Bolboceratinae

Odonteus armiger, copyright Udo Schmidt.

Belongs within: Geotrupidae.
Contains: Bolborhachium, Blackbolbus, Blackburnium, Australobolbus.

The Bolboceratinae are a cosmopolitan group of stout, strongly convex scarabaeoid beetles that dig deep burrows and feed on fungi or plant material. Males usually bear large cephalic and pronotal horns. They are most diverse in Australia where they are primarily found in coastal regions (Cassis & Weir 1992). Members of the tribe Athyreini have the mesocoxae much more widely separated that Bolboceratini (Jameson 2002).

<==Bolboceratinae [Bolboceratidae]
    |--BolboceratiniMF15
    |    |--Bolboceras Kirby 1819J02
    |    |    |--B. corniculatumM86
    |    |    |--B. filicornisJ02
    |    |    `--B. proboscidium Schreibers 1802M86
    |    `--Odonteus Samouelle 1819 [=Odontaeus Dejean 1821]J02
    |         `--O. armigerMF15
    `--Athyreini [Athyreinae]J02
         |--PseudoathyreusL96
         `--Neoathyreus Howden & Martínez 1963L96, J02
              |--N. fissicornisJ02
              `--N. mixtusJ02
Bolboceratinae incertae sedis:
  Bolbobaineus Howden & Cooper 1977CW92
    |--*B. planiceps (Macleay 1873) [=Bolboceras planiceps]CW92
    `--B. simpliciceps (Blackburn 1888) [=Bolboceras simpliciceps]CW92
  Eucanthus Westwood 1848CW92
    |--*E. meliboeus (Fabricius 1792) [=Scarabaeus meliboeus]CW92
    |--E. felschei Boucomont 1911CW92
    `--E. tricarinaticeps Lea 1924CW92
  Gilletinus Boucomont 1932CW92
    |--*G. multicostatus (Lansberge 1885) [=Bolboceras multicostatus]CW92
    |--G. bipagus Howden 1990CW92
    |--G. corrugatus (Lea 1924) [=Bolboceras corrugatum]CW92
    |--G. flavocastaneus (Lea 1924) [=Bolboceras corrugatum flavocastaneum]CW92
    `--G. williamsi Howden 1992G09
  Elephastomus Macleay 1819CW92
    |--*E. proboscideus (Schreibers 1802) [=Scarabaeus proboscideus]CW92
    |    |--E. p. proboscideus [incl. Bolboceras australasiae Kirby 1818]CW92
    |    `--E. p. kirbyanus Carne 1965 (see below for synonymy)CW92
    |--E. carnei Krikken 1976CW92
    |--E. gellarus Carne 1965CW92
    |--E. howdeni Nikolajev 1990CW92
    |--E. meraldus Carne 1965CW92
    `--E. terraereginae (Blackburn 1899) [=Bolboceras terraereginae]CW92
  Stenaspidius Westwood 1848CW92
    |--*S. nigricornis (Westwood 1848) [=Bolboceras (*Stenaspidius) nigricornis]CW92
    |--S. albosetosus Howden 1974CW92
    |--S. allsoppi Howden 1976CW92
    |--S. brittoni Howden 1974CW92
    |--S. houstoni Howden 1993G09
    |--S. lividus Howden 1992G09
    |--S. matthewsi Howden 1974CW92
    |--S. ruficornis (Boucomont 1906) [=Bolboceras (Stenaspidius) ruficornis]CW92
    `--S. spatuliferus Howden 1992G09
  Bolboleaus Howden & Cooper 1977CW92
    |--*B. truncatus (Blackburn 1904) [=Bolboceras truncatum]CW92
    |--B. froggatti (Blackburn 1904) [=Bolboceras froggatti]CW92
    |--B. hiaticollis Howden 1985CW92
    |--B. houstoni Howden 1985CW92
    |--B. ingens (Macleay 1888) [=Bolboceras ingens]CW92
    |--B. mimus Howden 1985CW92
    |--B. parvicollis Howden 1985CW92
    |--B. propinquus Howden 1985CW92
    |--B. quadriarmigerus (Howden 1954) (see below for synonymy)CW92
    |--B. quadrifoveatus (Lea 1919) [=Bolboceras quadrifoveatum]CW92
    |--B. storeyi Howden 1985CW92
    |--B. tenax (Blackburn 1904) [=Bolboceras tenax]CW92
    |--B. trifoveicollis (Lea 1916) [=Bolboceras trifoveicolle]CW92
    `--B. variolicollis (Lea 1915) [=Bolboceras variolicolle]CW92
  BolborhachiumCW92
  BlackbolbusCW92
  BlackburniumH79
  AustralobolbusCW92
  Bolbocerosoma Schaeffer 1906J02
    `--B. hamatumAT01
  Bolbelasmus Boucomont 1911 [incl. Kolbeus Boucomont 1911]J02
  Bolborhombus Cartwright 1953J02
  Bolbocerastes Cartwright 1953J02
  Bradycinetulus Cockerell 1906 [incl. Amechanus Horn 1870, Bradycinetus Horn 1871 non Sars 1866]J02

Bolboleaus quadriarmigerus (Howden 1954) [=Blackburnium quadriarmigerum, Bolboceras armigerum Macleay 1871 non Bo. armiger Scopoli 1772]CW92

Elephastomus proboscideus kirbyanus Carne 1965 [=Bolboceras (Elephastomus) kirbyi Westwood 1848 non Bainbridge in Hope 1841, B. kirbii]CW92

*Type species of generic name indicated

References

[AT01] Arnett, R. H., Jr & M. C. Thomas (eds) 2001. American Beetles vol. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC Press: Boca Raton.

[CW92] Cassis, G., & T. A. Weir. 1992. Geotrupidae. In: Houston, W. W. K. (ed.) Zoological Catalogue of Australia vol. 9. Coleoptera: Scarabaeoidea pp. 41–64. AGPS Press: Canberra.

[G09] Golding, M. R. 2009. A Pictorial Field Guide to the Beetles of Australia. Part 4: Bolboceratinae. Privately published.

[H79] Howden, H. F. 1979. A revision of the Australian genus Blackburnium Boucomont (Coleoptera: Scarabaeidae: Geotrupinae). Australian Journal of Zoology, Supplementary Series 72: 1–88.

[J02] Jameson, M. L. 2002. Geotrupidae 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. 23–27. CRC Press.

[L96] Lindquist, E. E. 1996. Antiquity of some symbiotic associations of heterostigmatic mite clades with beetle clades: phylogenetic and biogeographic patterns. In: Mitchell, R., D. J. Horn, G. R. Needham & W. C. Welbourn (eds) Acarology IX vol. 1. Proceedings pp. 301–303. Ohio Biological Survey: Columbus (Ohio).

[M86] 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.

[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.

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