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Highlights • Lowering density to cope with drier conditions might invalidate dendrometric rules. • We validated the density and site index-stand growth relationships at low density. • Top height growth increased with density, biasing estimates of site index. • In the productive area of sessile oak in France, climatic effects were weak. Abstract Context Lowering stand density has been suggested to adapt forests to warmer and drier conditions. Whether common dendrometric rules used to guide growth models and support silviculture are still valid at these densities lower than usual needs be tested. This includes (1) estimating the stand growth-density relationship over wider density gradients ('Langsaeter's rule'), (2) testing the stability of site index, and (3) 'Eichhorn's rule' (relating site index to stand growth) at lower densities. Additionally, environmental changes call for a more direct inclusion of environmental drivers in growth models. Material and methods We investigated the effects of relative stand density (RDI) on stand growth of even-aged stands of sessile oak. We also analyzed effects of climatic conditions to assess whether sessile oak is already at a critical stage that requires adapting forest management. We used a network of silvicultural trials designed to test the effects of maximum density gradients on stand dynamics along climatic gradients. Results As predicted by Langsaeter's rule, gross stand growth increased with RDI then saturated for RDI > 0.4–0.5, mortality began after crown closure, and diameter increment decreased non-linearly with RDI. Dominant height growth first increased with RDI then saturated for RDI > 0.8. Positive effects of RDI on both stand and height growth caused Eichhorn's rule to be stable over a large range of RDI (0.25–1). Effects of climate predictors were weak and only visible for radial growth indices. They however suggested antagonistic effects of water and temperature constraints in summer and autumn. Discussion Langsaeter's and Eichhorn's rules were found valid over a large range of RDI values. We suggest using them to guide growth models and management. However, assumptions behind the use of site index as a proxy for stand productivity were invalidated. Weak and opposed effect of temperature in summer and autumn makes the outcome of climate warming on sessile oak growth uncertain. [ABSTRACT FROM AUTHOR]

• Phenotypic variation in vigor and sensitivity is greater within than between stands. • Vigorous trees are more sensitive to drought stress. • Within-stand variances and correlations vary during stand development. • Phenotypic variation of sensitivity, but not vigor, is lower in stressful conditions. • Correlation between vigor and sensitivity decreases with higher soil water content. Within-population variation in individual tree growth and response to climate has an impact on forest dynamics, resilience and adaptation to environmental change. Combining dendrochronological analyses with a process-based ecophysiological model simulating drought stress at the stand scale, we studied the phenotypic variation of two growth-related traits within 22 pure stands of five contrasted tree species sampled in the RENECOFOR network over a wide range of ecological conditions. First, we computed the annual stress level from soil, climate and stand inventory data. Second, we computed individual sensitivity as the quantitative growth response to drought stress level and individual vigor as the capacity to grow in favorable years relative to an average stand-level growth model. We analyzed within-population variation and covariation of individual vigor and sensitivity, their temporal changes during stand development, as well as the effect of environmental conditions on population-level means, variances and correlation. Our results show that within-population variances in sensitivity and vigor exceed the between-population variances for all species. The populations located in more stressful environments, i.e., low summer precipitation and extractable soil water, showed lower mean and variance of sensitivity, suggesting possible multiscale adaptation at the population level and within populations. None of the environmental factors considered had an effect on the average population vigor or on the within-population variance of vigor. We found a general positive correlation between individual growth sensitivity and vigor in 17 out of 22 populations, potentially revealing a growth performance trade-off. The correlation was more pronounced in low extractable soil water environments, which may be related to a need for stressful conditions to reveal the trade-off or be the consequence of adaptive processes, i.e., acclimation and selection. If high within-population stand phenotypic variation in growth traits contributes to the resilience and adaptive capacity of forests to climate change, a trade-off could represent a constraint on selection. We provide genetic and environmental arguments supporting the hypothesis of a trade-off, then we highlight the importance of integrating it into the management process, especially during selective thinning, to avoid indirectly increasing population sensitivity by selecting the most vigorous trees. [ABSTRACT FROM AUTHOR]

Mixing species has been proposed as a forest management option to mitigate the effects of the increase in summer drought episodes predicted by climate change models in temperate zones. Studying how the relationship between tree rings and summer drought episodes is modified by species mixture is of primary importance in this context. Based on a dendroclimatology study, we (i) qualified and quantified the relationship between ring width (RW) or mean ring density (MRD) and soil water deficit (SWD) in a ring-porous species, Quercus petraea , and a coniferous species, Pinus sylvestris, and (ii) determined if the relationships between tree ring characteristics and SWD are modified in mixtures. We found inter-specific differences in RW patterns depending on the seasonality of the drought periods: RW in Q. petraea responded mainly to SWD from May to July, and RW in P. sylvestris to SWD from May to September. The ring-width of both species was strongly reduced by SWD; however, inter-specific differences in response to SWD seasonality triggered RW variations of opposite directions between the two species, for several years. In contrast, MRD for both species was dependent on the SWD from June to July. MRD in Q. petraea increased slightly with SWD whereas MRD in P. sylvestris decreased strongly with SWD. The relationship between RW and SWD was not modified in mixtures compared to pure stands. We found that the effect of SWD on MRD was stronger in mixtures. The impact of SWD on species may be worsened in mixtures, but mixture of species with different growth sensitivities to the seasonality of the drought periods and to the drought intensity might help to buffer the effects of climatic fluctuations on the overall mixed stands productivity compared to pure stands. [ABSTRACT FROM AUTHOR]

• Patterns of mixing effects on pine/beech growth-climate relationships were analyzed across Europe. • Tree growth-climate relationships were driven by the regional climate conditions. • Differences in climate-growth relationships between pure and mixed beech stands were evidenced in the driest climates. • On average, mixing had no significant effect on resistance to drought events. • Growth reduction during drought events was lower in mixed compared to pure pine stands in sites with higher water balance in autumn. Increasing species diversity is considered a promising strategy to mitigate the negative impacts of global change on forests. However, the interactions between regional climate conditions and species-mixing effects on climate-growth relationships and drought resistance remain poorly documented. In this study, we investigated the patterns of species-mixing effects over a large gradient of environmental conditions throughout Europe for European beech (Fagus sylvatica L.) and Scots pine (Pinus sylvestris L.), two species with contrasted ecological traits. We hypothesized that across large geographical scales, the difference of climate-growth relationships and drought resistance between pure and mixed stands would be dependent on regional climate. We used tree ring chronologies derived from 1143 beech and 1164 pine trees sampled in 30 study sites, each composed of one mixed stand of beech and pine and of the two corresponding pure stands located in similar site conditions. For each site and stand, we used Bootstrapped Correlation Coefficients (BCCs) on standardized chronologies and growth reduction during drought years on raw chronologies to analyze the difference in climate-tree growth relationships and resistance to drought between pure and mixed stands. We found consistent large-scale spatial patterns of climate-growth relationships. Those patterns were similar for both species. With the exception of the driest climates where pure and mixed beech stands tended to display differences in growth correlation with the main climatic drivers, the mixing effects on the BCCs were highly variable, resulting in the lack of a coherent response to mixing. No consistent species-mixing effect on drought resistance was found within and across climate zones. On average, mixing had no significant effect on drought resistance for neither species, yet it increased pine resistance in sites with higher climatic water balance in autumn. Also, beech and pine most often differed in the timing of their drought response within similar sites, irrespective of the regional climate, which might increase the temporal stability of growth in mixed compared to pure stands. Our results showed that the impact of species mixing on tree response to climate did not strongly differ between groups of sites with distinct climate characteristics and climate-growth relationships, indicating the interacting influences of species identity, stand characteristics, drought events characteristics as well as local site conditions. [ABSTRACT FROM AUTHOR]

To better understand how the radial growth of beech ( Fagus sylvatica L.) high forests has responded to climate change, we selected 12 sites (137 trees) with optimal growing conditions along a W–E altitudinal gradient (67–590 m) in Belgium. We evaluated temporal changes in growth response to climate by using pointer year analysis, moving mean sensitivities (1860–2011), and moving bootstrapped correlation coefficients (1952–2011). The strongest driving climatic variables were identified by using the partial least squares method. The common patterns of growth trends, pointer years, and mean sensitivities among sites provided evidences for the impact of environmental changes operating at a regional scale. The results of growth–climate analysis indicated that these changes were strongly influenced by the climatic conditions of the previous year. The climate sensitivity of beech increased progressively in response to more frequent and intense heat waves and warming-related droughts, especially during recent decades, leading to remarkable inter-site synchronization. The changes were much more pronounced for sites located in lowlands (<300 m). The differences in growth responses along the altitudinal gradient and the consequences of warming for beech growth and physiology are discussed. [ABSTRACT FROM AUTHOR]

Context: Growth partitioning among trees in forest stands is pivotal to silviculture, making it crucial to understand its control by factors such as stand development, stand density, or thinning. Since growth partitioning primarily depends on the partitioning of environmental resources among individuals, climatic change further calls for extending this framework to explicit climatic factors. Recent debate on adapting management to such changes also requires larger density gradients to be encompassed. Methods: We primarily aimed to investigate the effects of stand density and climatic factors on growth partitioning, in even-aged stands of sessile oak and Douglas-fir, two species currently managed under contrasted silvicultural regimes. We used two original permanent plot networks designed to explore effects of large density gradients, from open-grown to self-thinning situations. Growth partitioning was assessed on basal area growth, using both the growth dominance index, and the within-stand size-growth relationship. Their dependence on stand density, age, thinning, and climatic predictors was modeled statistically. A one-at-a-time sensitivity analysis of these models was performed to evaluate the magnitude of the effect of each predictor on growth partitioning. Simulations of the effect of extreme climatic conditions on stand growth, and on dominant, intermediate and close-to-suppressed trees growth were also performed. Results: For both species, stand density was found to strongly increase growth partitioning toward the biggest trees. Stand growth in sessile oak was reduced by high summer soil water deficit, with a particularly severe growth reduction for suppressed trees, suggesting asymmetric belowground competition for water in this species. In Douglas-fir, a stand growth reduction was found for high summer temperatures, with an increase in growth dominance that suggested a higher temperature-driven stress for suppressed trees. In addition, age slightly increased/decreased growth dominance in sessile oak/Douglas-fir, respectively. Conclusions: Growth dominance and size-growth relationships offered complementary insight into growth partitioning. Stand density appears to be the major driver of growth partitioning. Climatic factors were also shown to significantly affect growth partitioning, with species differences, in addition to stand density and ageing. These results suggest to maintain stands at medium density levels to reduce rotation length and minimize risk of exposure to extreme climatic events. [ABSTRACT FROM AUTHOR]

Abstract: Regional chronology and climate–growth relationships assessment are known to be sensitive to sampling effort. To disentangle the respective benefits of increasing whether the number of plots or the number of trees per plot when investigating climate–growth relationships under temperate conditions, we propose to simulate samples from a set of 84 plots established in plantations of Corsican pine (Pinus nigra Arnold ssp. laricio Poiret var. Corsicana), within which 10 dominant trees were cored in 1992. The effect of sampling effort was investigated through 20 modalities of sampled plots (from 3 to 84) and 10 modalities of sampled trees per plot (from 1 to 10). Regional chronology was studied using the mean effective correlation and the expressed population signal, while climate–growth relationships were evaluated through correlation functions. The calculation of the correlation functions also allowed testing the effect of sampling effort on various climatic regressors presenting different climate–growth correlation strengths. The accuracy of the dendroecological investigations increased with increasing sample size: estimating climate sensitivity with a regional chronology built from a small sample led to a general under-estimation of the climate–growth correlations. Decreasing sample size also increased the risk of estimating (i) “false” non-significant correlations for the most influencing climatic regressors, and (ii) “false” significant correlations for the less influencing ones. Increasing the number of plots was found of a greater interest than increasing the number of trees per plot to improve the climate–growth relationships assessment. Finally, the analysis revealed that the improvement of the expressed population signal with increasing sampling effort did not linearly mirror the bettering in the climate–growth correlations assessment. [Copyright &y& Elsevier]

Abstract: In most dendrochronological studies, climate–growth relationships are established on dominant trees to minimize non-climatic signals. However, response to environmental factors may be affected by tree-size, which begs the question of the representativeness of dominant trees on the stand level. To highlight the variations in climate–growth relationships among sizes and species, under a wide range of ecological conditions (climate and soil properties), 61 pure even-aged stands were sampled across France. At each stand, two tree-ring chronologies were established from 10 big- to 10 small-diameter trees. Our objectives were, (1) to assess variations in climate sensitivity between the two size-diameter classes, and (2) to investigate the role of species and ecological conditions on these variations. The climate-growth relationships were evaluated from 122 tree-ring chronologies (1220 trees) through extreme growth years and correlation function analyses. Sensitivity to climate of shade-intolerant and moderately shade-tolerant species (Picea abies (L.) Karst., Pinus sylvestris L. and Quercus petraea (Matt.) Liebl.) remained constant between the size-diameter classes for both temperature and hydric balance, while the shade-tolerant species Abies alba Mill. and Fagus sylvatica L. displayed significant differences, with larger trees being more sensitive to summer drought than smaller trees. This difference increased with increasing climatic xericity. Our results suggest that, for shade-tolerant species, (1) big trees could be more sensitive to climatic change especially under xeric climate, and (2) future tree ring studies should include trees stratified by size to produce unbiased estimation of sensitivity to climate. [Copyright &y& Elsevier]

The low-latitudinal range margins of many temperate and boreal tree species consist of scattered populations that persist locally in climate refugia. Recent studies have shown that such populations can be remarkably resilient, yet their past resilience does not imply that they are immune to threats from future climate change. The functioning of refugial tree populations therefore needs to be better understood if we are to anticipate their prospects correctly. We performed a detailed study of tree radial growth and vigor in a long-term climate refugial population of beech (Fagus sylvatica), comparing the observed trends with those of co-occurring pedunculate oak (Quercus robur). Annual growth rates (basal area increment, BAI) for both species were similar to those observed in range-core populations, but natural lifespan was half that in the mountains. The master chronologies spanning 1870–2015 revealed 22% (Fagus) and 20% (Quercus) increases in BAI until the 1980s and a smaller decrease (−6% for Fagus , −9% for Quercus) since then. Stable carbon isotope measurements (δ13C) revealed no effect of cambial age and an increase in water-use efficiency (iWUE) from 1870–2015 of about 50% for Fagus and 20% for Quercus. The trend continued until 2015 in Fagus , whereas Quercus reached its maximum in the 1980s. A detailed analysis of the relationship between climate and annual growth based on a 118-year meteorological record revealed a major role of water availability in the current and previous year. We used the observed climatic relationships to model future growth trends until 2100 for the IPCC scenarios RCP4.5 and RCP8.5. Most projections revealed no change in current growth rates, suggesting that this climate refugium will be able to provide suitable conditions for the persistence of Fagus and Quercus over the coming decades even under warmer and drier regional climate conditions. Overall, our study provides valuable insight into the precise climatic and biological mechanisms enhancing the persistence of refugial tree populations under ongoing climate change. [ABSTRACT FROM AUTHOR]