Belongs within: Onthophagini.
Onthophagus is a cosmopolitan genus of dung beetles with over 2000 species currently included (Philips 2016).
Characters (from Ratcliffe et al. 2002): Body 2–12 mm in length, oval, convex. Head and/or pronotum often with horns or carinae. Scutellum hidden. Elytra with seven discal striae. Third labial palpomere small or inconspicuous. Antenna with nine segments. Tarsi and tarsal claws present on all legs. Middle and posterior tibiae broadly expanded at apex.
<==Onthophagus Latreille 1802P16 (see below for synonymy) |--O. bunamin Matthews 1972M72 |--O. consentaneus Harold 1867M72 [=O. granulatus Macleay 1864 non Boheman 1858CW92] |--O. gandju Matthews 1972M72 |--O. rubrimaculatus Macleay 1864M72 (see below for synonymy) |--O. victoriensis Blackburn 1903M72 [incl. O. jungi Blackburn 1904CW92] |--O. wombalano Matthews 1972M72 |--+--O. blackwoodensis Blackburn 1892M72, CW92 [incl. O. pontilis Blackburn 1903CW92] | `--O. flavoapicalis Lea 1923M72 |--+--O. bornemisszai Matthews 1972M72 | `--O. granulatus Boheman 1858M72 |--+--O. gulmarri Matthews 1972M72 | `--O. pillara Matthews 1972M72 |--O. asper groupM72 | |--O. asper Macleay 1864 [incl. O. patruelis Harold 1886]CW92 | |--O. ocelliger Harold 1877M72 | `--O. rubicundulus Macleay 1871M72 |--O. pronus groupM72 | |--O. bornemisszanus Matthews 1972M72 | |--O. leanus Goidanich 1926M72 [=O. interruptus Lea 1923 non Raffrey 1877CW92] | `--O. pronus Erichson 1842CW92 [=*Psilax pronusM72] |--O. vilis groupM72 | |--O. geelongensis Blackburn 1892M72, CW92 [incl. O. aureoviridanus Blackburn 1903CW92] | |--O. vilis Harold 1877 [incl. O. microtrichius Lea 1923]CW92 | `--O. yiryoront Matthews 1972M72 |--O. dunningi groupM72 | |--O. anchommatus Lea 1923M72 | |--O. brooksi Matthews 1972M72 | |--O. dunningi Harold 1869 [incl. O. dumbrelli Blackburn 1903, O. sydneyicus Paulian 1937]CW92 | `--O. kumbaingeri Matthews 1972M72 |--O. planicollis groupM72 | |--O. clypealis Lea 1923M72 | |--O. macleayi Blackburn 1903M72 | |--O. planicollis Harold 1880CW92 | `--O. yunkara Matthews 1972M72 |--O. pexatus groupM72 | |--O. longipes Paulian 1937M72 | |--O. nammuldi Matthews 1972M72 | |--O. pexatus Harold 1869CW92 | `--O. squalidus Lea 1923M72 |--O. propinquus groupM72 | |--O. monteithi Matthews 1972M72 | |--O. propinquus Macleay 1888 [incl. O. acuticeps Macleay 1888, O. helmsi Blackburn 1903]CW92 | |--O. semimetallicus Lea 1923M72 | `--O. wagamen Matthews 1972M72 |--O. declivis groupM72 | |--O. alquirta Matthews 1972M72 | |--O. apterus Matthews 1972M72 | |--O. declivis Harold 1869CW92 | |--O. desectus Macleay 1871M72 [incl. O. howitti Blackburn 1903CW92] | `--O. devexus Macleay 1888M72 |--O. bicarinaticeps groupM72 | |--O. bicarinaticeps Lea 1923 [incl. O. pauperculus Frey 1963]CW92 | |--O. dummal Matthews 1972M72 | |--O. gidju Matthews 1972M72 | |--O. lamgalio Matthews 1972M72 | `--O. yeyeko Matthews 1972M72 |--O. posticus groupM72 | |--O. arrilla Matthews 1972M72 | |--O. incornutus Macleay 1871M72 [incl. O. semihirtus Frey 1963CW92] | |--O. millamilla Matthews 1972M72 | |--O. mulgravei Paulian 1937M72 | |--O. posticus Erichson 1842 [incl. O. flavolineatus Blanchard 1846, O. leechi Frey 1959]CW92 | `--O. turrbal Matthews 1972M72 |--O. latro groupM72 | |--O. bambra Matthews 1972M72 | |--O. endota Matthews 1972M72 | |--O. latro Harold 1877CW92 | |--O. margaretensis Blackburn 1903M72 | |--O. signaticollis Frey 1970M72 | `--O. varianus Lea 1923M72 |--O. erichsoni groupM72 | |--O. bicornis Macleay 1888M72 | |--O. capellinus Frey 1963M72 | |--O. erichsoni Hope 1842 [incl. O. quadrinodicollis Lea 1924]CW92 | |--O. picipennis Hope 1842M72, CW92 | |--O. tabellicornis Macleay 1864M72 [incl. O. hackeri Paulian 1937CW92] | `--O. wigmungan Matthews 1972M72 |--O. adelaidae groupM72 | |--O. adelaidae Hope 1847 [incl. O. hostilis Harold 1869]CW92 | |--O. duboulayi Waterhouse 1894M72 [incl. O. spissicollis Lea 1923CW92] | |--O. haagi Harold 1867M72 | |--O. jubatus Harold 1869M72 [incl. O. henleyensis Blackburn 1892CW92] | |--O. phoenicocerus Lea 1923M72 | |--O. rupicapra Waterhouse 1894M72 [=O. ruficapra (l. c.)M72; incl. O. granicollis Lea 1924CW92] | `--O. vermiculatus Frey 1970M72 |--O. quadripustulatus groupM72 | |--O. blackburni Shipp 1895M72 (see below for synonymy) | |--O. cruciger Macleay 1888M72 | |--O. evanidus Harold 1869M72 [incl. O. cupreopunctatus Lea 1923CW92, O. negatorius Blackburn 1903CW92] | |--O. fabricii Waterhouse 1894M72 (see below for synonymy) | |--O. kokereka Matthews 1972M72 | |--O. quadripustulatus (Fabricius 1775)CW92 (see below for synonymy) | `--O. subocelliger Blackburn 1903M72 (see below for synonymy) |--O. mutatus groupM72 | |--O. compositus Lea 1923M72 | |--O. frenchi Blackburn 1903M72 | |--O. jangga Matthews 1972M72 | |--O. mutatus Harold 1859 [=O. hirculus Erichson 1842 non Mannerheim 1829]CW92 | |--O. sydneyensis Blackburn 1903M72 [incl. O. monticola Paulian 1937CW92] | |--O. togeman Matthews 1972M72 | `--O. yungaburra Matthews 1972M72 |--O. perpilosus groupM72 | |--O. fletcheri Blackburn 1903M72 | |--O. gangulu Matthews 1972M72 | |--O. incanus Macleay 1888M72 [incl. O. leaniensis Paulian 1937CW92] | |--O. perpilosus Macleay 1871CW92 | |--O. tamworthi Blackburn 1903M72 | |--O. villosus Macleay 1888M72 | `--O. wakelbura Matthews 1972M72 |--O. auritus groupM72 | |--O. auritus Erichson 1842 (see below for synonymy)CW92 | |--O. cuniculus Macleay 1864M72 | |--O. dandalu Matthews 1972M72 | |--O. discolor Hope 1841M72 [incl. O. promptus Harold 1869CW92, O. viridiobscurus Blanchard 1846CW92] | |--O. furcaticeps Masters 1886M72 (see below for synonymy) | |--O. purpureicollis Macleay 1864M72 | |--O. rufosignatus Macleay 1864 [incl. O. kraatzi Harold 1880]M72 | `--O. walteri Macleay 1887M72 |--O. conspicuus groupM72 | |--O. bicavicollis Lea 1923M72 | |--O. conspicuus Macleay 1864 [incl. O. bovilli Blackburn 1903]CW92 | |--O. fissiceps Macleay 1888M72 (see below for synonymy) | |--O. jalamari Matthews 1972M72 | |--O. rugosicollis Gillet 1925M72 [=O. cribricollis Lea 1923 non Horn 1881CW92] | |--O. salebrosus Macleay 1888M72 | |--O. tenebrosus Harold 1871M72 [=O. rugosus Kirby 1818 non Poda 1761CW92] | `--O. tricavicollis Lea 1923M72 [incl. O. conspicuoformis Paulian 1937CW92] |--O. rubescens groupM72 | |--O. koebelei Blackburn 1903M72 | |--O. manya Matthews 1972M72 | |--O. parrumbal Matthews 1972M72 | |--O. rubescens Macleay 1888 [incl. O. imponderosus Lea 1923]CW92 | |--O. symbioticus (Arrow 1920)M72 [=Macropocopris symbioticusCW92] | |--O. tuckonie Matthews 1972M72 | |--O. waminda Matthews 1972M72 | `--O. wilgi Matthews 1972M72 |--O. australis groupM72 | |--O. anisocerus Erichson 1842M72 | |--O. australis Guérin-Méneville 1838CW92 (see below for synonymy) | |--O. fuliginosus Erichson 1842M72 | |--O. hoplocerus Lea 1923M72 | |--O. nurubuan Matthews 1972M72 | |--O. paluma Matthews 1972M72 | |--O. parallelicornis Macleay 1887M72 | |--O. thoreyi Harold 1868M72 [incl. O. mastersi Macleay 1871CW92, O. sutilistriatus Lea 1923CW92] | `--O. tweedensis Blackburn 1903M72 |--O. glabratus groupM72 | |--O. carmodensis Blackburn 1907 [=Macropocopris carmodensis]M72 | |--O. chepara Matthews 1972M72 | |--O. glabratus Hope 1842 [incl. O. chillagoensis Paulian 1937]CW92 | |--O. murchisoni Blackburn 1892M72 | |--O. muticus Macleay 1864M72 [incl. O. kingi Harold 1869CW92, Macropocopris kingiM72] | |--O. neboissi Frey 1970M72 | |--O. parvus Blanchard 1846M72, CW92 (see below for synonymy) | |--O. peramelinus (Lea 1923)M72 [=Macropocopris peramelinusCW92; incl. O. postcornutus Frey 1963CW92] | |--O. prehensilis (Arrow 1920)M72 [=*Macropocopris prehensilisCW92] | `--O. variolicollis Lea 1923M72 |--O. atrox groupM72 | |--O. atrox Harold 1867CW92 | |--O. capitosus Harold 1867M72 [=O. quadridentatus Hope 1842 non Fabricius 1798CW92] | |--O. demarzi Frey 1959M72 | |--O. ferox Harold 1867M72 [incl. O. inaequalisMas86] | |--O. laminatus Macleay 1864M72 [incl. O. cowleyi Blackburn 1903CW92, O. palmerstoni Blackburn 1903CW92] | |--O. leai Blackburn 1895M72 | |--O. mjobergi Gillet 1925M72 | |--O. mniszechi Harold 1869 [=*Monapus mniszechi, O. ferox var. mniszechi]M72 | |--O. pentacanthus Harold 1867M72 | |--O. pugnacior Blackburn 1903M72 | |--O. quinquetuberculatus Macleay 1871M72 | |--O. sloanei Blackburn 1903M72 | `--O. solidus Gillet 1927M72 `--O. capella groupM72 |--O. capella Kirby 1818 [incl. O. schmeltzi Harold 1869]CW92 |--O. capelliformis Gillet 1925M72 |--O. comperei Blackburn 1903M72 |--O. darlingtoni Matthews 1972M72 |--O. dicranocerus Gillet 1925M72 [incl. O. demarziellus Frey 1963CW92] |--O. ferrari Matthews 1972M72 |--O. macrocephalus Kirby 1818M72 |--O. mamillatus Lea 1923M72 |--O. mundill Matthews 1972M72 |--O. neostenocerus Goidanich 1926M72 [=O. stenocerus Lea 1923 non Harold 1867CW92] |--O. nodulifer Harold 1867M72 [incl. O. divaricatus Macleay 1871CW92] |--O. ouratita Matthews 1972M72 |--O. pugnax Harold 1868M72 [incl. O. opacipennis Lea 1923CW92] |--O. queenslandicus Blackburn 1903M72 [incl. O. fitzroyensis Blackburn 1903CW92] `--O. tabellifer Gillet 1927M72 Onthophagus incertae sedis: *O. taurus (Schreber 1759)CW92 [=Scarabaeus taurusCW92, Copris taurusL02] O. aleppensisC01 O. analis Lucas 1846E12 O. arkoola Storey & Weir 1990CW92 O. austriacusC01 O. beelarong Storey & Weir 1990CW92 O. bindaree Storey & Weir 1990CW92 O. binodis Thunberg 1818CW92 O. binyana Storey & Weir 1990CW92 O. bundara Storey & Weir 1990CW92 O. coenobitaC01 O. cooloola Storey & Weir 1990CW92 O. crinitisHG01 |--O. c. crinitisHG01 `--O. c. panamensisHG01 O. crotchi Harold 1871 (n. d.)CW92 O. damaB35 O. dinjerra Storey & Weir 1990CW92 O. femoralis Kirsch 1870FS90 O. fissicornisC01 O. flavipesG20 O. fracticornis (Preyssler 1790)A06 O. gazella (Fabricius 1787) [=Scarabaeus gazella]CW92 O. godarra Storey & Weir 1990CW92 O. granum Lansberge 1885 (n. d.)CW92 O. gurburra Storey & Weir 1990CW92 O. hecateP16 O. insensusWK86 O. kakadu Storey & Weir 1990CW92 O. kiambram Storey 1977CW92 O. kora Storey 1977CW92 O. marginalisC01 O. maurus Lucas 1846E12 O. mije Storey & Weir 1990CW92 O. mongana Storey & Weir 1990CW92 O. nigriventris d’Orbigny 1902 [=O. blanchardi nigriventris]CW92 O. nitidusG89 O. nuchicornisC01 O. nutansR26 O. obliquus (Olivier 1789) [=Scarabaeus obliquus]CW92 O. ovatusC01 O. oleipennis Macleay 1886Mac86 O. parryiMac86 O. pinaroo Storey & Weir 1990CW92 O. planiceps Macleay 1886Mac86 O. polyphemiRJS02 O. ponticusC01 O. punthari Storey 1977CW92 O. reticollis Macleay 1886Mac86 O. sagittarius (Fabricius 1781) [=Scarabaeus sagittarius]CW92 O. schreberiR26 O. striatulusRJS02 O. subaeneusRJS02 O. terrara Storey 1977CW92 O. transbaicalicus Csiki 1901C01 O. trawalla Storey & Weir 1990CW92 O. vaccaR26 O. verticicornisC01 O. wanappe Storey 1977CW92 O. weringerong Storey & Weir 1990CW92 O. williamsi Storey & Weir 1990CW92 O. worooa Storey & Weir 1990CW92 O. yackatoon Storey & Weir 1990CW92 O. yaran Storey & Weir 1990CW92 O. yarrumba Storey 1977CW92 O. yourula Storey & Weir 1990CW92 O. (Pseudonthophagus Kabakov & Shokhin 2014)P16
Onthophagus Latreille 1802P16 [incl. Chalcoderus Erichson 1848RJS02, Gonocyphus Lansberg 1885RJS02, Macropocopris Arrow 1920CW92, Monapus Erichson 1848M72, Psilax Erichson 1848M72, Tauronthophagus Shipp 1895RJS02]
Onthophagus auritus Erichson 1842 [incl. O. cereus Hope 1847, O. lucidicollis Boheman 1858, O. micans Sturm 1843, O. umbraculatus Blanchard 1846]CW92
Onthophagus australis Guérin-Méneville 1838CW92 [incl. O. capella Boisduval 1832 non Kirby 1818M72, O. cupreoviridis Blanchard 1846CW92]
Onthophagus blackburni Shipp 1895M72 [=O. nitidior Blackburn 1892 non Bates 1887CW92; incl. O. horianus Paulian 1937CW92, O. zietzi Blackburn 1903CW92]
Onthophagus fabricii Waterhouse 1894M72 [=O. humator Shipp 1895CW92, O. humeralis Macleay 1888 non Raffray 1877CW92; incl. O. strabonis Lea 1923CW92]
Onthophagus fissiceps Macleay 1888M72 [incl. O. darwini Paulian 1937CW92, O. integriceps Macleay 1888CW92, O. semirugosus Gillet 1927CW92]
Onthophagus furcaticeps Masters 1886M72 [=O. furcatus Macleay 1864 non Fabricius 1781CW92; incl. O. froggatti Macleay 1887CW92, O. lobicollis Macleay 1887CW92]
Onthophagus parvus Blanchard 1846M72, CW92 [=Macropocopris parvusM72; incl. O. inermis Macleay 1871CW92, O. submuticus Blackburn 1903CW92]
Onthophagus quadripustulatus (Fabricius 1775)CW92 [=Scarabaeus quadripustulatusCW92; incl. O. albertisi Harold 1877M72, S. bipustulatus Fabricius 1775CW92, O. bipustulatusM72, O. minusculus Macleay 1888CW92]
Onthophagus rubrimaculatus Macleay 1864M72 [incl. O. decurio Lansberge 1885CW92, O. emarginatus Macleay 1887CW92, O. pseudoemarginatus Goidanich 1926CW92]
Onthophagus subocelliger Blackburn 1903M72 [=O. nanus Waterhouse 1894 non Harold 1871CW92, O. waterhousei Boucomont & Gillet 1927CW92]
*Type species of generic name indicated
References
[A06] Ahrens, D. 2006. The phylogeny of Sericini and their position within the Scarabaeidae based on morphological characters (Coleoptera: Scarabaeidae). Systematic Entomology 31: 113–144.
[B35] Boisduval, J. B. 1835. Voyage de Découvertes de l’Astrolabe. Exécuté par ordre du Roi, pendant les années 1826–1827–1828–1829, sous le commandement de M. J. Dumont d’Urville. Faune entomologique de l’océan Pacifique, avec l’illustration des insectes nouveaux recueillis pendant le voyage vol. 2. Coléoptères et autres ordres. J. Tastu: Paris.
[CW92] Cassis, G., & T. A. Weir. 1992. Scarabaeinae. In: Houston, W. W. K. (ed.) Zoological Catalogue of Australia vol. 9. Coleoptera: Scarabaeoidea pp. 106–173. AGPS Press: Canberra.
[C01] Csiki, E. 1901. Bogarak [Coleopteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 75–120. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.
[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l’Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.
[FS90] Fleutiaux, E., & A. Sallé. 1890. Liste des coléoptères de la Guadeloupe et descriptions d’espèces nouvelles. Annales de la Société Entomologique de France, 6e série 9: 351–484.
[G89] Gestro, R. 1889. Viaggio ab Assab nel Mar Rosso dei signori G. Doria ed O. Beccari con il R. Avviso “Esploratore” dal 16 Novembre 1879 al 26 Febbraio 1880.—IV. Coleotteri. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a, 7: 5–72.
[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.
[HG01] Harvey, P. H., & C. J. Godfray. 2001. A horn for an eye. Science 291: 1505–1506.
[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.
[Mac86] Macleay, W. 1886. The insects of the Fly River, New Guinea, “Coleoptera”. Proceedings of the Linnean Society of New South Wales, series 2, 1 (1): 136–157.
[Mas86] Masters, G. 1886. Catalogue of the described Coleoptera of Australia. Part III. Proceedings of the Linnean Society of New South Wales, series 2, 1 (1): 21–126.
[M72] Matthews, E. G. 1972. A revision of the scarabaeine dung beetles of Australia. I. Tribe Onthophagini. Australian Journal of Zoology, Supplementary Series 9: 1–330.
[P16] Philips, T. K. 2016. Phylogeny of the Oniticellini and Onthophagini dung beetles (Scarabaeidae, Scarabaeinae) from morphological evidence. ZooKeys 579: 9–57.
[RJS02] Ratcliffe, B. C., M. L. Jameson & A. B. T. Smith. 2002. Scarabaeidae Latreille 1802. In: Arnett, R. H., Jr, M. C. Thomas, P. E. Skelley & J. H. Frank (eds) American Beetles vol. 2. Polyphaga: Scarabaeoidea through Curculionoidea pp. 39–81. CRC Press.
[R26] Risso, A. 1826. Histoire naturelle des principales productions de l’Europe méridionale et particulièrement de celles des environs de Nice et des Alpes maritimes vol. 5. F.-G. Levrault: Paris.
[WK86] Walter, D. E., & G. W. Krantz. 1986. Description of the Macrocheles kraepelini species complex (Acari: Macrochelidae) with two new species. Canadian Journal of Zoology 64: 212–217.