Silphidae

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Silphidae
Temporal range: Middle Jurassic–Present
Nicrophorus.vespillo.jpg
Nicrophorus vespillo
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Coleoptera
Suborder: Polyphaga
Infraorder: Staphyliniformia
Superfamily: Staphylinoidea
Family: Silphidae
Latreille, 1806 [1]
Subfamilies

Nicrophorinae Kirby, 1837
Silphinae Latreille, 1806

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.

Taxonomy, evolution, and etymology

The family Silphidae belongs to the order Coleoptera. They are commonly referred to as carrion beetles or burying beetles and are usually associated with carrion, fungi, and dung. In the past, members of the family Agyrtidae were included. This family has two subfamilies, Silphinae and Nicrophorinae. The antenna is made up of 15 segments and is capitate (ending in an abruptly capped club) in the Nicrophorinae and has a more gradual club shape in the Silphinae. The subfamilies also differ in behavior. Members of the subfamily Silphinae show little to no care for their young and breed on large carrion. Nicrophorinae breed on small animal carrion and will bury themselves and their food to rear their offspring in a bi-parental manner. [2] There are approximately 183 species in this family, which are found worldwide although they are commoner in temperate regions. Nicrophorus americanus , known as the American burying beetle, is an endangered species. [3]

The oldest fossils of silphids are known from the Middle Jurassic (~ 163 million years ago) Daohugou Bed in Northern China. [4] [5] Many Silphidae are flightless although they have wings. This loss is thought to be a result due to the changes in habitat over time. Researchers have found that most flight-capable species in this group feed on vertebrate carcasses, whereas flightless species will feed on soil invertebrates. They also found that egg production increased with flight loss because of a more limited food supply. [6]

Ridged Carrion Beetle (Oiceoptoma inaequale) Ridged Carrion Beetle (Oiceoptoma inaequale).jpg
Ridged Carrion Beetle (Oiceoptoma inaequale)

The word "silphid" or "sylph", first seen in the sixteenth century in Paracelsus' works, refers to any race of spirits inhabiting the air and is described as mortal, but lacking soul. The word is also related to the Latin word silva meaning "forests" or "of the woods". [7]

Diversity and distribution

Silphidae are ubiquitous and are most abundant in the temperate zone. The diversity is also greater in the temperate zone and they are quite rare in the tropics although there are species endemic to the region. It is thought that ants, flies and other carrion feeders outcompete them in these regions. [8] They vary in size from 7 to 45 mm. [9]

There are about 46 different species of Silphidae in North America which include Heterosilpha ramosa, Necrodes surinamensis, Necrophila americana , Nicrophorus americanus , Nicrophorus carolinus , Nicrophorus investigator , Nicrophorus marginatus , Nicrophorus orbicollis , Nicrophorus tomentosus , Oiceoptoma inaequale, Oiceoptoma noveboracense , Oiceoptoma rugulosum, Thanatophilus lapponicus. [10] [11] A species found in Great Britain is Oiceoptoma thoracicum.

One species of Nicrophorus beetle within the family Silphidae, Nicrophorus nepalensis, can be found primarily in the mountains of eastern Asia as well as along the Malay Archipelago. Nicrophorus nepalensis are found in the Indian subcontinent as well in the countries of India and Pakistan.

Development

Silphidae undergo holometabolous development. The development in the subfamily Silphinae proceeds at a slower rate than in Nicrophorinae. The Silphinae life cycle takes approximately twenty six to fifty eight days to go from an egg to adult. The breakdown of this process is essential to forensic entomologists. The cycle takes two to seven days after the egg is laid to hatch. The larvae will develop through three instars on the carrion lasting for ten to thirty days. After that time period is up the third instar larvae will venture away from the detritus to pupate. Pupation takes fourteen to twenty one days and is the major part of metamorphosis where a grandiose change occurs. During this stage the wings become fully developed and sexual maturity is reached, sometimes called the imago or adult stage where the cycle is then repeated. The Nicrophorinae cycle is generally quicker. Oviposition is done near the carcass and takes twelve to forty eight hours for the eggs to hatch into larvae. The amount of food and parental care exhibited help determine the length of the larval stage. Pupation in this subfamily lasts six to eight days and is completed in the soil. The adult Nicrophorinae will emerge from the soil and venture to find food and a mate. [10]

Reproduction

Male (left) and female Necrophila americana Necrophila americana (Linnaeus) pair.jpg
Male (left) and female Necrophila americana

Nicrophorinae are well known for the habit of locating a carcass and burying it by unearthing the soil underneath it. The burying behavior has seemingly evolved to prevent competition from other insects such as fly maggots. It has been observed that the cooperation of the two parent beetles leads to breeding success. More likely than not a breeding pair will work together, but in cases where there is large carrion males try to boost their reproduction by emitting pheromones. In this way, he will father more offspring, but the reproductive success of the primary female steadily declines. Sometimes, where there is a large carcass the likelihood of intense competition from flies leads to communal breeding. There appears to be a truce between females who would normally compete for the males, and in these cases cooperative behavior extends to females caring for each other's offspring. At the height of breeding season pairs of beetles may compete for the carrion. The losing pair will be ejected from the carrion and if any eggs have been laid they are killed so the new female can lay her own. [12]

Behavior and ecology

Food

Nicrophorus vespilloides colonizing a dead rodent. Nicrophorus vespilloides in dead rodent.jpg
Nicrophorus vespilloides colonizing a dead rodent.

Silphid adults feed in a saprophagous manner: they colonize the carrion during all four stages of decomposition, which are fresh, bloated, decay, and dry. The main areas of decomposition for adults are during both the bloated and decaying stages. Silphid larvae mainly inhabit during the decaying and dry stages of the carrion. The primary food source for the subfamily Silphinae is the maggot mass present on the detritus. Nicrophorinae will colonize the body earlier in decomposition in order to avoid competition with maggots. If there is a sufficiently large maggot mass they will not colonize the carcass. The parental care exhibited by this subfamily is that the adult beetles regurgitate food into the mouths of the young larvae until they are mature. Silphinae colonize later in the decaying process and the adults eat the maggot mass, sometimes leaving little maggot evidence left to estimate a post-mortem interval. In the case of the sexton or burying beetles, Nicrophorinae, the adults will bury small animal carcasses and lay their eggs on it. In some species, a slight depression is made on the detritus for maturing larvae that the adult beetles feed and protect. In both subfamilies the larvae are observed to eat the decaying organic material while the adults mainly consume the maggots. Flies are the major competitor of silphids for detritus. If a carcass is infested with maggots, many of the Nicrophorinae will abandon the carcass while members of Silphinae will feed on the maggots. [10]

Defense

Beetles have many different weapons available to protect them against predator attacks. The members of Silphidae have many different modifications that allow them to thrive in different ecological niches including colour warnings (from aposematism to Batesian mimicry), chemical defenses and parental care. Silphines are usually dark in color consisting of gold, black and brown. This dark coloring allows them to blend into their environment. Many nicrophorines have bright orange coloring on their elytra, which may serve as a warning to other predators. [13] Some species secrete a chemical from a rectal gland that consists of aliphatic acids and terpene alcohols. The secretion has a strong, foul odor and may be topically irritating to cockroaches and flies. The species Necrodes surinamensis ejects this secretion as a spray and can rotate the end of its abdomen to spray in all directions. [14]

Locomotion and navigation

Walking is the primary form of locomotion for Silphidae beetles. They are able to travel great distances to find carcasses to breed and feed on. Beetles also have two sets of wings, the elytra and the hind wings. The hind wings are membranous and are modified for flying or swimming. There are some Silphidae who are able to fly, but others have lost this ability throughout evolution. When an animal dies, hydrogen sulfide and some cyclic compounds are released. Silphidae use their sense of smell to locate carcasses from a long distance by chemoreceptors on their antennae, which are adapted to detect these chemicals. [15] At a short distance, the end organs of the palpi detect the odors. [13] Silphid beetles are usually more active at night, nocturnal, which may help reduce competition.

Competition

Competition from other organisms for the carrion puts the Silphids at both an advantage and a disadvantage. The advantage is that fly competitors lay eggs that result in maggots and supply food for Silphids. The disadvantage for the Nicrophorinae is that if the carcass is already “blown”, referring to Shakespearean time as being infected with fly maggots, they will not colonize. Nicrophorinae have adapted to these situations and will bury the carcass to remove it from other competitors. [16] With this technique, a steady food source is available for the larvae and procreation chances increase.

Silphidae compete for brooding areas. If an invader male overpowers the original male, it will mate with the original female and create a new brood. [17] If an invader female overpowers the original female, it will also create a new brood with the primary male.

Relationships

With humans

Silphids are usually not considered a nuisance to humans. They help the environment by laying their eggs on carcasses and the larvae break down the detritus, which prevents accumulation of deceased organisms.

Carcasses are kept out of sight and foul odors are prevented when Nicrophorinae bury it under the ground. This will also reduce the surface area for flies to lay their eggs and decrease fly population.

Some Silphidae occupy human-inhabited areas and become pests to farmers by using crops as a second source of nutrients. Aclypea opaca in Europe feed on beets. Necrophila americana feed on pumpkin, spinach, and sugar beet.

With other organisms

Members from the family Silphidae are known to have mutualistic relationships with other organisms. Nicrophorinae have a mutual relationship with phoretic mites. Mites from the genus Poecilochirus produce deutonymphs that crawl on Nicrophorinae and are transported to carrion. Once they arrive at the carrion, deutonymphs leave the adult Nicrophorinae and proceed to feed on nearby fly eggs and immature larvae. Mites help Nicrophorinae reduce the number of competitors on carrion. With less competition, both species are able to reproduce successfully underground. [18]

Silphidae are known for being hosts to juvenile nematodes. Nicrophorus vespilloides are closely associated with the nematode parasite. The parasites can be easily transmitted to other hosts during copulation.

Forensic research

Silphidae are one of several families of forensic importance in the order Coleoptera. They are a very important tool in determining a post-mortem interval by collecting Silphid progeny from the carcass, and determining the developmental rate. Based on the number of instars and the larval development stage, a time of death can be estimated. This is very useful in medicocriminal entomology, the emphasis on utilizing arthropods as evidence to aid in solving crimes. [10] Many of the methods in determining stages of development are subjective. However, recent studies have found a more precise way of determining the stage of development of Silphid larvae by measuring the maximum cranial width and other heavily sclerotized areas of the larvae instead of measuring just the length, which is subject to change with each larva, particularly in O. inaequale and N. surinamensis, which are more robust and have greater variations of length. The most accurate instar identification is possible by using distinct morphological features that are instar specific and cannot be affected by the size of the specimen. Although this is possible only for few species with described larvae of all instars, such as Thanatophilus rugosus . [19]

Silphidae are being studied to find more exact estimations of post-mortem intervals and possible manners of death. [20] Also, in the future, entomologists will explore the social behavior of the beetles to a greater degree. Members of family Silphidae are typically the first of the coleopterans to come in contact with carrion. Silphidae larvae are opportunistic predators that will feed on dipteran eggs, larvae, and on the carcass itself. [20] This presents a problem in the determination of post-mortem interval because Silphidae are known to eradicate other species from carrion. By eliminating the first colonizing species Silphidae can give an incorrect post-mortem interval.

Unlike most of the insects used as forensic markers, it is possible to assess if Silphidae adults have colonized a corpse and left it later. For this purpose, some Parasitidae mites (for example, Poecilochirus species) are used due to their phoront-host specificity. If a corpse is invaded by Poecilochirus specimens and no Silphidae were sampled by forensic entomologists, it is reasonable to suspect that some Silphidae carrying Poecilochirus phoretic instars arrived previously to the corpse and abandoned it some time later, leaving Poecilochirus mites in the process. [21]

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">Burying beetle</span> Genus of beetles

Burying beetles or sexton beetles, genus Nicrophorus, are the best-known members of the family Silphidae. Most of these beetles are black with red markings on the elytra (forewings). Burying beetles are true to their name—they bury the carcasses of small vertebrates such as birds and rodents as a food source for their larvae, this makes them carnivorous. They are unusual among insects in that both the male and female parents take care of the brood.

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

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 tergites exposed, and their geniculate (elbowed) antennae with clubbed ends. These predatory feeders are most active at night and will fake death if they feel threatened. This family of beetles will occupy almost any kind of niche throughout the world. Hister beetles have proved useful during forensic investigations to help in time of death estimation. Also, certain species are used in the control of livestock pests that infest dung and to control houseflies. Because they are predacious and will even eat other hister beetles, they must be isolated when collected.

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

Trogidae, sometimes called hide beetles, is a family of beetles with a distinctive warty or bumpy appearance. Found worldwide, the family includes about 300 species contained in four or five genera.

<i>Nicrophorus americanus</i> Species of beetle

Nicrophorus americanus, also known as the American burying beetle or giant carrion beetle, is a critically endangered species of beetle endemic to North America. It belongs to the order Coleoptera and the family Silphidae. The carrion beetle in North America is carnivorous, feeds on carrion and requires carrion to breed. It is also a member of one of the few genera of beetle to exhibit parental care. The decline of the American burying beetle has been attributed to habitat loss, alteration, and degradation, and they now occur in less than 10% of their historic range.

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>Nicrophorus vespilloides</i>

Nicrophorus vespilloides is a burying beetle described by Johann Friedrich Wilhelm Herbst in 1783. The beetles vary widely in size and can present with a range of anywhere from 12 mm to 20 mm in size. They have two conspicuous orange-yellow bands on the elytra. The color of the antennae are an important distinguishing feature, being totally black. The color of their orange and black markings is multifunctional, as they are conspicuous to avian predators. In general, they present a unique ecological niche, which is their evolution of aposematism, or the strategy they use to warn predators through their conspicuous signals. The wing cases of these beetles possess a squarish shape and are notably shorter in length than their abdomens, indicating a distinct physical characteristic of the species.

<i>Nicrophorus tomentosus</i> Species of beetle

Nicrophorus tomentosus is a species of burying beetle that was described by Friedrich Weber in 1801. The beetle belongs to the family Silphidae which are carrion beetles. The beetles have sensitive antennae that contain olfactory organs. Thus, the beetle can locate dead animals (carcass), and then as the name suggests, can bury them. However, unlike other burying beetles, N. tomentosus does not completely bury these brood carcasses. They instead dig a shallow hole under the carcass and cover it with leaf litter. Recognition of these beetles can be distinguished by its black color with orange markings on the wing covers (elytra).

<i>Nicrophorus nepalensis</i> Species of beetle

Nicrophorus nepalensis, commonly known as a burying beetle, is widespread across tropical and subtropical countries in Asia. It belongs to the order Coleoptera and the family Silphidae, and is part of the nepalensis species-group, which is the second largest species group within the genus Nicrophorus. N. nepalensis differs from some other beetles in that it exhibits biparental care. Its role as a decomposer is crucial in the energy cycle and energy transformation in the ecosystem.

<i>Nicrophorus pustulatus</i> Species of beetle

Nicrophorus pustulatus, also known as the pustulated carrion beetle or blistered burying beetle, is a species of burying beetle that was described by Johann Karl Wilhelm Illiger in 1807. This species is native to North America. N. pustulatus exhibits unique habitat utilization and breeding behaviour relative to other members of the genus. This species may be the only described example of a true parasitoid targeting a vertebrate host.

<i>Nicrophorus vespillo</i> Species of beetle

Nicrophorus vespillo is a burying beetle described by Carl Linnaeus in his landmark 1758 10th edition of Systema Naturae. It has a paleartic distribution and is commonly found across Europe and Asia, extending from Western Europe to Mongolia.

<i>Nicrophorus interruptus</i> Species of beetle

Nicrophorus interruptus is a species of burying beetle or sexton beetle belonging to the family Silphidae subfamily Nicrophorinae.

<i>Nicrophorus quadripunctatus</i> Species of beetle

Nicrophorus quadripunctatus is a species of burying beetle that predominates in East Asia. First described by German entomologist Ernst Kraatz in 1877, this beetle has since been the subject of much scientific inquiry—particularly concerning its parental care. Like other burying beetles, N. quadripunctatus inhabit small, vertebrate animal carcasses. This environment provides the beetles with the requisite nutrients for themselves and their offspring. To limit resource theft and predation, the carcass is buried underground. For additional protection, a single, dominant male-female pair guards the carcass cooperatively.

<i>Nicrophorus orbicollis</i> Species of beetle

Nicrophorus orbicollis is a nearctic burying beetle first described by Thomas Say in 1825. It is a member of the genus Nicrophorus or sexton beetles, comprising the most common beetles in the family Silphidae. This species is a decomposer feeding on carcasses of small dead animals. N. orbicollis can be used for scientific research both medically and forensically.

<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>Poecilochirus</i> Genus of mites

Poecilochirus is a Holarctic genus of mites in the family Parasitidae. They are relatively large and often found on rotting corpses, where they are transported by beetles. Deuteronymphs are characterized by two orange dorsal shields and in many species a transverse band on the sternal shield. The juvenile development consists of a larval stage, protonymph, and deuteronymph, but no tritonymph. Females are smaller than males. Males guard female deuteronymphs shortly before these mate, and pairs mate venter-to-venter.

<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.

<i>Oiceoptoma noveboracense</i> Species of beetle

Oiceoptoma noveboracense is a member of the family Silphidae, or carrion beetles, which feed on decaying organic matter such as dead animals. Its common name is the margined carrion beetle, from the orange-red margins on the pronotum, which are helpful when identifying this species. The larva is typically light brown to red and also has vertical ridges on its thorax like the adult. This diurnal beetle can be found mainly in the spring into the fall, and it has a strong preference towards a deciduous forest habitat. The primary forensic importance of this beetle is its ability to use the succession of insect fauna to provide confirmation of postmortem intervals.

<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.

References

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  3. ↑ Backlund, Douglas C.; Gary M. Marrone; Christopher K. Williams & Kelley Tilmon (2008). "Population Estimate of the Endangered American Burying Beetle, Nicrophorus americanus Olivier (Coleoptera: Silphidae) in South Dakota". The Coleopterists Bulletin. 62 (4): 9–15. doi:10.1649/982.1. S2CID   85841685.
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  19. ↑ Novák M.; Jakubec P.; Qubaiová J. Šuláková H.; Růžička J. (2017). "Revisited larval morphology of Thanatophilus rugosus (Coleoptera: Silphidae)". International Journal of Legal Medicine. 2017 (3): 939–954. doi:10.1007/s00414-017-1764-6. PMID   29270839. S2CID   206976798.
  20. 1 2 Watson E.J.; Carlton C.E. (2005). "Succession of Forensically Significant Carrion Beetle Larvae on Large Carcasses (Coleoptera: Silphidae)". Southeastern Naturalist. 4 (2): 335–346. doi:10.1656/1528-7092(2005)004[0335:sofscb]2.0.co;2. S2CID   86084430.
  21. ↑ González Medina A, González Herrera L, Perotti MA, Jiménez Ríos G (2013). "Occurrence of Poecilochirus austroasiaticus (Acari: Parasitidae) in forensic autopsies and its application on postmortem interval estimation". Experimental and Applied Acarology. 59 (3): 297–305. doi:10.1007/s10493-012-9606-1. PMID   22914911. S2CID   16228053.


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