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== Human interaction and cultural depictions == {{further |Human interactions with insects}} ===Symbolism=== [[File:Christus carthusian.jpg|thumb|left|upright |[[Petrus Christus]]'s 1446 painting ''Portrait of a Carthusian'' has a ''musca depicta'' (painted fly) on a ''[[trompe-l'œil]]'' frame.]] Flies play a variety of symbolic roles in different cultures. These include both positive and negative roles in religion. In the traditional [[Navajo]] religion, Big Fly is an important spirit being.<ref>{{cite book |title=Handbook of North American Indians |chapter-url=http://www.users.humboldt.edu/jwpowell/LelandWymanHONAI-NavajoCeremonialSystem,reduced.pdf |page=539 |chapter=Navajo Ceremonial System |last=Wyman |first=Leland Clifton |year=1983 |quote=Nearly every element in the universe may be thus personalized, and even the least of these such as tiny Chipmunk and those little insect helpers and mentors of deity and man in the myths, Big Fly (''Dǫ’soh'') and Ripener (Corn Beetle) Girl (''’Anilt’ánii ’At’ééd'') (Wyman and Bailey 1964:29–30, 51, 137–144), are as necessary for the harmonious balance of the universe as is the great Sun. |access-date=30 July 2015 |archive-url=https://web.archive.org/web/20160305073542/http://www.users.humboldt.edu/jwpowell/LelandWymanHONAI-NavajoCeremonialSystem,reduced.pdf |archive-date=5 March 2016 |url-status=dead|df=dmy-all }}</ref><ref>{{cite book |last1=Wyman |first1=Leland Clifton |last2=Bailey |first2=Flora L. |title=Navaho Indian Ethnoentomology|series=Anthropology Series |issue=12–13 |url=https://books.google.com/books?id=tHl1AAAAMAAJ&q=Big+Fly |date=1964 |publisher=University of New Mexico Press |isbn=9780826301109 |lccn=64024356}}</ref><ref>{{cite web |url=http://www.native-languages.org/legends-fly.htm |title=Native American Fly Mythology |publisher=Native Languages of the Americas website}}</ref> In [[Christian demonology]], [[Beelzebub]] is a demonic fly, the "Lord of the Flies", and a god of the [[Philistine]]s.<ref>"Βεελζεβούλ, ὁ indecl. (v.l. Βεελζεβούβ and Βεεζεβούλ W-S. §5, 31, cp. 27 n. 56) Beelzebul, orig. a Philistine deity; the name בַּעַל זְבוּב means Baal (lord) of the flies (4 Km 1:2, 6; Sym. transcribes βεελζεβούβ; Vulgate Beelzebub; TestSol freq. Βεελζεβούλ,-βουέλ).", Arndt, W., Danker, F. W., & Bauer, W. (2000). A Greek-English lexicon of the New Testament and other early Christian literature (3rd ed.) (173). Chicago: University of Chicago Press.</ref><ref>"1. According to 2 Kgs 1:2–6 the name of the Philistine god of Ekron was Lord of the Flies (Heb. ba‘al zeaûḇ), from whom Israel’s King Ahaziah requested an oracle.", Balz, H. R., & Schneider, G. (1990–). Vol. 1: Exegetical dictionary of the New Testament (211). Grand Rapids, Mich.: Eerdmans.</ref><ref>"For etymological reasons, Baal Zebub must be considered a Semitic god; he is taken over by the Philistine Ekronites and incorporated into their local cult.", Herrmann, "Baal Zebub", in Toorn, K., Becking, B., & Horst, P. W. (1999). Dictionary of deities and demons in the Bible DDD (2nd extensively rev. ed.) (154). Leiden; Boston; Grand Rapids, Mich.: Brill; Eerdmans.</ref> Flies have appeared in literature since ancient [[Sumer]].<ref name="BlackGreen1992">{{cite book |last1=Black |first1=Jeremy |first2=Anthony |last2=Green |title=Gods, Demons and Symbols of Ancient Mesopotamia: An Illustrated Dictionary |url=https://books.google.com/books?id=05LXAAAAMAAJ&q=Inana |publisher=[[The British Museum Press]] |year=1992 |isbn=978-0-7141-1705-8 |pages=84–85}}</ref> In a Sumerian poem, a fly helps the goddess [[Inanna]] when her husband [[Dumuzid]] is being chased by ''[[Gallu|galla]]'' demons.<ref name="BlackGreen1992"/> In the Mesopotamian versions of the [[flood myth]], the dead corpses floating on the waters are compared to flies.<ref name="BlackGreen1992"/> Later, the gods are said to swarm "like flies" around the hero [[Utnapishtim]]'s offering.<ref name="BlackGreen1992"/> Flies appear on [[Babylonia|Old Babylonian]] seals as symbols of [[Nergal]], the god of death.<ref name="BlackGreen1992"/> Fly-shaped [[lapis lazuli]] beads were often worn in ancient Mesopotamia, along with other kinds of fly-jewellery.<ref name="BlackGreen1992"/> In Ancient Egypt, flies appear in amulets and as a military award for bravery and tenacity, due to the fact that they always come back when swatted at. It is thought that flies may have also been associated with the departing spirit of the dead, as they are often found near dead bodies. In modern Egypt, a similar belief persists in some areas to not swat at shiny green flies, as they may be carrying the soul of a recently deceased person.<ref>{{Cite book |last=Haynes |first=Dawn |url=https://scholar.sun.ac.za/bitstream/10019.1/79920/1/haynes_symbolism_2013.pdf |title=The Symbolism and Significance of the Butterfly in Ancient Egypt}}</ref> In a little-known [[Greek myth]], a very chatty and talkative maiden named [[Myia (mythology)|Myia]] (meaning "fly") enraged the moon-goddess [[Selene]] by attempting to seduce her lover, the sleeping [[Endymion (mythology)|Endymion]], and was thus turned by the angry goddess into a fly, who now always deprives people of their sleep in memory of her past life.<ref>{{cite book | title = Metamorphosis in Greek Myths | first = Paul M. C. | last = Forbes Irving | publisher = [[Clarendon Press]] | date = 1990 | url = https://books.google.com/books?id=URvXAAAAMAAJ | isbn = 0-19-814730-9 | location = Oxford | page = 315}}</ref><ref>{{cite book | page = [https://books.google.com/books?id=oNWFDk5mOvQC&pg=PA5 5] | title = Lucian: Selected Dialogues | author1 = Lucian | author2 = C. D. N. Costa | author-link1 = Lucian | location = Oxford, New York | date = 2005 | publisher = [[Oxford University Press]] | url = https://books.google.com/books?id=oNWFDk5mOvQC | isbn = 978-0-19-925867-3}}</ref> In ''[[Prometheus Bound]]'', which is attributed to the Athenian tragic playwright [[Aeschylus]], a [[horse-fly|gadfly]] sent by [[Zeus]]'s wife [[Hera]] pursues and torments his mistress [[Io (mythology)|Io]], who has been transformed into a cow and is watched constantly by the hundred eyes of the herdsman [[Argus Panoptes|Argus]]:<ref>{{cite book |last1=Belfiore |first1=Elizabeth S. |title=Murder among Friends: Violation of Philia in Greek Tragedy |date=2000 |publisher=[[Oxford University Press]] |location=Oxford, England |isbn=978-0-19-513149-9 |page=47 |url=https://books.google.com/books?id=zhqCDBBEXSkC&q=Prometheus+Bound+Io+gadfly&pg=PA47}}</ref><ref name="Stagman2010">{{cite book |last=Stagman |first=Myron |title=Shakespeare's Greek Drama Secret |url=https://books.google.com/books?id=encnBwAAQBAJ&pg=PA207 |date=11 August 2010 |publisher=Cambridge Scholars Publishing |isbn=978-1-4438-2466-8 |pages=205–208}}</ref> "Io: Ah! Hah! Again the prick, the stab of gadfly-sting! O earth, earth, hide, the hollow shape—Argus—that evil thing—the hundred-eyed."<ref name="Stagman2010"/> [[William Shakespeare]], inspired by Aeschylus, has [[Tom o'Bedlam]] in ''[[King Lear]]'', "Whom the foul fiend hath led through fire and through flame, through ford and whirlpool, o'er bog and quagmire", driven mad by the constant pursuit.<ref name="Stagman2010"/> In ''[[Antony and Cleopatra]]'', Shakespeare similarly likens Cleopatra's hasty departure from the [[Battle of Actium|Actium battlefield]] to that of a cow chased by a gadfly.<ref name="Walker2002">{{cite book |last=Walker |first=John Lewis |title=Shakespeare and the Classical Tradition: An Annotated Bibliography, 1961–1991 |url=https://books.google.com/books?id=q3Voi0Dk_HwC&pg=PA363 |year=2002 |publisher=Taylor & Francis |isbn=978-0-8240-6697-0 |page=363}}</ref> More recently, in 1962 the biologist Vincent Dethier wrote ''To Know a Fly'', introducing the general reader to the behaviour and physiology of the fly.<ref>{{cite book |last1=Dethier |first1=Vincent G. |title=To Know a Fly |url=https://archive.org/details/toknowfly00deth |url-access=registration |date=1962 |publisher=Holden-Day |location=San Francisco}}</ref> ''[[Musca depicta]]'' ("painted fly" in Latin) is a depiction of a fly as an inconspicuous element of various paintings. This feature was widespread in 15th and 16th centuries paintings and its presence may be explained by various reasons.<ref name=eoi>''Encyclopedia of Insects'', [https://books.google.com/books?id=Jk0Hym1yF0cC&dq=%22musca+depicta%22&pg=PA242 p. 242]</ref> Flies appear in popular culture in concepts such as [[fly-on-the-wall]] [[documentary]]-making in [[filmmaking|film]] and [[television production]]. The [[metaphor]]ic name suggests that events are seen [[Candid photography|candidly]], as a fly might see them.<ref>{{cite web |url=http://www.screenonline.org.uk/tv/id/698785/ |publisher=British Film Institute |title=Fly on the Wall |access-date=21 July 2016}}</ref> Flies have inspired the design of miniature flying robots.<ref>{{cite journal |last1=Ma |first1=Kevin Y. |last2=Chirarattananon |first2=Pakpong |last3=Fuller |first3=Sawyer B. |last4=Wood |first4=Robert J. |date=2013-05-03 |title=Controlled flight of a biologically inspired, insect-scale robot |journal=Science |volume=340 |issue=6132 |pages=603–607 |doi=10.1126/science.1231806 |pmid=23641114|bibcode=2013Sci...340..603M |s2cid=21912409 }}</ref> [[Steven Spielberg]]'s 1993 film ''[[Jurassic Park]]'' relied on the idea that [[DNA]] could be preserved in the stomach contents of a blood-sucking fly [[fossil]]ised in [[amber]], though the mechanism has been discounted by scientists.<ref>{{cite news |last1=Gray |first1=Richard |title=Jurassic Park ruled out – dinosaur DNA could not survive in amber |url=https://www.telegraph.co.uk/news/science/dinosaurs/10303795/Jurassic-Park-ruled-out-dinosaur-DNA-could-not-survive-in-amber.html |access-date=21 July 2016 |agency=Daily Telegraph |date=12 September 2013}}</ref> ===Economic importance=== [[File:Anopheles stephensi.jpeg|thumb|An ''[[Anopheles stephensi]]'' mosquito drinking human blood. The species carries [[malaria]].]] Dipterans are an important group of insects and have a considerable impact on the environment. Some [[leaf-miner flies]] (Agromyzidae), [[Tephritidae|fruit flies]] (Tephritidae and Drosophilidae) and [[gall midges]] (Cecidomyiidae) are pests of agricultural crops; others such as [[tsetse flies]], [[screwworm]] and [[botflies]] (Oestridae) attack livestock, causing wounds, spreading disease, and creating significant economic harm. See article: [[Parasitic flies of domestic animals]]. A few can even cause [[myiasis]] in humans. Still others such as [[mosquito]]es (Culicidae), [[Black fly|blackflies]] (Simuliidae) and [[drain flies]] (Psychodidae) impact human health, acting as [[Vector (epidemiology)|vectors]] of major tropical diseases. Among these, ''[[Anopheles]]'' mosquitoes transmit [[malaria]], [[filariasis]], and [[arboviruses]]; ''[[Aedes aegypti]]'' mosquitoes carry [[dengue fever]] and the [[Zika virus]]; blackflies carry [[river blindness]]; sand flies carry [[leishmaniasis]]. Other dipterans are a nuisance to humans, especially when present in large numbers; these include houseflies, which contaminate food and spread food-borne illnesses; the [[Ceratopogonidae|biting midges and sandflies]] (Ceratopogonidae) and the [[Muscidae|houseflies and stable flies]] (Muscidae).<ref name=Resh>{{cite book |last1=Resh |first1=Vincent H. |last2=Cardé |first2=Ring T. |title=Encyclopedia of Insects |url=https://books.google.com/books?id=Jk0Hym1yF0cC&pg=PA284 |year=2009 |publisher=Academic Press |isbn=978-0-08-092090-0 |pages=284–297}}</ref> In tropical regions, eye flies ([[Chloropidae]]) which visit the eye in search of fluids can be a nuisance in some seasons.<ref>{{Cite journal |last1=Mulla |first1=Mir S. |last2=Chansang |first2=Uruyakorn |title=Pestiferous nature, resting sites, aggregation, and host-seeking behavior of the eye fly ''Siphunculina funicola'' (Diptera: Chloropidae) in Thailand |journal=Journal of Vector Ecology |volume=32 |issue=2 |doi=10.3376/1081-1710(2007)32[292:pnrsaa]2.0.co;2 |pages=292–301 |year=2007 |pmid=18260520 |s2cid=28636403 }}</ref> Many dipterans serve roles that are useful to humans. Houseflies, blowflies and [[Mycetophilidae|fungus gnats]] (Mycetophilidae) are scavengers and aid in decomposition. [[Robber flies]] (Asilidae), [[Tachinidae|tachinids]] (Tachinidae) and [[Empididae|dagger flies and balloon flies]] (Empididae) are predators and parasitoids of other insects, helping to control a variety of pests. Many dipterans such as [[bee flies]] (Bombyliidae) and [[hoverflies]] (Syrphidae) are [[Pollination|pollinators]] of crop plants.<ref name=Resh/> === Uses === [[File:Researcher beside Vials closed by cotton plugs and filled with Fruit fly larvae.jpg|thumb|left|Diptera in research: ''[[Drosophila melanogaster]]'' fruit fly larvae being bred in tubes in a [[genetics]] laboratory]] ''[[Drosophila melanogaster]]'', a fruit fly, has long been used as a [[model organism]] in research because of the ease with which it can be bred and reared in the laboratory, its small [[genome]], and the fact that many of its genes have counterparts in higher [[eukaryote]]s. A large number of genetic studies have been undertaken based on this species; these have had a profound impact on the study of [[gene expression]], [[Regulation of gene expression|gene regulatory mechanisms]] and [[mutation]]. Other studies have investigated [[physiology]], [[Pathogenesis|microbial pathogenesis]] and development among other research topics.<ref>{{cite web |url=https://www.yourgenome.org/theme/fruit-flies-in-the-laboratory/ |title=Fruit flies in the laboratory |date=19 June 2015 |website=YourGenome |access-date=24 March 2025}}</ref> The studies on dipteran relationships by [[Willi Hennig]] helped in the development of [[cladistics]], techniques that he applied to morphological characters but now adapted for use with molecular sequences in phylogenetics.<ref>{{Cite journal |last=Ashlock |first=P. D. |date=1974 |title=The Uses of Cladistics |journal=Annual Review of Ecology and Systematics |volume=5 |issue=1 |pages=81–99 |doi=10.1146/annurev.es.05.110174.000501}}</ref> [[Maggot]]s found on corpses are useful to [[forensic entomology|forensic entomologists]]. Maggot species can be identified by their anatomical features and by matching their [[DNA]]. Maggots of different species of flies visit corpses and carcases at fairly well-defined times after the death of the victim, and so do their predators, such as beetles in the family [[Histeridae]]. Thus, the presence or absence of particular species provides evidence for the time since death, and sometimes other details such as the place of death, when species are confined to particular habitats such as [[woodland]].<ref>{{cite journal |last1=Joseph |first1=Isaac |last2=Mathew |first2=Deepu G. |last3=Sathyan |first3=Pradeesh |last4=Vargheese |first4=Geetha |title=The use of insects in forensic investigations: An overview on the scope of forensic entomology |journal=Journal of Forensic Dental Sciences |date=2011 |volume=3 |issue=2 |pages=89–91 |doi=10.4103/0975-1475.92154 |pmc=3296382 |pmid=22408328 |doi-access=free }}</ref> [[File:Casu Marzu cheese.jpg|thumb|right|[[Casu marzu]] is a traditional [[Sardinia]]n sheep milk cheese that contains larvae of the [[cheese fly]], ''Piophila casei''.]] Some species of maggots such as [[Calliphoridae|blowfly larvae (gentles)]] and bluebottle larvae ([[caster (maggot)|casters]]) are bred commercially; they are sold as [[fishing bait|bait]] in [[angling]], and as food for carnivorous animals (kept as pets, in zoos, or for research) such as some [[mammal]]s,<ref>{{cite journal |last1=Ogunleye |first1=R. F. |last2=Edward |first2=J. B. |title=Roasted maggots (Dipteran larvae) as a dietary protein source for laboratory animals |journal=African Journal of Applied Zoology and Environmental Biology |date=2005 |volume=7 |pages=140–143 |url=http://www.ajol.info/index.php/ajazeb/article/view/41163}}</ref> [[fish]]es, [[reptile]]s, and [[bird]]s. It has been suggested that fly larvae could be used at a large scale as food for farmed chickens, pigs, and fish. However, consumers are opposed to the inclusion of insects in their food, and the use of insects in animal feed remains illegal in areas such as the [[European Union]].<ref>{{cite web |last1=Fleming |first1=Nic |title=How insects could feed the food industry of tomorrow |url=http://www.bbc.com/future/story/20140603-are-maggots-the-future-of-food |publisher=[[British Broadcasting Corporation]] |access-date=24 May 2016 |date=4 June 2014}}</ref><ref>{{cite web |title=Why are insects not allowed in animal feed? |url=http://www.protix.eu/wp-content/uploads/2014/07/20141029_Why-are-insects-not-allowed-in-animal-feed_Whitepaper_Insect_meal.pdf |publisher=All About Feed |access-date=24 May 2016 |date=August 2014 |url-status=dead|archive-url=https://web.archive.org/web/20160811001146/http://www.protix.eu/wp-content/uploads/2014/07/20141029_Why-are-insects-not-allowed-in-animal-feed_Whitepaper_Insect_meal.pdf |archive-date=11 August 2016 |df=dmy-all }}</ref> Fly larvae can be used as a [[biomedical]] tool for wound care and treatment. [[Maggot debridement therapy]] (MDT) is the use of [[Calliphoridae|blow fly]] larvae to remove the dead tissue from wounds, most commonly being amputations. Historically, this has been used for centuries, both intentional and unintentional, on battlefields and in early hospital settings.<ref>{{Cite journal |last1=Stegman |first1=Sylvia |last2=Steenvoorde |first2=Pascal |date=2011 |title=Maggot debridement therapy |url=https://www.researchgate.net/publication/260701991 |journal=Proceedings of the Netherlands Entomological Society Meeting |volume=22 |pages=61–66 }}</ref> Removing the dead tissue promotes cell growth and healthy wound healing. The larvae also have [[biochemical]] properties such as [[antibacterial]] activity found in their secretions as they feed.<ref>{{Cite journal |last1=Diaz-Roa |first1=A. |last2=Gaona |first2=M. A. |last3=Segura |first3=N. A. |last4=Suárez |first4=D. |last5=Patarroyo |first5=M.A. |last6=Bello |first6=F. J. |date=August 2014 |title=''Sarconesiopsis magellanica'' (Diptera: Calliphoridae) excretions and secretions have potent antibacterial activity |journal=Acta Tropica |pages=37–43 |doi=10.1016/j.actatropica.2014.04.018 |pmid=24754920 |volume=136}}</ref> These medicinal maggots are a safe and effective treatment for chronic wounds.<ref>{{Cite journal |last1=Gilead |first1=L. |last2=Mumcuoglu |first2=K. Y. |last3=Ingber |first3=A. |date=16 August 2013 |title=The use of maggot debridement therapy in the treatment of chronic wounds in hospitalised and ambulatory patients |journal=Journal of Wound Care |doi=10.12968/jowc.2012.21.2.78 |pmid=22584527 |volume=21 |issue=2 |pages=78–85}}</ref> The [[Sardinia]]n cheese [[casu marzu]] is exposed to flies known as [[cheese skipper]]s such as ''[[Piophila casei]]'', members of the family [[Piophilidae]].<ref>{{cite journal |year=2007 |title=A mite unappetizing |url=http://www.entsoc.org/PDF/Pubs/Periodicals/AE/AE-2007/Fall/Buzzwords.pdf |archive-url=https://web.archive.org/web/20101216084837/http://entsoc.org/PDF/Pubs/Periodicals/AE/AE-2007/Fall/Buzzwords.pdf |url-status=dead|archive-date=2010-12-16 |journal=American Entomologist |volume=53 |issue=3 |pages=132–133 |last=Berenbaum |first=May |doi=10.1093/ae/53.3.132 }}</ref> The digestive activities of the fly larvae soften the cheese and modify the aroma as part of the process of maturation. At one time European Union authorities banned sale of the cheese and it was becoming hard to find,<ref>{{cite web |last1=Colangelo |first1=Matt |title=A Desperate Search for Casu Marzu, Sardinia's Illegal Maggot Cheese |url=http://www.foodandwine.com/fwx/desperate-search-casu-marzu-sardinias-illegal-maggot-cheese |publisher=Food and Wine |access-date=24 May 2016 |date=9 October 2015}}</ref> but the ban has been lifted on the grounds that the cheese is a traditional local product made by traditional methods.<ref>{{cite news |title=Illegal food: step away from the cheese, ma'am |last=Brones |first=Anna |url=https://www.theguardian.com/lifeandstyle/wordofmouth/2013/apr/15/illegal-food-cheese-us-mimolette |newspaper=[[The Guardian]] |date=15 April 2013 |access-date=26 May 2016}}</ref>
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