Early evolution of wing scales prior to the rise of moths and butterflies.

Although scales are a defining and conspicuous feature of moths and butterflies (Lepidoptera), their earliest evolution predates the group but is shrouded by a dearth of fossil evidence. Herein, we report two new species in mid-Cretaceous Kachin amber, representing lineages closely related to Lepidoptera: one represents the extinct Tarachoptera, with dense scales on the fore- and hindwings, while the other is an early lineage of caddisflies, with a hindwing covered by a single layer of angustifoliate scales. A novel phylogenetic analysis of 174 morphological characters and 73 extant and fossil representatives of Mecopterida demonstrates a monophyletic origin of scales in the common ancestor of Tarachoptera, Trichoptera, and Lepidoptera; that Tarachoptera are monophyletic but their scale morphology is plesiomorphic for the whole group; and that scales were lost early in caddisfly evolution before reappearing multiple times within the clade. Collectively, these fossils provide clarity into the origin and early evolution of scales before their diversification among the moths and butterflies.