Flies areinsects of theorderDiptera, the name being derived from Ancient Greek δι- (di-)'two' and πτερόν (pteron)'wing'. Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advancedmechanosensory organs known ashalteres, which act as high-speed sensors of rotational movement and allow dipterans to perform advanced aerobatics. Diptera is a large order containing more than 150,000species includinghorse-flies,[a]crane flies,hoverflies,mosquitoes and others.
Flies have a mobile head, with a pair of largecompound eyes, and mouthparts designed for piercing and sucking (mosquitoes, black flies and robber flies), or for lapping and sucking in the other groups. Their wing arrangement gives them great manoeuvrability in flight, and claws and pads on their feet enable them to cling to smooth surfaces. Flies undergo completemetamorphosis; the eggs are often laid on the larval food-source and the larvae, which lack true limbs, develop in a protected environment, often inside their food source. Other species areovoviviparous, opportunistically depositing hatched or hatching larvae instead of eggs oncarrion, dung, decaying material, or open wounds of mammals. The pupa is a tough capsule from which the adult emerges when ready to do so; flies mostly have short lives as adults.
Diptera is one of the major insect orders and of considerableecological and human importance. Flies are major pollinators, second only to the bees and theirHymenopteran relatives. Flies may have been among the evolutionarily earliest pollinators responsible for early plantpollination.Fruit flies are used asmodel organisms in research, but less benignly,mosquitoes arevectors formalaria,dengue,West Nile fever,yellow fever,encephalitis, and otherinfectious diseases; andhouseflies, commensal with humans all over the world, spreadfoodborne 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 astsetse flies andscrewworms cause significant economic harm to cattle. Blowfly larvae, known asgentles, and other dipteran larvae, known more generally asmaggots, are used asfishing bait, as food for carnivorous animals, and in medicine indebridement,to clean wounds.
Taxonomy and phylogeny
Relationships to other insects
Dipterans areholometabolous, meaning that they undergo radical metamorphosis. They belong to theMecopterida, alongside theMecoptera,Siphonaptera,Lepidoptera andTrichoptera.[4][5] 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 asHippoboscidae (louse flies) have become secondarily wingless.[6][7]
Thecladogram represents the current consensus view.[8]
The first true dipterans known are from theMiddle Triassic (around 240 million years ago), and they became widespread during the Middle andLate Triassic.[9]Modern flowering plants did not appear until theCretaceous (around 140 million years ago), so the original dipterans must have had a different source of nutrition other thannectar. Based on the attraction of many modern fly groups to shiny droplets, it has been suggested that they may have fed onhoneydew produced bysap-sucking bugs which were abundant at the time, and dipteran mouthparts are well-adapted to softening and lapping up the crusted residues.[10] Thebasal clades in the Diptera include theDeuterophlebiidae and the enigmaticNymphomyiidae.[11] Three episodes ofevolutionary radiation are thought to have occurred based on the fossil record. Many new species of lower Diptera developed in theTriassic, about 220 million years ago. Many lower Brachycera appeared in theJurassic, some 180 million years ago. A third radiation took place among theSchizophora at the start of thePaleogene, 66 million years ago.[11]
Thephylogenetic position of Diptera has been controversial. Themonophyly ofholometabolous insects has long been accepted, with the main orders being established as Lepidoptera, Coleoptera, Hymenoptera and Diptera, and it is the relationships between these groups which has caused difficulties. Diptera is widely thought to be a member ofMecopterida, along with Lepidoptera (butterflies and moths), Trichoptera (caddisflies), Siphonaptera (fleas), Mecoptera (scorpionflies) and possiblyStrepsiptera (twisted-wing flies). Diptera has been grouped with Siphonaptera and Mecoptera in the Antliophora, but this has not been confirmed by molecular studies.[12]
Diptera were traditionally broken down into two suborders,Nematocera andBrachycera, distinguished by the differences in antennae. The Nematocera are identified 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.[13][14] Subsequent studies have identified the Nematocera as beingparaphyletic 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.[15][16]
Flies are often abundant and are found in almost all terrestrial habitats. They include many familiar insects such as house flies, blow flies, mosquitoes, gnats, black flies, midges and fruit flies. More than 150,000 have beenformally 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 isEuryplatea 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 includemolluscs,woodlice,millipedes, insects,mammals,[21] andamphibians.[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 evolvedtrap 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 is found among the flies, principally among the gall midges (Cecidomyiidae).[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 feed on fungi. These include the cave dwelling fungus gnats (Mycetophilidae) whose larvae are bioluminescent, and theSciaridae. Some plants are pollinated by fungus feeding flies that visit fungus-infected male flowers.[30]
The larvae ofMegaselia scalaris (Phoridae) are omnivorous and may consume such substances as paint and shoe polish.[31]Exorista mella is a generalist andparasitoid of multiple hosts.[32] The larvae of shore flies (Ephydridae) and somechironomid midges survive in extreme environments including glaciers (as inDiamesa sp.[33]), hot springs, geysers, saline pools, sulphur pools, septic tanks and even crude oil (Helaeomyia petrolei[33]).[21] Adulthoverflies (Syrphidae) are well known for theirmimicry and the larvae adopt diverse lifestyles including beinginquilinescavengers inside the nests of social insects.[34] Some brachycerans are agricultural pests, some bite animals and humans and suck their blood, and some transmit diseases.[21]
Flies are adapted for aerial movement and typically have short and streamlined bodies. The firsttagma of the fly, the head, bears the eyes, theantennae, and themouthparts (the labrum, labium, mandible, and maxilla make up the mouthparts). The second tagma, thethorax, bears the wings and contains the flight muscles on the second segment, which is greatly enlarged; the first and third segments have been reduced to collar-like structures, and the third segment bears thehalteres, which help to balance the insect during flight. The third tagma is theabdomen consisting of 11 segments, some of which may be fused, and with the three hindmost segments modified for reproduction.[35][36] Some Dipterans are mimics and can only be distinguished from their models by very careful inspection. An example of this isSpilomyia longicornis, which is a fly but mimics avespid wasp.[37]
Head ofa horse-fly showing largecompound eyes and stout piercingmouthpartsHead of a fly, showing the two compound eyes and three simple eyes
Flies have a mobile head with a pair of largecompound eyes on the sides of the head, and in most species, three smallocelli on the top. The compound eyes may be close together or widely separated, and in some instances are divided into a dorsal region and a ventral region, perhaps to assist in swarming behaviour. The antennae are well-developed but variable, being thread-like, feathery or comb-like in the different families. The mouthparts are adapted for piercing and sucking, as in the black flies, mosquitoes and robber flies, and for lapping and sucking as in many other groups.[36] Femalehorse-flies use knife-like mandibles and maxillae to make a cross-shaped incision in the host's skin and then lap up the blood that flows. The gut includes largediverticulae, allowing the insect to store small quantities of liquid after a meal.[38]
For visual course control, flies'optic flow field is analyzed by a set of motion-sensitive neurons.[39] 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.[40] 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.[41][42] TheH1 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.[43] The ocelli are concerned in the detection of changes in light intensity, enabling the fly to react swiftly to the approach of an object.[44]
Like other insects, flies havechemoreceptors that detect smell and taste, andmechanoreceptors 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,[45] 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.[44] Flies that feed on blood have special sensory structures that can detectinfrared emissions, and use them to home in on their hosts. Many blood-sucking flies can detect the raised concentration ofcarbon dioxide that occurs near large animals.[46] Some tachinid flies (Ormiinae) which are parasitoids ofbush crickets, have sound receptors to help them locate their singing hosts.[47]
Acrane fly, showing the hind wings reduced to drumstick-shapedhalteres
Diptera have one pair of forewings on themesothorax and a pair ofhalteres, or reduced hind wings, on themetathorax. A further adaptation for flight is the reduction in number of the neuralganglia, and concentration of nerve tissue in the thorax, a feature that is most extreme in the highly derived Muscomorpha infraorder.[38] Some flies such as the ectoparasiticNycteribiidae andStreblidae are exceptional in having lost their wings and become flightless. The only other order of insects bearing a single pair of true, functional wings, in addition to any form of halteres, are theStrepsiptera. In contrast to the flies, the Strepsiptera bear their halteres on the mesothorax and their flight wings on the metathorax.[48] Each of the fly's sixlegs has a typical insect structure of coxa, trochanter, femur, tibia and tarsus, with the tarsus in most instances being subdivided into fivetarsomeres.[36] At the tip of the limb is a pair of claws, and between these are cushion-like structures known aspulvilli which provide adhesion.[49]
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.[50] The last two or three segments are adapted for reproduction. Each segment is made up of a dorsal and a ventralsclerite, connected by an elastic membrane. In some females, the sclerites are rolled into a flexible, telescopicovipositor.[36]
Flies are capable of great manoeuvrability during flight due to the presence of the halteres. These act asgyroscopic organs and are rapidly oscillated in time with the wings; they act as a balance and guidance system by providing rapid feedback to the wing-steering muscles, and flies deprived of their halteres are unable to fly. The wings and halteres move in synchrony but the amplitude of each wing beat is independent, allowing the fly to turn sideways.[51] The wings of the fly are attached to two kinds of muscles, those used to power it and another set used for fine control.[52]
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 calledsaccades and typically involve an angle of 90°, being 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.[53]
Flies have rapid reflexes that aid their escape from predators but their sustained flight speeds are low.Dolichopodid flies in the genusCondylostylus respond in less than five milliseconds to camera flashes by taking flight.[54] 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 byCharles Townsend in 1927.[55] This claim, of speeds of 600 to 800 miles per hour, was regularly repeated until it was shown to be physically impossible as well as incorrect by Irving Langmuir. Langmuir suggested an estimated speed of 25 miles per hour.[56][57][58]
Although most flies live and fly close to the ground, a few are known to fly at heights and a few likeOscinella (Chloropidae) are known to be dispersed by winds at altitudes of up to 2,000 ft and over long distances.[59] Some hover flies likeMetasyrphus corollae have been known to undertake long flights in response to aphid population spurts.[60]
Males of fly species such asCuterebra, many hover flies,[61] bee flies (Bombyliidae)[62] and fruit flies (Tephritidae)[63] maintain territories within which they engage in aerial pursuit to drive away intruding males and other species.[64] While these territories may be held by individual males, some species, such asA. freeborni,[65] formleks with many males aggregating in displays.[63] 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.[66]
Diptera go through a complete metamorphosis with four distinct life stages – egg, larva, pupa and adult.
Larva
In many flies, the larval stage is long and adults may have a short life. Most dipteran larvae develop in protected environments; many are aquatic and others are found in moist places such as carrion, fruit, vegetable matter, fungi and, in the case of parasitic species, inside their hosts. They tend to have thin cuticles and become desiccated if exposed to the air. Apart from theBrachycera, most dipteran larvae have sclerotised head capsules, which may be reduced to remnant mouth hooks; the Brachycera, however, have soft, gelatinized head capsules from which the sclerites are reduced or missing. Many of these larvae retract their heads into their thorax.[36][67] Thespiracles in the larva and pupa do not have any internal mechanical closing device.[68]
Some other anatomical distinction exists between the larvae of theNematocera and theBrachycera. Especially in the Brachycera, little demarcation is seen between the thorax and abdomen, though the demarcation may be visible in many Nematocera, such as mosquitoes; in the Brachycera, the head of the larva is not clearly distinguishable from the rest of the body, and few, if any, sclerites are present. Informally, such brachyceran larvae are called maggots,[69] but the term is not technical and often applied indifferently to fly larvae or insect larvae in general. The eyes and antennae of brachyceran larvae are reduced or absent, and the abdomen also lacks appendages such ascerci. This lack of features is an adaptation to food such as carrion, decaying detritus, or host tissues surroundingendoparasites.[38] Nematoceran larvae generally have well-developed eyes and antennae, while those of Brachyceran larvae are reduced or modified.[70]
Dipteran larvae have no jointed, "true legs",[67] but some dipteran larvae, such as species ofSimuliidae,Tabanidae andVermileonidae, haveprolegs adapted to hold onto a substrate in flowing water, host tissues or prey.[71] The majority of dipterans areoviparous and lay batches of eggs, but some species areovoviviparous, 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.[72] This is widespread in some families such as the Sarcophagidae. InHylemya strigosa (Anthomyiidae) the larva moults to the second instar before hatching, and inTermitoxenia (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. Thetsetse fly (as well as other Glossinidae, Hippoboscidae, Nycteribidae and Streblidae) exhibitsadenotrophic 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 likeLundstroemia parthenogenetica (Chironomidae) reproduce bythelytokousparthenogenesis, and somegall midges have larvae that can produce eggs (paedogenesis).[73][74]
Pupa
Thepupae take various forms. In some groups, particularly the Nematocera, the pupa is intermediate between the larval and adult form; these pupae are described as "obtect", having the future appendages visible as structures that adhere to the pupal body. The outer surface of the pupa may be leathery and bear spines, respiratory features or locomotory paddles. In other groups, described as "coarctate", the appendages are not visible. In these, the outer surface is apuparium, formed from the last larval skin, and the actual pupa is concealed within. When the adult insect is ready to emerge from this tough, desiccation-resistant capsule, it inflatesa balloon-like structure on its head, and forces its way out.[36]
Adult
The adult stage is usually short, its function is only to mate and lay eggs. The genitalia of male flies are rotated to a varying degree from the position found in other insects.[75] 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 theanus 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.[38] More primitive groups mates in the air during swarming, but most of the more advanced species with a 360° torsion mate on a substrate.[76]
Ecology
Trophic levels
A calliphorid "bubbling"
As ubiquitous insects, dipterans play an important role at varioustrophic levels both as consumers and as prey. In some groups the larvae complete their development without feeding, and in others the adults do not feed. The larvae can be herbivores, scavengers, decomposers, predators or parasites, with the consumption of decaying organic matter being one of the most prevalent feeding behaviours. The fruit or detritus is consumed along with the associated micro-organisms, a sieve-like filter in the pharynx being used to concentrate the particles, while flesh-eating larvae have mouth-hooks to help shred their food. The larvae of some groups feed on or in the living tissues of plants and fungi, and some of these are serious pests of agricultural crops. Some aquatic larvae consume the films of algae that form underwater on rocks and plants. Many of the parasitoid larvae grow inside and eventually kill other arthropods, while parasitic larvae may attack vertebrate hosts.[36]
Habitats and ecological niches
Whereas many dipteran larvae are aquatic or live in enclosed terrestrial locations, the majority of adults live above ground and are capable of flight. Predominantly they feed on nectar or plant or animal exudates, such as honeydew, for which their lapping mouthparts are adapted. Some flies have functional mandibles that may be used for biting. The flies that feed on vertebrate blood have sharp stylets that pierce the skin, with some species having anticoagulant saliva that is regurgitated before absorbing the blood that flows; in this process, certain diseases can be transmitted. The bot flies (Oestridae) have evolved to parasitize mammals. Many species complete their life cycle inside the bodies of their hosts.[77] The larvae of a few fly groups (Agromyzidae, Anthomyiidae, Cecidomyiidae) are capable of inducing plant galls. Some dipteran larvae are leaf-miners. The larvae of many brachyceran families are predaceous. In many dipteran groups, swarming is a feature of adult life, with clouds of insects gathering in certain locations; these insects are mostly males, and the swarm may serve the purpose of making their location more visible to females.[36]
Most adult diptera have their mouthparts modified to sponge up fluid. The adults of many species of flies (e.g.Anthomyia sp.,Steganopsis melanogaster) that feed on liquid food willregurgitate fluid in a behaviour termed as "bubbling" which has been thought to help the insects evaporate water and concentrate food[78] or possibly to cool by evaporation.[79] Some adult diptera are known forkleptoparasitism such as members of the Sarcophagidae. The miltogramminae are known as "satellite flies" for their habit of following wasps and stealing their stung prey or laying their eggs into them. Phorids, milichids and the genusBengalia are known to steal food carried by ants.[80] Adults ofEphydra hians forage underwater, and have special hydrophobic hairs that trap a bubble of air that lets them breathe underwater.[81]
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.[82] Many flies have evolvedmimetic resemblances that aid their protection.Batesian mimicry is widespread with many hoverflies resembling bees and wasps,[83][84] ants[85] and some species of tephritid fruit fly resembling spiders.[86] Some species of hoverfly aremyrmecophilous—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.[87] Bombyliid bee flies such asBombylius major are short-bodied, round, furry, and distinctly bee-like as they visit flowers for nectar, and are likely also Batesian mimics of bees.[88]
In contrast,Drosophila subobscura, a species of fly in the genusDrosophila, lacks a category of hemocytes that are present in other studied species ofDrosophila, leading to an inability to defend against parasitic attacks, a form of innate immunodeficiency.[89]
Petrus Christus's 1446 paintingPortrait of a Carthusian has amusca depicta (painted fly) on atrompe-l'œil frame.
Flies play a variety of symbolic roles in different cultures. These include both positive and negative roles in religion. In the traditionalNavajo religion, Big Fly is an important spirit being.[90][91][92] InChristian demonology,Beelzebub is a demonic fly, the "Lord of the Flies", and a god of thePhilistines.[93][94][95]
Flies have appeared in literature since ancientSumer.[96] In a Sumerian poem, a fly helps the goddessInanna when her husbandDumuzid is being chased bygalla demons.[96] In the Mesopotamian versions of theflood myth, the dead corpses floating on the waters are compared to flies.[96] Later, the gods are said to swarm "like flies" around the heroUtnapishtim's offering.[96] Flies appear onOld Babylonian seals as symbols ofNergal, the god of death.[96] Fly-shapedlapis lazuli beads were often worn in ancient Mesopotamia, along with other kinds of fly-jewellery.[96]
In Ancient Egypt, flies appear in amulets and as a military award for bravery and tenacity, due to the fact that they always come back when swatted at. It is thought that flies may have also been associated with the departing spirit of the dead, as they are often found near dead bodies. In modern Egypt, a similar belief persists in some areas to not swat at shiny green flies, as they may be carrying the soul of a recently deceased person.[97]
In a little-knownGreek myth, a very chatty and talkative maiden namedMyia (meaning "fly") enraged the moon-goddessSelene by attempting to seduce her lover, the sleepingEndymion, and was thus turned by the angry goddess into a fly, who now always deprives people of their sleep in memory of her past life.[98][99] InPrometheus Bound, which is attributed to the Athenian tragic playwrightAeschylus, agadfly sent byZeus's wifeHera pursues and torments his mistressIo, who has been transformed into a cow and is watched constantly by the hundred eyes of the herdsmanArgus:[100][101] "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."[101]William Shakespeare, inspired by Aeschylus, hasTom o'Bedlam inKing 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.[101] InAntony and Cleopatra, Shakespeare similarly likens Cleopatra's hasty departure from theActium battlefield to that of a cow chased by a gadfly.[102] More recently, in 1962 the biologist Vincent Dethier wroteTo Know a Fly, introducing the general reader to the behaviour and physiology of the fly.[103]
Musca depicta ("painted fly" in Latin) is a depiction of a fly as an inconspicuous element of various paintings. This feature was widespread in 15th and 16th centuries paintings and its presence may be explained by various reasons.[104]
Many dipterans serve roles that are useful to humans. Houseflies, blowflies andfungus gnats (Mycetophilidae) are scavengers and aid in decomposition.Robber flies (Asilidae),tachinids (Tachinidae) anddagger flies and balloon flies (Empididae) are predators and parasitoids of other insects, helping to control a variety of pests. Many dipterans such asbee flies (Bombyliidae) andhoverflies (Syrphidae) arepollinators of crop plants.[36]
Drosophila melanogaster, a fruit fly, has long been used as amodel organism in research because of the ease with which it can be bred and reared in the laboratory, its smallgenome, and the fact that many of its genes have counterparts in highereukaryotes. A large number of genetic studies have been undertaken based on this species; these have had a profound impact on the study ofgene expression,gene regulatory mechanisms andmutation. Other studies have investigatedphysiology,microbial pathogenesis and development among other research topics.[109] The studies on dipteran relationships byWilli Hennig helped in the development ofcladistics, techniques that he applied to morphological characters but now adapted for use with molecular sequences in phylogenetics.[110]
Maggots found on corpses are useful toforensic entomologists. Maggot species can be identified by their anatomical features and by matching theirDNA. Maggots of different species of flies visit corpses and carcases at fairly well-defined times after the death of the victim, and so do their predators, such as beetles in the familyHisteridae. Thus, the presence or absence of particular species provides evidence for the time since death, and sometimes other details such as the place of death, when species are confined to particular habitats such aswoodland.[111]
Some species of maggots such asblowfly larvae (gentles) and bluebottle larvae (casters) are bred commercially; they are sold asbait inangling, and as food for carnivorous animals (kept as pets, in zoos, or for research) such as somemammals,[112]fishes,reptiles, andbirds. 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 theEuropean Union.[113][114]
Fly larvae can be used as abiomedical tool for wound care and treatment.Maggot debridement therapy (MDT) is the use ofblow 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.[115] Removing the dead tissue promotes cell growth and healthy wound healing. The larvae also havebiochemical properties such asantibacterial activity found in their secretions as they feed.[116] These medicinal maggots are a safe and effective treatment for chronic wounds.[117]
TheSardinian cheesecasu marzu is exposed to flies known ascheese skippers such asPiophila casei, members of the familyPiophilidae.[118] 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,[119] but the ban has been lifted on the grounds that the cheese is a traditional local product made by traditional methods.[120]
Notes
^Some authors draw a distinction in writing the common names of insects. True flies are in their view best written as two words, such ascrane fly,robber fly,bee fly,moth fly, and fruit fly. In contrast, common names of non-dipteran insects that have "fly" in their names are written as one word, e.g. butterfly, stonefly, dragonfly, scorpionfly, sawfly, caddisfly, whitefly.[1] In practice, however, this is a comparatively new convention; especially in older books, names like "saw fly" and "caddis fly", or hyphenated forms such ashouse-fly and dragon-fly are widely used.[2] Exceptions to this rule occur, such as thehoverfly (though some authors prefer "hover fly"[3]), which is a true fly, and theSpanish fly, a type ofblister beetle.
^Yeates, David K.; Weigmann, Brian M; Courtney, Greg W.; Meier, Rudolf; Lambkins, Christine; Pape, Thomas (2007). "Phylogeny and systematics of Diptera: Two decades of progress and prospects".Zootaxa.1668:565–590.Bibcode:2007Zoot.1668.1.27Y.doi:10.11646/zootaxa.1668.1.27.
^abcdHoell, H. V.; Doyen, J. T.; Purcell, A. H. (1998).Introduction to Insect Biology and Diversity (2nd ed.). Oxford University Press. pp. 493–499.ISBN978-0-19-510033-4.
^abRuppert, Edward E.; Fox, Richard, S.; Barnes, Robert D. (2004).Invertebrate Zoology, 7th edition. Cengage Learning. pp. 735–736.ISBN978-81-315-0104-7.{{cite book}}: CS1 maint: multiple names: authors list (link)
^Stocker, Reinhard F. (2005). "The organization of the chemosensory system inDrosophila melanogaster: a review".Cell and Tissue Research.275 (1):3–26.doi:10.1007/BF00305372.PMID8118845.S2CID23210046.
^Lakes-Harlan, Reinhard; Jacobs, Kirsten; Allen, Geoff R. (2007). "Comparison of auditory sense organs in parasitoid Tachinidae (Diptera) hosted by Tettigoniidae (Orthoptera) and homologous structures in a non-hearing Phoridae (Diptera)".Zoomorphology.126 (4):229–243.doi:10.1007/s00435-007-0043-3.S2CID46359462.
^Dickinson, Michael H; Tu, Michael S (1 March 1997). "The function of dipteran flight muscle".Comparative Biochemistry and Physiology Part A: Physiology.116 (3):223–238.doi:10.1016/S0300-9629(96)00162-4.
^Townsend, Charles H.T. (1927). "On theCephenemyia mechanism and the Daylight-Day circuit of the Earth by flight".Journal of the New York Entomological Society.35 (3):245–252.JSTOR25004207.
^Wellington, W. G.; Fitzpatrick, Sheila M. (2012). "Territoriality in the Drone Fly,Eristalis tenax (Diptera: Syrphidae)".The Canadian Entomologist.113 (8):695–704.doi:10.4039/Ent113695-8.S2CID86181761.
^Dodson, Gary; Yeates, David (1990). "The mating system of a bee fly (Diptera: Bombyliidae). II. Factors affecting male territorial and mating success".Journal of Insect Behavior.3 (5):619–636.Bibcode:1990JIBeh...3..619D.doi:10.1007/BF01052332.S2CID25061334.
^abBecerril-Morales, Felipe; Macías-Ordóñez, Rogelio (2009). "Territorial contests within and between two species of flies (Diptera: Richardiidae) in the wild".Behaviour.146 (2):245–262.doi:10.1163/156853909X410766.
^Marshall, S.A.; Kirk-Spriggs, A.H. (2017). "Natural history of Diptera". In Kirk-Spriggs, A.H.; Sinclair, B.J. (eds.).Manual of Afrotropical Diptera. Volume 1. Introductory chapters and keys to Diptera families. Suricata 4. Pretoria: South African National Biodiversity Institute. pp. 135–152.
^Pie, Marcio R.; Del-Claro, Kleber (2002). "Male-male agonistic behavior and ant-mimicry in a Neotropical richardiid (Diptera: Richardiidae)".Studies on Neotropical Fauna and Environment.37 (1):19–22.Bibcode:2002SNFE...37...19P.doi:10.1076/snfe.37.1.19.2114.S2CID84201196.
^Whitman, D. W.; Orsak, L.; Greene, E. (1988). "Spider mimicry in fruit flies (Diptera: Tephritidae): Further experiments on the deterrence of jumping spiders (Araneae: Salticidae) byZonosemata vittigera (Coquillett)".Annals of the Entomological Society of America.81 (3):532–536.doi:10.1093/aesa/81.3.532.
^Akre, Roger D.; Garnett, William B.; Zack, Richard S. (1990). "Ant hosts ofMicrodon (Diptera: Syrphidae) in the Pacific Northwest".Journal of the Kansas Entomological Society.63 (1):175–178.JSTOR25085158.
^Wyman, Leland Clifton (1983)."Navajo Ceremonial System"(PDF).Handbook of North American Indians. p. 539. Archived fromthe original(PDF) on 5 March 2016. Retrieved30 July 2015.Nearly every element in the universe may be thus personalized, and even the least of these such as tiny Chipmunk and those little insect helpers and mentors of deity and man in the myths, Big Fly (Dǫ'soh) and Ripener (Corn Beetle) Girl ('Anilt'ánii 'At'ééd) (Wyman and Bailey 1964:29–30, 51, 137–144), are as necessary for the harmonious balance of the universe as is the great Sun.
^"Βεελζεβούλ, ὁ indecl. (v.l. Βεελζεβούβ and Βεεζεβούλ W-S. §5, 31, cp. 27 n. 56) Beelzebul, orig. a Philistine deity; the name בַּעַל זְבוּב means Baal (lord) of the flies (4 Km 1:2, 6; Sym. transcribes βεελζεβούβ; Vulgate Beelzebub; TestSol freq. Βεελζεβούλ,-βουέλ).", Arndt, W., Danker, F. W., & Bauer, W. (2000). A Greek-English lexicon of the New Testament and other early Christian literature (3rd ed.) (173). Chicago: University of Chicago Press.
^"1. According to 2 Kgs 1:2–6 the name of the Philistine god of Ekron was Lord of the Flies (Heb. ba'al zeaûḇ), from whom Israel's King Ahaziah requested an oracle.", Balz, H. R., & Schneider, G. (1990–). Vol. 1: Exegetical dictionary of the New Testament (211). Grand Rapids, Mich.: Eerdmans.
^"For etymological reasons, Baal Zebub must be considered a Semitic god; he is taken over by the Philistine Ekronites and incorporated into their local cult.", Herrmann, "Baal Zebub", in Toorn, K., Becking, B., & Horst, P. W. (1999). Dictionary of deities and demons in the Bible DDD (2nd extensively rev. ed.) (154). Leiden; Boston; Grand Rapids, Mich.: Brill; Eerdmans.
^Mulla, Mir S.; Chansang, Uruyakorn (2007). "Pestiferous nature, resting sites, aggregation, and host-seeking behavior of the eye flySiphunculina funicola (Diptera: Chloropidae) in Thailand".Journal of Vector Ecology.32 (2):292–301.doi:10.3376/1081-1710(2007)32[292:pnrsaa]2.0.co;2.PMID18260520.S2CID28636403.
^Gilead, L.; Mumcuoglu, K. Y.; Ingber, A. (16 August 2013). "The use of maggot debridement therapy in the treatment of chronic wounds in hospitalised and ambulatory patients".Journal of Wound Care.21 (2):78–85.doi:10.12968/jowc.2012.21.2.78.PMID22584527.
Blagoderov, V.A., Lukashevich, E.D. & Mostovski, M.B. (2002)). "Order Diptera". In:Rasnitsyn, A.P. and Quicke, D.L.J.The History of Insects, Kluwer pp.–227–240.
Colless, D.H. & McAlpine, D.K. (1991).Diptera (flies), pp. 717–786. In: The Division of Entomology. Commonwealth Scientific and Industrial Research Organisation, Canberra (spons.),The Insects of Australia. Melbourne University Press.
Hennig, Willi. "Diptera (Zweifluger)".Handb. Zool. Berl. 4 (2) (31):1–337. General introduction with key to World Families(in German).
Oldroyd, Harold (1965).The Natural History of Flies. W. W. Norton.
Séguy, Eugène (1924–1953).Diptera: recueil d'etudes biologiques et systematiques sur les Dipteres du Globe (Collection of biological and systematic studies on Diptera of the World). 11 vols.Part of Encyclopedie Entomologique, Serie B II: Diptera.
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