Phylogeny of the tribe Empoascini (Hemiptera: Cicadellidae: Typhlocybinae) based on morphological characteristics, with reclassification of the Empoasca generic group

A morphology-based phylogenetic analysis of the tribe Empoascini (Hemiptera: Cicadellidae: Typhlocybinae) is presented for 58 of 83 formerly recognized genera based on 99 morphological characters of adults. The results support excluding the New World Beamerana generic group from Empoascini. The remaining genera of Empoascini were recovered as a monophyletic sister group of Dikraneurini. Previously recognized tribes Jorumini and Helionini are derived from within Empoascini and are considered synonyms of the latter tribe. Three previously recognized informal generic groups, the Empoasca group, Alebroides group and Usharia group were paraphyletic but the Ficiana group was recovered as monophyletic based on five synapomorphies. Genera previously placed in the Alebroides group represent at least six independent lineages, indicating that the hind wing character separating this group from the Empoasca group (CuA and MP veins free) is highly homoplasious. Empoasca (sensu lato) is also paraphyletic. Thus, twelve previously recognized subgenera of Empoasca are elevated to genus status and five species groups of Empoasca from the New World are recognized as separate new genera. Sikkimasca Dworakowska, 1993 is treated as synonym of Marolda Dworakowska, 1977 based on the phylogeny. Biogeographic analysis suggests that Empoascini most likely first evolved in the Oriental region and spread to other biogeographic realms more recently by multiple independent invasions.