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Temporal range: Carboniferous–Present
Insects, such as this scorpionfly, have a three-part body: head with large compound eyes and antennae, a thorax with 3 pairs of legs and often wings, and a segmented abdomen.
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Clade: Pancrustacea
Subphylum: Hexapoda
Class: Insecta
Linnaeus, 1758

See text.

  • Ectognatha
  • Entomida

Insects (from Latin insectum) are hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs of jointed legs, compound eyes, and a pair of antennae. Their blood is not totally contained in vessels; some circulates in an open haemocoel. Insects are the most diverse group of animals, with more than a million described species; they represent more than half of all animal species.

Nearly all insects hatch from eggs. Insect growth is constrained by the inelastic exoskeleton, so development involves a series of molts. The immature stages often differ from the adults in structure, habit and habitat. Groups that undergo four-stage metamorphosis often have a nearly immobile pupa. Insects that undergo three-stage metamorphosis lack a pupa, developing through a series of increasingly adult-like nymphal stages. The higher level relationship of the insects 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 to 28 in). The most diverse insect groups appear to have coevolved with flowering plants.

Adult insects typically move about by walking and flying; some can swim. Insects are the only invertebrates that can achieve sustained powered flight; insect flight evolved just once. Many insects spend part of their lives under water; their larvae have gills, while some have aquatic adults adapted to swimming. Some species, such as water striders, can walk on the surface of water. Insects are mostly solitary, but some, such as bees, ants and termites, are social and live in large, well-organized colonies. Others, such as earwigs, provide maternal care, guarding their eggs and young. Insects can communicate with each other in a variety of ways. Male moths can sense the pheromones of female moths over great distances. Other species communicate with sounds: crickets stridulate, or rub their wings together, to attract a mate and repel other males. Lampyrid beetles communicate with light.

Humans regard many insects as pests, especially those that damage crops, and attempt to control them using insecticides and other techniques. Others are parasitic, and may act as disease vectors. Others again have complex ecological roles; blow-flies consume carrion but spread diseases. Insect pollinators are essential to the reproduction of many flowering plants and so to their ecosystems. Many insects are ecologically beneficial as predators of pest insects, while a few provide direct economic benefit. Two species in particular are economically important and were domesticated many centuries ago: silkworms for silk and honey bees for honey. Insects are consumed as food in 80% of the world's nations, by people in roughly 3000 ethnic groups. Human activities are having serious effects on insect biodiversity.


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

Insects and other bugs

Distinguishing features

In common parlance, insects and other terrestrial arthropods are often called bugs (from Middle English bugge meaning "scarecrow, hobgoblin").[a] Entomologists to some extent reserve the name "bugs" for a narrow category of "true bugs", insects of the order Hemiptera, such as cicadas and shield bugs.[6] Other terrestrial arthropods, such as centipedes, millipedes, woodlice, spiders, mites and scorpions, are sometimes confused with insects, since they have a jointed exoskeleton.[7] Adult insects can be distinguished by their three-part body plan, with head, thorax, and abdomen. The thorax carries three pairs of legs, and up to two pairs of wings, unlike all other arthropods.[8]


Half of all eukaryotes are insects.

Estimates of the total number of insect species vary considerably, suggesting that there are perhaps some 5.5 million insect species in existence, of which about 1 million have been described and named.[9] These constitute around half of all eukaryote species, including animals, plants, and fungi.[10][11][12][13][14]

Phylogeny and evolution

External phylogeny

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


Collembola (springtails)

Protura (coneheads)

Diplura (two-pronged bristletails)

Insecta (=Ectognatha)

Internal phylogeny

The internal phylogeny is based on the works of Sroka, Staniczek & Bechly 2014,[17] Prokop et al. 2017[18] and Wipfler et al. 2019.[19] The numbers of described extant species (boldface for groups with over 100,000 species) are from Stork 2018.[9]


Archaeognatha (hump-backed/jumping bristletails, 513 spp)


Zygentoma (silverfish, firebrats, fishmoths, 560 spp)


Odonatoptera (dragonflies, 5,899 spp)

Panephemeroptera (mayflies, 3,240 spp)


Zoraptera (angel insects, 37 spp)

Dermaptera (earwigs, 1,978 spp)

Plecoptera (stoneflies, 3,743 spp)

Orthoptera (grasshoppers, crickets, katydids, 23,855 spp)


Mantodea (mantises, 2,400 spp)

Blattodea (cockroaches & termites, 7,314 spp)


Grylloblattodea (ice crawlers, 34 spp)

Mantophasmatodea (gladiators, 15 spp)


Phasmatodea (stick insects, 3,014 spp)

Embioptera (webspinners, 463 spp)


Psocodea (book lice, barklice & sucking lice, 11,000 spp[20])

Hemiptera (true bugs, 103,590 spp)

Thysanoptera (thrips, 5,864 spp)


Hymenoptera (sawflies, wasps, bees, ants, 116,861 spp)


Strepsiptera (twisted-wing flies, 609 spp)

Coleoptera (beetles, 386,500 spp)


Raphidioptera (snakeflies, 254 spp)

Neuroptera (lacewings, 5,868 spp)

Megaloptera (alderflies and dobsonflies, 354 spp)


Lepidoptera (butterflies & moths, 157,338 spp)

Trichoptera (caddisflies, 14,391 spp)


Diptera (true flies, 155,477 spp)


Mecoptera (scorpionflies, 757 spp)

Neomecoptera (winter scorpionflies)

Siphonaptera (fleas, 2,075 spp)

larvae, pupae
wings flex over abdomen


Traditional morphology-based systematics have usually given the Hexapoda the rank of superclass,[21] and identified four groups within it: insects (Ectognatha), springtails (Collembola), Protura, and Diplura, the latter three being grouped together as the Entognatha on the basis of internalized mouth parts. Relationships above the level of orders have undergone numerous changes with the use of phylogenetic data.[22] Insects can be divided into two groups historically treated as subclasses: wingless insects or Apterygota, and winged insects or Pterygota. The Apterygota traditionally consisted of the primitively wingless orders Archaeognatha (jumping bristletails) and Zygentoma (silverfish). However, Apterygota is not monophyletic, as Archaeognatha are sister to all other insects, based on the arrangement of their mandibles, while Zygentoma now forms part of the Dicondylia clade, with the Pterygota.[23]

Paleoptera and Neoptera are winged and have hardened body parts; the Neoptera have muscles that allow their wings to fold flat over the abdomen. Neoptera can be divided into groups with incomplete metamorphosis (Polyneoptera and Paraneoptera) and those with complete metamorphosis. The molecular finding that the traditional louse orders Mallophaga and Anoplura are within Psocoptera has led to the new taxon Psocodea.[24] Phasmatodea and Embiidina have been suggested to form the Eukinolabia.[25] Mantodea, Blattodea, and Isoptera form a monophyletic group, Dictyoptera.[26] Fleas are now thought to be closely related to boreid mecopterans.[27]

Evolutionary history

The oldest fossil that may be a primitive wingless insect is Leverhulmia from the Early Devonian Windyfield chert.[28] The oldest known flying insects are from the mid-Carboniferous, around 328–324 million years ago. The group subsequently underwent a rapid explosive diversification. Claims that they originated substantially earlier, during the Silurian or Devonian (some 400 million years ago) based on molecular clock estimates, are unlikely to be correct, given the fossil record.[29]

Four large-scale radiations of insects have occurred: beetles (from about 300 million years ago), flies (from about 250 million years ago), moths and wasps (both from about 150 million years ago).[30]

The remarkably successful Hymenoptera (wasps, bees, and ants) appeared some 200 million years ago in the Triassic period, but achieved their wide diversity more recently in the Cenozoic era, which began 66 million years ago. Some highly successful insect groups evolved in conjunction with flowering plants, a powerful illustration of coevolution. Insects were among the earliest terrestrial herbivores and acted as major selection agents on plants.[31] 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.[32]

Morphology and physiology


Insect morphology
A- Head B- Thorax C- Abdomen
  1. antenna
  2. ocellus (lower)
  3. ocellus (upper)
  4. compound eye
  5. brain (cerebral ganglia)
  6. prothorax
  7. dorsal blood vessel
  8. tracheal tubes (trunk with spiracle)
  9. mesothorax
  10. metathorax
  11. forewing
  12. hindwing
  13. midgut (stomach)
  14. dorsal tube (heart)
  15. ovary
  16. hindgut (intestine, rectum, anus)
  17. anus
  18. oviduct
  19. nerve cord (abdominal ganglia)
  20. Malpighian tubules
  21. tarsal pads
  22. claws
  23. tarsus
  24. tibia
  25. femur
  26. trochanter
  27. foregut (crop, gizzard)
  28. thoracic ganglion
  29. coxa
  30. salivary gland
  31. subesophageal ganglion
  32. mouthparts

Three-part body

Insects have segmented bodies supported by exoskeletons, the hard outer covering made mostly of chitin. The segments of the body are organized into three interconnected units: the head, thorax and abdomen. The head supports a pair of sensory antennae, a pair of compound eyes, zero to three simple eyes (or ocelli) and three sets of variously modified appendages that form the mouthparts. The thorax is made up of three segments: the prothorax, mesothorax and the metathorax. Each thoracic segment supports one pair of legs. The meso- and metathoracic segments may each have a pair of wings. The abdomen consists of eleven segments, sometimes fused together or reduced in size. The abdomen contains most of the digestive, respiratory, excretory and reproductive structures.[8]


The head is enclosed in a hard, heavily sclerotized, unsegmented head capsule, which contains most of the sensing organs, including the antennae, compound eyes, ocelli, and mouthparts.[34] The thorax is composed of three sections named (from front to back) the prothorax, mesothorax and metathorax. The prothorax carries the first pair of legs and the pronotum. The mesothorax carries the second pair of legs and the front wings. The metathorax carries the third pair of legs and the hind wings.[8][34] The abdomen is the largest part of the insect, typically with 11–12 segments, and is less strongly sclerotized than the head or thorax. Each segment of the abdomen has a sclerotized tergum and sternum, connected to adjacent sclerotized parts by membranes. Each segment carries a pair of spiracles.[34]


The outer skeleton, the cuticle, is made up of two layers: the epicuticle, a thin and waxy water-resistant outer layer without chitin, and a lower layer, the thick chitinous procuticle. The procuticle has two layers: an outer exocuticle and an inner endocuticle. The tough and flexible endocuticle is built from numerous layers of fibrous chitin and proteins, criss-crossing each other in a sandwich pattern, while the exocuticle is rigid and sclerotized.[35][36] As an adaptation to life on land, insects have evolved the enzyme multicopper oxidase-2 (MCO2). It uses atmospheric oxygen to harden their cuticle, unlike crustaceans which use heavy calcium compounds for the same purpose. This makes the insect exoskeleton a lightweight material.[37]


Nervous system

The nervous system of an insect consists of a brain and a ventral nerve cord. The head capsule is made up of six fused segments, each with either a pair of ganglia, or a cluster of nerve cells outside of the brain. The first three pairs of ganglia are fused into the brain, while the three following pairs are fused into a structure of three pairs of ganglia under the insect's esophagus, called the subesophageal ganglion.[38] The thoracic segments have one ganglion on each side, connected into a pair per segment. This arrangement is also seen in the first eight segments of the abdomen. Many insects have fewer ganglia than this.[39] Insects are capable of learning.[40]

Digestive system

An insect uses its digestive system to extract nutrients and other substances from the food it consumes.[41] There is extensive variation among different orders, life stages, and even castes in the digestive system of insects.[42] The gut runs lengthwise through the body. It has three sections, with paired salivary glands and salivary reservoirs.[43] By moving its mouthparts the insect mixes its food with saliva.[44][45] Some insects, like flies, expel digestive enzymes onto their food to break it down, but most insects digest their food in the gut.[46] The foregut is lined with cuticule as protection from tough food. It includes the mouth, pharynx, and crop which stores food.[47] Digestion starts in the mouth with enzymes in the saliva. Strong muscles in the pharynx pump fluid into the mouth, lubricating the food, and enabling certain insects to feed on blood or from the xylem and phloem vessels of plants.[48] Once food leaves the crop, it passes to the midgut, where the majority of digestion takes place. Microscopic projections, microvilli, increase the surface area of the wall to absorb nutrients.[49] In the hindgut, undigested food particles are joined by uric acid to form fecal pellets; most of the water is absorbed, leaving a dry pellet to be eliminated. Insects may have one to hundreds of Malpighian tubules. These remove nitrogenous wastes from the hemolymph of the insect and regulate osmotic balance. Wastes and solutes are emptied directly into the alimentary canal, at the junction between the midgut and hindgut.[50]

Reproductive system

The reproductive system of female insects consist of a pair of ovaries, accessory glands, one or more spermathecae, and ducts connecting these parts. The ovaries are made up of a variable number of egg tubes, ovarioles. Female insects make eggs, receive and store sperm, manipulate sperm from different males, and lay eggs. Accessory glands or glandular parts of the oviducts produce substances for sperm maintenance, transport and fertilization, as well as to protect the eggs. They can produce glue and protective substances for coating eggs or tough coverings for a batch of eggs called oothecae. Spermathecae are tubes or sacs in which sperm are stored between mating and fertilization.[51]

For males, the reproductive system is the testis, suspended in the body cavity by tracheae and the fat body. Most male insects have a pair of testes, containing sperm tubes or follicles enclosed in 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 to maintain the sperm. The ejaculatory duct is derived from 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.[52]

Respiratory system

The tube-like heart (green) of the mosquito Anopheles gambiae extends horizontally across the body, interlinked with the diamond-shaped wing muscles (also green) and surrounded by pericardial cells (red). Blue depicts cell nuclei.

Insect respiration is accomplished without lungs. Instead, insects have a system of internal tubes and sacs through which gases either diffuse or are actively pumped, delivering oxygen directly to tissues that need it via their tracheae. In most insects, air is taken in through paired spiracles, openings on the sides of the abdomen and thorax. The respiratory system limits the size of insects. As insects get larger, gas exchange via spiracles becomes less efficient, and thus the heaviest insect currently weighs less than 100 g. However, with increased atmospheric oxygen levels, as were present in the late Paleozoic, larger insects were possible, such as dragonflies with wingspans of more than two feet (60 cm).[53] Gas exchange patterns in insects range from continuous and diffusive ventilation, to discontinuous.[54][55][56][57]

Circulatory system

Because oxygen is delivered directly to tissues via tracheoles, the circulatory system is not used to carry oxygen, and is therefore greatly reduced. The insect circulatory system is open; it has no veins or arteries, and instead consists of little more than a single, perforated dorsal tube that pulses peristaltically. This dorsal blood vessel is divided into two sections: the heart and aorta. The dorsal blood vessel circulates the hemolymph, arthropods' fluid analog of blood, from the rear of the body cavity forward.[58][59] Hemolymph is composed of plasma in which hemocytes are suspended. Nutrients, hormones, wastes, and other substances are transported throughout the insect body in the hemolymph. Hemocytes include many types of cells that are important for immune responses, wound healing, and other functions. Hemolymph pressure may be increased by muscle contractions or by swallowing air into the digestive system to aid in molting.[60]

Reproduction and development


Butterflies mating

The majority of insects hatch from eggs. The fertilization and development takes place inside the egg, enclosed by a shell (chorion) that consists of maternal tissue. In contrast to eggs of other arthropods, most insect eggs are drought resistant. This is because inside the chorion two additional membranes develop from embryonic tissue, the amnion and the serosa. This serosa secretes a cuticle rich in chitin that protects the embryo against desiccation.[61] 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.[62] 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.[63] Some insects, like parasitoid wasps, are polyembryonic, meaning that a single fertilized egg divides into many separate embryos.[64] Insects may be univoltine, bivoltine or multivoltine, i.e. they may have one, two or many broods in a year.[65]

Aphid giving birth to live female young by parthenogenesis from unfertilized eggs

Other developmental and reproductive variations include haplodiploidy, polymorphism, paedomorphosis or peramorphosis, sexual dimorphism, parthenogenesis, and more rarely hermaphroditism.[66][67] In haplodiploidy, which is a type of sex-determination system, the offspring's sex is determined by the number of sets of chromosomes an individual receives. This system is typical in bees and wasps.[68]

Some insects are parthenogenetic, meaning that the female can reproduce and give birth without having the eggs fertilized by a male. Many aphids undergo a cyclical form of parthenogenesis in which they alternate between one or many generations of asexual and sexual reproduction.[69][70] 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 diploid females spawn many females and a few haploid males.[62]

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.[71] Still other insects have evolved mechanisms of diapause that allow eggs or pupae to survive harsh conditions.[72]


Metamorphosis in insects is the process of development that converts young to adults. There are two forms of metamorphosis: incomplete and complete.


Incomplete metamorphosis in a locust with multiple instar nymphs. Egg is not shown. The largest specimen is adult.

Hemimetabolous insects, those with incomplete metamorphosis, change gradually by undergoing a series of molts through stages called nymphs. 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 inside the old one. After this new epicuticle is secreted, the epidermis releases a mixture of enzymes that digests the endocuticle and thus detaches the old cuticle. When this stage is complete, the insect makes its body swell by taking in a large quantity of water or air, which makes the old cuticle split along predefined weaknesses where the old exocuticle was thinnest.[73][74]


Life-cycle of butterfly, undergoing complete metamorphosis from egg through caterpillar larvae to pupa and adult

Holometabolism, or complete metamorphosis, is where the insect changes in four stages, an egg or embryo, a larva, a pupa and the adult or imago. In these species, an egg hatches to produce a larva, which is generally worm-like in form. This can be 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. 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.[75] Insects undergo considerable change in form during the pupal stage, and emerge as adults. Butterflies are well-known for undergoing complete metamorphosis; most insects use this life cycle. Some insects have evolved this system to hypermetamorphosis. Complete metamorphosis is a trait of the most diverse insect group, the Endopterygota.[66]

Senses and communication

Many insects possess sensitive and specialized organs of perception. Some insects such as bees can perceive ultraviolet wavelengths, or detect polarized light, while the antennae of male moths can detect the pheromones of female moths over distances of many kilometers.[76] There is a trade-off between visual acuity and chemical or tactile acuity, such that most insects with well-developed eyes have reduced or simple antennae, and vice versa. 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. Most insects can hear only a narrow range of frequencies related to the frequency of the sounds they can produce. Mosquitoes can hear up to 2 kHz, and some grasshoppers can hear up to 50 kHz.[77] 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 bats, which helps them avoid predation.[78]

Light production and vision

Most insects have a pair of large compound eyes.

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.[79] 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.[80] The colors of emitted light vary from dull blue (Orfelia fultoni, Mycetophilidae) to the familiar greens and the rare reds (Phrixothrix tiemanni, Phengodidae).[81]

Most insects, except some cave crickets, are able to perceive light and dark. Many species have acute vision capable of detecting minute movements. The eyes may include simple eyes or ocelli as well as compound eyes of varying sizes. Many species can detect light in the infrared, ultraviolet and 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, some 400 million years ago.[82]

The individual lenses in compound eyes are immobile, and it was therefore presumed that insects were not able to focus. However, fruit flies have photoreceptor cells underneath each lens which move rapidly in and out of focus, in a series of movements called photoreceptor microsaccades. This gives them, and possibly many other insects, a much clearer image of the world than previously assumed.[83]

Sound production and hearing

Insects were the earliest organisms to produce and sense sounds. Hearing has evolved independently at least 19 times in different insect groups.[84] Insects make sounds mostly by mechanical action of appendages. In grasshoppers and crickets, this is achieved by stridulation. Cicadas make the loudest sounds among the insects by producing and amplifying sounds with special modifications to their body to form tymbals and associated musculature. The African cicada Brevisana brevis has been measured at 106.7 decibels at a distance of 50 cm (20 in).[85] Some insects, such as the Helicoverpa zea moths, hawk moths and Hedylid butterflies, can hear ultrasound and take evasive action when they sense that they have been detected by bats.[86][87] Some moths produce ultrasonic clicks that warn predatory bats of their unpalatability (acoustic aposematism),[88] while some palatable moths have evolved to mimic these calls (acoustic Batesian mimicry).[89] More recently, the claim that some moths can jam bat sonar has been revisited. Ultrasonic recording and high-speed infrared videography of bat-moth interactions suggest the palatable tiger moth really does defend against attacking big brown bats using ultrasonic clicks that jam bat sonar.[90]

Very low sounds are produced in various species of Coleoptera, Hymenoptera, Lepidoptera, Mantodea and Neuroptera. These low sounds are produced by the insect's movement, amplified by stridulatory structures on the insect's muscles and joints; these sounds can be used to warn or communicate with other insects. Most sound-making insects also have tympanal organs that can perceive airborne sounds. Some hemipterans, such as the water boatmen, communicate via underwater sounds.[91]

Cricket in garage with familiar call

Communication using surface-borne vibrational signals is more widespread among insects because of size constraints in producing air-borne sounds.[92] 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.[93]

Some species use vibrations for communicating, such as to attract mates as in the songs of the shield bug Nezara viridula.[94] Vibrations can also be used to communicate between species; lycaenid caterpillars, which form a mutualistic association with ants communicate with ants in this way.[95] The Madagascar hissing cockroach has the ability to press air through its spiracles to make a hissing noise as a sign of aggression;[96] the death's-head hawkmoth makes a squeaking noise by forcing air out of their pharynx when agitated, which may also reduce aggressive worker honey bee behavior when the two are close.[97]

Chemical communication

Social insects such as ants have multiple types of pheromonal glands, producing different semiochemicals for communication with other insects.[98]

Many insects have evolved chemical means for communication. These semiochemicals are often derived from plant metabolites including those meant to attract, repel and provide other kinds of information. Pheromones are used for attracting mates of the opposite sex, for aggregating conspecific individuals of both sexes, for deterring other individuals from approaching, to mark a trail, and to trigger aggression in nearby individuals. Allomones benefit their producer by the effect they have upon the receiver. Kairomones 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-developed in social insects.[98] Cuticular hydrocarbons are nonstructural materials produced and secreted to the cuticle surface to fight desiccation and pathogens. They are important, too, as pheromones, especially in social insects.[99]

Social behavior

A cathedral mound created by eusocial termites, which live underground.
Honey bee's figure-eight waggle dance. An orientation 45° to the right of ‘up' on the comb indicates food 45° to the right of the sun. The dancer's rapid waggling blurs her abdomen.

Social insects, such as termites, ants and many bees and wasps, are eusocial.[100] They live together in such large well-organized colonies of genetically similar individuals that they are sometimes considered superorganisms. Honey bees have evolved a system of abstract symbolic communication where a behavior is used to represent and convey specific information about the environment. In this communication system, called dance language, the angle at which a bee dances represents a direction relative to the sun, and the length of the dance represents the distance to be flown.[101] Bumblebees too have some social communication behaviors. Bombus terrestris, for example, more rapidly learns about visiting unfamiliar, yet rewarding flowers, when they can see a conspecific foraging on the same species.[102]

Only insects that live in nests or colonies possess fine-scale spatial orientation. Some can return unerringly to a single hole a few millimeters in diameter among thousands of similar holes, after a trip of several kilometers. In philopatry, insects that hibernate are able to recall a specific location up to a year after last viewing the area of interest.[103] A few insects seasonally migrate large distances between different geographic regions (e.g., the overwintering areas of the monarch butterfly).[104]

Care of young

Eusocial insects build nests, guard eggs, and provide food for offspring full-time. Most insects, however, lead short lives as adults, and rarely interact with one another except to mate or compete for mates. A small number provide parental care, where they at least guard their eggs, and sometimes guard their offspring until adulthood, possibly even feeding them. Many wasps and bees construct a nest or burrow, store provisions in it, and lay an egg upon those provisions, providing no further care.[105]



Insects such as hoverflies are capable of rapid and agile flight.

Insects are the only group of invertebrates to have developed flight. The evolution of insect wings has been a subject of debate. Some entomologists 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. Another theory suggest the wings are actually made up of both the notum and the pleuron.[106] 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.[107] In the Carboniferous age, some of the Meganeura dragonflies had as much as a 50 cm (20 in) wide wingspan. The appearance of gigantic insects is consistent with high atmospheric oxygen at that time, as the respiratory system of insects constrains their size.[108] The largest flying insects today are much smaller, with the largest wingspan belonging to the white witch moth (Thysania agrippina), at approximately 28 cm (11 in).[109]

Unlike birds, many small insects are swept along by the prevailing winds[110] although many of the larger insects are known to make migrations. Aphids are transported long distances by low-level jet streams.[111]


Spatial and temporal stepping pattern of walking desert ants performing an alternating tripod gait. Recording rate: 500 fps, Playback rate: 10 fps.

Many adult insects use six legs for walking, with an alternating tripod gait. This allows for rapid walking while always having a stable stance; it has been studied extensively in cockroaches and ants. For the first step, the middle right leg and the front and rear left legs are in contact with the ground and move the insect forward, while the front and rear right leg and the middle left leg are lifted and moved forward to a new 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.[112] 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, avoiding obstacles, climbing or slippery surfaces, four (tetrapodal) or more feet (wave-gait) may be touching the ground.[113] Cockroaches are among the fastest insect runners and, at full speed, adopt a bipedal run. More sedate locomotion is seen in the well-camouflaged stick insects (Phasmatodea). A few insects such 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.[114]


The backswimmer Notonecta glauca underwater, showing its paddle-like hindleg adaptation

A large number of insects live either part or the whole of their lives underwater. In many of the more primitive orders of insect, the immature stages are aquatic. Some groups of insects, like certain water beetles, have aquatic adults as well.[56]

Many of these species are adapted for under-water locomotion. Water beetles and water bugs have legs adapted into paddle-like structures. Dragonfly naiads use jet propulsion, forcibly expelling water out of their rectal chamber.[115] Some species like the water striders 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.[56] Other insects such as the rove beetle Stenus emit pygidial gland secretions that reduce surface tension; this enables them to move on the surface of water by Marangoni propulsion.[116][117]


Insect ecology is the scientific study of how insects, individually or as a community, interact with the surrounding environment or ecosystem.[118] Insects play one of the most important roles in their ecosystems, which includes many roles, such as soil turning and aeration, dung burial, pest control, pollination and wildlife nutrition. An example is the beetles, which are scavengers that feed on dead animals and fallen trees and thereby recycle biological materials into forms found useful by other organisms.[119] These insects, and others, are responsible for much of the process by which topsoil is created.[120]


Reduvius personatus, the masked hunter bug nymph, camouflages itself with sand grains to avoid predators.

Insects are mostly small, soft bodied, and fragile compared to larger lifeforms. The immature stages are small, move slowly or are immobile, and so all stages are exposed to predation and parasitism. Insects accordingly employ multiple defense strategies, including camouflage, mimicry, toxicity and active defense.[121]

Camouflage is an important defense strategy, which involves the use of coloration or shape to avoid being noticed.[122] It is common among beetles, especially those that feed on wood or vegetation, such as leaf beetles and weevils. In some of these species, sculpturing or various colored scales or hairs cause the beetle to mimic bird dung or other inedible objects.[121] Phasmids mimic the forms of sticks and leaves; the bodies of some species (such as Palophus centaurus) are covered in mossy outgrowths that supplement their disguise. A few species can change color as their surroundings shift (as in Bostra scabrinota). Some mantises sway from side to side, like leaves or twigs swaying in the breeze. Other insects feign death.[123]

Many insects use mimicry. In Batesian mimicry, edible species, such as of butterflies, longhorn beetles, and hoverflies (the mimics), gain a survival advantage by resembling inedible species (the models).[121][124] In Müllerian mimicry, inedible species, such as of wasps and bees, resemble each other so as to reduce the sampling rate by predators who need to learn about the insects' inedibility. Taxa from the toxic genus Heliconius form one of the most well known Müllerian complexes.[125]

Chemical defense is common among the Coleoptera and Lepidoptera, usually being advertised by bright warning colors (aposematism), such as in 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 the basis of Müllerian mimicry. A predator who has previously eaten a poisonous lepidopteran may avoid other species with similar markings in the future.[126] Some ground beetles of the family Carabidae can spray chemicals from their abdomen with great accuracy, to repel predators.[121]


European honey bee carrying pollen in a pollen basket back to the hive

Pollination is the process by which pollen is transferred in the reproduction of plants, thereby enabling fertilisation and sexual reproduction.[127] Most flowering plants require an animal to do the transportation. The majority of pollination is by insects.[128] Because insects usually receive benefit for the pollination in the form of energy rich nectar it is a mutualism. The various flower traits, such as bright colors and pheromones that coevolved with their pollinators, have been called pollination syndromes, though around one third of flowers cannot be assigned to a single syndrome.[129]


Many insects are parasitic. The largest group, with over 100,000 species[130] and perhaps over a million,[131] consists of a single clade of parasitoid wasps among the Hymenoptera.[132] These are parasites of other insects, eventually killing their hosts.[130] Some are hyper-parasites, as their hosts are other parasitoid wasps.[130][133] Several groups of insects can be considered as either micro-predators or external parasites;[134][135] for example, many hemipteran bugs have piercing and sucking mouthparts, adapted for feeding on plant sap,[136][137] while species in groups such as fleas, lice, and mosquitoes are hematophagous, feeding on the blood of animals.[135]

Relationship to humans

As pests

Aedes aegypti, a parasite, is the vector of dengue fever and yellow fever

Many insects are considered pests by humans. These include parasites of people and livestock, such as lice and bed bugs; many parasites such as mosquitoes also transmit diseases. Other pests include insects like termites that damage wooden structures; herbivorous insects such as locusts, aphids, and thrips that destroy agricultural crops, or like wheat weevils damage stored agricultural produce. Farmers have often attempted to control insects with chemical insecticides, but increasingly rely on biological pest control. This uses one organism to reduce the population density of a pest organism; it is a key element of integrated pest management.[139][140] Biological control is favored because insecticides can cause harm to ecosystems far beyond the intended pest targets.[141][142]

In beneficial roles

Silkworms were domesticated for silk for over 5000 years.[143][144] Here, silk cocoons are being unrolled.

Pollination of flowering plants by insects including bees, butterflies, flies, and beetles, is economically important.[145] The value of insect pollination of crops and fruit trees was estimated in 2021 to be about $34 billion in the US alone.[146]

Insects produce useful substances such as honey,[147] wax,[148][149] lacquer[150] and silk.[151] Honey bees have been cultured by humans for thousands of years for honey.[152] Beekeeping in pottery vessels began about 9,000 years ago in North Africa.[153] The silkworm has greatly affected human history, as silk-driven trade established relationships between China and the rest of the world.[154][155]

Insects that feed on or parasitise other insects are beneficial to humans if they thereby reduce damage to agriculture and human structures. For example, aphids feed on crops, causing economic loss, but ladybugs feed on aphids, and can be used to control them. Insects account for the vast majority of insect consumption.[156][157][158] Ants may help control animal populations by consuming small vertebrates.[159]

Fly larvae (maggots) were formerly used to treat wounds to prevent or stop gangrene, as they would only consume dead flesh. This treatment is finding modern usage in some hospitals. Recently insects have also gained attention as potential sources of drugs and other medicinal substances.[160] Adult insects, such as crickets and insect larvae of various kinds, are also commonly used as fishing bait.[161]

Population declines

At least 66 insect species extinctions have been recorded since 1500, many of them on oceanic islands.[162] Declines in insect abundance have been attributed to human activity in the form of artificial lighting,[163] land use changes such as urbanization or farming,[164][165] pesticide use,[166] and invasive species.[167] A 2019 research review suggested that a large proportion of insect species is threatened with extinction in the 21st century,[168] though the details have been disputed.[169] A larger 2020 meta-study, analyzing data from 166 long-term surveys, suggested that populations of terrestrial insects are indeed decreasing rapidly, by about 9% per decade.[170][171]

In research

The fruit fly Drosophila melanogaster is a widely used model organism.

Insects play important roles in biological research. For example, because of its small size, short generation time and high fecundity, the common fruit fly Drosophila melanogaster is a model organism for studies in the genetics of eukaryotes. 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.[172] The genome of D. melanogaster was sequenced in 2000, reflecting the organism's important role in biological research. It was found that 70% of the fly genome is similar to the human genome, supporting the evolution theory.[173]

As food

Witchetty grubs are prized as high-protein foods by Aboriginal Australians.[174]

Insects are consumed as food in 80% of the world's nations, by people in roughly 3000 ethnic groups.[175][176] In Africa, locally abundant species of locusts and termites are a common traditional human food source.[177] Some, especially deep-fried cicadas, are considered to be delicacies. Insects have a high protein content for their mass, and some authors suggest their potential as a major source of protein in human nutrition.[178] In most first-world countries, however, entomophagy (the eating of insects), is taboo.[179] They are also recommended by armed forces as a survival food for troops in adversity.[177] Because of the abundance of insects and a worldwide concern of food shortages, the Food and Agriculture Organization of the United Nations considers that people throughout the world may have to eat insects as a food staple. Insects are noted for their nutrients, having a high content of protein, minerals and fats and are already regularly eaten by one-third of the world's population.[180]

In other products

Black soldier fly larvae can provide protein and fats for use in cosmetics.[181] Insect cooking oil, insect butter and fatty alcohols can be made from such insects as the superworm (Zophobas morio).[182] Several insect species such as the black soldier fly or the housefly in their maggot forms, as well as beetle larvae such as mealworms can be processed and used as feed for farmed animals such as chicken, fish and pigs.[183] Many species of insects are sold and kept as pets.[184]

In religion and folklore

Ancient Egyptian scarab with separate wings, c. 712-342 BC

Scarab beetles held religious and cultural symbolism in ancient Egypt, Greece and some shamanistic Old World cultures. The ancient Chinese regarded cicadas as symbols of rebirth or immortality. In Mesopotamian literature, the epic poem of Gilgamesh has allusions to Odonata that signify the impossibility of immortality. Among the Aborigines of Australia of the Arrernte language groups, honey ants and witchetty grubs served as personal clan totems. In the case of the 'San' bush-men of the Kalahari, it is the praying mantis that holds much cultural significance including creation and zen-like patience in waiting.[185]

See also


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


  1. ^ a b Harper, Douglas; McCormack, Dan (November 2001). "Online Etymological Dictionary". p. 1. Archived from the original on 11 January 2012. Retrieved 1 November 2011.
  2. ^ Lewis, Charlton T.; Short, Charles (1879). "insĕco". A Latin Dictionary. Perseus Digital Library.
  3. ^ Liddell, Henry George; Scott, Robert (1940). "ἔντομος". A Greek-English Lexicon. Perseus Digital Library.
  4. ^ "insect translations".
  5. ^ "What is a bug? Insects, arachnids, and myriapods" at Museum of New Zealand Te Papa Tongarewa website. Accessed 10 March 2022.
  6. ^ a b Waldbauer, Gilbert (1998). The Handy Bug Answer Book. Visible Ink. pp. 1, 5–26. ISBN 9781578590490.
  7. ^ Chinery, Michael (1993). "Introduction". Insects of Britain & Northern Europe (3rd ed.). London: HarperCollins. pp. 11–13. ISBN 978-0-00-219918-6.
  8. ^ a b c Gullan & Cranston 2005, pp. 22–48.
  9. ^ a b Stork, Nigel E. (7 January 2018). "How Many Species of Insects and Other Terrestrial Arthropods Are There on Earth?". Annual Review of Entomology. 63 (1): 31–45. doi:10.1146/annurev-ento-020117-043348. PMID 28938083. S2CID 23755007.
  10. ^ Chapman, A. D. (2006). Numbers of living species in Australia and the World. Canberra: Australian Biological Resources Study. ISBN 978-0-642-56850-2. Archived from the original on 30 November 2012.
  11. ^ Wilson, E. O. "Threats to Global Diversity". Archived from the original on 20 February 2015. Retrieved 17 May 2009.
  12. ^ Novotny, Vojtech; Basset, Yves; Miller, Scott E.; Weiblen, George D.; Bremer, Birgitta; Cizek, Lukas; Drozd, Pavel (2002). "Low host specificity of herbivorous insects in a tropical forest". Nature. 416 (6883): 841–844. Bibcode:2002Natur.416..841N. doi:10.1038/416841a. PMID 11976681. S2CID 74583.
  13. ^ Erwin, Terry L. (1997). Biodiversity at its utmost: Tropical Forest Beetles (PDF). pp. 27–40. Archived (PDF) from the original on 9 November 2018. Retrieved 16 December 2017. In: Reaka-Kudla, M.L.; Wilson, D. E.; Wilson, E. O., eds. (1997). Biodiversity II. Joseph Henry Press, Washington, D.C. ISBN 9780309052276.
  14. ^ Erwin, Terry L. (1982). "Tropical forests: their richness in Coleoptera and other arthropod species" (PDF). The Coleopterists Bulletin. 36: 74–75. Archived (PDF) from the original on 23 September 2015. Retrieved 16 September 2018.
  15. ^ Misof, Bernhard; et al. (7 November 2014). "Phylogenomics resolves the timing and pattern of insect evolution". Science. 346 (6210): 763–767. Bibcode:2014Sci...346..763M. doi:10.1126/science.1257570. PMID 25378627. S2CID 36008925. Archived from the original on 18 October 2009. Retrieved 17 October 2009.
  16. ^ Kjer, Karl M.; Simon, Chris; Yavorskaya, Margarita; Beutel, Rolf G. (2016). "Progress, pitfalls and parallel universes: a history of insect phylogenetics". Journal of the Royal Society Interface. 13 (121): 121. doi:10.1098/rsif.2016.0363. PMC 5014063. PMID 27558853.
  17. ^ Sroka, Günter; Staniczek, Arnold H.; Bechly (December 2014). "Revision of the giant pterygote insect Bojophlebia prokopi Kukalová-Peck 1985 (Hydropalaeoptera: Bojophlebiidae) from the Carboniferous of the Czech Republic, with the first cladistic analysis of fossil palaeopterous insects". Journal of Systematic Palaeontology. 13 (11): 963–982. doi:10.1080/14772019.2014.987958. S2CID 84037275. Retrieved 21 May 2019.
  18. ^ Prokop, Jakub (2017). "Redefining the extinct orders Miomoptera & Hypoperlida as stem acercarian insects". BMC Evolutionary Biology. 17 (1): 205. doi:10.1186/s12862-017-1039-3. PMC 5574135. PMID 28841819.
  19. ^ Wipfler, B. (February 2019). "Evolutionary history of Polyneoptera & its implications for our understanding of early winged insects". Proceedings of the National Academy of Sciences. 116 (8): 3024–3029. Bibcode:2019PNAS..116.3024W. doi:10.1073/pnas.1817794116. PMC 6386694. PMID 30642969.
  20. ^ P. J. Gullan; P. S. Cranston (2010). "Taxobox 17 – Psocodea: "Psocoptera" (bark lice and book lice)". The Insects: an Outline of Entomology (4th ed.). John Wiley & Sons. p. 479. ISBN 9781444317671.
  21. ^ Gullan & Cranston 2005, p. 180.
  22. ^ Kendall, David A. (2009). "Classification of Bugs". Archived from the original on 20 May 2009. Retrieved 9 May 2009.
  23. ^ Blanke, Alexander; Machida, Ryuichiro; Szucsich, Nikolaus Urban; Wilde, Fabian; Misoe, Bernhard (15 October 2014). "Mandibles with two joints evolved much earlier in the history of insects: dicondyly is a synapomorphy of bristletails, silverfish and winged insects". Systematic Entomology. Wiley. 40 (2): 357–364. doi:10.1111/syen.12107. ISSN 0307-6970.
  24. ^ Johnson, K. P.; Yoshizawa, K.; Smith, V. S. (2004). "Multiple origins of parasitism in lice". Proceedings of the Royal Society of London. 271 (1550): 1771–1776. doi:10.1098/rspb.2004.2798. PMC 1691793. PMID 15315891.
  25. ^ Terry, M. D.; Whiting, M. F. (2005). "Mantophasmatodea and phylogeny of the lower neopterous insects". Cladistics. 21 (3): 240–257. doi:10.1111/j.1096-0031.2005.00062.x. S2CID 86259809.
  26. ^ Lo, Nathan; Tokuda, Gaku; Watanabe, Hirofumi; et al. (2000). "Evidence from multiple gene sequences indicates that termites evolved from wood-feeding cockroaches". Current Biology. 10 (13): 801–804. doi:10.1016/S0960-9822(00)00561-3. PMID 10898984. S2CID 14059547.
  27. ^ Whiting, M. F. (2002). "Mecoptera is paraphyletic: multiple genes and phylogeny of Mecoptera and Siphonaptera". Zoologica Scripta. 31 (1): 93–104. doi:10.1046/j.0300-3256.2001.00095.x. S2CID 56100681.
  28. ^ Ross, Andrew (August 2022). "Evolution: The origin of insect wings revisited". Current Biology. 32 (15): R851–R853. doi:10.1016/j.cub.2022.06.087. PMID 35944489. S2CID 251464185.
  29. ^ Schachat, Sandra R.; Goldstein, Paul Z.; Desalle, Rob; Bobo, Dean M.; Boyce, C. Kevin; Payne, Jonathan L.; Labandeira, Conrad C. (2 February 2023). "Illusion of flight? Absence, evidence and the age of winged insects". Biological Journal of the Linnean Society. 138 (2): 143–168. doi:10.1093/biolinnean/blac137. ISSN 0024-4066.
  30. ^ Wiegmann, Brian M.; Trautwein, Michelle D.; Winkler, Isaac S.; Barr, Norman B.; Kim, Jung-Wook; et al. (14 March 2011). "Episodic radiations in the fly tree of life". Proceedings of the National Academy of Sciences. 108 (14): 5690–5695. Bibcode:2011PNAS..108.5690W. doi:10.1073/pnas.1012675108. PMC 3078341. PMID 21402926.
  31. ^ Carter, J. Stein (29 March 2005). "Coevolution and Pollination". University of Cincinnati. Archived from the original on 30 April 2009. Retrieved 9 May 2009.
  32. ^ "Coevolution and Pollination". University of Cincinnati. Archived from the original on 30 April 2009. Retrieved 9 May 2009.
  33. ^ Taylor, Paul D.; Lewis, David N. (2007). Fossil Invertebrates (repeated ed.). Harvard University Press. p. 160. ISBN 978-0674025745.
  34. ^ a b c Resh & Carde 2009, p. 13.
  35. ^ Gullan & Cranston 2005, p. 22–24.
  36. ^ Belles, Xavier (14 October 2019). "The innovation of the final moult and the origin of insect metamorphosis". Philosophical Transactions of the Royal Society B: Biological Sciences. 374 (1783): 20180415. doi:10.1098/rstb.2018.0415. PMC 6711288. PMID 31438822.
  37. ^ Asano, Tsunaki; Hashimoto, Kosei; Everroad, R. Craig (2023). "Eco-evolutionary implications for a possible contribution of cuticle hardening system in insect evolution and terrestrialisation". Physiological Entomology. 48 (2–3): 55–60. doi:10.1111/phen.12406. S2CID 258209514.
  38. ^ Gullan & Cranston 2005, p. 57.
  39. ^ Schneiderman, Howard A. (1960). "Discontinuous respiration in insects: role of the spiracles". The Biological Bulletin. 119 (3): 494–528. doi:10.2307/1539265. JSTOR 1539265. Archived from the original on 25 June 2009. Retrieved 22 May 2009.
  40. ^ Dukas, Reuven (1 January 2008). "Evolutionary Biology of Insect Learning". Annual Review of Entomology. 53 (1): 145–160. doi:10.1146/annurev.ento.53.103106.093343. PMID 17803459.
  41. ^ "General Entomology – Digestive and Excritory system". NC state University. Archived from the original on 23 May 2009. Retrieved 3 May 2009.
  42. ^ Bueno, Odair Correa; Tanaka, Francisco André Ossamu; de Lima Nogueira, Neusa; Fox, Eduardo Gonçalves Paterson; Rossi, Mônica Lanzoni; Solis, Daniel Russ (1 January 2013). "On the morphology of the digestive system of two Monomorium ant species". Journal of Insect Science. 13 (1): 70. doi:10.1673/031.013.7001. PMC 3835044. PMID 24224520.
  43. ^ Gullan & Cranston 2005, pp. 70–77.
  44. ^ "General Entomology – Digestive and Excretory system". North Carolina State University. Archived from the original on 23 May 2009. Retrieved 3 May 2009.
  45. ^ Duncan, Carl D. (1939). A Contribution to The Biology of North American Vespine Wasps (1st ed.). Stanford: Stanford University Press. pp. 24–29.
  46. ^ Nation 2001, p. 31.
  47. ^ Gullan & Cranston 2005, p. 70.
  48. ^ Nation 2001, p. 30–31.
  49. ^ Nation 2001, p. 32.
  50. ^ Gullan & Cranston 2005, pp. 71–72, 78–80.
  51. ^ Resh, Carde & 2009, p. 880.
  52. ^ Resh, Carde & 2009, p. 885.
  53. ^ "What Keeps Bugs from Being Bigger?". Argonne National Laboratory. 8 August 2007. Archived from the original on 14 May 2017. Retrieved 15 July 2013.
  54. ^ Gullan & Cranston 2005, pp. 65–68.
  55. ^ Chown, S. L.; S. W. Nicholson (2004). Insect Physiological Ecology. New York: Oxford University Press. ISBN 978-0-19-851549-4.
  56. ^ a b c Richard W. Merritt; Kenneth W. Cummins; Martin B. Berg, eds. (2007). An Introduction to the Aquatic Insects of North America (4th ed.). Kendall Hunt Publishers. ISBN 978-0-7575-5049-2.
  57. ^ Merritt, R. W.; K. W. Cummins; M. B. Berg (2007). An Introduction To The Aquatic Insects Of North America. Kendall Hunt Publishing Company. ISBN 978-0-7575-4128-5.
  58. ^ Gullan & Cranston 2005, pp. 61–65.
  59. ^ Meyer, John R. (17 February 2006). "Circulatory System". NC State University: Department of Entomology, NC State University. p. 1. Archived from the original on 27 September 2009. Retrieved 11 October 2009.
  60. ^ Triplehorn, Charles (2005). Borror and DeLong's introduction to the study of insects. Johnson, Norman F., Borror, Donald J. (7th ed.). Belmont, California: Thompson Brooks/Cole. pp. 27–28. ISBN 978-0030968358. OCLC 55793895.
  61. ^ Jacobs, C. G.; Rezende, G. L.; Lamers, G. E.; van der Zee, M. (2013). "The extraembryonic serosa protects the insect egg against desiccation". Proceedings of the Royal Society of London B. 280 (1764): 20131082. doi:10.1098/rspb.2013.1082. PMC 3712428. PMID 23782888.
  62. ^ a b "insect physiology" McGraw-Hill Encyclopedia of Science and Technology, Ch. 9, p. 233, 2007
  63. ^ Gullan & Cranston 2005, pp. 129, 131, 134–135.
  64. ^ Gullan & Cranston 2005, pp. 136–137.
  65. ^ "Glossary of Lepidopteran and Odonate anatomy". Rare species atlas. Virginia Department of Conservation and Recreation. 2013. Archived from the original on 4 October 2013. Retrieved 14 June 2013.
  66. ^ a b Gullan & Cranston 2005, p. 143.
  67. ^ Judson, Olivia (14 August 2002). Dr. Tatiana's Sex Advice to All Creation: The Definitive Guide to the Evolutionary Biology of Sex. Macmillan. p. 198. ISBN 978-0-8050-6331-8.
  68. ^ Hughes, William O. H.; Oldroyd, Benjamin P.; Beekman, Madeleine; Ratnieks, Francis L. W. (2008). "Ancestral Monogamy Shows Kin Selection Is Key to the Evolution of Eusociality". Science. 320 (5880): 1213–1216. Bibcode:2008Sci...320.1213H. doi:10.1126/science.1156108. PMID 18511689. S2CID 20388889.
  69. ^ Nevo, E.; Coll, M. (2001). "Effect of nitrogen fertilization on Aphis gossypii (Homoptera: Aphididae): variation in size, color, and reproduction". Journal of Economic Entomology. 94 (1): 27–32. doi:10.1603/0022-0493-94.1.27. PMID 11233124. S2CID 25758038.
  70. ^ Jahn, G. C.; Almazan, L .P.; Pacia, J. (2005). "Effect of nitrogen fertilizer on the intrinsic rate of increase of the rusty plum aphid, Hysteroneura setariae (Thomas) (Homoptera: Aphididae) on rice (Oryza sativa L.)" (PDF). Environmental Entomology. 34 (4): 938–943. doi:10.1603/0046-225X-34.4.938. S2CID 1941852. Archived from the original (PDF) on 9 September 2010.
  71. ^ Hadley, Debbie. "Where do insects go in winter?". Archived from the original on 18 January 2012. Retrieved 19 April 2012.
  72. ^ Lee, Richard E. Jr. (1989). "Insect Cold-Hardiness: To Freeze or Not to Freeze" (PDF). BioScience. 39 (5): 308–313. doi:10.2307/1311113. JSTOR 1311113. Archived (PDF) from the original on 10 January 2011. Retrieved 2 December 2009.
  73. ^ Gullan & Cranston 2005, p. 142.
  74. ^ Ruppert, E. E.; Fox, R. S.; Barnes, R. D. (2004). Invertebrate Zoology (7th ed.). Brooks / Cole. pp. 523–524. ISBN 978-0-03-025982-1.
  75. ^ Gullan & Cranston 2005, p. 151.
  76. ^ "Insects" (PDF). Alien Life Forms. p. 4. Archived (PDF) from the original on 8 July 2011. Retrieved 17 May 2009.
  77. ^ Cator, L.J.; Arthur, B.J.; Harrington, L.C.; Hoy, R.R. (2009). "Harmonic convergence in the love songs of the dengue vector mosquito". Science. 323 (5917): 1077–1079. Bibcode:2009Sci...323.1077C. doi:10.1126/science.1166541. PMC 2847473. PMID 19131593.
  78. ^ Gullan & Cranston 2005, pp. 87–94.
  79. ^ Pugsley, Chris W. (1983). "Literature review of the New Zealand glowworm Arachnocampa luminosa (Diptera: Keroplatidae) and related cave-dwelling Diptera" (PDF). New Zealand Entomologist. 7 (4): 419–424. doi:10.1080/00779962.1983.9722435. Archived from the original (PDF) on 20 October 2007.
  80. ^ Lloyd, James E. (1984). "Occurrence of Aggressive Mimicry in Fireflies". The Florida Entomologist. 67 (3): 368–376. doi:10.2307/3494715. JSTOR 3494715. S2CID 86502129.
  81. ^ Lloyd, James E.; Gentry, Erin C. (2003). The Encyclopedia of Insects. Academic Press. pp. 115–120. ISBN 978-0-12-586990-4.
  82. ^ Briscoe, A. D.; Chittka, L. (2001). "The evolution of color vision in insects". Annual Review of Entomology. 46: 471–510. doi:10.1146/annurev.ento.46.1.471. PMID 11112177. S2CID 20894534.
  83. ^ Kemppainen, Joni; Scales, Ben; Razban Haghighi, Keivan; Takalo, Jouni; Mansour, Neveen; et al. (22 March 2022). "Binocular mirror–symmetric microsaccadic sampling enables Drosophila hyperacute 3D vision". Proceedings of the National Academy of Sciences. 119 (12): e2109717119. Bibcode:2022PNAS..11909717K. doi:10.1073/pnas.2109717119. PMC 8944591. PMID 35298337.
  84. ^ Warren, Ben; Nowotny, Manuela (11 April 2021). "Bridging the Gap Between Mammal and Insect Ears – A Comparative and Evolutionary View of Sound-Reception". Frontiers in Ecology and Evolution. 9. doi:10.3389/fevo.2021.667218.
  85. ^ "The University of Florida Book of Insect Records". Department of Entomology & Nematology, UF/IFAS. Retrieved 13 January 2022.
  86. ^ Kay, Robert E. (1969). "Acoustic signalling and its possible relationship to assembling and navigation in the moth, Heliothis zea". Journal of Insect Physiology. 15 (6): 989–1001. doi:10.1016/0022-1910(69)90139-5.
  87. ^ Spangler, Hayward G. (1988). "Moth hearing, defense, and communication". Annual Review of Entomology. 33 (1): 59–81. doi:10.1146/annurev.ento.33.1.59.
  88. ^ Hristov, N. I.; Conner, William E. (2005). "Sound strategy: acoustic aposematism in the bat–tiger moth arms race". Naturwissenschaften. 92 (4): 164–169. Bibcode:2005NW.....92..164H. doi:10.1007/s00114-005-0611-7. PMID 15772807. S2CID 18306198.
  89. ^ Barber, J. R.; Conner, W. E. (2007). "Acoustic mimicry in a predator–prey interaction". Proceedings of the National Academy of Sciences. 104 (22): 9331–9334. Bibcode:2007PNAS..104.9331B. doi:10.1073/pnas.0703627104. PMC 1890494. PMID 17517637.
  90. ^ Corcoran, Aaron J.; Barber, Jesse R.; Conner, William E. (2009). "Tiger Moth Jams Bat Sonar". Science. 325 (5938): 325–327. Bibcode:2009Sci...325..325C. doi:10.1126/science.1174096. PMID 19608920. S2CID 206520028.
  91. ^ Theiss, Joachim (1982). "Generation and radiation of sound by stridulating water insects as exemplified by the corixids". Behavioral Ecology and Sociobiology. 10 (3): 225–235. doi:10.1007/BF00299689. S2CID 10338592.
  92. ^ Virant-Doberlet, M.; Čokl, Andrej (2004). "Vibrational communication in insects". Neotropical Entomology. 33 (2): 121–134. doi:10.1590/S1519-566X2004000200001.
  93. ^ Bennet-Clark, H. C. (1998). "Size and scale effects as constraints in insect sound communication". Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences. 353 (1367): 407–419. doi:10.1098/rstb.1998.0219. PMC 1692226.
  94. ^ Miklas, Nadège; Stritih, Nataša; Čokl, Andrej; Virant-Doberlet, Meta; Renou, Michel (2001). "The Influence of Substrate on Male Responsiveness to the Female Calling Song in Nezara viridula". Journal of Insect Behavior. 14 (3): 313–332. doi:10.1023/A:1011115111592. S2CID 11369425.
  95. ^ DeVries, P. J. (1990). "Enhancement of symbiosis between butterfly caterpillars and ants by vibrational communication". Science. 248 (4959): 1104–1106. Bibcode:1990Sci...248.1104D. doi:10.1126/science.248.4959.1104. PMID 17733373. S2CID 35812411.
  96. ^ Nelson, Margaret C.; Fraser, Jean (1980). "Sound production in the cockroach, Gromphadorhina portentosa: evidence for communication by hissing". Behavioral Ecology and Sociobiology. 6 (4): 305–314. doi:10.1007/BF00292773. S2CID 9637568.
  97. ^ Moritz, R. F. A.; Kirchner, W. H.; Crewe, R. M. (1991). "Chemical camouflage of the death's head hawkmoth (Acherontia atropos L.) in honeybee colonies". Naturwissenschaften. 78 (4): 179–182. Bibcode:1991NW.....78..179M. doi:10.1007/BF01136209. S2CID 45597312.
  98. ^ a b Gullan & Cranston 2005, pp. 96–105.
  99. ^ Yan, Hua; Liebig, Jürgen (1 April 2021). "Genetic basis of chemical communication in eusocial insects". Genes & Development. Cold Spring Harbor Laboratory Press & The Genetics Society. 35 (7–8): 470–482. doi:10.1101/gad.346965.120. PMC 8015721. PMID 33861721.
  100. ^ Brewer, Gary. "Social insects". North Dakota State University. Archived from the original on 21 March 2008. Retrieved 6 May 2009.
  101. ^ Gullan & Cranston 2005, pp. 309–311.
  102. ^ Leadbeater, E.; Chittka, L. (2007). "The dynamics of social learning in an insect model, the bumblebee (Bombus terrestris)". Behavioral Ecology and Sociobiology. 61 (11): 1789–1796. doi:10.1007/s00265-007-0412-4. S2CID 569654.
  103. ^ Salt, R. W. (1961). "Principles of Insect Cold-Hardiness". Annual Review of Entomology. 6: 55–74. doi:10.1146/annurev.en.06.010161.000415.
  104. ^ Gullan & Cranston 2005, p. 14.
  105. ^ "Social Insects". North Dakota State University. Archived from the original on 21 March 2008. Retrieved 12 October 2009.
  106. ^ The Complete Insect: Anatomy, Physiology, Evolution, and Ecology
  107. ^ Jockusch, E. L.; Ober, K. A. (September 2004). "Hypothesis testing in evolutionary developmental biology: a case study from insect wings". Journal of Heredity. 95 (5): 382–396. doi:10.1093/jhered/esh064. PMID 15388766.
  108. ^ Dudley, R (1998). "Atmospheric oxygen, giant Paleozoic insects and the evolution of aerial locomotor performance" (PDF). Journal of Experimental Biology. 201 (8): 1043–1050. doi:10.1242/jeb.201.8.1043. PMID 9510518. Archived (PDF) from the original on 24 January 2013. Retrieved 8 December 2012.
  109. ^ "Chapter 32: Largest Lepidopteran Wing Span | The University of Florida Book of Insect Records | Department of Entomology & Nematology | UF/IFAS". Retrieved 13 January 2022.
  110. ^ Yates, Diana. "Birds migrate together at night in dispersed flocks, new study indicates". Retrieved 13 January 2022.
  111. ^ Drake, V. A.; Farrow, R. A. (1988). "The Influence of Atmospheric Structure and Motions on Insect Migration". Annual Review of Entomology. 33: 183–210. doi:10.1146/annurev.en.33.010188.001151.
  112. ^ Biewener, Andrew A. (2003). Animal Locomotion. Oxford University Press. ISBN 978-0-19-850022-3.[page needed]
  113. ^ Grabowska, Martyna; Godlewska, Elzbieta; Schmidt, Joachim; Daun-Gruhn, Silvia (2012). "Quadrupedal gaits in hexapod animals – inter-leg coordination in free-walking adult stick insects". Journal of Experimental Biology. 215 (24): 4255–4266. doi:10.1242/jeb.073643. PMID 22972892.
  114. ^ Ikawa, Terumi; Okabe, Hidehiko; Hoshizaki, Sugihiko; Kamikado, Takahiro; Cheng, Lanna (2004). "Distribution of the oceanic insects Halobates (Hemiptera: Gerridae) off the south coast of Japan". Entomological Science. 7 (4): 351–357. doi:10.1111/j.1479-8298.2004.00083.x. S2CID 85017400.
  115. ^ Mill, P. J.; Pickard, R. S. (1975). "Jet-propulsion in anisopteran dragonfly larvae". Journal of Comparative Physiology A. 97 (4): 329–338. doi:10.1007/BF00631969. S2CID 45066664.
  116. ^ Linsenmair, K.; Jander, R. (1976). "Das "entspannungsschwimmen" von Velia and Stenus". Naturwissenschaften. 50 (6): 231. Bibcode:1963NW.....50..231L. doi:10.1007/BF00639292. S2CID 40832917.
  117. ^ Bush, J. W. M.; Hu, David L. Hu (2006). "Walking on Water: Biolocomotion at the Interface" (PDF). Annual Review of Fluid Mechanics. 38 (1): 339–369. Bibcode:2006AnRFM..38..339B. doi:10.1146/annurev.fluid.38.050304.092157. Archived from the original (PDF) on 10 July 2007.
  118. ^ Schowalter 2006, pp. 3, 572.
  119. ^ Losey, John E.; Vaughan, Mace (2006). "The Economic Value of Ecological Services Provided by Insects". BioScience. 56 (4): 311–323(13). doi:10.1641/0006-3568(2006)56[311:TEVOES]2.0.CO;2. Archived from the original on 12 January 2012. Retrieved 8 November 2011.
  120. ^ Gullan & Cranston 2005, pp. 3, 218–228.
  121. ^ a b c d Evans, Arthur V.; Charles Bellamy (2000). An Inordinate Fondness for Beetles. University of California Press. p. 31. ISBN 978-0-520-22323-3.
  122. ^ "Photos: Masters of Disguise – Amazing Insect Camouflage". 24 March 2014. Archived from the original on 12 June 2015. Retrieved 11 June 2015.
  123. ^ Bedford, Geoffrey O. (1978). "Biology and Ecology of the Phasmatodea". Annual Review of Entomology. 23: 125–149. doi:10.1146/annurev.en.23.010178.001013.
  124. ^ Ritland, D. B.; L. P. Brower (1991). "The viceroy butterfly is not a Batesian mimic". Nature. 350 (6318): 497–498. Bibcode:1991Natur.350..497R. doi:10.1038/350497a0. S2CID 28667520. Viceroys are as unpalatable as monarchs, and significantly more unpalatable than queens from representative Florida populations.
  125. ^ Meyer, A. (2006). "Repeating Patterns of Mimicry". PLOS Biology. 4 (10): e341. doi:10.1371/journal.pbio.0040341. PMC 1617347. PMID 17048984.
  126. ^ Kricher, John (1999). "6". A Neotropical Companion. Princeton University Press. pp. 157–158. ISBN 978-0-691-00974-2.
  127. ^ Wigglesworth, Vincent Brian. "Insect". Encyclopædia Britannica online. Archived from the original on 4 May 2012. Retrieved 19 April 2012.
  128. ^ "Pollinator Factsheet" (PDF). United States Forest Service. Archived from the original (PDF) on 10 April 2008. Retrieved 19 April 2012.
  129. ^ Ollerton, Jeff; Alarcón, Ruben; Waser, Nickolas M.; Price, Mary V.; et al. (14 February 2009). "A global test of the pollination syndrome hypothesis". Annals of Botany. Oxford University Press. 103 (9): 1471–1480. doi:10.1093/aob/mcp031. ISSN 1095-8290.
  130. ^ a b c Polaszek, Andrew; Vilhemsen, Lars (2023). "Biodiversity of hymenopteran parasitoids". Current Opinion in Insect Science. 56: 101026. doi:10.1016/j.cois.2023.101026. PMID 36966863. S2CID 257756440.
  131. ^ Forbes, Andrew A.; Bagley, Robin K.; Beer, Marc A.; et al. (12 July 2018). "Quantifying the unquantifiable: why Hymenoptera, not Coleoptera, is the most speciose animal order". BMC Ecology. 18 (1): 21. doi:10.1186/s12898-018-0176-x. ISSN 1472-6785. PMC 6042248. PMID 30001194.
  132. ^ Zhang, Qi; Kopylov, Dmitry S.; Rasnitsyn, Alexandr P.; Zheng, Yan; Zhang, Haichun (November 2020). Smith, Andrew (ed.). "Burmorussidae, a new family of parasitic wasps (Insecta, Hymenoptera) from mid‐Cretaceous Burmese amber". Papers in Palaeontology. 6 (4): 593–603. doi:10.1002/spp2.1312. ISSN 2056-2802. S2CID 219039881.
  133. ^ Tanaka, S.; Ohsaki, N. (2006). "Behavioral manipulation of host caterpillars by the primary parasitoid wasp Cotesia glomerata (L.) to construct defensive webs against hyperparasitism". Ecological Research. 21 (4): 570. doi:10.1007/s11284-006-0153-2. S2CID 23457678.
  134. ^ Poulin, Robert (2011). Rollinson, D.; Hay, S. I. (eds.). "The Many Roads to Parasitism: A Tale of Convergence". Advances in Parasitology. Academic Press. 74: 27–28. doi:10.1016/B978-0-12-385897-9.00001-X. ISBN 978-0-12-385897-9. PMID 21295676.
  135. ^ a b Poulin, Robert; Randhawa, Haseeb S. (February 2015). "Evolution of parasitism along convergent lines: from ecology to genomics". Parasitology. 142 (Suppl 1): S6–S15. doi:10.1017/S0031182013001674. PMC 4413784. PMID 24229807. open access
  136. ^ Gullan, P.J.; Cranston, P.S. (2014). The Insects: An Outline of Entomology, 5th Edition. Wiley. pp. 80–81, 790–. ISBN 978-1-118-84616-2.
  137. ^ Labandeira, Conrad C. (May 1998). "Early History of Arthropod and Vascular Plant Associations". Annual Review of Earth and Planetary Sciences. 26: 329–377. Bibcode:1998AREPS..26..329L. doi:10.1146/
  138. ^ Wilson, C. G.; Swincer, D. E.; Walden, K. J. (1982). "The Introduction of Trioxys Complanatus Quilis (Hymenoptera: Aphidiidae), an Internal Parasite of the Spotted Alfalfa Aphid, into South Australia". Australian Journal of Entomology. 21 (1): 13–27. doi:10.1111/j.1440-6055.1982.tb01758.x.
  139. ^ Bale, J. S.; van Lenteren, J. C.; Bigler, F. (27 February 2008). "Biological control and sustainable food production". Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences. 363 (1492): 761–776. doi:10.1098/rstb.2007.2182. PMC 2610108. PMID 17827110.
  140. ^ Davidson, E. (2006). Big Fleas Have Little Fleas: How Discoveries of Invertebrate Diseases Are Advancing Modern Science. Tucson, Ariz.: University of Arizona Press. ISBN 978-0-8165-2544-7.
  141. ^ Colborn, T.; vom Saal, F. S.; Soto, A. M. (October 1993). "Developmental effects of endocrine-disrupting chemicals in wildlife and humans". Environmental Health Perspectives. 101 (5): 378–384. doi:10.2307/3431890. JSTOR 3431890. PMC 1519860. PMID 8080506.
  142. ^ Nakamaru, M.; Iwasab, Y.; Nakanishic, J. (October 2003). "Extinction risk to bird populations caused by DDT exposure". Chemosphere. 53 (4): 377–387. Bibcode:2003Chmsp..53..377N. doi:10.1016/S0045-6535(03)00010-9. PMID 12946395.
  143. ^ Hong-Song Yu1; Yi-Hong Shen; Gang-Xiang Yuan; et al. (2011). "Evidence of selection at melanin synthesis pathway loci during silkworm domestication". Molecular Biology and Evolution. 28 (6): 1785–99. doi:10.1093/molbev/msr002. PMID 21212153.{{cite journal}}: CS1 maint: numeric names: authors list (link)
  144. ^ Dennis Normile (2009). "Sequencing 40 Silkworm Genomes Unravels History of Cultivation". Science. 325 (5944): 1058–1059. Bibcode:2009Sci...325.1058N. doi:10.1126/science.325_1058a. PMID 19713499.
  145. ^ Holldobler, Wilson (1994). Journey to the ants: a story of scientific exploration. Cambridge, Massachusetts: Belknap Press. pp. 196–199. ISBN 978-0-674-48525-9.
  146. ^ Jordan, Alex; Patch, Harland M.; Grozinger, Christina M.; Khanna, Vikas (26 January 2021). "Economic Dependence and Vulnerability of United States Agricultural Sector on Insect-Mediated Pollination Service". Environmental Science & Technology. 55 (4): 2243–2253. Bibcode:2021EnST...55.2243J. doi:10.1021/acs.est.0c04786. PMID 33496588. S2CID 231710967.
  147. ^ Crane, E. (1990). "Honey from honeybees and other insects". Ethology Ecology & Evolution. 3 (sup1): 100–105. doi:10.1080/03949370.1991.10721919.
  148. ^ Sanford, M.T.; Dietz, A. (1976). "The fine structure of the wax gland of the honey bee (Apis mellifera L.)". Apidologie. 7 (3): 197–207. doi:10.1051/apido:19760301.
  149. ^ "Wax Rendering". Bee Culture. 23 March 2016. Retrieved 26 October 2018.
  150. ^ "How Shellac Is Manufactured". The Mail (Adelaide, SA : 1912 – 1954). 18 December 1937. Retrieved 17 July 2015.
  151. ^ Bezzina, Neville. "Silk Production Process". Sense of Nature Research. Archived from the original on 29 June 2012.
  152. ^ Dams, M.; Dams, L. (21 July 1977). "Spanish Rock Art Depicting Honey Gathering During the Mesolithic". Nature. 268 (5617): 228–230. Bibcode:1977Natur.268..228D. doi:10.1038/268228a0. S2CID 4177275.
  153. ^ Roffet-Salque, Mélanie; et al. (14 June 2016). "Widespread exploitation of the honeybee by early Neolithic farmers". Nature. 534 (7607): 226–227. doi:10.1038/nature18451. PMID 26560301.
  154. ^ Vainker, Shelagh (2004). Chinese Silk: A Cultural History. Rutgers University Press. p. 20. ISBN 0813534461.
  155. ^ Christian, David (2000). "Silk Roads or Steppe Roads? The Silk Roads in World History". Journal of World History. 2 (1): 1. doi:10.1353/jwh.2000.0004. S2CID 18008906.
  156. ^ Gullan & Cranston 2005, p. 328–348, 400.
  157. ^ "Biocontrol Network – Beneficial Insects". Biocontrol Network. Archived from the original on 28 February 2009. Retrieved 9 May 2009.
  158. ^ Davidson, R. H.; Lyon, William F. (1979). Insect Pests of Farm, Garden, and Orchard. John Wiley & Sons. p. 38. ISBN 978-0-471-86314-4.
  159. ^ Camargo, Rafael; Oliveira, Paulo (2011). "Natural history of the Neotrobical arboreal ant, Odontomachus hastatus: Nest sites, foraging schedule, and diet". Journal of Insect Science. 12 (18): 48. doi:10.1673/031.012.4801. PMC 3476954. PMID 22957686.
  160. ^ Dossey, Aaron T. (December 2010). "Insects and their chemical weaponry: New potential for drug discovery". Natural Product Reports. 27 (12): 1737–1757. doi:10.1039/c005319h. PMID 20957283.
  161. ^ Sherman, Ronald A.; Pechter, Edward A. (1987). "Maggot therapy: a review of the therapeutic applications of fly larvae in human medicine, especially for treating osteomyelitis". Medical and Veterinary Entomology. 2 (3): 225–230. doi:10.1111/j.1365-2915.1988.tb00188.x. PMID 2980178. S2CID 44543735.
  162. ^ Briggs, John C (October 2017). "Emergence of a sixth mass extinction?". Biological Journal of the Linnean Society. 122 (2): 243–248. doi:10.1093/biolinnean/blx063.
  163. ^ Owens, Avalon C. S.; Lewis, Sara M. (November 2018). "The impact of artificial light at night on nocturnal insects: A review and synthesis". Ecology and Evolution. 8 (22): 11337–11358. Bibcode:2018EcoEv...811337O. doi:10.1002/ece3.4557. PMC 6262936. PMID 30519447.
  164. ^ Tscharntke, Teja; Klein, Alexandra M.; Kruess, Andreas; Steffan-Dewenter, Ingolf; Thies, Carsten (August 2005). "Landscape perspectives on agricultural intensification and biodiversity and ecosystem service management". Ecology Letters. 8 (8): 857–874. Bibcode:2005EcolL...8..857T. doi:10.1111/j.1461-0248.2005.00782.x. S2CID 54532666.
  165. ^ Insect-plant interactions in a crop protection perspective. Academic Press. 19 January 2017. pp. 313–320. ISBN 978-0-12-803324-1.
  166. ^ Braak, Nora; Neve, Rebecca; Jones, Andrew K.; Gibbs, Melanie; Breuker, Casper J. (November 2018). "The effects of insecticides on butterflies – A review". Environmental Pollution. 242 (Pt A): 507–518. doi:10.1016/j.envpol.2018.06.100. PMID 30005263. S2CID 51625489.
  167. ^ Wagner, David L.; Van Driesche, Roy G. (January 2010). "Threats Posed to Rare or Endangered Insects by Invasions of Nonnative Species". Annual Review of Entomology. 55 (1): 547–568. doi:10.1146/annurev-ento-112408-085516. PMID 19743915.
  168. ^ Sánchez-Bayo, Francisco; Wyckhuys, Kris A.G. (April 2019). "Worldwide decline of the entomofauna: A review of its drivers". Biological Conservation. 232: 8–27. doi:10.1016/j.biocon.2019.01.020.
  169. ^ Saunders, Manu (16 February 2019). "Insectageddon is a great story. But what are the facts?". Ecology is not a dirty word. Archived from the original on 25 February 2019. Retrieved 24 February 2019.
  170. ^ van Klink, Roel (24 April 2020), "Meta-analysis reveals declines in terrestrial but increases in freshwater insect abundances", Science, 368 (6489): 417–420, Bibcode:2020Sci...368..417V, doi:10.1126/science.aax9931, PMID 32327596, S2CID 216106896
  171. ^ McGrath, Matt (23 April 2020). "'Insect apocalypse' more complex than thought". BBC News. Retrieved 24 April 2020.
  172. ^ Pierce, B. A. (2006). Genetics: A Conceptual Approach (2nd ed.). New York: W.H. Freeman and Company. p. 87. ISBN 978-0-7167-8881-2.
  173. ^ Adams, M. D.; Celniker, S. E.; Holt, R. A.; et al. (24 March 2000). "The genome sequence of Drosophila melanogaster". Science. 287 (5461): 2185–2195. Bibcode:2000Sci...287.2185.. CiteSeerX doi:10.1126/science.287.5461.2185. PMID 10731132.
  174. ^ Ceurstemont, Sandrine (6 July 2013). "Inevitable insectivores? Not so fast". New Scientist. 219 (2924): 35. doi:10.1016/S0262-4079(13)61691-7. Retrieved 3 December 2021.
  175. ^ "Insects could be the key to meeting food needs of growing global population". the Guardian. 31 July 2010. Retrieved 13 January 2022.
  176. ^ Ramos-Elorduy, Julieta; Menzel, Peter (1998). Creepy crawly cuisine: the gourmet guide to edible insects. Inner Traditions / Bear & Company. p. 44. ISBN 978-0-89281-747-4. Retrieved 23 April 2014.
  177. ^ a b "Insects as Food for Humans". Retrieved 14 September 2022.
  178. ^ Gullan & Cranston 2005, pp. 10–13.
  179. ^ Michels, John (1880). John Michels (ed.). Science. Vol. 1. New York: American Association for the Advance of Science. p. 69.
  180. ^ Maierbrugger, Arno (14 May 2013). "UN: Insects are 'food of the future' (video)". Inside Investor. Archived from the original on 10 September 2013. Retrieved 17 May 2013.
  181. ^ Verheyen, Geert; Ooms, Tom; Vogels, Liesbeth; Vreysen, Steven; Bovy, Ann; Van Miert, Sabine; Meersman, Filip (1 May 2018). "Insects as an Alternative Source for the Production of Fats for Cosmetics". Journal of Cosmetic Science. 69 (3): 187–202. PMID 30052193.
  182. ^ "From Pest to Pot: Can Insects Feed the World?". National Geographic Culture. 15 August 2016. Archived from the original on 10 April 2021. Retrieved 13 January 2022.
  183. ^ "How AgriProtein makes chicken food from maggots". Wired UK. Retrieved 13 January 2022.
  184. ^ Bugs. "Bugs – das Wirbellosenmagazin". NTV Verlag. Retrieved 7 March 2021.
  185. ^ Gullan & Cranston 2005, p. 9.


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