Belongs within: Scarabaeidae.
Contains: Hyboma, Ateuchus, Lepanus, Demarziella, Copris, Amphistomus, Coptodactyla, Sisyphini, Eurysternus, Onitini, Onthophagini, Oniticellini, Phanaeini, Gymnopleurini, Deltochilini, Eucraniini, Circellium, Scarabaeini.
The Scarabaeinae include the true dung beetles, so named for their use of dung as a food source for developing larvae. Dung beetles can be divided between tunnelling species (that burrow into and bury dung below or beside where they find it) and rolling species (that collect dung and roll it away from its original location). Phylogenetic analysis indicates that the tunnelling behaviour is ancestral with multiple origins of rolling behaviour (Philip et al. 2004). The previously recognised forest-dwelling tribe Dichotomiini is paraphyletic to the remaining scarabaeines, and the fungus-feeding ‘dichotomiins’ Coptorhina, Bdelyrus and Bdelyropsis occupy relatively basal positions in the tree. As related subfamilies of Scarabaeidae include fungus feeders, it is possible that these genera have retained an ancestral food preference with dung-feeding being derived with Scarabaeinae (Philips et al. 2004).
In many Scarabaeinae, the tarsi of the fore legs have become greatly reduced or completely lost. Cephalodesmius is an eastern Australian genus with a smooth dorsum and four to eight teeth on the clypeus, the median pair of which is often very prominent. Malagoniella is a Neotropical genus in which the scutellum remains visible (Ratcliffe et al. 2002). Heliocopris is an Old World genus of often large dung beetles, with males often bearing horns on the head or pronotum.
Synapomorphies (from Philips et al. 2004): Wing with RA distinctly sigmoidal, ‘Z’ vein long, extending up to middle of wing; epipharynx with anterior median dorsal process apically rounded, basal lines of fine setae present laterally opposite to anterior median process on ventral surface; maxilla without anterolateral projection on ventral articulatory sclerite; labium with two distinct anteriorly directly lobes with median emargination on glossae; scutum and scutellum on distinct levels with a strong declivity; alar ridge of scutellum distinctly converging in dorsal view; pygidium with dorsolateral edge angled; aedeagus with parameres somewhat blunt or truncate; clypeus with two distinct well-developed teeth; midcoxae slightly oblique; metendosternite narrow near posterior attachment, ventral edge (posterior of lateral chitinous projection) concave.
Scarabaeinae [Canthonini, Coprides, Coprinae, Coprini, Dichotomiini, Mentophilina]
|--+--DelopleurusPPS04
| `--CoptorhinaPPS04
| |--C. africanaLC40
| `--C. klugiPPS04
`--+--Anomiopus panamensisPPS04
`--+--Canthidium Erichson 1847PPS04, FS90
| |--C. melanocephalum Ol. 1789FS90
| `--C. perceptiblePPS04
`--+--AteuchusPPS04
|--+--+--+--OnychothecusPPS04
| | | `--ParaphytusPPS04 [incl. FreyusP16]
| | `--+--+--Scatimus quadridentatusPPS04
| | | |--Bdelyropsis bowditchiPPS04
| | | `--BdelyrusPPS04
| | `--+--+--LepanusPPS04
| | | `--ArachnodesPPS04
| | `--+--DemarziellaPPS04
| | `--+--PedaridiumPPS04
| | `--SarophorusPPS04
| `--+--+--Xinidium dentilabrumPPS04
| | `--Macroderes Westwood 1843PPS04, D56
| `--+--EpirinusPPS04
| | |--E. hilarisP16
| | `--E. silvestrisP16
| `--+--+--PseudopedariaPPS04
| | `--CoprisPPS04
| `--+--Metacatharsius opacusPPS04
| `--+--CopridaspidusPPS04
| `--CatharsiusPPS04
| |--C. molossus (Linnaeus 1758)A06
| `--C. sagaxWK86a
`--+--+--Anachalcos convexusPPS04
| `--AmphistomusPPS04
|--+--CoptodactylaPPS04
| `--Thyregis Blackburn 1904PPS04, CW92 [incl. Arrowianella Paulian 1933CW92]
| |--*T. kershawi Blackburn 1904 [incl. Arrowianella pauliani Frey 1970]CW92
| |--T. monteithi Matthews 1976CW92
| |--T. relictus Matthews 1976CW92
| `--T. tarsatus (Paulian 1933) [=*Arrowianella tarsata]CW92
`--+--+--SisyphiniPPS04
| `--+--EurysternusPPS04
| `--+--OnitiniPPS04
| `--+--OnthophaginiPPS04
| `--OniticelliniPPS04
`--+--+--PhanaeiniPPS04
| `--+--+--ChalcocoprisPPS04
| | `--Dichotomius Hope 1838PPS04, RJS02 (see below for synonymy)
| | `--D. satanusPPS04
| `--+--Synapsis tmolusPPS04
| `--HeliocoprisPPS04
| |--H. andersoniB14
| |--H. bucephalusWK86a
| |--H. gigas (Linnaeus 1758)B14
| |--H. hamadryasPPS04
| |--H. midasB14
| `--H. tyrannusWK86b
`--+--Malagoniella Martínez 1961PPS04, RJS02
| `--M. astyanaxRJS02
| |--M. a. astyanaxRJS02
| `--M. a. yucatecaRJS02
`--+--+--GymnopleuriniPPS04
| `--DeltochiliniPPS04
`--+--Cephalodesmius Westwood 1841PPS04, CW92
| |--*C. armiger Westwood 1841 [incl. C. minor Blackburn 1900]CW92
| |--C. laticollis Pascoe 1863 (see below for synonymy)CW92
| `--C. quadridens Macleay 1871CW92
`--+--EucraniiniPPS04
`--+--CircelliumPPS04
`--ScarabaeiniPPS04
Scarabaeinae incertae sedis:
AphengiumPPS04
GromphasPPS04
ChironitisPPS04
Uroxys gorgonPPS04, A71
Ontherus Erichson 1847RJS02
`--O. mexicanusRJS02
SaphobiusPPS04
ParachoriusP16
ScatrichusP16
Melanocanthon Halffter 1958RJS02
Pseudocanthon Bates 1887 [incl. Opiocanthon Paulian 1947]RJS02
`--P. perplexusRJS02
PachysomaLC40
|--P. (Pachysoma)LC40
| |--P. (P.) aesculapiusLC40
| |--P. (P.) macleayi Laporte de Castelnau 1840LC40
| |--P. (P.) hippocratesLC40
| `--P. (P.) striatum Laporte de Castelnau 1840LC40
`--P. (Cyclodema Laporte de Castelnau 1840)LC40
`--P. (*C.) lacordairei Laporte de Castelnau 1840LC40
Scatonomus viridisLC40
HybomaLC40
EnicotarsusLC40
|--E. aterLC40
|--E. quadratusLC40
`--E. viridipennisLC40
ChaeridiumLC40
|--C. capistratumLC40
|--C. collareLC40
|--C. elegansLC40
|--C. fasciatum Bonfils in Laporte de Castelnau 1840LC40
`--C. nitidulum Klug in Laporte de Castelnau 1840LC40
PedariaLC40
|--P. nigraLC40
`--P. tuberculata Laporte de Castelnau 1840LC40
Cephalodesmius laticollis Pascoe 1863 [incl. C. magnini Paulian 1935, C. teissieri Paulian 1935]CW92
Dichotomius Hope 1838PPS04, RJS02 [incl. Brachycopris Haldeman 1845RJS02, Pinotus Erichson 1847RJS02]
*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.
[A71] Askew, R. R. 1971. Parasitic Insects. Heinemann Educational Books: London.
[B14] Bouchard, P. (ed.) 2014. The Book of Beetles: A lifesize guide to six hundred of nature’s gems. Ivy Press: Lewes (United Kingdom).
[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.
[D56] Dawes, B. 1956. The Trematoda with special reference to British and other European forms. University Press: Cambridge.
[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.
[LC40] Laporte de Castelnau, F. 1840. Histoire Naturelle des Insectes Coléoptères vol. 2. P. Duménil: Paris.
[P16] Philips, T. K. 2016. Phylogeny of the Oniticellini and Onthophagini dung beetles (Scarabaeidae, Scarabaeinae) from morphological evidence. ZooKeys 579: 9–57.
[PPS04] Philips, T. K., E. Pretorius & C. H. Scholtz. 2004. A phylogenetic analysis of dung beetles (Scarabaeinae: Scarabaeidae): unrolling an evolutionary history. Invertebrate Systematics 18: 53–88.
[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.
[WK86a] Walter, D. E., & G. W. Krantz. 1986a. Description of the Macrocheles kraepelini species complex (Acari: Macrochelidae) with two new species. Canadian Journal of Zoology 64: 212–217.
[WK86b] Walter, D. E., & G. W. Krantz. 1986b. A review of the glaber-group (s. str.) species of the genus Macrocheles (Acari: Macrochelidae), and a discussion of species complexes. Acarologia 27 (4): 277–294.