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.

Checklist of the Hymenoptera of the British Isles

“…) luteiceps (Kieffer, 1902, Allotria victrix var. ) luteicornis (Kieffer, 1902, Allotria) areolata (Kieffer, 1909, Charips) grandicornis (Kieffer, 1904, Allotria) io (Girault, 1932 piciceps (Thomson, 1862, Allotria) collina (Cameron, 1889, Allotria) ambrosiae (Ashmead, 1898, Allotria) carpentieri (Kieffer, 1902, Allotria) foveigera (Kieffer, 1902, Allotria) curvata (Kieffer, 1902, Allotria) recticornis (Kieffer, 1902, Allotria) subaptera (Kieffer, 1904, Alloxysta) campyla (Kieffer, 1904, Alloxysta) necans (Kieffer, 1909, Glyptoxysta) numidica (Kieffer, 1909, Bothrioxysta) bifoveata (Girault, 1931, Glyptoxysta) islandica (Hellén, 1931, Alloxysta) flavipes (Ionescu, 1963 Notes: Eucoilinae has been a subfamily of Figitidae rather than a family of its own since Ronquist (1999). Tribal classification follows Forshage and Nordlander (2008).…”

Revision of Canadian Eurytomidae (Hymenoptera, Chalcidoidea) associated with galls induced by cynipid wasps of the genus Diplolepis Geoffroy (Hymenoptera, Cynipidae) and description of a new species

“…One particularly species-rich, ecologically-closed model system for studies of host-parasitoid relationships is the community of gall wasps (Hymenoptera: Cynipidae) and their associated inquilines (e.g. Periclistus spp., Cynipidae: Diastrophini) and parasitoids (primarily Hymenoptera: Ichneumonoidea, Chalcidoidea) on oaks and roses (Csóka et al 2005, Nicholls et al 2010, Ronquist et al 2015. Inquilines have lost their ability to induce galls, but still retain the ability to modify gall tissue (Brooks and Shorthouse 1997, Ronquist et al 2015, Pujade-Villar et al 2016.…”

“…Periclistus spp., Cynipidae: Diastrophini) and parasitoids (primarily Hymenoptera: Ichneumonoidea, Chalcidoidea) on oaks and roses (Csóka et al 2005, Nicholls et al 2010, Ronquist et al 2015. Inquilines have lost their ability to induce galls, but still retain the ability to modify gall tissue (Brooks and Shorthouse 1997, Ronquist et al 2015, Pujade-Villar et al 2016. While entirely phytophagous, inquilines usually result in the death of inducers either through adult oviposition, or through competition for the same resources in the gall (Shorthouse 1973, 2010, Pujade-Villar et al 2016.…”

Primera cita del género <em>Neuroterus Hartig</em> (Hymenoptera, Cynipidae, Cynipini) para América Central, con descripción de tres especies nuevas de Panamá y Costa Rica

“…They either induce the galls themselves or live inside galls caused by other insects, most frequently other cynipids but also Chalcidoidea and Lepidoptera (Nieves-Aldrey, 2001;Van Noort et al, 2007;NievesAldrey & San Blas, 2015;Ronquist et al, 2015). According to Ronquist et al (2015), the family is currently divided into 12 tribes: Aylacini, Aulacideini, Ceroptresini, Cynipini, Diastrophini, Diplolepidini, Eschatocerini, Paraulacini, Pediaspidini, Phanacidini, Qwaqwaini and Synergini. Of these, Cynipini is the most species-rich and diverse tribe, with approximately 1,000 species of so-called "oak gall wasps," cynipids associated with oaks (Quercus species) and other plants of the Fagaceae family (Csóka et al, 2005;Stone et al, 2009).…”