Editing Piophilidae

{{Short description|Family of flies}}
{{Use dmy dates|date=May 2024}}
{{automatic taxobox
 | image               = Piophila casei british entymology detail.jpg
 | image_caption       = ''[[Piophila casei]]''
 | name                = Cheese flies
 | taxon = Piophilidae
 | authority = [[Pierre-Justin-Marie Macquart|Macquart]], 1835
 | synonyms            = *Neottiophilidae
*Thyreophoridae
 | subdivision_ranks   = Subfamilies
 | subdivision         = *[[Neottiophilinae]]
*[[Piophilinae]]
*[[Thyreophorinae]]
}}

The '''Piophilidae''' are a [[Family (biology)|family]] of "true flies", in the [[Order (biology)|order]] [[Diptera]]. The so-called '''cheese flies''' are the best-known members, but most species of the Piophilidae are scavengers in animal products, [[carrion]], and fungi. They may accordingly be important in [[forensic entomology]]<ref name="Gennard2012">{{cite book |first=Dorothy |last=Gennard |title=Forensic Entomology: An Introduction|url=https://books.google.com/books?id=zeWwA0h2zIAC&pg=PA67 |date=2012 |publisher=Wiley |isbn=978-1-119-94543-7|page=67}}</ref> and [[medical entomology]].<ref name="MullenDurden2002">{{cite book|author1=Gary R. Mullen|author2=Lance A. Durden|title=Medical and Veterinary Entomology|url=https://books.google.com/books?id=6R1v9o-uaI4C&pg=PA139|access-date=13 April 2013|date=27 September 2002|publisher=Academic Press|isbn=978-0-08-053607-1|page=139}}</ref> For a fly maggot, the larvae of many species have an unusually well-developed ability to leap when alarmed or when abandoning their larval food to pupate; they accordingly may be known as '''cheese skippers''' or other kinds of skippers according to their food source.<ref name="GordhHeadrick2001">{{cite book|first1=G. |last1=Gordh|first2=David H. |last2=Headrick|title=A dictionary of entomology|url=https://books.google.com/books?id=d0XSwMJLDg4C&pg=PA701 |year=2001|publisher=CABI|isbn=978-0-85199-291-4|page=701}}</ref>

==Overview==
The most notorious member of the family is the [[cheese fly]], ''Piophila casei''; it is [[cosmopolitan distribution|cosmopolitan]], and a typical member of the family. It is a small species, about {{convert|4|mm|in|abbr=on}} long. The fly's [[larva]]e infest cured meats, smoked or salted fish, cheeses, and [[carrion]]. The mature larva is about {{convert|8|mm|in|abbr=on}} long and is sometimes called the cheese skipper because of its leaping ability - when disturbed, this tiny maggot can hop some 15&nbsp;cm (6&nbsp;in) into the air.<ref name="ohio">{{cite journal| author = Don C. Mote | year = 1914 | title = The cheese skipper (''Piophila casei'' Linne) | journal = The Ohio Naturalist | volume = 14 | issue = 7 | pages = 309–315 | url = https://kb.osu.edu/dspace/bitstream/1811/1821/1/V14N07_309.pdf }}</ref> Adults are also known as bacon flies and their larvae as bacon skippers, ham skippers, cheese maggots, cheese hoppers, etc. In the [[Mediterranean Sea|Mediterranean]] island of [[Sardinia]], the larvae are intentionally introduced into [[pecorino]] [[cheese]] to produce the characteristic ''[[casu marzu]]'' ("rotten cheese" in [[Sardinian language|Sardinian]]).

The adult cheese fly's body is black, blue-black, or bronze, with some yellow on the head, antennae, and legs. The wings are faintly [[Iridescence|iridescent]] and lie flat upon the fly's [[abdomen]] when at rest. At {{convert|4|mm|in|abbr=on}} long, the fly is one-third to one-half as long as the common [[housefly]].

==Behaviour patterns==
Like the larvae of various fly families, including the family [[Tephritidae]], the larvae of typical piophilids are notorious for jumping or "skipping", especially in their final [[instar]].<ref>{{cite journal |first=David P. |last=Maitland |title=Locomotion by jumping in the Mediterranean fruit-fly larva ''Ceratitis capitata'' |journal=Nature |volume=355 |issue= 6356|pages=159–161 |year=1992 |doi=10.1038/355159a0 |s2cid=4365010 }}</ref><ref>{{Cite journal |last=Bonduriansky |first=Russell |date=October 2002 |title=Leaping behaviour and responses to moisture and sound in larvae of piophilid carrion flies |url=https://www.cambridge.org/core/product/identifier/S0008347X00005629/type/journal_article |journal=The Canadian Entomologist|volume=134 |issue=5 |pages=647–656 |doi=10.4039/Ent134647-5 |s2cid=86676206 |issn=0008-347X}}</ref> The larvae accomplish their jumps by bending over, grabbing onto the rears of their own bodies with their mouth hooks, and tensing their muscles in a manner that increases the pressure on their own blood and internal organs. When they release their grip, the internal pressure straightens out the tubular body, propelling the forequarters upwards, the rest of the body following.<ref name="ohio" /> Jumping is performed most typically when the larva is alarmed by a disturbance, or when it is abandoning its feeding site in preparation for pupation.

The tiny piophilid species ''[[Protopiophila litigata]]'', commonly known as the antler fly,<ref>{{Cite web |last=Bonduriansky |first=Russell |title=Antler flies |url=http://bonduriansky.net/antlerflies.htm |access-date=2022-05-14 |website=bonduriansky.net}}</ref> breeds on discarded antlers of moose and other deer. On discarded antlers, the males form complex, highly structured [[Aggregation (ethology)|aggregations]] in which a great deal of [[Territory (animal)|territorial competition]] occurs. In prime areas of the antler, near oviposition sites (cracks in the antler surface), males spend much of their time battling rival males. Males spend their entire lives competing on the same antler (only leaving to spend the night in nearby vegetation), making it possible to mark flies individually and obtain longitudinal field data on these tiny insects. This unique ecology made it possible to document senescence in wild insects for the first time.<ref>{{cite journal|last1=Bonduriansky|first1=R.|last2=Brassil|first2=C.E.|title=Rapid and costly ageing in wild male flies|journal=Nature|date=2002|volume=420|issue=6914|page=377|doi=10.1038/420377a|pmid=12459773|doi-access=free}}</ref>

The waltzing fly, ''[[Prochyliza xanthostoma]]'', occurs in North America. It is one of the carrion-feeding piophilids and is remarkable for its [[sexual dimorphism]] and its patterns of [[Adaptation|behavioural adaptation]] and associated [[Morphology (biology)|morphological adaptations]]. In particular, the antennae, forelegs, and heads of the males are adapted in unusual ways to their behaviour in combat and courtship.<ref>{{Cite web |last=Bonduriansky |first=Russell |title=Waltzing flies |url=http://www.bonduriansky.net/waltzingflies.htm |access-date=2022-05-14 |website=bonduriansky.net}}</ref> A male courts a female by dancing side-to-side, forequarters held high, displaying his elongated antennae and vibrating his elongated forelegs.<ref name="Bonduriansky5">{{cite journal |first=R. |last=Bonduriansky |title=Layered sexual selection: a comparative analysis of sexual behaviour within an assemblage of piophilid flies |journal=Canadian Journal of Zoology |volume=81 |issue=3 |pages=479–491 |year=2003 |doi=10.1139/z03-031 }}</ref>

==Medical and forensic significance==
If swallowed (whether accidentally or otherwise), the larvae sometimes survive in the [[intestine]]s and pass through the digestive system alive. Such behaviour is known as [[enteric]] or [[intestinal]] [[myiasis]]. In the gut, the larvae may cause serious lesions by attempting to bore through the intestinal walls. Symptoms include [[nausea]], [[vomiting]], pain in the abdomen, and bloody [[diarrhea]]. Both living and dead larvae may pass in the [[feces|stool]].<ref name="WarrellCox2003">{{cite book|author1=David Alan Warrell|author2=Timothy M. Cox|author3=John D. Firth|title=Oxford Textbook of Medicine: Sections 1-10|url=https://books.google.com/books?id=EhjX517cGVsC&pg=PA853 |year=2003|publisher=Oxford University Press|isbn=978-0-19-857014-1|page=853}}</ref> Some species also have been known to cause naso-oral and urogenital myiasis.<ref name="Gennard2012"/>

In [[forensic entomology]], the presence of ''P. casei'' larvae may be useful in estimating the date of death for human remains because they do not take up residence in a corpse until three to six months after death.<ref>{{cite web | title=Discovery Channel: You're on the Case | url=http://dsc.discovery.com/fansites/onthecase/toolbox/tool_05.html | access-date=14 November 2005 |archive-url = https://web.archive.org/web/20060316000328/http://dsc.discovery.com/fansites/onthecase/toolbox/tool_05.html <!-- Bot retrieved archive --> |archive-date = 16 March 2006}}</ref> However, ''P. casei'' is not the only piophilid species to attack human corpses, so caution is appropriate in identification of the species found and in interpretation of their significance.<ref name="Gennard2012"/><ref>{{Cite journal |last=Martín-Vega |first=Daniel |date=2011-10-10 |title=Skipping clues: Forensic importance of the family Piophilidae (Diptera) |url=https://linkinghub.elsevier.com/retrieve/pii/S0379073811002970 |journal=Forensic Science International|volume=212 |issue=1–3 |pages=1–5 |doi=10.1016/j.forsciint.2011.06.016|pmid=21802229 }}</ref>

==Description==
<!-- poorly written
{{tone|section|date=December 2014}}
For terms see [[Morphology of Diptera]].

Small and mostly deeply pigmented flies. The head is round.The face is not carinate. There are always two pairs of vertical bristles.The postvertical bristles on the head are divergent. Vibrissae on head are present.The fronto-orbital bristles vary from two pairs to none.The antennae are short and decumbent, the third segment elongate oval (face strongly receding and the antennae variable in length in ''Amphipogon'').The palpi are well developed.The mesonotum is almost always finely pubescent and polished.The sternopleura are never pruinose. There is one or two pairs of dorsocentral bristles and four scutellar bristles.The legs of the male never toothed or "deformed" and the front femora usually have long but delicate bristles.The costa is interrupted near the subcosta, close to radial vein 1. Anal vein of wing arcuate (continues parallel to postalar margin). The discal cell is usually large, with the posterior crossvein usually long.
The abdomen is more or less polished and pubescent but lacks bristles and is broad, depressed, not constricted at the base genitalia of the male more or less hidden and asymmetrical. The female ovipositor is extensile. -->
[[File:WillistonPiophilidae.jpg|thumb|230px|right|Morphological details of the Piophilidae]]
Piophilidae are small flies, often dark in color and shiny. The wings are usually clear and unmarked, with the exception of the genera ''Mycetaulus'', ''Neottiophilum'', ''Pseudoseps'', and ''Thyreophora'', which have brown wing markings.<ref name="Mcalpine-piophilidae">{{cite journal | last1 = McAlpine | first1 = JF | year = 1977 | title = A revised classification of the Piophilidae, including 'Neottiophilidae' and 'Thyreophoridae' (Diptera: Schizophora) | journal = Memoirs of the Entomological Society of Canada | volume = 103 | pages = 1–66 | doi = 10.4039/entm109103fv }}</ref> Like all [[Tephritoidea|tephritoid]] flies, female piophilids have an extensible [[Ovipositor#In insects|ovipositor]].

The family differs from the similar looking family Sepsidae in several characters, particularly in having the costa broken at the end of the subcosta, the setulose mesonotum and the absence of a hair or fine bristle arising on the posterior edge of the posterior spiracle of the thorax.

== Classification ==

The Piophilidae are a small family of less than 100 [[Species description|described]] species in 21 genera, mainly [[Holarctic]] in distribution, though some species are [[Cosmopolitan distribution|cosmopolitan]]. The nomenclature is volatile, with two subfamily names (Neottiophilinae and Thyreophorinae) in use recently, having been subsumed in the subfamily Piophilinae.

The genera of Piophilidae are:<ref>{{Cite journal |last=Ozerov |first=A. L. |date=2004 |title=On the classification of the family Piophilidae (Diptera) |journal=Entomological Review |volume=84 |issue=5 |pages=600–608}}</ref>
{{Div col|colwidth=22em}}
* ''[[Actenoptera]]''
* ''[[Allopiophila]]''
* ''[[Amphipogon (fly)|Amphipogon]]''
* ''[[Bocainamyia]]''
* ''[[Centrophlebomyia]]''
* ''[[Dasyphlebomyia]]''
* ''[[Diacanthomyia]]''
* ''[[Lasiopiophila]]''
* ''[[Mycetaulus]]''
* ''[[Neopiophila]]''
* ''[[Neottiophilum]]''
* ''[[Piophila]]''
* ''[[Piophilosoma]]''
* ''[[Prochyliza]]''
* ''[[Protopiophila]]''
* ''[[Protothyreophora]]''
* ''[[Pseudoseps]]''
* ''[[Pygopiophila]]''
* ''[[Stearibia]]''
* ''[[Thyreolepida]]''
* ''[[Thyreophora cynophila|Thyreophora]]''
{{Div col end}}

Recent works containing keys for identification of the Piophilidae include:
* {{cite journal | last1 = McAlpine | first1 = J. F. | year = 1977 | title = A revised classification of the Piophilidae, including 'Neottiophilidae' and 'Thyreophoridae' (Diptera: Schizophora) | journal = Memoirs of the Entomological Society of Canada | volume = 103 | pages = 1–66 | doi = 10.4039/entm109103fv }}
* {{cite book |editor-last=McAlpine |editor-first=J. F. |year=1989 |title=Manual of Nearctic Diptera |volume=2 |series=Agriculture Canada Monograph No. 32}}
* {{cite book |last=Ozerov |first=A. L. |year=2000 |chapter=Piophilidae |editor1-last=Papp |editor-first=L. |editor2-last=Darvas |editor2-first=A. |title=Contributions to a Manual of Palaearctic Diptera |volume=Appendix |publisher=Science Herald |location=Budapest |pages=355–365 |isbn=963-04-8840-X}}
* {{cite book |author-link=Aleksandr Stackelberg |last=Stackelberg |first=A. A. |chapter=Family Piophilidae |editor-last=Bei-Bienko |editor-first=G. Ya. |year=1988 |title=Keys to the Insects of the European Part of the USSR |volume=5 (Diptera) |at=Part 2 |edition=English}}
*{{cite journal | last1 = Rochefort | first1 = S. |display-authors=etal | year = 2015 | title = Key to forensically important Piophilidae (Diptera) in the Nearctic Region | journal = Canadian Journal of Arthropod Identification | volume = 27 | pages = 1–37 }}

A still useful older work is Séguy, E. (1934) [http://faunedefrance.org/bibliotheque/docs/E.SEGUY(FdeFr28)Dipt.Brachyceres.pdf {{lang|fr|cat=no|Diptères: Brachycères}}]. II. ''Muscidae acalypterae, Scatophagidae''. {{lang|fr|italic=no|Éditions Faune de France}} 28. Paris: Paul Lechevalier et fils – via {{lang|fr|italic=no|Bibliotheque Virtuelle Numerique}}.

== See also ==
* ''[[Thyreophora cynophila]]''
* [[Cheese mite]]
* [[Casu marzu]]

== Genera ==
* [https://web.archive.org/web/20051031235458/http://www.faunaeur.org/full_results.php?id=10941 The Western Palearctic subfamilies, tribes and genera of Piophilidae - ''Fauna Europaea'']

=== Species lists ===
* [https://web.archive.org/web/20051031235458/http://www.faunaeur.org/full_results.php?id=10941 Palaearctic]
* [http://www.nearctica.com/nomina/main.htm Nearctic]
* [http://hbs.bishopmuseum.org/aocat/piophilidae.html Australasian/Oceanian]
* [https://web.archive.org/web/20190527005729/http://konchudb.agr.agr.kyushu-u.ac.jp/mokuroku/ Japan]
* [http://eol.org/pages/80292/overview World list]

== Image galleries ==
* {{cite web |url=http://www.bonduriansky.net/antlerflies.htm |title=Antler Flies (''Protopiophila litigata'') |work=Evolutionary Biology Lab |publisher=University of New South Wales }}
* {{cite web |url=http://www.bonduriansky.net/waltzingflies.htm |title=Waltzing Flies (''Prochyliza xanthostoma'') |work=Evolutionary Biology Lab |publisher=University of New South Wales }}
* [http://www.diptera.info/photogallery.php?album_id=37 Images at Diptera.info]

== References ==
{{Reflist}}
* {{cite book | author=Berenbaum, May R. | title=Ninety-Nine More Maggots, Mites, and Munchers | publisher=University of Illinois Press | year=1993 | isbn=978-0-252-06322-0 | url-access=registration | url=https://archive.org/details/ninetyninemorema00bere }}
* Lieutenant Brian F. Prendergast, USN (2001). [https://web.archive.org/web/20060214194219/http://nepmu6.med.navy.mil/Entomology/filth%20flies.pdf Filth Flies: Significance, Surveillance and Control in Contingency Operations (.pdf format)]. Retrieved 1 October 2005.
* {{cite book | author=Robinson, W H | title=Handbook of Urban Insects And Arachnids: A Handbook of Urban Entomology | publisher=Cambridge University Press | year=2005 | isbn=978-0-521-81253-5 |pages=180–1 |chapter-url=https://books.google.com/books?id=aluUgDVYJ8wC&pg=PA180 |chapter=Diptera, Ephemeroptera: Piophilidae: Cheese skipper (''Piophila casei'')}}

== External links ==
*{{Wikispecies-inline|Piophilidae}}
* [http://delta-intkey.com/britin/dip/www/piophili.htm Family description and images]
* [http://www.bonduriansky.net/resources.htm An extensive bibliography on the Piophilidae]
*[http://entnemdept.ifas.ufl.edu/creatures/urban/flies/cheese_skipper.htm Piophila casei] on the [[University of Florida|UF]] / [[Institute of Food and Agricultural Sciences|IFAS]] Featured Creatures Web site (note that many of the images on this site show other species misidentified as Piophila casei)
*{{cite web |title=''Piophilidae'': Skipper Flies |work=Atlas of Living Australia |url=https://bie.ala.org.au/species/urn:lsid:biodiversity.org.au:afd.taxon:3304e8db-3fb7-4cd5-9750-1e71089ff09e}}

{{Diptera|2}}
{{Taxonbar|from=Q135117}}

[[Category:Piophilidae| ]]
[[Category:Brachycera families]]
[[Category:Taxa named by Pierre-Justin-Marie Macquart]]