Evolution of the planthoppers (Insecta: Hemiptera: Fulgoroidea).

The planthopper superfamily Fulgoroidea (Insecta: Hemiptera) comprises approximately 20 described insect families, depending on which classification is followed. Multiple competing hypotheses of fulgoroid phylogeny have been published, based on either morphological character coding or DNA sequence data; however, those hypotheses disagree in several key aspects regarding the evolution of planthoppers. The current paper seeks to test these hypotheses, including the Asche (Asche, M. 1987. Preliminary thoughts on the phylogeny of Fulgoromorpha (Homoptera Auchenorrhyncha). In: Proceedings of the 6th Auchenorrhyncha Meeting, Turin, Italy, 7-11 September, 1987, pp. 47-53.) hypothesis of a trend in ovipositor structure, which may be correlated with planthopper feeding ecology. Presented here are phylogenetic reconstructions of Fulgoroidea based on analysis of DNA nucleotide sequence data from four loci (18S rDNA, 28S rDNA, Histone 3, and Wingless) sequenced from 83 exemplar taxa representing 18 planthopper families and outgroups. Data sets were analyzed separately and in various combinations under the maximum parsimony criterion, and the total combined dataset was analyzed via both maximum parsimony and partitioned Bayesian criteria; results of the combined analyses were concordant across reconstruction paradigms. Relationships recovered suggest several major planthopper lineages, including: (1) Delphacidae+Cixiidae; (2) Kinnaridae+Meenoplidae; (3) Fulgoridae+Dictyopharidae; (4) Lophopidae+Eurybrachidae (possibly+Flatidae); (5) Ricaniidae+Caliscelidae (possibly+Tropiduchidae). Results also suggest the placement of Achilixiidae outside of Cixiidae and of Tettigometridae as one of the more recently diversified lineages within Fulgoroidea. The resulting phylogeny supports Asche's (1987) hypothesis of a functional trend in ovipositor structure across families.