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'{{for||fly (disambiguation)|flies (disambiguation)}} {{Automatic taxobox |fossil_range= [[Middle Triassic]] - Recent {{fossilrange|245|0}} | image = Diptera1.jpg | image_caption = A poster with sixteen different species of flies | taxon = Diptera | authority = [[Carolus Linnaeus|Linnaeus]], [[Systema Naturae|1758]] | subdivision_ranks = Suborders | subdivision = [[Nematocera]] (includes [[Eudiptera]])<br> [[Brachycera]] }} True '''flies''' are [[insect]]s of the [[Order (biology)|order]] '''Diptera''' (from the Greek ''di'' = two, and ''ptera'' = wings). They possess a pair of [[insect wing|wing]]s on the [[mesothorax]] and a pair of [[haltere]]s, 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. == 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 [[mayfly|mayflies]], [[dragonfly|dragonflies]], [[damselfly|damselflies]], [[stonefly|stoneflies]], [[whitefly|whiteflies]], [[firefly|fireflies]], [[alderfly|alderflies]], [[dobsonfly|dobsonflies]], [[snakefly|snakeflies]], [[sawfly|sawflies]], [[caddisfly|caddisflies]], [[butterfly|butterflies]] or [[scorpionfly|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.<ref>[http://bugguide.net/node/view/55 Bug guide]</ref> 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, [[midge (insect)|midges]] and others, although under half of these (about 120,000 species) have been described.<ref>{{cite web |author=B. M. Wiegmann & D. K. Yeates |year=1996 |title=Tree of Life: Diptera |url=http://www2.ncsu.edu/unity/lockers/ftp/bwiegman/fly_html/diptera.html#about}}</ref> 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 [[Vector (epidemiology)|vectors]] for [[malaria]], [[dengue]], [[West Nile virus]], [[yellow fever]], [[encephalitis]] and other [[infectious diseases]]. ==Anatomy and biology== [[File:Musca domestica Portrait.jpg|thumb|left|Portrait of a [[flesh fly]] (Sarcophagidae)]] [[File:FLY EYE.jpg|thumb|right|An image of a house fly [[compound eye|eye]] surface by using [[scanning electron microscope]] at 450× magnification|300px]] 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 eye]]s, and the [[insect mouthparts|mouthparts]] (the labrum, labium, mandible and maxilla make up the mouthparts). The second tagma, the [[Thorax (insect anatomy)|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 [[ganglion|ganglia]], and concentration of nerve tissue in the thorax, a feature that is most extreme in the highly derived Muscomorpha infraorder.<ref name=IIBD>{{cite book |author= Hoell, H.V., Doyen, J.T. & Purcell, A.H. |year=1998 |title=Introduction to Insect Biology and Diversity, 2nd ed. |publisher= Oxford University Press |pages= 493–499|isbn= 0-19-510033-6}}</ref> [[File:Fly Eye 30wd 3x3.JPG#file|thumb|right|A scan of a house fly taken at 40 magnifications under a [[scanning electron microscope]]]] Flies have a mobile head with eyes and in most cases have large [[compound eye]]s on the sides of the head, with three small [[ocelli]] on the top. The [[antenna (biology)|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 [[diverticula]]e, allowing the insect to store small quantities of liquid after a meal.<ref name=IIBD/> ===Reproduction and development=== [[File:Anthomyiidae sp. 1 (aka).jpg|thumb|200px|Mating [[Anthomyiidae|anthomyiid flies]]]] 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.<ref name=IIBD/> 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.<ref name=IIBD/> [[File:Larve de moustique.JPG|thumb|Mosquito larva. A typical Nematoceran. Note the head capsule and clearly defined three segments of the thorax]] [[File:Syrphid.maggot3554.5.13.08cw.jpg|thumb|Syrphid larva. Maggot-like anatomy, typical of a Brachycerid fly, lacking both a conspicuously demarcated thorax and head capsule]] 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.<ref>{{cite book | author = Chapman, R. F.| title = The Insects; Structure & Function | publisher = Cambridge University Press | location = Cambridge | year = 1998 | isbn = 9780521578905 }}</ref> 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 (insect anatomy)|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,<ref name="isbn0-19-861271-0">{{cite book |author=Brown, Lesley |title=The New shorter Oxford English dictionary on historical principles |publisher=Clarendon |location=Oxford [Eng.] |year=1993 |pages= |isbn=0-19-861271-0 |oclc= |doi= |accessdate=}}</ref> 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 [[cercus|cerci]]. This lack of features is an adaptation to food such as carrion, decaying detritus, or host tissues surrounding [[Parasitism|endoparasites]].<ref name=IIBD/> 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. ==Classification == {{See also|List of families of Diptera}} [[File:Fly cleaning.ogv|thumb|right|200px|Cleaning]] [[File:Closeup black fly.jpg|thumb|right|200px| A fly]] The [[Nematocera]] 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]].<ref>B.B. Rohdendorf. 1964. Trans. Inst. Paleont., Acad. Sci. USSR, Moscow, v. 100</ref> #Suborder [[Nematocera]] (77 families, 35 of them extinct) &ndash; long antennae, [[pronotum]] distinct from [[mesonotum]], in Nematocera, larvae are either eucephalic or hemicephalic and often aquatic. #Suborder [[Brachycera]] (141 families, 8 of them extinct) &ndash; short antennae, the [[pupa]] is inside a puparium formed from the last [[larva]]l skin, they are generally robust flies with larvae having reduced mouthparts. ##Infraorders [[Tabanomorpha]] and [[Asilomorpha]] &ndash; these comprise the majority of what was the [[Orthorrhapha]] under older classification schemes. The antennae are short, but differ in structure from those of the Muscomorpha. ##Infraorder [[Muscomorpha]] &ndash; (largely the [[Cyclorrhapha]] of older schemes). Muscomorpha have three-segmented, [[Arista (biology)|aristate]] (with a bristle) [[antenna (biology)|antennae]] and larvae with three acephalic [[instar]]s (maggots). 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.<ref>{{cite web |url=http://www.taxonomy.nl/Taxonomicon/TaxonTree.aspx?id=102510 |title=Taxon: Superorder Antliophora |work=The Taxonomicon |accessdate=2007-08-21}}</ref> ==Evolution== Diptera derive from [[Mecoptera]] or a strictly related group. The first true dipterans are known from the Middle [[Triassic]], becoming widespread during the Middle and Late Triassic&nbsp;.<ref>{{cite book |url=http://palaeoentomolog.ru/New/diptera.html |chapter=Order Diptera Linné, 1758. The true flies |author=V. A. Blagoderov, E. D. Lukashevich & M. B. Mostovski |title=History of Insects |publisher=[[Kluwer Academic Publishers]] |isbn=1-4020-0026-X |editor=A. P. Rasnitsyn & D. L. J. Quicke |year=2002}}</ref> The basal clades in the Diptera are the [[Deuterophlebiidae]] and [[Nymphomyiidae]].<ref name="Wiegmann2011">{{cite journal |last=Wiegmann |first=B. M. |last2=''et al.'' |year=2011 |title=Episodic radiations in the fly tree of life |journal=[[Proceedings of the National Academy of Sciences|PNAS]] |volume=108 |issue=14 |pages=5690–5695 |doi=10.1073/pnas.1012675108 |first2=M. D. |last3=Winkler |first3=I. S. |last4=Barr |first4=N. B. |last5=Kim |first5=J.-W. |last6=Lambkin |first6=C. |last7=Bertone |first7=M. A. |last8=Cassel |first8=B. K. |last9=Bayless |first9=K. M. |pmid=21402926}}</ref> 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).<ref name="Wiegmann2011" /> ==Maggots== {{further|[[Maggot]]}} [[File:Maggots.jpg|right|220px|thumb|[[Maggot]]s being used to treat a wound]] Maggots found on corpses are useful to [[forensic science|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 [[Use of DNA in forensic entomology|identified]] using their [[DNA]]. The [[housefly]] maggot measures 10–20&nbsp;mm (⅜–¾&nbsp;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 [[maggot therapy|used in medicine]] to clean out [[necrosis|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. ==Flies in culture== Flies have been depicted in [[mythology]] and [[literature]]. In the Biblical [[Plagues of Egypt|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 (Mountain)|Mount Olympus]]. In the traditional [[Navajo people|Navajo]] religion, [http://www.coyoteandanotherone.com/insect_folder/fly_page.html 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.<ref>{{cite web |url=http://www.metmuseum.org/toah/hd/optg/hod_49.7.19.htm |title=Portrait of a Carthusian, 1446 |work=Timeline of Art History |publisher=[[The Metropolitan Museum of Art]] |month=October | year=2006}}</ref> [[Emily Dickinson|Emily Dickinson's]] [[1955 in poetry|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|William Golding's]] [[1954 in literature|1954]] [[novel]] ''[[Lord of the Flies]]'', the fly is a symbol of the children involved. In the [[1958 in cinema|1958]] science fiction film ''[[The Fly (1958 film)|The Fly]]'' (and its [[1986 in cinema|1986]] [[The Fly (1986 film)|remake]]), a scientist accidentally exchanges parts of his body with those of a fly. Musical works that mention flies: [[Yoko Ono]]'s [[1971 in music|1971]] album ''[[Fly (Yoko Ono album)|Fly]]'', [[U2]]'s [[1991 in music|1991]] song "[[The Fly (song)|The Fly]]", [[Wire (band)|Wire]]'s [[1978 in music|1978]] song "I Am The Fly", [[Dave Matthews]]'s [[2007 in music|2007]] song "[[The Fly (Dave Matthews song)|The Fly]]" and [[Béla Bartók|Béla Bartók's]] 1920s piano work [[Mikrokosmos (Béla Bartók)|"From the Diary of a Fly"]]. [[Damien Hirst]]'s provocative [[1990 in art|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.<ref>http://www.conceptlab.com/fly/</ref> The ability of flies to cling to surfaces has also inspired the title of "[[Human Fly (disambiguation)|Human Fly]]" for real and fictional stunt performers whose feats involve climbing buildings. ==See also== * [[Female sperm storage]] == Notes ==<!-- Syst. Biol. 55(5):715 --> {{Reflist}} == References == ===Biology=== *[[Harold Oldroyd]] ''The Natural History of Flies''. New York: W. W. Norton. 1965. *[[Eugène Séguy]] ''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.&nbsp;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 ''[http://pubs.nrc-cnrc.gc.ca/eng/books/books/9780660198330.html 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.&nbsp;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 Hennig]] ''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). *{{cite web |url=http://www.sel.barc.usda.gov/diptera/names/BDWDsour.pdf |title=Sources for the Biosystematic Database of World Diptera (Flies) |author=F. Christian Thompson |publisher=[[United States Department of Agriculture]], Systematic Entomology Laboratory|format=PDF}} *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. [http://palaeoentomolog.ru/New/diptera.html Order Diptera]. In: [[Alex Rasnitsyn|Rasnitsyn, A.P.]] and Quicke, D.L.J. The History of Insects, Kluwer Publ., Dordrecht, Boston, London, pp.&nbsp;227–240. ==External links== {{wikiquote|Flies}} {{Commons category|Diptera}} {{Wikispecies|Diptera}} {{Wikisource1911Enc|Fly}} *[http://www.sel.barc.usda.gov/Diptera/biosys.htm The Diptera Site] *[http://www.dipteristsforum.org.uk/ The Dipterists Forum] - The Society for the study of flies *[http://hbs.bishopmuseum.org/fossilcat/ The Bishop Museum Catalog of Fossil Diptera] *[http://www.diptera.info The Diptera.info Portal] *[http://tolweb.org/tree?group=Diptera The Tree of Life Project] *{{dmoz|Science/Biology/Flora_and_Fauna/Animalia/Arthropoda/Insecta/Diptera/|Diptera}} *[http://afrotropicalmanual.net Manual of Afrotropical Diptera] *[http://bugguide.net/node/view/55 BugGuide] {{Diptera}} {{Orders of Insects}} [[Category:Flies| ]] [[Category:Orders of insects]] [[Category:Pollinators]] [[ar:ذوات الجناحين]] [[gn:Mberu]] [[ay:Chhichhillanka]] [[az:İkiqanadlılar]] [[bn:মাছি]] [[zh-min-nan:Hô͘-sîn]] [[be:Мухі]] [[be-x-old:Мухі]] [[bg:Двукрили]] [[ca:Dípter]] [[cs:Dvoukřídlí]] [[da:Tovinger]] [[de:Zweiflügler]] [[nv:Tsʼíʼii dóó Tséʼédǫ́ʼii]] [[et:Kahetiivalised]] [[el:Μύγα]] [[es:Diptera]] [[eo:Dipteroj]] [[eu:Diptera]] [[fa:دوبالان]] [[fr:Diptera]] [[ga:Cuileog]] [[gl:Mosca]] [[ko:파리목]] [[hi:मक्खी]] [[hr:Dvokrilci]] [[io:Diptero]] [[id:Lalat]] [[is:Tvívængjur]] [[it:Diptera]] [[he:זבובאים]] [[jv:Laler]] [[ka:ორფრთიანები]] [[kk:Қосқанаттылар]] [[kv:Гут]] [[ht:Mouch]] [[ku:Mêş]] [[lv:Divspārņu kārta]] [[lt:Dvisparniai]] [[hu:Kétszárnyúak]] [[mg:Lalitra]] [[ml:ഡിപ്‌റ്റെറ]] [[xmf:ჭანდი]] [[ms:Lalat]] [[nl:Tweevleugeligen]] [[ja:ハエ目]] [[ce:Moza]] [[frr:Twiijügeten]] [[no:Tovinger]] [[nn:Tovengjer]] [[mhr:Карме]] [[koi:Гут]] [[pl:Muchówki]] [[pt:Diptera]] [[ro:Diptera]] [[qu:Iskaylla rapra]] [[ru:Двукрылые]] [[simple:Fly]] [[sk:Dvojkrídlovce]] [[sl:Dvokrilci]] [[sh:Dvokrilci]] [[fi:Kaksisiipiset]] [[sv:Tvåvingar]] [[ta:இருசிறகிப் பூச்சிகள்]] [[te:ఈగ]] [[th:แมลงวัน]] [[tr:Sinek]] [[uk:Двокрилі]] [[vi:Ruồi]] [[war:Langaw]] [[yi:פליג]] [[yo:Esinsin]] [[zh:雙翅目]]'