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.
This study presents the first molecular phylogenetic analysis of the Clinocerinae, challenging the traditionally accepted monophyly of this subfamily. DNA was extracted from fresh and museum specimens representing all biogeographical regions. Maximum likelihood (ML) and Bayesian inference (BI) phylogenetic analyses were performed based on sequences from two mitochondrial genes, cytochrome c oxidase subunit I (COI) and cytochrome β, and three nuclear genes, carbomoylphosphate synthase domain of rudimentary, elongation factor-1α and isocitrate dehydrogenase. Through molecular data and morphological examination, our results reveal a division within Clinocerinae, distinguishing ‘typical’ or Clinocerinae (s.s.) from several genera, specifically Afroclinocera Sinclair, Asymphyloptera Collin and Proagomyia Collin, possibly lending support for a reclassification of these genera outside Clinocerinae. Bergenstammia Mik is proposed as a junior synonym of Phaeobalia Mik, syn. n., and the following new combinations are recognized: Phaeobalia albanica (Wagner) comb. n., Phaeobalia aurinae (Pusch & Wagner) comb. n., Phaeobalia carniolica (Horvat) comb. n., Phaeobalia frigida (Vaillant) comb. n., Phaeobalia glacialis (Palaczyk & Słowińska) comb. n., Phaeobalia multiseta (Strobl) comb. n., Phaeobalia nudimana (Vaillant) comb. n., Phaeobalia nudipes (Loew) comb. n., Phaeobalia pulla (Vaillant & Wagner) comb. n., Phaeobalia pyrenaica (Vaillant & Vinçon) comb. n., Phaeobalia slovaca (Wagner) comb. n. and Phaeobalia thomasi (Vaillant & Vinçon) comb. n. Re-evaluation of the genus Roederiodes resulted in the following new combinations: Clinocerella macedonicus (Wagner & Horvat) comb. n. and Clinocerella montenegrinus (Wagner & Horvat) comb. n. The origins of Clinocerinae (s.s.) are traced back to the Holarctic region, Laurasian origin, with a likely complex history of dispersal events into the Southern Hemisphere. Based on current knowledge, the greatest generic and species richness is confined to the Palaearctic Region. These findings provide valuable insights into the evolutionary relationships and distribution patterns of Clinocerinae (s.s.), challenging existing taxonomic classifications and shedding light on their historical biogeography.