Sarcophaga bullata
Encyclopedia
Sarcophaga bullata, or the grey flesh fly, is a species of fly
belonging to the family
Sarcophagidae. It is 7.5 to 15 millimeters in length and is very similar in appearance and behavior to a closely related species, Sarcophaga haemorrhoidalis
. S. bullata is a common scavenger species in the Eastern United States
, but is found throughout the Nearctic
region. Identification down to the species level in the Sarcophagidae family is notably difficult and relies primarily on the male genitalia. Though limited information is available regarding S. bullata, it has gained increasing recognition in the field of forensic entomology as a forensically-relevant fly species, as it may be among the first species to colonize human remains. In these instances, recovered maggots may be analyzed for post-mortem interval (PMI) estimations, which may be used as evidence in courts of law. Current studies regarding S. bullata have revealed a maternal effect operating in these flies that prevents pupal diapause
under certain environmental conditions, which is an important factor to be considered during forensic analyses.
and some Tachinidae
. Most, however, consider S. bullata to be a member of the Sarcophaginae subfamily within Sarcophagidae. Beyond that, the family can be split into tribes before defining genus. S. bullata is widely accepted to be a member of the Sarcophaga genus, but it is also included by some into a subgenus, Neobellieria.
(hair) of the antenna is plumose (feather-like) only at the base, unlike a Calliphorid fly, whose arista is plumose the entire length. The eyes of S. bullata are bright red in color, and are rather widely separated on the top of the head. On the genae, or cheeks, are long bristly hairs. Two orbital bristles are present on the females, while the orbital bristles are absent on the males.
are three contrasting black and grey longitudinal stripes; these stripes do not continue onto the abdomen. Sarcophaga
have 4, rather than the normal 2 or 3, notopleural bristles, and 3 sternopleural bristles. A prominent row of black bristles (setae) can be found on each side of the thorax, just above the base of the hind leg and just under the base of the wing. These two sets of bristles are what differentiate Sarcophagidae from Muscidae
. S. bullata lacks the postscutellum, the large swelling underneath the scutellum of the thorax, which distinguishes the flesh fly from Tachinid. S. bullata have black legs, translucent wings, and do not have a costal spine.
larvae. This must be accounted for when using the species in forensic work to calculate a post-mortem interval.
First instar:
First instar Sarcophagidae larvae are 0.5 to 5.0 millimeters long. The Sarcophaginae larvae, specifically, usually lack a labral sclerite, but in some cases it is present as a small, triangular plate located between the mandibles. The first segment is at times sclerotized on the dorsal surface, which can form a partial head capsule. The Sarcophaginae have a hypopharyngeal sclerite that is sometimes fused with the large tentoropharyngeal sclerite. They also lack an anterior spiracle, while the posterior spiracle usually consists of two lobes; rarely it may be bispinose.
Second instar:
Second instar larvae of the Sarcophagidae family are 4.0 to 10.0 millimeters long and closely resemble the third instar with the exception being that the posterior spiracle has only two openings.
Third instar:
Third instar larvae are 9.5 to 20 millimeters long. The mandibles are by now large and strongly curved. The dorsal cornu of the tentorpharyngeal sclerite appears incised due to a narrow apical window. The anterior spiracle is present with a short stalk that branches into several nodes, resembling a fan. The posterior spiracles now consist of three vertical slits within a sunken oval cavity. This cavity is surrounded by 8 to 12 tubercles, of which the dorsal ones are often sclerotized and have toothed processes, while the ventral tubercles are often covered in nipple-like projections. The anal opening is located in a fairly prominent posteroventral lobe that also has lateral tubercles.
region, or the part of North America that is north of Mexico. It is most common in the eastern and southern United States, but can be found from the western cost of the United States to Canada. S. bullata are commonly found in rural and urban environments, and it is commonly found in houses and indoor dwellings, especially during the summer months. The flies prefer the warmer months, and are active from May to September, with the majority of the activity in August and September.
concentrations. Extensive research has suggested that S. bullata females in particular favor amino acid-containing nectars as opposed to strictly sugary nectars, perhaps to secure healthy levels of protein for successful reproduction.
Adult flesh flies are attracted to animal remains in both early and late decomposition stages, and have been known to fly through even the most extreme weather conditions to reach carrion. Though adult flies of the family Calliphoridae typically arrive on human remains before those of Sarcophagidae, sarcophagid adults are unhindered by nearly all climatic conditions and may reach the carrion first during particularly stormy weather. Rather than laying eggs, S. bullata females retain fertilized eggs internally until they develop into first-instar larvae, which are then deposited directly onto carrion. As sarcophagid larvae are larger than those of most other colonizing flies, they oftentimes present significant larval competition for other species. Though sarcophagid larvae have been attributed to causing myiasis
, a condition in which maggots feed on the tissue of a living human or animal, they are most commonly discovered feeding on animal remains. In the southeastern United States, species of the genus Sarcophaga are the principal flies found during the warm summer months on bodies located indoors, as females frequently enter dwellings to deposit its larvae.
In terms of relevance to decomposing tissue analysis, forensic entomologists must disregard an egg development time period in post-mortem interval (PMI) calculations, as S. bullata and other sarcophagids deposit live larvae on carrion as opposed to eggs. For the same reason, fly egg masses discovered on human remains can be automatically ruled-out as belonging to sarcophagid flies.
, and occasionally consume the smaller larvae present on animal remains.
, Sceloporus undulatus or Eastern Fence Lizard
, and the Perimyotis subflavus (Eastern Pipistrelle
).
Another important consideration with the grey flesh flies concerns maggot mass temperature. "Maggot mass" refers to a large group of maggots clustered together at a particular location on a carcass. Reporting an accurate temperature is critical to properly calculating accumulated degree hours in the estimation of a post-mortem interval. Accumulated degree hours and similar measures are determined to attempt to trace back the "age" of the maggots and determine the time of exposure of decaying tissue. Some scientists claim that maggot mass temperature is generally significantly above ambient air temperature. Others maintain that while maggot mass temperature may be elevated in a laboratory setting, in the field the maggots self-regulate their temperature by moving in and out of the feeding mass. Obviously, more research is needed to determine the general effects of maggot mass temperature.
Other work has been reported concerning tanning in S. bullata with the age pigment lipofuscin
. Lipofuscin accumulation in larvae is a linear function of time. Some of the pigment is disposed of at the pupal stage and then linear accumulation resumes in adults. Many projects have focused on an assay of this pigment as it could hold a rather accurate technique for aging S. bullata and other forensically important insects.
Accumulated degree days can be determined by subtracting the minimum threshold from the average ambient temperature over a 24-hour period and multiplying the difference by the corresponding number of days. Accumulated degree hours, which are equally useful, are similarly determined by subtracting the minimum threshold from the ambient temperature over a 1-hour period and multiplying the difference by the corresponding number of hours. The term "minimum threshold" refers to the lowest temperature at which an insect species will develop; any temperatures below this threshold will result in delayed development and probable death of the insect in question.
It is important to note that similar calculations may be carried out to determine “accumulated degree minutes,” “accumulated degree seconds,” etc., though ADD and ADH are typically used in forensic analyses.
For S. bullata, the following temperature data has been determined in a laboratory setting. These calculations correspond to a standardized temperature of 27 °C.
when reared under short day conditions. It was determined that certain environmental stressors or chemical treatments on the mother, such as extreme temperature shocks, food deprivation and reducing maternal size, were ineffective in altering the diapause-suppressant. However, there were several chemical agents that proved effective. Gamma-aminobutyric acid
, octopamine
and pilocarpine
all countered the maternal affect and permitted the expression of diapause in the flesh fly progeny. The ability to express diapause is an important consideration for forensic entomologists as diapause allows the pupae to become metabollically "dormant" until environmental conditions improve; this would significantly affect pupal time estimations and subsequent post-mortem interval estimations.
Fly
True flies are insects of the order Diptera . They possess a pair of wings on the mesothorax and a pair of halteres, derived from the hind wings, on the metathorax...
belonging to the family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...
Sarcophagidae. It is 7.5 to 15 millimeters in length and is very similar in appearance and behavior to a closely related species, Sarcophaga haemorrhoidalis
Sarcophaga haemorrhoidalis
Sarcophaga haemorrhoidalis, also known as the red-tailed flesh fly, is a fly in the Sarcophagidae family. This fly often breeds in carrion and feces, making it a possible vector for disease. The larvae of this species can cause myiasis, as well as accidental myiasis...
. S. bullata is a common scavenger species in the Eastern United States
United States
The United States of America is a federal constitutional republic comprising fifty states and a federal district...
, but is found throughout the Nearctic
Nearctic
The Nearctic is one of the eight terrestrial ecozones dividing the Earth's land surface.The Nearctic ecozone covers most of North America, including Greenland and the highlands of Mexico...
region. Identification down to the species level in the Sarcophagidae family is notably difficult and relies primarily on the male genitalia. Though limited information is available regarding S. bullata, it has gained increasing recognition in the field of forensic entomology as a forensically-relevant fly species, as it may be among the first species to colonize human remains. In these instances, recovered maggots may be analyzed for post-mortem interval (PMI) estimations, which may be used as evidence in courts of law. Current studies regarding S. bullata have revealed a maternal effect operating in these flies that prevents pupal diapause
Diapause
Diapause is the delay in development in response to regularly and recurring periods of adverse environmental conditions. It is considered to be a physiological state of dormancy with very specific initiating and inhibiting conditions...
under certain environmental conditions, which is an important factor to be considered during forensic analyses.
Taxonomy
Classification of the species has also proved difficult, as it has for much of the Sarcophagidae family. C.H. Murran, for instance, included Sarcophagidae in a larger family, Metopidae, that added some MuscidaeMuscidae
Muscidae are a family of flies found in the superfamily Muscoidea. The apical segment of the antennae of Muscidae are plumose, and the basal portion is smooth....
and some Tachinidae
Tachinidae
Tachinidae is a large and rather variable family of true flies within the insect order Diptera, with more than 8,200 known species and many more to be discovered. There are over 1300 species in North America. Insects in this family are commonly called tachina flies or simply tachinids...
. Most, however, consider S. bullata to be a member of the Sarcophaginae subfamily within Sarcophagidae. Beyond that, the family can be split into tribes before defining genus. S. bullata is widely accepted to be a member of the Sarcophaga genus, but it is also included by some into a subgenus, Neobellieria.
Head
Sarcophaga bullata is approximately 7.5 to 15 millimeters in length. The head is colored ashen grey, and the aristaArista (biology)
In insect anatomy the arista is a simple or variously modified apical or subapical bristle, arising from the third antennal segment. It is the evolutionary remains of antennal segments, and may sometimes show signs of segmentation...
(hair) of the antenna is plumose (feather-like) only at the base, unlike a Calliphorid fly, whose arista is plumose the entire length. The eyes of S. bullata are bright red in color, and are rather widely separated on the top of the head. On the genae, or cheeks, are long bristly hairs. Two orbital bristles are present on the females, while the orbital bristles are absent on the males.
Thorax
Present on the thoraxThorax
The thorax is a division of an animal's body that lies between the head and the abdomen.-In tetrapods:...
are three contrasting black and grey longitudinal stripes; these stripes do not continue onto the abdomen. Sarcophaga
Sarcophaga
Sarcophaga is a genus of true flies, the type of the flesh-fly family .This genus occurs essentially worldwide. These flies are generally well-sized and of a greyish color; like many of their relatives, the typical patterns are lengthwise darker stripes on the thorax and dark and light square dots...
have 4, rather than the normal 2 or 3, notopleural bristles, and 3 sternopleural bristles. A prominent row of black bristles (setae) can be found on each side of the thorax, just above the base of the hind leg and just under the base of the wing. These two sets of bristles are what differentiate Sarcophagidae from Muscidae
Muscidae
Muscidae are a family of flies found in the superfamily Muscoidea. The apical segment of the antennae of Muscidae are plumose, and the basal portion is smooth....
. S. bullata lacks the postscutellum, the large swelling underneath the scutellum of the thorax, which distinguishes the flesh fly from Tachinid. S. bullata have black legs, translucent wings, and do not have a costal spine.
Abdomen
The abdomen is grey and slender, with a median black stripe. It is checkered with a shifting pattern and black reflecting spots.Sarcophaga most resemble the blow fly, but never have metallic colored abdomens. There are no bristles on the middle tibia of the fly, but the hind tibia has fringed or long hairs irregularly placed. The genitalia is located on the end of the abdomen and in the males is a bright red color. The genitalia is one of the main identification characteristics used to identify the different genera of Sarcophagidae.Development
The life cycle of Sarcophaga bullata is characterized by holometabolous development, though, like all known sarcophagids, the species is larviparous, meaning that the egg develops internally and females then give birth to first-instarInstar
An instar is a developmental stage of arthropods, such as insects, between each molt , until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or assume a new form. Differences between instars can often be seen in altered body proportions, colors, patterns, or...
larvae. This must be accounted for when using the species in forensic work to calculate a post-mortem interval.
Eggs
Though not much has been written specifically on the reproductive cycle of S. bullata, Sarcophagids are thought to be consistent within the family when it comes to life cycle. The eggs themselves are about 0.5 to 3.5 millimeters in length and 0.12 to 0.8 millimeters in width, though hatching occurs within the uterus just before the deposition of larvae.Larvae
Sarcophagidae larvae are white or pale yellow, cylindrical, and tapered anteriorly. All segments beyond the first have anterior and posterior bands of hairs. The mandibles are usually strong and curved, resembling a hook. Posterior spiracles are sunken in, which is a characteristic that can be used to distinguish between flesh fly and blow fly larvae. Larvae are most often found in decaying animal matter.First instar:
First instar Sarcophagidae larvae are 0.5 to 5.0 millimeters long. The Sarcophaginae larvae, specifically, usually lack a labral sclerite, but in some cases it is present as a small, triangular plate located between the mandibles. The first segment is at times sclerotized on the dorsal surface, which can form a partial head capsule. The Sarcophaginae have a hypopharyngeal sclerite that is sometimes fused with the large tentoropharyngeal sclerite. They also lack an anterior spiracle, while the posterior spiracle usually consists of two lobes; rarely it may be bispinose.
Second instar:
Second instar larvae of the Sarcophagidae family are 4.0 to 10.0 millimeters long and closely resemble the third instar with the exception being that the posterior spiracle has only two openings.
Third instar:
Third instar larvae are 9.5 to 20 millimeters long. The mandibles are by now large and strongly curved. The dorsal cornu of the tentorpharyngeal sclerite appears incised due to a narrow apical window. The anterior spiracle is present with a short stalk that branches into several nodes, resembling a fan. The posterior spiracles now consist of three vertical slits within a sunken oval cavity. This cavity is surrounded by 8 to 12 tubercles, of which the dorsal ones are often sclerotized and have toothed processes, while the ventral tubercles are often covered in nipple-like projections. The anal opening is located in a fairly prominent posteroventral lobe that also has lateral tubercles.
Distribution
Sarcophaga bullata do not have wide natural distribution, and can only be found in the NearcticNearctic
The Nearctic is one of the eight terrestrial ecozones dividing the Earth's land surface.The Nearctic ecozone covers most of North America, including Greenland and the highlands of Mexico...
region, or the part of North America that is north of Mexico. It is most common in the eastern and southern United States, but can be found from the western cost of the United States to Canada. S. bullata are commonly found in rural and urban environments, and it is commonly found in houses and indoor dwellings, especially during the summer months. The flies prefer the warmer months, and are active from May to September, with the majority of the activity in August and September.
Feeding habits
Adults of the family Sarcophagidae do not feed exclusively on carrion, despite the weighty implications of the name “flesh flies;” in fact, adult flies are frequently discovered feeding on sweet substances, such as sap and flower nectar. The flesh-eating character of the family refers primarily to sarcophagid maggots, which feed predominantly on carrion, exposed meat, and excrement. Sarcophaga bullata, in addition to all species of the family Sarcophagidae, may be considered “specialized flies,” a term coined by Baker and Baker to describe flies that feed on carrion and dung. Specialized flies have been proven capable of detecting and selectively favoring plant nectars with high amino acidAmino acid
Amino acids are molecules containing an amine group, a carboxylic acid group and a side-chain that varies between different amino acids. The key elements of an amino acid are carbon, hydrogen, oxygen, and nitrogen...
concentrations. Extensive research has suggested that S. bullata females in particular favor amino acid-containing nectars as opposed to strictly sugary nectars, perhaps to secure healthy levels of protein for successful reproduction.
Adult flesh flies are attracted to animal remains in both early and late decomposition stages, and have been known to fly through even the most extreme weather conditions to reach carrion. Though adult flies of the family Calliphoridae typically arrive on human remains before those of Sarcophagidae, sarcophagid adults are unhindered by nearly all climatic conditions and may reach the carrion first during particularly stormy weather. Rather than laying eggs, S. bullata females retain fertilized eggs internally until they develop into first-instar larvae, which are then deposited directly onto carrion. As sarcophagid larvae are larger than those of most other colonizing flies, they oftentimes present significant larval competition for other species. Though sarcophagid larvae have been attributed to causing myiasis
Myiasis
Myiasis is a general term for infection by parasitic fly larvae feeding on the host's necrotic or living tissue. Colloquialisms for myiasis include flystrike, blowfly strike, and fly-blown. In Greek, "myia" means fly....
, a condition in which maggots feed on the tissue of a living human or animal, they are most commonly discovered feeding on animal remains. In the southeastern United States, species of the genus Sarcophaga are the principal flies found during the warm summer months on bodies located indoors, as females frequently enter dwellings to deposit its larvae.
In terms of relevance to decomposing tissue analysis, forensic entomologists must disregard an egg development time period in post-mortem interval (PMI) calculations, as S. bullata and other sarcophagids deposit live larvae on carrion as opposed to eggs. For the same reason, fly egg masses discovered on human remains can be automatically ruled-out as belonging to sarcophagid flies.
Social behavior
It is difficult to quantify the level and nature of “communication” that takes place among sarcophagid adults, though much is known about the role of pheromones in their reproductive physiologies. An experiment by Girard et al. in the 1970s demonstrated that S. bullata males release the sex pheromone hexanal, which was shown to attract over 65% of the females tested from over long distances. In studies regarding the social behavior of adult female sarcophagids larvipositing on animal remains, it has been suggested that adult females prefer to aggregate with other carrion flies and larviposit on carcasses that already have larvae present. A possible explanation for this selective preference is that an increased number of larvae results in a greater concentration of larval enzymatic secretions on the carrion, aiding in digestion. Sarcophagid larvae are also known to outcompete the larvae of other species and cause their extinctionExtinction
In biology and ecology, extinction is the end of an organism or of a group of organisms , normally a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and recover may have been lost before this point...
, and occasionally consume the smaller larvae present on animal remains.
Natural predators
The Sarcophaga bullata has few, if any, specific natural predators due to their dispersal throughout the North American continent. Predators of S. bullata are carnivores, ranging from mammals and reptiles to carnivorous plants. However, a few primary predators are the Stagmomantis californica or the California Mantis, Catopsis berteronianaCatopsis berteroniana
Catopsis berteroniana is an epiphytic bromeliad thought to be a possible carnivorous plant, similar to Brocchinia reducta, although the evidence is equivocal. Its native range is from southern Florida to southern Brazil. It generally grows on the unshaded twigs of trees, and appears to trap more...
, Sceloporus undulatus or Eastern Fence Lizard
Eastern fence lizard
The eastern fence lizard is a medium-sized species of lizard found along forest edges, rock piles, and rotting logs or stumps in the eastern United States...
, and the Perimyotis subflavus (Eastern Pipistrelle
Eastern Pipistrelle
The Tricolored Bat is a species of bat that is widely distributed throughout the eastern parts of North America, ranging west until Kansas and Texas, from Honduras up north until southern Ontario...
).
Forensic related research
As members of Sarcophaga bullata are found from coast to coast in the United States and in parts of Canada, these flesh flies have been the focus of many forensically related studies. Most of these studies deal with the immature stages, as maggots are a helpful tool in the forensic world for determining post-mortem interval estimations. One such study concerning S. bullata determined that when evaluating a post-mortem interval, temperature plays a major role in maturation of the maggots and the decomposition of the surrounding tissue. This is an important consideration for forensic entomologists, as the ambient temperature at the crime scene before and during the colonization of human remains by arthropods must be accounted for to ensure that post-mortem interval estimations are accurate.Another important consideration with the grey flesh flies concerns maggot mass temperature. "Maggot mass" refers to a large group of maggots clustered together at a particular location on a carcass. Reporting an accurate temperature is critical to properly calculating accumulated degree hours in the estimation of a post-mortem interval. Accumulated degree hours and similar measures are determined to attempt to trace back the "age" of the maggots and determine the time of exposure of decaying tissue. Some scientists claim that maggot mass temperature is generally significantly above ambient air temperature. Others maintain that while maggot mass temperature may be elevated in a laboratory setting, in the field the maggots self-regulate their temperature by moving in and out of the feeding mass. Obviously, more research is needed to determine the general effects of maggot mass temperature.
Other work has been reported concerning tanning in S. bullata with the age pigment lipofuscin
Lipofuscin
Lipofuscin is the name given to finely granular yellow-brown pigment granules composed of lipid-containing residues of lysosomal digestion. It is considered one of the aging or "wear-and-tear" pigments, found in the liver, kidney, heart muscle, adrenals, nerve cells, and ganglion cells...
. Lipofuscin accumulation in larvae is a linear function of time. Some of the pigment is disposed of at the pupal stage and then linear accumulation resumes in adults. Many projects have focused on an assay of this pigment as it could hold a rather accurate technique for aging S. bullata and other forensically important insects.
Temperature data
To estimate insect age, forensic entomologists must determine the amount of time a particular insect spends in each stage of development and apply this information to temperature data at a crime scene to determine a post-mortem interval estimation. Timing of various insect stages is determined in a laboratory at standardized temperatures so that ranges may be applied to the different temperature conditions encountered in the field. Post-mortem intervals are estimated by determining the accumulated degree days (ADD) or accumulated degree hours (ADH) acquired by each insect stage on the remains; this information is then used to calculate the overall degree days or degree hours accumulated during the insect’s entire life cycle and association with the carcass, representing the minimal amount of time the body has been available for colonization by insects. This estimation does not claim to represent an accurate time of death, but rather a minimal time of colonization. This is an important distinction that must be considered with the use of forensic evidence in courts of law.Accumulated degree days can be determined by subtracting the minimum threshold from the average ambient temperature over a 24-hour period and multiplying the difference by the corresponding number of days. Accumulated degree hours, which are equally useful, are similarly determined by subtracting the minimum threshold from the ambient temperature over a 1-hour period and multiplying the difference by the corresponding number of hours. The term "minimum threshold" refers to the lowest temperature at which an insect species will develop; any temperatures below this threshold will result in delayed development and probable death of the insect in question.
It is important to note that similar calculations may be carried out to determine “accumulated degree minutes,” “accumulated degree seconds,” etc., though ADD and ADH are typically used in forensic analyses.
For S. bullata, the following temperature data has been determined in a laboratory setting. These calculations correspond to a standardized temperature of 27 °C.
| Stage | Time Spent in Stage (27 °C) |
|---|---|
| Egg | --- |
| 1st Instar Larvae | 26 hrs |
| 2nd Instar Larvae | 18 hrs |
| 3rd Instar Larvae | 54 hrs |
| Prepupae | 112 hrs |
| Pupae | 12 days |
| Total | 17 days |
Other research
Another study was conducted concerning an effect by the mothers of the flesh fly population. There is a maternal effect that operates in S. bullata that can prevent pupal diapauseDiapause
Diapause is the delay in development in response to regularly and recurring periods of adverse environmental conditions. It is considered to be a physiological state of dormancy with very specific initiating and inhibiting conditions...
when reared under short day conditions. It was determined that certain environmental stressors or chemical treatments on the mother, such as extreme temperature shocks, food deprivation and reducing maternal size, were ineffective in altering the diapause-suppressant. However, there were several chemical agents that proved effective. Gamma-aminobutyric acid
Gamma-aminobutyric acid
γ-Aminobutyric acid is the chief inhibitory neurotransmitter in the mammalian central nervous system. It plays a role in regulating neuronal excitability throughout the nervous system...
, octopamine
Octopamine
Octopamine is an endogenous biogenic amine that is closely related to norepinephrine, and has effects on the adrenergic and dopaminergic systems. It is also found naturally in numerous plants, including bitter orange. Biosynthesis of the D--enantiomer of octopamine is by β-hydroxylation of...
and pilocarpine
Pilocarpine
Pilocarpine is a parasympathomimetic alkaloid obtained from the leaves of tropical American shrubs from the genus Pilocarpus. It is a non-selective muscarinic receptor agonist in the parasympathetic nervous system, which acts therapeutically at the muscarinic acetylcholine receptor M3 due to its...
all countered the maternal affect and permitted the expression of diapause in the flesh fly progeny. The ability to express diapause is an important consideration for forensic entomologists as diapause allows the pupae to become metabollically "dormant" until environmental conditions improve; this would significantly affect pupal time estimations and subsequent post-mortem interval estimations.
External links
- Pape, T. & Carlberg, U. (2001 onwards), A pictorial guide to the Sarcophagidae of the world (Insecta: Diptera).
- "Sarcophagidae Central" Overview,Identification Annotated bibliography etc.
- The Definitive website Thomas Pape
- Flesh Fly: Sarcophaga sp. Diagnostic photographs, descriptions and information; female specimen
- Flesh Fly: Bellieria sp. Photographs, descriptions and information; female specimen
- Family description and images
- Images from Diptera info
- Fly morphology
- Key to the Calyptrate families