Intraspecific perspective of phenotypic coordination of functional traits in Scots pine.

Functional traits have emerged as a key to understand species responses to environmental conditions. The concerted expression of multiple traits gives rise to the phenotype of each individual, which is the one interacting with the environment and evolving. However, patterns of trait covariation and how they vary in response to environmental conditions remain poorly understood, particularly at the intraspecific scale. Here, we have measured traits at different scales and in different organs, and analysed their covariation in a large number of conspecifics distributed in two contrasting environments. We expected significant correlations among traits, not only within clusters of traits as found in global, multispecies studies, but also among clusters, with more relationships within clusters, due to genetic constraints, and among clusters due to more coordinated phenotypes than community level, multispecies studies. We surveyed 100 Pinus sylvestris trees in a Mediterranean mountainous area distributed in two contrasting elevations. We measured 13 functional traits, in three clusters (leaf, stem and whole-plant traits), and analysed their variation and coordination. We found significant coordination among traits belonging to different clusters that reveals coordinated phenotypes. However, we found fewer correlations within trait clusters than initially expected. Trait correlation structures (number, intensity and type of correlations among traits) differed among individuals at different elevations. We observed more correlations within trait clusters at low elevation compared to those at high elevation. Moreover, the higher number of correlations among different trait clusters and the lower trait variation at the higher elevation suggests that variability decreases under more stressful conditions. Altogether, our results reveal that traits at intraspecific scale are coordinated in a broad network and not only within clusters of traits but also that this trait covariation is significantly affected by environmental conditions.