A phylogeny of the families of Scarabaeoidea (Coleoptera)

A study, based on examination of thirteen scarabaeoid families, was made of 134 adult and larval characters from the following character suites: 105 a dult characters of the antennae, eye, epipharynx, mandible, maxillae, labiu m, tentorium, trochantin, procoxae, mesocoxae, mesothoracic spiracles, hind wing articulation hind wing base, hind wing venation, hind wing folding, a bdominal sternites, abdominal spiracles, male genitalia, ovarioles and kary otype; twenty larval characters of the antennae, fronto-clypeal suture, ste mmata, labial palpi, maxillae, mandibles, legs, stridulatory apparatus, spi racles and ecdysial process; and nine adult and larval biological character s. In order to assess the reliability of different characters in resolving scarabaeoid family relationships, six data sets were subjected to cladistic analysis: the total evidence character set (134 characters), restricted ad ult character set (thirty-two characters, not including those of the wings) , wing character set (seventy-three characters), larval character set (twen ty characters), biological character set (nine characters) and re-coded How den (1982) character set (thirty-nine characters). The complete character s et and wing character set both produced phylograms with all nodes resolved; the restricted adult data set, larval data set, Howden (1982) data set and biological data set produced phylograms with diminishing levels of node re solution. The reconstructed phylogeny, from the preferred phylogram of the total evidence character set, shows that the Scarabaeoidea comprises three major lineages; a glaresid, passalid and scarabaeid lineage. The glaresid l ineage consists only of the Glaresidae. The passalid lineage comprises two major lines; a glaphyrid Line (containing Glaphyridae, Passalidae, Lucanida e, Diphyllostomatidae, Trogidae, Bolboceratidae and Pleocomidae) and a geot rupid line (containing Geotrupidae, Ochodaeidae, Ceratocanthidae and Hyboso ridae). The scarabaeid lineage contains those taxa traditionally included w ithin the Scarabaeidae (Aphodiinae, Scarabaeinae, Orphninae, Melolonthinae, Acoma, Chasmatopterinae, Hopliinae, Oncerinae, Rutelinae, Dynastinae, Tric hiinae, Cetoniinae and Valginae).