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Temporal range: 240 –0 Ma
Middle Triassic – Recent
Bessenbandzweefvlieg Vrouwtje (2).JPG
The hoverfly Syrphus ribesii, suborder Brachycera, showing characteristic dipteran features: large eyes, small antennae, sucking mouthparts, single pair of flying wings, hindwings reduced to clublike halteres
Scientific classification edit
Kingdom: Animalia
Phylum: Euarthropoda
Class: Insecta
Superorder: Panorpida
(unranked): Antliophora
Order: Diptera
Linnaeus, 1758

Flies are insects with a pair of functional wings for flight and a pair of vestigial hindwings called halteres for balance. They are classified as an order called Diptera, that name being derived from the Greek δι- di- "two", and πτερόν pteron "wings". The order Diptera is divided into two suborders (although one suborder is non-monophyletic), with about 110 families divided between them; the families contain an estimated one million named species, including the familiar housefly, horse-fly, crane fly, and hoverfly;[a] although only about 125,000 species have a species description published.[3] The earliest fly fossils found so far are from the Triassic, about 240 million years ago; phylogenetic analysis suggests that flies originated in the Permian, about 260 million years ago.

Many insects, such as the butterfly, contain the word fly in their name, but are not Dipterans. Also, the word "fly" is sometimes used colloquially and non-scientifically as a name for any small flying insect: the term "true fly" is sometimes invoked to make clear the insect being referenced is a Dipteran.

Flies have a mobile head, with a pair of large compound eyes, and mouthparts designed for piercing and sucking (mosquitoes, black flies and robber flies), or for lapping and sucking in the other groups. The suborder Nematocera (from Greek, "thready-horns") have thin, long antennae; while the suborder Brachycera (from Greek "short-horns") have short antennae. Flies have only a single pair of wings to fly; their arrangement gives them great maneuverability in flight. The hindwings (halteres) evolved into advanced mechanosensory organs, which act as high-speed sensors of rotational movement and allow them to perform advanced aerobatics.[4] Claws and pads on their feet enable them to cling to smooth surfaces.

The life cycle of flies consists of the eggs, larva, pupa, and the adult. Flies undergo complete metamorphosis; the eggs are laid on the larval food-source, and the larvae (which lack true limbs) develop in a protected environment, often inside their food source. The pupa in higher dipterans is a tough capsule from which the adult emerges when ready to do so. Flies have short lives: for example, the adult housefly lives about a month; the mayfly about a year. The source of nutrition for adult flies is liquified food, including nectar.

Flies are of considerable ecological and human importance. They are important pollinators, second only to the bees and their Hymenopteran relatives. They may have been responsible for the first plant pollination in the Triassic. Mosquitoes are vectors for malaria, dengue, West Nile fever, yellow fever, encephalitis, and other infectious diseases; and houseflies, commensal with humans all over the world, spread food-borne illnesses. Flies can be annoyances, especially in some parts of the world where they can occur in large numbers, buzzing and settling on the skin or eyes to bite or seek fluids. Larger flies such as tsetse flies and screwworms cause significant economic harm to cattle. Blowfly larvae, known as gentles, and other dipteran larvae, known more generally as maggots, are used as fishing bait, as food for carnivorous animals, and in medicine for debridement to clean wounds. Fruit flies are used as model organisms in research. In culture, the subject of flies appears in religion, literature, cinema, and music.

Taxonomy and phylogeny[edit]

Relationships to other insects[edit]

Dipterans are endopterygotes, insects that undergo radical metamorphosis. They belong to the Mecopterida, alongside the Mecoptera, Siphonaptera, Lepidoptera and Trichoptera.[5][6] The possession of a single pair of wings distinguishes most true flies from other insects with "fly" in their names. However, some true flies such as Hippoboscidae (louse flies) have become secondarily wingless.[7]

The cladogram represents the current consensus view.[8]

part of Endopterygota

Diptera Common house fly, Musca domestica.jpg


Boreidae (snow scorpionflies, 30 spp.) Boreus hiemalis2 detail.jpg

Siphonaptera (fleas, 2500 spp.) Pulex irritans female ZSM.jpg

Trichoptera (caddisflies) Sericostoma.personatum.jpg

Lepidoptera (butterflies and moths) Tyria jacobaeae-lo.jpg



Relationships between fly subgroups and families[edit]


The first true dipterans known are from the Middle Triassic around 240 million years ago, and they became widespread during the Middle and Late Triassic.[9] Phylogenetic analysis of times of divergence suggests that dipterans originated in the Permian, some 260 million years ago.[10] Modern flowering plants did not appear until the Cretaceous (around 140 million years ago), so the original dipterans must have had a different source of nutrition other than nectar. Based on the attraction of many modern fly groups to shiny droplets, it has been suggested that they may have fed on honeydew produced by sap-sucking bugs which were abundant at the time, and dipteran mouthparts are well-adapted to softening and lapping up the crusted residues.[11] The basal clades in the Diptera include the Deuterophlebiidae and the enigmatic Nymphomyiidae.[12] Three episodes of evolutionary radiation are thought to have occurred based on the fossil record. Many new species of lower Diptera developed in the Triassic, about 220 million years ago. Many lower Brachycera appeared in the Jurassic, some 180 million years ago. A third radiation took place among the Schizophora at the start of the Paleogene, 66 million years ago.[12]


Diptera were traditionally broken down into two suborders, Nematocera and Brachycera, distinguished by the differences in antennae. The Nematocera are recognized by their elongated bodies and many-segmented, often feathery antennae as represented by mosquitoes and crane flies. The Brachycera have rounder bodies and much shorter antennae.[14][15] Subsequent studies have identified the Nematocera as being non-monophyletic with modern phylogenies placing the Brachycera within grades of groups formerly placed in the Nematocera. The construction of a phylogenetic tree has been the subject of ongoing research. The following cladogram is based on the FLYTREE project.[12][16][17]


Ptychopteromorpha (phantom and primitive crane-flies) Ptychoptera contaminata.jpg

Culicomorpha (mosquitoes) AnophelesGambiaemosquito.jpg

Blephariceromorpha (net-winged midges, etc) Imago of Blepharicera fasciata as Asthenia fasciata in Westwood 1842, plate 94.png

Bibionomorpha (gnats) Bibio marci02.jpg

Psychodomorpha (drain flies, sand flies, etc) Clogmia Albipunctata or moth fly.jpg



Stratiomyomorpha (soldier flies, etc) Hermetia illucens Black soldier fly edit1.jpg

Xylophagomorpha (stink flies, etc) Stinkfliege Coenomyia ferruginea male.jpg






Empidoidea (dance flies, etc) Empis.tessellata.male.jpg


Aschiza (in part)


Syrphoidea (hoverflies) Mosca cernidora de la grosella.jpg


Hippoboscoidea (louse flies, etc) CrataerhinaPallida.jpg



Acalyptratae (marsh flies, etc) Marsh fly01.jpg

Abbreviations used in the cladogram:


Geographic distribution[edit]

Gauromydas heros is the largest fly in the world.

Flies are often abundant and are found in almost all terrestrial habitats in the world apart from Antarctica. They include many familiar insects such as house flies, blow flies, mosquitoes, gnats, black flies, midges and fruit flies. More than 150,000 have been formally described and the actual species diversity is much greater, with the flies from many parts of the world yet to be studied intensively.[18][19] The suborder Nematocera include generally small, slender insects with long antennae such as mosquitoes, gnats, midges and crane-flies, while the Brachycera includes broader, more robust flies with short antennae. Many nematoceran larvae are aquatic.[20] There are estimated to be a total of about 19,000 species of Diptera in Europe, 22,000 in the Nearctic region, 20,000 in the Afrotropical region, 23,000 in the Oriental region and 19,000 in the Australasian region.[21] While most species have restricted distributions, a few like the housefly (Musca domestica) are cosmopolitan.[22] Gauromydas heros (Asiloidea), with a length of up to 7 cm (2.8 in), is generally considered to be the largest fly in the world,[23] while the smallest is Euryplatea nanaknihali, which at 0.4 mm (0.016 in) is smaller than a grain of salt.[24]


Brachycera are ecologically very diverse, with many being predatory at the larval stage and some being parasitic. Animals parasitised include molluscs, woodlice, millipedes, insects, mammals,[21] and amphibians.[25] Flies are the second largest group of pollinators after the Hymenoptera (bees, wasps and relatives). In wet and colder environments flies are significantly more important as pollinators. Compared to bees, they need less food as they do not need to provision their young. Many flowers that bear low nectar and those that have evolved trap pollination depend on flies.[26] It is thought that some of the earliest pollinators of plants may have been flies.[27]


The greatest diversity of gall forming insects are found among the flies, principally in the family Cecidomyiidae (gall midges).[28] Many flies (most importantly in the family Agromyzidae) lay their eggs in the mesophyll tissue of leaves with larvae feeding between the surfaces forming blisters and mines.[29] Some families are mycophagous or fungus feeding. These include the cave dwelling Mycetophilidae (fungus gnats) whose larvae are the only diptera with bioluminescence. The Sciaridae are also fungus feeders. Some plants are pollinated by fungus feeding flies that visit fungus infected male flowers.[30]

The larvae of Megaselia scalaris (Phoridae) are almost omnivorous and consume such substances as paint and shoe polish.[31] The larvae of the shore flies (Ephydridae) and some Chironomidae survive in extreme environments including glaciers (Diamesa sp., Chironomidae[32]), hot springs, geysers, saline pools, sulphur pools, septic tanks and even crude oil (Helaeomyia petrolei[32]).[21] Adult hoverflies (Syrphidae) are well known for their mimicry and the larvae adopt diverse lifestyles including being inquiline scavengers inside the nests of social insects.[33] Some brachycerans are agricultural pests, some bite animals and humans and suck their blood, and some transmit diseases.[21]







For visual course control, flies' optic flow field is analyzed by a set of motion-sensitive neurons.[36] A subset of these neurons is thought to be involved in using the optic flow to estimate the parameters of self-motion, such as yaw, roll, and sideward translation.[37] Other neurons are thought to be involved in analyzing the content of the visual scene itself, such as separating figures from the ground using motion parallax.[38][39] The H1 neuron is responsible for detecting horizontal motion across the entire visual field of the fly, allowing the fly to generate and guide stabilizing motor corrections midflight with respect to yaw.[40] The ocelli are concerned in the detection of changes in light intensity, enabling the fly to react swiftly to the approach of an object.[41]

Other sensory receptors[edit]

Like other insects, flies have chemoreceptors that detect smell and taste, and mechanoreceptors that respond to touch. The third segments of the antennae and the maxillary palps bear the main olfactory receptors, while the gustatory receptors are in the labium, pharynx, feet, wing margins and female genitalia,[42] enabling flies to taste their food by walking on it. The taste receptors in females at the tip of the abdomen receive information on the suitability of a site for ovipositing.[41] Flies that feed on blood have special sensory structures that can detect infrared emissions, and use them to home in on their hosts, and many blood-sucking flies can detect the raised concentration of carbon dioxide that occurs near large animals.[43] Some tachinid flies (Ormiinae) which are parasitoids of bush crickets, have sound receptors to help them locate their singing hosts.[44]



Diptera have one pair of fore wings on the mesothorax and a pair of halteres, or reduced hind wings, on the metathorax. A further adaptation for flight is the reduction in number of the neural ganglia, and concentration of nerve tissue in the thorax, a feature that is most extreme in the highly derived Muscomorpha infraorder.[35] Some species of flies are exceptional in that they are secondarily flightless. The only other order of insects bearing a single pair of true, functional wings, in addition to any form of halteres, are the Strepsiptera. In contrast to the flies, the Strepsiptera bear their halteres on the mesothorax and their flight wings on the metathorax.[45] Each of the fly's six legs has a typical insect structure of coxa, trochanter, femur, tibia and tarsus, with the tarsus in most instances being subdivided into five tarsomeres.[34] At the tip of the limb is a pair of claws, and between these are cushion-like structures known as pulvilli which provide adhesion.[46]


The abdomen shows considerable variability among members of the order. It consists of eleven segments in primitive groups and ten segments in more derived groups, the tenth and eleventh segments having fused.[47] The last two or three segments are adapted for reproduction. Each segment is made up of a dorsal and a ventral sclerite, connected by an elastic membrane. In some females, the sclerites are rolled into a flexible, telescopic ovipositor.[34]



Flies tend to fly in a straight line, then make a rapid change in direction before continuing on a different straight path. The directional changes are called saccades and typically involve an angle of 90°, achieved in 50 milliseconds. They are initiated by visual stimuli as the fly observes an object; nerves then activate steering muscles in the thorax that cause a small change in wing stroke, which generate sufficient torque to turn. Detecting this within four or five wingbeats, the halteres trigger a counter-turn and the fly heads off in a new direction.[50]

Flies have rapid reflexes that aid their escape from predators but their sustained flight speeds are low. Dolichopodid flies in the genus Condylostylus respond in less than 5 milliseconds to camera flashes by taking flight.[51] In the past, the deer bot fly, Cephenemyia, was claimed to be one of the fastest insects on the basis of an estimate made visually by Charles Townsend in 1927.[52] This claim, of speeds of 600 to 800 miles per hour, was regularly repeated until Irving Langmuir showed it to be physically impossible as well as incorrect. Langmuir suggested an estimated speed of 25 miles per hour.[53][54][55]

Although most flies live and fly close to the ground, a few are known to fly at heights and a few like Oscinella (Chloropidae) are known to be dispersed by winds at altitudes of up to 2000 ft and over long distances.[56] Some hover flies like Metasyrphus corollae have been known to undertake long flights in response to aphid population spurts.[57]

Males of fly species such as Cuterebra, many hover flies,[58] bee flies (Bombyliidae)[59] and fruit flies (Tephritidae)[60] maintain territories within which they engage in aerial pursuit to drive away intruding males and other species.[61] While these territories may be held by individual males, some species form leks with many males aggregating in displays.[60] Some flies maintain an airspace and still others form dense swarms that maintain a stationary location with respect to landmarks. Many flies mate in flight while swarming.[62]

Life cycle and development[edit]






Dipteran larvae have no jointed, "true legs",[63] but some dipteran larvae, such as species of Simuliidae, Tabanidae and Vermileonidae, have prolegs adapted to hold onto a substrate in flowing water, host tissues or prey.[66] The majority of dipterans are oviparous and lay batches of eggs, but some species are ovoviviparous, where the larvae starting development inside the eggs before they hatch or viviparous, the larvae hatching and maturing in the body of the mother before being externally deposited. These are found especially in groups that have larvae dependent on food sources that are short-lived or are accessible for brief periods.[67] This is widespread in some families such as the Sarcophagidae. In Hylemya strigosa (Anthomyiidae) the larva moults to the second instar before hatching, and in Termitoxenia (Phoridae) females have incubation pouches, and a full developed third instar larva is deposited by the adult and it almost immediately pupates with no freely feeding larval stage. The tsetse fly (as well as other Glossinidae, Hippoboscidae, Nycteribidae and Streblidae) exhibits adenotrophic viviparity; a single fertilised egg is retained in the oviduct and the developing larva feeds on glandular secretions. When fully grown, the female finds a spot with soft soil and the larva works its way out of the oviduct, buries itself and pupates. Some flies like Lundstroemia parthenogenetica (Chironomidae) reproduce by thelytokous parthenogenesis, and some gall midges have larvae that can produce eggs (paedogenesis).[68][69]




The adult stage is usually short, its function only to mate and lay eggs. The genitalia of male flies are rotated to a varying degree from the position found in other insects.[70] In some flies, this is a temporary rotation during mating, but in others, it is a permanent torsion of the organs that occurs during the pupal stage. This torsion may lead to the anus being below the genitals, or, in the case of 360° torsion, to the sperm duct being wrapped around the gut and the external organs being in their usual position. When flies mate, the male initially flies on top of the female, facing in the same direction, but then turns around to face in the opposite direction. This forces the male to lie on his back for his genitalia to remain engaged with those of the female, or the torsion of the male genitals allows the male to mate while remaining upright. This leads to flies having more reproduction abilities than most insects, and much quicker. Flies occur in large populations due to their ability to mate effectively and quickly during the mating season.[35]


Feeding and distribution[edit]




In many dipteran groups, swarming is a feature of adult life, with clouds of insects gathering in certain locations. These swarming insects are mostly males, and the swarm may serve the purpose of making their location more visible to females.[34]

Anti-predator adaptations[edit]


Flies are eaten by other animals at all stages of their development. The eggs and larvae are parasitised by other insects and are eaten by many creatures, some of which specialise in feeding on flies but most of which consume them as part of a mixed diet. Birds, bats, frogs, lizards, dragonflies and spiders are among the predators of flies.[72] Many flies have evolved mimetic resemblances that aid their protection. Batesian mimicry is widespread with many hoverflies resembling bees and wasps,[73][74] ants[75] and some species of tephritid fruit fly resembling spiders.[76] Some species of hoverfly are myrmecophilous, their young live and grow within the nests of ants. They are protected from the ants by imitating chemical odours given by ant colony members.[77] Bombyliid bee flies such as Bombylius major are short-bodied, round, furry, and distinctly bee-like as they visit flowers for nectar, and are likely also Batesian mimics of bees.[78]

In culture[edit]



Flies play a variety of symbolic roles in different cultures. These include both positive and negative roles in religion. In the traditional Navajo religion, Big Fly is an important spirit being.[79][80][81] In Christian demonology, Beelzebub is a demonic fly, the "Lord of the Flies", and a god of the Philistines.[82][83][84]


Flies have appeared in literature since ancient Sumer.[85] In a Sumerian poem, a fly helps the goddess Inanna when her husband Dumuzid is being chased by galla demons.[85] In the Mesopotamian versions of the flood myth, the dead corpses floating on the waters are compared to flies.[85] Later, the gods are said to swarm "like flies" around the hero Utnapishtim's offering.[85] Flies appear on Old Babylonian seals as symbols of Nergal, the god of death.[85] Fly-shaped lapis lazuli beads were often worn in ancient Mesopotamia, along with other kinds of fly-jewellery.[85]

In Prometheus Bound, which is attributed to the Athenian tragic playwright Aeschylus, a gadfly sent by Zeus's wife Hera pursues and torments his mistress Io, who has been transformed into a cow and is watched constantly by the hundred eyes of the herdsman Argus:[86][87] "Io: Ah! Hah! Again the prick, the stab of gadfly-sting! O earth, earth, hide, the hollow shape—Argus—that evil thing—the hundred-eyed."[87] William Shakespeare, inspired by Aeschylus, has Tom o'Bedlam in King Lear, "Whom the foul fiend hath led through fire and through flame, through ford and whirlpool, o'er bog and quagmire", driven mad by the constant pursuit.[87] In Antony and Cleopatra, Shakespeare similarly likens Cleopatra's hasty departure from the Actium battlefield to that of a cow chased by a gadfly.[88] More recently, in 1962 the biologist Vincent Dethier wrote To Know a Fly, introducing the general reader to the behaviour and physiology of the fly.[89]


Damien Hirst's provocative[90] 1990 artwork, titled A Thousand Years, featured a severed cow's head contained in a glass box with thousands of flies and a bug zapper, showing the life-cycle of the fly as well as the decomposition of the cow.[91]


Flies appear in popular culture in concepts such as fly-on-the-wall documentary-making in film and television production. The metaphoric name suggests that events are seen candidly, as a fly might see them.[92] Flies have inspired the design of miniature flying robots.[93] Steven Spielberg's 1993 film Jurassic Park relied on the idea that DNA could be preserved in the stomach contents of a blood-sucking fly fossilised in amber, though the mechanism has been discounted by scientists.[94]


Musical works that feature the fly:

Economic importance[edit]





Blowflies feeding on the fresh corpse of a porcupine, Hystrix africaeaustralis



Maggots used as animal feed at London Zoo

Some species of maggots such as blowfly larvae (gentles) and bluebottle larvae (casters) are bred commercially; they are sold as bait in angling, and as food for carnivorous animals (kept as pets, in zoos, or for research) such as some mammals,[99] fishes, reptiles, and birds. It has been suggested that fly larvae could be used at a large scale as food for farmed chickens, pigs, and fish. However, consumers are opposed to the inclusion of insects in their food, and the use of insects in animal feed remains illegal in areas such as the European Union.[100][101]


Fly larvae can be used as a biomedical tool for wound care and treatment. Maggot debridement therapy (MDT) is the use of blow fly larvae to remove the dead tissue from wounds, most commonly being amputations. Historically, this has been used for centuries, both intentional and unintentional, on battlefields and in early hospital settings.[102] Removing the dead tissue promotes cell growth and healthy wound healing. The larvae also have biochemical properties such as antibacterial activity found in their secretions as they feed.[103] These medicinal maggots are a safe and effective treatment for chronic wounds.[104]

The Sardinian cheese casu marzu is exposed to flies known as cheese skippers such as Piophila casei, members of the family Piophilidae.[105] The digestive activities of the fly larvae soften the cheese and modify the aroma as part of the process of maturation. At one time European Union authorities banned sale of the cheese and it was becoming hard to find,[106] but the ban has been lifted on the grounds that the cheese is a traditional local product made by traditional methods.[107]






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Further reading[edit]

  • Séguy, Eugène La Biologie des Dipteres 1950.

External links[edit]