Nuclear Ribosomal DNA Restriction-Site Variation in Penstemon Section Peltanthera (Scrophulariaceae): An Evaluation of Diploid Hybrid Speciation and Evidence for Introgression

Abstract

Penstemon spectabilis is a putative stabilized diploid hybrid of P. centranthifolius and P. grinnellii. It is morphologically intermediate, and all three species have different pollinators. Penstemon centranthifolius and P. spectabilis have been proposed as parents of P. clevelandii, which is purportedly isolated by ecological factors. Although hybridization between the proposed parental species has been reported, hybrid swarm formation has not been observed and introgression is purported to be minimal. We tested hypotheses of diploid hybrid speciation and introgression among these species based on rDNA restrictionsite and length variation for 56 populations within and outside of the hybrid complex. Unambiguous molecular markers clearly differentiated P. centranthifolius, P. grinnellii, and P. spectabilis, whereas P. spectabilis and P. clevelandii had the same rDNA type. The P. centranthifolius rDNA type was found in some populations of P. spectabilis and P. clevelandii, but there was no evidence of the additive profile documented for other recent diploid hybrid species. In contrast, the rDNA profile of P. × parishii had a completely additive profile of its proposed parental species P. centranthifolius and P. spectabilis. Ribosomal DNA markers for P. grinnellii were restricted to populations within the species and were not found in any population of P. spectabilis. Our data did not support hybrid-speciation hypotheses, but were in accord with allozyme data that provided evidence for introgression between P. centranthifolius and P. grinnellii, P. spectabilis and P. clevelandii. These results were used to propose criteria to differentiate ancient diploid hybrid speciation from patterns of introgression.