Phylogeny of the families of Scarabaeoidea (Coleoptera) based on characters of the hindwing articulation, hindwing base and wing venation

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Abstract
A study is made of the articulation and base of the hindwings of Scarabaeoidea. The survey is based on an examination of over 250 genera from thirteen scarabaeoid families. Relationships among all families of Scarabaeoidea are examined here for the first time. The reconstructed phylogeny shows that the Scarabaeoidea is comprised of three major lineages: a glaresid, passalid and scarabaeid lineage. The glaresid lineage consists only of the Glaresidae. The passalid lineage is comprised of two major lines: a passalid line (containing Passalidae, Lucanidae, Diphyllostomatidae, Glaphyridae, Trogidae, Bolboceratidae and Pleocomidae) and a geotrupid line (containing Geotrupidae, Ochodaeidae, Ceratocanthidae and Hybosoridae). The scarabaeid lineage contains those taxa traditionally included within the Scarabaeidae (Aegialiinae, Aulonocneminae, Aphodiinae, Scarabaeinae, Orphninae, Melolonthinae,Acoma, Chasmatopterinae, Hopliinae, Oncerinae, Rutelinae, Dynastinae, Trichiinae, Cetoniinae and Valginae). Additional evidence, from other character suites, supporting this breakdown of the Scarabaeoidea is given.