Mitogenome sequences stabilize the phylogenetics of weevils (Curculionoidea) and establish the monophyly of larval ectophagy

The weevils and their relatives (superfamily Curculionoidea) constitute a huge radiation of beetles, but basal relationships in this group remain controversial, in particular within the largest family, Curculionidae. We used next-generation sequencing to generate mitochondrial genome data comprising 12 protein coding genes for 27 taxa. Together with two published ingroup sequences, (sub)family relationships were assessed with Bayesian and ML searches under various models and partitioning schemes. Forward and reverse strands of the mitochondrial genome differed in nucleotide skew. Consequently synonymous codon usage differed substantially on either strand for each amino acid, whereby codons ending in AT and GC were favored on the forward and reverse strands, respectively. Data partitioning by forward/reverse strand and codon position greatly improved likelihood scores and nodal support, whereas the tree topology was largely stable. The analysis generally supports the basal position of several 'orthocerus' lineages with straight antennae and male genitalia of an ancestral type; a paraphyletic mixed group of Heteromorphi exhibiting mixed ancestral and derived antennal and genitalic characters; and the derived 'gonathocerous' lineages with kinked antennae corresponding to the strongly supported Curculionidae. The latter did not include the wood boring Platypodinae that was recovered as sister to Dryophthoridae, while the Scolytinae and Cossoninae formed two independent lineages of wood borers within Curculionidae. A basal split in Curculionidae placed the Entiminae and Hyperinae as sister to all other subfamilies with high support, which provides a new ecological concept for structuring the Curculionidae according to the ectophagous larval life style in the former versus endophagous larvae in all others. This basal split is also supported by gene order rearrangements in a tRNA cluster. Recent studies supporting the monophyly of wood boring weevils may be attributable to long-branch attraction, as molecular rates in their mitochondrial genomes were found to be higher than in other lineages, but this did not confound tree searches under combined analysis of mitochondrial protein coding genes.