Hemipteroid insects (Paraneoptera), with over 10% of all known insect diversity, are a major component of terrestrial and aquatic ecosystems. Previous phylogenetic analyses have not consistently resolved the relationships among major hemipteroid lineages. We provide maximum likelihood-based phylogenomic analyses of a taxonomically comprehensive dataset comprising sequences of 2,395 single-copy, protein-coding genes for 193 samples of hemipteroid insects and outgroups. These analyses yield a well-supported phylogeny for hemipteroid insects. Monophyly of each of the three hemipteroid orders (Psocodea, Thysanoptera, and Hemiptera) is strongly supported, as are most relationships among suborders and families. Thysanoptera (thrips) is strongly supported as sister to Hemiptera. However, as in a recent large-scale analysis sampling all insect orders, trees from our data matrices support Psocodea (bark lice and parasitic lice) as the sister group to the holometabolous insects (those with complete metamorphosis). In contrast, four-cluster likelihood mapping of these data does not support this result. A molecular dating analysis using 23 fossil calibration points suggests hemipteroid insects began diversifying before the Carboniferous, over 365 million years ago. We also explore implications for understanding the timing of diversification, the evolution of morphological traits, and the evolution of mitochondrial genome organization. These results provide a phylogenetic framework for future studies of the group.

A data set comprising DNA sequences from 388 loci and >99,000 aligned nucleotide positions, generated using anchored hybrid enrichment, was used to estimate relationships among 138 leafhoppers and treehoppers representative of all major lineages of Membracoidea, the most diverse superfamily of hemipteran insects. Phylogenetic analysis of the concatenated nucleotide sequence data set using maximum likelihood produced a tree with most branches receiving high support. A separate coalescent gene tree analysis of the same data generally recovered the same strongly supported clades but was less well resolved overall. Several nodes pertaining to relationships among leafhopper subfamilies currently recognized based on morphological criteria were separated by short internodes and received low support. Although various higher taxa were corroborated with improved branch support, relationships among some major lineages of Membracoidea are only somewhat more resolved than previously published phylogenies based on single gene regions or morphology. In agreement with previous studies, the present results indicate that leafhoppers (Cicadellidae) are paraphyletic with respect to the three recognized families of treehoppers (Aetalionidae, Melizoderidae, and Membracidae). Divergence time estimates indicate that most of the poorly resolved divergence events among major leafhopper lineages occurred during the lower to middle Cretaceous and that most modern leafhopper subfamilies, as well as the lineage comprising the three recognized families of treehoppers, also arose during the Cretaceous.

Abstract. The classification of the largest subfamily of leafhoppers, Deltocephalinae, including 38 tribes, 923 genera, and 6683 valid species, is reviewed and revised. An updated phylogeny of the subfamily based on molecular (28S, Histone H3) and morphological data and an expanded taxon sample (37 taxa not included in previous analyses) is presented. Based on the results of these analyses and on the morphological examination of many representatives of the subfamily, the classification of the tribes and subtribes of Deltocephalinae is revised. Complete morphological descriptions, illustrations, lists of the included genera, and notes on their distribution, ecology, and important vector species are provided for the 38 recognized tribes and 18 subtribes. A dichotomous key to the tribes is provided. All names in the taxonomic treatments are hyperlinked to online resources for individual taxa which are supported by a comprehensive database for Deltocephalinae compiled using the taxonomic database software package 3I. The online functionality includes an interactive key to tribes and subtribes and advanced database searching options. Each taxon (subspecies through subfamily) has a unique taxon webpage providing nomenclatural information, lists of included taxa, an automated description (if available), images (if available), distributional information, bibliographic references and links to outside resources.

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