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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

Tree species mixing of oak (Quercus petraea (Matt.) Liebl./Quercus robur L.) and pine (Pinus sylvestris L.) has been shown to have positive effects on ecosystem service provision. From a management perspective, however, it is still uncertain which thinning regime provides the highest possible productivity of mixed oak–pine forests in the long term. Because of a lack of empirical studies dealing with thinning and species mixing effects on oak–pine forests, we simulated forest growth in order to test which thinning type and intensity may provide the highest productivity in the long-term. To achieve this, we simulated the growth of pure and mixed stands of oak and pine for 100 years in 23 triplets located on an ecological gradient across Europe. For this purpose, we applied four different growth simulators and compared their results: the distance-independent single-tree simulator PROGNAUS, the distance-dependent single-tree simulator SILVA, the gap model ForCEEPS, and the process-based simulator 3D-CMCC-FEM. We investigated the effects of species mixing and thinning from the upper (thinning from above) and lower tail (thinning from below) of the diameter distribution by reducing the stand basal area to 50 and 80% of the maximum basal area. We compared simulated results of the relative volume productivity of mixed versus pure stands and of thinned versus unthinned stands to empirical results previously obtained on the same set of triplets. Simulated relative volume productivity ranged between 61 and 156%, although extremes of 10% and of 300% could be observed. We found the relative volume productivity to be influenced by stand age, but not by stand density, except for PROGNAUS. Relative volume productivity did not increase with the site water supply of the triplet location. Highest long-term productivity for oak, pine and oak–pine stands can be expected in consequence of thinning from above, but the effect of thinning intensity differed between simulators. Thinning effe

The increasing disturbances in monocultures around the world are testimony to their instability under global change. Many studies have claimed that temporal stability of productivity increases with species richness, although the ecological fundamentals have mainly been investigated through diversity experiments. To adequately manage forest ecosystems, it is necessary to have a comprehensive understanding of the effect of mixing species on the temporal stability of productivity and the way in which it is influenced by climate conditions across large geographical areas. Here, we used a unique dataset of 261 stands combining pure and two-species mixtures of four relevant tree species over a wide range of climate conditions in Europe to examine the effect of species mixing on the level and temporal stability of productivity. Structural equation modelling was employed to further explore the direct and indirect influence of climate, overyielding, species asynchrony and additive effect (i.e. temporal stability expected from the species growth in monospecific stands) on temporal stability in mixed forests. We showed that by adding only one tree species to monocultures, the level (overyielding: +6%) and stability (temporal stability: +12%) of stand growth increased significantly. We identified the key effect of temperature on destabilizing stand growth, which may be mitigated by mixing species. We further confirmed asynchrony as the main driver of temporal stability in mixed stands, through both the additive effect and species interactions, which modify between-species asynchrony in mixtures in comparison to monocultures. Synthesis and applications. This study highlights the emergent properties associated with mixing two species, which result in resource efficient and temporally stable production systems. We reveal the negative impact of mean temperature on temporal stability of forest productivity and how the stabilizing effect of mixing two species can counterbalance this

Heterogeneity of structure can increase mechanical stability, stress resistance and resilience, biodiversity and many other functions and services of forest stands. That is why many silvicultural measures aim at enhancing structural diversity. However, the effectiveness and potential of structuring may depend on the site conditions. Here, we revealed how the stand structure is determined by site quality and results from site-dependent partitioning of growth and mortality among the trees. We based our study on 90 mature, even-aged, fully stocked monocultures of Scots pine (Pinus sylvestris L.) sampled in 21 countries along a productivity gradient across Europe. A mini-simulation study further analyzed the site-dependency of the interplay between growth and mortality and the resulting stand structure. The overarching hypothesis was that the stand structure changes with site quality and results from the site-dependent asymmetry of competition and mortality. First, we show that Scots pine stands structure across Europe become more homogeneous with increasing site quality. The coefficient of variation and Gini coefficient of stem diameter and tree height continuously decreased, whereas Stand Density Index and stand basal area increased with site index. Second, we reveal a site-dependency of the growth distribution among the trees and the mortality. With increasing site index, the asymmetry of both competition and growth distribution increased and suggested, at first glance, an increase in stand heterogeneity. However, with increasing site index, mortality eliminates mainly small instead of all-sized trees, cancels the size variation and reduces the structural heterogeneity. Third, we modelled the site-dependent interplay between growth partitioning and mortality. By scenario runs for different site conditions, we can show how the site-dependent structure at the stand level emerges from the asymmetric competition and mortality at the tree level and how the interplay chang

Heterogeneity of structure can increase mechanical stability, stress resistance and resilience, biodiversity and many other functions and services of forest stands. That is why many silvicultural measures aim at enhancing structural diversity. However, the effectiveness and potential of structuring may depend on the site conditions. Here, we revealed how the stand structure is determined by site quality and results from site-dependent partitioning of growth and mortality among the trees. We based our study on 90 mature, even-aged, fully stocked monocultures of Scots pine (Pinus sylvestris L.) sampled in 21 countries along a productivity gradient across Europe. A mini-simulation study further analyzed the site-dependency of the interplay between growth and mortality and the resulting stand structure. The overarching hypothesis was that the stand structure changes with site quality and results from the site-dependent asymmetry of competition and mortality. First, we show that Scots pine stands structure across Europe become more homogeneous with increasing site quality. The coefficient of variation and Gini coefficient of stem diameter and tree height continuously decreased, whereas Stand Density Index and stand basal area increased with site index. Second, we reveal a site-dependency of the growth distribution among the trees and the mortality. With increasing site index, the asymmetry of both competition and growth distribution increased and suggested, at first glance, an increase in stand heterogeneity. However, with increasing site index, mortality eliminates mainly small instead of all-sized trees, cancels the size variation and reduces the structural heterogeneity. Third, we modelled the site-dependent interplay between growth partitioning and mortality. By scenario runs for different site conditions, we can show how the site-dependent structure at the stand level emerges from the asymmetric competition and mortality at the tree level and how the interplay chang

The increasing disturbances in monocultures around the world are testimony to their instability under global change. Many studies have claimed that temporal stability of productivity increases with species richness, although the ecological fundamentals have mainly been investigated through diversity experiments. To adequately manage forest ecosystems, it is necessary to have a comprehensive understanding of the effect of mixing species on the temporal stability of productivity and the way in which it is influenced by climate conditions across large geographical areas. Here, we used a unique dataset of 261 stands combining pure and two-species mixtures of four relevant tree species over a wide range of climate conditions in Europe to examine the effect of species mixing on the level and temporal stability of productivity. Structural equation modelling was employed to further explore the direct and indirect influence of climate, overyielding, species asynchrony and additive effect (i.e. temporal stability expected from the species growth in monospecific stands) on temporal stability in mixed forests. We showed that by adding only one tree species to monocultures, the level (overyielding: +6%) and stability (temporal stability: +12%) of stand growth increased significantly. We identified the key effect of temperature on destabilizing stand growth, which may be mitigated by mixing species. We further confirmed asynchrony as the main driver of temporal stability in mixed stands, through both the additive effect and species interactions, which modify between-species asynchrony in mixtures in comparison to monocultures. Synthesis and applications. This study highlights the emergent properties associated with mixing two species, which result in resource efficient and temporally stable production systems. We reveal the negative impact of mean temperature on temporal stability of forest productivity and how the stabilizing effect of mixing two species can counterbalance this

The increasing disturbances in monocultures around the world are testimony to their instability under global change. Many studies have claimed that temporal stability of productivity increases with species richness, although the ecological fundamentals have mainly been investigated through diversity experiments. To adequately manage forest ecosystems, it is necessary to have a comprehensive understanding of the effect of mixing species on the temporal stability of productivity and the way in which it is influenced by climate conditions across large geographical areas. Here, we used a unique dataset of 261 stands combining pure and two-species mixtures of four relevant tree species over a wide range of climate conditions in Europe to examine the effect of species mixing on the level and temporal stability of productivity. Structural equation modelling was employed to further explore the direct and indirect influence of climate, overyielding, species asynchrony and additive effect (i.e. temporal stability expected from the species growth in monospecific stands) on temporal stability in mixed forests. We showed that by adding only one tree species to monocultures, the level (overyielding: +6%) and stability (temporal stability: +12%) of stand growth increased significantly. We identified the key effect of temperature on destabilizing stand growth, which may be mitigated by mixing species. We further confirmed asynchrony as the main driver of temporal stability in mixed stands, through both the additive effect and species interactions, which modify between-species asynchrony in mixtures in comparison to monocultures. Synthesis and applications. This study highlights the emergent properties associated with mixing two species, which result in resource efficient and temporally stable production systems. We reveal the negative impact of mean temperature on temporal stability of forest productivity and how the stabilizing effect of mixing two species can counterbalance th

The increasing disturbances in monocultures around the world are testimony to their instability under global change. Many studies have claimed that temporal stability of productivity increases with species richness, although the ecological fundamentals have mainly been investigated through diversity experiments. To adequately manage forest ecosystems, it is necessary to have a comprehensive understanding of the effect of mixing species on the temporal stability of productivity and the way in which it is influenced by climate conditions across large geographical areas. Here, we used a unique dataset of 261 stands combining pure and two-species mixtures of four relevant tree species over a wide range of climate conditions in Europe to examine the effect of species mixing on the level and temporal stability of productivity. Structural equation modelling was employed to further explore the direct and indirect influence of climate, overyielding, species asynchrony and additive effect (i.e. temporal stability expected from the species growth in monospecific stands) on temporal stability in mixed forests. We showed that by adding only one tree species to monocultures, the level (overyielding: +6%) and stability (temporal stability: +12%) of stand growth increased significantly. We identified the key effect of temperature on destabilizing stand growth, which may be mitigated by mixing species. We further confirmed asynchrony as the main driver of temporal stability in mixed stands, through both the additive effect and species interactions, which modify between-species asynchrony in mixtures in comparison to monocultures. Synthesis and applications. This study highlights the emergent properties associated with mixing two species, which result in resource efficient and temporally stable production systems. We reveal the negative impact of mean temperature on temporal stability of forest productivity and how the stabilizing effect of mixing two species can counterbalance this

The increasing disturbances in monocultures around the world are testimony to their instability under global change. Many studies have claimed that temporal stability of productivity increase with species richness, although the ecological fundaments have mainly been investigated through diversity experiments. To adequately manage forest ecosystems, it is necessary to have a comprehensive understanding of the effect of mixing species on the temporal stability of productivity and the way in which this it is influenced by climate conditions across large geographical areas. Here, we used a unique dataset of 261 stands combining pure and two-species mixtures of four relevant tree species over a wide range of climate conditions in Europe to examine the effect of species mixing on the level and temporal stability of productivity. Structural equation modelling was employed to further explore the direct and indirect influence of climate, overyielding, species asynchrony and additive effect (i.e. temporal stability expected from the species growth in monospecific stands) on temporal stability in mixed forests. We showed that by adding only one tree species to monocultures, the level (overyielding: +6%) and stability (temporal stability: +12%) of stand growth increased significantly. We identified the key effect of temperature on destabilizing stand growth, which may be mitigated by mixing species. We further confirmed asynchrony as the main driver of temporal stability in mixed stands, through both the additive effect and species interactions, which modify between-species asynchrony in mixtures in comparison to monocultures. Synthesis and applications. This study highlights the emergent properties associated with mixing two-species, which result in resource efficient and temporally stable production systems. We reveal the negative impact of mean temperature on temporal stability of forest productivity and how the stabilizing effect of mixing two species can counterbalance thi

Heterogeneity of structure can increase mechanical stability, stress resistance and resilience, biodiversity and many other functions and services of forest stands. That is why many silvicultural measures aim at enhancing structural diversity. However, the effectiveness and potential of structuring may depend on the site conditions. Here, we revealed how the stand structure is determined by site quality and results from site-dependent partitioning of growth and mortality among the trees. We based our study on 90 mature, even-aged, fully stocked monocultures of Scots pine (Pinus sylvestris L.) sampled in 21 countries along a productivity gradient across Europe. A mini-simulation study further analyzed the site-dependency of the interplay between growth and mortality and the resulting stand structure. The overarching hypothesis was that the stand structure changes with site quality and results from the site-dependent asymmetry of competition and mortality. First, we show that Scots pine stands structure across Europe become more homogeneous with increasing site quality. The coefficient of variation and Gini coefficient of stem diameter and tree height continuously decreased, whereas Stand Density Index and stand basal area increased with site index. Second, we reveal a site-dependency of the growth distribution among the trees and the mortality. With increasing site index, the asymmetry of both competition and growth distribution increased and suggested, at first glance, an increase in stand heterogeneity. However, with increasing site index, mortality eliminates mainly small instead of all-sized trees, cancels the size variation and reduces the structural heterogeneity.Third, we modelled the site-dependent interplay between growth partitioning and mortality. By scenario runs for different site conditions, we can show how the site-dependent structure at the stand level emerges from the asymmetric competition and mortality at the tree level and how the interplay change

Tree species mixing of oak (Quercus petraea (Matt.) Liebl./Quercus robur L.) and pine (Pinus sylvestris L.) has been shown to have positive effects on ecosystem service provision. From a management perspective, however, it is still uncertain which thinning regime provides the highest possible productivity of mixed oak–pine forests in the long term. Because of a lack of empirical studies dealing with thinning and species mixing effects on oak–pine forests, we simulated forest growth in order to test which thinning type and intensity may provide the highest productivity in the long-term. To achieve this, we simulated the growth of pure and mixed stands of oak and pine for 100 years in 23 triplets located on an ecological gradient across Europe. For this purpose, we applied four different growth simulators and compared their results: the distance-independent single-tree simulator PROGNAUS, the distance-dependent single-tree simulator SILVA, the gap model ForCEEPS, and the process-based simulator 3D-CMCC-FEM. We investigated the effects of species mixing and thinning from the upper (thinning from above) and lower tail (thinning from below) of the diameter distribution by reducing the stand basal area to 50 and 80% of the maximum basal area. We compared simulated results of the relative volume productivity of mixed versus pure stands and of thinned versus unthinned stands to empirical results previously obtained on the same set of triplets. Simulated relative volume productivity ranged between 61 and 156%, although extremes of 10% and of 300% could be observed. We found the relative volume productivity to be influenced by stand age, but not by stand density, except for PROGNAUS. Relative volume productivity did not increase with the site water supply of the triplet location. Highest long-term productivity for oak, pine and oak–pine stands can be expected in consequence of thinning from above, but the effect of thinning intensity differed between simulators. Thinning effe