Forensic entomology
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Forensic entomology is the scientific study of the colonization of a dead body by
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s. This includes the study of insect types commonly associated with
cadavers A cadaver or corpse is a dead human body that is used by medical students, physicians and other scientists to study anatomy, identify disease sites, determine causes of death, and provide tissue to repair a defect in a living human being. Stud ...
, their respective life cycles, their ecological presences in a given environment, as well as the changes in insect assemblage with the progression of
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 ...
. 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. 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 Urban means "related to a city". In that sense, the term may refer to: * Urban area, geographical area distinct from rural areas * Urban culture, the culture of towns and cities Urban may also refer to: General * Urban (name), a list of people ...
, stored-product and medico-legal/medico-criminal entomology.


History

Historically, there have been several accounts of applications for, and experimentation with, forensic entomology. Aside from an initial case report in China from the 13th century, the primitive observation and correlation between arthropods and forensic contexts were documented in Germany and France. This was conducted during a mass exhumation in the late 1880s by Hofmann and Reinhard. However, only in the last 30 years has forensic entomology been systematically explored as a feasible source for evidence 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 preservat ...
s. Through their own experiments and interest in arthropods and death, Sung Tzu, Francesco Redi, Bergeret d’Arbois, Jean Pierre Mégnin and the physiologist Hermann Reinhard have helped to lay the foundations for today's modern forensic entomology.


Song Ci

The Song Dynasty (960–1279) forensic science book '' Collected Cases of Injustice Rectified'' published by court judge, physician, medical scientist and writer Song Ci in 1247 contains the oldest known case of forensic entomology.Haskell (2006), 432. In a murder case of 1235, a villager was stabbed to death and authorities determined that his wounds were inflicted by a
sickle A sickle, bagging hook, reaping-hook or grasshook is a single-handed agricultural tool designed with variously curved blades and typically used for harvesting, or reaping, grain crops or cutting Succulent plant, succulent forage chiefly for feed ...
; this was a tool used for cutting rice at harvest time, a fact which led them to suspect a fellow peasant worker was involved. The local magistrate had the villagers assemble in the town square where they would temporarily relinquish their sickles. Within minutes, a mass of blow flies gathered around one sickle and none other, attracted to the scent of traces of
blood Blood is a body fluid in the circulatory system of humans and other vertebrates that delivers necessary substances such as nutrients and oxygen to the cells, and transports metabolic waste products away from those same cells. Blood in the cir ...
unseen by the naked eye. It became apparent to all that the owner of that sickle was the culprit, the latter pleading for mercy as he was detained by authorities. Song Ci (sometimes referred to as Sung Tzu) was a judicial intendant who lived in China 1188-1251 AD. In 1247 AD Song Ci wrote a book entitled '' Washing Away of Wrongs'' as a handbook for coroners. In this book Song Ci depicts several cases in which he took notes on how a person died and elaborates on probable causes. He explains in detail on how to examine a corpse both before and after burial. He also explains the process of how to determine a probable cause of death. The main purpose of this book was to be used as a guide for other investigators so they could assess the scene of the crime effectively. His level of detail in explaining what he observed in all his cases laid down the fundamentals for modern forensic entomologists and is the first recorded account in history of someone using forensic entomology for judicial means.


Francesco Redi

In 1668, Italian physician Francesco Redi disproved the theory of
spontaneous generation Spontaneous generation is a superseded scientific theory that held that living creatures could arise from nonliving matter and that such processes were commonplace and regular. It was hypothesized that certain forms, such as fleas, could arise f ...
. The accepted theory of Redi's day claimed that
maggots 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. Entom ...
developed spontaneously from rotting meat. In an experiment, he used samples of rotting meat that were either fully exposed to the air, partially exposed to the air, or not exposed to air at all. Redi showed that both fully and partially exposed rotting meat developed fly
maggots 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. Entom ...
, whereas rotting meat that was not exposed to air did not develop maggots. This discovery completely changed the way people viewed the decomposition of organisms and prompted further investigations into insect life cycles and into entomology in general.


Bergeret d'Arbois

Dr. Louis François Etienne Bergeret (1814–1893) was a French hospital physician, and was the first to apply forensic entomology to a case. In a case report published in 1855 he stated a general life cycle for insects and made many assumptions about their mating habits. Nevertheless, these assumptions led him to the first application of forensic entomology in an estimation of
post-mortem interval The post-mortem interval (PMI) is the time that has elapsed since an individual's death. When the time of death is not known, the interval may be estimated, and so an approximate time of death established. Postmortem interval estimations can ra ...
(PMI). His report used forensic entomology as tool to prove his hypothesis on how and when the person had died.


Hermann Reinhard

The first systematic study in forensic entomology was conducted in 1881 by Hermann Reinhard, a German medical doctor who played a vital role in the history of forensic entomology. He exhumed many bodies and demonstrated that the development of many different types of insect species could be tied to buried bodies. Reinhard conducted his first study in east Germany, and collected many Phorid flies from this initial study. He also concluded that the development of only some of the insects living with corpses underground were associated with them, since there were 15-year-old beetles who had little direct contact with them. Reinhard's works and studies were used extensively in further forensic entomology studies.


Jean Pierre Mégnin

French veterinarian and entomologist Jean Pierre Mégnin (1828–1905), published many articles and books on various subjects including the books ''Faune des Tombeaux'' and ''La Faune des Cadavres'', which are considered to be among the most important forensic entomology books in history. In his second book he did revolutionary work on the theory of predictable waves, or successions of insects onto corpses. By counting numbers of live and dead mites that developed every 15 days and comparing this with his initial count on the infant, he was able to estimate how long that infant was dead. In this book, he asserted that exposed corpses were subject to eight successional waves, whereas buried corpses were only subject to two waves. Mégnin made many great discoveries that helped shed new light on many of the general characteristics of decaying flora and fauna. Mégnin's work and study of the larval and adult forms of insect families found in cadavers sparked the interest of future entomologists and encouraged more research in the link between arthropods and the deceased, and thereby helped to establish the scientific discipline of forensic entomology.


Forensic entomology subfields


Urban forensic entomology

Urban forensic entomology typically concerns pests infestations in buildings gardens or that may be the basis of litigation between private parties and service providers such as landlords or exterminators. Urban forensic entomology studies may also indicate the appropriateness of certain pesticide treatments and may also be used in stored products cases where it can help to determine chain of custody, when all points of possible infestation are examined in order to determine who is at fault.


Stored-product forensic entomology

Stored-product forensic entomology is often used in litigation over insect infestation or contamination of commercially distributed foods.


Medico-legal forensic entomology

Medicolegal forensic entomology covers evidence gathered through arthropod studies at the scenes of murder, suicide, rape, physical abuse and contraband trafficking. In murder investigations it deals with which
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three ...
s eggs appear, their location on the body and in what order they appear. This can be helpful in determining a
post mortem interval The post-mortem interval (PMI) is the time that has elapsed since an individual's death. When the time of death is not known, the interval may be estimated, and so an approximate time of death established. Postmortem interval estimations can ra ...
(PMI) and location of a
death Death is the irreversible cessation of all biological functions that sustain an organism. For organisms with a brain, death can also be defined as the irreversible cessation of functioning of the whole brain, including brainstem, and brain ...
in question. Since many insects exhibit a degree of
endemism Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found else ...
(occurring only in certain places), or have a well-defined phenology (active only at a certain season, or time of day), their presence in association with other evidence can demonstrate potential links to times and locations where other events may have occurred. This discipline also assists in helping associate a victim, suspect and scene together. This is possible due to the different insect species located in specific geographical locations. Another area covered by medicolegal forensic entomology is the relatively new field of entomotoxicology. This particular branch involves the utilization of entomological specimens found at a scene in order to test for different drugs that may have possibly played a role in the death of the victim. The analytical perspective behind these methods essentially relies upon the fact that the presence of drugs within the carcass has an effect on the growth and morphology of the insects subsequently ingesting the toxins from the corpse. However, due to these morphological alterations resulting on the insects in question, this can potentially lead to an erroneous postmortem interval interpretation when basing the PMI estimation on the specimens physical development. Entomology can also be of aid in medico-legal cases when depicting the site of an injury. This can be said due to the species preference of alimentation. When eggs laid on the corpse by blow flies subsequently hatch into first instar (first stage larvae), they require a liquid protein meal. However, due to their minute size and fragility they are unable to break through the individuals skin themselves to attain this nutrition. Therefore, the female ensures to lay her eggs near a wound site or natural orifice to provide blood, mucosal layer and body fluids that are accessible for an easier feed.


Myiasis

In cases of abandonment or neglect, the presence of
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 spe ...
may be noted. The latter can be described as blow flies colonizing a human or animal in life. In other words, it is the infestation of living vertebrate animals with dipteran larvae. Essentially, this insect succession will feed on the organisms living tissue, ingested food or liquid body substances. However, it is imperative to be cautious of the confusion myiasis may cause in a forensic context. This said, if the victim or found remains were colonized when alive, prior to being discovered, the entomological evidence would be indicating time of neglect or injury, rather than time of death.


Invertebrate types


Scorpionflies

Scorpionflies (order
Mecoptera Mecoptera (from the Greek: ''mecos'' = "long", ''ptera'' = "wings") is an order of insects in the superorder Endopterygota with about six hundred species in nine families worldwide. Mecopterans are sometimes called scorpionflies after their lar ...
) were the first insects to arrive at a donated human cadaver observed (by the entomologist Natalie Lindgren) at the Southeast Texas Applied Forensic Science Facility near
Huntsville, Texas Huntsville is a city in and the county seat of Walker County, Texas. The population was 45,941 as of the 2020 census. It is the center of the Huntsville micropolitan area. Huntsville is in the East Texas Piney Woods on Interstate 45 and home ...
, and remained on the corpse for one and a half days, outnumbering flies during that period. The presence of scorpionflies thus indicates that a body must be fresh.


Flies

Flies 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 ...
(order Diptera) are often first on the scene. They prefer a moist corpse for their offspring,
maggots 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. Entom ...
to feed on. The most significant types of fly include: *Blow flies – 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 ...
- Flies in this family are often metallic in appearance and between 10 and 14 mm in length. In addition to the name blow-fly, some members of this family are known as blue bottle fly, cluster flies, greenbottles, or black blowfly. A characteristic of the blow-fly is its 3-segmented antennae. Hatching from an egg to the first larval stage takes from eight hours to one day.
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. ...
e have three stages of development (called
instars 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 ...
); each stage is separated by a molting event. Molting can be defined as the process of new cubicle production while subsequently shedding the old cuticle. Larvae's ideal habitat in regard to pupation are locations providing access to loose, damp soil and litter. The latter consists of temperate and rather tropical areas. Worldwide, there are 1100 known species of blowflies, with 228 species in the
Neotropics The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In bioge ...
, and a large number of species in
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
and Southern
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a subcontinent of Eurasia and it is located entirel ...
. The most common area to find Calliphoridae species are in the countries of
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...
, Japan,
Central America Central America ( es, América Central or ) is a subregion of the Americas. Its boundaries are defined as bordering the United States to the north, Colombia to the south, the Caribbean Sea to the east, and the Pacific Ocean to the west. ...
, and in the southern United States. The forensic importance of this fly is that it is the first
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three ...
to come in contact with carrion because they have the ability to smell death from up to ten miles (16 km) away. Some prominent species of Calliphoridae are '' Calliphora vomitoria'' and '' Calliphora vicina.'' * Flesh flies – Family
Sarcophagidae Sarcophagidae () are a family of flies commonly known as flesh flies. They differ from most flies in that they are ovoviviparous, opportunistically depositing hatched or hatching maggots instead of eggs on carrion, dung, decaying material, or o ...
- Most flesh flies breed in carrion, dung, garbage, or decaying material, but a few species lay their eggs in the open wounds of mammals; hence their common name. Characteristics of the flesh-fly is its 3-segmented antennae. Most holarctic Sarcophagidae vary in size from 4 to 18 mm in length (Tropical species can be larger) with black and gray longitudinal stripes on the
thorax 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 cre ...
and checkering on the
abdomen The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the to ...
. Flesh-flies, being viviparous, frequently give birth to live young on corpses of human and other animals, at any stage of decomposition, from newly dead through to bloated or decaying (though the latter is more common). Sarcophaga barbata are specifically useful since they deposit maggots directly onto the decomposing body, their larger, visible size, and difference in activity during different stages. Their main limitation, however, is due to lack of information surrounding their geographic distribution and taxonomic features. * House fly – Family
Muscidae Muscidae are a family of flies found in the superfamily Muscoidea. Muscidae, some of which are commonly known as house flies or stable flies due to their synanthropy, are worldwide in distribution and contain almost 4,000 described species i ...
- is the most common of all
flies 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 ...
found in homes, and indeed one of the most widely distributed insects; it is often considered a pest that can carry serious diseases. The adults are 6–9 mm long. Their
thorax 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 cre ...
is gray, with four longitudinal dark lines on the back. The underside of their abdomen is yellow, and their whole body is covered with hair. Each female fly can lay up to 500 eggs in several batches of about 75 to 150
eggs Humans and human ancestors have scavenged and eaten animal eggs for millions of years. Humans in Southeast Asia had domesticated chickens and harvested their eggs for food by 1,500 BCE. The most widely consumed eggs are those of fowl, especial ...
. Genus '' Hydrotaea'' are of particular forensic importance. *Cheese flies – Family Piophilidae - Most are scavengers in animal products and fungi. The best-known member of the family is '' Piophila casei''. It is a small fly, about four mm (1/6 inch) long, found worldwide. This fly's larva infests cured meats, smoked fish, cheeses, and decaying animals and is sometimes called the cheese skipper for its leaping ability. Forensic entomology uses the presence of Piophila casei larvae to help estimate the date of death for human remains. They do not take up residence in a corpse until three to six months after death. The adult fly's body is black, blue-black, or bronze, with some yellow on the head, antennae, and legs. The wings are faintly iridescent and lie flat upon the fly's
abdomen The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the to ...
when at rest. At four mm (1/6 inch) long, the fly is one-third to one-half as long as the common housefly. *Coffin flies –
Phoridae The Phoridae are a family of small, hump-backed flies resembling fruit flies. Phorid flies can often be identified by their escape habit of running rapidly across a surface rather than taking to the wing. This behaviour is a source of one of thei ...
– a/k/a Humpbacked flies - Larvae feed on decaying bodies. Some species can burrow to a depth of 50 cm over 4 days. Important in buried bodies. *Lesser corpse flies –
Sphaeroceridae Sphaeroceridae are a family of true flies in the order Diptera, often called small dung flies, lesser dung flies or lesser corpse flies due to their saprophagous habits. They belong to the typical fly suborder Brachycera as can be seen by their ...
*Lesser house flies – Fanniidae *Black scavenger flies –
Sepsidae The Sepsidae are a family of flies, commonly called the black scavenger flies or ensign flies. Over 300 species are described worldwide. They are usually found around dung or decaying plant and animal material. Many species resemble ants, having ...
*Sun flies - Heleomyzidae *
Black soldier fly ''Hermetia illucens'', the black soldier fly, is a common and widespread fly of the family Stratiomyidae. Distribution This species is native to the Neotropical realm, but in recent decades has spread across all continents, becoming virtually ...
- Stratiomyidae - have potential for use in forensic entomology. The larvae are common
scavengers Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding b ...
in compost heaps, are found in association with carrion, can be destructive pests in honey bee hives, and are used in
manure Manure is organic matter that is used as organic fertilizer in agriculture. Most manure consists of animal feces; other sources include compost and green manure. Manures contribute to the Soil fertility, fertility of soil by adding organic ma ...
management (for both house fly control and reduction in manure volume). The larvae range in size from 1/8 to 3/4 of an inch (3 to 19 millimeters). The adult fly is a mimic, very close in size, color, and appearance to the
organ pipe mud dauber The organ pipe mud dauber (''Trypoxylon politum'') is a predatory wasp in the family Crabronidae. They are fairly large wasps, ranging from 3.9–5.1 cm, and have been recorded to fly from May to September. Female and male are similar in co ...
wasp and its relatives. *Non-biting midges -
Chironomidae The Chironomidae (informally known as chironomids, nonbiting midges, or lake flies) comprise a family of nematoceran flies with a global distribution. They are closely related to the Ceratopogonidae, Simuliidae, and Thaumaleidae. Many specie ...
- these flies have a complex life cycle. While adults are terrestrial and phytophagous, larvae are aquatic and detritivorous. Immature instars have been used as forensic markers in several cases where submerged corpses were found.


Beetles

Beetle Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
s (Order Coleoptera) are generally found on the corpse when it is more decomposed. In drier conditions, the beetles can be replaced by moth flies (
Psychodidae Psychodidae, called drain flies, sink flies, filter flies, sewer flies, or sewer gnats, is a family of Fly, true flies. Some genera have short, hairy bodies and wings giving them a "furry" moth-like appearance, hence one of their common names, m ...
). *Rove beetles – family Staphylinidae – are elongate beetles with small
elytra An elytron (; ; , ) is a modified, hardened forewing of beetles (Coleoptera), though a few of the true bugs (Hemiptera) such as the family Schizopteridae are extremely similar; in true bugs, the forewings are called hemelytra (sometimes alterna ...
(wing covers) and large jaws. Rove beetles have a four stage life cycle; egg, larvae, pupa and adult. Creophilus species are common predators of carrion, and since they are large, are a very visible component of the fauna of corpses. Some adult Staphylinidae are early visitors to a corpse, feeding on larvae of all species of fly, including the later predatory fly larvae. They lay their eggs in the corpse, and the emerging larvae are also predators. Some species have a long development time in the egg, and are common only during the later stages of decomposition. Staphylinids can also tear open the
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 thei ...
l cases of flies, to sustain themselves at a corpse for long periods. *Hister beetles – family
Histeridae Histeridae is a family of beetles commonly known as clown beetles or Hister beetles. This very diverse group of beetles contains 3,900 species found worldwide. They can be easily identified by their shortened elytra that leaves two of the seven ...
. Adult histerids are usually shiny beetles (black or metallic-green) which have an introverted head. The carrion-feeding species only become active at night when they enter the maggot-infested part of the corpse to capture and devour their maggot prey. During daylight they hide under the corpse unless it is sufficiently decayed to enable them to hide inside it. They have fast larval development with only two larval stages. Among the first beetles to arrive at a corpse are Histeridae of the genus '' Saprinus''. ''Saprinus'' adults feed on both the larvae and pupae of blowflies, although some have a preference for fresh pupae. The adults lay their eggs in the corpse, inhabiting it in the later stages of decay. *Carrion beetles – family
Silphidae 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. Nicrophorines are sometimes known as sexton beetles. The number of ...
- Adult Silphidae have an average size of about 12 mm. They are also referred to as burying beetles because they dig and bury small carcasses underground. Both parents tend to their young and exhibit communal breeding. The male carrion beetle's job in care is to provide protection for the breed and carcass from competitors. *Ham beetles – family Cleridae *Carcass beetles – family Trogidae * Skin/hide beetles – family
Dermestidae Dermestidae are a family of Coleoptera that are commonly referred to as skin beetles. Other common names include larder beetle, hide or leather beetles, carpet beetles, and khapra beetles. There are over 1,100 species described. Dermestids have ...
. Hide beetles are important in the final stages of decomposition of a carcass. The adults and larvae feed on the dried skin, tendons and bone left by fly larvae. Hide beetles are the only beetle with the enzymes necessary for breaking down
keratin Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. Alpha-keratin (α-keratin) is a type of keratin found in vertebrates. It is the key structural material making up scales, hair, nails, feathers, ho ...
, a protein component of hair. *Scarab beetles – family
Scarabaeidae The family Scarabaeidae, as currently defined, consists of over 30,000 species of beetles worldwide; they are often called scarabs or scarab beetles. The classification of this family has undergone significant change in recent years. Several sub ...
- Scarab beetles may be any one of around 30,000 beetle species worldwide that are compact, heavy-bodied and oval in shape. The flattened plates, which each antenna terminates, are fitted together to form a club. The outer edges of the front legs may also be toothed or scalloped. Scarab beetles range from in length. These species are known for being one of the heaviest insect species. *Sap beetles – family Nitidulidae


Mites

Many
mite Mites are small arachnids (eight-legged arthropods). Mites span two large orders of arachnids, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari, but genetic analysis does not show clear e ...
s (class
Acari Mites are small arachnids (eight-legged arthropods). Mites span two large orders of arachnids, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari, but genetic analysis does not show clear evid ...
, not insects) feed on corpses with ''Macrocheles'' mites common in the early stages of decomposition, while Tyroglyphidae and Oribatidae mites such as Rostrozetes feed on dry skin in the later stages of decomposition. '' Nicrophorus'' beetles often carry on their bodies the mite '' Poecilochirus'' which feed on fly eggs. If they arrive at the corpse before any fly eggs hatch into maggots, the first eggs are eaten and maggot development is delayed. This may lead to incorrect PMI estimates. ''Nicrophorus'' beetles find the
ammonia Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogenous wa ...
excretions of blowfly maggots toxic, and the ''Poecilochirus'' mites, by keeping the maggot population low, allow ''Nicrophorus'' to occupy the corpse.


Moths

Moth Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of w ...
s (order Lepidoptera) specifically clothes-moths – Family
Tineidae Tineidae is a family of moths in the order Lepidoptera described by Pierre André Latreille in 1810. Collectively, they are known as fungus moths or tineid moths. The family contains considerably more than 3,000 species in more than 300 genera. ...
– are closely related to butterflies. Most species of moth are nocturnal, but there are crepuscular and diurnal species. During their larval stages, clothes moths tend to feed on mammalian hair. They are amongst the final animals contributing to the decomposition of a corpse. This said, adult moths lay their legs on a carcass ''subsequently'' to fly larvae having had their presence on it.


Wasps, ants, and bees

Wasps, ants, and bees (order Hymenoptera) are not necessarily necrophagous. While some feed on the body, some are also predatory, and eat the insects feeding on the body. Thus meaning they are parasitoids (
Parasitoid wasp Parasitoid wasps are a large group of hymenopteran superfamilies, with all but the wood wasps (Orussoidea) being in the wasp-waisted Apocrita. As parasitoids, they lay their eggs on or in the bodies of other arthropods, sooner or later causin ...
). These Hymenoptera lay their eggs within the eggs or pupae of other insects; essentially causing the death of host insects. Wasps can also be of association to the family Pteromalidae. The latter can lay single or multiple eggs. They oviposit in pupae of muscoid flies (blow flies). Subsequently to the wasp eggs hatching, larvae will feed on the fly developing within the puparium; leading to its death. Bees and
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. ...
s have been seen feeding on the body during the early stages. This may cause problems for murder cases in which larval flies are used to estimate the post mortem interval since eggs and larvae on the body may have been consumed prior to the arrival on scene of investigators.


Factors


Moisture levels

Rain and
humidity Humidity is the concentration of water vapor present in the air. Water vapor, the gaseous state of water, is generally invisible to the human eye. Humidity indicates the likelihood for precipitation, dew, or fog to be present. Humidity dep ...
levels in the area where the body is found can affect the time for insect development. In most species, large amounts of rain will indirectly cause slower development due to drop in temperature. Light rain or a very humid environment, by acting as an insulator, will permit a greater core temperature within the maggot mass, resulting in faster development.


Submerged corpses

M. Lee Goff, a noted and well respected forensic entomologist, was assigned to a case involving the discovery of a decomposing body found on a boat half a mile from shore. Upon collection of the maggot mass, only one insect, ''
Chrysomya megacephala ''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 so ...
'', was discovered. He concluded that the water barrier accounted for the scarcity of other flies. He also noted that flies will not attempt to trek across large bodies of water unless there is a substantially influential attractant. In addition, the amount of time a maggot mass has been exposed to salt water can affect its development. From the cases Goff observed he found that if subjected for more than 30 minutes, there was a 24‑hour developmental delay. Not many more studies have been conducted and thus a specific amount of delay time is difficult to estimate.Goff, M. L. A Fly for the Prosecution. Cambridge, Massachusetts: Harvard University Press, 2000. The main focus of a study accomplished by Payne and King using fetal pigs, was the insect succession regarding carcass decomposition in an aquatic setting. Their results concluded that in the early floating stages of the cadaver, eggs were laid by blowflies. Moreover, by the bloating stage, most of the exposed flesh was absent and maggots migrated from the body. Many of the latter were present below the water line and fed on the carcass; with only their
Spiracle (arthropods) A spiracle or stigma is the opening in the exoskeletons of insects and some spiders to allow air to enter the trachea. In the respiratory system of insects, the tracheal tubes primarily deliver oxygen directly into the animals' tissues. The spi ...
protruding the surface.


Sun exposure

"Because insects are cold-blooded animals, their rate of development is more or less dependent on ambient temperature." Bodies exposed to large amounts of sunlight will heat up, giving the insects a warmer area to develop, reducing their development time. An experiment conducted by Bernard Greenberg and John Charles Kunich with the use of rabbit carcasses to study accumulation of degree days found that with temperature ranging in the mid 70s to high 80s the amount of developmental time for maggots was significantly reduced. In contrast, bodies found in shaded areas will be cooler, and insects will require longer growth periods. In addition, if temperatures reach extreme levels of cold, insects instinctively know to prolong their development time in order to hatch into a more accepting and viable climate in order to increase the chance of survival and reproduction.


Air exposure

Hanged bodies can be expected to show their own quantity and variety of flies. Also, the amount of time flies will stay on a hanged body will vary in comparison to one found on the ground. A hanged body is more exposed to air and thus will dry out faster, leaving less food source for the maggots. As the body begins to decompose, a compilation of fluids will leak to the ground. This area is where most of the expected fauna can be found. Also, it is more likely that rove beetles and other non-flying insects will be found here instead of directly on the body. Fly maggots, initially deposited on the body, may also be found below.


Geography

According to Jean Pierre Mégnin's book ''La Faune des Cadavres'' there are eight distinct faunal successions attracted to a corpse. While most
beetle Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
s and
flies 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 ...
of forensic importance can be found worldwide, a portion of them are limited to a specific range of habitats. It is forensically important to know the geographical distribution of these insects is order to determine information such as post mortem interval or whether a body has been moved from its original place of death.
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 ...
is arguably the most important family concerning forensic entomology given that they are the first to arrive on the corpse. The family can be found worldwide. ''Chrysomya rufifaces'', the hairy maggot blow fly, is a forensically important member of the family Calliphoridae and is widespread, however it is not prevalent in the Southern
California California is a state in the Western United States, located along the Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the most populous U.S. state and the 3rd largest by area. It is also the m ...
,
Arizona Arizona ( ; nv, Hoozdo Hahoodzo ; ood, Alĭ ṣonak ) is a state in the Southwestern United States. It is the 6th largest and the 14th most populous of the 50 states. Its capital and largest city is Phoenix. Arizona is part of the Fou ...
,
New Mexico ) , population_demonym = New Mexican ( es, Neomexicano, Neomejicano, Nuevo Mexicano) , seat = Santa Fe , LargestCity = Albuquerque , LargestMetro = Tiguex , OfficialLang = None , Languages = English, Spanish ( New Mexican), Navajo, Ke ...
,
Louisiana Louisiana , group=pronunciation (French: ''La Louisiane'') is a state in the Deep South and South Central regions of the United States. It is the 20th-smallest by area and the 25th most populous of the 50 U.S. states. Louisiana is borde ...
,
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
, or
Illinois Illinois ( ) is a state in the Midwestern United States. Its largest metropolitan areas include the Chicago metropolitan area, and the Metro East section, of Greater St. Louis. Other smaller metropolitan areas include, Peoria and Rockf ...
regions. Flesh flies fall under the family Sacrophagidae and generally arrive at a corpse following Calliphoridae. Unlike Calliphoridae, however, members of this family are able to fly in heavy rain. This key advantage enables them to occasionally reach a body before Calliphoridae, affecting the maggot mass that will be discovered. Flesh flies are globally distributed including habitats in the United States, Europe, Asia, and the Middle East. Beetles are representative of the order Coleoptera which accounts for the largest of the insect orders. Beetles are very adaptive and can be found in almost all environments with the exception of
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest cont ...
and high mountainous regions. The most diverse beetle fauna can be found in the tropics. In addition, beetles are less submissive to temperatures. Thus, if a carcass has been found in cold temperatures, the beetle will be prevalent over Calliphoridae.


Weather

Various weather conditions in a given amount of time cause certain pests to invade human households. This is because the insects are in search of food, water, and shelter. Damp weather causes reproduction and growth enhancement in many insect types, especially when coupled with warm temperatures. Most pests concerned at this time are
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of ...
s,
spider Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
s,
cricket Cricket is a bat-and-ball game played between two teams of eleven players on a field at the centre of which is a pitch with a wicket at each end, each comprising two bails balanced on three stumps. The batting side scores runs by str ...
s,
cockroach Cockroaches (or roaches) are a Paraphyly, paraphyletic group of insects belonging to Blattodea, containing all members of the group except termites. About 30 cockroach species out of 4,600 are associated with human habitats. Some species are we ...
es, ladybugs,
yellowjacket Yellowjacket or yellowjacket is the common name in North America for predatory social wasps of the genus, genera ''Vespula'' and ''Dolichovespula''. Members of these genera are known simply as "wasps" in other English-speaking countries. Most of ...
s, hornets, mice, and rats. When conditions are dry, the deprivation of moisture outside drives many pests inside searching for water. While the rainy weather increases the numbers of insects, this dry weather causes pest invasions to increase. The pests most commonly known during dry conditions are
scorpion Scorpions are predatory arachnids of the order Scorpiones. They have eight legs, and are easily recognized by a pair of grasping pincers and a narrow, segmented tail, often carried in a characteristic forward curve over the back and always en ...
s, ants,
pillbug Armadillidiidae is a family of woodlice, a terrestrial crustacean group in the order Isopoda. Unlike members of some other woodlice families, members of this family can roll into a ball, an ability they share with the outwardly similar but unre ...
s,
millipede Millipedes are a group of arthropods that are characterised by having two pairs of jointed legs on most body segments; they are known scientifically as the class Diplopoda, the name derived from this feature. Each double-legged segment is a resu ...
s, crickets, and spiders. Extreme drought does kill many populations of insects, but also drives surviving insects to invade more often. Cold temperatures outside will cause invasions beginning in the late summer months and early fall.
Box elder bug The boxelder bug (''Boisea trivittata'') is a North American species of true bug. It is found primarily on boxelder trees, as well as maple and ash trees.
s, cluster flies, ladybugs, and silverfish are noticed some of the most common insects to seek the warm indoors."Dead rat tips"
. ''Terminix.com''. 19 March 2008.
In general, insects are poikilothermic animals; thus meaning their level of activity is substantially depended upon their surrounding environmental conditions. An increase in the temperature will result in an accelerated metabolism of the insect; hence resulting in an increased activity.


Modern techniques

Many new techniques have been developed and are used in order to more accurately gather evidence, or reevaluate old information. The use of these newly developed techniques and evaluations have become relevant in litigation and appeals. Forensic entomology not only uses arthropod biology, but it pulls from other sciences, introducing fields like chemistry and genetics, exploiting their inherent synergy through the
use of DNA in forensic entomology Forensic entomology has three sub-fields: urban, stored product and medico-criminal entomologies. This article focuses on medico-criminal entomology and how DNA is analyzed with various blood-feeding insects. Forensic entomology can be an import ...
.


Scanning electron microscopy

Fly larvae and fly eggs are used to aid in the determination of a PMI. In order for the data to be useful the larvae and eggs must be identified down to a species level to get an accurate estimate for the PMI. There are many techniques currently being developed to differentiate between the various species of forensically important insects. A study in 2007 demonstrates a technique that can use scanning electron microscopy (SEM) to identify key morphological features of eggs and maggots. Some of the morphological differences that can help identify the different species are the presence/absence of anastomosis, the presence/absence of anterior and posterior spiracles, the cephalopharyngeal skeleton as well as the shape and length of the median area. The SEM method provides an array of morphological features for use in identifying fly eggs; however, this method does have some disadvantages. The main disadvantage is that it requires expensive equipment and can take time to identify the species from which the egg originated, so it may not be useful in a field study or to quickly identify a particular egg. The SEM method is effective provided there is ample time and the proper equipment and the particular fly eggs are plentiful. The ability to use these morphological differences gives forensic entomologists a powerful tool that can help with estimating a post mortem interval, along with other relevant information, such as whether the body has been disturbed post mortem.


Potassium permanganate staining

When scanning electron microscopy is not available, a faster, lower cost technique is
potassium permanganate Potassium permanganate is an inorganic compound with the chemical formula KMnO4. It is a purplish-black crystalline salt, that dissolves in water as K+ and , an intensely pink to purple solution. Potassium permanganate is widely used in the c ...
staining. The collected eggs are rinsed with a normal saline solution and placed in a glass petri dish. The eggs are soaked in a 1% potassium permanganate solution for one minute and then dehydrated and mounted onto a slide for observation. These slides can be used with any light microscope with a calibrated eyepiece to compare various morphological features. The most important and useful features for identifying eggs are the size, length, and width of the plastron, as well as the morphology of the plastron in the area around the micropyle. The various measurements and observations when compared to standards for forensically important species are used to determine the species of the egg.


Mitochondrial DNA

In 2001, a method was devised by Jeffrey Wells and Felix Sperling to use mitochondrial DNA to differentiate between different species of the subfamily Chrysomyinae. This is particularly useful when working to determine the identity of specimens that do not have distinctive morphological characteristics at certain life stages.


Mock crime scenes

A valuable tool that is becoming very common in the training of forensic entomologists is the use of mock crime scenes using pig carcasses. The pig carcass represents a human body and can be used to illustrate various environmental effects on both arthropod succession and the estimate of the post mortem interval. Pigs are the most frequently utilised model in an attempt to gather data regarding forensic experimental analysis. The latter is highly proportionate to human nature due to our overlapping characteristics with the mentioned species. These interrelated components include: subcutaneous fat stores, skin thickness, range of adult body mass, hair covering, and omnivorous diets.


Gene expression studies

Although physical characteristics and sizes at various
instars 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 ...
have been used to estimate fly age, a more recent study has been conducted to determine the age of an egg based on the expression of particular genes. This is particularly useful in determining developmental stages that are not evidenced by change in size; such as the egg or pupa and where only a general time interval can be estimated based on the duration of the particular developmental stage. This is done by breaking the stages down into smaller units separated by predictable changed in gene expression. Three genes were measured in an experiment with ''Drosophila melanogaster'': bicoid (bcd), slalom (sll), and chitin synthase (cs). These three genes were used because they are likely to be in varied levels during different times of the egg development process. These genes all share a linear relationship in regards to age of the egg; that is, the older the egg is the more of the particular gene is expressed. However, all of the genes are expressed in varying amounts. Different genes on different loci would need to be selected for another fly species. The genes expressions are mapped in a control sample to formulate a developmental chart of the gene expression at certain time intervals. This chart can then be compared to the measured values of gene expression to accurately predict the age of an egg to within two hours with a high confidence level. Even though this technique can be used to estimate the age of an egg, the feasibility and legal acceptance of this must be considered for it to be a widely utilized forensic technique. One benefit of this would be that it is like other DNA-based techniques so most labs would be equipped to conduct similar experiments without requiring new capital investment. This style of age determination is in the process of being used to more accurately find the age of the instars and pupa; however, it is much more complicated, as there are more genes being expressed during these stages. The hope is that with this and other similar techniques a more accurate PMI can be obtained.


Insect activity case study

A preliminary investigation of insect colonization and succession on remains in
New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island count ...
revealed the following results on decay and insect colonization.


Open field habitat

This environment had a daily average maximum temperature of and a daily minimum temperature of . The average rainfall for the first 3 weeks in this environment was 3.0 mm/day. Around days 17–45, the body began to start active decay. During this stage, the insect successions started with '' Calliphora stygia'', which lasted until day 27. The larvae of '' Chrysomya rufifacies'' were present between the day 13 and day 47. '' Hydrotaea rostrata'', larvae of '' Lucilia sericata'', family
Psychodidae Psychodidae, called drain flies, sink flies, filter flies, sewer flies, or sewer gnats, is a family of Fly, true flies. Some genera have short, hairy bodies and wings giving them a "furry" moth-like appearance, hence one of their common names, m ...
, and ''
Sylvicola ''Sylvicola'' is a genus of wood gnats in the family Anisopodidae. There are more than 80 described species in ''Sylvicola''. Species These 89 species belong to the genus ''Sylvicola'': *'' Sylvicola adornatus'' Yang & Cui, 1998 *'' Sylvicola a ...
'' were found to be present relatively late in the body's decay.


Coastal sand-dune habitat

This environment had an average daily maximum temperature of and minimum of . The daily average rainfall was recorded as 1.4 mm/day for the first 3 weeks. The post-decay time interval, beginning at day six after death and ending around day 15 after death, is greatly reduced from the average post-decay time, due to the high average temperature of this environment. Insects obtained late in the post-active stage include the ''Calliphora quadrimaculata'', adult Sphaeroceridae,
Psychodidae Psychodidae, called drain flies, sink flies, filter flies, sewer flies, or sewer gnats, is a family of Fly, true flies. Some genera have short, hairy bodies and wings giving them a "furry" moth-like appearance, hence one of their common names, m ...
and Piophilidae (no larvae from this last family were obtained in recovery).


Native bush habitat

This environment had recorded daily average maximum and minimum temperatures were and , respectively. The average rainfall in this habitat was recorded at 0.4 mm/day. After the bloat stage, which lasted until day seven after death, post-active decay began around day 14. In this habitat, the ''H. rostrata'', adult
Phoridae The Phoridae are a family of small, hump-backed flies resembling fruit flies. Phorid flies can often be identified by their escape habit of running rapidly across a surface rather than taking to the wing. This behaviour is a source of one of thei ...
, Sylvicola larvae and adult were the predominant species remaining on the body during the pre-
skeletonization Skeletonization is the state of a dead organism after undergoing decomposition.The Australian Museum. (2018). Decomposition-Body Changes. Retrieved from: https://australianmuseum.net.au/about/history/exhibitions/death-the-last-taboo/decomposit ...
stages.


In literature

Throughout its history the study of forensic entomology has not remained an esoteric science reserved only for entomologists and forensic scientists. Early twentieth-century popular scientific literature began to pique a broader interest in entomology. The very popular ten-volume book series, Alfred Brehem's ''Thierleben'' (Life of Animals, 1876–1879) expounded on many
zoological Zoology ()The pronunciation of zoology as is usually regarded as nonstandard, though it is not uncommon. is the branch of biology that studies the animal kingdom, including the structure, embryology, evolution, classification, habits, and dis ...
topics, including arthropods. The accessible writing style of French entomologist
Jean-Henri Fabre Jean-Henri Casimir Fabre (21 December 1823 – 11 October 1915) was a French naturalist, entomologist, and author known for the lively style of his popular books on the lives of insects. Biography Fabre was born on 21 December 1823 in Saint-L ...
was also instrumental in the popularization of entomology. His collection of writings ''Souvenirs Entomologique'', written during the last half of the 19th century, is especially useful because of the meticulous attention to detail to the observed insects' behaviors and life cycles. The real impetus behind the modern cultural fascination with solving crime using entomological evidence can be traced back to the works ''Faune des Tombeaux'' (Fauna of the Tombs, 1887) and ''Les Faunes des Cadavres'' (Fauna of Corpses, 1894) by French
veterinarian A veterinarian (vet), also known as a veterinary surgeon or veterinary physician, is a medical professional who practices veterinary medicine. They manage a wide range of health conditions and injuries in non-human animals. Along with this, vet ...
and entomologist Jean Pierre Mégnin. These works made the concept of the process of insect ecological succession on a corpse understandable and interesting to an ordinary reader in a way that no other previous scientific work had done. It was after the publication of Mégnin's work that the studies of forensic science and entomology became an established part of Western popular culture, which in turn inspired other scientists to continue and expand upon his research.Benecke, M. (2001). A brief history of forensic entomology. Forensic Entomology International, 120, page 5


See also

* Forensic entomology and the law * Insect indicators of abuse or neglect


Notes


Further reading

* * * Spiralbound also aimed at professional entomologists, but shorter and with a popular style. * * *Gennard, Dorothy E., 2007 ''Forensic entomology : an introduction''

assumes no prior knowledge * * * * * * * A technical hardback designed for professional entomologists. *


External links

*
www.forensic-entomology.com by Dr. Jason H. ByrdEuropean Association for Forensic Entomology homepageCollection of original articles about Forensic Entomology. Many downloadable illustrated pdf's of cases, including neglect cases.
* ttp://www.deathonline.net/decomposition/index.htm What happens to the body after deathbr>Pastoral putrefaction down on the Body Farm: Autopsy, HBO Documentaries

DNA techniques for forensic entomologypdf about forensic entomology Deon Canyon''Survey on carrion-visiting beetles''
– Research into the fauna of carrion-visiting beetles in Erlangen, Bavaria (2006): Carrion is an ecosystem of its own. Diverse species of beetles – belonging to different guilds – arrive and depart from carrion at different times – succession occurs. The arrival time and growth rates of beetles inhabiting corpses depending on weather, location and type of carrion have been studied.
History of Forensic Entomology (many illustrations)Diagnosis and Keys to South American Coleoptera of Forensic ImportanceForensic entomology - use of insects to help solve crimeA Forensic Entomology Case from the Amazon Rain Forest of Brazil''Grounds Maintenance'' magazine

Medical History
Annick Opinel 2008, The Emergence of French Medical Entomology: The Influence of Universities, the Institut Pasteur and Military Physicians (1890–c.1938) ''Med Hist.'' 2008 July; 52(3): 387–405. {{Authority control Chinese inventions French inventions German inventions Italian inventions Subfields of entomology