazoan life forms on Earth. Insects may be found in nearly all environments
Natural environment
The natural environment encompasses all living and non-living things occurring naturally on Earth or some region thereof. It is an environment that encompasses the interaction of all living species....
, although only a small number of species occur in the oceans, a habitat dominated by another arthropod group, the crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...
s.
The life cycles of insects vary but most hatch from eggs
Egg (biology)
An egg is an organic vessel in which an embryo first begins to develop. In most birds, reptiles, insects, molluscs, fish, and monotremes, an egg is the zygote, resulting from fertilization of the ovum, which is expelled from the body and permitted to develop outside the body until the developing...
. Insect growth is constrained by the inelastic exoskeleton
Exoskeleton
An exoskeleton is the external skeleton that supports and protects an animal's body, in contrast to the internal skeleton of, for example, a human. In popular usage, some of the larger kinds of exoskeletons are known as "shells". Examples of exoskeleton animals include insects such as grasshoppers...
and development involves a series of molts. The immature stages can differ from the adults in structure, habit and habitat and can include a passive pupa
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...
l stage in those groups that undergo complete metamorphosis. Insects that undergo incomplete metamorphosis
Hemimetabolism
Hemimetabolism or hemimetaboly, also called incomplete metamorphosis, is a term used to describe the mode of development of certain insects that includes three distinct stages: the egg, nymph, and the adult stage, or imago. These groups go through gradual changes; there is no pupal stage...
lack a pupal stage and adults develop through a series of nymph
Nymph (biology)
In biology, a nymph is the immature form of some invertebrates, particularly insects, which undergoes gradual metamorphosis before reaching its adult stage. Unlike a typical larva, a nymph's overall form already resembles that of the adult. In addition, while a nymph moults it never enters a...
al stages. The higher level relationship of the hexapoda
Hexapoda
The subphylum Hexapoda constitutes the largest grouping of arthropods and includes the insects as well as three much smaller groups of wingless arthropods: Collembola, Protura, and Diplura . The Collembola are very abundant in terrestrial environments...
is unclear. Fossilized insects of enormous size have been found from the Paleozoic Era
Paleozoic
The Paleozoic era is the earliest of three geologic eras of the Phanerozoic eon, spanning from roughly...
, including giant dragonflies with wingspans of 55 to 70 cm (22–28 in). The most diverse insect groups appear to have coevolved with flowering plant
Plant
Plants are living organisms belonging to the kingdom Plantae. Precise definitions of the kingdom vary, but as the term is used here, plants include familiar organisms such as trees, flowers, herbs, bushes, grasses, vines, ferns, mosses, and green algae. The group is also called green plants or...
s.
Insects typically move about by walking, flying or occasionally swimming. As it allows for rapid yet stable movement, many insects adopt a tripedal gait in which they walk with their legs touching the ground in alternating triangles. Insects are the only invertebrates to have evolved flight. Many insects spend at least part of their life underwater, with larval adaptations that include gill
Gill
A gill is a respiratory organ found in many aquatic organisms that extracts dissolved oxygen from water, afterward excreting carbon dioxide. The gills of some species such as hermit crabs have adapted to allow respiration on land provided they are kept moist...
s and some adult insects are aquatic and have adaptations for swimming. Some species, like water striders, are capable of walking on the surface of water. Insects are mostly solitary, but some insects, such as certain bee
Bee
Bees are flying insects closely related to wasps and ants, and are known for their role in pollination and for producing honey and beeswax. Bees are a monophyletic lineage within the superfamily Apoidea, presently classified by the unranked taxon name Anthophila...
s, ant
Ant
Ants are social insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than...
s, and termite
Termite
Termites are a group of eusocial insects that, until recently, were classified at the taxonomic rank of order Isoptera , but are now accepted as the epifamily Termitoidae, of the cockroach order Blattodea...
s are social and live in large, well-organized colonies. Some insects, like earwig
Earwig
Earwigs make up the insect order Dermaptera, found throughout the Americas, Africa, Eurasia, Australia and New Zealand. With 1,800 species in 12 families, they are one of the smaller insect orders...
s, show maternal care, guarding their eggs and young. Insects can communicate with each other in a variety of ways. Male moth
Moth
A moth is an insect closely related to the butterfly, both being of the order Lepidoptera. Moths form the majority of this order; there are thought to be 150,000 to 250,000 different species of moth , with thousands of species yet to be described...
s can sense the pheromone
Pheromone
A pheromone is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting outside the body of the secreting individual to impact the behavior of the receiving individual...
s of female moths over distances of many kilometers. Other species communicate with sounds: crickets
Cricket (insect)
Crickets, family Gryllidae , are insects somewhat related to grasshoppers, and more closely related to katydids or bush crickets . They have somewhat flattened bodies and long antennae. There are about 900 species of crickets...
stridulate
Stridulation
Stridulation is the act of producing sound by rubbing together certain body parts. This behavior is mostly associated with insects, but other animals are known to do this as well, such as a number of species of fishes, snakes and spiders...
, or rub their wings together, to attract a mate and repel other males. Lampyridae
Firefly
Lampyridae is a family of insects in the beetle order Coleoptera. They are winged beetles, and commonly called fireflies or lightning bugs for their conspicuous crepuscular use of bioluminescence to attract mates or prey. Fireflies produce a "cold light", with no infrared or ultraviolet frequencies...
in the beetle order Coleoptera
Beetle
Coleoptera is an order of insects commonly called beetles. The word "coleoptera" is from the Greek , koleos, "sheath"; and , pteron, "wing", thus "sheathed wing". Coleoptera contains more species than any other order, constituting almost 25% of all known life-forms...
communicate with light.
Humans regard certain insects as pests and attempt to control them using insecticide
Insecticide
An insecticide is a pesticide used against insects. They include ovicides and larvicides used against the eggs and larvae of insects respectively. Insecticides are used in agriculture, medicine, industry and the household. The use of insecticides is believed to be one of the major factors behind...
s and a host of other techniques. Some insects damage crops by feeding on sap, leaves or fruit
Fruit
In broad terms, a fruit is a structure of a plant that contains its seeds.The term has different meanings dependent on context. In non-technical usage, such as food preparation, fruit normally means the fleshy seed-associated structures of certain plants that are sweet and edible in the raw state,...
s, a few bite humans and livestock, alive and dead, to feed on blood and some are capable of transmitting disease
Disease
A disease is an abnormal condition affecting the body of an organism. It is often construed to be a medical condition associated with specific symptoms and signs. It may be caused by external factors, such as infectious disease, or it may be caused by internal dysfunctions, such as autoimmune...
s to humans, pets and livestock. Nevertheless, without insects to pollinate flowers, the human race would soon run out of food because many of the crop plants that we rely on would not be able to reproduce. Many other insects are considered ecologically beneficial as predators and a few provide direct economic benefit. Silkworms and bees have been used extensively by humans for the production of silk
Silk
Silk is a natural protein fiber, some forms of which can be woven into textiles. The best-known type of silk is obtained from the cocoons of the larvae of the mulberry silkworm Bombyx mori reared in captivity...
and honey
Honey
Honey is a sweet food made by bees using nectar from flowers. The variety produced by honey bees is the one most commonly referred to and is the type of honey collected by beekeepers and consumed by humans...
, respectively.
Etymology
The word insect dates back to 1600, from the Latin word meaning "with a notched or divided body," literally "cut into," from neuter plural of "to cut into, to cut up," from in- "into" + secare "to cut"; from the fact that insects are "cut into" three sections. The word created by Pliny the ElderPliny the Elder
Gaius Plinius Secundus , better known as Pliny the Elder, was a Roman author, naturalist, and natural philosopher, as well as naval and army commander of the early Roman Empire, and personal friend of the emperor Vespasian...
's loan-translation of the Greek
Greek language
Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...
word or "insect" (as in entomology
Entomology
Entomology is the scientific study of insects, a branch of arthropodology...
), which was Aristotle's term for this class of life, also in reference to their "notched" bodies, first documented in English in 1601 in Holland's translation of Pliny. Translations of Aristotle's term also form the usual word for "insect" in Welsh
Welsh language
Welsh is a member of the Brythonic branch of the Celtic languages spoken natively in Wales, by some along the Welsh border in England, and in Y Wladfa...
, Serbo-Croatian
Serbo-Croatian
Serbo-Croatian or Serbo-Croat, less commonly Bosnian/Croatian/Serbian , is a South Slavic language with multiple standards and the primary language of Serbia, Croatia, Bosnia and Herzegovina, and Montenegro...
(zareznik, from rezati "cut"), Russian
Russian language
Russian is a Slavic language used primarily in Russia, Belarus, Uzbekistan, Kazakhstan, Tajikistan and Kyrgyzstan. It is an unofficial but widely spoken language in Ukraine, Moldova, Latvia, Turkmenistan and Estonia and, to a lesser extent, the other countries that were once constituent republics...
( nasekomoe, from sekat "cut"), etc.
Distribution and diversity
Even though the true dimensions of species diversity remain uncertain, estimates are ranging from 1.4 to 1.8 million species. This probably represents less than 20% of all species on Earth, and with only about 20,000 new species of all organisms being described each year, it seems that most species will not remain described for many years unless there is a rapid increase in species descriptions. About 850,000–1,000,000 of all described species are insects. Of the 30 or so orders of insects, four dominate in terms of numbers of described species, with an estimated 600,000–795,000 species include Coleoptera, DipteraDiptera
Diptera , or true flies, is the order of insects possessing only a single pair of wings on the mesothorax; the metathorax bears a pair of drumstick like structures called the halteres, the remnants of the hind wings. It is a large order, containing an estimated 240,000 species, although under half...
, Hymenoptera
Hymenoptera
Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...
, and Lepidoptera
Lepidoptera
Lepidoptera is a large order of insects that includes moths and butterflies . It is one of the most widespread and widely recognizable insect orders in the world, encompassing moths and the three superfamilies of butterflies, skipper butterflies, and moth-butterflies...
. There are almost as many named species of beetle as there are of all other insects added together, or all other non-insects (plant
Plant
Plants are living organisms belonging to the kingdom Plantae. Precise definitions of the kingdom vary, but as the term is used here, plants include familiar organisms such as trees, flowers, herbs, bushes, grasses, vines, ferns, mosses, and green algae. The group is also called green plants or...
s and animal
Animal
Animals are a major group of multicellular, eukaryotic organisms of the kingdom Animalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their life. Most animals are motile, meaning they can move spontaneously and...
s).
{| class="wikitable" style="text-align: center; width:80%;"
|+ Comparison of the Estimated Number of Species for Vertebrates and the Four Most Species Order
|-
|
! scope="col" | Described Species
! scope="col" | Average description rate (species per year)
! scope="col" | Publication effort
|-
! scope="row" | Coleoptera
| 300,000–400,000
| 2308
| 0.01
|-
! scope="row" | Lepidoptera
| 110,000–120,000
| 642
| 0.03
|-
! scope="row" | Diptera
| 90,000–150,000
| 1048
| 0.04
|-
! scope="row" | Hymenoptera
| 100,000–125,000
| 1196
| 0.02
|-
|}
External
Insects have segmentSegmentation (biology)
Segmentation in biology refers to either a type of gastrointestinal motility or the division of some animal and plant body plans into a series of repetitive segments. This article will focus on the segmentation of animal body plans, specifically using the examples of the phyla Arthropoda,...
ed bodies supported by an exoskeleton
Exoskeleton
An exoskeleton is the external skeleton that supports and protects an animal's body, in contrast to the internal skeleton of, for example, a human. In popular usage, some of the larger kinds of exoskeletons are known as "shells". Examples of exoskeleton animals include insects such as grasshoppers...
, a hard outer covering made mostly of chitin
Chitin
Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world...
. The segments of the body are organized into three distinctive but interconnected units, or tagmata
Tagma (biology)
In invertebrate biology, a tagma is a specialized grouping of arthropod segments, such as the head, the thorax, and the abdomen with a common function. The segments of a tagma may be either fused or moveable.-Tagmata:...
: a head, a thorax
Thorax (insect anatomy)
The thorax is the mid section of the insect body. It holds the head, legs, wings and abdomen. It is also called mesosoma in other arthropods....
, and an abdomen
Abdomen
In vertebrates such as mammals the abdomen constitutes the part of the body between the thorax and pelvis. The region enclosed by the abdomen is termed the abdominal cavity...
. The head supports a pair of sensory antennae
Antenna (biology)
Antennae in biology have historically been paired appendages used for sensing in arthropods. More recently, the term has also been applied to cilium structures present in most cell types of eukaryotes....
, a pair of compound eyes, and, if present, one to three simple eyes (or ocelli) and three sets of variously modified appendages that form the mouthpart
Insect mouthparts
Insects exhibit a range of mouthparts, adapted to particular modes of feeding. The earliest insects had chewing mouthparts...
s. The thorax has six segmented leg
Arthropod leg
The arthropod leg is a form of jointed appendage of arthropods, usually used for walking. Many of the terms used for arthropod leg segments are of Latin origin, and may be confused with terms for bones: coxa , trochanter , femur, tibia, tarsus, ischium, metatarsus, carpus, dactylus ,...
s—one pair each for the prothorax, mesothorax and the metathorax segments making up the thorax—and, if present in the species, two or four wing
Insect wing
Insects are the only group of invertebrates known to have evolved flight. Insects possess some remarkable flight characteristics and abilities, still far superior to attempts by humans to replicate their capabilities. Even our understanding of the aerodynamics of flexible, flapping wings and how...
s. The abdomen consists of eleven segments, though in a few species of insects these segments may be fused together or reduced in size. The abdomen also contains most of the digestive
Digestion
Digestion is the mechanical and chemical breakdown of food into smaller components that are more easily absorbed into a blood stream, for instance. Digestion is a form of catabolism: a breakdown of large food molecules to smaller ones....
, respiratory
Respiration (physiology)
'In physiology, respiration is defined as the transport of oxygen from the outside air to the cells within tissues, and the transport of carbon dioxide in the opposite direction...
, excretory
Excretion
Excretion is the process by which waste products of metabolism and other non-useful materials are eliminated from an organism. This is primarily carried out by the lungs, kidneys and skin. This is in contrast with secretion, where the substance may have specific tasks after leaving the cell...
and reproductive internal structures. There is considerable variation and many adaptations in the body parts of insects especially wings, legs, antenna, mouth-parts etc.
Segmentation
The headHead
In anatomy, the head of an animal is the rostral part that usually comprises the brain, eyes, ears, nose and mouth . Some very simple animals may not have a head, but many bilaterally symmetric forms do....
: is enclosed in a hard, heavily sclerotized, unsegmented, exoskeletal head capsule, or epicranium
Epicranium
The Epicranium is the medical term for the collection of structures covering the cranium. It consists of the muscles, aponeurosis, and skin....
, which contains most of the sensing organs, including the antennae, ocellus or eyes, and the mouthparts. Out of all the insect orders, Orthoptera displays the most features found in other insects, including the sutures and sclerite
Sclerite
A sclerite is a hardened body part. The term is used in various branches of biology for various structures including hardened portions of sponges, but it is most commonly used for the hardened portions of arthropod exoskeletons....
s. here, the vertex
Vertex (anatomy)
In arthropod and vertebrate anatomy, the vertex refers to the upper surface of the head.In humans, the vertex is formed by four bones of the skull: the frontal bone, the two parietal bones, and the occipital bone...
, or the apex (dorsal region), is situated between the compound eyes for insects with a hypognathous and opisthognathous head. In prognathous insects, the vertex is not found between the compound eyes, but rather, where the ocelli
Ocellus
A simple eye refers to a type of eye design or optical arrangement that contains a single lens which detect light. A "simple eye" is so-called in distinction from a multi-lensed "compound eye", and is not necessarily at all simple in the usual sense of the word...
are normally. This is because the primary axis of the head is rotated 90 degrees to become parallel to the primary axis of the body. In some species this region is modified and assumes a different name.
The thorax
Thorax
The thorax is a division of an animal's body that lies between the head and the abdomen.-In tetrapods:...
: is a segment composed of three sections, the prothorax
Prothorax
The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites are the pronotum , the prosternum , and the propleuron on each side. The prothorax never bears wings in extant insects, though some fossil groups possessed...
, mesothorax
Mesothorax
The mesothorax is the middle of the three segments in the thorax of an insect, and bears the second pair of legs. Its principal sclerites are the mesonotum , the mesosternum , and the mesopleuron on each side...
, and the metathorax
Metathorax
The metathorax is the posterior of the three segments in the thorax of an insect, and bears the third pair of legs. Its principal sclerites are the metanotum , the metasternum , and the metapleuron on each side...
. The anterior segment, closest to the head, is the prothorax, with the major features being the first pair of legs and the pronotum. The middle segment is the mesothorax with the major features being the second pair of legs and the anterior wings. The third and most posterior segment, abutting the abdomen, is the metathorax, which features the third pair of legs and the posterior wings. Each segment is dilineated by an intersegmental suture. Each segment has four basic regions. The dorsal surface is called the tergum (or notum) to distinguish them from the abdominal terga. The two lateral regions are called the pleura (singular: pleuron) and the ventral aspect is called the sternum. In turn, the notum of the prothorax is called the pronotum, the notum for the mesothorax is called the mesonotum and the notum for the metathorax is called the metanotum. Continuing with this logic, there is also the mesopleura and metapleura as well as the mesosternum and metasternum.
The abdomen
Abdomen
In vertebrates such as mammals the abdomen constitutes the part of the body between the thorax and pelvis. The region enclosed by the abdomen is termed the abdominal cavity...
: is a the last segment of the insect, which typically consists of 11–12 segments and is less strongly sclerotized than the head or thorax. Each segment of the abdomen is represented by a sclerotized tergum, sternum, and perhaps a pleurite. Terga are separated from each other and from the adjacent sterna or pleura by a membrane. Spiracles are located in the pleural area. Variation of this ground plan includes the fusion of terga or terga and sterna to form continuous dorsal or ventral shields or a conical tube. Some insects bear a sclerite in the pleural area called a laterotergite. Ventral sclerites are sometimes called laterosternites. During the embryonic stage of many insects and the postembryonic stage of primitive insects, 11 abdominal segments are present. In modern insects there is a tendency toward reduction in the number of the abdominal segments, but the primitive number of 11 is maintained during embryogenesis.Variation in abdominal segment number is considerable. If the Apterygota are considered to be indicative of the ground plan for pterygotes, confusion reigns: adult Protura have 12 segments, Collembola have 6. The orthopteran family Acrididae has 11 segments, and a fossil specimen of Zoraptera has a 10-segmented abdomen.
Exoskeleton
Insect outer skeleton, the cuticle, is made up of two layers: the epicuticle, which is a thin and waxy water resistant outer layer and contains no chitinChitin
Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world...
, and a lower layer called the procuticle. The procuticle is chitinous and much thicker than the epicuticle and has two layers: an outer layer known as the exocuticle and an inner layer known as the endocuticle. The tough and flexible endocuticle is built from numerous layers of fibrous chitin and proteins, criss-crossing each others in a sandwich pattern, while the exocuticle is rigid and hardened. The exocuticle is greatly reduced in many soft-bodied insects (e.g., caterpillar
Caterpillar
Caterpillars are the larval form of members of the order Lepidoptera . They are mostly herbivorous in food habit, although some species are insectivorous. Caterpillars are voracious feeders and many of them are considered to be pests in agriculture...
s), especially during their larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...
l stages.
Insects are the only invertebrate
Invertebrate
An invertebrate is an animal without a backbone. The group includes 97% of all animal species – all animals except those in the chordate subphylum Vertebrata .Invertebrates form a paraphyletic group...
s to have developed active flight capability, and this has played an important role in their success. Their muscles are able to contract multiple times for each single nerve impulse, allowing the wings to beat faster than would ordinarily be possible. Having their muscles attached to their exoskeletons is more efficient and allows more muscle connections; crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...
s also use the same method, though all spider
Spider
Spiders are air-breathing arthropods that have eight legs, and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other groups of organisms...
s use hydraulic pressure to extend their legs, a system inherited from their pre-arthropod ancestors. Unlike insects, though, most aquatic crustaceans are biomineralized with calcium carbonate
Calcium carbonate
Calcium carbonate is a chemical compound with the formula CaCO3. It is a common substance found in rocks in all parts of the world, and is the main component of shells of marine organisms, snails, coal balls, pearls, and eggshells. Calcium carbonate is the active ingredient in agricultural lime,...
extracted from the water.
Nervous system
The nervous systemNervous system
The nervous system is an organ system containing a network of specialized cells called neurons that coordinate the actions of an animal and transmit signals between different parts of its body. In most animals the nervous system consists of two parts, central and peripheral. The central nervous...
of an insect can be divided into a brain and a ventral nerve cord
Ventral nerve cord
The ventral nerve cord makes up the nervous system of some phyla of the invertebrates, particularly within the nematodes, annelids and the arthropods. It usually consists of cerebral ganglia anteriorly with the nerve cords running down the ventral plane of the organism...
. The head capsule is made up of six fused segments, each with a pair of ganglia
Ganglion
In anatomy, a ganglion is a biological tissue mass, most commonly a mass of nerve cell bodies. Cells found in a ganglion are called ganglion cells, though this term is also sometimes used to refer specifically to retinal ganglion cells....
, 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
Esophagus
The esophagus is an organ in vertebrates which consists of a muscular tube through which food passes from the pharynx to the stomach. During swallowing, food passes from the mouth through the pharynx into the esophagus and travels via peristalsis to the stomach...
, called the subesophageal ganglion
Subesophageal ganglion
The subesophageal ganglion of insects is composed of three pairs of fused ganglia. It controls the mouthparts, the salivary glands and certain muscles....
.
The thoracic segments have one ganglion on each side, which are connected into a pair, one pair per segment. This arrangement is also seen in the abdomen but only in the first eight segments. Many species of insects have reduced numbers of ganglia due to fusion or reduction. Some cockroaches have just six ganglia in the abdomen, whereas the wasp Vespa crabro has only two in the thorax and three in the abdomen. Some insects, like the house fly Musca domestica
Housefly
The housefly , Musca domestica, is a fly of the suborder Cyclorrhapha...
, have all the body ganglia fused into a single large thoracic ganglion.
At least a few insects have nociceptor
Nociceptor
A nociceptor is a sensory receptor that responds to potentially damaging stimuli by sending nerve signals to the spinal cord and brain. This process, called nociception, usually causes the perception of pain.-History:...
s, cells that detect and transmit sensations of pain
Pain in animals
Pain is a sensory and emotional experience often caused by intense or damaging stimuli. The International Association for the Study of Pain says pain is a conscious experience involving unpleasantness, i.e. suffering...
. This was discovered in 2003 by studying the variation in reactions of larvae
Larvae
In Roman mythology, lemures were shades or spirits of the restless or malignant dead, and are probably cognate with an extended sense of larvae as disturbing or frightening...
of the common fruitfly Drosophila
Drosophila
Drosophila is a genus of small flies, belonging to the family Drosophilidae, whose members are often called "fruit flies" or more appropriately pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many species to linger around overripe or rotting fruit...
to the touch of a heated probe and an unheated one. The larvae reacted to the touch of the heated probe with a stereotypical rolling behavior that was not exhibited when the larvae were touched by the unheated probe. Although nociception has been demonstrated in insects, there is not a consensus that insects feel pain consciously but see Pain in invertebrates
Pain in invertebrates
Pain in invertebrates is a problematic issue. Although there are numerous definitions of pain, almost all involve two key components. First, nociception is required. This is the ability to detect noxious stimuli which evokes a reflex response that moves the entire animal, or the affected part of...
.
Digestive system
An insect uses its digestive system to extract nutrients and other substances from the food it consumes. Most of this food is ingested in the form of macromoleculeMacromolecule
A macromolecule is a very large molecule commonly created by some form of polymerization. In biochemistry, the term is applied to the four conventional biopolymers , as well as non-polymeric molecules with large molecular mass such as macrocycles...
s and other complex substances like protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...
s, polysaccharide
Polysaccharide
Polysaccharides are long carbohydrate molecules, of repeated monomer units joined together by glycosidic bonds. They range in structure from linear to highly branched. Polysaccharides are often quite heterogeneous, containing slight modifications of the repeating unit. Depending on the structure,...
s, fat
Fat
Fats consist of a wide group of compounds that are generally soluble in organic solvents and generally insoluble in water. Chemically, fats are triglycerides, triesters of glycerol and any of several fatty acids. Fats may be either solid or liquid at room temperature, depending on their structure...
s, and nucleic acid
Nucleic acid
Nucleic acids are biological molecules essential for life, and include DNA and RNA . Together with proteins, nucleic acids make up the most important macromolecules; each is found in abundance in all living things, where they function in encoding, transmitting and expressing genetic information...
s. These macromolecules must be broken down by catabolic reaction
Catabolism
Catabolism is the set of metabolic pathways that break down molecules into smaller units and release energy. In catabolism, large molecules such as polysaccharides, lipids, nucleic acids and proteins are broken down into smaller units such as monosaccharides, fatty acids, nucleotides, and amino...
s into smaller molecules like amino acid
Amino acid
Amino acids are molecules containing an amine group, a carboxylic acid group and a side-chain that varies between different amino acids. The key elements of an amino acid are carbon, hydrogen, oxygen, and nitrogen...
s and simple sugars before being used by cells of the body for energy, growth, or reproduction. This break-down process is known as digestion.
The main structure of an insect's digestive system is a long enclosed tube called the alimentary canal, which runs lengthwise through the body. The alimentary canal directs food unidirectionally from the mouth
Mouth
The mouth is the first portion of the alimentary canal that receives food andsaliva. The oral mucosa is the mucous membrane epithelium lining the inside of the mouth....
to the anus
Anus
The anus is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, unwanted semi-solid matter produced during digestion, which, depending on the type of animal, may be one or more of: matter which the animal cannot digest,...
. It has three sections, each of which performs a different process of digestion. In addition to the alimentary canal, insects also have paired salivary glands and salivary reservoirs. These structures usually reside in the thorax, adjacent to the foregut.
The salivary gland
Salivary gland
The salivary glands in mammals are exocrine glands, glands with ducts, that produce saliva. They also secrete amylase, an enzyme that breaks down starch into maltose...
s (element 30 in numbered diagram) in an insect's mouth produce saliva. The salivary ducts lead from the glands to the reservoirs and then forward through the head to an opening called the salivarium, located behind the hypopharynx. By moving its mouthparts (element 32 in numbered diagram) the insect can mix its food with saliva. The mixture of saliva and food then travels through the salivary tubes into the mouth, where it begins to break down. Some insects, like flies
Diptera
Diptera , or true flies, is the order of insects possessing only a single pair of wings on the mesothorax; the metathorax bears a pair of drumstick like structures called the halteres, the remnants of the hind wings. It is a large order, containing an estimated 240,000 species, although under half...
, have extra-oral digestion
Mastication
Mastication or chewing is the process by which food is crushed and ground by teeth. It is the first step of digestion and it increases the surface area of foods to allow more efficient break down by enzymes. During the mastication process, the food is positioned between the teeth for grinding by...
. Insects using extra-oral digestion expel digestive enzymes onto their food to break it down. This strategy allows insects to extract a significant proportion of the available nutrients from the food source. The gut is where almost all of insects' digestion takes place. It can be divided into the foregut
Foregut
The foregut is the anterior part of the alimentary canal, from the mouth to the duodenum at the entrance of the bile duct. At this point it is continuous with the midgut...
, midgut
Stomach
The stomach is a muscular, hollow, dilated part of the alimentary canal which functions as an important organ of the digestive tract in some animals, including vertebrates, echinoderms, insects , and molluscs. It is involved in the second phase of digestion, following mastication .The stomach is...
and hindgut
Hindgut
The hindgut is the posterior part of the alimentary canal. In mammals, it includes the distal third of the transverse colon and the splenic flexure, the descending colon, sigmoid colon and rectum.-Blood flow:...
.
Foregut
The first section of the alimentary canal is the foregut
Foregut
The foregut is the anterior part of the alimentary canal, from the mouth to the duodenum at the entrance of the bile duct. At this point it is continuous with the midgut...
(element 27 in numbered diagram), or stomodaeum. The foregut is lined with a cuticular lining made of chitin
Chitin
Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world...
and protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...
s as protection from tough food. The foregut includes the buccal cavity (mouth), pharynx
Pharynx
The human pharynx is the part of the throat situated immediately posterior to the mouth and nasal cavity, and anterior to the esophagus and larynx. The human pharynx is conventionally divided into three sections: the nasopharynx , the oropharynx , and the laryngopharynx...
, esophagus
Esophagus
The esophagus is an organ in vertebrates which consists of a muscular tube through which food passes from the pharynx to the stomach. During swallowing, food passes from the mouth through the pharynx into the esophagus and travels via peristalsis to the stomach...
, and Crop
Crop (anatomy)
A crop is a thin-walled expanded portion of the alimentary tract used for the storage of food prior to digestion that is found in many animals, including gastropods, earthworms, leeches, insects, birds, and even some dinosaurs.- Bees :Cropping is used by bees to temporarily store nectar of flowers...
and proventriculus
Proventriculus
The proventriculus is part of the digestive system of birds, invertebrates and insects.-Birds:The proventriculus is a standard part of avian anatomy...
(any part may be highly modified) which both store food and signify when to continue passing onward to the midgut.
Digestion starts in buccal cavity (mouth) as partially chewed food is broken down by saliva from the salivary glands. As the salivary glands produce fluid and carbohydrate-digesting enzyme
Enzyme
Enzymes are proteins that catalyze chemical reactions. In enzymatic reactions, the molecules at the beginning of the process, called substrates, are converted into different molecules, called products. Almost all chemical reactions in a biological cell need enzymes in order to occur at rates...
s (mostly amylase
Amylase
Amylase is an enzyme that catalyses the breakdown of starch into sugars. Amylase is present in human saliva, where it begins the chemical process of digestion. Food that contains much starch but little sugar, such as rice and potato, taste slightly sweet as they are chewed because amylase turns...
s), strong muscles in the pharynx pump fluid into the buccal cavity, lubricating the food like the salivarium does, and helping blood feeders, and xylem and phloem feeders.
From there, the pharynx passes food to the esophagus, which could be just a simple tube passing it on to the crop and proventriculus, and then onward to the midgut, as in most insects. Alternately, the foregut may expand into a very enlarged crop and proventriculus, or the crop could just be a diverticulum
Diverticulum
A diverticulum is medical or biological term for an outpouching of a hollow structure in the body. Depending upon which layers of the structure are involved, they are described as being either true or false....
, or fluid filled structure, as in some Diptera species.
Midgut
Once food leaves the crop, it passes to the midgut
Stomach
The stomach is a muscular, hollow, dilated part of the alimentary canal which functions as an important organ of the digestive tract in some animals, including vertebrates, echinoderms, insects , and molluscs. It is involved in the second phase of digestion, following mastication .The stomach is...
(element 13 in numbered diagram), also known as the mesenteron, where the majority of digestion takes place. Microscopic projections from the midgut wall, called microvilli
Microvillus
Microvilli are microscopic cellular membrane protrusions that increase the surface area of cells, and are involved in a wide variety of functions, including absorption, secretion, cellular adhesion, and mechanotransduction....
, increase the surface area of the wall and allow more nutrients to be absorbed; they tend to be close to the origin of the midgut. In some insects, the role of the microvilli and where they are located may vary. For example, specialized microvilli producing digestive enzymes may more likely be near the end of the midgut, and absorption near the origin or beginning of the midgut.
Hindgut
In the hindgut
Hindgut
The hindgut is the posterior part of the alimentary canal. In mammals, it includes the distal third of the transverse colon and the splenic flexure, the descending colon, sigmoid colon and rectum.-Blood flow:...
(element 16 in numbered diagram), or proctodaeum, undigested food particles are joined by uric acid
Uric acid
Uric acid is a heterocyclic compound of carbon, nitrogen, oxygen, and hydrogen with the formula C5H4N4O3. It forms ions and salts known as urates and acid urates such as ammonium acid urate. Uric acid is created when the body breaks down purine nucleotides. High blood concentrations of uric acid...
to form fecal pellets. The rectum absorbs 90% of the water in these fecal pellets, and the dry pellet is then eliminated through the anus (element 17), completing the process of digestion. The uric acid is formed using hemolymph waste products diffused from the Malpighian tubule
Malpighian tubule system
The Malpighian tubule system is a type of excretory and osmoregulatory system found in some Uniramia , arachnids and tardigrades....
s (element 20). It is then emptied directly into the alimentary canal, at the junction between the midgut and hindgut. The number of Malpighian tubules possessed by a given insect varies between species, ranging from only two tubules in some insects to over 100 tubules in others.
Endocrine system
The salivary glandSalivary gland
The salivary glands in mammals are exocrine glands, glands with ducts, that produce saliva. They also secrete amylase, an enzyme that breaks down starch into maltose...
s (element 30 in numbered diagram) in an insect's mouth produce saliva. The salivary ducts lead from the glands to the reservoirs and then forward through the head to an opening called the salivarium, located behind the hypopharynx. By moving its mouthparts (element 32 in numbered diagram) the insect can mix its food with saliva. The mixture of saliva and food then travels through the salivary tubes into the mouth, where it begins to break down. Some insects, like flies
Diptera
Diptera , or true flies, is the order of insects possessing only a single pair of wings on the mesothorax; the metathorax bears a pair of drumstick like structures called the halteres, the remnants of the hind wings. It is a large order, containing an estimated 240,000 species, although under half...
, have extra-oral digestion
Mastication
Mastication or chewing is the process by which food is crushed and ground by teeth. It is the first step of digestion and it increases the surface area of foods to allow more efficient break down by enzymes. During the mastication process, the food is positioned between the teeth for grinding by...
. Insects using extra-oral digestion expel digestive enzymes onto their food to break it down. This strategy allows insects to extract a significant proportion of the available nutrients from the food source.
Reproductive system
The reproductive system of female insects consist of a pair of ovaries, accessory glands, one or more spermathecaSpermatheca
The spermatheca , also called receptaculum seminis , is an organ of the female reproductive tract in insects, some molluscs, oligochaeta worms and certain other invertebrates and vertebrates...
e, and ducts connecting these parts. The ovaries are made up of a number of egg tubes, called ovarioles
Ovarioles
An ovariole is one of the tubes of which the ovaries of most insects are composed. Typically an insect will have two ovaries. The constituent ovarioles lead to two oviducts, which converge into a single oviduct. The ovarioles are composed of a germarium and a set of ovarial follicles.-In D...
, which vary in size and number by species. The number of eggs that the insect is able to make vary by the number of ovarioles with the rate that eggs can be develop being also influenced by ovariole design. Female insects are able make eggs, receive and store sperm, manipulate sperm from different males, and lay eggs. Accessory glands or glandular parts of the oviducts produce a variety of substances for sperm maintenance, transport, and fertilization, as well as for protection of eggs. They can produce glue and protective substances for coating eggs or tough coverings for a batch of eggs called ootheca
Ootheca
An ootheca is a type of egg mass made by any member of a variety of species .The word is a latinized combination of oo-, meaning "egg", from the Greek word ōon , and theca, meaning a "cover" or "container", from the Greek theke...
e. Spermathecae are tubes or sacs in which sperm can be stored between the time of mating and the time an egg is fertilized.
For males, the reproductive system is the testis, suspended in the body cavity by trachea
Invertebrate trachea
The invertebrate trachea refers to the open respiratory system composed of spiracles, tracheae, and tracheoles that terrestrial arthropods have to transport metabolic gases to and from tissues....
e and the fat body. Most male insects have a pair of testes, inside of which are sperm tubes or follicles that are enclosed within a membranous sac. The follicles connect to the vas deferens by the vas efferens, and the two tubular vasa deferentia connect to a median ejaculatory duct that leads to the outside. A portion of the vas deferens is often enlarged to form the seminal vesicle, which stores the sperm before they are discharged into the female. The seminal vesicles have glandular linings that secrete nutrients for nourishment and maintenance of the sperm. The ejaculatory duct is derived from an invagination of the epidermal cells during development and, as a result, has a cuticular lining. The terminal portion of the ejaculatory duct may be sclerotized to form the intromittent organ, the aedeagus. The remainder of the male reproductive system is derived from embryonic mesoderm, except for the germ cells, or spermatogonia, which descend from the primordial pole cells very early during embryogenesis.
Respiratory and circulatory systems
Insect respiration is accomplished without lungLung
The lung is the essential respiration organ in many air-breathing animals, including most tetrapods, a few fish and a few snails. In mammals and the more complex life forms, the two lungs are located near the backbone on either side of the heart...
s. Instead, the insect respiratory system
Respiratory system of insects
The respiratory system of insects is the means by which insects obtain oxygen from the air.-The Respiratory System:Air enters the respiratory systems of most insects through a series of external openings called spiracles. These external openings, which act as muscular valves in some insects, lead...
uses 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 trachea
Invertebrate trachea
The invertebrate trachea refers to the open respiratory system composed of spiracles, tracheae, and tracheoles that terrestrial arthropods have to transport metabolic gases to and from tissues....
(element 8 in numbered diagram). Since oxygen is delivered directly, the circulatory system is not used to carry oxygen, and is therefore greatly reduced. The insect circulatory system has no vein
Vein
In the circulatory system, veins are blood vessels that carry blood towards the heart. Most veins carry deoxygenated blood from the tissues back to the heart; exceptions are the pulmonary and umbilical veins, both of which carry oxygenated blood to the heart...
s or arterie
Artery
Arteries are blood vessels that carry blood away from the heart. This blood is normally oxygenated, exceptions made for the pulmonary and umbilical arteries....
s, and instead consists of little more than a single, perforated dorsal tube which pulses peristaltically
Peristalsis
Peristalsis is a radially symmetrical contraction and relaxation of muscles which propagates in a wave down the muscular tube, in an anterograde fashion. In humans, peristalsis is found in the contraction of smooth muscles to propel contents through the digestive tract. Earthworms use a similar...
. Toward the thorax, the dorsal tube (element 14) divides into chambers and acts like the insect's heart. The opposite end of the dorsal tube is like the aorta of the insect circulating the hemolymph
Hemolymph
Hemolymph, or haemolymph, is a fluid in the circulatory system of some arthropods and is analogous to the fluids and cells making up both blood and interstitial fluid in vertebrates such as birds and mammals...
, arthropods' fluid analog of blood
Blood
Blood is a specialized bodily fluid in animals that delivers necessary substances such as nutrients and oxygen to the cells and transports metabolic waste products away from those same cells....
, inside the body cavity. Air is taken in through openings on the sides of the abdomen called spiracle
Spiracle
Spiracles are openings on the surface of some animals that usually lead to respiratory systems.-Vertebrates:The spiracle is a small hole behind each eye that opens to the mouth in some fishes. In the primitive jawless fish the first gill opening immediately behind the mouth is essentially similar...
s.
There are many different patterns of gas exchange
Gas exchange
Gas exchange is a process in biology where gases contained in an organism and atmosphere transfer or exchange. In human gas-exchange, gases contained in the blood of human bodies exchange with gases contained in the atmosphere. Human gas-exchange occurs in the lungs...
demonstrated by different groups of insects. Gas exchange patterns in insects can range from continuous and diffusive
Diffusion
Molecular diffusion, often called simply diffusion, is the thermal motion of all particles at temperatures above absolute zero. The rate of this movement is a function of temperature, viscosity of the fluid and the size of the particles...
ventilation, to discontinuous gas exchange
Discontinuous gas exchange
Discontinuous gas-exchange cycles , also called discontinuous ventilation or discontinuous ventilatory cycles, follow one of several patterns of arthropod gas exchange that have been documented primarily in insects; they occur when the insect is at rest...
. During continuous gas exchange, oxygen
Oxygen
Oxygen is the element with atomic number 8 and represented by the symbol O. Its name derives from the Greek roots ὀξύς and -γενής , because at the time of naming, it was mistakenly thought that all acids required oxygen in their composition...
is taken in and carbon dioxide
Carbon dioxide
Carbon dioxide is a naturally occurring chemical compound composed of two oxygen atoms covalently bonded to a single carbon atom...
is released in a continuous cycle. In discontinuous gas exchange, however, the insect takes in oxygen while it is active and small amounts of carbon dioxide are released when the insect is at rest. Diffusive ventilation is simply a form of continuous gas exchange that occurs by diffusion
Diffusion
Molecular diffusion, often called simply diffusion, is the thermal motion of all particles at temperatures above absolute zero. The rate of this movement is a function of temperature, viscosity of the fluid and the size of the particles...
rather than physically taking in the oxygen. Some species of insect that are submerged also have adaptations to aid in respiration. As larvae, many insects have gills that can extract oxygen dissolved in water, while others need to rise to the water surface to replenish air supplies which may be held or trapped in special structures.
Reproduction and development
Egg (biology)
An egg is an organic vessel in which an embryo first begins to develop. In most birds, reptiles, insects, molluscs, fish, and monotremes, an egg is the zygote, resulting from fertilization of the ovum, which is expelled from the body and permitted to develop outside the body until the developing...
s. The fertilization and development takes place inside the egg, enclosed by a shell (chorion
Chorion
The chorion is one of the membranes that exist during pregnancy between the developing fetus and mother. It is formed by extraembryonic mesoderm and the two layers of trophoblast and surrounds the embryo and other membranes...
). Some species of insects, like the cockroach Blaptica dubia
Blaptica dubia
The Dubia Cockroach , also known as Orange-spotted Cockroach or Guyana Spotted Cockroach, is a medium/large species of cockroach, measuring about long. They are sexually dimorphic; the males have wings while the females have only tiny wing stubs...
, as well as juvenile aphids and tsetse flies, are ovoviviparous
Ovoviviparity
Ovoviviparity, ovovivipary, or ovivipary, is a mode of reproduction in animals in which embryos develop inside eggs that are retained within the mother's body until they are ready to hatch...
. The eggs of ovoviviparous animals develop entirely inside the female, and then hatch immediately upon being laid. Some other species, such as those in the genus of cockroaches known as Diploptera
Diploptera
Diploptera is a genus of cockroach, used particularly in molecular research into allatostatins. The species Diploptera punctata, the Pacific beetle cockroach, is the only known viviparous cockroach ....
, are viviparous, and thus gestate inside the mother and are born alive
Vivipary
Vivipary has two different meanings. In animals, it means development of the embryo inside the body of the mother, eventually leading to live birth, as opposed to laying eggs...
. Some insects, like parasitic wasps, show polyembryony
Polyembryony
Polyembryony is the phenomenon of two or more embryos developing from a single fertilized egg . Polyembryony occurs regularly in many plants and animals. The nine banded armadillo, for instance, usually gives birth to four identical young...
, where a single fertilized egg divides into many and in some cases thousands of separate embryos.
Polymorphism (biology)
Polymorphism in biology occurs when two or more clearly different phenotypes exist in the same population of a species — in other words, the occurrence of more than one form or morph...
, paedomorphosis or peramorphosis
Peramorphosis
In developmental biology, peramorphosis is a phylogenetic change in which individuals of a species mature past adulthood and take on hitherto unseen traits...
, sexual dimorphism
Sexual dimorphism
Sexual dimorphism is a phenotypic difference between males and females of the same species. Examples of such differences include differences in morphology, ornamentation, and behavior.-Examples:-Ornamentation / coloration:...
, parthenogenesis
Parthenogenesis
Parthenogenesis is a form of asexual reproduction found in females, where growth and development of embryos occur without fertilization by a male...
and more rarely hermaphroditism. In haplodiploidy, which is a type of sex-determination system
Sex-determination system
A sex-determination system is a biological system that determines the development of sexual characteristics in an organism. Most sexual organisms have two sexes. In many cases, sex determination is genetic: males and females have different alleles or even different genes that specify their sexual...
, the offspring's sex is determined by the number of sets of chromosome
Chromosome
A chromosome is an organized structure of DNA and protein found in cells. It is a single piece of coiled DNA containing many genes, regulatory elements and other nucleotide sequences. Chromosomes also contain DNA-bound proteins, which serve to package the DNA and control its functions.Chromosomes...
s an individual receives. This system is typical in bees and wasps. Polymorphism is the where a species may have different morphs or forms, as in the oblong winged katydid
Amblycorypha oblongifolia
The Oblong-Winged katydid is a species of katydid, or bush-cricket, of the family Tettigoniidae .-Appearance:...
, which has four different varieties: green, pink, and yellow or tan. Some insects may retain phenotype
Phenotype
A phenotype is an organism's observable characteristics or traits: such as its morphology, development, biochemical or physiological properties, behavior, and products of behavior...
s that are normally only seen in juveniles; this is called paedomorphosis. In peramorphosis, an opposite sort of phenomenon, insects take on previously unseen traits after they have matured into adults. Many insects display sexual dimorphism, in which males and females have notably different appearances, such as the moth Orgyia recens
Orgyia recens
The Scarce Vapourer is a species of moth of the Lymantriidae family. It is found in Europe.The wingspan is 35-40 mm for the males, the females are wingless. The moth flies from June to July depending on the location....
as an exemplar of sexual dimorphism in insects.
Some insects use parthenogenesis
Parthenogenesis
Parthenogenesis is a form of asexual reproduction found in females, where growth and development of embryos occur without fertilization by a male...
, a process in which the female can reproduce and give birth without having the eggs fertilized by a male
Male
Male refers to the biological sex of an organism, or part of an organism, which produces small mobile gametes, called spermatozoa. Each spermatozoon can fuse with a larger female gamete or ovum, in the process of fertilization...
. Many aphids undergo a form of parthenogenesis, called cyclical parthenogenesis, in which they alternate between one or many generations of asexual and sexual reproduction. 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 which they spawn males. However, overall, most individuals are female, which are produced by fertilization. The males are haploid and the females are diploid. More rarely, some insects display hermaphroditism, in which a given individual has both male and female reproductive organs.
Insect life-histories show adaptations to withstand cold and dry conditions. Some temperate region insects are capable of activity during winter, while some others migrate to a warmer climate or go into a state of torpor
Torpor
Torpor, sometimes called temporary hibernation is a state of decreased physiological activity in an animal, usually characterized by a reduced body temperature and rate of metabolism. Animals that go through torpor include birds and some mammals such as mice and bats...
. Still other insects have evolved mechanisms of diapause
Diapause
Diapause is the delay in development in response to regularly and recurring periods of adverse environmental conditions. It is considered to be a physiological state of dormancy with very specific initiating and inhibiting conditions...
that allow eggs or pupae to survive these conditions.
Metamorphosis
MetamorphosisMetamorphosis
Metamorphosis is a biological process by which an animal physically develops after birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation...
in insects is the biological process of development all insects must undergo. There are two forms of metamorphosis: incomplete metamorphosis and complete metamorphosis.
Incomplete metamorphosis
Insects that show hemimetabolismHemimetabolism
Hemimetabolism or hemimetaboly, also called incomplete metamorphosis, is a term used to describe the mode of development of certain insects that includes three distinct stages: the egg, nymph, and the adult stage, or imago. These groups go through gradual changes; there is no pupal stage...
, or incomplete metamorphosis, change gradually by undergoing a series of molt
Ecdysis
Ecdysis is the moulting of the cuticula in many invertebrates. This process of moulting is the defining feature of the clade Ecdysozoa, comprising the arthropods, nematodes, velvet worms, horsehair worms, rotifers, tardigrades and Cephalorhyncha...
s. An insect 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. 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, which makes the old cuticle split along predefined weaknesses where the old exocuticle was thinnest. Other arthropods have a much different process and only molt; though must accommodate for the difference in exoskeleton structure and make up with other enzymes.
Immature insects that go through incomplete metamorphosis are called nymph
Nymph (biology)
In biology, a nymph is the immature form of some invertebrates, particularly insects, which undergoes gradual metamorphosis before reaching its adult stage. Unlike a typical larva, a nymph's overall form already resembles that of the adult. In addition, while a nymph moults it never enters a...
s or in the case of dragonflies and damselflies as naiads. Nymphs are similar in form to the adult except for the presence of wings, which are not developed until adulthood. With each molt, nymphs grow larger and become more similar in appearance to adult insects.
Complete metamorphosis
Holometabolism
Holometabolism, also called complete metamorphism, is a term applied to insect groups to describe the specific kind of insect development which includes four life stages - as an embryo or egg, a larva, a pupa and an imago or adult. Holometabolism is a monophyletic trait that all insects in the...
, or complete metamorphosis, is where the insect changes all in four stages, an egg
Egg (biology)
An egg is an organic vessel in which an embryo first begins to develop. In most birds, reptiles, insects, molluscs, fish, and monotremes, an egg is the zygote, resulting from fertilization of the ovum, which is expelled from the body and permitted to develop outside the body until the developing...
or embryo
Embryo
An embryo is a multicellular diploid eukaryote in its earliest stage of development, from the time of first cell division until birth, hatching, or germination...
, a larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...
, a pupa
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...
, and the adult
Adult
An adult is a human being or living organism that is of relatively mature age, typically associated with sexual maturity and the attainment of reproductive age....
or imago
Imago
In biology, the imago is the last stage of development of an insect, after the last ecdysis of an incomplete metamorphosis, or after emergence from the pupa where the metamorphosis is complete...
. In these species, egg hatches to produce a larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...
, which is generally worm-like in form. This worm-like form can be one of several varieties: eruciform (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
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...
, a stage marked by reduced movement and often sealed within a cocoon
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...
. 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. Insects undergo considerable change in form during the pupal stage, and emerge as adults. Butterflies are a well known example of an insects that undergo complete metamorphosis, although most insects use this life cycle. Some insects have evolved this system to hypermetamorphosis
Hypermetamorphosis
Hypermetamorphosis is a kind of complete insect metamorphosis in which the different larval instars represent two or more different forms of larva. As the larva molts its morphology can change from that of a campodeiform to scarabaeiform or to vermiform...
.
Some of the oldest and most successful insect groups, such Endopterygota
Endopterygota
The Endopterygota, also known as Holometabola, are insects of the subclass Pterygota which go through distinctive larval, pupal, and adult stages. They undergo a radical metamorphosis, with the larval and adult stages differing considerably in their structure and behaviour...
, use a system of complete metamorphosis. Strangely though, complete metamorphosis is unique to certain insect orders, like Diptera
Diptera
Diptera , or true flies, is the order of insects possessing only a single pair of wings on the mesothorax; the metathorax bears a pair of drumstick like structures called the halteres, the remnants of the hind wings. It is a large order, containing an estimated 240,000 species, although under half...
, Lepidoptera
Lepidoptera
Lepidoptera is a large order of insects that includes moths and butterflies . It is one of the most widespread and widely recognizable insect orders in the world, encompassing moths and the three superfamilies of butterflies, skipper butterflies, and moth-butterflies...
, and Hymenoptera
Hymenoptera
Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...
, and no other arthropods undergo it, but incomplete metamorphosis.
Senses and communication
Many insects possess very sensitive and/or specialized organs of perceptionSense
Senses are physiological capacities of organisms that provide inputs for perception. The senses and their operation, classification, and theory are overlapping topics studied by a variety of fields, most notably neuroscience, cognitive psychology , and philosophy of perception...
. Some insects such as bees can perceive ultraviolet
Ultraviolet
Ultraviolet light is electromagnetic radiation with a wavelength shorter than that of visible light, but longer than X-rays, in the range 10 nm to 400 nm, and energies from 3 eV to 124 eV...
wavelengths, or detect polarized light, while the antennae
Antenna (biology)
Antennae in biology have historically been paired appendages used for sensing in arthropods. More recently, the term has also been applied to cilium structures present in most cell types of eukaryotes....
of male moths can detect the pheromone
Pheromone
A pheromone is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting outside the body of the secreting individual to impact the behavior of the receiving individual...
s of female moths over distances of many kilometers. There is a pronounced tendency for there to be 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. There are a variety of different mechanisms by which insects perceive sound, while the patterns are not universal, insects can generally hear sound if they can produce it. Different insect species can have varying hearing, though most insects can hear only a narrow range of frequencies related to the frequency of the sounds they can produce. Mosquitoes have been found to hear up to 2 MHz., and some grasshoppers can hear up to 50 MHz. Certain predatory and parasitic insects can detect the characteristic sounds made by their prey or hosts, respectively. For instance, some nocturnal moths can perceive the ultrasonic
Ultrasound
Ultrasound is cyclic sound pressure with a frequency greater than the upper limit of human hearing. Ultrasound is thus not separated from "normal" sound based on differences in physical properties, only the fact that humans cannot hear it. Although this limit varies from person to person, it is...
emissions of bat
Bat
Bats are mammals of the order Chiroptera "hand" and pteron "wing") whose forelimbs form webbed wings, making them the only mammals naturally capable of true and sustained flight. By contrast, other mammals said to fly, such as flying squirrels, gliding possums, and colugos, glide rather than fly,...
s, which helps them avoid predation. Insects that feed on blood have special sensory structures that can detect infrared
Infrared
Infrared light is electromagnetic radiation with a wavelength longer than that of visible light, measured from the nominal edge of visible red light at 0.74 micrometres , and extending conventionally to 300 µm...
emissions, and use them to home in on their hosts.
Some insects display a rudimentary sense of numbers
Numeracy
Numeracy is the ability to reason with numbers and other mathematical concepts. A numerically literate person can manage and respond to the mathematical demands of life...
, such as the solitary wasps that prey upon a single species. The mother wasp lays her eggs in individual cells and provides each egg with a number of live caterpillars on which the young feed when hatched. Some species of wasp always provide five, others twelve, and others as high as twenty-four caterpillars per cell. The number of caterpillars is different among species, but always the same for each sex of larva. The male solitary wasp in the genus Eumenes
Eumenes (genus)
Eumenes is the type genus of the subfamily Eumeninae of Vespidae. It is a large and widespread genus, with over 100 taxa , mostly occurring in the temperate portions of the Northern Hemisphere. Most species are black or brown, and commonly marked with strikingly contrasting patterns of yellow,...
is smaller than the female, so the mother of one species supplies him with only five caterpillars; the larger female receives ten caterpillars in her cell.
Light production and vision
Mycetophilidae
Mycetophilidae is a family of small flies, forming the bulk of those species known as fungus gnats. There are approximately 3000 described species in 150 genera but the true number of species is undoubtedly much higher...
(Diptera), and the beetle families Lampyridae, Phengodidae
Phengodidae
The beetle family Phengodidae is known also as glowworm beetles, whose larvae are known as glowworms. The females and larvae have bioluminescent organs. They occur throughout the New World from extreme southern Canada to Chile...
, Elateridae and Staphylinidae are bioluminescent. The most familiar group are the fireflies
Fireflies
Fireflies is a novel by Shiva Naipaul originally published in 1970. It was his first book, a comic novel set in Trinidad. In an essay in An Unfinished Journey, Naipaul described how in 1968 as a final year student at Oxford University studying Chinese, he had been moved to write down a sentence,...
, 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
Arachnocampa
Arachnocampa is a genus of four fungus gnat species which are, in their larval stage, glowworms. They are found mostly in New Zealand and Australia in caves and grottos, or sheltered places in forests....
(Mycetophilidae, Fungus gnats) glow to lure small flying insects into sticky strands of silk.
Some fireflies of the genus Photuris mimic the flashing of female Photinus
Photinus (beetle)
The rover fireflies are a genus of fireflies . They are the type genus of tribe Photinini in subfamily Lampyrinae. This genus contains, for example, the common eastern firefly , the state insect of Tennessee.Male Photinus beetles emit a flashing light pattern to signal for females...
species to attract males of that species, which are then captured and devoured. The colors of emitted light vary from dull blue (Orfelia fultoni, Mycetophilidae) to the familiar greens and the rare reds (Phrixothrix tiemanni, Phengodidae).
Most insects, except some species of cave crickets
Rhaphidophoridae
The orthopteran family Rhaphidophoridae includes the cave wetas, cave crickets, camelback crickets, camel crickets, spider crickets and sand treaders, of the suborder Ensifera; in some regions, such as Missouri and Virginia, these crickets are referred to as "Cricket Spiders"...
, are able to perceive light and dark. Many species have acute vision capable of detecting minute movements. The eyes include simple eyes or ocelli
Ocellus
A simple eye refers to a type of eye design or optical arrangement that contains a single lens which detect light. A "simple eye" is so-called in distinction from a multi-lensed "compound eye", and is not necessarily at all simple in the usual sense of the word...
as well as compound eyes of varying sizes. Many species are able to detect light in the infrared, ultraviolet and the visible light wavelengths. Color vision has been demonstrated in many species and phylogenetic analysis suggests that UV-green-blue trichromacy existed from at least the Devonian
Devonian
The Devonian is a geologic period and system of the Paleozoic Era spanning from the end of the Silurian Period, about 416.0 ± 2.8 Mya , to the beginning of the Carboniferous Period, about 359.2 ± 2.5 Mya...
period between 416 and 359 million years ago.
Sound production and hearing
Insects were the earliest organisms to produce and sense sounds. Insects make sounds mostly by mechanical action of appendages. In grasshopperGrasshopper
The grasshopper is an insect of the suborder Caelifera in the order Orthoptera. To distinguish it from bush crickets or katydids, it is sometimes referred to as the short-horned grasshopper...
s and crickets, this is achieved by stridulation
Stridulation
Stridulation is the act of producing sound by rubbing together certain body parts. This behavior is mostly associated with insects, but other animals are known to do this as well, such as a number of species of fishes, snakes and spiders...
. Cicada
Cicada
A cicada is an insect of the order Hemiptera, suborder Auchenorrhyncha , in the superfamily Cicadoidea, with large eyes wide apart on the head and usually transparent, well-veined wings. There are about 2,500 species of cicada around the world, and many of them remain unclassified...
s make the loudest sounds among the insects by producing and amplifying sounds with special modifications to their body and musculature. The African cicada
Cicada
A cicada is an insect of the order Hemiptera, suborder Auchenorrhyncha , in the superfamily Cicadoidea, with large eyes wide apart on the head and usually transparent, well-veined wings. There are about 2,500 species of cicada around the world, and many of them remain unclassified...
Brevisana brevis
Brevisana brevis
Brevisana brevis is a cicada found in Africa and is the loudest insect on record. It has been recorded producing sounds with pressure levels of 106.7 decibels at a distance of 50 cm....
has been measured at 106.7 decibel
Decibel
The decibel is a logarithmic unit that indicates the ratio of a physical quantity relative to a specified or implied reference level. A ratio in decibels is ten times the logarithm to base 10 of the ratio of two power quantities...
s at a distance of 50 cm (19.7 in). Some insects, such as the hawk moths and Hedylid
Hedylidae
Hedylidae, the "American moth-butterflies", is a family of insects in the lepidopteran order, representing the superfamily Hedyloidea. They are an extant sister group of the butterfly superfamilies Papilionoidea and Hesperioidea...
butterflies, can hear ultrasound
Ultrasound
Ultrasound is cyclic sound pressure with a frequency greater than the upper limit of human hearing. Ultrasound is thus not separated from "normal" sound based on differences in physical properties, only the fact that humans cannot hear it. Although this limit varies from person to person, it is...
and take evasive action when they sense that they have been detected by bats. Some moths produce ultrasonic clicks that were once thought to have a role in jamming bat echolocation
Animal echolocation
Echolocation, also called biosonar, is the biological sonar used by several kinds of animals.Echolocating animals emit calls out to the environment and listen to the echoes of those calls that return from various objects near them. They use these echoes to locate and identify the objects...
. The ultrasonic clicks were subsequently found to be produced mostly by unpalatable moths to warn bats, just as warning coloration
Aposematism
Aposematism , perhaps most commonly known in the context of warning colouration, describes a family of antipredator adaptations where a warning signal is associated with the unprofitability of a prey item to potential predators...
s are used against predators that hunt by sight. Some otherwise palatable moths have evolved to mimic
Batesian mimicry
Batesian mimicry is a form of mimicry typified by a situation where a harmless species has evolved to imitate the warning signals of a harmful species directed at a common predator...
these calls.
Very low sounds are also produced in various species of Coleoptera, Hymenoptera
Hymenoptera
Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...
, Lepidoptera
Lepidoptera
Lepidoptera is a large order of insects that includes moths and butterflies . It is one of the most widespread and widely recognizable insect orders in the world, encompassing moths and the three superfamilies of butterflies, skipper butterflies, and moth-butterflies...
, Mantodea, and Neuroptera
Neuroptera
The insect order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order contains some 6,010 species...
. These low sounds are simply the sounds made by the insect's movement. Through microscopic stridulatory structures located on the insect's muscles and joints, the normal sounds of the insect moving are amplified and can be used to warn or communicate with other insects. Most sound-making insects also have tympanal organ
Tympanal organ
A tympanal organ is a hearing organ in insects, consisting of a membrane stretched across a frame backed by an air sac and associated sensory neurons...
s that can perceive airborne sounds. Some species in Hemiptera
Hemiptera
Hemiptera is an order of insects most often known as the true bugs , comprising around 50,000–80,000 species of cicadas, aphids, planthoppers, leafhoppers, shield bugs, and others...
, such as the corixids (water boatmen), are known to communicate via underwater sounds. Most insects are also able to sense vibration
Vibration
Vibration refers to mechanical oscillations about an equilibrium point. The oscillations may be periodic such as the motion of a pendulum or random such as the movement of a tire on a gravel road.Vibration is occasionally "desirable"...
s transmitted through surfaces. For example, an insect is caught in a spider web
Spider web
A spider web, spiderweb, spider's web or cobweb is a device built by a spider out of proteinaceous spider silk extruded from its spinnerets....
and struggles to escape. The vibrations it produces are sensed by the spider
Spider
Spiders are air-breathing arthropods that have eight legs, and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other groups of organisms...
, who is alerted to its presence. Through these vibrations, the spider can tell where on the web the insect is located, as well as how big it is.
Communication using surface-borne vibrational signals is more widespread among insects because of size constraints in producing air-borne sounds. 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. The mechanisms of production of vibrational signals are just as diverse as those for producing sound in insects.
Some species use vibrations for communicating within members of the same species, such as to attract mates as in the songs of the shield bug
Shield bug
Pentatomoidea is a superfamily of insects in the Heteroptera suborder of the Hemiptera order and, as such, share a common arrangement of sucking mouthparts...
Nezara viridula
Nezara viridula
Nezara viridula, commonly known as the southern green stink bug or green vegetable bug , is a plant-feeding stink bug. Although believed to have originated in Ethiopia, it can now be found around the world...
. Vibrations can also be used to communicate between entirely different species; lycaenid (gossamer-winged butterfly) caterpillars which are myrmecophilous (living in a mutualistic association with ants) communicate with ants in this way. The Madagascar hissing cockroach
Madagascar hissing cockroach
The Madagascar hissing Cockroach , also known as the Hissing Cockroach or simply Hisser, is one of the largest species of cockroach, reaching 2–4 inches at maturity. They are from the island of Madagascar off the African coast, where they can be found in rotting logs.Unlike most cockroaches, they...
has the ability to press air through its spiracles to make a hissing noise as a sign of aggression; the Death's-head Hawkmoth
Death's-head Hawkmoth
The name Death's-head Hawkmoth refers to any one of the three species of moth in the genus Acherontia. The former species is primarily found in Europe, the latter two are Asian, and most uses of the common name refer to the European species...
makes a squeaking noise by forcing air out of their pharynx
Pharynx
The human pharynx is the part of the throat situated immediately posterior to the mouth and nasal cavity, and anterior to the esophagus and larynx. The human pharynx is conventionally divided into three sections: the nasopharynx , the oropharynx , and the laryngopharynx...
when agitated, which may also reduce aggressive worker honey bee behavior when the two are in close proximity.
Chemical communication
In addition to the use of sound for communication, a wide range of insects have evolved chemical means for communication. These chemicals, termed semiochemicalSemiochemical
A semiochemical is a generic term used for a chemical substance or mixture that carries a message. These chemicals acts as messengers within or between species...
s, are often derived from plant metabolites include those meant to attract, repel and provide other kinds of information. Pheromone
Pheromone
A pheromone is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting outside the body of the secreting individual to impact the behavior of the receiving individual...
s, a type of semiochemical, are used for attracting mates of the opposite sex, for aggregating conspecific
Conspecificity
Conspecificity is a concept in biology. Two or more individual organisms, populations, or taxa are conspecific if they belong to the same species....
individuals of both sexes, for deterring other individuals from approaching, to mark a trail, and to trigger aggression in nearby individuals. Allomone
Allomone
An allomone is any chemical substance produced and released by an individual of one species that affects the behaviour of a member of another species to the benefit of the originator but not the receiver...
a benefit their producer by the effect they have upon the receiver. Kairomone
Kairomone
A kairomone is a semiochemical, emitted by an organism, which mediates interspecific interactions in a way that benefits an individual of another species which receives it, without benefiting the emitter. This "eavesdropping" is often disadvantageous to the producer...
s benefit their receiver instead of their producer. Synomones benefit the producer and the receiver. While some chemicals are targeted at individuals of the same species, others are used for communication across species. The use of scents is especially well known to have developed in social insects.
Social behavior
Termite
Termites are a group of eusocial insects that, until recently, were classified at the taxonomic rank of order Isoptera , but are now accepted as the epifamily Termitoidae, of the cockroach order Blattodea...
s, ant
Ant
Ants are social insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than...
s and many bee
Bee
Bees are flying insects closely related to wasps and ants, and are known for their role in pollination and for producing honey and beeswax. Bees are a monophyletic lineage within the superfamily Apoidea, presently classified by the unranked taxon name Anthophila...
s and wasp
Wasp
The term wasp is typically defined as any insect of the order Hymenoptera and suborder Apocrita that is neither a bee nor an ant. Almost every pest insect species has at least one wasp species that preys upon it or parasitizes it, making wasps critically important in natural control of their...
s, are the most familiar species of eusocial
Eusociality
Eusociality is a term used for the highest level of social organization in a hierarchical classification....
animal. They live together in large well-organized colonies that may be so tightly integrated and genetically similar that the colonies of some species are sometimes considered superorganism
Superorganism
A superorganism is an organism consisting of many organisms. This is usually meant to be a social unit of eusocial animals, where division of labour is highly specialised and where individuals are not able to survive by themselves for extended periods of time. Ants are the best-known example of...
s. It is sometimes argued that the various species of honey bee
Honey bee
Honey bees are a subset of bees in the genus Apis, primarily distinguished by the production and storage of honey and the construction of perennial, colonial nests out of wax. Honey bees are the only extant members of the tribe Apini, all in the genus Apis...
are the only invertebrates (and indeed one of the few non-human groups) to have evolved a system of abstract symbolic communication where a behavior is used to represent and convey specific information about something in the environment. In this communication system, called dance language
Bee learning and communication
Honey bees learn and communicate in order to find food sources and for other means.-Learning:Learning is essential for efficient foraging. Honey bees are unlikely to make many repeat visits if a plant provides little in the way of reward...
, 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.
Only insects which live in nests or colonies demonstrate any true capacity for fine-scale spatial orientation or homing. This can allow an insect to return unerringly to a single hole a few millimeters in diameter among thousands of apparently identical holes clustered together, after a trip of up to several kilometers' distance. In a phenomenon known as philopatry
Philopatry
Broadly, philopatry is the behaviour of remaining in, or returning to, an individual's birthplace. More specifically, in ecology philopatry is the behaviour of elder offspring sharing the parental burden in the upbringing of their siblings, a classic example of kin selection...
, insects that hibernate
Hibernation
Hibernation is a state of inactivity and metabolic depression in animals, characterized by lower body temperature, slower breathing, and lower metabolic rate. Hibernating animals conserve food, especially during winter when food supplies are limited, tapping energy reserves, body fat, at a slow rate...
have shown the ability to recall a specific location up to a year after last viewing the area of interest. A few insects seasonally migrate large distances between different geographic regions (e.g., the overwintering areas of the Monarch butterfly
Monarch butterfly
The Monarch butterfly is a milkweed butterfly , in the family Nymphalidae. It is perhaps the best known of all North American butterflies. Since the 19th century, it has been found in New Zealand, and in Australia since 1871 where it is called the Wanderer...
).
Care of young
Some bees, ants, termites and some wasps are eusocial, build nest, guard eggs and provide food for offsprings fulltime (see EusocialityEusociality
Eusociality is a term used for the highest level of social organization in a hierarchical classification....
).
Most insects, however, lead short lives as adults, and rarely interact with one another except to mate or compete for mates. A small number exhibit some form of parental care, where they will at least guard their eggs, and sometimes continue guarding their offspring until adulthood, and possibly even feeding them. Another simple form of parental care is to construct a nest (a burrow or an actual construction, either of which may be simple or complex), store provisions in it, and lay an egg upon those provisions. The adult does not contact the growing offspring, but it nonetheless does provide food. This sort of care is typical for most species of bees and various types of wasps.
Flight
Insects are the only group of invertebrates to have developed flight. The evolution of insect wings has been a subject of debate. Some entomologistEntomology
Entomology is the scientific study of insects, a branch of arthropodology...
s suggest that the wings are from paranotal lobes, or extensions from the insect's exoskeleton called the nota
Notum
The notum is the dorsal portion of an insect's thoracic segment. It is typically a single large sclerite. It can be further divided into the alinotum which bears the wings and the postnotum which bears the phragma....
, called the paranotal theory. Other theories are based on a pleural origin. These theories include suggestions that wings originated from modified gills, spiracular flaps or as from an appendage of the epicoxa. The epicoxal theory suggests the insect wings are modified epicoxal exites, a modified appendage at the base of the legs or coxa
Arthropod leg
The arthropod leg is a form of jointed appendage of arthropods, usually used for walking. Many of the terms used for arthropod leg segments are of Latin origin, and may be confused with terms for bones: coxa , trochanter , femur, tibia, tarsus, ischium, metatarsus, carpus, dactylus ,...
. In the Carboniferous
Carboniferous
The Carboniferous is a geologic period and system that extends from the end of the Devonian Period, about 359.2 ± 2.5 Mya , to the beginning of the Permian Period, about 299.0 ± 0.8 Mya . The name is derived from the Latin word for coal, carbo. Carboniferous means "coal-bearing"...
age, some of the Meganeura
Meganeura
Meganeura is a genus of extinct insects from the Carboniferous period approximately 300 million years ago, which resembled and are related to the present-day dragonflies. With wingspans of more than 75 cm , M. monyi is one of the largest known flying insect species; the Permian Meganeuropsis...
dragonflies had as much as a 50 cm (19.7 in) wide wingspan. The appearance of gigantic insects has been found to be consistent with high atmospheric oxygen. The respiratory system of insects constrains their size, however the high oxygen in the atmosphere allowed larger sizes. The largest flying insects today are much smaller and include several moth species such as the Atlas moth and the White Witch (Thysania agrippina). Insect flight has been a topic of great interest in aerodynamics
Aerodynamics
Aerodynamics is a branch of dynamics concerned with studying the motion of air, particularly when it interacts with a moving object. Aerodynamics is a subfield of fluid dynamics and gas dynamics, with much theory shared between them. Aerodynamics is often used synonymously with gas dynamics, with...
due partly to the inability of steady-state theories to explain the lift generated by the tiny wings of insects.
Unlike bird
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...
s, many small insects are swept along by the prevailing winds although many of the larger insects are known to make migrations. Aphid
Aphid
Aphids, also known as plant lice and in Britain and the Commonwealth as greenflies, blackflies or whiteflies, are small sap sucking insects, and members of the superfamily Aphidoidea. Aphids are among the most destructive insect pests on cultivated plants in temperate regions...
s are known to be transported long distances by low-level jet stream
Jet stream
Jet streams are fast flowing, narrow air currents found in the atmospheres of some planets, including Earth. The main jet streams are located near the tropopause, the transition between the troposphere and the stratosphere . The major jet streams on Earth are westerly winds...
s. As such, fine line patterns associated with converging winds within weather radar
Weather radar
Weather radar, also called weather surveillance radar and Doppler weather radar, is a type of radar used to locate precipitation, calculate its motion, estimate its type . Modern weather radars are mostly pulse-Doppler radars, capable of detecting the motion of rain droplets in addition to the...
imagery, like the WSR-88D radar network, often represent large groups of insects.
Walking
Many adult insects use six legs for walking and have adopted a tripedalTripedal
Tripedal is the term used for any animal that stands on three legs. The terms bipedal and quadrupedal are used more commonly when referring to animals that either walk on two legs Tripedal (from the Latin tri = three + ped = foot) is the term used for (or would be used for) any animal that stands...
gait
Gait
Gait is the pattern of movement of the limbs of animals, including humans, during locomotion over a solid substrate. Most animals use a variety of gaits, selecting gait based on speed, terrain, the need to maneuver, and energetic efficiency...
. The tripedal gait allows for rapid walking while always having a stable stance and has been studied extensively in cockroach
Cockroach
Cockroaches are insects of the order Blattaria or Blattodea, of which about 30 species out of 4,500 total are associated with human habitations...
es. The legs are used in alternate triangles touching the ground. 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 position. When they touch the ground to form a new stable triangle the other legs can be lifted and brought forward in turn and so on. The purest form of the tripedal gait is seen in insects moving at high speeds. However, this type of locomotion is not rigid and insects can adapt a variety of gaits. For example, when moving slowly, turning, or avoiding obstacles, four or more feet may be touching the ground. Insects can also adapt their gait to cope with the loss of one or more limbs.
Cockroaches are among the fastest insect runners and, at full speed, adopt a bipedal run to reach a high velocity in proportion to their body size. As cockroaches move very quickly, they need to be video recorded at several hundred frames per second to reveal their gait. More sedate locomotion is seen in the stick insects or walking sticks (Phasmatodea
Phasmatodea
The Phasmatodea are an order of insects, whose members are variously known as stick insects , walking sticks or stick-bugs , phasmids, ghost insects and leaf insects...
). A few insects have evolved to walk on the surface of the water, especially the bugs of the Gerridae
Water strider
Gerridae is a family of true bugs in the order Hemiptera, commonly known as water striders, water bugs, magic bugs, pond skaters, skaters, skimmers, water scooters, water skaters, water skeeters, water skimmers, water skippers, water spiders, or Jesus bugs...
family, commonly known as water striders. A few species of ocean-skaters in the genus Halobates
Halobates
Halobates or sea skaters are a genus with over 40 species of water striders. While many are coastal, about five of these are able to survive and stand on the surface of the open ocean, a habitat containing very few insect species. They are predators, coastal species feeding mainly on fallen...
even live on the surface of open oceans, a habitat that has few insect species.
Use in robotics
Insect walking is of particular interest as an alternative form of locomotion in robotRobot
A robot is a mechanical or virtual intelligent agent that can perform tasks automatically or with guidance, typically by remote control. In practice a robot is usually an electro-mechanical machine that is guided by computer and electronic programming. Robots can be autonomous, semi-autonomous or...
s. The study of insects and biped
Biped
Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs, or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning "two feet"...
s has a significant impact on possible robotic methods of transport. This may allow new robots to be designed that can traverse terrain
Terrain
Terrain, or land relief, is the vertical and horizontal dimension of land surface. When relief is described underwater, the term bathymetry is used...
that robots with wheel
Wheel
A wheel is a device that allows heavy objects to be moved easily through rotating on an axle through its center, facilitating movement or transportation while supporting a load, or performing labor in machines. Common examples found in transport applications. A wheel, together with an axle,...
s may be unable to handle.
Swimming
Water beetle
A water beetle is a beetle adapted to living in water. Water beetles rise to the water surface and take atmospheric air into their tracheal systems. There are approximately 2000 species of water beetles. The rest marine species tend to live in the intertidal zone...
s, have aquatic adults as well.
Many of these species have adaptations to help in under-water locomotion. Water beetles and water bugs have legs adapted into paddle-like structures. Dragonfly naiad
Naiad
In Greek mythology, the Naiads or Naiades were a type of nymph who presided over fountains, wells, springs, streams, and brooks....
s use jet propulsion, forcibly expelling water out of their rectal chamber. Some species like the water strider
Water strider
Gerridae is a family of true bugs in the order Hemiptera, commonly known as water striders, water bugs, magic bugs, pond skaters, skaters, skimmers, water scooters, water skaters, water skeeters, water skimmers, water skippers, water spiders, or Jesus bugs...
s are capable of walking on the surface of water. They can do this because their claws are not at the tips of the legs as in most insects, but recessed in a special groove further up the leg; this prevents the claws from piercing the water's surface film. Other insects such as the Rove beetle Stenus
Stenus
Stenus is a genus of semiaquatic rove beetles in the subfamily Steninae. They are known for swimming by emitting pygidial gland secretions that reduce surface tension and allow them to move on the surface of water....
are known to emit pygidial gland secretions that reduce surface tension making it possible for them to move on the surface of water by Marangoni propulsion
Marangoni effect
The Marangoni effect is the mass transfer along an interface between two fluids due to surface tension gradient...
(also known by the German
German language
German is a West Germanic language, related to and classified alongside English and Dutch. With an estimated 90 – 98 million native speakers, German is one of the world's major languages and is the most widely-spoken first language in the European Union....
term Entspannungsschwimmen).
Phylogeny and systematics
The evolutionEvolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...
ary relationships of insects to other animal groups remain unclear. Although more traditionally grouped with millipede
Millipede
Millipedes are arthropods that have two pairs of legs per segment . Each segment that has two pairs of legs is a result of two single segments fused together as one...
s and centipede
Centipede
Centipedes are arthropods belonging to the class Chilopoda of the subphylum Myriapoda. They are elongated metameric animals with one pair of legs per body segment. Despite the name, centipedes can have a varying number of legs from under 20 to over 300. Centipedes have an odd number of pairs of...
s, evidence has emerged favoring closer evolutionary ties
Common descent
In evolutionary biology, a group of organisms share common descent if they have a common ancestor. There is strong quantitative support for the theory that all living organisms on Earth are descended from a common ancestor....
with crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...
s. In the Pancrustacea
Pancrustacea
Pancrustacea is a clade, comprising all crustaceans and hexapods. This grouping is contrary to the Atelocerata hypothesis, in which Myriapoda and Hexapoda are sister taxa, and Crustacea are only more distantly related. As of 2010, the Pancrustacea taxon is considered well accepted. The clade has...
theory, insects, together with Remipedia
Remipedia
Remipedia is a class of blind crustaceans found in coastal aquifers which contain saline groundwater, with populations identified in almost every ocean basin so far explored, including in Australia, the Caribbean Sea, the Atlantic Ocean...
and Malacostraca
Malacostraca
Malacostraca is the largest of the six classes of crustaceans, containing over 25,000 extant species, divided among 16 orders. Its members display a greater diversity of body forms than any other class of animals, and include crabs, lobsters, shrimp, krill, woodlice, scuds , mantis shrimp and many...
, make up a natural clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...
. Other terrestrial arthropods, such as centipede
Centipede
Centipedes are arthropods belonging to the class Chilopoda of the subphylum Myriapoda. They are elongated metameric animals with one pair of legs per body segment. Despite the name, centipedes can have a varying number of legs from under 20 to over 300. Centipedes have an odd number of pairs of...
s, millipede
Millipede
Millipedes are arthropods that have two pairs of legs per segment . Each segment that has two pairs of legs is a result of two single segments fused together as one...
s, scorpion
Scorpion
Scorpions are predatory arthropod animals of the order Scorpiones within the class Arachnida. They have eight legs and are easily recognized by the pair of grasping claws and the narrow, segmented tail, often carried in a characteristic forward curve over the back, ending with a venomous stinger...
s and spider
Spider
Spiders are air-breathing arthropods that have eight legs, and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other groups of organisms...
s, are sometimes confused with insects since their body plans can appear similar, sharing (as do all arthropods) a jointed exoskeleton. However, upon closer examination their features differ significantly; most noticeably they do not have the six legs characteristic of adult insects.
A phylogenetic tree of the arthropods and related groups
The higher-level phylogeny of the arthropods continues to be a matter of debate and research. In 2008, researchers at Tufts University
Tufts University
Tufts University is a private research university located in Medford/Somerville, near Boston, Massachusetts. It is organized into ten schools, including two undergraduate programs and eight graduate divisions, on four campuses in Massachusetts and on the eastern border of France...
uncovered what they believe is the world's oldest known full-body impression of a primitive flying insect, a 300 million-year-old specimen from the Carboniferous Period. The oldest definitive insect fossil is the Devonian
Devonian
The Devonian is a geologic period and system of the Paleozoic Era spanning from the end of the Silurian Period, about 416.0 ± 2.8 Mya , to the beginning of the Carboniferous Period, about 359.2 ± 2.5 Mya...
Rhyniognatha hirsti
Rhyniognatha
Rhyniognatha hirsti is the world’s oldest known insect. It emerged very early during the Early Devonian Period, around 400 million years ago, when earth’s first terrestrial ecosystems were being formed.-Evidence:...
, from the 396 million year old Rhynie chert
Rhynie chert
The Rhynie chert is an Early Devonian sedimentary deposit exhibiting extraordinary fossil detail or completeness . It is exposed near the village of Rhynie, Aberdeenshire, Scotland; a second unit, the Windyfield chert, is located some 700 m away...
. It may have superficially resembled a modern-day silverfish
Silverfish
Lepisma saccharina, frequently called silverfish, fishmoths, carpet sharks or paramites, are small, wingless insects in the order Thysanura...
insect. This species already possessed dicondylic mandibles (two articulations in the mandible), a feature associated with winged insects, suggesting that wings may already have evolved at this time. Thus, the first insects probably appeared earlier, in the Silurian
Silurian
The Silurian is a geologic period and system that extends from the end of the Ordovician Period, about 443.7 ± 1.5 Mya , to the beginning of the Devonian Period, about 416.0 ± 2.8 Mya . As with other geologic periods, the rock beds that define the period's start and end are well identified, but the...
period.
There have been four super radiations of insects: beetle
Beetle
Coleoptera is an order of insects commonly called beetles. The word "coleoptera" is from the Greek , koleos, "sheath"; and , pteron, "wing", thus "sheathed wing". Coleoptera contains more species than any other order, constituting almost 25% of all known life-forms...
s (evolved ~300 million years ago), flies
Fly
True flies are insects of the order Diptera . They possess a pair of wings on the mesothorax and a pair of halteres, derived from the hind wings, on the metathorax...
(evolved ~250 million years ago), moth
Moth
A moth is an insect closely related to the butterfly, both being of the order Lepidoptera. Moths form the majority of this order; there are thought to be 150,000 to 250,000 different species of moth , with thousands of species yet to be described...
s and wasp
Wasp
The term wasp is typically defined as any insect of the order Hymenoptera and suborder Apocrita that is neither a bee nor an ant. Almost every pest insect species has at least one wasp species that preys upon it or parasitizes it, making wasps critically important in natural control of their...
s (evolved ~150 million years ago). These four groups account for the majority of described species. The flies and moths along with the flea
Flea
Flea is the common name for insects of the order Siphonaptera which are wingless insects with mouthparts adapted for piercing skin and sucking blood...
s evolved from the Mecoptera
Mecoptera
Mecoptera are an order of insects with about 550 species in nine families worldwide. Mecoptera are sometimes called scorpionflies after their largest family, Panorpidae, in which the males have enlarged genitals that look similar to the stinger of a scorpion...
.
The origins of insect flight
Insect flight
Insects are the only group of invertebrates known to have evolved flight. Insects possess some remarkable flight characteristics and abilities, still far superior to attempts by humans to replicate their capabilities. Even our understanding of the aerodynamics of flexible, flapping wings and how...
remain obscure, since the earliest winged insects currently known appear to have been capable fliers. Some extinct insects had an additional pair of winglets attaching to the first segment of the thorax, for a total of three pairs. As of 2009, there is no evidence that suggests that the insects were a particularly successful group of animals before they evolved to have wings.
Late Carboniferous
Pennsylvanian
The Pennsylvanian is, in the ICS geologic timescale, the younger of two subperiods of the Carboniferous Period. It lasted from roughly . As with most other geochronologic units, the rock beds that define the Pennsylvanian are well identified, but the exact date of the start and end are uncertain...
and Early Permian
Permian
The PermianThe term "Permian" was introduced into geology in 1841 by Sir Sir R. I. Murchison, president of the Geological Society of London, who identified typical strata in extensive Russian explorations undertaken with Edouard de Verneuil; Murchison asserted in 1841 that he named his "Permian...
insect orders include both extant groups and a number of Paleozoic species, now extinct. During this era, some giant dragonfly-like forms reached wingspans of 55 to 70 cm (21.7 to 27.6 in) making them far larger than any living insect. This gigantism may have been due to higher atmospheric oxygen levels that allowed increased respiratory efficiency relative to today. The lack of flying vertebrates could have been another factor. Most extinct orders of insects developed during the Permian period that began around 270 million years ago. Many of the early groups became extinct during the Permian-Triassic extinction event
Permian-Triassic extinction event
The Permian–Triassic extinction event, informally known as the Great Dying, was an extinction event that occurred 252.28 Ma ago, forming the boundary between the Permian and Triassic geologic periods, as well as the Paleozoic and Mesozoic eras...
, the largest mass extinction in the history of the Earth, around 252 million years ago.
The remarkably successful Hymenoptera
Hymenoptera
Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...
ns appeared as long as 146 million years ago in the Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...
period, but achieved their wide diversity more recently in the Cenozoic
Cenozoic
The Cenozoic era is the current and most recent of the three Phanerozoic geological eras and covers the period from 65.5 mya to the present. The era began in the wake of the Cretaceous–Tertiary extinction event at the end of the Cretaceous that saw the demise of the last non-avian dinosaurs and...
era, which began 66 million years ago. A number of highly successful insect groups evolved in conjunction with flowering plants, a powerful illustration of coevolution.
Many modern insect genera
Genera
Genera is a commercial operating system and development environment for Lisp machines developed by Symbolics. It is essentially a fork of an earlier operating system originating on the MIT AI Lab's Lisp machines which Symbolics had used in common with LMI and Texas Instruments...
developed during the Cenozoic. Insects from this period on are often found preserved in amber
Amber
Amber is fossilized tree resin , which has been appreciated for its color and natural beauty since Neolithic times. Amber is used as an ingredient in perfumes, as a healing agent in folk medicine, and as jewelry. There are five classes of amber, defined on the basis of their chemical constituents...
, often in perfect condition. The body plan, or morphology
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....
, of such specimens is thus easily compared with modern species. The study of fossilized insects is called paleoentomology
Entomology
Entomology is the scientific study of insects, a branch of arthropodology...
.
Evolutionary relationships
Insects are prey for a variety of organisms, including terrestrial vertebrates. The earliest vertebrates on land existed 400 million years ago and were large amphibious piscivoresPiscivore
A piscivore is a carnivorous animal which eats primarily fish. Piscivory was the diet of early tetrapods , insectivory came next, then in time reptiles added herbivory....
, through gradual evolutionary change, insectivory was the next diet type to evolve.
Insects were among the earliest terrestrial herbivore
Herbivore
Herbivores are organisms that are anatomically and physiologically adapted to eat plant-based foods. Herbivory is a form of consumption in which an organism principally eats autotrophs such as plants, algae and photosynthesizing bacteria. More generally, organisms that feed on autotrophs in...
s and acted as major selection agents on plants. Plants evolved chemical defenses against this herbivory
Plant defense against herbivory
Plant defense against herbivory or host-plant resistance describes a range of adaptations evolved by plants which improve their survival and reproduction by reducing the impact of herbivores. Plants use several strategies to defend against damage caused by herbivores...
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. This successful evolutionary pattern has also been utilized by mimic
Mimic
In evolutionary biology, mimicry is the similarity of one species to another which protects one or both. This similarity can be in appearance, behaviour, sound, scent and even location, with the mimics found in similar places to their models....
s. Over time, this has led to complex groups of coevolved species. Conversely, some interactions between plants and insects, like pollination
Pollination
Pollination is the process by which pollen is transferred in plants, thereby enabling fertilisation and sexual reproduction. Pollen grains transport the male gametes to where the female gamete are contained within the carpel; in gymnosperms the pollen is directly applied to the ovule itself...
, are beneficial to both organisms. Coevolution has led to the development of very specific mutualisms in such systems.
Taxonomy
{| align="right" style="background:Transparent; border:solid 0px #503DF9;"! colspan="4" style="background:#E6D09D"| Classification
|-
|
|-
{|
| rowspan="4" style="background:#ECF4ED"| Insecta
| colspan="3" style="background:#ECF4ED"|
{| align="right" class="collapsible collapsed" style="background:Transparent"
! Monocondylia
|-
| style="border-top: 1px solid black;"| -Archaeognatha
Archaeognatha
Archaeognatha is an order of wingless insects, also known as jumping bristletails. They are among the least evolutionarily changed insects, appearing in the Devonian period along with the arachnids. The name Archaeognatha is derived from Greek Archaeos meaning "ancient" and gnatha meaning "jaw"...
- 470
|}
|-
| rowspan="3" style="background:#ECF4ED"| Dicondylia
| colspan="2" style="background:#ECF4ED"|
{| align="right" class="collapsible collapsed" style="background:Transparent;"
! Apterygota
Apterygota
The name Apterygota is sometimes applied to a subclass of small, agile insects, distinguished from other insects by their lack of wings in the present and in their evolutionary history...
|-
| style="border-top: 1px solid black;"| -Thysanura
Thysanura
Thysanura is an order of insects, encompassing silverfish and firebrats, known for their three long caudal filaments.The families Machilidae and Meinertellidae of the jumping bristletails were once included with Thysanura....
<200
|-
| -Monura
Monura
Monura is an extinct order of wingless insects in the subclass Apterygota. They resembled their modern relatives, the bristletails, and had a single lengthy filament projecting from the end of the abdomen. They also had a pair of leg-like cerci and some non-ambulatory abdominal appendages. The...
|}
|-
| rowspan="2" style="background:#ECF4ED"| Pterygota
Pterygota
Pterygota is a subclass of insects that includes the winged insects. It also includes insect orders that are secondarily wingless ....
| style="background:#ECF4ED"|
{| align="right" class="collapsible collapsed" style="background:Transparent"
! Paleoptera
Paleoptera
The name Palaeoptera has been traditionally applied to those primitive groups of winged insects that lacked the ability to fold the wings back over the abdomen as characterizes the Neoptera...
|-
| style="border-top: 1px solid black;"| -Ephemeroptera- 2,500–<3,000
|-
| -Odonata
Odonata
Odonata is an order of insects, encompassing dragonflies and damselflies . The word dragonfly is also sometimes used to refer to all Odonata, but the back-formation odonate is a more correct English name for the group as a whole...
- 6,500
|}
|-
| style="background:#ECF4ED"|
{| align="right" class="collapsible collapsed" style="background:Transparent"
! Neoptera
Neoptera
Neoptera is a classification group that includes almost all the winged insects, specifically those that can flex their wings over their abdomens...
|-
| style="border-top: 1px solid black;"| -Blattodea - 3,684–4,000
|-
| -Coleoptera - 360,000–400,000
|-
| -Dermaptera - 1,816
|-
| -Diptera
Diptera
Diptera , or true flies, is the order of insects possessing only a single pair of wings on the mesothorax; the metathorax bears a pair of drumstick like structures called the halteres, the remnants of the hind wings. It is a large order, containing an estimated 240,000 species, although under half...
- 152,956
|-
| -Embioptera
Embioptera
The order Embioptera, commonly known as webspinners, are a small group of mostly tropical and subtropical insects, classified under the subclass Pterygota. The order has also been referred to as Embiodea or Embiidina...
- 200–300
|-
| -Grylloblattaria - 24
|-
| -Hymenoptera
Hymenoptera
Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...
- 115,000
|-
| -Lepidoptera
Lepidoptera
Lepidoptera is a large order of insects that includes moths and butterflies . It is one of the most widespread and widely recognizable insect orders in the world, encompassing moths and the three superfamilies of butterflies, skipper butterflies, and moth-butterflies...
- 174,250
|-
| -Mantodea - 2,200
|-
| -Mecoptera
Mecoptera
Mecoptera are an order of insects with about 550 species in nine families worldwide. Mecoptera are sometimes called scorpionflies after their largest family, Panorpidae, in which the males have enlarged genitals that look similar to the stinger of a scorpion...
- 481
|-
| -Megaloptera
Megaloptera
Megaloptera is an order of insects. It contains the alderflies, dobsonflies and fishflies, and there are about 300 known species.The Megaloptera were formerly considered part of a group then called Neuroptera, together with lacewings and snakeflies, but these are now generally considered to be...
- 250–300
|-
| -Neuroptera
Neuroptera
The insect order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order contains some 6,010 species...
- 5,000
|-
| -Orthoptera
Orthoptera
Orthoptera is an order of insects with paurometabolous or incomplete metamorphosis, including the grasshoppers, crickets and locusts.Many insects in this order produce sound by rubbing their wings against each other or their legs, the wings or legs containing rows of corrugated bumps...
- 24,380
|-
| -Phasmatodea
Phasmatodea
The Phasmatodea are an order of insects, whose members are variously known as stick insects , walking sticks or stick-bugs , phasmids, ghost insects and leaf insects...
- 2,500–3,300
|-
| -Phthiraptera - 3,000–3,200
|-
| -Plecoptera
Plecoptera
Plecoptera are an order of insects, commonly known as stoneflies. There are some 3,500 described species worldwide, with new species still being discovered. Stoneflies are found worldwide, except Antarctica...
- 2,274
|-
| -Siphonaptera - 2,525
|-
| -Strepsiptera
Strepsiptera
The Strepsiptera are an order of insects with ten families making up about 600 species...
- 596
|-
| -Trichoptera
Trichoptera
The caddisflies are an order, Trichoptera, of insects with approximately 12,000 described species. Also called sedge-flies or rail-flies, they are small moth-like insects having two pairs of hairy membranous wings...
- 12,627
|-
| -Zoraptera
Zoraptera
The insect order Zoraptera contains a single family, the Zorotypidae, which in turn contains one extant genus with 34 species, Zorotypus as well as 9 extinct species.-Phylogeny:...
- 28
|-
| -Zygentoma - 370
|-
|}
|}
{| align="right" style="background:Transparent; border:solid 0px #503DF9;"
! colspan="4" style="background:Transparent"|Cladogram
Cladogram
A cladogram is a diagram used in cladistics which shows ancestral relations between organisms, to represent the evolutionary tree of life. Although traditionally such cladograms were generated largely on the basis of morphological characters, DNA and RNA sequencing data and computational...
of living insect groups, with numbers of species in each group. Note that Apterygota
Apterygota
The name Apterygota is sometimes applied to a subclass of small, agile insects, distinguished from other insects by their lack of wings in the present and in their evolutionary history...
, Palaeoptera and Exopterygota
Exopterygota
The Exopterygota, also known as Hemipterodea, are a superorder of insects of the subclass Pterygota in the infraclass Neoptera, in which the young resemble adults but have externally-developing wings. They undergo a modest change between immature and adult, without going through a pupal stage...
are possibly paraphyletic groups.
|}
Traditional morphology-based or appearance-based systematics
Systematics
Biological systematics is the study of the diversification of terrestrial life, both past and present, and the relationships among living things through time. Relationships are visualized as evolutionary trees...
has usually given Hexapoda
Hexapoda
The subphylum Hexapoda constitutes the largest grouping of arthropods and includes the insects as well as three much smaller groups of wingless arthropods: Collembola, Protura, and Diplura . The Collembola are very abundant in terrestrial environments...
the rank of superclass, and identified four groups within it: insects (Ectognatha), springtails (Collembola
Springtail
Springtails form the largest of the three lineages of modern hexapods that are no longer considered insects...
), Protura and Diplura, the latter three being grouped together as Entognatha
Entognatha
The Entognatha is a class of ametabolous arthropods, which, together with insects, makes up the subphylum Hexapoda. Their mouthparts are entognathous, meaning they are retracted within the head. Entognatha are apterous, meaning they lack wings. The class contains three orders: Collembola , Diplura...
on the basis of internalized mouth parts. Supraordinal relationships have undergone numerous changes with the advent of methods based on evolutionary history and genetic data. A recent theory is that Hexapoda is polyphyletic (where the last common ancestor was not a member of the group), with the entognath classes having separate evolutionary histories from Insecta. Many of the traditional appearance-based taxa have been shown to be paraphyletic, so rather than using ranks like subclass
Class (biology)
In biological classification, class is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, order, family, genus, and species, with class fitting between phylum and order...
, superorder
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...
and infraorder
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...
, it has proved better to use monophyletic groupings (in which the last common ancestor is a member of the group). The following represents the best supported monophyletic groupings for the Insecta.
Insects can be divided into two groups historically treated as subclasses: wingless insects, known as Apterygota, and winged insects, known as Pterygota. The Apterygota consist of the primitively wingless order of the silverfish (Thysanura). Archaeognatha make up the Monocondylia based on the shape of their mandibles, while Thysanura and Pterygota are grouped together as Dicondylia. It is possible that the Thysanura themselves are not monophyletic, with the family Lepidotrichidae
Lepidotrichidae
Lepidotrichidae is a family of basal insects belonging to the order Thysanura. The family is considered to contain two genera: the extinct Lepidotrix, known from specimens preserved in Baltic amber, and the extant Tricholepidion, which contains a single species, Tricholepidion gertschi...
being a sister group to the Dicondylia (Pterygota and the remaining Thysanura).
Paleoptera and Neoptera are the winged orders of insects differentiated by the presence of hardened body parts called sclerite
Sclerite
A sclerite is a hardened body part. The term is used in various branches of biology for various structures including hardened portions of sponges, but it is most commonly used for the hardened portions of arthropod exoskeletons....
s; also, in Neoptera, muscles that allow their wings to fold flatly over the abdomen. Neoptera can further be divided into incomplete metamorphosis-based (Polyneoptera and Paraneoptera
Paraneoptera
Paraneoptera is a monophyletic superorder of insects which includes four orders, the bark lice, true lice, thrips, and hemipterans, the true bugs. The mouthparts of the Paraneoptera reflect diverse feeding habits...
) and complete metamorphosis-based groups. It has proved difficult to clarify the relationships between the orders in Polyneoptera because of constant new findings calling for revision of the taxa. For example, Paraneoptera has turned out to be more closely related to Endopterygota than to the rest of the Exopterygota. The recent molecular finding that the traditional louse orders Mallophaga and Anoplura are derived from within Psocoptera
Psocoptera
Psocoptera are an order of insects that are commonly known as booklice, barklice or barkflies. They first appeared in the Permian period, 295–248 million years ago. They are often regarded as the most primitive of the hemipteroids. Their name originates from the Greek word psokos meaning...
has led to the new taxon Psocodea. Phasmatodea
Phasmatodea
The Phasmatodea are an order of insects, whose members are variously known as stick insects , walking sticks or stick-bugs , phasmids, ghost insects and leaf insects...
and Embiidina have been suggested to form Eukinolabia. Mantodea, Blattodea and Isoptera are thought to form a monophyletic group termed Dictyoptera
Dictyoptera
Dictyoptera includes three groups of polyneopterous insects - cockroaches , termites and mantids...
.
It is likely that Exopterygota is paraphyletic in regard to Endopterygota. Matters that have had a lot of controversy include Strepsiptera and Diptera grouped together as Halteria based on a reduction of one of the wing pairs – a position not well-supported in the entomological community. The Neuropterida are often lumped or split on the whims of the taxonomist. Fleas are now thought to be closely related to boreid mecopterans. Many questions remain to be answered when it comes to basal relationships amongst endopterygote orders, particularly Hymenoptera.
The study of the classification or taxonomy of any insect is called systematic entomology
Entomology
Entomology is the scientific study of insects, a branch of arthropodology...
. If one works with a more specific order or even a family, the term may also be made specific to that order or family, for example systematic dipterology.
Ecology
Insect ecology is the scientific study of how insectInsect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...
s, individually or as a community, interact with the surrounding environment
Natural environment
The natural environment encompasses all living and non-living things occurring naturally on Earth or some region thereof. It is an environment that encompasses the interaction of all living species....
or ecosystem
Ecosystem
An ecosystem is a biological environment consisting of all the organisms living in a particular area, as well as all the nonliving , physical components of the environment with which the organisms interact, such as air, soil, water and sunlight....
. Insects play one of the most important roles in their ecosystems, which includes many roles, such as dung burial, pest control, pollination, and wildlife nutrition. An example is beetle
Beetle
Coleoptera is an order of insects commonly called beetles. The word "coleoptera" is from the Greek , koleos, "sheath"; and , pteron, "wing", thus "sheathed wing". Coleoptera contains more species than any other order, constituting almost 25% of all known life-forms...
s, which are scavenger
Scavenger
Scavenging is both a carnivorous and herbivorous feeding behavior in which individual scavengers search out dead animal and dead plant biomass on which to feed. The eating of carrion from the same species is referred to as cannibalism. Scavengers play an important role in the ecosystem by...
s that feed on dead animals and fallen trees and thereby recycle
Recycling
Recycling is processing used materials into new products to prevent waste of potentially useful materials, reduce the consumption of fresh raw materials, reduce energy usage, reduce air pollution and water pollution by reducing the need for "conventional" waste disposal, and lower greenhouse...
biological materials into forms found useful by other organism
Organism
In biology, an organism is any contiguous living system . In at least some form, all organisms are capable of response to stimuli, reproduction, growth and development, and maintenance of homoeostasis as a stable whole.An organism may either be unicellular or, as in the case of humans, comprise...
s. These insects, and others, are responsible for much of the process by which topsoil
Topsoil
Topsoil is the upper, outermost layer of soil, usually the top to . It has the highest concentration of organic matter and microorganisms and is where most of the Earth's biological soil activity occurs.-Importance:...
is created.
Defense and predation
Insects are mostly soft bodied, fragile and almost defenseless compared to other, larger lifeforms, while the immature stages are small, move slowly or are immobile, hence all stages are exposed to predationPredation
In ecology, predation describes a biological interaction where a predator feeds on its prey . Predators may or may not kill their prey prior to feeding on them, but the act of predation always results in the death of its prey and the eventual absorption of the prey's tissue through consumption...
and parasitism
Parasitism
Parasitism is a type of symbiotic relationship between organisms of different species where one organism, the parasite, benefits at the expense of the other, the host. Traditionally parasite referred to organisms with lifestages that needed more than one host . These are now called macroparasites...
. So insects have a variety of strategies to avoid being attacked by predators or parasitoid
Parasitoid
A parasitoid is an organism that spends a significant portion of its life history attached to or within a single host organism in a relationship that is in essence parasitic; unlike a true parasite, however, it ultimately sterilises or kills, and sometimes consumes, the host...
s. These include camouflage
Camouflage
Camouflage is a method of concealment that allows an otherwise visible animal, military vehicle, or other object to remain unnoticed, by blending with its environment. Examples include a leopard's spotted coat, the battledress of a modern soldier and a leaf-mimic butterfly...
, mimicry, toxicity, and active defense.
Camouflage
Camouflage
Camouflage is a method of concealment that allows an otherwise visible animal, military vehicle, or other object to remain unnoticed, by blending with its environment. Examples include a leopard's spotted coat, the battledress of a modern soldier and a leaf-mimic butterfly...
is also important defense strategies, which involves the use of coloration or shape to blend into the surrounding environment. This sort of protective coloration is common and widespread among beetle families, especially those that feed on wood or vegetation, such as many of the leaf beetle
Leaf beetle
Beetles in the family Chrysomelidae are commonly known as leaf beetles. This is a family of over 35,000 species in more than 2,500 genera, one of the largest and most commonly encountered of all beetle families....
s (family Chrysomelidae) or weevil
Weevil
A weevil is any beetle from the Curculionoidea superfamily. They are usually small, less than , and herbivorous. There are over 60,000 species in several families, mostly in the family Curculionidae...
s. In some of these species, sculpturing or various colored scales or hairs cause the beetle to resemble bird dung or other inedible objects. Many of those that live in sandy environments blend in with the coloration of the substrate. Most phasmids are known for effectively replicating the forms of sticks and leaves, and the bodies of some species (such as O. macklotti and Palophus centaurus) are covered in mossy or lichenous
Lichen
Lichens are composite organisms consisting of a symbiotic organism composed of a fungus with a photosynthetic partner , usually either a green alga or cyanobacterium...
outgrowths that supplement their disguise. Some species have the ability to change color as their surroundings shift (B. scabrinota, T. californica). In a further behavioral adaptation to supplement crypsis
Crypsis
In ecology, crypsis is the ability of an organism to avoid observation or detection by other organisms. It may be either a predation strategy or an antipredator adaptation, and methods include camouflage, nocturnality, subterranean lifestyle, transparency, and mimicry...
, a number of species have been noted to perform a rocking motion where the body is swayed from side to side that is thought to reflect the movement of leaves or twigs swaying in the breeze. Another method by which stick insects avoid predation and resemble twigs is by feigning death (catalepsy
Catalepsy
Catalepsy is also a term used by hypnotists to refer to the state of making a hypnotised subject's arm, leg or back rigid. "Arm catalepsy" is often a pre-hypnotic test performed prior to an induction into a full trance.-Causes:...
), where the insect enters a motionless state that can be maintained for a long period. The nocturnal feeding habits of adults also aids Phasmatodea in remaining concealed from predators.
Another defense that often uses color or shape to deceive potential enemies is mimicry. A number of longhorn beetle
Longhorn beetle
The longhorn beetles are a cosmopolitan family of beetles, typically characterized by extremely long antennae, which are often as long as or longer than the beetle's body...
s (family Cerambycidae) bear a striking resemblance to wasp
Wasp
The term wasp is typically defined as any insect of the order Hymenoptera and suborder Apocrita that is neither a bee nor an ant. Almost every pest insect species has at least one wasp species that preys upon it or parasitizes it, making wasps critically important in natural control of their...
s, which helps them avoid predation even though the beetles are in fact harmless. Batesian
Batesian mimicry
Batesian mimicry is a form of mimicry typified by a situation where a harmless species has evolved to imitate the warning signals of a harmful species directed at a common predator...
and Müllerian
Müllerian mimicry
Müllerian mimicry is a natural phenomenon when two or more harmful species, that may or may not be closely related and share one or more common predators, have come to mimic each other's warning signals...
mimicry complexes are commonly found in Lepidoptera. Genetic polymorphism and natural selection give rise to otherwise edible species (the mimic) gaining a survival advantage by resembling inedible species (the model). Such a mimicry complex is referred to as Batesian and is most commonly known by the mimicry by the limenitidine
Limenitidinae
Limenitidinae is a subfamily of butterflies that includes the admirals and relatives. The common names of many species and genera reference military ranks or – namely the Adoliadini – titles of nobility , in reference to these butterflies' large size, bold pattern and dashing flight...
Viceroy butterfly
Viceroy butterfly
The Viceroy Butterfly is a North American butterfly with a range from the Northwest Territories along the eastern edges of the Cascade Range and Sierra Nevada mountains, southwards into central Mexico....
of the inedible danaine Monarch. Later research has discovered that the Viceroy is, in fact more toxic than the Monarch and this resemblance should be considered as a case of Müllerian mimicry. In Müllerian mimicry, inedible species, usually within a taxonomic order, find it advantageous to resemble each other so as to reduce the sampling rate by predators who need to learn about the insects' inedibility. Taxa from the toxic genus Heliconius
Heliconius
Heliconius comprise a colorful and widespread brush-footed butterfly genus distributed throughout the tropical and subtropical regions of the New World. These butterflies utilize Passion flower plants as their larval food source and rely on bright wing color patterns to signal their distastefulness...
form one of the most well known Müllerian complexes.
Chemical defense is another important defense found amongst species of Coleoptera, usually being advertised by bright colors, such as the Monarch butterfly. They obtain their toxicity by sequestering the chemicals from the plants they eat into their own tissues. Some Lepidoptera manufacture their own toxins. Predators that eat poisonous butterflies and moths may become sick and vomit violently, learning not to eat those types of species; this is actually the basis of Müllerian mimicry. A predator who has previously eaten a poisonous lepidopteran may avoid other species with similar markings in the future, thus saving many other species as well. Chemical defense is another important defense found amongst species of Coleoptera, with some species releasing chemicals in the form of a spray with surprising accuracy, such as ground beetle
Ground beetle
Ground beetles are a large, cosmopolitan family of beetles, Carabidae, with more than 40,000 species worldwide, approximately 2,000 of which are found in North America and 2,700 in Europe.-Description and ecology:...
s (Carabidae), may spray chemicals from their abdomen to repel predators.
As pests
Pest (animal)
A pest is an animal which is detrimental to humans or human concerns. It is a loosely defined term, often overlapping with the related terms vermin, weeds, parasites and pathogens...
by humans. Insects commonly regarded as pests include those that are parasitic (mosquito
Mosquito
Mosquitoes are members of a family of nematocerid flies: the Culicidae . The word Mosquito is from the Spanish and Portuguese for little fly...
es, lice, bed bugs), transmit diseases (mosquitoes, flie
Fly
True flies are insects of the order Diptera . They possess a pair of wings on the mesothorax and a pair of halteres, derived from the hind wings, on the metathorax...
s), damage structures (termite
Termite
Termites are a group of eusocial insects that, until recently, were classified at the taxonomic rank of order Isoptera , but are now accepted as the epifamily Termitoidae, of the cockroach order Blattodea...
s), or destroy agricultural goods (locust
Locust
Locusts are the swarming phase of short-horned grasshoppers of the family Acrididae. These are species that can breed rapidly under suitable conditions and subsequently become gregarious and migratory...
s, weevil
Weevil
A weevil is any beetle from the Curculionoidea superfamily. They are usually small, less than , and herbivorous. There are over 60,000 species in several families, mostly in the family Curculionidae...
s). Many entomologists are involved in various forms of pest control, as in research for companies to produce insecticides, but increasingly relying on methods of biological pest control
Biological pest control
Biological control of pests in agriculture is a method of controlling pests that relies on predation, parasitism, herbivory, or other natural mechanisms...
, or biocontrol. Biocontrol uses one organism to reduce the population density of another organism — the pest — and is considered a key element of integrated pest management
Integrated Pest Management
Integrated pest management is an ecological approach to agricultural pest control that integrates pesticides/herbicides into a management system incorporating a range of practices for economic control of a pest...
.
Despite the large amount of effort focused at controlling insects, human attempts to kill pests with insecticides can backfire. If used carelessly the poison can kill all kinds of organisms in the area, including insects' natural predators such as birds, mice, and other insectivores. The effects of DDT
DDT
DDT is one of the most well-known synthetic insecticides. It is a chemical with a long, unique, and controversial history....
's use exemplifies how some insecticides can threaten wildlife beyond intended populations of pest insects.
As beneficial
Environment (biophysical)
The biophysical environment is the combined modeling of the physical environment and the biological life forms within the environment, and includes all variables, parameters as well as conditions and modes inside the Earth's biosphere. The biophysical environment can be divided into two categories:...
and to human
Human
Humans are the only living species in the Homo genus...
s. Some insects, like wasp
Wasp
The term wasp is typically defined as any insect of the order Hymenoptera and suborder Apocrita that is neither a bee nor an ant. Almost every pest insect species has at least one wasp species that preys upon it or parasitizes it, making wasps critically important in natural control of their...
s, bee
Bee
Bees are flying insects closely related to wasps and ants, and are known for their role in pollination and for producing honey and beeswax. Bees are a monophyletic lineage within the superfamily Apoidea, presently classified by the unranked taxon name Anthophila...
s, butterflie
Butterfly
A butterfly is a mainly day-flying insect of the order Lepidoptera, which includes the butterflies and moths. Like other holometabolous insects, the butterfly's life cycle consists of four parts: egg, larva, pupa and adult. Most species are diurnal. Butterflies have large, often brightly coloured...
s, and ant
Ant
Ants are social insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than...
s, pollinate
Pollination
Pollination is the process by which pollen is transferred in plants, thereby enabling fertilisation and sexual reproduction. Pollen grains transport the male gametes to where the female gamete are contained within the carpel; in gymnosperms the pollen is directly applied to the ovule itself...
flowering plant
Flowering plant
The flowering plants , also known as Angiospermae or Magnoliophyta, are the most diverse group of land plants. Angiosperms are seed-producing plants like the gymnosperms and can be distinguished from the gymnosperms by a series of synapomorphies...
s. Pollination is a mutualistic relationship between plants and insects. As insects gather nectar from different plants of the same species, they also spread pollen
Pollen
Pollen is a fine to coarse powder containing the microgametophytes of seed plants, which produce the male gametes . Pollen grains have a hard coat that protects the sperm cells during the process of their movement from the stamens to the pistil of flowering plants or from the male cone to the...
from plants on which they have previously fed. This greatly increases plants' ability to cross-pollinate, which maintains and possibly even improves their evolutionary fitness
Fitness (biology)
Fitness is a central idea in evolutionary theory. It can be defined either with respect to a genotype or to a phenotype in a given environment...
. This ultimately affects humans since ensuring healthy crops is critical to agriculture
Agriculture
Agriculture is the cultivation of animals, plants, fungi and other life forms for food, fiber, and other products used to sustain life. Agriculture was the key implement in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that nurtured the...
. A serious environmental problem is the decline of populations of pollinator
Pollinator decline
The term pollinator decline refers to the reduction in abundance of pollinators in many ecosystems worldwide during the end of the twentieth century....
insects, and a number of species of insects are now cultured primarily for pollination management
Pollination management
Pollination Management is the label for horticultural practices that accomplish or enhance pollination of a crop, to improve yield or quality, by understanding of the particular crop's pollination needs, and by knowledgeable management of pollenizers, pollinators, and pollination...
in order to have sufficient pollinators in the field, orchard
Orchard
An orchard is an intentional planting of trees or shrubs that is maintained for food production. Orchards comprise fruit or nut-producing trees which are grown for commercial production. Orchards are also sometimes a feature of large gardens, where they serve an aesthetic as well as a productive...
or greenhouse
Greenhouse
A greenhouse is a building in which plants are grown. These structures range in size from small sheds to very large buildings...
at bloom
Flowering plant
The flowering plants , also known as Angiospermae or Magnoliophyta, are the most diverse group of land plants. Angiosperms are seed-producing plants like the gymnosperms and can be distinguished from the gymnosperms by a series of synapomorphies...
time.
Insects also produce useful substances such as honey
Honey
Honey is a sweet food made by bees using nectar from flowers. The variety produced by honey bees is the one most commonly referred to and is the type of honey collected by beekeepers and consumed by humans...
, wax
Wax
thumb|right|[[Cetyl palmitate]], a typical wax ester.Wax refers to a class of chemical compounds that are plastic near ambient temperatures. Characteristically, they melt above 45 °C to give a low viscosity liquid. Waxes are insoluble in water but soluble in organic, nonpolar solvents...
, lacquer
Lacquer
In a general sense, lacquer is a somewhat imprecise term for a clear or coloured varnish that dries by solvent evaporation and often a curing process as well that produces a hard, durable finish, in any sheen level from ultra matte to high gloss and that can be further polished as required...
and silk
Silk
Silk is a natural protein fiber, some forms of which can be woven into textiles. The best-known type of silk is obtained from the cocoons of the larvae of the mulberry silkworm Bombyx mori reared in captivity...
. Honey bee
Honey bee
Honey bees are a subset of bees in the genus Apis, primarily distinguished by the production and storage of honey and the construction of perennial, colonial nests out of wax. Honey bees are the only extant members of the tribe Apini, all in the genus Apis...
s have been cultured by humans for thousands of years for honey, although contracting for crop pollination is becoming more significant for beekeeper
Beekeeper
A beekeeper is a person who keeps honey bees for the purposes of securing commodities such as honey, beeswax, pollen, royal jelly; pollinating fruits and vegetables; raising queens and bees for sale to other farmers; and/or for purposes satisfying natural scientific curiosity...
s. The silkworm has greatly affected human history, as silk-driven trade
Silk Road
The Silk Road or Silk Route refers to a historical network of interlinking trade routes across the Afro-Eurasian landmass that connected East, South, and Western Asia with the Mediterranean and European world, as well as parts of North and East Africa...
established relationships between China and the rest of the world.
Insectivore
An insectivore is a type of carnivore with a diet that consists chiefly of insects and similar small creatures. An alternate term is entomophage, which also refers to the human practice of eating insects....
insects, or insects which feed on other insects, are beneficial to humans because they eat insects that could cause damage to agriculture and human structures. For example, aphid
Aphid
Aphids, also known as plant lice and in Britain and the Commonwealth as greenflies, blackflies or whiteflies, are small sap sucking insects, and members of the superfamily Aphidoidea. Aphids are among the most destructive insect pests on cultivated plants in temperate regions...
s feed on crops and cause problems for farmers, but ladybugs feed on aphids, and can be used as a means to get significantly reduce pest aphid populations. While bird
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...
s are perhaps more visible predators of insects, insects themselves account for the vast majority of insect consumption. Without predators to keep them in check, insects can undergo almost unstoppable population explosions.
Insects are also used in medicine, for example fly larvae (maggot
Maggot
In everyday speech the word maggot means the larva of a fly ; it is applied in particular to the larvae of Brachyceran flies, such as houseflies, cheese flies, and blowflies, rather than larvae of the Nematocera, such as mosquitoes and Crane flies...
s) were formerly used to treat wounds
Maggot therapy
Maggot therapy is a type of biotherapy involving the intentional introduction of live, disinfected maggots into the non-healing skin and soft tissue wound of a human or animal for the purpose of cleaning out the...
to prevent or stop gangrene
Gangrene
Gangrene is a serious and potentially life-threatening condition that arises when a considerable mass of body tissue dies . This may occur after an injury or infection, or in people suffering from any chronic health problem affecting blood circulation. The primary cause of gangrene is reduced blood...
, as they would only consume dead flesh. This treatment is finding modern usage in some hospitals. Recently insects have also gained attention as potential sources of drugs and other medicinal substances. Also adult insects, such as crickets, and insect larvae of various kinds are also commonly used as fishing bait.
In research
Fecundity
Fecundity, derived from the word fecund, generally refers to the ability to reproduce. In demography, fecundity is the potential reproductive capacity of an individual or population. In biology, the definition is more equivalent to fertility, or the actual reproductive rate of an organism or...
, the common fruit fly Drosophila melanogaster
Drosophila melanogaster
Drosophila melanogaster is a species of Diptera, or the order of flies, in the family Drosophilidae. The species is known generally as the common fruit fly or vinegar fly. Starting from Charles W...
is a model organism
Model organism
A model organism is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the organism model will provide insight into the workings of other organisms. Model organisms are in vivo models and are widely used to...
for studies in the genetics
Genetics
Genetics , a discipline of biology, is the science of genes, heredity, and variation in living organisms....
of higher eukaryote
Eukaryote
A eukaryote is an organism whose cells contain complex structures enclosed within membranes. Eukaryotes may more formally be referred to as the taxon Eukarya or Eukaryota. The defining membrane-bound structure that sets eukaryotic cells apart from prokaryotic cells is the nucleus, or nuclear...
s. D. melanogaster has been an essential part of studies into principles like genetic linkage
Genetic linkage
Genetic linkage is the tendency of certain loci or alleles to be inherited together. Genetic loci that are physically close to one another on the same chromosome tend to stay together during meiosis, and are thus genetically linked.-Background:...
, interactions between genes
Epistasis
In genetics, epistasis is the phenomenon where the effects of one gene are modified by one or several other genes, which are sometimes called modifier genes. The gene whose phenotype is expressed is called epistatic, while the phenotype altered or suppressed is called hypostatic...
, chromosomal
Chromosome
A chromosome is an organized structure of DNA and protein found in cells. It is a single piece of coiled DNA containing many genes, regulatory elements and other nucleotide sequences. Chromosomes also contain DNA-bound proteins, which serve to package the DNA and control its functions.Chromosomes...
genetics, development, behavior, and evolution
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...
. Because genetic systems are well conserved among eukaryotes, understanding basic cellular processes like DNA replication
DNA replication
DNA replication is a biological process that occurs in all living organisms and copies their DNA; it is the basis for biological inheritance. The process starts with one double-stranded DNA molecule and produces two identical copies of the molecule...
or transcription
Transcription (genetics)
Transcription is the process of creating a complementary RNA copy of a sequence of DNA. Both RNA and DNA are nucleic acids, which use base pairs of nucleotides as a complementary language that can be converted back and forth from DNA to RNA by the action of the correct enzymes...
in fruit flies can help to understand those processes in other eukaryotes, including humans. The genome
Genome
In modern molecular biology and genetics, the genome is the entirety of an organism's hereditary information. It is encoded either in DNA or, for many types of virus, in RNA. The genome includes both the genes and the non-coding sequences of the DNA/RNA....
of D. melanogaster was sequenced
Genome project
Genome projects are scientific endeavours that ultimately aim to determine the complete genome sequence of an organism and to annotate protein-coding genes and other important genome-encoded features...
in 2000, reflecting the organism's important role in biological research.
As food
In some parts of the world, insects are used for human foodEntomophagy
Entomophagy is the consumption of insects as food. Insects are eaten by many animals, but the term is generally used to refer to human consumption of insects; animals that eat insects are known as insectivores...
, while being a taboo
Taboo
A taboo is a strong social prohibition relating to any area of human activity or social custom that is sacred and or forbidden based on moral judgment, religious beliefs and or scientific consensus. Breaking the taboo is usually considered objectionable or abhorrent by society...
in other places. In some cultures, insects, especially deep-fried cicada
Cicada
A cicada is an insect of the order Hemiptera, suborder Auchenorrhyncha , in the superfamily Cicadoidea, with large eyes wide apart on the head and usually transparent, well-veined wings. There are about 2,500 species of cicada around the world, and many of them remain unclassified...
s, are considered to be delicacie
Delicatessen
Delicatessen is a term meaning "delicacies" or "fine foods". The word entered English via German,with the old German spelling , plural of Delikatesse "delicacy", ultimately from Latin delicatus....
s, while in other places they form part of the normal diet as they have a high protein content for their mass. In most first-world countries, however, entomophagy
Entomophagy
Entomophagy is the consumption of insects as food. Insects are eaten by many animals, but the term is generally used to refer to human consumption of insects; animals that eat insects are known as insectivores...
, or the consumption of insects, is taboo
Taboo
A taboo is a strong social prohibition relating to any area of human activity or social custom that is sacred and or forbidden based on moral judgment, religious beliefs and or scientific consensus. Breaking the taboo is usually considered objectionable or abhorrent by society...
.
There are proponents of developing this use to provide a major source of protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...
in human nutrition
Nutrition
Nutrition is the provision, to cells and organisms, of the materials necessary to support life. Many common health problems can be prevented or alleviated with a healthy diet....
. Since it is impossible to entirely eliminate pest insects from the human food chain, insects are present in many foods, especially grains. Food safety
Food safety
Food safety is a scientific discipline describing handling, preparation, and storage of food in ways that prevent foodborne illness. This includes a number of routines that should be followed to avoid potentially severe health hazards....
laws in many countries do not prohibit insect parts in food, but rather limit the quantity. According to cultural materialist
Cultural materialism (anthropology)
Cultural materialism is an anthropological research orientation first introduced by Marvin Harris in his 1968 book The Rise of Anthropological Theory, as a theoretical paradigm and research strategy. Indeed it is said to be the most enduring achievement of that work. Harris subsequently developed a...
anthropologist Marvin Harris
Marvin Harris
Marvin Harris was an American anthropologist. He was born in Brooklyn, New York. A prolific writer, he was highly influential in the development of cultural materialism...
, the eating of insects is taboo in cultures that have other protein sources such as fish or livestock.
In culture
Scarab beetlesScarabaeidae
The family Scarabaeidae as currently defined consists of over 30,000 species of beetles worldwide. The species in this large family are often called scarabs or scarab beetles. The classification of this family is fairly unstable, with numerous competing theories, and new proposals appearing quite...
held religious and cultural symbolism in Old Egypt, Greece
Greece
Greece , officially the Hellenic Republic , and historically Hellas or the Republic of Greece in English, is a country in southeastern Europe....
and some shamanistic Old World cultures. The ancient Chinese
China
Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...
regarded cicada
Cicada
A cicada is an insect of the order Hemiptera, suborder Auchenorrhyncha , in the superfamily Cicadoidea, with large eyes wide apart on the head and usually transparent, well-veined wings. There are about 2,500 species of cicada around the world, and many of them remain unclassified...
s as symbols of rebirth or immortality. In Mesopotamia
Mesopotamia
Mesopotamia is a toponym for the area of the Tigris–Euphrates river system, largely corresponding to modern-day Iraq, northeastern Syria, southeastern Turkey and southwestern Iran.Widely considered to be the cradle of civilization, Bronze Age Mesopotamia included Sumer and the...
n literature, the epic poem of Gilgamesh
Gilgamesh
Gilgamesh was the fifth king of Uruk, modern day Iraq , placing his reign ca. 2500 BC. According to the Sumerian king list he reigned for 126 years. In the Tummal Inscription, Gilgamesh, and his son Urlugal, rebuilt the sanctuary of the goddess Ninlil, in Tummal, a sacred quarter in her city of...
has allusions to Odonata
Odonata
Odonata is an order of insects, encompassing dragonflies and damselflies . The word dragonfly is also sometimes used to refer to all Odonata, but the back-formation odonate is a more correct English name for the group as a whole...
which signify the impossibility of immortality.
Amongst the Aborigines
Indigenous Australians
Indigenous Australians are the original inhabitants of the Australian continent and nearby islands. The Aboriginal Indigenous Australians migrated from the Indian continent around 75,000 to 100,000 years ago....
of Australia
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...
of the Arrernte language groups, honey ants and witchety grubs served as personal clan totems. In the case of the 'San' bush-men of the Kalahari, it is the praying mantis which holds much cultural significance including creation and zen-like patience in waiting.
See also
- EntomologyEntomologyEntomology is the scientific study of insects, a branch of arthropodology...
- EthnoentomologyEthnoentomologyEthnoentomology is the study of the relationship between insects and people. The name is derived from "ethno" - study of people and "entomology" - study of insects. The focus of ethnoentomology is on how insects have been or are being used in human societies around the world...
- Flying and gliding animalsFlying and gliding animalsA number of animals have evolved aerial locomotion, either by powered flight or by gliding. Flying and gliding animals have evolved separately many times, without any single ancestor. Flight has evolved at least four times, in the insects, pterosaurs, birds, and bats. Gliding has evolved on many...
- :Category: Insect-borne diseases
- Prehistoric insectsPrehistoric insectsPrehistoric insects are various groups of insects that lived before recorded history. Their study is the field of paleoentomology. Insects inhabited Earth since before the time of the dinosaurs. The earliest identifiable insect is the Devonian Rhyniognatha hirsti, estimated at...
- Ecological importance of bees
- Insect ecologyInsect ecologyInsect ecology is the scientific study of how insects, individually or as a community, interact with the surrounding environment or ecosystem....
- Chemical ecologyChemical ecologyChemical ecology is the study of the chemicals involved in the interactions of living organisms. It focuses on the production of and response to signaling molecules and toxins. Chemical ecology is of particular importance among ants and other social insects – including bees, wasps, and termites –...
- Insect biodiversityInsect biodiversityInsect biodiversity accounts for a large proportion of all biodiversity on the planet, with over 1,000,000 insect species described.-Agriculture:...
- Defense in insectsDefense in insectsInsects have a wide variety of predators, including birds, reptiles, amphibians, mammals, carnivorous plants, and other arthropods. The great majority of individuals born do not survive to reproductive age, with perhaps 50% of this mortality rate attributed to predation. In order to deal with this...
External links
- Insects of North America
- Overview of Orders of Insects
- Manual for Entomology and Arachnology
- Tree of Life Project – Insecta, Insecta Movies
- Insects of Brazil
- Insect Morphology Overview of insect external and internal anatomy
- Fossil Insect Database International Palaeoentological Society
- UF Book of Insect Records
- InsectImages.org 24,000 high resolution insect photographs
- BBC Nature: Insect news, and video clips from BBC programmes past and present.
- The Nature Explorers Many insect video clips.
