Phenotypic evidence for an extensive mosaic hybrid zone between two species of columbine, Aquilegia flavescens and A. formosa

Describing the structure and dynamics of hybrid zones has important consequences for our understanding of lineage diversification. Herbarium collections constitute an extensive, though often underutilized, morphological resource that can potentially be leveraged to identify and describe hybrid zones. We demonstrate the use of herbarium collections to describe the structure of a hybrid zone between two North American columbines, Aquilegia flavescens S. Watson and A. formosa Fisch. ex DC. Hybrid indices were calculated from floral morphology measurements and mapped using geolocation data, revealing an extensive and consistent pattern of floral intermediacy in several contact zones. Population phenotype samples from contact zones corroborate these findings and strongly suggest introgression. At one locality with a confirmed hybrid population, floral phenotypes show clinal variation in parental-type morphology along an elevational gradient, in the direction predicted by the divergent elevational distributions of these taxa. We argue that these patterns together reflect a habitat-associated mosaic hybrid zone, consistent with divergent ecological selection maintaining the structure of the hybrid zone.