Morphology-based Phylogenetic Analysis of Membracoidea (Hemiptera: Cicadomorpha) With Placement of Fossil Taxa and Description of a New Subfamily

Recently discovered amber-preserved fossil Cicadellidae exhibit combinations of morphological traits not observed in the modern fauna and have the potential to shed new light on the evolution of this highly diverse family. To place the fossils explicitly within a phylogenetic context, representatives of five extinct genera from Cretaceous Myanmar amber, and one from Eocene Baltic amber were incorporated into a matrix comprising 229 discrete morphological characters and representatives of all modern subfamilies. Phylogenetic analyses yielded well resolved and largely congruent estimates that support the monophyly of most previously recognized cicadellid subfamilies and indicate that the treehoppers are derived from a lineage of Cicadellidae. Instability in the morphology-based phylogenies is mainly confined to deep internal splits that received low branch support in one or more analyses and also were not consistently resolved by recent phylogenomic analyses. Placement of fossil taxa is mostly stable across analyses. Three new Cretaceous leafhopper genera, Burmotettix gen. nov., Kachinella gen nov., and Viraktamathus gen. nov., consistently form a monophyletic group distinct from extant leafhopper subfamilies and are placed in Burmotettiginae subfam. nov. Extinct Cretaceous fossils previously placed in Ledrinae and Signoretiinae are recovered as sister to modern representatives of these groups. Eomegophthalmus Dietrich and Gonçalves from Baltic amber consistently groups with a lineage comprising treehoppers, Megophthalminae, Ulopinae, and Eurymelinae but its position is unstable. Overall, the morphology-based phylogenetic estimates agree with recent phylogenies based on molecular data alone suggesting that morphological traits recently used to diagnose subfamilies are generally informative of phylogenetic relationships within this group.