Cleridae

Superregnum: Eukaryota
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Cladus: Metapterygota
Infraclassis: Neoptera
Cladus: Eumetabola
Cladus: Endopterygota
Superordo: Coleopterida
Ordo: Coleoptera
Subordo: Polyphaga
Infraordo: Cucujiformia
Superfamilia: Cleroidea

Familia: Cleridae
Subfamiliae (11): Clerinae - Enopliinae - Epiclininae - Epiphloeinae - Hydnocerinae - Isoclerinae - Korynetinae - Neorthopleurinae - Peloniinae - Tarsosteninae - Tillinae
Overview of genera

Aberrokorynetes Winkler, 1990
Abiliella Peracchi, 1964
Abrosius Fairmaire, 1902
Achlamys Waterhouse, 1879
Agapetilus Opitz, 2012
Agaphalera Opitz, 2009
Agnatis Opitz, 2014
Akonesis Opitz, 2014
Allelidea Waterhouse, 1839
Allelidia
Allochotes Westwood, 1875
Allonyx Duval, 1861
Allonyx Jacquelin du Val, 1860
Allothaneroclerus Corporaal, 1939
Amboakis Opitz, 2006
Amphelissus Opitz, 2017
Ampleris
Anebomorpha Poinar Jr. & Brown, 2020
Anisophyllus Westwood, 1876
Antenius Fairmaire, 1902
Antennosus Opitz, 2014
Anthicoclerus Schenkling, 1906
Aphelocerus Kirsch, 1871
Aphelochroa Quedenfeldt, 1885
Apolopha Spinola, 1841
Apopempsis Schenkling, 1903
Apopylus Kolibac, 2003
Apteroclerus Wollaston, 1867
Apteropilo Lea, 1908
Aptinoclerus Kuwert, 1893
Arachnoclerus Fairmaire, 1902
Aradamicula Sedlacek & Winkler, 1975
Araeodontia Barr, 1952-11
Arawakis Opitz, 2007
Archalius Fairmaire, 1903
Arenaria Opitz, 1997
Aroterus Schenkling, 1906
Artoadenus Opitz, 2015
Astigmus Kuwert, 1894
Aulicus Spinola, 1841
Avena Opitz, 2011
Axina Kirby, 1819
Balcus Sharp, 1877
Barriella Opitz, 2003
Barrotillus Rifkind, 1996
Basilewskyus Pic, 1950
Biflabellotillus Pic, 1949
Bilbotillus Kolibac, 1997
Blackburniella Chapin, 1924
Blaesiophthalmus Schenkling, 1903
Blaxima Gorham, 1882
Bogcia Barr, 1978-01
Boschella Barr, 1980
Bostrichoclerus Van Dyke, 1938
Bousquetoclerus Menier, 1997
Brachycallimerus Chapin, 1924
Brinckodes Winkler, 1960
Burgeoneus Pic, 1950
Caestron Dupont, 1844
Calendyma Lacordaire, 1857
Callimerus Gorham, 1876
Callotillus Wolcott, 1911
Canariclerus Winkler, 1982
Cardiostichus Quedenfeldt, 1885
Careydon
Caridopus Schenkling, 1908
Cephaloclerus Kuwert, 1893
Ceratocopus Hintz, 1902
Chariessa Perty, 1832
Chilioclerus Solervicens, 1976
Cladiscopallenis Pic, 1949
Cladiscus Chevrolat, 1843
Cladomorpha Pic, 1949
Cleromorpha Gorham, 1876
Cleropiestus Fairmaire, 1889
Clerus Geoffroy, 1762
Clytomadius Corporaal, 1949
Colobotis Opitz, 2009
Colyphus Spinola, 1841
Coptoclerus Chapin, 1924
Corinthiscus Fairmaire & Germain, 1861
Cormodes Pascoe, 1860
Corynetinus Reitter, 1894
Corynommadius Schenkling, 1899
Cregya LeConte, 1861
Crobenia Blackburn, 1891
Crusbatus Opitz, 2014
Ctenaxina Schenkling, 1906
Cteniopachys Fairmaire, 1889
Ctenoclerus Solervicens, 1997
Cucujocallimerus Pic, 1929
Cylidroctenus Kraatz, 1899
Cylidru
Cylidrus Latreille, 1829
Cymatodera Gray, 1832
Cymatoderella Barr, 1962
Cyrtinoclerus Chapin, 1924
Damiricus Fairmaire, 1903
Dasyceroclerus Kuwert, 1894
Dasyteneclines Pic, 1941
Decicornis Opitz, 2009
Decorosa Opitz, 2008
Dedana Fairmaire, 1888
Denops Fischer Von Waldheim, 1829
Dermestoides Houlbert & Bétis, 1905
Dermestoides J.C.Schaeffer, 1769
Diapromeces Opitz, 1997
Dieropsis Gahan, 1908
Diplocladus Fairmaire, 1885
Diplopherusa Heller, 1922
Diutius Opitz, 2014
Dolichopsis Gorham, 1878
Dologenitus Opitz, 2009
Dozocolletus Chevrolat, 1842
Eburiphora Spinola, 1841
Eburneocladiscus Pic, 1955
Eburneoclerus Pic, 1950
Egenocladiscus Corporaal & Van der Wiel, 1949
Ekisius Winkler, 1987
Elasmocylidrus Corporaal, 1940
Eleale Newman, 1841
Ellipotoma Spinola, 1844
Emmepus Motschoulsky, 1845
Enoclerus Gahan, 1910
Enoplioides Fairmaire, 1886
Enopliomorpha Pic, 1932
Enoplium Latreille, 1802
Enoploclerus Hintz, 1902
Epiclines Chevrolat, 1838
Epiphloeus Spinola, 1841
Erymanthus Spinola, 1841
Eucymatodera Schenkling, 1899
Eunatalis Schenkling, 1909
Eurymetomorphon Pic, 1950
Eurymetopum Blanchard, 1844
Evenoclerus Corporaal, 1950
Fallopylus Opitz, 2012
Falsocorynetes Pic, 1932
Falsoerymanthus Pic, 1941
Falsolissaulicus Pic, 1939
Falsomadius Gerstmeier, 2002
Falsoorthopleuroides Pic, 1939
Falsoorthrius Pic, 1940
Falsopallenis Pic, 1926
Falsopelonium Pic, 1950
Falsotillus Gerstmeier & Kuff, 1992
Falsotricladus Pic, 1939
Flabellotilloidea Gerstmeier & Kuff, 1992
Funicula Opitz, 2009
Gastrocentrum Gorham, 1876
Gerstmeierius Opitz, 2011
Globoclava Opitz, 2012
Gnidmus Bartlett, 2013
Goyasanum Pic, 1936
Gracilotillus Pic, 1933
Graptoclerus Gorham, 1901
Gyponyx Gorham, 1883
Hapsidopteris Opitz, 1997
Hautenerus Bartlett, 2013
Hemitrachys Gorham, 1876
Homalopilo Schenkling, 1915
Ichnea Laporte, 1836
Impressopallenis Pic, 1953
Inconnexus Opitz, 2017
Incorynetes Pic, 1941
Inhumeroclerus Pic, 1955
Iontoclerus Opitz, 1997
Isocymatodera Hintz, 1902
Isoforma Opitz, 2015
Isohydnocera E.A.Chapin, 1917
Isolemidia Gorham, 1877
Jenjouristia Fursov, 1936
Kanaliella Gerstmeier & Seitner, 2013
Katamyurus Opitz, 1997
Kataspinula Opitz, 2009
Kolibacius Opitz, 2011
Korynetes Herbst, 1792
Lacordarius Kuwert, 1893
Laiomorphus Pic, 1927
Languropilus Pic, 1940
Lasiocallimerus Corporaal, 1939
Lasiodera Gray, 1832
Lebasiella Spinola, 1844
Lecontella Wolcott & Chapin, 1918
Lemidia Spinola, 1841
Leptoclerus Kraatz, 1899
Lesneoclerus Corporaal, 1936
Liostylus Fairmaire, 1886
Lissaulicus Waterhouse, 1879
Loedelia Lucas, 1920
Lyctosoma Lewis, 1892
Macilentus Opitz, 2017
Macroliostylus Pic, 1939
Madoniella Pic, 1935
Magnotillus Pic, 1936
Megafodina Opitz, 2009
Megaphloeus Opitz, 2010
Megatrachys Opitz, 1997
Melanoclerus Chapin, 1919
Memorthrius Pic, 1940
Menieroclerus
Metademius Schenkling, 1899
Metaxina Broun, 1909
Microclerus Wollaston, 1867
Micropteroclerus Chapin, 1920
Microstigmatium Kraatz, 1899
Microtillus Pic, 1950
Mimenoplioides Pic, 1939
Mimolesterus Gerstmeier, 1991
Mimopallenis Pic, 1941
Miohydnocera Mawdsley, 1992
Mitrandiria Kolibac, 1997
Monophylla Spinola, 1841
Muisca Spinola, 1844
Musamuisca Zappi, 2021
Myrmecomaea Fairmaire, 1886
Natalis Laporte, 1836
Necrobia Olivier, 1800
Necrobinus Reitter, 1894
Nelsonoplium Barr, 2006
Nelsopelonium
Neocallotillus Burke, 2016
Neogyponyx Schenkling, 1906
Neohydnus Gorham, 1892
Neopylus Solervicens, 1989
Neorthopleura Barr, 1976
Neorthrius Gerstmeier & Eberle, 2011
Neoscrobiger Blackburn, 1900
Neotenerus Schenkling, 1906
Nodepus Gorham, 1892
Nolafigura Opitz, 2018
Nonalatus Gerstmeier, 2011
Notocymatodera Schenkling, 1907
Notostenus Dejean, 1833
Novemera Opitz, 2009
Odontophlogitus
Ohanlonella Rifkind, 2008
Olesterus Spinola, 1841
Omadius Laporte de Castelnau, 1836
Oncochelyna Opitz, 2017
Onerunka Kolibác, 2012
Onychotillus Chapin, 1945
Oodontophlogistus Elston, 1923
Operculiphorus Kuwert, 1894
Opetiopalpus Spinola, 1844
Opilo Latreille, 1802
Opitzia Nemesio, 2006
Opitzius Barr, 2006
Orthocladiscus Corporaal & van der Wiel, 1949
Orthopleuroides Kuwert, 1893
Orthrius Gorham, 1876
Oxystigmatium Kraatz, 1899
Pallenis Laporte de Castelnau, 1836
Pallenothriocera Pic, 1949
Paracladiscus Miyatake, 1965
Paradoxocerus Kraatz, 1899
Parapelonides Barr, 1980-01
Parapylus Blackburn, 1891
Paraspinoza Corporaal, 1942
Paratenerus Chapin, 1924
Paratillus Gorham, 1878
Parmius Sharp, 1877
Parvochaetus Opitz, 2006
Patuleius Fairmaire, 1903
Paupris Sharp, 1877
Pectobullus Opitz, 2011
Pelmatus Opitz, 2017
Pelonides Kuwert, 1894
Pelonium Spinola, 1844
Pennasolis Opits, 2008-21
Pericales Opitz, 2008
Perilypus Spinola, 1841
Philocalus Klug, 1842
Phlogistomorpha Hintz, 1908
Phlogistus Gorham, 1876
Phloiocopus Spinola, 1841
Phonius Chevrolat, 1843
Phyllobaenus Dejean, 1833
Phymatophaea Pascoe, 1876
Picoclerus Corporaal, 1936
Pieleus Pic, 1940
Pilosirus Opitz, 1997
Placocerus Klug, 1837
Placopterus Wolcott, 1910
Plastophagus Zhang, 1989
Plathanocera Schenkling, 1902
Platyclerus Spinola, 1841
Platynoptera Chevrolat, 1834
Platytenerus Miyatake, 1985
Plocamocera Spinola, 1844
Priocera Kirby, 1819
Priocleromorphus Pic, 1950
Prioclerus Hintz, 1902
Prionodera Wolcott, 1910
Propekorynetes Winkler, 1960
Prosymnus Laporte, 1836
Protoclerus Kolibáč & Huang, 2016
Pseudachlamys Duvivier, 1892
Pseudichnea Schenkling, 1901
Pseudoastigmus Eberle, 2011
Pseudogyponix Pic, 1939
Pseudolesterus Miyatake, 1968
Pseudomadius Chapin, 1924
Pseudopallenis Kuwert, 1893
Pseudopylus Opitz, 2012
Pseudoteloclerus Pic, 1932
Pteroferus Opitz, 2008
Pujoliclerus Pic, 1950
Pylus Newman, 1842
Pylusopsis Elston, 1929
Pyrrhoclerus Zhang, 1989
Pyticara Spinola, 1841
Pyticeroides Kuwert, 1894
Pyticeropsis Schenkling, 1916
Quasibrinckodes Winkler, 1960
Reptopetalis Opitz, 2011
Rhopaloclerus Fairmaire, 1886
Ridica
Rifkindius Opitz, 2009
Romanaeclerus Winkler, 1960
Salarium Fairmaire, 1903
Sallea Chevrolat, 1874
Scrobiger Spinola, 1841
Sedlacekius Winkler, 1972
Sedlacekvia
Shartegopsis Kirejtshuk, 2012
Sicamus
Sikorius Kuwert, 1893
Silvanoclerus Opitz, 2017
Silveirasia Nemesio, 2006
Silviella Solervicens, 1987
Sirpa
Smudlotillus Kolibac, 1997
Solervicensia Barr, 1979
Spinolarius Kuwert, 1893
Spinoza Lewis, 1892
Stenocallimerus Corporaal & Pic, 1940
Stenocylidrus Spinola, 1844
Stenoplium Fairmaire, 1903
Stigmatium Gray, 1832
Strotocera Schenkling, 1902
Sulcopatuleius Pic, 1941
Synellapotillus Pic, 1939
Synellapus Fairmaire, 1903
Syriopelta Winkler, 1984
Systenoderes Spinola, 1841
Tanocleria Hong, 2002
Tapainca
Tarsostenodes Blackburn, 1900
Tarsostenus Spinola, 1844
Teloclerus Schenkling, 1903
Tenerastes Lesne, 1932
Teneroides Gahan, 1910
Teneromimus Gahan, 1910
Teneropsis Chapin, 1924
Tenerus Laporte, 1836
Teutonia Opitz, 1997
Thalerocnemis Lohde, 1900
Thanasimodes Murray, 1867
Thanasimomorpha Blackburn, 1891
Thanasimus Latreille, 1806
Thaneroclerus Westwood, 1838
Theano Laporte, 1836
Theresamora Pic, 1950
Thriocera Gorham, 1878
Thriocerodes Wolcott & Dybas, 1947
Tillicera Spinola, 1841
Tilloclerus White, 1849
Tillodadiscus Pic, 1953
Tillodenops Hintz, 1905
Tilloidea Castelnau, 1832
Tilloidea Laporte, 1833
Tillus Olivier, 1790
Tilus
Titanoclerus Zhang, 1989
Trichodes Herbst, 1792
Trichodes Walckenaer, 1802
Tricladus Fairmaire, 1903
Trogodendron Spinola, 1841
Turbophloeus Opitz, 2008
Tylotosoma Hintz, 1902
Undecima Opitz, 2014
Viticlerus Miyatake, 1977
Wangweiella Kolibáč & Huang, 2016
Winklerius Menier, 1986
Wittmeridecus Winkler, 1981
Wolcottella Lucas, 1920
Wolcottia Chapin, 1917
Xenorthrius Gorham, 1892
Xylobius Guérin, 1841
Zahradnikius Winkler, 1992
Zebuclerus Opitz, 2011
Zenithicola Spinola, 1841

Name

Cleridae Latreille, 1802
References
Primary references

Latreille, P.A. 1802. Histoire naturelle, générale et particulière des crustacés et des insectes. Ouvrage faisant suite à l’histoire naturelle générale et particulière, composée par Leclerc de Buffon, et rédigée par C.S. Sonnini, membre de plusieurs sociétés savantes. Familles naturelles des genres. Tome troisième. F. Dufart, Paris, xii + pp. 13–467 + [1 (errata)]. BHL Reference page.

Additional references

Bartlett, J.S. 2009: The Cleridae of Lord Howe Island, with descriptions of two new species (Coleoptera: Cleroidea). Records of the Australian Museum, 61: 225–228. DOI: 10.3853/j.0067-1975.61.2009.1530
Bartlett, J.S. 2015. New synonymies for Australian Cleridae (Coleoptera). Zootaxa 4057(2): 281–287. DOI: 10.11646/zootaxa.4057.2.9. Preview (PDF) Reference page.
Corporaal, J.B. 1937: Check list of the Cleridae (Coleoptera) of Oceania. Occasional papers of the Bernice P. Bishop Museum, 13(3): 11–26. PDF
Faisal, M.; Singh, S.; Yousuf, M. 2014: Cleridae (Insecta: Coleoptera) type collection at National Forest Insect Collection (NFIC), Forest Research Institute, Dehradun (India). Zootaxa 3846(1): 105–118. DOI: 10.11646/zootaxa.3846.1.5 Reference page.
Gerstmeier, R. 2014: An overview of taxonomy and biology of the Cleridae (Coleoptera, Cleroidea, Cleridae). Giornale italiano di entomologia 13(59): 481–494. Abstract only seen Reference page.
Gunter, N.L. et al. 2013: A molecular phylogeny of the checkered beetles and a description of Epiclininae a new subfamily (Coleoptera: Cleroidea: Cleridae). Systematic entomology, DOI: 10.1111/syen.12019
Kolibáč, J. 1997(1996): Classification of the subfamilies of Cleridae (Coleoptera: Cleroidea). Acta Musei Moraviae, scientiae naturales (Brno), 81: 307–361.
Kolibáč, J. 2010: 9.6. Cleridae Latreille, 1802. Pp. 257-261 in: Leschen, R.A.B.; Beutel, R.G.; Lawrence, J.F. (volume eds.) Coleoptera, beetles. Volume 2: Morphology and systematics (Elateroidea, Bostrichiformia, Cucujiformia partim). In: Kristensen, N.P. & Beutel, R.G. (eds.) Handbook of zoology. A natural history of the phyla of the animal kingdom. Volume IV. Arthropoda: Insecta. Part 38. Berlin, New York: Walter de Gruyter. ISBN 3110190753 ISBN 9783110190755 DOI: 10.1515/9783110911213.257
Kolibáč, J. & Huang, D. 2016. The oldest known clerid fossils from the Middle Jurassic of China, with a review of Cleridae systematics (Coleoptera). Systematic Entomology 41(4): 808–823. DOI: 10.1111/syen.12192 Paywall. Reference page.
Leavengood, J.M., jr.; Garner, B.H. 2014: Nomenclatural notes on some checkered beetle (Coleoptera: Cleridae) types of the Natural History Museum, London (BMNH). Zootaxa 3760(3): 301–335. DOI: 10.11646/zootaxa.3760.3.1 Reference page.
I.Löbl & A.Smetana (eds). 2007 Catalogue of Palearctic Coleoptera. Vol. 4: Elateroidea, Derodontoidea, Bostrichoidea, Lymexyloidea, Cleroidea and Cucujoidea. Apollo Books, Stenstrup, Denmark ISBN 87-88757-67-6, p. 367
Opitz, W. 2009: Classification, natural history, and evolution of Neorthopleurinae subfam. nov. (Coleoptera, Cleridae). Part I. Generic composition of the subfamily and key to genera. Entomologica Basiliensia et Collectionis Frey, 31: 135–207.
Opitz, W. 2010: Classification, natural history, phylogeny, and subfamily composition of the Cleridae and generic content of the subfamilies (Coleoptera: Cleroidea). Entomologica Basiliensia et Collectionis Frey, 32: 31–128. [not seen]
Zappi, I. & Ghahari, H. 2016. A checklist of the Cleridae of Iran with new data (Coleoptera: Cleroidea). Zootaxa 4147(4): 403–420. DOI: 10.11646/zootaxa.4147.4.3. Reference page.

Links

Australian Faunal Directory
Cleridae in SYNOPSIS OF THE DESCRIBED COLEOPTERA OF THE WORLD
Atlas of checkered beetles (Cleridae) of Russia

Vernacular names
العربية: بوقيٌات ضاريٌة
беларуская: Стракачы
čeština: Pestrokrovečníkovití
Deutsch: Buntkäfer
English: checkered beetles
eesti: Sipelgmardiklased
suomi: Kirjokuoriaiset
עברית: קלריתיים
magyar: Szúfarkasfélék
日本語: カッコウムシ科
lietuvių: Keršvabaliai
Nederlands: Bonte kevers
norsk: Maurbiller
polski: Przekraskowate
русский: Пестряки
svenska: Brokbaggar
中文: 郭公蟲科

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 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.[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 the 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. The 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 the Phyllobaenus spp. are wood borers, immature weevils, and hymenoptera larvae.[9] One of the more diverse genera is Trichodes, the 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 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 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 lifespaned 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 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.[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. 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.[14] N. rufipes has been recorded to have fed 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 beetles 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]

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 Olivier, 1795
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

Johnson, Norman F.; Triplehorn, Charles A. (2004). Borror and DeLong's Introduction to the Study of Insects (7th ed.). Belmont: Brooks/Cole. pp. 365–400, 428–429. ISBN 0-03-096835-6.
White, Richard E. (1998). A Field Guide to the Beetles of North America (illustrated and revised ed.). New York: Houghton Mifflin Harcourt. pp. 208–209. ISBN 978-0-395-91089-4. Retrieved 22 March 2009.
Byrd, Jason H. (2001). James L., Castner (ed.). Forensic Entomology: The Utility of Arthropods in Legal Investigations (2nd ed.). Boca Raton: CRC Press. ISBN 0-8493-8120-7.
Gerstmeier, Ronald (1998). Checkered Beetles: Illustrated Key to the Cleridae and Thanerocleridae of the Western Palaearctic. Weikersheim, Germany: Margraf Verlag. pp. 12–15.
Bellows, Thomas S.; Fisher, T. W.; Caltagirone, L. E. (1999). Handbook of biological control. San Diego: Academic Press. pp. 429–430. ISBN 0-12-257305-6. Retrieved 19 March 2009.
Insects of eastern forests. Misc. Publ. 1426. U.S. Forestry Service. Washington D.C: U.S. Department of Agriculture, Forest Service. 1985. pp. 274–275. Retrieved 22 March 2009.
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.
Costello, Sheryl (2003). "Clerid Beetles- Voracious Predators" (PDF). Colorado State University Department of Entomology. pp. 1–15. Archived from the original (PDF) on 24 July 2011. Retrieved 19 March 2009.
Majka, Christopher (20 June 2006). "A guide to the Cleridae of Atlantic Canada". Chebucto Community Net. Retrieved 19 March 2009.
Gredilha, R.; Lima, A. F. (February 2007). "First record of Necrobia rufipes (De Geer, 1775) (Coleoptera; Cleridae) associated with pet food in Brazil" (PDF). Brazilian Journal of Biology. 1. 67 (1): 187. doi:10.1590/S1519-69842007000100026. PMID 17505768.
Mahr, Susan (May 2000). "Know Your Friends Checkered Beetles". Biological Control News. University of Wisconsin-Madison. 7 (1). Retrieved 20 March 2009.
Linsley, Earle G. (1936). "Studies in the genus Aulicus Spinola (Coleoptera-Cleridae)". University of California Publications in Entomology. Berkeley, California: University of California Press. 6 (9): 249–262.
Hue, Yi (28 June 2008). "Life Cycle of Thanasimus formicarius (Coleoptera: Cleridae) in Southern Norway". Insect Science. Institute of Zoology, Chinese Academy of Sciences. 5 (1): 55–62. doi:10.1111/j.1744-7917.1998.tb00298.x. S2CID 85273860. Archived from the original on 5 January 2013.
Ebeling, Walter (23 August 2002). "Chapter 7 Pests of Stored Food Products". Urban Entomology. University of California Riverside. Archived from the original on 8 November 2015. Retrieved 22 March 2009.
Souza, Adriana; Linhares, Aricio (28 June 2008). "Diptera and Coleoptera of potential forensic importance in southeastern Brazil: relative abundance and seasonality". Medical and Veterinary Entomology. Royal Entomological Society. 11 (1): 8–12. doi:10.1111/j.1365-2915.1997.tb00284.x. PMID 9061672. S2CID 30551315. Archived from the original on 5 January 2013.
Poland, Therese M.; Borden, John H. (December 1997). "Attraction of bark beetle predator, Thanasimus undatulus (Coleoptera: Cleridae), to pheromones of the spruce beetle and two secondary bark beetles (Coleoptera: Scolytidae)" (PDF). Journal of the Entomological Society of British Columbia. 94: 35–41. Retrieved 20 March 2009.
Mawdsley, Jonathan R. (2002). "Ecological notes on species of Cleridae (Insecta: Coleoptera) associated with the prairie flora of central North America". The Great Lakes Entomologist. 35 (1): 15–22. Retrieved 20 March 2009.
Kolibáč, Jiří; Huang, Diying (October 2016). "The oldest known clerid fossils from the Middle Jurassic of China, with a review of Cleridae systematics (Coleoptera)". Systematic Entomology. 41 (4): 808–823. doi:10.1111/syen.12192.