Cleridae

Cleridae Temporal range: Middle Jurassic–Recent PreꞒ Ꞓ O S D C P T J K Pg N
Some checkered beetles described in the mid-19th century
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Coleoptera
Suborder:	Polyphaga
Infraorder:	Cucujiformia
Superfamily:	Cleroidea
Family:	Cleridae Latreille, 1802
Subfamilies
Clerinae Latreille, 1802 Enopliinae Gistel, 1856 (disputed) Epiphloeinae Gistel, 1856 (disputed) Hydnocerinae Spinola, 1844 Korynetinae Laporte, 1836 Tarsosteninae Jacquelin du Val, 1861 (disputed) Thaneroclerinae Chapin, 1921[ verification needed ] (but see text) Tillinae Leach, 1815 and see below

Cleridae are a family of beetles of the superfamily Cleroidea. They are commonly known as checkered beetles. The family Cleridae has a worldwide distribution, and a variety of habitats and feeding preferences.

Cleridae have many niches and feeding habits. Most genera are predaceous and feed on other beetles and larvae; however other genera are scavengers or pollen feeders. Clerids have elongated bodies with bristly hairs, are usually bright colored, and have variable antennae. Checkered beetles range in length between 3 and 24 millimetres (0.12 and 0.94 in). 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 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. 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

Narrow pronotum in Enoclerus ichneumoneus (Clerinae)

Appearance

Generally, checkered beetles are elongated and oval in shape and range from 3–24 millimetres (0.12–0.94 in) in length. ^[1] Their entire bodies are covered with bristly hairs and many display an ornate body color pattern. ^[1] These often brightly color patterns can be red, yellow, orange, or blue. ^[2] The antennae are clubbed at the tip for most species, but others can be "clubbed, saw-tooth, or thread-like." ^{[1] [2]} The pronotum region is nearly cylindrical and characteristically narrower than the elytra (special hardened front wings), while the head is as wide or wider than the pronotum. ^[2] Their elytra have tiny pits or depressions, and never expose more than two tergites (dorsal plates). ^[1]

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. Also, the notopleural suture (found under the pronotal shield) is not present. ^[1] To further identify Clerid beetles, a few additional characteristics need to be examined.

5 rear leg tarsomeres of Tillus elongatus (Tillinae)

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). ^[1] One or more of these subsegments on each leg is typically lobed, and the 4th tarsomere 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. ^[1]

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 and pronotum. This characteristic distinguishes them from a similar family Melyridae which sometimes has these glands. ^[1] This trait is very important in correctly differentiating checkered beetles from Melyridae.

Distribution and ecology

Trichodes ornatus (Clerinae) on a flower

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. ^[3] 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 visiting insects and also feed on pollen. These species are found in moist, sunny environments where flowering plants are found in abundance. ^[4]

Another habitat commonly inhabited by clerid beetles is trees. 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. ^[4] 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. ^[4] Not all nest robbing species actively hunt live prey, some species for example prefer to feed only on dead honey bee larvae and adults. ^[4]

Feeding habits

The Cleridae contains many species of predaceous beetles that feed on other beetles and beetle larvae in their natural habitat. ^[3] The most common prey item for checkered beetles are bark beetles and wood boring beetles. ^[5]

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. ^[6]

Although most species of checkered beetles are predaceous in nature, some are scavengers and others have been found feeding on flower pollen. ^[7] Because of the checkered beetles predaceous nature and insatiable appetite, they are often key players in the biological control of other insects. The checkered beetles have also developed a unique adaptation to aid in their quest for prey. The beetles use pheromones to help them locate, kill, and consume their prey. ^[8]

Necrobia rufipes (Korynetinae)

The diversity of checkered beetle's feeding habits is quite evident when different species are examined. Necrobia spp. are attracted to dry carrion and other decomposing animal matter such as bones and skin as well as various meat products. ^[9] Thanasimus spp. are found in woodland areas where bark beetle species constitute their main source of prey. ^[9] The primary source of prey for Phyllobaenus spp. are wood borers, immature weevils, and hymenoptera larvae. ^[9] One of the more diverse genera is Trichodes , in which larvae feed on the pollen of flowering plants and adults prey upon grasshoppers and wasps. ^[9]

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. ^[10] 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 life cycle, and the cooler the temperature is the slower the life cycle. 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 life cycle: the egg hatches into a larva, which grows and feeds, changing its skin to form a pupa, 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. ^[11]

Larva of Thanasimus dubius

Copulation takes place while the female feeds, because females need a large amount of food for egg development. ^[12] 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. ^[11] She may lay 28–42 eggs at a time. ^[12] For the longer-living species such as Thanasimus, this occurs in late summer or early fall to give the larvae enough time for proper growth before having to overwinter. ^[13]

When larvae hatch from their eggs, they are either red or yellow. ^[11] 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 area of the last body segment. ^[11] 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. ^[11] 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. ^[13] The pupal stage can last from 6 weeks to one year depending on the need to overwinter, and how short the overall life cycle 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. ^[11] The rest remain in pupal cells. Adult beetles emerge from pupation and spend a variable time of their life maturing, and eventually oviposit. Sexually mature adults or imagos of Thanasiumus overwinter inside the wood-borer-infested trees and oviposit during the spring. ^[12]

Forensic relevance

Stored product entomology

Necrobia rufipes , commonly known as the red-legged ham beetle, is of particular importance in stored product entomology. It 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. ^[14] Necrobia rufipes has been recorded feeding upon a large variety of items ranging from hides and dried figs to Egyptian mummies. ^[14] In addition, products such as wool and silk can become infested, but not destroyed. ^[14]

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. ^[9] 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 or PMI. Although the checkered beetle is not the most significant insect on carrion, the beetle's predaceous nature and its ability to reproduce in carrion that is exposed to the environment provides some forensic importance. ^[15]

Thanasimus dubius attacking bark beetle prey

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 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 can become pests to the lumber industry because in large numbers they can cause damage and kill live trees. Thanasimus undatulus has been researched as a possible biological control 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. ^[16] There is also additional research being done pertaining to the impact of clerids on pollination in flowers. ^[17]

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 . 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) Daohugou bed in Inner Mongolia, China. ^[18]

Clerus mutillarius (Clerinae)

Phymatophaea guttigera (Enopliinae)

Lemidia aptera (Hydnocerinae)

Necrobia violacea (Korynetinae)

Tarsostenus univittatus (Tarsosteninae)

Diplocladus kuwerti (Tillinae)

Dermestoides sanguinicollis ( incertae sedis )

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) 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 [19] 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 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 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 Spinola, 1844 (Clerinae? Enopliinae?) Parapelonides Barr, 1980 (Korynetinae s.l.?) Perilypus Spinola, 1841 (Clerinae? Tillinae?) Syriopelta Winkler, 1984 (Korynetinae s.l.?)

References

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External links

• Atlas of checkered beetles (Cleridae) of Russia
• Roland Gerstmeier publications on Cleridae
• Cleridae of Atlantic Canada