Analyses of expanded anchored-hybrid phylogenomic datasets support revisions to the higher classification of leafhoppers (Hemiptera: Cicadellidae)

Previous phylogenomic analyses of Membracoidea have strongly supported the monophyly of most currently recognized subfamilies of leafhoppers (Cicadellidae) but suggested that some revisions to the higher classification are needed. However, the most taxon-rich previous analysis included very few representatives of some family-group taxa. Here, a dataset comprising sequences from 269 taxa, ca. twice as many loci and taxa as the largest previous phylogenomic dataset for Membracoidea and representing all extant families, subfamilies and over 96% of the extant tribes, was analysed using concatenated and coalescent methods to test previous results and provide a stronger basis for revisions to the higher classification. The following taxonomic changes are proposed based on the consistently strong branch support for several lineages in both concatenated maximum likelihood and coalescent gene tree analyses. Cicadellinae is expanded to include tribes Makilingiini and Tungurahualini, both previously included in Mileewinae. Mileewinae is therefore restricted to include the nominotypical tribe and Tinteromini. Errhomeninae, most recently treated as a tribe of Aphrodinae, is elevated to subfamily status and includes tribes Bathysmatophorini, Euacanthellini and Malmaemichungiini. Portanini is transferred from Aphrodinae to Neocoelidiinae. The concept of Aphrodinae is consequently narrowed to include only the nominotypical tribe, Sagmatiini and Xestocephalini. Neobalinae is expanded to include Equeefini, a tribe previously included in Coelidiinae. The morphologically aberrant soil-dwelling genus Paulianiana from Madagascar, previously placed in Sagmatiini (Aphrodinae), is transferred to Ledrinae. The Australian endemic tribes Thymbrini and Stenocotini, most recently placed in Tartessinae, are transferred to Ledrinae. Tartessinae is restricted to include the nominotypical tribe, Neopsini and Tardrabassus (new placement, previously unplaced to a tribe within Iassinae). Although most leafhopper subfamilies, as currently defined, are consistently recovered as monophyletic with strong support, these major lineages are connected to each other by very short, deep internal branches and their relationships remain unstable. Aphrodinae, Eurymelinae, Iassinae, Mileewinae, Ledrinae and Tartessinae, as currently defined, were not consistently recovered as monophyletic, but support for their non-monophyly was also low, so their current definitions are retained, pending further analyses. Keys to the families of Membracoidea and to the 19 currently recognized leafhopper subfamilies are provided. Divergence time estimates suggest that Cicadellidae originated during the Middle Triassic (~224 mya) and began to diverge at ~217 mya, with most modern leafhopper subfamilies arising during the Jurassic and Early Cretaceous periods.