Fly

Flies are insects of the order Diptera, the name being derived
from the Greek δι- di- "two", and πτερόν pteron "wing". Fly
Insects of this order use only a single pair of wings to fly, the Temporal range: Middle Triassic – Recent
hindwings having evolved into advanced mechanosensory
organs known as halteres, which act as high-speed sensors of
rotational movement and allow dipterans to perform
advanced aerobatics.[1] Diptera is a large order containing an
estimated 1,000,000 species including horse-flies,[a] crane
flies, hoverflies and others, although only about 125,000
species have been described.[4]

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. Their wing arrangement gives
them great maneuverability in flight, and claws and pads on
their feet enable them to cling to smooth surfaces. Flies
undergo complete metamorphosis; the eggs are often laid on Diptera from different families:
the larval food-source and the larvae, which lack true limbs,
develop in a protected environment, often inside their food Housefly (Muscidae) (top left)
source. Other species like Metopia argyrocephala are Haematopota pluvialis (Tabanidae)
ovoviviparous, opportunistically depositing hatched or (top right)
hatching maggots instead of eggs on carrion, dung, decaying Ctenophora pectinicornis (Tipulidae)
material, or open wounds of mammals. The pupa is a tough (mid left)
capsule from which the adult emerges when ready to do so; Ochlerotatus notoscriptus
flies mostly have short lives as adults. (Culicidae) (mid right)
Milesia crabroniformis (Syrphidae)
Diptera is one of the major insect orders and of considerable (bottom left)
ecological and human importance. Flies are important Holcocephala fusca (Asilidae)
pollinators, second only to the bees and their Hymenopteran (bottom right)
relatives. Flies may have been among the evolutionarily
earliest pollinators responsible for early plant pollination.
Scientific classification
Fruit flies are used as model organisms in research, but less
benignly, mosquitoes are vectors for malaria, dengue, West Kingdom: Animalia
Nile fever, yellow fever, encephalitis, and other infectious
Phylum: Arthropoda
diseases; and houseflies, commensal with humans all over the
world, spread food-borne illnesses. Flies can be annoyances Class: Insecta
especially in some parts of the world where they can occur in
Superorder: Panorpida
large numbers, buzzing and settling on the skin or eyes to bite
or seek fluids. Larger flies such as tsetse flies and (unranked): Antliophora
screwworms cause significant economic harm to cattle.
Order: Diptera
Blowfly larvae, known as gentles, and other dipteran larvae,
Linnaeus, 1758

Suborders
known more generally as maggots, are used as fishing bait Nematocera (includes Eudiptera)
and as food for carnivorous animals. They are also used in Brachycera
medicine in debridement to clean wounds.

Contents
Taxonomy and phylogeny
Relationships to other insects
Relationships between subgroups and families
Diversity
Anatomy and morphology
Flight
Life cycle and development An Anthomyiidae species showing
characteristic dipteran features: large
Larva
eyes, small antennae, sucking
Pupa
mouthparts, single pair of flying
Adult wings, hindwings reduced to clublike
Ecology halteres
Anti-predator adaptations
Human interaction and cultural depictions
Symbolism
Economic importance
Uses
Hazards
Notes
References
Further reading
External links

Taxonomy and phylogeny

Relationships to other insects

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]

   Diptera
 

Mecoptera (scorpionflies, hangingflies, 400

 
  spp.) (exc. Boreidae)
Antliophora
 

  Boreidae (snow scorpionflies, 30 spp.)

   
 
Mecopterida
   
 
part of Endopterygota
    Siphonaptera (fleas, 2500 spp.)
 

  Trichoptera (caddisflies)
 
 
 
  Lepidoptera (butterflies and moths)
 

  Hymenoptera (sawflies, wasps, ants, bees)

 

Relationships between subgroups and families

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] 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
Fossil brachyceran in fed on honeydew produced by sap-sucking bugs which were abundant at the
Baltic amber. Lower time, and dipteran mouthparts are well-adapted to softening and lapping up the
Eocene, c. 50 million crusted residues.[10] The basal clades in the Diptera include the
years ago Deuterophlebiidae and the enigmatic Nymphomyiidae.[11] 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.[11]
The phylogenetic position of Diptera has been controversial. The monophyly of holometabolous 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 of Mecopterida, along with Lepidoptera (butterflies and moths), Trichoptera
(caddisflies), Siphonaptera (fleas), Mecoptera (scorpionflies) and possibly Strepsiptera (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 and
Brachycera, 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 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 Fossil nematoceran in
subject of ongoing research. The following cladogram is based on the Dominican amber.
FLYTREE project.[11][15] Sandfly, Lutzomyia
adiketis (Psychodidae),
Early Miocene, c. 20
million years ago
   Ptychopteromorpha (phantom and primitive crane-flies)
 
 
 

  Culicomorpha (mosquitoes)
 

Nematocera
  

  Blephariceromorpha (net-winged midges, etc)

 

 
Abbreviations used in the cladogram:

Cal=Calyptratae
Cyc=Cyclorrhapha
Ere=Eremoneura
Mus=Muscomorpha
Sch=Schizophora
  
Tab=Tabanomorpha
  Bibionomorpha (gnats)
 
Diversity

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.[16][17] The suborder Nematocera include generally
small, slender insects with long antennae such as mosquitoes, gnats,
midges and crane-flies, while the Brachycera includes broader, Gauromydas heros is the largest fly
more robust flies with short antennae. Many nematoceran larvae are in the world.
aquatic.[18] 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.[19] While
most species have restricted distributions, a few like the housefly (Musca domestica) are cosmopolitan.[20]
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,[21] while the smallest is Euryplatea nanaknihali, which at 0.4  mm (0.016  in) is smaller
than a grain of salt.[22]

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,[19] and
amphibians.[23] 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.[24] It is thought that some of the earliest
pollinators of plants may have been flies.[25]

The greatest diversity of gall forming insects are found among the flies, principally in the family
Cecidomyiidae (gall midges).[26] 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.[27]
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.[28]

The larvae of Megaselia scalaris (Phoridae) are almost omnivorous and consume such substances as paint
and shoe polish.[29] The Exorista mella (Walker) fly are considered generalists and parasitoids of a variety
of hosts.[30] The larvae of the shore flies (Ephydridae) and some Chironomidae survive in extreme
environments including glaciers (Diamesa sp., Chironomidae[31]), hot springs, geysers, saline pools,
sulphur pools, septic tanks and even crude oil (Helaeomyia petrolei[31]).[19] 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.[32] Some brachycerans are agricultural pests, some bite animals
and humans and suck their blood, and some transmit diseases.[19]

Anatomy and morphology

   aerial movement and typically have short and streamlined bodies. The first tagma of
Flies are adapted for
the fly, the head, bears  the eyes, the antennae, and the mouthparts (the labrum, labium, mandible, and
Psychodomorpha
maxilla make up the mouthparts). The second(drain flies,
tagma, the sand flies,
thorax, etc)
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 the halteres, which help to balance the insect during
flight. The third tagma is the abdomen consisting of 11 segments, some of which may be fused, and with
the 3 hindmost segments modified for reproduction.[33] Some Dipterans are mimics and can only be
distinguished from their models by very careful inspection. An example of this is Spilomyia longicornis,
which is a fly but mimics a vespid wasp.

Flies have a mobile head with a pair of large compound eyes on the
sides of the head, and in most species, three small ocelli 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.[33]
Female horse-flies use knife-like mandibles and maxillae to make a
Head of a horse-fly showing large
cross-shaped incision in the host's skin and then lap up the blood
compound eyes and stout piercing
mouthparts
that flows. The gut includes large diverticulae, allowing the insect
to store small quantities of liquid after a meal.[34]

For visual course control, flies' optic flow field is analyzed by a set
of motion-sensitive neurons.[35] 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.[36] 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.[37][38] 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.[39] The
 
ocelli are concerned in the detection of changes in light intensity,
enabling the fly to react swiftly to the approach of an object.[40]
A head of a fly, showing the two
Like other insects, flies have chemoreceptors that detect smell and
compound eyes and three simple
taste, and mechanoreceptors that respond to touch. The third
eyes clearly.
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,[41] 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.[40] 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.[42] Some tachinid flies (Ormiinae) which are parasitoids of
bush crickets, have sound receptors to help them locate their singing hosts.[43]

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
   [34] Some species of flies are exceptional
Muscomorpha infraorder.
in that they are secondarily flightless.
  The only (crane
Tipuloidea other order
flies) 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.[44] 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.[33]
At the tip of the limb is a pair of claws, and between these are
cushion-like structures known as pulvilli which provide A cranefly, showing the hind wings
adhesion.[45] reduced to drumstick-shaped
halteres
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.[46] 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.[33]

Flight

Flies are capable of great manoeuvrability during flight due to the

presence of the halteres. These act as gyroscopic 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.[47] The
wings of the fly are attached to two kinds of muscles, those used to
power it and another set used for fine control.[48] Tabanid fly in flight
 
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°, 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.[49]

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.[50] 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.[51] 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.[52][53][54]

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.[55] Some hover flies like Metasyrphus corollae have been known to undertake long flights in
response to aphid population spurts.[56]
Males of fly species such  as Cuterebra, many hover flies,[57] bee flies (Bombyliidae)[58] and fruit flies
(Tephritidae)[59] maintain territories within which they engage in aerial pursuit to drive away intruding
  Stratiomyomorpha (soldier flies, etc)
males and other species.[60] While these territories
  may be held by individual males, some species, such as
A. freeborni, [61] form leks with many males aggregating in displays.[59] 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]
  Xylophagomorpha (stink flies, etc)
Tab  
Life cycle and development  

 
Diptera go through a complete metamorphosis  withTabanomorpha
four distinct life (horse flies, snipe flies, etc)
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, Mating anthomyiid flies
inside their hosts. They tend to have thin cuticles and become
desiccated if exposed to the air. Apart from the Brachycera, most
dipteran larvae have sclerotinised 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.[33][63]

Some other anatomical distinction exists between the larvae of the

Brachycera mosquitoes; in the Brachycera, the head of the larva is not clearly
Life cycle of stable fly Stomoxys
calcitrans, showing eggs, 3 larval
instars, pupa, and adult
Nematocera and the Brachycera. Especially in the Brachycera, little
demarcation is seen between the thorax and abdomen, though the
demarcation may be visible in many Nematocera, such as
distinguishable from the rest of the body, and few, if any, sclerites
are present. Informally, such brachyceran larvae are called
maggots,[64] 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 as cerci. This lack of features
is an adaptation to food such as carrion, decaying detritus, or host
tissues surrounding endoparasites.[34] Nematoceran larvae
generally have well-developed eyes and antennae, while those of
Brachyceran larvae are reduced or modified.[65]

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  inNemestrinoidea
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]
  Asiloidea (robber flies, bee flies, etc)
 
Pupa

The pupae take various forms. In some groups, particularly   Empidoidea

the Nematocera,
(dance flies,the etc)
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    Aschiza (in part)
other groups, described as "coarctate", the
appendages are not visible. In these, the outer surface is a puparium, 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 inflates a balloon-like structure on its head,  and way out.[33]flies, etc)
forces its(flat-footed
Phoroidea
 

Adult
  Syrphoidea (hoverflies)
The adult stage is usually short, its function is only to mate and  lay eggs. The genitalia of male flies are
Mus found in other insects.[70] In some flies, this is a temporary
rotated to a varying degree from the position 
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,(louse
Hippoboscoidea 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,Ere   flies,
facing in the same direction, butetc)
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
Cyc
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. (house
Muscoidea Flies occur in
flies,
  [34]
large populations due to their ability to mate effectively and quickly during
Cal the   mating season.
  dung flies, etc)
   
Ecology Sch  
   
Oestroidea (blow flies,
As ubiquitous insects, dipterans play an important role at various
 
trophic levels both as consumers and  flesh
as prey. In etc)
flies, 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 oneAcalyptratae
of the most(marsh flies,
prevalent etc)
feeding
 
behaviours. The fruit or detritus
  is consumed along with the
associated micro-organisms, a sieve-like filter in the pharynx being
A calliphorid "bubbling" 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.[33]
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.[71] 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.[33]

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 will regurgitate fluid in a
behaviour termed as "bubbling" which has been thought to help the insects evaporate water and
concentrate food[72] or possibly to cool by evaporation.[73] Some adult diptera are known for
kleptoparasitism 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 genus Bengalia are known to steal food carried by ants.[74] Adults of Ephydra hians
forage underwater, and have special hydrophobic hairs that trap a bubble of air that lets them breathe
underwater.[75]

Anti-predator adaptations

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.[76] Many flies have evolved mimetic resemblances that aid
their protection. Batesian mimicry is widespread with many
hoverflies resembling bees and wasps,[77][78] ants[79] and some
species of tephritid fruit fly resembling spiders.[80] Some species of
hoverfly are myrmecophilous, their young live and grow within the
nests of ants. They are protected from the ants by imitating The large bee-fly, Bombylius major,
chemical odours given by ant colony members.[81] Bombyliid bee is a Batesian mimic of bees.
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.[82]

In contrast, Drosophila subobscura, a species of fly in the genus Drosophila, lacks a category of hemocytes
that are present in other studied species of Drosophila, leading to an inability to defend against parasitic
attacks, a form of innate immunodeficiency.[83]

Human interaction and cultural depictions

Symbolism
 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.[84][85][86] In Christian
demonology, Beelzebub is a demonic fly, the "Lord of the Flies", and a god
of the Philistines.[87][88][89]

Flies have appeared in literature since ancient Sumer.[90] In a Sumerian

poem, a fly helps the goddess Inanna when her husband Dumuzid is being
chased by galla demons.[90] In the Mesopotamian versions of the flood
myth, the dead corpses floating on the waters are compared to flies.[90]
Later, the gods are said to swarm "like flies" around the hero Utnapishtim's
offering.[90] Flies appear on Old Babylonian seals as symbols of Nergal,
Petrus Christus's 1446
the god of death.[90] Fly-shaped lapis lazuli beads were often worn in
painting Portrait of a ancient Mesopotamia, along with other kinds of fly-jewellery.[90]
Carthusian has a musca
In Prometheus Bound, which is attributed to the Athenian tragic playwright
depicta (painted fly) on a
Aeschylus, a gadfly sent by Zeus's wife Hera pursues and torments his
trompe-l'œil frame.
mistress Io, who has been transformed into a cow and is watched
constantly by the hundred eyes of the herdsman Argus:[91][92] "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." [92] 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.[92] In Antony and Cleopatra, Shakespeare similarly
likens Cleopatra's hasty departure from the Actium battlefield to that of a cow chased by a gadfly.[93] 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.[94]

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.[95]

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.[96] Flies have inspired the design of miniature flying robots.[97] 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.[98]

Economic importance

Dipterans are an important group of insects and have a considerable

impact on the environment. Some leaf-miner flies (Agromyzidae),
fruit flies (Tephritidae and Drosophilidae) and gall midges
(Cecidomyiidae) are pests of agricultural crops; others such as
tsetse flies, screwworm and botflies (Oestridae) attack livestock,
causing wounds, spreading disease, and creating significant
economic harm. See article: Parasitic flies of domestic animals. A
few can even cause myiasis in humans. Still others such as
An Anopheles stephensi mosquito
mosquitoes (Culicidae), blackflies (Simuliidae) and drain flies
drinking human blood. The species
(Psychodidae) impact human health, acting as vectors of major carries malaria.
tropical diseases. Among these, Anopheles mosquitoes transmit
malaria, filariasis, and arboviruses; Aedes aegypti mosquitoes carry dengue fever and the Zika virus;
blackflies carry river blindness; sand flies carry leishmaniasis. Other dipterans are a nuisance to humans,
especially when present in large numbers; these include houseflies, which contaminate food and spread
food-borne illnesses; the biting midges and sandflies (Ceratopogonidae) and the houseflies and stable flies
(Muscidae).[33] In tropical regions, eye flies (Chloropidae) which visit the eye in search of fluids can be a
nuisance in some seasons.[99]

Many dipterans serve roles that are useful to humans. Houseflies, blowflies and fungus gnats
(Mycetophilidae) are scavengers and aid in decomposition. Robber flies (Asilidae), tachinids (Tachinidae)
and dagger flies and balloon flies (Empididae) are predators and parasitoids of other insects, helping to
control a variety of pests. Many dipterans such as bee flies (Bombyliidae) and hoverflies (Syrphidae) are
pollinators of crop plants.[33]

Uses

Drosophila melanogaster, a fruit fly, has long been used as a model

Diptera in research: Drosophila
melanogaster fruit fly larvae being
bred in tubes in a genetics laboratory
organism in research because of the ease with which it can be bred
and reared in the laboratory, its small genome, and the fact that
many of its genes have counterparts in higher eukaryotes. A large
number of genetic studies have been undertaken based on this
species; these have had a profound impact on the study of gene
expression, gene regulatory mechanisms and mutation. Other
studies have investigated physiology, microbial pathogenesis and
development among other research topics.[100] The studies on
dipteran relationships by Willi Hennig helped in the development of
cladistics, techniques that he applied to morphological characters
but now adapted for use with molecular sequences in
phylogenetics.[101]

Maggots found on corpses are useful to forensic entomologists. Maggot species can be identified by their
anatomical features and by matching their DNA. 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 family Histeridae. 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 as woodland.[102]

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,[103] 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.[104][105]
Casu marzu is a traditional Sardinian
Fly larvae can be used as a biomedical tool for wound care and sheep milk cheese that contains
treatment. Maggot debridement therapy (MDT) is the use of blow larvae of the cheese fly, Piophila
fly larvae to remove the dead tissue from wounds, most commonly casei.
being amputations. Historically, this has been used for centuries,
both intentional and unintentional, on battlefields and in early
hospital settings.[106] 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.[107] These medicinal maggots are a safe and effective treatment for chronic wounds.[108]

The Sardinian cheese casu marzu is exposed to flies known as cheese skippers such as Piophila casei,
members of the family Piophilidae.[109] 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,[110] but the ban has been lifted on the grounds that the
cheese is a traditional local product made by traditional methods.[111]

Hazards
Flies are a health hazard and are attracted to toilets because of their smell. The New Scientist magazine
suggested a trap for these flies. A pipe acting as a chimney was fitted to the toilet which let in some light to
attract these flies up to the end of this pipe where a gauze prevented escape to the air outside so that they
were trapped and died. Toilets are generally dark inside particularly if the door is closed.

Notes
a. Some authors draw a distinction in writing the common names of insects. True flies are in
their view best written as two words, such as crane 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.[2] In practice, however, this is a comparatively new convention; especially in older
books, names like "saw fly" and "caddis fly", or hyphenated forms such as house-fly and
dragon-fly are widely used.[3] In any case, non-entomologists cannot, in general, be
expected to tell dipterans, "true flies", from other insects, so it would be unrealistic to expect
rigour in the use of common names. Also, exceptions to this rule occur, such as the hoverfly,
which is a true fly, and the Spanish fly, a type of blister beetle.

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Further reading
Blagoderov, V.A., Lukashevich, E.D. & Mostovski, M.B. 2002. Order Diptera (http://palaeoent
omolog.ru/New/diptera.html). 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 The Natural History of Flies. W. W. Norton. 1965.
Séguy, Eugène 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. 1924–1953.
Séguy, Eugène La Biologie des Dipteres 1950.
Thompson, F. Christian. "Sources for the Biosystematic Database of World Diptera (Flies)"
(https://web.archive.org/web/20150918204612/http://www.sel.barc.usda.gov/diptera/names/
 BDWDsour.pdf) (PDF). United States Department of Agriculture, Systematic Entomology
Laboratory. Archived from the original on 18 September 2015.

External links
General

The Systema Dipterorum Database site (https://web.archive.org/web/20100405091526/htt

p://www.diptera.org/)
The Diptera.info portal with galleries and discussion forums (http://www.diptera.info)
FLYTREE – dipteran phylogeny (http://wwx.inhs.illinois.edu/research/flytree/) Archived (http
s://web.archive.org/web/20200413095925/https://www.inhs.illinois.edu/research/flytree/) 13
April 2020 at the Wayback Machine
The Dipterists Forum (http://www.dipteristsforum.org.uk/) – The Society for the study of flies
BugGuide (http://bugguide.net/node/view/55)
The World Catalog of Fossil Diptera (http://hbs.bishopmuseum.org/fossilcat/)
The Tree of Life Project (http://tolweb.org/tree?group=Diptera)

Anatomy

Fly: Anatomical Atlas (https://web.archive.org/web/20120118040549/http://www.ces.csiro.au/

biology/fly/fly.html) at CSIRO
Drawing Wing venation (http://www.drawwing.org/insects/diptera)

Describers

Authors of fly names (http://hbs.bishopmuseum.org/publications/pdf/tr51.pdf)

Systema Dipterorum Nomenclator (https://web.archive.org/web/20100405091526/http://ww
w.diptera.org/)

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