RT Journal Article SR Electronic T1 Compositional phylogenomic modelling resolves the ‘Zoraptera problem’: Zoraptera are sister to all other polyneopteran insects JF bioRxiv FD Cold Spring Harbor Laboratory SP 2021.09.23.461539 DO 10.1101/2021.09.23.461539 A1 Erik Tihelka A1 Michael S. Engel A1 Jesus Lozano-Fernandez A1 Mattia Giacomelli A1 Ziwei Yin A1 Omar Rota-Stabelli A1 Diying Huang A1 Davide Pisani A1 Philip C.J. Donoghue A1 Chenyang Cai YR 2021 UL http://biorxiv.org/content/early/2021/09/24/2021.09.23.461539.abstract AB The evolution of wings propelled insects to their present mega-diversity. However, interordinal relationships of early-diverging winged insects and the timescale of their evolution are difficult to resolve, in part due to uncertainties in the placement of the enigmatic and species-poor order Zoraptera. The ‘Zoraptera problem’ has remained a contentious issue in insect evolution since its discovery more than a century ago. This is a key issue because different placements of Zoraptera imply dramatically different scenarios of diversification and character evolution among polyneopteran. Here, we investigate the systematic placement of Zoraptera using the largest protein-coding gene dataset available to date, deploying methods to mitigate common sources of error in phylogenomic inference, and testing historically proposed hypotheses of zorapteran evolution. We recover Zoraptera as the earliest-diverging polyneopteran order, while earwigs (Dermaptera) and stoneflies (Plecoptera) form a monophyletic clade (Dermoplectopterida) sister to the remainder of Polyneoptera. The morphology and palaeobiology of stem-zorapterans are informed by Mesozoic fossils. The gut content and mouthparts of a male specimen of Zorotypus nascimbenei from Kachin amber (Cretaceous) reveal a fungivorous diet of Mesozoic zorapterans, akin to extant species. Based on a set of 42 justified fossil and stratigraphic calibrations, we recover a Devonian origin of winged insects and Polyneoptera, suggesting that these groups coincided with the rise of arborescence during the diversification of early terrestrial plants, fungi, and animals. Our results provide a robust framework for understanding the pattern and timescale of early winged insect diversification.Competing Interest StatementThe authors have declared no competing interest.