Fly
Encyclopedia
True flies are insect
s of the order
Diptera (from the Greek di = two, and ptera = wings). They possess a pair of wing
s on the mesothorax
and a pair of halteres, derived from the hind wings, on the metathorax
. Apart from secondarily flightless insects (including some flies), the only other order of insects with any form of halteres are the Strepsiptera
, and theirs are on the mesothorax, with the flight wings on the metathorax.
, dragonflies
, damselflies
, stoneflies, whiteflies
, fireflies
, alderflies
, dobsonflies
, snakeflies
, sawflies
, caddisflies, butterflies
or scorpionflies. Some true flies have become secondarily wingless, especially in the superfamily Hippoboscoidea
, or among those that are inquiline
s in social insect colonies.
Some authors draw a distinction in writing the common names of insects: true flies are written as two words, e.g., crane fly, robber fly, bee fly, moth fly, fruit fly. In contrast, common names of nondipteran 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, and, especially in older books, one commonly might see the likes of "saw fly" and "caddis fly". In any case, non-entomologists cannot in general be expected to tell dipterans, "true flies", from other insects.
The Diptera comprise a large order, containing an estimated 240,000 species
of mosquito
es, gnat
s, midges
and others, although under half of these (about 120,000 species) have been described. It is one of the major insect orders both in terms of ecological and human (medical and economic) importance. The Diptera, in particular the mosquitoes (Culicidae), are of great importance as disease transmitters, acting as vectors for malaria
, dengue, West Nile virus
, yellow fever
, encephalitis
and other infectious diseases.
Flies are adapted for aerial movement, and typically have short and streamlined bodies. The first tagma
of the fly, the head, consists of ocelli, antennae, compound eyes, 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. The third segment of the thorax bears the halteres, which help to balance the insect during flight. 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.
Flies have a mobile head with eyes and in most cases have large compound eyes on the sides of the head, with three small ocelli on the top. The antennae
take a variety of forms, but are often short, which reduces drag while flying.
Because no species of fly has teeth or any other organ or limb that allows them to eat solid foods, flies consume only liquid food, and their mouthparts and digestive tracts show various modifications for this diet. Female Tabanidae use knife-like mandibles and maxillae to make a cross-shaped incision in the hosts' skin and then lap up the blood. The gut includes large diverticulae, allowing the insect to store small quantities of liquid after a meal.
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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 located below the genitals, or, in the case of 360° torsion, to the sperm duct being wrapped around the gut, despite 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 round 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 at a much quicker rate. Flies occur in great populations due to their ability to mate effectively and in a short period of time during the mating season.
The female lays her eggs as close to the food source as possible, and development is rapid, allowing the larvae to consume as much food as possible in a short period of time before transforming into adults. The eggs hatch immediately after being laid, or the flies are ovoviviparous, with the larvae hatching inside the mother.
Larval flies have no true legs. Some Dipteran larvae, such as species of Simuliidae, Tabanidae, and Vermileonidae
, have proleg
s adapted to such functions as holding onto a substrate in flowing water, holding onto host tissues, or holding prey. Roughly speaking, there is some anatomical distinction between the larvae of the Nematocera
and the Brachycera
(see Classification section, below); especially in the Brachycera, there is little demarcation between the thorax
and abdomen
, though the demarcation may be very visible in many Nematocera, such as mosquitoes (see image, both here and in the mosquito
es article); in the Brachycera, the head of the larva is not clearly distinguishable from the rest of the body, and there are few, if any, sclerite
s. Informally, such Brachyceran larvae are called maggots, but the term is nontechnical 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 visible eyes and antennae, though usually small and of limited function.
The pupa
e take various forms, and in some cases develop inside a silk cocoon. After emerging from the pupa, the adult fly rarely lives more than a few days, and serves mainly to reproduce and to disperse in search of new food sources.
are recognized by their elongated bodies and feathery antennae as represented by mosquitoes and crane flies. The Brachycera
have a more roundly proportioned body and much shorter antennae. In 1964, B.B. Rohdendorf proposed a classification in which the Nematocera is split into two suborders, the Archidiptera
and the Eudiptera
.
Most of the Muscomorpha are further subdivided into the Acalyptratae
and Calyptratae
based on whether or not they have a calypter (a wing flap that extends over the halteres).
Beyond that, considerable revision in the taxonomy of the flies has taken place since the introduction of modern cladistic techniques, and much remains uncertain. The secondary ranks between the suborders and the families are out of practical or historical considerations than out of strict respect for phylogenetic classifications (modern cladists spurn the use of Linnaean rank names). All classifications in use now, including this article, contain some paraphyletic groupings; this is emphasized where the numerous alternative systems are most greatly at odds. See list of families of Diptera.
Dipterans belong to the taxon Mecopterida, that also contains Mecoptera
, Siphonaptera, Lepidoptera
(butterflies and moth
s) and Trichoptera
. Inside it, they are classified closely together with Mecoptera
and Siphonaptera in the superorder Antliophora.
or a strictly related group. The first true dipterans are known from the Middle Triassic
, becoming widespread during the Middle and Late Triassic .
The basal clades in the Diptera are the Deuterophlebiidae and Nymphomyiidae
. The Bibionomorpha
are a sister clade to Brachycera
. The branching order of the remaining clades of the lower Diptera - infraorders Culicomorpha
, Psychodomorpha
and Tipulomorpha
- has yet to be resolved.
Within the Brachycera, several progressively nested groups exist: Eremoneura (three larval instars), Cyclorrhapha
(pupation occurs within a puparium), Schizophora
(flies that escape from their puparium using the ptilinal sac, an evertable frontal pouch) and Calyptratae
(larger flies with wings that have the calypter
, an enlarged basal lobe).
The Schizophora include most of the family-level diversity in Diptera (∼85 families) and more than 50,000 species. The Calyptratae form a monophyletic superfamily. Other monophyletic superfamilies include the Ephydroidea
, Lauxanioidea
, Nerioidea
, Sciomyzoidea
and Tephritoidea
. The relationships between the remaining families have yet to be clarified.
Sister groups to the Drosophilidae
consist of two families, Braulidae
and Cryptochetidae.
There were three episodes of rapid evolution in the lower Diptera (~220 million years ago), lower Brachycera (180 million years ago) and Schizophora (65 million years ago).
Maggots found on corpses are useful to forensic scientists; specifically forensic entomology
. By their stage of development, these maggots indicate the time elapsed since death, as well as the place of death.
Maggot species can be identified
using their DNA
. The housefly
maggot measures 10–20 mm (⅜–¾ in) in size. At the height of the summer season, a generation of flies (egg to adult) may be produced in 12 to 14 days. Other insect families, such as Histeridae, feed on maggots. Thus the lack of maggots would increase the estimated time of death.
Maggots are bred commercially, as a popular bait in angling
, and as a food for carnivorous pets such as reptiles or birds.
Maggots are used in medicine
to clean out necrotic
wound
s and in food production, particularly of cheeses such as casu marzu
designed to rot as part of their aging process.
Flies are reared in large numbers in Japan
to serve as pollinator
s of sunflower
s in greenhouses, especially the maggots.
and literature
. In the Biblical fourth plague of Egypt
, flies represent death and decay. Myiagros
was a god in Greek mythology
who chased away flies during the sacrifices to Zeus
and Athena
, and Zeus
sent a fly to bite Pegasus
, causing Bellerophon
to fall back to Earth when he attempted to ride the winged steed to Mount Olympus. In the traditional Navajo
religion, Big Fly is an important spirit being.
In the 15th-century trompe l'oeil
painting Portrait of a Carthusian (1446) by Petrus Christus
, a fly sits on a fake frame.
Emily Dickinson's
1955
poem "I Heard a Fly Buzz When I Died" also makes reference to flies in the context of death. In fact, flies such as the genus Hydrotaea
are used in forensic cases to determine time of death for corpses. In William Golding's
1954
novel
Lord of the Flies
, the fly is a symbol of the children involved.
In the 1958 science fiction film The Fly
(and its 1986 remake
), a scientist accidentally exchanges parts of his body with those of a fly.
Musical works that mention flies: Yoko Ono
's 1971
album Fly
, U2
's 1991
song "The Fly
", Wire
's 1978
song "I Am The Fly", Dave Matthews
's 2007
song "The Fly" and Béla Bartók's
1920s piano work "From the Diary of a Fly".
Damien Hirst
's provocative 1990
artwork, titled A Thousand Years, featured a severed cow's head contained in a box with thousands of flies and a bug zapper
, creating an entire life cycle within a glass box. In 2001 Dr. Garnet Hertz implanted a complete web server
into a dead fly.
The ability of flies to cling to surfaces has also inspired the title of "Human Fly" for real and fictional stunt performers whose feats involve climbing buildings.
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...
s of the order
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...
Diptera (from the Greek di = two, and ptera = wings). They possess a pair of wing
Insect wing
Insects are the only group of invertebrates known to have evolved flight. Insects possess some remarkable flight characteristics and abilities, still far superior to attempts by humans to replicate their capabilities. Even our understanding of the aerodynamics of flexible, flapping wings and how...
s on the mesothorax
Mesothorax
The mesothorax is the middle of the three segments in the thorax of an insect, and bears the second pair of legs. Its principal sclerites are the mesonotum , the mesosternum , and the mesopleuron on each side...
and a pair of halteres, derived from the hind wings, on the metathorax
Metathorax
The metathorax is the posterior of the three segments in the thorax of an insect, and bears the third pair of legs. Its principal sclerites are the metanotum , the metasternum , and the metapleuron on each side...
. Apart from secondarily flightless insects (including some flies), the only other order of insects with any form of halteres are the Strepsiptera
Strepsiptera
The Strepsiptera are an order of insects with ten families making up about 600 species...
, and theirs are on the mesothorax, with the flight wings on the metathorax.
The order Diptera
The presence of a single pair of patent, metathoracic flight wings distinguishes true flies from other insects with "fly" in their names, such as mayfliesMayfly
Mayflies are insects which belong to the Order Ephemeroptera . They have been placed into an ancient group of insects termed the Palaeoptera, which also contains dragonflies and damselflies...
, dragonflies
Dragonfly
A dragonfly is a winged insect belonging to the order Odonata, the suborder Epiprocta or, in the strict sense, the infraorder Anisoptera . It is characterized by large multifaceted eyes, two pairs of strong transparent wings, and an elongated body...
, damselflies
Damselfly
Damselflies are insects in the order Odonata. Damselflies are similar to dragonflies, but the adults can be distinguished by the fact that the wings of most damselflies are held along, and parallel to, the body when at rest...
, stoneflies, whiteflies
Whitefly
The whiteflies, comprising only the family Aleyrodidae, are small hemipterans. More than 1550 species have been described. Whiteflies typically feed on the underside of plant leaves.-Agricultural threat:...
, fireflies
Firefly
Lampyridae is a family of insects in the beetle order Coleoptera. They are winged beetles, and commonly called fireflies or lightning bugs for their conspicuous crepuscular use of bioluminescence to attract mates or prey. Fireflies produce a "cold light", with no infrared or ultraviolet frequencies...
, alderflies
Alderfly
Alderflies are megalopteran insects of the family Sialidae. They are closely related to the dobsonflies and fishflies as well as to the prehistoric Euchauliodidae. All living alderflies - about 66 species altogether - are part of the subfamily Sialinae, which contains between one and seven extant...
, dobsonflies
Dobsonfly
A Dobsonfly is any insect of the subfamily Corydalinae, part of the megalopteran family Corydalidae. There are over 220 species of dobsonflies. Dobsonflies are found throughout the Americas and Asia, as well as South Africa...
, snakeflies
Snakefly
Snakeflies are a group of insects in the order Raphidioptera, consisting of about 210 extant species. Together with the Megaloptera they were formerly placed within the Neuroptera, but now these two are generally regarded as separate orders....
, sawflies
Sawfly
Sawfly is the common name for insects belonging to suborder Symphyta of the order Hymenoptera. Sawflies are distinguishable from most other Hymenoptera by the broad connection between the abdomen and the thorax, and by their caterpillar-like larvae...
, caddisflies, butterflies
Butterfly
A butterfly is a mainly day-flying insect of the order Lepidoptera, which includes the butterflies and moths. Like other holometabolous insects, the butterfly's life cycle consists of four parts: egg, larva, pupa and adult. Most species are diurnal. Butterflies have large, often brightly coloured...
or scorpionflies. Some true flies have become secondarily wingless, especially in the superfamily Hippoboscoidea
Hippoboscoidea
Hippoboscoidea is a superfamily of Calyptratae. The flies in this superfamily are blood-feeding obligate parasites of their hosts. Five families are often placed here:* Glossinidae* Hippoboscidae...
, or among those that are inquiline
Inquiline
In zoology, an inquiline is an animal that lives commensally in the nest, burrow, or dwelling place of an animal of another species. For example, some organisms such as insects may live in the homes of gophers and feed on debris, fungi, roots, etc...
s in social insect colonies.
Some authors draw a distinction in writing the common names of insects: true flies are written as two words, e.g., crane fly, robber fly, bee fly, moth fly, fruit fly. In contrast, common names of nondipteran 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, and, especially in older books, one commonly might see the likes of "saw fly" and "caddis fly". In any case, non-entomologists cannot in general be expected to tell dipterans, "true flies", from other insects.
The Diptera comprise a large order, containing an estimated 240,000 species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...
of mosquito
Mosquito
Mosquitoes are members of a family of nematocerid flies: the Culicidae . The word Mosquito is from the Spanish and Portuguese for little fly...
es, gnat
Gnat
A gnat is any of many species of tiny flying insects in the Dipterid suborder Nematocera, especially those in the families Mycetophilidae, Anisopodidae and Sciaridae.In British English the term applies particularly to Nematocerans of the family Culicidae...
s, midges
Midge (insect)
Midges comprise many kinds of very small two-winged flies found world-wide. The term does not encapsulate a well-defined taxonomic group, but includes animals in several families of Nematoceran Diptera. While some midges are vectors for disease, many others play useful roles as prey items for...
and others, although under half of these (about 120,000 species) have been described. It is one of the major insect orders both in terms of ecological and human (medical and economic) importance. The Diptera, in particular the mosquitoes (Culicidae), are of great importance as disease transmitters, acting as vectors for malaria
Malaria
Malaria is a mosquito-borne infectious disease of humans and other animals caused by eukaryotic protists of the genus Plasmodium. The disease results from the multiplication of Plasmodium parasites within red blood cells, causing symptoms that typically include fever and headache, in severe cases...
, dengue, West Nile virus
West Nile virus
West Nile virus is a virus of the family Flaviviridae. Part of the Japanese encephalitis antigenic complex of viruses, it is found in both tropical and temperate regions. It mainly infects birds, but is known to infect humans, horses, dogs, cats, bats, chipmunks, skunks, squirrels, domestic...
, yellow fever
Yellow fever
Yellow fever is an acute viral hemorrhagic disease. The virus is a 40 to 50 nm enveloped RNA virus with positive sense of the Flaviviridae family....
, encephalitis
Encephalitis
Encephalitis is an acute inflammation of the brain. Encephalitis with meningitis is known as meningoencephalitis. Symptoms include headache, fever, confusion, drowsiness, and fatigue...
and other infectious diseases.
Anatomy and biology
Tagma
A tagma may refer to:* Tagma , a grouping of segments, usually in arthropod anatomy* tagma , a subdivision of the Byzantine army...
of the fly, the head, consists of ocelli, antennae, compound eyes, and the mouthparts
Insect mouthparts
Insects exhibit a range of mouthparts, adapted to particular modes of feeding. The earliest insects had chewing mouthparts...
(the labrum, labium, mandible and maxilla make up the mouthparts). The second tagma, the thorax
Thorax (insect anatomy)
The thorax is the mid section of the insect body. It holds the head, legs, wings and abdomen. It is also called mesosoma in other arthropods....
, 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. The third segment of the thorax bears the halteres, which help to balance the insect during flight. A further adaptation for flight is the reduction in number of the neural ganglia
Ganglion
In anatomy, a ganglion is a biological tissue mass, most commonly a mass of nerve cell bodies. Cells found in a ganglion are called ganglion cells, though this term is also sometimes used to refer specifically to retinal ganglion cells....
, and concentration of nerve tissue in the thorax, a feature that is most extreme in the highly derived Muscomorpha infraorder.
Flies have a mobile head with eyes and in most cases have large compound eyes on the sides of the head, with three small ocelli on the top. The antennae
Antenna (biology)
Antennae in biology have historically been paired appendages used for sensing in arthropods. More recently, the term has also been applied to cilium structures present in most cell types of eukaryotes....
take a variety of forms, but are often short, which reduces drag while flying.
Because no species of fly has teeth or any other organ or limb that allows them to eat solid foods, flies consume only liquid food, and their mouthparts and digestive tracts show various modifications for this diet. Female Tabanidae use knife-like mandibles and maxillae to make a cross-shaped incision in the hosts' skin and then lap up the blood. The gut includes large diverticulae, allowing the insect to store small quantities of liquid after a meal.
Reproduction and development
Anus
The anus is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, unwanted semi-solid matter produced during digestion, which, depending on the type of animal, may be one or more of: matter which the animal cannot digest,...
being located below the genitals, or, in the case of 360° torsion, to the sperm duct being wrapped around the gut, despite 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 round 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 at a much quicker rate. Flies occur in great populations due to their ability to mate effectively and in a short period of time during the mating season.
The female lays her eggs as close to the food source as possible, and development is rapid, allowing the larvae to consume as much food as possible in a short period of time before transforming into adults. The eggs hatch immediately after being laid, or the flies are ovoviviparous, with the larvae hatching inside the mother.
Larval flies have no true legs. Some Dipteran larvae, such as species of Simuliidae, Tabanidae, and Vermileonidae
Vermileonidae
The Brachyceran family Vermileonidae is a small family of uncertain affinities and unusual biology, containing fewer than 80 rare species in 10 genera. Historically the Vermileonids had been regarded as belonging to the family Rhagionidae, possibly in a subfamily Vermileoninae...
, have proleg
Proleg
A Proleg is the small fleshy, stub structure found on the ventral surface of the abdomen of most larval forms of insects of the order Lepidoptera, though they can also be found on other larval insects such as sawflies and a few types of flies....
s adapted to such functions as holding onto a substrate in flowing water, holding onto host tissues, or holding prey. Roughly speaking, there is some anatomical distinction between the larvae of the Nematocera
Nematocera
Nematocera , is a suborder of elongated flies with thin, segmented antennae and mostly aquatic larvae, consisting of the mosquitoes, crane flies, gnats, and midges....
and the Brachycera
Brachycera
Brachycera is a suborder of Diptera. It is a major suborder consisting of around 120 families. The most distinguishing characteristic of the suborder is reduced antenna segmentation...
(see Classification section, below); especially in the Brachycera, there is little demarcation between the thorax
Thorax (insect anatomy)
The thorax is the mid section of the insect body. It holds the head, legs, wings and abdomen. It is also called mesosoma in other arthropods....
and abdomen
Abdomen
In vertebrates such as mammals the abdomen constitutes the part of the body between the thorax and pelvis. The region enclosed by the abdomen is termed the abdominal cavity...
, though the demarcation may be very visible in many Nematocera, such as mosquitoes (see image, both here and in the mosquito
Mosquito
Mosquitoes are members of a family of nematocerid flies: the Culicidae . The word Mosquito is from the Spanish and Portuguese for little fly...
es article); in the Brachycera, the head of the larva is not clearly distinguishable from the rest of the body, and there are few, if any, sclerite
Sclerite
A sclerite is a hardened body part. The term is used in various branches of biology for various structures including hardened portions of sponges, but it is most commonly used for the hardened portions of arthropod exoskeletons....
s. Informally, such Brachyceran larvae are called maggots, but the term is nontechnical 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
Cercus
Cerci are paired appendages on the rear-most segments of many arthropods, including insects and arachnids but not crustaceans. Cerci often serve as sensory organs, but they may also be used as weapons or copulation aids, or they may simply be vestigial structures.Typical cerci may appear to be...
. This lack of features is an adaptation to food such as carrion, decaying detritus, or host tissues surrounding endoparasites
Parasitism
Parasitism is a type of symbiotic relationship between organisms of different species where one organism, the parasite, benefits at the expense of the other, the host. Traditionally parasite referred to organisms with lifestages that needed more than one host . These are now called macroparasites...
. Nematoceran larvae generally have visible eyes and antennae, though usually small and of limited function.
The pupa
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...
e take various forms, and in some cases develop inside a silk cocoon. After emerging from the pupa, the adult fly rarely lives more than a few days, and serves mainly to reproduce and to disperse in search of new food sources.
Classification
The NematoceraNematocera
Nematocera , is a suborder of elongated flies with thin, segmented antennae and mostly aquatic larvae, consisting of the mosquitoes, crane flies, gnats, and midges....
are recognized by their elongated bodies and feathery antennae as represented by mosquitoes and crane flies. The Brachycera
Brachycera
Brachycera is a suborder of Diptera. It is a major suborder consisting of around 120 families. The most distinguishing characteristic of the suborder is reduced antenna segmentation...
have a more roundly proportioned body and much shorter antennae. In 1964, B.B. Rohdendorf proposed a classification in which the Nematocera is split into two suborders, the Archidiptera
Archidiptera
Archidiptera is a suborder of Diptera under an alternative classification based largely on fossil taxa; it has not gained wide acceptance among non-paleontological dipterists....
and the Eudiptera
Eudiptera
Eudiptera is a suborder of Diptera under an alternative subordinal classification based largely on fossil taxa; it has not gained wide acceptance among non-paleontological dipterists and is presented here for comparative purposes only....
.
- Suborder NematoceraNematoceraNematocera , is a suborder of elongated flies with thin, segmented antennae and mostly aquatic larvae, consisting of the mosquitoes, crane flies, gnats, and midges....
(77 families, 35 of them extinct) – long antennae, pronotum distinct from mesonotum, in Nematocera, larvae are either eucephalic or hemicephalic and often aquatic. - Suborder BrachyceraBrachyceraBrachycera is a suborder of Diptera. It is a major suborder consisting of around 120 families. The most distinguishing characteristic of the suborder is reduced antenna segmentation...
(141 families, 8 of them extinct) – short antennae, the pupaPupaA pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...
is inside a puparium formed from the last larvaLarvaA larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...
l skin, they are generally robust flies with larvae having reduced mouthparts.- Infraorders TabanomorphaTabanomorphaThe Brachyceran infraorder Tabanomorpha is a small group that consists primarily of two large families, the Tabanidae and Rhagionidae , and an assortment of very small affiliated families, most of which have been included within the Rhagionidae...
and AsilomorphaAsilomorphaThe Brachyceran infraorder Asilomorpha is a large and diverse group of flies, containing the bulk of the non-muscoid Brachycera.The larvae of asilomorphs are extremely diverse in habits, as well.-Classification:...
– these comprise the majority of what was the OrthorrhaphaOrthorrhaphaOrthorrhapha is a circumscriptional name which historically was used for an infraorder of Brachycera, one of the two suborders into which the order Diptera, the flies, are divided. As the group was paraphyletic, it has not been used in classifications in the last decade, and is effectively...
under older classification schemes. The antennae are short, but differ in structure from those of the Muscomorpha. - Infraorder MuscomorphaMuscomorphaThe Brachyceran infraorder Muscomorpha is a large and diverse group of flies, containing the bulk of the Brachycera, and, in fact, most of the known flies. It includes a number of the most familiar flies, such as the housefly, the fruit fly and the blowfly. The antennae are short, usually...
– (largely the CyclorrhaphaCyclorrhaphaCyclorrhapha is an unranked taxon within the infraorder Muscomorpha. They are called "Cyclorrhapha" with reference to the circular aperture through which the adult escapes the puparium...
of older schemes). Muscomorpha have three-segmented, aristateArista (biology)In insect anatomy the arista is a simple or variously modified apical or subapical bristle, arising from the third antennal segment. It is the evolutionary remains of antennal segments, and may sometimes show signs of segmentation...
(with a bristle) antennaeAntenna (biology)Antennae in biology have historically been paired appendages used for sensing in arthropods. More recently, the term has also been applied to cilium structures present in most cell types of eukaryotes....
and larvae with three acephalic instarInstarAn instar is a developmental stage of arthropods, such as insects, between each molt , until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or assume a new form. Differences between instars can often be seen in altered body proportions, colors, patterns, or...
s (maggots).
- Infraorders Tabanomorpha
Most of the Muscomorpha are further subdivided into the Acalyptratae
Acalyptratae
Acalyptratae is a subsection of Schizophora, commonly referred to as the acalyptrate muscoids . It is a very large assemblage, exhibiting very diverse habits, with one notable and perhaps surprising exception; there are no known acalyptrates that are obligate blood-feeders , though this is a life...
and Calyptratae
Calyptratae
Calyptratae is a subsection of Schizophora in the insect order Diptera, commonly referred to as the calyptrate muscoids...
based on whether or not they have a calypter (a wing flap that extends over the halteres).
Beyond that, considerable revision in the taxonomy of the flies has taken place since the introduction of modern cladistic techniques, and much remains uncertain. The secondary ranks between the suborders and the families are out of practical or historical considerations than out of strict respect for phylogenetic classifications (modern cladists spurn the use of Linnaean rank names). All classifications in use now, including this article, contain some paraphyletic groupings; this is emphasized where the numerous alternative systems are most greatly at odds. See list of families of Diptera.
Dipterans belong to the taxon Mecopterida, that also contains Mecoptera
Mecoptera
Mecoptera are an order of insects with about 550 species in nine families worldwide. Mecoptera are sometimes called scorpionflies after their largest family, Panorpidae, in which the males have enlarged genitals that look similar to the stinger of a scorpion...
, Siphonaptera, Lepidoptera
Lepidoptera
Lepidoptera is a large order of insects that includes moths and butterflies . It is one of the most widespread and widely recognizable insect orders in the world, encompassing moths and the three superfamilies of butterflies, skipper butterflies, and moth-butterflies...
(butterflies and moth
Moth
A moth is an insect closely related to the butterfly, both being of the order Lepidoptera. Moths form the majority of this order; there are thought to be 150,000 to 250,000 different species of moth , with thousands of species yet to be described...
s) and Trichoptera
Trichoptera
The caddisflies are an order, Trichoptera, of insects with approximately 12,000 described species. Also called sedge-flies or rail-flies, they are small moth-like insects having two pairs of hairy membranous wings...
. Inside it, they are classified closely together with Mecoptera
Mecoptera
Mecoptera are an order of insects with about 550 species in nine families worldwide. Mecoptera are sometimes called scorpionflies after their largest family, Panorpidae, in which the males have enlarged genitals that look similar to the stinger of a scorpion...
and Siphonaptera in the superorder Antliophora.
Evolution
Diptera derive from MecopteraMecoptera
Mecoptera are an order of insects with about 550 species in nine families worldwide. Mecoptera are sometimes called scorpionflies after their largest family, Panorpidae, in which the males have enlarged genitals that look similar to the stinger of a scorpion...
or a strictly related group. The first true dipterans are known from the Middle Triassic
Triassic
The Triassic is a geologic period and system that extends from about 250 to 200 Mya . As the first period of the Mesozoic Era, the Triassic follows the Permian and is followed by the Jurassic. Both the start and end of the Triassic are marked by major extinction events...
, becoming widespread during the Middle and Late Triassic .
The basal clades in the Diptera are the Deuterophlebiidae and Nymphomyiidae
Nymphomyiidae
The Nymphomyiidae are a family of slender, delicate flies . Larvae are found among aquatic mosses in small, rapid streams in northern regions of the world. Many fossil species and a few extant species are known...
. The Bibionomorpha
Bibionomorpha
Bibionomorpha is an infraorder of Nematocera. One of its constituent families, the Anisopodidae, is the presumed sister taxon to the entire suborder Brachycera. Several of the remaining families in the infraorder are former subfamilies of the Mycetophilidae, which has been recently subdivided...
are a sister clade to Brachycera
Brachycera
Brachycera is a suborder of Diptera. It is a major suborder consisting of around 120 families. The most distinguishing characteristic of the suborder is reduced antenna segmentation...
. The branching order of the remaining clades of the lower Diptera - infraorders Culicomorpha
Culicomorpha
Culicomorpha is an infraorder of Nematocera. It includes mosquitoes, black flies, and several extant and extinct families of insect.-Classification:Extant families* Superfamily Culicoidea** Dixidae -meniscus midges** Corethrellidae -frog biting midges...
, Psychodomorpha
Psychodomorpha
The Nematoceran infraorder Psychodomorpha includes two common families, Psychodidae and Scatopsidae, and other very small, rare families. In some classifications , the group is paraphyletic....
and Tipulomorpha
Tipulomorpha
Tipulomorpha is an infraorder of Nematocera, containing the crane flies, a very large group, and several allied families, three of which have historically been included within the Tipulidae....
- has yet to be resolved.
Within the Brachycera, several progressively nested groups exist: Eremoneura (three larval instars), Cyclorrhapha
Cyclorrhapha
Cyclorrhapha is an unranked taxon within the infraorder Muscomorpha. They are called "Cyclorrhapha" with reference to the circular aperture through which the adult escapes the puparium...
(pupation occurs within a puparium), Schizophora
Schizophora
Schizophora is a section of true flies containing 78 families, which are collectively referred to as muscoids, even though - technically - the term "muscoid" should be limited to flies in the superfamily Muscoidea; this is an example of informal, historical usage persisting in the vernacular...
(flies that escape from their puparium using the ptilinal sac, an evertable frontal pouch) and Calyptratae
Calyptratae
Calyptratae is a subsection of Schizophora in the insect order Diptera, commonly referred to as the calyptrate muscoids...
(larger flies with wings that have the calypter
Calypter
A calypter is either of two posterior lobes of the posterior margin of the forewing of flies between the extreme posterior wing base and the alula, which covers the halteres....
, an enlarged basal lobe).
The Schizophora include most of the family-level diversity in Diptera (∼85 families) and more than 50,000 species. The Calyptratae form a monophyletic superfamily. Other monophyletic superfamilies include the Ephydroidea
Ephydroidea
Ephydroidea is a superfamily of flies....
, Lauxanioidea
Lauxanioidea
Lauxanioidea is a superfamily of flies that includes the two large families, the Lauxaniidae and Chamaemyiidae, and the small families Eurychoromyiidae and Celyphidae. Generally they are small to medium, densely populated, coloured flies. Chamaemyiidae and Cremifaniidae live as parasites on insects...
, Nerioidea
Nerioidea
Nerioidea is a superfamily of Acalyptratae flies....
, Sciomyzoidea
Sciomyzoidea
Sciomyzoidea is a superfamily of Acalyptratae flies.The families placed here are:* Coelopidae – seaweed flies* Dryomyzidae* Helosciomyzidae * Ropalomeridae* Heterocheilidae* Sepsidae – scavenger flies...
and Tephritoidea
Tephritoidea
The Tephritoidea are a superfamily of flies. The following families are included:* Lonchaeidae - lance flies* Pallopteridae - flutter flies* Piophilidae - skippers* Platystomatidae - signal flies* Pyrgotidae* Richardiidae* Tephritidae - fruit flies...
. The relationships between the remaining families have yet to be clarified.
Sister groups to the Drosophilidae
Drosophilidae
Drosophilidae is a diverse, cosmopolitan family of flies, which includes fruit flies. Another family of flies called Tephritidae also includes fruit flies. The best known species of Drosophilidae is Drosophila melanogaster, within the genus Drosophila, and this species Is used extensively for...
consist of two families, Braulidae
Braulidae
Braulidae, or bee lice, is a flies family that contains eight species in two genera, Braula and Megabraula. These are very unusual flies, wingless and flattened, and barely recognizable as Diptera....
and Cryptochetidae.
There were three episodes of rapid evolution in the lower Diptera (~220 million years ago), lower Brachycera (180 million years ago) and Schizophora (65 million years ago).
Maggots
Forensic entomology
Forensic entomology is the application and study of insect and other arthropod biology to criminal matters. It is primarily associated with death investigations; however, it may also be used to detect drugs and poisons, determine the location of an incident, and find the presence and time of the...
. By their stage of development, these maggots indicate the time elapsed since death, as well as the place of death.
Maggot species can be identified
Use of DNA in forensic entomology
Forensic entomology contains three aspects: medicocriminal entomology, urban entomology, and stored product entomology. This article focuses more on the medicocriminal aspect and how DNA is analyzed with various blood feeding insects.- Blood meal extraction :...
using their DNA
DNA
Deoxyribonucleic acid is a nucleic acid that contains the genetic instructions used in the development and functioning of all known living organisms . The DNA segments that carry this genetic information are called genes, but other DNA sequences have structural purposes, or are involved in...
. The housefly
Housefly
The housefly , Musca domestica, is a fly of the suborder Cyclorrhapha...
maggot measures 10–20 mm (⅜–¾ in) in size. At the height of the summer season, a generation of flies (egg to adult) may be produced in 12 to 14 days. Other insect families, such as Histeridae, feed on maggots. Thus the lack of maggots would increase the estimated time of death.
Maggots are bred commercially, as a popular bait in angling
Angling
Angling is a method of fishing by means of an "angle" . The hook is usually attached to a fishing line and the line is often attached to a fishing rod. Fishing rods are usually fitted with a fishing reel that functions as a mechanism for storing, retrieving and paying out the line. The hook itself...
, and as a food for carnivorous pets such as reptiles or birds.
Maggots are used in medicine
Maggot therapy
Maggot therapy is a type of biotherapy involving the intentional introduction of live, disinfected maggots into the non-healing skin and soft tissue wound of a human or animal for the purpose of cleaning out the...
to clean out necrotic
Necrosis
Necrosis is the premature death of cells in living tissue. Necrosis is caused by factors external to the cell or tissue, such as infection, toxins, or trauma. This is in contrast to apoptosis, which is a naturally occurring cause of cellular death...
wound
Wound
A wound is a type of injury in which skin is torn, cut or punctured , or where blunt force trauma causes a contusion . In pathology, it specifically refers to a sharp injury which damages the dermis of the skin.-Open:...
s and in food production, particularly of cheeses such as casu marzu
Casu marzu
Casu marzu is a traditional Sardinian sheep milk cheese, notable for containing live insect larvae...
designed to rot as part of their aging process.
Flies are reared in large numbers in Japan
Japan
Japan is an island nation in East Asia. Located in the Pacific Ocean, it lies to the east of the Sea of Japan, China, North Korea, South Korea and Russia, stretching from the Sea of Okhotsk in the north to the East China Sea and Taiwan in the south...
to serve as pollinator
Pollinator
A pollinator is the biotic agent that moves pollen from the male anthers of a flower to the female stigma of a flower to accomplish fertilization or syngamy of the female gamete in the ovule of the flower by the male gamete from the pollen grain...
s of sunflower
Sunflower
Sunflower is an annual plant native to the Americas. It possesses a large inflorescence . The sunflower got its name from its huge, fiery blooms, whose shape and image is often used to depict the sun. The sunflower has a rough, hairy stem, broad, coarsely toothed, rough leaves and circular heads...
s in greenhouses, especially the maggots.
Flies in culture
Flies have been depicted in mythologyMythology
The term mythology can refer either to the study of myths, or to a body or collection of myths. As examples, comparative mythology is the study of connections between myths from different cultures, whereas Greek mythology is the body of myths from ancient Greece...
and literature
Literature
Literature is the art of written works, and is not bound to published sources...
. In the Biblical fourth plague of Egypt
Plagues of Egypt
The Plagues of Egypt , also called the Ten Plagues or the Biblical Plagues, were ten calamities that, according to the biblical Book of Exodus, Israel's God, Yahweh, inflicted upon Egypt to persuade Pharaoh to release the ill-treated Israelites from slavery. Pharaoh capitulated after the tenth...
, flies represent death and decay. Myiagros
Myiagros
Myiagros, or Myagron, in Greek mythology was a god who chased away flies and mice before the sacrifices to Zeus and Athena....
was a god in Greek mythology
Greek mythology
Greek mythology is the body of myths and legends belonging to the ancient Greeks, concerning their gods and heroes, the nature of the world, and the origins and significance of their own cult and ritual practices. They were a part of religion in ancient Greece...
who chased away flies during the sacrifices to Zeus
Zeus
In the ancient Greek religion, Zeus was the "Father of Gods and men" who ruled the Olympians of Mount Olympus as a father ruled the family. He was the god of sky and thunder in Greek mythology. His Roman counterpart is Jupiter and his Etruscan counterpart is Tinia.Zeus was the child of Cronus...
and Athena
Athena
In Greek mythology, Athena, Athenê, or Athene , also referred to as Pallas Athena/Athene , is the goddess of wisdom, courage, inspiration, civilization, warfare, strength, strategy, the arts, crafts, justice, and skill. Minerva, Athena's Roman incarnation, embodies similar attributes. Athena is...
, and Zeus
Zeus
In the ancient Greek religion, Zeus was the "Father of Gods and men" who ruled the Olympians of Mount Olympus as a father ruled the family. He was the god of sky and thunder in Greek mythology. His Roman counterpart is Jupiter and his Etruscan counterpart is Tinia.Zeus was the child of Cronus...
sent a fly to bite Pegasus
Pegasus
Pegasus is one of the best known fantastical as well as mythological creatures in Greek mythology. He is a winged divine horse, usually white in color. He was sired by Poseidon, in his role as horse-god, and foaled by the Gorgon Medusa. He was the brother of Chrysaor, born at a single birthing...
, causing Bellerophon
Bellerophon
Bellerophon or Bellerophontes is a hero of Greek mythology. He was "the greatest hero and slayer of monsters, alongside of Cadmus and Perseus, before the days of Heracles", and his greatest feat was killing the Chimera, a monster that Homer depicted with a lion's head, a goat's body, and a...
to fall back to Earth when he attempted to ride the winged steed to Mount Olympus. In the traditional Navajo
Navajo people
The Navajo of the Southwestern United States are the largest single federally recognized tribe of the United States of America. The Navajo Nation has 300,048 enrolled tribal members. The Navajo Nation constitutes an independent governmental body which manages the Navajo Indian reservation in the...
religion, Big Fly is an important spirit being.
In the 15th-century trompe l'oeil
Trompe l'oeil
Trompe-l'œil, which can also be spelled without the hyphen in English as trompe l'oeil, is an art technique involving extremely realistic imagery in order to create the optical illusion that the depicted objects appear in three dimensions.-History in painting:Although the phrase has its origin in...
painting Portrait of a Carthusian (1446) by Petrus Christus
Petrus Christus
Petrus Christus was an Early Netherlandish painter active in Bruges from 1444.-Life:Christus was born in Baarle, near Antwerp and Breda. Long considered a student of and successor to Jan van Eyck, his paintings have sometimes been confused with those of Van Eyck. At the death of Van Eyck in 1441,...
, a fly sits on a fake frame.
Emily Dickinson's
Emily Dickinson
Emily Elizabeth Dickinson was an American poet. Born in Amherst, Massachusetts, to a successful family with strong community ties, she lived a mostly introverted and reclusive life...
1955
1955 in poetry
Nationality words link to articles with information on the nation's poetry or literature .-Events:* The Group, a British poetry movement, starts meeting in London with gatherings taking place once a week, on Friday evenings, at first at Hobsbaum's flat and later at the house of Edward Lucie-Smith...
poem "I Heard a Fly Buzz When I Died" also makes reference to flies in the context of death. In fact, flies such as the genus Hydrotaea
Hydrotaea
Hydrotaea are insects located in most regions of the world, but are more populous in warmer climates. They are often found on excrement in summer months, and are therefore generally found in close proximity to livestock. Among the 130 known species in this genus, the most commonly recognized is the...
are used in forensic cases to determine time of death for corpses. In William Golding's
William Golding
Sir William Gerald Golding was a British novelist, poet, playwright and Nobel Prize for Literature laureate, best known for his novel Lord of the Flies...
1954
1954 in literature
The year 1954 in literature involved some significant events and new books.-Events:*Jack Kerouac reads Dwight Goddard's A Buddhist Bible, which will influence him greatly.*John Updike graduates from Harvard with a thesis on George Herbert....
novel
Novel
A novel is a book of long narrative in literary prose. The genre has historical roots both in the fields of the medieval and early modern romance and in the tradition of the novella. The latter supplied the present generic term in the late 18th century....
Lord of the Flies
Lord of the Flies
Lord of the Flies is a novel by Nobel Prize-winning author William Golding about a group of British boys stuck on a deserted island who try to govern themselves, with disastrous results...
, the fly is a symbol of the children involved.
In the 1958 science fiction film The Fly
The Fly (1958 film)
The Fly is a 1958 American science-fiction horror film, directed by Kurt Neumann. The screenplay was written by James Clavell , from the short story "The Fly" by George Langelaan...
(and its 1986 remake
The Fly (1986 film)
The Fly is a 1986 science fiction horror film co-written and directed by David Cronenberg. Produced by 20th Century Fox, and Brooksfilms, the film stars Jeff Goldblum, Geena Davis and John Getz. It is a remake of the 1958 film of the same name, but retains only the basic premise of a scientist...
), a scientist accidentally exchanges parts of his body with those of a fly.
Musical works that mention flies: Yoko Ono
Yoko Ono
is a Japanese artist, musician, author and peace activist, known for her work in avant-garde art, music and filmmaking as well as her marriage to John Lennon...
's 1971
1971 in music
-Events:*February 1 – after months of feuding in the press, Ginger Baker and Elvin Jones hold a "drum battle" at The Lyceum.*February 8 – Bob Dylan's hour-long documentary film, Eat the Document, is premièred at New York's Academy of Music...
album Fly
Fly (Yoko Ono album)
Fly is the second album by Yoko Ono. It was produced by John Lennon and Ono, and released in 1971. It was a complete avant-garde/Fluxus package in a gatefold sleeve that came with a full-size poster and a postcard to order Ono's book Grapefruit. Notable songs include the singles "Midsummer New...
, U2
U2
U2 are an Irish rock band from Dublin. Formed in 1976, the group consists of Bono , The Edge , Adam Clayton , and Larry Mullen, Jr. . U2's early sound was rooted in post-punk but eventually grew to incorporate influences from many genres of popular music...
's 1991
1991 in music
See also:* 1991 in music Record labels established in 1991-Summary:The year 1991 is the year that grunge music made its popular breakthrough. Nirvana's Nevermind, led by the surprise hit single "Smells Like Teen Spirit", becomes the most popular U.S. album of the year...
song "The Fly
The Fly (song)
"The Fly" is a song by rock band U2. It is the seventh track from their 1991 album Achtung Baby and it was released as the album's first single on 12 October 1991. "The Fly" introduced a more abrasive sounding U2, as the song featured hip-hop and industrial beats, distorted vocals, and an elaborate...
", Wire
Wire (band)
Wire are an English rock band, formed in London in October 1976 by Colin Newman , Graham Lewis , Bruce Gilbert , and Robert Gotobed...
's 1978
1978 in music
This is a list of notable events in music that took place in the year 1978.-January–April:*January 14 – The Sex Pistols play their final show at San Francisco's Winterland Ballroom....
song "I Am The Fly", Dave Matthews
Dave Matthews
David John "Dave" Matthews is a South African–born American musician and occasional actor, best known as the lead vocalist, songwriter, and guitarist for the Dave Matthews Band...
's 2007
2007 in music
This is a list of notable events in music that took place in the year 2007.-January:*January 1 - George Shearing is knighted for services to music in the Queen's New Year Honours List. Evelyn Glennie becomes a Dame...
song "The Fly" and Béla Bartók's
Béla Bartók
Béla Viktor János Bartók was a Hungarian composer and pianist. He is considered one of the most important composers of the 20th century and is regarded, along with Liszt, as Hungary's greatest composer...
1920s piano work "From the Diary of a Fly".
Damien Hirst
Damien Hirst
Damien Steven Hirst is an English artist, entrepreneur and art collector. He is the most prominent member of the group known as the Young British Artists , who dominated the art scene in Britain during the 1990s. He is internationally renowned, and is reportedly Britain's richest living artist,...
's provocative 1990
1990 in art
-Events:*18 March – Twelve paintings, collectively worth from $100 to $300 million, are stolen from the Isabella Stewart Gardner Museum in Boston, Massachusetts by 2 thieves posing as police officers...
artwork, titled A Thousand Years, featured a severed cow's head contained in a box with thousands of flies and a bug zapper
Bug zapper
A bug zapper, or more formally an electrical discharge insect control system is a device that attracts and kills flying insects that are attracted by light. A light source attracts insects to an electrical grid, where they are electrocuted by touching two wires with a high voltage between them...
, creating an entire life cycle within a glass box. In 2001 Dr. Garnet Hertz implanted a complete web server
Web server
Web server can refer to either the hardware or the software that helps to deliver content that can be accessed through the Internet....
into a dead fly.
The ability of flies to cling to surfaces has also inspired the title of "Human Fly" for real and fictional stunt performers whose feats involve climbing buildings.
Biology
- Harold OldroydHarold OldroydHarold Oldroyd was a British entomologist, born in 1914. He specialised in the biology of flies, and wrote many books, especially popular science that helped entomology to reach a broader public. His "The Natural History of Flies" is considered to be the "fly Bible"...
The Natural History of Flies. New York: W. W. Norton. 1965. - Eugène SéguyEugène SéguyEugène Séguy was a French entomologist who specialised in Diptera. He held a chair of entomology at the Muséum national d'histoire naturelle in Paris from 1956 -1960.Portraits and Number 7, bottom row.-Works:...
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. Text figs. Part of Encyclopedie Entomologique, Serie B II: Diptera. 1924–1953. - Eugène Seguy. La Biologie des Dipteres 1950. pp. 609. 7 col + 3 b/w plates, 225 text figs.
Classification
- Brown, B.V., Borkent, A., Cumming, J.M., Wood, D.M., Woodley, N.E., and Zumbado, M. (Editors) 2009 Manual of Central American Diptera. Volume 1 NRC Research Press, Ottawa ISBN 978-0-660-19833-0
- 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 Univ. Press, Melbourne.
- Griffiths, G.C.D. The phylogenetic classification of Diptera Cyclorrhapha, withspecial reference to the structure of the male postabdomen. Ser. Ent. 8, 340 pp. [Dr. W. Junk, N. V., The Hague] (1972).
- Willi HennigWilli HennigEmil Hans Willi Hennig was a German biologist who is considered the founder of phylogenetic systematics, also known as cladistics. With his works on evolution and systematics he revolutionised the view of the natural order of beings...
Die Larvenformen der Dipteren. 3. Teil. Akad.-Verlag, Berlin. 185 pp., 3 pls. 1948 - Willi Hennig (1954) Flugelgeader und System der Dipteren unter Berucksichtigung der aus dem Mesozoikum beschriebenen Fossilien. Beitr. Ent. 4: 245-388 (1954).
- Willi Hennig: Diptera (Zweifluger). Handb. Zool. Berl. 4 (2 ) (31):1–337. General introduction with key to World Families. In German.
Evolution
- Blagoderov, V.A., Lukashevich, E.D. & Mostovski, M.B. 2002. Order Diptera. In: Rasnitsyn, A.P.Alex RasnitsynRasnitsyn, Alexandr Pavlovich — Soviet and Russian entomologist, expert in palaeoentomology. Honored Scientist of Russian Federation ....
and Quicke, D.L.J. The History of Insects, Kluwer Publ., Dordrecht, Boston, London, pp. 227–240.
External links
- The Diptera Site
- The Dipterists Forum - The Society for the study of flies
- The Bishop Museum Catalog of Fossil Diptera
- The Diptera.info Portal
- The Tree of Life Project
- Manual of Afrotropical Diptera
- BugGuide