Variable wing venation in<i>Agathiphaga</i>(Lepidoptera: Agathiphagidae) is key to understanding the evolution of basal moths

Details of the ancestral groundplan of wing venation in moths remain uncertain, despite approximately a century of study. Here, we describe a 3-branched subcostal vein, a 5-branched medial vein and a 2-branched cubitus posterior vein on the forewing of <i>Agathiphaga vitiensis</i> Dumbleton 1952 from Vanuatu. Such veins had not previously been described in any Lepidoptera. Because wing veins are typically lost during lepidopteran evolutionary history, rarely-if ever-to be regained, the venation of <i>A. vitiensis</i> probably represents the ancestral character state for moths. Wing venation is often used to identify fossil insects as moths, because wing scales are not always preserved; the presence of a supposedly trichopteran 3-branched subcostal vein in crown Lepidoptera may decrease the certainty with which certain amphiesmenopteran fossils from the Mesozoic can be classified. And because plesiomorphic veins can influence the development of lepidopteran wing patterns even if not expressed in the adult wing, the veins described here may determine the location of wing pattern elements in many lepidopteran taxa.