Major tree species of Central European forests differ in their proportion of positive, negative, and nonstationary growth trends.

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.

Ecological and methodological drivers of non-stationarity in tree growth response to climate.

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.

Large old trees increase growth under shifting climatic constraints: Aligning tree longevity and individual growth dynamics in primary mountain spruce forests.

In a world of accelerating changes in environmental conditions driving tree growth, tradeoffs between tree growth rate and longevity could curtail the abundance of large old trees (LOTs), with potentially dire consequences for biodiversity and carbon storage. However, the influence of tree-level tradeoffs on forest structure at landscape scales will also depend on disturbances, which shape tree size and age distribution, and on whether LOTs can benefit from improved growing conditions due to climate warming. We analyzed temporal and spatial variation in radial growth patterns from ~5000 Norway spruce (Picea abies [L.] H. Karst) live and dead trees from the Western Carpathian primary spruce forest stands. We applied mixed-linear modeling to quantify the importance of LOT growth histories and stand dynamics (i.e., competition and disturbance factors) on lifespan. Finally, we assessed regional synchronization in radial growth variability over the 20th century, and modeled the effects of stand dynamics and climate on LOTs recent growth trends. Tree age varied considerably among forest stands, implying an important role of disturbance as an age constraint. Slow juvenile growth and longer period of suppressed growth prolonged tree lifespan, while increasing disturbance severity and shorter time since last disturbance decreased it. The highest age was not achieved only by trees with continuous slow growth, but those with slow juvenile growth followed by subsequent growth releases. Growth trend analysis demonstrated an increase in absolute growth rates in response to climate warming, with late summer temperatures driving the recent growth trend. Contrary to our expectation that LOTs would eventually exhibit declining growth rates, the oldest LOTs (>400 years) continuously increase growth throughout their lives, indicating a high phenotypic plasticity of LOTs for increasing biomass, and a strong carbon sink role of primary spruce forests under rising temperatures, intensifying droughts, and increasing bark beetle outbreaks.

Importance of conserving large and old trees to continuity of tree-related microhabitats.

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.

Importancia de conservar los árboles viejos y grandes para la continuidad de los microhábitats relacionados Resumen La protección de las características estructurales, como los microhábitats relacionados a los árboles (MhAr), es una herramienta económica importante para la conservación de la biodiversidad que puede aplicarse en los paisajes boscosos extensos. Aunque el diámetro, especie y vitalidad del árbol influyen sobre el desarrollo de los MhAr, todavía se sabe poco sobre las relaciones entre la edad del árbol y el perfil. Modelamos los efectos de la edad y diámetro del árbol y las características del sitio sobre la riqueza y presencia de los MhAr en algunos de los bosques primarios más preservados de Europa, incluyendo los bosques mixtos de hayas y abetos, con una estrategia basada en los anillos de crecimiento y un conjunto con datos de 8038 árboles. Observamos un incremento generalizado en la riqueza de MhAr en los árboles viejos y grandes en ambos tipos de bosques. La presencia de grupos específicos de MhAr tuvo una relación variada con el diámetro y la edad del árbol, aunque algunos grupos de MhAr (p. ej.: epífitas) tuvieron una relación positiva más fuerte con la elevación y la especie del árbol. Mientras que muchos grupos de MhAr estuvieron asociados positivamente con la edad y diámetro del árbol, sólo dos grupos de MhAr en los abetos reaccionaron exclusivamente a la edad del árbol (galerías de insectos y savia expuesta) sin responder al diámetro. Por lo tanto, la retención de los árboles con fines de conservación basada en los diámetros parece ser una estrategia plausible con un riesgo bajo de subrepresentación de los MhAr. Ya que a mayor edad y diámetro del árbol hubo efectos positivos en el desarrollo de los MhAr, poner un mayor énfasis sobre la conservación de los árboles grandes y permitirles alcanzar una edad mayor, por ejemplo, a través del establecimiento de reservas de conservación, mantendría de mejor manera la continuidad del MhAr y de la biodiversidad asociada. Sin embargo, esta estrategia puede ser complicada debido a la intensificación generalizada de la gestión forestal y el cambio climático mundial.

Late-spring frost risk between 1959 and 2017 decreased in North America but increased in Europe and Asia.

Late-spring frosts (LSFs) affect the performance of plants and animals across the world's temperate and boreal zones, but despite their ecological and economic impact on agriculture and forestry, the geographic distribution and evolutionary impact of these frost events are poorly understood. Here, we analyze LSFs between 1959 and 2017 and the resistance strategies of Northern Hemisphere woody species to infer trees' adaptations for minimizing frost damage to their leaves and to forecast forest vulnerability under the ongoing changes in frost frequencies. Trait values on leaf-out and leaf-freezing resistance come from up to 1,500 temperate and boreal woody species cultivated in common gardens. We find that areas in which LSFs are common, such as eastern North America, harbor tree species with cautious (late-leafing) leaf-out strategies. Areas in which LSFs used to be unlikely, such as broad-leaved forests and shrublands in Europe and Asia, instead harbor opportunistic tree species (quickly reacting to warming air temperatures). LSFs in the latter regions are currently increasing, and given species' innate resistance strategies, we estimate that ∼35% of the European and ∼26% of the Asian temperate forest area, but only ∼10% of the North American, will experience increasing late-frost damage in the future. Our findings reveal region-specific changes in the spring-frost risk that can inform decision-making in land management, forestry, agriculture, and insurance policy.

Trends in climatically driven extreme growth reductions of Picea abies and Pinus sylvestris in Central Europe.

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.

Natural disturbance regimes as a guide for sustainable forest management in Europe.

In Europe, forest management has controlled forest dynamics to sustain commodity production over multiple centuries. Yet over-regulation for growth and yield diminishes resilience to environmental stress as well as threatens biodiversity, leading to increasing forest susceptibility to an array of disturbances. These trends have stimulated interest in alternative management systems, including natural dynamics silviculture (NDS). NDS aims to emulate natural disturbance dynamics at stand and landscape scales through silvicultural manipulations of forest structure and landscape patterns. We adapted a "Comparability Index" (CI) to assess convergence/divergence between natural disturbances and forest management effects. We extended the original CI concept based on disturbance size and frequency by adding the residual structure of canopy trees after a disturbance as a third dimension. We populated the model by compiling data on natural disturbance dynamics and management from 13 countries in Europe, covering four major forest types (i.e., spruce, beech, oak, and pine-dominated forests). We found that natural disturbances are highly variable in size, frequency, and residual structure, but European forest management fails to encompass this complexity. Silviculture in Europe is skewed toward even-aged systems, used predominately (72.9% of management) across the countries assessed. The residual structure proved crucial in the comparison of natural disturbances and silvicultural systems. CI indicated the highest congruence between uneven-aged silvicultural systems and key natural disturbance attributes. Even so, uneven-aged practices emulated only a portion of the complexity associated with natural disturbance effects. The remaining silvicultural systems perform poorly in terms of retention compared to tree survivorship after natural disturbances. We suggest that NDS can enrich Europe's portfolio of management systems, for example where wood production is not the primary objective. NDS is especially relevant to forests managed for habitat quality, risk reduction, and a variety of ecosystem services. We suggest a holistic approach integrating NDS with more conventional practices.

Natural disturbance impacts on trade-offs and co-benefits of forest biodiversity and carbon.

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.

SoilTemp: A global database of near-surface temperature.

Current analyses and predictions of spatially explicit patterns and processes in ecology most often rely on climate data interpolated from standardized weather stations. This interpolated climate data represents long-term average thermal conditions at coarse spatial resolutions only. Hence, many climate-forcing factors that operate at fine spatiotemporal resolutions are overlooked. This is particularly important in relation to effects of observation height (e.g. vegetation, snow and soil characteristics) and in habitats varying in their exposure to radiation, moisture and wind (e.g. topography, radiative forcing or cold-air pooling). Since organisms living close to the ground relate more strongly to these microclimatic conditions than to free-air temperatures, microclimatic ground and near-surface data are needed to provide realistic forecasts of the fate of such organisms under anthropogenic climate change, as well as of the functioning of the ecosystems they live in. To fill this critical gap, we highlight a call for temperature time series submissions to SoilTemp, a geospatial database initiative compiling soil and near-surface temperature data from all over the world. Currently, this database contains time series from 7,538 temperature sensors from 51 countries across all key biomes. The database will pave the way toward an improved global understanding of microclimate and bridge the gap between the available climate data and the climate at fine spatiotemporal resolutions relevant to most organisms and ecosystem processes.

Quantifying natural disturbances using a large-scale dendrochronological reconstruction to guide forest management.

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.

The climatic drivers of primary Picea forest growth along the Carpathian arc are changing under rising temperatures.

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.

Large-scale disturbance legacies and the climate sensitivity of primary Picea abies forests.

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).

EFO-LCI: A New Life Cycle Inventory Database of Forestry Operations in Europe.

Life cycle assessment (LCA) has become a common methodology to analyze environmental impacts of forestry systems. Although LCA has been widely applied to forestry since the 90s, the LCAs are still often based on generic Life Cycle Inventory (LCI). With the purpose of improving LCA practices in the forestry sector, we developed a European Life Cycle Inventory of Forestry Operations (EFO-LCI) and analyzed the available information to check if within the European forestry sector national differences really exist. We classified the European forests on the basis of "Forest Units" (combinations of tree species and silvicultural practices). For each Forest Unit, we constructed the LCI of their forest management practices on the basis of a questionnaire filled out by national silvicultural experts. We analyzed the data reported to evaluate how they vary over Europe and how they affect LCA results and made freely available the inventory data collected for future use. The study shows important variability in rotation length, type of regeneration, amount and assortments of wood products harvested, and machinery used due to the differences in management practices. The existing variability on these activities sensibly affect LCA results of forestry practices and raw wood production. Although it is practically unfeasible to collect site-specific data for all the LCAs involving forest-based products, the use of less generic LCI data of forestry practice is desirable to improve the reliability of the studies. With the release of EFO-LCI we made a step toward the construction of regionalized LCI for the European forestry sector.

Life and death of Picea abies after bark-beetle outbreak: ecological processes driving seedling recruitment.

The severity and spatial extent of bark-beetle outbreaks substantially increased in recent decades worldwide. The ongoing controversy about natural forest recovery after these outbreaks highlights the need for individual-based long-term studies, which disentangle processes driving forest regeneration. However, such studies have been lacking. To fill this gap, we followed the fates of 2,552 individual seedlings for 12 years after a large-scale bark-beetle outbreak that caused complete canopy dieback in mountain Norway spruce (Picea abies) forests in southeast Germany. We explore the contribution of advance, disturbance-related, and post-disturbance regeneration to forest recovery. Most seedlings originated directly within the three-year dieback of canopy trees induced by bark-beetle outbreak. After complete canopy dieback, the establishment of new seedlings was minimal. Surprisingly, advance regeneration formed only a minor part of all regeneration. However, because it had the highest survival rate, its importance increased over time. The most important factor influencing the survival of seedlings after disturbance was their height. Survival was further modified by microsite: seedlings established on dead wood survived best, whereas almost all seedlings surrounded by graminoids died. For 5 cm tall seedlings, annual mortality ranged from 20 to 50% according to the rooting microsite. However, for seedlings taller than 50 cm, annual mortality was below 5% at all microsites. While microsite modified seedling mortality, it did not affect seedling height growth. A model of regeneration dynamics based on short-term observations accurately predicts regeneration height growth, but substantially underestimates mortality rate, thus predicting more surviving seedlings than were observed. We found that P. abies forests were able to regenerate naturally even after severe bark-beetle outbreaks owing to advance and particularly disturbance-related regeneration. This, together with microsite-specific mortality, yields structurally and spatially diverse forests. Our study thus highlights the so far unrecognized importance of disturbance-related regeneration for stand recovery after bark-beetle outbreaks.

A walk on the wild side: Disturbance dynamics and the conservation and management of European mountain forest ecosystems.

Mountain forests are among the most important ecosystems in Europe as they support numerous ecological, hydrological, climatic, social, and economic functions. They are unique relatively natural ecosystems consisting of long-lived species in an otherwise densely populated human landscape. Despite this, centuries of intensive forest management in many of these forests have eclipsed evidence of natural processes, especially the role of disturbances in long-term forest dynamics. Recent trends of land abandonment and establishment of protected forests have coincided with a growing interest in managing forests in more natural states. At the same time, the importance of past disturbances highlighted in an emerging body of literature, and recent increasing disturbances due to climate change are challenging long-held views of dynamics in these ecosystems. Here, we synthesize aspects of this Special Issue on the ecology of mountain forest ecosystems in Europe in the context of broader discussions in the field, to present a new perspective on these ecosystems and their natural disturbance regimes. Most mountain forests in Europe, for which long-term data are available, show a strong and long-term effect of not only human land use but also of natural disturbances that vary by orders of magnitude in size and frequency. Although these disturbances may kill many trees, the forests themselves have not been threatened. The relative importance of natural disturbances, land use, and climate change for ecosystem dynamics varies across space and time. Across the continent, changing climate and land use are altering forest cover, forest structure, tree demography, and natural disturbances, including fires, insect outbreaks, avalanches, and wind disturbances. Projected continued increases in forest area and biomass along with continued warming are likely to further promote forest disturbances. Episodic disturbances may foster ecosystem adaptation to the effects of ongoing and future climatic change. Increasing disturbances, along with trends of less intense land use, will promote further increases in coarse woody debris, with cascading positive effects on biodiversity, edaphic conditions, biogeochemical cycles, and increased heterogeneity across a range of spatial scales. Together, this may translate to disturbance-mediated resilience of forest landscapes and increased biodiversity, as long as climate and disturbance regimes remain within the tolerance of relevant species. Understanding ecological variability, even imperfectly, is integral to anticipating vulnerabilities and promoting ecological resilience, especially under growing uncertainty. Allowing some forests to be shaped by natural processes may be congruent with multiple goals of forest management, even in densely settled and developed countries.

The historical disturbance regime of mountain Norway spruce forests in the Western Carpathians and its influence on current forest structure and composition.

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.

Structural chromosomal aberrations as potential risk markers in incident cancer patients.

Epidemiological prospective studies have shown that increased chromosomal aberrations (CAs) in peripheral blood lymphocytes may predict cancer risk. Here, we report CAs in newly diagnosed 101 colorectal, 87 lung and 158 breast cancer patients and corresponding healthy controls. Strong differences in distributions of aberrant cells (ACs), CAs, chromatid-type aberrations (CTAs) and chromosome-type aberrations (CSAs) were observed in lung and breast cancer patients as compared to healthy controls. In colorectal cancer (CRC) patients, only CTAs were significantly elevated. Binary logistic regression, adjusted for main confounders, indicates that all the analysed cytogenetic parameters along with smoking were significantly associated with breast and lung cancer risks. Significant differences in terminal deletions between breast cancer patients and corresponding female controls were recorded (0.39 vs. 0.18; P ≤ 0.05). We did not find any association of CAs with TNM (tumor nodus metastasis) stages or histopathological grade in either cancer type. CAs were neither associated with additional tumor characteristics-invasivity, ductal and lobular character, estrogene/progesterone receptors in breast tumors nor with non-small/small cell and bronchogenic/pulmonary types of lung tumors. Our study demonstrates that CAs serve as a predictive marker for breast and lung cancer, whereas only CTAs were elevated in incident CRC patients.

HOTAIR long non-coding RNA is a negative prognostic factor not only in primary tumors, but also in the blood of colorectal cancer patients.

Colorectal cancer (CRC) is one of the main causes of death of neoplasia. Demand for predictive and prognostic markers to reverse this trend is increasing. Long non-coding RNA HOTAIR (Homeobox Transcript Antisense Intergenic RNA) overexpression in tumors was previously associated with poor prognosis and higher mortality in different carcinomas. We analyzed HOTAIR expression levels in tumor and blood of incident sporadic CRC patients in relation to their overall survival with the aim to evaluate surrogate prognostic marker for CRC. Tissue donor group consisted of 73 CRC patients sampled for tumor and normal tissue. Blood donor group was represented by 84 CRC patients compared with 40 healthy controls. Patients were characterized for tumor-node-metastasis stage, tumor grade, microsatellite instability and tumor penetration by stromal cells. HOTAIR levels were assessed by real-time quantitative PCR. CRC patients had higher HOTAIR expression in blood than healthy controls (P = 0.0001), whereas there was no difference in HOTAIR levels between tumor and adjacent mucosa of CRC patients. HOTAIR levels positively correlated between blood and tumor (R = 0.43, P = 0.03). High HOTAIR levels in tumors were associated with higher mortality of patients [Cox's proportional hazard, hazard ratio = 4.4, 95% confidence interval: 1.0-19.2, P = 0.046]. The hazard ratio was even higher when blood HOTAIR levels were taken into account (hazard ratio = 5.9, 95% confidence interval: 1.3-26.1, P = 0.019). Upregulated HOTAIR relative expression in primary tumors and in blood of CRC patients is associated with unfavorable prognosis. Our data suggest that HOTAIR blood levels may serve as potential surrogate prognostic marker in sporadic CRC.

Disturbance, life history traits, and dynamics in an old-growth forest landscape of southeastern Europe.

Much of our understanding of natural forest dynamics in the temperate region of Europe is based on observational studies in old-growth remnants that have emphasized small-scale gap dynamics and equilibrium stand structure and composition. Relatively little attention has been given to the role of infrequent disturbance events in forest dynamics. In this study, we analyzed dendroecological data from four stands and three windthrow patches in an old-growth landscape in the Dinaric Mountains of Bosnia and Herzegovina to examine disturbance history, tree life history traits, and compositional dynamics. Over all stands, most decades during the past 340 years experienced less than 10% canopy loss, yet each stand showed evidence of periodic intermediate-severity disturbances that removed > 40% of the canopy, some of which were synchronized over the study area landscape. Analysis of radial growth patterns indicated several life history differences among the dominant canopy trees; beech was markedly older than fir, while growth patterns of dead and dying trees suggested that fir was able to tolerate longer periods of suppressed growth in shade. Maple had the fastest radial growth and accessed the canopy primarily through rapid early growth in canopy gaps, whereas most beech and fir experienced a period of suppressed growth prior to canopy accession. Peaks in disturbance were roughly linked to increased recruitment, but mainly of shade-tolerant beech and fir; less tolerant species (i.e., maple, ash, and elm) recruited successfully on some of the windthown sites where advance regeneration of beech and fir was less abundant. The results challenge the traditional notions of stability in temperate old-growth forests of Europe and highlight the nonequilibrial nature of canopy composition due to unique histories of disturbance and tree life history differences. These findings provide valuable information for developing natural disturbance-based silvicultural systems, as well as insight into maintaining less shade-tolerant, but valuable broadleaved trees in temperate forests of Europe.