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Radial tree growth is sensitive to environmental conditions, making observed growth increments an important indicator of climate change effects on forest growth. However, unprecedented climate variability could lead to non‐stationarity, that is, a decoupling of tree growth responses from climate over time, potentially inducing biases in climate reconstructions and forest growth projections. Little is known about whether and to what extent environmental conditions, species, and model type and resolution affect the occurrence and magnitude of non‐stationarity. To systematically assess potential drivers of non‐stationarity, we compiled tree‐ring width chronologies of two conifer species, Picea abies and Pinus sylvestris, distributed across cold, dry, and mixed climates. We analyzed 147 sites across the Europe including the distribution margins of these species as well as moderate sites. We calibrated four numerical models (linear vs. non‐linear, daily vs. monthly resolution) to simulate growth chronologies based on temperature and soil moisture data. Climate–growth models were tested in independent verification periods to quantify their non‐stationarity, which was assessed based on bootstrapped transfer function stability tests. The degree of non‐stationarity varied between species, site climatic conditions, and models. Chronologies of P. sylvestris showed stronger non‐stationarity compared with Picea abies stands with a high degree of stationarity. Sites with mixed climatic signals were most affected by non‐stationarity compared with sites sampled at cold and dry species distribution margins. Moreover, linear models with daily resolution exhibited greater non‐stationarity compared with monthly‐resolved non‐linear models. We conclude that non‐stationarity in climate–growth responses is a multifactorial phenomenon driven by the interaction of site climatic conditions, tree species, and methodological features of the modeling approach. Given the existence of multiple drivers and the frequent occurrence of non‐stationarity, we recommend that temporal non‐stationarity rather than stationarity should be considered as the baseline model of climate–growth response for temperate forests. [ABSTRACT FROM AUTHOR]

With ongoing global warming, increasing water deficits promote physiological stress on forest ecosystems with negative impacts on tree growth, vitality, and survival. How individual tree species will react to increased drought stress is therefore a key research question to address for carbon accounting and the development of climate change mitigation strategies. Recent tree‐ring studies have shown that trees at higher latitudes will benefit from warmer temperatures, yet this is likely highly species‐dependent and less well‐known for more temperate tree species. Using a unique pan‐European tree‐ring network of 26,430 European beech (Fagus sylvatica L.) trees from 2118 sites, we applied a linear mixed‐effects modeling framework to (i) explain variation in climate‐dependent growth and (ii) project growth for the near future (2021–2050) across the entire distribution of beech. We modeled the spatial pattern of radial growth responses to annually varying climate as a function of mean climate conditions (mean annual temperature, mean annual climatic water balance, and continentality). Over the calibration period (1952–2011), the model yielded high regional explanatory power (R2 = 0.38–0.72). Considering a moderate climate change scenario (CMIP6 SSP2‐4.5), beech growth is projected to decrease in the future across most of its distribution range. In particular, projected growth decreases by 12%–18% (interquartile range) in northwestern Central Europe and by 11%–21% in the Mediterranean region. In contrast, climate‐driven growth increases are limited to around 13% of the current occurrence, where the historical mean annual temperature was below ~6°C. More specifically, the model predicts a 3%–24% growth increase in the high‐elevation clusters of the Alps and Carpathian Arc. Notably, we find little potential for future growth increases (−10 to +2%) at the poleward leading edge in southern Scandinavia. Because in this region beech growth is found to be primarily water‐limited, a northward shift in its distributional range will be constrained by water availability. [ABSTRACT FROM AUTHOR]

Temperate forests are undergoing significant transformations due to the influence of climate change, including varying responses of different tree species to increasing temperature and drought severity. To comprehensively understand the full range of growth responses, representative datasets spanning extensive site and climatic gradients are essential. This study utilizes tree‐ring data from 550 sites from the temperate forests of Czechia to assess growth trends of six dominant Central European tree species (European beech, Norway spruce, Scots pine, silver fir, sessile and pedunculate oak) over 1990–2014. By modeling mean growth series for each species and site, and employing principal component analysis, we identified the predominant growth trends. Over the study period, linear growth trends were evident across most sites (56% increasing, 32% decreasing, and 10% neutral). The proportion of sites with stationary positive trends increased from low toward high elevations, whereas the opposite was true for the stationary negative trends. Notably, within the middle range of their distribution (between 500 and 700 m a.s.l.), Norway spruce and European beech exhibited a mix of positive and negative growth trends. While Scots pine growth trends showed no clear elevation‐based pattern, silver fir and oaks displayed consistent positive growth trends regardless of site elevation, indicating resilience to the ongoing warming. We demonstrate divergent growth trajectories across space and among species. These findings are particularly important as recent warming has triggered a gradual shift in the elevation range of optimal growth conditions for most tree species and has also led to a decoupling of growth trends between lowlands and mountain areas. As a result, further future shifts in the elevation range and changes in species diversity of European temperate forests can be expected. [ABSTRACT FROM AUTHOR]

Assessing the impacts of natural disturbance on the functioning of complex forest systems are imperative in the context of global change. The unprecedented rate of contemporary species extirpations, coupled with widely held expectations that future disturbance intensity will increase with warming, highlights a need to better understand how natural processes structure habitat availability in forest ecosystems. Standardised typologies of tree-related microhabitats (TreMs) have been developed to facilitate assessments of resource availability for multiple taxa. However, natural disturbance effects on TreM diversity have never been assessed. We amassed a comprehensive dataset of TreM occurrences and a concomitant 300-year disturbance history reconstruction that spanned large environmental gradients in temperate primary forests. We used nonlinear analyses to quantify relations between past disturbance parameters and contemporary patterns of TreM occurrence. Our results reveal that natural forest dynamics, characterised by fluctuating disturbance intervals and variable severity levels, maintained structurally complex landscapes rich in TreMs. Different microhabitat types developed over time in response to divergent disturbance histories. The relative abundance of alternate TreMs was maximised by unique interactions between past disturbance severity and elapsed time. Despite an unequal distribution of individual TreMs, total microhabitat diversity was maintained at constant levels, suggesting that spatially heterogeneous disturbances maintained a shifting mosaic of habitat types over the region as a whole. Our findings underscore the fundamental role of natural processes in promoting conditions that maximise biodiversity potential. Strict conservation and management systems that preserve natural disturbance outcomes, including associated biological legacies, may therefore safeguard biodiversity at large scales. [ABSTRACT FROM AUTHOR]

Protecting structural features, such as tree‐related microhabitats (TreMs), is a cost‐effective tool crucial for biodiversity conservation applicable to large forested landscapes. Although the development of TreMs is influenced by tree diameter, species, and vitality, the relationships between tree age and TreM profile remain poorly understood. Using a tree‐ring‐based approach and a large data set of 8038 trees, we modeled the effects of tree age, diameter, and site characteristics on TreM richness and occurrence across some of the most intact primary temperate forests in Europe, including mixed beech and spruce forests. We observed an overall increase in TreM richness on old and large trees in both forest types. The occurrence of specific TreM groups was variably related to tree age and diameter, but some TreM groups (e.g., epiphytes) had a stronger positive relationship with tree species and elevation. Although many TreM groups were positively associated with tree age and diameter, only two TreM groups in spruce stands reacted exclusively to tree age (insect galleries and exposed sapwood) without responding to diameter. Thus, the retention of trees for conservation purposes based on tree diameter appears to be a generally feasible approach with a rather low risk of underrepresentation of TreMs. Because greater tree age and diameter positively affected TreM development, placing a greater emphasis on conserving large trees and allowing them to reach older ages, for example, through the establishment of conservation reserves, would better maintain the continuity of TreM resource and associated biodiversity. However, this approach may be difficult due to the widespread intensification of forest management and global climate change. [ABSTRACT FROM AUTHOR]

Extreme tree growth reductions represent events of abrupt forest productivity decline and carbon sequestration reduction. An increase in their magnitude can represent an early warning signal of impending tree mortality. Yet the long‐term trends in extreme growth reductions remain largely unknown. We analyzed the trends in the proportion of trees exhibiting extreme growth reductions in two Central‐European conifer species—Pinus sylvestris (PISY) and Picea abies (PCAB)—between 1901 and 2018. We used a novel approach for extreme growth reduction quantification by relating their size to their mean recurrence interval. Twenty‐eight sites throughout Czechia and Slovakia with 1120 ring width series representing high‐ and low‐elevation forests were inspected for extreme growth reductions with recurrence intervals of 15 and 50 years along with their link to climatic drivers. Our results show the greatest growth reductions at low‐elevation PCAB sites, indicating high vulnerability of PCAB to drought. The proportions of trees exhibiting extreme growth reductions increased over time at low‐elevation PCAB, decreased recently following an abrupt increase in the 1970–1980s at high‐elevation PCAB, and showed nonsignificant trends in high‐ and low‐elevation PISY. Climatic drivers of extreme growth reductions, however, shifted over time for all site categories as the proportion of low‐temperature‐induced extreme growth reductions declined since the 1990s, whereas events caused by drought consistently increased in frequency during the same period. We observed higher growth volatility at the lower range of distribution compared with the upper range margin of PISY and PCAB. This will undoubtedly considerably impact tree growth and vitality as temperatures and incidence of drought in Central Europe are expected to further increase with ongoing climate change. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Aim: Natural disturbances influence forest structure, successional dynamics and consequently, the distribution of species through time and space. We quantified the long‐term impacts of natural disturbances on lichen species richness and composition in primary mountain forests, with a particular focus on the occurrence of endangered species. Location: Ten primary mountain spruce forest stands across five mountain chains of the Western Carpathians, a European hotspot of biodiversity. Methods: Living trees, snags and downed logs were surveyed for epiphytic and epixylic lichens in 57 plots. Using reconstructed disturbance history, we tested how lichen species richness and composition was affected by the current forest structure and disturbance regimes in the past 250 years. We also examined differences in community composition among discrete microhabitats. Results: Dead standing trees as biological legacies of natural disturbances promoted lichen species richness and the occurrence of threatened species at the plot scale, suggesting improved growing conditions for rare and common lichens during the early stages of recovery post disturbance. However, high‐severity disturbances compromised plot‐scale species richness. Both species richness and the number of old‐growth specialists increased with time since disturbance (i.e., long‐term uninterrupted succession). No lichen species was strictly dependent on live trees as a habitat, but numerous species showed specificity to logs, standing objects or an admixture of tree species. Conclusions: Lichen species richness was lower in regenerating, young and uniform plots compared with overmature and recently disturbed areas. Natural forest dynamics and its legacies are critical to the diversity and species composition of lichens. Spatio‐temporal consequences of natural dynamics require a sufficient area of protected forests for provisioning continual habitat variability at the landscape scale. Ongoing climatic changes may further accentuate this necessity. Hence, we highlighted the need to protect the last remaining primary forests to ensure the survival of regionally unique species pools of lichens. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

• Substantial growth variance of spruce trees in the Carpathian arc is non-climatic. • Primary Carpathian Spruce forests exhibit mixed and asymptotic climate response. • Southern Carpathian spruce forests are distinct from Northern tracts in their response. 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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Abstract: Determining the drivers of shifting forest disturbance rates remains a pressing global change issue. Large‐scale forest dynamics are commonly assumed to be climate driven, but appropriately scaled disturbance histories are rarely available to assess how disturbance legacies alter subsequent disturbance rates and the climate sensitivity of disturbance. We compiled multiple tree ring‐based disturbance histories from primary Picea abies forest fragments distributed throughout five European landscapes spanning the Bohemian Forest and the Carpathian Mountains. The regional chronology includes 11,595 tree cores, with ring dates spanning the years 1750–2000, collected from 560 inventory plots in 37 stands distributed across a 1,000 km geographic gradient, amounting to the largest disturbance chronology yet constructed in Europe. Decadal disturbance rates varied significantly through time and declined after 1920, resulting in widespread increases in canopy tree age. Approximately 75% of current canopy area recruited prior to 1900. Long‐term disturbance patterns were compared to an historical drought reconstruction, and further linked to spatial variation in stand structure and contemporary disturbance patterns derived from LANDSAT imagery. Historically, decadal Palmer drought severity index minima corresponded to higher rates of canopy removal. The severity of contemporary disturbances increased with each stand's estimated time since last major disturbance, increased with mean diameter, and declined with increasing within‐stand structural variability. Reconstructed spatial patterns suggest that high small‐scale structural variability has historically acted to reduce large‐scale susceptibility and climate sensitivity of disturbance. Reduced disturbance rates since 1920, a potential legacy of high 19th century disturbance rates, have contributed to a recent region‐wide increase in disturbance susceptibility. Increasingly common high‐severity disturbances throughout primary Picea forests of Central Europe should be reinterpreted in light of both legacy effects (resulting in increased susceptibility) and climate change (resulting in increased exposure to extreme events). [ABSTRACT FROM AUTHOR]

Questions How do canopy-understorey interactions respond to variation in disturbance severity over extended periods of time? For forests with different disturbance histories, do light availability and understorey cohort densities converge towards a common old-growth structure, or do historical legacies influence populations indefinitely? Locations Remnants of primary spruce ( Picea abies (L.) Karst.) forests throughout Germany, Slovakia, Ukraine and Romania. Methods A disturbance history of >200 yr was reconstructed from 11 278 tree cores collected from forest plots ( n = 520). Understorey tree densities of two size classes and hemispherical photo-based light availabilities were inventoried and modelled as functions of the severity of the main disturbance and time since the event. Results Variation in understorey tree densities had a hump-shaped distribution through time. Stem densities were approximately static in the least disturbed sites, and declined in relation to disturbance severity over approx. 100 yr. Similar to patterns of stem densities, initially high understorey light availability also reached a minimum at 100 yr, which indicated crown closure. Following this, light availability and stem densities both increased as stands transitioned from stem exclusion to understorey re-initiation. The effect of disturbance severity on understorey densities and patchiness in light availability persisted for >200 yr. Conclusions Long-term trends in canopy-understorey interactions validate current conceptual models of forest development. Furthermore, we empirically validate that these conceptual models generalized over gradients in disturbance severity. Higher disturbance sites exhibited a more even-aged character with more pronounced periods of stem exclusion, canopy closure and understorey re-initiation; forests with low-severity disturbance histories yielded a more stationary uneven-aged structure. The model identified the extent of variation in disturbance severity within which these P. abies forests are able to regenerate and retain their monospecific character, which is increasingly relevant as disturbance regimes continue to shift under global climate change. [ABSTRACT FROM AUTHOR]

Tree mortality is a key factor influencing forest functions and dynamics, but our understanding of the mechanisms leading to mortality and the associated changes in tree growth rates are still limited. We compiled a new pan-continental tree-ring width database from sites where both dead and living trees were sampled (2970 dead and 4224 living trees from 190 sites, including 36 species), and compared early and recent growth rates between trees that died and those that survived a given mortality event. We observed a decrease in radial growth before death in ca. 84% of the mortality events. The extent and duration of these reductions were highly variable (1-100 years in 96% of events) due to the complex interactions among study species and the source(s) of mortality. Strong and long-lasting declines were found for gymnosperms, shade- and drought-tolerant species, and trees that died from competition. Angiosperms and trees that died due to biotic attacks (especially bark-beetles) typically showed relatively small and short-term growth reductions. Our analysis did not highlight any universal trade-off between early growth and tree longevity within a species, although this result may also reflect high variability in sampling design among sites. The intersite and interspecific variability in growth patterns before mortality provides valuable information on the nature of the mortality process, which is consistent with our understanding of the physiological mechanisms leading to mortality. Abrupt changes in growth immediately before death can be associated with generalized hydraulic failure and/or bark-beetle attack, while long-term decrease in growth may be associated with a gradual decline in hydraulic performance coupled with depletion in carbon reserves. Our results imply that growth-based mortality algorithms may be a powerful tool for predicting gymnosperm mortality induced by chronic stress, but not necessarily so for angiosperms and in case of intense drought or bark-beetle outbreaks. [ABSTRACT FROM AUTHOR]

Background: Severe canopy-removing disturbances are native to many temperate forests and radically alter stand structure, but biotic legacies (surviving elements or patterns) can lend continuity to ecosystem function after such events. Poorly understood is the degree to which the structural complexity of an old-growth forest carries over to the next stand. We asked how pre-disturbance spatial pattern acts as a legacy to influence post-disturbance stand structure, and how this legacy influences the structural diversity within the early-seral stand. Methods: Two stem-mapped one-hectare forest plots in the Czech Republic experienced a severe bark beetle outbreak, thus providing before-and-after data on spatial patterns in live and dead trees, crown projections, down logs, and herb cover. Results: Post-disturbance stands were dominated by an advanced regeneration layer present before the disturbance. Both major species, Norway spruce (Picea abies) and rowan (Sorbus aucuparia), were strongly self-aggregated and also clustered to former canopy trees, pre-disturbance snags, stumps and logs, suggesting positive overstory to understory neighbourhood effects. Thus, although the disturbance dramatically reduced the stand’s height profile with ~100% mortality of the canopy layer, the spatial structure of post-disturbance stands still closely reflected the pre-disturbance structure. The former upper tree layer influenced advanced regeneration through microsite and light limitation. Under formerly dense canopies, regeneration density was high but relatively homogeneous in height; while in former small gaps with greater herb cover, regeneration density was lower but with greater heterogeneity in heights. Conclusion: These findings suggest that pre-disturbance spatial patterns of forests can persist through severe canopy-removing disturbance, and determine the spatial structure of the succeeding stand. Such patterns constitute a subtle but key legacy effect, promoting structural complexity in early-seral forests as well as variable successional pathways and rates. This influence suggests a continuity in spatial ecosystem structure that may well persist through multiple forest generations. [ABSTRACT FROM AUTHOR]

Questions How have the historical frequency and severity of natural disturbances in primary Picea abies forests varied at the forest stand and landscape level during recent centuries? Is there a relationship between physiographic attributes and historical patterns of disturbance severity in this system? Location Primary P. abies forests of the Eastern Carpathian Mountains, Romania; a region thought to hold the largest concentration of primary P. abies forests in Europe's temperate zone. Methods We used dendrochronological methods applied to many plots over a large area (132 plots representing six stands in two landscapes), thereby providing information at both stand and landscape levels. Evidence of past canopy disturbance was derived from two patterns of radial growth: (1) abrupt, sustained increases in growth (releases) and (2) rapid early growth rates (gap recruitment). These methods were augmented with non-metric multidimensional scaling to facilitate the interpretation of factors influencing past disturbance. Results Of the two growth pattern criteria used to assess past disturbance, gap recruitment was the most common, representing 80% of disturbance evidence overall. Disturbance severities varied over the landscape, including stand-replacing events, as well as low- and intermediate-severity disturbances. More than half of the study plots experienced extreme-severity disturbances at the plot level, although they were not always synchronized across stands and landscapes. Plots indicating high-severity disturbances were often spatially clustered (indicating disturbances up to 20 ha), while this tendency was less clear for low- and moderate-severity disturbances. Physiographic attributes such as altitude and land form were only weakly correlated with disturbance severity. Historical documents suggest windstorms as the primary disturbance agent, while the role of bark beetles ( Ips typographus) remains unclear. Conclusions The historical disturbance regime revealed in this multi-scale study is characterized by considerable spatial and temporal heterogeneity, which could be seen among plots within stands, among stands within landscapes and between the two landscapes. When the disturbance regime was evaluated at these larger scales, the entire range of disturbance severity was revealed within this landscape. [ABSTRACT FROM AUTHOR]

Highlights: [•] Forest management has negatively affected biodiversity in northern European forests. [•] Ecological restoration has been introduced to safeguard forest biodiversity. [•] Feasible restoration actions should be conducted in well-known target ecosystems. [•] Successful restoration must maintain connectivity and assess functionality. [•] Restoration must cope with uncertainty and manage conflicts/restrictions. [Copyright &y& Elsevier]