Higher susceptibility of beech to drought in comparison to oak

The expected increase in drought severity and frequency as a result of anthropogenic climate change leads to concerns about the ability of native tree species to cope with these changes. To determine the susceptibility of Fagus sylvatica (European beech) and Quercus robur (pedunculate oak) – the two dominant deciduous tree species in Central Europe – to drought, we quantified the climate sensitivity and drought-response of radial growth for both species using an array of dendroecological methods. Tree-ring data were collected from a site east of Coburg, Bavaria which had shown pronounced stress-symptoms (early leaf coloration) during the record drought of 2018. Climate-growth relationships were used to establish the sensitivity of radial growth to multiple climatic variables. The impact of specific drought events on tree growth was quantified using tolerance indices. In addition, we employed a Principal Component Gradient Analysis (PCGA) and remote sensing data (MODIS Normalized Difference Vegetation Index (NDVI)) to delineate the species specific drought responses. Using these methods we were able to show a clear difference in drought susceptibility between beech and oak. Beech displayed a higher sensitivity to temperature and the standardized precipitation evapotranspiration index (SPEI) and showed lower resistance and resilience to drought events than oak. In particular, beech was unable to fully recover from the 2003 drought, after which it expressed a stark growth decline, i.e. drought legacies, which was not observed for oak. The PCGA revealed a clear differentiation in the grouping of drought responses between beech and oak, supporting the findings of the climate-growth analysis and the tolerance indices. Correlations of NDVI and ring-width indices (RWI) indicated that under normal climatic conditions NDVI variability is linked to the start of the growing season. This is in contrast to drought years, such as 2003, where summer NDVI mirrored the drought response of beech and oak. These results reveal beech to have both a higher sensitivity to summer temperature and SPEI and a higher susceptibility to drought events. Although, in the past high plasticity and adaptability to drought have been attributed to both beech and oak, our study assigns beech a higher risk than oak to suffer from anticipated increases in drought frequency and intensity as a consequence of climate change.     

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.     

Resistance of European tree species to drought stress in mixed versus pure forests: evidence of stress release by inter‐specific facilitation

While previous studies focused on tree growth in pure stands, we reveal that tree resistance and resilience to drought stress can be modified distinctly through species mixing. Our study is based on tree ring measurement on cores from increment boring of 559 trees of Norway spruce (Picea abies [L.] Karst.), European beech (Fagus sylvatica [L.]) and sessile oak (Quercus petraea (Matt.) Liebl.) in South Germany, with half sampled in pure, respectively, mixed stands. Indices for resistance, recovery and resilience were applied for quantifying the tree growth reaction on the episodic drought stress in 1976 and 2003. The following general reaction patterns were found. (i) In pure stands, spruce has the lowest resistance, but the quickest recovery; oak and beech were more resistant, but recover was much slower and they are less resilient. (ii) In mixture, spruce and oak perform as in pure stands, but beech was significantly more resistant and resilient than in monoculture. (iii) Especially when mixed with oak, beech is facilitated. We hypothesise that the revealed water stress release of beech emerges in mixture because of the asynchronous stress reaction pattern of beech and oak and a facilitation of beech by hydraulic lift of water by oak. This facilitation of beech in mixture with oak means a contribution to the frequently reported overyield of beech in mixed versus pure stands. We discuss the far‐reaching implications that these differences in stress response under intra‐ and inter‐specific environments have for forest ecosystem dynamics and management under climate change.     

Climate response of cork growth in the Mediterranean oak (Quercus suber L.) woodlands of southwestern Portugal

In the Mediterranean climate regions, drought events are expected to affect the growth of forests ecosystems by changing trees growth rates and eventually inducing shifts in their growth patterns. Cork oak (Quercus suber L.) is a strictly western Mediterranean tree species periodically harvested for its bark, the cork. So far, cork oak has received limited attention for dendroclimatological studies due to its typical faint and erratic tree wood rings. Moreover, its distinct cork rings chronologies have been completely neglected. In this study we introduce an approach using cork ring chronologies dated back 9–10 years for climate response. Despite enhancing interannual variability and increasing statistical response to short-term climatic variability, still poorly understood, this study will possibly allow infer long-term climate response. We analyzed the cork ring chronologies of 55 cork samples collected in mature (under exploitation) trees in three distinct locations in southwestern Portugal. Cork growth recorded a high climate signal, with highly significant and coherent responses to the yearly climate-related sources of variation. We successfully assessed trends of cork growth via correlation analysis including selected climate variables among mean monthly temperature, monthly precipitation and, on an annual basis, eight precipitation indices. The high mean sensitivities and inter-series correlations found for cork ring chronologies combined with the significant variance explained by climate variables suggest that climate is likely one dominant signal that affects cork growth, but local environmental stresses can decisively affect this (climate) signal. Assuming cork growth as a proxy for cork oak growth, it seems conceivable that despite the trees being highly resistant to drought stress, cork oak woodlands in southwestern Portugal would have to face lesser growth in a global warming scenario.     

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.     

Shifts in the height‐related competitiveness of tree species following recent climate warming and implications for tree community composition: the case of common beech and sessile oak as predominant broadleaved species in Europe

Height growth is a trait that contributes to tree species fitness. How height growth responds to environmental changes may therefore provide indications on species ability to compete and maintain, and on changes in tree community composition. Common beech Fagus sylvatica and sessile oak Quercus petraea are the predominant late‐successional broadleaved species in Europe, and they differ in their shade‐tolerance. On common beech (a shade tolerant species), recent observations across Europe have shown a growth decline during recent climate warming. Because sessile oak is a warmth‐ and light‐demanding species, we therefore hypothesised that it may gain in competitiveness relative to common beech. We conducted analyses of historical height growth in several regions spanning the distributional range of the two species across a temperate‐continental gradient in France. Common beech and sessile oak were sampled in two and four regions, respectively, and were compared in two neighbouring regions. We documented the climatic and nutritional conditions of regional samples. Height growth of 408 trees of various ages was reconstituted from stem analyses. We estimated 20th‐century regional chronologies of height growth using a statistical modelling approach that filtered out the effects of ontogeny and site fertility. In regions where both species were sampled, modelled height trajectories were compared at different periods over the 20th century. Growth chronologies revealed 1) long‐term growth rate increases of a magnitude of 50–100% over 100 years in both species, more acute in the continental domain, 2) recurrent historical inversions in growth fluctuations between species, 3) a recent divergence, with growth decline in common beech versus a dramatic growth increase in sessile oak, more acute in colder regions. The analysis of height trajectories indicated a recent reduction in common beech competitiveness relative to sessile oak. In the face of future climate warming, we conclude that increased prevalence of beech–oak mixtures may arise.     

Climate sensitivity of radial growth in European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) at different aspects in southwestern Germany

The climate sensitivity of radial growth in European beech (Fagus sylvatica L.) was analyzed within a narrow valley in the Swabian Alb (southwestern Germany). We collected stem disks from three aspects (NE, NW and SW) of trees belonging to different social classes. Common climatic factors limiting growth across the valley were identified using a principal component analysis (PCA). Further, we performed hierarchical cluster analysis (HCA), redundancy analysis (RDA) and bootstrapped correlation analysis to reveal differences in chronologies and climate-growth relationships between aspect and social class. Climatic variables considered in our analyses were monthly and seasonal data on temperature and precipitation, as well as a self-calibrating Palmer drought severity index (sc-PDSI). We identified drought in the period June–August as the most prominent factor limiting growth across the valley. Dominant and co-dominant trees at the NW and SW aspects were found to be particularly drought sensitive, whereas intermediate trees were less susceptible to drought. Underlying causes of established climate–growth relationships are discussed in the context of drought susceptibility, tree-size modulation and tree physiological processes.     

Frost and drought: Effects of extreme weather events on stem carbon dynamics in a Mediterranean beech forest

The effects of short-term extreme events on tree functioning and physiology are still rather elusive. European beech is one of the most sensitive species to late frost and water shortage. We investigated the intra-annual C dynamics in stems under such conditions. Wood formation and stem CO₂ efflux were monitored in a Mediterranean beech forest for 3 years (2015–2017), including a late frost (2016) and a summer drought (2017). The late frost reduced radial growth and, consequently, the amount of carbon fixed in the stem biomass by 80%. Stem carbon dioxide efflux in 2016 was reduced by 25%, which can be attributed to the reduction of effluxes due to growth respiration. Counter to our expectations, we found no effects of the 2017 summer drought on radial growth and stem carbon efflux. The studied extreme weather events had various effects on tree growth. Even though late spring frost had a strong impact on beech radial growth in the current year, trees fully recovered in the following growing season, indicating high resilience of beech to this stressful event.     

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.     

Spatial and altitudinal bioclimatic zones of the Italian peninsula identified from a beech (Fagus sylvatica L.) tree-ring network

A network of 24 beech (Fagus sylvatica L.) tree-ring chronologies has been developed for the Italian peninsula. Principal component and cluster analyses were used to identify geographical and altitudinal patterns of tree growth. Correlations and response functions were then applied to the main modes of tree-ring variability to uncover climatic signals. In a landscape occupied by humans for millennia, this approach provided a detailed quantitative ecological characterization of forest types. Altitude was significantly correlated with dendrochronological parameters. The Alps and northern Apennines could be distinguished from the central-southern Apennines. In central Italy, we recognized three different vegetation belts occupied by beech forests, from low- to high-elevation sites. Summer drought impacted beech growth with different intensity at different elevations, depending on the onset and duration of the growing season. Moreover, low-elevation beech forests showed a distinct late spring climate signal, which was opposite to that of high-elevation sites. The coherent geographical and ecological patterns of tree-ring variability suggest that dendrochronological networks help define bioclimatic zones and forest types.     

Branch growth and leaf numbers of red maple (Acer rubrum L.) and red oak (Quercus rubra L.): response to defoliation

Summary Branch growth and leaf formation from terminal and from lateral buds of red maple (Acer rubrum L.) and red oak (Quercus rubra L.) were measured in response to simulated insect defoliation. A single large branch representative of the crown of each tree was used for enumeration of growth and of bud numbers throughout three successive years of 0, 50, 75, and 100% leaf removal for the entire tree. Leaf number per tree for both species after the last year of defoliation was reduced in direct proportion to the severity of defoliation, in comparison to the predefoliation status of the trees. Bud number per tree for red maple, but not for red oak, was also reduced in proportion to severity of defoliation.Averaged over all defoliation treatments, defoliation reduced branch growth more than leaf production. Furthermore, the reduction in branch growth and leaf production was greater in red oak than in red maple. Three years of successive defoliation reduced the mean lateral plus terminal branch growth by 40% in red oak and by 23% in red maple, while leaf number was reduced 22% in red oak and remained unchanged in red maple. In red maple, 100% defoliation caused greater branch death than the 50 or 75% defoliation treatments, and the amount of death was greater after each successive year of defoliation. In contrast to red maple, undefoliated red oak incurred a substantial amount of branch death throughout the study which was little affected by defoliation treatment.     

Drought alters timing, quantity, and quality of wood formation in Scots pine

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.     

Evidence of hydraulic lift in a young beech and oak mixed forest using 18O soil water labelling

Hydraulic lift (HL) by tree roots in a young, broad-leaved, mixed temperate European forest was investigated during the 2008 growing season by injecting ¹⁸O-enriched soil water at a depth of 75–90 cm under drought conditions experimentally imposed in a rain-exclusion system. Based on sap flow, leaf water potential, 2-D root distribution measurements, soil isotope profiles, and xylem water isotope composition, water acquisition and use by two tree species, beech (Fagus sylvatica) and oak (Quercus petraea) was compared. We showed that, unlike oak, beech experienced a marked decrease in sap flow and predawn leaf water potential with increasing soil drought. This behaviour was logical considering the shallower root system in beech than in oak. Six days after ¹⁸O-labelling, we observed isotopic enrichment in the shallower soil layers. Since the intermediate soil layers did not display any enrichment, our results clearly pointed to hydraulic lift by tree roots. The superficial enrichment that was observed in the vicinity of oak trunks and the increase in the isotopic signature of xylem sap in the oak trees but not in the beech trees confirmed the predominant role of oak in the hydraulic lift at our site. Even though facilitation for water acquisition among species was not observed here, our results suggest a potential positive contribution of species like oak toward maintaining species diversity in mixed forest ecosystems submitted to severe drought events.     

Seasonal variations of gas exchange, photosynthetic pigments, and antioxidants in Turkey oak (Quercus cerris L.) and Hungarian oak (Quercus frainetto Ten.) of different age

Seasonal changes in physiological and biochemical parameters were studied in 35-, 55- and 140-year-old trees of Turkey oak (Quercus cerris L.) and Hungarian oak (Q. frainetto Ten.), growing in natural stands in Eastern Balkan Mountains (Bulgaria). During the seasonal drought period (August), assimilation activity, transpiration rate, stomatal conductance and water potential had a seasonal minimum in all the studied tree ages and species. The foliar concentrations of glutathione, ascorbate, α-tocopherol, as well as photosynthetic pigments in oak leaves were significantly affected by season. With the increasing age of the studied trees, we observed a decrease of the physiological activity and an increase of the antioxidants’ accumulation. Both the species were drought tolerant and anisohydric, where Q. frainetto exhibited higher rates of gas exchange than Q. cerris. Moreover, they differed in the extent of increase in the foliar antioxidants and carotenoids.     

Effect of ecological factors on intra-annual stem girth increment of holm oak

Key message

The intra-annual stem girth increment of Quercus ilex is mainly driven by water availability and secondly by temperature. Tree size and competition modulate the growth response to climate.

Abstract

Holm oak (Quercus ilex ssp. ballota [Desf.] Samp.) is the most widespread species in the Iberian peninsula, being one of the most representative trees in forests and open woodlands. The analysis of stem girth increment of holm oak may provide valuable information about how Mediterranean ecosystems will respond to the forecasted climate changes. However, due to the variability of the Mediterranean climate, the knowledge of intra-annual patterns of growth is needed for a better understanding of the influence of the climatic variables at this scale. To this end, we used band dendrometers to measure monthly stem girth increments of 96 holm oak trees from 2003 to 2010, located in open woodlands and dense Mediterranean forests in southwestern Spain. We assessed the effects of climate, competition, topography, and initial stem diameter on stem girth increment. The major stem increment periods were in spring and autumn whereas increment rates were very low or even negative in winter and summer. Spring was not every year the season with the higher stem increments, but autumn when spring was very dry. Higher precipitation, soil moisture, and relative humidity had significant positive effects on stem increment, whereas higher temperature, reference evapotranspiration, and solar radiation had significant negative effects. Initial tree diameter and competition from nearby trees partly explained significant differences in stem increment of individual trees. Therefore, the forecasted climatic changes, in which decreased rainfall in spring and increased summer drought are expected in the Mediterranean region, may be a significant threat to the Q. ilex ecosystems.     

Drought-driven growth reduction in old beech (Fagus sylvatica L.) forests of the central Apennines, Italy

Productivity of old‐growth beech forests in the Mediterranean Basin was measured by average stem basal area increment (BAI) of dominant trees at two mountain sites in the Italian Apennines. Both forests could be ascribed to the old‐growth stage, but they differed markedly with regard to elevation (1000 vs. 1725 m a.s.l.), soil parent material (volcanic vs. calcareous), mean tree age (less than 200 years vs. 300 years), and stand structure (secondary old‐growth vs. primary old‐growth forest). Drought at the two sites was quantified by the self‐calibrated Palmer Moisture Anomaly Index (Z‐index), and by the self‐calibrating Palmer Drought Severity Index (PDSI) for summer (June through August) and the growing season (May through September). Dendroclimatological analyses revealed a moisture limitation of beech BAI at interannual (water availability measured by Z‐index) and decadal scales (water availability measured by PDSI). Both BAI and water availability increased from 1950 to 1970, and decreased afterwards. Trees were grouped according to their BAI trends in auxological groups (growth‐type chronologies), which confirmed that growth of most trees at both sites declined in recent decades, in agreement with increased drought. Because BAI is not expected to decrease without an external forcing, the patterns we uncovered suggest that long‐term drought stress has reduced the productivity of beech forests in the central Apennines, in agreement with similar trends identified in other Mediterranean mountains, but opposite to growth trends reported for many forests in central Europe.     

Can tree species richness attenuate the effect of drought on organic matter decomposition and stabilization in young plantation forests?

Changes in precipitation due to climate change are likely to influence soil organic matter (SOM) decomposition and stabilization. In forests, increased tree species diversity could modulate the effects of drought on SOM decomposition and stabilization. We addressed this issue by a decomposition study under simulated drought (through precipitation reduction at Zedelgem, Belgium) and natural drought (ORPHEE, southern France) in young experimental plantations (tree species richness 1 to 5). In Zedelgem, the study focused on tree species richness around oak and beech trees. Two tea bag indices (TBI) – decomposition rate (k) and stabilization factor (S) – were calculated by measuring the decay of green and rooibos tea in soils. Overall, TBI's were higher in Zedelgem than at ORPHEE. In Zedelgem, k increased with tree species richness under drought around oak, indicating that tree species richness modulated the effects of drought on decomposition. Under beech, k decreased with drought while no effect of tree species richness was detected. S increased with drought under both oak and beech, without any effect of tree species richness. In ORPHEE, we did not detect any tree species richness effect on both TBIs. S decreased significantly, while k was marginally reduced under drought. The higher S under drought in Zedelgem and under control in OPRHEE suggests that the carbon sequestration potential under climate change would be dependent on the environmental context. Further, in young plantations, high species richness may modulate the drought effect on SOM decomposition, but not on stabilization.     

Spread of walnut (Juglans regia L.) in an Alpine valley is correlated with climate warming

Since the Roman time, the walnut tree (Juglans regia L.) has been cultivated in gardens and orchards in large parts of Europe, but only since Karl The Great (742–814) the walnut tree has been introduced to Tyrol (Austria) and other Alpine regions in Central Europe. As late as the beginning of the 20th century no wild populations but only cultivated trees are reported for inner Alpine valleys of Tyrol. Recently, however a striking increase of growing sites and individuals of J. regia has been detected within the Inn valley and other inner Alpine valleys of the Eastern Alps. In addition, climate warming, specifically milder winters since 1970 have been recorded within the study area (Innsbruck, Tyrol, Austria). Therefore, a dendroecological study at four study sites in surroundings of Innsbruck dominated by Scots pine (Pinus sylvestris L.) and/or Norway spruce (Picea abies (L.) Karst.) forests was initiated to examine more closely, if influence of climate warming favoured spread of this deciduous tree.Together, the four study sites encompassed 30 ha, where in total 651 J. regia trees were analysed. Mapping of trees indicated that >90% grew on south and south-west-facing slopes. Evaluation of the age structure revealed that >48% of the analysed J. regia trees were younger than 10 years, >72% younger than 20 years and >92% younger than 30 years.Analysis of climate conditions in years of germination revealed a striking correspondence between the number of germinated trees and winter (December–February) minimum temperatures (r=0.43; p<0.01) and the number of frost days (December–February) <−5 °C (r=0.40; p<0.01), which can be related to frost sensitivity during seedling stage. In summary, results of this study provide evidence that climate warming after 1970 favoured establishment of J. regia within this inner Alpine valley.     

Silver fir and Douglas fir are more tolerant to extreme droughts than Norway spruce in south‐western Germany

Improving our understanding of the potential of forest adaptation is an urgent task in the light of predicted climate change. Long‐term alternatives for susceptible yet economically important tree species such as Norway spruce (Picea abies) are required, if the frequency and intensity of summer droughts will continue to increase. Although Silver fir (Abies alba) and Douglas fir (Pseudotsuga menziesii) have both been described as drought‐tolerant species, our understanding of their growth responses to drought extremes is still limited. Here, we use a dendroecological approach to assess the resistance, resilience, and recovery of these important central Europe to conifer species the exceptional droughts in 1976 and 2003. A total of 270 trees per species were sampled in 18 managed mixed‐species stands along an altitudinal gradient (400–1200 m a.s.l.) at the western slopes of the southern and central Black Forest in southwest Germany. While radial growth in all species responded similarly to the 1976 drought, Norway spruce was least resistant and resilient to the 2003 summer drought. Silver fir showed the overall highest resistance to drought, similarly to Douglas fir, which exhibited the widest growth rings. Silver fir trees from lower elevations were more drought prone than trees at higher elevations. Douglas fir and Norway spruce, however, revealed lower drought resilience at higher altitudes. Although the 1976 and 2003 drought extremes were quite different, Douglas fir maintained consistently the highest radial growth. Although our study did not examine population‐level responses, it clearly indicates that Silver fir and Douglas fir are generally more resistant and resilient to previous drought extremes and are therefore suitable alternatives to Norway spruce; Silver fir more so at higher altitudes. Cultivating these species instead of Norway spruce will contribute to maintaining a high level of productivity across many Central European mountain forests under future climate change.     

Growth reactions of Pinus sylvestris L. and Quercus pubescens Willd. to drought years at a xeric site in Valais, Switzerland

In Valais, an inner-Alpine dry valley in Switzerland, low-elevation Scots pine (Pinus sylvestris L.) forests are changing. While pine shows high mortality rates, deciduous species, in particular pubescent oak (Quercus pubescens Willd.), are becoming more abundant. We hypothesise that increasing drought and the species-specific drought tolerance are key factors in these processes. In this study, the growth reaction to drought years of pine and oak growing at a xeric site in Valais was analysed using dendrochronological and wood anatomical methods. Congruent with theoretical expectations, the tree-ring widths of both species, the mean lumen area of earlywood vessels in oak and the number of tracheids in a radial row in pine decreased in response to dry conditions. However, both species also showed reactions deviating from those known from mesic sites: In oak, the mean lumen area of latewood vessels increased in drought years. In pine, in the driest year of the period (1976), the mean radial diameter increased in latewood and decreased only slightly in earlywood. These results emphasises that the process of wood formation and cell functionality at xeric sites is not completely understood yet. Both species seem to have difficulties to adapt the size of their water-conducting cells to strongly reduced water availability in drought years. Additionally, the cell number is strongly reduced. Thus it remains unclear if both species can maintain sufficient water transport under increasingly dry conditions.