Functional traits and adaptive capacity of European forests to climate change

Plants distributed across a wide range of environmental conditions are submitted to differential selective pressures. Long-term selection can lead to the development of adaptations to the local environment, generating ecotypic differentiation. Additionally, plant species can cope with this environmental variability by phenotypic plasticity. In this study, we examine the importance of both processes in coping with environmental heterogeneity in the Mediterranean sclerophyllous cork oak Quercus suber. For this purpose, we measured growth and key functional traits at the leaf level in 9-year-old plants across 2 years of contrasting precipitation (2005 and 2006) in a common garden. Plants were grown from acorns originated from 13 populations spanning a wide range of climates along the distribution range of the species. The traits measured were: leaf size (LS), specific leaf area (SLA), carbon isotope discrimination (Delta(13)C) and leaf nitrogen content per unit mass (N(mass)). Inter-population differences in LS, SLA and Delta(13)C were found. These differences were associated with rainfall and temperature at the sites of origin, suggesting local adaptation in response to diverging climates. Additionally, SLA and LS exhibited positive responses to the increase in annual rainfall. Year effect explained 28% of the total phenotypic variance in LS and 2.7% in SLA. There was a significant genotype x environment interaction for shoot growth and a phenotypic correlation between the difference in shoot growth among years and the annual mean temperature at origin. This suggests that populations originating from warm sites can benefit more from wet conditions than populations from cool sites. Finally, we investigated the relationships between functional traits and aboveground growth by several regression models. Our results showed that plants with lower SLA presented larger aboveground growth in a dry year and plants with larger leaf sizes displayed larger growth rates in both years. Overall, the study supports the adaptive value of SLA and LS for cork oak under a Mediterranean climate and their potentially important role for dealing with varying temperature and rainfall regimes through both local adaptation and phenotypic plasticity.

/pdf/phenotypic-plasticity-and-local-adaptation-in-leaf-eygpz6oprn.pdf

Phenotypic plasticity and local adaptation in leaf ecophysiological traits of 13 contrasting cork oak populations under different water availabilities

The genus Quercus is among the most widespread and species-rich tree genera in the northern hemisphere. The extraordinary species diversity in America and Asia together with the continuous continental distribution of a limited number of European species raise questions about how macro- and microevolutionary processes made the genus Quercus an evolutionary success. Synthesizing conclusions reached during the past three decades by complementary approaches in phylogenetics, phylogeography, genomics, ecology, paleobotany, population biology and quantitative genetics, this review aims to illuminate evolutionary processes leading to the radiation and expansion of oaks. From opposing scales of time and geography, we converge on four overarching explanations of evolutionary success in oaks: accumulation of large reservoirs of diversity within populations and species; ability for rapid migration contributing to ecological priority effects on lineage diversification; high rates of evolutionary divergence within clades combined with convergent solutions to ecological problems across clades; and propensity for hybridization, contributing to adaptive introgression and facilitating migration. Finally, we explore potential future research avenues, emphasizing the integration of microevolutionary and macroevolutionary perspectives.

Oaks: an evolutionary success story.

Increased drought severity is expected in the Mediterranean Basin over the twenty-first century, but our understanding of the potential of most forest tree species to cope with it remains uncertain. In this study, (1) we examined the potential effects of long-term selection and the capacity to respond to future changes in selective pressures in three populations of cork oak (Quercus suber L.). For this purpose, we evaluated the response to dry conditions of 45 open-pollinated trees originating from populations in Morocco, Portugal, and Spain. Growth, leaf size, specific leaf area (SLA), carbon isotope discrimination (Δ13C), leaf nitrogen content (Nmass), and total chlorophyll content (Chlmass) were measured in 9-year-old plants. (2) We also investigated the relationships between functional traits and aboveground growth by regression models. Plants presenting larger and more sclerophyllous leaves (low SLA and high leaf thickness) exhibited higher growths, with results suggesting that these traits are subjected to divergent selection in this species. Heritability estimates were moderately high for Δ13C (0.43 ± 0.25–0.83 ± 0.31) and stem diameter (0.40 ± 0.15–0.71 ± 0.28) for the tree populations. For the rest of the traits (except for annual growth), heritability values varied among populations, particularly for height, leaf size, leaf thickness, and Nmass. Our results suggest that natural selection has led to local adaptations and has also affected the genetic variance intrapopulation in these cork oak populations, although studies with a higher number of populations should be carried out across different years. Additionally, the absence of significant genetic correlations and the fact that correlated traits did not undergo opposing selection provided little evidence for constraints on evolution caused by genetic correlations.

Factors affecting cork oak growth under dry conditions: local adaptation and contrasting additive genetic variance within populations

We have studied in this work the natural variability in Holm oak (Quercus ilex subsp. ballota) by analyzing acorn morphometry and chemical composition in 13 populations from the Andalusia region (southern Spain). Acorn morphometry showed statistic significant differences between the populations with a higher acorn weight, length, and diameter in the northern provenances (TJA, GSE, CTH, CHU, APS, and PCO), while the Southern provenances (RMA, SAA, CCO, BCA, and SCA) showed lower value that even decreased with the altitude. Acorn chemical composition also showed statistic significant differences between populations. Morphometry and chemical composition data were subjected to clustering analysis, and two major clusters were detected. This analysis showed that northern populations (CTH, TJA, SCA, CHU, PCO, GSE, and APS) showed a tendency to present values highest in acorn weight (5.42 g vs. 3.21 g); acorn length (37.21 mm vs. 28.73 mm); acorn diameter (16.12 mm vs. 13.79 mm); ash (1.88% vs. 1.55%); protein (4.82% vs. 4.38%); fat (12.55% vs. 9.92%); and oleic acid (66.93 vs. 66.4%). On the contrary, Southern populations (SSA, VJA, CCO, BCA, RG, and RMA) showed a tendency to present values lowest in sugar (10.28% vs. 11.25%); energy (20,076.71 kJ/kg vs. 20,149.26 kJ/kg); palmitic (16.11% vs. 16.39%) and linoleic acids (13.07% vs. 13.11%). The populations studied here seemed to integrate acorn morphology and chemical composition related to environmental conditions of the original provenances. These data have provided biochemical markers that help establish phylogenic associations between populations and also reveal potentially account as an alternative source for dietary nutrition.

Population variability based on the morphometry and chemical composition of the acorn in Holm oak (Quercus ilex subsp. ballota [Desf.] Samp.)

Twenty-six provenances (2 340 plants) of cork oak (Quercus suber spp.) originating from Portugal, Spain, Italy, Morocco, Algeria, and Tunisia were tested for genetic variation among and within provenances by growth traits. Seven morphometrical characters were measured in 90 plants from each provenance. Analysis of variance showed highly significant differences for all characters. The phenotypic coefficient of differentiation reached 0.24 for the form and 0.22 for height, thus revealing a strong structuring between the provenances. Comparative study of growth among the provenances revealed more vigorous growth and better survival rate for those from Morocco, Spain, and Portugal, which may constitute better materials for afforestation. Furthermore, this variability appeared to be geographically structured and would be mainly genetically controlled, as cork oak provenances were cultivated under the same environmental conditions. Our results should be helpful for guide forest managers in afforestation.

/pdf/morphological-evaluation-of-cork-oak-quercus-suber-2vq51upgit.pdf

Lamjed Toumi

Papers

Morphological evaluation of cork oak (Quercus suber): Mediterranean provenance variability in Tunisia