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{{Short description|Family of flies}}
{{Use dmy dates|date=August 2013}}
{{Use dmy dates|date=May 2024}}
{{automatic taxobox
{{automatic taxobox
| image = Piophila casei british entymology detail.jpg
| image = Piophila casei british entymology detail.jpg
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| name = Cheese flies
| name = Cheese flies
| taxon = Piophilidae
| taxon = Piophilidae
| authority = [[Macquart]], 1835
| authority = [[Pierre-Justin-Marie Macquart|Macquart]], 1835
| synonyms = *Neottiophilidae
| synonyms = *Neottiophilidae
*Thyreophoridae
*Thyreophoridae
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}}
}}


The '''Piophilidae''' are a [[Family (biology)|family]] of "true flies", in the [[Order (biology)|order]] [[fly|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|pages=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|accessdate=13 April 2013|date=27 September 2002|publisher=Academic Press|isbn=978-0-08-053607-1|pages=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|first=G. |last=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|pages=701–}}</ref>
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==
==Overview==
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==Behaviour patterns==
==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 |url=}}</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" /> A series of photos illustrating this remarkable behaviour and a video can be seen at [http://www.bonduriansky.net/antlerflies.htm]. 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.
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>[http://www.bonduriansky.net/antlerflies.htm Antler fly (University of New South Wales)]</ref> breeds on discarded antlers of moose and other deer. As adaptations to such a scarce and scattered resource, they have several unusual behaviour patterns. For one thing, the males are astonishingly [[Wiktionary:bellicose|bellicose]]; the [[Specific name (zoology)|specific epithet]] ''litigata'' reflects the fact &mdash; in context it means "aggressive", as in the English word "[[Wiktionary:litigious|litigious]]". On discarded antlers, the males form complex, highly structured [[Aggregation (ethology)|aggregations]] in which a great deal of [[Territory (animal)|territorial competition]] occurs. Some individuals defend stable territories, while others simply wander in search of females that arrive on antlers to feed, mate, and [[Oviparity|oviposit]]. In prime areas of the antler, near oviposition sites (cracks in the antler surface), males spend much of their time battling rival males. They even attack insects much larger than themselves.<ref name="Bonduriansky2">{{cite journal |last=Bonduriansky |first=R. |last2=Brooks |first2=R.J. |title=Why do male antler flies (''Protopiophila litigata'') fight? |journal=Ethology Ecology & Evolution |volume=11 |issue=3 |pages=287–301 |year=1999 |doi=10.1080/08927014.1999.9522829 }}</ref> Another adaptation to life on discarded antlers is an astonishing degree of site fidelity: 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}}</ref>
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 | url=http://www.bonduriansky.net/waltzingflies.htm | title=Waltzing flies}}</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>

Waltzing flies breed in early spring, generally on the carcasses of moose. By midspring of a good year, they are one of the most abundant insect species in the local forests. Nubile females bask on vegetation around carcasses, and males aggregate in their sun to court and to defend territories. Competing males usually begin by spreading their forelegs and holding each other's fore[[Insect morphology|tarsi]]. This recalls the competitive behaviour and adaptations of another fly family, the [[Diopsidae]], in which the males match the spread of each other's eyes. Such behaviour may permit the adversaries to assess each other's size, thereby avoiding futile and possibly damaging conflict with a larger opponent. When neither of two competing males of ''P. xanthostoma'' is intimidated, they fight, each striking the other with his head and antennae. Such fights may last for several minutes, a long conflict for such small, active, and delicate animals.

A male courts a female by dancing side-to-side, forequarters held high, displaying his elongated antennae and vibrating his elongated forelegs. A receptive female responds by spreading her forelegs to contact the male's long forelegs. The male somersaults over her forequarters and flips over to land on her back and lock onto her genitals. After several minutes of copulation, the female ejects a large part of the semen and swallows it. Such behaviour seems to [[Correlation and dependence|correlate]] with the success of fertile egg production. Given such elaborate behavioural and morphological adaptations, waltzing flies are of interest as a model system for studying of the evolution of sexual dimorphism.<ref name="Bonduriansky3">{{cite journal |last=Bonduriansky |first=Russell |last2=Wheeler |first2=Jill |first3=Locke |last3=Rowe |title=Ejaculate feeding and female fitness in the sexually dimorphic fly ''Prochyliza xanthostoma'' (Diptera: Piophilidae) |journal=Animal Behaviour |volume=69 |issue= 2|pages=489–497 |year=2005 |doi=10.1016/j.anbehav.2004.03.018 |url=}}</ref><ref name="Bonduriansky4">{{cite journal |last=Bonduriansky |first=R. |last2=Rowe |first2=L. |title=Interactions among mechanisms of sexual selection on male body size and head shape in a sexually dimorphic fly |journal=Evolution |volume=57 |issue=9 |pages=2046–53 |year=2003 |doi= 10.1554/03-047}}</ref>

Females oviposit on carrion, and larvae develop largely in sheltered parts of the carcass; depending on the degree of decomposition, they often develop inside the bone marrow. As in other piophilid species, final-instar larvae leap off the surface of the carcass where they had fed, and pupate in the soil.<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==
==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|pages=853–}}</ref> Some species also have been known to cause naso-oral and urogenital myiasis.<ref name="Gennard2012"/>
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 | accessdate=14 November 2005 |archiveurl = https://web.archive.org/web/20060316000328/http://dsc.discovery.com/fansites/onthecase/toolbox/tool_05.html <!-- Bot retrieved archive --> |archivedate = 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"/>


⚫
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>
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[[File:WillistonPiophilidae.jpg|thumb|230px|left|Morphological details of the Piopilidae]]


<!-- poorly written ==Description==
==Description==
<!-- poorly written
{{tone|section|date=December 2014}}
{{tone|section|date=December 2014}}
For terms see [[Morphology of Diptera]].
For terms see [[Morphology of Diptera]].
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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.
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. -->
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]]
⚫
The family differs from the closely related 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.
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 ==
== Classification ==


The Piophilidae are a small family of less than 100 [[Species description|described]] species in 23 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 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:
Recent works containing keys for identification of the Piophilidae include:
* {{cite journal | last1 = McAlpine | first1 = JF | year = 1977 | title = A revised classification of the Piophilidae, including 'Neottiophilidae' and 'Thyreophoridae' (Diptera: Schizophora) | url = | journal = Memoirs of the Entomological Society of Canada | volume = 103 | issue = | pages = 1–66 }}
* {{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 }}
* McAlpine JF. (ed.) 1989. ''Manual of Nearctic Diptera'' (Vol. 3). Agriculture Canada Monograph No. 32.
* McAlpine JF. (ed.) 1989. ''Manual of Nearctic Diptera'' (Vol. 2). Agriculture Canada Monograph No. 32.
* Ozerov, 2000. ''Piophilidae''. In: Papp, L. & Darvas, A. (eds). ''Contributions to a Manual of Palaearctic Diptera''. Appendix Volume. Science Herald, Budapest.pp 355–365. {{ISBN|963-04-8840-X}}
* Ozerov, 2000. ''Piophilidae''. In: Papp, L. & Darvas, A. (eds). ''Contributions to a Manual of Palaearctic Diptera''. Appendix Volume. Science Herald, Budapest.pp 355–365. {{ISBN|963-04-8840-X}}
*[[Aleksandr Aleksandrovich Stackelberg|Shtakel'berg, A.A.]] Family Piophilidae in Bei-Bienko, G. Ya, 1988 ''Keys to the insects of the European Part of the USSR'' Volume 5 (Diptera) Part 2 English edition
*[[Aleksandr Stackelberg|Stackelberg, A.A.]] Family Piophilidae in Bei-Bienko, G. Ya, 1988 ''Keys to the insects of the European Part of the USSR'' Volume 5 (Diptera) Part 2 English edition
*{{cite journal | last1 = Rochefort | first1 = S. |display-authors=et al | year = 2015 | title = Key to forensically important Piophilidae (Diptera) in the Nearctic Region | url = | journal = Canadian Journal of Arthropod Identification | volume = 27 | issue = | pages = 1–37 }}
*{{cite journal | last1 = Rochefort | first1 = S. |display-authors=et al | 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) ''Diptères: Brachycères''. II. ''Muscidae acalypterae, Scatophagidae''. Paris: Éditions Faune de France 28 [http://faunedefrance.org/bibliotheque/docs/E.SEGUY(FdeFr28)Dipt.Brachyceres.pdf Bibliotheque Virtuelle Numerique pdf]
A still useful older work is Séguy, E. (1934) ''Diptères: Brachycères''. II. ''Muscidae acalypterae, Scatophagidae''. Paris: Éditions Faune de France 28 [http://faunedefrance.org/bibliotheque/docs/E.SEGUY(FdeFr28)Dipt.Brachyceres.pdf Bibliotheque Virtuelle Numerique pdf]
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== Genera ==
== Genera ==
* [http://www.faunaeur.org/full_results.php?id=10941 The Western Palearctic subfamilies, tribes and genera of Piophilidae - ''Fauna Europaea'']
* [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 ===
=== Species lists ===
* [http://www.faunaeur.org/full_results.php?id=10941 Palaearctic]
* [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://www.nearctica.com/nomina/main.htm Nearctic]
* [http://hbs.bishopmuseum.org/aocat/piophilidae.html Australasian/Oceanian]
* [http://hbs.bishopmuseum.org/aocat/piophilidae.html Australasian/Oceanian]
* [http://konchudb.agr.agr.kyushu-u.ac.jp/mokuroku/ Japan]
* [https://web.archive.org/web/20190527005729/http://konchudb.agr.agr.kyushu-u.ac.jp/mokuroku/ Japan]
* [http://eol.org/pages/80292/overview World list]
* [http://eol.org/pages/80292/overview World list]


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* {{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 }}
* {{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.
* 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 |chapterurl=https://books.google.com/books&id=aluUgDVYJ8wC&pg=PA180 |chapter=Diptera, Ephemeroptera: Piophilidae: Cheese skipper (''Piophila casei'')}}
* {{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 ==
== External links ==
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{{Taxonbar|from=Q135117}}
{{Taxonbar|from=Q135117}}


{{DEFAULTSORT:Cheese Fly}}
⚫
[[Category:Brachycera families]]
[[Category:Piophilidae| ]]
[[Category:Piophilidae| ]]
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[[Category:Brachycera families]]
[[Category:Taxa named by Pierre-Justin-Marie Macquart]]

Latest revision as of 02:36, 9 May 2024

Cheese flies
Piophila casei
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Diptera
Superfamily: Tephritoidea
Family: Piophilidae
Macquart, 1835
Subfamilies
Synonyms
  • Neottiophilidae
  • Thyreophoridae

The Piophilidae are a family of "true flies", in the 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[1] and medical entomology.[2] 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.[3]

Overview[edit]

The most notorious member of the family is the cheese fly, Piophila casei; it is cosmopolitan, and a typical member of the family. It is a small species, about 4 mm (0.16 in) long. The fly's larvae infest cured meats, smoked or salted fish, cheeses, and carrion. The mature larva is about 8 mm (0.31 in) long and is sometimes called the cheese skipper because of its leaping ability - when disturbed, this tiny maggot can hop some 15 cm (6 in) into the air.[4] Adults are also known as bacon flies and their larvae as bacon skippers, ham skippers, cheese maggots, cheese hoppers, etc. In the Mediterranean island of Sardinia, the larvae are intentionally introduced into pecorino cheese to produce the characteristic casu marzu ("rotten cheese" in 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 iridescent and lie flat upon the fly's abdomen when at rest. At 4 mm (0.16 in) long, the fly is one-third to one-half as long as the common housefly.

Behaviour patterns[edit]

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.[5][6] 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.[4] 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,[7] breeds on discarded antlers of moose and other deer. On discarded antlers, the males form complex, highly structured aggregations in which a great deal of 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.[8]

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 behavioural adaptation and associated morphological adaptations. In particular, the antennae, forelegs, and heads of the males are adapted in unusual ways to their behaviour in combat and courtship.[9] A male courts a female by dancing side-to-side, forequarters held high, displaying his elongated antennae and vibrating his elongated forelegs.[10]

Medical and forensic significance[edit]

If swallowed (whether accidentally or otherwise), the larvae sometimes survive in the intestines 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 stool.[11] Some species also have been known to cause naso-oral and urogenital myiasis.[1]

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.[12] 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.[1][13]

Description[edit]

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.[14] Like all tephritoid flies, female piophilids have an extensible 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[edit]

The Piophilidae are a small family of less than 100 described species in 21 genera, mainly Holarctic in distribution, though some species are 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:[15]

Recent works containing keys for identification of the Piophilidae include:

  • McAlpine, JF (1977). "A revised classification of the Piophilidae, including 'Neottiophilidae' and 'Thyreophoridae' (Diptera: Schizophora)". Memoirs of the Entomological Society of Canada. 103: 1–66. doi:10.4039/entm109103fv.
  • McAlpine JF. (ed.) 1989. Manual of Nearctic Diptera (Vol. 2). Agriculture Canada Monograph No. 32.
  • Ozerov, 2000. Piophilidae. In: Papp, L. & Darvas, A. (eds). Contributions to a Manual of Palaearctic Diptera. Appendix Volume. Science Herald, Budapest.pp 355–365. ISBN 963-04-8840-X
  • Stackelberg, A.A. Family Piophilidae in Bei-Bienko, G. Ya, 1988 Keys to the insects of the European Part of the USSR Volume 5 (Diptera) Part 2 English edition
  • Rochefort, S.; et al. (2015). "Key to forensically important Piophilidae (Diptera) in the Nearctic Region". Canadian Journal of Arthropod Identification. 27: 1–37.

A still useful older work is Séguy, E. (1934) Diptères: Brachycères. II. Muscidae acalypterae, Scatophagidae. Paris: Éditions Faune de France 28 Bibliotheque Virtuelle Numerique pdf

See also[edit]

Genera[edit]

Species lists[edit]

Image galleries[edit]

References[edit]

  1. ^ a b c Gennard, Dorothy (2012). Forensic Entomology: An Introduction. Wiley. p. 67. ISBN 978-1-119-94543-7.
  2. ^ Gary R. Mullen; Lance A. Durden (27 September 2002). Medical and Veterinary Entomology. Academic Press. p. 139. ISBN 978-0-08-053607-1. Retrieved 13 April 2013.
  3. ^ Gordh, G.; Headrick, David H. (2001). A dictionary of entomology. CABI. p. 701. ISBN 978-0-85199-291-4.
  4. ^ a b Don C. Mote (1914). "The cheese skipper (Piophila casei Linne)" (PDF). The Ohio Naturalist. 14 (7): 309–315.
  5. ^ Maitland, David P. (1992). "Locomotion by jumping in the Mediterranean fruit-fly larva Ceratitis capitata". Nature. 355 (6356): 159–161. doi:10.1038/355159a0. S2CID 4365010.
  6. ^ Bonduriansky, Russell (October 2002). "Leaping behaviour and responses to moisture and sound in larvae of piophilid carrion flies". The Canadian Entomologist. 134 (5): 647–656. doi:10.4039/Ent134647-5. ISSN 0008-347X. S2CID 86676206.
  7. ^ Bonduriansky, Russell. "Antler flies". bonduriansky.net. Retrieved 14 May 2022.
  8. ^ Bonduriansky, R.; Brassil, C.E. (2002). "Rapid and costly ageing in wild male flies". Nature. 420 (6914): 377. doi:10.1038/420377a. PMID 12459773.
  9. ^ Bonduriansky, Russell. "Waltzing flies". bonduriansky.net. Retrieved 14 May 2022.
  10. ^ Bonduriansky, R. (2003). "Layered sexual selection: a comparative analysis of sexual behaviour within an assemblage of piophilid flies". Canadian Journal of Zoology. 81 (3): 479–491. doi:10.1139/z03-031.
  11. ^ David Alan Warrell; Timothy M. Cox; John D. Firth (2003). Oxford Textbook of Medicine: Sections 1-10. Oxford University Press. p. 853. ISBN 978-0-19-857014-1.
  12. ^ "Discovery Channel: You're on the Case". Archived from the original on 16 March 2006. Retrieved 14 November 2005.
  13. ^ Martín-Vega, Daniel (10 October 2011). "Skipping clues: Forensic importance of the family Piophilidae (Diptera)". Forensic Science International. 212 (1–3): 1–5. doi:10.1016/j.forsciint.2011.06.016. PMID 21802229.
  14. ^ McAlpine, JF (1977). "A revised classification of the Piophilidae, including 'Neottiophilidae' and 'Thyreophoridae' (Diptera: Schizophora)". Memoirs of the Entomological Society of Canada. 103: 1–66. doi:10.4039/entm109103fv.
  15. ^ Ozerov, A.L. (2004). "On the classification of the family Piophilidae (Diptera)". Entomological Review. 84 (5): 600–608.

External links[edit]

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