22Hybridization is differentially manifested across the genome, with observed introgression 23 representing a balance between selection and migration. The capacity to quantify introgression 24 and subsequently pinpoint the constituent genetic elements governing cross-species exchange has 25 been promoted by the unprecedented resolution of contemporary sequencing technologies.
26Furthermore, the availability of annotated reference genomes has allowed genomic patterns to be 27 associated with ecologically relevant phenotypes. We followed this pattern herein by harnessing 28 genomic resources to decipher the role of selection in shaping hybrid zones at the interface of 29 species-boundaries in North American box turtles (Terrapene). By so doing, we identified 30 adaptive divergence in genes related to immune system function and intrinsic thermal 31 adaptations. These, in turn, impact temperature-dependent sex determination and hypoxia 32 tolerance. Their patterns were then contrasted among inter-and intra-specific hybrid zones that 33 differed in a temporal and biogeographic context. Our results demonstrate that hybridization is 34 broadly apparent in Terrapene, but with varying levels of divergence at loci that impinge upon 35 thermal adaptation. These loci displayed signatures of adaptive introgression across intraspecific 36 boundaries, and do so despite a genome-wide selective trend against intergrades. By contrast, 37 interspecific comparisons at the same loci retained evidence of divergence. Importantly, 38 adaptations that shape species-boundaries in Terrapene not only underscore climatic boundaries 39 for these terrestrial ectotherms, but also bookmark their vulnerability to anthropogenic pressures. 40