Neottiophilidae
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The Piophilidae are a
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
of "true flies", in the
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
Diptera Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...
. The so-called cheese flies are the best-known members, but most species of the Piophilidae are scavengers in animal products,
carrion Carrion () is the decaying flesh of dead animals, including human flesh. Overview Carrion is an important food source for large carnivores and omnivores in most ecosystems. Examples of carrion-eaters (or scavengers) include crows, vultures, c ...
, and fungi. They may accordingly be important in
forensic entomology Forensic entomology is the scientific study of the colonization of a dead body by arthropods. This includes the study of insect types commonly associated with cadavers, their respective life cycles, their ecological presences in a given environme ...
and
medical entomology The discipline of medical entomology, or public health entomology, and also ''veterinary entomology'' is focused upon insects and arthropods that impact human health. Veterinary entomology is included in this category, because many animal disease ...
. 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.


Overview

The most notorious member of the family is the
cheese fly The cheese fly (''Piophila casei'') is a species of fly whose larvae are known for infesting human foodstuffs including cured meats, smoked or salted fish, cheeses and also carrion. The larvae of this fly are known as cheese skippers, bacon sk ...
, ''Piophila casei''; it is
cosmopolitan Cosmopolitan may refer to: Food and drink * Cosmopolitan (cocktail), also known as a "Cosmo" History * Rootless cosmopolitan, a Soviet derogatory epithet during Joseph Stalin's anti-Semitic campaign of 1949–1953 Hotels and resorts * Cosmopoli ...
, and a typical member of the family. It is a small species, about long. The fly's
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
e infest cured meats, smoked or salted fish, cheeses, and
carrion Carrion () is the decaying flesh of dead animals, including human flesh. Overview Carrion is an important food source for large carnivores and omnivores in most ecosystems. Examples of carrion-eaters (or scavengers) include crows, vultures, c ...
. The mature larva is about 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. Adults are also known as bacon flies and their larvae as bacon skippers, ham skippers, cheese maggots, cheese hoppers, etc. In the
Mediterranean The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on the e ...
island of
Sardinia Sardinia ( ; it, Sardegna, label=Italian, Corsican and Tabarchino ; sc, Sardigna , sdc, Sardhigna; french: Sardaigne; sdn, Saldigna; ca, Sardenya, label=Algherese and Catalan) is the second-largest island in the Mediterranean Sea, after ...
, the larvae are intentionally introduced into
pecorino Pecorino cheeses are hard Italian cheeses made from sheep's milk. The name "" derives from ''pecora'' which means sheep in Italian. Overview Of the six main varieties of pecorino, all of which have Protected Designation of Origin (PDO) status u ...
cheese Cheese is a dairy product produced in wide ranges of flavors, textures, and forms by coagulation of the milk protein casein. It comprises proteins and fat from milk, usually the milk of cows, buffalo, goats, or sheep. During production, ...
to produce the characteristic ''
casu marzu (; literally 'rotten/putrid cheese'), sometimes spelled , and also called , and in Sardinian, is a traditional Sardinian sheep milk cheese that contains live insect larvae (maggots). A variation of this cheese exists also in Corsica (Franc ...
'' ("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 Iridescence (also known as goniochromism) is the phenomenon of certain surfaces that appear to gradually change color as the angle of view or the angle of illumination changes. Examples of iridescence include soap bubbles, feathers, butterfl ...
and lie flat upon the fly's
abdomen The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the torso. ...
when at rest. At long, the fly is one-third to one-half as long as the common
housefly The housefly (''Musca domestica'') is a fly of the suborder Cyclorrhapha. It is believed to have evolved in the Cenozoic Era, possibly in the Middle East, and has spread all over the world as a commensal of humans. It is the most common fl ...
.


Behaviour patterns

Like the larvae of various fly families, including the family
Tephritidae The Tephritidae are one of two fly families referred to as fruit flies, the other family being the Drosophilidae. The family Tephritidae does not include the biological model organisms of the genus ''Drosophila'' (in the family Drosophilidae), w ...
, the larvae of typical piophilids are notorious for jumping or "skipping", especially in their final
instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ass ...
. 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. 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 ''Protopiophila'' is a genus of cheese skippers (insects in the family Piophilidae). There are eleven described species in ''Protopiophila''. Species These species belong to the genus ''Protopiophila'': * '' Protopiophila aethiopica'' (Hennig, 19 ...
'', commonly known as the antler fly, 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. The waltzing fly, ''
Prochyliza xanthostoma The waltzing fly ''(Prochyliza xanthostoma)'' is a species of "carrion-feeding" cheese skippers, insects in the family Piophilidae and the order Diptera. ''P. xanthostoma'' is a member of the genus Prochyliza, which contains eleven species. The ...
'', occurs in North America. It is one of the carrion-feeding piophilids and is remarkable for its
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
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. A male courts a female by dancing side-to-side, forequarters held high, displaying his elongated antennae and vibrating his elongated forelegs.


Medical and forensic significance

If swallowed (whether accidentally or otherwise), the larvae sometimes survive in the
intestine The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans ...
s and pass through the digestive system alive. Such behaviour is known as
enteric The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans and ...
or
intestinal The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans ...
myiasis Myiasis is the parasitic infestation of the body of a live animal by fly larvae (maggots) which grow inside the host while feeding on its tissue. Although flies are most commonly attracted to open wounds and urine- or feces-soaked fur, some spec ...
. In the gut, the larvae may cause serious lesions by attempting to bore through the intestinal walls. Symptoms include
nausea Nausea is a diffuse sensation of unease and discomfort, sometimes perceived as an urge to vomit. While not painful, it can be a debilitating symptom if prolonged and has been described as placing discomfort on the chest, abdomen, or back of the ...
,
vomiting Vomiting (also known as emesis and throwing up) is the involuntary, forceful expulsion of the contents of one's stomach through the mouth and sometimes the Human nose, nose. Vomiting can be the result of ailments like Food-poisoning, foo ...
, pain in the abdomen, and bloody
diarrhea Diarrhea, also spelled diarrhoea, is the condition of having at least three loose, liquid, or watery bowel movements each day. It often lasts for a few days and can result in dehydration due to fluid loss. Signs of dehydration often begin wi ...
. Both living and dead larvae may pass in the stool. Some species also have been known to cause naso-oral and urogenital myiasis. In
forensic entomology Forensic entomology is the scientific study of the colonization of a dead body by arthropods. This includes the study of insect types commonly associated with cadavers, their respective life cycles, their ecological presences in a given environme ...
, 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. 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.


Description

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. Like all tephritoid flies, female piophilids have an extensible
ovipositor The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
. 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 described species in 21 genera, mainly
Holarctic The Holarctic realm is a biogeographic realm that comprises the majority of habitats found throughout the continents in the Northern Hemisphere. It corresponds to the floristic Boreal Kingdom. It includes both the Nearctic zoogeographical region ...
in distribution, though some species are
cosmopolitan Cosmopolitan may refer to: Food and drink * Cosmopolitan (cocktail), also known as a "Cosmo" History * Rootless cosmopolitan, a Soviet derogatory epithet during Joseph Stalin's anti-Semitic campaign of 1949–1953 Hotels and resorts * Cosmopoli ...
. 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: * '' Actenoptera'' * '' Allopiophila'' * ''
Amphipogon ''Amphipogon'', the greybeard grasses, is a genus of Australian plants in the grass family. ; Species ; formerly included see ''Diplopogon Melanocenchris '' * ''Amphipogon humilis - Melanocenchris jacquemontii'' * ''Amphipogon setaceus - Mel ...
'' * '' Bocainamyia'' * '' Centrophlebomyia'' * '' Dasyphlebomyia'' * '' Diacanthomyia'' * '' Lasiopiophila'' * '' Mycetaulus'' * '' Neopiophila'' * '' Neottiophilum'' * ''
Piophila ''Piophila'' is a genus of small flies which includes the species known as the cheese fly. Both ''Piophila'' species feed on carrion, including human corpses. Description ''Piophila'' are small dark flies with unmarked wings. The setulae (fine ...
'' * '' Piophilosoma'' * '' Prochyliza'' * '' Protopiophila'' * '' Protothyreophora'' * '' Pseudoseps'' * '' Pygopiophila'' * '' Stearibia'' * '' Thyreolepida'' * ''
Thyreophora Thyreophora ("shield bearers", often known simply as "armored dinosaurs") is a group of armored ornithischian dinosaurs that lived from the Early Jurassic until the end of the Cretaceous. Thyreophorans are characterized by the presence of body ...
'' Recent works containing keys for identification of the Piophilidae include: * * 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. * 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 * A still useful older work is Séguy, E. (1934) ''Diptères: Brachycères''. II. ''Muscidae acalypterae, Scatophagidae''. Paris: Éditions Faune de France 2
Bibliotheque Virtuelle Numerique pdf


See also

* '' Thyreophora cynophila'' *
Cheese mite Cheese mites (for instance '' Tyrophagus casei'' or other species) are mites that are used to produce such cheeses as ', Cantal and Mimolette. The action of the living mites on the surface of these cheeses contributes to the flavor and gives them ...
*
Casu marzu (; literally 'rotten/putrid cheese'), sometimes spelled , and also called , and in Sardinian, is a traditional Sardinian sheep milk cheese that contains live insect larvae (maggots). A variation of this cheese exists also in Corsica (Franc ...


Genera


The Western Palearctic subfamilies, tribes and genera of Piophilidae - ''Fauna Europaea''


Species lists


Palaearctic





Japan

World list


Image galleries

* *
Images at Diptera.info


References

* * Lieutenant Brian F. Prendergast, USN (2001)
Filth Flies: Significance, Surveillance and Control in Contingency Operations (.pdf format)
Retrieved 1 October 2005. *


External links

*



on the UF /
IFAS IFAS may refer: * Institute of Food and Agricultural Sciences * Integrated Fixed-Film Activated Sludge, a sewage treatment process * International French adjectival system In rock climbing, mountaineering, and other climbing disciplines, clim ...
Featured Creatures Web site (note that many of the images on this site show other species misidentified as Piophila casei) * {{Taxonbar, from=Q135117 Brachycera families Taxa named by Pierre-Justin-Marie Macquart