Gymnopleurus flagellatus, copyright leucospis.

Belongs within: Scarabaeoidea.
Contains: Lepanus, Demarziella, Copris, Amphistomus, Coptodactyla, Onitini, Onthophagini, Oniticellini, Eucraniini, 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. 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. The Gymnopleurini, found in Africa and Eurasia, have the sides of the elytra emarginate to expose the underlying pleural sclerites.

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]
    |  i. s.: Aphengium PPS04
    |         Gromphas PPS04
    |         Chironitis PPS04
    |         Uroxys gorgon PPS04, A71
    |         Ontherus PPS04
    |         Saphobius PPS04
    |         Boreocanthon praticola C90
    |         Parachorius P16
    |         Scatrichus P16
    |--+--Coptorhina klugi PPS04
    |  `--Delopleurus PPS04
    `--+--Anomiopus panamensis PPS04
       `--+--Canthidium perceptible PPS04
          `--+--Ateuchus PPS04
             |    |--A. cicatricosus Lucas 1846 E12
             |    |--A. hollandiae [=Aulacium hollandiae, Scarabaeus hollandiae] B35
             |    |--A. lecomtei PPS04
             |    `--A. sacer L02
             |--+--+--+--Onychothecus PPS04
             |  |  |  `--Paraphytus PPS04 [incl. Freyus P16]
             |  |  `--+--+--Scatimus quadridentatus PPS04
             |  |     |  |--Bdelyropsis bowditchi PPS04
             |  |     |  `--Bdelyrus PPS04
             |  |     `--+--+--Lepanus PPS04
             |  |        |  `--Arachnodes PPS04
             |  |        `--+--Demarziella PPS04
             |  |           `--+--Pedaridium PPS04
             |  |              `--Sarophorus PPS04
             |  `--+--+--Xinidium dentilabrum PPS04
             |     |  `--Macroderes Westwood 1843 PPS04, D56
             |     `--+--Epirinus PPS04
             |        |    |--E. hilaris P16
             |        |    `--E. silvestris PPS04
             |        `--+--+--Pseudopedaria PPS04
             |           |  `--Copris PPS04
             |           `--+--Metacatharsius opacus PPS04
             |              `--+--Copridaspidus PPS04
             |                 `--Catharsius PPS04
             |                      |--C. molossus (Linnaeus 1758) A06
             |                      `--C. sagax WK86a
             `--+--+--Anachalcos convexus PPS04
                |  `--Amphistomus PPS04
                |--+--Coptodactyla PPS04
                |  `--Thyregis Blackburn 1904 PPS04, CW92 [incl. Arrowianella Paulian 1933 CW92]
                |       |--*T. kershawi Blackburn 1904 [incl. Arrowianella pauliani Frey 1970] CW92
                |       |--T. monteithi Matthews 1976 CW92
                |       |--T. relictus Matthews 1976 CW92
                |       `--T. tarsatus (Paulian 1933) [=*Arrowianella tarsata] CW92
                `--+--+--Sisyphini PPS04
                   |  |    |--Neosisyphus regardi PPS04
                   |  |    `--Sisyphus Latreille 1807 CW92
                   |  |         |--*S. schaefferi (Linnaeus 1758) [=Scarabaeus schaefferi] CW92
                   |  |         |--S. infuscatus Klug 1855 CW92
                   |  |         |--S. rubrus Paschalidis 1974 CW92
                   |  |         `--S. spinipes (Thunberg 1818) [=Ateuchus spinipes] CW92
                   |  `--+--Eurysternus [Eurysternini] PPS04
                   |     |    |--E. confusus PPS04
                   |     |    `--E. velutinus PPS04
                   |     `--+--Onitini PPS04
                   |        `--+--Onthophagini PPS04
                   |           `--Oniticellini PPS04
                   `--+--+--Phanaeini PPS04
                      |  |    |--Sulcophanaeus PPS04
                      |  |    |    |--S. imperator (Chevrolat 1844) B14
                      |  |    |    `--S. velutinus PPS04
                      |  |    `--+--Oxysternon festivum PPS04
                      |  |       `--Phanaeus PPS04
                      |  |            |--P. difformis PPS04
                      |  |            |--P. milon R13
                      |  |            `--P. vindex RD77
                      |  `--+--+--Dichotomius satanus PPS04
                      |     |  `--Chalcocopris PPS04
                      |     `--+--Synapsis tmolus PPS04
                      |        `--Heliocopris PPS04
                      |             |--H. andersoni B14
                      |             |--H. bucephalus WK86a
                      |             |--H. gigas (Linnaeus 1758) B14
                      |             |--H. hamadryas PPS04
                      |             |--H. midas B14
                      |             `--H. tyrannus WK86b
                      `--+--Malagoniella PPS04
                         `--+--+--Cephalodesmius Westwood 1841 PPS04, CW92
                            |  |    |--*C. armiger Westwood 1841 [incl. C. minor Blackburn 1900] CW92
                            |  |    |--C. laticollis Pascoe 1863 (see below for synonymy) CW92
                            |  |    `--C. quadridens Macleay 1871 CW92
                            |  `--+--Eucraniini PPS04
                            |     `--+--Circellium bacchus PPS04
                            |        `--Scarabaeini PPS04
                            `--+--Canthon PPS04
                               |    |--C. cyanellus PPS04
                               |    |--C. humectus B70
                               |    |--C. imitator PPS04
                               |    `--C. obliquus PPS04
                               `--Gymnopleurini PPS04
                                    |--Garreta nitens PPS04
                                    `--Gymnopleurus PPS04
                                         |--G. cantharus C01
                                         |--G. elegans G89
                                         |--G. flagellatus C01
                                         `--G. pilularius C01

Cephalodesmius laticollis Pascoe 1863 [incl. C. magnini Paulian 1935, C. teissieri Paulian 1935] CW92

*Type species of generic name indicated


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[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.

[B14] Bouchard, P. (ed.) 2014. The Book of Beetles: A lifesize guide to six hundred of nature's gems. Ivy Press: Lewes (United Kingdom).

[B70] Britton, E. B. 1970. Coleoptera (beetles). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 495–621. Melbourne University Press.

[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.

[C90] Crawford, C. S. 1990. Scorpiones, Solifugae, and associated desert taxa. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 421-475. John Wiley & Sones: New York.

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[D56] Dawes, B. 1956. The Trematoda with special reference to British and other European forms. University Press: Cambridge.

[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.

[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.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: 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.

[R13] Reuter, O. M. 1913. Lebensgewohnheiten und Instinkte der Insekten bis zum Erwachen der sozialen Instinkte. R. Friedländer & Sohn: Berlin.

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms' General Textbook of Entomology 10th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

[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.

Last updated: 2 May 2021.

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