The parasitoid wasp family Ichneumonidae is among the most diverse groups of organisms, with conservative estimates suggesting that it contains more species than all vertebrates together. However, ichneumonids are also among the most severely understudied groups, and our understanding of their evolution is hampered by the lack of a robust higher-level phylogeny of this group. Based on newly generated transcriptome sequence data, which were filtered according to several criteria of phylogenetic informativeness, we developed target DNA enrichment baits to capture 93 genes across species of Ichneumonidae. The baits were applied to DNA of 55 ichneumonids, with a focus on Pimpliformes, an informal group containing nine subfamilies. Phylogenetic trees were inferred under maximum likelihood and Bayesian approaches, at both the nucleotide and amino acid levels. We found maximum support for the monophyly of Pimpliformes but low resolution and very short branches close to its base, strongly suggesting a rapid radiation. Two genera and one genus-group were consistently recovered in unexpected parts of the tree, prompting changes in their higher-level classification: Pseudorhyssa Merrill, currently classified in the subfamily Poemeniinae, is transferred to the tribe Delomeristini within Pimplinae, and Hemiphanes Förster is moved from Orthocentrinae to Cryptinae. Likewise, the tribe Theroniini is resurrected for the Theronia group of genera (stat. rev.). Phylogenetic analyses, in which we gradually increased the numbers of genes, revealed that the initially steep increase in mean clade support slows down at around 40 genes, and consideration of up to 93 genes still left various nodes in the inferred phylogenetic tree poorly resolved. It remains to be shown whether more extensive gene or taxon sampling can resolve the early evolution of the pimpliform subfamilies.

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Parasitoid wasps of the family Ichneumonidae are one of the most diverse and species-rich groups of organisms with a worldwide distribution. We here describe seven new ichneumonid fossil species and two new genera from a remarkable insect fossil site, the Eocene Messel Pit in Germany (~47Ma). The unique fossil preservation allows us to place five out of the seven new species unequivocally in extant subfamilies and genera. For the first time, lobed claws which are a clear synapomorphy for the subfamily Pimplinae, are observed in a fossil, making the newly described Scambus fossilobus sp. nov. the oldest unequivocal representative of the group. We also describe a fossil of Labeninae (Trigonator macrocheirus gen. et sp. nov.), an ichneumonid subfamily that was until now believed to be an exclusively Gondwanan element. Furthermore, the newly described Rhyssella vera sp. nov., Xanthopimpla messelensis sp. nov., and X. praeclara sp. nov. provide evidence that these extant genera date back as far as the Early/Middle Eocene. In contrast to the clear placement of most of the newly described species, we were unable to place Polyhelictes bipolarus gen. et sp. nov. and Mesornatus markovici gen. et sp. nov. in an ichneumonid subfamily, mostly due to the high levels of homoplasy found in this group. These findings on the one hand demonstrate the need for a more rigorous approach in the taxonomic placement of fossil ichneumonids, and on the other hand provide more precise minimum ages for several ichneumonid genera and subfamilies.

Taxon sampling is a central aspect of phylogenetic study design, but it has received limited attention in the context of total-evidence dating, a widely used dating approach that directly integrates molecular and morphological information from extant and fossil taxa. We here assess the impact of commonly employed outgroup sampling schemes and missing morphological data in extant taxa on age estimates in a total-evidence dating analysis under the uniform tree prior. Our study group is Pimpliformes, a highly diverse, rapidly radiating group of parasitoid wasps of the family Ichneumonidae. We analyze a data set comprising 201 extant and 79 fossil taxa, including the oldest fossils of the family from the Early Cretaceous and the first unequivocal representatives of extant subfamilies from the mid Paleogene. Based on newly compiled molecular data from ten nuclear genes and a morphological matrix that includes 222 characters, we show that age estimates become both older and less precise with the inclusion of more distant and more poorly sampled outgroups. These outgroups not only lack morphological and temporal information, but also sit on long terminal branches and considerably increase the evolutionary rate heterogeneity. In addition, we discover an artefact that might be detrimental for total-evidence dating: “bare-branch attraction”, namely high attachment probabilities of certain fossils to terminal branches for which morphological data are missing. Using computer simulations, we confirm the generality of this phenomenon and show that a large phylogenetic distance to any of the extant taxa, rather than just older age, increases the risk of a fossil being misplaced due to bare-branch attraction. After restricting outgroup sampling and adding morphological data for the previously attracting, bare branches, we recover a Jurassic origin for Pimpliformes and Ichneumonidae. This first age estimate for the group not only suggests an older origin than previously thought, but also that diversification of the crown group happened well before the Cretaceous-Paleogene boundary. Our case study demonstrates that in order to obtain robust age estimates, total-evidence dating studies need to be based on a thorough and balanced sampling of both extant and fossil taxa, with the aim of minimizing evolutionary rate heterogeneity and missing morphological information.

The fossil record constitutes the primary source of information about the evolutionary history of extant and extinct groups, and many analyses of macroevolution rely on fossils that are accurately placed within phylogenies. To avoid misinterpretation of the fossil record, especially by non-palaeontologists, the proper assessment and communication of uncertainty in fossil placement is crucial. We here use Bayesian morphological phylogenetics to evaluate the classifications of fossil parasitoid wasps (Hymenoptera, Ichneumonidae) and introduce ‘RoguePlots’ to illustrate placement uncertainty on the phylogeny of extant taxa. Based on an extensive, newly constructed morphological matrix of 222 characters in 24 fossil and 103 extant taxa, we test three different aspects of models of morphological evolution. We find that a model that includes ordered characters, among-character rate variation, and a state-space restricted to observed states achieves the highest marginal likelihoods. The individual RoguePlots reveal large differences in confidence in the placement of the different fossils and allow some refinements to their classification: Polyhelictes bipolarus and Ichninsum appendicrassum are moved from an uncertain subfamily placement to Pimplinae, Plectiscidea lanhami is transferred to Allomacrus in Cylloceriinae ( Allomacrus lanhami , comb. nov.), Lithotorus cressoni is moved from Diplazontinae to Orthocentrinae, and we note uncertainty in the generic placements of Rhyssella ? vera and Xanthopimpla ? messelensis . We discuss potential artefacts that might result in biased posterior probabilities in Bayesian morphological phylogenetic analyses, pertaining to character and taxon sampling, fossilization biases, and model misspecification. Finally, we suggest future directions both in ichneumonid palaeontology and in the way RoguePlots can improve both assessment and representation of placement uncertainty, both in fossils and other rogue taxa.

The parasitoid wasp family Ichneumonidae is arguably one of the groups for which current knowledge lags most strongly behind their enormous diversity. In a five-day meeting in Basel (Switzerland) in June 2019, 22 researchers from 14 countries met to discuss the most important issues in ichneumonid research, including increasing the speed of species discovery, resolving higher-level relationships, and studying the radiation of these parasitoids onto various host groups through time. All agreed that it is time to advertise ichneumonid research more broadly in the scientific community and thereby attract young talents to this group for which specialists are sorely lacking. In order to popularize the group, we here suggest a new name for the family, “Darwin wasps”, to reflect the pivotal role they played in convincing Charles Darwin that not all of creation could have been created by a benevolent god. We hope that the name catches on, and that Darwin wasps start buzzing more loudly across all disciplines of biology.

Calibrating the molecular clock is the most contentious step in every dating analysis, but the emerging total-evidence dating approach promises increased objectivity. It combines molecular and morphological data of extant and fossil taxa in a Bayesian framework.Information about absolute node ages stems from the inferred fossil placements and associated branch lengths, under the assumption of a morphological clock. We here use computer simulations to assess the impact of mismatch of the morphology model, such as misspecification of character states and transition rates, non-stationarity of the evolutionary process, and extensive variation of evolutionary rates among branches. Comparisons with published datasets suggest that, at least for evolutionary rates typically observed in discrete morphological characters, the total-evidence dating framework is surprisingly robust to these factors. We show that even with relatively low numbers of morphological characters sampled, extensive model mismatch is mostly irrelevant for the performance of the method.The only exception we found are cases of highly asymmetric state frequencies and thus transition rates, but these can be accounted for by appropriate morphology models. In contrast, we find that the temporal scope of fossil sampling has a major impact on divergence time estimates, with the time signal quickly eroding if only rather young fossils are included in an analysis. Our results suggest that total-evidence dating might work even without a good understanding of morphological evolution and that study design should instead focus on an adequate sampling of all relevant fossils, even those with highly incomplete preservation.

Ichneumonidae constitute one of the largest families of organisms, but their fossil record is severely understudied, which precludes inferences about their early evolutionary history. We here describe a new genus and two new species of ichneumonids from the Early Eocene Tadushi Formation (Russian Far East). The new taxa are Crusopimpla tadushensis gen. et sp. nov. and Dolichomitus? saxeus sp. nov. Pimpla rediviva (Brues) is redescribed and placed in the genus Crusopimpla. This new genus is unique within Pimplinae in having a completely developed propodeal carination and probably represents one of its most basal lineages. Using a phylogenetic hypothesis of the subfamily, we discuss potential pathways of morphological evolution in the group, which will provide a basis for future placement of pimpline fossil taxa.