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A phylogenetic study was conducted to hypothesize relationships of most of the genera of the   Oniticellini   and   Onthophagini   for the first time using morphological characters from a diverse array of external and internal sclerites. The monophyly and sister relationship of both tribes was found using Bayesian and parsimony analyses with heavily to moderately weighted data. An alternative hypothesis based on parsimony analyses of unweighted or slightly weighted data show a paraphyletic   Oniticellini   without the   Onthophagini  , although recognition of the subtribe   Helictopleurina   as a tribe would eliminate non-monophyly.Of the three   Oniticellini   subtribes, the   Helictopleurina   and   Drepanocerina   are monophyletic. There is no support for the monophyly of the   Oniticellina   or the   Onthophagini   subtribe   Alloscelina  , as currently defined. The genus    Liatongus    is paraphyletic, while strong support was found for monophyly of the Madagascan genus,    Helictopleurus   . The genus    Onthophagus    is never monophyletic in any analysis performed. Two new subtribes are also proposed:   Liatongina   subtr. n.  including the genus    Liatongus    and   Attavicina   subtr. n.  including the genera   Attavicinus   and   Paroniticellus  .Topological evidence shows that the ancestral oniticellines and onthophagines were all coprophagous with alternative food sources evolving relatively recently. Both myrmecophily and termitophily probably evolved only once in the onthophagines. The phylogenetic analysis supports an African origin for the two tribes, with a relatively early age for the split of the Madagascar helictopleurines from the remaining oniticellines via dispersal. Furthermore, the presence of the oniticellines in the New World is hypothesized to be due to two relatively old dispersal events via Beringia and two relatively recent trans-Atlantic invasions of the Caribbean.

Phylogeny of the Oniticellini and Onthophagini dung beetles ( Scarabaeidae , Scarabaeinae ) from morphological evidence