Phylogeny and biogeography of Neotropical spittlebugs (Hemiptera: Cercopidae: Ischnorhininae): revised tribal classification based on morphological data

The spittlebug family Cercopidae is currently divided into two subfamilies: the paraphyletic Old World Cercopinae and the monophyletic New World Ischnorhininae. The most recent classification scheme proposed by Fennah in 1968 divided the New World Cercopidae into four tribes: Tomaspidini, Ischnorhinini, Hyboscartini and Neaenini. Herein we present a phylogenetic analysis of Ischnorhininae using 108 morphological characters and including 53 of the 59 recognized genera, to evaluate the tribal‐level classification and understand the processes underlying the current distributional patterns of these genera. We found significant support for the monophyly of many Neotropical genera, but Fennah's tribal classification is revised because tribes Neaenini, Ischnorhinini and Tomaspidini were recovered as polyphyletic. Hyboscartini was synonymized with Tomaspidini. A taxonomic key to tribes and genera of Neotropical spittlebugs is provided based mostly on recovered apomorphies. The biogeographical analysis suggests a Neotropical origin of ischnorhinines, more specifically in northwestern South America. This was possibly coincident spatially and temporally with the origin of grasses, with ancestral range expansions southward to the Amazonian and Paraná regions, and posterior vicariant events, possibly related to the expansion of forests in the Chacoan region, the South America diagonal of open formations. Dispersals to the Chacoan region and to the Nearctic region are hypothesized to have occurred only within genera. In the Chacoan region it is associated with more recent events, such as the diversification of C4 grasses and establishment of the savannas and seasonally dry forests.