Summer drought exposure,stand structure,and soil properties jointly control the growth of European beech along a steep precipitation gradient in northern Germany

Increasing exposure to climate warming-related drought and heat threatens forest vitality in many regions on earth, with the trees' vulnerability likely depending on local climatic aridity, recent climate trends, edaphic conditions, and the drought acclimatization and adaptation of populations. Studies exploring tree species' vulnerability to climate change often have a local focus or model the species' entire distribution range, which hampers the separation of climatic and edaphic drivers of drought and heat vulnerability. We compared recent radial growth trends and the sensitivity of growth to drought and heat in central populations of a widespread and naturally dominant tree species in Europe, European beech (Fagus sylvatica), at 30 forest sites across a steep precipitation gradient (500–850 mm year⁻¹) of short length to assess the species' adaptive potential. Size-standardized basal area increment remained more constant during the period of accelerated warming since the early 1980s in populations with >360 mm growing season precipitation (April–September), while growth trends were negative at sites with <360 mm. Climatic drought in June appeared as the most influential climatic factor affecting radial growth, with a stronger effect at drier sites. A decadal decrease in the climatic water balance of the summer was identified as the most important factor leading to growth decline, which is amplified by higher stem densities. Inter-annual growth variability has increased since the early 1980s, and variability is generally higher at drier and sandier sites. Similarly, within-population growth synchrony is higher at sandier sites and has increased with a decrease in the June climatic water balance. Our results caution against predicting the drought vulnerability of trees solely from climate projections, as soil properties emerged as an important modulating factor. We conclude that beech is facing recent growth decline at drier sites in the centre of its distribution range, driven by climate change-related climate aridification.     

Site‐adapted admixed tree species reduce drought susceptibility of mature European beech

Some forest‐related studies on possible effects of climate change conclude that growth potential of European beech (Fagus sylvatica L.) might be impaired by the predicted increase in future serious drought events during the growing season. Other recent research suggests that not only multiyear increment rates but also growth resistance and recovery of beech during, respectively, after dry years may differ between pure and mixed stands. Thus, we combined dendrochronological investigations and wood stable isotope measurements to further investigate the impact of neighborhood diversity on long‐term performance, short‐term drought response and soil water availability of European beech in three major geographic regions of Germany. During the last four decades, target trees whose competitive neighborhood consisted of co‐occurring species exhibited a superior growth performance compared to beeches in pure stands of the same investigation area. This general pattern was also found in exceptional dry years. Although the summer droughts of 1976 and 2003 predominantly caused stronger relative growth declines if target trees were exposed to interspecific competition, with few exceptions they still formed wider annual rings than beeches growing in close‐by monocultures. Within the same study region, recovery of standardized beech target tree radial growth was consistently slower in monospecific stands than in the neighborhood of other competitor species. These findings suggest an improved water availability of beech in mixtures what is in line with the results of the stable isotope analysis. Apparently, the magnitude of competitive complementarity determines the growth response of target beech trees in mixtures. Our investigation strongly suggest that the sensitivity of European beech to environmental constrains depends on neighborhood identity. Therefore, the systematic formation of mixed stands tends to be an appropriate silvicultural measure to mitigate the effects of global warming and droughts on growth patterns of Fagus sylvatica.     

Weather or weathering? Growth of Nothofagus dombeyi on volcanic soils differing in nitrogen and phosphorus concentrations

Aims We tested whether—in addition to weather conditions—the concentrations of nitrogen and phosphorus in the substrate have an effect on the radial stem increment of Nothofagus dombeyi trees in old-growth forest stands on volcanic soil at the western slopes of the Andes in South-Central Chile.Methods We took soil samples and tree increment cores from five proximate sites (1000–1300 m a.s.l.) that are located in the volcanic region of the Conguillío National Park and differ in the age of the substrate (Miocene—3500 years B.P.) and in its concentrations of nitrogen (N) and phosphorus (P). The soil samples were also analysed on their concentrations of other plant mineral nutrients, carbon (C) and nitrogen isotope ratios (δ 15 N). Tree-ring widths and the stem basal area increment (BAI) were related to climate parameters. In selected tree rings, the stable isotope ratios of carbon (δ 13 C) and oxygen (δ 18 O) were determined and related to growth and climate parameters.Important findings Consistent with theory, the soils on the oldest substrate showed the highest (least negative) δ 15 N values, but mineral N was the only nutrient whose concentration exhibited a straightforward (increasing) relationship with increasing substrate age. The BAI was largest on the soil with the highest concentration of plant-available P. In contrast to BAI, tree-ring chronologies did not differ among the study sites. However, tree-ring chronologies and BAI exhibited significantly positive correlations with summer precipitation, and negative correlations, with summer (December) temperature. A negative correlation was found between δ 13 C and precipitation anomalies in the growing season (November–March). We interpret the negative correlations between growth and temperature, and between δ 13 C and δ 18 O in the tree rings, as an impairment of net carbon assimilation by anomalously warm weather conditions during the growing season. We conclude that the growth of N. dombeyi is mainly affected by high temperature and low precipitation in spring and summer irrespective of the substrate's age, and enhanced by higher concentrations of plant-available P in the soil. Our results may be representative of N. dombeyi stands on volcanic substrate within their principal distribution range along the Andes of South America.     

European beech in its northeasternmost stands in Europe: Varying climate-growth relationships among generations and diameter classes

Age, genetics and social status of trees affect their sensitivity to environmental factors, and information about such effects is needed for comprehensive assessment of growth potential. Climatic sensitivity of radial increment (i.e., tree-ring width) of introduced European beech (Fagus sylvatica L.) of different generations and social status, growing in its northeasternmost stands in Europe, was studied by dendroclimatological methods. At present, the studied stands occur outside of the natural distribution area of the species, providing opportunity to study adaptability and potential growth of beech in novel environments under changing climate. The sensitivity of radial growth to climatic factors was modulated by the generation and social status (size) of trees. The first generation trees, which were propagated from the material transferred from the northern Germany, were highly sensitive to climatic factors and showed wide spectrum of responses. The dominant trees were particularly sensitive to June precipitation, indicating sensitivity to water deficit in summer. The suppressed trees were mainly sensitive to temperature in the dormant period. Tree-ring width of the second generation trees, which were propagated from the first generation stands, was mainly affected by water deficit in summer, yet the local factors, modulated the mechanisms of response. In one stand, tree-ring width was affected by conditions during the formation of tree-ring, indicating direct influence of weather conditions on xylogenesis. In the other stand, tree-ring width was correlated to weather conditions in the preceding year, suggesting influence via carbohydrate reserves. The effect of social status on climatic sensitivity in the second generation stands was considerably weaker, likely due to the natural and anthropogenic selection of the material best adapted for local conditions. The effect of climatic factors on radial growth of beech has shifted during the 20th century. The effect of autumn temperature has weakened, likely due to warming; the effect of factors related to water deficit in summer has intensified that could be related to both, changes in climate and ageing. The observed climate-growth relationships suggested that conditions in winter have become suitable for beech, yet careful selection of sites/regions with appropriate hydrological conditions appear necessary to counteract the increasing effect of water deficit, hence to ensure productivity of future beech sites in Latvia.     

Growth-climate responses indicate shifts in the competitive ability of European beech and Norway spruce under recent climate warming in East-Central Europe

Long-term changes in climate substantially affect the tree growth and species distribution in Europe. In the presented study, the radial growth of Fagus sylvatica (L.) and Picea abies ((L.) Karst.) has been studied along an altitudinal gradient covering six vegetation formations characteristic for sub-montane, montane and high-montane conditions of the western Carpathians. Tree growth responses to temperature and precipitation changes have been analysed based on the sample of increment cores and standard dendroclimatic methods in two time periods, the reference period 1961–1990 and the recent period 1991–2012. The growth responses of spruce and beech to recent changes in climate were similar up to high-montane zones, where the beech shows significantly larger improvements of radial increments in comparison to spruce. The growth responses were mainly temperature driven. In the sub-montane area, the increased effect of precipitation in the recent period was overridden by the negative effects of warming, and the alleviated temperature limitation had an evidently supportive effect on tree growth in montane and high-montane areas. In the near future, the warming will likely cause decline in radial increments of beech and spruce in sub-montane areas due to expected landscape drying. At the same time, the improved competitive ability of beech in the high-montane zones suggests a shift in the leading edge of beech distribution into higher altitudes in East-Central Europe.     

How climate,migration ability and habitat fragmentation affect the projected future distribution of European beech

Recent efforts to incorporate migration processes into species distribution models (SDMs) are allowing assessments of whether species are likely to be able to track their future climate optimum and the possible causes of failing to do so. Here, we projected the range shift of European beech over the 21st century using a process‐based SDM coupled to a phenomenological migration model accounting for population dynamics, according to two climate change scenarios and one land use change scenario. Our model predicts that the climatically suitable habitat for European beech will shift north‐eastward and upward mainly because (i) higher temperature and precipitation, at the northern range margins, will increase survival and fruit maturation success, while (ii) lower precipitations and higher winter temperature, at the southern range margins, will increase drought mortality and prevent bud dormancy breaking. Beech colonization rate of newly climatically suitable habitats in 2100 is projected to be very low (1–2% of the newly suitable habitats colonised). Unexpectedly, the projected realized contraction rate was higher than the projected potential contraction rate. As a result, the realized distribution of beech is projected to strongly contract by 2100 (by 36–61%) mainly due to a substantial increase in climate variability after 2050, which generates local extinctions, even at the core of the distribution, the frequency of which prevents beech recolonization during more favourable years. Although European beech will be able to persist in some parts of the trailing edge of its distribution, the combined effects of climate and land use changes, limited migration ability, and a slow life‐history are likely to increase its threat status in the near future.     

Effects of changes in tree species composition on water flow dynamics – Model applications and their limitations

Water-plant relations play a key role in the water cycling in terrestrial ecosystems. Consequently, changes in tree species composition may have distinct effects on the water retention capacity as well as on the pattern of streamflow generation. Such changes may result from modified interception properties and transpiration related to differences in canopy properties and root distribution. In order to evaluate the potential hydrological effects of the current silvicultural conversion from monocultural conifer stands into mixed or pure deciduous stands the hydrological model BROOK90 was applied to two forested upland catchments in Germany. The Rotherdbach catchment (9.4 ha, 93 yr-old Norway spruce) is situated in the Eastern Ore Mountains. The Schluchsee catchment (11 ha, 55-yr-old Norway spruce) is located in the higher altitudes of the Black Forest. The calibrated model is capable to describe rather well the temporal variation of streamflow but also the portions of the individual flow components. Data for a beech scenario were adapted for each site using a standard parameter set for deciduous trees provided by BROOK90. The annual discharge in the fictional beech stand at Rotherdbach is 30 to 50% higher compared to spruce with an increase of soil moisture and especially the slow streamflow components. This mainly results from low interception rates during winter time. In contrast, the spruce stand has a permanently higher interception rate. Effects of tree species conversion are moderate at Schluchsee. The annual discharge of a fictional beech stand at Schluchsee is 7 to 14% higher compared to spruce. There in contrast to Rotherdbach, effects of tree species conversion on soil moisture dynamics are small since vertical percolation in the highly permeable soil dominates and precipitation is abundant. Practical forestry will favorably establish mixed beech–spruce rather than pure beech stands. However, it is critical to simulate mixed stands with BROOK90. Therefore, a simple summation of model results from spruce and beech according to their respective area in a fictional mixed stand can only be a first approximation. Advanced hydrological simulation of mixed stand conditions should regard interactions of tree species and spatial parameter distribution. However, this is not yet feasible due to a distinct lack of information. As a consequence, there is a strong need to collect relevant hydrological and ecophysiological data in mixed stands in the future.     

Survival of Norway spruce remains higher in mixed stands under a dryer and warmer climate

Shifts in tree species distributions caused by climatic change are expected to cause severe losses in the economic value of European forestland. However, this projection disregards potential adaptation options such as tree species conversion, shorter production periods, or establishment of mixed species forests. The effect of tree species mixture has, as yet, not been quantitatively investigated for its potential to mitigate future increases in production risks. For the first time, we use survival time analysis to assess the effects of climate, species mixture and soil condition on survival probabilities for Norway spruce and European beech. Accelerated Failure Time (AFT) models based on an extensive dataset of almost 65 000 trees from the European Forest Damage Survey (FDS) – part of the European‐wide Level I monitoring network – predicted a 24% decrease in survival probability for Norway spruce in pure stands at age 120 when unfavorable changes in climate conditions were assumed. Increasing species admixture greatly reduced the negative effects of unfavorable climate conditions, resulting in a decline in survival probabilities of only 7%. We conclude that future studies of forest management under climate change as well as forest policy measures need to take this, as yet unconsidered, strongly advantageous effect of tree species mixture into account.     

The effects of genetic diversity,climate and defoliation events on trembling aspen growth performance across Canada

Tree genetic makeup may provide an important control of growth dynamics; however, no studies have previously attempted to evaluate its effects in natural trembling aspen stands. In this study, we examined the relative contribution of genetics (i.e. clonal diversity, observed heterozygosity) and environmental conditions (i.e. insects, climate) on aspen growth as represented by mean inter-tree correlation (RBAR), tree basal area increment (TBAI) and inter-annual growth variability (MS). We sampled 440 trees in 22 even-aged natural stands dominated by aspen along an east-west continental gradient of decreasing annual precipitation in the Canadian boreal forest. Linear and mixed-effect models tested the relationships between tree growth, genetics and environmental factors. We showed that clonal diversity and number of years with forest tent caterpillar (FTC) defoliation (N_FTC) reduced and increased the level of growth synchronicity (RBAR), respectively. Clonal diversity explained 30 % of variation in RBAR among sites. TBAI was positively influenced by high moisture conditions while N_FTC and climate explained the variation in MS among trees for each site. No genetic effect could explain either TBAI or the MS variation. Climate and N_FTC drive annual growth variability in trembling aspen at stand and subcontinental scales. Tree genetic makeup contributed to these dynamics, the annual growth dynamics of multi-clonal stands being less homogeneous than those of monoclonal stands. Maintaining diverse aspen stands may ensure a wider range of growth responses to environmental variability, which in turn may help maintain resilience of aspen stands under future climate.     

Nutrient dynamics along a precipitation gradient in European beech forests

Precipitation as a key determinant of forest productivity influences forest ecosystems also indirectly through alteration of the nutrient status of the soil, but this interaction is not well understood. Along a steep precipitation gradient, we studied the consequences of reduced precipitation for the soil and biomass nutrient pools and dynamics in 14 mature European beech (Fagus sylvatica L.) forests on Triassic sandstone. We tested the hypotheses that lowered summer precipitation (1) is associated with less acid soils and (2) a reduced accumulation of organic matter on the forest floor, and (3) reduces nutrient supply from the soil and leads to decreasing foliar and root nutrient concentrations. Soil acidity, the amount of forest floor organic matter, and the associated organic matter N and P pools decreased to about a half from wet to dry sites; the C/P and N/P ratios, but not the C/N ratio, of forest floor organic matter were reduced as well. Net N mineralization and P and K pools in the mineral soil did not change with decreasing precipitation. Foliar P and K concentrations (beech sun leaves) increased while N remained constant, resulting in decreasing foliar N/P and N/K ratios. Estimated N resorption efficiency increased toward the dry sites. We conclude that a reduction in summer rainfall significantly reduces the soil C, N and P pools but does not result in decreasing foliar N and P contents in beech. However, the decreasing foliar N/P ratios towards the dry stands indicate that the importance of P limitation for tree growth declines with decreasing precipitation.     

Warmer winters result in reshaping of the European beech forest soil microbiome (bacteria,archaea and fungi)—With potential implications for ecosystem functioning

In temperate regions, climate warming alters temperature and precipitation regimes. During winter, a decline in insulating snow cover changes the soil environment, where especially frost exposure can have severe implications for soil microorganisms and subsequently for soil nutrient dynamics. Here, we investigated winter climate change responses in European beech forests soil microbiome. Nine study sites with each three treatments (snow exclusion, insolation, and ambient) were investigated. Long-term adaptation to average climate was explored by comparing across sites. Triplicated treatment plots were used to evaluate short-term (one single winter) responses. Community profiles of bacteria, archaea and fungi were created using amplicon sequencing. Correlations between the microbiome, vegetation and soil physicochemical properties were found. We identify core members of the forest-microbiome and link them to key processes, for example, mycorrhizal symbiont and specialized beech wood degraders (fungi) and nitrogen cycling (bacteria, archaea). For bacteria, the shift of the microbiome composition due to short-term soil temperature manipulations in winter was similar to the community differences observed between long-term relatively cold to warm conditions. The results suggest a strong link between the changes in the microbiomes and changes in environmental processes, for example, nitrogen dynamics, driven by variations in winter climate.     

Long-Term Trends in Tree-Ring Width and Isotope Signatures (δ13C, δ15N) of Fagus sylvatica L. on Soils with Contrasting Water Supply

We investigated long-term responses (since 1850) of Fagus sylvatica (Luxembourg; central Europe) to shifts in temperature, precipitation, and nitrogen deposition by analyzing diameter at breast height (DBH) increment, basal area increment (BAI), and tree-ring stable isotopes (δ¹³C, δ¹⁵N). We compared stands on soils with contrasting water supply (Regosols and Cambisols with an available water capacity of ca. 40 and 170 mm, respectively) and of two different age classes (ca. 60 vs. 200 years). All stands showed a peak in DBH increment in the decade 1978–1987, but a decline in increment growth in subsequent decades. In addition, BAI declined in mature stands in the last two decades. Decreasing increment rates were attributable to an increasing drought limitation of stands, mainly induced by increasing temperatures in the last two decades. Contrary to our expectations, stands on Cambisols showed a similar susceptibility to shifts in temperature and precipitation as stands on Regosols, suggesting a strong adaptation of the respective ecotypes grown at dryer sites. This result was in line with long-term trends for tree-ring δ¹³C signatures, which did not differ significantly between stands on Cambisols and Regosols. Climate impacts on tree-ring δ¹⁵N signatures were low. High spring precipitation and temperatures caused increasing and decreasing δ¹⁵N values, respectively, but only in mature stands on Cambisols. Stands on Regosols tended to have lower δ¹⁵N values than stands on Cambisols. Decreasing δ¹⁵N values in recent decades suggest an increasing impact of allochthonous N loads with isotopically lighter N.     

Potential risks for European beech (Fagus sylvatica L.) in a changing climate

Over large areas of Europe, coniferous monocultures are being transformed into mixed forests by the re-introduction of broadleaf tree species belonging to the potential natural vegetation. One important species of interest in this changing forest policy is European beech (Fagus sylvatica). However, at present, this forest management directive has ignored potential adverse effects of global climate change on wide-spread re-introduction of beech to these areas. Average global surface temperatures have risen by approx. 0.8°C in the period between 1861 and 2005 and are expected to continue to increase until the end of this century by 1.5–5.8°C above the 1990 value. To estimate the climate change in the southern part of central Europe in future, we reviewed calculations from regional climate models. Temperature increase for the southern part of central Europe is projected to be up to 2°C within the next 40 years. In contrast, the annual precipitation will most likely remain constant over the same time period, but will experience significant changes in seasonal patterns. Rising intensities of individual precipitation events may result in increasing number and intensities of flooding events and reduced precipitation during the growing season in a higher frequency of summer droughts. Growth and competitive ability of European beech will not, necessarily, respond to increasing CO₂ concentrations but may be strongly impacted by intensive drought that occurs during the growing season. Seedlings as well as adult trees may suffer from xylem embolism, restricted nutrient uptake capacity and reduced growth under limited water availability. However, it remains uncertain to what extent other environmental factors (e.g. soil properties, competitive interactions) may modify the drought response of beech, thus either enhancing susceptibility or increasing drought tolerance and resilience potential. Water-logged soils, predicted during the spring for several regions due to higher than average precipitation, could negatively impact nutrient uptake and growth of beech. Whereas other dominant species as, e.g. oak are well adapted to that environmental stress, beech is known to be sensitive to water-logging and flooding. Thus, the competitive capacity of beech might—depending on the other environmental conditions—be reduced under the expected future climate conditions. Silvicultural practices must be aware today of the potential risks which a changing climate may impose on sustainable forest development.     

Is the spatial distribution of European beech (Fagus sylvatica L.) limited by its potential height growth?

Aim To improve our understanding of species range limits by studying how height growth, a trait related to plant survival, varies throughout the geographic range of Fagus sylvatica L. in France. Location The geographic range of beech in France, representing the western area of its European distribution, within which this species exhibits range distribution limits in both plains and mountainous areas. Methods A generalized linear regression model was used to link beech growth performance to environmental variables using data from 819 plots of the French National Forest Inventory (IFN) database. This model was applied to predict potential growth on 97,281 IFN plots covering the geographic range of beech in France. A kriging technique was used to interpolate estimated growth potential. Finally, the performance of plot‐based predictions of potential growth from the map (i.e. map quality) was evaluated against an independent data set. Results The beech growth performance model highlighted the major impact of climate on potential tree growth at a broad spatial scale. The relevant climatic factors were related mainly to spring cold, summer heat, and winter temperatures and rainfall. The study also revealed the predictive power of soil parameters, which explained a large proportion of the variation in potential beech growth (c. 30%). Analyses of height growth patterns near the boundary of the species range in France showed that the limit only partly coincides with the growth decline caused by climatic and soil factors. Along parts of the range limit, the predicted potential for growth was high, suggesting that in these areas the limit of the range could be explained by other factors, such as competition or constraints on reproduction. Main conclusions The spatial variation in the potential height growth of Fagus sylvatica can be explained by environmental factors and is partly correlated with its regional range limits. By identifying areas where growth potential constrains the geographic range of species, environmental growth models can help to improve our knowledge of the spatial drivers of species geographic range limits and shed light on their response to future environmental changes.     

Patterns of post-glacial spread and the extent of glacial refugia of European beech (Fagus sylvatica)

Aim  Establishing possible relationships between the magnitudes of the glacial distribution of the European beech, Fagus sylvatica L., and its post-glacial spread.
Location  Europe.
Methods  A database of over 400 pollen records has been used to locate Fagus populations at the end of the last glacial and during the post-glacial in Europe and to assess the areal extent of their past distribution.
Results  The rate of late-glacial and post-glacial increase in the number of pollen sites where Fagus was locally present conforms well to a logistic model of population growth. This suggests that the area occupied by beech populations expanded exponentially from the glacial refugia for a duration of over 10,000 years, until about 3500 yr bp . In the past three millennia beech populations increased at a slower rate, tending towards an equilibrium value.
Main conclusions  The conformity of the increase in beech distribution to the classical logistic model of population growth indicates that: (1) a multiplicative biological process was the main factor shaping the pattern of the post-glacial expansion of F. sylvatica in Europe, (2) climate conditions, human activity and competition may have influenced its rate of spread, and (3) beech populations did not expand with a moving closed front, but with a diffuse spread from scattered nuclei. The distribution of Fagus in Europe at the end of the last glacial appears to have been of two orders of magnitude less extensive than at present. Pleistocene refugia were likely to have been a mosaic of sparse stands of small populations scattered in multiple regions. Fagus populations appear to have increased very slowly and to a moderate extent in southern Europe, where they are now declining slightly. The central European populations increased quickly and extensively, reaching northern Europe, and are now approaching their carrying capacity.     

Increasing climate sensitivity of beech and pine is not mediated by adaptation and soil characteristics along a precipitation gradient in northeastern Germany

Rising temperature and altered precipitation regimes will lead to severe droughts and concomitant extreme events in the future. Forest ecosystems have shown to be especially prone to climate change. In assessing climate change impacts, many studies focus on high altitude or ecological edge populations where a climate signal is supposedly most pronounced. While these studies represent only a fraction of the forest ecosystems throughout Europe, findings on climate sensitivity of lowland core populations remain comparatively underrepresented.By using tree-ring widths of a large region-wide network of European beech and Scots pine populations along a precipitation gradient in northeastern Germany, we identify main climatic drivers and spatio-temporal patterns in climate sensitivity. Further, we analyze the resistance of tree growth towards drought. Detailed data on soil characteristics was used to interpret climate-growth relationships.Beech was found to be most sensitive to summer drought during early summer at dry sites, whereas pine displayed highest sensitivity for winter temperature at wet sites. The resistance to extreme drought was lower for beech. By splitting the observation period (1964–2017) into an early and late period, we found non-stationary climate-growth relationships for both study species with beech showing an increase in drought sensitivity and pine in winter temperature sensitivity.Overall, beech populations seem to be especially endangered by prospective climate changes, whereas climate-growth relationships of pine seem more ambiguous with a possible trade-off between enhanced photosynthetic activity caused by early photosynthesis in late winter and reduced activity due to summer drought.     

Growth responses of Pinus sylvestris L. to climate from the southeastern limit of its natural distribution area,Turkey

Pinus sylvestris, the most widely distributed pine species, is commonly used in dendrochronological studies. Based on a lack of studies at its southeastern distribution, we analysed the growth responses of P. sylvestris to temperature and precipitation. We selected 13 sites to study the effects of climate on the growth of Scots pine stands throughout a geographic gradient over time. Trees were sampled from pure stands at different elevations and landscape conditions. The linear and non-linear associations between tree-ring widths and climate variables were calculated with locally specific linear correlation analysis and a mixed generalised additive model. Moving window correlation function was also performed to understand the temporal stability of limiting factors on growth from 1930 to 2013. Our findings showed that early spring temperature (March-April) and late spring-early summer precipitation (May-June) are the major drivers of growth at all sites, where high temperature constraints and high precipitation enhances the growth. Moving window correlation analysis highlighted that the response to precipitation was stationary while temperature changed over time. Our non-linear analysis provided a threshold for March-April temperature. The threshold indicates that the relative additional increment sharply increases up to 7 °C and then slightly decreases.     

Growth patterns in relation to drought-induced mortality at two Scots pine (Pinus sylvestris L.) sites in NE Iberian Peninsula

Drought-related tree mortality has become a widespread phenomenon. Scots pine (Pinus sylvestris L.) is a boreal species with high ecological amplitude that reaches its southwestern limit in the Iberian Peninsula. Thus, Iberian Scots pine populations are particularly good models to study the effects of the increase in aridity predicted by climate change models. A total of 78 living and 39 dead Scots pines trees were sampled at two sites located in the NE of the Iberian Peninsula, where recent mortality events have been recorded. Annual tree rings were used to (1) date dead trees; (2) investigate if there was an association between the occurrence of tree death and severe drought periods characterized by exceptionally low ratios of summer precipitation to potential evapotranspiration (P/PET); and (3) to compare the growth patterns of trees that died with those of surviving ones. Mixed models were used to describe the relationships between tree growth (in terms of basal area increment, BAI, and the percentage of latewood, LW%) and climate variables. Our results showed a direct association between Scots pine mortality and severe drought periods characterized by low summer water availability. At the two sites, the growth patterns of dead trees were clearly distinguishable from those of the trees that survived. In particular, the BAI of dead trees was more sensitive to climate dryness (low P/PET_summer, high temperatures) and started to decline below the values of surviving neighbors 15–40 years before the time of death, implying a slow process of growth decline preceding mortality.     

Quantitative losses vs. qualitative stability of ectomycorrhizal community responses to 3 years of experimental summer drought in a beech–spruce forest

Forest ecosystems in central Europe are predicted to face an increasing frequency and severity of summer droughts because of global climate change. European beech and Norway spruce often coexist in these forests with mostly positive effects on their growth. However, their different below‐ground responses to drought may lead to differences in ectomycorrhizal (ECM) fungal community composition and functions which we examined at the individual root and ecosystem levels. We installed retractable roofs over plots in Kranzberg Forest (11°39′42″E, 48°25′12″N; 490 m a.s.l.) to impose repeated summer drought conditions and assigned zones within each plot where trees neighboured the same or different species to study mixed species effects. We found that ECM fungal community composition changed and the numbers of vital mycorrhizae decreased for both tree species over 3 drought years (2014–2016), with the ECM fungal community diversity of beech exhibiting a faster and of spruce a stronger decline. Mixed stands had a positive effect on the ECM fungal community diversity of both tree species after the third drought year. Ectomycorrhizae with long rhizomorphs increased in both species under drought, indicating long‐distance water transport. However, there was a progressive decline in the number of vital fine roots during the experiment, resulting in a strong reduction in enzyme activity per unit volume of soil. Hydrolytic enzyme activities of the surviving ectomycorrhizae were stable or stimulated upon drought, but there was a large decline in ECM fungal species with laccase activity, indicating a decreased potential to exploit nutrients bound to phenolic compounds. Thus, the ectomycorrhizae responded to repeated drought by maintaining or increasing their functionality at the individual root level, but were unable to compensate for quantitative losses at the ecosystem level. These findings demonstrate a strong below‐ground impact of recurrent drought events in forests.     

Plant-soil feedback in spruce (Picea abies) and mixed spruce-beech (Fagus sylvatica) stands as indicated by dendrochemistry

Ten pairs of secondary pure spruce (Picea abies) and adjacent mixed spruce-beech (Fagus sylvatica) stands on comparable sites were selected on two different bedrocks for soil formation (Flysch: nutrient rich and high soil pH; Molasse: poor nutrient supply and acidic) to study how an admixture of beech to spruce stands affects nutrient cycling and consequently soil chemistry. Soil analyses indicated accumulation of Ca under the mixed stands while the top soil under pure spruce was acidified. It was hypothesized that changes of soil chemical properties due to species composition over the last six decades are reflected in the stem wood of spruce. Three healthy dominant spruce trees per plot were selected for coring. Cores were crossdated and half-decadal samples were analyzed for Ca, Mg, Mn and Al. Calcium and Mg concentrations in stem wood of spruce were significantly higher for the pure spruce than for the mixed stands in spite of lower Ca and Mg stores in the soil. We assume that acidification caused by pure spruce mobilized these cations temporarily, increasing soil solution contents and consequently stem wood concentrations. It was possible to reconstruct soil pH from the element ratios Ca/Al (pure stands) and Ca/Mg (mixed stands), since these ratios in the stem wood of the last half-decade did correlate with soil pH for selected soil depths. Reconstructed soil pH showed a decline over the last 60 years under both species compositions due to accumulation of base cations in the increasing biomass. Comparisons of reconstructed soil pH in 0–5 and 10–20 cm soil depth indicated more pronounced top soil acidification (lower soil pH in 0–5 cm) by spruce on the nutrient rich soil (Flysch) than on the acidic soil (Molasse). However, admixture of beech caused higher pH values in 0–5 cm than in 10–20 cm soil depth on Flysch due to the observed Ca-pump effect of beech (uptake of Ca from deeper soil horizons).