Cleridae
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. 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 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 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, while the head is as wide or wider than the pronotum. Their elytra have tiny pits or depressions, and never expose more than two tergites.Identification
Clerid beetles fall under the suborder Polyphaga. Key characteristics of Polyphaga are that the hind coxa, do not divide the first and second abdominal/ventral plates which are known as sternites. Also, the notopleural suture 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 five tarsomeres. One or more of these subsegments on each leg is typically lobed, and the fourth tarsomere is normally difficult to distinguish. Furthermore, an important feature that eliminates many other families of beetles is that clerids' front coxae 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 on their abdomen and pronotum. This characteristic distinguishes them from a similar family Melyridae 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 visiting insects 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. 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 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. 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.
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. Thanasimus spp. are found in woodland areas where bark beetle species constitute their main source of prey. The primary source of prey for Phyllobaenus spp. are wood borers, immature weevils, and hymenoptera larvae. 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.
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 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.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-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.
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 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 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. 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.
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. Necrobia rufipes has been recorded feeding upon a large variety of items ranging from hides and dried figs to Egyptian mummies. In addition, products such as wool and silk 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 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.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.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. 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 Daohugou bed in Inner Mongolia, China.ClerinaeAllonyx Jacquelin du Val, 1860Anthicoclerus Schenkling, 1906Aphelochroa Quedenfeldt, 1885Apopempsis Schenkling, 1903Apteroclerus Wollaston, 1867Aptinoclerus Kuwert, 1893Aradamicula Sedlacek & Winkler, 1975
- †Arawakis Astigmus Kuwert, 1894Aulicus Spinola, 1841Axina Kirby, 1818BalcusBarriella Opitz, 2003Barrotillus Rifkind, 1996Blaxima Gorham, 1882Bousquetoclerus Menier, 1997Burgeoneus Pic, 1950Caestron Dupont in Spinola, 1844Calendyma Lacordaire, 1857Canariclerus Winkler, 1982Cardiostichus Quedenfeldt, 1885Caridopus Schenkling in Sjöstedt, 1908Cleromorpha Gorham, 1876Cleropiestus Fairmaire, 1889Clerus Fabricius, 1775Clytomadius Corporaal, 1949Colyphus Spinola, 1841Coptoclerus Chapin, 1924Cormodes Pascoe, 1860Corynommadius Schenkling, 1899Ctenaxina Schenkling, 1906Ctenoclerus Solervicens, 1997Dasyceroclerus Kuwert, 1894Dasyteneclines Pic, 1941Dieropsis Gahan, 1908Dologenitus Opitz, 2009Dozocolletus Chevrolat, 1842Eburiphora Spinola, 1841Eburneoclerus Pic, 1950Ekisius Winkler, 1987Eleale Newman, 1840Enoclerus Gahan, 1910Epiclines Chevrolat in Guérin-Ménéville, 1839Eunatalis Schenkling, 1909
- * Eunatalis porcataErymanthus Spinola, 1841Eurymetomorphon Pic, 1950Falsomadius Gerstmeier, 2002Falsoorthrius Pic, 1940Graptoclerus Gorham, 1901Gyponyx Gorham, 1883Hemitrachys Gorham, 1876Homalopilo Schenkling, 1915Inhumeroclerus Pic, 1955Jenjouristia Fursov, 1936Languropilus Pic, 1940Lissaulicus C.O.Waterhouse, 1879Memorthrius Pic, 1940Metademius Schenkling, 1899Microclerus Wollaston, 1867Micropteroclerus Chapin, 1920Microstigmatium Kraatz, 1899Mimolesterus Gerstmeier, 1991Mitrandiria Kolibac, 1997Myrmecomaea Fairmaire, 1886Natalis Laporte de Castelnau, 1836Neogyponyx Schenkling, 1906Neoscrobiger Blackburn, 1900Ohanlonella Rifkind, 2008Olesterus Spinola, 1841Omadius Laporte de Castelnau, 1836Oodontophlogistus Elston, 1923Operculiphorus Kuwert, 1894Opilo Latreille, 1802Orthrius Gorham, 1876Oxystigmatium Kraatz, 1899Phlogistomorpha Hintz, 1908Phlogistus Gorham, 1876Phloiocopus Spinola, 1841Phonius Chevrolat, 1843Pieleus Pic, 1940Placocerus Klug, 1837Placopterus Wolcott, 1910Plathanocera Schenkling, 1902Platyclerus Spinola, 1841Priocera Kirby, 1818Priocleromorphus Pic, 1950Prioclerus Hintz, 1902Pseudolesterus Miyatake, 1968Pseudomadius Chapin, 1924Pujoliclerus Pic, 1947Sallea Chevrolat, 1874Scrobiger Spinola, 1841Sedlacekius Winkler, 1972Sikorius Kuwert, 1893Stigmatium Gray in Griffith, 1832Systenoderes Spinola, 1841Tanocleria Hong, 2002Thalerocnemis Lohde, 1900Thanasimodes Murray, 1867Thanasimus Latreille, 1806
- * Thanasimus formicarius - Ant BeetleTillicera Spinola, 1841Trichodes Herbst, 1792
- * Trichodes alvearius
- * Trichodes apiarius
- * Trichodes leucopsideusTrogodendron Spinola, 1841
- * Trogodendron fasciculatum - Yellow-horned CleridWilsonoclerusWinklerius Menier, 1986Wittmeridecus Winkler, 1981Xenorthrius Gorham, 1892Zahradnikius Winkler, 1992Zenithicola Spinola, 1841
Epiphloeinae AcanthocollumAmboakisDecaphloeusDecorosa Opitz, 2008Diapromeces Opitz, 1997Ellipotoma Spinola, 1844Epiphloeus Spinola, 1841Hapsidopteris Opitz, 1997Ichnea Laporte de Castelnau, 1836Iontoclerus Opitz, 1997Katamyurus Opitz, 1997Madoniella Pic, 1935MegaphloeusMegatrachys Opitz, 1997Opitzius Barr, 2006Parvochaetus Opitz, 2006Pennasolis Opitz, 2008PericalesPilosirus Opitz, 1997Plocamocera Spinola, 1844PteroferusPyticeroides Kuwert, 1894SilveirasiaStegnoclavaTurbophloeus
Hydnocerinae Abrosius Fairmaire, 1902Achlamys C.O.Waterhouse, 1879Allelidea G.R.Waterhouse, 1839Blaesiophthalmus Schenkling, 1903Brachycallimerus Chapin, 1924BrachyptevenusCallimerus Gorham, 1876Cephaloclerus Kuwert, 1893Cucujocallimerus Pic, 1929Emmepus Motschulsky, 1845Eurymetopum Blanchard, 1842Isohydnocera Chapin, 1917Isolemidia Gorham, 1877Laiomorphus Pic, 1927Lasiocallimerus Corporaal, 1939Lemidia Spinola, 1841Neohydnus Gorham, 1892Parmius Sharp, 1877Paupris Sharp, 1877Phyllobaenus Dejean, 1837Silviella Solervicens, 1987SolemidiaStenocallimerus Corporaal& Pic, 1940Theano Laporte de Castelnau, 1836Wolcottia Chapin, 1917
KorynetinaeChariessa Perty in Spix & Martius, 1830Korynetes Herbst, 1792
- * Korynetes caeruleus - steely blue beetleLebasiella Spinola, 1844Loedelia R.Lucas, 1918Necrobia Latreille, 1797
- * Necrobia ruficollis - red-shouldered ham beetle
- * Necrobia rufipes - red-legged ham beetleNeorthopleura Barr, 1976Opetiopalpus Spinola, 1844Romanaeclerus Winkler, 1960
Thaneroclerinae Cleridopsis Champion, 1913Compactoclerus Pic, 1939Cyrtinoclerus Chapin, 1924Isoclerus Lewis, 1892Meprinogenus Kolibáč, 1992Neoclerus Lewis, 1892Onerunka KolibáčThaneroclerus Lefebvre, 1838ViticlerusZenodosus Wolcott, 1910
TillinaeAntenius Fairmaire, 1903Arachnoclerus Fairmaire, 1902Araeodontia Barr, 1952Archalius Fairmaire, 1903Aroterus Schenkling, 1906Basilewskyus Pic, 1950Biflabellotillus Pic, 1949Bilbotillus Kolibac, 1997Bogcia Barr, 1978Bostrichoclerus Van Dyke, 1938Callotillus Wolcott, 1911Ceratocopus Hintz, 1902Chilioclerus Solervicens, 1976Cladiscopallenis Pic, 1949Cladiscus Chevrolat, 1843Cladomorpha Pic, 1949Cteniopachys Fairmaire, 1889Cylidroctenus Kraatz, 1899Cylidrus Latreille, 1825Cymatodera Gray in Griffith, 1832Cymatoderella Barr, 1962Dedana Fairmaire, 1888Denops Fischer von Waldheim, 1829Diplocladus Fairmaire, 1885Diplopherusa Heller, 1921Eburneocladiscus Pic, 1955Egenocladiscus Corporaal& van der Wiel, 1949Elasmocylidrus Corporaal, 1939Enoploclerus Hintz, 1902Eucymatodera Schenkling, 1899Falsopallenis Pic, 1926Falsotillus Gerstmeier & Kuff, 1992Flabellotilloidea Gerstmeier & Kuff, 1992Gastrocentrum Gorham, 1876Gracilotillus Pic, 1933Impressopallenis Pic, 1953Isocymatodera Hintz, 1902Lecontella Wolcott & Chapin, 1918Leptoclerus Kraatz, 1899Liostylus Fairmaire, 1886Macroliostylus Pic, 1939Magnotillus Pic, 1936Melanoclerus Chapin, 1919Microtillus Pic, 1950Monophylla Spinola, 1841Neocallotillus Burke, 2016Nodepus Gorham, 1892Notocymatodera Schenkling, 1907Onychotillus Chapin, 1945Orthocladiscus Corporaal& van der Wiel, 1949Pallenis Laporte de Castelnau, 1836Paracladiscus Miyatake, 1965Paradoxocerus Kraatz, 1899Paraspinoza Corporaal, 1942Philocalus Klug, 1842Picoclerus Corporaal, 1936
- †Prospinoza Pseudachlamys Duvivier, 1892Pseudogyponix Pic, 1939Pseudopallenis Kuwert, 1893Pseudoteloclerus Pic, 1932Rhopaloclerus Fairmaire, 1886Smudlotillus Kolibac, 1997Spinoza Lewis, 1892Stenocylidrus Spinola, 1844Strotocera Schenkling, 1902Synellapotillus Pic, 1939Synellapus Fairmaire, 1903Teloclerus Schenkling, 1903Tilloclerus White, 1849Tillodadiscus Pic, 1953Tillodenops Hintz, 1905Tilloidea Laporte de Castelnau, 1832Tillus Olivier, 1790Tylotosoma Hintz, 1902