Phylogeny and historical biogeography of the leafhopper subfamily Coelidiinae (Hemiptera: Cicadellidae) based on morphological and molecular data

The phylogenetic relationships among major lineages of the globally distributed leafhopper subfamily Coelidiinae were reconstructed by analysis of 2903 nucleotide positions from two mitochondrial genes (COI and 16S), four nuclear genes (28S, H3, H2A and Wingless) and 102 discrete morphological characters, compiled for 86 species representing 52 genera within 9 coelidiine tribes broadly representative of the world fauna in addition to 12 outgroup taxa. Maximum likelihood and Bayesian analyses yielded well-resolved phylogenetic estimates that were highly congruent with most branches receiving strong support. The results indicate Coelidiinae sensu stricto (=Coelidiinae sensu lato without Equeefini and Macroceratogoniini), Thagriini, Tharrini, Tinobregmini and Equeefini are monophyletic. However, the two largest tribes, Coelidiini and Teruliini, are paraphyletic, as well as Youngolidiini and most genera of Oriental Coelidiini. Fossil-calibrated molecular divergence time analysis indicates that the Coelidiinae sensu stricto originated ca. 149 Ma, prior to the complete separation of the Gondwanan continents, and the tribe-level lineages diverged between the Lower and Upper Cretaceous (92.77–138.03 Ma). The major lineages of Coelidiinae arose in the Oriental and Neotropical regions, and Oriental Coelidiini arose as a result of dispersal and colonisation from the Afrotropical region.