The onset of phylogenomics has contributed to the resolution of numerous challenging evolutionary questions while offering new perspectives regarding biodiversity. However, in some instances, analyses of large genomic datasets can also result in conflicting estimates of phylogeny. Here, we present the first phylogenomic scale study of a dipteran parasitoid family, built upon anchored hybrid enrichment and transcriptomic data of 240 loci of 43 ingroup acrocerid taxa. A new hypothesis for the timing of spider fly evolution is proposed, wielding recent advances in divergence time dating, including the fossilized birth-death process to show that the origin of Acroceridae is younger than previously proposed. To test the robustness of our phylogenetic inferences, we analyzed our datasets using different phylogenetic estimation criteria, including supermatrix and coalescent-based approaches, maximum-likelihood and Bayesian methods, combined with other approaches such as permutations of the data, homogeneous versus heterogeneous models, and alternative data and taxon sets. Resulting topologies based on amino acids and nucleotides are both strongly supported but critically discordant, primarily in terms of the monophyly of Panopinae. Conflict was not resolved by controlling for compositional heterogeneity and saturation in third codon positions, which highlights the need for a better understanding of how different biases affect different data sources. In our study, results based on nucleotides were both more robust to alterations of the data and different analytical methods and more compatible with our current understanding of acrocerid morphology and patterns of host usage.

Fossils not only provide unique opportunity to understand the "tempo and mode" of evolution but are essential for modeling lineage-contingent diversification histories. Here, we interrogate the Mesozoic fossil record of the Aculeata, with emphasis on the ants (Formicidae), and conduct an extended series of ancestral state estimation exercises on distributions of tip-dated combined-evidence phylogenies. We developed and illustrated from ground-up a set of 576 morphological characters which we scored for 144 extant and 431 fossil taxa; we used average posterior probability support to filter this to a target matrix of 303 taxa, for which we integrated strongly filtered ultraconserved element (UCE) data for 115 living species. We also implemented reversible jump MCMC (rjMCMC) and hidden state methods to model complex behavioral characters to test hypotheses about the pathway to obligate eusociality. In addition to revising the higher classification of all sampled groups to family or subfamily level using estimated character polarities to diagnose nodes across the phylogeny, we find that the mid-Cretaceous genera †Camelomecia and †Camelosphecia form a clade which is robustly supported as sister to the total clade Formicidae. For this reason, we name this extinct clade as †@@@idae fam. nov. and provide a definition for the expanded Formicoidea. Based on our results, we recognize three major phases in the early evolution of the ants: (1) origin of ants as running-adapted huntresses during the Late Jurassic in the "stinging aggressor" guild (Aculeata) among various lineages of "sneaking parasitoids" (non-aculeate Vespina); (2) the first formicoid radiation during the Early Cretaceous, by the end of which all major extant ant linages had originated; and (3) turnover of the Formicoidea at the end-Cretaceous leading to the second formicoid radiation, i.e., the Cenozoic formicid diversification. We conclude with a concentrated series of considerations for future directions of study with this dataset and beyond.

Scoliid wasps comprise a clade of aculeate insects whose larvae are parasitoids of scarabaeid beetle grubs. While scoliids have been studied and used as biological control agents, research into the group's evolution, as well as the stability of scoliid taxonomy, has been limited by a lack of reliable phylogenies. We use ultraconserved element (UCE) data under concatenation and the multispecies coalescent to infer a phylogeny of the Scoliidae. In order to mitigate potential issues arising from model misspecification, we perform data filtering experiments using posterior predictive checks and matched-pairs tests of symmetry. Our analyses confirm the position of Proscolia as sister to all other extant scoliids. We also find strong support for a sister group relationship between the campsomerine genus Colpa and the Scoliini, rendering the Campsomerini non-monophyletic. Campsomerini excluding Colpa (hereafter Campsomerini sensu stricto) is inferred to be monophyletic, with the Australasian genus Trisciloa recovered as sister to the remaining members of the group. Many sampled genera, including Campsomeriella, Dielis, Megascolia, and Scolia are inferred to be non-monophyletic. Analyses incorporating fossil data indicate an Early Cretaceous origin of the crown Scoliidae, with the split between Scoliini + Colpa and Campsomerini s.s. most probably occurring in the Late Cretaceous. Posterior means of Scoliini + Colpa and Campsomerini s.s. crown ages are estimated to be in the Paleogene, though age 95% HPD intervals extend slightly back past the K-Pg boundary, and analyses including fossils of less certain placement result in more posterior mass on older ages. Our estimates of the stem ages of Nearctic scoliid clades are consistent with dispersal across Beringia during the Oligocene or later Eocene. Our study provides a foundation for future research into scoliid wasp evolution and biogeography by being the first to leverage genome-scale data and model-based methods. However, the precision of our dating analyses is constrained by the paucity of well-preserved fossils reliably attributable to the scoliid crown group. Despite concluding that the higher-level taxonomy of the Scoliidae is in dire need of revision, we recommend that taxonomic changes be predicated on datasets that extend the geographic and taxonomic sampling of the current study.