Coleoptera

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
Ordo: Coleoptera
Subordines (4 + 1†): Adephaga – Archostemata – Myxophaga – Polyphaga – †Protocoleoptera
[source: Bouchard et al., 2011]

Genera incertae sedis: †Adiphlebia – †Coleopteron – †Grahamelytron
Overview of familiae (170)

Acanthocnemidae – Aderidae – Agapythidae – Agyrtidae – Akalyptoischiidae – Alexiidae – Amphizoidae – Anthicidae – Anthribidae – Archeocrypticidae – Artematopodidae – Aspidytidae – Attelabidae – Belidae – Biphyllidae – Boganiidae – Boridae – Bostrichidae – Bothrideridae – Brachyceridae – Brachypsectridae – Brentidae – Buprestidae – Byrrhidae – Byturidae – Callirhipidae – Cantharidae – Carabidae – Caridae – Cavognathidae – Cerambycidae – Cerophytidae – Cerylonidae – Chaetosomatidae – Chalcodryidae – Chelonariidae – Chrysomelidae – Ciidae – Clambidae – Cleridae – Cneoglossidae – Coccinellidae – Corylophidae – Crowsoniellidae – Cryptophagidae – Cucujidae – Cupedidae – Curculionidae – Cybocephalidae – Cyclaxyridae – Dascillidae – Decliniidae – Dermestidae – Derodontidae – Discolomatidae – Disteniidae – Dryophthoridae – Dryopidae – Dytiscidae – Elateridae – Elmidae – Endecatomidae – Endomychidae – Epimetopidae – Erotylidae – Eucinetidae – Eucnemidae – Eulichadidae – Georissidae – Gyrinidae – Haliplidae – Helophoridae – Helotidae – Heteroceridae – Histeridae – Hobartiidae – Hydraenidae – Hydrochidae – Hydrophilidae – Hydroscaphidae – Hygrobiidae – Ischaliidae – Jacobsoniidae – Jurodidae – Kateretidae – Laemophloeidae – Lamingtoniidae – Lampyridae – Latridiidae – Leiodidae – Lepiceridae – Lophocateridae – Limnichidae – Lutrochidae – Lycidae – Lymexylidae – Mauroniscidae – Megalopodidae – Melandryidae – Meloidae – Melyridae – Meruidae – Micromalthidae – Monotomidae – Mordellidae – Mycetophagidae – Mycteridae – Myraboliidae – Nemonychidae – Nitidulidae – Nosodendridae – Noteridae – Oedemeridae – Omalisidae – Omethidae – Ommatidae – Orsodacnidae – Oxypeltidae – Passandridae – Peltidae – Phalacridae – Phengodidae – Phloeostichidae – Phloiophilidae – Phycosecidae – Plastoceridae – Priasilphidae – Prionoceridae – Promecheilidae – Propalticidae – Prostomidae – Protocucujidae – Protopeltidae – Psephenidae – Pterogeniidae – Ptiliidae – Ptilodactylidae – Ptinidae – Pyrochroidae – Pythidae – Rhadalidae – Rentoniidae – Rhagophthalmidae – Rhinorhipidae – Rhipiceridae – Rhysodidae – Ripiphoridae – Salpingidae – Schizopodidae – Scirtidae – Scraptiidae – Silphidae – Silvanidae – Smicripidae – Spercheidae – Sphaeritidae – Sphaeriusidae – Sphindidae – Staphylinidae – Stenotrachelidae – Synchroidae – Synteliidae – Tasmosalpingidae – Telegeusidae – Tenebrionidae – Tetratomidae – Thanerocleridae – Throscidae – Thymalidae – Torridincolidae – Trachelostenidae – Trachypachidae – Trictenotomidae – Trogossitidae – Ulodidae – Vesperidae – Zopheridae

Fossil: †Ademosynidae – †Asiocoleidae – †Berendtimiridae – †Colymbotethidae – †Coprinisphaeridae – †Coptoclavidae – †Elodophthalmidae – †Labradorocoleidae – †Liadytidae – †Magnocoleidae – †Mesocinetida – †Oborocoleidae – †Obrieniidae – †Pallichnidae – †Parahygrobiidae – †Parandrexidae – †Permocupedidae – †Permosynidae – †Phoroschizidae – †Praelateriidae – †Rhombocoleidae – †Schizocoleidae – †Sinisilvanidae – †Taldycupedidae – †Tshekardocoleidae – †Triadocupedinae – †Triaplidae – †Tritarsidae – †Ulyanidae
Name

Coleoptera Linnaeus, 1758
References
Primary references

Linnaeus, C. 1758. Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio Decima, Reformata. Tomus I. Holmiæ (Stockholm): impensis direct. Laurentii Salvii. 824 pp. DOI: 10.5962/bhl.title.542 BHL Reference page.

Additional references

Abdel-Dayem, M.S., Abu El-Ghiet, U.M., El-Sheikh, T.M., Elgharbawy, A.A., al-Fifi, Z.I.A. & Al Dhafer, H.M. 2020. The first survey of the beetles (Coleoptera) of the Farasan Archipelago of the southern Red Sea, Kingdom of Saudi Arabia. ZooKeys, 959: 17–86. DOI: 10.3897/zookeys.959.51224 Open access Reference page.
Alekseev, V.I. 2013. The beetles (Insecta: Coleoptera) of Baltic amber: the checklist of described species and preliminary analysis of biodiversity. Zoology and ecology 23: 5–12. DOI: 10.1080/21658005.2013.769717 Reference page.
de Araujo, B.C., Schmidt, S., Schmidt, O., von Rintelen, T., von Rintelen, K., Floren, A., Ubaidillah, R., Peggie, P. & Balke, M. 2019. DNA barcoding data release for Coleoptera from the Gunung Halimun canopy fogging workpackage of the Indonesian Biodiversity Information System (IndoBioSys) project. Biodiversity Data Journal, 7: e31432. DOI: 10.3897/BDJ.7.e31432 Reference page.
Arnett, R.H., jr.; Thomas, M.C.; Skelley, P.E.; Frank, J.H. (eds.) 2002: American beetles. Volume 2. Polyphaga: Scarabaeoidea through Curculionoidea. CRC Press LLC, Boca Raton, Florida. limited preview
Béthoux, O. 2009: The earliest beetle identified. Journal of paleontology, 83: 931–937. DOI: 10.1666/08-158.1
Beutel, R.G.; Ge, S.-Q.; Hörnschemeyer, T. 2008: On the head morphology of Tetraphalerus, the phylogeny of Archostemata and the basal branching events in Coleoptera. Cladistics, 24: 270–298. DOI: 10.1111/j.1096-0031.2007.00186.x
Beutel, R.G.; Haas, F. 2000: Phylogenetic relationships of the suborders of Coleoptera (Insecta). Cladistics, 16: 103–141.
Beutel, R.G.; Haas, F. 2003: Basal splitting events in Coleoptera. Pp. 160-161 in: Klass, K.D. (ed.) Proceedings of the 1st Dresden meeting on insect phylogeny: “Phylogenetic relationships within the insect orders” (Dresden, September 19–21, 2003). Entomologische Abhandlungen, 61: 119–172.[1]
Beutel, R.G.; Leschen, R.A.B. (volume eds.) 2005: Coleoptera, beetles. Volume 1: Morphology and systematics (Archostemata, Adephaga, Myxophaga, Polyphaga 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 3110171309 DOI: 10.1515/9783110904550
Bishop, D.J.; Majka, C.G.; Bondrup-Nielsen, S.; Peck, S.B. 2009: Deadwood and saproxylic beetle diversity in naturally disturbed and managed spruce forests in Nova Scotia. In: Majka, C.G.; Klimaszewski, J. (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera II. ZooKeys, 22: 309–340. DOI: 10.3897/zookeys.22.144
Blackmon, H. & Demuth, J.P. 2015. Coleoptera karyotype database. The Coleopterists Bulletin 69(1): 174–175. DOI: 10.1649/0010-065X-69.1.174 Paywall. Reference page.
Bocák, L., Barton, C., Crampton-Blatt, A., Chesters, D., Ahrens, D. & Vogler, A.P. 2014. Building the Coleoptera tree-of-life for >8000 species: composition of public DNA data and fit with Linnaean classification. Systematic Entomology 39(1): 97–110. DOI: 10.1111/syen.12037 Paywall. Reference page.
Bouchard, P. et al. 2009: Biodiversity of Coleoptera. Pp. 265-301 in: Foottit, R.G.; Adler, P.H. (eds.) 2009: Insect biodiversity: science and society. Blackwell Publishing Ltd. ISBN 9781405151429
Bouchard, P., Bousquet, Y., Davies, A.E., Alonso-Zarazaga, M.A., Lawrence, J.F., Lyal, C.H.C., Newton, A.F., Reid, C.A.M., Schmitt, M., Ślipiński, S.A. & Smith, A.B.T. 2011. Family-group names in Coleoptera (Insecta). ZooKeys 88: 1–972. DOI: 10.3897/zookeys.88.807 Open access. PMC: 3088472 Open access. Reference page.
Bousquet, Y., Bouchard, P., Davies, A.E. & Sikes, D.S. 2013. Checklist of beetles (Coleoptera) of Canada and Alaska. Second edition. ZooKeys 360: 1–44. DOI: 10.3897/zookeys.360.4742 Open access. Reference page.
Brunke, A.J., Bouchard, P., Douglas, H.B. & Pentinsaari, M. 2019. Coleoptera of Canada. Pp 361–376 In Langor, D.W. & Sheffield, C.S. (eds.). The Biota of Canada – A Biodiversity Assessment. Part 1: The Terrestrial Arthropods. ZooKeys 819: 520 pp. Reference page. DOI: 10.3897/zookeys.819.24724 Open access. Reference page.
Caterino, M.S.; Shull, V.L.; Hammond, P.M.; Vogler, A.P. 2002: Basal relationships of Coleoptera inferred from 18S rDNA sequences. Zoologica scripta, 31: 41–49.
Crotch, G.R. 1871: List of all the Coleoptera described A.D. 1758-1821, referred to their modern genera. Cambridge. BUGZ
Cuccodoro, G.; Leschen, R.A.B. (eds.) 2003: Systematics of Coleoptera: papers celebrating the retirement of Ivan Löbl. Memoirs on entomology international, 17. contents
Egorov, L.V., Ruchin, A.B., Semenov, V.B., Semionenkov, O.I. & Semishin, G.B. 2020. Checklist of the Coleoptera of Mordovia State Nature Reserve, Russia. ZooKeys, 962: 13–122. DOI: 10.3897/zookeys.962.54477 Open access Reference page.
Fikáček, M.; Skuhrovec, J.; Šípek, P. (eds.) 2010: Abstracts of the Immature Beetles Meeting 2009, October 1–2, Prague, Czech Republic. Acta Entomologica Musei Nationalis Pragae, 50: 323–342. ISSN: 0374-1036 PDF
Friedrich, F.; Beutel, R.G. 2006: The pterothoracic skeletomuscular system of Scirtoidea (Coleoptera: Polyphaga) and its implications for the high-level phylogeny of beetles. Journal of zoological systematics & evolutionary research, 44: 290–315.
Friedrich, F.; Farrell, B.D.; Beutel, R.G. 2008: The thoracic morphology of Archostemata and the relationships of the extant suborders of Coleoptera (Hexapoda). Cladistics, 24: 1–37. DOI: 10.1111/j.1096-0031.2008.00233.x
Gerlach, J. (ed.) 2009. The Coleoptera of the Seychelles Islands. Series Faunistica 88. Sofia-Moscow: Pensoft Publishers. 266 pp. Reference page.
Grebennikov, V.V.; Leschen, R.A.B. 2010: External exoskeletal cavities in beetles (Insecta: Coleoptera) and their possible mycangial functions. Entomological science, 13: 81–98. DOI: 10.1111/j.1479-8298.2009.00351.x PDF
Hangay, G.; Zborowski, P. 2010: A guide to the beetles of Australia. CSIRO Publishing: Collingwood VIC, Australia. ISBN 9780643094871 ISBN 0643094873
Hunt, T.; Bergsten, J.; Levkanicova, Z.; Papadopoulou, A.; St. John, O.; Wild, R.; Hammond, P.M.; Ahrens, D.; Balke, M.; Caterino, M.S.; Gomez-Zurita, J.; Ribera, I.; Barraclough, T.G.; Bocakova, M.; Bocak, L.; Vogler, A.P. 2007: A comprehensive phylogeny of beetles reveals the evolutionary origins of a superradiation. Science, 318: 1913–1916. DOI: 10.1126/science.1146954
Jäch, M.A.; Balke, M. (eds.) 2010: Water beetles of New Caledonia (part 1). Monographs on Coleoptera, 3. ISSN: 1027-8869 [not seen]
Jäch, M.A.; Balke, M. 2010: Introduction. Pp. 1-30 in: Jäch, M.A. & Balke, M. (eds.): Water beetles of New Caledonia (part 1). Monographs on Coleoptera, 3. ISSN: 1027-8869 PDF(in .zip file)
Kirby, W.F. 1881. Coleoptera. The Zoological record 17: 11–99. BHL. Reference page.
Kirejtshuk, A.G. & Prokin, A.A. 2018. The Position of the Palaeozoic Genus: Tunguskagyrus Yan, Beutel et Lawrence in the Family Triaplidae sensu n. (Coleoptera, Archostemata: Schizophoroidea). Entomological Review 98: 872–882. DOI: 10.1134/S0013873818070084 Paywall. Reference page.
Lamelas-López, L., Raposeiro, P.M., Borges, P.A.V. & Florencio, M. 2017. Annotated checklist of aquatic beetles (Coleoptera) and true bugs (Heteroptera) in the Azores Islands: new records and corrections of colonization status. Zootaxa 4353(1): 117–132. DOI: 10.11646/zootaxa.4353.1.7. Reference page.
Lawrence, J.F. 1982: Coleoptera. Pp. 482-553 in Parker, S.P. (ed.) Synopsis and classification of living organisms, 2. McGraw-Hill, USA.
Lawrence, J.F. (coordinator) 1991: Order Coleoptera, in: Stehr, F.W. (ed.) Immature insects, 2. Kendall/Hunt Publishing Company, Iowa, USA.
Lawrence, J.F. 1999: The Australian Ommatidae (Coleoptera: Archostemata): new species, larva and discussion of relationships. Invertebrate taxonomy, 13: 369–390. DOI: 10.1071/IT99008
Lawrence, J.F.; Beutel, R.G.; Leschen, R.A.B.; Ślipiński, A. 2010: 1. Changes in classification and list of families and subfamilies. Pp. 1-7 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
Lawrence, J.F.; Beutel, R.G.; Leschen, R.A.B.; Ślipiński, A. 2010: 2. Glossary of morphological terms. Pp. 9-20 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
Lawrence, J.F. et al. 2011: Phylogeny of the Coleoptera based on morphological characters of adults and larvae. Annales zoologici, 61(1): 1–217. abstract only seen
Lawrence, J.F.; Newton, A.F., jr. 1982: Evolution and classification of beetles. Annual review of ecology and systematics, 13: 261–290. [2]
Lawrence, J.F. & Newton, A.F. 1995. Families and subfamilies of Coleoptera (with selected genera, notes, references and data on family-group names). Pp. 779–1006. In: Pakaluk, J. & Ślipiński, S.A. (eds.) Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson. Volume 2. Museum i Instytut Zoologii PAN, Warszawa. x + 1092 pp. in 2 vols. ISBN 83-85192-34-4. ResearchGate Open access green. Reference page.
Lawrence, J.F.; Ślipiński, S.A.; Pakaluk, J. 1995: From Latreille to Crowson: a history of the higher-level classification of beetles. Pp. 87-154 in Pakaluk, J.; Ślipiński, S.A. (eds.) Biology, phylogeny, and classification of Coleoptera: papers celebrating the 80th birthday of Roy A. Crowson. Museum i Instytut Zoologii PAN, Warszawa.
Leschen, R.A.B. & Beutel, R.G. (volume eds.) 2014. Arthropoda: Insecta: Coleoptera Volume 3: Morphology and Systematics (Phytophaga). 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 978-3-11-027446-2 DOI: 10.1515/9783110274462 Reference page.
Leschen, R.A.B.; Beutel, R.G.; Lawrence, J.F. (volume eds.) 2010: 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
Löbl, I. & Leschen, R.A.B. 2013 (Online) 2014 (Print). Misinterpreting global species numbers: examples from Coleoptera. Systematic Entomology 39(1): 2–6. DOI: 10.1111/syen.12042 Reference page.
Maddison, P. (compiler) 2010: Order Coleoptera. Pp. 409-436, in: Checklist of New Zealand Hexapoda. Pp. 396-467 in:
Macfarlane, R.P., Maddison, P.A., Andrew, I.G., Berry, J.A., Johns, P.M., Hoare, R.J.B., Larivière, M.-C., Greenslade, P., Henderson, R.C., Smithers, C.N., Palma, R.L., Ward, J.B., Pilgrim, R.L.C., Towns, D.R., McLellan, I., Teulon, D.A.J., Hitchings, T.R., Eastop, V.F., Martin, N.A., Fletcher, M.J., Stufkens, M.A.W., Dale, P.J., Burckhardt, D., Buckley, T.R. & Trewick, S.A. 2010. Phylum Arthropoda subphylum Hexapoda: Protura, springtails, Diplura, and insects. Pp. 233–467 in: Gordon, D.P. (ed.) 2010. New Zealand inventory of biodiversity. Volume 2. Kingdom Animalia. Chaetognatha, Ecdysozoa, ichnofossils. Canterbury University Press, Christchurch, New Zealand. Reference page. PDF. Reference page.
Majka, C.G.; Sikes, D.S. 2009: Thomas L. Casey and Rhode Island's precinctive beetles: taxonomic lessons and the utility of distributional checklists. In: Majka, C.G.; Klimaszewski, J. (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera II. ZooKeys, 22: 267–283. DOI: 10.3897/zookeys.22.93
McKenna, D.D.; Farrell, B.D. 2009: Beetles (Coleoptera). Pp. 278-289 in Hedges, S.B. & Kumar, S. (eds.) The Timetree of Life. Oxford University Press. PDF
Pentinsaari, M., Anderson, R., Borowiec, L., Bouchard, P., Brunke, A., Douglas, H., Smith, A. & Hebert, P. 2019. DNA barcodes reveal 63 overlooked species of Canadian beetles (Insecta, Coleoptera). ZooKeys 894: 53–150. DOI: 10.3897/zookeys.894.37862 Open access Reference page.
Ponomarenko, A.G. 1995: The geological history of beetles. Pp. 155-171 in Pakaluk, J.; Ślipiński, S.A. (eds.) Biology, phylogeny, and classification of Coleoptera: papers celebrating the 80th birthday of Roy A. Crowson. Museum i Instytut Zoologii PAN, Warszawa.
Ponomarenko, A.G. 2002: 2.2.1.3.2. Superorder Scarabaeidea Laicharting, 1781. Order Coleoptera Linné, 1758. The beetles. Pp. 164-176 in Rasnitsyn, A.P.; Quicke D.L.J. (eds.) History of insects. Dordrecht: Kluwer. ISBN 140200026X [3]
Ponomarenko, A.G. 2003: Ecological evolution of beetles (Insecta: Coleoptera). Acta zoologica cracoviensia, 46(suppl.-Fossil Insects): 319–328. [4]
Ponomarenko, A.G. 2011. New beetles (Insecta, Coleoptera) from Vyazniki Locality, Terminal Permian of European Russia. Paleontologicheskii Zhurnal 2011(4): 55–63. [in Russian, English translation in Paleontological Journal 45(4): 414–422. (2011) DOI: 10.1134/S0031030111040095 Paywall.]
Sánchez-Fernández, D., Millán, A., Abellán, P., Picazo, F., Carbonell, J.A. & Ribera, I. 2015. Atlas of Iberian water beetles (ESACIB database). Zookeys 520: 147-154. DOI: 10.3897/zookeys.520.6048 Preview (PDF) Reference page.
Seago, A.E.; Brady, P.; Vigneron, J.-P.; Schultz, T.D. 2009: Gold bugs and beyond: a review of iridescence and structural colour mechanisms in beetles (Coleoptera). Journal of the Royal Society Interface, 6, S165–S184. DOI: 10.1098/rsif.2008.0354.focus
Slipinski, S.A., Leschen, R.A.B. & Lawrence, J.F. 2011. Order Coleoptera Linnaeus, 1758. Pp 203–208 In
Zhang, Z.-Q. (ed.) 2011. Animal biodiversity: an outline of higher-level classification and survey of taxonomic richness. Zootaxa 3148: 1–237. Open access. Reference page . (PDF) Reference page.
Tamutis, V.; Tamutė, B.; Ferenca, R. 2011: A catalogue of Lithuanian beetles (Insecta, Coleoptera). ZooKeys, 121: 1–494. DOI: 10.3897/zookeys.121.732
Vogler, A.P.; Caterino, M.S. 2003: The basal relationships of Coleoptera based on 18S rRNA sequences. Pp. 161-162 in: Klass, K.D. (ed.) Proceedings of the 1st Dresden meeting on insect phylogeny: “Phylogenetic relationships within the insect orders” (Dresden, September 19–21, 2003). Entomologische Abhandlungen, 61: 119–172.[5]
Watt, J.C. 1975: Notes on priority of family-group names in Coleoptera. Coleopterists bulletin, 29: 31–34.

Links

Assembling the Beetle Tree of Life (BToL)
www.coleoptera.org
Coleoptera - Australian Faunal Directory
Catalogue of Palaearctic Coleoptera. Edited by I. Löbl & A. Smetana
SYNOPSIS OF THE DESCRIBED COLEOPTERA OF THE WORLD
Coleoptera incertae sedis in SYNOPSIS OF THE DESCRIBED COLEOPTERA OF THE WORLD
Maddison, David R. 2000. Coleoptera. Beetles. Version 11 September 2000 (under construction) in The Tree of Life Web Project
Beetles and coleopterologists Russian site with English version, with information about biology, systematics and paleontology of beetles
Coleoptera - Coleoptera-atlas (Beautiful Pictures of coleoptera - picture Gallery (over 4800 records)

Vernacular names
Alemannisch: Chäfer
беларуская: Жукі
български: Твърдокрили
বাংলা: গুবরে পোকা
čeština: Brouci
dansk: Biller
Deutsch: Käfer
Ελληνικά: Κολεόπτερα
English: Beetles
Esperanto: Koleopteroj
español: Escarabajo
eesti: Mardikalised
suomi: Kovakuoriaiset
Nordfriisk: Kraaben
français: Coléoptères
עברית: חיפושיות
हिन्दी: कंचुकपक्ष
magyar: Bogarak
íslenska: Bjöllur
italiano: Coleotteri
日本語: コウチュウ目 (鞘翅目)
한국어: 딱정벌레목 (초시목)
Līvõ kēļ: Būmbald
lietuvių: Vabalai
latviešu: Vaboles
Malagasy: Borera
македонски: Тврдокрилци
Nederlands: Kevers
norsk: Biller
polski: Chrząszcze
português: Coleópteros
русский: Жесткокрылые
davvisámegiella: Goppát
srpskohrvatski / српскохрватски: Tvrdokrilci
slovenčina: Chrobáky
slovenščina: Hrošči
српски / srpski: Тврдокрилци
svenska: Skalbaggar
தமிழ்: வண்டு (vandu)
ไทย: แมลงปีกแข็ง
Türkçe: Kın kanatlılar
українська: Твердокрилі
Tiếng Việt: Bọ cánh cứng
中文: 鞘翅目

Subordines: Adephaga - Archostemata - Myxophaga - Polyphagastem group

Coleoptera (pronounced /koʊliːˈɒptərə/), species commonly called Beetles, is an order of insects; from Greek κολεός, koleos, "sheath"; and πτερόν, pteron, "wing", thus "sheathed wing"), which contains more species than any other order in the animal kingdom, constituting almost 25% of all known life-forms.[1] About 40% of all described insect species are beetles (about 400,000 species[2]), and new species are discovered frequently. Some estimates put the total number of species, described and undescribed, at as high as 100 million, but 1 million is a more likely figure.[3] The largest taxonomic family, the Curculionidae (the weevils or snout beetles), also belongs to this order .

The diversity of beetles is very wide-ranging. Being found in almost all habitats, but are not known to occur in the sea or in the polar regions. They interact with their ecosystems in several ways. They often feed on plants and fungi, break down animal and plant debris, and eat other invertebrates. Some species are prey of various animals including birds and mammals. Certain species are agricultural pests, such as the Colorado potato beetle Leptinotarsa decemlineata, the boll weevil Anthonomus grandis, the red flour beetle Tribolium castaneum, and the mungbean or cowpea beetle Callosobruchus maculatus, while other species of beetles are important controls of agricultural pests. For example, beetles in the family Coccinellidae ("ladybirds" or "ladybugs") consume aphids, scale insects, thrips, and other plant-sucking insects that damage crops.

Species in the order Coleoptera are generally characterized by a particularly hard exoskeleton and hard forewings (elytra). This elytra separates it from most other insect species, except for a few Hemiptera species. The beetle's exoskeleton is made up of numerous plates called sclerites, separated by thin sutures. This design creates the armored defenses of the beetle while maintaining flexibility. The general anatomy of a beetle is quite uniform, although specific organs and appendages may vary greatly in appearance and function between the many families in the order. Like all insects, beetles' bodies are divided into three sections: the head, the thorax, and the abdomen. The internal morphology is like other insects, however some factors are unique. Such as species of diving beetles, who use air bubbles in order to dive under the water, which could last a respectable period of time due to passive diffusion allowing oxygen to transfer from the water into the bubble.

Beetles are endopterygotes, which means that they undergo complete metamorphosis, a biological process by which an animal physically develops after birth or hatching, undergoing a series of conspicuous and relatively abrupt change in the its body structure. Coleopteran species have an extremely intricate behavior when mating, using such methods as pheromones for communication to locate potential mates. Males may fight for females using very elongated mandibles, causing a strong divergence between males and females in sexual dimorphism.

Etymology

Coleoptera comes from the Greek koleopteros, literally "sheath-wing," from "koleos" meaning "sheath," and pteron, meaning "wing. The name "Coleoptera" was given by Aristotle for the elytra, a hardened shield-like forewing. According to common vernacular, species of Coleoptera may go by an array of names, including fireflies, june bug, ladybugs, weevils, and the obvious beetles.[1] The word beetle comes from the Old English word bitela, literally meaning small biter, deriving from the word bitel, which means biting.[4]

Distribution and diversity

With one of the largest order of insects, with 350,000 - 400,000, including 24,000 in North America, and 20,000 in Australia, species described in four suborders (Adephaga, Archostemata, Myxophaga, and Polyphaga), which makes up about 40% of all insect species described. Even though at the family level of classification is a bit unstable, there are about 500 recognized families and subfamilies.[1][5] Species of coleoptera are found in an array of natural habitats, including nearly all of them. Including water and marine habitats, and every habitat with vegetative foliage, from trees and there bark to flower, leaves and underground near roots. even being found inside the plants in galls in ever plant tissue including dead or decaying ones.[6]

External Morphology
The morphology of a beetle, with a fiddler beetle as an example species.

Beetles are generally characterized by a particularly hard exoskeleton and hard forewings (elytra). The beetle's exoskeleton is made up of numerous plates called sclerites, separated by thin sutures. This design creates the armored defenses of the beetle while maintaining flexibility. The general anatomy of a beetle is quite uniform, although specific organs and appendages may vary greatly in appearance and function between the many families in the order. Like all insects, beetles' bodies are divided into three sections: the head, the thorax, and the abdomen.[1]
Scarabaeus viettei (syn. Madateuchus viettei, Scarabaeidae) showing a "shovel head" adaptation.
Head of Cephalota circumdata, showing the compound eyes and mouthparts

Head

The head, having a projecting forward mouthparts or sometimes downturned, is usually heavily sclerotized and varying in size.[5] The eyes are compound and may display remarkable adaptability, as in the case of whirligig beetles (family Gyrinidae), in which the eyes are split to allow a view both above and below the waterline. Other species also have divided eyes – some longhorn beetles (family Cerambycidae) and weevils – while many beetles have eyes that are notched to some degree. A few beetle genera also possess ocelli, which are small, simple eyes usually situated farther back on the head (on the vertexes).

Beetles' antennae are primarily organs of smell, but may also be used to feel out a beetle's environment physically. They may also be used in some families during mating, or among a few beetles for defence. Antennae vary greatly in form within the Coleoptera, but are often similar within any given family. In some cases, males and females of the same species will have different antennal forms. Antennae may be clavate (flabellate and lamellate are sub-forms of clavate, or clubbed antennae), filiform, geniculate, moniliform, pectinate, or serrate.

Beetles have mouthparts similar to those of grasshoppers. Of these parts, the most commonly known are probably the mandibles, which appear as large pincers on the front of some beetles. The mandibles are a pair of hard, often tooth-like structures that move horizontally to grasp, crush, or cut food or enemies (see defence, below). Two pairs of finger-like appendages are found around the mouth in most beetles, serving to move food into the mouth. These are the maxillary and labial palpi. In many species the mandibles are sexually dismorphic, with the males having theirs enlarged, with some being enormosly enlarged compared to the females of the same species.[5]

Thorax

The thorax is segmented into the two discernible parts, the pro- and pterathorax. The pterathorax is the fused meso- and metathorax, which are commonly separate in other insect species, although being flexibly articulate from the prothorax. When viewed from below, the thorax is that part from which all three pairs of legs and both pairs of wings arise. The abdomen is everything posterior to the thorax.[1] When viewed from above, most beetles appear to have three clear sections, but this is deceptive: on the beetle's upper surface, the middle "section" is a hard plate called the pronotum, which is only the front part of the thorax; the back part of the thorax is concealed by the beetle's wings. This further segmentation is usually best seen on the abdomen.
Acilius sulcatus, a diving beetle showing hind legs adapted for life in water

The legs, which are multi-segmented, end in two to five small segments called tarsi. Like many other insect orders beetles bear claws, usually one pair, on the end of the last tarsal segment of each leg. While most beetles use their legs for walking, legs may be variously modified and adapted for other uses. Among aquatic families – Dytiscidae, Haliplidae, many species of Hydrophilidae and others – the legs, most notably the last pair, are modified for swimming and often bear rows of long hairs to aid this purpose. Other beetles have fossorial legs that are widened and often spined for digging. Species with such adaptations are found among the scarabs, ground beetles, and clown beetles (family Histeridae). The hind legs of some beetles, such as flea beetles (within Chrysomelidae) and flea weevils (within Curculionidae), are enlarged and designed for jumping.

The elytra is connected to the pterathorax; being called as such because it is where the wings are connected (pteron meaning "wing" in Greek).[1] The elytra are not used for flight, but tend to cover the hind part of the body and protect the second pair of wings (alae). The elytra must be raised in order to move the hind flight wings. A beetle's flight wings are crossed with veins and are folded after landing, often along these veins, and are stored below the elytra. In some beetles, the ability to fly has been lost. These include some ground beetles (family Carabidae) and some "true weevils" (family Curculionidae), but also some desert and cave-dwelling species of other families. Many of these species have the two elytra fused together, forming a solid shield over the abdomen. In a few families, both the ability to fly and the elytra have been lost, with the best known example being the glow-worms of the family Phengodidae, in which the females are larviform throughout their lives.

Abdomen

The abdomen is the section behind the metathorax, made up of a series of ring, each with a hole for breathing and respiration, called spiracles; composing three different segmented sclerites: the tergum, pleura, and the sternum. The tergum in almost all species is membranous, or usually soft and concealed by the wings and elytra when not in flight. The pleura (singular: pleuron) are usually small or hidden in some species, with each pleuron having a single spiracle. The sternum is the most widely visible part of the abdomen, being a more or less scelortized segment. The abdomen itself does not have any appendages, however some species (e.g., Mordellidae) have articulating sternal lobes.[7]

Internal Morphology
A diagram showing the general internal anatomy of species of coleoptera.

The digestive system of beetles is primarily based on plants which they for the most part feed upon, with mostly the anterior midgut performing digestion. Although, in predatory species (e.g., Carabidae) most digestion occurs in the crop by means of midgut enzymes. In Elateridae species, the predatory larvae defecate enzymes on their prey, with digestion being extraorally.[1] The alimentary canal basically comprises of a short narrow pharynx, a widened expansion, the crop and a poorly developed gizzard. After there is a midgut, that varies in dimensions between species, with a large amount of cecum, with a hingut, with varying lengths. There are typically four to six Malpighian tubules. [5]

The nervous system in beetles contains all the types found in insects, varying between different species. With three thoracic and seven or eight abdominal ganglia can be distinguished to that in which all the thoracic and abdominal ganglia are fused to form a composite structure. Oxygen is obtained via a tracheal system. Air enters a series of tubes along the body through openings called spiracles, and is then taken into increasingly finer fibers.[1] Pumping movements of the body force the air through the system. Beetles have hemolymph instead of blood like other insect species, the open circulatory system of the beetle is driven by a tube-like heart attached to the top inside of the thorax. Some species of diving beetles (Dytiscidae) carry a bubble of air with them whenever they dive beneath the water surface. This bubble may be held under the elytra or it may be trapped against the body using specialized hairs. The bubble usually covers one or more spiracles so the insect can breathe air from the bubble while submerged. An air bubble provides an insect with only a short-term supply of oxygen, but thanks to its unique physical properties, oxygen will diffuse into the bubble and displacing the nitrogen, called passive diffusion, however the volume of the bubble eventually diminishes and the beetle will have to return to the surface.[8]

Specialized organs

Different glands specialize for different pheromones produced for finding mates. Pheromones from species of Rutelinea are produced from epithelial cells lining the inner surface of the apical abdominal segments or amino acid based pheromones of Melolonthinae from eversible glands on the abdominal apex. Other species produce different types of pheromones. Dermestids produce esters, and species of Elateridae produce fatty-acid-derived aldehydes and acetates.[1] For means of finding a mate also, fireflies (Lampyridae) utilized modified fat body cells woth transparent surfaces backed with reflective uric acid crystals to biosynthetically produce light, or bioluminescence. The light produce is highly efficient, as it is produced by oxidation of luciferin by the enzymes luciferase in the presence of ATP (adenosine triphospate) and oxygen, producing oxyluciferin, carbon dioxide, and light.[1]

A notable number of species have developed special glands that produce chemicals for deterring predators (see Defense and predation). The Ground beetle's (of Carabidae) defensive glands, located at the posterior, produce a variety of hydrocarbons, aldehydes, phenols, quinones, esters, and acids released from an opening at the end of the abdomen. While african carabid beetles (e.g., Anthia and Thermophilium) employ the same chemicals as ants: formic acid.[9] While Bombardier beetles have well developed, like other carabid beetles, pygidial glands that empty from the lateral edges of the intersegment membranes between the seventh and eighth abdominal segments. The gland is made of two containing chambers. The first holds hydroquinones and hydrogen peroxide, with the second holding just hydrogen peroxide plus catalases. These chemicals mix and result in an explosive ejection, forming temperatures of around 100 C, with the brake down of hydroquinone to H2 + O2 + quinone, with the O2 propelling the excretion.[1]

Tympanal organs or hearing organs, which is a membrane (tympanum) stretched across a frame backed by an air sac and associated sensory neurons, are described in two families.[10] Several species of the genus Cicindela (Cicindelidae) have ears on the dorsal surface of the first abdominal segment beneath the wing; two tribes tribes in the family Dynastinae (Scarabaeidae) have ears just beneath the pronotal shield or neck membrane. The ears of both families are to ultrasonic frequencies, with strong evidence that they function to detect the presence of bats via there ultrasonic echolocation. Even though beetles constitute a large order and live in a variety of niches, examples of hearing is surprisingly lacking in species, though it is likely that most are just undiscovered.[1]

Reproduction and development

Beetles are members of Endopterygota, which means like most other insects under complete metamorphosis, which consists of four main stages: the egg, the larva, the pupa, and the imago or adult. The larvae are commonly called grubs and the pupa are called cocoons.
[edit] Mating
Punctate flower chafers (Neorrhina punctata, Scarabaeidae) mating

Beetles may display extremely intricate behavior when mating. Pheromone communication is likely to be important in the location of a mate. Different species use different chemicals for their pheromones. Some scarab beetles (e.g., Rutelinae) utilize pheromones derived from fatty acid synthesis, while other scarab beetles use amino acids and terpenoid compounds (e.g., Melolonthinae). Another way species of Coleoptera find mates is the use of biosynthesized light, or bioluminescence. This special form of a mating call is confined to fireflies (Lampyridae) by the use of abdominal light producing organs. The males and females engage in complex dialogue before mating, identifying different species by differences in duration, flight patterns, composition, and intensity.[1]

Before mating male ans females may engage in various forms of behavior. Males and females may stridulate, or vibrate the object they are on. In some species (e.g., Meloidae) the male climbs onto the dorsum of the female and stroke his antennae on her head, palps and antennae. In the genus Eupompha of said family, the males draws the antennae alonga the longitudinal vertex on the male. They may not mate at all if they do not perform the precopulatory ritual.[1]

Conflict can play a part in the mating rituals of species such as burying beetles (genus Nicrophorus) where conflicts between males and females rage until only one of each is left, thus ensuring reproduction by the strongest and fittest. Many male beetles are territorial and will fiercely defend their small patch of territory from intruding males. In such species, the males may often have horns on the head and/or thorax, making their overall body lengths greater than those of the females, unlike most insects. Pairing is generally short but in some cases will last for several hours. During pairing sperm cells are transferred to the female to fertilize the egg.[5]

Life Cycle
The life cycle of the stag beetle, including the 3 instars.

Egg

A single female may lay from several dozen to several thousand eggs during her lifetime. Eggs are usually laid according to the substrate the larva will feed on upon hatching. Among others, they can be laid loose in the substrate (e.g. flour beetle), laid in clumps on leaves (e.g. Colorado potato beetle), or individually attached (e.g. mungbean beetle and other seed borers) or buried in the medium (e.g. carrot weevil).

Parental care varies between species, ranging from the simple laying of eggs under a leaf to certain scarab beetles, which construct underground structures complete with a supply of dung to house and feed their young.[1] Other beetles are leaf rollers, biting sections of leaves to cause them to curl inwards, then laying their eggs, thus protected, inside.[1]

Larva
A scarabaeiform larvae known as a Curl grub.

The larva is usually the principal feeding stage of the beetle life cycle. Larvae tend to feed voraciously once they emerge from their eggs. Some feed externally on plants, such as those of certain leaf beetles, while others feed within their food sources. Examples of internal feeders are most Buprestidae and longhorn beetles. The larvae of many beetle families are predatory like the adults (ground beetles, ladybirds, rove beetles). The larval period varies between species but can be as long as several years. The larva are highly varied amongst species, with a well developed and sclerotized head and have distinguishable thoracic and abdominal segments (usually the tenth, though sometimes the eight or ninth).[5]

Beetle larvae can be differentiated from other insect larvae by their hardened, often darkened head, the presence of chewing mouthparts, and spiracles along the sides of the body. Like adult beetles, the larvae are varied in appearance, particularly between beetle families. Beetles whose larvae are somewhat flattened and are highly mobile are the ground beetles, some rove beetles, and others; their larvae are described as campodeiform. Some beetle larvae resemble hardened worms with dark head capsules and minute legs. These are elateriform larvae, and are found in the click beetle (Elateridae) and darkling beetle (Tenebrionidae) families. Some elateriform larvae of click beetles are known as wireworms. Beetles in the families of the Scarabaeoidea have short, thick larvae described as scarabaeiform, but more commonly known as grubs.

All beetle larvae go through several instars, which are the developmental stages between each moult. In many species the larvae simply increase in size with each successive instar as more food is consumed. In some cases, however, more dramatic changes occur. Among certain beetle families or genera, particularly those that exhibit parasitic lifestyles, the first instar (the planidium) is highly mobile in order to search out a host, while the following instars are more sedentary and remain on or within their host. This is known as hypermetamorphosis; examples include the blister beetles (family Meloidae) and some rove beetles, particularly those of the genus Aleochara.

Pupa

As with all endopterygotes, beetle larvae pupate, and from this pupa emerges a fully formed, sexually mature adult beetle, or imago. Adults have an extremely variable lifespan, from weeks to years, depending on the species. In some species the pua may go through all four forms during its development, called hypermetamorphosis (e.g., Meloidae). Pupa always have no mandibles, or abecticous. In most, the appendags are not attached to the pupa, or they are exarate; with most being obtect in form. [5]

Behavior

Locomotion
Photinus pyralis in midflight.Photinus pyralis Firefly

Aquatic beetles use several techniques for retaining air beneath the water's surface. Beetles of the family Dytiscidae hold air between the abdomen and the elytra when diving. Hydrophilidae have hairs on their under surface that retain a layer of air against their bodies. Adult crawling water beetles use both their elytra and their hind coxae (the basal segment of the back legs) in air retention,[11] while whirligig beetles simply carry an air bubble down with them whenever they dive.

The elytra allows beetles and weevils to both fly and move through confined spaces. Doings so by folding the delicate wings under the elytra while not flying, and folding their wings out just before take off. The unfolding and folding of the wings is operated by muscles attached to the wing base; as long as the tension on the radial and cubital veins remains, the wings remain straight. In day-flying species (e.g. Buprestidae, Scarabaeidae), flight does not include large amounts of lifting of the elytra, having the metathorac wings extended under the lateral elytra margins.[1]

Communication

Beetles have a variety of ways to communicate. Some of which include a sophisticated chemical language through the use of pheromones. From the host tree, the mountain pine beetle have many forms of communication. They can emit both an aggregative pheromone and an anti-aggregative pheramone. The aggregative pheromone attracts other beetles to the tree, and the anti-aggregative pheromone neutralizes the aggregative pheromone. This helps to avoid the harmful effects of having too many beetles on one tree competing for resources. The mountain pine beetle can also stridulate to communicate, or rub body parts together to create sound, having a “scraper” on their abdomen that they rub against a grooved surface on the underside of their left wing cover to create a sound that is not audible to humans. Once the female beetles have arrived on a suitable pine tree host, they begin to stridulate and produce aggregative pheromones to attract other unmated males and females. New females arrive and do the same as they land and bore into the tree. As the males arrive, they enter the galleries that the females have tunneled, and begin to stridulate to let the females know they have arrived, and to also warn others that the female in that gallery is taken. At this point, the female stops producing aggregative pheromones and starts producing anti-aggregative pheromone to deter more beetles from coming.[12]

Since species of Coleoptera use environmental stimuli to communicate, they are affected by the climate. Microclimates, such as wind or temperature, can disturb the use of pheromones; wind would blow the pheromones while they ravel through the air. Stridulating can be interrupted when the stimulus is vibrated by something else.[12]

Parental care
a dung beetle rolling dung, near the giant tomb Sa Ena 'e Thomes, Sardinia, Italy

Among insect, parental care is very uncommon, only found in a few species. Some beetles also display this unique social behavior.[1] One theory states why there is parental care is that it is necessary for the survival of the larvae, protecting them from adverse environmental conditions and predators. One species, a rover beetle (Bledius spectabilis) displays both causes for parental care: physical and biotic environmental factors. Said species lives in salt marshes, so the eggs and/or larvae are endangered by the rising tide. The maternal beetle will patrol the eggs and larva and apply the appropriate burrowing behavior the keep them from flooding and from asphyxiating. Another advantage is that the mother protects the eggs and larvae from the predatory carabid beetles species Dicheirotrichus gustavi and from the parasitoid wasp species Barycnemis blediator. Up to 15% of larvae are killed by this parasitoid wasp, being only protected by maternal beetles in their dens.[13]

Some species of Dung beetles also display a form of parental care. Dung beetles, from which their name is derived, collect the feces, or "dung" and roll it into a ball, sometimes being up to 50 times their own wait; albeit sometimes it is also used to store food. Usually it is the male that rolls the ball, with the female hitch-hiking or simply following behind. In some cases the male and the female roll together. When a spot with soft soil is found, they stop and bury the dung ball. They will then mate underground. After the mating, both or one of them will prepare the brooding ball. When the ball is finished, the female lays eggs inside it, a form of mass provisioning. Some species do not leave after this stage, but remain to safeguard their offspring.[14]
Mylabris pustulata (Meloidae) feeding on the petals of Ipomoea carnea

Feeding

Besides being abundant and varied, beetles are able to exploit the widediversity of food sources available in their many habitats. Some are omnivores, eating both plants and animals. Other beetles are highly specialized in their diet. Many species of leaf beetles, longhorn beetles, and weevils are very host specific, feeding on only a single species of plant. Ground beetles and rove beetles (family Staphylinidae), among others, are primarily carnivorous and will catch and consume many other arthropods and small prey such as earthworms and snails. While most predatory beetles are generalists, a few species have more specific prey requirements or preferences.[15]

Decaying organic matter is a primary diet for many species. This can range from dung, which is consumed by coprophagous species such as certain scarab beetles (family Scarabaeidae), to dead animals, which are eaten by necrophagous species such as the carrion beetles (family Silphidae). Some of the beetles found within dung and carrion are in fact predatory, such as the clown beetles, preying on the larvae of coprophagous and necrophagous insects.

Ecology

Defense and predation
Beetles may be preyed upon by other insects such as robber flies

Beetles and their larvae have a variety of strategies to avoid being attacked by predators or parasitoids. These include camouflage, mimicry, toxicity, and active defense.

Camouflage involves the use of coloration or shape to blend into the surrounding environment. This sort of protective coloration is common and widespread among beetle families, especially those that feed on wood or vegetation, such as many of the leaf beetles (family Chrysomelidae) or weevils. In some of these species, sculpturing or various colored scales or hairs cause the beetle to resemble bird dung or other inedible objects. Many of those that live in sandy environments blend in with the coloration of the substrate.[9]:126 For example, the Giant African longhorn beetle (Petrognatha gigas) which resembles the mossa nd bark of the tree from which it feeds on.

Another defense that often uses color or shape to deceive potential enemies is mimicry. A number of longhorn beetles (family Cerambycidae) bear a striking resemblance to wasps, which helps them avoid predation even though the beetles are in fact harmless. This defense can be found to a lesser extent in other beetle families, such as the scarab beetles. Beetles may combine their color mimicry with behavioral mimicry, acting like the wasps they already closely resemble. Many beetle species, including ladybirds, blister beetles, and lycid beetles can secrete distasteful or toxic substances to make them unpalatable or even poisonous. These same species often exhibit aposematism, where bright or contrasting color patterns warn away potential predators, and there are, not surprisingly, a great many beetles and other insects that mimic these chemically protected species.[9]
Clytus arietis (Cerambycidae), a wasp mimic

Chemical defense is another important defense found amongst species of Coleoptera, usually being advertised by bright colors. Others may utilize behaviors that would be done when releasing noxious chemicals (e.g., Tenebrionidae). Chemical defense may serve purposes other then just protection from vertebrates, such as protection from a wide range of microbes, and repellents. Some species release chemicals in the form of a spray with surprising accuracy, such as ground beetles (Carabidae), may spray chemicals from their abdomen to repel predators. Some species take advantage of the plants from which they feed, and sequester the chemicals from the plant that would protect it and incorporate into their own defense. African carabid beetles (e.g., Anthia and Thermophilium) employ the same chemicals sued by ants, while Bombardier beetles have a their own unique separate gland, spraying potential predators from far distances.[9]:126

Large ground beetles and longhorn beetles may defend themselves using strong mandibles and/or spines or horns to forcibly persuade a predator to seek out easier prey.[9]:126 Many species have large protrusions from their thorax and head such as the Rhinoceros beetle, which can be used to defended themselves from predators. Many species of weevil that feed out in the open on leaves of plants react to attack by employing a "drop-off reflex." Even further, some will combine it with thanatosis, which they will close up their legs, antennae, mandibles, ect. and use their cryptic coloration to blend in with the background. Species with varied coloration do not do this as they can not camaflouge.[1]:199

Parasitism

There are over 1000 known species of beetles to be either parasitic, predatory, or commensals in the nests of ants.[16] Most beetle larvae can be considered parasites, as they feed on the plants and live inside the bark of trees and plants. Though there are a few species that are ectoparasitic to mammals, such as Platypsyllus castoris, which is affects beavers (Castor spp.). This beaver beetle parasite is a parasite as both an adult and a larva, with the appropriate modifications. They are wingless and eyeless with a striking dorso-ventral flattening. Additionally, P. castoris, has a remarkably modified antennal club, with antennomeres 3-11 shortened, globularly compacted, and partly enclosed in a scoop shaped antennomere 2, as is also found in Gyrinidae and Dryopidae.[17]

other parasites include those who are parasitoids of other invertebrates, such as the small hive beetle (Aethina tumida) infecting Honey bee hives. The larvae tunnel through comb towards stored honey or pollen, damaging or destroying cappings and comb in the process. Larvae defecate in honey and the honey becomes discolored from the feces, which causes fermentation and a frothiness in the honey; the honey develops a characteristic odor of decaying oranges. Damage and fermentation cause honey to run out of combs, destroing large portions in hives and sometimes the extracting rooms. Heavy infestations cause bees to abscond; some beekeepers have reported the rapid collapse of even strong colonies.[18]

Mutualism

Amongst most orders of insects, mutualism is not common, however there are some examples in species of Coleoptera. Such as the Ambrosia beetle, the Ambrosia fungus, and probably bacteria. The beetles excavate tunnels in dead trees in which they cultivate fungal gardens, their sole source of nutrition. After landing on a suitable tree, an ambrosia beetle excavates a tunnel in which it releases spores of its fungal symbiont. The fungus penetrates the plant's xylem tissue, digests it, and concentrates the nutrients on and near the surface of the beetle gallery; so the weevils and the fungus both benefit. The beetles can not eat due to toxins, which uses its relationship with fungi to help overcome it's host tree defenses and to provide nutrition for their larvae.[19]
1: The adult beetle burrows hole into wood and lays eggs, carrying fungal spores in their mycangia
2: The larva feeds on the fungus which digest the wood, removing toxins: they mutually benefit
3: The larva pupates and then ecloses.

The beetle-fungal mutualism is chemically mediated by a bacterially produced polyunsaturated peroxide. The molecule's selective toxicity toward the beetle's fungal antagonist, combined with the prevalence and localization of its bacterial source, indicates an insect-microbe association that is both mutualistic and coevolved. This unexpected finding in a well-studied system indicates that mutualistic associations between insects and antibiotic-producing bacteria are more common than currently recognized and that identifying their small-molecule mediators can provide a powerful search strategy for therapeutically useful antimicrobial compounds.[19][20]

Commensalism

Psuedoscorpions are small arachnids with a flat, pear-shaped body and pincers that resemble those of scorpions though are not, usually ranging from 2 to 8 millimetres (0.08 to 0.31 in) in length.[21] Their small size allows them to hitch rides under the elytra of a giant harlequin beetle to be dispersed over wide areas while simultaneously being protected from predators. They may also find mating partners as other individuals join them on the beetle. This would be a form of parasitism if the beetle was harmed in the process, however the beetle is, presumably, unaffected by the presence of the hitchhikers.[22][23]

Phylogeny and systematics
Baltic amber inclusions, from the Eucene era, 50 million years old (Coleoptera, Scraptiidae)

Fossil record

A 2007 study based on DNA of living beetles and maps of likely beetle evolution indicated that beetles may have originated during the Lower Permian, up to 299 million years ago.[24] In 2009, a fossil beetle was described from the Pennsylvanian of Mazon Creek, Illinois, pushing the origin of the beetles to an earlier date, 318 to 299 million years ago.[25]Fossils from this time have been found in Asia and Europe, for instance in the red slate fossil beds of Niedermoschel near Mainz, Germany.[26] Further fossils have been found in Obora, Czechia and Tshekarda in the Ural mountains, Russia.[27]However, there are only a few fossils from North America before the middle Permian, although both Asia and North America had been united to Euramerica. The first discoveries from North America were made in the Wellington formation of Oklahoma and were published in 2005 and 2008.[28][29]

As a consequence of the P-Tr Mass Extinction at the border of Permian and Triassic, there is only little fossil record of insects including beetles from the Lower Triassic.[30] However, there are a few exemptions, like in Eastern Europe: At the Babiy Kamen site in the Kuznetsk Basin numerous beetle fossils were discovered, even entire specimen of the infraorders Archostemata (e.i., Ademosynidae, Schizocoleidae), Adephaga (e.i., Triaplidae, Trachypachidae) and Polyphaga ( e.i., Hydrophilidae, Byrrhidae, Elateroidea) and in nearly a perfectly preserved condition.[31] However, species from the families Cupedidae and Schizophoroidae are not present at this site, whereas they dominate at other fossil sites from the Lower Triassic. Further records are known from Khey-Yaga, Russia in the Korotaikha Basin.[28] There are many important sites from the Jurassic, with more than 150 important sites with beetle fossils, the majority being situated in Eastern Europe and North Asia. In North America and especially in South America and Africa the number of sites from that time period is smaller and the sites have not been exhaustively investigated yet. Outstanding fossil sites include [[Solnhofen] in Upper Bavaria, Germany,[32] Karatau in South Kazakhstan,[33] the Yixian formation in Liaoning, North China[34] as well as the Jiulongshan formation and further fossil sites in Mongolia. In North America there are only a few sites with fossil records of insects from the Jurassic, namely the shell limestone deposits in the Hartford basin, the Deerfield basin and the Newark basin.[35][28]
Fossil buprestid beetle from the Eocene Messel pit, which retains its structural color

There is a large number of important fossil sites worldwide containing beetles from the Cretaceous. Most of of them are located in Europe and Asia and belong to the temperate climate zone during the Cretaceous. A few of the fossil sites mentioned in the chapter Jurassic also shed some light on the early cretaceous beetle fauna (e.g. the Yixian formation in Liaoning, North China).[34] Further important sites from the Lower Cretaceous include the Crato Fossil Beds in the Araripe basin in the Ceará, North Brazil as well as overlying Santana formation, with the latter was situated near the paleoequator, or the position of the earth's equator in the geologic past as defined for a specific geologic period. In Spain there are important sites near Montsec and Las Hoyas. In Australia the Koonwarra fossil beds of the Korumburra group, South Gippsland, Victoria is noteworthy. Important fossil sites from the Upper Cretaceous are Kzyl-Dzhar in South Kazakhstan and Arkagala in Russia.[28]
The phylogenetic growth of three different trophic levels in Coleoptera by number of genera.

Evolution

The oldest known insect that resembles species of Coleoptera date back to the Lower Permian (270 mya), though they instead have 13-segmented antennae, elytra with more fully developed venation and more irregular longitudinal ribbing, and an abdomen and ovipositor extending beyond the apex of the elytra. The oldest true beetle, that is having features that include 11-segmented antennae, regular longitudinal ribbing on the elytra, and having genitalia that are internal.[1]:186 At the end of the Permian, the biggest mass extinction in the history history took place, collectively called the Permian–Triassic extinction event: 30% of all insect species became extinct, however, it is the only mass extinction of insects in Earth's history until today.[28]

Consequence do to the P-Tr extinction, there is only little fossil record of insects including beetles from the Lower Triassic (220 mya). Around this time, during the Late Triassic, mycetophagous, or fungus feeding species (e.i. Cupedidae) appear in the fossil record. In the stages of the Upper Triassic representatives of the algophagous, or algae feeding species (e.i. Triaplidae and Hydrophilidae) begin to appear, as well as predatory water beetles. The first primitive weevils appear (e.i. Obrienidae), as well as the first representatives of the rove beetles (e.i. Staphylinidae), which show no marked difference in physique compared to recent species.[28]

During the Jurassic (210-145 mya) there was a dramatic increase in the known diversity of family-level Coleoptera.[28] This includes the development and growth of carnivorous and herbivorous species. Species of the superfamily Chrysomeloidea are believed to have developed around the same time, which include a wide array of plant host ranging from cycads and conifers, to angiosperms.[1]:186 Close to the Upper Jurassic, the portion of the Cupedidae decreased, however at the same time the diversity of the early plant eating, or phytophagous species increased. Most of the recent phytophagous species of Coleoptera feed on flowering plants or angiosperms. It is believed that the increase in diversity of the angiosperms also influenced the diversity of the phytophagous species, which doubled during the Middle Jurassic. However, recently doubts have been raised since the increase of the number of beetle families during the Cretaceous does not correlate with the increase of the number of angiosperm species.[36] Also around the same time, numerous primitive weevils (e.i., Curculionoidea) and click beetles (e.i. Elateroidea) appeared. Also first jewel beetles (e.i., Buprestidae) are present, however, they were rather rare until the Cretaceous.[37][38][39] The first scarab beetles would apear around this time, however they where not coprophagous, or feeding upon fecal matter, presumably feeding upon the rotting wood with the help of fungus, and early example of a mutualistic relationship (see the Mutualism section ).

The Cretaceous witness the initiation of the most recent round of southern landmass fragmentation, via the opening of the southern Atlantic ocean and the isolation of New Zealand, while the South America, Antarctica, and Australia grew more distant.[1]:186 During the Cretaceous the diversity of Cupedidae and Archostemata decreased considerably. Predatory ground beetles (Carabidae) and rove beetles (Staphylinidae) began to distribute into different patterns: whereas the Carabidae predominantly occurred in the warm regions, the Staphylinidae and click beetles (Elateridae) preferred many areas with temperate climate. Likewise, predatory species of Cleroidea and Cucujoidea, hunted their prey under the bark of trees together with the jewel beetles (Buprestidae). The jewel beetles diversity increased rapidly during the Cretaceous, as they were the primary consumers of wood,[40] while longhorn beetles (Cerambycidae) were rather rare and their diversity increased only towards the end of the Upper Cretaceous.[28] The first coprophagous beetles have been recorded from the Upper Cretaceous,[41] and are believed to have lived on the excrement of herbivorous dinosaurs, however there is still a discussion, whether the beetles where always tied to mammals during its development.[42] Also, the first species with an adaption of both larvae and adults to the aquatic lifestyle are found. Whirligig beetles (Gyrinidae) were moderately diverse, although other early beetles (e.i., Dytiscidae) where less, with the most widespread being the species of Coptoclavidae, which preyed on aquatic fly larvae.[28]

The time between the Paleogene and the Neogene, or more recent history is where today's beetles developed. During this time, the continents began to situate themselves to where we see them today. Around 5 million years ago the land bridge between South America and North America was formed, and this is when fauna exchange between Asia and North America started. Even though many recent genera and species already existed during the Miocene, however, their distribution differed considerably from today's.[28]

Phylogeny

These suborders diverged in the Permian and Triassic. Their phylogenetic relationship is uncertain, with the most popular hypothesis being that Polyphaga and Myxophaga are most closely related, with Adephaga as the sister group to those two, and Archostemata as sister to the other three collectively.[1]:186[43] Although there are six other competing hypotheses, the other most widely discussed hypothesis is Myxophaga is the sister group of all remaining beetles rather then just Polyphaga.[44] Evidence for a close relationship of the two suborders, Polyphaga and Myxophaga, includes the shared reduction in the number of larval leg articles. further considered the Adephaga as sister to Myxophaga and Polyphaga, based on their completely sclerotized elytra, reduced number of crossveins in the hind wings, and folded (as opposed to rolled) hind wings of those three suborders.

Recent cladistic analysis of some of the structural characteristics supports the Polyphaga and Myxophaga hypothesis.[43] The composition of the clade Coleoptera is not in dispute, with the exception of the twisted-wing parasites, Strepsiptera. These odd insects have been regarded as related to the beetle families Rhipiphoridae and Meloidae, with which they share first instar larvae that are active, host-seeking triungulins and later instar larvae that are endoparasites of other insects, or as the sister group of beetles, or more distantly related to insects.[44][45]

Taxonomy

There are about 450,000 species of beetles – representing about 40% of all known insects. Such a large number of species poses special problems for classification, with some families consisting of thousands of species and needing further division into subfamilies and tribes. This immense number of species allegedly led evolutionary biologist J. B. S. Haldane to quip, when some theologians asked him what could be inferred about the mind of the Creator from the works of His Creation, that God displayed "an inordinate fondness for beetles".[46]

* Polyphaga is the largest suborder, containing more than 300,000 described species in more than 170 families, including rove beetles (Staphylinidae), scarab beetles (Scarabaeidae), blister beetles (Meloidae), stag beetles (Lucanidae) and true weevils (Curculionidae). These beetles can be identified by the cervical sclerites (hardened parts of the head used as points of attachment for muscles) absent in the other suborders.
* Adephaga contains about 10 families of largely predatory beetles, includes ground beetles (Carabidae), Dytiscidae and whirligig beetles (Gyrinidae). In these beetles, the testes are tubular and the first abdominal sternum (a plate of the exoskeleton) is divided by the hind coxae (the basal joints of the beetle's legs).
* Archostemata contains four families of mainly wood-eating beetles, including reticulated beetles (Cupedidae) and the telephone-pole beetle.
* Myxophaga contains about 100 described species in four families, mostly very small, including Hydroscaphidae and the genus Sphaerius.

Relationship to people

As pests
Larvae of the Colorado potato beetle, Leptinotarsa decemlineata

Many agricultural, forestry, and household insect pests are beetles. These include the following:

* The Colorado potato beetle, Leptinotarsa decemlineata, is a notorious pest of potato plants. Crops are destroyed and the beetle can only be treated by employing expensive pesticides, many of which it has begun to develop resistance to. As well as potatoes, suitable hosts can be a number of plants from the potato family (Solanaceae), such as nightshade, tomato, aubergine and capsicum.
* The boll weevil, Anthonomus grandis, has cost cotton producers in the United States billions of dollars since it first entered that country.
* The bark beetles Hylurgopinus rufipes and Scolytus multistriatus, the elm leaf beetle, Pyrrhalta luteola, and other beetles attack elm trees. The bark beetles are important elm pests because they carry Dutch elm disease as they move from infected breeding sites to feed on healthy elm trees. The spread of the fungus by the beetle has led to the devastation of elm trees in many parts of the Northern Hemisphere, notably in Europe and North America.

Red flour beetle, Tribolium castaneum

* Flour beetles are pests of cereal silos. They feed on wheat and other grains and are adapted to survive in very dry environments. They are a major pest in the agricultural industry and are highly resistant to insecticides.
* The death watch beetle, Xestobium rufovillosum, (family Anobiidae) is of considerable importance as a pest of older wooden buildings in Great Britain. It attacks hardwoods such as oak and chestnut, always where some fungal decay has taken or is taking place. It is thought that the actual introduction of the pest into buildings takes place at the time of construction.
* Coconut hispine beetle, Brontispa longissima, feeds on young leaves and damages seedlings and mature coconut palms. On September 27, 2007, Philippines' Metro Manila and 26 provinces were quarantined due to having been infested with this pest (to save the $800-million Philippine coconut industry).[47]
* The mountain pine beetle normally attacks mature or weakened lodgepole pine. It can be the most destructive insect pest of mature pine forests. The current infestation in British Columbia is the largest Canada has ever seen.[48]

As beneficial
Coccinella septempunctata, a beneficial beetle

A number of insects are beneficial to humans, usually by controlling the populations of pests.

Both the larvae and adults of some ladybugs (family Coccinellidae) are found in aphid colonies. Other ladybugs feed on scale insects and mealybugs. If normal food sources are scarce, they may feed on other things, such as small caterpillars, young plant bugs, honeydew and nectar.

Ground beetles (family Carabidae) are common predators of many different insects and other arthropods, including fly eggs, caterpillars, wireworms and others.

Plant-feeding beetles are often important beneficial insects, controlling problem weeds. Some flea beetles of the genus Aphthona feed on Euphorbia esula (leafy spurge, Euphorbiaceae), a considerable weed of rangeland in western North America.

Dung beetles (Coleoptera, Scarabidae) have been successfully used to reduce the populations of pestilent flies and parasitic worms that breed in cattle dung. The beetles make the dung unavailable to breeding pests by quickly rolling and burying it in the soil, with the added effect of improving soil fertility and nutrient cycling. The Australian Dung Beetle Project (1965–1985), led by Dr. George Bornemissza of the Commonwealth Scientific and Industrial Research Organization introduced species of dung beetle to Australia from South Africa and Europe and effectively reduced the bush fly (Musca vetustissima) population by 90%.

Some farmers develop beetle banks to foster and provide cover for beneficial beetles.

Some beetles of the family Dermestidae are often used in taxidermy to clean bones of remaining flesh.

As food

In ancient culture
Ancient Egyptian scene depicting a scarab beetle

Several species of dung beetle, most notably Scarabaeus sacer (often referred to as "scarab"), enjoyed a sacred status among the ancient Egyptians, as the creatures were likened to the major god Khepri. Many thousands of amulets and stamp seals have been excavated that depict the scarab. In many artifacts, the scarab is depicted pushing the sun along its course in the sky, much as scarabs push or roll balls of dung to their brood sites. During and following the New Kingdom, scarab amulets were often placed over the heart of the mummified deceased.

Some tribal groups, particularly in tropical parts of the world, use the colorful, iridescent elytra of certain beetles, especially certain Scarabaeidae, in ceremonies and as adornment.

In modern culture

The study of beetles is called coleopterology (from Coleoptera, see above, and Greek -λογία, -logia), and its practitioners are coleopterists. Coleopterists have formed organizations to facilitate the study of beetles. Among these is The Coleopterists Society, an international organization based in the United States. Such organizations may have both professionals and amateurs interested in beetles as members.

Research in this field is often published in peer-reviewed journals specific to the field of coleopterology, though journals dealing with general entomology also publish many papers on various aspects of beetle biology. Some of the journals specific to beetle research are:

* The Coleopterist (United Kingdom beetle fauna)
* The Coleopterists Bulletin (published by The Coleopterists Society)
* Elytron (published by the European Association of Coleopterology)

References

General references

* Poul Beckmann, Living Jewels: The Natural Design of Beetles ISBN 3-7913-2528-0
* Arthur V. Evans, Charles Bellamy, and Lisa Charles Watson, An Inordinate Fondness for Beetles ISBN 0-520-22323-3
* Cooter J. & Barclay M.V.L. (eds.) (2006) A Coleopterist’s Handbook. Amateur Entomological Society. 439 pages. ISBN 0-900054-70-0
* Entomological Society of America, Beetle Larvae of the World ISBN 0-643-05506-1
* David Grimaldi, Michael S. Engel, Evolution of the Insects ISBN 0-521-82149-5
* Ross H. Arnett, Jr. and Michael C. Thomas, American Beetles (CRC Press, 2001–2002). ISBN 0-8493-1925-0
* K. W. Harde, A Field Guide in Color to Beetles ISBN 0-7064-1937-5 Pages 7–24
* White, R.E. 1983. Beetles. Houghton Mifflin Company, New York, NY. ISBN 0-395-91089-7

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