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Aim: Primary forests have high conservation value but are rare in Europe due to historic land use. Yet many primary forest patches remain unmapped, and it is unclear to what extent they are effectively protected. Our aim was to (1) compile the most comprehensive European-scale map of currently known primary forests, (2) analyse the spatial determinants characterizing their location and (3) locate areas where so far unmapped primary forests likely occur. Location: Europe. Methods: We aggregated data from a literature review, online questionnaires and 32 datasets of primary forests. We used boosted regression trees to explore which biophysical, socio-economic and forest-related variables explain the current distribution of primary forests. Finally, we predicted and mapped the relative likelihood of primary forest occurrence at a 1-km resolution across Europe. Results: Data on primary forests were frequently incomplete or inconsistent among countries. Known primary forests covered 1.4 Mha in 32 countries (0.7% of Europe's forest area). Most of these forests were protected (89%), but only 46% of them strictly. Primary forests mostly occurred in mountain and boreal areas and were unevenly distributed across countries, biogeographical regions and forest types. Unmapped primary forests likely occur in the least accessible and populated areas, where forests cover a greater share of land, but wood demand historically has been low. Main conclusions: Despite their outstanding conservation value, primary forests are rare and their current distribution is the result of centuries of land use and forest management. The conservation outlook for primary forests is uncertain as many are not strictly protected and most are small and fragmented, making them prone to extinction debt and human disturbance. Predicting where unmapped primary forests likely occur could guide conservation efforts, especially in Eastern Europe where large areas of primary forest still exist but are being lost at an alarming pace.

Aims Primary forests are critical for forest biodiversity and provide key ecosystem services. In Europe, these forests are particularly scarce and it is unclear whether they are sufficiently protected. Here we aim to: (a) understand whether extant primary forests are representative of the range of naturally occurring forest types, (b) identify forest types which host enough primary forest under strict protection to meet conservation targets and (c) highlight areas where restoration is needed and feasible. Location Europe. Methods We combined a unique geodatabase of primary forests with maps of forest cover, potential natural vegetation, biogeographic regions and protected areas to quantify the proportion of extant primary forest across Europe's forest types and to identify gaps in protection. Using spatial predictions of primary forest locations to account for underreporting of primary forests, we then highlighted areas where restoration could complement protection. Results We found a substantial bias in primary forest distribution across forest types. Of the 54 forest types we assessed, six had no primary forest at all, and in two‐thirds of forest types, less than 1% of forest was primary. Even if generally protected, only ten forest types had more than half of their primary forests strictly protected. Protecting all documented primary forests requires expanding the protected area networks by 1,132 km2 (19,194 km2 when including also predicted primary forests). Encouragingly, large areas of non‐primary forest existed inside protected areas for most types, thus presenting restoration opportunities. Main conclusion Europe's primary forests are in a perilous state, as also acknowledged by EU's “Biodiversity Strategy for 2030.” Yet, there are considerable opportunities for ensuring better protection and restoring primary forest structure, composition and functioning, at least partially. We advocate integrated policy reforms that explicitly account for the irreplaceable nature of primary forests and ramp up protection and restoration efforts alike.

QUESTIONS: Are the structural characteristics of natural beech forests predominant on small monitoring plots representative of the large‐scale features of these forests? Do the findings of our large‐scale investigation support the hypothesis that this primeval beech forest is shaped by fine‐scale processes, or is there evidence that high‐severity disturbance events have affected its structure? LOCATION: Ukrainian Carpathians, Uholka‐Shyrokyi Luh, the largest primeval beech forest in Europe, covering 102.8 km².METHODS: On 314 (500 m²) circular plots, systematically distributed across the forest, all living and dead trees with a DBH ≥6 cm were assessed. Lying deadwood, tree regeneration, the size of canopy gaps and the number of canopy layers were recorded. Spatial analyses were conducted using Moran's I. Dendrochronological analysis was used to reconstruct tree ages and growth patterns. RESULTS: The forest is characterized overall by a density of 435.0 ± 12.2 ha⁻¹(mean ± SE) living trees, a basal area of 36.6 ± 0.8 m²·ha⁻¹, a volume of living trees of 582.1 ± 13.5 m³·ha⁻¹and a total deadwood volume of 162.5 ± 8.4 m³·ha⁻¹. Beech is the dominant species (97.3 ± 0.7%, by basal area), interspersed with mostly deciduous species with moderate shade tolerance. The forest canopy is multi‐layered, with a high abundance of old trees, and with canopy gaps rarely larger than the crown projection area of a few trees. CONCLUSIONS: The results lend support to findings from small monitoring plots, but our landscape approach allows a reliable estimation of key forest characteristics such as basal area and standing volume, which tend to be overestimated in studies on subjectively placed small monitoring plots. We conclude that disturbance events of moderate or higher severity have left only few discernible traces in forest structure or species composition. All findings suggest that this forest is characterized by a mainly small‐scale disturbance regime, leading to a homogeneity of forest characteristics of the living stand at larger spatial scales and causing only minor fluctuations around the average values of forest parameters observed.

Here we present the first reliable record of the rare Hyperoscelis veternosa Mamaev and Krivosheina, 1969 from the headwaters area of the Someșul River, Apuseni Natural Park, Romania. Species level identification was made by detailed examination of the single female genitalia, supported by DNA barcode data. The new finding confirms the presence of the species in Romania, whereas a very old, single and doubtful record from the Bucharest area dates back to 1900 needs further confirmation. The presence of the second species of genus Hyperoscelis from Romania is of high conservation value, and confirm the persistence of a relict biodiversity of an old-growth forests-spot in the Carpathians.

Research Highlights: Carpathian forests hold high ecological and economic value while generating conservation concerns, with some of these forests being among the few remaining temperate virgin forests in Europe. Carpathian forests partially lost their original integrity due to their management. Climate change has also gradually contributed to forest changes due to its modification of the environmental conditions. Background and Objectives: Understanding trees’ responses to past climates and forms of management is critical in foreseeing the responses of forests to future conditions. This study aims (1) to determine the sensitivity of Carpathian forests to past climates using dendrochronological records and (2) to describe the effects that climate change and management will have on the attributes of Carpathian forests, with a particular focus on the different response of pure and mixed forests. Materials and Methods: To this end, we first analysed the past climate-induced growth change in a dendrochronological reference series generated for virgin forests in the Romanian Curvature Carpathians and then used the obtained information to calibrate spatially explicit forest Landis-II models for the same region. The model was used to project forest change under four climate change scenarios, from mild to extreme. Results: The dendrochronological analysis revealed a climate-driven increase in forest growth over time. Landis-II model simulations also indicate that the amount of aboveground forest biomass will tend to increase with climate change. Conclusions: There are differences in the response of pure and mixed forests. Therefore, suitable forest management is required when forests change with the climate.

We analyzed growth trajectories recorded in the tree-ring series of Norway spruce Picea abies (L.) H. Karst. and silver fir Abies alba Mill. co-occuring with European beech Fagus sylvatica L. in old-growth forests driven by natural stand dynamics. The study sites were localized in five old-growth stands in the Western Carpathians (Central Europe). We characterized the life histories of 218 individual trees by using 25 metrics which refer to the age of the trees, number of release events, reaction to releases, radial increment and basal area increment. We found a large variation in the dbh-age relationship. The life histories of most trees included two or three (in single cases up to seven) release events. We did not find tree age as an important factor in determining post-release growth response. The maximal basal area increment was frequently registered at the terminal stage of tree life. The life histories of spruces and firs did not significantly differ. We concluded that in natural forests driven by gap-phase dynamics, the history of release events is the primary driver of tree growth and that tree age plays a secondary role. After reaching the overstory, trees can continue growing without significant symptoms of aging until extrinsic forces cause their death. Our results indicate that in the studied ecosystem the persistence of less shade-tolerant spruce is not attributable to the niche differentiation and site partitioning mechanism. An alternative hypothesis based on shifts in stand densities, species composition and climatic factors is more probable.

Despite extensive forest destruction in the Middle Ages and later intensive commercial forest management, remnants of virgin forests remained spared in some Central, Eastern and South-Eastern European countries. These virgin forests are the last examples of original forests in this part of Europe. That is why their protection becomes an important issue of current European forestry and nature protection policy. But the knowledge about the location and the area of virgin forests in these countries is incomplete up till now. This article has the prime goal to present a conceptual framework what virgin forests might be (“A conceptual framework for defining of virgin forests” section). Based on this framework, a working methodology has been tested in Bulgaria and Romania (“Results of the two national projects in Romania and in Bulgaria” section and further). For this reason two projects have been carried out by the Royal Dutch Society of Nature Conservation (KNNV) in close co-operation with the Forestry Institutes in Romania and in Bulgaria. The results of these projects are described in general terms and further analysis in the future is necessary to describe specific features like forest structure and spatial heterogeneity of these forests. Based on the results of the inventory, principles of sustainable protection and management of the mapped virgin forests were defined and described in the research reports. The usefulness of the inventory became evident already during the EU pre-accession period of both countries while preparing the NATURA 2000 network. The remaining virgin forests of temperate Europe are an inexhaustible source of ecological information about biodiversity, structure, natural processes and overall functioning of undisturbed forest ecosystems. Their research will reveal information which can be used for ecological restoration of man-made forests which are degraded through intensive forestry practices over the last centuries. The last virgin forests of temperate Europe represent an irreplaceable part of the natural capital of Europe and are worth to be protected by law. Their last remnants in South-Eastern and Eastern Europe are endangered by commercial activities. A full inventory of remaining virgin forests in all countries of temperate Europe is a matter of highest urgency. A representative selection of virgin forest sites should be declared by UNESCO as World Heritage Sites.

The soil carbon pool holds an enormous amount of carbon, making it the largest reservoir in the terrestrial ecosystem. However, there is growing concern that unsustainable logging methods damage the soil ecosystem, thus triggering the release of soil carbon into the atmosphere hence contributing to ongoing climate change. This study uses a replicated (n = 4) logging experiment to examine the impact of supervised logging with climber cutting (SLCC) and conventional logging (CL) on basic soil characteristics, litter input to soils, soil carbon pools, and soil respiration in a mixed dipterocarp forest 26 years after logging. This study found that there was no significant difference observed in the soil physicochemical properties and total carbon pools between the logging treatments and the virgin forest. Soil carbon pools dominated the total carbon pools, and the highest mean value was recorded in SLCC (87.95 ± 13.67 Mg C ha−1). Conventional logging had a lower mean value (71.17 ± 12.09 Mg C ha−1) than virgin forest (83.20 ± 11.97 Mg C ha−1). SLCC also shows a higher value of soil respiration rate (161.75 ± 21.67 mg C m−2 h−1) than CL (140.54 ± 12.54 mg C m−2 h−1). These findings highlight the importance of accurate quantification of the effect of different logging methods on the forest’s carbon pools.

Little is known about the gap pattern of primeval beech forests, since large-scale studies with continuous coverage are lacking. Analyses of forest structural patterns have benefitted from advances in remote sensing, especially with the launch of satellites providing data of submetric ground resolution. These developments can strongly advance our knowledge of natural forest dynamics and disturbance regimes. The Uholka-Shyrokyi Luh forest in the Ukrainian Carpathians, the largest remnant of primeval European beech ( Fagus sylvatica L.) covering 102.8 km 2 , is an outstanding object to analyze the frequency distribution of gap sizes and to infer processes of forest dynamics. A stereo pair of very high-resolution WorldView-2 satellite images was used to characterize the forest's gap pattern. Canopy gaps were first digitized stereoscopically based on the image pair. In a second step, spectral properties in the red and yellow frequency bands were used to distinguish the stereoscopically mapped gap areas from non-gap areas, which enabled gap mapping over the entire study area. To validate the spectral gap mapping 338 randomly distributed samples were assigned manually to gap and non-gap areas based on the ortho-images. We found excellent agreement except for an overestimation of gaps close to clouds due to diffuse image areas. The frequency distribution of gap size revealed the forest to be structured by a small-scale mosaic of gaps mainly <200 m 2 (98% of the gaps). Only a few large, stand-replacing events were detected, most probably caused by a wind storm in March 2007 and a heavy wet snow fall in October 2009. The small canopy gaps reflect fine-scale processes shaping forest structure, i.e., the death of single trees or groups of a few trees and is in line with the findings of the terrestrial forest inventory. We conclude that remote sensing approaches based on very high-resolution satellite images are highly useful to characterize even small-scale forest disturbance regimes and to study long-term gap dynamics. Stereo satellite images provide two viewing angles of the study area, thus allowing for a highly accurate mapping of canopy gaps in forests with a complex topography.

Despite the sharply growing interest in the disturbances occurring in primary forests, little is known about the response of European virgin forests to anthropogenic disturbance. The present study investigated the effect of the first silvicultural interventions that took place nine years earlier in a former virgin forest (FVF). Changes in the stand structure, environmental characteristics, and diversity of ground vegetation were studied in comparison with a nearby virgin forest (VF), both consisting of a mixture of European beech and silver fir. While the tree density did not differ significantly between the two forests, the number of large trees, the basal area, and the stand volume were significantly reduced in the FVF. The deadwood volume was twice as great in the VF as in the FVF and was found in both forests, particularly from silver fir. Despite significantly better light conditions in the FVF, natural regeneration was not significantly higher than in the VF. However, a slight improvement in the proportion of silver fir and other tree species into total regeneration was reported. The soil temperature was significantly higher in the FVF, independent of the measurement season, while the soil moisture showed a higher value in the VF only in spring. The FVF is characterized by a greater soil CO2 emission, which is especially significant in summer and fall. The diversity of the ground vegetation did not yet react significantly to the silvicultural intervention. These preliminary findings are important in drawing suitable forest management practices that need to be applied in mixed beech–silver fir stands, especially in terms of maintaining species diversity. However, the short time frame since the intervention obliges further research on this VF–FVF pair over the next 10–20 years, at least regarding silver fir dynamics.