Gymnopleurus flagellatus, copyright leucospis.

Belongs within: Scarabaeoidea.
Contains: Onthophagini, Oniticellini, 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).

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, Coprinae, Coprini, Dichotomiini, Mentophilina]
    |  i. s.: Aphengium PPS04
    |         Gromphas PPS04
    |         Chironitis PPS04
    |         Uroxys gorgon PPS04, A71
    |         Ontherus PPS04
    |         Saphobius PPS04
    |         Aptenocanthon AY04
    |         Temnoplectron AY04
    |         Boreocanthon praticola C90
    |--+--Coptorhina klugi PPS04
    |  `--Delopleurus PPS04
    `--+--Anomiopus panamensis PPS04
       `--+--Canthidium perceptible PPS04
          `--+--Ateuchus PPS04
             |    |--A. cicatricosus Lucas 1847 E12
             |    `--A. lecomtei PPS04
             |--+--+--+--Onychothecus PPS04
             |  |  |  `--Paraphytus PPS04
             |  |  `--+--+--Scatimus quadridentatus PPS04
             |  |     |  |--Bdelyropsis bowditchi PPS04
             |  |     |  `--Bdelyrus PPS04
             |  |     `--+--+--Lepanus ustulatus PPS04
             |  |        |  `--Arachnodes PPS04
             |  |        `--+--Demarziella imitatrix PPS04
             |  |           `--+--Pedaria PPS04
             |  |              `--+--Pedaridium PPS04
             |  |                 `--Sarophorus PPS04
             |  `--+--+--Xinidium dentilabrum PPS04
             |     |  `--Macroderes Westwood 1843 PPS04, D56
             |     `--+--Epirinus silvestris PPS04
             |        `--+--+--Metacatharsius opacus PPS04
             |           |  `--+--Catharsius PPS04
             |           |     `--Copridaspidus PPS04
             |           `--+--Pseudopedaria PPS04
             |              `--Copris PPS04
             |                   |--C. brachypterus Nomura 1964 I92
             |                   |--C. hispanus WP99
             |                   |--C. lunaris C01b
             |                   `--C. pristinus Z02
             `--+--+--Coptodactyla lesnei PPS04
                |  `--Thyregis PPS04
                |--+--Anachalcos convexus PPS04
                |  `--Amphistomus PPS04
                `--+--+--Sisyphini PPS04
                   |  |    |--Neosisyphus regardi PPS04
                   |  |    `--Sisyphus PPS04
                   |  `--+--Eurysternus [Eurysternini] PPS04
                   |     |    |--E. confusus PPS04
                   |     |    `--E. velutinus PPS04
                   |     `--+--+--Stiptopodius PPS04
                   |        |  `--+--Onthophagini PPS04
                   |        |     `--Oniticellini PPS04
                   |        `--Onitini PPS04
                   |             |  i. s.: Platyonitis smeenkorum PPS04
                   |             |--Onitis PPS04
                   |             |    |--O. chevrolatii Lucas 1847 E12
                   |             |    `--O. fulgidus PPS04
                   |             `--+--Bubas bubalus PPS04
                   |                `--Heteronitis castelnaui PPS04
                   `--+--+--Phanaeini PPS04
                      |  |    |--Sulcophanaeus velutinus PPS04
                      |  |    `--+--Phanaeus difformis PPS04
                      |  |       `--Oxysternon festivum PPS04
                      |  `--+--+--Dichotomius satanus PPS04
                      |     |  `--Chalcocopris PPS04
                      |     `--+--Synapsis tmolus PPS04
                      |        `--Heliocopris hamadryas PPS04
                      `--+--Malagoniella PPS04
                         `--+--+--Cephalodesmius laticollis PPS04
                            |  `--+--+--Circellium bacchus PPS04
                            |     |  `--Scarabaeini PPS04
                            |     `--Eucraniini PPS04
                            |          |--Glyphoderus sterquilinus PPS04
                            |          |--Anomiopsoides cavifrons PPS04
                            |          |--Eucranium PPS04
                            |          `--Ennearabdus lobocephalus PPS04
                            `--+--Canthon PPS04
                               |    |--C. cyanellus PPS04
                               |    |--C. imitator PPS04
                               |    `--C. obliquus PPS04
                               `--Gymnopleurini PPS04
                                    |--Garreta nitens PPS04
                                    `--Gymnopleurus PPS04
                                         |--G. cantharus C01b
                                         |--G. elegans G89
                                         |--G. flagellatus C01a
                                         `--G. pilularius C01b

*Type species of generic name indicated


[A71] Askew, R. R. 1971. Parasitic Insects. Heinemann Educational Books: London.

[AY04] Austin, A. D., D. K. Yeates, G. Cassis, M. J. Fletcher, J. La Salle, J. F. Lawrence, P. B. McQuillan, L. A. Mound, D. J. Bickel, P. J. Gullan, D. F. Hales & G. S. Taylor. 2004. Insects ‘Down Under’—diversity, endemism and evolution of the Australian insect fauna: examples from select orders. Australian Journal of Entomology 43 (3): 216-234.

[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.

[C01a] Csiki, E. 1901a. Utivázlat [Reiseskizze]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy], Band II. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy], pp. xii-xli. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[C01b] Csiki, E. 1901b. Bogarak [Coleopteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy], Band II. 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.

[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.

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[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.

[WP99] Walter, D. E. & H. C. Proctor. 1999. Mites: Ecology, Evolution and Behaviour. CABI Publishing: Wallingford (UK).

[Z02] Zherikhin, V. V. 2002. Ecological history of the terrestrial insects. In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 331-388. Kluwer Academic Publishers: Dordrecht.

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