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For the Romanian theater troupe "Cărăbuş", see Constantin Tănase.

Carabus
Carabus glabratus
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Coleoptera
Suborder: Adephaga
Family: Carabidae
Subfamily: Carabinae
Tribe: Carabini
Genus: Carabus
Linnaeus, 1758
Synonyms[1]

Leptinocarabus Reitter, 1895

Carabus ulrichii, Romania
Carabus auratus, Germany
Carabus granulatus, Canada
Carabus glabratus, UK
Carabus nemoralis, Canada
Carabus lineatus lateralis, Spain
Carabus nymph, Switzerland
Carabus rossii, Italy

Carabus is a genus of beetles in the family Carabidae,[2] and was first described in 1758 by Carl Linnaeus.[2][3]

The genus is highly diverse with 94 subgenera, over 900 species and 2300 subspecies, thus is the largest genus in the subfamily Carabinae.[4][5][6] The vast majority are native to the Palearctic, but 16 Nearctic species are also known.[4][6] Carabus species are 12–50 mm (0.47–1.97 in) long, and most species are wingless and often very colourful.[4] These are nocturnal, predatory beetles that feed on snails, earthworms, and caterpillars.[4][7] Most Carabus species were thought to have inhabited the Eurasian forest, but their low dispersal abilities altered the distribution of lineages within the genus.[4]

Diet

[edit]

Adult Carabus feed on both small live and dead animals such as slugs, snails, earthworms, and insects in all stages, sometimes dead vertebrates. The ways of feeding on snails are different for their adaptations, as macrocephalic beetles feed on snails by crushing the snail shell, and stenocephalic beetles feed on snails by inserting their heads into the snail shell.[8]

Species

[edit]
Subgenus Acathaicus Reitter, 1896
Carabus alexandrae Semenov, 1887
Subgenus Acoptolabrus A.Morawitz, 1886
Carabus changgeonleei (Ishikawa & Kim, 1983)
Carabus constricticollis (Kraatz, 1886)
Carabus gehinii Fairmaire, 1876
Carabus haeckeli Brezina & Imura, 1997
Carabus jingkeanus A.Müller, 2007
Carabus leechi (Bates, 1888)
Carabus lopatini A.Morawitz, 1886
Carabus mirabilissimus (Ishikawa & Deuve, 1982)
Carabus planicranion Rapuzzi, 2015
Carabus schrenckii (Motschulsky, 1860)
Subgenus Acoptopterus Lapouge, 1927
Carabus agamemnon Breuning, 1943
Carabus andreiianus Deuve, 2007
Carabus anhuinus Imura, 1996
Carabus baimanorum Deuve, 1999
Carabus baogai Häckel & Sehnal, 2009
Carabus battoniensis Deuve, 1991
Carabus bornianus G.Hauser, 1922
Carabus dabashanus Imura, 1995
Carabus daiyunshan Kleinfeld, 1998
Carabus dongchuanicus Deuve, 1994
Carabus donjuan (Imura, 2011)
Carabus dreuxioides Deuve, 1998
Carabus emanuelei Imura, 1997
Carabus fontellus Deuve, 1997
Carabus jinnanicus Deuve & Tian, 2006
Carabus koidei Imura, 1996
Carabus kucerai Deuve, 1997
Carabus kweitshauensis Mandl, 1975
Carabus lacertosus Cavazzuti, 2008
Carabus latipennis Breuning, 1932
Carabus ludingensis Deuve & Vigna Taglianti, 1992
Carabus maowenensis Deuve, 2001
Carabus mianningensis Kleinfeld, 2000
Carabus mirandus Cavazzuti & Rapuzzi, 2005
Carabus morphocaraboides Deuve, 1989
Carabus nanosomus (G.Hauser, 1931)
Carabus nestor (Breuning, 1934)
Carabus oblongior Deuve, 1992
Carabus ohomopteroides Deuve & Tian, 2004
Carabus paris Breuning, 1932
Carabus platyfluvius Imura, 2012
Carabus pseudolatipennis Deuve, 1991
Carabus relictus Semenov, 1898
Carabus roccaianus Deuve, 2011
Carabus tianbaoshan Kleinfeld, 1998
Carabus tieguanzi Imura, 1990
Carabus vigil Semenov, 1898
Carabus vigilax Bates, 1890
Carabus wutaishanicus A.Müller, 2000
Carabus xupuensis Kleinfeld, 1998
Carabus yunnanus Fairmaire, 1886
Subgenus Acrocarabus Lapouge, 1930
Carabus callisthenoides Semenov, 1888
Carabus guerini Fischer von Waldheim, 1842
Subgenus Alipaster Reitter, 1896
Carabus barovskii Semenov & Znojko, 1932
Carabus dengyuani Deuve & Tian, 2012
Carabus dshungaricola Deuve, 1990
Carabus eokirgisicus Kabak, 1990
Carabus eous A.Morawitz, 1889
Carabus eugeniellus Obydov, 2000
Carabus foreli G.Hauser, 1922
Carabus hiekei Kabak & Kryzhanovskij, 1990
Carabus infantulus A.Morawitz, 1886
Carabus karkarensis Kabak & Ovtshinnikov, 1994
Carabus khalyktauensis Kabak, 2005
Carabus kiritschenkoi Breuning, 1934
Carabus malkovskyi Kabak, 1990
Carabus manap Brezina & Kabak, 1993
Carabus pauliani Deuve, 2004
Carabus pupulus A.Morawitz, 1889
Carabus rueckbeili Breuning, 1932
Carabus semenovianus Breuning, 1932
Carabus semenoviellus Breuning, 1932
Carabus shirtalensis Gottwald, 1990
Carabus thianshanskii Breuning, 1932
Carabus tianshanicola Deuve & Tian, 2003
Carabus valikhanovi Kabak, 1990
Subgenus Apoplesius Deuve, 1990
Carabus omphreodes Reitter, 1898
Subgenus Apotomopterus Hope, 1838
Carabus adolescens (G.Hauser, 1925)
Carabus aeneocupreus (G.Hauser, 1932)
Carabus ajax Breuning, 1933
Carabus angermaieri Kleinfeld, 2008
Carabus anlongensis Deuve & Tian, 2006
Carabus antoniettae Cavazzuti, 2002
Carabus anxiensis Kleinfeld, 1998
Carabus arrowi G.Hauser, 1913
Carabus benardi Breuning, 1931
Carabus birmanus Andrewes, 1929
Carabus boulbenianus Deuve, 1996
Carabus bousquetellus Deuve, 1998
Carabus breuningianus Lemoult, 1930
Carabus candidiequus (Imura, 2009)
Carabus cantonensis (G.Hauser, 1918)
Carabus caoyutangicus Deuve, 2009
Carabus casaleianus Deuve, 1994
Carabus cavazzutiellus Deuve, 2002
Carabus cenwangensis Deuve & Tian, 2002
Carabus chadianus Cavazzuti & Ratti, 1999
Carabus cheni Deuve, 1992
Carabus chenianus Deuve & Tian, 1999
Carabus circe Cavazzuti & Ratti, 1998
Carabus clermontianus Breuning, 1933
Carabus cyanopterus G.Hauser, 1919
Carabus dabamontanus Imura, 1996
Carabus dapanshanicus Deuve, 2013
Carabus datianshanicus Kleinfeld, 1997
Carabus davidis H.Deyrolle, 1878
Carabus dechambreianus Deuve & Li, 2000
Carabus delavayi Fairmaire, 1886
Carabus dilatatipennis Rapuzzi & Cavazzuti, 2006
Carabus ebian Cavazzuti & Rapuzzi, 2010
Carabus eccoptopteroides Rapuzzi, 2012
Carabus eccoptopterus (Kraatz, 1894)
Carabus elektra Kleinfeld, 2004
Carabus elliptipennis Deuve, 1995
Carabus fallettiellus Deuve, 1998
Carabus fanjinensis Deuve & Tian, 2001
Carabus feae Gestro, 1888
Carabus flavihervosus (Imura, 2009)
Carabus foveatus (G.Hauser, 1913)
Carabus francottei Rapuzzi, 2012
Carabus fushuangensis Deuve, 1997
Carabus gracilithorax Deuve, 1989
Carabus grossefoveatus (G.Hauser, 1913)
Carabus guangxicus Deuve, 1989
Carabus guibeicus Deuve & Tian, 1999
Carabus guzhangensis Kleinfeld, 1998
Carabus hera Kleinfeld, 2000
Carabus hiekeianus Deuve, 1991
Carabus huangianus Deuve & Tian, 2003
Carabus hubeicus Deuve, 1991
Carabus ichangensis Bates, 1889
Carabus inagakii Deuve, 1991
Carabus infirmior (G.Hauser, 1924)
Carabus inventoides Deuve & Li in Deuve, 2000
Carabus inventus Cavazzuti, 1999
Carabus jingliae Deuve & Tian, 2019
Carabus jingzhongensis Deuve & Tian, 2005
Carabus kafka Kleinfeld & Reuter, 2010
Carabus keithi Deuve, 1995
Carabus koiwayai Deuve & Imura, 1990
Carabus korellianus Kleinfeld, 2002
Carabus kouanping Maindron, 1906
Carabus kouichii Imura & Mizusawa, 1997
Carabus kryzhanovskianus Deuve, 1992
Carabus lamarcki Deuve, 1994
Carabus lambrechti Kleinfeld, 2006
Carabus laoshanicus Imura, 1995
Carabus leda Kleinfeld, 2000
Carabus lizizhongi Deuve & Tian, 2007
Carabus longeantennatus (G.Hauser, 1931)
Carabus ludivinae Deuve, 1996
Carabus luschanensis (G.Hauser, 1919)
Carabus madefactus Cavazzuti, 1997
Carabus malaisei Breuning, 1947
Carabus maoershanicus Cavazzuti, 1995
Carabus marcilhacianus Deuve & Tian, 2010
Carabus masuzoi (Imura & Satô, 1989)
Carabus mecynodes Bates, 1890
Carabus melli (Born, 1923)
Carabus menelaus Breuning, 1951
Carabus microtatos Cavazzuti, 1997
Carabus ngi Deuve, 1994
Carabus nigrovagans Deuve, 2016
Carabus novenumus Deuve, 1995
Carabus odysseus Breuning, 1932
Carabus pangi Deuve & Tian, 1999
Carabus patroclus Breuning, 1933
Carabus penelope Kleinfeld, 1997
Carabus pineticola Deuve & Mourzine, 2004
Carabus prattianus Bates, 1890
Carabus prodigus Erichson, 1834
Carabus protenes Bates, 1889
Carabus pseudocenwangensis Deuve & Tian, 2005
Carabus pseudoguizhouensis Kleinfeld, 2006
Carabus pseudotorquatus Deuve, 1995
Carabus rapuzzi Kleinfeld, 2000
Carabus reni Deuve & Tian, 2009
Carabus rugulothorax Deuve, 2016
Carabus saga Cavazzuti, 1997
Carabus sauteri Roeschke, 1912
Carabus schuetzei Kleinfeld, 1998
Carabus semelai Deuve, 1995
Carabus shun Deuve, 1995
Carabus solidior Deuve & Imura, 1990
Carabus tanakaianus (Imura, 2005)
Carabus tarasovinus Deuve, 2009
Carabus telemachus (G.Hauser, 1925)
Carabus tengchongicola Deuve, 1999
Carabus thilliezi Deuve, 1994
Carabus thoraciculus Cavazzuti, 1998
Carabus tiani Deuve, 2013
Carabus tibeticus Cavazzuti & Rapuzzi, 2005
Carabus tientei J.Thomson, 1857
Carabus tonkinensis Deuve, 1990
Carabus torquatus Cavazzuti, 1995
Carabus toulgoeti Deuve, 1989
Carabus tuxeni Mandl, 1979
Carabus vitalisi Lapouge, 1918
Carabus vogtae Beheim & Breuning, 1943
Carabus wangziensis Deuve & Tian, 2005
Carabus wumingensis Deuve, 1992
Carabus xianhensis Deuve & Tian, 2007
Carabus xiaoxiangensis Deuve, 1995
Carabus yaophilus Deuve, 1990
Carabus yaorenensis Deuve, 2011
Carabus yinjiangicus Deuve & Tian, 2001
Carabus yonganensis Deuve, 2008
Carabus yuae Deuve, 1989
Carabus yuanbaoensis Deuve, 1994
Carabus yuanshanensis Kleinfeld, 1996
Carabus yunanensis (Born, 1905)
Carabus yundongbeicus Deuve, 2002
Carabus yunlingensis Deuve, 1991
Carabus yunnanicola Deuve, 1989
Carabus zengae Deuve & Tian, 2000
Subgenus Archicarabus Seidlitz, 1887
Carabus alysidotus Illiger, 1798
Carabus monticola Dejean, 1826
Carabus montivagus Palliardi, 1825
Carabus nemoralis O.F.Müller, 1764
Carabus phoenix Lapouge, 1925
Carabus pseudomonticola Lapouge, 1908
Carabus rossii Dejean, 1826
Carabus steuartii A.Deyrolle, 1852
Carabus victor Fischer von Waldheim, 1836
Carabus wiedemanni Ménétriés, 1836
Subgenus Aristocarabus Semenov, 1897
Carabus romanowi Semenov, 1897
Carabus viridifossulatus Fairmaire, 1887
Subgenus Aulonocarabus Reitter, 1896
Carabus canaliculatus M.Adams, 1812
Carabus gaschkewitschi Motschulsky, 1859
Carabus gossarei Haury, 1879
Carabus kabakovi Lafer, 1989
Carabus kolymensis Lafer, 1989
Carabus kurilensis Lapouge, 1913
Carabus mouthiezianus Deuve, 1991
Carabus nangnimicus Blumenthal & Deuve, 1984
Carabus pseudokoreanus Breuning, 1932
Carabus truncaticollis Eschscholtz, 1833
Subgenus Axinocarabus A.Morawitz, 1886
Carabus fedtschenkoi Solsky, 1874
Subgenus Calocarabus Semenov, 1887
Carabus aristochroides Deuve, 1992
Carabus aurocostulus Deuve & Kalab, 2018
Carabus benpo Kalab, 2007
Carabus deuvianus Cavazzuti & Casale, 2006
Carabus dietererberi Heinz, 2001
Carabus gratus Semenov, 1887
Carabus guinanensis Deuve, 1991
Carabus janatai Brezina, 1996
Carabus juengerianus Kleinfeld, 1995
Carabus kalabi Deuve, 1990
Carabus linxiaensis Deuve, 1992
Carabus luzeicola Kalab, 2005
Carabus mandarin Kalab, 2002
Carabus miraculix Kleinfeld & Puchner, 2011
Carabus nevestimus (Imura, 2005)
Carabus przewalskii A.Morawitz, 1886
Carabus punctatocostulus Deuve & Kalab, 2018
Carabus sementivus (Imura, 2005)
Carabus sifanicus Semenov, 1898
Carabus trichothorax Brezina & Imura, 1997
Carabus turnaianus Deuve, 1995
Carabus worahniki Kleinfeld & Puchner, 2011
Subgenus Carabus Linnaeus, 1758
Carabus angustus Roeschke, 1898
Carabus arcensis Herbst, 1784
Carabus billbergi Mannerheim, 1827
Carabus cartereti Deuve, 1982
Carabus catenulatus Scopoli, 1763
Carabus cumanus Fischer von Waldheim, 1823
Carabus deyrollei Gory, 1839
Carabus goryi Dejean, 1831
Carabus granulatus Linnaeus, 1758
Carabus italicus Dejean, 1826
Carabus leplati Deuve & Li, 2008
Carabus manfredeggeri A.Müller, 2012
Carabus manifestus Kraatz, 1881
Carabus menetriesi Hummel, 1827
Carabus mianyangensis Deuve & Li, 1998
Carabus namhaedoensis Kwon & Lee, 1984
Carabus nitididorsus Ishikawa & Kim, 1983
Carabus obsoletus Sturm, 1815
Carabus paraysi Palliardi, 1825
Carabus pawlowskianus Deuve, 1989
Carabus pseudomanifestus Rapuzzi, 2009
Carabus sculpturatus Ménétriés, 1832
Carabus sternbergi Roeschke, 1898
Carabus stscheglowi Mannerheim, 1827
Carabus szeli Deuve, 1994
Carabus ulrichii Germar, 1823
Carabus vanvolxemi Putzeys, 1875
Carabus vinctus (Weber, 1801)
Carabus xiuyanensis Deuve & Li, 1998
Subgenus Cathaicus Bates, 1870
Carabus brandti Faldermann, 1835
Subgenus Cathoplius C.G.Thomson, 1875
Carabus aliai (Escalera, 1944)
Carabus asperatus (Dejean, 1826)
Carabus stenocephalus Lucas, 1866
Subgenus Cechenochilus Motschulsky, 1850
Carabus adangensis Gottwald, 1983
Carabus boeberi M.Adams, 1817
Carabus heydenianus (Starck, 1889)
Carabus kokujewi Semenov, 1898
Carabus lazorum Belousov & Zamotajlov, 1999
Subgenus Cechenotribax Semenov & Znojko, 1932
Carabus petri Semenov & Znojko, 1932
Subgenus Cephalornis Semenov, 1889
Carabus potanini (Semenov, 1887)
Subgenus Chaetocarabus C.G.Thomson, 1875
Carabus arcadicus Gistel, 1848
Carabus intricatus Linnaeus, 1760
Carabus krueperi (Reitter, 1896)
Carabus lefebvrei Dejean, 1826
Carabus merlini Schaum, 1861
Subgenus Chaetomelas C.G.Thomson, 1875
Carabus piochardi (Géhin, 1883)
Subgenus Chrysocarabus C.G.Thomson, 1875
Carabus auronitens Fabricius, 1792
Carabus basilicus Chevrolat, 1836
Carabus hispanus Fabricius, 1787
Carabus olympiae Sella, 1855
Carabus punctatoauratus Germar, 1823
Carabus rutilans Dejean, 1826
Carabus solieri Dejean, 1826
Carabus splendens Olivier, 1790
Carabus strasseri (Lauffer, 1905)
Subgenus Cratocarabus Reitter, 1896
Carabus gorodinskii Obydov, 1998
Carabus kryzhanovskii Bogachev, 1965
Carabus puer A.Morawitz, 1886
Subgenus Cratocechenus Reitter, 1896
Carabus akinini A.Morawitz, 1886
Carabus jacobsoni Semenov, 1908
Carabus ovtshinnikovi Gottwald, 1987
Carabus redikortzevi Semenov & Breuning, 1931
Carabus tshistjakovae Kabak, 2001
Carabus znojkoi Semenov & Breuning, 1931
Subgenus Cratocephalus Kirsch, 1859
Carabus balassogloi C.A.Dohrn, 1881
Carabus chan Breuning, 1932
Carabus cicatricosus Fischer von Waldheim, 1842
Carabus corrugis C.A.Dohrn, 1882
Carabus narinensis Csiki, 1927
Carabus solskyi (Ballion, 1878)
Subgenus Cratophyrtus Reitter, 1896
Carabus hauseri (Reitter in F.Hauser, 1894)
Carabus katajevi Gottwald, 1989
Carabus kaufmanni Solsky, 1874
Carabus nikolajevi Kabak, 1998
Carabus turcosinensis Mandl, 1955
Subgenus Cryptocarabus Reitter, 1896
Carabus iliensis Kabak, 1994
Carabus kadyrbekovi Kabak, 1994
Carabus lindemanni Ballion, 1878
Carabus merkensis Kabak, 1992
Carabus mullerellus Beheim & Breuning, 1943
Carabus munganasti Reitter, 1909
Carabus sacarum Kabak, 1998
Carabus sororius A.Morawitz, 1886
Carabus subparallelus Ballion, 1878
Carabus tsharynensis Kabak, 1994
Carabus turkestanus Breuning, 1928
Subgenus Ctenocarabus C.G.Thomson, 1875
Carabus galicianus Gory, 1839
Carabus melancholicus Fabricius, 1798
Subgenus Cupreocarabus Deuve, 1997
Carabus balangicus Cavazzuti, 2002
Carabus brezinai Deuve, 1994
Carabus concursans Deuve & Kalab, 2010
Carabus drahoslavae Brezina & Häckel, 2006
Carabus huangi Deuve, 1992
Carabus laevithorax Breuning, 1935
Carabus lixianensis Deuve, 1990
Carabus melii Cavazzuti & Rapuzzi, 2005
Carabus minimocupreus Deuve, 2003
Carabus miroslavi Deuve, 2000
Carabus mollardianus Deuve, 2016
Carabus pseudohuangi Deuve, 1995
Carabus quintus Cavazzuti & Rapuzzi, 2009
Carabus sackenioides Deuve, 1991
Carabus sculptior Deuve, 1992
Carabus siguniangensis Deuve, 1997
Carabus verecundus Deuve & Kalab, 2010
Carabus xichang Kleinfeld & Puchner, 2013
Carabus zhegushanus Cavazzuti & Rapuzzi, 2005
Subgenus Cychrostomus Reitter, 1896
Carabus anchocephalus Reitter, 1896
Carabus pseudoprosodes Semenov & Znojko, 1932
Subgenus Cyclocarabus Reitter, 1896
Carabus aulacocnemus Semenov, 1897
Carabus belousovi Kabak, 1992
Carabus brosciformis Semenov, 1897
Carabus karaterekensis Kalab, 1996
Carabus mniszechii Chaudoir, 1852
Carabus namanganensis Heyden, 1886
Carabus pseudolamprostus Kalab, 1996
Carabus pskemicus Deuve & Kalab, 1993
Carabus pullus Semenov & Znojko, 1932
Carabus zarudnyi Semenov & Znojko, 1932
Subgenus Damaster Kollar, 1836
Carabus augustus Bates, 1888
Carabus blaptoides (Kollar, 1836)
Carabus elysii J.Thomson, 1856
Carabus formosus Semenov, 1887
Carabus fruhstorferi (Roeschke, 1900)
Carabus gemmifer (Fairmaire, 1887)
Carabus guerryi Born, 1903
Carabus ignigena (G.Hauser, 1914)
Carabus ignimitella Bates, 1888
Carabus ishizukai Deuve & Ohshima, 1989
Carabus jankowskii (Oberthür, 1883)
Carabus kubani Deuve, 1990
Carabus lafossei Feisthamel, 1845
Carabus liianus Deuve, 2008
Carabus monilifer Tatum, 1847
Carabus nankotaizanus Kano, 1932
Carabus osawai Imura; Zhou & Su, 1999
Carabus principalis Bates, 1889
Carabus pustulifer Lucas, 1869
Carabus smaragdinus Fischer, 1823
Subgenus Deroplectes Reitter, 1895
Carabus arcanus Semenov, 1898
Carabus dokhtouroffi Ganglbauer, 1887
Carabus klapperichianus Mandl, 1955
Carabus sphinx Reitter, 1895
Carabus staudingeri Ganglbauer, 1886
Subgenus Diocarabus Reitter, 1896
Carabus aurocinctus Motschulsky, 1844
Carabus beybienkoi Kryzhanovskij, 1973
Carabus caustomarginatus Imura & Mizusawa, 1994
Carabus chamissonis Fischer von Waldheim, 1820
Carabus dorogostaiskii Shilenkov, 1983
Carabus fraterculus Reitter, 1895
Carabus loschnikovii Fischer von Waldheim, 1823
Carabus lucepunctus Cavazzuti & Rapuzzi, 2005
Carabus massagetus Motschulsky, 1844
Carabus opaculus Putzeys, 1875
Carabus slovtzovi Mannerheim, 1849
Subgenus Eccoptolabrus Semenov, 1898
Carabus exiguus Semenov, 1898
Carabus sunwukong Imura, 1993
Subgenus Euleptocarabus Nakane, 1955
Carabus porrecticollis Bates, 1883
Subgenus Eupachys Chaudoir, 1857
Carabus acutithorax Deuve, 1989
Carabus glyptopterus Fischer von Waldheim, 1828
Subgenus Eurycarabus Géhin, 1876
Carabus faminii Dejean, 1826
Carabus favieri Fairmaire, 1859
Carabus genei Gené, 1839
Subgenus Fulgenticarabus Deuve & Li, 1998
Carabus flutschi Deuve & Li, 1998
Subgenus Gnathocarabus Deuve, 1991
Carabus kusnetzovi Semenov, 1903
Subgenus Goniocarabus Géhin, 1885
Carabus caerulans A.Morawitz, 1886
Carabus grombczewskii Semenov, 1891
Carabus gussakovskii Kryzhanovskij, 1971
Carabus perelloi Casale, 1979
Carabus sogdianus Semenov, 1898
Carabus stackelbergi Kryzhanovskij, 1971
Carabus tadzhikistanus Kryzhanovskij, 1968
Subgenus Hemicarabus Géhin, 1876
Carabus macleayi Dejean, 1826
Carabus nitens Linnaeus, 1758
Carabus serratus Say, 1823
Carabus tuberculosus Dejean, 1829
Subgenus Heterocarabus A.Morawitz, 1886
Carabus bischoffii Chaudoir, 1848
Carabus marietti Cristofori & Jan, 1837
Carabus muchei Breuning, 1961
Subgenus Homoeocarabus Reitter, 1896
Carabus maeander Fischer von Waldheim, 1820
Subgenus Hygrocarabus C.G.Thomson, 1875
Carabus nodulosus Creutzer, 1799
Carabus variolosus Fabricius, 1787
Subgenus Hypsocarabus Semenov, 1898
Carabus laotse Breuning, 1943
Carabus latro Semenov, 1898
Carabus mikhaili Deuve & Mourzine, 1997
Carabus qinlingensis Imura, 1993
Carabus taibaiensis Kleinfeld, 2001
Carabus tewoensis Deuve, 1992
Subgenus Imaibius Bates, 1889
Carabus anami (Ledoux, 1977)
Carabus baronii Heinertz, 1977
Carabus barysomus Bates, 1889
Carabus boysii Tatum, 1851
Carabus caschmirensis Kollar & L.Redtenbacher, 1844
Carabus cavifrons Mandl, 1974
Carabus dardiellus Bates, 1889
Carabus epipleuralis Semenov, 1907
Carabus erberi Heinz, 1983
Carabus gandharae Heinertz, 1978
Carabus gridellii Breuning, 1959
Carabus isabellae Lassalle, 1985
Carabus kaghanensis Heinertz, 1978
Carabus leepai Heinz, 1993
Carabus nouristani (Ledoux, 1977)
Carabus olafi Deuve & J.Schmidt, 2007
Carabus pachtoun (Ledoux, 1975)
Carabus piffli Mandl, 1961
Carabus rostianus Semenov, 1907
Carabus stoliczkanus Bates, 1878
Carabus wittmerorum Heinertz, 1978
Subgenus Iniopachus Solier, 1848
Carabus auriculatus Putzeys, 1872
Carabus pyrenaeus Audinet-Serville, 1821
Subgenus Ischnocarabus Kraatz, 1877
Carabus cychropalpus Peyron, 1858
Carabus tenuitarsis (Kraatz, 1877)
Subgenus Isiocarabus Reitter, 1896
Carabus dargei Deuve, 1988
Carabus fiduciarius J.Thomson, 1856
Carabus gressittianus Mandl, 1975
Carabus heterodynamus Deuve, 1991
Carabus hienfoungii J.Thomson, 1857
Carabus hunanicola Deuve & Yu, 1992
Carabus kiukiangensis Bates, 1888
Carabus miaorum Lassalle & Prunier, 1993
Carabus pseudohunanicola Deuve & Tian, 2009
Carabus pustululatus Deuve, 1993
Carabus strandiellus Breuning, 1934
Subgenus Lamprostus Motschulsky, 1866
Carabus calleyi Fischer von Waldheim, 1823
Carabus cylindricior Deuve, 1994
Carabus erenleriensis Schweiger, 1964
Carabus guycolasianus Deuve, 2001
Carabus hemprichii Dejean, 1826
Carabus ledouxi Deuve, 2001
Carabus punctatus Laporte, 1834
Carabus renardi Chaudoir, 1846
Carabus robustus E.Deyrolle, 1869
Carabus rostandianus Deuve, 2005
Carabus saulcyi Piochard de la Brûlerie, 1875
Carabus seroulibkin Cavazzuti, 2008
Carabus spinolae Cristofori & Jan, 1837
Carabus staveni Heinz, 2002
Carabus syrus Roeschke, 1898
Carabus torosus I.Frivaldszky von Frivald, 1835
Subgenus Leptocarabus Géhin, 1885
Carabus arboreus Lewis, 1882
Carabus harmandi Lapouge, 1909
Carabus hiurai Kamiyoshi & Mizoguchi, 1960
Carabus koreanus Reitter, 1895
Carabus kumagaii (Kimura & Komiya, 1974)
Carabus kyushuensis Nakane, 1961
Carabus marcilhaci Deuve, 1992
Carabus procerulus Chaudoir, 1862
Carabus seishinensis (Lapouge, 1932)
Carabus semiopacus Reitter, 1895
Carabus vogtianus Beheim & Breuning, 1943
Carabus yokoae Deuve, 1988
Subgenus Leptoplesius Reitter, 1898
Carabus dolini Deuve, 1992
Carabus dolonicus Obydov, 1996
Carabus horoshanensis Deuve, 2002
Carabus itshkibashi Kabak, 2004
Carabus kleinfeldorum Kabak & Putchkov, 1995
Carabus marquardti Reitter, 1898
Carabus merzbacheri G.Hauser, 1922
Carabus shokalskii Semenov & Breuning, 1931
Carabus subtilistriatus G.Hauser, 1922
Subgenus Limnocarabus Géhin, 1876
Carabus clatratus Linnaeus, 1760
Subgenus Lipaster Motschulsky, 1866
Carabus gordius Reitter, 1898
Carabus stjernvalli Mannerheim, 1830
Subgenus Macrothorax Chenu, 1851
Carabus aumontii Lucas, 1850
Carabus meurguesianus Ledoux, 1990
Carabus morbillosus Fabricius, 1792
Carabus planatus Chaudoir, 1843
Carabus rugosus Fabricius, 1792
Subgenus Meganebrius Kraatz, 1895
Carabus alanstivelli (Morvan, 1981)
Carabus arunensis Heinertz, 1980
Carabus deliae (Morvan, 1972)
Carabus dilatotarsalis Mandl, 1979
Carabus epsteini Heinertz, 1978
Carabus everesti Andrewes, 1929
Carabus franzi Mandl, 1974
Carabus granulatocostatus Mandl, 1965
Carabus indicus Fairmaire, 1889
Carabus kadoudali (Morvan, 1982)
Carabus koganae Colas, 1961
Carabus lebretae Colas, 1961
Carabus montreuili Deuve, 2003
Carabus pseudoharmandi Mandl, 1965
Carabus queinneci Deuve, 1983
Carabus quinlani Mandl, 1965
Carabus salpansis Deuve, 1984
Carabus santostamangi Deuve & J.Schmidt, 2017
Carabus scheibei Eidam, 1937
Carabus swatensis Heinertz, 1979
Carabus tamang Deuve, 1983
Carabus thudamensis Deuve & J.Schmidt, 2017
Carabus tuberculipennis Mandl, 1974
Carabus wallichii Hope, 1831
Subgenus Megodontoides Deuve, 1991
Carabus erwini Mandl, 1975
Carabus promachus Bates, 1891
Carabus qiangding Kleinfeld & Puchner, 2007
Carabus thibetanus Breuning, 1950
Subgenus Megodontus Solier, 1848
Carabus aurolimbatus Dejean, 1830
Carabus avinovi Semenov & Znojko, 1932
Carabus blakistoni Newman, 1858
Carabus bonvouloirii Chaudoir, 1863
Carabus caelatus Fabricius, 1801
Carabus croaticus Dejean, 1826
Carabus danilevskii Obydov, 1993
Carabus exaratus Quensel, 1806
Carabus germarii Sturm, 1815
Carabus imperialis Fischer von Waldheim, 1823
Carabus kantaikensis Géhin, 1885
Carabus kolbei Roeschke, 1897
Carabus leachii Fischer von Waldheim, 1823
Carabus obovatus Fischer von Waldheim, 1828
Carabus persianus Roeschke, 1896
Carabus planicollis Küster, 1846
Carabus purpurascens Fabricius, 1787
Carabus schoenherri Fischer von Waldheim, 1820
Carabus septemcarinatus Motschulsky, 1840
Carabus stroganowi Zoubkoff, 1837
Carabus tuvensis Shilenkov, 1996
Carabus vietinghoffii M.Adams, 1812
Carabus violaceus Linnaeus, 1758
Subgenus Mesocarabus C.G.Thomson, 1875
Carabus dufourii Dejean, 1829
Carabus lusitanicus Fabricius, 1801
Carabus problematicus Herbst, 1786
Carabus riffensis Fairmaire, 1872
Subgenus Microplectes Reitter, 1896
Carabus convallium (Starck, 1889)
Carabus riedelii Ménétriés, 1832
Subgenus Mimocarabus Géhin, 1876
Carabus elbursensis Breuning, 1946
Carabus khorasanensis Deuve, 1993
Carabus maurus (M.Adams, 1817)
Carabus pumilio Küster, 1846
Carabus roseni Reitter, 1897
Subgenus Morphocarabus Géhin, 1876
Carabus aeruginosus Fischer von Waldheim, 1820
Carabus brinevi Kabak, 2014
Carabus chaudoirii Gebler, 1847
Carabus excellens Fabricius, 1798
Carabus gebleri Fischer von Waldheim, 1817
Carabus henningi Fischer von Waldheim, 1817
Carabus hummelii Fischer von Waldheim, 1823
Carabus karpinskii Kryzhanovskij & Matveev, 1993
Carabus kollari Palliardi, 1825
Carabus kozhantschikowi Lutshnik, 1924
Carabus mestscherjakovi Lutshnik, 1924
Carabus michailovi Kabak, 1992
Carabus monilis Fabricius, 1792
Carabus odoratus Motschulsky, 1844
Carabus praecellens Palliardi, 1825
Carabus regalis Fischer von Waldheim, 1820
Carabus rothii Dejean, 1829
Carabus scheidleri Panzer, 1799
Carabus spasskianus Fischer von Waldheim, 1823
Carabus tarbagataicus Kraatz, 1878
Carabus venustus A.Morawitz, 1862
Carabus versicolor I.Frivaldszky von Frivald, 1835
Carabus wulffiusi A.Morawitz, 1862
Carabus zawadzkii Kraatz, 1854
Carabus zhubajie Imura, 1993
Subgenus Neoplectes Reitter, 1885
Carabus chaudoirianus Lapouge, 1909
Carabus iberus (Fischer von Waldheim, 1823)
Carabus lafertei Chaudoir, 1846
Carabus martviliensis Retezar & Djavelidze, 1992
Carabus mellyi Chaudoir, 1846
Carabus prunierianus Deuve, 2012
Carabus szekelyii Retezar, 2011
Carabus titarenkoi Zamotajlov & Fominykh, 2014
Subgenus Neoplesius Reitter, 1896
Carabus alexpuchneri Kleinfeld & Puchner, 2011
Carabus alpherakii Semenov, 1898
Carabus borodini Heinz, 1996
Carabus bruggeianus Deuve, 1992
Carabus chomae (Imura, 2002)
Carabus chuandonzicus (Imura, 2011)
Carabus danae Kalab, 1995
Carabus ditomoides Deuve, 1991
Carabus edacis Cavazzuti, 2006
Carabus glinkai Heinz, 1996
Carabus hummelioides Deuve, 1989
Carabus kamensis Semenov, 1903
Carabus kaznakovi Semenov & Znojko, 1932
Carabus kozloviellus Semenov & Znojko, 1932
Carabus lama Semenov, 1898
Carabus leptoplesioides Deuve, 1992
Carabus mouthiezianoides Deuve & Kalab, 1992
Carabus nanschanicus Semenov, 1898
Carabus noctivagus Deuve, 1992
Carabus panda Deuve, 1988
Carabus paulusi Kalab, 1995
Carabus pseudomarkamensis Deuve, 1992
Carabus puetzi Kleinfeld, 2000
Carabus sinotibeticola Mandl, 1975
Carabus takashimai Deuve & Imura, 1993
Carabus tsogoensis Deuve, 1997
Carabus wagae Fairmaire, 1882
Carabus wrzecionkoianus Deuve, 2007
Carabus yushuensis Deuve, 1991
Carabus zhangjieae Deuve & Li, 2020
Subgenus Nesaeocarabus Bedel, 1895
Carabus abbreviatus Brullé, 1835
Carabus coarctatus Brullé, 1836
Carabus faustus Brullé, 1836
Carabus gomerae A.Müller, 2004
Subgenus Ohomopterus Reitter, 1896
Carabus albrechti A.Morawitz, 1862
Carabus arrowianus Breuning, 1934
Carabus chugokuensis (Nakane, 1961)
Carabus daisen (Nakane, 1953)
Carabus dehaanii Chaudoir, 1848
Carabus esakii Csiki, 1927
Carabus insulicola Chaudoir, 1869
Carabus iwawakianus (Nakane, 1953)
Carabus japonicus Motschulsky, 1858
Carabus kimurai (Ishikawa, 1966)
Carabus komiyai (Ishikawa, 1966)
Carabus lewisianus Nakane, 1953
Carabus maiyasanus Bates, 1873
Carabus sue Imura, 2012
Carabus uenoi (Ishikawa, 1960)
Carabus yaconinus Bates, 1873
Carabus yamato (Nakane, 1953)
Subgenus Ophiocarabus Reitter, 1896
Carabus aeneolus A.Morawitz, 1886
Carabus arshanicus Kabak, 2005
Carabus ballionis Kraatz, 1879
Carabus confinis Semenov, 1888
Carabus ernsti Kabak, 2001
Carabus imperfectus Semenov, 1887
Carabus iteratus Breuning, 1934
Carabus juldusanus Breuning, 1933
Carabus kalabellus Deuve, 1993
Carabus kurdaiensis Kabak, 2016
Carabus latiballioni Deuve, 1993
Carabus politulus A.Morawitz, 1886
Carabus praecox Semenov, 1898
Carabus regeli A.Morawitz, 1886
Carabus rufocuprescens Deuve, 1993
Carabus successor Reitter, 1896
Carabus tekesensis Deuve & Tian, 2004
Carabus variabilis Ballion, 1878
Carabus wusunshanicus Kabak, 2015
Subgenus Oreocarabus Géhin, 1876
Carabus ghilianii LaFerté-Sénectère, 1847
Carabus guadarramus LaFerté-Sénectère, 1847
Carabus luetgensi Beuthin, 1886
Carabus reitterianus Breuning, 1932
Subgenus Orinocarabus Kraatz, 1878
Carabus alpestris Sturm, 1815
Carabus bertolinii (Kraatz, 1878)
Carabus bremii Stierlin, 1881
Carabus carinthiacus Sturm, 1815
Carabus castanopterus A. & G.B.Villa, 1833
Carabus concolor Fabricius, 1792
Carabus fairmairei C.G.Thomson, 1875
Carabus linnei Panzer, 1810
Carabus pedemontanus Ganglbauer, 1891
Carabus sylvestris Panzer, 1793
Subgenus Oxycarabus Semenov, 1898
Carabus saphyrinus Cristofori & Jan, 1837
Subgenus Pachycarabus Géhin, 1876
Carabus imitator Reitter, 1883
Carabus koenigi Ganglbauer, 1887
Carabus roseri Faldermann, 1836
Carabus staehlini M.Adams, 1817
Carabus swaneticus Reitter, 1883
Subgenus Pachystus Motschulsky, 1866
Carabus cavernosus I.Frivaldszky von Frivald, 1838
Carabus cribratus Quensel, 1806
Carabus gemellatus Ménétriés, 1832
Carabus glabratus Paykull, 1790
Carabus graecus Dejean, 1826
Carabus hortensis Linnaeus, 1758
Carabus hungaricus Fabricius, 1792
Carabus morio Mannerheim, 1830
Carabus pisidicus (Peyron, 1855)
Carabus preslii Dejean, 1830
Carabus tamsii Ménétriés, 1832
Subgenus Pagocarabus A.Morawitz, 1886
Carabus caelestinus (Imura, 2007)
Carabus crassesculptus Kraatz, 1881
Carabus draco Brezina, 1999
Carabus hengduanicola Deuve, 1996
Carabus ludmilae Deuve, 1992
Carabus sichuanicola Deuve, 1989
Carabus trachynodes Bates, 1891
Carabus tryznai Brezina & Häckel, 2006
Subgenus Pantophyrtus Thieme, 1881
Carabus alajensis Semenov, 1897
Carabus brachypedilus A.Morawitz, 1886
Carabus debilis Semenov, 1897
Carabus evstigneevi Obydov, 1997
Carabus ferghanicus Breuning, 1933
Carabus longipedatus Belousov & Kabak, 1993
Carabus turcomanorum (Thieme, 1881)
Carabus validus (Kraatz, 1884)
Subgenus Piocarabus Reitter, 1896
Carabus blumenthaliellus Deuve, 1988
Carabus choui Deuve, 1989
Carabus kitawakianus Imura, 1993
Carabus nanwutai Kleinfeld; Korell & Wrase, 1996
Carabus ohshimaianus Deuve, 1988
Carabus sui Imura & Zhou, 1998
Carabus titanus Breuning, 1933
Carabus vladsimirskyi Dejean, 1830
Subgenus Platycarabus A.Morawitz, 1886
Carabus creutzeri Fabricius, 1801
Carabus cychroides Baudi di Selve, 1864
Carabus depressus Bonelli, 1810
Carabus fabricii Panzer, 1810
Carabus irregularis Fabricius, 1792
Subgenus Procerus Dejean, 1821
Carabus bulgharmaadensis (E.Bodemeyer, 1915)
Carabus caucasicus M.Adams, 1817
Carabus elbursianus Mandl, 1958
Carabus gigas Creutzer, 1799
Carabus scabrosus Olivier, 1790
Carabus syriacus (Kollar in L.Redtenbacher, 1843)
Subgenus Procrustes Bonelli, 1810
Carabus anatolicus (Chaudoir, 1857)
Carabus banonii Dejean, 1829
Carabus chevrolati Cristofori & Jan, 1837
Carabus clypeatus M.Adams, 1817
Carabus coriaceus Linnaeus, 1758
Carabus hybridus (Ganglbauer, 1887)
Carabus mulsantianus A.Morawitz, 1886
Carabus paiafa White, 1845
Carabus talychensis (Ménétriés, 1832)
Subgenus Pseudocoptolabrus Reitter, 1896
Carabus armiger Imura, 1997
Carabus belousovianus (Imura, 2011)
Carabus branaungi Imura, 1999
Carabus burmanensis Breuning, 1932
Carabus businskyi Deuve, 1990
Carabus chortenensis Cavazzuti, 2005
Carabus firmatus Cavazzuti, 1997
Carabus masahiroi (Imura, 2006)
Carabus nosei Imura, 1997
Carabus pseudochortenensis Deuve & Tian, 2020
Carabus taliensis (Fairmaire, 1886)
Carabus watanabei (Imura, 2003)
Subgenus Pseudocranion Reitter, 1896
Carabus aba Kalab, 2002
Carabus absonus Cavazzuti & Rapuzzi, 2005
Carabus baxianus Cavazzuti & Rapuzzi, 2007
Carabus brachygnathus Deuve, 2002
Carabus flavigenua Cavazzuti, 2002
Carabus fumigatus Semenov, 1898
Carabus gansuensis Semenov, 1887
Carabus gonggaicus Deuve, 1989
Carabus jiudingensis Deuve, 1994
Carabus kitawakiellus Imura, 1995
Carabus labrangicus Deuve, 1992
Carabus lazikouensis Deuve, 1997
Carabus meditabundus Deuve, 1992
Carabus pseudosackeni Deuve, 1989
Carabus sackeni Semenov, 1898
Carabus shuamaluko Deuve, 1991
Carabus sinicus Breuning, 1950
Carabus taibaishanicus Deuve, 1989
Carabus tibetanophilus Deuve, 1991
Carabus viatorum Deuve, 1992
Carabus wenxianicola Deuve, 1996
Carabus zhanglaensis Deuve, 1991
Subgenus Rhigocarabus Reitter, 1896
Carabus allegroi Cavazzuti & Rapuzzi, 2007
Carabus batangicoides Deuve & Tian, 2011
Carabus boanoi Cavazzuti, 2003
Carabus broukpytlik Brezina & Häckel, 2004
Carabus buddaicus Semenov, 1887
Carabus casanova (Imura & Brezina, 2008)
Carabus cateniger A.Morawitz, 1886
Carabus cechenoides Deuve & Kalab, 2014
Carabus dacatraianus Deuve, 1995
Carabus dubifer Deuve & Kalab, 2014
Carabus dungchen Deuve & Kalab, 2019
Carabus fubianensis Deuve & Kalab, 2007
Carabus gentleman Brezina & Häckel, 2004
Carabus gigolo Heinz & Brezina, 1996
Carabus gigoloides Cavazzuti, 2000
Carabus gracilicollis Semenov, 1887
Carabus handelmazzettii Mandl, 1955
Carabus humilior Deuve & Kalab, 2012
Carabus impavidus Cavazzuti, 2008
Carabus indigestus Semenov, 1898
Carabus itzingeri (Breuning, 1934)
Carabus jintangicus Deuve, 2001
Carabus jiulongensis Deuve, 1994
Carabus ladygini Semenov, 1903
Carabus legrandianus Deuve & Tian, 2007
Carabus maleki Deuve, 1991
Carabus mifan Kalab, 2005
Carabus morawitzianus Semenov, 1887
Carabus nianjuaensis Imura, 2016
Carabus paraxiei Kleinfeld & Puchner, 2013
Carabus pepek (Imura & Kalab, 2006)
Carabus poeta Semenov, 1898
Carabus propiorthais Cavazzuti, 2000
Carabus pseudopusio Deuve, 1996
Carabus pusioides Cavazzuti & Rapuzzi, 2007
Carabus quindecim Cavazzuti, 2002
Carabus rhododendron Deuve & Imura, 1991
Carabus roborowskii Semenov, 1887
Carabus sehnali Brezina & Häckel, 2006
Carabus shaluishan Kleinfeld & Puchner, 2011
Carabus sininensis Semenov, 1898
Carabus thais Heinz, 1997
Carabus turnai Deuve, 1994
Carabus wengdaensis Kleinfeld & Puchner, 2009
Carabus xiei Deuve, 1992
Subgenus Scambocarabus Reitter, 1896
Carabus auritus Cavazzuti, 2000
Carabus kruberi Fischer von Waldheim, 1820
Carabus modestulus Semenov, 1887
Carabus sculptipennis Chaudoir, 1877
Carabus shaanxiensis Deuve, 1991
Subgenus Semnocarabus Reitter, 1896
Carabus carbonicolor A.Morawitz, 1886
Carabus cicatricosulus A.Morawitz, 1886
Carabus erosus Motschulsky, 1866
Carabus perminimus Deuve, 1994
Carabus regulus C.A.Dohrn, 1882
Carabus rustemi Kabak, 2010
Carabus tekeliensis Kabak, 2001
Carabus transiliensis Semenov, 1897
Subgenus Shunichiocarabus Imura, 1995
Carabus uenoianus Imura, 1995
Subgenus Sphodristocarabus Géhin, 1885
Carabus adamsi M.Adams, 1817
Carabus biroi Csiki, 1927
Carabus bohemani Ménétriés, 1832
Carabus coruhnehriensis Cavazzuti, 1990
Carabus elegantulus Motschulsky, 1850
Carabus enigmaticus Heinz, 1980
Carabus georgia Cavazzuti, 1984
Carabus gilnickii E.Deyrolle, 1869
Carabus heinzi Breuning, 1964
Carabus karasudominus Cavazzuti, 1991
Carabus kurdicus Heinz, 1975
Carabus macrogonus Chaudoir, 1847
Carabus pavesii Cavazzuti, 1992
Carabus scovitzii Faldermann, 1836
Carabus separatus Lapouge, 1907
Carabus tokatensis Roeschke, 1898
Carabus varians Fischer von Waldheim, 1823
Subgenus Stephanocarabus Imura, 1995
Carabus fraterculoides Breuning, 1961
Subgenus Tachypus Weber, 1801
Carabus auratus Linnaeus, 1760
Carabus cancellatus Illiger, 1798
Carabus vagans Olivier, 1795
Subgenus Tanaocarabus Reitter, 1896
Carabus finitimus Haldeman, 1852
Carabus forreri Bates, 1882
Carabus hendrichsi Bolivar y Pieltain; Rotger & Coronado, 1967
Carabus sylvosus Say, 1823
Carabus taedatus Fabricius, 1787
Subgenus Teratocarabus Semenov & Znojko, 1932
Carabus azrael Semenov & Znojko, 1932
Subgenus Tmesicarabus Reitter, 1896
Carabus cristoforii Spence, 1833
Subgenus Tomocarabus Reitter, 1896
Carabus bessarabicus Fischer von Waldheim, 1823
Carabus convexus Fabricius, 1775
Carabus decolor Fischer von Waldheim, 1823
Carabus marginalis Fabricius, 1794
Carabus rumelicus Chaudoir, 1867
Carabus scabripennis Chaudoir, 1850
Subgenus Trachycarabus Géhin, 1876
Carabus besseri Fischer von Waldheim, 1820
Carabus coriaceipennis Chaudoir, 1863
Carabus estreicheri Fischer von Waldheim, 1820
Carabus latreillei Fischer von Waldheim, 1820
Carabus mandibularis Fischer von Waldheim, 1828
Carabus perrini Dejean, 1831
Carabus planarius Obydov, 1994
Carabus scabriusculus Olivier, 1795
Carabus sibiricus Fischer von Waldheim, 1820
Subgenus Tribax Fischer von Waldheim, 1817
Carabus adelphus (Rost, 1892)
Carabus agnatus (Ganglbauer, 1889)
Carabus apollo (Zolotarev, 1913)
Carabus apschuanus (Rost, 1893)
Carabus balkaricus Belousov & Abdurakhmanov, 1991
Carabus biebersteini Ménétriés, 1832
Carabus certus (Reitter, 1896)
Carabus circassicus (Ganglbauer, 1886)
Carabus compressus Chaudoir, 1846
Carabus curlettii Cavazzuti, 1984
Carabus daphnis Kurnakov, 1962
Carabus edithae Reitter, 1893
Carabus edmundi Semenov, 1897
Carabus jason Semenov, 1898
Carabus kasakorum Semenov, 1897
Carabus kasbekianus Kraatz, 1877
Carabus koltzei (Rost, 1889)
Carabus komarowi Reitter, 1882
Carabus kraatzi Chaudoir, 1877
Carabus kratkyi Ganglbauer, 1891
Carabus lailensis Belousov, 1992
Carabus lederi Reitter, 1882
Carabus macropus Chaudoir, 1877
Carabus merdeniki Cavazzuti & Korell, 1992
Carabus nothus M.Adams, 1817
Carabus osseticus M.Adams, 1817
Carabus plasoni (Ganglbauer, 1886)
Carabus polychrous (Rost, 1892)
Carabus prometheus Reitter, 1887
Carabus protensus Schaum, 1864
Carabus puschkini M.Adams, 1817
Carabus reitteri (Retowski, 1885)
Carabus satyrus Kurnakov, 1962
Carabus schamylii Hampe, 1852
Carabus shtchurovi Belousov & Zamotajlov, 1993
Carabus starcki (Heyden, 1885)
Carabus starckianus (Ganglbauer, 1886)
Carabus titan (Zolotarev, 1913)
Subgenus Ulocarabus Reitter, 1896
Carabus stschurowskii Solsky, 1874
Carabus theanus Reitter, 1895
Extinct, not assigned to subgenus
†Carabus ceresti Nel, 1987
†Carabus dzieduszyckii Lomnicki, 1894
†Carabus elongatus Brodie, 1845
†Carabus foveolatus Piton & Theobald, 1935
†Carabus jeffersoni Scudder, 1900
†Carabus maeandroides Lomnicki, 1894
†Carabus mecothoracus Zhang: Liu & Shangguan, 1989
†Carabus neli Deuve, 1998
†Carabus novalensis Omboni, 1886
†Carabus ovalis Hong & Wang, 1986
†Carabus praearvensis Lomnicki, 1894
†Carabus praeviolaceus Lomnicki, 1894
†Carabus westwoodii Giebel, 1856
†Carabus winkleri Weyenbergh, 1869

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Carabus

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Carabus is a of large, predatory ground beetles in the family Carabidae (subfamily Carabinae, tribe Carabini), comprising approximately 970–1,000 characterized by their robust bodies, often metallic or iridescent coloration, and nocturnal habits. These beetles, established taxonomically by in 1758, are primarily distributed across the Holarctic region, with the vast majority of species confined to and a few adventive in . Members of the genus Carabus are typically 15–35 mm in length, featuring elongated legs suited for rapid , powerful mandibles for subduing prey, and elytra (wing covers) that may display punctures, ridges, or metallic sheens ranging from green and blue to bronze and black. They inhabit diverse terrestrial environments, including forests, grasslands, alpine meadows, and riverbanks, often preferring moist, vegetated s under leaf litter or stones where they hunt at night. Ecologically, Carabus are carnivorous predators, feeding on a variety of such as (many are helicophagous, specializing in breaking snail shells), earthworms, caterpillars, and other , thereby serving as natural biocontrol agents against agricultural pests. Larvae are also predacious, developing in and contributing to the genus's role in regulation, though some exhibit habitat specificity and vulnerability to . The is highly diverse, organized into about 90 subgenera, reflecting adaptations to varied ecological niches across temperate zones, with ongoing highlighting evolutionary radiations and phylogenetic relationships through molecular studies. Certain Carabus species, such as those in the , demonstrate specialized morphologies like flattened bodies and long legs for alpine or riparian habitats, underscoring their adaptability and conservation importance in European hotspots.

Taxonomy and phylogeny

Classification

Carabus is classified in the kingdom Animalia, phylum Arthropoda, class Insecta, order Coleoptera, family Carabidae, subfamily Carabinae, tribe Carabini, and genus Carabus Linnaeus, 1758. As the type genus of Carabidae, Carabus occupies a basal phylogenetic position within the tribe Carabini and is closely related to other genera in the tribe, particularly its sister genus Calosoma. The encompasses approximately 970 arranged across 91 subgenera, making it the most species-rich in Carabinae. Key examples include Leptinocarabus (distinguished by fine elytral punctures and slender body form), Ulocarabus (characterized by robust build and pronounced pronotal margins), and Archicarabus (noted for archaic elytral microsculpture and relictual morphology). Carabus originated during the early Eocene, with major diversification occurring in the mid-Miocene. Its evolutionary is marked by low dispersal capabilities, often due to flightlessness in many lineages, resulting in fragmented, relictual distributions across Holarctic temperate habitats.

Etymology and

The genus name Carabus derives from the Ancient Greek term kárabos (κάραβος), referring to a "horned " or "crawfish," an allusion to the elongated, segmented body shape of these ground beetles resembling crustaceans. This nomenclature was formally established by in the 10th edition of Systema Naturae (1758), where he designated Carabus as a within Coleoptera and described initial such as C. granulatus. Following Linnaeus's foundational work, the classification of Carabus expanded during the 18th and 19th centuries through contributions by European entomologists. Pierre François Marie Auguste Dejean cataloged numerous Carabus in his multi-volume Catalogue de Coléoptères (1821–1836), emphasizing morphological variations and geographic distributions across the Palearctic region. Similarly, Maximilien de Chaudoir advanced the in monographs like Monographie des Carabus (e.g., 1840s–1850s publications), introducing subgeneric divisions based on elytral punctation and pronotal features, which laid groundwork for recognizing diverse lineages within the genus. In the , revisions by Severin Breuning in his comprehensive Monographie der Gattung Carabus (1932–1937) synthesized prior descriptions, incorporating around 50 subgenera while focusing on Palearctic species. Subsequent updates, such as those by Michael Luff and others, refined these groupings amid growing collections. Post-2000 phylogenetic studies integrating molecular data, including mitochondrial ND5 sequences (Imura et al., 2003) and multi-locus analyses (Deuve et al., 2012), prompted major reevaluations, confirming approximately 91 monophyletic subgenera through evidence of evolutionary divergences. Recent advances as of 2024, using museomics and population genomics, further support an early Eocene origin and radiations in alpine and Himalayan regions. These debates highlighted shifts from morphology-driven hierarchies to phylogenetically informed ones, resolving longstanding uncertainties in subgeneric boundaries.

Physical description

Morphology

Carabus beetles exhibit a robust, elongated oval body shape, typically measuring 12–50 mm in length, which supports their terrestrial predatory . The overall form is dorsoventrally flattened, with a hardened providing protection against environmental hazards and predators. Key morphological features include the elytra, which are hardened forewings that cover and protect the and hindwings; these are often metallic or iridescent, displaying colors such as green, blue, or copper that arise from structural interference in the layers. Most are brachypterous, possessing reduced or absent hindwings, which limits flight capability and emphasizes ground-dwelling adaptations. The mandibles are powerful and sickle-shaped, enabling effective predation on soft-bodied , while the pronotum—the dorsal plate of the —features distinct lateral grooves that contribute to structural rigidity. The head is large and prognathous, equipped with prominent compound eyes for detecting movement and filiform (thread-like) antennae consisting of 11 segments for sensory perception. The thorax bears three pairs of long, legs adapted for rapid running on the ground surface, with segmented tarsi comprising five articles. The abdomen comprises multiple sclerotized tergites and sternites, covered dorsally by the elytra when at rest, and features coloration ranging from matte black to vibrant metallics, serving roles in against soil or leaf litter and potential aposematic warning signals to deter attackers. These powerful mandibles allow Carabus to tackle prey such as snails by crushing shells.

Intraspecific variation

Intraspecific variation within the genus Carabus manifests prominently through and geographic morphs, reflecting adaptations to mating, reproduction, and environmental pressures. Males typically exhibit enlarged fore-tarsi, which are dilated and equipped with adhesive structures to facilitate grasping females during copulation. In contrast, females often display broader and more elongated abdomens, enabling greater capacity for development and laying. Subtle differences in elytral sculpture, such as variations in ridge patterns or microsculpture density, have also been observed between sexes, potentially linked to or structural reinforcement. Geographic and habitat-related variation further diversifies Carabus morphology, particularly in body size and coloration. Across many species, body size exhibits a latitudinal cline, with individuals decreasing in size toward higher latitudes; for instance, Carabus populations in southern regions tend to be larger than those in northern areas, possibly due to climatic influences on growth rates. Color morphs show habitat-specific patterns, where melanic (dark, non-reflective) forms predominate in shaded, forested environments for crypsis against predators, while metallic, iridescent variants are more common in open, sunny habitats, enhancing thermoregulation or signaling. These variations often correlate with local environmental gradients, such as altitude or vegetation cover, contributing to adaptive divergence within species. The extensive subspecific diversity in Carabus, with numerous described across its range, underscores the role of geographic isolation in driving intraspecific differentiation. For example, in Carabus auratus, the nominate subspecies C. a. auratus exhibits a golden metallic sheen typical of central European populations, differing from regional variants like C. a. lotharingus in eastern and , which show subtle shifts in coloration and sculpture due to and reduced . Such isolation, often promoted by landscape barriers like rivers or mountains, fosters localized adaptations and has led to the recognition of numerous in European C. auratus, highlighting the genus's propensity for rapid morphological .

Distribution and habitat

Geographic range

The genus Carabus exhibits a predominantly Palearctic distribution, with the highest species diversity concentrated in temperate regions of , encompassing over 900 species across , , , , the , , and Mediterranean islands. In the Nearctic region, only about 12 native species occur, primarily as relict populations in northern , reflecting ancient dispersals via Beringian land bridges during the . Additionally, several Palearctic species have been introduced to , bringing the total to approximately 16 species, though these introductions are mostly adventive and not fully naturalized. The expansion of Carabus species has been shaped by post-glacial recolonization patterns following the , with populations radiating northward from multiple refugia in (such as the Iberian, Italian, and Balkan peninsulas) and southern . This recolonization was facilitated by warming climates and the retreat of ice sheets, allowing low-dispersal, often flightless species to repopulate temperate zones at rates limited by their brachypterous morphology. The genus is confined to the , with no native occurrences in tropical regions, , or , due to historical barriers like the Turgai Sea and climatic unsuitability for their cold-adapted ecologies. Endemism in Carabus is pronounced in montane habitats, driven by the genus's limited dispersal capabilities and isolation in heterogeneous highland environments; for instance, numerous species are endemic to the and in , while in , local endemics persist in the Himalayan-Tibetan orogen, including central Tibetan highlands. Recent human-mediated introductions, such as Carabus granulatus via shipping ballast and nursery stock from since the late , have expanded the genus's footprint in , particularly along northern coastlines and waterways in states like , , and .

Habitat preferences

Carabus species predominantly inhabit moist, temperate environments such as forests, meadows, and wetlands, where they favor soil-rich areas providing ample leaf litter or cover for concealment and foraging. These beetles exhibit a strong preference for habitats with high humidity and moderate shade, including forests and edges, which support their predatory lifestyle by maintaining stable microclimates and prey availability. For instance, species like Carabus granulatus are hygrophilous and eurytopic, thriving in both open and shaded moist settings from lowlands to montane regions. Within these environments, Carabus beetles are primarily nocturnal, seeking refuge during the day in microhabitats such as under stones, logs, loose bark, or within soil crevices to avoid and predators. Some , such as Carabus variolosus, occupy specialized niches like peat bogs or alpine zones with high , while the generally avoids arid deserts or heavily disturbed agricultural lands lacking structural cover. This habitat selectivity is influenced by physiological adaptations to gradients, with many showing reduced activity in dry conditions. Adaptations to these habitats include burrowing behaviors for overwintering, where adults and larvae excavate in well-aerated, soils to endure periods, enhancing survival in temperate climates. Preference for soils high in not only facilitates burrowing and oviposition but also sustains abundant prey, underscoring the genus's reliance on nutrient-dense, undisturbed substrates. Such traits contribute to their role as indicators of habitat quality in and ecosystems.

Ecology and behavior

Diet and feeding

Carabus beetles are predominantly nocturnal predators that primarily consume live prey such as slugs, snails, earthworms, caterpillars, and other , reflecting their role as generalist carnivores in terrestrial ecosystems. These exhibit polyphagous feeding habits, with a marked preference for soft-bodied , though they occasionally scavenge carrion, including portions of dead vertebrates when available. generally provide high nutritional value, while earthworms serve as an intermediate food source, and slugs rank lower in dietary quality for like Carabus nemoralis. Feeding adaptations in Carabus are particularly specialized for consuming mollusks, with two distinct morphological strategies observed among snail-feeding species. Macrocephalic forms, characterized by enlarged heads and robust mandibles, crush snail shells directly to access the soft tissues within. In contrast, stenocephalic species possess slender heads and elongated mouthparts, enabling them to insert into shell apertures and extract prey without breaking the shell, a strategy suited to smaller or more protected snails. Complementing these structures, Carabus employ powerful jaws to inject salivary fluids rich in , facilitating extra-oral digestion that liquefies prey tissues for easier consumption. Foraging in Carabus involves active nocturnal hunting, where individuals patrol leaf litter and surfaces under cover of to locate prey. They rely on chemoreceptors located on antennae and palps to detect chemical cues from potential prey, enabling targeted tracking rather than random searching, which enhances efficiency in detecting soft-bodied . This behavior aligns with prey availability in moist, vegetated habitats, where such food sources are abundant.

Life cycle and reproduction

Carabus species exhibit holometabolous , progressing through distinct , larval, pupal, and stages. Eggs are typically laid singly by s in small chambers excavated in moist , with each female producing a variable number of eggs, ranging from a few to several hundred over the reproductive season depending on , environmental conditions, and reproductive . The larvae are campodeiform, characterized by an elongated, flattened body with well-developed thoracic legs and urogomphi; they undergo three s, during which they remain predatory, feeding on small while burrowing in . Pupation occurs in a soil chamber formed by the final instar larva, lasting several days to weeks before the emergence of the . Reproduction in Carabus is generally univoltine, with one per year in most , though timing varies: spring breeders mate from late winter to early summer, while autumn breeders do so in late summer or fall. often involves pheromonal attraction, where both sexes release chemical signals to locate partners, facilitating in suitable habitats. Females select oviposition sites based on and prey availability, laying eggs over several weeks. longevity spans 1–3 years, allowing iteroparity in many cases, where individuals reproduce in multiple seasons. Population dynamics are shaped by overwintering strategies and high juvenile mortality. Many species overwinter as late-instar larvae or diapausing adults in refugia, emerging in spring to reproduce or . Juvenile stages face high mortality due to predation by vertebrates and , as well as in dry conditions, which particularly affects third-instar larvae and pupae. These factors contribute to low recruitment rates, with population stability maintained through adult longevity and environmental cues regulating reproductive output.

Diversity and species

Species diversity

The genus Carabus encompasses approximately 970–1,000 and more than 2300 subspecies, organized into about 90–94 subgenera, rendering it the most speciose genus in the subfamily Carabinae. This extraordinary diversity underscores the genus's evolutionary success within the Carabidae family, with the vast majority of taxa concentrated in the Holarctic region, particularly exhibiting peak richness in and . In , approximately 132 are documented, while East Asia hosts even greater numbers, reflecting adaptations to varied temperate and montane environments. Subgenera in Carabus are delineated primarily through morphological traits, such as elytral sculpture and genital structures, supplemented by phylogenetic analyses that reveal evolutionary lineages. These groupings facilitate taxonomic organization amid the genus's complexity; for instance, supports 14 species across 9 subgenera, including 3 adventive introductions from that have established populations. Such regional patterns highlight how historical dispersal and local adaptation contribute to subgeneric variation. Conservation challenges threaten this diversity, with several Carabus species classified as endangered owing to habitat fragmentation from urbanization, agriculture, and forestry practices. Notable examples include Carabus nitens, vulnerable in Central European heathlands due to shrinking open habitats, and Carabus variolosus, endangered in fragmented montane wetlands of Central and Southeastern Europe due to and dispersal barriers. Fossil records include extinct Carabus species from various geological periods, illustrating the genus's vulnerability to past climatic shifts and the need for ongoing protection of extant taxa.

Notable species

Carabus auratus, commonly known as the , is a striking species characterized by its metallic green coloration and body length of 20-30 mm. Native to and introduced to , it is widespread in gardens, fields, and woodlands where it preys on slugs, , caterpillars, and other pests, including the . This predatory behavior makes C. auratus valuable in biological control, particularly against invasive slugs, and its iridescent appearance also contributes to its ornamental appeal in natural settings. Carabus nemoralis, the bronze , measures 20-25 mm and exhibits variable bronze-black coloration with purplish iridescence on the pronotum and elytra margins. It is commonly found in European forests and agricultural areas, where it nocturnally hunts pests such as the invasive slug Arion vulgaris. Recent studies have explored its potential as a biocontrol agent in large-scale farming to manage multiple invertebrate pests. Carabus glabratus, or the smooth , is a large (25-32 mm), dull metallic black species with subtle blue flashes, inhabiting montane bogs and damp hummocks across Central and , extending to the . It is considered rare in northwestern Europe due to habitat loss in , serving as an indicator of bog . As a specialized wetland predator, its populations have declined with the degradation of bogs, highlighting its vulnerability. Carabus variolosus is a flightless, hygrophilous species endemic to mountain regions of Central and Southeastern , including the Carpathians and , where it prefers moist alpine habitats at high altitudes. Reaching up to 30 mm, it is stenotopic and endangered in certain populations due to sensitivity to changes and climate shifts. This acts as a for montane integrity, with its restricted range underscoring threats from . The extinct Carabus jeffersoni, a fossil species from the Eocene Florissant Formation in Colorado, USA, represents an early member of the genus known only from paleontological records dating to approximately 37 million years ago. Named in honor of , it provides insights into the ancient diversity of Carabus in but has no modern ecological significance. These notable species exemplify the genus's roles in pest control—particularly C. auratus and C. nemoralis against agricultural threats—ornamental value through vivid morphologies, and as indicators of habitat health in sensitive environments like bogs and mountains.

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