Recent discoveries have shown that symbiotic relationships of viruses with their hosts are pervasive, confer major benefits to their hosts, and have played a major role in the evolution of life on earth. Understanding how these viral associations evolve is thus essential for a holistic view on the evolution of a broad diversity of organisms. Parasitoid wasps (wasps that grow within and kill other insects) are a natural laboratory for understanding the evolution of viral symbiosis because they are highly diverse and have evolved repeated associations with viruses. These viruses are able to suppress the immune defense mechanisms of the insect hosts that are parasitized by the wasps, enabling the wasps to survive and reproduce. Parasitoid wasps also provide substantial economic benefits to society because they are natural enemies of insect pests that cause damage to forests and agricultural crops. This project investigates the evolution and interactions between one of the most astonishing radiations on Earth (braconid parasitoid wasps, with 19,000 species) and the massive untapped diversity of their symbiotic viruses. The long-term goal is to understand the rules that govern the associations of the viruses, the wasps, and the hosts of the wasps. This research project advances knowledge in several ways: (1) discovery of new biodiversity in two poorly studied areas of the tree of life--viruses and wasps; (2) resolution of the evolutionary history and age of wasp-viral symbioses; (3) discovery of predictive genomic traits for viral symbiosis to understand the rules that govern these interactions; (4) characterization of the function of viruses; and (5) generate publicly available genome data. The proposed work will substantially transform understanding of symbiotic viral evolution, which will provide general information about virus evolution. The researchers will integrate wasp and viral research with education and public outreach by working closely with teachers in local schools and developing new educational resources. Furthermore, the program will improve STEM education and educator development and increase public literacy and engagement in STEM.<br/><br/>This project integrates phylogenetic, functional, and genomics approaches to discover the rules that govern mutualistic associations between viruses and wasps and, in turn, the interactions between these parasitoid wasps and their hosts. The overall objectives are to resolve and date the evolutionary relationships of the wasp family Braconidae using new methods of DNA analysis, discover which wasp lineages are associated with mutualistic viruses and the timing of their domestication, determine how viruses affect the antagonistic interactions between the wasps and their hosts, and examine genome features related to viral mutualism. The researchers aim to: (1) Resolve and date deep and shallow nodes for one of the most diverse lineages (19,000 species) on Earth; (2) Discover and characterize the multiple origins of viral symbiosis in Braconidae; and (3) decipher the roles of viral symbionts in the biology of the parasitoid wasps and their hosts. The outreach plan has two main components: 1) Providing research internships in the lab for local K-12 teachers so that they may learn new research techniques that will improve their teaching; and 2) using immersive virtual reality and animation experiences to engage the public and enhance learning in undergraduate education. The researchers will also provide integrated research and training opportunities to underrepresented mentees (2 postdoctoral researchers, 7 undergraduate and 2 graduate students) across the disciplines of phylogenetics, functional and comparative genomics, and revisionary systematics.<br/><br/>This award reflects NSF's statutory mission and has been deemed worthy of support through evaluation using the Foundation's intellectual merit and broader impacts review criteria.