Cleridae
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Cleridae are a family 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 of the superfamily Cleroidea. They are commonly known as checkered beetles. The family Cleridae has a worldwide distribution, and a variety of
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
s and feeding preferences. Cleridae have many niches and feeding habits. Most genera are
predaceous 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 the ...
and feed on other beetles and larvae; however other genera are
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 ...
or pollen feeders. Clerids have elongated bodies with bristly hairs, are usually bright colored, and have variable antennae. Checkered beetles range in length between . Cleridae can be identified based on their 5–5–5 tarsal formula, division of sternites, and the absence of a special type of vesicle. Female Cleridae lay between 28–42
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 ...
at a time predominately under the bark of trees. Larvae are predaceous and feed vigorously before pupation and subsequently emergence as adults. Clerids have a minor significance in
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 ...
. Some species are occasionally found on carrion in the later dry stages of decay. Also, some species are pests ( stored product entomology) and are found infesting various food products. Research efforts related to Cleridae have focused primarily on using certain species as biological controls. This is a very effective technique for controlling bark beetles due to the voracious appetite of many clerid species.


Description


Appearance

Generally, checkered beetles are elongated and
oval An oval () is a closed curve in a plane which resembles the outline of an egg. The term is not very specific, but in some areas (projective geometry, technical drawing, etc.) it is given a more precise definition, which may include either one ...
in shape and range from in length. Their entire bodies are covered with bristly hairs and many display an ornate body color pattern. These often brightly color patterns can be red, yellow, orange, or blue. The antennae are clubbed at the tip for most species, but others can be "clubbed, saw-tooth, or thread-like." The pronotum region is nearly cylindrical and characteristically narrower than 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 ...
(special hardened front wings), while the head is as wide or wider than the
pronotum 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) on e ...
. Their elytra have tiny pits or depressions, and never expose more than two
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 ...
(
dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage * Dorsal c ...
plates).


Identification

Clerid beetles fall under the suborder Polyphaga. Key characteristics of Polyphaga are that the hind coxa (base of the leg), do not divide the first and second abdominal/ventral plates which are known as
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 ...
. Also, the notopleural suture (found under the pronotal shield) is not present. To further identify Clerid beetles, a few additional characteristics need to be examined. Clerid beetles have unique legs that help to distinguish them from other families. Their tarsal formula is 5–5–5, meaning that on each of the front, middle and hind legs there are 5 tarsomeres (individual subsegments of the feet/tarsi). One or more of these subsegments on each leg is typically lobed, and the 4th tarsi is normally difficult to distinguish. Furthermore, an important feature that eliminates many other families of beetles is that clerids' front coxae (base of the leg) expose the second segment of the legs known as the trochanter. The second defining characteristic of the family Cleridae is that clerids never have eversible vesicles (small usually hidden balloon-like structures thought to be scent glands) on their
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 ...
and
pronotum 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) on e ...
. This characteristic distinguishes them from a similar family
Melyridae Melyridae (common name: soft-winged flower beetles) are a family of beetles of the superfamily Cleroidea. Description Most are elongate-oval, soft-bodied beetles 10 mm long or less. Many are brightly patterned in black and brown, yellow, ...
which sometimes has these glands. This trait is very important in correctly differentiating checkered beetles from Melyridae.


Distribution and ecology

Cleridae can be found in the Americas, Africa, Europe, the Middle East and even in Australia. There are approximately 3,500 species in the world and about 500 species in North America. Due to this wide distribution there are many different habitats in which the checkered beetles can be found. Many of the species are known as "flower visitors", that prey on other
flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechani ...
visiting
insects 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 pairs of j ...
and also feed on pollen. These species are found in moist, sunny environments where flowering plants are found in abundance. Another habitat commonly inhabited by clerid beetles is
trees In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are u ...
. These "tree living species" are found in forests across the world with various climates and an array of easily preyed upon insects. They seek protection under the bark and hunt for other insects above and below the bark. The primary source of prey for these bark living hunters is bark beetles. The third type of clerid beetles is the "nest robbing species" which live in shrubbery and in trees. Unlike the tree living species these species do not actually burrow into the bark. Nest robbing species typically hunt termite, bee, and wasp larvae, and one particular species has been noted to prey primarily on grasshopper egg masses. Not all nest robbing species actively hunt live prey, some species for example prefer to feed only on dead honey bee larvae and adults.


Feeding habits

The Cleridae contains many species of predaceous beetles that feed on other beetles and beetle larvae in their natural habitat. The most common prey item for checkered beetles are the bark beetles and wood boring beetles. In general, the bulk of adult Cleridae feed mainly on other adult beetles while the larvae stage feed on other beetle larvae. Some checkered beetles are known to have an extremely voracious appetite with some larvae able to consume "several times their own body weight" in a day. Although most species of checkered beetles are predaceous in nature, some are scavengers and others have been found feeding on flower pollen.McNamara, J. (1991
"Family Cleridae: checkered beetles"
(PDF). ''In'': Bousquet, Y. (Ed.). ''Checklist of the Beetles of Canada and Alaska''. Agriculture Canada Publication 1861/E. 208–211.
Because of the checkered beetles predaceous nature and insatiable appetite, they are often key players in the
biological control Biological control or biocontrol is a method of controlling pests, such as insects, mites, weeds, and plant diseases, using other organisms. It relies on predation, parasitism, herbivory, or other natural mechanisms, but typically also i ...
of other insects. The checkered beetles have also developed a unique adaptation to aid in their quest for prey. The beetles use
pheromones A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
to help them locate, kill, and consume their prey. The diversity of checkered beetle's feeding habits is quite evident when different species are examined. The '' Necrobia''
spp. 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 se ...
are attracted to dry carrion and other decomposing animal matter such as bones and skin as well as various meat products. '' Thanasimus'' spp. are found in
woodland A woodland () is, in the broad sense, land covered with trees, or in a narrow sense, synonymous with wood (or in the U.S., the ''plurale tantum'' woods), a low-density forest forming open habitats with plenty of sunlight and limited shade (se ...
areas where bark beetle species constitute their main source of
prey 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 ...
. The primary source of prey for the '' Phyllobaenus'' spp. are
wood borer The term woodboring beetle encompasses many species and families of beetles whose larval or adult forms eat and destroy wood (i.e., are xylophagous). In the woodworking industry, larval stages of some are sometimes referred to as woodworms. The t ...
s, immature
weevils Weevils are beetles belonging to the superfamily Curculionoidea, known for their elongated snouts. They are usually small, less than in length, and herbivorous. Approximately 97,000 species of weevils are known. They belong to several families, ...
, and hymenoptera larvae. One of the more diverse genera is '' Trichodes'', the larvae feed on the pollen of flowering plants and adults prey upon grasshoppers and wasps.


Life cycle

The general life cycle of clerids has been known to last anywhere from 35 days to more than 3 years, and is strongly dependent on the life cycle of their prey. While the life cycle can vary in length between genus and species, temperature is also a major determinant in the length of time spent in each stage of development. The warmer the temperature is, the quicker the lifecycle, and the cooler the temperature is the slower the lifecycle. If temperatures dip below a threshold temperature for an extended period of time clerids and most other insects will have growth and developmental progress arrested. Like all beetles, Cleridae follow a
holometabolous Holometabolism, also called complete metamorphosis, is a form of insect development which includes four life stages: egg, larva, pupa, and imago (or adult). Holometabolism is a synapomorphic trait of all insects in the superorder Endopterygota. ...
life cycle: the egg hatches into a
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. ...
, which grows and feeds, changing its skin to form a
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 ...
, and the pupa shedding its skin to emerge as an adult. The larvae of the majority of the known species of Cleridae feed upon the eggs and young of wood-boring beetles, while the adults feed on the adult bark beetles. Copulation takes place while the female feeds, because females need a large amount of food for egg development. The female lays her eggs 36–72 hours after copulation. The eggs are laid in between pieces of bark on wood-borer-infested trees or under stones in the soil. She may lay 28–42 eggs at a time. For the longer lifespaned species such as '' Thanasimus'' this occurs in late summer or early fall to give the
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 enough time for proper growth before having to
overwinter Overwintering is the process by which some organisms pass through or wait out the winter season, or pass through that period of the year when "winter" conditions (cold or sub-zero temperatures, ice, snow, limited food supplies) make normal acti ...
. When larvae hatch from their eggs, they are either red or yellow. Their bodies have a slender and flat appearance with short legs due to their minimal movement. The larvae are covered in hair and have two horn-type projections on the
dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage * Dorsal c ...
area of the last body segment. Immediately after birth, they start searching for food close to where they hatched. They feed on wood-borer insects on trees, or feed on their species' substrate or prey of choice. Feeding is the main purpose of the larval stage to prepare for pupation. Once their larval stage is complete the tree dwelling species make their way to the bottom of the tree to pupate. 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 stage can last from 6 weeks to one year depending on the need to overwinter, and how short the overall lifecycle is for a particular species. A majority of clerid species pupate in earthen cells which are made from soil and certain enzymes secreted from their mouths. The rest remain in pupal cells. Adult beetles emerge from pupation and spend a variable time of their life maturing, and eventually
oviposit The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
. Sexually mature adults or
imago In biology, the imago (Latin for "image") is the last stage an insect attains during its metamorphosis, its process of growth and development; it is also called the imaginal stage, the stage in which the insect attains maturity. It follows the f ...
s of ''Thanasiumus'' overwinter inside the wood-borer-infested trees and oviposit during the spring.


Forensic relevance


Stored product entomology

'' Necrobia rufipes'', commonly known as the red-legged ham beetle, is of particular importance in stored product entomology. ''N. rufipes'' infests dried or smoked meats, especially those products that are stored unwrapped for long periods of time. Adults feed on the surface of the products, while the larvae damage the meat by boring down usually in the fatty parts. ''N. rufipes'' has been recorded to have fed upon a large variety of items ranging from hides and dried figs to Egyptian mummies. In addition, products such as
wool Wool is the textile fibre obtained from sheep and other mammals, especially goats, rabbits, and camelids. The term may also refer to inorganic materials, such as mineral wool and glass wool, that have properties similar to animal wool. ...
and
silk Silk is a natural protein fiber, some forms of which can be woven into textiles. The protein fiber of silk is composed mainly of fibroin and is produced by certain insect larvae to form cocoons. The best-known silk is obtained from the ...
can become infested, but not destroyed.


Medico-legal entomology

Since clerids are predaceous in nature, they have been found feeding on fly larvae as well as the skin and bones of carrion. Most clerids are not useful in forensics because of their food choice, but some species such as ''Necrobia rufipes'' can be useful. ''Necrobia rufipes'' is attracted towards carrion in the later stages of decomposition, so its arrival on carrion can help provide an estimate for the
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 ...
or PMI. Although the checkered beetle is not the most significant insect on carrion, the beetles predaceous nature and its ability to reproduce in carrion that is exposed to the environment provides some forensic importance.


Ongoing research

There is ongoing research with some clerid species. Forensic research is limited because of their late arrival on carrion, but members such as '' Thanasimus undatulus'' have been researched as a possible role in
integrated pest management Integrated pest management (IPM), also known as integrated pest control (IPC) is a broad-based approach that integrates both chemical and non-chemical practices for economic control of pests. IPM aims to suppress pest populations below the econ ...
or IPM. ''Thanasimus undatulus'' is a predator of bark beetles. Some species of bark beetles such as the southern pine beetle and the
mountain pine beetle The mountain pine beetle (''Dendroctonus ponderosae'') is a species of bark beetle native to the forests of western North America from Mexico to central British Columbia. It has a hard black exoskeleton, and measures approximately , about the siz ...
can become pests to the
lumber Lumber is wood that has been processed into dimensional lumber, including beams and planks or boards, a stage in the process of wood production. Lumber is mainly used for construction framing, as well as finishing (floors, wall panels, wi ...
industry because in large numbers they can cause damage and kill live trees. ''Thanasimus undatulus'' has been researched as a possible
biological control Biological control or biocontrol is a method of controlling pests, such as insects, mites, weeds, and plant diseases, using other organisms. It relies on predation, parasitism, herbivory, or other natural mechanisms, but typically also i ...
agent for these pests. Researchers and forestry officials have used bark beetle aggregation pheromones to attract the checkered beetle to specific trees. This causes the bark beetles to be overwhelmed, extensively preyed upon by the clerid beetles, and typically eliminated. There is also additional research being done pertaining to the impact of clerids on pollination in flowers.


Systematics

The genera of Cleridae are divided among several subfamilies, though some genera still defy easy classification. Several taxonomic schemes exist, recognizing for example a group around '' Neorthopleura'' as distinct subfamily Neorthopleurinae, or splitting off the Thaneroclerinae as distinct family, or circumscribing the Korynetinae '' sensu stricto'' or ''
sensu lato ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular c ...
''. The following list of tribes and selected genera is thus preliminary. Some notable species are also listed. The oldest members of the family are '' Protoclerus'' and '' Wangweiella'' the late Middle Jurassic (
Callovian In the geologic timescale, the Callovian is an age and stage in the Middle Jurassic, lasting between 166.1 ± 4.0 Ma (million years ago) and 163.5 ± 4.0 Ma. It is the last stage of the Middle Jurassic, following the Bathonian and preceding the ...
) Daohugou bed in Inner Mongolia, China.
Clerinae * '' Allonyx'' Jacquelin du Val, 1860 * '' Anthicoclerus'' Schenkling, 1906 * '' Aphelochroa'' Quedenfeldt, 1885 * '' Apopempsis'' Schenkling, 1903 * '' Apteroclerus'' Wollaston, 1867 * '' Aptinoclerus'' Kuwert, 1893 * '' Aradamicula'' Sedlacek & Winkler, 1975 * †'' Arawakis'' (
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
) * '' Astigmus'' Kuwert, 1894 * '' Aulicus'' Spinola, 1841 * '' Axina'' Kirby, 1818 * '' Balcus'' * '' Barriella'' Opitz, 2003 * '' Barrotillus'' Rifkind, 1996 * '' Blaxima'' Gorham, 1882 * '' Bousquetoclerus'' Menier, 1997 * '' Burgeoneus'' Pic, 1950 * '' Caestron'' Dupont in Spinola, 1844 * '' Calendyma'' Lacordaire, 1857 * '' Canariclerus'' Winkler, 1982 * '' Cardiostichus'' Quedenfeldt, 1885 * '' Caridopus'' Schenkling in Sjöstedt, 1908 * '' Cleromorpha'' Gorham, 1876 * '' Cleropiestus'' Fairmaire, 1889 * '' Clerus'' Fabricius, 1775 * '' Clytomadius'' Corporaal, 1949 * '' Colyphus'' Spinola, 1841 * '' Coptoclerus'' Chapin, 1924 * '' Cormodes'' Pascoe, 1860 * '' Corynommadius'' Schenkling, 1899 * '' Ctenaxina'' Schenkling, 1906 * '' Ctenoclerus'' Solervicens, 1997 * '' Dasyceroclerus'' Kuwert, 1894 * '' Dasyteneclines'' Pic, 1941 * '' Dieropsis'' Gahan, 1908 * '' Dologenitus'' Opitz, 2009 * '' Dozocolletus'' Chevrolat, 1842 * '' Eburiphora'' Spinola, 1841 * '' Eburneoclerus'' Pic, 1950 * '' Ekisius'' Winkler, 1987 * '' Eleale'' Newman, 1840 * '' Enoclerus'' Gahan, 1910 * '' Epiclines'' Chevrolat in Guérin-Ménéville, 1839 * '' Eunatalis'' Schenkling, 1909 ** '' Eunatalis porcata'' * '' Erymanthus'' Spinola, 1841 * '' Eurymetomorphon'' Pic, 1950 * '' Falsomadius'' Gerstmeier, 2002 * '' Falsoorthrius'' Pic, 1940 * '' Graptoclerus'' Gorham, 1901 * '' Gyponyx'' Gorham, 1883 * '' Hemitrachys'' Gorham, 1876 * '' Homalopilo'' Schenkling, 1915 * '' Inhumeroclerus'' Pic, 1955 * '' Jenjouristia'' Fursov, 1936 * '' Languropilus'' Pic, 1940 * '' Lissaulicus'' C.O.Waterhouse, 1879 * '' Memorthrius'' Pic, 1940 * '' Metademius'' Schenkling, 1899 * '' Microclerus'' Wollaston, 1867 * '' Micropteroclerus'' Chapin, 1920 * '' Microstigmatium'' Kraatz, 1899 * '' Mimolesterus'' Gerstmeier, 1991 * '' Mitrandiria'' Kolibac, 1997 * '' Myrmecomaea'' Fairmaire, 1886 * '' Natalis'' Laporte de Castelnau, 1836 * '' Neogyponyx'' Schenkling, 1906 * '' Neoscrobiger'' Blackburn, 1900 * '' Ohanlonella'' Rifkind, 2008 * '' Olesterus'' Spinola, 1841 * '' Omadius'' Laporte de Castelnau, 1836 * '' Oodontophlogistus'' Elston, 1923 * '' Operculiphorus'' Kuwert, 1894 * '' Opilo'' Latreille, 1802 * '' Orthrius'' Gorham, 1876 * '' Oxystigmatium'' Kraatz, 1899 * '' Phlogistomorpha'' Hintz, 1908 * '' Phlogistus'' Gorham, 1876 * '' Phloiocopus'' Spinola, 1841 * '' Phonius'' Chevrolat, 1843 * '' Pieleus'' Pic, 1940 * '' Placocerus'' Klug, 1837 * '' Placopterus'' Wolcott, 1910 * '' Plathanocera'' Schenkling, 1902 * '' Platyclerus'' Spinola, 1841 * '' Priocera'' Kirby, 1818 * '' Priocleromorphus'' Pic, 1950 * '' Prioclerus'' Hintz, 1902 * '' Pseudolesterus'' Miyatake, 1968 * '' Pseudomadius'' Chapin, 1924 * '' Pujoliclerus'' Pic, 1947 * '' Sallea'' Chevrolat, 1874 * '' Scrobiger'' Spinola, 1841 * '' Sedlacekius'' Winkler, 1972 * '' Sikorius'' Kuwert, 1893 * '' Stigmatium'' Gray in Griffith, 1832 * '' Systenoderes'' Spinola, 1841 * '' Tanocleria'' Hong, 2002 * '' Thalerocnemis'' Lohde, 1900 * '' Thanasimodes'' Murray, 1867 * '' Thanasimus'' Latreille, 1806 ** '' Thanasimus formicarius'' – Ant Beetle * '' Tillicera'' Spinola, 1841 * '' Trichodes'' Herbst, 1792 ** '' Trichodes alvearius'' ** '' Trichodes apiarius'' ** '' Trichodes leucopsideus'' * '' Trogodendron'' Spinola, 1841 ** '' Trogodendron fasciculatum'' – Yellow-horned Clerid * '' Wilsonoclerus'' * '' Winklerius'' Menier, 1986 * '' Wittmeridecus'' Winkler, 1981 * '' Xenorthrius'' Gorham, 1892 * '' Zahradnikius'' Winkler, 1992 * '' Zenithicola'' Spinola, 1841 Enopliinae (sometimes in Korynetinae) * '' Antygodera'' * '' Apolopha'' Spinola, 1841 * '' Corinthiscus'' Fairmaire & Germain, 1861 * '' Cregya'' LeConte, 1861 * '' Curacavi'' * '' Enoplium'' Latreille, 1802 * '' Exochonotus'' * '' Hublella'' * '' Lasiodera'' Gray in Griffith, 1832 * '' Neopylus'' Solervicens, 1989 * '' Paracregya'' * '' Pelonium'' * '' Phymatophaea'' Pascoe, 1876 * '' Platynoptera'' Chevrolat, 1834 * '' Pseudichnea'' Schenkling, 1900 * '' Pylus'' Newman, 1840 * '' Pyticara'' Spinola, 1841 (including ''Pelonides'') * '' Teneroides'' Gahan, 1910 * '' Tenerus'' Laporte de Castelnau, 1836 * '' Thriocerodes'' Wolcott & Dybas, 1947 Epiphloeinae (sometimes in Korynetinae) * '' Acanthocollum'' * '' Amboakis'' * '' Decaphloeus'' * '' Decorosa'' Opitz, 2008 * '' Diapromeces'' Opitz, 1997 * '' Ellipotoma'' Spinola, 1844 * '' Epiphloeus'' Spinola, 1841 * '' Hapsidopteris'' Opitz, 1997 * '' Ichnea'' Laporte de Castelnau, 1836 * '' Iontoclerus'' Opitz, 1997 * '' Katamyurus'' Opitz, 1997 * '' Madoniella'' Pic, 1935 * '' Megaphloeus'' * '' Megatrachys'' Opitz, 1997 * '' Opitzius'' Barr, 2006 * '' Parvochaetus'' Opitz, 2006 * '' Pennasolis'' Opitz, 2008 * '' Pericales'' * '' Pilosirus'' Opitz, 1997 * '' Plocamocera'' Spinola, 1844 * '' Pteroferus'' * '' Pyticeroides'' Kuwert, 1894 * '' Silveirasia'' * '' Stegnoclava'' * '' Turbophloeus'' Hydnocerinae (including Phyllobaeninae) * '' Abrosius'' Fairmaire, 1902 * '' Achlamys'' C.O.Waterhouse, 1879 * '' Allelidea'' G.R.Waterhouse, 1839 * '' Blaesiophthalmus'' Schenkling, 1903 * '' Brachycallimerus'' Chapin, 1924 * '' Brachyptevenus'' * '' Callimerus'' Gorham, 1876 * '' Cephaloclerus'' Kuwert, 1893 * '' Cucujocallimerus'' Pic, 1929 * '' Emmepus'' Motschulsky, 1845 * '' Eurymetopum'' Blanchard, 1842 * '' Isohydnocera'' Chapin, 1917 * '' Isolemidia'' Gorham, 1877 * '' Laiomorphus'' Pic, 1927 * '' Lasiocallimerus'' Corporaal, 1939 * '' Lemidia'' Spinola, 1841 * '' Neohydnus'' Gorham, 1892 * '' Parmius'' Sharp, 1877 * '' Paupris'' Sharp, 1877 * '' Phyllobaenus'' Dejean, 1837 * '' Silviella'' Solervicens, 1987 * '' Solemidia'' * '' Stenocallimerus'' Corporaal & Pic, 1940 * '' Theano'' Laporte de Castelnau, 1836 * '' Wolcottia'' Chapin, 1917 Korynetinae * '' Chariessa'' Perty in Spix & Martius, 1830 * '' Korynetes'' Herbst, 1792 ** '' Korynetes caeruleus'' – steely blue beetle * '' Lebasiella'' Spinola, 1844 * '' Loedelia'' R.Lucas, 1918 * '' Necrobia'' Latreille, 1797 ** '' Necrobia ruficollis'' – red-shouldered ham beetle ** '' Necrobia rufipes'' – red-legged ham beetle * '' Neorthopleura'' Barr, 1976 * '' Opetiopalpus'' Spinola, 1844 * '' Romanaeclerus'' Winkler, 1960 Tarsosteninae (sometimes in Korynetinae) * '' Paratillus'' Gorham, 1876 * '' Tarsostenodes'' Blackburn, 1900 * '' Tarsostenus'' Spinola, 1844 Thaneroclerinae (tentatively placed here) * '' Cleridopsis'' Champion, 1913 * '' Compactoclerus'' Pic, 1939 * '' Cyrtinoclerus'' Chapin, 1924 * '' Isoclerus'' Lewis, 1892 * '' Meprinogenus'' Kolibáč, 1992 * '' Neoclerus'' Lewis, 1892 * '' Onerunka'' Kolibáč * '' Thaneroclerus'' Lefebvre, 1838 * '' Viticlerus'' * '' Zenodosus'' Wolcott, 1910 Tillinae * '' Antenius'' Fairmaire, 1903 * '' Arachnoclerus'' Fairmaire, 1902 * '' Araeodontia'' Barr, 1952 * '' Archalius'' Fairmaire, 1903 * '' Aroterus'' Schenkling, 1906 * '' Basilewskyus'' Pic, 1950 * '' Biflabellotillus'' Pic, 1949 * '' Bilbotillus'' Kolibac, 1997 * '' Bogcia'' Barr, 1978 * '' Bostrichoclerus'' Van Dyke, 1938 * '' Callotillus'' Wolcott, 1911 * '' Ceratocopus'' Hintz, 1902 * '' Chilioclerus'' Solervicens, 1976 * '' Cladiscopallenis'' Pic, 1949 * '' Cladiscus'' Chevrolat, 1843 * '' Cladomorpha'' Pic, 1949 * '' Cteniopachys'' Fairmaire, 1889 * '' Cylidroctenus'' Kraatz, 1899 * '' Cylidrus'' Latreille, 1825 * '' Cymatodera'' Gray in Griffith, 1832 * '' Cymatoderella'' Barr, 1962 * '' Dedana'' Fairmaire, 1888 * '' Denops'' Fischer von Waldheim, 1829 * '' Diplocladus'' Fairmaire, 1885 * '' Diplopherusa'' Heller, 1921 * '' Eburneocladiscus'' Pic, 1955 * '' Egenocladiscus'' Corporaal & van der Wiel, 1949 * '' Elasmocylidrus'' Corporaal, 1939 * '' Enoploclerus'' Hintz, 1902 * '' Eucymatodera'' Schenkling, 1899 * '' Falsopallenis'' Pic, 1926 * '' Falsotillus'' Gerstmeier & Kuff, 1992 * '' Flabellotilloidea'' Gerstmeier & Kuff, 1992 * '' Gastrocentrum'' Gorham, 1876 * '' Gracilotillus'' Pic, 1933 * '' Impressopallenis'' Pic, 1953 * '' Isocymatodera'' Hintz, 1902 * '' Lecontella'' Wolcott & Chapin, 1918 * '' Leptoclerus'' Kraatz, 1899 * '' Liostylus'' Fairmaire, 1886 * '' Macroliostylus'' Pic, 1939 * '' Magnotillus'' Pic, 1936 * '' Melanoclerus'' Chapin, 1919 * '' Microtillus'' Pic, 1950 * '' Monophylla'' Spinola, 1841 * '' Neocallotillus'' Burke, 2016 * '' Nodepus'' Gorham, 1892 * '' Notocymatodera'' Schenkling, 1907 * '' Onychotillus'' Chapin, 1945 * '' Orthocladiscus'' Corporaal & van der Wiel, 1949 * '' Pallenis'' Laporte de Castelnau, 1836 * '' Paracladiscus'' Miyatake, 1965 * '' Paradoxocerus'' Kraatz, 1899 * '' Paraspinoza'' Corporaal, 1942 * '' Philocalus'' Klug, 1842 * '' Picoclerus'' Corporaal, 1936 * †'' Prospinoza'' (fossil) * '' Pseudachlamys'' Duvivier, 1892 * '' Pseudogyponix'' Pic, 1939 * '' Pseudopallenis'' Kuwert, 1893 * '' Pseudoteloclerus'' Pic, 1932 * '' Rhopaloclerus'' Fairmaire, 1886 * '' Smudlotillus'' Kolibac, 1997 * ''
Spinoza Baruch (de) Spinoza (born Bento de Espinosa; later as an author and a correspondent ''Benedictus de Spinoza'', anglicized to ''Benedict de Spinoza''; 24 November 1632 – 21 February 1677) was a Dutch philosopher of Portuguese-Jewish origin, ...
'' Lewis, 1892 * '' Stenocylidrus'' Spinola, 1844 * '' Strotocera'' Schenkling, 1902 * '' Synellapotillus'' Pic, 1939 * '' Synellapus'' Fairmaire, 1903 * '' Teloclerus'' Schenkling, 1903 * '' Tilloclerus'' White, 1849 * '' Tillodadiscus'' Pic, 1953 * '' Tillodenops'' Hintz, 1905 * '' Tilloidea'' Laporte de Castelnau, 1832 * '' Tillus'' Olivier, 1790 * '' Tylotosoma'' Hintz, 1902 ''
Incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertain ...
'' * '' Aphelocerus'' Kirsch, 1871 (Clerinae? Tillinae?) * '' Apteropilo'' Lea, 1908 (Clerinae? Enopliinae?) * '' Cleropiestus'' Fairmaire, 1889 (Clerinae? Hydnocerinae?) * '' Dermestoides'' Schaeffer, 1771 (Korynetinae '' s.l.''?) * '' Evenoclerus'' Corporaal, 1950 (Clerinae? Hydnocerinae?) * ''
Muisca The Muisca (also called Chibcha) are an indigenous people and culture of the Altiplano Cundiboyacense, Colombia, that formed the Muisca Confederation before the Spanish conquest. The people spoke Muysccubun, a language of the Chibchan langu ...
'' Spinola, 1844 (Clerinae? Enopliinae?) * '' Parapelonides'' Barr, 1980 (Korynetinae ''s.l.''?) * '' Perilypus'' Spinola, 1841 (Clerinae? Tillinae?) * '' Syriopelta'' Winkler, 1984 (Korynetinae ''s.l.''?)


References


External links


Atlas of checkered beetles (Cleridae) of Russia




{{Taxonbar, from=Q377988 Polyphaga families Taxa named by Pierre André Latreille