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Background: Understanding the role of species identity in interactions among individuals is crucial for assessing the productivity and stability of mixed forests over time. However, there is limited knowledge concerning the variation in competitive effect and response of different species along climatic gradients. In this study, we investigated the importance of climate, tree size, and competition on the growth of three tree species: spruce (Picea abies), fir (Abies alba), and beech (Fagus sylvatica), and examined their competitive response and effect along a climatic gradient. Methods: We selected 39 plots distributed across the European mountains with records of the position and growth of 5,759 individuals. For each target species, models relating tree growth to tree size, climate and competition were proposed. Competition was modelled using a neighbourhood competition index that considered the effects of inter- and intraspecific competition on target trees. Competitive responses and effects were related to climate. Likelihood methods and information theory were used to select the best model. Results: Our findings revealed that competition had a greater impact on target species growth than tree size or climate. Climate did influence the competitive effects of neighbouring species, but it did not affect the target species' response to competition. The strength of competitive effects varied along the gradient, contingent on the identity of the interacting species. When the target species exhibited an intermediate competitive effect relative to neighbouring species, both higher inter- than intraspecific competitive effects and competition reduction occurred along the gradient. Notably, species competitive effects were most pronounced when the target species’ growth was at its peak and weakest when growing conditions were far from their maximum. Conclusions: Climate modulates the effects of competition from neighbouring trees on the target tree and not the susceptibil

Spatial and temporal variation in functional traits allows trees to adjust to shifting environmental conditions such as water stress. However, the change of traits, both mean and variances, along water availability gradients and across growing seasons, as well as their covariation with tree performance, have been rarely assessed. We examined intraspecific trait variation in coexisting evergreen (Quercus ilex ssp. ilex and Q. ilex ssp. ballota) and deciduous (Quercus faginea and Quercus humilis) Mediterranean oaks along a wide water availability gradient in northeastern Spain during six years. We measured leaf area (LA), shoot twig mass (Sm), leaf mass per area (LMA) and the ratio of shoot twig to leaf biomass (Sm:Lm). We characterized tree performance through basal area increment (BAI) and drought resilience indices. Higher variation was found within individuals than between individuals across populations and years. Within species, we found trait adjustments toward more conservative water-use (low LA and Sm and high LMA) with increasing drier conditions. Intraspecific trait variation was constrained by water availability, particularly on the deciduous species. In Q. ilex, trait variance of LMA positively covaried with annual BAI, whereas variance of LA, Sm and Sm:Lm was positively related to resistance and resilience against the severe 2012 drought in deciduous oaks. Our results support a tradeoff between the ability to tolerate drought and the capacity to cope with unpredictable changes in the environment through increased intraspecific trait variation, which may have implications on tree performance in the face of increased extreme events.

Process-based models and empirical modelling techniques are frequently used to (i) explore the sensitivity of tree growth to environmental variables, and (ii) predict the future growth of trees and forest stands under climate change scenarios. However, modelling approaches substantially influence predictions of the sensitivity of trees to environmental factors. Here, we used tree-ring width (TRW) data from 1630 beech trees from a network of 70 plots established across European mountains to build empirical predictive growth models using various modelling approaches. In addition, we used 3-PG and Biome-BGCMuSo process-based models to compare growth predictions with derived empirical models. Results revealed similar prediction errors (RMSE) across models ranging between 3.71 and 7.54 cm2 of basal area increment (BAI). The models explained most of the variability in BAI ranging from 54 % to 87 %. Selected explanatory variables (despite being statistically highly significant) and the pattern of the growth sensitivity differed between models substantially. We identified only five factors with the same effect and the same sensitivity pattern in all empirical models: tree DBH, competition index, elevation, Gini index of DBH, and soil silt content. However, the sensitivity to most of the climate variables was low and inconsistent among the empirical models. Both empirical and process-based models suggest that beech in European mountains will, on average, likely experience better growth conditions under both 4.5 and 8.5 RCP scenarios. The process-based models indicated that beech may grow better across European mountains by 1.05 to 1.4 times in warmer conditions. The empirical models identified several drivers of tree growth that are not included in the current process-based models (e.g., different nutrients) but may have a substantial effect on final results, particularly if they are limiting factors. Hence, future development of process-based models may build upon our findi

Quercus robur/Quercus petraea and Pinus sylvestris are widely distributed and economically important tree species in Europe co-occurring on mesotrophic, xeric and mesic sites. Increasing dry conditions may reduce their growth, but growth reductions may be modified by mixture, competition and site conditions. The annual diameter growth in monospecific and mixed stands along an ecological gradient with mean annual temperatures ranging from 5.5 °C to 11.5 °C was investigated in this study. On 36 triplets (108 plots), trees were cored and the year-ring series were cross-dated, resulting in year-ring series of 785 and 804 trees for Q. spp. and P. sylvestris, respectively. A generalized additive model with a logarithmic link was fit to the data with random effects for the intercept at the triplet, year and tree level and a random slope for the covariate age for each tree; the Tweedie-distribution was used. The final model explained 87 % of the total variation in diameter increment for both tree species. Significant covariates were age, climate variables (long-term mean, monthly), local competition variables, relative dbh, mixture, stand structure and interactions thereof. Tree growth declined with age and local density and increased with social position. It was positively influenced by mixture and structural diversity (Gini coefficient); mixture effects were significant for P. sylvestris only. The influence of potential evapotranspiration (PET) in spring and autumn on tree growth was positive and non-linear, whereas tree growth sharply decreased with increasing PET in June, which proved to be the most influential month on tree growth along the whole ecological gradient. Interactions of PET with tree social position (relative dbh) were significant in July and September for Q. spp. and in April for P. sylvestris. Interactions of climate with density or mixture were not significant. Climatic effects found agree well with previous results from intra-annual growth studies and

Quercus robur/Quercus petraea and Pinus sylvestris are widely distributed and economically important tree species in Europe co-occurring on mesotrophic, xeric and mesic sites. Increasing dry conditions may reduce their growth, but growth reductions may be modified by mixture, competition and site conditions. The annual diameter growth in monospecific and mixed stands along an ecological gradient with mean annual temperatures ranging from 5.5 °C to 11.5 °C was investigated in this study. On 36 triplets (108 plots), trees were cored and the year-ring series were cross-dated, resulting in year-ring series of 785 and 804 trees for Q. spp. and P. sylvestris, respectively. A generalized additive model with a logarithmic link was fit to the data with random effects for the intercept at the triplet, year and tree level and a random slope for the covariate age for each tree; the Tweedie-distribution was used. The final model explained 87 % of the total variation in diameter increment for both tree species. Significant covariates were age, climate variables (long-term mean, monthly), local competition variables, relative dbh, mixture, stand structure and interactions thereof. Tree growth declined with age and local density and increased with social position. It was positively influenced by mixture and structural diversity (Gini coefficient); mixture effects were significant for P. sylvestris only. The influence of potential evapotranspiration (PET) in spring and autumn on tree growth was positive and non-linear, whereas tree growth sharply decreased with increasing PET in June, which proved to be the most influential month on tree growth along the whole ecological gradient. Interactions of PET with tree social position (relative dbh) were significant in July and September for Q. spp. and in April for P. sylvestris. Interactions of climate with density or mixture were not significant. Climatic effects found agree well with previous results from intra-annual growth studies and

Quercus robur/Quercus petraea and Pinus sylvestris are widely distributed and economically important tree species in Europe co-occurring on mesotrophic, xeric and mesic sites. Increasing dry conditions may reduce their growth, but growth reductions may be modified by mixture, competition and site conditions. The annual diameter growth in monospecific and mixed stands along an ecological gradient with mean annual temperatures ranging from 5.5 °C to 11.5 °C was investigated in this study. On 36 triplets (108 plots), trees were cored and the year-ring series were cross-dated, resulting in year-ring series of 785 and 804 trees for Q. spp. and P. sylvestris, respectively. A generalized additive model with a logarithmic link was fit to the data with random effects for the intercept at the triplet, year and tree level and a random slope for the covariate age for each tree; the Tweedie-distribution was used. The final model explained 87 % of the total variation in diameter increment for both tree species. Significant covariates were age, climate variables (long-term mean, monthly), local competition variables, relative dbh, mixture, stand structure and interactions thereof. Tree growth declined with age and local density and increased with social position. It was positively influenced by mixture and structural diversity (Gini coefficient); mixture effects were significant for P. sylvestris only. The influence of potential evapotranspiration (PET) in spring and autumn on tree growth was positive and non-linear, whereas tree growth sharply decreased with increasing PET in June, which proved to be the most influential month on tree growth along the whole ecological gradient. Interactions of PET with tree social position (relative dbh) were significant in July and September for Q. spp. and in April for P. sylvestris. Interactions of climate with density or mixture were not significant. Climatic effects found agree well with previous results from intra-annual growth studies a