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Forest models are instrumental for understanding and projecting the impact of climate change on forests. A considerable number of forest models have been developed in the last decades. However, few systematic and comprehensive model comparisons have been performed in Europe that combine an evaluation of modelled carbon and water fluxes and forest structure. We evaluate 13 widely used, state-of-the-art, stand-scale forest models against field measurements of forest structure and eddy-covariance data of carbon and water fluxes over multiple decades across an environmental gradient at nine typical European forest stands. We test the models' performance in three dimensions: accuracy of local predictions (agreement of modelled and observed annual data), realism of environmental responses (agreement of modelled and observed responses of daily gross primary productivity to temperature, radiation and vapour pressure deficit) and general applicability (proportion of European tree species covered). We find that multiple models are available that excel according to our three dimensions of model performance. For the accuracy of local predictions, variables related to forest structure have lower random and systematic errors than annual carbon and water flux variables. Moreover, the multi-model ensemble mean provided overall more realistic daily productivity responses to environmental drivers across all sites than any single individual model. The general applicability of the models is high, as almost all models are currently able to cover Europe's common tree species. We show that forest models complement each other in their response to environmental drivers and that there are several cases in which individual models outperform the model ensemble. Our framework provides a first step to capturing essential differences between forest models that go beyond the most commonly used accuracy of predictions. Overall, this study provides a point of reference for future model work aimed at predicting climate impacts and supporting climate mitigation and adaptation measures in forests., Global Change Biology, 28 (23), ISSN:1354-1013, ISSN:1365-2486

While enhanced tree growth over the last decades has been reported in forests across the globe, it remains unclear whether it drives persistent biomass increases of the stands, particularly in mature forests. Enhanced tree growth and stand-level biomass are often linked with a simultaneous increase in density-driven mortality and a reduction in tree longevity. Identifying empirical evidence regarding the balance between these processes is challenging due to the confounding effects of stand history, management, and environmental changes. Here, we investigate the link between growth and biomass via the shift in the negative relationship between average tree size and stand density (tree number). We find increasing stand density for a given tree size in unmanaged closed-canopy forests in Switzerland over the past six decades and a positive relationship between growth and stand density - qualitatively consistent with simulations by a mechanistic, cohort-resolving ecosystem model (LM3-PPA). Model simulations show that, in the absence of other disturbances, enhanced growth persistently increases biomass stocks despite simultaneous decreases in carbon residence time and tree longevity, independent of assumptions about the drivers of tree mortality. However, the magnitude of simulated changes critically depends on the shape of the mortality parameterizations. Our analyses reconcile reports of growth-induced reductions of tree longevity with model predictions of persistent biomass increases, and with our finding of a trend towards denser forests in response to growth - also in mature stands.

Process-based forest models are important tools for predicting forest growth and their vulnerability to factors such as climate change or responses to management. One of the most widely used stand-level process-based models is the 3-PG model (Physiological Processes Predicting Growth), which is used for applications including estimating wood production, carbon budgets, water balance and susceptibility to climate change. Few 3-PG parameter sets are available for central European species and even fewer are appropriate for mixed-species forests. Here we estimated 3-PG parameters for twelve major central European tree species using 1418 long-term permanent forest monitoring plots from managed forests, 297 from un-managed forest reserves and 784 Swiss National Forest Inventory plots. A literature review of tree physiological characteristics, as well as regression analyses and Bayesian inference, were used to calculate the 3-PG parameters. The Swiss-wide calibration, based on monospecific plots, showed a robust performance in predicting forest stocks such as stem, foliage and root biomass. The plots used to inform the Bayesian calibration resulted in posterior ranges of the calibrated parameters that were, on average, 69% of the prior range. The bias of stem, foliage and root biomass predictions was generally less than 20%, and less than 10% for several species. The parameter sets also provided reliable predictions of biomass and mean tree sizes in mixed-species forests. Given that the information sources used to develop the parameters included a wide range of climatic, edaphic and management conditions and long time spans (from 1930 to present), these species parameters for 3-PG are likely to be appropriate for most central European forests and conditions., European Journal of Forest Research, 5 (1), ISSN:1612-4677, ISSN:1612-4669

Central Europe has been experiencing unprecedented droughts during the last decades, stressing the decrease in tree water availability. However, the assessment of physiological drought stress is challenging, and feedback between soil and vegetation is often omitted because of scarce belowground data. Here we aimed to model Swiss forests' water availability during the 2015 and 2018 droughts by implementing the mechanistic soil-vegetation-atmosphere-transport (SVAT) model LWF-Brook90 taking advantage of regionalized depth-resolved soil information. We calibrated the model against soil matric potential data measured from 2014 to 2018 at 44 sites along a Swiss climatic and edaphic drought gradient. Swiss forest soils' storage capacity of plant-available water ranged from 53 mm to 341 mm, with a median of 137 ± 42 mm down to the mean potential rooting depth of 1.2 m. Topsoil was the primary water source. However, trees switched to deeper soil water sources during drought. This effect was less pronounced for coniferous trees with a shallower rooting system than for deciduous trees, which resulted in a higher reduction of actual transpiration (transpiration deficit) in coniferous trees. Across Switzerland, forest trees reduced the transpiration by 23% (compared to potential transpiration) in 2015 and 2018, maintaining annual actual transpiration comparable to other years. Together with lower evaporative fluxes, the Swiss forests did not amplify the blue water deficit. The 2018 drought, characterized by a higher and more persistent transpiration deficit than in 2015, triggered widespread early wilting across Swiss forests that was better predicted by the SVAT-derived mean soil matric potential in the rooting zone than by climatic predictors. Such feedback-driven quantification of ecosystem water fluxes in the soil–plant-atmosphere continuum will be crucial to predicting physiological drought stress under future climate extremes., Global Change Biology, 28 (20), ISSN:1354-1013, ISSN:1365-2486

Process-based forest models (PBMs) are important tools for quantifying forest growth and vulnerability, particularly under climate change. The 3-PG model (Physiological Processes Predicting Growth) is one of the most widely used forest growth simulators for this purpose worldwide. Here, we presentr3PG, a newFortranimplementation of 3-PG, wrapped into anrpackage.r3PGcan simulate monospecific as well as mixtures of evergreen and deciduous tree species in even-aged or uneven-aged stands. The combination ofFortranfunctions with anrinterface makes the model extremely fast. This facilitates the use ofr3PGfor extensive computer experiments and sensitivity analysis. We demonstrate this in a case study including (a) single model runs; (b) a sensitivity analysis and a full Bayesian calibration of the model and (c) spatial simulations of forest growth across Switzerland. r3PGis faster and easier to use than previous implementations of 3-PG in visual basic. We believe that this will make 3-PG even more useful and popular for ecologists and climate change scientists.

As currently framed, the forest cycle model that underlies close-to-nature management in temperate beech forests throughout the globe specifies an orderly sequence of temporal development within even-aged patches comprising the forest mosaic. Although this model has been widely applied to European beech (Fagus sylvatica L.) forests, the underlying assumptions of disturbance-induced even-agedness (i.e., within-patch age homogeneity) and competition-induced size differentiation (i.e., within-patch size heterogeneity) have not been tested in natural beech forests due to prohibitions on tree coring in primeval forest reserves. In a rare and unprecedented test dataset of spatially explicit tree ages in an old-growth European beech forest, we employed triangulated irregular networks of Delaunay triangles to objectively identify natural tree neighborhoods to determine if neighboring (i.e., within-patch) trees were even- or, at most, two-aged. Age differences among neighboring trees (summarized in 25-yr age classes) were rarely 50 yrs, while the few "even-aged" patches were very small (100 m2) and relatively young (

3-PG: ein physiologisches Waldwachstumsmodell Ertragstafeln waren die ersten Waldwachstumsmodelle, die in Europa entwickelt wurden. Diese empirischen Modelle sind für die Anwendung unter zukünftigen klimatischen Bedingungen, für Mischbestände oder für die Abschätzung der Effekte neuer waldbaulicher Systeme ungeeignet. In den letzten Jahrzehnten wurden zunehmend Alternativen zu Ertragstafeln entwickelt, unter anderem prozessbasierte Modelle, die auch Zustände ausserhalb der Beobachtungsdaten abbilden können. Solche Modelle berücksichtigen Wachstumsdaten, aber ihre Formulierungen basieren meist auf physiologischen und allometrischen Prinzipien des Pflanzenwachstums. Zu dieser Modellfamilie gehört 3-PG (Physiological Processes Predicting Growth), das in Australien und Nordamerika für gleichaltrige Reinbestände von immergrünen Laub- und Nadelbäumen entwickelt wurde. In einer kürzlich erfolgten Weiterentwicklung (Version 3-PGmix) können auch Wechselwirkungen zwischen Bäumen verschiedener Grösse, verschiedenen Alters und verschiedener Arten simuliert werden. Der Artikel beschreibt das Modell, das aus sechs Untermodellen besteht und trotz seiner physiologischen Grundlagen Kenngrössen liefert, die für die praktische Forstwirtschaft von grosser Bedeutung sind. Zwei Beispiele von Anwendungen von 3-PG für die Schweiz werden vorgestellt. Im ersten Beispiel wird die Veränderung des Wachstums in Mischbeständen aus Waldföhre (Pinus sylvestris) und Buche (Fagus sylvatica) aufgrund des Klimawandels dargestellt. Dabei wird gezeigt, dass das Wachstum der Buche durch erhöhte Trockenheit wesentlich stärker eingeschränkt wird als jenes der Waldföhre, die zwar ebenfalls durch die Trockenheit stärker limitiert wird, aber gleichzeitig vom geringeren Konkurrenzdruck durch die Buche profitiert. Im zweiten Beispiel wird 3-PG angewendet, um den klimatischen Stress für die Buche als Funktion der Standorteigenschaften schweizweit zu quantifizieren. Beide Beispiele zeigen, dass 3-PG standortspezifische Informationen liefert, wie die Waldwachstumsdynamik durch Klima, Artenzusammensetzung und Waldbau beeinflusst wird.

The expected future intensification of forest disturbance as a consequence of ongoing anthropogenic climate change highlights the urgent need to more robustly quantify associated biotic responses. Saproxylic beetles are a diverse group of forest invertebrates representing a major component of biodiversity that is associated with the decomposition and cycling of wood nutrients and carbon in forest ecosystems. Disturbance-induced declines or shifts in their diversity indicate the loss of key ecological and/or morphological species traits that could change ecosystem functioning. Functional and phylogenetic diversity of biological communities is commonly used to link species communities to ecosystem functions. However, our knowledge on how disturbance intensity alters functional and phylogenetic diversity of saproxylic beetles is incomplete. Here, we analyzed the main drivers of saproxylic beetle abundance and diversity using a comprehensive dataset from montane primary forests in Europe. We investigated cascading relationships between 250 years of historical disturbance mechanisms, forest structural attributes and the taxonomic, phylogenetic and functional diversity of present-day beetle communities. Our analyses revealed that historical disturbances have significant effects on current beetle communities. Contrary to our expectations, different aspects of beetle communities, that is, abundance, taxonomic, phylogenetic and functional diversity, responded to different disturbance regime components. Past disturbance frequency was the most important component influencing saproxylic beetle communities and habitat via multiple temporal and spatial pathways. The quantity of deadwood and its diameter positively influenced saproxylic beetle abundance and functional diversity, whereas phylogenetic diversity was positively influenced by canopy openness. Analyzing historical disturbances, we observed that current beetle diversity is far from static, such that the importance of various drivers might change during further successional development. Only forest landscapes that are large enough to allow for the full range of temporal and spatial patterns of disturbances and post-disturbance development will enable long-term species coexistence and their associated ecosystem functions.

With accelerating environmental change, understanding forest disturbance impacts on trade-offs between biodiversity and carbon dynamics is of high socio-economic importance. Most studies, however, have assessed immediate or short-term effects of disturbance, while long-term impacts remain poorly understood. Using a tree-ring-based approach, we analysed the effect of 250 years of disturbances on present-day biodiversity indicators and carbon dynamics in primary forests. Disturbance legacies spanning centuries shaped contemporary forest co-benefits and trade-offs, with contrasting, local-scale effects. Disturbances enhanced carbon sequestration, reaching maximum rates within a comparatively narrow post-disturbance window (up to 50 years). Concurrently, disturbance diminished aboveground carbon storage, which gradually returned to peak levels over centuries. Temporal patterns in biodiversity potential were bimodal; the first maximum coincided with the short-term post-disturbance carbon sequestration peak, and the second occurred during periods of maximum carbon storage in complex old-growth forest. Despite fluctuating local-scale trade-offs, forest biodiversity and carbon storage remained stable across the broader study region, and our data support a positive relationship between carbon stocks and biodiversity potential. These findings underscore the interdependencies of forest processes, and highlight the necessity of large-scale conservation programmes to effectively promote both biodiversity and long-term carbon storage, particularly given the accelerating global biodiversity and climate crises.

Climatic constraints on tree growth mediate an important link between terrestrial and atmospheric carbon pools. Tree rings provide valuable information on climate-driven growth patterns, but existing data tend to be biased toward older trees on climatically extreme sites. Understanding climate change responses of biogeographic regions requires data that integrate spatial variability in growing conditions and forest structure. We analyzed both temporal (c. 1901-2010) and spatial variation in radial growth patterns in 9,876 trees from fragments of primary Picea abies forests spanning the latitudinal and altitudinal extent of the Carpathian arc. Growth was positively correlated with summer temperatures and spring moisture availability throughout the entire region. However, important seasonal variation in climate responses occurred along geospatial gradients. At northern sites, winter precipitation and October temperatures of the year preceding ring formation were positively correlated with ring width. In contrast, trees at the southern extent of the Carpathians responded negatively to warm and dry conditions in autumn of the year preceding ring formation. An assessment of regional synchronization in radial growth variability showed temporal fluctuations throughout the 20th century linked to the onset of moisture limitation in southern landscapes. Since the beginning of the study period, differences between high and low elevations in the temperature sensitivity of tree growth generally declined, while moisture sensitivity increased at lower elevations. Growth trend analyses demonstrated changes in absolute tree growth rates linked to climatic change, with basal area increments in northern landscapes and lower altitudes responding positively to recent warming. Tree growth has predominantly increased with rising temperatures in the Carpathians, accompanied by early indicators that portions of the mountain range are transitioning from temperature to moisture limitation. Continued warming will alleviate large-scale temperature constraints on tree growth, giving increasing weight to local drivers that are more challenging to predict.

A tree’s radial growth sequence can be thought of as an aggregate of different growth components such as age and size limitations, presence or absence of disturbance events, continuous impact of climate variability and variance induced by unknown origin. The potentially very complex growth patterns with prominent temporal and spatial variability imply that our understanding of climate-vegetation feedbacks essentially benefits from the expansion of large tree ring networks into data-poor regions, and our ability to disentangle growth constraints by comparing ring series at multiple scales. In this study, we analyze Central-Eastern Europe’s most substantial assemblage of primary Norway spruce forests found in the Carpathian arc. The vast data set, >10,000 tree-ring series, is stratified along a prominent gradient in climate response space over four separate landscapes. We integrated curve intervention detection and dendroclimatic standardization to decompose tree growth variance into climatic, disturbance and residual components to explore the behavior of the components over increasingly larger spatial hierarchies. We show that the residual variance of unknown origin is the most prominent variance in individual Carpathian spruce trees, but at larger spatial hierarchies, climate variance dominates. The variance induced by climate was further explored with common correlation analyses, growth response to extreme climate years and forward modeling of tree growth to identify leading modes of climate response, and potentially non-linear and mixed climate response patterns. We find that the climatic response of the different forest landscapes overall can be described as an asymptotic response to June and July temperatures, most likely intermixed with influence from winter precipitation. In the collection of landscapes, Southern Romania stands out as being the least temperature sensitive and most likely exhibiting the most complicated mixed temperature and moisture limitation.

Winter drought becomes a limiting factor of forest stand growth by the end of the twentieth century. Disturbances strongly influence the structure of natural forests. The frequency and severity of natural disturbances, as well as drought events, are expected to increase with climate change. Our study investigated if forests with differing forest structures related to disturbance histories also differed in sensitivity to drought. In a natural forest landscape in the Calimani Mountains of the Eastern Carpathians, Romania, we used six forest patches to represent different successional stages, from early- to late-successional stages. We used two temporal resolutions of the Standardized Precipitation-Evapotranspiration Index to describe short and long water resource dynamics within or between hydrological years, respectively. We detected an increase in the importance of winter drought across all successional stages; it was first identified in the oldest patch in the 1970s, and it consecutively affected younger patches. We observed that different forest structures do not lead to substantial differences in trends in drought–growth relationships. A shift in sensitivity to water availability from early spring to winter occurred over the twentieth century. These findings suggest that the impact of climate change on Norway spruce forest ecosystems of the Eastern Carpathians will likely be difficult to mitigate at a local scale using traditional forestry practices.

Disentangling the importance of developmental vs. environmental drivers of variation in forest biomass is key to predicting the future of forest carbon sequestration. At coarse scales, forest biomass is likely to vary along major climatic and physiographic gradients. Natural disturbance occurs along these broad biophysical gradients, and depending on their extent, severity and frequency, could either amplify or dampen spatial heterogeneity in forest biomass. Here we evaluate spatial variation in the basal area of late-successional Picea abies (L./Karst.) forests across the Carpathian Mountain Range of central Europe and compare the roles of coarse-scale biophysical gradients and natural disturbances in driving that variation across a hierarchy of scales (landscapes, stands, and plots). We inventoried forest composition and structure, and reconstructed disturbance histories using tree cores collected from 472 plots nested within 30 late-successional stands, spanning the Carpathian Mountains (approximately 4.5 degrees of latitude). We used linear mixed-effects models to compare the effect of disturbance regimes and site conditions on stand basal area at three hierarchical scales. We found that the basal area of late-successional Picea abies forests varied across a range of spatial scales, with climatic drivers being most important at coarse scales and natural disturbances acting as the primary driver of forest heterogeneity at fine scales. For instance, the stand-level basal area varied among landscapes, with the highest values (48–68 m2 ha−1) in the warmer southern Carpathian Mountains, and lower values (37–52 m2 ha−1 on average) in cooler areas of the eastern and western Carpathians. Finer-scale variation was driven by local disturbances (mainly bark beetle and windstorms) and the legacies of disturbances that occurred more than a century ago. Our findings suggest that warming could increase the basal area of northern sites, but potential increasing disturbances could disrupt these environmental responses.

Forest demographic processes - growth, recruitment and mortality - are being altered by global change. The changing balance between growth and mortality strongly influences forest dynamics and the carbon balance. Elevated atmospheric carbon dioxide (eCO2) has been reported to enhance photosynthesis and tree growth rates by increasing both light-use efficiency (LUE) and water-use efficiency (WUE). Tree growth enhancement could be translated into an increase in biomass stocks or could be associated with a reduction in the longevity of trees, thus reducing the ability of forest ecosystems to act as carbon sinks over long timescales. These links between growth and mortality, and the implications for forest stand density and self-thinning relationships are still debated. Scarce empirical evidence exists for how changing drivers affect tree mortality due to existing data and modelling limitations. Understanding the causes of observed mortality trends and the mechanisms underlying these processes is critical for accurate projections of global terrestrial carbon storage and its feedbacks to anthropogenic climate change.Here, we combine a mechanistic model with empirical forest data to better understand the causes of changes in tree mortality and the implications for past and future trends in forest tree density. Specifically, we test the Grow-Fast-Die-Young hypothesis to investigate if a leaf-level CO2 fertilization effect may lead to an increase in the biomass stock in forest stands. We use a novel vegetation demography model (LM3-PPA) which includes vegetation dynamics with biogeochemical processes allowing for explicit representation of individuals and a mechanistic treatment of tree mortality. The key links between leaf-level assimilation and stand dynamics depend on the carbon turnover time. In this sense, we investigate alternative mortality assumptions about the functional dependence of mortality on tree size, tree carbon balance or growth rate. These formulations represent typical approaches to simulate mortality in mechanistic forest models. Model simulations show that increasing photosynthetic LUE leads to higher biomass stocks, with contrasting behavior among mortality assumptions. Empirical data from Swiss forest inventories support the results from the model simulations showing a shift upwards in the self-thinning relationships, with denser stands and bigger trees. This data-supported mortality-modelling helps to identify links between forest responses and environmental changes at the leaf, tree and stand levels and yields new insight into the causes of currently observed terrestrial carbon sinks and future responses.

Restoring the structural characteristics of secondary old-growth forests that were previously managed is increasingly debated to help increase the area of more complex forests which provide a broader array of forest services and functions. The paucity of long-term data sets in Central Europe has limited our ability to understand the ongoing ecological processes required for effective restoration programs for old-growth forests. To address this, we used repeated census data from eight permanent plots to evaluate forest structural dynamics over a 12-year period in the largest complex of European beech (Fagus sylvatica L.) forests in the Czech Highlands without intensive forestry intervention for almost 50 years. Our results showed that previously managed forests can exhibit structural qualities typically associated with old-growth forests after management has ceased for a period. The stand structural characteristics (e.g., density of large and old trees) is comparable with protected reserves of old-growth European beech-dominated forests. The average stand age was 196 years, but the oldest tree was 289 years old. The annual mortality rate was 0.43% for all species, and the U-shaped distribution indicating size-dependent mortality is likely an important process that is balanced by the turnover of new tree recruitment. During the study period, we detected that the diameter distribution tended towards a rotated sigmoid distribution. The lasting effects of the most recent forest management are evident in the scarcity of dead wood, and a prolonged process of dead wood accumulation has begun. Thus, the abandonment of all management activities in near-natural forest reserves, including dead wood removal, will ensure that the forests will develop characteristics typical of old-growth forests.

Warming alters the variability and trajectories of tree growth around the world by intensifying or alleviating energy and water limitation. This insight from regional to global-scale research emphasizes the susceptibility of forest ecosystems and resources to climate change. However, globally-derived trends are not necessarily meaningful for local nature conservation or management considerations, if they lack specific information on present or prospective tree species. This is particularly the case towards the edge of their distribution, where shifts in growth trajectories may be imminent or already occurring.Importantly, the geographic and bioclimatic space (or “niche”) occupied by a tree species is not only constrained by climate, but often reflects biotic pressure such as competition for resources with other species. This aspect is underrepresented in many species distribution models that define the niche as a climatic envelope, which is then allowed to shift in response to changes in ambient conditions. Hence, distinguishing climatic from competitive niche boundaries becomes a central challenge to identifying areas where tree species are most susceptible to climate change.Here we employ a novel concept to characterize each position within a species’ bioclimatic niche based on two criteria: a climate sensitivity index (CSI) and a habitat suitability index (HSI). The CSI is derived from step-wise multiple linear regression models that explain variability in annual radial tree growth as a function of monthly climate anomalies. The HSI is based on an ensemble of five species distribution models calculated from a combination of observed species occurrences and twenty-five bioclimatic variables. We calculated these two indices for 11 major tree species across the Northern Hemisphere.The combination of climate sensitivity and habitat suitability indicated hotspots of change, where tree growth is mainly limited by competition (low HSI and low CSI), as well as areas that are particularly sensitive to climate variability (low HSI and high CSI). In the former, we expect that forest management geared towards adjusting the competitive balance between several candidate species will be most effective under changing environmental conditions. In the latter areas, selecting particularly drought-tolerant accessions of a given species may reduce forest susceptibility to the predicted warming and drying.

Under unprecedent climate change and increased frequency of extreme events, e.g. drought, it is important to assess and forecast forest ecosystem vulnerability and stability. Large volumes of data from observational and experimental networks, increases in computational power, advances in ecological models, and optimization methodologies are the main measures to improve quantitative forecasting in ecology. Data assimilation is a key tool to improve ecosystem state prediction and forecasting by combining model simulations and observations. We assimilated observations of carbon stocks and fluxes from 271 permanent long-term forest monitoring plots across Switzerland into the 3-PG forest ecosystem model using Bayesian inference, reducing the bias of model predictions from 14% to 5% for forest stem carbon stocks and from 45% to 9% for stem carbon stock changes, respectively. We then estimated the productivity of forests dominated by Picea abies and Fagus sylvatica for the period of 1960-2018 and tested for climate-induced shifts in productivity along elevational gradient and in extreme years. Overall, we demonstrated a high potential of using data assimilation to improve predictions of forest ecosystem productivity. Furthermore, our calibrated model simulations suggest that climate extremes affect forest productivity in more complex ways than by simply shifting the response upwards in elevation.

International audience; Tree-related microhabitats (TreMs) are important features for the conservation of biodiversity in forest ecosystems. Although other structural indicators of forest biodiversity have been extensively studied in recent decades, TreMs have often been overlooked, either due to the absence of a consensual definition or a lack of knowledge. Despite the increased number of TreM studies in the last decade, the role of drivers of TreM profile in primary forests and across different geographical regions is still unknown. To evaluate the main drivers of TreM density and diversity, we conducted the first large-scale study of TreMs across European primary forests. We established 146 plots in eight primary forests dominated by European beech (Fagus sylvatica L.) in the Carpathian and Dinaric mountain ranges. Generalized linear mixed effect models were used to test the effect of local plot characteristics and spatial variability on the density and diversity (alpha, beta, and gamma) of TreMs. Total TreM density and diversity were significantly positively related with tree species richness and the proportion of snags. Root mean square tree diameters were significantly related to alpha and gamma diversity of TreMs. Both regions reached similarly high values of total TreM densities and total TreM densities and diversity were not significantly different between the two regions; however, we observed between the two regions significant differences in the densities of two TreM groups, conks of fungi and epiphytes. The density and diversity of TreMs were very high in beech-dominated mountain primary forests, but their occurrence and diversity was highly variable within the landscapes over relatively short spatial gradients (plot and stand levels). Understanding these profile provides a benchmark for further comparisons, such as with young forest reserves, or for improving forest management practices that promote biodiversity.

The study was supported by the institutional project MSMT (CZ.02.1.01/0.0/0.0/16_019/0000803) and the Czech Ministry of Education (Project INTER-COST No. LCT17055). Accurately capturing medium- to low-frequency trends in tree-ring data is vital to assessing climatic response and developing robust reconstructions of past climate. Non-climatic disturbance can affect growth trends in tree-ring-width (RW) series and bias climate information obtained from such records. It is important to develop suitable strategies to ensure the development of chronologies that minimize these medium- to low-frequency biases. By performing high density sampling (760 trees) over a ~40-ha natural high-elevation Norway spruce (Picea abies) stand in the Romanian Carpathians, this study assessed the suitability of several sampling strategies for developing chronologies with an optimal climate signal for dendroclimatic purposes. There was a roughly equal probability for chronologies (40 samples each) to express a reasonable (r = 0.3?0.5) to non-existent climate signal. While showing a strong high-frequency response, older/larger trees expressed the weakest overall temperature signal. Although random sampling yielded the most consistent climate signal in all sub-chronologies, the outcome was still sub-optimal. Alternative strategies to optimize the climate signal, including very high replication and principal components analysis, were also unable to minimize this disturbance bias and produce chronologies adequately representing climatic trends, indicating that larger scale disturbances can produce synchronous pervasive disturbance trends that affect a large part of a sampled population. The Curve Intervention Detection (CID) method, used to identify and reduce the influence of disturbance trends in the RW chronologies, considerably improved climate signal representation (from r = 0.28 before correction to r = 0.41 after correction for the full 760 sample chronology over 1909?2009) and represents a potentially important new approach for assessing disturbance impacts on RW chronologies. Blue intensity (BI) also shows promise as a climatically more sensitive variable which, unlike RW, does not appear significantly affected by disturbance. We recommend that studies utilizing RW chronologies to investigate medium- to long-term climatic trends also assess disturbance impact on those series. Postprint

Natural disturbances strongly influence forest structural dynamics, and subsequently stand structural heterogeneity, biomass, and forest functioning. The impact of disturbance legacies on current forest structure can greatly influence how we interpret drivers of forest dynamics. However, without clear insight into forest history, many studies default to coarse assumptions about forest structure, for example, whether forests are even or unevenly aged. The aim of this study was to analyze the effects of past disturbances on the current diameter distributions of Norway spruce (Picea abies (L.) Karst.)-dominated landscapes throughout the Carpathian Mountains. Our dendroecological dataset comprises tree cores from 339 plots (7,845 total tree cores), nested within 28 primary forest stands, known to vary greatly in the severity of historical disturbances. Our analyses revealed that historical disturbances had a strong and significant effect on the current diameter distribution shapes at the plot level. We demonstrated that mixed-severity disturbance regimes were more frequent and create a complex pattern of diameter distributions at the plot and stand scale. Here, we show that high severity disturbance was associated with unimodal diameter distributions, while low and moderate severity was associated with the reverse J-shaped distribution. This is a result of complex disturbance patterns, with structural biological legacies. Our results will have important management implication in the context of tree size heterogeneity, biomass storage, and productivity as influenced by natural disturbances. Lastly, these results demonstrate that structural changes may arise as consequences of changing disturbance regime associated with global change.

Mixed-severity disturbance regimes are prevalent in temperate forests worldwide, but key uncertainties remain regarding the variability of disturbance-mediated structural development pathways. This study investigates the influence of disturbance history on current structure in primary, unmanaged Norway spruce (Picea abies) forests throughout the Carpathian Mountains of central and eastern Europe, where windstorms and native bark beetle outbreaks are the dominant natural disturbances. We inventoried forest structure on 453 plots (0.1 ha) spanning a large geographical gradient (>1,000 km), coring 15–25 canopy trees per plot for disturbance history reconstruction (tree core total n = 11,309). Our specific objectives were to: (1) classify sub-hectare-scale disturbance history based on disturbance timing and severity; (2) classify current forest structure based on tree size distributions (live, dead, standing, downed); (3) characterize structural development pathways as revealed by the association between disturbance history and current forest structural complexity. We used hierarchical cluster analysis for the first two objectives and indicator analysis for the third. The disturbance-based cluster analysis yielded six groups associated with three levels of disturbance severity (low, moderate, and high canopy loss) and two levels of timing (old, recent) over the past 200 years. The structure-based cluster analysis yielded three groups along a gradient of increasing structural complexity. A large majority of plots exhibited relatively high (53%) or very high (26%) structural complexity, indicated by abundant large live trees, standing and downed dead trees, and spruce regeneration. Consistent with conventional models of structural development, some disturbance history groups were associated with specific structural complexity groups, particularly low-severity/recent (very high complexity) and high-severity/recent (moderate complexity) disturbances. In other cases, however, the distribution of plots among disturbance history and structural complexity groups indicated either divergent or convergent pathways. For example, multiple disturbance history groups were significantly associated with the high complexity group, demonstrating structural convergence. These results illustrate that complex forest structure – including features nominally associated with old-growth – can be associated as much with disturbance severity as it is with conventional notions of forest age. Because wind and bark beetles are natural disturbance processes that can induce relatively high levels of tree mortality while simultaneously contributing to structural complexity and heterogeneity, we suggest that forest management plans allow for the stochastic occurrence of disturbance and variable post-disturbance development trajectories. Such applications are especially appropriate in conservation areas where biodiversity and forest resilience are management objectives, particularly given projections of increasing disturbance activity under global change.

Primary forests are characterized by high vertical and horizontal stand diversity, which provides habitat for a diverse range of species with complex habitat requirements. Detailed knowledge of related ecological processes and habitat development of primary forest species are essential to inform forest management and biodiversity conservation decisions, but relationships are not well documented. We collected dendrochronological data and inventoried numerous structural elements in permanent plots throughout the primary temperate forests within the Carpathian Mountains. We fit and compared multiple predictive models to quantify the importance of 200 years of natural disturbance dynamics on the occurrence probability of an umbrella species – the capercaillie (Tetrao urogallus). We showed that a mixed-severity disturbance regime ranging from low through moderate to high severity disturbances is required to generate diverse forest habitats suitable for capercaillie. The variation in natural disturbance severity and its timing promoted key structural habitat elements, such as low natural regeneration density, low mature tree density, high ground vegetation cover, availability of forest gaps, and abundance of standing deadwood. This study demonstrates the importance of natural disturbance in maintaining the variety of conditions necessary to support primary forest specialist species. Managers of protected areas should be mindful that natural disturbances generate habitat for the capercaillie in mountain Norway spruce forests. Further intervention is unnecessary. Conservation planning and forest reserve design should shift focus to the large-scale spatial requirements needed to ensure that a wide range of forest developmental phases are represented in protected areas.

European natural mountain Norway spruce (Picea abies) forests are currently subject to extensive disturbances. An improved understanding of the self-regulated regenerative capacity of this forest type is therefore needed. We used the last remnant of natural mountain Norway spruce forests in central northwestern Europe (BNF Brocken natural forest), to analyze (1) the diversity of structure and age distribution of the tree population and (2) the effect of disturbances on self-regulated tree regeneration over the last 264 years. To this end, we combined an assessment of stand structure with dendrochronological investigations and a review of disturbance history. We hypothesized that BNF exhibits a high diversity of tree ages and dimensions and that recruitment and survival of tree regeneration were largely independent from disturbances. BNF showed a high structural and age diversity. Disturbances exhibited no regular temporal pattern. Their effect on tree regeneration was rather complex and changed with observation period. Impeding and facilitating effects of past disturbances on recruitment were significant from 1736 to 1910. From 1911 until 2000, recruitment decoupled from preceding disturbances. Subsequent disturbances facilitated survival of established trees from 1736 to 1820, while afterward no significant influence could be proved. Our study showed that in the course of self-regulated development the tree population of BNF has gradually acquired, or maintained, a diverse structure. Disturbances served as an important driver of diversification. We concluded that increasing deadwood availability and limiting browsing are the key to securing immediate regeneration.

Forest disturbances are sensitive to climate. However, our understanding of disturbance dynamics in response to climatic changes remains incomplete, particularly regarding large-scale patterns, interaction effects and dampening feedbacks. Here we provide a global synthesis of climate change effects on important abiotic (fire, drought, wind, snow and ice) and biotic (insects and pathogens) disturbance agents. Warmer and drier conditions particularly facilitate fire, drought and insect disturbances, while warmer and wetter conditions increase disturbances from wind and pathogens. Widespread interactions between agents are likely to amplify disturbances, while indirect climate effects such as vegetation changes can dampen long-term disturbance sensitivities to climate. Future changes in disturbance are likely to be most pronounced in coniferous forests and the boreal biome. We conclude that both ecosystems and society should be prepared for an increasingly disturbed future of forests. © 2017 Macmillan Publishers Limited, part of Springer Nature. All rights reserved.

Habitat loss and fragmentation can negatively impact the persistence of dispersal-limited lichen species with narrow niches. Rapid change in microclimate due to canopy dieback exposes species to additional stressors that may limit their capacity to survive and colonize. We studied the importance of old trees as micro-refuges and microclimate stability in maintaining lichen survival and diversity. The study was situated in mountain Norway spruce (Picea abies) forests of the Gorgany Mountains of the Ukrainian Carpathian mountain belt. Lichens were collected on 13 circular study plots (1000 m2). Dendrochronological methods were used to reconstruct age structure and maximum disturbance event history. A linear mixed effects model and general additive models were used to explain patterns and variability of lichens based on stand age and disturbance history for each plot. Tree age was the strongest variable influencing lichen diversity and composition. Recent ( 200 years old) that survived the disturbances served as microrefuges for the habitat-specialized and/or dispersal limited species, thus epiphytic lichen biodiversity was markedly higher on those plots in comparison to plots without any old trees. Most species were able to survive microclimatic change after disturbances, or recolonize disturbed patches from surrounding old-growth forests. We concluded that the survival of old trees after disturbances could maintain and/or recover large portions of epiphytic lichen biodiversity even in altered microclimates.

In order to gauge ongoing and future changes to disturbance regimes, it is necessary to establish a solid baseline of historic disturbance patterns against which to evaluate these changes. Further, understanding how forest structure and composition respond to variation in past disturbances may provide insight into future resilience to climate-driven alterations of disturbance regimes. We established 184 plots (mostly 1000 m2) in 14 primary mountain Norway spruce forests in the Western Carpathians. On each plot we surveyed live and dead trees and regeneration, and cored around 25 canopy trees. Disturbance history was reconstructed by examining individual tree growth trends. The study plots were further aggregated into five groups based on disturbance history (severity and timing) to evaluate and explain its influence on forest structure. These ecosystems are characterized by a mixed severity disturbance regime with high spatiotemporal variability in severity and frequency. However, periods of synchrony in disturbance activity were also found. Specifically, a peak of canopy disturbance was found for the mid-19th century across the region (about 60% of trees established), with the most important periods of disturbance in the 1820s and from the 1840s to the 1870s. Current stand size and age structure were strongly influenced by past disturbance activity. In contrast, past disturbances did not have a significant effect on current tree density, the amount of coarse woody debris, and regeneration. High mean densities of regeneration with height >50 cm (about 1400 individuals per ha) were observed. Extensive high severity disturbances have recently affected Central European forests, spurring a discussion about the causes and consequences. We found some evidence that forests in the Western Carpathians were predisposed to recent severe disturbance events as a result of synchronized past disturbance activity, which partly homogenized size and age structure and made recent stands more vulnerable to bark beetle outbreak. Our data suggest that these events are still part of the range of natural variability. The finding that regeneration density and volume of coarse woody debris were not influenced by past disturbance illustrates that vastly different past disturbance histories are not likely to change the future trajectories of these forests. These ecosystems currently have high ecological resilience to disturbance. In conclusion, we suggest that management should recognize disturbances as a natural part of ecosystem dynamics in the mountain forests of Central Europe, account for their stochastic occurrence in management planning, and mimic their patterns to foster biodiversity in forest landscapes.

During the growing season, trees allocate photoassimilates to increase their aboveground woody biomass in the stem (ABIstem). This ‘carbon allocation’ to structural growth is a dynamic process influenced by internal and external (e.g., climatic) drivers. While radial variability in wood formation and its resulting structure have been intensively studied, their variability along tree stems and subsequent impacts on ABIstem remain poorly understood. We collected wood cores from mature trees within a fixed plot in a well-studied temperate Fagus sylvatica L. forest. For a subset of trees, we performed regular interval sampling along the stem to elucidate axial variability in ring width (RW) and wood density (ρ), and the resulting effects on tree- and plot-level ABIstem. Moreover, we measured wood anatomical traits to understand the anatomical basis of ρ and the coupling between changes in RW and ρ during drought. We found no significant axial variability in ρ because an increase in the vessel-to-fiber ratio with smaller RW compensated for vessel tapering towards the apex. By contrast, temporal variability in RW varied significantly along the stem axis, depending on the growing conditions. Drought caused a more severe growth decrease, and wetter summers caused a disproportionate growth increase at the stem base compared with the top. Discarding this axial variability resulted in a significant overestimation of tree-level ABIstem in wetter and cooler summers, but this bias was reduced to ~2% when scaling ABIstem to the plot level. These results suggest that F. sylvatica prioritizes structural carbon sinks close to the canopy when conditions are unfavorable. The different axial variability in RW and ρ thereby indicates some independence of the processes that drive volume growth and wood structure along the stem. This refines our knowledge of carbon allocation dynamics in temperate diffuse-porous species and contributes to reducing uncertainties in determining forest carbon fixation.

The response of forest productivity to climate extremes strongly depends on ambient environmental and site conditions. To better understand these relationships at a regional scale, we used nearly 800 observation years from 271 permanent long‐term forest monitoring plots across Switzerland, obtained between 1980 and 2017. We assimilated these data into the 3‐PG forest ecosystem model using Bayesian inference, reducing the bias of model predictions from 14% to 5% for forest stem carbon stocks and from 45% to 9% for stem carbon stock changes. We then estimated the productivity of forests dominated by Picea abies and Fagus sylvatica for the period of 1960–2018, and tested for productivity shifts in response to climate along elevational gradient and in extreme years. Simulated net primary productivity (NPP) decreased with elevation (2.86 ± 0.006 Mg C ha−1 year−1 km−1 for P. abies and 0.93 ± 0.010 Mg C ha−1 year−1 km−1 for F. sylvatica). During warm–dry extremes, simulated NPP for both species increased at higher and decreased at lower elevations, with reductions in NPP of more than 25% for up to 21% of the potential species distribution range in Switzerland. Reduced plant water availability had a stronger effect on NPP than temperature during warm‐dry extremes. Importantly, cold–dry extremes had negative impacts on regional forest NPP comparable to warm–dry extremes. Overall, our calibrated model suggests that the response of forest productivity to climate extremes is more complex than simple shift toward higher elevation. Such robust estimates of NPP are key for increasing our understanding of forests ecosystems carbon dynamics under climate extremes., We assimilated extensive and long‐term monitoring data into the 3‐PG forest ecosystem model to assess how forest productivity responds to climate extremes across environmental gradients in Switzerland. Data assimilation using Bayesian inference significantly improved model predictions. During warm–dry extremes, net primary productivity increased at higher and decreased at lower elevations, with reductions in NPP of more than 25% for one‐fifth of the potential species distribution range in Switzerland.

Estimates of historical disturbance patterns are essential to guide forest management aimed at ensuring the sustainability of ecosystem functions and biodiversity. However, quantitative estimates of various disturbance characteristics required in management applications are rare in longer-term historical studies. Thus, our objectives were to (1) quantify past disturbance severity, patch size, and stand proportion disturbed and (2) test for temporal and subregional differences in these characteristics. We developed a comprehensive dendrochronological method to evaluate an approximately two-century-long disturbance record in the remaining Central and Eastern European primary mountain spruce forests, where wind and bark beetles are the predominant disturbance agents. We used an unprecedented large-scale nested design data set of 541 plots located within 44 stands and 6 subregions. To quantify individual disturbance events, we used tree-ring proxies, which were aggregated at plot and stand levels by smoothing and detecting peaks in their distributions. The spatial aggregation of disturbance events was used to estimate patch sizes. Data exhibited continuous gradients from low- to high-severity and small- to large-size disturbance events. In addition to the importance of small disturbance events, moderate-scale (25-75% of the stand disturbed, >10 ha patch size) and moderate-severity (25-75% of canopy disturbed) events were also common. Moderate disturbances represented more than 50% of the total disturbed area and their rotation periods ranged from one to several hundred years, which is within the lifespan of local tree species. Disturbance severities differed among subregions, whereas the stand proportion disturbed varied significantly over time. This indicates partially independent variations among disturbance characteristics. Our quantitative estimates of disturbance severity, patch size, stand proportion disturbed, and associated rotation periods provide rigorous baseline data for future ecological research, decisions within biodiversity conservation, and silviculture intended to maintain native biodiversity and ecosystem functions. These results highlight a need for sufficiently large and adequately connected networks of strict reserves, more complex silvicultural treatments that emulate the natural disturbance spectrum in harvest rotation times, sizes, and intensities, and higher levels of tree and structural legacy retention.

International audience; Transpiration in humid tropical forests modulates the global water cycle and is a key driver of climate regulation. Yet, our understanding of how tropical trees regulate sap flux in response to climate variability remains elusive. With a progressively warming climate, atmospheric evaporative demand [i.e., vapor pressure deficit (VPD)] will be increasingly important for plant functioning, becoming the major control of plant water use in the twenty-first century. Using measurements in 34 tree species at seven sites across a precipitation gradient in the neotropics, we determined how the maximum sap flux velocity (vmax) and the VPD threshold at which vmax is reached (VPDmax) vary with precipitation regime [mean annual precipitation (MAP); seasonal drought intensity (PDRY)] and two functional traits related to foliar and wood economics spectra [leaf mass per area (LMA); wood specific gravity (WSG)]. We show that, even though vmax is highly variable within sites, it follows a negative trend in response to increasing MAP and PDRY across sites. LMA and WSG exerted little effect on vmax and VPDmax, suggesting that these widely used functional traits provide limited explanatory power of dynamic plant responses to environmental variation within hyper-diverse forests. This study demonstrates that long-term precipitation plays an important role in the sap flux response of humid tropical forests to VPD. Our findings suggest that under higher evaporative demand, trees growing in wetter environments in humid tropical regions may be subjected to reduced water exchange with the atmosphere relative to trees growing in drier climates.

© 2019 The Authors FORMIT-M is a widely applicable, open-access, simple and flexible, climate-sensitive forest management simulator requiring only standard forest inventory data as input. It combines a process-based carbon balance approach with a strong inventory-based empirical component. The model has been linked to the global forest sector model EFI-GTM to secure consistency between timber cutting and demand, although prescribed harvest scenarios can also be used. Here we introduce the structure of the model and demonstrate its use with example simulations until the end of the 21st century in Europe, comparing different management scenarios in different regions under climate change. The model was consistent with country-level statistics of growing stock volumes (R 2 = 0.938) and its projections of climate impact on growth agreed with other studies. The management changes had a greater impact on growing stocks, harvest potential and carbon balance than projected climate change, at least in the absence of increased disturbance rates. ispartof: Environmental Modelling and Software vol:115 pages:128-143 status: published

Stand-level process-based models have rarely been applied to uneven-aged forests that contain many size classes and negative exponential shaped size distributions. However, the relative simplicity of such models, in terms of parameterisation, use and interpretation, could make them valuable tools for studying and managing such forests. In particular, the effects of climate change on the stand-level growth of forests with negative exponential shaped size distributions has received very little attention compared with even-aged forests. The first objective of this study was to validate 3-PG, a stand-level process-based model, for five types of uneven-aged forests in Switzerland; (1) Fagus sylvatica dominated, (2) Picea abies dominated, (3) mixtures of Picea abies and Abies alba, (4) mixtures of Picea abies, Abies alba and Fagus sylvatica, and (5) mixtures of Larix decidua, Pinus cembra and P. abies. The second objective was to use 3-PG to examine how climate change has influenced the growth of these forests since the 1930s. 3-PG predictions of biomass, biomass partitioning in above- and belowground components, and light absorption were validated using inventory data from 23 plots, which had been monitored for an average of 81 years (15 to 112 years). For all species and size classes (2–3 per species), 3-PG produced accurate predictions of root biomass, stem biomass and outputs derived from it such as mean diameter, basal area and height, which were all highly correlated with the observed values (R2 > 0.86). The slope of predicted versus observed values was often not significantly different to 1 (averaged 1.13) and the bias averaged −1.2%. 3-PG simulations to examine the effects of climate change without the confounding effects of stand structure and management, showed that the growth of the five forests types has, on average, increased by 17% since the 1930s. The growth was mainly influenced by temperature, while in the case of A. alba, growth was largely influenced by vapour pressure deficit. The accelerated growth rates imply that thinning intensities also need to increase to prevent high stand densities from inhibiting regeneration in these uneven-aged forests. This study shows that 3-PG can be used to predict the growth dynamics of uneven-aged mixed-species forests, and to our knowledge, this is the first time a stand-level process-based forest growth model has been used and validated for such forests.

Disturbances, both natural and human induced, influence forest dynamics, ecosystem functioning, and ecosystem services. Here, we aim to evaluate the consequences of natural disturbances on the magnitude and dynamics of tree- and stand-level biomass accumulation from decadal to centennial scales. We use tree-ring data from 2301 trees and biometric data from 4909 trees sampled in 96 plots (each 1000 m 2 ) to quantify the influence of mixed severity disturbance regimes on annual aboveground biomass increment (AGBI) and total aboveground biomass accumulation (AGB) across a mountainous monotypic Norway spruce ( Picea abies (L.) Karst.) primary forest. We hypothesise that the multiple internal and external factors constraining tree growth will cause differences in tree and stand-level biomass trajectories in these natural forests. Although we found that tree-level AGB growth increases with tree size, we also found that tree age and disturbance legacies plays a crucial role for AGB in the investigated Norway spruce forests. Importantly, while younger trees of the same diameter class have an average current AGBI rate that is ∼225% higher than older trees (300–400 years), we find trees that have been suppressed for up to 120 years can respond vigorously when competition is reduced. On average, post disturbance AGBI was ∼400% greater than pre-disturbance AGBI. Growth of suppressed trees, independent of their age, followed similar trajectories after canopy accession. While aboveground biomass generally increased through time, the time since disturbance and disturbance severity are important co-predictors for stand-level AGBI and AGB. These forests regained most of the above ground living biomass over short interval (∼50 years) after low intensity disturbances. The highest stand-level living AGB was observed on plots that experienced >40% canopy removal 160–190 years ago, whereas the highest AGBI occurred in plots disturbed recently within the past 40–50 years. Our results emphasize the importance of including both individual tree age and disturbance legacies to accurately characterize biomass dynamics and trajectories in forest ecosystems. Importantly, the period of time that a tree is in the canopy, and not tree age, modulates the trajectory of tree level AGBI. Growth rates begin to decline after ∼30 years (tree-rings width) and ∼100 years (AGBI) in the canopy. We demonstrate that even late-seral forests can rapidly regain biomass lost to low intensity disturbance.

Understanding the processes shaping the composition of assemblages in response to disturbance events is crucial for preventing ongoing biodiversity loss in forest ecosystems. However, studies of forest biodiversity responses to disturbance typically analyze immediate or short-term impacts only, while studies relating long-term disturbance history to biodiversity assemblage dynamics are rare. To address this important knowledge gap, we used a dendroecological approach to link natural disturbance history of 250 years (1750–2000) to structural habitat elements and, in turn, to breeding bird assemblages. We used data collected in 2017 and 2018 from 58 permanent study plots within 10 primary spruce forest stands distributed across the Western Carpathian Mountains of Europe. This dataset contained breeding bird counts and environmental variables describing forest density, tree diameter distribution, tree height, tree microhabitats, deadwood quantity and quality, and regeneration. Bird assemblages were significantly influenced by forest structure which was in turn shaped by disturbance dynamics (disturbance frequency, time since the last disturbance and its severity). Early successional species associated with more open habitats were positively influenced by disturbance-related structure (i.e. deadwood-related variables, canopy cover), while some species responded negatively. At the same time, overall abundance, species richness and Shannon diversity of the bird assemblage remained unchanged under variable disturbance histories. Our results support a view of primary spruce forests as a highly dynamic ecosystem, harbouring populations of bird species at all stages of succession despite significant structural changes and shifting patch mosaics over time due to natural disturbances.

Natural disturbances are key factors in the formation of forest ecosystem structure. Evaluation of the spatial and temporal extent of disturbance regimes is critical for understanding forest dynamics, forest structural heterogeneity, and biodiversity habitats. Quantifying disturbance regimes is therefore imperative for appropriate management of forests and protected areas. However, natural disturbance regimes have rarely been assessed using dendrochronological methods on a regional scale across primary mixed beech-fir forest stands - one of the dominant forest vegetation types in Europe. To study the natural disturbance regimes of beech-dominated mixed-forest stands, we established 42 permanent study plots with an area of 0.1 ha across three primary forest stands in the Western Carpathians, a region that still contains large areas of primary forest. We reconstructed each stand-level disturbance history using a tree-ring based approach. The temporal synchronicity of disturbance events was then evaluated by delineating stand-level disturbance events using a kernel density function, and through the detection of plot-level disturbances with severities greater than 10 percent. The results obtained from the chronologies showed substantial variability in time and space, especially in the mid-19th century. Low- and moderate-severity plot-level disturbance events were most common, but high- and extremely high-severity plot-level disturbance events also occurred. The observed spatial and temporal variability suggests that the beech-dominated forests were primarily driven by mixed-severity disturbance regimes, with windstorms as the main disturbance agent. This reconstruction of the disturbance regime provided unique insight into the scale of mortality processes in these beech-dominated mixed forests. This information can help guide ecological forestry in areas where both wood production and biodiversity preservation are simultaneous goals, such as by employing more spatiotemporally-complex silvicultural systems that resemble natural disturbance patterns and facilitate heterogeneous forest structures.

Soil vegetation atmosphere transport (SVAT) models are important for the quantification of water fluxes, soil water availability, drought stress and their uncertainties under climate change. We present LWFBrook90R , an enhanced implementation of the well-established, process-based SVAT model LWF-Brook90 for the R environment for statistical computing. The package provides new functions and sub-models for model parameterization, and facilitates parallel computing, sensitivity analysis and inverse calibration of the model. A case study comprising i) basic forward water balance simulations for temperate grassland vegetation, deciduous and evergreen forest, ii) a parallelized sensitivity analysis, and iii) Bayesian calibrations based on soil water storage observed in a poplar (Populus nigra × P. maximowiczii) Short Rotation Forest (SRF) demonstrates the utility of the R package. The sensitivity analysis revealed parameters affecting plant-available soil water storage capacity and the vegetation's timing and level of water demand to be most important for the annual course of simulated soil water storage, with seasonal and interannual differences in parameter importance rankings. The subsequent calibration yielded a very high agreement between daily simulated and observed soil water storage (0-200 cm soil depth) for the calibration and validation datasets, with Nash-Sutcliffe efficiencies of 0.97 and 0.95, respectively. The final model predicted high though realistic rates of annual evapotranspiration (2011: 844 ± 3.8 mm y-1, 2012: 733 ± 4.5 mm y-1) for the poplar SRF, regularly exceeding grass reference evaporation (ET0) by 20-47% during the months of the growing season. However, basing calibrations solely on observed soil water storage probably resulted in biased partitioning of evapotranspiration towards interception losses. The integration of the LWF-Brook90 hydrological model into R with its wide variety of extensions was successfully tested and may provide efficient, reliable and reproducible water balance predictions by facilitating complex statistical analyses and large-scale applications of the model.

While shifting disturbance rates and climate change have major implications for the structure of contemporary forests through their effects on adult tree mortality, the responses of regenerating trees to disturbances and environmental variation will ultimately determine the structure and functioning of forests in the future. Assessing the resilience of forests to changing conditions requires information on what constrains tree performance during recruitment and whether recruitment dynamics have changed throughout history. We analyzed growth patterns in a large sample of tree cores (n = 14 793) collected from primary Picea forests throughout the Carpathian Mountains. Growth rate anomalies recorded in tree-rings permitted the reconstruction of several key recruitment and disturbance parameters: (1) whether individuals were recruited after a period of competitive suppression (Released Trees; RT; 66% of trees) or immediately following gap formation (Gap Recruited Trees; GRT; 33%), (2) growth rates during recruitment, (3) the duration of recruitment and (4) historical disturbance severity variation. High neighborhood density led to lower growth rates in RTs, but favored a higher growth rate in GRTs. Winter temperatures were positively correlated with Picea growth during recruitment, GRTs were also more sensitive to winter precipitation. Recent increases in growth during recruitment and reductions in recruitment intervals suggest that rates of canopy replacement have increased over recent decades. Assessments of forest resilience must recognize that constraints on tree growth differ during recruitment and interact with disturbance severity. An individual's experience prior to competitive release and factors altering the immediate abiotic conditions of a recruiting individual (competition and disturbance severity) are important determinants of canopy replacement rates; these recruitment parameters will certainly interact with shifting disturbance regimes. Ultimately, increasing growth rates and decreasing recruitment intervals suggest that forest dynamics are accelerating, and are potentially compensating for recent increases in tree mortality rates.

FB acknowledges funding from the Swiss National Science Foundation (P300P2_154543) and the EU Horizon-2020 project “BACI” (grant 640176). SK, FJ, RS and DCF are supported by the SNF iTREE sinergia project 136295. FJ, SL, and RS acknowledge support by the Swiss National Science Foundation (#200020_172476), OB acknowledges funding from UEFISCDI project PN-II-ID-PCE-2011-3-07 and VT is supported by the GACR 15-14840S and CIGA 20154316. The response of forest growth to climate variability varies along environmental gradients. A growth increase and decrease with warming is usually observed in cold‐humid and warm‐dry regions, respectively. However, it remains poorly known where the sign of these temperature effects switches. Here we introduce a newly developed European tree‐ring network that has been specifically collected to reconstruct forest aboveground biomass increment (ABI). We quantify, how the long‐term (1910‐2009) inter‐annual variability of ABI depends on local mean May‐August temperature and test, if a dynamic global vegetation model (DGVM) ensemble reflects the resulting patterns. We find that sites at 8°C mean May‐August temperature increase ABI on average by 5.7 ± 1.3 %, whereas sites at 20°C decrease ABI by 3.0±1.8 % m‐2 y‐1 ∆°C‐1. A threshold temperature between beneficial and detrimental effects of warming and the associated increase in water demand on tree growth emerged at 15.9 ± 1.4°C mean May‐August temperature. Because inter‐annual variability increases proportionally with mean growth rate – i.e. the coefficient of variation stays constant – we were able to validate these findings with a much larger tree‐ring dataset that had been established following classic dendrochronological sampling schemes. While the observed climate sensitivity pattern is well reflected in the DGVM ensemble, there is a large spread of threshold temperatures between the individual models. Also, individual models disagree strongly on the magnitude of climate impact at the coldest and warmest locations, suggesting where model improvement is most needed to more accurately predict forest growth and effectively guide silvicultural practices. Publisher PDF

In a recent opinion paper, Schulze et al. (2014) compared an-imal and plant richness of protected areas with intensivelyand extensively managed forests in Germany and in Roma-nia. Schulze et al. hypothesised that “differences in manage-ment practice have contributed to the observed differences indiversity of central versus southeastern Europe”. The studyconcludes with a hypothesis that “clear-felling followed bynatural succession may even be superior to the protectionof old-growth forests, regarding biodiversity”. We wish tocontinue the dialog on this important topic, and express oursomewhat different perspective on the conclusions and im-plications of this particular paper (Schulze et al., 2014). Weagree in principal that emulating natural disturbance regimesand creating openings via clear-fellings can be one of sev-eral management tools to introduce heterogeneity in other-wise homogenous mid-seral stands that have resulted frompast management (Franklin and Johnson, 2012). However,we believe it is misleading to compare clear-fellings to pro-tected areas dominated by old-growth or primary forests us-ing a simplistic measure of biodiversity and without a land-scape perspective on the role of different habitat types (suc-cessional stages) to sustain biodiversity over time and space.We identify some critical limitations of the original opinionpaper and offer an alternative perspective to the authors’ con-clusions regarding protected forests.We highlight three major issues of the opinion paper ofSchulze et al. (2014): limited scope, an incomplete concep-tual framework, and conjectural interpretations unsupportedby solid data.

Stands and trees may exhibit different climate–growth responses compared to neighbouring forests and individuals. The study of these differences is crucial to understanding the effects of climate change on the growth and vulnerability of forests and trees. In this research we analyse the growth responsiveness of primary Norway spruce forests to climate as a function of different stand (elevation, aspect, slope, crowding, historic disturbance regime) and tree (age, tree-to-tree competition) features in the Romanian Carpathians. Climate–growth relationships were analysed using Pearson correlation coefficients between ring-width indices (RWIs) and climate variables. The influence of stand and tree characteristics on the RWI responses to climate were investigated using linear mixed-effects models. Elevation greatly modulated the climate–growth associations and it frequently interacted with competition intensity or tree age to differentially influence growth responsiveness to climate. Old trees were more sensitive to climate than young trees, but while old tree’s response to climate highly depended on elevation (e.g. positive influence of summer temperature on old trees’ RWIs at high elevations, but negative effect at low elevations), differences of the young trees’ response across the elevation gradient were less evident. The severity of the past disturbance also modified the climate–growth associations because of contrasting canopy structures. Our results suggest that although an increase in temperature might enhance growth at high elevations, it may also induce growth declines due to drought stress at lower elevations, particularly for old trees or trees growing under high levels of competition, which may increase their vulnerability to disturbances.

The retrospective study of abrupt and sustained increases in the radial growth of trees (hereinafter ‘releases’) by tree-ring analysis is an approach widely used for reconstructing past forest disturbances. Despite the range of dendrochronological methods used for release-detection, a lack of in-depth comparison between them can lead researchers to question which method to use and, potentially, increases the uncertainties of disturbance histories derived with different methods. Here, we investigate the efficacy and sensitivity of four widely used release detection methods using tree-ring width series and complete long-term inventories of forest stands with known disturbances. We used support vector machine (SVM) analysis trained on long-term forest census data to estimate the likelihood that Acer rubrum trees experiencing reductions in competition show releases in their tree-ring widths. We compare methods performance at the tree and stand level, followed by evaluation of method sensitivity to changes in their parameters and settings. Disturbance detection methods agreed with 60–76% of the SVM-identified growth releases under high canopy disturbance and 80–94% in a forest with canopy disturbance of low severity and frequency. The median competition index change (CIC) of trees identified as being released differed more than two-fold between methods, from −0.33 (radial-growth averaging) to −0.68 (time-series). False positives (type I error) were more common in forests with low severity disturbance, whereas false negatives (type II error) occurred more often in forests with high severity disturbance. Sensitivity analysis indicated that reductions of the detection threshold and the length of the time window significantly increased detected stand-level disturbance severity across all methods. Radial-growth averaging and absolute-increase methods had lower levels of type I and II error in detecting disturbance events with our datasets. Parameter settings play a key role in the accuracy of reconstructing disturbance history regardless of the method. Time-series and radial-growth averaging methods require the least amount of a priori information, but only the time-series method quantified the subsequent growth increment related to a reduction in competition. Finally, we recommend yearly binning of releases using a kernel density estimation function to identify local maxima indicating disturbance. Kernel density estimation improves reconstructions of forest history and, thus, will further our understanding of past forest dynamics.