True flies are insects of the order Diptera, the name being derived
from the Greek δι- di- "two", and πτερόν pteron "wings".
Insects of this order use only a single pair of wings 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 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 maneuverability in flight, and
claws and pads on their feet enable them to cling to smooth surfaces.
Flies undergo complete metamorphosis; the eggs are laid on the larval
food-source and the larvae, which lack true limbs, develop in a
protected environment, often inside their food source. The pupa 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 considerable
ecological and human importance. Flies are important pollinators,
second only to the bees and their Hymenopteran relatives. Flies may
have been among the evolutionarily earliest pollinators responsible
for early plant pollination. Fruit flies are used as model organisms
in research, but less benignly, mosquitoes are vectors for malaria,
dengue, West Nile fever, yellow fever, encephalitis, and other
infectious diseases; and houseflies, commensal with humans all over
the world, spread food-borne illnesses. Flies can be annoyances
especially in some parts of the world where they can occur in large
numbers, buzzing and settling on the skin or eyes to bite or seek
fluids. Larger flies such as tsetse flies and screwworms cause
significant economic harm to cattle. Blowfly larvae, known as gentles,
and other dipteran larvae, known more generally as maggots, are used
as fishing bait and as food for carnivorous animals. They are also
used in medicine in debridement to clean wounds.
1 Taxonomy and phylogeny
1.1 Relationships to other insects
1.2 Relationships between fly subgroups and families
2 Anatomy and morphology
3 Life cycle and development
4.1 Anti-predator adaptations
5 In culture
5.2 Economic importance
8 Further reading
9 External links
Taxonomy and phylogeny
Relationships to other insects
Dipterans are endopterygotes, insects that undergo radical
metamorphosis. They belong to the Mecopterida, alongside the
Lepidoptera and Trichoptera. The
possession of a single pair of wings distinguishes most true flies
from other insects with "fly" in their names. However, some true flies
Hippoboscidae (louse flies) have become secondarily
The cladogram represents the current consensus view.
part of Endopterygota
Mecoptera (scorpionflies, hangingflies, 400 spp.) (exc. Boreidae)
Boreidae (snow scorpionflies, 30 spp.)
Siphonaptera (fleas, 2500 spp.)
Lepidoptera (butterflies and moths)
Hymenoptera (sawflies, wasps, ants, bees)
Fossil nematoceran in Dominican amber. Sandfly, Lutzomyia adiketis
(Psychodidae), Early Miocene, c. 20 million years ago
Relationships between fly subgroups and families
Fossil brachyceran in Baltic amber. Lower Eocene, c. 50 million years
The first true dipterans known are from the Middle
240 million years ago), and they became widespread during the Middle
and Late Triassic. Modern flowering plants did not appear until the
Cretaceous (around 140 million years ago), so the original dipterans
must have had a different source of nutrition other than nectar. Based
on the attraction of many modern fly groups to shiny droplets, it has
been suggested that they may have fed on honeydew produced by
sap-sucking bugs which were abundant at the time, and dipteran
mouthparts are well-adapted to softening and lapping up the crusted
residues. The basal clades in the
Diptera include the
Deuterophlebiidae and the enigmatic Nymphomyiidae. 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
in the Jurassic, some 180 million years ago. A third radiation took
place among the
Schizophora at the start of the Paleogene, 66 million
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),
Siphonaptera (fleas), Mecoptera
(scorpionflies) and possibly
Strepsiptera (twisted-wing flies).
Diptera has been grouped with
Mecoptera in the
Antliophora, but this has not been confirmed by molecular studies.
Diptera were traditionally broken down into two suborders, Nematocera
and Brachycera, distinguished by the differences in antennae. The
Nematocera are recognized by their elongated bodies and
many-segmented, often feathery antennae as represented by mosquitoes
and crane flies. The
Brachycera have rounder bodies and much shorter
antennae. Subsequent studies have identified the
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
Ptychopteromorpha (phantom and primitive crane-flies)
Blephariceromorpha (net-winged midges, etc)
Psychodomorpha (drain flies, etc)
Tipuloidea (crane flies)
Stratiomyomorpha (soldier flies, etc)
Xylophagomorpha (stink flies, etc)
Tabanomorpha (horse flies, snipe flies, etc)
Asiloidea (robber flies, bee flies, etc)
Empidoidea (dance flies, etc)
Aschiza (in part)
Phoroidea (flat-footed flies, etc)
Hippoboscoidea (louse flies, etc)
Muscoidea (house flies, dung flies, etc)
Oestroidea (blow flies, flesh flies, etc)
Acalyptratae (marsh flies, etc)
Abbreviations used in the cladogram:
Gauromydas heros is the largest fly in the world.
Flies are often abundant and are found in almost all terrestrial
habitats in the world apart from Antarctica. They include many
familiar insects such as house flies, blow flies, mosquitoes, gnats,
black flies, midges and fruit flies. More than 150,000 have been
formally described and the actual species diversity is much greater,
with the flies from many parts of the world yet to be studied
intensively. 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.
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. While most species have restricted distributions, a few
like the housefly (Musca domestica) are cosmopolitan. 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, while the
smallest is Euryplatea nanaknihali, which at 0.4 mm
(0.016 in) is smaller than a grain of salt.
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, and
amphibians. Flies are the second largest group of pollinators
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. It is thought that some
of the earliest pollinators of plants may have been flies.
The greatest diversity of gall forming insects are found among the
flies, principally in the family
Cecidomyiidae (gall midges). 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. Some families are mycophagous
or fungus feeding. These include the cave dwelling Mycetophilidae
(fungus gnats) whose larvae are the only diptera with bioluminescence.
Sciaridae are also fungus feeders. Some plants are pollinated by
fungus feeding flies that visit fungus infected male flowers.
The larvae of
Megaselia scalaris (Phoridae) are almost omnivorous and
consume such substances as paint and shoe polish. The larvae of
the shore flies (Ephydridae) and some
Chironomidae survive in extreme
environments including glaciers (
Diamesa sp., Chironomidae), hot
springs, geysers, saline pools, sulphur pools, septic tanks and even
crude oil (Helaeomyia petrolei). 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. Some brachycerans are agricultural pests, some
bite animals and humans and suck their blood, and some transmit
Anatomy and morphology
Morphology of Diptera
Morphology of Diptera and Biology of Diptera
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 hindermost segments modified for
Head of a horse-fly showing large compound eyes and stout piercing
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. Female horse-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 large diverticulae, allowing the insect to store small
quantities of liquid after a meal.
For visual course control, flies' optic flow field is analyzed by a
set of motion-sensitive neurons. 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. 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. 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. The ocelli are
concerned in the detection of changes in light intensity, enabling the
fly to react swiftly to the approach of an object.
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, 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. 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.
Some tachinid flies (Ormiinae) which are parasitoids of bush crickets,
have sound receptors to help them locate their singing hosts.
A cranefly, showing the hind wings reduced to drumstick-shaped
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
Some species of flies are exceptional in that they are secondarily
flightless. The only other order of insects bearing a single pair of
true, functional wings, in addition to any form of halteres, are the
Strepsiptera. In contrast to the flies, the
Strepsiptera bear their
halteres on the mesothorax and their flight wings on the
metathorax. 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. At
the tip of the limb is a pair of claws, and between these are
cushion-like structures known as pulvilli which provide adhesion.
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.
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.
Tabanid fly in 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. The wings
of the fly are attached to two kinds of muscles, those used to power
it and another set used for fine control.
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.
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. 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. 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
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. Some hover flies like Metasyrphus corollae have been
known to undertake long flights in response to aphid population
Males of fly species such as Cuterebra, many hover flies, bee
flies (Bombyliidae) and fruit flies (Tephritidae) maintain
territories within which they engage in aerial pursuit to drive away
intruding males and other species. While these territories may be
held by individual males, some species form leks with many males
aggregating in displays. 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.
Life cycle and development
Mating anthomyiid flies
Diptera go through a complete metamorphosis with four distinct life
stages – egg, larva, pupa and adult. 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
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
Life cycle of stable fly Stomoxys calcitrans, showing eggs, 3 larval
instars, pupa, and adult
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, 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.
Nematoceran larvae generally have well-developed eyes and antennae,
while those of Brachyceran larvae are reduced or modified.
Dipteran larvae have no jointed, "true legs", 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. 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. 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).
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.
The adult stage is usually short, its function only to mate and lay
eggs. The genitalia of female flies are rotated to a varying degree
from the position found in other insects. 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.
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.
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. The flies that feed on vertebrate blood have sharp
stylets that pierce the skin, the insects inserting anticoagulant
saliva and 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. 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.
The large bee-fly, Bombylius major, is a Batesian mimic of bees.
Further information: Anti-predator adaptation
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.
Many flies have evolved mimetic resemblances that aid their
Batesian mimicry is widespread with many hoverflies
resembling bees and wasps, ants and some species of
tephritid fruit fly resembling spiders. Some species of hoverfly
are myrmecophilous, their young live and grow within the nests of
ants. They are protected from the ants by imitating chemical odours
given by ant colony members. 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
Further information: Insects in culture
Petrus Christus's 1446 painting Portrait of a Carthusian has a fly
painted on a trompe l'oeil frame.
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. In Christian demonology,
Beelzebub is a demonic
fly, the "Lord of the Flies", and a god of the
Flies have appeared in literature since ancient Sumer. In a
Sumerian poem, a fly helps the goddess
Inanna when her husband Dumuzid
is being chased by galla demons. In the Mesopotamian versions of
the flood myth, the dead corpses floating on the waters are compared
to flies. Later, the gods are said to swarm "like flies" around
the hero Utnapishtim's offering. Flies appear on Old Babylonian
seals as symbols of Nergal, the god of death. Fly-shaped lapis
lazuli beads were often worn in ancient Mesopotamia, along with other
kinds of fly-jewellery.
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:
"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." 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. In Antony and Cleopatra,
Shakespeare similarly likens Cleopatra's hasty departure from the
Actium battlefield to that of a cow chased by a gadfly. More
recently, in 1962 the biologist Vincent Dethier wrote To Know a Fly,
introducing the general reader to the behaviour and physiology of the
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. Flies have inspired the design of miniature flying
robots. Steven Spielberg's 1993 film
Jurassic Park relied on the
DNA could be preserved in the stomach contents of a
blood-sucking fly fossilised in amber, though the mechanism has been
discounted by scientists.
Anopheles stephensi mosquito drinking human blood. The species
Dipterans are an important group of insects and have a considerable
impact on the environment. Some leaf-miner flies (Agromyzidae), fruit
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 mosquitoes (Culicidae),
blackflies (Simuliidae) and drain flies (Psychodidae) impact human
health, acting as vectors of 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).
In tropical regions, eye flies (Chloropidae) which visit the eye in
search of fluids can be a nuisance in some seasons.
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.
Diptera in research:
Drosophila melanogaster fruit fly larvae being
bred in tubes in a genetics laboratory
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. 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.
Blowflies feeding on the fresh corpse of a porcupine, Hystrix
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
Maggots used as animal feed at London Zoo
Some species of maggots such as blowfly larvae (gentles) and
bluebottle larvae (casters) are bred commercially; they are sold as
bait in angling, and as food for carnivorous animals (kept as pets, in
zoos, or for research) such as some mammals, 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
Casu marzu is a traditional Sardinian sheep milk cheese that contains
larvae of the cheese fly, Piophila casei.
Fly larvae can be used as a biomedical tool for wound care and
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. 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. These medicinal maggots are a safe and effective treatment
for chronic wounds.
The Sardinian cheese casu marzu is exposed to flies known as cheese
skippers such as Piophila casei, members of the family
Piophilidae. 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, but the ban has been lifted on the
grounds that the cheese is a traditional local product made by
^ 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. 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. 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.
^ Dickinson, Michael H. (1999-05-29). "Haltere–mediated equilibrium
reflexes of the fruit fly, Drosophila melanogaster". Philosophical
Transactions of the Royal Society of London B: Biological Sciences.
354 (1385): 903–916. doi:10.1098/rstb.1999.0442.
PMC 1692594 . PMID 10382224.
^ "Order Diptera: Flies". BugGuide. Iowa State University. Retrieved
26 May 2016.
^ Comstock, John Henry (1949). An Introduction to Entomology. Comstock
Publishing. p. 773.
^ Mayhew, Peter J. (2007). "Why are there so many insect species?
Perspectives from fossils and phylogenies". Biological Reviews. 82
(3): 425–454. doi:10.1111/j.1469-185X.2007.00018.x.
^ Peters, Ralph S.; Meusemann, Karen; Petersen, Malte; Mayer,
Christoph; Wilbrandt, Jeanne; Ziesmann, Tanja; Donath, Alexander;
Kjer, Karl M.; Aspöck, Ulrike; Aspöck, Horst; Aberer, Andre;
Stamatakis, Alexandros; Friedrich, Frank; Hünefeld, Frank; Niehuis,
Oliver; Beutel, Rolf G.; Misof, Bernhard (2014). "The evolutionary
history of holometabolous insects inferred from transcriptome-based
phylogeny and comprehensive morphological data". BMC Evolutionary
Biology. 14 (1): 52. doi:10.1186/1471-2148-14-52. PMC 4000048 .
^ "Taxon: Superorder Antliophora". The Taxonomicon. Retrieved 21
^ Hutson, A. M. (1984). Diptera: Keds, flat-flies & bat-flies
Hippoboscidae & Nycteribiidae). Handbooks for the Identification
of British Insects. 10 pt 7. Royal Entomological Society of London.
^ Yeates, David K.; Wiegmann, Brian. "
Endopterygota Insects with
complete metamorphosis". Tree of Life. Retrieved 24 May 2016.
^ Blagoderov, V. A.; Lukashevich, E. D.; Mostovski, M. B. (2002).
Diptera Linné, 1758. The true flies". In Rasnitsyn, A. P.;
Quicke, D. L. J. History of Insects. Kluwer Academic Publishers.
^ Downes, William L. Jr.; Dahlem, Gregory A. (1987). "Keys to the
Evolution of Diptera: Role of Homoptera". Environmental Entomology. 16
(4): 847–854. doi:10.1093/ee/16.4.847.
^ a b c Wiegmann, B. M.; Trautwein, M. D.; Winkler, I. S.; Barr, N.
B.; Kim, J.-W.; Lambkin, C.; Bertone, M. A.; Cassel, B. K.; et al.
(2011). "Episodic radiations in the fly tree of life". PNAS. 108 (14):
5690–5695. Bibcode:2011PNAS..108.5690W. doi:10.1073/pnas.1012675108.
PMC 3078341 . PMID 21402926.
^ Wiegmann,Brian; Yeates, David K. (2012). The Evolutionary Biology of
Flies. Columbia University Press. pp. 4–6.
^ B.B. Rohdendorf. 1964. Trans. Inst. Paleont., Acad. Sci. USSR,
Moscow, v. 100
^ Wiegmann, Brian M.; Yeates, David K. (29 November 2007). "Diptera
True Flies". Tree of Life. Retrieved 25 May 2016.
^ Yeates, David K.; Meier, Rudolf; Wiegmann, Brian. "Phylogeny of True
Flies (Diptera): A 250 Million Year Old Success Story in Terrestrial
Diversification". Flytree. Retrieved 24 May 2016.
^ "FLYTREE". Illinois Natural History Survey. Retrieved
^ Pape, Thomas; Bickel, Daniel John; Meier, Rudolf (2009). Diptera
Diversity: Status, Challenges and Tools. BRILL. p. 13.
^ Yeates, D. K.; Wiegmann, B. M. (1999). "Congruence and controversy:
toward a higher-level phylogeny of diptera". Annual Review of
Entomology. 44: 397–428. doi:10.1146/annurev.ento.44.1.397.
^ Wiegmann, Brian M.; Yeates, David K. (2007). "Diptera: True flies".
Tree of Life Web Project. Retrieved 27 May 2016.
^ a b c d Pape, Thomas; Beuk, Paul; Pont, Adrian Charles; Shatalkin,
Anatole I.; Ozerov, Andrey L.; Woźnica, Andrzej J.; Merz, Bernhard;
Bystrowski, Cezary; Raper, Chris; Bergström, Christer; Kehlmaier,
Christian; Clements, David K.; Greathead, David; Kameneva, Elena
Petrovna; Nartshuk, Emilia; Petersen, Frederik T.; Weber, Gisela;
Bächli, Gerhard; Geller-Grimm, Fritz; Van de Weyer, Guy; Tschorsnig,
Hans-Peter; de Jong, Herman; van Zuijlen, Jan-Willem; Vaňhara,
Jaromír; Roháček, Jindřich; Ziegler, Joachim; Majer, József;
Hůrka, Karel; Holston, Kevin; Rognes, Knut; Greve-Jensen, Lita;
Munari, Lorenzo; de Meyer, Marc; Pollet, Marc; Speight, Martin C. D.;
Ebejer, Martin John; Martinez, Michel; Carles-Tolrá, Miguel;
Földvári, Mihály; Chvála, Milan; Barták, Miroslav; Evenhuis, Neal
L.; Chandler, Peter J.; Cerretti, Pierfilippo; Meier, Rudolf;
Rozkosny, Rudolf; Prescher, Sabine; Gaimari, Stephen D.; Zatwarnicki,
Tadeusz; Zeegers, Theo; Dikow, Torsten; Korneyev, Valery A.; Richter,
Vera Andreevna; Michelsen, Verner; Tanasijtshuk, Vitali N.; Mathis,
Wayne N.; Hubenov, Zdravko; de Jong, Yde (2015). "Fauna Europaea:
Diptera – Brachycera". Biodiversity Data Journal. 3 (3): e4187.
doi:10.3897/BDJ.3.e4187. PMC 4339814 .
^ Marquez, J. G.; Krafsur, E. S. (2002-07-01). "Gene Flow Among
Housefly Populations (Musca domestica L.): A
Worldwide Survey of Mitochondrial Diversity". Journal of Heredity. 93
(4): 254–259. doi:10.1093/jhered/93.4.254. PMID 12407211.
^ Owen, James (10 December 2015). "World's Biggest
Fly Faces Two New
Challengers". National Geographic. Retrieved 21 July 2016.
^ Welsh, Jennifer (2 July 2012). "World's Tiniest
Fly May Decapitate
Ants, Live in Their Heads". Livescience. Retrieved 21 July 2016.
^ Strijbosch, H. (1980). "Mortality in a population of Bufo bufo
resulting from the fly Lucilia bufonivora". Oecologia. 45 (2):
^ Ssymank, Axel; Kearns, C. A.; Pape, Thomas; Thompson, F. Christian
(2008-04-01). "Pollinating Flies (Diptera): A major contribution to
plant diversity and agricultural production". Biodiversity. 9 (1–2):
^ Labandeira, Conrad C. (1998-04-03). "How Old Is the Flower and the
Fly?". Science. 280 (5360): 57–59.
^ Price, Peter W. (2005). "Adaptive radiation of gall-inducing
insects". Basic and Applied Ecology. 6 (5): 413–421.
^ Scheffer, Sonja J.; Winkler, Isaac S.; Wiegmann, Brian M. (2007).
"Phylogenetic relationships within the leaf-mining flies (Diptera:
Agromyzidae) inferred from sequence data from multiple genes".
Molecular Phylogenetics and Evolution. 42 (3): 756–75.
doi:10.1016/j.ympev.2006.12.018. PMID 17291785.
^ Sakai, Shoko; Kato, Makoto; Nagamasu, Hidetoshi (2000). "Artocarpus
Pollination Mutualism Mediated by a Male-Flower
Parasitic Fungus". American Journal of Botany. 87 (3): 440–445.
^ Disney, R.H.L. (2007). "Natural History of the Scuttle Fly,
Megaselia scalaris". Annual Review of Entomology. 53: 39–60.
doi:10.1146/annurev.ento.53.103106.093415. PMID 17622197.
^ a b Foote, B A (1995). "Biology of Shore Flies". Annual Review of
Entomology. 40: 417–442.
^ Gullan, P.J.; Cranston, P.S. (2009). The Insects: An Outline of
Entomology. John Wiley & Sons. p. 320.
^ a b c d e f g h i j Resh, Vincent H.; Cardé, Ring T. (2009).
Encyclopedia of Insects. Academic Press. pp. 284–297.
^ a b c d Hoell, H. V.; Doyen, J. T.; Purcell, A. H. (1998).
Insect Biology and Diversity (2nd ed.). Oxford
University Press. pp. 493–499. ISBN 0-19-510033-6.
^ Haag, Juergen; Borst, Alexander (2002). "Dendro-dendritic
interactions between motion-sensitive large-field neurons in the fly".
The Journal of Neuroscience. 22 (8): 3227–33.
^ Hausen, Klaus; Egelhaaf, Martin (1989). "Neural Mechanisms of Visual
Course Control in Insects". In Stavenga, Doekele Gerben; Hardie, Roger
Clayton. Facets of Vision. pp. 391–424.
doi:10.1007/978-3-642-74082-4_18. ISBN 978-3-642-74084-8.
^ Egelhaaf, Martin (1985). "On the neuronal basis of figure-ground
discrimination by relative motion in the visual system of the fly".
Biological Cybernetics. 52 (3): 195–209. doi:10.1007/BF00339948
^ Kimmerle, Bernd; Egelhaaf, Martin (2000). "Performance of fly visual
interneurons during object fixation". The Journal of Neuroscience. 20
(16): 6256–66. PMID 10934276.
^ Eckert, Hendrik (1980). "Functional properties of the H1-neurone in
the third optic
Ganglion of the Blowfly, Phaenicia". Journal of
Comparative Physiology. 135 (1): 29–39.
^ a b Ruppert, Edward E.; Fox, Richard, S.; Barnes, Robert D. (2004).
Invertebrate Zoology, 7th edition. Cengage Learning.
pp. 735–736. ISBN 978-81-315-0104-7.
^ Stocker, Reinhard F. (2005). "The organization of the chemosensory
system in Drosophila melanogaster: a rewiew". Cell and Tissue
Research. 275 (1): 3–26. doi:10.1007/BF00305372.
^ Zhu, Junwei J; Zhang, Qing-he; Taylor, David B; Friesen, Kristina A
(2016-09-01). "Visual and olfactory enhancement of stable fly
trapping". Pest Management Science. 72 (9): 1765–1771.
^ 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.
^ "Strepsiptera: Stylops". Insects and their Allies. CSIRO. Retrieved
25 May 2016.
^ Langer, Mattias G.; Ruppersberg, J. Peter; Gorb, Stanislav N.
(2004). "Adhesion Forces Measured at the Level of a Terminal Plate of
the Fly's Seta". Proceedings of the Royal Society B. 271 (1554):
2209–2215. doi:10.1098/rspb.2004.2850. JSTOR 4142949.
PMC 1691860 . PMID 15539345.
^ Gibb, Timothy J.; Oseto, Christian (2010).
Arthropod Collection and
Identification: Laboratory and Field Techniques. Academic Press.
p. 189. ISBN 978-0-08-091925-6.
^ Deora, Tanvi; Singh, Amit Kumar; Sane, Sanjay P. (3 February 2015).
"Biomechanical basis of wing and haltere coordination in flies".
Proceedings of the National Academy of Sciences. 112 (5): 1481–1486.
doi:10.1073/pnas.1412279112. PMC 4321282 .
^ Dickinson, Michael H; Tu, Michael S (1997-03-01). "The Function of
Dipteran Flight Muscle". Comparative
A: Physiology. 116 (3): 223–238.
^ Dickinson, Michael H. (2005). "The Initiation and Control of Rapid
Flight Maneuvers in Fruit Flies". Integrated Comparative Biology. 45
(2): 274–281. doi:10.1093/icb/45.2.274.
^ Sourakov, Andrei (2011). "Faster than a Flash: The Fastest Visual
Startle Reflex Response is Found in a Long-Legged Fly,Condylostylussp.
(Dolichopodidae)". Florida Entomologist. 94 (2): 367–369.
^ Townsend, Charles H.T. (1927). "On the Cephenemyia Mechanism and the
Daylight-Day Circuit of the Earth by Flight". Journal of the New York
Entomological Society. 35 (3): 245–252. JSTOR 25004207.
^ Langmuir, Irving (1938). "The speed of the deer fly". Science. 87
(2254): 233–234. doi:10.1126/science.87.2254.233.
^ Townsend, Charles H.T. (1939). "Speed of Cephenemyia". Journal of
the New York Entomological Society. 47 (1): 43–46.
^ Berenbaum, M. (1999). "Getting Up to Speed". American Entomologist.
45: 4–5. doi:10.1093/ae/45.1.4.
^ Johnson, C.G.; Taylor, L.R.; T.R.E. Southwood (1962). "High Altitude
Migration of Oscinella frit L. (Diptera: Chloropidae)". Journal of
Animal Ecology. 31 (2): 373–383. doi:10.2307/2148.
^ Svensson, BO G.; Janzon, Lars-ÅKE (1984). "Why does the hoverfly
Metasyrphus corollae migrate?". Ecological Entomology. 9 (3):
^ 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.
^ 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.
^ a b Becerril-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.
^ Alcock, John; Schaefer, John E. (1983). "Hilltop territoriality in a
Sonoran desert bot fly (Diptera: Cuterebridae)".
Animal Behaviour. 31
(2): 518. doi:10.1016/S0003-3472(83)80074-8.
^ Downes, J. A. (1969). "The Swarming and Mating Flight of Diptera".
Annual Review of Entomology. 14: 271–298.
^ a b Gullan, P.J.; Cranston, P.S. (2005). The Insects: An Outline of
Entomology 3rd Edition. John Wiley & Sons. pp. 499–505.
^ Brown, Lesley (1993). The New shorter Oxford English dictionary on
historical principles. Clarendon. ISBN 0-19-861271-0.
^ Lancaster, Jill; Downes, Barbara J. (2013). Aquatic Entomology.
Oxford University Press. p. 16.
^ Chapman, R. F. (1998). The Insects; Structure & Function.
Cambridge: Cambridge University Press.
^ Meier, Rudolf; Kotrba, Marion; Ferrar, Paul (August 1999).
Ovoviviparity and viviparity in the Diptera". Biological Reviews. 74
(3): 199–258. doi:10.1111/j.1469-185X.1999.tb00186.x.
^ Mcmahon, Dino P.; Hayward, Alexander (April 2016). "Why grow up? A
perspective on insect strategies to avoid metamorphosis". Ecological
Entomology: n/a–n/a. doi:10.1111/een.12313.
^ Gillott, Cedric (2005). Entomology (3 ed.). Springer.
^ Papavero, N. (1977). The World Oestridae (Diptera), Mammals and
Continental Drift. Springer. doi:10.1007/978-94-010-1306-2.
^ Collins, Robert (2004). What Eats Flies for Dinner?. Shortland
Mimosa. ISBN 978-0-7327-3471-8.
^ Gilbert, Francis (2004). The evolution of imperfect mimicry in
Insect Evolution. CABI.
^ Rashed, A.; Khan, M. I.; Dawson, J. W.; Yack, J. E.; Sherratt, T. N.
(2008). "Do hoverflies (Diptera: Syrphidae) sound like the Hymenoptera
they morphologically resemble?". Behavioral Ecology. 20 (2):
^ 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:
^ Whitman, D. W.; Orsak, L.; Greene, E. (1988). "Spider
Fruit Flies (Diptera: Tephritidae): Further Experiments on the
Deterrence of Jumping Spiders (Araneae: Salticidae) by Zonosemata
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 of Microdon (Diptera: Syrphidae) in the Pacific Northwest".
Journal of the Kansas Entomological Society. 63 (1): 175–178.
^ Godfray, H. C. J. (1994). Parasitoids: Behavioral and Evolutionary
Ecology. Princeton University Press. p. 299.
^ Leland Clifton Wyman (1983). "Navajo Ceremonial System". Handbook of
North American Indians (PDF). Humboldt State University. p. 539.
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
^ Leland Clifton Wyman; Flora L. Bailey (1964). Navaho Indian
Ethnoentomology. Anthropology Series. University of New Mexico Press.
^ "Native American
Fly Mythology". Native Languages of the Americas
^ "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.
^ a b c d e f Black, Jeremy; Green, Anthony (1992). Gods, Demons and
Symbols of Ancient Mesopotamia: An Illustrated Dictionary. The British
Museum Press. pp. 84–85. ISBN 0-7141-1705-6.
^ Belfiore, Elizabeth S. (2000). Murder among Friends: Violation of
Philia in Greek Tragedy. Oxford, England: Oxford University Press.
p. 47. ISBN 0-19-513149-5.
^ a b c Stagman, Myron (11 August 2010). Shakespeare's Greek Drama
Secret. Cambridge Scholars Publishing. pp. 205–208.
^ Walker, John Lewis (2002). Shakespeare and the Classical Tradition:
An Annotated Bibliography, 1961–1991. Taylor & Francis.
p. 363. ISBN 978-0-8240-6697-0.
^ Dethier, Vincent G. (1962). To Know a Fly. San Francisco:
Fly on the Wall". British Film Institute. Retrieved 21 July
^ Ma, Kevin Y.; Chirarattananon, Pakpong; Fuller, Sawyer B.; Wood,
Robert J. (2013-05-03). "Controlled Flight of a Biologically Inspired,
Insect-Scale Robot". Science. 340 (6132): 603–607.
doi:10.1126/science.1231806. PMID 23641114.
^ Gray, Richard (12 September 2013). "
Jurassic Park ruled out –
DNA could not survive in amber". Daily Telegraph. Retrieved
21 July 2016.
^ Mulla, Mir S.; Chansang, Uruyakorn. "Pestiferous nature, resting
sites, aggregation, and host-seeking behavior of the eye fly
Siphunculina funicola (Diptera: Chloropidae) in Thailand". Journal of
Vector Ecology. 32 (2): 292.
^ "Why use the fly in research?". YourGenome. 19 June 2015. Retrieved
27 May 2016.
^ Ashlock, P. D. (1974). "The Uses of Cladistics". Annual Review of
Ecology and Systematics. 5 (1): 81–99.
^ Joseph, Isaac; Mathew, Deepu G.; Sathyan, Pradeesh; Vargheese,
Geetha (2011). "The use of insects in forensic investigations: An
overview on the scope of forensic entomology". Journal of Forensic
Dental Sciences. 3 (2): 89–91. doi:10.4103/0975-1475.92154.
PMC 3296382 . PMID 22408328.
^ Ogunleye, R. F.; Edward, J. B. (2005). "Roasted maggots (Dipteran
larvae) as a dietary protein source for laboratory animals". African
Journal of Applied Zoology and Environmental Biology. 7:
^ Fleming, Nic (4 June 2014). "How insects could feed the food
industry of tomorrow". British Broadcasting Corporation. Retrieved 24
^ "Why are insects not allowed in animal feed?" (PDF). All About Feed.
August 2014. Archived from the original (PDF) on 11 August 2016.
Retrieved 24 May 2016.
^ Stegman, Sylvia; Steenvoorde, Pascal (2011). "
therapy" (PDF). Proceedings of the Netherlands Entomological Society
Meeting. 22: 61–66.
^ Diaz-Roa, A.; Gaona, M. A.; Segura, N. A.; Suárez, D.; Patarroyo,
M.A.; Bello, F. J. (August 2014). "Sarconesiopsis magellanica
(Diptera: Calliphoridae) excretions and secretions have potent
antibacterial activity". Acta Tropica. 136: 37–43.
doi:10.1016/j.actatropica.2014.04.018. PMID 24754920.
^ 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:
^ Berenbaum, May (2007). "A mite unappetizing" (PDF). American
Entomologist: 132–133. [permanent dead link]
^ Colangelo, Matt (9 October 2015). "A Desperate Search for Casu
Marzu, Sardinia's Illegal
Maggot Cheese". Food and Wine. Retrieved 24
^ Brones, Anna (15 April 2013). "Illegal food: step away from the
cheese, ma'am". The Guardian. Retrieved 26 May 2016.
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
pp. 717–786. In: The Division of Entomology. Commonwealth
Scientific and Industrial Research Organisation, Canberra (spons.),
The insects of Australia. Melbourne University Press.
Diptera (Zweifluger). Handb. Zool. Berl. 4 (2)
(31):1–337. General introduction with key to World Families. In
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
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
Diptera (Flies)" (PDF). United States Department of Agriculture,
Systematic Entomology Laboratory. Archived from the original on 18
September 2015. CS1 maint: BOT: original-url status unknown
Wikiquote has quotations related to: Flies
Wikimedia Commons has media related to Diptera.
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Wikisource has the text of the 1911 Encyclopædia Britannica article
The Systema Dipterorum Database site
The Diptera.info portal with galleries and discussion forums
FLYTREE – dipteran phylogeny
The Dipterists Forum – The Society for the study of flies
The World Catalog of
The Tree of Life Project
Fly: Anatomical Atlas at CSIRO
Authors of fly names
Systema Dipterorum Nomenclator
Dixidae (meniscus midges)
Corethrellidae (frog-biting midges)
Chaoboridae (phantom midges)
Thaumaleidae (solitary midges)
Simuliidae (black flies)
Ceratopogonidae (biting midges)
Chironomidae (non-biting midges)
Blephariceridae (net-winged midges)
Deuterophlebiidae (mountain midges)
Bibionidae (march flies, lovebugs)
Anisopodidae (wood gnats)
Sciaridae (dark-winged fungus gnats)
Cecidomyiidae (gall midges)
Scatopsidae (minute black scavenger flies, or dung midges)
Psychodidae (moth flies)
Ptychopteridae (phantom crane flies)
Tanyderidae (primitive crane flies)
Trichoceridae (winter crane flies)
Pediciidae (hairy-eyed craneflies)
Tipulidae (crane flies)
Apioceridae (flower-loving flies)
Asilidae (robber flies)
Bombyliidae (bee flies)
Hilarimorphidae (hilarimorphid flies)
Mydidae (mydas flies)
Scenopinidae (window flies)
Therevidae (stiletto flies)
Hybotidae (dance flies)
Dolichopodidae (long-legged flies)
Empididae (dagger flies, balloon flies)
Acroceridae (small-headed flies)
Nemestrinidae (tangle-veined flies)
Phoridae (scuttle flies, coffin flies, humpbacked flies)
Opetiidae (flat-footed flies)
Ironomyiidae (ironic flies)
Lonchopteridae (spear-winged flies)
Platypezidae (flat-footed flies)
Pipunculidae (big-headed flies)
Conopidae (thick-headed flies)
Pallopteridae (flutter flies)
Piophilidae (cheese flies)
Platystomatidae (signal flies)
Tephritidae (peacock flies)
Ulidiidae (picture-winged flies)
Micropezidae (stilt-legged flies)
Neriidae (cactus flies, banana stalk flies)
Diopsidae (stalk-eyed flies)
Psilidae (rust flies)
Coelopidae (kelp flies)
Sepsidae (black scavenger flies)
Sciomyzidae (marsh flies)
Sphaeroceridae (small dung flies)
Celyphidae (beetle-backed flies)
Chamaemyiidae (aphid flies)
Agromyzidae (leaf miner flies)
Aulacigastridae (sap flies)
Clusiidae (lekking, or druid flies)
Neurochaetidae (upside-down flies)
Curtonotidae (quasimodo flies)
Diastatidae (bog flies)
Ephydridae (shore flies)
Drosophilidae (vinegar and fruit flies)
Braulidae (bee lice)
Canacidae (beach flies)
Chloropidae (frit flies)
Milichiidae (freeloader flies)
Lonchaeidae (lance flies)
Anthomyiidae (cabbage flies)
Fanniidae (little house flies)
Muscidae (house flies, stable flies)
Scathophagidae (dung flies)
Calliphoridae (blow-flies: bluebottles, greenbottles)
Mystacinobiidae (New Zealand batfly)
Sarcophagidae (flesh flies)
Tachinidae (tachina flies)
Glossinidae (tsetse flies)
Hippoboscidae (louse flies)
Mormotomyiidae (frightful hairy fly)
Nycteribiidae (bat flies)
Streblidae (bat flies)
Pantophthalmidae (timber flies)
Stratiomyidae (soldier flies)
Xylomyidae (wood soldier flies)
Rhagionidae (snipe flies)
Athericidae (water snipe flies)
Tabanidae (horse and deer flies)
Xylophagidae (awl flies)
List of families of Diptera
Archaeognatha (jumping bristletails)
Thysanura (Zygentoma) (silverfish, firebrats)
Odonata (dragonflies, damselflies)
Phasmatodea (stick and leaf insects)
Notoptera (ice-crawlers, gladiators)
Orthoptera (crickets, wetas, grasshoppers, locusts)
Zoraptera (angel insects)
Blattodea (cockroaches, termites)
Psocodea (barklice, lice)
Hemiptera (cicadas, aphids, true bugs)
Hymenoptera (sawflies, wasps, ants, bees)
Strepsiptera (twisted-winged parasites)
Megaloptera (alderflies, dobsonflies, fishflies)
Neuroptera (net-winged insects: lacewings, mantidflies, antlions)
Mecoptera (scorpionflies) +
Diptera (gnats, mosquitoes, flies)
Lepidoptera (moths, butterflies)
Four most speciose orders are marked in bold
Italic are paraphyletic groups
Based on Sasaki et al. (2013)
Extinct incertae sedis families and genera are marked in italic
Insects in culture
In the arts
Insects in art
Insects in film
Insects in literature
Insects in music
List of insect-inspired songs
Insects on stamps
Insects in religion
Colorado potato beetle
Cottony cushion scale
Western corn rootworm
Insect bites and stings
Insect sting allergy
House longhorn beetle
Home-stored product entomology
Alfred Russel Wallace
Hans Zinsser (Rats, Lice and History)
Lafcadio Hearn (
Living things in culture
Fauna Europaea: 10877