Carabidae

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
Subordo: Adephaga

Familia: Carabidae
Subfamiliae (22 + 2†): Apotominae – Brachininae – Broscinae – Carabinae – Cicindelinae – Cicindinae – Elaphrinae – Gehringiinae – Harpalinae – Hiletinae – Loricerinae – Melaeninae – Migadopinae – Nebriinae – Nototylinae – Omophroninae – Patrobinae – Paussinae – Psydrinae – Scaritinae – Siagoninae – Trechinae – †Conjunctiinae – †Protorabinae

Genera overview (1997 + 13†):

A

Abacaelostus – Abacetodes – Abacetus – Abacidus – Abacillius – Abacillodes – Abacoleptus – Abacomorphus – Abacophrastus – Abacops – Abaditicus – Abaris – Abax – Abrodiella – Abropus – Abroscelis – Acalathus – Acallistus – Acanthoferonia – Acanthoodes – Acanthoscelis – Accoella – Achaetocephala – Acheroniotes – Acinopus – Acrogenys – Actenoncus – Actenonyx – Actodus – Acupalpus – Acutosternus – Adelopomorpha – Adelotopus – Adotela – Adriaphaenops – Adrimus – Aenigma – Aeolodermus – Aepiblemus – Aepopsis – Aepsera – Aepus – Afroclivina – Afrodromius – Afrogehringia – Afromizonus – Afrosyleter – Afrotachys – Afrotarus – Agastus – Agelaea – Agonica – Agonidium – Agonidus – Agonobembix – Agonocheila – Agonops – Agonoriascus – Agonorites – Agonotrechiotes – Agonotrechus – Agonum – Agostinia – Agra – Akephorus – Alachnothorax – Alanorites – Albanotrechus – Alkestis – Allardina – Allegrettia – Allendia – Allocinopus – Allocota – Allophanes – Allophanopsis – Allosiopelus – Allotrechiama – Allotriopus – Alpiodytes – Altagonum – Amara – Amaroschema – Amarotypus – Amblops – Amblycheila – Amblycoleus – Amblygnathus – Amblystogenium – Amblystomus – Amblytelus – Amelus – Amerinus – Amerizus – Ameroduvalius – Ametroglossus – Amorphomerus – Amphasia – Amphimasoreus – Amphimenes – Amphimenoides – Amphithasus – Amphithasus – Analoma – Anasis – Anatrichis – Anaulacus – Anchista – Anchomenidius – Anchomenus – Anchonoderus – Anchonoderus – Anchotrechus – Ancus – Ancystroglossus – Andinodontis – Andinorites – Andinotrechus – Andrewesia – Andrewesinulus – Andrewesius – Androzelma – Anentmetus – Angionychus – Anheterus – Aniara – Anillidius – Anillinus – Anillodes – Anillopsidius – Anillopsis – Anillotarsus – Anillus – Anisocnemus – Anisodactylus – Anisostichus – Anomophaenus – Anomostomus – Anomotachys – Anomotariella – Anomotarus – Anoncopeucus – Anophthalmus – Anoplogenius – Antarctiola – Antarctonomus – Antennaria – Anthia – Anthracus – Antilliscaris – Antimerina – Antipionycha – Antireicheia – Antoinella – Aparupa – Apatrobus – Apenes – Aphaenopidius – Aphaenops – Aphaenopsis – Aphaenotyphlus – Aplothorax – Apocimmerites – Apoduvalius – Apophylon – Apoplotrechus – Aporesthus – Aporesthus – Apotomus – Apristomimus – Apristus – Apsaustodon – Apsidocnemus – Apterodromites – Apteroessa – Aptinoderus – Aptinus – Aquilex – Arame – Archaeocindis – Archagonum – Archastes – Archicolliuris – Archicolpodes – Archidela – Archileistobrius – Archipatrobus – Arctaphaenops – Ardistomis – Ardistomopsis – Argentinatachoides – Argiloborus – Argutoridius – Arhytinus – Aristochroa – Aristochroodes – Aristolebia – Aristopus – Arsinoe – Arthropterus – Asakalaphium – Asaphidion – Asioreicheia – Asios – Asklepia – Aspasiola – Aspidoglossa – Assadecma – Astastus – Athrostictus – Atlantomasoreus – Atongolium – Atractonotus – Atranopsis – Atranus – Atrotus – Auchmerus – Aulacolius – Aulacopodus – Australomasoreus – Australovelinda – Austranillus – Austroglyptolenus – Austrotrechus – Awatrechus – Axinidium – Axinopalpus – Axinotoma – Axonya – Aztecarpalus
B

Caecidium – Caecocaelus – Caecoparvus – Caelostomus – Cainogenion – Calathidius – Calathocosmus – Calathosoma – Calathus – Caledonica – Caledonomorpha – Caledyschirius – Calleida – Calleidomorpha – Callidadelpha – Callidagonum – Callistomimus – Callistus – Callytron – Calodromius – Calomera – Calophaena – Calophaenoidea – Calosoma – Calybe – Calybe – Calyptoglossa – Calyptogonia – Cameroniola – Camptidius – Camptodontus – Camptogenys – Camptoscelis – Camptotoma – Canarobius – Capabatus – Carabidomemnus – Carabus – Carayonites – Carbonellia – Cardiaderus – Cardiomera – Carenochyrus – Carenum – Carterus – Casaleius – Cascellius – Castelnaudia – Catacolpodes – Catadromus – Catalanodytes – Catapiesis – Catascopellus – Catascopus – Cathaiaphaenops – Caucasaphaenops – Caucasorites – Cavicarabus – Cedrorum – Celanida – Celioinkosa – Cenogmus – Cenothyla – Cephalosdrophus – Cephalostichus – Cephalota – Cerabilia – Cerapterus – Ceratoderus – Ceroglossus – Cerotalis – Chaetagonum – Chaetobroscus – Chaetocrepis – Chaetodactyla – Chaetodera – Chaetoduvalius – Chaetogenys – Chaetomargoreicheia – Chaetopatrobus – Chaetosaurus – Chaetotrechiama – Chalcochrous – Chaltenia – Chaudoirella – Chaudoirina – Cheilonycha – Cheiloxya – Chiapadytes – Chilotomus – Chinapenetretus – Chlaeminus – Chlaenius – Chydaeus – Chylnus – Cicindela – Cicindelites – Cicindis – Cillenus – Cimmerites – Cimmeritodes – Cinctagonum – Cintaroa – Cistelagonum – Clarencia – Climax – Clivina – Clivinarchus – Clivinopsis – Cnemalobus – Cnides – Coarazuphium – Colasidia – Coleolissus – Colfax – Collagonum – Colliuris – Collyris – Colpodes – Colpodichius – Colpoides – Colpomimus – Colposphodrus – Colpostoma – Columbitrechus – Conchitella – Conopterum – Cophosomorpha – Coptia – Coptocarpus – Coptodera – Coptoglossus – Coptolobus – Coptoptera – Coptopterella – Coptopterina – Corcyranillus – Coreoblemus – Corintascaris – Corsyra – Coryza – Cosmodiscus – Cosnania – Costitachys – Cothresia – Crasodactylus – Craspedonotus – Craspedophorus – Crassacantha – Crassagena – Cratacanthus – Cratocerus – Cratoferonia – Cratogaster – Cratognathus – Cratohaerea – Cratosoma – Creagris – Crenulostrigus – Creobius – Crepidogaster – Crepidogastrillus – Crepidogastrinus – Crepidolomus – Crepidonellus – Crepidopterus – Crepidozaena – Cribrodyschirius – Criniventer – Croatotrechus – Cryptobatis – Cryptocephalomorpha – Cryptocharidius – Cryptocolliuris – Cryptomma – Cryptorites – Cryptoscaphus – Cteatus – Ctenodactyla – Ctenognathus – Ctenostoma – Cuneipectus – Cyanotarus – Cychropsis – Cychrus – Cycloloba – Cyclosomus – Cyclotrachelus – Cylindera – Cylindrocranius – Cylindronotum – Cylindropectus – Cymbionotum – Cymenopterus – Cymindis – Cymindoidea – Cymindoides – Cynthidia – Cyphocoleus – Cypholoba – Cyphotrechodes – Cyptomicrus – Cyrtolaus – Cyrtomoscelis – Cyrtopilus
D

Dabatrechus – Dactyleurys – Dactylinius – Daer – Daiconotrechus – Dailodontus – Dalatagonum – Dalmataphaenops – Dalmatoreicheia – Dalyat – Daptus – Darlingtonea – Darlingtonica – Darodilia – Dasiosoma – Decogmus – Deipyrodes – Delarouzee – Deliaesianum – Delinius – Deltocolpodes – Deltomerodes – Deltomerus – Demetrias – Demetrida – Demetriola – Dendragonum – Dendromigadops – Dercylinus – Dercylus – Derocrania – Derostichus – Deuveotrechus – Dhanya – Diacanthostylus – Diacheila – Diachipteryx – Diachromus – Diamella – Diastrophella – Dicaelindus – Dicaelus – Diceromerus – Dicheirotrichus – Dicheirus – Dicranoglossus – Dicranoncus – Dicraspeda – Dicrochile – Dicrodontus – Dicropterus – Diglymma – Dilatotarsa – Dilonchus – Dimorphopatrobus – Dimorphoreicheia – Dinocolpodes – Dinopelma – Dinoscaris – Diodercarus – Dioryche – Diplochaetus – Diplocheila – Diploharpus – Diplous – Dirotus – Dischissus – Discoderus – Discoptera – Disphericus – Distichus – Distipsidera – Distrigidius – Ditomus – Dixus – Do – Dobodura – Doderotrechus – Dolichocolpodes – Dolichoctis – Dolichodes – Dolichus – Doliodactyla – Dongodytes – Dontolobus – Dregus – Dromica – Dromicoida – Dromidea – Dromiops – Dromistomus – Dromius – Dromoceryx – Dromochorus – Drymatus – Drymonaxus – Drypta – Duvalioblemus – Duvaliomimus – Duvaliopsis – Duvalius – Dyscaris – Dyscherinus – Dyscherus – Dyschiridium – Dyschiriodes – Dyschirius – Dyscolus – Dystrichothorax
E

Ebertius – Eccoptomenus – Eccoptoptera – Ecnomolaus – Ectenognathus – Ectinothorax – Egadyla – Elaphropus – Elaphrotites – Elaphrus – Elgonotyphlus – Ellipsoptera – Elliptoleus – Enantiola – Enceladus – Endynomena – Enoicus – Entomoantyx – Eobroscus – Eocarterus – Eocnides – Eohomopterus – Eopaussus – Eosphodrus – Eotrechodes – Epaphiopsis – Epelyx – Epicolpodes – Epigraphodes – Epigraphus – Epikastea – Epilectus – Epimicodema – Epomis – Erebotrechus – Erectocolliuris – Eremolestes – Eremosphodrus – Erephognathus – Eriophonus – Eripus – Erwiniana – Eucaerus – Eucaerus – Eucallia – Eucamaragnathus – Eucamptognathus – Eucarterus – Eucheila – Euchroa – Eucolliuris – Eudalia – Eudromus – Eujalmenus – Euleptus – Eumara – Eunostus – Eunota – Euphorticus – Euphorticus – Euplatyrhopalus – Euplynes – Euproctinus – Euprosopus – Euryabax – Euryaptus – Euryarthron – Eurycoleus – Eurydera – Euryderus – Eurygnathus – Eurylychnus – Eurymorpha – Euryscaphus – Eurystomis – Euschizomerus – Eustra – Euthenarus – Eutogeneius – Eutrechopsis – Eutrechus – Euzona – Evolenes
F

Fangshania – Falsodromius – Feostoma – Feroniascus – Feroniola – Feronista – Filicerozaena – Forcipator – Formosiella – Fortagonum – Foveocrotaphus
G

Gaioxenus – Galerita – Galiciotyphlotes – Gallerucidia – Gastragonum – Gastrellarius – Gastrogmus – Geballusa – Gehringia – Geobaenus – Geobius – Geocharidius – Geocharis – Geodromus – Geoffreyella – Geopinus – Geoscaptus – Geotrechus – Gestroania – Giachinoana – Gibbozaena – Gidda – Gigadema – Ginema – Giraffaphaenops – Glenopterus – Glycia – Glyphodactyla – Glyptogrus – Glyptolenoides – Glyptolenus – Glyptus – Gnaphon – Gnathaphanus – Gnathoxys – Gnopheroides – Goedetrechus – Gomerina – Goniocellus – Gonogenia – Gotoblemus – Gouleta – Gourlayia – Grandopronotalia – Graniger – Granulopaussus – Graphipterus – Grundmannius – Grundmannius – Guatemalteca – Guatemalteca – Gugheorites – Guineica – Guiodytes – Guizhaphaenops – Gulaphaenops – Gunvorita – Guyanemorpha – Gynandromorphus
H

Habragonum – Habrodera – Habroscelimorpha – Habutarus – Hakaharpalus – Halocoryza – Hannaphota – Haplaner – Haplanister – Haplobothynus – Haplocolpodes – Haploferonia – Haplogaster – Haplopeza – Haplotrachelus – Haptoderidius – Haptoderodes – Harpaglossus – Harpalinus – Harpalobrachys – Harpalodiodes – Harpalomimetes – Harpalomorphus – Harpalus – Harpathaumas – Harponixus – Hartonymus – Hebeicarabus – Helenaea – Helluapterus – Helluarchus – Helluo – Helluobrochus – Helluocolpodes – Helluodema – Helluodes – Helluomorpha – Helluomorphoides – Helluonidius – Helluopapua – Helluosoma – Hemiaulax – Hemiplatynus – Hemitelestus – Henrotius – Henvelik – Heptodonta – Herculagonum – Heteracantha – Heteropaussus – Hexachaetus – Hexagonia – Hexaplatarthrus – Hikosanoagonum – Hiletus – Himalaphaenops – Himalotrechodes – Himalopenetretus – Himalosphodrus – Hippoloetis – Holaxinidium – Holcaspis – Holcocoleus – Holcoderus – Holconotus – Hololeius – Holoponerus – Holoprizus – Holosoma – Homaloderodes – Homalomorpha – Homalonesiota – Homethes – Homopterus – Honduranillus – Hoplizomenus – Hoplolenus – Hormacrus – Horniulus – Horologion – Hyboptera – Hybopteroides – Hybothecus – Hydraphaenops – Hygranillus – Hylopaussus – Hylotorus – Hypaetha – Hyperectenus – Hyperion – Hyphaereon – Hypharpax – Hypotyphlus – Hystrichopus – Hystricosphodrus
I

Iason – Iberanillus – Iberodytes – Iberotrechus – Idacarabus – Idiagonum – Idiastes – Idiocolpodes – Idiomelas – Idiomorphus – Ifridytes – Iga – Ildobates – Illaphanus – Incagonum – Incatrechus – Indiophonus – Infernophilus – Inflatozaena – Inkosa – Inotrechus – Inpa – Iresia – Iridagonum – Ischnagonum – Ishidatrechus – Ishikawatrechus – Italaphaenops – Itamus
J

Ja – Jalzicaphaenops – Jansenia – Jeannelius – Jiangxiaphaenops – Jocqueius – Jujiroa – Junaphaenops – Junnanotrechus
K

Kaffovatus – Kalchdigor – Kar – Kashmirobroscus – Kaszabellus – Kearophus – Kenodactylus – Kenyacus – Kenyoreicheia – Kettlotrechus – Kiwiharpalus – Kiwitachys – Kiwitrechus – Klapperichella – Klepteromimus – Kosswigia – Kuceraianum – Kultianella – Kupeharpalus – Kupetrechus – Kurasawatrechus – Kuschelinus – Kusumia
L

Labocephalus – Laccocenus – Laccopterum – Lachenus – Lachnocrepis – Lachnoderma – Lachnolebia – Lachnophorus – Lachnophorus – Lachnothorax – Lacordairia – Laemostenus – Laevagonum – Langea – Laoreicheia – Laosaphaenops – Lasiocera – Lassalleianum – Lebia – Lebidema – Lebidia – Lebioderus – Lebiomorphica – Lebistina – Lebistinida – Lecanomerus – Ledouxius – Leiradira – Leistus – Leleupanillus – Leleupaussus – Leleupidia – Leleuporella – Leleuporites – Lelis – Lepcha – Leptagonum – Leptocolpodes – Leptognatha – Leptosarcus – Leptotrachelon – Leptotrachelus – Lessinodytes – Lesticus – Lestignathus – Letouzeya – Leuropus – Liagonum – Liamegalonychus – Libotrechus – Lichnasthenus – Licinopsis – Licinus – Limnastis – Lindrothius – Liochirus – Liocolpodes – Liodaptus – Lioholus – Lionedya – Lionychus – Liopasa – Lioptera – Liotachys – Lipostratia – Lissopogonus – Lissopterus – Litarthrum – Lithagonum – Lobatodes – Lobobrachus – Lobocolpodes – Lobodontidius – Lobodontulus – Lobodontus – Lonchosternus – Lophidius – Lophocoryza – Lophoglossus – Lophyra – Lorestania – Loricera – Lorostema – Lovricia – Loxandrus – Loxocrepis – Loxodactylus – Loxogenius – Loxomerus – Loxoncus – Lucicolpodes – Luoxiaotrechus – Luperca – Luraphaenops – Luyatrechus – Luzonotrechus
M

Macfarlandia – Machozetus – Macrocheilus – Macromorphus – Macroprotus – Maculagonum – Madapelmus – Madascaris – Madecassellus – Madecassina – Mamboicus – Manautea – Mantica – Manticora – Manumorpha – Maoriharpalus – Maoripamborus – Maoritrechus – Marsyas – Martyr – Mascarenhia – Masoreus – Mastax – Masuzoa – Masuzonoblemus – Matabele – Mateuellus – Mayaphaenops – Mecodema – Mecyclothorax – Mecynognathus – Mecynoscaris – Megacephala – Megadromus – Megalebia – Megalomma – Megalonychus – Megaloodes – Megalopaussus – Megamorio – Meganophthalmus – Meladroma – Melaenus – Melanchiton – Melanchrous – Melanospilus – Meleagros – Meleagros – Melisodera – Menarus – Menigius – Meonis – Meonochilus – Meotachys – Merizodus – Merizomena – Meroctenus – Meropalpus – Mesarthropterus – Mesocolpodes – Mesolestes – Mesorabus – Mesus – Metabacetus – Metablus – Metacolpodes – Metadromius – Metaglymma – Metallanchista – Metallica – Metascopus – Metaxellus – Metaxidius – Metaxinidium – Metaxymorphus – Metaxys – Metazuphium – Metius – Metoncidus – Metriocheila – Metrius – Mexanillus – Mexaphaenops – Mexisphodrus – Mexitrechus – Meileyingia – Micratopus – Micrixys – Microcephalus – Microcheila – Microcosmodes – Microdaccus – Microderes – Microdipnites – Microdipnodes – Microdipnus – Microferonia – Microlestes – Microlestodes – Micromentignatha – Microodes – Microthylax – Microtyphlus – Microzaena – Microzargus – Migadopidius – Migadopiella – Migadops – Miltodes – Mimanillus – Mimocolliuris – Mimodromius – Mimophilorhizus – Mimotrechus – Mimovelindopsis – Mimozaena – Minimaphaenops – Minipenetretus – Minosaphaenops – Minuphloeus – Minuthodes – Minypatrobus – Miolebidia – Mioptachys – Miquihuana – Mirachlaenius – Mireius – Miscelus – Mischocephalus – Miscodera – Mizotrechus – Mnuphorus – Mochtheroides – Mochtherus – Moirainpa – Molopidius – Molopinus – Molops – Molopsida – Monnea – Monocentrum – Monodryxus – Monolobus – Montagonum – Morimotoidius – Moriodema – Moriomorpha – Morion – Morionidius – Moriosomus – Mormolyce – Mormolycina – Morphaeus – Morphodaclyla – Mouhotia – Mouhotia – Myas – Myriochila – Myrmecodemus – Mystroceridius – Mystropomus
N

Nannoryctes – Nannotrechus – Nanodiodes – Naviauxella – Naxipenetretus – Neanops – Neaphaenops – Nebria – Nebriagonum – Nebrioferonia – Nebriosoma – Negrea – Negreum – Nelidus – Nelsonites – Nemaglossa – Nemotarsus – Neoaulacoryssus – Neobatenus – Neoblemus – Neocarenum – Neochila – Neochryopus – Neocicindela – Neocollyris – Neocolpodes – Neocoptodera – Neodendragonum – Neodiachipteryx – Neodyschirius – Neoeudalia – Neoferonia – Neoglyptus – Neohelluo – Neohyparpalus – Neolaphyra – Neoleleupidia – Neolovricia – Neomegalonychus – Neonomius – Neoodes – Neoorthogonius – Neophygas – Neoscaphus – Neotalus – Neothanes – Neotrechus – Nepalorthogonius – Nesamblyops – Nesarpalus – Nesiocolpodes – Nesiotrechus – Nesites – Netrodera – Nickerlea – Nipponagonum – Nipponaphaenops – Nipponoharpalus – Nippononebria – Nipponosynuchus – Nirmala – Nomius – Nornalupia – Notabax – Notagonum – Nothanillus – Nothobroscus – Nothocascellius – Nothodaptus – Notiobia – Notiokasis – Notiophilus – Notocolpodes – Notolestus – Notonomus – Notoplatynus – Notopseudomorpha – Notospeophonus – Notospira – Nototrechus – Nototylus – Novillidius – Nurus – Nycteis – Nyctosyles
O

Oarotrechus – Obadius – Oberthueria – Ochtozetus – Ochyropus – Ocys – Odacantha – Odontocarus – Odontocheila – Odontomasoreus – Oechalius – Oecornis – Oedesis – Oesyperus – Ogmophora – Oilea – Olegius – Olisthopus – Omalodera – Omestes – Omobrus – Omophagus – Omophron – Omostropus – Omphra – Omphreoides – Omphreus – Omus – Onota – Onotokiba – Onycholabis – Onypterygia – Oodes – Oodinkosa – Oodinotrechus – Oodinus – Ooidius – Oopterus – Oosoma – Ophonichius – Ophoniscus – Ophonus – Opilidia – Opisthencentrus – Opisthius – Oregus – Oreodicastes – Oribazus – Orictites – Orientoreicheia – Orionella – Oroblemites – Oroblemus – Orophicus – Orotrechus – Orthobasis – Orthocerodus – Orthocindela – Orthogenium – Orthoglymma – Orthogonius – Orthomus – Orthotrichus – Orthotyphlus – Orzolina – Oscadytes – Otoglossa – Oxoides – Oxycentrus – Oxycheila – Oxycrepis – Oxydrepanus – Oxyglychus – Oxygnathopsis – Oxygnathus – Oxygonia – Oxygoniola – Oxygonium – Oxylobus – Oxyodontus – Oxypselaphus – Oxytrechus – Ozaena
P

Pachybatenus – Pachycaecus – Pachycallida – Pachycarus – Pachydesus – Pachyferonia – Pachylebia – Pachylebioides – Pachymorphus – Pachyodontus – Pachyroxochus – Pachyteles – Pachythecus – Pachytrachelus – Pachytrechodes – Pamborus – Panagaeus – Panagrius – Pangus – Paniestichus – Parabaris – Parabroscus – Parachlaenius – Paracimmerites – Paracolpodes – Paracoryza – Paradromius – Paradyscherus – Paraeutrichopus – Paraglycia – Paragonotrechus – Paraleleupidia – Paraliagonum – Paralophidius – Paralovricia – Paramecus – Paramegalonychus – Paranchodemus – Paranchus – Paranillopsis – Paranillus – Paranurus – Parapenetretus – Paraphaenops – Paraphysodeutera – Parapionycha – Parascopodes – Parasiopelus – Paratachys – Parathlibops – Paratrechiotes – Paratrechodes – Paratrechus – Paratrichothorax – Paraxinidium – Parazuphium – Paregaploa – Paregraphus – Parena – Pareuryaptus – Pareurydera – Parhypates – Paridiagonum – Parophonus – Paropisthius – Parorthomus – Parvocaecus – Pasimachus – Passalidius – Patanitretus – Patrobus – Paulianites – Paulianolebia – Paulianoscirtus – Paussomorphus – Paussotropus – Paussus – Pawgammm – Pectinitarsus – Pediomorphus – Pedius – Pelecium – Peliocypas – Pelmatellus – Pelocharis – Pelodiaetodes – Pelodiaetus – Pelonomites – Pelophila – Penetretus – Pentacomia – Pentagonica – Pentaplatarthrus – Penthophonus – Penthus – Pephrica – Percodermus – Percolaus – Percolestus – Percosoma – Percus – Pericalus – Pericompsus – Peridexia – Perigona – Perileptus – Peripristus – Perissostomus – Perochnoristhus – Peronomerus – Perucharidius – Petrimagnia – Peyrieraselus – Phacocerus – Phaenaulax – Phanagnathus – Pheggomisetes – Pheropsophus – Philipis – Philophlaeus – Philorhizus – Philoscaphus – Phloeoxena – Phobophorus – Pholeodytes – Phorticosomus – Phrypeus – Phyllodroma – Phyrometus – Physea – Physeomorpha – Physocrotaphus – Physodera – Physodeutera – Physolaesthus – Physomerus – Picnochile – Piesmus – Piezia – Pilades – Pionycha – Pioprosopus – Piosoma – Plagiopyga – Plagiotelum – Planetes – Platidiolus – Platyagonum – Platycerozaena – Platychile – Platycoelus – Platyderus – Platyhelluo – Platylytron – Platymetopsis – Platymetopus – Platynodes – Platynus – Platypatrobus – Platyrhopalopsis – Platyrhopalus – Platysmodes – Platysphyrus – Platytarus – Platyxythrius – Plaumannium – Pleuracanthus – Plicagonum – Plocamostethus – Plochionus – Poecilus – Pogonistes – Pogonodaptus – Pogonoglossus – Pogonopsis – Pogonoschema – Pogonostoma – Pogonus – Polistichus – Pollicobius – Polpochila – Polyaulacus – Polychaetus – Polydamasium – Polyderis – Polyrhanis – Pometon – Ponapagonum – Pontodytes – Pontonoa – Porocara – Porocimmerites – Porotachys – Potamagonum – †Praeanchodemus – Praepristus – Prakasha – Prepusa – Prioniomus – Prionognathus – Pristacrus – Pristosia – Probstia – Procletodema – Procletus – Prodiplous – Proditomus – Prodyscherodes – Prodyscherus – Progonochaetus – Promecognathus – Promecoptera – Promegalonychus – Pronyssa – Pronyssiformia – Proozaena – Prophenorites – Propionycha – Prosopogmus – Prosphodrus – Prostalomus – Prothyma – Prothymidia – Protocolliuris – Protocollyris – Protocolpodes – Protopaussus – Protopidius – Psammocoryza – Psammodromius – Psecadius – Psegmatopterus – Pseudabacetus – Pseudabarys – Pseudagonica – Pseudamara – Pseudamblytelus – Pseudanchomenus – Pseudanillus – Pseudanisotarsus – Pseudanophthalmus – Pseudaphaenops – Pseudaptinus – Pseudaristus – Pseudobatenus – Pseudocalleida – Pseudoceneus – Pseudoclivina – Pseudocnides – Pseudodiachipteryx – Pseudognathaphanus – Pseudohyparpalus – Pseudomasoreus – Pseudomegalonychus – Pseudomenarus – Pseudometabletus – Pseudomigadops – Pseudomonnea – Pseudomorpha – Pseudomyas – Pseudopachylebia – Pseudophorticus – Pseudophorticus – Pseudoplatyderus – Pseudorhysopus – Pseudoselenophorus – Pseudosinurus – Pseudosphaerodes – Pseudotaphoxenus – Pseudotoglossa – Pseudotrechisibus – Pseudotrechus – Pseudoxycheila – Pseudozaena – Psilidius – Psychristus – Psydrus – Pterocyrtus – Pterogmus – Pterorhopalus – Pterostichus – Pterostillichus – Puertrechus – Punctoduvalius – Putzeysius – Pylartesius – Pyramoides
Q

Qianaphaenops – Qiangopatrobus – Qianotrechus – Quammenis – Quammenis – Quasipenetretus – Queinnectrechus
R

Rakantrechus – Raphetis – Rathymus – Rawlinsius – Reflexisphodrus – Reicheadella – Reicheia – Reicheidius – Reicheiodes – Renneria – Rhabdotus – Rhabidius – Rhadine – Rhaebolestes – Rhagadillius – Rhegmatobius – Rhopalomelus – Rhopalostyla – Rhysocara – Rhysopleura – Rhysopus – Rhytidognathus – Rhytidophaena – Rhytiferonia – Rhytisternus – Ripogenites – Rivacindela – Robustopenetretus – Ronhuberia – Rossjoycea – Rugiluclivina – Rupa – Ryugadous
S

Sakagutia – Salcedia – Salpingophora – Samiriamorpha – Sardaphaenops – Sarothrocrepis – Sarticus – Sbordoniella – Scalidion – Scaphinotus – Scapterus – Scaraphites – Scarites – Scaurotrechodes – Schidonychus – Schizogenius – Schuelea – Scopodes – Scotodipnus – Scotoplanetes – Scototrechus – Scybalicus – Secatophus – Selenochilus – Selenophorus – Selenotichnus – Selina – Selina – Semiardistomis – Semiclivina – Sericoda – Serranillus – Serratozaena – Serrimargo – Setalidius – Setalimorphus – Setalis – Setitachys – Setodyschirius – Setolebia – Sfitakantha – Shenaphaenops – Shilinotrechus – Shuaphaenops – Siagona – Siagonyx – Sichuanotrechus – Sidublemus – Simodontus – Simous – Sinaphaenops – Sinechostictus – Sinesetosa – Singiliomimus – Singilis – Sinobrosculus – Sinocolpodes – Sinometrius – Sinotrechiama – Sinotroglodytes – Sinozolus – Sinurus – Siopelus – Sitaphe – Skoeda – Skorlagad – Skouedhirraad – Sloaneana – Sloanella – Smeringocera – Socotrana – Soesilascarites – Sofota – Solenogenys – Somalodromius – Somoplatodes – Somoplatus – Somotrichus – Sophroferonia – Sparostes – Speagonum – Spelaeodytes – Spelaeovulcania – Speluncarius – Speocolpodes – Speokokosia – Speomolops – Speotarus – Speothalpius – Speotrechus – Speozuphium – Sphaeroderus – Sphaerodes – Sphaerodinus – Sphaerostylus – Sphallomorpha – Sphodropsis – Sphodroschema – Sphodrosomus – Sphodrus – Sporades – Steganomma – Stegazopteryx – Stenidia – Stenocheila – Stenocnemus – Stenocosmia – Stenocrepis – Stenognathus – Stenolepta – Stenolophidius – Stenolophus – Stenomorphus – Stenoodes – Stenotelus – Stereocerus – Stereostoma – Steropanus – Stevensius – Sthenocranion – Stichonotus – Stigmaphorus – Stilboma – Stolonis – Stomis – Stomonaxellus – Storthodontus – Straneoa – Straneoites – Straneostichus – Straneotia – Stratiotes – Stricklandiana – Stricteripus – Strigia – Strigomerodes – Strigomerus – Stuhlmannium – Stygiotrechus – Stylulus – Styphlodromus – Styphlomerus – Styracoderus – Subilsia – Sugimotoa – Sumlinia – Superbotrechus – Suzuka – Syleter – Syletor – Syllectus – Synteratus – Syntomus – Synuchidius – Synuchus – Syrdenoidius – Syrdenus – Systolocranius
T

Tachypeles – Tachys – Tachysbembix – Tachyta – Taenarthrus – Taicona – Taiwanotrechus – Takasagoagonum – Taniatrechus – Tantillus – Tanystoma – Taphoxenus – Tapinopterus – Tapinoscaris – Tarastethus – Taridius – Tarsagonum – Tasmanillus – Tasmanitachoides – Tasmanorites – Tasmanotrechus – Taurocimmerites – Tecnophilus – Tefflus – Teiresia – Temnostega – Teradaia – Teraphis – Teratotarsa – Teropha – Tetracha – Tetragonoderus – Tetraleucus – Teukrus – Thaioodes – Thalassobius – Thalassoduvalius – Thalassophilus – Thalassotrechus – Thaumastaphaenops – Theprisa – Therates – Thermoscelis – Thliboclivina – Thlibops – Thoasia – Thopeutica – Thryptocerus – Thyreochaetus – Thyreopterus – Thysanotus – Tibioscarites – Tienmutrechus – Tiferonia – Tilius – Tinoderus – Tinognathus – Titaresius – Tonkinoscaris – Torretassoa – Toshiaphaenops – Tostkar – Trachelocyphoides – Trachelocyphus – Trachysarus – Trechiama – Trechiamiotes – Trechicomimus – Trechiella – Trechimorphus – Trechinotus – Trechiotes – Trechisibus – Trechistus – Trechobembix – Trechoblemus – Trechodes – †Trechoides – Trechosia – Trechosiella – Trechus – Trephionus – Trephisa – Triaenogenius – Trichamblytelus – Trichaphaenops – Tricharnhemia – Trichidema – Trichillinus – Trichis – Trichisia – Trichocarenum – Trichocoptodera – Trichognathus – Trichopiezia – Trichopselaphus – Trichopsida – Trichosternus – Trichotichnus – Trichoxycentrus – Tricondyla – Trigonognatha – Trigonothops – Trigonotoma – Trilophidius – Trilophus – Triplosarus – Trirammatus – Trogloagonum – Troglocimmerites – Tropicoritus – Tropidotrechus – Tropopterus – Trymosternus – Tschitscherinellus – Tuiharpalus – Turkanillus – Tuxtlamorpha – Typhlocharis – Typhlonesiotes – Typhloreicheia – Typhloscaris – Typhlotrechus – Typsiharpalus
U

Uenanthracus – Uenotrechus – Ushijimaella
V

Vachinius – Valeriaaschero – Vata – Velebitaphaenops – Velinda – Velindastus – Velindomimus – Velindopsis – Veradia – Vianasia – Vietotrechus – Violagonum – Viridagonum – Vitagonum – Vulcanophilus
W

Wahlbergiana – Waiputrechus – Waltherhornia – Wate – Whiteheadiana – Winklerites
X

Xanthomelina – Xenaroswelliana – Xenion – Xenitenopsis – Xenitenus – Xenodochus – Xenodromius – Xenophonus – Xenotrechus – Xestagonum – Xestonotus – Xestopus – Xystosomus
Y

Yamautidius – Yasunimorpha – Yukihikous
Z

Zabrus – Zacotus – Zargus – Zariquieya – Zeanillus – Zecicindela – Zeodera – Zeopoecilus – Zhijinaphaenops – Zoianillus – Zolotarevskyella – Zuphium
Extinct and questionable genera

?Amphitasus – ?Diaphoromerus – ?Oreonebria – ?Orientonebria – ?Stigmaphorus – †Aethocarabus – †Cordorabus – †Cretorabus – †Lithorabus – †Nebrorabus – †Obesofemoria – †Ovrabites – †Oxycheilopsis – †Palaeoaxinidium – †Penecupes – †Praeanchodemus – †Protorabus – †Sinocarabus
Name

Carabidae Latreille, 1802: 80

Type genus: Carabus Linnaeus, 1758 [Stem: Carab-]

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. [see p. 80]

Additional references

Ariza, G.M., Jácome, J., Esquivel, H.E. & Kotze, D.J. 2021. Early successional dynamics of ground beetles (Coleoptera, Carabidae) in the tropical dry forest ecosystem in Colombia. Pp 877–906 In Spence, J.R., Casale, A., Assmann, T., Liebherr, J.K & Penev, L. (eds.). Systematic Zoology and Biodiversity Science: A tribute to Terry Erwin (1940-2020). ZooKeys 1044: 1–1042. Reference page. DOI: 10.3897/zookeys.1044.59475 Open access Reference page.
Arndt, E., Beutel, R.G. & Will, K.W. 2005. 7.8. Carabidae Latreille, 1802. Pp. 119–146 in Kristensen, N.P. & Beutel, R.G. (series eds.) Handbook of Zoology, Vol. IV Arthropoda: Insecta. Part 38: Coleoptera, Vol. 1: Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga (partim) (vol. eds. Beutel, R.G. and Leschen, R.A.B.). Walter De Gruyter, Berlin, New York. PDF
Azadbakhsh, S. & Nozari, J. 2015. Checklist of the Iranian Ground Beetles (Coleoptera; Carabidae). Zootaxa 4024(1): 1–108. DOI: 10.11646/zootaxa.4024.1.1. Preview (PDF) ISBN 978-1-77557-801-7 (paperback); ISBN 978-1-77557-802-4 (Online edition) Reference page.
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.
Faisal, M.; Singh, S. 2014: Carabid (Coleoptera) type collection at National Forest Insect Collection (NFIC), Forest Research Institute, Dehradun (India). Zootaxa 3786(3): 331–358. DOI: 10.11646/zootaxa.3786.3.5 Reference page.
Fuller, L.; Johns, P.M.; Ewers, R.M. 2013: Assessment of protected area coverage of threatened ground beetles (Coleoptera: Carabidae): a new analysis for New Zealand. New Zealand journal of ecology, 37(2): 184–192. abstract and full article (PDF) Reference page.
Giachino, P.M. 2009. Family Carabidae. Pp. 37–45 in: Gerlach, J. (ed.) The Coleoptera of the Seychelles Islands. Pensoft Publishers, Sofia-Moscow. ISBN 9789546424983
Guéorguiev, B.V. & Sciaky, R. 2015. A new genus and two new species of Pterostichini from China, with “sphodrine-like” parameres (Coleoptera, Carabidae). Deutsche Entomologische Zeitschrift 62(2): 225–237. DOI: 10.3897/dez.62.5493 Reference page.
Hammond, H.E.J., García-Tejero, S., Pohl, G.H., Langor, D.W. & Spence, J.R. 2021. Spatial and temporal variation of epigaeic beetle assemblages (Coleoptera, Carabidae, Staphylinidae) in aspen-dominated mixedwood forests across north-central Alberta. Pp 951–991 In Spence, J.R., Casale, A., Assmann, T., Liebherr, J.K & Penev, L. (eds.). Systematic Zoology and Biodiversity Science: A tribute to Terry Erwin (1940-2020). ZooKeys 1044: 1–1042. Reference page. DOI: 10.3897/zookeys.1044.65776 Open access Reference page.
Holland, J.M. (ed.) 2002. The agroecology of carabid beetles. Intercept: Andover, UK. ISBN 1-898298-76-9
Hristovski, S. & Guéorguiev, B. 2015. Annotated catalogue of the carabid beetles of the Republic of Macedonia (Coleoptera: Carabidae). Zootaxa 4002(1): 1–190. DOI: 10.11646/zootaxa.4002.1.1. (PDF) ISBN 978-1-77557-767-6 (paperback); ISBN 978-1-77557-768-3 (Online edition) Reference page.
Kostova, R. & Guéorguiev, B. 2016. The ground beetles (Coleoptera: Carabidae) of the Strandzha Mountain and adjacent coastal territories (Bulgaria and Turkey). Biodiversity Data Journal 4: e8135. DOI: 10.3897/BDJ.4.e8135. Reference page.
Kryzhanovskij, O.L. et al. 1995. A Checklist of the Ground-Beetles of Russia and Adjacent Lands. Pensoft Series Faunistica (3) ISBN 954 642 004 2 PDF Google books
Lohaj, R. & Delić, T. 2019. Playing hard to get: two new species of subterranean Trechini beetles (Coleoptera, Carabidae, Trechinae) from the Dinaric Karst. Deutsche Entomologische Zeitschrift, 66(1): 1–15. DOI: 10.3897/dez.66.31754 Reference page.
Looney, C., Zack, R.S. & LaBonte, J.R. 2014. Ground beetles (Coleoptera, Carabidae) of the Hanford Nuclear Site in south-central Washington State. ZooKeys 396: 13–42. DOI: 10.3897/zookeys.396.6936 Reference page.
Lorenz, W. 2005. A systematic list of extant ground beetles of the World (Coleoptera "Geadephaga": Trachypachidae and Carabidae, incl. Paussinae, Cicindelinae, Rhysodinae). Second edition. (Tutzing) ISBN 3-933 896-06-1 First page of page by pagePDF's
Maddison, D.R., Baker, M.D. & Ober, K.A. 1999. Phylogeny of carabid beetles as inferred from 18S ribosomal DNA (Coleoptera: Carabidae). Systematic entomology 24: 103–138. DOI: 10.1046/j.1365-3113.1999.00088.x PDF
Madge, R.B. 1975. The type-species of Bonelli's genera of Carabidae (Coleoptera). Quaestiones entomologicae 11: 579–586.
Moravec, J. & Vybíra, J. 2018. New or rare Madagascar tiger beetles—17. Description of Pogonostoma (Microstenocera) noheli sp. nov. and redescription of P. (M.) flavomaculatum W. Horn (Coleoptera: Cicindelidae). Zootaxa 4388(1): 76–88. DOI: 10.11646/zootaxa.4388.1.5 Reference page.
Osawa, S., Su, Z.-H. & Inmura, Y. 2004. Molecular phylogeny and evolution of carabid ground beetles. Springer. ISBN 4-431-00487-4 Google books
Peterson, K.N.R., Browne, R.A. & Erwin, T.L. 2021. Carabid beetle (Coleoptera, Carabidae) richness, diversity, and community structure in the understory of temporarily flooded and non-flooded Amazonian forests of Ecuador. Pp 831–876 In Spence, J.R., Casale, A., Assmann, T., Liebherr, J.K & Penev, L. (eds.). Systematic Zoology and Biodiversity Science: A tribute to Terry Erwin (1940-2020). ZooKeys 1044: 1–1042. Reference page. DOI: 10.3897/zookeys.1044.62340 Open access Reference page.
Raupach, M.J., Hannig, K., Moriniere, J. & Hendrich, L. 2016. A DNA barcode library for ground beetles (Insecta, Coleoptera, Carabidae) of Germany: The genus Bembidion Latreille, 1802 and allied taxa. ZooKeys 592: 121-141. DOI: 10.3897/zookeys.592.8316. Reference page.
Schmidt, J. 2015. On the Eocene age of Limodromus Motschulsky, 1850, with description of L. hoffeinsorum sp. n. from Baltic Amber (Coleoptera: Carabidae: Platynini). Zootaxa 3974(4): 573–581. DOI: 10.11646/zootaxa.3974.4.8. Preview (PDF) Reference page.
Taglianti, A.V. 2009. An updated checklist of the ground beetles (Coleoptera: Carabidae) of Sardinia. Pp. 169–196 in: Cerretti, P., Mason, F., Minelli, A., Nardi, G. & Whitmore, D. (eds), Research on the terrestrial arthropods of Sardinia (Italy). Zootaxa 2318: 1–602. Abstract & excerpt (PDF). PDF
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Tsafack, N., Wang, X-P., Xie, Y-Z. & Fattorini, S. 2021. Niche overlap and species co-occurrence patterns in carabid communities of the northern Chinese steppes. Pp 929–949 In Spence, J.R., Casale, A., Assmann, T., Liebherr, J.K & Penev, L. (eds.). Systematic Zoology and Biodiversity Science: A tribute to Terry Erwin (1940-2020). ZooKeys 1044: 1–1042. Reference page. DOI: 10.3897/zookeys.1044.62478 Open access Reference page.
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Zumstein, P., Bruelheide, H., Fichtner, A., Schuldt, A., Staab, M., Härdtle, W., Zhou, H-Z. & Assmann, T. 2021. What shapes ground beetle assemblages in a tree species-rich subtropical forest? Pp 907–927 In Spence, J.R., Casale, A., Assmann, T., Liebherr, J.K & Penev, L. (eds.). Systematic Zoology and Biodiversity Science: A tribute to Terry Erwin (1940-2020). ZooKeys 1044: 1–1042. Reference page. DOI: 10.3897/zookeys.1044.63803 Open access Reference page.

Links

Trees of family Carabidae
Atlas of ground beetles (Carabidae) of Russia
Australian Faunal Directory
Carabidae in: Synopsis of the Described Coleoptera of the World
Carabidae in the Carabidae of the World database.
Maddison, David R. 2006. Carabidae. Ground beetles and tiger beetles. Version 11 April 2006 (under construction) in The Tree of Life Web Project
EoL

Vernacular names
беларуская: Жужалі
čeština: Střevlíkovití
dansk: Løbebiller
Deutsch: Laufkäfer
English: Ground beetle
suomi: Maakiitäjäiset
Nordfriisk: Skelebleeden
français: Carabes
עברית: רצניתיים
magyar: Futrinkafélék, futóbogárfélék
日本語: オサムシ科
한국어: 딱정벌레과
lietuvių: Žygiai
македонски: Тркачи
Nederlands: Loopkevers
polski: Biegaczowate (owady)
română: Carabidae
русский: Жужелицы
slovenčina: Bystruškovité
српски / srpski: Бауљари
svenska: Jordlöpare
українська: Жужелиці, туруни
中文: 步甲科

Ground beetles are a large, cosmopolitan family of beetles,[2] the Carabidae, with more than 40,000 species worldwide, around 2,000 of which are found in North America and 2,700 in Europe.[3] As of 2015, it is one of the 10 most species-rich animal families.
Description and ecology

Although their body shapes and coloring vary somewhat, most are shiny black or metallic and have ridged wing covers (elytra). The elytra are fused in some species, particularly the large Carabinae, rendering the beetles unable to fly. The species Mormolyce phyllodes is known as violin beetle due to their peculiarly shaped elytra. All carabids except the quite primitive flanged bombardier beetles (Paussinae) have a groove on their fore leg tibiae bearing a comb of hairs used for cleaning their antennae.[4]
A Brachinus species typical bombardier beetle (Brachininae: Brachinini) from North Carolina
Defensive secretions

Typical for the ancient beetle suborder Adephaga to which they belong, they have paired pygidial glands in the lower back of the abdomen. These are well developed in ground beetles, and produce noxious or even caustic secretions used to deter would-be predators. In some, commonly known as bombardier beetles, these secretions are mixed with volatile compounds and ejected by a small combustion, producing a loud popping sound and a cloud of hot and acrid gas that can injure small mammals, such as shrews, and is liable to kill invertebrate predators outright.

To humans, getting "bombed" by a bombardier beetle is a decidedly unpleasant experience. This ability has evolved independently twice, as it seems, in the flanged bombardier beetles (Paussinae), which are among the most ancient ground beetles, and in the typical bombardier beetles (Brachininae), which are part of a more "modern" lineage. The Anthiini, though, can mechanically squirt their defensive secretions for considerable distances and are able to aim with a startling degree of accuracy; in Afrikaans, they are known as oogpisters ("eye-pissers"). In one of the very few known cases of a vertebrate mimicking an arthropod, juvenile Heliobolus lugubris lizards are similar in color to the aposematic oogpister beetles, and move in a way that makes them look surprisingly similar to the insects at a casual glance.[5]

A folk story claims that Charles Darwin once found himself on the receiving end of a bombardier beetle's attack, based on a passage in his autobiography.[6][7] Darwin stated in a letter to Leonard Jenyns that a beetle had attacked him on that occasion, but he did not know what kind:

A Cychrus rostratus once squirted into my eye & gave me extreme pain; & I must tell you what happened to me on the banks of the Cam in my early entomological days; under a piece of bark I found two carabi (I forget which) & caught one in each hand, when lo & behold I saw a sacred Panagæus crux major; I could not bear to give up either of my Carabi, & to lose Panagæus was out of the question, so that in despair I gently seized one of the carabi between my teeth, when to my unspeakable disgust & pain the little inconsiderate beast squirted his acid down my throat & I lost both Carabi & Panagæus![8]

A Lophyra sp. tiger beetle from Tanzania
Ecology

Common habitats are under the bark of trees, under logs, or among rocks[2] or sand by the edge of ponds and rivers. Most species are carnivorous and actively hunt for any invertebrate prey they can overpower.[2] Some run swiftly to catch their prey; tiger beetles (Cicindelinae) can sustain speeds of 9 km/h (5.6 mph)[9] – in relation to their body length they are among the fastest land animals on Earth. Unlike most Carabidae, which are nocturnal, the tiger beetles are active diurnal hunters and often brightly coloured; they have large eyes and hunt by sight. Ground beetles of the species Promecognathus laevissimus are specialised predators of the cyanide millipede Harpaphe haydeniana, countering the hydrogen cyanide that makes these millipedes poisonous to most carnivores.
Relationship with humans

As predators of invertebrates, including many pests, most ground beetles are considered beneficial organisms. The caterpillar hunters (Calosoma) are famous for their habit of devouring prey in quantity, eagerly feeding on tussock moth (Lymantriidae) caterpillars, processionary caterpillars (Thaumetopoeidae) and woolly worms (Arctiidae), which, due to their urticating hairs, are avoided by most insectivores. Large numbers of the forest caterpillar hunter (C. sycophanta), native to Europe, were shipped to New England for biological control of the gypsy moth (Lymantria dispar) as early as 1905.

A few species are nuisance pests. Zabrus is one of the few herbivorous ground beetle genera, and on rare occasions Zabrus tenebrioides, for example, occurs abundantly enough to cause some damage to grain crops.[10] Large species, usually the Carabinae, can become a nuisance if present in large numbers, particularly during outdoor activities such as camping; they void their defensive secretions when threatened, and in hiding among provisions, their presence may spoil food. Since ground beetles are generally reluctant or even unable to fly, mechanically blocking their potential routes of entry is usually easy. The use of insecticides specifically for carabid intrusion may lead to unfortunate side effects, such as the release of their secretions, so it generally is not a good idea unless the same applications are intended to exclude ants, parasites or other crawling pests.
A crucifix ground beetle (Panagaeus cruxmajor) got Charles Darwin into trouble in 1828.

Especially in the 19th century and to a lesser extent today, their large size and conspicuous coloration, as well as the odd morphology of some (e.g. the Lebiini), made many ground beetles a popular object of collection and study for professional and amateur coleopterologists. High prices were paid for rare and exotic specimens, and in the early to mid-19th century, a veritable "beetle craze" occurred in England. As mentioned above, Charles Darwin was an ardent collector of beetles when he was about 20 years old, to the extent that he would rather scour the countryside for rare specimens with William Darwin Fox, John Stevens Henslow, and Henry Thompson than to study theology as his father wanted him to do. In his autobiography, he fondly recalled his experiences with Licinus and Panagaeus, and wrote:

No poet ever felt more delight at seeing his first poem published than I did at seeing in Stephen's Illustrations of British Insects the magic words, "captured by C. Darwin, Esq."[7]

Evolution and systematics

The Adephaga are documented since the end of the Permian, about 250 million years ago (Mya). Ground beetles evolved in the latter Triassic, having separated from their closest relatives by 200 Mya. The family diversified throughout the Jurassic, and the more advanced lineages, such as the Harpalinae, underwent a vigorous radiation starting in the Cretaceous. The closest living relatives of the ground beetles are the false ground beetles (Trachypachidae) and the wrinkled bark beetles (Rhysodidae). They are sometimes even included in the Carabidae as subfamilies or as tribes incertae sedis, but more preferably they are united with the ground beetles in the superfamily Caraboidea.

Much research has been done on elucidating the phylogeny of the ground beetles and adjusting systematics and taxonomy accordingly. While no completely firm consensus exists, a few points are generally accepted: The ground beetles seemingly consist of a number of more basal lineages and the extremely diverse Harpalinae, which contain over half the described species and into which several formerly independent families had to be subsumed.[11]
Subfamilies and selected genera
Main article: List of ground beetle genera

The taxonomy used here is based on the Catalogue of Palaearctic Coleoptera[12] and the Carabidae of the World Database.[13] Other classifications, while generally agreeing with the division into a basal radiation of more primitive lineages and the more advanced group informally called "Carabidae Conjunctae",[14] differ in details. For example, the system used by the Tree of Life Web Project makes little use of subfamilies, listing most tribes as incertae sedis as to subfamily.[15] Fauna Europaea, though, splits rather than lumps the Harpalinae, restricting them to what in the system used here is the tribe Harpalini.[16]

All the approaches mentioned above are legitimate, as they agree with the phylogeny as far as it has been resolved. The inclusive Harpalinae presented here are used for two reasons, one scientific and one practical – first, the majority of authors presently use this system, following the Catalogue of Palaearctic Coleoptera. Second, the MediaWiki markup cannot at present adequately represent the relationships of the ground beetle subgroups in detail if the restricted view of the Harpalinae is chosen.
Basal ground beetles
Carabus lateralis (Carabinae: Carabini)
Notiophilus substriatus (Nebriinae: Notiophilini)
Clivina fossor (Scaritinae: Clivinini)
Amblytelus sp. (Amblytelinae: Amblytelini)

Carabinae Latreille, 1802 – including Agoninae and Callistinae

Altagonum
Aplothorax (monotypic genus)
Calosoma - including Callisthenes
Carabus
Cychrus
Homothes
Fortagonum
Laemostenus
Notagonum

Cicindelinae – tiger beetles (roughly 2,600 species; sometimes included in the Carabidae)

Cicindinae

Elaphrinae Latreille, 1802

Blethisa
Diacheila Motschulsky, 1844
Elaphrus

Hiletinae

Loricerinae Bonelli, 1810

Loricera

Migadopinae

Nebriinae (includes Notiophilinae, often included in Carabinae)

Leistus
Nebria
Notiophilus
Pelophila Dejean, 1821
Nippononebria

Nototylinae

Omophroninae Bonelli, 1810 – round sand beetles

Omophron

Paussinae – ant nest beetles, flanged bombardier beetles

Promecognathinae

Scaritinae Bonelli, 1810 – pedunculate ground beetles

Clivina
Dyschirius Bonelli, 1810
Scarites

Siagoninae Bonelli, 1810
Carabidae Conjunctae
Dixus sphaerocephalus (Harpalinae: Harpalini)
Mormolyce phyllodes (Lebiinae: Lebiini)

Amblytelinae Sloane, 1898[Note 1]

Amblytelus

Apotominae

Brachininae – typical bombardier beetles

Broscinae

Dryptinae (sometimes in Harpalinae)

Gineminae (sometimes in Harpalinae)

Harpalinae (over 20,000 species[verification needed])

Lebiinae – including Cyclosominae, Mormolycinae, Odacanthinae, Perigoninae (sometimes in Harpalinae)

Licininae – including Chlaeniinae, Oodinae (sometimes in Harpalinae)

Melaeninae

Migadopinae

Orthogoniinae (sometimes in Harpalinae)

Panagaeinae (sometimes in Harpalinae)

Platyninae (sometimes in Harpalinae)

Pseudomorphinae (sometimes in Harpalinae)

Psydrinae

Pterostichinae – including Zabrinae (sometimes in Harpalinae)

Trechinae – including Bembidiinae, Patrobinae

Tribes incertae sedis

Amarotypini – Migadopinae or a distinct subfamily?
Gehringiini – Psydrinae, Trechinae, or a distinct subfamily?

Notes

Usually placed in the Psydrinae or Trechinae, they seem to represent a distinct lineage related to Brachininae and Harpalinae, and in the system used here would consequently be eligible for subfamily status.[17]

References

"Carabidae Taxa". Carabidae of the World. 2011. Archived from the original on 2012-11-02. Retrieved 6 Nov 2017.
Gomes Gonçalves, Marcos Paulo (December 2017). "Relationship Between Meteorological Conditions and Beetles in Mata de Cocal". Revista Brasileira de Meteorologia. 32 (4): 543–554. doi:10.1590/0102-7786324003. ISSN 0102-7786.
B. Kromp (1999). "Carabid beetles in sustainable agriculture: a review on pest control efficacy, cultivation aspects and enhancement". Agriculture, Ecosystems and Environment. 74 (1–3): 187–228. doi:10.1016/S0167-8809(99)00037-7.
John L. Capinera. Encyclopedia of Entomology. p. 1746.
R. B. Huey & E. R. Pianka (1977). "Natural selection for juvenile lizards mimicking noxious beetles". Science. 195 (4274): 201–203. doi:10.1126/science.831272. PMID 831272.
"Young Naturalist, A Lifelong Passion". Darwin. American Museum of Natural History. 2005. Archived from the original on December 21, 2010. Retrieved February 16, 2011.
Nora Barlow, ed. (1958). "Cambridge, 1828–1831". The Autobiography of Charles Darwin. pp. 56–71.
Charles Darwin (1846). "Letter to Leonard Jenyns, October 17, 1846". Archived from the original on September 22, 2009. Retrieved July 24, 2008.
"When tiger beetles chase prey at high speeds they go blind temporarily, Cornell entomologists learn - Cornell Chronicle". www.news.cornell.edu.
"Damage to winter cereals by Zabrus tenebrioides (Goeze) (Coleoptera: Carabidae)". CABI. Retrieved 2021-11-11.
Shōzō Ōsawa, Zhi-Hui Su & Yūki Inmura (2004). Molecular Phylogeny and Evolution of Carabid Ground Beetles. Springer. ISBN 4-431-00487-4.
I. Löbl & A. Smetana, ed. (2003). Catalogue of Palaearctic Coleoptera. I. Stenstrup, Denmark: Apollo Books.
"Trees of family Carabidae". Carabidae of the World Database. 2008. Archived from the original on September 19, 2008. Retrieved July 24, 2008.
David R. Maddison (January 1, 1995). "Carabidae Conjunctae". Tree of Life Web Project. Retrieved July 24, 2008.
David R. Maddison (April 11, 2006). "Carabidae. Ground beetles and tiger beetles". Tree of Life Web Project. Retrieved July 24, 2008.
"Harpalinae". Fauna Europaea. 2004. Retrieved February 16, 2011.

David R. Maddison (January 1, 1999). "Amblytelini". Tree of Life Web Project. Retrieved July 24, 2008.

Further reading

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E. Csiki (1946). Die Käferfauna des Karpaten-Beckens [The beetle fauna of the Carparthian basin] (in German). Budapest. pp. 71–546.
K. Kult (1947). Klíč k určování brouků čeledi Carabidae Československé republiky [Key to the beetles of family Carabidae of the Czech Republic] (in Czech). Prague.
C. H. Lindroth (1942). Coleoptera, Carabidae. Svensk Insectenfauna, Vol. 9 (in Swedish). Stockholm. pp. 1–260.
Edmund Reitter (1908–1917). Die Käfer des Deutschen Reiches [The beetles of the German Empire] (in German). Stuttgart: K. G. Lutz.