Divergent selection on flowering phenology but not on floral morphology between two closely related orchids

Closely related species often differ in traits that influence reproductive success, suggesting that divergent selection on such traits contribute to the maintenance of species boundaries.
Gymnadenia conopsea ss. and Gymnadenia densiflora are two closely related,  perennial orchid species that differ in (1) floral traits important for pollination, including flowering phenology, floral display and spur length, and (2) dominant pollinators. If plant-pollinator interactions contribute to the maintenance of trait differences between these two taxa, we expect current divergent selection on flowering phenology and floral morphology between the two species.
We quantified phenotypic selection via female fitness in one year on flowering start, three floral display traits (plant height, number of flowers and corolla size) and spur length, in six populations of G. conopsea s.s. and in four populations of G. densiflora. There was indication of divergent selection on flowering start in the exp...