Genetic drift versus natural selection affecting the evolution of spectral and functional traits of two key macrophytes: Phragmites australis and Nuphar lutea

Both genetic and phenotypic intraspecific diversity play a crucial role in the ecological and evolutionary dynamics of organisms. Several studies have compared phenotypic divergence (P-st) and differentiation of neutral loci (F-st) to infer the relative roles of genetic drift and natural selection in population differentiation (P-st-F-st comparison). For the first time, we have assessed and compared the genetic variation and differentiation at the leaf trait level in two key macrophytes, Phragmites australis and Nuphar lutea. To this aim, we quantified and described the genetic structure and phenotypic diversity of both species in five lake systems in north-central Italy. We then investigated the relative roles of genetic drift and natural selection on leaf trait differentiation (P-st-F-st), assuming that F-st reflects divergence caused only by genetic drift while P-st also incorporates the effects of selective dynamics on the phenotype. In terms of genetic structure, the results for P. australis were in line with those observed for other Italian and European conspecific populations. Conversely, N. lutea showed a more complex genetic structure than expected at the site level, probably due to the combined effect of genetic isolation and its mixed mating system. Both species exhibited high variability in leaf functional traits within and among sites, highlighting a high degree of phenotypic plasticity. P-st-F-st comparisons showed a general tendency towards directional selection in P. australis and a more complex pattern in N. lutea. Indeed, the drivers of phenotypic differentiation in N. lutea showed a variable mix of stabilising and directional selection or neutral divergence at most sites. The prevalence of vegetative over generative reproduction leads P. australis populations to be dominated by a few clones that are well adapted to local conditions, including phenotypes that respond plastically to the environment. In contrast, in N. lutea, the interaction of a mixed mating system and geographical isolation among distant sites tends to reduce the effect of outbreeding depression and provides the genetic basis for adaptive capacity. The first joint analysis of the genetic structure of these two key macrophytes allowed a better understanding of the relative roles of genetic drift and natural selection in the diversification of phenotypic traits within habitats dominated by P. australis and N. lutea.