Copris Geoffroy, 1762

Zidek, Jiri, 2020, Catalogue of species-group names assigned to Copris Geoffroy, Coptodactyla Burmeister, Litocopris Waterhouse, Microcopris Balthasar, Paracopris Balthasar, Pseudocopris Ferreira, Pseudopedaria Felsche, Sinocopris Ochi, Kon and Bai, and Thyregis Blackburn (Coleoptera: Scarabaeidae: Scarabaeinae: Coprini), Insecta Mundi 756, pp. 1-24 : 3-12

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 ; see Microcopris

– hintoni Saylor 1933: 238 ; = klugi

– hispanicus ( Poiret 1789: 297) , as Scarabaeus ; = hispanus hispanus

hispanus hispanus ( Linnaeus 1764: 12) , as Scarabaeus ; western ssp.: s. France incl. Corsica, Sardinia ( Italy), Iberian peninsula, Morocco, Tunisia; introduced into Australia (WA).

hispanus cavolini ( Petagna 1792: 140) , as sp. of Scarabaeus ; eastern ssp.: continental Italy + Sicily, Balkans, Turkey, Israel, n. Iran, c. Asia. [For C. hispanus and C. cavolini as separate species see Falahee and Angus 2010.]

– hispanus persianus Mikšić 1958: 1 ; = hispanus cavolini

hosakai Ochi and Kon ; see Microcopris

howdeni Matthews and Halffter 1959: 200 ; USA (Florida, Georgia?, South Carolina).

humilis Gillet 1908 a: 71 ; Tanzania.

hybridus Gillet 1910: 11 ; Tanzania.

igualensis Warner 1990: 236 ; Mexico (Guerrero).

illotus Balthasar 1942b: 189 ; China (Fujian).

imitans (Felsche) ; see Paracopris

– imitatrix Balthasar ; see Coptodactyla

inaequabilis Zhang 1997: 758 ; China (Hubei).

incertus Say 1835: 175 ; Colombia, Ecuador, Mexico (Hidalgo, Puebla, San Luis Potosí, Tamaulipas, Yucatán), Panama; introduced into Hawaii, New Zealand, Vanuatu.

– indicus Gillet 1910: 5 ; = fricator fricator

inemarginatus Blatchley 1918: 54 ; USA (Florida).

inhalatus inhalatus Quedenfeldt 1884: 272 ; Angola, DRC, Mozambique, Namibia, RSA (Cape), Zimbabwe.

inhalatus grandis Nguyen-Phung and Cambefort 1986b: 102 ; Zimbabwe.

inhalatus perturbator Péringuey 1901: 352 , as sp.; Botswana, Namibia, RSA, Zimbabwe.

inhalatus sanctaeluciae Nguyen-Phung and Cambefort 1986b:102; RSA (Natal).

insidiosus Péringuey 1901: 346 , 366; Burundi, DRC, Rwanda, Tanzania, Uganda, Zimbabwe.

integer Reiche 1850: 312 ; Burkina Faso, Burundi, DRC, Ethiopia, Kenya, Rwanda, Tanzania, Zimbabwe.

– integer Harold 1869a: 61 ; = fallaciosus

interioris Kolbe 1897: 143 ; Burundi, Cameroon, DRC, Gambia, Guinea, Nigeria, Senegal, Uganda, Tanzania.

iris Sharp 1875: 45 ; Laos, Malaysia, Myanmar, Thailand, Vietnam.

* isthmiensis Matthews; ssp. of lecontei

jacchoides Nguyen-Phung and Cambefort 1987: 70 ; Lesotho, RSA.

jacchus ( Fabricius 1775: 20) , as Scarabaeus ; Lesotho, Namibia, RSA, Zimbabwe.

jahi Nguyen-Phung and Cambefort 1986a: 413 ; Cameroon.

javanensis (Bui, Ziegler and Bonkowski) ; see Paracopris

– jenisonianus Gistel 1857: 586 ; = lunaris

jucundus Gillet 1933: 326 ; Cameroon, Guinea.

kachinensis Ochi, Kon and Kawahara 2008: 245 ; Myanmar.

kasagii (Ochi and Kon) ; see Paracopris

katangae Cambefort and Nguyen-Phung 1996: 286 ; DRC.

keralensis Gill 1986: 632 ; India (Kerala).

kershawi Blackburn ; see Thyregis

kiuchii (Masumoto) ; see Sinocopris

klapperichi Balthasar 1942a: 116 ; China (Fujian).

klugi klugi Harold 1869c: 498 ; Mexico (most states), Central America.

klugi sierrensis Matthews 1961: 72 ; Mexico.

kusakabei (Ochi, Kon and Kawahara) ; see Sinocopris

laeviceps Harold 1869c: 498 ; Belize, Colombia, Costa Rica, Guatemala, Honduras, Mexico (Chiapas, DF, Oaxaca, Veracruz).

– laevicollis Mulsant 1842: 69 , as var. of hispanus ; = hispanus hispanus

laevigatus (Gillet) ; see Sinocopris

laiiformis Nguyen-Phung and Cambefort 1987: 72 ; Zimbabwe.

laioides Boucomont 1932: 46 ; DRC, Namibia, Zambia, Zimbabwe.

laius laius Harold 1868a: 81 ; Burkina Faso, DRC, Gambia, Guinea Bissau, Senegal, Sudan.

laius angusticornis Nguyen-phung and Cambefort 1987: 73 ; Cameroon, Chad, Sudan, Uganda.

lannathai Hanboonsong, Masumoto and Ochi 2003: 108 ; Thailand.

– lar ( Fabricius 1787: 13) , as Scarabaeus ; = minutus

– laticornis Boheman 1857: 226 ; = elphenor

– laticornis Wallengren 1881: 20 ; = jacchus

– laticornis Péringuey 1901: 343 , 347; = corniger , fallaciosus

lecontei lecontei Matthews 1961: 98 ; USA (Arizona, New Mexico), Mexico (Chihuahua, Colima, Guerrero, Jalisco, Nayarit, Oaxaca, Sinaloa, Sonora).

lecontei isthmiensis Matthews 1961: 102; Mexico.

lesnei Paulian ; see Coptodactyla

– lineatus Wiedemann 1823: 9 ; = sexdentatus

lugubris Boheman 1858: 42 ; Belize, Costa Rica, El Salvador, Guatemala, Honduras, Mexico (most states), Nicaragua, Panama.

– lunarioides Waterhouse 1891a: 513 ; = corniger

lunaris ( Linnaeus 1758: 346) , as Scarabaeus ; UK, s. Sweden, c.+s.+e. Europe, w. China, w. Siberia, Turkey, Syria, Iran, c. Asia, Kazakhstan; introduced to Australia.

lunaticeps (Gillet) ; see Pseudocopris

– lunus ( Schrank 1798: 393) , as Pilularius ; = lunaris

macacus Lansberge 1886: 14 ; Indonesia (Sulawesi).

macclevei Warner 1990: 232 ; Mexico (Sonora), USA (Arizona).

macer Péringuey 1901: 343 , 346; Angola, Mozambique, Namibia, RSA (Transvaal), Swaziland, Tanzania, Zimbabwe.

maesi Ratcliffe 1998: 93 ; Nicaragua.

magicus Harold 1881: 88 ; China (Guizhou, Hubei, Hunan, Sichuan, Xinjiang, Yunnan), India (Sikkim), Laos, Myanmar, Thailand, Vietnam.

manni (Ochi, Kon and Bai); see Sinocopris marcoberrai Ochi, Kon and Pham 2019: 435; c. Vietnam (Quang Nam).

– marcus (Gillet) ; see Microcopris

– marianus ( Petiver 1764: 27) , as Scarabaeus ; = hispanus hispanus martinae Nguyen-Phung 1988a: 257 ; Tanzania. martinezi Matthews and Halffter 1968: 154 ; Mexico (Durango).

* masayukii Ochi and Kon; ssp. of erratus

matthewsi Reid ; see Coptodactyla

matthewsi matthewsi Delgado and Kohlmann 2001: 344 ; Guatemala, Mexico (Chiapas).

matthewsi pacificus Delgado and Kohlmann 2001: 348 ; El Salvador, Guatemala, Mexico.

medogensis Zhang 1988: 228 ; China (Xizang).

megaceratoides Waterhouse 1891a: 511 ; DRC, Gambia, Guinea, Nigeria, Senegal.

megasoma Matthews and Halffter 1959: 194 ; Mexico (Durango, Michoacán).

merdeka Reid ; see Coptodactyla

meridionalis Matthews ; see Coptodactyla

mesacanthus mesacanthus Harold 1878: 45 ; Angola, DRC, Malawi, Mozambique, Namibia, RSA, Zambia, Zimbabwe.

mesacanthus transvaalensis Nguyen-Phung 1988c: 10 ; RSA.

mexicanus Matthews and Halffter 1959: 194 ; Mexico (Michoacán).

– minator Harold 1880: 152 ; = anceus [valid in Gillet 1910] mindorensis (Ochi) ; see Microcopris

– minutissimus (Kryzhanovskii and Medvedev) ; see Microcopris minutus (Drury 1770: 78) , as Scarabaeus ; e.+c. USA. misellus Péringuey 1901: 344 , 349; Botswana, DRC, Mozambique, Namibia, RSA, Zambia, Zimbabwe.

– modestus Boheman 1857: 242 ; = evanidus moechus LeConte 1854: 222 ; Mexico (Chihuahua, Durango), USA (Texas). moffartsi Gillet 1907: 177 ; Burundi, Cameroon, DRC, Kenya, Rwanda, Uganda. mongkhoni Hanboonsong, Masumoto and Ochi 2003: 114 ; Thailand. monstrosa Felsche ; see Coptodactyla monteithi Matthews ; see Thyregis

* montivagus Gillet 1908 a: 80 , as var. of harrisi [valid sp. in Gillet 1910, var. of harrisi in Ferreira 1961: 209 ]. montreuili (Ochi, Kon and Pham) ; see Paracopris

– morgani Waterhouse 1891a: 514 ; = carmelita morphaeus Gillet 1932: 325 ; Kenya. mourgliai Zelenka 1993: 239 ; Somalia.

– muelleri Balthasar 1939b: 42 ; = acutidens muticus (Boheman) ; see Litocopris

– muticus (Péringuey) ; see Litocopris neglectus Moxey 1963: 25 ; India (Tamil Nadu). nepos Gillet 1908 a: 76 ; Burundi, DRC, Kenya, Rwanda, Tanzania, Uganda, Zimbabwe.

– nepticulus Kolbe 1897: 144 ; = diversus

– neptis Reiche 1850: 314 ; = orphanus Guérin-Méneville nevinsoni Waterhouse 1891a: 519 ; Cambodia, Laos, Malaysia, Thailand, Vietnam. nitida Paulian ; see Coptodactyla novaki Zelenka 1992: 167 ; Malaysia. nubilosus Kohlmann, Cano and Delgado 2003: 4 ; Guatemala, Honduras. numa Lansberge 1886: 19 ; India (Assam), Indonesia (Borneo, Sumatra), Malaysia, Myanmar. obenbergeri (Balthasar) ; see Sinocopris obesus Boheman 1857: 237 ; Lesotho, Mozambique, RSA, Tanzania, Zimbabwe.

– obliteratus Mulsant 1842: 73 , as var. of lunaris ; = lunaris ochus (Motschulsky) ; see Sinocopris

– oedipus ( Fabricius 1775: 20) , as Scarabaeus ; = jacchus

olivieri (Ochi, Kon and Kawahara) ; see Paracopris

onitoides Gillet ; see Coptodactyla

– orientalis Fabricius 1792: 52 ; = repertus

orion orion Klug 1835: 34 ; Angola, Burkina Faso, Burundi, Cameroon, DRC, Gambia, Kenya, Mozambique, Namibia, RSA, Rwanda, Senegal, Tanzania.

orion caffer Gillet 1908 a: 70 , as var.; RSA.

orion centralis Gillet 1908 a: 70, as var.; DRC.

orion overlaeti ( Balthasar 1961: 98), as sp.; DRC, Gabon, Rwanda (?).

orphanus Guérin-Méneville 1847: 308 ; Angola, Burundi, DRC, Ethiopia, Guinea, Kenya, Rwanda, Tanzania, Uganda.

– orphanus Péringuey 1901: 344 , 350; = evanidus

– orphanus Ferreira 1958b: 501 ; = gracilis

– oryctes (Herbst) ; see Microcopris

* overlaeti (Balthasar); ssp. of orion

* pacificus Delgado and Kohlmann ; ssp. of matthewsi

– paniscus ( Fabricius 1775: 123) , as Scarabaeus ; = hispanus hispanus

papua Lansberge ; see Coptodactyla

parapecuarius (Ochi, Kon and Kawahara) ; see Sinocopris

parasonensis Ochi, Kon and Pham 2019: 437; Laos (Hua Phan).

pauliani Nguyen-Phung and Cambefort 1986a: 413 ; Cameroon.

– pauliani (Frey) ; see Thyregis

pecuarius pecuarius (Lewis) ; see Sinocopris

pecuarius continentalis Ochi and Kon; see Sinocopris

pedarioides (Lansberge) ; see Paracopris

– persianus Mikšić 1958:1 ; = hispanus cavolini

– persimilis Felsche 1908: 273 ; = fallaciosus

* perturbator Péringuey ; ssp. of inhalatus

phungae phungae Cambefort 1992b: 181 ; Gabon, Ivory Coast.

phungae gabonicus Cambefort 1992b: 182; Gabon.

phylax Gillet 1908 b: 59 ; Cameroon, DRC, Ivory Coast.

– pluridens Gillet 1908 a: 70 ; = interioris

– plutus ( Fabricius 1794: 436) , as Scarabaeus ; = fidius

poggii Ochi and Kon 2005: 69 ; Malaysia (Sabah).

– pompilius Waterhouse 1875: 75 ; = sarpedon

popaensis (Ochi, Kon and Kawahara); see Paracopris

* poringensis Ochi and Kon; ssp. of fujiokai ; see Microcopris

potanini Semenov 1891: 310 ; China (Gansu).

– prociduus Say 1835: 176; = incertus [var. of incertus in Gillet 1910 ]

– pronus Gerstaecker 1884: 50 ; = integer

propinquus (Felsche) ; see Microcopris

– pseudobootes Ferreira 1962a: 104 ; = typhoeus

pseudochus pseudochus (Ochi and Kon) ; see Sinocopris

pseudochus chuduraziensis Ochi, Kon and Kawahara; see Sinocopris

– pseudolunaris Ferreira 1969: 3 ; = hispanus hispanus

pseudosinicus Balthasar 1958: 475 ; “ China ”.

pueli Mollandin de Boissy 1905: 110 ; Algeria, Tunisia, Armenia?, Iran?, Turkey?

pumiliaris Wallengren 1881: 20; RSA (Transvaal).

punctatus punctatus Gillet 1910: 4 ; China (Fujian, Sichuan), Myanmar, Thailand.

punctatus formosanus Gillet 1921: 126 ; Taiwan.

puncticollis Boheman 1857: 243 ; Mozambique, Zimbabwe.

punctipennis Boucomont 1914: 333 , 335; Indonesia (Sumatra).

punctiventris Waterhouse ; see Litocopris

punctulatus punctulatus (Wiedemann) ; see Paracopris

punctulatus septentrionalis (Ochi, Kon and Pham) ; see Paracopris

punjabensis (Gillet) ; see Paracopris

– quadridentatus ( DeGeer 1778: 638) , as Scarabaeus ; = lunaris

quasilaevigatus (Ochi, Kon and Bai); see Sinocopris

ramosiceps (Gillet) ; see Paracopris

rebouchei Harold 1869c: 497 ; Mexico (Guerrero, Jalisco, México, Michoacán, Morelos, Nuevo León, Puebla, Veracruz).

reflexus (Fabricius) ; see Microcopris

– reflexus ( Panzer 1794: 7) , as Scarabaeus ; = minutus

relictus Matthews ; see Thyregis

remotus remotus LeConte 1866: 381 ; Mexico (Coahuila, Nuevo León, Tamaulipas), USA (Oklahoma, Texas).

remotus dicyrtus Matthews and Halffter 1959: 198 ; Mexico.

renwarti Nguyen-Phung and Cambefort 1986a: 407 ; Ivory Coast.

repertus Walker 1858: 208 ; China (Xizang, Yunnan), India (Himachal Pradesh, Sikkim, Uttar Pradesh), Nepal, Sri Lanka, Thailand.

– retusus Mulsant 1842: 68 , as var. of C. paniscus ; = hispanus hispanus

ribbei Lansberge 1886 a: 12 ; Indonesia (Sulawesi).

riekoae Ochi and Kon 2003: 51 ; Laos.

ritsemae Harold 1875: 137 ; Lesotho, RSA.

ruandanus (Balthasar) ; see Litocopris

rugosus Gillet 1908 a: 77 : Uganda.

ruricola Balthasar 1933: 266 ; China (Sichuan).

sabinus (Gillet) ; see Sinocopris

sacontala (Kollar and Redtenbacher) [in Redtenbacher]; see Sinocopris

sallei Harold 1869c: 496 ; Mexico (Veracruz to Chiapas).

* sanctaeluciae Nguyen-Phung and Cambefort; ssp. of inhalatus

sarpedon Harold 1869b: 104 [replacement name for sexdentatus Kollar and Redtenbacher ]; India (Himachal Pradesh, Uttar Pradesh), Nepal, Thailand, Vietnam–Laos ( Annam).

saundersi Harold 1868b: 99 ; Sulawesi.

schoolmeestersi Ochi and Kon 2010: 127 ; Nepal.

– schoutedeni Boucomont 1928: 419 ; = wiesei

scorpio Zelenka 1993: 241 ; Tanzania.

senegalensis Gillet 1910: 15 ; Senegal.

* septentrionalis (Ochi, Kon and Pham) ; ssp. of punctulatus ; see Paracopris

serius Nguyen-Phung 1987: 271 ; Tanzania.

– servius Harold 1877a: 46 ; = agnus

sexdentatus Thunberg 1818: 404 ; RSA (Cape), Zimbabwe.

– sexdentatus Kollar and Redtenbacher 1844: 520 [in Redtenbacher]; = sarpedon

siamensis Gillet 1910: 12 ; Thailand.

siangensis Biswas 1979: 339 ; India (Arunachal Pradesh).

* sierrensis Matthews ; ssp. of klugi

signatus (Walker) ; see Paracopris

– silenus ( Fabricius 1775: 21) , as Scarabaeus ; = minutus

– similis Balthasar 1939a: 4 , as Paracopris; = mesacanthus

– simillimus Felsche 1910: 346 [erroneously as from “ Sierra Leone ”]; = minutus

simonettai Cambefort 1992b: 179 ; Somalia.

simplex (Harold) ; see Litocopris

simulator Nguyen-Phung and Cambefort 1986a: 409 ; Chad.

singularis Nguyen-Phung and Cambefort 1986a: 406 ; Ethiopia.

sinicus Hope 1842a: 60 ; China (Fujian, Hubei, Jiangxi, Sichuan, Shanghai, Yunnan), Cambodia, Indonesia (Java), Malaysia, Thailand, Vietnam–Laos ( Annam).

sinon ( Olivier 1789: 123) , as Scarabaeus ; Gambia, Mauritania, Senegal.

– sinuatus Mulsant 1842: 68 , as var. of C. paniscus ; = hispanus cavolini sodalis Walker 1858: 208 ; Sri Lanka. sorex Balthasar 1942a: 117 ; China (Fujian). sphaeropterus Harold 1877b: 98 ; RSA (Cape), Zimbabwe. spinator Harold 1881: 89 ; India (Nicobar Is.), Indonesia (Borneo, Sumatra), Malaysia. stereocera Gillet ; see Coptodactyla storeyi Reid ; see Coptodactyla subaenea Harold ; see Coptodactyla subdolus Balthasar 1958: 476 ; China (Fujian). subpunctatus Gillet 1910: 3 ; Costa Rica, Panama. subsidens Péringuey 1901: 344 , 355; Botswana, Namibia.

– sulcicollis Lansberge 1886: 10 ; = sinicus [valid in Gillet 1910] sumatrensis Gillet 1921: 123 ; Sumatra. surdus (Arrow) ; see Paracopris szechouanicus Balthasar 1958: 479 ; China (Fujian, Guizhou, Hunan, Sichuan, Zhejiang).

– taprobanus Gillet 1910: 9 ; = sodalis tarsatus (Paulian) ; see Thyregis tetraodon Gillet 1910: 11 ; Philippines.

– tiberius Schaufuss 1885: 188; = saundersi

* toguchii Ochi and Kawahara; ssp. of brachypterus torresica Matthews ; see Coptodactyla

* transvaalensis Nguyen-Phung ; ssp. of mesacanthus tridens Felsche 1901: 150 ; Cameroon, DRC.

– tridens Mulsant 1842: 68 , as var. of paniscus ; = hispanus cavolini tridentatus Solís and Kohlmann 2003: 2 ; Costa Rica. tripartitus Waterhouse 1875: 74 ; China (Fujian, Sichuan, Yunnan), Taiwan, Japan, Korean peninsula.

– troglodytarum Roth 1851: 124 ; = integer truncatus Felsche 1901: 150 ; Ivory Coast. tsukamotoi (Ochi, Kon and Kawahara) ; see Sinocopris tuberculata Gillet ; see Coptodactyla tubericollis (Felsche) ; see Pseudocopris

– tullius ( Olivier 1789: 118) , as Scarabaeus ; = fricator fricator

– tullius Harold 1877a: 45 ; = spinator typhoeus Gerstaecker 1884: 50 ; Kenya. uenoi (Hanboonsong, Masumoto and Ochi) ; see Sinocopris umbilicatus Abeille de Perrin 1901: 68 ; Albania, Bulgaria, Croatia, France (incl. Corsica), Greece, Hungary, Italy, Romania, Slovakia, Turkey. urus Boheman 1857: 229 ; Mozambique, RSA, Swaziland. usambaricus Gillet 1908 a: 80 ; Kenya, Tanzania. vankhaii vankhaii Nguyen-Phung 1988c: 12 ; Ethiopia, Somalia. vankhaii genopunctatus Nguyen-Phung 1988c: 15; Kenya, Tanzania. victorini Boheman 1857: 234 ; RSA (Cape, Natal).

– victorini Péringuey 1901: 344 , 354; = ritsemae vietnamicus Kabakov 1994: 300 ; Vietnam. vilhenai Ferreira 1962b: 154 ; Angola. villiersi Walter ; see Pseudopedaria vitalisi (Gillet) ; see Microcopris vrydaghi Ferreira 1962a: 105 ; DRC, RSA (Natal). warneri McCleve and Kohlmann 2005: 18 ; Mexico (Chihuahua, Sonora). wiesei Kolbe 1914: 310 ; Burundi, DRC, RSA (Cape), Rwanda, Tanzania, Uganda.

– yamamotoi (Tesař) ; see Sinocopris yangi (Ochi, Kon and Bai); see Sinocopris yokoae Ochi, Kusui and Pham; see Microcopris youngai Balthasar 1967: 49 , 50; PRC (Brazzaville).

zhangi (Ochi, Kon and Bai); see Sinocopris