In xeric environments, an increase in drought is related to reduced forest productivity and to enhanced mortality. However, predictions of future forest development remain difficult as the mechanisms underlying the responses of mature trees to long-term variations in water availability are not well understood. Here, we aimed to compare the adjustments in radial growth and morphological needle and shoot traits of mature Scots pine (Pinus sylvestris L.) growing along open water channels with those of control trees growing under naturally dry conditions at three sites in Valais, an inner-Alpine dry valley of Switzerland. The trees growing along two channels had been irrigated since germination (>70 years), whereas those along another previously drained channel had been irrigated only from 2010 to 2012, when the channel was re-established, and could thus be used to quantify the short-term effects of re-irrigation. Linear mixed models revealed that needle and shoot lengths as well as early- and late-wood basal area increments (BAIs) were most responsive to short-term and long-term irrigation. However, the magnitude of the response to the short-term irrigation exceeded that of the long-term irrigation. An extreme drought during the first half of 2011 led to an immediate decrease in the needle length, needle width, and early- and late-wood BAIs of the control trees, whereas the shoot length and needle numbers of control trees reacted with a 1-year delay to the extreme drought, as the shoots were responding to water availability of previous year's summer. Such negative responses to dry climatic conditions were even found in irrigated trees at one of our sites, which might be linked to tree growth becoming more sensitive to drought with increasing tree height and leaf area. In order to improve predictions of future forest development, long-term studies are necessary that consider lagged responses and adjustment processes of trees to changes in water availability.

Higher atmospheric CO2 concentrations (c(a)) can under certain conditions increase tree growth by enhancing photosynthesis, resulting in an increase of intrinsic water-use efficiency (i WUE) in trees. However, the magnitude of these effects and their interactions with changing climatic conditions are still poorly understood under xeric and mesic conditions. We combined radial growth analysis with intra- and interannual δ(13)C and δ(18)O measurements to investigate growth and physiological responses of Larix decidua, Picea abies, Pinus sylvestris, Pinus nigra and Pseudotsuga menziesii in relation to rising c(a) and changing climate at a xeric site in the dry inner Alps and at a mesic site in the Swiss lowlands. (i)WUE increased significantly over the last 50 yr by 8-29% and varied depending on species, site water availability, and seasons. Regardless of species and increased (i)WUE, radial growth has significantly declined under xeric conditions, whereas growth has not increased as expected under mesic conditions. Overall, drought-induced stomatal closure has reduced transpiration at the cost of reduced carbon uptake and growth. Our results indicate that, even under mesic conditions, the temperature-induced drought stress has overridden the potential CO2 'fertilization' on tree growth, hence challenging today's predictions of improved forest productivity of temperate forests.

Planting provenances originating from southern to northern locations has been discussed as a strategy to speed up species migration and mitigate negative effects of climate change on forest stability and productivity. Especially for drought-susceptible species such as European beech (Fagus sylvatica L.), the introduction of drought-tolerant provenances from the south could be an option. Yet, beech has been found to respond plastically to environmental conditions, suggesting that the climate on the plantation site might be more important for tree growth than the genetic predisposition of potentially drought-adapted provenances. In this study, we compared the radial growth, wood-anatomical traits and leaf phenology of four beech provenances originating from southern (Bulgaria, France) and northern locations (Sweden, the Netherlands) and planted in a provenance trial in the Netherlands. The distribution of precipitation largely differs between the sites of origin. The northern provenances experience a maximum and the southern provenances experience a minimum of rainfall in summer. We compared tree productivity and the anatomy of the water-conducting system for the period from 2000 to 2010, including the drought year 2003. In addition, tree mortality and the timing of leaf unfolding in spring were analysed for the years 2001, 2007 and 2012. Comparison of these traits in the four beech provenances indicates the influence of genetic predisposition and local environmental factors on the performance of these provenances under moderate site conditions. Variation in radial growth was controlled by environment, although the growth level slightly differed due to genetic background. The Bulgarian provenance had an efficient water-conducting system which was moreover unaffected by the drought in 2003, pointing to a high ability of this provenance to cope well with dry conditions. In addition, the Bulgarian provenance showed up as most productive in terms of height and radial growth. Altogether, we conclude that the similarity in ring-width variation among provenances points to environmental control of this trait, whereas the differences encountered in wood-anatomical traits between the well-performing Bulgarian provenance and the other three provenances, as well as the consistent differences in flushing pattern over 3 years under various environmental conditions, support the hypothesis of genetic control of these features.

Our knowledge on tree responses to drought is mainly based on short-term manipulation experiments which do not capture any possible long-term adjustments in this response. Therefore, historical water channels in inner-Alpine dry valleys were used as century-long irrigation experiments to investigate adjustments in tree growth to contrasting water supply. This involved quantifying the tree-ring growth of irrigated and non-irrigated (control) Scots pine (Pinus sylvestris L.) in Valais (Switzerland), as well as European larch (Larix decidua Mill.) and black pine (Pinus nigra Arnold) in Vinschgau (Italy). Furthermore, the adjustments in radial growth of Scots pine and European larch to an abrupt stop in irrigation were analyzed.Irrigation promoted the radial growth of all tree species investigated compared to the control: (1) directly through increased soil water availability, and (2) indirectly through increased soil nutrients and humus contents in the irrigated plots. Irrigation led to a full elimination of growth responses to climate for European larch and black pine, but not for Scots pine, which might become more sensitive to drought with increasing tree size in Valais. For the control trees, the response of the latewood increment to water availability in July/August has decreased in recent decades for all species, but increased in May for Scots pine only. The sudden irrigation stop caused a drop in radial growth to a lower level for Scots pine or similar level for larch compared to the control for up to ten years. However, both tree species were then able to adjust to the new conditions and subsequently grew with similar (Scots pine) or even higher growth rates(larch) than the control.To estimate the impact of climate change on future forest development, the duration of manipulation experiments should be on longer time scales in order to capture adjustment processes and feedback mechanisms of forest ecosystems. (C) 2014 Published by Elsevier B.V.

Forests of the future should be resistant to exacerbating climatic conditions, especially to increasing drought, but at the same time provide a sufficient amount and quality of timber. In this context coastal Douglas-fir (Pseudotsuga menziesii (Mirb.)) is a promising species since it remains productive even under chronic drought. By choosing suitable provenances within the range of Douglas-fir (P. menziesii (Mirb.)) for a given site we can further optimise tree fitness under dry conditions or even increase timber yield.Eighteen coastal Douglas-fir (Pseudotsuga menziesii (Mirb.) var. menziesii) provenances were tested for seedling survival, yield, wood quality, and drought tolerance by taking advantage of a Dutch provenance trial, established in 1971 within the framework of the 1966/1967 IUFRO seed collection program. The site of the Dutch trial is representative for many sites in Central Europe and is characterised by a moderate precipitation and temperature regime. Measurements on height and diameter growth were combined with a dendrochronological study on growth response to drought years.We found a clear latitudinal trend indicating that Douglas-fir provenances from the northern part of the species-distribution range are generally more productive than provenances from the south. In contrast, drought tolerance increased towards the south. This suggests that it is impossible to identify provenances combining maximum productivity with lowest susceptibility towards drought. However, based on the results from the trial we can give recommendations on suitable provenances that are expected to perform best under future conditions in Central Europe. On sites where severe drought events are unlikely to occur in future, fast growing provenances from the north, like Nimkish, should be planted. These provenances respond plastically to drought years, but the strong reduction of tree growth in the drought year itself indicates that these provenances will be harmed by an increasing frequency of drought events. However, on sites where water availability is likely to decrease, provenances from the Olympic Peninsula like Forks and Matlock are very promising since they showed still relatively high yield in combination with a high potential to cope with drought.If summer drought increases in frequency and severity as expected, the latewood/earlywood ratio will be drastically reduced with negative consequences for wood quality and cavitation resistance. However, some provenances, like Marblemount or Matlock, might compensate for the negative effect of summer drought on latewood/earlywood ratio by the contribution of photosynthesis in winter to whole-year carbon stock. (C) 2013 Elsevier B.V. All rights reserved.

One short-term adjustment of trees to drought is the reduction of photosynthetic tissues via leaf shedding. But in conifers, it usually takes several years to fully restore needle mass and assimilation capacity.This study aims to evaluate whether leaf shedding sustainably damages conifers or if these trees still have the ability to recover from drought with respect to their foliage and wood formation.An irrigation experiment was established in a mature dry forest to test the growth reactions of Scots pine (Pinus sylvestris L.) differing in crown transparency (low, medium, high) to a drought release by irrigation in comparison with equivalent control trees growing under naturally dry conditions on the same site.Drought and high crown transparency had a combined negative effect on radial tree growth: Control trees with medium to high crown transparencies showed a substantially shorter growth period and a long-lasting growth depression in response to the severe summer drought in 2003. However, all trees benefited from irrigation, irrespective of their crown status, and immediately increased growth in response to irrigation.The progressed drought-induced defoliation seemed to be a weakening factor for trees suffering from drought, but this can be reversed if the water supply is improved.

Due to projected global warming, there is a great concern about the ability of European beech to adapt to future climate conditions. Provenance trials provide an excellent basis to assess the potential of various provenances to adjust to given climate conditions. In this study we compared the performance of four European beech (Fagus sylvatica L.) provenances growing in a provenance trial at the Fruska Gora Mountain, Serbia. Three of the investigated provenances (Hollerbach and Hasbruch from Germany and Vrani Kamen from Croatia) originate from moist sites, with annual precipitation sums being twice as high as at the provenance trial in Serbia. The performance of these provenances are compared to the growth of the local provenance Fruska Gora which is well adapted to dry site conditions. We analysed tree-ring width, mean vessel area, vessel density and water-conductive area for the period from 2006 to 2012. In spite of differences in climate conditions at their place of origin all beech provenances showed a similar pattern in radial increment. Also the wood- anatomical variables showed similar inter-annual patterns for all provenances and no significant differences between the provenances. This indicates that beech provenances from moist environments can adjust to the relatively dry temperate climate in Serbia.

The ability of tree species to cope with anticipated decrease in water availability is still poorly understood. We evaluated the potential of Norway spruce, Scots pine, European larch, black pine, and Douglas‐fir to withstand drought in a drier future climate by analyzing their past growth and physiological responses at a xeric and a mesic site in Central Europe using dendroecological methods. Earlywood, latewood, and total ring width, as well as the δ 13 C and δ 18 O in early‐ and latewood were measured and statistically related to a multiscalar soil water deficit index from 1961 to 2009. At the xeric site, δ 13 C values of all species were strongly linked to water deficits that lasted longer than 11 months, indicating a long‐term cumulative effect on the carbon pool. Trees at the xeric site were particularly sensitive to soil water recharge in the preceding autumn and early spring. The native species European larch and Norway spruce, growing close to their dry distribution limit at the xeric site, were found to be the most vulnerable species to soil water deficits. At the mesic site, summer water availability was critical for all species, whereas water availability prior to the growing season was less important. Trees at the mesic were more vulnerable to water deficits of shorter duration than the xeric site. We conclude that if summers become drier, trees growing on mesic sites will undergo significant growth reductions, whereas at their dry distribution limit in the Alps, tree growth of the highly sensitive spruce and larch may collapse, likely inducing dieback and compromising the provision of ecosystem services. However, the magnitude of these changes will be mediated strongly by soil water recharge in winter and thus water availability at the beginning of the growing season.

Wood-anatomical parameters form a valuable archive to study past limitations on tree growth and act as a link between dendrochronology and ecophysiology. Yet, analysing these parameters is a time-consuming procedure and only few long chronologies exist. To increase measurement efficiency of wood-anatomical parameters, novel tools like the automated image-analysis system ROXAS were developed. So far, ROXAS has only been applied to measure large earlywood vessels in ring-porous species.In this study, we evaluate if ROXAS is also suitable for efficient and accurate detection and measurement of vessels in diffuse-porous European beech. To do so, we compared the outcome of ROXAS with that of the established measurement programme Image-Pro Plus in terms of efficiency and accuracy.The two methods differed substantially in efficiency with automatic measurements using ROXAS being 19 times faster than with Image-Pro Plus. Although the procedures led to similar patterns in annual variation of mean vessel area and vessel density, the absolute values differed. Image-Pro Plus measured systematically lower mean vessel areas and higher vessel densities than ROXAS. This was attributed to the species-specific technical settings in ROXAS, leading to more realistic results than those obtained using the default settings in Image-Pro Plus. A shortcoming of ROXAS was, however, that small vessels (<100 μm²) could not be detected with sufficient accuracy. Yet, based on thin sections of European beech, it is generally difficult to distinguish such small vessels from parenchyma cells. Moreover, these small vessels do not contribute substantially to conductive efficiency. Therefore, we do not foresee any problems for most studies if the lower vessel area threshold to be measured is set to 100 μm². Overall, ROXAS proved to be useful for measuring vessel parameters in diffuse-porous tree species, allowing accurate and efficient analyses of large numbers of samples.

Studies on climate impacts on tree annual growth are mainly restricted to marginal sites. To date, the climate effects on annual growth of trees in favorable environments remain therefore unclear despite the importance of these sites in terms of forest productivity. Because species respond differently to climate, comparing a multitude of species further enhances our knowledge on climate impacts on tree growth and forest productivity.We present a first study that reveals to what extent radial growth is limited by climate and the groundwater table across 10 temperate tree species growing on a uniform, productive soil (i.e. high nutrient and water contents) in the Netherlands. We ranked our study species according to their shade tolerance, which is associated with species' resource requirements and growth rates, and examined their annual growth using tree-ring analysis. This allowed us to investigate how these species with diverse ecological backgrounds differ in their growth response to precipitation, temperature, irradiance, potential evapotranspiration (PET), and the groundwater table, when growing under similar and favorable site conditions.Nine out of 10 species had strikingly similar radial growth rates (on average 3.9-4.8 mm yr(-1)), which contradicts the widely established trade-off between shade tolerance and growth. Populus trichocarpa, the least shade-tolerant of our species, however grew much faster (on average 6.8 mm yr(-1)). Trees of all species reduced their growth significantly during dry summers, driven by low rainfall and high PET. Yet, the magnitude of their growth responses to climate conditions and especially the groundwater table differed across species. Receding groundwater tables significantly and strongly reduced radial growth of the shade-intolerant species, but not of the shade-tolerant species. The climate impacts on growth were not associated with variation in shade tolerance across our species.Our study demonstrates that even on a productive forest site, summer droughts reduced radial growth across a multitude of common tree species, whereas lower groundwater tables only affected the shade-intolerant species significantly. Thus, the productivity of forests in favorable environments may be seriously affected when summers in north western Europe become hotter and drier as predicted by climate models. (C) 2013 Elsevier B.V. All rights reserved.

An increasing number of studies have reported on forest declines and vegetation shifts triggered by drought. In the Swiss Rhone valley (Valais), one of the driest inner-Alpine regions, the species composition in low elevation forests is changing: The sub-boreal Scots pine (Pinus sylvestris L.) dominating the dry forests is showing high mortality rates. Concurrently the sub-Mediterranean pubescent oak (Quercus pubescens Willd.) has locally increased in abundance. However, it remains unclear whether this local change in species composition is part of a larger-scale vegetation shift. To study variability in mortality and regeneration in these dry forests we analysed data from the Swiss national forest inventory (NFI) on a regular grid between 1983 and 2003, and combined it with annual mortality data from a monitoring site. Pine mortality was found to be highest at low elevation (below 1000 m a.s.l.). Annual variation in pine mortality was correlated with a drought index computed for the summer months prior to observed tree death. A generalized linear mixed-effects model indicated for the NFI data increased pine mortality on dryer sites with high stand competition, particularly for small-diameter trees. Pine regeneration was low in comparison to its occurrence in the overstorey, whereas oak regeneration was comparably abundant. Although both species regenerated well at dry sites, pine regeneration was favoured at cooler sites at higher altitude and oak regeneration was more frequent at warmer sites, indicating a higher adaptation potential of oaks under future warming. Our results thus suggest that an extended shift in species composition is actually occurring in the pine forests in the Valais. The main driving factors are found to be climatic variability, particularly drought, and variability in stand structure and topography. Thus, pine forests at low elevations are developing into oak forests with unknown consequences for these ecosystems and their goods and services.

Climate change is challenging forestry management and practices. Among other things, tree species with the ability to cope with more extreme climate conditions have to be identified. However, while environmental factors may severely limit tree growth or even cause tree death, assessing a tree species' potential for surviving future aggravated environmental conditions is rather demanding. The aim of this study was to find a tree-ring-based method suitable for identifying very drought-tolerant species, particularly potential substitute species for Scots pine (Pinus sylvestris L.) in Valais. In this inner-Alpine valley, Scots pine used to be the dominating species for dry forests, but today it suffers from high drought-induced mortality. We investigate the growth response of two native tree species, Scots pine and European larch (Larix decidua Mill.), and two non-native species, black pine (Pinus nigra Arnold) and Douglas fir (Pseudotsuga menziesii Mirb. var. menziesii), to drought. This involved analysing how the radial increment of these species responded to increasing water shortage (abandonment of irrigation) and to increasingly frequent drought years. Black pine and Douglas fir are able to cope with drought better than Scots pine and larch, as they show relatively high radial growth even after irrigation has been stopped and a plastic growth response to drought years. European larch does not seem to be able to cope with these dry conditions as it lacks the ability to recover from drought years. The analysis of trees' short-term response to extreme climate events seems to be the most promising and suitable method for detecting how tolerant a tree species is towards drought. However, combining all the methods used in this study provides a complete picture of how water shortage could limit species.

Forest and climate change in the inner-Alpine dry region of Visp Over the past decades, observed increases in temperature have been particularly pronounced in mountain regions. If this trend should continue in the 21st Century, frequency and intensity of droughts will increase, and will pose major challenges for forest management. Under current conditions drought-related tree mortality is already an important factor of forest ecosystems in dry inner-Alpine valleys. Here we assess the sensitivity of forest ecosystems to climate change and evaluate alternative forest management strategies in the Visp region. We integrate data from forest monitoring plots, field experiments and dynamic forests models to evaluate how the forest ecosystem services timber production, protection against natural hazards, carbon storage and biodiver-sity will be impacted. Our results suggest that at dry low elevation sites the drought tolerance of native tree species will be exceeded so that in the longer term a transition to more drought-adapted species should be considered. At medium elevations, drought and insect disturbances as by bark beetles are projected to be important for forest development, while at high elevations forests are projected to expand and grow better. All of the ecosystem services that we considered are projected to be impacted by changing forest conditions, with the specific impacts often being elevation-dependent. In the medium term, forest management that aims to increase the resilience of forests to drought can help maintain forest ecosystem services temporarily. However, our results suggest that relatively rigid management interventions are required to achieve significant effects. By using a combination of environmental monitoring, field experiments and modeling, we are able to gain insight into how forest ecosystem, and the services they provide, will respond to future changes.

Drought has been frequently discussed as a trigger for forest decline. Today, large-scale Scots pine decline is observed in many dry inner-Alpine valleys, with drought discussed as the main causative factor. This study aimed to analyse the impact of drought on wood formation and wood structure. To study tree growth under contrasting water supply, an irrigation experiment was installed in a mature Scots pine (Pinus sylvestris L.) forest at a xeric site in a dry inner-Alpine valley. Inter- and intra-annual radial increments as well as intra-annual variations in wood structure of pine trees were studied. It was found that non-irrigated trees had a noticeably shorter period of wood formation and showed a significantly lower increment. The water conduction cells were significantly enlarged and had significantly thinner cell walls compared with irrigated trees. It is concluded that pine trees under drought stress build a more effective water-conducting system (larger tracheids) at the cost of a probably higher vulnerability to cavitation (larger tracheids with thinner cell walls) but without losing their capability to recover. The significant shortening of the growth period in control trees indicated that the period where wood formation actually takes place can be much shorter under drought than the 'potential' period, meaning the phenological growth period.