MORPHOLOGICAL VARIATION, TAXONOMIC DISTRIBUTION, AND PHYLOGENETIC SIGNIFICANCE OF CORNUTI IN TORTRICINAE (LEPIDOPTERA: TORTRICIDAE)

Based on the examination of 4,218 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 versus 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, between other sister groups, features of the cornuti (presence/absence, attachment, shape, size, etc.) provide little or no evidence of putative relationships (e.g., Ceracini and Archipini). Our studies suggest that if deciduous cornuti are homologous throughout Tortricidae, 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) possess typical deciduous cornuti as do most Archipini, Sparganothini, and Atteriini, and some Epitymbiini (Tortricinae), as well as some Eucosmini and Grapholitini (Olethreutinae).