A multi-gene phylogenetic analysis of the leafhopper subfamily Typhlocybinae (Hemiptera: Cicadellidae) challenges the traditional view of the evolution of wing venation

Abstract

Genera of the diverse leafhopper subfamily Typhlocybinae have traditionally been classified into tribes primarily based on characters of the wing venation and an intuitive phylogeny of this group was previously proposed based on the hypothetical pattern of wing vein evolution. Some recent authors suggested that wing vein characters are not always reliable but few attempts have been made to examine the status and relationships of typhlocybine tribes using rigorous phylogenetic analyses. Phylogenetic analysis of a dataset comprising DNA sequences from five gene regions (H3, H2A, 28S rDNA D2, 16S rDNA, and COI with a total length of 2413 bps) and 61 species representing six previously recognized tribes of Typhlocybinae provides strong support for the monophyly of the subfamily and five of the previously recognized tribes. Most branches received moderate to strong maximum likelihood bootstrap support. The following intertribal relationships were recovered: (Alebrini + Empoascini) + ((Dikraneurini + Erythroneurini) + Typhlocybini). Maximum likelihood analysis recovered Zyginellini (treated as a separate tribe by some authors) as sister to Typhlocybini with low branch support, but the former tribe was derived from within the latter in Bayesian analysis of the same dataset and relationships within Typhlocybini (sensu lato) were generally poorly resolved in both analyses. The relationship of Typhlocybini to other tribes is also unstable, suggesting that more data are needed to resolve the position of this tribe with confidence. Parts of the phylogeny receiving strong support in both analyses contradict the traditional view that Alebrini, the only tribe retaining an appendix in the forewing, is the earliest diverging lineage and possibly gave rise to the other tribes. Ancestral state reconstructions indicate that characters of the wing venation traditionally used for diagnosing typhlocybine tribes are generally conservative but exhibit some homoplasy and may not, by themselves, be reliable for recognizing monophyletic groups within this subfamily.