Reproductive isolation and introgression between sympatric Mimulus species.

Incompletely isolated species provide an opportunity to investigate the genetic mechanisms and evolutionary forces that maintain distinct species in the face of ongoing gene flow. Here, we use field surveys and reduced representation sequencing to characterize the patterns of reproductive isolation, admixture and genomic divergence between populations of the outcrossing wildflower Mimulus guttatus and selfing M. nasutus. Focusing on a single site where these two species have come into secondary contact, we find that phenological isolation is strong, although incomplete, and is likely driven by divergence in response to photoperiod. In contrast to previous field studies, which have suggested that F1 -hybrid formation might be rare, we discover patterns of genomic variation consistent with ongoing introgression. Strikingly, admixed individuals vary continuously from highly admixed to nearly pure M. guttatus, demonstrating ongoing hybridization and asymmetric introgression from M. nasutus into M. guttatus. Patterns of admixture and divergence across the genome show that levels of introgression are more variable than expected by chance. Some genomic regions show a reduced introgression, including one region that overlaps a critical photoperiod QTL, whereas other regions show elevated levels of interspecific gene flow. In addition, we observe a genome-wide negative relationship between absolute divergence and the local recombination rate, potentially indicating natural selection against M. nasutus ancestry in M. guttatus genetic backgrounds. Together, our results suggest that Mimulus speciation is both ongoing and dynamic and that a combination of divergence in phenology and mating system, as well as selection against interspecific alleles, likely maintains these sympatric species.