Taxonomic review of immature dung beetles of the subfamily Scarabaeinae (Coleoptera: Scarabaeidae)

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Abstract
Discussion of the taxonomy of scarabaeine larvae includes: (i) a redescription of the subfamily Scarabaeinae based upon larval characters, (ii) descriptions of known genera based upon larvae, and (iii) a key to known genera. Twenty‐five genera representing each of the six currently recognized tribes and most subtribes are considered:Onthophagus, Liatongus, Oniticellus, Onitis, Bubas, Dichotomius, Ontherus, Ateuchus, Scatimus, Canthidium, Neocanthidium, Copris, Heliocopris, Phanaeus, Coprophanaeus, Sulcophanaeus, Sisyphus, Gymnopleurus, Scarabaeus, Megathopa, Canthon. Deltochilum, Canthochilum, Arachnodes and Eurysternus.Four others are treated briefly:Chironitis, Synapsis, Catharsius, Nesosisyphus.On morphological and other grounds,ScatimusandHeliocoprisare removed from the subtribe Coprina and reassigned to Dichotomiina. There is little about the variation among larvae studied which supports or weakens the current tribal classification based upon adults as there emerge no combinations of larval characters which satisfactorily define generic groupings. Discussion of scarabaeine pupae is limited to description of pupal support projections of known genera and to definition of the support system characteristic of the subfamily, which is functionally related to nesting behaviour culminating in metamorphosis within a spheroid pupation chamber.