The legacy of climate variability over the last century on populations' phenotypic variation in tree height.

Phenotypic plasticity and local adaptation are the two main processes underlying trait variability. Under rapid environmental change, phenotypic plasticity, if adaptive, could increase the odds for organisms to persist. However, little is known on how environmental variation has shaped plasticity across species ranges over time. Here, we assess whether the portion of phenotypic variation of tree populations linked to the environment is related to the inter-annual climate variability of the last century and how it varies among populations across species ranges and age. To this aim, we used 372,647 individual tree height measurements of three pine species found in low elevation forests in Europe: Pinus nigra Arnold, P. pinaster Aiton and P. pinea L. Measurements were taken in a network of 38 common gardens established in Europe and North Africa with 315 populations covering the distribution range of the species. We fitted linear mixed-effect models of tree height as a function of age, population, climate and competition effects. Models allowed us to estimate tree height response curves at the population level and indexes of populations' phenotypic variation, as a proxy of phenotypic plasticity, at 4, 8 and 16 years old, and relate these indexes to the inter-annual climate variability of the last century. We found that phenotypic variation in tree height was higher in young trees than in older ones. We also found that P. pinea showed the highest phenotypic variation in tree height compared with P. pinaster and P. nigra. Finally, phenotypic variation in tree height may be partly adaptive, and differently across species, as climate variability during the last century at the origin of the populations explained between 51 and 69% of the current phenotypic variation of P. nigra and P. pinea, almost twice of the levels of P. pinaster. MAIN CONCLUSIONS: Populations' phenotypic variation in tree height is largely explained by the climate variability that the populations experienced during the last century, which we attribute to the genetic diversity among populations.

Population divergence for heteroblasty in the Canary Island pine (Pinus canariensis, Pinaceae).

A heteroblastic (or vegetative phase) change is an abrupt manifestation in the general heteroblastic development during the ontogeny of plants. The Canary Island pine undergoes an especially marked and delayed heteroblastic change, including both the formation of secondary needles on dwarf shoots and the onset of preformed growth. To assess genetic and environmental effects on the heteroblastic change in this species, we followed plants from 19 populations at a dry site and a wetter site. Comparing juvenile and adult needles from the same individuals, the adult had a significantly lower rate of water loss and higher leaf mass per area. Pooling data from all seed sources, the heteroblastic change took place when plants reached a critical height, on average, at 4 years of age at the dry site and 1 year earlier at the wet site. Within a subsample of individuals of equal size, mortality was significantly higher in juvenile plants than in mature plants. However, the juvenile phase was longer in plants from dry regions when compared to plants from highly productive, wet regions. This apparent contradiction might be explained through differential resource allocation and the cost of sclerophylly and resprouting ability. Considering the life strategy of the Canary Island pine, we interpret the prolonged juvenile phase as an unavoidable trade-off for the high tolerance of adults to harsh environments.