The frenulum is a wing coupling structure that is found on the wings of most families of Lepidoptera. It is a single bristle or set of bristles that originate from the base of the hindwing that often interlocks with the forewing during flight. This wing coupling mechanism is thought to have been a major evolutionary innovation that allowed for enhanced flight in Lepidoptera. The sack-bearer moths (Mimallonidae) are unusual among Lepidoptera in that not all species within the family have a frenulum. We test the hypothesis that the frenulum is not necessary and is therefore lost in mimallonids that have longer male forewings because such wings are perhaps better suited to be coupled by other means. To understand the evolution of the frenulum, we inferred the most taxonomically and genetically sampled anchored hybrid enrichment-based phylogeny of Mimallonidae, including 604 loci from all 41 genera and from 120 species, covering about 40% of the described species in the family. The maximum likelihood tree robustly supports major relationships within the family, and ancestral state reconstruction clearly recovers the frenulum as the plesiomorphic condition in Mimallonidae. Our results show that the frenulum is more often observed in species that have shorter, rather than longer, male forewings. The frenulum has historically been used as an important character for intrafamilial classification in Mimallonidae, but our results conclusively show that this character system is more variable than previously thought. Based on our results, we erect two new subfamilies, Roelofinae St Laurent & Kawahara, subfam. n. and Meneviinae St Laurent, Herbin, & Kawahara, subfam. n., for four genera previously considered incertae sedis. In the predominantly frenulum-lacking clade Cicinninae, we describe a new genus, Cerradocinnus St Laurent, Mielke, & Kawahara, gen. n., and the genus Gonogramma stat. rev. is revalidated to include many species previously placed in Cicinnus sensu lato. With these changes, Cicinnus can now be considered monophyletic. Thirty-three species are transferred to Gonogramma from Cicinnus sensu lato.

Anchored Hybrid Enrichment (AHE) using the LEP1 Lepidoptera-specific probe set, raw sequences processed with the Lepidoptera-specific pipeline as published by Breinholt et al. (2018).

The ReadMe file will further explain the contents of this submission, but in summary:

We are providing:

-raw Illumina reads for newly sequenced Mimallonidae species

-individual probe region nucleotide alignments for the 604 loci used in this study, after running the raw reads through the pipeline of Breinholt et al. (2018)

-a concatenated supermatrix of the 604 loci

-a full AMAS summary file

-A tree file (NEWICK) for the best of 100 individual runs for the "part1" partitioning scheme inferred in IQ-TREE, the best of 100 part2 tree file

-A tree file (NEWICK) for the best of 100 individual runs for the "part2" partitioning scheme inferred in IQ-TREE, the best of 100 part2 tree file

-A tree file (NEWICK) for the part2 partitioned dataset, but with problematic taxa removed

-R stats results and relevant code

-A csv file for input into the various ancestral state reconstructions and statistical tests performed in R