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Abstract

A phylogenetic analysis of Hormaphidinae is presented based on a total-evidence approach. Four genes (two mitochondrial, COI and CytB, and two nuclear, EF-1α and LWO) are combined with 65 morphological and seven biological characters. Sixty-three hormaphidine species representing three tribes and 36 genera as well as nine outgroups are included. Parsimony and model-based approaches are used, and several support values and implied weighting schemes are explored to assess clade stability. The monophyly of Hormaphidinae and Nipponaphidini is supported, but Cerataphidini and Hormaphidini are not recovered as monophyletic. Based on the parsimony hypothesis from the total-evidence analysis, the phylogenetic relationships within Hormaphidinae are discussed. Cerataphidini is re-delimited to exclude Doraphis and Tsugaphis, and Hormaphidini is redefined to include Doraphis. Ceratocallis Qiao & Zhang is established as a junior synonym of Ceratoglyphina van der Goot, syn. nov. Lithoaphis quercisucta Qiao, Guo & Zhang is transferred to the genus Neohormaphis Noordam as Neohormaphis quercisucta (Qiao, Guo & Zhang) comb. nov. Galls have evolved independently within three tribes of Hormaphidinae. In Cerataphidini, pseudogalls are ancestral, both single-cavity and multiple-cavity galls have evolved once, and galls appear to have evolved towards greater complexity. Galling on secondary hosts has evolved twice in hormaphidines.