Beech Mortality Research Articles

The Future of European Beech in Northern Germany—Climate Change Vulnerability and Adaptation Potential

Earth’s forests are increasingly exposed to climate risks through climate change-related drought and heat waves. Here, we review the climate vulnerability of European beech (Fagus sylvatica), the most abundant tree species of Central Europe’s temperate forests, and its adaptation potential to a hotter and drier climate in northern Germany, close to the center of the species’ distribution range. About two-thirds of the studied beech forests show persistent negative growth trends of their dominant trees since the onset of rapid warming in the early 1980s, driven primarily by long-term deterioration of the climatic water balance (CWB) in summer and, locally, long-term decrease in June precipitation. Lower water storage capacity of the soil increases the climate sensitivity of growth. Even though beech populations of drier sub-regions reveal some acclimation to drought (reduced stomatal conductance, turgor loss point reduction, higher C allocation to roots), they are more climate-vulnerable than populations at moister sites, visible in larger growth decreases and greater CWB sensitivity of growth. Even though beech mortality is still lower in the study region than in other parts of Central Europe, our results identify beech forest regions with <350 mm growing-season precipitation as vulnerable to recent climate conditions, which refers to half of the study region. With further climate warming and aridification, most of northern Germany likely will become unfavorable for beech growth within the next tree generation, which suggests choosing in many production forests alternative, more drought- and heat-resistant timber species.

Effects of Norway spruce shelter density and soil scarification method on mortality and height growth of underplanted European beech, European silver fir and Douglas fir

AbstractIn Europe, 6–7 million ha even-aged, pure stands of Norway spruce grown outside its natural range are prone to butt rot, windthrow, and bark beetle infestation. Climate change may well accelerate these problems and there is an urgent need to develop methods for converting Norway spruce plantations into more heterogeneous forest with larger resistance and resilience towards natural disturbances. Based on a field experiment located on former heathland on an outwash plain in western Denmark, we examined the mortality and height growth of European beech (Fagus sylvatica L.), European silver fir (Abies alba Mill.), and Douglas fir (Pseudotsuga menziesii (Mirb.) Franco) planted under a shelterwood of 17–19 m tall Norway spruce (Picea abies (L.) Karst.) at shelter densities initially ranging by basal area from 10 to 36 m2 ha−1 (271–1099 stems ha−1). Underplanting included plots of beech (100 per cent) and silver fir (100 per cent) as well as plots with two-species mixtures of silver fir/beech (66 per cent/33 per cent and 33 per cent/66 per cent) and Douglas fir/beech (66 per cent/33 per cent). The experiment also included two soil scarification methods (patch and strip scarification). The experiment was laid out in an unbalanced factorial split-plot design, replicated in time and space, and totaling 144 plots in four blocks. The experiment was followed for 19 years during which a gradual reduction in shelter density took place although in some plots the shelterwood was removed abruptly by windthrow and bark beetle infestation. Shelter densities larger than 10 m2 ha−1 had only a minor effect on the mortality of beech and silver fir, while shelter densities larger than 20 m2 ha−1 resulted in increased mortality of Douglas fir. Subsequent abrupt reduction in shelter density resulted in increased mortality, especially of silver fir. Height growth decreased with increasing shelter density, more for Douglas fir than for beech, and marginally more for beech than for silver fir. For shelter densities <15 m2 ha−1, height growth of silver fir was unaffected. Beech grew faster when mixed with Douglas fir. The soil scarification method did not influence mortality, but strip scarification resulted in marginally larger height growth than patch scarification. We hypothesize that the shelter effect was related mainly to competition for light, while late frost and browsing were important factors in open shelters (<10 m2 ha−1). Based on these results, recommended initial shelter densities are at or <10 m2 ha−1 for the establishment of Douglas fir, at ~15–20 m2 ha−1 for silver fir, and around 15 m2 ha−1 for beech.

European beech dieback after premature leaf senescence during the 2018 drought in northern Switzerland.

During the particularly severe hot summer drought in 2018, widespread premature leaf senescence was observed in several broadleaved tree species in Central Europe, particularly in European beech (Fagus sylvatica L.). For beech, it is yet unknown whether the drought evoked a decline towards tree mortality or whether trees can recover in the longer term. In this study, we monitored crown dieback, tree mortality and secondary drought damage symptoms in 963 initially live beech trees that exhibited either premature or normal leaf senescence in 2018 in three regions in northern Switzerland from 2018 to 2021. We related the observed damage to multiple climate- and stand-related parameters. Cumulative tree mortality continuously increased up to 7.2% and 1.3% in 2021 for trees with premature and normal leaf senescence in 2018, respectively. Mean crown dieback in surviving trees peaked at 29.2% in 2020 and 8.1% in 2019 for trees with premature and normal leaf senescence, respectively. Thereafter, trees showed first signs of recovery. Crown damage was more pronounced and recovery was slower for trees that showed premature leaf senescence in 2018, for trees growing on drier sites, and for larger trees. The presence of bleeding cankers peaked at 24.6% in 2019 and 10.7% in 2020 for trees with premature and normal leaf senescence, respectively. The presence of bark beetle holes peaked at 22.8% and 14.8% in 2021 for trees with premature and normal leaf senescence, respectively. Both secondary damage symptoms occurred more frequently in trees that had higher proportions of crown dieback and/or showed premature senescence in 2018. Our findings demonstrate context-specific differences in beech mortality and recovery reflecting the importance of regional and local climate and soil conditions. Adapting management to increase forest resilience is gaining importance, given the expected further beech decline on dry sites in northern Switzerland.

Response and mortality of beech, fir, spruce and sycamore to rapid light exposure after large-scale disturbance

Physiological response to rapid light exposure due to canopy disintegration in young beech, fir, Norway spruce and sycamore trees was measured in three consecutive years after the severe ice storm in 2014 and after windthrow in 2017. Nitrogen amount (Ntot), maximum assimilation response to light (Amax) and quantum yield (Φ) were measured in three categories of different light intensities under closed canopy with indirect site factor (ISF) < 15%, at the forest edge (15% <ISF < 25%) and in the open (ISF > 25%). Tree responses with number of seedlings per hectare were compared between damaged and undamaged sites, with young trees gradually adapting to light conditions in the two years following the two disturbance events.Nitrogen levels were in the optimal range for all species studied. Rapid exposure to elevated light reduced efficiency in fir and increased efficiency in beech and especially sycamore. No differences in response were observed in spruce. Assimilation efficiency, where both tree species were equal, shifted towards shade. Recovery was similar for all species studied after both disturbances: better after the windstorm, compared to the ice storm, indicating the severity of the event. Reductions in seedling numbers followed the pattern observed for physiological traits.

Tree mortality of European beech and Norway spruce induced by 2018-2019 hot droughts in central Germany

Anthropogenic climate change pushes forest ecosystems globally beyond their limits. Widespread events of forest die-off have been attributed to direct and indirect impacts of increasingly frequent and intense droughts. Here, we focus on an extensive mortality event in Norway spruce (Picea abies (L.) H. Karst.) and European beech (Fagus sylvatica (L.)) forests in Germany, following the successive 2018 – 2019 hot droughts. To examine whether this die-off indeed attributed to observed trends in drought occurrence and intensity, we sampled 143 beech and 186 spruce trees at three low-elevation sites (Spessart, Hassberge, Fichtelberg) with different edaphic properties in northern Bavaria. We analysed long-term hydroclimatic sensitivity and growth responses to extreme events of five site- and species-specific tree-ring width chronologies, including a reference site for each species. Growth of beech was sensitive to drought in April to June, whereas spruce growth was strongly related to drought during June to August, except at slightly higher elevations at the Fichtelberg site, where a summer temperature signal was observed. Trees at the Spessart and Hassberge sites showed an increased response to hydroclimatic conditions in April following the extreme drought in 1976 and from the 1990s onwards at the Fichtelberg site. Spruce bark beetle (Ips typographus L.) outbreaks during the 2018 drought accelerated the high mortality rates in around 50% of the trees at the Spessart and Hassberge site. In 2018, around 7% of all beech trees died at the Hassberge site, the site with the highest clay content. Our results suggest that these widespread mortality events can be attributed to an increasing drought sensitivity and were accelerated by the consecutive recent drought years. Sustainable forest management practices for these ecologically and economically important tree species are required to mitigate the effects of global warming in the future.

Effect magnitudes of operational-scale partial harvesting on residual tree growth and mortality of ten major tree species in Maine USA

Silvicultural systems based on partial harvesting (PH) have become increasingly common in areas historically dominated by clearcut-based systems in response to ecological and social concerns. Current knowledge about the effects of PH is based primarily on stand responses from controlled experiments from limited number of sites. A broader scope of inference is needed to fully understand the effects of PH when applied operationally at a landscape scale. We used 835 permanent monitoring plots from the US Forest Service Forest Inventory and Analysis network to quantify the effect magnitude (EM) of low- (5–40% basal area removal) and high-intensity (41–80% basal area removal) PH on the periodic diameter growth and mortality of residual trees of ten major tree species in Maine, USA. Our results showed that the EM and timing of statistically significant effect (p < 0.05) of the two intensities of PH varies across species. Tree diameter growth responses to high-intensity PH was rapid and prolonged for sugar maple (Acer saccharum) and American beech (Fagus grandifolia), while other species did not show significant responses for at least five years. High-intensity PH increased mortality of American beech, balsam fir (Abies balsamea), red spruce (Picea rubens), northern white cedar (Thuja occidentalis), and red maple (Acer rubrum), but not for other tree species. No relationship was found between species shade tolerance and species-specific response magnitude to PH in terms of diameter growth and tree- or stand-level (absolute or relative) mortality. This result suggests that species shade tolerance does not always determine the type or magnitude of response that a given species has to increased resource availability following PH and that other functional traits may be more important. Overall, the findings provide strong evidence that subsequent tree responses to PH are not consistent among contrasting species for at least 15 years following harvest and longer-term regional assessments are needed.

Gains or Losses in Forest Productivity under Climate Change? The Uncertainty of CO2 Fertilization and Climate Effects

Global warming poses great challenges for forest managers regarding adaptation strategies and species choices. More frequent drought events and heat spells are expected to reduce growth and increase mortality. Extended growing seasons, warming and elevated CO2 (eCO2) can also positively affect forest productivity. We studied the growth, productivity and mortality of beech (Fagus sylvatica L.) and fir (Abies alba Mill.) in the Black Forest (Germany) under three climate change scenarios (representative concentration pathways (RCP): RCP2.6, RCP4.5, RCP8.5) using the detailed biogeochemical forest growth model GOTILWA+. Averaged over the entire simulation period, both species showed productivity losses in RCP2.6 (16–20%) and in RCP4.5 (6%), but productivity gains in RCP8.5 (11–17%). However, all three scenarios had a tipping point (between 2035–2060) when initial gains in net primary productivity (NPP) (6–29%) eventually turned into losses (1–26%). With eCO2 switched off, the losses in NPP were 26–51% in RCP2.6, 36–45% in RCP4.5 and 33–71% in RCP8.5. Improved water-use efficiency dampened drought effects on NPP between 4 and 5%. Tree mortality increased, but without notably affecting forest productivity. Concluding, cultivation of beech and fir may still be possible in the study region, although severe productivity losses can be expected in the coming decades, which will strongly depend on the dampening CO2 fertilization effect.

Impact of the Invasive Beech Leaf-Mining Weevil, Orchestes fagi, on American Beech in Nova Scotia, Canada

The beech leaf-mining weevil, Orchestes fagi (L.), is native to Europe where it commonly attacks European beech. The weevil was discovered infesting American beech in Halifax and Cape Breton Island, Nova Scotia, Canada in 2012, but anecdotal reports of defoliated beech in the Halifax area as early as 2006 suggest it established 5–10 years prior to its discovery. Our objectives were to estimate the impact of O. fagi on American beech in forested sites and urban areas, as well as its economic impact on owners of residential properties with mature American beech. In 2014, we established fifteen 11.28 m fixed radius plots in forested sites containing a total of 260 American beech at Sandy Lake, Oakfield, and Mt. Uniacke (n=5 plots per site), where weevil infestation levels were high, moderate, and nil, respectively. At the same time we recorded the degree of cankering by beech bark disease on the main stems of each tree. Plots were visited annually to record tree mortality (2014–2019) and percentage of leaves with larval mines or adult feeding (2016–2019). We surveyed residents of Halifax in 2016 and 2018 to determine the rate of beech mortality and costs of tree removal in urban residential areas. Between 2016 and 2019, the percentage of leaves mined by weevil larvae increased from 6% to 59% at Mt. Uniacke, from 48% to 83% at Oakfield, and from 87% to nil at Sandy Lake (because there were too few leaves to sample). During the same period, cumulative beech mortality increased from 35% to 48% at Mt. Uniacke, from 10% to 70% at Oakfield, and from 88% to 94% at Sandy Lake. Tree mortality was not associated with severity of beech bark disease, except at Mt. Uniacke which had the fewest years of defoliation by the leaf-mining weevil. Mortality rates were lower in urban areas (32% in 2016, 44% in 2018) but direct costs to property owners who hired arborists to remove dead beech trees were significant, averaging $1934 ($300–$6600) per resident in 2018. Options for mitigating the impact of O. fagi on American beech are briefly discussed.

Growth and mortality of Norway spruce and European beech in monospecific and mixed-species stands under natural episodic and experimentally extended drought. Results of the KROOF throughfall exclusion experiment

Key messageUnder severe drought, growth of Norway spruce suffered much more than European beech. Norway spruce benefited from growing in the environment of beech, and both species acclimated slightly to 5 years of experimentally extended drought.Recent studies show that the detrimental effects of drought on stand growth are mitigated when the stand contains mixed tree species. We analysed the growth responses of Norway spruce and European beech to episodic and experimentally extended drought in intra- and inter-specific mature stands. We used annual diameter growth records dating back to 1998 to determine the impact of the natural episodic drought in 2003 and 2015. To analyse extended drought, spruce and beech trees were exposed to extreme drought under automatic throughfall exclusion roofs from 2014 to 2018. The growth of spruce in an inter-specific environment with beech was 20–50% less affected by natural episodic drought compared with an intra-specific constellation. When beech grew in an inter-specific environment, it was by 23% more affected by drought compared to intra-specific conditions, but seemed to recover faster. The induced drought from 2014 to 2018 resulted in a strong growth reduction in the first year particularly for spruce, followed by a slight acclimation to the dry conditions. Beech acclimated and recovered faster than spruce across all growing conditions, while spruce only acclimatized faster in the environment of beech. Both species showed a higher mortality under induced drought compared with the controls; for spruce, the mortality rate was fivefold higher compared to the long-term mortality. The long-term moderate-growth stabilization and the growth increase after the 5-year exposure to drought suggest a gradual acclimation to drought by beech. The resistance and acclimation to drought of spruce when growing in mixture should be considered when designing resource efficient and productive mixed conifer-broadleaved stands for future climates.

Similar patterns of background mortality across Europe are mostly driven by drought in European beech and a combination of drought and competition in Scots pine

Background tree mortality is a complex demographic process that affects structure and long-term forest dynamics. Here we investigated how climatic drought intensity interacts with interspecific and intraspecific competition (or facilitation) in shaping mortality patterns across tree species ranges. To this aim, we used data from five European national forest inventories to perform logistic regression models based on individual tree mortality in Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.). We computed the relative importance of climatic drought intensity, basal area of conspecific and heterospecific trees (proxy of indirect intra- and interspecific competition or facilitation) and the effects of their interactions on mortality along the entire European latitudinal gradient of both species range. Increase in climatic drought intensity over the study period was associated with higher mortality rates in both species. Climatic drought intensity was the most important driver of beech mortality at almost all latitudes while Scots pine mortality was mainly driven by basal area. High conspecific basal area was associated with high mortality rates in both species while high heterospecific basal area was correlated with mortality rates that were high in Scots pine but low in beech. Overall, beech mortality was directly affected by climatic drought intensity while Scots pine mortality was indirectly affected by climatic drought intensity through interactions with basal area. Despite their different sensitivity to drought and basal area, the highest predicted mortality rates for both species were at the ecotone between the cool temperate and Mediterranean biomes, which can be explained by the combined effect of climatic drought intensity and competition. In the context of global warming, which is expected to be particularly strong in the Mediterranean biome, our results suggest that populations at the southern limit of species ranges may experience increased mortality rates in the near future.

Drivers of persistent post-fire recruitment in European beech forests

Climate change is expected to alter disturbance regimes including fires in European beech (Fagus sylvatica L.) forests. Regarding the resilience of beech forests to fire it is questionable whether seeds of this non-serotinous obligate masting seeder find advantageous conditions in a post-fire environment. The probability of recruitment success has been shown to increase when fire coincides with a mast year. However, the fire-induced recruitment window is poorly defined, and it is unclear how other interacting factors influence its duration. We used a space-for-time approach to model the relationships between post-fire beech recruitment, timing of seed mast events, and interacting environmental conditions using a zero-inflated model. Our results show that recruitment peaks 5–12 years after a fire, and continues throughout three decades post-fire. Beech recruitment in the post-fire period is driven by mast intensity interacting with (i) canopy opening as a consequence of progressive post-fire tree mortality and (ii) coverages of competing ground vegetation. Spring-summer moisture showed a weak positive effect on beech recruitment. We conclude that fires increase light availability, which in coincidence with a mast event results in pulses of beech recruitment. The delayed post-fire mortality of beech creates a recruitment window lasting for up to three decades, resulting in a higher-than-expected resilience of beech to individual fire disturbances.

Not dead yet: Beech trees can survive nearly three decades in the aftermath phase of a deadly forest disease complex

Abstract Management efforts often focus on preventing the arrival of destructive insects and pathogens or mitigating damage in forests experiencing heavy mortality. This need for management often abates after the mortality event due to reduced causal agent presence. However, the persistence of causal agents in beech bark disease (BBD)-impacted forests typically results in repeated tree mortality; this cycle has cascading impacts on forest biota and timber regeneration. We analyzed remeasurement data on BBD severity and tree death collected from 1988 to 2016 in disease aftermath stands in the Adirondack Mountains of New York to quantify the survival of BBD-affected trees over time. We found that while BBD severity has a strong influence on the yearly probability of tree survival, affected trees can survive for at least 28 years. However, the probability a tree will survive this long declined with more severe initial disease status regardless of how BBD severity changed over time. These findings could help inform management efforts to ameliorate the indirect impacts of BBD on forest wildlife populations, plant community diversity, and the regeneration of desirable timber species over time. Furthermore, our results underscore the continued need to manage BBD-induced beech mortality and disease-associated agents in aftermath stands.

Gap dynamics in the Western Carpathian mixed beech old-growth forests affected by spruce bark beetle outbreak

Understanding how species-specific disturbances affect the dynamics of mixed forests is becoming increasingly important due to rapidly changing disturbance regimes. This study estimated the effect of Norway spruce (Picea abies (L.) Karst.) mortality on the disturbance processes in two mixed beech stands of the Western Carpathians that were affected by a bark beetle outbreak. We evaluated the size distribution, fraction of canopy and expanded gaps, the characteristics of gapmakers and the contribution of different species to gap size. The measured canopy gap fraction was <5%, and most canopy gaps were small (<100 m2). Spruce was the most abundant gapmaker, and its share among gapmakers was 3–6 times higher than its share in the canopy. We found that the increase in spruce mortality due to the outbreak resulted in a fine-scale mortality pattern. However, spruce gapmakers did not contribute much to gap area size, as shown by a weak correlation between the number of spruce gapmakers and the area of expanded gaps. Diameter distribution of living versus recently dead trees showed that beech mortality occurred disproportionately in large size classes. However, dead spruce trees were equally frequent in all diameter classes, which means beetles did not exclusively attack larger trees in these stands during the outbreak. We conclude that spruce mortality may have influenced successional processes by giving a competitive advantage to two other species that were not affected by the outbreak, provided that a high deer browsing intensity does not hinder the regeneration of new seedlings.

Beech Bark Disease ReducesSus scrofa(Boar) Rooting Intensity in Great Smoky Mountains National Park

Abstract This study examines the effect of beech bark disease (BBD) on Sus scrofa (European Wild Boar) rooting in high-elevation Beech gaps of Great Smoky Mountains National Park. In 2011, we sampled vegetative cover by stratum (canopy, shrub, herb) and European Wild Boar rooting extent in pre-existing fenced boar-exclosure plots and corresponding unfenced plots. We also used data from previous studies to compare frequencies of individual herbaceous species collected pre-BBD to those collected post-BBD. Our results indicate that mortality of Fagus grandifolia (American Beech) trees due to BBD and the consequent growth of a dense shrub-layer significantly reduced boar rooting in gaps within the Beech stands. We found that herbs were affected by both European Wild Boar and the dense shrub-cover following American Beech mortality; however, some plant species remained abundant, possibly because they were protected from detection within the shrubs.

Does one model fit all? Patterns of beech mortality in natural forests of three European regions.

Large uncertainties characterize forest development under global climate change. Although recent studies have found widespread increased tree mortality, the patterns and processes associated with tree death remain poorly understood, thus restricting accurate mortality predictions. Yet, projections of future forest dynamics depend critically on robust mortality models, preferably based on empirical data rather than theoretical, not well-constrained assumptions. We developed parsimonious mortality models for individual beech (Fagus sylvatica L.) trees and evaluated their potential for incorporation in dynamic vegetation models (DVMs). We used inventory data from nearly 19,000 trees from unmanaged forests in Switzerland, Germany, and Ukraine, representing the largest dataset used to date for calibrating such models. Tree death was modelled as a function of size and growth, i.e., stem diameter (dbh) and relative basal area increment (relBAI), using generalized logistic regression accounting for unequal re-measurement intervals. To explain the spatial and temporal variability in mortality patterns, we considered a large set of environmental and stand characteristics. Validation with independent datasets was performed to assess model generality. Our results demonstrate strong variability in beech mortality that was independent of environmental or stand characteristics. Mortality patterns in Swiss and German strict forest reserves were dominated by competition processes as indicated by J-shaped mortality over tree size and growth. The Ukrainian primeval beech forest was additionally characterized by windthrow and a U-shaped size-mortality function. Unlike the mortality model based on Ukrainian data, the Swiss and German models achieved good discrimination and acceptable transferability when validated against each other. We thus recommend these two models to be incorporated and examined in DVMs. Their mortality predictions respond to climate change via tree growth, which is sufficient to capture the adverse effects of water availability and competition on the mortality probability of beech under current conditions.

What drives European beech (Fagus sylvatica L.) mortality after forest fires of varying severity?

Predicting the timing and amount of tree mortality after a forest fire is of paramount importance for post-fire management decisions, such as salvage logging or reforestation. Such knowledge is particularly needed in mountainous regions where forest stands often serve as protection against natural hazards (e.g., snow avalanches, rockfalls, landslides). In this paper, we focus on the drivers and timing of mortality in fire-injured beech trees (Fagus sylvatica L.) in mountain regions. We studied beech forests in the southwestern European Alps, which burned between 1970 and 2012. The results show that beech trees, which lack fire-resistance traits, experience increased mortality within the first two decades post-fire with a timing and amount strongly related to the burn severity. Beech mortality is fast and ubiquitous in high severity sites, whereas small- (DBH <12cm) and intermediate-diameter (DBH 12–36cm) trees face a higher risk to die in moderate-severity sites. Large-diameter trees mostly survive, representing a crucial ecological legacy for beech regeneration. Mortality remains low and at a level similar to unburnt beech forests for low burn severity sites. Beech trees diameter, the presence of fungal infestation and elevation are the most significant drivers of mortality. The risk of beech to die increases toward higher elevation and is higher for small-diameter than for large-diameter trees. In case of secondary fungi infestation beech faces generally a higher risk to die. Interestingly, fungi that initiate post-fire tree mortality differ from fungi occurring after mechanical injury. From a management point of view, the insights about the controls of post-fire mortality provided by this study should help in planning post-fire silvicultural measures in montane beech forests.

Climate modifies tree interactions in terms of basal area growth and mortality in monospecific and mixed Fagus sylvatica and Pinus sylvestris forests

Net interactions between trees vary depending on environmental conditions and other factors such as stand density, age, or between-species complementarity and/or facilitation. According to the stress gradient hypothesis, positive or facilitative interactions are more frequent in high-stress environments whereas negative or competitive interactions occur in benign environments, although recent studies highlight the influence of species composition, type of stress, ontogeny, etc. on the interaction–stress gradient pattern. The aim of this paper is to analyze whether site climatic variables are a key factor in tree interactions in mixed stands of beech (Fagus sylvatica L.) and Scots pine (Pinus sylvestris L.). To test how site climatic conditions modify the effect of inter-specific competition on tree basal area growth and tree mortality, growth and mortality models were fitted using monospecific and mixed sample plots located in matching site conditions selected from the Spanish National Forest Inventory in the Navarra Province. Tree competition status was broken down into four terms according to size-symmetry (size-symmetric vs. size-asymmetric) and species identity (intra-specific vs. inter-specific). The results showed that the size-symmetric inter-specific component was non-significant on beech basal area growth and had a negative effect on pine growth. The effect of size-asymmetric inter-specific competition was always non-significant, resulting in a higher basal area growth when the admixed species are of a larger size. The interaction between annual precipitation and this inter-specific competition effect was much more pronounced on beech than on pine. Inter-specific competition had a negative effect on pine growth under better climatic conditions and a positive effect at dryer sites, while beech always benefited from the presence of pine, although the benefit was greater where climatic site conditions were better. In tree mortality models, pine mortality increased with the proportion of beech, while beech mortality was lower as the proportion of pine increased. Precipitation modified the inter-specific competition effect on tree mortality although the site influence was less relevant than on tree growth. For pine mortality, the negative effect of beech admixture was stronger at lower mean annual precipitation, while in the case of beech the positive effect of pine increased at higher levels of precipitation. The influence of climate on the effect of competition, the variation in their strength depending on the mode of competition (size-symmetric or size-asymmetric), along with the inter-specific competition component, highlight the importance of considering the effect of site conditions on between-species interactions when modeling tree growth and mortality. The species-specific patterns of variation in tree interactions along climatic gradients and the differences in tree growth and mortality corroborate the need to consider the nature of stress-limiting factors and species composition and the importance of analyzing both dynamic processes simultaneously.

Indikatoren zur Schätzung des Trockenheitsrisikos in Buchen- und Fichtenwäldern

Indicators for quantitative assessment of drought risks in beech and Norway spruce forests The application of climate models to forecast future forest development asks for quantitative drought-response relationships, with the term “drought” first needing a definition. The long-term data series of an intercantonal forest observation program allowed to test various drought indicators, to compare them and to derive quantitative relationships for beech and Norway spruce. For basal area increment of both tree species indicators of the site water balance (SWB) performed best. For beech mortality site water balance and several indicators basing on the ratio between actual and potential evapo-transpiration (ETa/ETp) were equivalent, whereas for spruce mortality ETa/ETp during the first 80 days of the season was the best indicator. With these indicators the average drought related growth reduction after 2003 – a year with extreme drought – was estimated to amount to 32% for beech and 37% for Norway spruce. Mortality of Norway spruce increased by 130%, whereas the estimates for the increase of beech mortality vary between 54 and 110%, depending on the indicator. The observed quantitative relationships for growth were applied to map drought responses of growth for Switzerland. The maps clearly show the dry regions of Switzerland (northern Switzerland, southern Jura foothills, Lemanic region, Valais and Rhine valley around Chur), where basal area increment of beech and Norway spruce was reduced by more than 40%.

Fine root dynamics in Slovenian beech forests in relation to soil temperature and water availability

Fine root ingrowth and mortality of European beech are related to evapotranspiration, cumulative forest floor precipitation, soil temperature and water content, which are affected by forest management and gap creation. The ingrowth and mortality of European beech (Fagus sylvatica L.) fine roots (diameters <2 mm) were studied in relation to environmental variables describing temperature and water availability at four sites, covering a range in environmental conditions likely to be encountered in Slovenian beech forests. Minirhizotron images were used to determine fine root dynamics in a stand and gap in each of the sites for 12 periods during the 2007–2009 growing seasons. The environmental variables included air and soil temperatures, precipitation, forest floor precipitation, evapotranspiration and soil water contents. For data analysis, the daily mean values for each period for all variables were used. Fine root ingrowth and mortality were higher in the managed stand and gap compared to the old-growth stand and gap, but only significantly correlated with each other in the case of the managed stand. Forest floor precipitation and soil temperature were significant in explaining fine root ingrowth, whereas maximal evapotranspiration, soil temperature and soil water content were more important for fine root mortality. However, the correlations were weak and inconsistent among the four sites. By including site as predictor as well as environmental variables, R 2 values of 0.49 and 0.55 for ingrowth and mortality, respectively, were achieved. Despite this, the relationships between the fine root dynamics and selected environmental factors appeared relatively weak and complex, especially for fine root ingrowth and might be partially related also to differences in successional stages of the forests under study.

Low relative growth rates predict future mortality of common beech (Fagus sylvatica L.)

For many tree species, growth patterns derived from tree-ring time series have been shown to be good indicators of tree mortality. Although tree rings of common beech (Fagus sylvatica) have been widely used to answer complex questions of forest ecology, there are only few studies using growth characteristics, such as growth decline or growth variability, as informative predictors of tree mortality in old-growth beech forests. To do this, we used dendrochronological data of living and dead trees from a nature reserve in eastern Germany. Using a logistic regression model, we predicted tree mortality based on growth characteristics (basal area increment; variance, autocorrelation, mean sensitivity of ring widths; growth trends) over different time horizons. Beech mortality could reliably be predicted up to 20years in advance on the basis of relative basal area growth. Trees that grew slower in relative basal area than 0.95cm2/cmy on average over a 20-year-period had a much higher mortality risk than faster-growing trees. Other statistical characteristics of the dendrochronological time series, such as the variance or autocorrelation of ring widths, the mean sensitivity or growth trends, did not convey significant additional information and did not lead to better mortality predictions.