Larvae of Ceratocanthidae and Hybosoridae (Coleoptera: Scarabaeoidea): Study of morphology, phylogenetic analysis and evidence of paraphyly of Hybosoridae

Larvae of the scarabaeoid genera Germarostes Paulian, Cyphopisthes Gestro, Paulianostes Ballerio, Ceratocanthus White, Pterorthochaetes Gestro, Madrasostes Paulian, Astaenomoechus Martı ́nez & Pereira (Ceratocanthidae) and Hybosorus Macleay, Phaeochrous Castelnau, and Anaides Westwood (Hybo- soridae) are described, keyed and illustrated with fifty-seven drawings. A phylo- genetic analysis of these two families based on larval morphology is presented. Fifty-four larval morphological and three biological characters from twenty-seven taxa revealed nineteen equally parsimonious cladograms. The monophyly of (Ceratocanthidae þ Hybosoridae) is supported by four unambiguous unique syna- pomorphies: dorsal medial endocarina on cranium extended anteriorly into fron- tal sclerite; presence of large membranous spot on apical antennomere; labium dorsally with four pores in centre (secondarily reduced to two pores in some groups); and presence of stridulatory organ on fore- and middle legs (secondarily reduced in some groups). Our analysis suggests that the family Hybosoridae is paraphyletic with respect to Ceratocanthidae. The clade comprising the hybosorid genera Hybosorus and Phaeochrous is the sister group of the remaining Hybo- soridae plus Ceratocanthidae. It is supported by two unambiguous synapomorphies: two apical antennomeres completely joined and the stridulatory organ represented by seven to nine large teeth anteriorly on the middle leg. The hybosorid genus Anaides is a sister group to the remaining Hybosoridae plus Ceratocanthidae (without Hybosorus and Phaeochrous) and the ceratocanthid genus Germarostes is a sister group to the remaining Hybosoridae plus Ceratocanthidae (without Hybosorus, Phaeochrous and Anaides). The ceratocanthid genera Cyphopisthes, Astaenomoechus, Paulianostes, Pterorthochaetes, and Madrasostes constitute a sister group to the hybosorid genus Cryptogenius and are supported by the presence of two reversions: two dorsal pores on labium and completely reduced stridulatory organs on fore- and middle legs.