Calliphora vomitoria | |
---|---|
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Diptera |
Family: | Calliphoridae |
Genus: | Calliphora |
Species: | C. vomitoria |
Binomial name | |
Calliphora vomitoria | |
Synonyms [1] [2] | |
|
Calliphora vomitoria, known as the blue bottle fly, [3] orange-bearded blue bottle, [4] or bottlebee, is a species of blow fly, a species in the family Calliphoridae. Calliphora vomitoria is the type species of the genus Calliphora . It is common throughout many continents including Europe, Americas, and Africa. They are fairly large flies, nearly twice the size of the housefly, with a metallic blue abdomen and long orange setae on the gena.
While adult flies feed on nectar, females deposit their eggs on rotting corpses, making them important forensic insects, as their eggs and timing of oviposition can be used to estimate time of death.
Blue bottle flies are typically 10–14 mm (3⁄8–9⁄16 in) long, almost twice the size of a housefly. The head and thorax are dull gray, and the back of the head has long yellow-orange setae. [5] [6] The abdomen is bright metallic blue with black markings. Its body and legs are covered with black bristly hairs. It has short, aristate antennae and four tarsi per leg. The eyes are red and the wings are transparent. The legs and antennae are black and pink. The chest is bright purple and has spikes for protection from other flies. [7] [8] To differentiate C. vomitoria from other closely related species such as Calliphora vicina , C. vomitoria can be identified by characteristic "orange cheeks", which are the orange hairs below the eyes. Additionally, C. vomitoria has a dark basicosta (base of the wing) while C. vicina has a yellow basicosta. All these characteristics can be identified through a simple photograph. [9] [10]
Calliphora vomitoria can be found throughout the world, including most of Europe, Alaska, Greenland, the south of Mexico, United States, and southern Africa. [11] [12] It prefers higher elevations relative to other Calliphoridae species, such as Lucilia sericata and Chrysomya albiceps . They are among the most abundant flies found in these regions. [13]
Temperature has a significant effect on distribution. As is the case with most flies, C. vomitoria are found most abundantly during spring and summer, and least abundant during fall and winter. [14] The preferred habitat of C. vomitoria varies depending on the season. During winter and summer, they can be found mostly in rural areas (and riparian areas to a lesser extent). During spring and fall, they are found in riparian areas. [15]
Blue bottle flies have the complete cycle of egg, larva, pupa, and adult. Development usually takes around 2 weeks. [16] Larvae are protein-rich and can theoretically be used as feed. A female blue bottle fly lays her eggs where she feeds, usually in decaying meat, garbage, or feces. Pale whitish larvae, commonly called maggots, soon hatch from the eggs and immediately begin feeding on carcasses of dead animals and on the decomposing matter where they were hatched. [17] After a few days of feeding, they are fully grown. At that time they crawl away to a drier place where they burrow into soil or similar matter and pupate into tough brown cocoons. The pupal stage is the longest stage of the development cycle. [14]
After two or three weeks, the adults emerge from the pupal stage to mate, beginning the cycle again. The normal duration spent in adult form averages 10–14 days, however, during cold weather, pupae and adults can hibernate until higher temperatures revive them. [8]
Undergoing metamorphosis requires a tremendous amount of change for the fly, such as cell death. While it is commonly believed that programmed cell death and apoptosis are the same, they are not always so. At the beginning of metamorphosis during the larvae stage, salivary gland cells of Calliphora vomitoria larvae are programmed to self-destruct. After enough feeding, the larvae come to rest and an initial protein synthesis stage occurs, culminating in the production of high amounts of protein. This occurs from day 1 to about day 8. Then, on day 9, cell death of salivary gland cells occurs. This pattern of synthesis and destruction is not to be confused with apoptosis, as no DNA degeneration is seen and cells are shown to vacuolate and swell (instead of condensing and shrinking, as in the case of apoptosis). Instead, selective expression and DNA synthesis occur during programmed cell death of salivary gland cells. [18]
Like other blowflies, C. vomitoria colonize animal remains, including humans. While adult C. vomitoria feed on nectar, the larvae feed on corpses, the medium in which they grow. However, it has been shown that feeding on processed substrates (food that are modified for human consumption by increasing shelf life and taste through salting, curing, smoking, etc) provided much better growth than unprocessed substrates such as raw unmodified liver. Because different substrates drastically affected growth, C. vomitoria is best characterized as a specialist that best utilizes processed substrates (minced meats, for example). Its close relative, Calliphora vicina , is a generalist, being able to utilize mixed substrates with equal growth rates. [19] In the case of overcrowding, C. vomitoria competition results in compensation by increased speed of development, leading to smaller larvae and adults. This has complications in forensics because different parts of the body would grow at different rates. [20] Additionally, it has been shown that the fly larvae are able to colonize even buried remains. Growth rates are similar between surface and buried larvae. [21] Usually, these flies lay their eggs around wounds on fresh corpses shortly after death. Right before the pupal stage, the fly larvae that leaves the carrion can burrow into the soil in order to pupate. Then, adult flies emerge. [14] In decaying carcasses, it was found that Calliphoridae flies dominate, especially C. vomitoria. In both spring and fall, C. vomitoria is the primary species found on carcasses. In some cases, C. vomitoria shares carcasses with other calliphorid species such as Lucilia caesar. [22]
Bluebottle fly adults feed on nectar, and they are pollinators of flowers. They are especially attracted to flowers that have strong odors, such as those that have adapted to smell like rotting meat. Plants pollinated by the fly include the skunk cabbage ( Symplocarpus foetidus ), American pawpaw ( Asimina triloba ), dead horse arum ( Helicodiceros muscivorus ), goldenrod and some species of the carrot family. [23] These insects tend to fly in packs in order to detect possible food sources more efficiently. If one fly detects food, it disperses a pheromone, which will alert the others to the meal. [8]
Blow flies like C. vomitoria lay their eggs at carrion sites, which are scarce in most places so these corpses end up with many eggs of various species. As a result, high larval density arises. In fact, when there are many other individuals around the site, pregnant females increase oviposition rate (which increases number of offspring), likely triggered by contact and chemical stimulation. [24] However, the large number of larvae ends up being beneficial for each individual. The larvae feed by secretion of enzymes that break down tissues of the corpse, so by aggregating in large numbers these secretions are more effective, leading to easier feeding. Additionally, the large aggregation helps generate heat and keep the larvae warm, as the flies generally prefer warmer temperature. One complication with the high number of individuals is that competition is still a factor, as larvae on the periphery may be left out of the feeding, and by the end of the developmental cycle they emerge undernourished and undersized. [20]
It has been suggested that C. vomitoria rarely fly at night, regardless of the presence of an existing corpse. They thus may not deposit eggs on corpses during the night. This is relevant for forensic science, as the approximate time of oviposition would be during the daytime. [25]
The median neuro-secretory cells (MNC) of the brain of Calliphora species contain peptide hormones that resemble insulin. This was proven when researchers were able to bind these insulin-like peptides with antibodies of bovine insulin. This shows that an insect hormone can be structurally analogous to a prominent mammalian hormone [20] and it raises the possibility of these insulin-like or polypeptide-like materials serving as central nervous system regulatory hormones before they were metabolic regulatory hormones. [26] [27]
On the terminal region of the 5th tarsal segment, the C. vomitoria contain pulvilli, which are the cushion-like hairy feet on insects and many arthropods located at base of their two claws. The hair that project from the ventral surface is the key for the adhesion abilities of these flies. Additionally, they have large claws that help to hold on to irregular surfaces to prevent falling. Calliphora vomitoria, like other blowflies, also secrete non-volatile lipids through the hairs that are important for further adhesion. By a combination of the physical grip of the claws and hairs and the surface tension created by the lipid secretions, they are able to adhere to smooth surfaces with ease. [28] [29]
These flies are among the most important insect evidence in forensic science, specifically for obtaining time of colonization (TOC) and post mortem interval (PMI). [14] Calliphora species are the most important in temperate regions because of their growth rate in accordance to temperature. By knowing the temperature, the amount of time since the eggs were laid can be estimated. In addition, C. vomitoria has higher threshold temperature for growth than many species; likewise, it is present in many regions. There is a limit to their usage, though, as few species can survive in cold temperatures; most cannot continue development unless it is warmer than roughly 2 °C (36 °F). [30]
Degradation of carcasses can be divided into six separate stages: stage of decomposition, fresh stage, bloated stage, active decay stage, advanced decay stage, and remains stage. Adult C. vomitoria first starts to appear at carcasses during the bloated stage, followed by larvae 1 to 3 days after. During the active decay stage, the blow fly larvae population reaches its peak. [31]
In buried corpses, information of time since burial and how the body was kept (above/below ground before burial) can also be collected through the identification of C. vomitoria. [21] The study of these flies, however, is limited to areas where entomologists are readily available, as life histories can differ in separate regions. These life histories differ in subtle ways due to differences in climate such as temperature and elevation. These restrictions should be thus considered so the proper time of colonization (TOC) and post mortem interval (PMI) can be established. [14]
This bluebottle fly can also cause human or animal myiasis (parasitization in a living individual). Forensic scientists sometimes identify it in the course of their work, such as in one case of an autopsy of a neglected child. [32] [33]
Calliphora vomitoria is often not the only species present at carrion, so some process of identification of the correct species is needed in order to avoid false estimates of the time of death due to their having different developmental cycles. In the past, simple morphological differences are used to differentiate between species. However, it is very difficult in crime scenes because more often than not these sites are not ideal, with preservation of insect species far from good. Methods that can best differentiate between the species are DNA, mitochondrial DNA, and the COI gene. The COI gene used in conjunction with restriction enzymes has been shown to be a relatively fast and simple method of distinguishing between blowfly species with good accuracy. [34]
Post mortem interval (PMI) is the time between death and discovery of a corpse. Calliphora vomitoria is important for PMI estimations because it is among the first species to lay eggs on the corpse. There are two ways of estimating PMI. One is killing the larvae, and then comparing the larvae's length and temperature to those in the standardized data. Another way to calculate PMI is to calculate the accumulated degree hours/days (ADH/D) that a larva needs to reach a certain developmental stage. The later method is the more widely accepted way to estimate PMI. [20]
As one of the most abundant flies and their tendency to be first on the case (carrion), they are very useful in legal investigations. Other Calliphora species, while important as parasites of humans, are not as important simply because they are less often found. However, there is not a clear consensus on fly distribution, as different areas attract different species of flies, and so field research should be conducted in local areas to confirm the presence or absence of these important forensic resources. [15]
Calliphora vomitoria can sometimes pollinate crops, working especially well with strongly scented crops. However, it can also transmit pathogenic bacteria such as Xanthomonas campestris pv. campestris to flowers, resulting in infected seeds. [35]
Forensic entomology is a branch of forensic science that uses insects found on corpses to help solve criminal cases. This includes studying the types of insects commonly found on cadavers, their life cycles, their presence in different environments, and how insect assemblages change with decomposition.
The Calliphoridae are a family of insects in the order Diptera, with almost 1,900 known species. The maggot larvae, often used as fishing bait, are known as gentles. The family is known to be polyphyletic, but much remains disputed regarding proper treatment of the constituent taxa, some of which are occasionally accorded family status.
The common green bottle fly is a blowfly found in most areas of the world and is the most well-known of the numerous green bottle fly species. Its body is 10–14 mm (0.39–0.55 in) in length – slightly larger than a house fly – and has brilliant, metallic, blue-green or golden coloration with black markings. It has short, sparse, black bristles (setae) and three cross-grooves on the thorax. The wings are clear with light brown veins, and the legs and antennae are black. The larvae of the fly may be used for maggot therapy, are commonly used in forensic entomology, and can be the cause of myiasis in livestock and pets. The common green bottle fly emerges in the spring for mating.
Chrysomya putoria, also known as the tropical African latrine blowfly, is a fly species belonging to the blowfly family, Calliphoridae.C. putoria is native to Africa and has recently spread to the Americas. These flies pose significant health risks, especially due to their close association with human settlements. Adult flies can carry pathogens, while larvae may cause myiasis by growing and feeding on the flesh of domestic animals and humans. Other myiasis-causing flies in the same genus are C. bezziana and C. megacephala. C. putoria and other flies that feed on decomposing tissue are used as important tools in forensic entomology to establish the post-mortem interval, or the time elapsed since death.
Cynomya mortuorum belongs to the order Diptera, sometimes referred to as "true flies". In English, the only common name occasionally used is "fly of the dead". It has a bluish-green appearance, similar to other Calliphoridae and is found in multiple geographic locations with a preference for colder regions. Belonging to the family Calliphoridae, it has been shown to have forensically relevant implications due to its appearance on carrion. Current research is being done to determine C. mortuorum's level of importance and usage within forensic entomology.
Forensic entomological decomposition is how insects decompose and what that means for timing and information in criminal investigations. Medicolegal entomology is a branch of forensic entomology that applies the study of insects to criminal investigations, and is commonly used in death investigations for estimating the post-mortem interval (PMI). One method of obtaining this estimate uses the time and pattern of arthropod colonization. This method will provide an estimation of the period of insect activity, which may or may not correlate exactly with the time of death. While insect successional data may not provide as accurate an estimate during the early stages of decomposition as developmental data, it is applicable for later decompositional stages and can be accurate for periods up to a few years.
Calliphora vicina is a member of the family Calliphoridae, which includes blow flies and bottle flies. These flies are important in the field of forensic entomology, being used to estimate the time of a person's death when a corpse is found and then examined. C. vicina is currently one of the most entomologically important fly species for this purpose because it arrives at and colonizes a body following death in consistent timeframes.
Phormia regina, the black blow fly, belongs to the blow fly family Calliphoridae and was first described by Johann Wilhelm Meigen.
Chrysomya villeneuvi, or hairy maggot, is a South East Asian fly species of forensic importance because the maggots of this species have been collected from human corpses.
Chrysomya megacephala, more commonly known as the oriental latrine fly or oriental blue fly, is a member of the family Calliphoridae (blowflies). It is a warm-weather fly with a greenish-blue metallic box-like body. The fly infests corpses soon after death, making it important to forensic science. This fly is implicated in some public health issues; it can be the cause of myiasis, and also infects fish and livestock.
Compsomyiops callipes, previously known as Paraluclia wheeleri, is a member of the blowfly family Calliphoridae. It is a warm weather fly that can be found in southwestern parts of the United States and parts of South America. This species can be identified by its chaetotaxy, metallic blue color, club-shaped palp, and brown calypters.
Cynomya cadaverina, also known as the shiny blue bottle fly, is a member of the family Calliphoridae, which includes blow flies as well as bottle flies. In recent years, this family has become a forensically important facet in many medicocriminal investigations in the growing field of forensic entomology. C. cadaverina is specifically important in determining a post-mortem interval, as well as other important factors.
Calliphora livida is a member of the family Calliphoridae, the blow flies. This large family includes the genus Calliphora, the "blue bottle flies". This genus is important in the field of forensic entomology because of its value in post-mortem interval estimation.
Lucilia coeruleiviridis, formerly Phaenecia coeruleiviridis, is commonly known as a green bottle fly, because of its metallic blue-green thorax and abdomen. L. coeruleiviridis was first discovered by French entomologist Pierre-Justin-Marie Macquart in 1855. It belongs to the family Calliphoridae and is one of many forensically important Diptera, as it is often found on decaying substances. L. coeruleiviridis is one of the most ubiquitous blow fly species in the southeastern United States, particularly in the spring and fall months.
Protophormia terraenovae is commonly called northern blowfly, blue-bottle fly or blue-assed fly. It is distinguished by its deep blue coloration and large size and is an important species throughout the Northern Hemisphere. This fly is notable for its economic effect as a myiasis pest of livestock and its antibiotic benefits in maggot therapy. Also of interest is P. terraenovae’s importance in forensic investigations: because of their temperature-dependent development and their prominent presence on corpses, the larvae of this species are useful in minimum post-mortem interval (mPMI) determination.
Calliphora latifrons is a species of blue bottle fly.
Calliphora loewi is part of the family Calliphoridae, which consisted of bottle flies and blowflies, and in the genus Calliphora, blue bottle flies. The genus can be deceiving since C. loewi is not blue. Though this species is rare, it can play an important part in forensic entomology, spreading disease, and decomposing carrion. The life cycle of C. loewi is similar to the life cycle of the genus Calliphora. Since this species is rare there has not been much research on it.
Carrion insects are insects associated with decomposing remains. The processes of decomposition begin within a few minutes of death. Decomposing remains offer a temporary, changing site of concentrated resources which are exploited by a wide range of organisms, of which arthropods are often the first to arrive and the predominant exploitive group. However, not all arthropods found on or near decomposing remains will have an active role in the decay process.
Calliphora stygia, commonly known as the brown blowfly, or rango tumaro in Māori, is a species of blow-fly that is found in Australia and New Zealand. The brown blowfly has a grey thorax and yellow-brown abdomen.
Calliphora quadrimaculata, commonly known as the New Zealand blue blowfly and by its Māori name rango pango, is an insect in the genus Calliphora of family Calliphoridae in the order Diptera. This particular blowfly is found throughout New Zealand as well as on Chatham, Auckland, Stewart and Campbell Islands that surround New Zealand. Generally blowfly maggots in New Zealand have to feed on animal tissue or faeces to develop into adult blowflies. However the New Zealand blue blowfly larvae can survive on decaying leaves of snow tussock in alpine regions and reach adult maturity without feeding on any animal tissue.