Fly

Flies are insects of the order Diptera, the name being

derived from the Greek δι- di- "two", and πτερόν pteron Fly
"wing". Insects of this order use only a single pair of wings Temporal range: Middle Triassic – Recent
to fly, the 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. Diptera is a large
order containing an estimated 1,000,000 species including
horse-flies,[a] crane flies, hoverflies, mosquitoes and others,
although only about 125,000 species have been described.

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

Taxonomy and phylogeny

Relationships to other insects

Dipterans are holometabolans, insects that undergo radical
metamorphosis. They belong to the Mecopterida, alongside the
Mecoptera, Siphonaptera, Lepidoptera and Trichoptera.[3][4] 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
An Anthomyiidae species showing
wingless.[5][6] characteristic dipteran features:
large eyes, small antennae, sucking
The cladogram represents the current consensus view.[7] mouthparts, single pair of flying
wings, hindwings reduced to clublike
halteres
⁠ Hymenoptera (sawflies, wasps
⁠

⁠
⁠

⁠Neuropterida
⁠
⁠
⁠
⁠Neuropteroidea
⁠

⁠
⁠
⁠Coleopterida
⁠ ⁠
⁠

⁠Panorpida
⁠Holometabola
⁠
⁠Aparaglossata
⁠
⁠Amphiesmeno
 ⁠

⁠(Mecopterida)

⁠Antliophor
⁠

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.[8] 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
Fossil brachyceran in may have fed on honeydew produced by sap-sucking bugs which were
Baltic amber. Lower abundant at the time, and dipteran mouthparts are well-adapted to softening
Eocene, c. 50 million and lapping up the crusted residues.[9] The basal clades in the Diptera include
years ago the Deuterophlebiidae and the enigmatic Nymphomyiidae.[10] 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.[10]

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.[11]

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.[12][13] 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.[14][15]

⁠Ptychopteromorpha (phantom and primitive crane-flies)

Fossil nematoceran in
⁠⁠
Dominican amber.
⁠ Sandfly, Lutzomyia
⁠Culicomorpha (mosquitoes, blackflies and midges) adiketis (Psychodidae),
⁠ Early Miocene, c. 20
⁠ million years ago

⁠Blephariceromorpha (net-winged midges, etc)

⁠

⁠
⁠Bibionomorpha (gnats)
⁠

⁠
⁠Psychodomorpha (drain flies, sand flies, etc)
⁠

⁠
⁠ Tipuloidea (crane flies)
⁠
⁠Brachycera Stratiomyomorpha (soldier flies
⁠
⁠

⁠Xylophagomorpha (stink flies, etc)

⁠
⁠
⁠
⁠ Tabanomorpha (horse flies, snipe fl
⁠
⁠
⁠

⁠Mus
⁠Nemestrinoidea
⁠
⁠ Asiloidea (robber flies, bee flie
⁠
⁠Diptera ⁠
⁠
 ⁠

⁠ ⁠ Empidoidea (dance flies, etc)

⁠
⁠
⁠Aschiza (in part)
⁠ ⁠
⁠ ⁠Phoroidea (flat-footed flies
⁠

⁠ Syrphoidea (hoverflie
⁠
⁠
Hippoboscoidea
⁠ ⁠
⁠
⁠Ere
⁠ ⁠Cyc
⁠ ⁠ Muscoidea (ho
⁠ ⁠
⁠ ⁠Cal ⁠etc)
⁠ ⁠
⁠Sch ⁠
⁠ Oestroidea (blo
⁠
⁠
⁠

⁠ Acalyptratae (ma
⁠

Abbreviations used in the cladogram:

Cal=Calyptratae
Cyc=Cyclorrhapha
Ere=Eremoneura
Mus=Muscomorpha
Sch=Schizophora

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, more robust flies with short antennae.
Many nematoceran larvae are 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]
Gauromydas heros is the largest fly
Brachycera are ecologically very diverse, with many being
in the world.
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

Flies are adapted for aerial movement and typically have short and streamlined bodies. The first tagma of
the fly, the head, bears the eyes, the antennae, and the mouthparts (the labrum, labium, mandible, and
maxilla make up the mouthparts). The second tagma, the thorax, 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][34] 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
Head of a horse-fly showing large
groups.[34] Female horse-flies use knife-like mandibles and
compound eyes and stout piercing
mouthparts maxillae to make a cross-shaped incision in the host's skin and
then lap up the blood that flows. The gut includes large
diverticulae, allowing the insect to store small quantities of liquid
after a meal.[35]

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
A head of a fly, showing the two
stabilizing motor corrections midflight with respect to yaw.[40]
compound eyes and three simple The ocelli are concerned in the detection of changes in light
eyes clearly. intensity, enabling the fly to react swiftly to the approach of an
object.[41]

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,
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 flies such as the ectoparasitic Nycteribiidae and Streblidae 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 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
A crane fly, showing the hind wings
adapted for reproduction. Each segment is made up of a dorsal and
reduced to drumstick-shaped
a ventral sclerite, connected by an elastic membrane. In some halteres
females, the sclerites are rolled into a flexible, telescopic
ovipositor.[34]

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.[48]
The wings of the fly are attached to two kinds of muscles, those
used to power it and another set used for fine control.[49] 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.[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 it was shown to be physically impossible as well as
incorrect by Irving Langmuir. 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, such
as A. freeborni,[62] 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.[63]

Life cycle and development

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 Mating anthomyiid flies

become desiccated if exposed to the air. Apart from the

Brachycera, 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.[34][64] The spiracles in the larva and pupa do not have any internal mechanical closing
device.[65]

Some other anatomical distinction exists between the larvae of the

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
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,[66] 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
Life cycle of stable fly Stomoxys
is an adaptation to food such as carrion, decaying detritus, or host
calcitrans, showing eggs, 3 larval
instars, pupa, and adult
tissues surrounding endoparasites.[35] Nematoceran larvae
generally have well-developed eyes and antennae, while those of
Brachyceran larvae are reduced or modified.[67]

Dipteran larvae have no jointed, "true legs",[64] 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.[68] 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.[69] 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).[70][71]

Pupa
The pupae 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 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 forces its way out.[34]

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.[72] 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] More primitive groups mates in the air during swarming, but most of the more advanced
species with a 360° torsion mate on a substrate.[73]

Ecology
As ubiquitous insects, dipterans play an important role at various trophic 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.[34]

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
A calliphorid "bubbling" 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.[74] 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.[34]

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[75] or possibly to cool by evaporation.[76] 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.[77] Adults of Ephydra
hians forage underwater, and have special hydrophobic hairs that trap a bubble of air that lets them
breathe underwater.[78]

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.[79] Many flies have evolved mimetic
resemblances that aid their protection. Batesian mimicry is
widespread with many hoverflies resembling bees and
wasps,[80][81] ants[82] and some species of tephritid fruit fly
resembling spiders.[83] Some species of hoverfly are
The large bee-fly, Bombylius major,
myrmecophilous—their young live and grow within the nests of
is a Batesian mimic of bees.
ants. They are protected from the ants by imitating chemical
odours given by ant colony members.[84] 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.[85]
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.[86]

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.[87][88][89] In Christian
demonology, Beelzebub is a demonic fly, the "Lord of the Flies", and a
god of the Philistines.[90][91][92]

Flies have appeared in literature since ancient Sumer.[93] In a Sumerian

poem, a fly helps the goddess Inanna when her husband Dumuzid is being
chased by galla demons.[93] In the Mesopotamian versions of the flood
myth, the dead corpses floating on the waters are compared to flies.[93]
Later, the gods are said to swarm "like flies" around the hero
Utnapishtim's offering.[93] Flies appear on Old Babylonian seals as
Petrus Christus's 1446
painting Portrait of a
symbols of Nergal, the god of death.[93] Fly-shaped lapis lazuli beads
Carthusian has a musca were often worn in ancient Mesopotamia, along with other kinds of fly-
depicta (painted fly) on a jewellery.[93]
trompe-l'œil frame.
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.[94]

In a little-known Greek myth, a very chatty and talkative maiden named Myia (meaning "fly") enraged
the moon-goddess Selene by attempting to seduce her lover, the sleeping Endymion, 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.[95][96] 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:[97][98] "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."[98] 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.[98] In Antony and Cleopatra, Shakespeare similarly likens
Cleopatra's hasty departure from the Actium battlefield to that of a cow chased by a gadfly.[99] 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.[100]
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.[101]

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.[102] Flies have inspired the design of miniature flying robots.[103] 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.[104]

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 An Anopheles stephensi mosquito
such as mosquitoes (Culicidae), blackflies (Simuliidae) and drain drinking human blood. The species
flies (Psychodidae) impact human health, acting as vectors of carries malaria.
major 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).[34] In tropical regions, eye
flies (Chloropidae) which visit the eye in search of fluids can be a nuisance in some seasons.[105]

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.[34]

Uses
Drosophila melanogaster, a fruit fly, has long been used as a model 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.[106] 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.[107]
 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
Diptera in research: Drosophila
particular habitats such as woodland.[108]
melanogaster fruit fly larvae being
bred in tubes in a genetics
Some species of maggots
laboratory 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,[109] 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 Casu marzu is a traditional
to the inclusion of insects in their food, and the use of insects in Sardinian sheep milk cheese that
animal feed remains illegal in areas such as the European contains larvae of the cheese fly,
Union.[110][111] Piophila casei.

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.[112] 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.[113] These medicinal maggots are a safe and
effective treatment for chronic wounds.[114]

The Sardinian cheese casu marzu is exposed to flies known as cheese skippers such as Piophila casei,
members of the family Piophilidae.[115] 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,[116] but the ban has been lifted on the grounds that
the cheese is a traditional local product made by traditional methods.[117]

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 covering 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.[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 as house-fly and
dragon-fly are widely used.[2] 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://palaeo
entomolog.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 (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.
Séguy, Eugène (1950). La Biologie des Dipteres.
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 (https://www.sel.barc.usda.gov/diptera/names/BDWDs
our.pdf) (PDF) 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://www.ento.csiro.au/biology/fly/fly.php) at CSIRO

Drawing Wing venation (http://www.drawwing.org/insects/diptera)
Describers

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

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

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