Copris Geoffroy 1762

Copris Geoffroy Geoffroy 1762: 87; type sp. Scarabaeus lunaris Linnaeus, 1758. – Insulicopris Tsukamoto 1974: 1; type sp. Copris brachypterus Nomura, 1964. – Pilularius Schrank 1798: 394; type sp. Pilularius lunus Schrank, 1798 (= Scarabaeus lunaris Linnaeus, 1758). – Palaeocopris Pierce 1946: 130; type sp. Palaeocopris labreae Pierce, 1946 = Phanaeus labreae (Pierce) [late Pleistocene of California, see Miller et al. 1981: 627, Edmonds 1994: 91]. Species-group taxa – achates Péringuey 1901: 346, 365; = cornifrons – acuticeps Felsche; see Coptodactyla – acuticollis (Motsch.) of Kabakov 2006: 87 [key]; = acutidens acutidens Motschulsky 1860: 13; China (Fujian, Hubei, Sichuan), Japan. afgoi Cambefort 1992b: 181; Somalia. agnus Sharp 1875: 47; Borneo, Java, Malaysia, Singapore. algol Nguyen-Phung 1989: 202; Kenya, Tanzania. – amabile Ferreira 1958a: 3; = amabilis Kolbe amabilis Kolbe 1914: 311; DRC, Uganda. – ammon (Fabricius 1781: 24), as Scarabaeus; = minutus amyntor Klug 1855: 655; Kenya, Mozambique, RSA, Zimbabwe. – amyntor Harold 1869a: 61; = harrisi – anaglypticus Say 1823: 204; = fricator fricator anceus (Olivier 1789: 116), as Scarabaeus; RSA (Cape). andrewesi (Waterhouse); see Paracopris * angolensis Ferreira 1962b: 144; ssp. of cassius angusticornis Arrow 1933: 427; China (Fujian, Hubei, Sichuan, Yunnan), Thailand, Vietnam. * angusticornis Nguyen-Phung and Cambefort; ssp. of laius angustus Nguyen-phung and Cambefort 1986: 407; Benin, Ivory Coast, Mali, Senegal. anomiopseoides (Boucomont); see Paracopris antares Ferreira 1958a: 5; RSA (Free State). apicepunctatus Balthasar; see Microcopris arcturus Gillet 1907: 179; DRC, Gabon. ares Zelenka 1993: 242; RSA. arimotoi Ochi and Araya 1992: 83; Indonesia (Sumatra). arizonensis Schaeffer 1906: 254, 256; Mexico (Chihuahua), USA (Arizona, New Mexico, Texas). armatus Harold 1869c: 498; Mexico (DF, Guerrero, Hidalgo, Jalisco, México, Michoacán). armeniacus Faldermann 1835: 238; Armenia, Azerbaijan, Georgia, Iran, Turkey. [sp. inquirenda in Ziani 2017] armiger Gillet 1910: 1; Burundi, DRC, Kenya, Mozambique, RSA (Transvaal), Tanzania, Zambia, Zimbabwe. arrowi Felsche 1910: 350; “South China ”, Thailand. aspericollis Gillet 1910: 3; Costa Rica, El Salvador, Guatemala, Mexico (Chiapas). atropolitus Gillet 1910: 7; DRC, Kenya, Tanzania. – baileyi Blackburn; see Coptodactyla barclayi Ochi, Kon and Bai; see Sinocopris bartolozzii (Ochi, Kon and Pham); see Paracopris basilewskyi Ferreira 1962a: 101; DRC. basipunctatus (Balthasar); see Sinocopris – belisama (Schrank 1798: 394), as Pilularius; = lunaris bellator Chevrolat in Guérin-Méneville 1844: 81; Indonesia (Java), Malaysia. bengalensis Gillet 1911a: 289; China (Guizhou, Xizang, Yunnan), Myanmar, Thailand. bidens Kolbe 1893: 200; Cameroon. – bifidus (Poda von Neuhaus 1761: 18), as Scarabaeus; = lunaris – bihamatus Balthasar 1965: 20, as Paracopris; = fidius bootes Klug 1855: 655; Botswana, Kenya, Mozambique, RSA (Transvaal), Tanzania. * borneensis Ochi and Kon; ssp. of gibbulus – bornemisszai Ferreira 1972: 167, 783; = bootes boucardi Harold 1869c: 497; El Salvador, Guatemala, Mexico (Guerrero, Oaxaca). bovinus Gillet 1908 a: 68; Angola, DRC. brachypterus brachypterus Nomura 1964: 50; Japan (Okinawa: Amami-Ôshima). brachypterus toguchii Ochi and Kawahara 2012: i; Japan (Tokunoshima). brittoni Ferreira 1962a: 103; DRC, Tanzania. brooksi Matthews; see Coptodactyla caelatus Fabricius 1794: 435; Lesotho, Mozambique, RSA. * caffer Gillet; ssp. of orion caliginosus Kohlmann, Cano and Delgado 2003: 2; Guatemala. – calvus Sharp 1875: 46; = saundersi cambeforti Nguyen-Phung 1988c: 12; RSA (Transvaal). cambodiensis Ochi, Kon and Kawahara 2008: 243; Cambodia. camerunus Felsche 1904: 402; Cameroon. capensis Waterhouse 1891a: 513; Lesotho, RSA (Cape). cariniceps (Felsche); see Paracopris carinicus Gillet 1910: 8; China (Yunnan), India (Assam), Laos, Myanmar, Thailand. carmelita Fabricius 1801: 39; Benin, Burkina Faso, DRC, Gambia, Guinea, Sierra Leone, Senegal, Uganda. * cartwrighti Robinson 1941: 131; = ssp. of fricator cassius Péringuey 1901: 344, 351; Angola, Botswana, Mozambique, RSA (Cape), Zimbabwe. – castaneus Mulsant 1842: 73, as var. of C. lunaris; = lunaris * cavolini (Petagna); ssp. of hispanus [Prevailing usage of the subsequent spelling cavolinii has not been officially demonstrated; S. Ziani and T. Branco, pers. comm.] celebensis Lansberge 1886: 9; Indonesia (Sulawesi). * centralis Gillet; ssp. of orion cheni (Ochi, Kon and Bai); see Sinocopris chinmontanus (Ochi, Kon and Kawahara); see Paracopris * chuduraziensis Ochi, Kon and Kawahara; ssp. of pseudochus; see Sinocopris – ciprianii Balthasar 1939a: 3, as Paracopris; = coriarius – claudius (Harold 1877a: 48), as Catharsius; = repertus – clavicornis Matthews and Halffter 1959: 191; = moechus colmanti Gillet 1908 b: 56; Cameroon, DRC, Gabon. complexus Nguyen-Phung 1989: 206; Burundi, Cameroon, DRC, Kenya, Malawi, Rwanda, Tanzania, Uganda. compressipennis (Gillet); see Paracopris confucius Harold 1877a: 48; Cambodia, China (Fujian, Guangdong, Guizhou, Hongkong, Sichuan, Yunnan), Laos, Malaysia, Myanmar, Taiwan, Thailand, Vietnam. – confusus Boheman 1857: 240; = amyntor – consobrinus (Lansberge); see Microcopris * continentalis Ochi and Kon; ssp. of pecuarius; see Sinocopris – contractus Boheman 1857: 236; = caelatus – contractus Péringuey 1901: 344, 352; = obesus convexiusculus Shipp 1897: 450; Somaliland. coriarius Gillet 1907: 178; DRC, Eritrea, Ethiopia, Guinea, Ivory Coast, Malawi, Nigeria, RSA, Tanzania, Uganda, Zimbabwe [var. of mesacanthus in Ferreira 1961: 156]. – corniculatus Mulsant 1842: 73, as ab. of lunaris; = lunaris cornifrons Boheman 1860: 110; Botswana, Mozambique, RSA (Cape). corniger Sahlberg 1823: 6; Ethiopia, Kenya, Lesotho, RSA. – corniger Harold 1869c: 1014; = bellator – coronatus (Harold 1877b: 98), as Catharsius; = signatus corpulentus Gillet 1910: 13; China (Fujian, Sichuan, Yunnan), India (Arunachal Pradesh), Laos, Myanmar, Thailand, Vietnam. costaricensis costaricensis Gahan 1894: 116; Costa Rica, Mexico (Chiapas), Panama. costaricensis dolichocerus Matthews 1961: 97; Guatemala, Mexico (Chiapas). cribratus (Gillet); see Paracopris cribricollis Gillet 1910: 6; Tanzania. – curvicornis Boheman 1860: 110; = cornifrons ♀ – davidis (Deyrolle 1878: 95), as Catharsius; = Catharsius molossus (syn. Cheung et al. 2018) davisi Nguyen-Phung and Cambefort 1986b: 104; Angola. davisoni (Waterhouse); see Paracopris decellei Cambefort 1992a: 29; DRC, Guinea, Rwanda, Uganda. – deletus Mulsant 1842: 73, as var. of lunaris; = lunaris delicatus Arrow 1931: 117; China (Yunnan), India (Arunachal Pradesh, Assam, Sikkim). denticulatus Nguyen-Phung 1988b: 216; Mozambique, RSA, Zimbabwe. depressa Paulian; see Coptodactyla * dicyrtus Matthews and Halffter; ssp. of remotus diversus Waterhouse 1891a: 518; Kenya, Somalia, Sudan, Tanzania, Madagascar; introduced into Australia (Queensland, NT, WA). * dolichocerus Matthews; ssp. of costaricensis doriae (Harold); see Microcopris draco Arrow 1906: 127; Angola. dracunculus Ferreira 1959: 13; DRC. ducalis Blackburn; see Coptodactyla dudleyi Cambefort 1992a: 30; Malawi. eburneus Cambefort 1992a: 28; Ivory Coast. elpheneroides Felsche 1910: 345; Tanzania. elphenor Klug 1855: 655; Angola, Burkina Faso, Botswana, Cameroon, DRC, Ethiopia, Kenya, Mozambique, Namibia, RSA, Tanzania, Zimbabwe; introduced into Australia (Queensland). – emarginatus (Olivier 1789: 115), as Scarabaeus; = lunaris erratus erratus Lansberge 1886: 13; Sulawesi. erratus masayukii Ochi and Kon 2004a: 28; Indonesia (Sulawesi: Peleng I.). evanidus Klug 1855: 655; Botswana, Ethiopia, Kenya, Mozambique, RSA, Tanzania, Uganda, Zimbabwe. – excavatus Klug 1855: 655; = bootes excisus (Waterhouse); see Paracopris fallaciosus Gillet 1907: 180; Botswana, Ethiopia, Kenya, Mozambique, Namibia, RSA, Somalia, Uganda, Zimbabwe; introduced into Australia (Queensland). fallax Felsche 1910: 349; Namibia. felschei Reitter 1892: 216 [= 1893: 95]; Armenia, Azerbaijan, Georgia, Iran, e. Turkey. – ferreirae Frey 1974: 273; = megaceratoides fidius (Olivier 1789: 184), as Scarabaeus; DRC, Mozambique, Namibia, RSA. * formosanus Gillet; ssp. of punctatus – frankenbergeri Balthasar 1934: 149; = acutidens frankenbergerianus Balthasar 1958: 477; China (Sichuan). freyi Ferreira 1961a: 138; Gabon. fricator fricator (Fabricius 1787: 15), as Scarabaeus; India, Sri Lanka; introduced into much of e.+c. USA and se. Canada (Ontario). fricator cartwrighti Robinson 1941: 131, as sp.; USA (South Carolina). – fricator Harold 1877a: 46; = erratus fujiokai fujiokai Ochi and Kon; see Microcopris fujiokai poringensis Ochi and Kon; see Microcopris fukiensis Balthasar 1953: 225; China (Fujian, Zhejiang), Taiwan. furciceps (Felsche); see Paracopris – furcillatus Felsche 1910: 345; = costaricensis * gabonicus Cambefort; ssp. of phungae garambae Cambefort 1992a: 27; DRC gazellarum Gillet 1918: 148; Burkina Faso, DRC, Sudan, Uganda. * genopunctatus Nguyen-Phung; ssp. of vankhaii gibbulus gibbulus Lansberge 1886: 17; Malaysia, Borneo, Java, Sumatra. gibbulus borneensis Ochi and Kon 2005: 71; Malaysia (Sabah). gilleti Kolbe 1907: 25; Tanzania. – gistelianus Gistel 1857: 587; = lunaris –glabriceps Felsche 1910: 346; = acutidens glabricollis (Hope); see Coptodactyla gladiator Arrow 1933: 425; Myanmar. – globulipennis Waterhouse 1891a: 512; = sexdentatus Thunberg gopheri Hubbard 1894: 310; USA (Florida). gracilis Waterhouse 1891a: 518; Namibia. * grandis Nguyen-Phung and Cambefort; ssp. of inhalatus graueri Kolbe 1914: 312; Tanzania. grossa (Thomson); see Pseudopedaria halffteri Matthews 1959: 133; Mexico (Guerrero, Morelos, Puebla, San Luis Potosí). – haroldi Lansberge 1886: 15; = spinator harrisi Waterhouse 1891a: 515; Benin, CAR, DRC, Eritrea, Ethiopia, Kenya, Sierra Leone, Somalia, Sudan, Tanzania, Uganda, Zimbabwe. hartli Frey 1975: 294; Tanzania. hatayamai Ochi and Araya 1992: 79; Malaysia. hidakai Ochi and Kon; s

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