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Histeridae is a
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
of
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 commonly known as clown beetles or Hister beetles. This very diverse group of beetles contains 3,900
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
found worldwide. They can be easily identified by their shortened
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 ...
that leaves two of the seven
tergites A ''tergum'' (Latin for "the back"; plural ''terga'', associated adjective tergal) is the dorsal ('upper') portion of an arthropod segment other than the head. The anterior edge is called the 'base' and posterior edge is called the 'apex' or 'mar ...
exposed, and their geniculate (elbowed) antennae with clubbed ends. These
predatory Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
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 Niche may refer to: Science *Developmental niche, a concept for understanding the cultural context of child development *Ecological niche, a term describing the relational position of an organism's species *Niche differentiation, in ecology, the ...
throughout the world. Hister beetles have proved useful during
forensic Forensic science, also known as criminalistics, is the application of science to criminal and civil laws, mainly—on the criminal side—during criminal investigation, as governed by the legal standards of admissible evidence and criminal p ...
investigations to help in time of death estimation. Also, certain
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
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.


Characteristics

The Hister beetles are easily identified by their shiny
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 ...
which is typically shiny black or metallic green. The two main shapes for this family are oval and flat. The elytra is shorter than 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 ...
with typically two of the seven
tergites A ''tergum'' (Latin for "the back"; plural ''terga'', associated adjective tergal) is the dorsal ('upper') portion of an arthropod segment other than the head. The anterior edge is called the 'base' and posterior edge is called the 'apex' or 'mar ...
exposed. Hister beetles have specialized heads that can retract into their
prothorax The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites (exoskeletal plates) are the pronotum ( dorsal), the prosternum ( ventral), and the propleuron ( lateral) o ...
and two geniculate (elbowed) antennae with clubbed ends. As
predatory Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
feeders, the Hister beetles commonly feed on the egg, larval, and adult stages of other insects. Certain species are also used to control
livestock Livestock are the domesticated animals raised in an agricultural setting to provide labor and produce diversified products for consumption such as meat, eggs, milk, fur, leather, and wool. The term is sometimes used to refer solely to animal ...
pests that infest dung or to control house flies. Histeridae are most active at night and they will play dead if they feel threatened.


Etymology

Histeridae was named by
Leonard Gyllenhaal Leonard Gyllenhaal (3 December 1752 – 13 May 1840) was a Sweden, Swedish military officer and entomology, entomologist. Born on the Ribbingsberg Manorialism, manor in Västergötland in west Sweden, Leonard Gyllenhaal was son of an army of ...
. Histeridae has two common names, the “Clown” beetle and the “Hister” beetle. There have been several theories explaining the origin of these common names.Dillon, Elizabeth S., and Lawrence R. Dillon. A Manual of Common Beetles of Eastern North America. Peterson and Co., 1961. One theory for the “Hister” nickname comes from the work of
Juvenal Decimus Junius Juvenalis (), known in English as Juvenal ( ), was a Roman poet active in the late first and early second century CE. He is the author of the collection of satirical poems known as the '' Satires''. The details of Juvenal's life ...
, a Roman poet. Juvenal used the word “hister” to mean a dirty, lowly being. Another theory for the origin of this beetle's name stems from the fact that in Latin, “hister” means actor. Many people believe the name is associated with the Hister beetles’ ability to imitate death when they are disturbed. Still, some believe this family of beetles was named for its physical characteristics. The Clown beetle has flattened legs, which can be compared to a clown's flat shoes or loose-fitting pants.


Anatomy

The
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 ...
and forewings are normally well developed in beetles, but in Histeridae the elytra are shortened and rectangular. The shortened elytra expose the final two of the seven
tergites A ''tergum'' (Latin for "the back"; plural ''terga'', associated adjective tergal) is the dorsal ('upper') portion of an arthropod segment other than the head. The anterior edge is called the 'base' and posterior edge is called the 'apex' or 'mar ...
. The head has compound eyes, a
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
mouthpart, and shortened antennae. A distinctive characteristic is that the antennae are elbowed and contain three antennomeres that form a club at the end. Clown beetles have an open
circulatory system The blood circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascular system, tha ...
within its hemocoel, also known as a body cavity. They have a tube shaped heart that spans the length of the body, and use
hemolymph Hemolymph, or haemolymph, is a fluid, analogous to the blood in vertebrates, that circulates in the interior of the arthropod (invertebrate) body, remaining in direct contact with the animal's tissues. It is composed of a fluid plasma in which ...
as blood. This blood does not contain any oxygen, but carries nutrients throughout the body. Spiracles are located on the abdomen and are the tracheal system of the beetle. Oxygen is brought into the body by spiracles and tiny sacs exchange the oxygen like a mammal's lung.


Habitat

Hister beetles are found throughout the world in various habitats. Histeridae have been located in North America, Central America, Europe, Asia, and Australia, but each Hister beetle occupies certain niches. The beetles live in dung, carrion, dead vegetation, sandy areas, under tree bark, mammal burrows, and ant/termite colonies. Charactertistics of the Hister beetle are dependent upon its habitat. For example, the flat Hister beetles are found under bark, while the cylindrical beetles are not. Shape of the beetle will vary from species to species. A remarkable ability of the Hister beetle is their capacity to live in close proximity with
ants Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Creta ...
(myrmecophiles) and
termites Termites are small insects that live in colonies and have distinct castes (eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blattod ...
(termitophiles). The Hister beetles can live in harmony with the ants or prey on the ants, depending on the species. Histeridae's habitat is widespread because they feed on other arthropods. As new food sources are introduced into an environment, the predators of that food source will soon follow. Histeridae live in areas where their prey feed. Some examples are mammalian nests where other arthropods scavenge for food or carrion where maggots will arrive.Mazur, Slawomir. "Ecology and Habitats of Histeridae." 13 Mar. 2009 .


Evolutionary history

The oldest record of the family is '' Antigracilus'' from the Aptian aged
Yixian Formation The Yixian Formation (; formerly transcribed as Yihsien Formation) is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans the late Barremian and early Aptian stages of the Early Cretaceous. It is known for its exq ...
of China, which was resolved as the sister species to all living members of the family. The oldest crown group representatives are known from the
Burmese amber Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. The ...
, around 99 million years old, including those belonging to the extant subfamily Haeteriinae, and the living genus '' Onthophilus.''


Feeding habits

The larvae and adult forms of Histeridae have been known to feed on dung, carrion, decomposing vegetation, other insects, larvae, and pupae. The Hister beetles are able to locate both dung and carrion through olfaction. When found on dung, carrion and vegetation, the Hister beetles will feed on the fly larvae found there. The predacious Hister beetle will feed on soft-bodied insect eggs and larvae, Diptera in particular. Some species of Histeridae will even feed on other Histeridae. The majority of Histeridae species prefer dried, decaying habitats. Some Histeridae species live in an integrated nest with ants and termites. Some species have been found to be fed by the ants, while others simply feed on leftover insect larvae that the ants do not want. In contrast, in their adult stage ''Psiloscelis'' will actually feed on adult ants. Because the Histeridae are predacious they can be utilized as control agents, but must be stored in isolation. The Hister beetles have proven useful in both the control of pest flies in poultry houses and pastures, and against pest beetles of stored food products.


Development

Histeridae goes through holometabolous development. In this type of development the larval form does not resemble the adult form and the pupae has internally developing
wing A wing is a type of fin that produces lift while moving through air or some other fluid. Accordingly, wings have streamlined cross-sections that are subject to aerodynamic forces and act as airfoils. A wing's aerodynamic efficiency is e ...
s. They also undergo an immobile pupal stage where they do not eat. After the pupal stage they emerge in their adult form. The Histeridae life cycle consists of egg, larval,
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 and adult stages of development:


Egg

The average time of development from Egg to adult at 30 °C is 20.5 days. The eggs of most species are off-white and oval in shape. The egg takes on average 3.8±0.02 days to develop into the first
instar 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 ...
. The
chorion The chorion is the outermost fetal membrane around the embryo in mammals, birds and reptiles (amniotes). It develops from an outer fold on the surface of the yolk sac, which lies outside the zona pellucida (in mammals), known as the vitell ...
is shiny and smooth. In certain species like ''Epierus'' or ''Platylomalus'' it can look pale brown and be leathery in texture.


Larval

The larval stage of the beetle typically goes through two instars, the second
instar 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 ...
is the longest stage of its entire development, taking up 39% of the overall development time. It takes 5.1±0.1 days on average for the first
instar 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 ...
to develop into the second. The larval form of the insect will range in length from three millimeters to several centimeters. They have a membranous body with a limited amount of
sclerotization Sclerotization is a biochemical process that produces the rigid shell of sclerotin that comprises an insect's chitinous exoskeleton. It is prominent in the thicker, armored parts of insects and arachnids, especially in the biting mouthparts and s ...
around the head. There is some
pigmentation A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compoun ...
around the body and it is horizontally segmented. The legs are short and do not help much in
locomotion Locomotion means the act or ability of something to transport or move itself from place to place. Locomotion may refer to: Motion * Motion (physics) * Robot locomotion, of man-made devices By environment * Aquatic locomotion * Flight * Locomo ...
. They move mostly through muscular contraction.


Pupal

The pupal form of the beetle is similar in appearance to the adult form. They have outer cells produced in the larval stages that are reinforced with proteinaceous cement. This makes their outer shell harder and protects them during this vulnerable stage. While they pupate they breathe through spiracles on the abdomen. The beetle is non-feeding and immobile in this stage, as their internal structure is breaking down and rebuilding to its adult form. Under good temperature conditions the Hister beetle will stay in the pupal stage for about a week.


Male and Female Structures

The male and female reproductive organs are hidden underneath the last few
sternites The sternum (pl. "sterna") is the ventral portion of a segment of an arthropod thorax or abdomen. In insects, the sterna are usually single, large sclerites, and external. However, they can sometimes be divided in two or more, in which case the ...
on the mesosternal ( mesosternum) side. The female's structure is modified to work as an ovipositor while the male's is adapted as a copulatory structure. The female has oviducts that carry the developed eggs from the
ovaries The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. T ...
to the ovipositor. The males also have a duct that carries the sperm from the
testes A testicle or testis (plural testes) is the male reproductive gland or gonad in all bilaterians, including humans. It is homologous to the female ovary. The functions of the testes are to produce both sperm and androgens, primarily testoste ...
to the copulatory structure, which stays concealed until copulation. When
fertilization Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Proce ...
takes place, the male leaves enough sperm in the female to fertilize all the eggs in the female's ovaries. The excess sperm is kept in a special structure called
spermatheca The spermatheca (pronounced plural: spermathecae ), also called receptaculum seminis (plural: receptacula seminis), is an organ of the female reproductive tract in insects, e.g. ants, bees, some molluscs, oligochaeta worms and certain other in ...
which holds the sperm until the eggs are fully developed.


Subclades

There are four subclades of Histeridae that are
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
s worth mentioning. These subclades are known as: 1. Dendrobites They have two common body shapes. The first type has a flatter shape; the second has a more cylindrical appearance. The former usually lives near tree barks. This is because the prey they feed on, fly eggs, are found near tree bark. The latter also feed on insects and prefer to live in a forested area. Species of the cylindrical Dendrobites usually hunt prey that is unique for that species. 2. Geobiotes This subclade is the most diverse and the largest of the Hister predator subclasses. The Geobites body structures are generally circular, and they are known for their digging tendencies. This subclass is separated into five more divisions. The members of this subclass live anywhere from the soil, desert, and coast to caves, mammalian burrows, and vegetation. They live in accordance with where their prey lives. One division of the Geobiote feeds on maggots and eggs that are found in forest vegetation or in carrion. Naturally, this Geobiote is found in heavily forested areas. The second and third division hunts for arthropods that feed on dead plant matter. Thus, these types of Geobiotes are found in the sand and burrowed in soil. The fourth type of Geobiotes feed on fly eggs that grow on fresh dung. This division of Geobiotes is found near animal homes such as nests and burrows. The last type of Geobiotes lives in caves. They feed on the mites and other arthropods that occupy the vegetation and fungi found there. This type of Geobiote is known to include species that become blind. 3. Microhisterids This subclade is the smallest of the four. They live on plant litter and feed on the tiny arthropods found there. Microhisterids, like all other Histeridae, become specialized to hunt their prey and live in their habitats. Like the fifth division of Geobiotes, some Microhisterids are known to be blind as well. 4. Inquilines This division consists of those Histeridae that live in close proximity with social colonies of arthropods such as ants and termites. Histeridae that live near ants can live in a harmonious or hostile relationship. The hostile Hister beetles feed on the ants. The harmonious Hister beetles eat the same food as the ants, however they may not be in direct competition for the food. These beetles have an excretory organ that produces an odor telling the ants they mean no harm.


Current research

Because members of the family Histeridae can be found on carrion, they have proven to be important in certain forensic investigations. The predacious Hister beetles will feed on the various insects on the body, primarily Diptera. To estimate a person's time of death, forensic investigators must look at the insects on the body and determine time of colonization. If the Histeridae beetles are present, the investigator can assume that some of the other insects have been eaten by the Hister beetles. Due to their importance in forensic entomology, research is continually being conducted on this family's life cycle and development, their prevalence in specific locations, and their geographical distribution. A recent study was conducted at Hacettepe University in Ankara, Turkey. Forty species of Coleoptera, including the Hister beetles, were observed on twelve pig carcasses over a one-year period. The beetles’ distribution and their time of colonization in the various decomposition stages over the year were examined and recorded.


Importance in forensics

Forensic Entomology Forensic entomology is the scientific study of the colonization of a dead body by arthropods. This includes the study of insect types commonly associated with cadavers, their respective life cycles, their ecological presences in a given environme ...
is the study of insects related to crime scenes. Insects can be very helpful at a crime scene and give people an idea of what happened and when it happened. Flesh eating
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 arrive at the dead body within a few hours. Certain species of the Hister beetles follow shortly behind and prey on the
maggot 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. ...
s and other
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 present. Insects that feed on dead bodies increase the rate 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 ...
and their mandibular mouth parts can cause excess damage to the body. Insects can also help determine the season the body died in. The Hister beetle is more prevalent in Spring and Summer. The family, Histeridae, is very diverse, so different species can be found on the body at different times because of their various feeding habits. This needs to be noted when examining a dead body. They are predacious beetles and arrive when there is material to feed on, like other beetles or maggots. They do not actually feed on the carrion. For example, ''Hister quadrinotatus'' and ''Hister sedakovi'' arrive to feed when the carrion is bloat to dry (full of maggots to no maggots). In contrast, ''Saprinus pennsylvanicus'' arrive at the body first, when it is fresh to late (beginning of decomposition to advanced decomposition). The Hister beetles lay their eggs in dead bodies and over a short period of time they develop into various stages. The Histeridae
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 are destructive and when they hatch they eat the maggots in the dead body. The stage of the beetle and other insects in the body at the time of investigation helps to determine the time of death. "Beetle larvae are ... helpful when determining post-mortem time interval. Beetle larvae often reside in fly breeding resources, and they can be distinguished from fly larvae by the following: Beetle larvae possess a hard, head capsule, often brown in colour. Fly larvae lack a head capsule, instead having distinct, internal, black mouth hooks (cephalopharyngeal skeleton of mouthparts) at the anterior end of their body." Understanding how long ago the eggs were laid and the time period of the developmental stages is important for determining the time of death. The Histers beetles will hide under the dead body in the soil during the day and come out at night to feed. For this reason, the dead body must be examined at different times of the day. After collecting the Hister beetles from a body, isolate them because they are predacious beetles and eat anything in their way.


Genera and species

Histeridae is a large and diverse family. There are more than 410 genera and 4,800 described species in Histeridae worldwide, with more than 500 species in North America. The beetles range in size, shape and color. Many of the adults are predacious.Myers, P., R. Espinosa, and C. S. Parr. "Family Histeridae." Animal Diversity Web. 1998. University of Michigan Museum of Zoology. 22 Mar. 2009 .


See also

* List of Histeridae genera


References


Further references (not necessarily used inline)

*Achiano, K. A., and J. H. Giliomee. "Biology of the house fly predator ''Carcinops pumilio'' (Erichson) (Coleoptera:Histeridae)." BioControl (2005). 14 Mar. 2005. 20 Mar. 2009 . *"Hister beetles - Family Histeridae." Decomposition: Hister Beetles. 2003. Australian Museum. 11 Mar. 2009 . *Stephens, Stephanie A. "An Overview of the Coleopteran Family Histeridae and its Significance to Forensic Entomology." 2003. 13 Mar. 2009 . *Summerlin, J. W., and G. T. Fincher. "Laboratory observations on the life cycle of ''Hister nomas'' (Coleoptera: Histeridae)." Journal of Entomological Science. CABI. Vet. Toxicology & Entomology Res. Lab., College station. 20 Mar. 2009.


External links

*
Tree of lifeHisteridae de
German site
Histeridae of Europe

Overview of HisteridaeBug GuideLectures on Forensically Important Insects
{{Taxonbar, from=Q596252 Histeroidea Beetle families Taxa named by Leonard Gyllenhaal