Scarabaeinae

Gymnopleurus flagellatus, copyright leucospis.

Belongs within: Scarabaeidae.
Contains: Lepanus, Demarziella, Copris, Amphistomus, Coptodactyla, Onitini, Onthophagini, Oniticellini, Phanaeini, 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 (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. The Gymnopleurini, found in Africa and Eurasia, have the sides of the elytra emarginate to expose the underlying pleural sclerites. Members of the tribe Sisyphini are characterised by a laterally compressed body and long, slender mid and hind legs (Ratcliffe et al. 2002).

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]
    |--+--Coptorhina klugiPPS04
    |  `--DelopleurusPPS04
    `--+--Anomiopus panamensisPPS04
       `--+--Canthidium Erichson 1847PPS04, FS90
          |    |--C. melanocephalum Ol. 1789FS90
          |    `--C. perceptiblePPS04
          `--+--Ateuchus Weber 1801PPS04, RJS02 [incl. Choeridium LePeletier & Serville 1828RJS02]
             |    |--‘Choeridium’ breveFS90
             |    |--A. cicatricosus Lucas 1846E12
             |    |--A. flagellatusR26
             |    |--A. hollandiae [=Aulacium hollandiae, Scarabaeus hollandiae]B35
             |    |--‘Choeridium’ insulare Chv. in Fleutiaux & Sallé 1890FS90
             |    |--A. laticollisR26
             |    |--A. lecomteiPPS04
             |    `--A. sacerL02
             |--+--+--+--OnychothecusPPS04
             |  |  |  `--ParaphytusPPS04 [incl. FreyusP16]
             |  |  `--+--+--Scatimus quadridentatusPPS04
             |  |     |  |--Bdelyropsis bowditchiPPS04
             |  |     |  `--BdelyrusPPS04
             |  |     `--+--+--LepanusPPS04
             |  |        |  `--ArachnodesPPS04
             |  |        `--+--DemarziellaPPS04
             |  |           `--+--PedaridiumPPS04
             |  |              `--SarophorusPPS04
             |  `--+--+--Xinidium dentilabrumPPS04
             |     |  `--Macroderes Westwood 1843PPS04, D56
             |     `--+--EpirinusPPS04
             |        |    |--E. hilarisP16
             |        |    `--E. silvestrisPPS04
             |        `--+--+--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
                `--+--+--+--Eurysternus [Eurysternini]PPS04
                   |  |  |    |--E. confususPPS04
                   |  |  |    `--E. velutinusPPS04
                   |  |  `--+--OnitiniPPS04
                   |  |     `--+--OnthophaginiPPS04
                   |  |        `--OniticelliniPPS04
                   |  `--SisyphiniPPS04
                   |       |--Neosisyphus regardiPPS04
                   |       `--Sisyphus Latreille 1807CW92
                   |            |--*S. schaefferi (Linnaeus 1758) [=Scarabaeus schaefferi]CW92
                   |            |--S. infuscatus Klug 1855CW92
                   |            |--S. rubrus Paschalidis 1974CW92
                   |            |--S. spinipes (Thunberg 1818) [=Ateuchus spinipes]CW92
                   |            `--S. submonticolusRJS02
                   `--+--+--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
                         `--+--+--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
                            |     `--+--Circellium bacchusPPS04
                            |        `--ScarabaeiniPPS04
                            `--+--GymnopleuriniPPS04
                               |    |--Garreta nitensPPS04
                               |    `--GymnopleurusPPS04
                               |         |--G. cantharusC01
                               |         |--G. elegansG89
                               |         |--G. flagellatusC01
                               |         `--G. pilulariusC01
                               `--DeltochiliniMF15
                                    |--Deltochilum Eschscholtz 1822RJS02
                                    `--Canthon Hoffmansegg 1817PPS04, FS90 (see below for synonymy)
                                         |--C. cyanellusPPS04
                                         |--C. humectusB70
                                         |--C. imitatorPPS04
                                         |--C. obliquusPPS04
                                         |--C. pilulariusRJS02
                                         |--‘Boreocanthon’ praticolaC90
                                         `--C. vitraci Fleutiaux & Sallé 1890FS90
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

Canthon Hoffmansegg 1817PPS04, FS90 [incl. Boreocanthon Halffter 1958RJS02, Geocanthon Pereira & Martínez 1956RJS02, Glaphyrocanthon Martínez 1948RJS02]

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.

[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.

[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.

[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.

[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.

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

[MF15] McKenna, D. D., B. D. Farrell, M. S. Caterino, C. W. Farnum, D. C. Hawks, D. R. Maddison, A. E. Seago, A. E. Z. Short, A. F. Newton & M. K. Thayer. 2015. Phylogeny and evolution of Staphyliniformia and Scarabaeiformia: forest litter as a stepping stone for diversification of nonphytophagous beetles. Systematic Entomology 40: 35–60.

[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.

[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.

[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.

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