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The geographic mosaic of coevolution predicts parasite virulence should be locally adapted to the host community.  Cotesia  parasitoid wasps adapt to local lepidopteran species possibly through their symbiotic bracovirus. The virus, essential for the parasitism success, is at the heart of the complex coevolutionary relationship linking the wasps and their hosts. The large segmented genome contained in the virus particles encodes virulence genes involved in host immune and developmental suppression. Coevolutionary arms race should result in the positive selection of particular beneficial alleles. To understand the global role of bracoviruses in the local adaptation or specialization of parasitoid wasps to their hosts, we studied the molecular evolution of four bracoviruses associated with wasps of the genus  Cotesia,  including  C congregata, C vestalis  and new data and annotation on two ecologically differentiated populations of  C sesamie ,  Kitale  and  Mombasa.  Paired orthologs analyses revealed more genes under positive selection when comparing the two  C sesamiae  bracoviruses belonging to the same species, and more genes under strong evolutionary constraint between species. Furthermore branch-site evolutionary models showed that 17 genes, out of the 54 currently available shared by the four bracoviruses, harboured sites under positive selection including: the  histone H4-like , a  C-type lectin , two  ep1-like ,  ep2 , a  viral ankyrin ,  CrV1 , a  ben-domain , a  Serine-rich , and eight unknown genes. Lastly the phylogenetic analyses of the  histone, ep2  and  CrV1  genes in different African  C sesamiae  populations showed that each gene described differently the individual relationships. In particular we found recombination had happened between the  ep2  and  CrV1  genes, which are localized 37.5 kb apart on the wasp chromosomes. Involved in multidirectional coevolutionary interactions,  C sesamiae  wasps rely on different bracovirus mediated molecular pathways to overcome local host resistance.

Adaptive Selection on Bracovirus Genomes Drives the Specialization of Cotesia Parasitoid Wasps