Necrophage

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A shrew corpse (Soricidae) surrounded by multiple necrophages, including members of the Orders Diptera and Coleoptera. Carronneros1.jpg
A shrew corpse ( Soricidae ) surrounded by multiple necrophages, including members of the Orders Diptera and Coleoptera.

Necrophages are organisms that obtain nutrients by consuming decomposing dead animal biomass, such as the muscle and soft tissue of carcasses and corpses. [1] [2] [3] The term derives from Greek nekros, meaning 'dead', and phagein, meaning 'to eat.' [1] Mainly, necrophages are species within the phylum Arthropoda; however, other animals, such as gastropods and Accipitrimorphae birds have been noted to engage in necrophagy. [2] [4]

Contents

Necrophages play a critical role in the study of forensic entomology, as certain Arthropoda, such as Diptera larvae, engage in myiasis and colonization of the human body. [2]

Invertebrates

Diptera

Members of the order Diptera, such as Nematocera, Calliphoridae, Sacrophagidae, and Muscidae, as well as semi-aquatic Diptera larvae, such as Simuliidae and Chironomidae, are the most common necrophages within the Animalia kingdom. [5] [2] Diptera species play a critical role in forensic entomology, as they tend to colonize the human body during the early floating phase of decomposition. [5] The flies utilize the submerged corpse as a source of food as well as an attachment site. Notably, Diptera do not specifically colonize and feed on human carcasses. [6] Diptera species, such as Musca domestica and Chloroprocta idioidea have been observed feeding on the carcasses of other mammalian carcasses, including the Mona monkey, the European rabbit, and the Giant cane rat, as well as fish carrion. [6] [7] [8] The carcass' appeal is characterized by the putridness of the odour it emits; thus, the olfactory system of Diptera species plays a role in their food selectivity. [9] In addition, the diversity and abundance of Diptera species vary both spatially and temporally. [10] Necrophagous Diptera, such as Calliphora vicina, tend to be concentrated in urban areas and rare in more rural areas. However, some researchers oppose this notion and claim anthropogenic impacts are negligible regarding species richness. [9] Temporally, the necrophagous Diptera are observed in higher abundances in the summer season than the winter season. The presence of specific Diptera abundance depends on the reproductive strategy retained by the family. [9] For example, Calliphoridae have a high larval growth rate accompanied by a low survival rate, while Sarcophagidae produce little offspring with longer life cycles. The latter is said to benefit necrophagous Diptera in particular environments, such as forests and urbanized areas.

Hymenoptera

Particular Hymenoptera, such as members of the genus Trigona are obligate necrophages. [3] [11] Trigona worker bees play a similar role to the Apis genus; however, along with collecting pollen, nectar, and plant resins, Trigona workers also collect carrion from vertebrate carcasses. [3] Although pollen is associated with higher energy value, dead tissue from vertebrate carcasses is preferred by Trigona bees because it is biochemically easier to extract energy from. [11] The dead animal tissue and muscle replace pollen as the primary protein source. Cerumen pots are utilized by some Trigona species, such as T. necrophaga, as vesicles to store foodstuff. [12] The foodstuff of T. necrophaga consists of both honey and carrion from vertebrate carcasses. [3] Ultimately, the stored food is utilized by developing larvae and the worker bee itself as a source of nutrition and energy. Due to the rapid decomposition of carrion, especially in warm temperatures, the bees must efficiently metabolize the carrion to avoid rotten carrion in their cerumen pots. [3] Trigona hypogea communicate the presence of a valuable carcass through olfactory signals. [3] The bees create an odour trail between their nest and the prospective animal carcass; thus, the bees recruit the other nest members to respond and exploit the corpse's resources rapidly. Additionally, interspecific competition is observed in Trigona hypogea bees. The bees are observed to defend their colonized food item, including but not limited to a monkey, lizard, fish, or snake carcass, from competing necrophages, such as flies.

Coleoptera

The larval stage of the Dytiscidae life cycle (left), and the adult stage (right). Dytiscidae life stages.jpg
The larval stage of the Dytiscidae life cycle (left), and the adult stage (right).

Coleoptera, specifically Cleridae, Dytiscidae, Scarabaeidae, Hydrophilidae, are common necrophages and, like Diptera, tend to play critical roles in forensic entomology. [6] [5] [13] [14] Dytiscidae are aquatic in both the adult and larval stages of their life cycles; thus, the beetles play a role in colonizing submerged human corpses. [5] Through colonization, the beetles assume a predacious role and feed on the dead tissue of the body. Dytiscidae species, such as Rhantus validus, retain mouthparts characterized by curved, asymmetrical and highly sclerotized mandibles. The mouthparts have a cutting edge and a groove that allows the insect to release digestive enzymes into its prey item and maxillae with sharp teeth. Researchers matched the post-mortem skin injuries of a human corpse to the mouthparts of such beetles and, in doing so, revealed necrophagous activity in the dermis and epidermis. Notably, the necrophagy elicited by Rhantus validus also created microhabitats for other, smaller necrophages by allowing access to freshly dead internal tissue. Necrophagy amongst Coleoptera is not confined to mammalian carcasses. [13] Necrophagous activity has been observed in Scarabaeidae species, such as Scybalocanthon nigriceps and on the fresh carcasses of Tree frogs. The beetle is observed to use its front legs and clypeus to shape the frog carrion into pellets for eventual consumption. In aquatic environments, Dytiscidae and Hydrophilidae species have also been observed to engage in the necrophagous activity of Granular toads. [14] Necrophagous beetles like Coprophanaeus ensifer may also build their burrows near carcasses for easier transportation of pieces of carrion into the burrow. [15]

Gastropoda

Nassariidae, such as Nassarius festivus and Nassarius clarus scavenge upon dead or decaying animal matter in the intertidal zone of eulittoral soft shores. [4] [16] On a sandflat in Monkey Mia in the World Heritage Site of Shark Bay, Australia, Nassarius clarus acts as a necrophage and feeds on the carrion of fishes and bivalves. [16] In the presence of carrion, the animal's proboscis performs a search reaction followed by a quick onset of feeding. When faced with a competitor, such as a hermit crab, at the site of the carrion, the Nassarius clarus attack the competition to defend their meal. Nassarius clarus are attracted to fish and bivalve carrion to a distance of 26 miles and have a heightened interest in areas where the sand has been disturbed; thus, indicating the potential presence of organic detritus or damaged fauna.

Vertebrates

Egyptian vultures (Neophron percnopterus) surrounding a mammalian carcass. 20191213 Neophron percnopterus, Jor Beed Bird Sanctuary, Bikaner 0926 8272.jpg
Egyptian vultures ( Neophron percnopterus ) surrounding a mammalian carcass.

Accipitrimorphae

Necrophagy has been observed in members of the clade Accipitrimorphae, including the Egyptian vulture, Eurasian griffon, Cinereous vulture, Black vulture, Turkey vulture, and the King vulture. [17] The birds, mainly vultures, have been noted to feed on the fleshy tissue and muscle of mammalian vertebrates, such as cows, pigs, and rabbits, as well as other birds. [17] [18] Additionally, the preyed upon carcasses have been recorded to be naturally deceased or the product of anthropogenic events, such as roadkill; thus, the means of prey retrieval may differ depending on spatial circumstances, such as the urbanization of a particular area. [18] Notably, anthropogenic impacts have had adverse impacts on the biological parameters of necrophagous birds, specifically in the territory of the Azerbaijan Republic. [19] The adverse effects include but are not limited to shortage of food, shootings of birds and nests, and removal of nestlings from nests. Due to the advances of cattle-breeding, involving indoor breeding and the utilization of dead cattle, many necrophagous birds, such as the Eurasian griffon, are losing access to nutritionally valuable cow carcasses. Therefore, the feeding ecology and the interspecific relationships of necrophagous birds are both directly and indirectly impacted by humans. [19] [18]

Role in Forensic Entomology

A case of human breast myiasis initiated by the Australian sheep blowfly (Lucila cuprina). Lucilia cuprina - A case of human breast myiasis - 01.png
A case of human breast myiasis initiated by the Australian sheep blowfly ( Lucila cuprina ).
Australian sheep blowfly (Lucilla cuprina) adult and larvae removed from a colonized human breast. Lucilia cuprina - A case of human breast myiasis - 02.png
Australian sheep blowfly ( Lucilla cuprina ) adult and larvae removed from a colonized human breast.

Necrophagous Diptera and Coleoptera play vital roles in the applications of forensic entomology. [5] Necrophagy is critical to forensic scientists because of the production of post-mortem changes on human corpses through myiasis. [5] [20] For example, a forensic medical examiner must determine which soft tissues have been removed as a result of necrophages post-mortem. [5] Entomological data can be utilized in homicide cases to determine the post-mortem interval (PMI) and localize the crime. [21] This information is indicated by the time it takes the necrophage larvae to reach a particular developmental stage; thus, the life cycle of the necrophage provides detail regarding the interval between the initial myiasis and the discovery of the corpse. Notably, the colonization of a human corpse by Diptera necrophages is positively correlated with particular injuries, such as gunshot wounds and lacerations. The nature of these injuries provides higher levels of accessibility for the necrophagous Diptera and allows them to colonize the corpse rapidly. Seasonal changes, particularly temperature changes, significantly affect the abundance and degree of myiasis of necrophages. [22] [23] The increase or decrease in temperature that accompanies seasonal changes determines the rate of development retained by necrophages, particularly Diptera. [23] High temperatures lead to an exponential increase in the rate of development in Diptera. Therefore, the body temperature of a corpse is of utmost importance to necrophages, as they prefer fresh internal tissue and sexually thrive in warm environments. However, the corpse's temperature may have adverse effects on the determination of the post-mortem interval (PMI) because the necrophages' development may be rapid due to body's temperature and not the passage of time.

In addition, Coleoptera provide valuable information to forensic entomologists. [5] [24] Specifically, in the later stages of decomposition, Dermestidae and Cleridae have been recorded to colonize human corpses and provide insight regarding the post-mortem interval (PMI). [24] However, researchers note that there are spatial differences that affect the latency of necrophagous Coleoptera presence.

See also

Related Research Articles

<span class="mw-page-title-main">Forensic entomology</span> Application of insect and other arthropod biology to forensics

Forensic entomology is a field of forensic science that uses insects found on corpses to help solve criminal cases. This includes the study of insect types commonly associated with cadavers, their respective life cycles, their ecological presences in a given environment, as well as the changes in insect assemblage with the progression of decomposition. Insect succession patterns are identified based on the time a given species of insect spends in a given developmental stage, and how many generations have been produced since the insects introduction to a given food source. Insect development alongside environmental data such as temperature and vapor density, can be used to estimate the time since death, due to the fact that flying insects are attracted to a body immediately after death, determine any possible movement of the body after death, and the determination of antemortem trauma. The identification of postmortem interval to aid in death investigations is the primary scope of this scientific field. However, forensic entomology is not limited to homicides, it has also been used in cases of neglect and abuse, in toxicology contexts to detect the presence of drugs, and in dry shelf food contamination incidents. Equally, insect assemblages present on a body, can be used to approximate a given location, as certain insects may be unique to certain areas. Therefore, forensic entomology can be divided into three subfields: urban, stored-product and medico-legal/medico-criminal entomology.

<span class="mw-page-title-main">Silphidae</span> Family of beetles

Silphidae is a family of beetles that are known commonly as large carrion beetles, carrion beetles or burying beetles. There are two subfamilies: Silphinae and Nicrophorinae. Members of Nicrophorinae are sometimes known as burying beetles or sexton beetles. The number of species is relatively small, at around two hundred. They are more diverse in the temperate region although a few tropical endemics are known. Both subfamilies feed on decaying organic matter such as dead animals. The subfamilies differ in which uses parental care and which types of carcasses they prefer. Silphidae are considered to be of importance to forensic entomologists because when they are found on a decaying body they are used to help estimate a post-mortem interval.

<span class="mw-page-title-main">Maggot</span> Larva of a fly

A maggot is the larva of a fly ; 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.

<span class="mw-page-title-main">Common green bottle fly</span> Species of insect

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.

Vulture bees, also known as carrion bees, are a small group of three closely related South American stingless bee species in the genus Trigona which feed on rotting meat. Some vulture bees produce a substance similar to royal jelly which is not derived from nectar, but rather from protein-rich secretions of the bees' hypopharyngeal glands. These secretions are likely derived from the bees' diet, which consists of carrion eaten outside the nest, and resulted in the belief that they produce what is known as “meat honey”. This unusual behavior was only discovered in 1982, nearly two centuries after the bees were first classified.

Trigona hypogea is a species of stingless bee from the Neotropics; it is unusual in that it is one of only three known species of bee that exclusively uses carrion as a protein source, rather than pollen, earning it the nickname "vulture bee".

<i>Calliphora vomitoria</i> Species of fly

Calliphora vomitoria, known as the blue bottle fly, orange-bearded blue bottle, 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.

<i>Chrysomya</i> Genus of flies

Chrysomya is an Old World blow fly genus of the family Calliphoridae. The genus Chrysomya contains a number of species including Chrysomya rufifacies and Chrysomya megacephala. The term “Old World blow fly” is a derivative of both the associated family, Calliphoridae, and the belief that the genus Chrysomya originated in Asia and migrated to North America only relatively recently. Chrysomya’s primary importance to the field of medico-criminal forensic entomology is due to the genus’ reliable life cycle, allowing investigators to accurately develop a postmortem interval. Chrysomya adults are typically metallic colored with thick setae on the meron and plumose arista. The name comes from the word chrysos, meaning “golden” in reference to the metallic sheen of the genus’ species, and -mya, a derivation from the word myia, meaning “fly”.

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.

<i>Phormia regina</i> Species of fly

Phormia regina, the black blow fly, belongs to the blow fly family Calliphoridae and was first described by Johann Wilhelm Meigen.

In forensic entomology, entomotoxicology is the analysis of toxins in arthropods that feed on carrion. Using arthropods in a corpse or at a crime scene, investigators can determine whether toxins were present in a body at the time of death. This technique is a major advance in forensics; previously, such determinations were impossible in the case of severely decomposed bodies devoid of intoxicated tissue and bodily fluids. Ongoing research into the effects of toxins on arthropod development has also allowed better estimations of postmortem intervals.

<span class="mw-page-title-main">American carrion beetle</span> Species of beetle

The American carrion beetle is a North American beetle of the family Silphidae. It lays its eggs in, and its larvae consume, raw flesh and fungi. The larvae and adults also consume fly larvae and the larvae of other carrion beetles that compete for the same food sources as its larvae.

<i>Chrysomya albiceps</i> Species of fly

Chrysomya albiceps is a species belonging to the blow fly family, Calliphoridae.

<i>Chrysomya megacephala</i> Species of fly

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.

<i>Sarcophaga pernix</i> Species of fly

Sarcophaga pernix, 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. It is potentially useful in forensic entomology.

<i>Sarcophaga bullata</i> Species of fly

Sarcophaga bullata, or the grey flesh fly, is a species of fly belonging to the family Sarcophagidae. It varies in size from small to large, 8 to 17 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 family Sarcophagidae 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.

Lucilia thatuna belongs to the family Calliphoridae, the species most commonly referred to as the blowflies, and the genus Lucilia. Along with several other species of Lucilia, L. thatuna is commonly referred to as a green bottle fly. L. thatuna is very scarce and not much is known about this particular fly. It has been noted to reside in mountainous regions of the northwestern United States.

<i>Cynomya cadaverina</i> Species of fly

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.

<span class="mw-page-title-main">Carrion insects</span> Insects associated with decomposing remains

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.

<i>Necrodes littoralis</i> Species of beetle

Necrodes littoralis, also known as the short sexton beetle, is a species of carrion beetle of the genus Necrodes, found in countries across Europe. As a carrion beetle, it feeds on decaying vertebrate remains and maggots. This species' feeding behaviors make it an important asset to forensic entomology.

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