First comprehensive higher level phylogeny of Zygaenidae (Lepidoptera) including estimated ages of the major lineages and a review of known zygaenid fossils

Zygaenidae, also known as burnet, forester, smoky, or leaf skeletonizer moths, are a family of mainly diurnal moths well known for their aposematic colouration and the ability to release hydrogen cyanide as a defence mechanism. So far, few attempts have been made to understand the evolutionary history of the global zygaenid fauna. Here, we inferred the most comprehensive molecular phylogeny for Zygaenidae to date and estimated the lineage timing-of-divergence with a Bayesian approach. Building on earlier work, we significantly increased the taxon and gene sampling for the family, which here included data from 30 gene fragments, recovered from public databases or newly sequenced, for almost 30% of the species representing 92 genera (49%) and all five subfamilies. We recovered strong support for the monophyly of Zygaenidae, Chalcosiinae, and Zygaeninae. Procridinae were recovered as monophyletic with low support, whereas the monophyly of Callizygaeninae remains untested as we sampled only one of the two genera. In the core dataset, we recovered Procridinae as sister to Callizygaeninae + Chalcosiinae. This large clade is the sister lineage to Zygaeninae. The position of Inouelinae could not be resolved. The lineage leading to the extant Zygaenidae appears to have diverged in Late Cretaceous (ca. 86 Ma), while the divergence among the subfamilies occurred several million years before the Cretaceous-Paleogene mass extinction event (ca. 66 Ma). Additionally, we provide a review of known fossil Zygaenidae as Appendix S1. Our results form a strong basis for future studies of zygaenid biosystematics, including their ecology, evolution, and behaviour. Using 30 gene fragments and sampling nearly 50% of the genera, we provide a well-resolved phylogeny of Zygaenidae; Zygaenidae are recovered as monophyletic as are most of the subfamilies. The lineage leading to the extant Zygaenidae diverged in Late Cretaceous, while the divergence among the subfamilies occurred a little before the Cretaceous-Paleogene mass extinction event. In the Appendix S1, we provide a re-evaluation of all known zygaenid fossils and an assessment of whether observable morphological features warrant confident assignment of these specimens to Zygaenidae. image