Phylogeny and diversification analyses of extant planthopper families (Hemiptera: Fulgoromorpha) based on a mitochondrial dataset reveal ancient lineages originating during the Jurassic and originally feeding on gymnosperms

Phylogenetic analyses of planthoppers were conducted using a mitogenome dataset to examine the evolutionary relationships of current families based on both maximum likelihood (ML) and Bayesian inference methods. Delphacoidea, including Cixiidae and Delphacidae, and Fulgoroidea, including all other families, were consistently identified as monophyletic groups. However, Nogodinidae displayed a polyphyletic pattern, with various lineages emerging sister to several issidoid families. Variations were observed across different analyses in the positioning of (Dictyopharidae + Fulgoridae) and the relationships of Ricaniidae sister either to Issidae or to (Flatidae + Acanaloniidae). Several statistical tests indicate that the ML topology has the highest level of statistical confidence.The (Dictyopharidae + Fulgoridae) clade is best positioned as sister to (De rbidae + (Achilidae + (Tropiduchidae + other “higher” planthoppers families))) and the Ricaniidae as sister to (Flatidae + Acanaloniidae).With 13 fossil calibration points, origin times for Delphacoidea and Fulgoroidea were dated back to the Late Permian, approximately 257.46 million years ago. Delphacidae and Cixiidae split during the LateTriassic, while Fulgoroidea diversified earlier during the MiddleTriassic. Divergence and diversification times are also provided for all other extant planthopper families.These results suggest that Delphacoidea and Fulgoroidea likely diverged from the older planthopper fossil families prior to the emergence of angiosperms and may have initially fed on gymnosperms and ancient ferns.The diversification of extant planthopper families occurred during the Jurassic and Cretaceous periods, influenced by evolving climatic conditions, the decline in gymnosperms, and the increasing diversity of angiosperm plants.