Variation in cornuti in the leaf-roller moths (Lepidoptera: Tortricidae: Tortricinae)

Based on the examination of over 4,000 slide-mounted preparations of male and female genitalia of tortricine moths, representing all major clades of the subfamily worldwide, we propose a classification system for cornuti based on four criteria: (1) presence/absence; (2) deciduous/non-deciduous; (3) type of attachment; and (4) shape. In general, the taxonomic distribution of deciduous vs. non-deciduous cornuti is in conformance with a recent phylogenetic hypothesis of the family. Some sister groups (i.e., tribes) have remarkably similar cornuti (e.g., Atteriini and Sparganothini); however, in others, features of the cornuti (presence/absence, attachment, shape, size, etc.) provide little or no evidence of these relationships (e.g., Ceracini and Archipini). Our studies suggest that if deciduous cornuti are homologous throughout Tortrticidae, which seems likely, this feature arose near the base of the tree at the branch that supports the sister groups Olethreutinae + Tortricinae. The least derived Tortricinae (i.e., Phricanthini) posses typical deciduous cornuti as do most Archipini, Epitymbiini, Sparganothini, and Atteriini in Tortricinae, and many Eucosmini and Grapholitini in Olethretinae.