One hundred and sixty years of taxonomic confusion resolved: Belonocnema (Hymenoptera: Cynipidae: Cynipini) gall wasps associated with live oaks in the USA

Gall wasps (Hymenoptera: Cynipidae) in the genus Belonocnema induce galls on live oaks (Quercus series Virentes), forming multilocular root galls in the sexual generation and unilocular leaf galls in the asexual generation. Using morphological characters, host records, museum specimens, flight propensity and phylogenetic analysis of published cytochrome c oxidase subunit I (COI) and nuclear SNP data, we resolve the long-standing taxonomic confusion within Belonocnema and recognize three distinct species that are distributed throughout the southern and south-eastern USA: B. fossoria (rev. stat.), B. kinseyi (rev. stat.) and B. treatae, while B. quercusvirens is treated as species inquirenda. The presence of mitonuclear discordance results in the failure of a mitochondrial DNA barcode region to distinguish between B. fossoria and B. treatae, while recognizing B. kinseyi, despite the three species being clearly separated based on morphology and phylogenetic analysis of SNP data. We provide re-descriptions and an updated dichotomous key for both asexual and sexual generations of these widespread species. Finally, as Belonocnema has emerged as a model organism for ecological and evolutionary studies, we clarify the species examined in published studies to date.