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