title: Phylogeny, evolution and classification of gall wasps: the plot thickens.
abstract: Gall wasps (Cynipidae) represent the most spectacular radiation of gall-inducing insects. In addition to true gall formers, gall wasps also include phytophagous inquilines, which live inside the galls induced by gall wasps or other insects. Here we present the first comprehensive molecular and total-evidence analyses of higher-level gall wasp relationships. We studied more than 100 taxa representing a rich selection of outgroups and the majority of described cynipid genera outside the diverse oak gall wasps (Cynipini), which were more sparsely sampled. About 5 kb of nucleotide data from one mitochondrial (COI) and four nuclear (28S, LWRh, EF1alpha F1, and EF1alpha F2) markers were analyzed separately and in combination with morphological and life-history data. According to previous morphology-based studies, gall wasps evolved in the Northern Hemisphere and were initially herb gallers. Inquilines originated once from gall inducers that lost the ability to initiate galls. Our results, albeit not conclusive, suggest a different scenario. The first gall wasps were more likely associated with woody host plants, and there must have been multiple origins of gall inducers, inquilines or both. One possibility is that gall inducers arose independently from inquilines in several lineages. Except for these surprising results, our analyses are largely consistent with previous studies. They confirm that gall wasps are conservative in their host-plant preferences, and that herb-galling lineages have radiated repeatedly onto the same set of unrelated host plants. We propose a revised classification of the family into twelve tribes, which are strongly supported as monophyletic across independent datasets. Four are new: Aulacideini, Phanacidini, Diastrophini and Ceroptresini. We present a key to the tribes and discuss their morphological and biological diversity. Until the relationships among the tribes are resolved, the origin and early evolution of gall wasps will remain elusive.
authors.affiliation: Department of Bioinformatics and Genetics, Swedish Museum of Natural History, Stockholm, Sweden.
authors.forename: Fredrik
authors.initials: F
authors.lastname: Ronquist
authors.affiliation: Departamento de Biodiversidad y Biología Evolutiva, Museo Nacional de Ciencias Naturales (CSIC), Madrid, Spain.
authors.forename: José-Luis
authors.initials: JL
authors.lastname: Nieves-Aldrey
authors.affiliation: Systematic Entomology Lab, USDA, c/o Smithsonian Institution, Washington DC, United States of America.
authors.forename: Matthew L
authors.initials: ML
authors.lastname: Buffington
authors.affiliation: Department of Biological Sciences, Eastern Illinois University, Charleston, IL, United States of America.
authors.forename: Zhiwei
authors.initials: Z
authors.lastname: Liu
authors.affiliation: Swedish Species Information Centre, Swedish University of Agricultural Sciences, Uppsala, Sweden.
authors.forename: Johan
authors.initials: J
authors.lastname: Liljeblad
authors.affiliation: Department of Bioinformatics and Genetics, Swedish Museum of Natural History, Stockholm, Sweden; Bioinformatics Infrastructure for Life Sciences, Linköping University, Linköping, Sweden.
authors.forename: Johan A A
authors.initials: JA
authors.lastname: Nylander
mesh_terms: D000818:Animals; D019659:Asteraceae / Q000254:growth & development / Q000469:parasitology*; D005075:Biological Evolution; D006790:Host-Parasite Interactions; D008018:Life Cycle Stages; D010802:Phylogeny; D014863:Wasps / Q000145:classification* / Q000502:physiology*