Fly Flies Are Insects of The Order Diptera, The Name Being Derived
Fly Flies Are Insects of The Order Diptera, The Name Being Derived
Flies are insects of the order Diptera, the name being derived
from the Greek δι- di- "two", and πτερόν pteron "wing". Fly
Insects of this order use only a single pair of wings to fly, the Temporal range: Middle Triassic – Recent
hindwings having evolved into advanced mechanosensory
organs known as halteres, which act as high-speed sensors of
rotational movement and allow dipterans to perform
advanced aerobatics.[1] Diptera is a large order containing an
estimated 1,000,000 species including horse-flies,[a] crane
flies, hoverflies and others, although only about 125,000
species have been described.[4]
Suborders
known more generally as maggots, are used as fishing bait Nematocera (includes Eudiptera)
and as food for carnivorous animals. They are also used in Brachycera
medicine in debridement to clean wounds.
Contents
Taxonomy and phylogeny
Relationships to other insects
Relationships between subgroups and families
Diversity
Anatomy and morphology
Flight
Life cycle and development An Anthomyiidae species showing
characteristic dipteran features: large
Larva
eyes, small antennae, sucking
Pupa
mouthparts, single pair of flying
Adult wings, hindwings reduced to clublike
Ecology halteres
Anti-predator adaptations
Human interaction and cultural depictions
Symbolism
Economic importance
Uses
Hazards
Notes
References
Further reading
External links
Dipterans are endopterygotes, insects that undergo radical metamorphosis. They belong to the Mecopterida,
alongside the Mecoptera, Siphonaptera, Lepidoptera and Trichoptera.[5][6] The possession of a single pair
of wings distinguishes most true flies from other insects with "fly" in their names. However, some true flies
such as Hippoboscidae (louse flies) have become secondarily wingless.[7]
Trichoptera (caddisflies)
Lepidoptera (butterflies and moths)
The first true dipterans known are from the Middle Triassic (around 240
million years ago), and they became widespread during the Middle and Late
Triassic.[9] Modern flowering plants did not appear until the Cretaceous
(around 140 million years ago), so the original dipterans must have had a
different source of nutrition other than nectar. Based on the attraction of many
modern fly groups to shiny droplets, it has been suggested that they may have
Fossil brachyceran in fed on honeydew produced by sap-sucking bugs which were abundant at the
Baltic amber. Lower time, and dipteran mouthparts are well-adapted to softening and lapping up the
Eocene, c. 50 million crusted residues.[10] The basal clades in the Diptera include the
years ago Deuterophlebiidae and the enigmatic Nymphomyiidae.[11] Three episodes of
evolutionary radiation are thought to have occurred based on the fossil record.
Many new species of lower Diptera developed in the Triassic, about 220
million years ago. Many lower Brachycera appeared in the Jurassic, some 180 million years ago. A third
radiation took place among the Schizophora at the start of the Paleogene, 66 million years ago.[11]
The phylogenetic position of Diptera has been controversial. The monophyly of holometabolous insects has
long been accepted, with the main orders being established as Lepidoptera, Coleoptera, Hymenoptera and
Diptera, and it is the relationships between these groups which has caused difficulties. Diptera is widely
thought to be a member of Mecopterida, along with Lepidoptera (butterflies and moths), Trichoptera
(caddisflies), Siphonaptera (fleas), Mecoptera (scorpionflies) and possibly Strepsiptera (twisted-wing flies).
Diptera has been grouped with Siphonaptera and Mecoptera in the Antliophora, but this has not been
confirmed by molecular studies.[12]
Diptera were traditionally broken down into two suborders, Nematocera and
Brachycera, distinguished by the differences in antennae. The Nematocera are
identified by their elongated bodies and many-segmented, often feathery
antennae as represented by mosquitoes and crane flies. The Brachycera have
rounder bodies and much shorter antennae.[13][14] Subsequent studies have
identified the Nematocera as being non-monophyletic with modern
phylogenies placing the Brachycera within grades of groups formerly placed
in the Nematocera. The construction of a phylogenetic tree has been the Fossil nematoceran in
subject of ongoing research. The following cladogram is based on the Dominican amber.
FLYTREE project.[11][15] Sandfly, Lutzomyia
adiketis (Psychodidae),
Early Miocene, c. 20
million years ago
Ptychopteromorpha (phantom and primitive crane-flies)
Culicomorpha (mosquitoes)
Nematocera
Abbreviations used in the cladogram:
Cal=Calyptratae
Cyc=Cyclorrhapha
Ere=Eremoneura
Mus=Muscomorpha
Sch=Schizophora
Tab=Tabanomorpha
Bibionomorpha (gnats)
Diversity
Flies are often abundant and are found in almost all terrestrial
habitats in the world apart from Antarctica. They include many
familiar insects such as house flies, blow flies, mosquitoes, gnats,
black flies, midges and fruit flies. More than 150,000 have been
formally described and the actual species diversity is much greater,
with the flies from many parts of the world yet to be studied
intensively.[16][17] The suborder Nematocera include generally
small, slender insects with long antennae such as mosquitoes, gnats,
midges and crane-flies, while the Brachycera includes broader, Gauromydas heros is the largest fly
more robust flies with short antennae. Many nematoceran larvae are in the world.
aquatic.[18] There are estimated to be a total of about 19,000
species of Diptera in Europe, 22,000 in the Nearctic region, 20,000
in the Afrotropical region, 23,000 in the Oriental region and 19,000 in the Australasian region.[19] While
most species have restricted distributions, a few like the housefly (Musca domestica) are cosmopolitan.[20]
Gauromydas heros (Asiloidea), with a length of up to 7 cm (2.8 in), is generally considered to be the largest
fly in the world,[21] while the smallest is Euryplatea nanaknihali, which at 0.4 mm (0.016 in) is smaller
than a grain of salt.[22]
Brachycera are ecologically very diverse, with many being predatory at the larval stage and some being
parasitic. Animals parasitised include molluscs, woodlice, millipedes, insects, mammals,[19] and
amphibians.[23] Flies are the second largest group of pollinators after the Hymenoptera (bees, wasps and
relatives). In wet and colder environments flies are significantly more important as pollinators. Compared to
bees, they need less food as they do not need to provision their young. Many flowers that bear low nectar
and those that have evolved trap pollination depend on flies.[24] It is thought that some of the earliest
pollinators of plants may have been flies.[25]
The greatest diversity of gall forming insects are found among the flies, principally in the family
Cecidomyiidae (gall midges).[26] Many flies (most importantly in the family Agromyzidae) lay their eggs in
the mesophyll tissue of leaves with larvae feeding between the surfaces forming blisters and mines.[27]
Some families are mycophagous or fungus feeding. These include the cave dwelling Mycetophilidae
(fungus gnats) whose larvae are the only diptera with bioluminescence. The Sciaridae are also fungus
feeders. Some plants are pollinated by fungus feeding flies that visit fungus infected male flowers.[28]
The larvae of Megaselia scalaris (Phoridae) are almost omnivorous and consume such substances as paint
and shoe polish.[29] The Exorista mella (Walker) fly are considered generalists and parasitoids of a variety
of hosts.[30] The larvae of the shore flies (Ephydridae) and some Chironomidae survive in extreme
environments including glaciers (Diamesa sp., Chironomidae[31]), hot springs, geysers, saline pools,
sulphur pools, septic tanks and even crude oil (Helaeomyia petrolei[31]).[19] Adult hoverflies (Syrphidae)
are well known for their mimicry and the larvae adopt diverse lifestyles including being inquiline
scavengers inside the nests of social insects.[32] Some brachycerans are agricultural pests, some bite animals
and humans and suck their blood, and some transmit diseases.[19]
Flies have a mobile head with a pair of large compound eyes on the
sides of the head, and in most species, three small ocelli on the top.
The compound eyes may be close together or widely separated,
and in some instances are divided into a dorsal region and a ventral
region, perhaps to assist in swarming behaviour. The antennae are
well-developed but variable, being thread-like, feathery or comb-
like in the different families. The mouthparts are adapted for
piercing and sucking, as in the black flies, mosquitoes and robber
flies, and for lapping and sucking as in many other groups.[33]
Female horse-flies use knife-like mandibles and maxillae to make a
Head of a horse-fly showing large
cross-shaped incision in the host's skin and then lap up the blood
compound eyes and stout piercing
mouthparts
that flows. The gut includes large diverticulae, allowing the insect
to store small quantities of liquid after a meal.[34]
For visual course control, flies' optic flow field is analyzed by a set
of motion-sensitive neurons.[35] A subset of these neurons is
thought to be involved in using the optic flow to estimate the
parameters of self-motion, such as yaw, roll, and sideward
translation.[36] Other neurons are thought to be involved in
analyzing the content of the visual scene itself, such as separating
figures from the ground using motion parallax.[37][38] The H1
neuron is responsible for detecting horizontal motion across the
entire visual field of the fly, allowing the fly to generate and guide
stabilizing motor corrections midflight with respect to yaw.[39] The
ocelli are concerned in the detection of changes in light intensity,
enabling the fly to react swiftly to the approach of an object.[40]
A head of a fly, showing the two
Like other insects, flies have chemoreceptors that detect smell and
compound eyes and three simple
taste, and mechanoreceptors that respond to touch. The third
eyes clearly.
segments of the antennae and the maxillary palps bear the main
olfactory receptors, while the gustatory receptors are in the labium,
pharynx, feet, wing margins and female genitalia,[41] enabling flies to taste their food by walking on it. The
taste receptors in females at the tip of the abdomen receive information on the suitability of a site for
ovipositing.[40] Flies that feed on blood have special sensory structures that can detect infrared emissions,
and use them to home in on their hosts, and many blood-sucking flies can detect the raised concentration of
carbon dioxide that occurs near large animals.[42] Some tachinid flies (Ormiinae) which are parasitoids of
bush crickets, have sound receptors to help them locate their singing hosts.[43]
Diptera have one pair of fore wings on the mesothorax and a pair of halteres, or reduced hind wings, on the
metathorax. A further adaptation for flight is the reduction in number of the neural ganglia, and
concentration of nerve tissue in the thorax, a feature that is most extreme in the highly derived
[34] Some species of flies are exceptional
Muscomorpha infraorder.
in that they are secondarily flightless.
The only (crane
Tipuloidea other order
flies) of
insects bearing a single pair of true, functional wings, in addition to
any form of halteres, are the Strepsiptera. In contrast to the flies, the
Strepsiptera bear their halteres on the mesothorax and their flight
wings on the metathorax.[44] Each of the fly's six legs has a typical
insect structure of coxa, trochanter, femur, tibia and tarsus, with the
tarsus in most instances being subdivided into five tarsomeres.[33]
At the tip of the limb is a pair of claws, and between these are
cushion-like structures known as pulvilli which provide A cranefly, showing the hind wings
adhesion.[45] reduced to drumstick-shaped
halteres
The abdomen shows considerable variability among members of
the order. It consists of eleven segments in primitive groups and ten
segments in more derived groups, the tenth and eleventh segments having fused.[46] The last two or three
segments are adapted for reproduction. Each segment is made up of a dorsal and a ventral sclerite,
connected by an elastic membrane. In some females, the sclerites are rolled into a flexible, telescopic
ovipositor.[33]
Flight
Flies have rapid reflexes that aid their escape from predators but their sustained flight speeds are low.
Dolichopodid flies in the genus Condylostylus respond in less than 5 milliseconds to camera flashes by
taking flight.[50] In the past, the deer bot fly, Cephenemyia, was claimed to be one of the fastest insects on
the basis of an estimate made visually by Charles Townsend in 1927.[51] This claim, of speeds of 600 to
800 miles per hour, was regularly repeated until it was shown to be physically impossible as well as
incorrect by Irving Langmuir. Langmuir suggested an estimated speed of 25 miles per hour.[52][53][54]
Although most flies live and fly close to the ground, a few are known to fly at heights and a few like
Oscinella (Chloropidae) are known to be dispersed by winds at altitudes of up to 2000 ft and over long
distances.[55] Some hover flies like Metasyrphus corollae have been known to undertake long flights in
response to aphid population spurts.[56]
Males of fly species such as Cuterebra, many hover flies,[57] bee flies (Bombyliidae)[58] and fruit flies
(Tephritidae)[59] maintain territories within which they engage in aerial pursuit to drive away intruding
Stratiomyomorpha (soldier flies, etc)
males and other species.[60] While these territories
may be held by individual males, some species, such as
A. freeborni, [61] form leks with many males aggregating in displays.[59] Some flies maintain an airspace
and still others form dense swarms that maintain a stationary location with respect to landmarks. Many flies
mate in flight while swarming.[62]
Xylophagomorpha (stink flies, etc)
Tab
Life cycle and development
Diptera go through a complete metamorphosis withTabanomorpha
four distinct life (horse flies, snipe flies, etc)
stages – egg, larva, pupa and adult.
Larva
In many flies, the larval stage is long and adults may have a short
life. Most dipteran larvae develop in protected environments; many
are aquatic and others are found in moist places such as carrion,
fruit, vegetable matter, fungi and, in the case of parasitic species, Mating anthomyiid flies
inside their hosts. They tend to have thin cuticles and become
desiccated if exposed to the air. Apart from the Brachycera, most
dipteran larvae have sclerotinised head capsules, which may be reduced to remnant mouth hooks; the
Brachycera, however, have soft, gelatinized head capsules from which the sclerites are reduced or missing.
Many of these larvae retract their heads into their thorax.[33][63]
Adult
Syrphoidea (hoverflies)
The adult stage is usually short, its function is only to mate and lay eggs. The genitalia of male flies are
Mus found in other insects.[70] In some flies, this is a temporary
rotated to a varying degree from the position
rotation during mating, but in others, it is a permanent torsion of the organs that occurs during the pupal
stage. This torsion may lead to the anus being below the genitals, or, in the case of 360° torsion,(louse
Hippoboscoidea to the
sperm duct being wrapped around the gut and the external organs being in their usual position. When flies
mate, the male initially flies on top of the female,Ere flies,
facing in the same direction, butetc)
then turns around to face
in the opposite direction. This forces the male to lie on his back for his genitalia to remain engaged with
Cyc
those of the female, or the torsion of the male genitals allows the male to mate while remaining upright.
This leads to flies having more reproduction abilities than most insects, and much quicker. (house
Muscoidea Flies occur in
flies,
[34]
large populations due to their ability to mate effectively and quickly during
Cal the mating season.
dung flies, etc)
Ecology Sch
Oestroidea (blow flies,
As ubiquitous insects, dipterans play an important role at various
trophic levels both as consumers and flesh
as prey. In etc)
flies, some groups the
larvae complete their development without feeding, and in others
the adults do not feed. The larvae can be herbivores, scavengers,
decomposers, predators or parasites, with the consumption of
decaying organic matter being oneAcalyptratae
of the most(marsh flies,
prevalent etc)
feeding
behaviours. The fruit or detritus
is consumed along with the
associated micro-organisms, a sieve-like filter in the pharynx being
A calliphorid "bubbling" used to concentrate the particles, while flesh-eating larvae have
mouth-hooks to help shred their food. The larvae of some groups
feed on or in the living tissues of plants and fungi, and some of
these are serious pests of agricultural crops. Some aquatic larvae consume the films of algae that form
underwater on rocks and plants. Many of the parasitoid larvae grow inside and eventually kill other
arthropods, while parasitic larvae may attack vertebrate hosts.[33]
Whereas many dipteran larvae are aquatic or live in enclosed terrestrial locations, the majority of adults live
above ground and are capable of flight. Predominantly they feed on nectar or plant or animal exudates,
such as honeydew, for which their lapping mouthparts are adapted. Some flies have functional mandibles
that may be used for biting. The flies that feed on vertebrate blood have sharp stylets that pierce the skin,
with some species having anticoagulant saliva that is regurgitated before absorbing the blood that flows; in
this process, certain diseases can be transmitted. The bot flies (Oestridae) have evolved to parasitize
mammals. Many species complete their life cycle inside the bodies of their hosts.[71] The larvae of a few fly
groups (Agromyzidae, Anthomyiidae, Cecidomyiidae) are capable of inducing plant galls. Some dipteran
larvae are leaf-miners. The larvae of many brachyceran families are predaceous. In many dipteran groups,
swarming is a feature of adult life, with clouds of insects gathering in certain locations; these insects are
mostly males, and the swarm may serve the purpose of making their location more visible to females.[33]
Most adult diptera have their mouthparts modified to sponge up fluid. The adults of many species of flies
(e.g. Anthomyia sp., Steganopsis melanogaster) that feed on liquid food will regurgitate fluid in a
behaviour termed as "bubbling" which has been thought to help the insects evaporate water and
concentrate food[72] or possibly to cool by evaporation.[73] Some adult diptera are known for
kleptoparasitism such as members of the Sarcophagidae. The miltogramminae are known as "satellite flies"
for their habit of following wasps and stealing their stung prey or laying their eggs into them. Phorids,
milichids and the genus Bengalia are known to steal food carried by ants.[74] Adults of Ephydra hians
forage underwater, and have special hydrophobic hairs that trap a bubble of air that lets them breathe
underwater.[75]
Anti-predator adaptations
In contrast, Drosophila subobscura, a species of fly in the genus Drosophila, lacks a category of hemocytes
that are present in other studied species of Drosophila, leading to an inability to defend against parasitic
attacks, a form of innate immunodeficiency.[83]
Symbolism
Flies play a variety of symbolic roles in different cultures. These include
both positive and negative roles in religion. In the traditional Navajo
religion, Big Fly is an important spirit being.[84][85][86] In Christian
demonology, Beelzebub is a demonic fly, the "Lord of the Flies", and a god
of the Philistines.[87][88][89]
Musca depicta ("painted fly" in Latin) is a depiction of a fly as an inconspicuous element of various
paintings. This feature was widespread in 15th and 16th centuries paintings and its presence may be
explained by various reasons.[95]
Flies appear in popular culture in concepts such as fly-on-the-wall documentary-making in film and
television production. The metaphoric name suggests that events are seen candidly, as a fly might see
them.[96] Flies have inspired the design of miniature flying robots.[97] Steven Spielberg's 1993 film
Jurassic Park relied on the idea that DNA could be preserved in the stomach contents of a blood-sucking
fly fossilised in amber, though the mechanism has been discounted by scientists.[98]
Economic importance
Many dipterans serve roles that are useful to humans. Houseflies, blowflies and fungus gnats
(Mycetophilidae) are scavengers and aid in decomposition. Robber flies (Asilidae), tachinids (Tachinidae)
and dagger flies and balloon flies (Empididae) are predators and parasitoids of other insects, helping to
control a variety of pests. Many dipterans such as bee flies (Bombyliidae) and hoverflies (Syrphidae) are
pollinators of crop plants.[33]
Uses
Maggots found on corpses are useful to forensic entomologists. Maggot species can be identified by their
anatomical features and by matching their DNA. Maggots of different species of flies visit corpses and
carcases at fairly well-defined times after the death of the victim, and so do their predators, such as beetles
in the family Histeridae. Thus, the presence or absence of particular species provides evidence for the time
since death, and sometimes other details such as the place of death, when species are confined to particular
habitats such as woodland.[102]
The Sardinian cheese casu marzu is exposed to flies known as cheese skippers such as Piophila casei,
members of the family Piophilidae.[109] The digestive activities of the fly larvae soften the cheese and
modify the aroma as part of the process of maturation. At one time European Union authorities banned sale
of the cheese and it was becoming hard to find,[110] but the ban has been lifted on the grounds that the
cheese is a traditional local product made by traditional methods.[111]
Hazards
Flies are a health hazard and are attracted to toilets because of their smell. The New Scientist magazine
suggested a trap for these flies. A pipe acting as a chimney was fitted to the toilet which let in some light to
attract these flies up to the end of this pipe where a gauze prevented escape to the air outside so that they
were trapped and died. Toilets are generally dark inside particularly if the door is closed.
Notes
a. Some authors draw a distinction in writing the common names of insects. True flies are in
their view best written as two words, such as crane fly, robber fly, bee fly, moth fly, and fruit
fly. In contrast, common names of non-dipteran insects that have "fly" in their names are
written as one word, e.g. butterfly, stonefly, dragonfly, scorpionfly, sawfly, caddisfly,
whitefly.[2] In practice, however, this is a comparatively new convention; especially in older
books, names like "saw fly" and "caddis fly", or hyphenated forms such as house-fly and
dragon-fly are widely used.[3] In any case, non-entomologists cannot, in general, be
expected to tell dipterans, "true flies", from other insects, so it would be unrealistic to expect
rigour in the use of common names. Also, exceptions to this rule occur, such as the hoverfly,
which is a true fly, and the Spanish fly, a type of blister beetle.
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Further reading
Blagoderov, V.A., Lukashevich, E.D. & Mostovski, M.B. 2002. Order Diptera (http://palaeoent
omolog.ru/New/diptera.html). In: Rasnitsyn, A.P. and Quicke, D.L.J. The History of Insects,
Kluwer pp.–227–240.
Colless, D.H. & McAlpine, D.K. 1991 Diptera (flies), pp. 717–786. In: The Division of
Entomology. Commonwealth Scientific and Industrial Research Organisation, Canberra
(spons.), The insects of Australia. Melbourne University Press.
Hennig, Willi Diptera (Zweifluger). Handb. Zool. Berl. 4 (2) (31):1–337. General introduction
with key to World Families. In German.
Oldroyd, Harold The Natural History of Flies. W. W. Norton. 1965.
Séguy, Eugène Diptera: recueil d'etudes biologiques et systematiques sur les Dipteres du
Globe (Collection of biological and systematic studies on Diptera of the World). 11 vols. Part
of Encyclopedie Entomologique, Serie B II: Diptera. 1924–1953.
Séguy, Eugène La Biologie des Dipteres 1950.
Thompson, F. Christian. "Sources for the Biosystematic Database of World Diptera (Flies)"
(https://web.archive.org/web/20150918204612/http://www.sel.barc.usda.gov/diptera/names/
BDWDsour.pdf) (PDF). United States Department of Agriculture, Systematic Entomology
Laboratory. Archived from the original on 18 September 2015.
External links
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