Diptera
Encyclopedia
Diptera or true flies, is 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...

 of insect
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 possessing only a single pair of wings 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...

; 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...

 bears a pair of drumstick like structures called the halteres
Halteres
Halteres are small knobbed structures modified from the hind wings in some two-winged insects. They are flapped rapidly and function as gyroscopes, informing the insect about rotation of the body during flight....

, the remnants of the hind wings. It is 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...

, 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 phenomenal historical and current 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....

 and other infectious diseases. The study of the Diptera is called dipterology.

Diptera include flies
Fly
True flies are insects of the order Diptera . They possess a pair of wings on the mesothorax and a pair of halteres, derived from the hind wings, on the metathorax...

, mosquitoes
Mosquito
Mosquitoes are members of a family of nematocerid flies: the Culicidae . The word Mosquito is from the Spanish and Portuguese for little fly...

, maggots, 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 and 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...

. In compound names containing "fly" for members of this order, the name is normally written as two words, as in "crane fly
Crane fly
A crane fly is an insect in the family Tipulidae. Adults are very slender, long-legged flies that may vary in length from though tropical species may exceed to ....

", though there are a few historical exceptions, such as "sandfly
Sandfly
thumb|Sandfly biting a human's little fingerthumb|Sandfly biteSandfly is a colloquial name for any species or genus of flying, biting, blood-sucking Dipteran encountered in sandy areas...

" and "blackfly
Blackfly
Blackfly, black-fly, or black fly may refer to:*Black fly, a fly of the family Simuliidae*Blackfly , a 2001 Canadian comedy series*Aphid or blackfly...

". For insects that are members of other orders the name should always be written as a single word such as in "butterfly
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...

", "scorpionfly".

Ecology and life history

Diptera are a diverse order with an enormous range of ecological roles. Every type of trophic level
Trophic level
The trophic level of an organism is the position it occupies in a food chain. The word trophic derives from the Greek τροφή referring to food or feeding. A food chain represents a succession of organisms that eat another organism and are, in turn, eaten themselves. The number of steps an organism...

 pattern can be seen in the Diptera. Dipteran predators include the robber fly (Asilidae) . A variety of herbivore
Herbivore
Herbivores are organisms that are anatomically and physiologically adapted to eat plant-based foods. Herbivory is a form of consumption in which an organism principally eats autotrophs such as plants, algae and photosynthesizing bacteria. More generally, organisms that feed on autotrophs in...

s can be found in the Diptera, such as the economically important fruit flies (Tephritidae
Tephritidae
Tephritidae is one of two fly families referred to as "fruit flies", the other family being Drosophilidae. Tephritidae does not include the biological model organisms of the genus Drosophila , which is often called the "common fruit fly". There are nearly 5,000 described species of tephritid...

). Flies are often parasites, including internal parasites endo- such as the bot fly and external parasites ectoparasite such as 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...

, black fly
Black fly
A black fly is any member of the family Simuliidae of the Culicomorpha infraorder. They are related to the Ceratopogonidae, Chironomidae, and Thaumaleidae. There are over 1,800 known species of black flies . Most species belong to the immense genus Simulium...

, sand fly or louse fly
Hippoboscidae
Hippoboscidae, the louse flies or keds are obligate parasites of mammals and birds. In this family there are winged species which can fly at least reasonably well, as well as others with vestigial or no wings which are flightless and highly apomorphic...

. Myiasis
Myiasis
Myiasis is a general term for infection by parasitic fly larvae feeding on the host's necrotic or living tissue. Colloquialisms for myiasis include flystrike, blowfly strike, and fly-blown. In Greek, "myia" means fly....

 is the special term for flies infecting living tissue (such as the screw worm fly
Cochliomyia hominivorax
Cochliomyia hominivorax, the New World screw-worm fly, or screw-worm for short, is a species of parasitic fly that is well known for the way in which its larvae eat the living tissue of warm-blooded animals. It is present in the New World tropics...

. Many flies eat dead organic matter detritovores, plant or animal remains. This is especially common in the larval stage, seen in the filter-feeding mosquitoes and black flies to the dung-feeding blow flies (Calliphoridae) or the organic deposit feeding rat-tailed maggot
Rat-tailed maggot
Rat-tailed maggots are the larvae of some species of hoverfly, commonly Eristalis tenax. A characteristic feature of rat-tailed maggots is a tube-like, three segmented, telescoping breathing siphon located at their posterior end. This acts like a snorkel, allowing the larva to breathe air while...

.

The basic fly life cycle is egg, larvae, pupa and adult (winged stage), called holometabolism
Holometabolism
Holometabolism, also called complete metamorphism, is a term applied to insect groups to describe the specific kind of insect development which includes four life stages - as an embryo or egg, a larva, a pupa and an imago or adult. Holometabolism is a monophyletic trait that all insects in the...

. There is often a different in food sources for larvae versus adult dipterans of the same species. For example, [mosquito] larvae live in standing water and feed on detritus while the adults feed on nectar as their energy source while females utilize blood as their energy source for egg production.

Anatomy

CSIRO provides an introduction to the external anatomy of the Diptera. Various species are 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 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).

Classification overview

There are two generally accepted suborders of Diptera. 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....

 are usually 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...

 tend to have a more roundly proportioned body and very short antennae. A more recent classification has been proposed 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....

, but this has not yet gained widespread acceptance among dipterists.
  1. Suborder Nematocera – long antennae, pronotum distinct from mesonotum. In Nematocera larvae are either eucephalic or hemicephalic and often aquatic.
  2. Suborder Brachycera – short antennae, 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...

     is inside a puparium formed from the last larva
    Larva
    A 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. Brachycera are generally robust flies with larvae having reduced mouthparts.
    1. Infraorders Tabanomorpha
      Tabanomorpha
      The 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 Asilomorpha
      Asilomorpha
      The 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 Orthorrhapha
      Orthorrhapha
      Orthorrhapha 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
      Muscomorpha
      The 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...

      .
    2. Infraorder Muscomorpha
      Muscomorpha
      The 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 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...

       of older schemes). Muscomorpha have 3-segmented, aristate (with a bristle) 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....

       and larvae with three instar
      Instar
      An 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 that are acephalic (maggots).

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 more out of practical or historical considerations than out of any strict respect for phylogenetic classifications. (Modern cladists tend to spurn the use of Linnaean rank names.) Nearly 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.

Current suborders and families

There are two suborders: the Nematocera divided into 77 families of which 35 are extinct ( fossil
Fossil
Fossils are the preserved remains or traces of animals , plants, and other organisms from the remote past...

 only ) and the Brachycera with 141 families of which 8 are extinct. See Families of Diptera
Families of Diptera
-"Nematocera":*Anisopodidae Edwards , 1921 Synonyms: Phryneidae, Rhyphidae, Sylvicolidae, Mycetobiidae, Protolbiogastridae.*Ansorgiidae Extinct.*Antefungivoridae Extinct...

.

Obsolete names in diptera

There are many names at the family level or above that have been used historically, even some of recent vintage, that are already obsolete or simply not recognized or universally accepted by the dipterological community. For a rather exhaustive list of such names (esp. for those readers who may be wondering why they cannot find a familiar name), please see the List of obsolete names in Diptera.

Biology

  • Harold Oldroyd
    Harold Oldroyd
    Harold 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 Séguy
    Eugène Séguy
    Eugène 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

  • 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).
  • Hendel, F. 1935 Bemerkungen zu "The families and genera of North American Diptera" by C. H.Curran,New-York 1934. Konowia 14: 51-57. [1935.03.15]
  • Willi Hennig
    Willi Hennig
    Emil 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 and
  • Flugelgeader und System der Dipteren unter Berucksichtigung der aus dem Mesozoikum beschriebenen Fossilien. Beitr. Ent. 4: 245-388 (1954).

A very well-researched reference list of works on the Phylogeny (Classification and Identification of Diptera) is provided by Thompson as a pdf http://www.sel.barc.usda.gov/diptera/names/BDWDsour.pdf

Evolution

Blagoderov, V.A., Lukashevich, E.D. & Mostovski, M.B. 2002. Order Diptera. In: Rasnitsyn, A. P.
Alex Rasnitsyn
Rasnitsyn, 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.

Identification

  • Erwin Lindner
    Erwin Lindner
    Erwin Lindner was a German entomologist mainly interested in Diptera.He was born in Böglins, Memmingen , 7 April 1888 and died in Stuttgart 30 November 1988, aged 100 years....

     Die Fliegen der Paläarktischen Region. The available parts of this very large work are listed on the publishers website http://www.schweizerbart.de/pubs/series/lindner-erwin:-die-fliegen-der-palaearktischen-region132.html. Band 1 Handbuch, by Lindner himself, contains a history of dipterology, with sections on morphology
    Morphology (biology)
    In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....

    , physiology
    Physiology
    Physiology is the science of the function of living systems. This includes how organisms, organ systems, organs, cells, and bio-molecules carry out the chemical or physical functions that exist in a living system. The highest honor awarded in physiology is the Nobel Prize in Physiology or...

     and taxonomy
    Taxonomy
    Taxonomy is the science of identifying and naming species, and arranging them into a classification. The field of taxonomy, sometimes referred to as "biological taxonomy", revolves around the description and use of taxonomic units, known as taxa...

     with a key to families and short notes on each. The work is finished and almost all parts are out of date but it remains the essential work.
  • Faune de France Lechevalier, Paris. The many parts of this work are listed on the publishers website http://www.insects.demon.co.uk/faune-france.html. Less comprehensive than Lindner and , similarly, out of date but more convenient to use. The best parts are Parent on Dolichopodidae
    Dolichopodidae
    Dolichopodidae, the long-legged flies, make up a large family of true flies with more than 7,000 described species in about 230 genera distributed worldwide. The genus Dolichopus is the most speciose, with some 600 species. They are generally small flies with large, prominent eyes and a metallic...

     and Séguy’s large part 28 on Muscidae
    Muscidae
    Muscidae are a family of flies found in the superfamily Muscoidea. The apical segment of the antennae of Muscidae are plumose, and the basal portion is smooth....

     (including Anthomyiidae
    Anthomyiidae
    Anthomyiidae is a large and diverse family of Muscoidea flies. Name came from Greek "anthos" + "myia" . Some species are commonly called "root-maggots", as the larvae are found in the stems and roots of various plants...

    ) Scathophagidae
    Scathophagidae
    The Scathophagidae is a small family of Muscoidea which are often known as "Dung-flies" although this name is not appropriate except for a few species of the genus Scathophaga which do indeed pass their larval stages in animal dung. The name probably derives from the "Common Yellow Dung-fly", S...

     and 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...

    ).
  • Diptera of European part of Russia and Diptera of Far East of the U.S.S.R. in Fauna of the U.S.S.R. (New Series Fauna SSSR)

(Nasekomye dvukrylye) Chief Editor: B.E. Bykhovskii; Editorial board: I.M. Gromov, A.S. Monchadskii, A .A. Shtakel’berg, 0-A. Skarlato, and A.A. Strelkov (Volume editor) Izdatel’stvo “Nauka”Leningradskoe Otdelenie Leningrad (St. Petersburg). In Russian but with very useful figures. Some parts have been translated. Can anyone add a full list and availalibility details?
  • Willi Hennig
    Willi Hennig
    Emil 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...

     Diptera (Zweifluger). Handb. Zool. Berl. 4 (2 ) (31):1-337. General introduction with key to World Families. In German.
  • László Papp and Béla Darvas Contributions to a Manual of Palaearctic Diptera. This book series was written by a group of international scientists. (1992-2000).
  • McAlpine, J. P. (ed.) Manual of Nearctic Diptera. Research Branch, Agriculture Canada Monograph 27, 28 & 32 1981-89. Very good family and other keys.
  • Smith, KGV 1989 An introduction to the immature stages of British flies. Diptera (14). Handbks. Ident. Br. Insects 10(14). RESL Excellent introduction to Diptera larvae.

Famous dipterists

A list of famous Dipterists, the term for entomologists who specialised in the order Diptera, is found under Dipterists
Early entomologists who described Diptera as parts of general entomological works are listed under Entomologists examples are:

External links

The source of this article is wikipedia, the free encyclopedia.  The text of this article is licensed under the GFDL.
 
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