Hybosoridae Erichson, 1847

Costa-Silva, Vinícius da, Ferreira, André da Silva, Bordin, Bruna R., Basílio, Daniel S., Rodrigues, Diego F., Gama, Emanuel R. R., Fuhrmann, Juares, Mariano, Júlia, Bevilaqua, Marcus, Cherman, Mariana A., Duarte, Paulo R. M., Grossi, Paschoal C. & Vaz-de-Mello, Fernando Z., 2024, Brazilian Scarabaeoidea (Insecta: Coleoptera) in the Taxonomic Catalogue of the Brazilian Fauna, with a key for families and subfamilies, Zoologia (e 23075) 41, pp. 1-37 : 11

Hybosoridae Erichson, 1847

Fig. 4A–D

Diagnosis. Hybosoridae are a morphologically heterogeneous family. Most species are commonly recognized by the presence of a prominent mandible and antennal club formed by three antennomeres, with the basal antennomere cupuliform, sheltering the other two ( Ocampo 2006 a, BasÍlio et al. 2023). The species of Ceratocanthinae do not present this antennal pattern, but they can be easily identified either by having a body capable of conglobation ( Ceratocanthini ) or by being very small in size (less than 6 mm) and having a transverse and deep excavation in the pronotum (Ivieolini and Scarabatermitini ) ( Ballerio and Grebennikov 2016). The only common characteristic to all Hybosoridae is the tarsal insertion before the apex of the protibia (BasÍlio et al. 2023).

Remarks. Hybosoridae have a worldwide distribution, comprising 96 genera and about 723 species distributed in six subfamilies, one extinct ( Mimaphodiinae ) and five extant ( Anaidinae , Ceratocanthinae, Hybosorinae , Liparochrinae , and Pachyplectrinae ) (BasÍlio et al. 2023). In Brazil, there are currently 19 genera and 85 species recorded in four of the five extant subfamilies – except Liparochrinae ( Table 2) ( BasÍlio and Vaz-de-Mello 2023). However, the number of genera and species, both in Brazil and worldwide, has been constantly increasing. Species of Hybosoridae have been recorded in almost all Brazilian states, and it is likely that there are species of this family in the five states that lack records so far (Tocantins, PiauÍ, Rio Grande do Norte, Alagoas, and Sergipe).

Hybosoridae were historically classified as a subfamily of Scarabaeidae (i.e., Hybosorinae and Ceratocanthinae) ( Paulian 1982, Allsopp 1984, Howden and Gill 2000). However, even after Hybosoridae being elevated to the rank of family (with Ceratocanthinae treated as a subfamily of Hybosoridae ) and being consensually accepted, these two taxa continued to be treated as separated families ( Table 1) ( Paulian 1988). In 2004, morphological data from larvae and adults recovered Hybosoridae with the inclusion of Ceratocanthidae , which started to be treated as Ceratocanthinae ( Grebennikov et al. 2004) . This classification is currently the most accepted, and is supported by morphological ( Ocampo 2006b, Ballerio and Grebennikov 2016) and molecular data ( Ocampo and Hawks 2006, Grebennikov and Smith 2021).

Regarding their biology, they can be necrophagous, coprophagous, fungivores, or feed on rotting wood. Many species have the habit of burying themselves, and some have stridulatory behavior. Associations with ants and termites have also been recorded ( Jameson 2002b, Ocampo 2006 b, BasÍlio et al. 2023). Representatives of Ceratocanthinae and Liparochrinae have the ability to roll their bodies into a ball ( Ocampo 2006b, Ballerio and Grebennikov 2016). Dichotomous keys have been proposed to identify genera of Anaidinae ( Ocampo 2006b) , Ceratocanthinae ( Paulian 1982, Ballerio and Grebennikov 2016), and Neotropical Hybosorinae ( BasÍlio et al. 2022) .