Expressed sequence tags reveal Proctotrupomorpha (minus Chalcidoidea) as sister to Aculeata (Hymenoptera: Insecta)

Hymenoptera is one of the most diverse groups of animals on the planet and have vital importance for ecosystem function as pollinators and parasitoids. Higher-level relationships among Hymenoptera have been notoriously difficult to resolve with both morphological and traditional molecular approaches. Here we examined the utility of expressed sequence tags for resolving relationships among hymenopteran superfamilies. Transcripts were assembled for 6 disparate Hymenopteran taxa with additional sequences added from public databases for a final dataset of 24 genes for 16 taxa and over 10 kb of sequence data. The concatenated dataset recovered a robust and well-supported topology demonstrating the monophyly of Holometabola, Hymenoptera, Apocrita, Aculeata, Ichneumonoidea, and a sister relationship between the two most closely related proctotrupomorphs in the dataset (Cynipoidea + Proctotrupoidea). The data strongly supported a sister relationship between Aculeata and Proctotrupomorpha, contrary to previously proposed hypotheses. Additionally there was strong evidence indicating Ichneumonoidea as sister to Aculeata + Proctotrupomorpha. These relationships were robust to missing data, nucleotide composition biases, low taxonomic sampling, and conflicting signal across gene trees. There was also strong evidence indicating that Chalcidoidea is not contained within Proctotrupomorpha.