Ichneumonoidea phylogenomics using anchored target enrichment

Subfamily relationships of the Vespidae were inferred through analysis of phenotypic characters (269 morphological and 66 behavioral) and molecular data (CO1, 28S, 16S, 12S) across 74 taxa. Maximum-Likelihood, Bayesian Inference, and Maximum Parsimony were employed and analyses were done for individual and combined subsets of data. This approach allowed us to evaluate incongruence among different subsets of data (i.e. biological drivers of conflict) and the effect of methodological choice on estimation. Simultaneous analysis of all evidence supports a (Stenogastrinae + (Polistinae + Vespinae)) clade and thus a single origin of eusociality. Phylogenies derived strictly from molecular evidence consistently support a dual origin of eusociality. We argue that previous studies recovering diphyly of eusocial wasps may withstand the effects of low taxon sampling, suboptimal alignments and a two-step approach causing phylogenetic error; we achieved similar topologies using ~three-fold more terminal ingroups and a different suite of loci. Three aspects of the tree are incongruent when comparing molecular and phenotypic evidence: the placement of Stenogastrinae; the relationship between Euparagiinae and Masarinae; and Polistinae tribal relationships. We conclude that vespid subfamily relationships are obfuscated by the fact that phenotypic and molecular evidence are at odds. This study highlights the confounding effects of biological processes on tree reconstruction and the importance of elucidating drivers of conflict when formulating phylogenetic hypotheses. A current working hypothesis is that a period of rapid adaptive divergence characterizes the early history of Vespidae, resulting in random sorting of ancestral polymorphisms for both molecular and phenotypic characters.