''Phormia regina'', the black blow fly, belongs to the
blow fly family
Calliphoridae
The Calliphoridae (commonly known as blow flies, blow-flies, carrion flies, bluebottles, greenbottles, or cluster flies) are a family of insects in the order Diptera, with almost 1,900 known species. The maggot larvae, often used as fishing ba ...
and was first described by
Johann Wilhelm Meigen
Johann Wilhelm Meigen (3 May 1764 – 11 July 1845) was a German entomologist famous for his pioneering work on Diptera.
Life
Early years
Meigen was born in Solingen, the fifth of eight children of Johann Clemens Meigen and Sibylla Margaretha ...
.
The black blow fly's wings are specialized with a sharp bend. These flies are also have well-developed
calypter
A calypter is either of two posterior lobes of the posterior margin of the forewing of flies between the extreme posterior wing base and the alula, which covers the halteres.
The lower calypter is the proximal calypter (synonyms: squama (of som ...
s. Blow flies generally are about the size of a house fly or a little larger, and many are metallic blue or green in color. Key characteristics of this species include black gena, mostly white
calypters, and
anterior
Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
thoracic
The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the crea ...
spiracle Spiracle or spiraculum may refer to:
* Spiracle (arthropods), opening in the exoskeletons of some arthropods
* Spiracle (vertebrates), openings on the surface of some vertebrates
* Spiraculum, a genus of land snails in family Cyclophoridae
Cycl ...
s that appear to be orange yellow due to being surrounded by bright orange
setae
In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms.
Animal setae
Protostomes
Annelid setae are stiff bristles present on the body. Th ...
.
''P. regina'' is especially important to
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 ...
. Female ''P. regina'', like other blow flies in the family Calliphoridae,
oviposit
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 ...
their eggs on
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 ...
, where they hatch. The
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 develop through three
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 ...
stages until
pupa
A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
tion.
Adult black blow flies aggregate on feces in order to mate. The success of these mating interactions has been studied appears related to size and diet of the adult flies. Following successful mating encounters, the adult females are then attracted to decaying material rather quickly for oviposition,
allowing forensic entomologists to use development stages of larva as a clue in determining an approximate time of death. ''P. regina'' are extremely common in the United States and other areas in North America.
Combined with their importance in forensic entomology, these flies have been relatively well studied, particularly for variables affecting their development.
Taxonomy
''Phormia regina'' was described by the German
entomologist
Entomology () is the scientific study of insects, a branch of zoology. In the past the term "insect" was less specific, and historically the definition of entomology would also include the study of animals in other arthropod groups, such as arach ...
Johann Wilhelm Meigen in 1826. Its specific
epithet
An epithet (, ), also byname, is a descriptive term (word or phrase) known for accompanying or occurring in place of a name and having entered common usage. It has various shades of meaning when applied to seemingly real or fictitious people, di ...
is derived from the
Latin
Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
word ''regina'' meaning 'queen'.
Description
''Phormia regina'' adults are metallic blue or green in color and have a distinctive set of orange setae near the anterior thoracic spiracle.
They are also recognizable by their black gena (the side of the head below eye level), mostly white calypteres, and the distinctive bend in their wings.
Distribution
''P. regina'' is commonly found across the entirety of the United States as well as across other regions in North America
Habitat
''P. regina'' can be found on human corpses, animal carcasses, and feces. Researchers have observed these flies on a wide variety of animal excrement, but at least in the northwestern United States, they exhibit a preference for human, mink, and swine feces.
''P. regina'' is predominantly found in the northern United States during spring and summer months, but in the winter they are localized to more southern regions.
This localization is tentatively linked to the importance of temperature on their development.
These flies prefer rural areas, especially near water sources, as opposed to urban spaces—at least in the Bay Area.
Life history
The life cycle and development of ''Phormia regina'' is similar to that of most other
Dipteran
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 m ...
species, in which females oviposit their eggs onto a nutrient substrate. Then, after hatching, the
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 feed throughout three
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 ...
stages until they have stored up enough calories to commence pupation and finally emerge as adult blow flies. Each transition from first, second, and third instar is marked by a
molt
In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is the manner in which an animal routinely casts off a part of its body (often, but not always, an outer ...
, and eventually the third-instar larvae develop
sclerotized (hardened) casings which envelop and protect them throughout
metamorphosis
Metamorphosis is a biological process by which an animal physically develops including birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation. Some inse ...
.
Larval development
Relatively few studies have been conducted on the adults of this species in comparison to those on larval development, mostly due to the importance of blow fly larvae in determining the
''post mortem'' interval (PMI) of corpses during investigations by
forensic
Forensic science, also known as criminalistics, is the application of science to Criminal law, criminal and Civil law (legal system), civil laws, mainly—on the criminal side—during criminal investigation, as governed by the legal standard ...
entomologists. For this reason, many researchers have conducted experiments to investigate the effect of various environmental factors on the duration time of larval development in this species (as well as many others).
Because the larval life cycle of this species is dependent on a climate with temperatures ranging from 12.7
°C
The degree Celsius is the unit of temperature on the Celsius scale (originally known as the centigrade scale outside Sweden), one of two temperature scales used in the International System of Units (SI), the other being the Kelvin scale. The ...
(55
°F) to 35 °C (95 °F), it tends to inhabit the northern regions of the United States during summer months and southern regions in the winter. Researchers have discovered that at 40 to 45 °C, larval development occurs normally until the
prepupal stage, at which point a majority of the larvae die. The few able to
pupate
A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
at these high temperatures do not emerge as adults. The lowest temperature threshold for this species was found to be 12.5 °C below which females will not oviposit. The highest rate of development (with survival into adulthood) was seen at a constant temperature of 35 °C, where the average time of adult emergence was 265 hours (about 11 days). Flies at constant temperatures between 15 and 30 °C (at 5-degree increments) developed slower, with flies at the coolest temperatures taking the longest. Cyclic temperatures ranges of 25 to 35 °C and 15 to 25 °C proved to decrease the rate of development when compared to constant temperatures. (The cyclic temperature data were collected by placing specimens in an incubator which steadily alternated between the maximum and minimum temperatures of a particular 10-degree range (e.g. 25 to 35 °C). Each 10-degree fluctuation took place over a 12-hour span.)
Also, studies have been conducted to assess the effects of light exposure on developmental variability in larvae. Larvae exposed to cyclic photoperiods (shifting intermittently between 12 hours of light to 12 hours of darkness) have higher rates of development than larvae exposed to constant photoperiods (24 hours of light per day). These findings suggest that darkness may be a stimulus for larval growth. However, these variations in light photoperiods failed to influence pupal duration times.
Adult development
While this species is in the wild, dung constitutes a majority of the
nutrition
Nutrition is the biochemical and physiological process by which an organism uses food to support its life. It provides organisms with nutrients, which can be metabolized to create energy and chemical structures. Failure to obtain sufficient n ...
al intake used for sexual development in both male and female adults, but diets with higher
protein
Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
intake better facilitate
mating
In biology, mating is the pairing of either opposite-sex or hermaphroditic organisms for the purposes of sexual reproduction. ''Fertilization'' is the fusion of two gametes. ''Copulation'' is the union of the sex organs of two sexually reproduc ...
ability of both sexes. Female sexual maturity requires the completion of 10 stages of
follicle development in the ovaries to produce eggs that are completely mature, and ovaries in females which have been deprived of a high-protein diet do not develop fully. Although it is possible for females on a diet of only dung to reach the final stage of sexual maturity, it takes much more time than if they were to feed exclusively on
beef liver; even then, a lower percentage of those feeding on dung will have fully developed. An experiment by Stoffolano demonstrated this, revealing that 100% of females feeding exclusively on beef liver were able to reach the final stage of sexual development after 13 days, while only 78% of females were able to do so when feeding exclusively on pig dung over a 20-day period. Although some Dipteran species oviposit on dung, ''P. regina'' females exclusively deposit their eggs on carrion.
The
neuroendocrine
Neuroendocrine cells are cells that receive neuronal input (through neurotransmitters released by nerve cells or neurosecretory cells) and, as a consequence of this input, release messenger molecules ( hormones) into the blood. In this way they b ...
system in adult males, which controls their mating behavior, must be stimulated before they will mate with a female. Protein in the male diet is not necessarily needed for this stimulation, but Stoffolano also saw that higher percentages of females were successfully
inseminate
Insemination is the introduction of sperm into a female’s reproductive system for the purpose of impregnating, also called fertilizing, the female for sexual reproduction. The sperm is introduced into the uterus of a mammal or the oviduct of an ...
d by male specimens which had been fed either dung or beef liver versus specimens with a diet of only sugar. Protein is not generally necessary for
spermatogenesis
Spermatogenesis is the process by which haploid spermatozoa develop from germ cells in the seminiferous tubules of the testis. This process starts with the mitotic division of the stem cells located close to the basement membrane of the tubule ...
in male flies, but it is paramount for accessory reproductive gland development, higher rates of copulation, and the capability of impregnating females.
An additional study found a positive correlation between male head size and the size of the
aedeagi
An aedeagus (plural aedeagi) is a reproductive organ of male arthropods through which they secrete sperm from the testes during copulation (zoology), copulation with a female. It can be thought of as the insect equivalent of a mammal's penis, t ...
(the external reproductive organs), which has been proposed as a possible reason for lower percentages of insemination between small males and large females within this species.
Food sources
Feeding habits vary greatly throughout the life cycle of ''P. regina''. The feeding habits of adult flies depend both on sex and reproduction status of the fly. This is particularly important with respect to their protein intake, which literature suggests has an important role in sexual maturation and copulation ability in adult ''P. regina.''
Like most other flies, the black blow fly feeds via sponging. Though adults require various types of nutrients, in the wild they are typically able to procure all their nutrition from feces.
Larvae
Adult females oviposit eggs on carrion. These eggs hatch into larvae that feed on the decaying tissue.
Adult females
Virgin adult females, when given access to protein and carbohydrate sources, utilize both for food. However, after their emergence as adults, they will gradually increase their protein intake until they are approximately 8 to 10 days old. From here they will continue to intake protein at a relatively constant rate that is lower than their required carbohydrate intake. Once a female has mated, either for the first time or all subsequent times, a spike in protein intake occurs following oviposition of her eggs.
Adult males
Protein intake in adult male black blow flies follows similar trends to that of virgin adult females; however, they require less total protein than females. The males will gradually increase their protein intake for the first 8–10 days of adulthood. They will then drop their protein intake to a relatively low and constant amount.
Mating
Adult black blow flies aggregate on feces, particularly animal dung pats, not only because they act as a food source, but also in order to mate. Some research has suggested a preference for settling on human, swine, and mink feces, but this may be localized to the northwestern region of the United States.
''P. regina'' is a
panmictic
Panmixia (or panmixis) means random mating. A panmictic population is one where all individuals are potential partners. This assumes that there are no mating restrictions, neither genetic nor behavioural, upon the population and that therefore all ...
species.
Research supports mating success of adult males positively correlates to their size. Larger males have larger
aedeagus which leads to more successful insemination of females. The mechanism of how and why larger aedeagus lead to better insemination is unknown.
Female reproductive system
Female black blow flies each have a pair of ovaries and lateral
oviduct
The oviduct in mammals, is the passageway from an ovary. In human females this is more usually known as the Fallopian tube or uterine tube. The eggs travel along the oviduct. These eggs will either be fertilized by spermatozoa to become a zygote, o ...
s, one common oviduct, a vagina, and a bursa copulatrix, which is simply a sac-like cavity within the female that is roughly equivalent to a uterus. Each ovary has a lateral oviduct; these two lateral oviducts meet to form the common oviduct and connect the ovaries to the burs copulatrix. Three
spermatheca
The spermatheca (pronounced plural: spermathecae ), also called receptaculum seminis (plural: receptacula seminis), is an organ of the female reproductive tract in insects, e.g. ants, bees, some molluscs, oligochaeta worms and certain other ...
e are connected to the bursa copulatrix via spermathecal ducts, and two accessory glands attach to the bursa copulatrix near the spermathecae entries. The vagina is connected to the bursa copulatrix on the opposite end of the ovaries and allows the male aedeagus access to the bursa copulatrix and spermathecae.
Male reproductive system
Male black blow flies have an external phallus that is a tube-like structure with unique flaps centrally located. The phallus can be divided up into several different sections. There is the hypophallus, which protrudes from the abdomen of the fly. The cornua, which is the name given to the centrally located wing-like flaps on the phallus, protrude from the hypophallus. A pair of paraphallic bulbs are embedded in the cornua with only their tips jutting out. These bulbs are sclerotized, meaning they are hardened, as are a variety of spines on the outside of the cornua. From the cornua a narrower tube structure, the acrophallus, extends with a terminal fluted opening, referred to as the gonophore, from which sperm and accessory secretions flood.
Copulation
During mating the tip of the male's phallic structure is near the spermathecal ducts of the female's reproductive system. The wing-like flap structures on the male's phallus are situated in grooves within the bursa copulatrix of the female's reproductive system. The males also have appendages used to grasp and hold the female during copulation.
Parental care
Similarly to other flies in the family Calliphoridae, ''P. regina'' exhibit little or no parental care. Females, however, oviposit their eggs after mating on carrion, leaving the eggs to hatch and develop on their own.
Oviposition
Mature ''P. regina'' females are known to oviposit their eggs on animal carcasses following copulation. Interestingly, it has been seen that the olfactory senses of female flies play some role in their decision of where and when to oviposit. Research suggests adult female ''P. regina'' will increase their ovipositing in the presence of a medium that provides odor stimuli to the fly.
Social behavior
Maggot mass effect
![Decomposition00](https://upload.wikimedia.org/wikipedia/commons/c/c0/Decomposition00.jpg)
Black blow fly larvae like many other blow flies exhibit the
maggot
A maggot is the larva of a fly (order Diptera); it is applied in particular to the larvae of Brachycera flies, such as houseflies, cheese flies, and blowflies, rather than larvae of the Nematocera, such as mosquitoes and crane flies.
...
mass effect: a raise in temperature due to the aggregation of maggots. This rise in temperature may be beneficial to the larvae by aiding in feeding and development as well as protection from dangerously cold temperatures, predators, and parasites. Studies with other blow flies suggest that the change in temperature seen in the maggot mass effect functions well at a wide range of temperatures and that the larvae do not reduce their heat production at higher temperatures.
Genetics
The entire genome of ''P. regina'' has been sequenced so that it may serve as a reference genome that can be used in comparison with other Calliphoridae species. This fly has a
chromosome number
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectivel ...
of 2n=12. The sex chromosomes of ''P. regina'' are a heteromorphic chromosome pair, meaning they have two morphologically distinct chromosomes that remain a
homologous chromosome
A couple of homologous chromosomes, or homologs, are a set of one maternal and one paternal chromosome that pair up with each other inside a cell during fertilization. Homologs have the same genes in the same loci where they provide points alon ...
pair.
Picard and Wells used
amplified fragment length polymorphism
AFLP-PCR or just AFLP is a PCR-based tool used in genetics research, DNA fingerprinting, and in the practice of genetic engineering. Developed in the early 1990s by KeyGene, AFLP uses restriction enzymes to digest genomic DNA, followed by liga ...
s (AFLP) to create genetic profiles of ''P. regina'' flies collected from different regions to compare both within region and across region relatedness. They discovered adult flies take from the same corpse showed high relatedness, but between regions genetic variability was high. They suggest these findings may be especially important to forensic entomology with respect to determining if a recovered body has moved locations by comparing AFLP data across individual ''P. regina''.
Physiology
''P. regina'', like other flies, is
poikilothermic
A poikilotherm () is an animal whose internal temperature varies considerably. Poikilotherms have to survive and adapt to environmental stress. One of the most important stressors is temperature change, which can lead to alterations in membrane ...
: the growth and development of the fly is dependent on temperature. At room temperature, the egg to pupal stage lasts about 6–11 hours. With an increase in temperature of the surrounding environment, metabolic rates of the blow fly typically increase, causing an increase in the rate of growth and development.
However, this species is intolerant to warmer temperatures, unlikely to survive to adulthood at temperatures of 40 °C or greater. This observation provides an explanation as to why the species cannot survive the summer months in southern regions of the US. The adult organism's activity becomes inhibited by temperatures lower than 12.5 °C.
In addition to an increase in the growth and development, temperature also has a profound impact on female oviposition.
It is key to note the fluctuations between diurnal and nocturnal temperatures.
It is important to note the larval stage of the blowfly, due to its importance in forensics. The larvae have posterior spiracles, small openings on the back used for the intake of oxygen. Larvae are also equipped with mouth hooks used for the physical breakdown of proteins when feeding, while
proteolytic
Proteolysis is the breakdown of proteins into smaller polypeptides or amino acids. Uncatalysed, the hydrolysis of peptide bonds is extremely slow, taking hundreds of years. Proteolysis is typically catalysed by cellular enzymes called proteases, ...
enzyme
Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products. A ...
s are used for chemical breakdown of these proteins.
Interactions with humans
Myiasis
''P. regina'' larvae have been shown to be an important organism in
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 ...
of both humans and animals, particularly in the southern region of the United States. Often in a myiatic infection where ''P. regina'' larvae are found, they are the only species involved; however, they have also been found to be present in a wound with ''
Cochliomyia americana'' as well.
Forensic importance
''P. regina'' is a very important species in medico-criminal entomology, an area 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 ...
which uses
entomologists
Entomology () is the scientific study of insects, a branch of zoology. In the past the term "insect" was less specific, and historically the definition of entomology would also include the study of animals in other arthropod groups, such as arach ...
to aid with
arthropod
Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
evidence
Evidence for a proposition is what supports this proposition. It is usually understood as an indication that the supported proposition is true. What role evidence plays and how it is conceived varies from field to field.
In epistemology, evidenc ...
in
criminal investigation
Criminal investigation is an applied science that involves the study of facts that are then used to inform criminal trials. A complete criminal investigation can include searching, interviews, interrogations, evidence collection and preservatio ...
s.
[Byrd, J. H., and J. L. Castner. 2001. Forensic entomology : The utility of arthropods in legal investigations. CRC Press, New York.] This aspect of forensic science stresses using arthropod evidence in solving crimes, often of a violent nature, through two ideal approaches to estimate
''post mortem'' interval (PMI). The first approach accounts for the general succession of arthropod communities, and the second considers environmental influences on the development of arthropods.
With knowledge of the regional insect fauna and times of carrion colonization, the insect gathering associated with the remains can be analyzed to determine a window of time in which death took place. PMI is a critical aspect of criminal investigations following recovery of a corpse. PMI relies on an entomologist’s ability to correlate the species or stage of development of arthropods, in this case ''P. regina'', to an approximation of the elapsed period between a person's death and the discovery of his/her body.
Blow flies are usually the first insects to colonize a body, frequently within minutes after death. ''P. regina'' adults and larvae are attracted to the body because, during
decomposition
Decomposition or rot is the process by which dead organic substances are broken down into simpler organic or inorganic matter such as carbon dioxide, water, simple sugars and mineral salts. The process is a part of the nutrient cycle and is e ...
, the remains go through rapid physical, biological, and chemical changes. If a corpse is found, the presence of Dipteran larvae may be used to suggest PMI by estimating the time it would have taken for eggs to have been deposited and larvae to have developed to the stage they were collected, while taking into account environmental factors. Blow flies will lay their eggs on the corpse, usually in a wound, if present, or in any of the natural
orifices
A body orifice is any opening in the body of an animal.
External
In a typical mammalian body such as the human body, the external body orifices are:
* The nostrils, for breathing and the associated sense of smell
* The mouth, for eating, bre ...
, and the maggot age can give a date of death accurate to a day or less, and is used in the first few weeks after death.
[Anderson, G.S. Forensic Entomology: the use of insects in death investigations]
/ref>
Research is being conducted to further perfect the dating of a post mortem interval, PMI. One study suggests that ''P. regina'' occasionally oviposits on carrion at night only when certain conditions are met. A similar study found a combination of artificial lighting, warm temperatures, and the onset of low-pressure atmospheric conditions encourages nocturnal oviposition in ''P. regina'' Other studies suggest ''P. regina'' preferentially distribute their eggs during the day. Confirming the preference for daytime or nighttime oviposition in ''P. regina'' could aid in increasing the accuracy of PMI determination, and the disagreement necessitates more experimentation to investigate this environmental effect.
Medical Importance
''Phormia regina'' is a flesh-eating fly not usually used for medical gain. However, the maggots from the black blow fly are used in maggot therapy
Maggot therapy (also known as larval therapy) is a type of biotherapy involving the introduction of live, disinfected maggots (fly larvae) into non-healing skin and soft-tissue wounds of a human or other animal for the purpose of cleaning out th ...
, a type of biotherapy involving the intentional introduction of live, disinfected maggots into the skin and soft tissue wound of a human or animal for the purpose of selectively cleaning out only the necrotic tissue to promote healing.Hays, A. History of Medicine Part 1: Maggot Therapy. 15 March 2007
/ref>
References
Further reading
*Anderson, G.S
/small>
*
*Byrd, J. H., and J. L. Castner. 2001. Forensic entomology : The utility of arthropods in legal investigations. CRC Press, New York.
*
*
*
*
*
*Rognes, Knut. 1991. Blowflies (Diptera, Calliphoridae) of Fennoscandia and Denmark. E.J. Brill/Scandinavian Science Ltd., 272pp.
*
*
*Watson L., Dallwitz M.J.
/small>
*
{{Taxonbar, from=Q7187389
Calliphoridae
Insects described in 1826
Diptera of North America
Taxa named by Johann Wilhelm Meigen