A PHYLOGENETIC ANALYSIS OF HIGHER-LEVEL GALL WASP RELATIONSHIPS (HYMENOPTERA, CYNIPIDAE)

We present the most comprehensive analysis of higher-level relationshi ps in gall wasps conducted thus far. The analysis was based on detaile d study of the skeletal morphology of adults, resulting in 164 phyloge netically informative characters, complemented with a few biological c haracters. Thirty-seven cynipid species from thirty-one genera, includ ing four genera of the apparently monophyletic Cynipini and almost all of the genera in the other tribes, were examined. The outgroup includ ed exemplar species from three successively more distant cynipoid fami lies: Figitidae (the sister group of the Cynipidae), Liopteridae and I baliidae. There was considerable homoplasy in the data, but many group ings in the shortest tree were nonetheless well supported, as indicate d by bootstrap proportions and decay indices. Partitioning of the data suggested that the high level of homoplasy is characteristic of the C ynipidae and not the result of the amount of available phylogeneticall y conservative characters being exhausted. The analysis supported the monophyly of the Cynipini (oak gall wasps) which, together with the Rh oditini (the rose gall wasps), Eschatocerini and Pediaspidini formed a larger monophyletic group of gall inducers restricted to woody repres entatives of the eudicot subclass Rosidae. The inquilines (Synergini) were indicated to be monophyletic, whereas the Aylacini, primarily her b gall inducers, appeared as a paraphyletic assemblage of basal cynipi d groups. The shortest tree suggests that the Cynipidae can be divided into three major lineages: one including the inquilines, the Aylacini genera associated with Rosaceae, and Liposthenes; one consisting enti rely of Aylacini genera, among them Aulacidea, Isocolus and Neaylax; a nd one comprising the woody rosid gallers (the oak and rose gall wasps and allies), the Phanacis-Timaspis complex and the Aylacini genera as sociated with Papaveraceae.