Elevation-dependent tree growth response to climate in a natural Scots pine/downy birch forest in northern Sweden

Forests dominate the landscape at high latitudes in the boreal regions and contribute significantly to the global carbon stock. Large areas are protected and provide possibilities to analyze natural forest dynamics including resilience to climate change. In Fennoscandia, Scots pine (Pinus sylvestris L.) and downy birch (Betula pubescens Ehrh.) often coexist in natural forests close to the limits of their ecological ranges. Tree growth in these forests is generally thought to be limited by temperature, and changes in growth trends can therefore serve as early indicators of the impact of global warming on natural ecosystems. We sampled 592 Scots pine and downy birch trees along two elevational gradients spanning the transition from the forest zone to the coniferous treeline in Tjeggelvas nature reserve, northern Sweden. Based on the tree-ring data, we compared annual basal area increment (BAI) trends from 1902 to 2021, analyzed the ring-width indices (RWI) in relation to local climate data, and investigated trends in climate–growth relationships. We found that the mean annual growth of both species was higher in more recent years than at the beginning of the 20th century. The RWI were positively correlated with summer temperatures, however, we found a much stronger relationship for Scots pine than downy birch. We noticed a decrease in the importance of summer temperature for Scots pine growth, whereas the importance of late spring temperatures increased over the 120-year-long study period. Due to strongly positive BAI trends combined with a decrease in temperature sensitivity, the overall conclusion of our study is that the influence of increasing temperatures is still positive and outweighs the negative impacts of climate change on Scots pine growth in natural forests in northern Sweden, particularly at higher elevations. Natural forests are important natural experiments that contrast the managed forests and are key to understanding the latter

How to Calibrate Historical Aerial Photographs: A Change Analysis of Naturally Dynamic Boreal Forest Landscapes

Time series of repeat aerial photographs currently span decades in many regions. However, the lack of calibration data limits their use in forest change analysis. We propose an approach where we combine repeat aerial photography, tree-ring reconstructions, and Bayesian inference to study changes in forests. Using stereopairs of aerial photographs from five boreal forest landscapes, we visually interpreted canopy cover in contiguous 0.1-ha cells at three time points during 1959-2011. We used tree-ring measurements to produce calibration data for the interpretation, and to quantify the bias and error associated with the interpretation. Then, we discerned credible canopy cover changes from the interpretation error noise using Bayesian inference. We underestimated canopy cover using the historical low-quality photographs, and overestimated it using the recent high-quality photographs. Further, due to differences in tree species composition and canopy cover in the cells, the interpretation bias varied between the landscapes. In addition, the random interpretation error varied between and within the landscapes. Due to the varying bias and error, the magnitude of credibly detectable canopy cover change in the 0.1-ha cells depended on the studied time interval and landscape, ranging from -10 to -18 percentage points (decrease), and from +10 to +19 percentage points (increase). Hence, changes occurring at stand scales were detectable, but smaller scale changes could not be separated from the error noise. Besides the abrupt changes, also slow continuous canopy cover changes could be detected with the proposed approach. Given the wide availability of historical aerial photographs, the proposed approach can be applied for forest change analysis in biomes where tree-rings form, while accounting for the bias and error in aerial photo interpretation.Peer reviewe

Moisture content variation of ground vegetation fuels in boreal mesic and sub-xeric mineral soil forests in Finland

Forest fire risk in Finland is estimated with the Finnish Forest Fire Index (FFI), which predicts the fuel moisture content (FMC) of the forest floor. We studied the FMC variation of four typical ground vegetation fuels, Pleurozium schreberi, Hylocomium splendens, Dicranum spp., and Cladonia spp., and raw humus in mature and recently clear-cut stands. Of these, six were sub-xeric Pinus sylvestris stands, and six mesic Picea abies stands. We analysed the ability of the FFI to predict FMC and compared it with the widely applied Canadian Fire Weather Index (FWI). We found that in addition to stand characteristics, ground layer FMC was highly dependent on the species so that Dicranum was the moistest, and Cladonia the driest. In the humus layer, the differences among species were small. Overall, the FWI was a slightly better predictor of FMC than the FFI. While the FFI generally predicted ground layer FMC well, the shape of the relationship varied among the four species. The use of auxiliary variables thus has potential in improving predictions of ignitions and forest fire risk. Knowledge of FMC variation could also benefit planning and timing of prescribed burns.Peer reviewe

Dead standing pine trees in a boreal forest landscape in the Kalevala National park, northern Fennoscandia : amount, population characteristics and spatial pattern

Background: After their death, Scots pine trees can remain standing for decades and sometimes up to 200 years, forming long-lasting and ecologically important structures in boreal forest landscapes. Standing dead pines decay very slowly and with time develop into 'kelo' trees, which are characterized by hard wood with silvery-colored appearance. These kelo trees represent an ecologically important, long lasting and visually striking element of the structure of natural pine-dominated forests in boreal Fennoscandia that is nowadays virtually absent from managed forest landscapes. Methods: We examined and mapped the amount, structural features, site characteristics and spatial distribution of dead standing pine trees over a ten hectare area in an unmanaged boreal forest landscape in the Kalevala National Park in Russian Viena Karelia. Results: The mean basal area of dead standing pine trees in the forested part of the landscape was 1.7 m(2).ha(-1) and the estimated volume 12.7 m(3).ha(-1). From the total number of standing dead pine trees 65% were kelo trees, with a basal area of 1.1 m(2).ha(-1) and volume of 8.0 m(3).ha(-1), the remainder consisting of standing dead pines along the continuum between a recently dead tree and a kelo tree. Overall, standing dead pines were distributed throughout the study area, but there was a tendency towards spatial clustering up to <100 m distances. Standing dead pines were most commonly situated on flat ground or in the mid slope in the local topography. In addition, standing dead pines contributed to substrate diversity also by commonly having charred wood and broken tops. Based on the presence of dead pine snags in different stage of transition from a recently dead pine to a kelo with silvery surface, it seems evident that the process of kelo recruitment was continuously in action in the studied landscape. Conclusions: Kelo trees are an omnipresent feature in natural pine-dominated forest landscapes with important contribution to forest structural and substrate diversity. Because of their longevity and extremely slow turnover dynamics and importance for biodiversity, protection of vulnerable kelo tree populations, and ensuring their continuous recruitment, should be of high priority in forest restoration and sustainable management.Peer reviewe

Changes in nitrogen content and isotopic composition in subarctic Empetrum nigrum seeds in the period 1976–2010

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North Fennoscandian mountain forests : History, composition, disturbance dynamics and the unpredictable future

North Fennoscandian mountain forests are distributed along the Scandes Mountains between Sweden and Norway, and the low-mountain regions of northern Norway, Sweden and Finland, and the adjacent northwestern Russia. Regionally, these forests are differentiated into spruce, pine or birch dominance due to climatic differences. Variation in tree species dominance within these regions is generally caused by a combination of historical and prevailing disturbance regimes, including both chronic and episodic disturbances, their magnitude and frequency, as well as differences in edaphic conditions and topography. Because of their remoteness, slow growth and restrictions of use, these mountain forests are generally less affected by human utilization than more productive and easily utilizable forests at lower elevations and/or latitudes. As a consequence, these northern forests of Europe are often referred to as "Europe's last wilderness", even if human influence of varying intensity has been ubiquitous through historical time. Because of their naturalness, the North Fennoscandian mountain forests are of paramount importance for biodiversity conservation, monitoring of ecosystem change and for their sociocultural values. As such, they also provide unique reference areas for basic and applied research, and for developing methods of forest conservation, restoration and ecosystem-based management for the entire Fennoscandia. However, the current rapid change in climate is predicted to profoundly affect the ecology and dynamics of these forests in the future. (C) 2016 Elsevier B.V. All rights reserved.Peer reviewe

Increase in dead wood, large living trees and tree diversity, yet decrease in understory vegetation cover: The effect of three decades of biodiversity-oriented forest policy in Swedish forests

In Sweden, the majority of forest area has been altered by industrial forestry over the decades. Almost 30 years ago, a shift towards biodiversity-oriented forest management practices occurred. Here we took advantage of long-term data collected by the Swedish National Forest Inventory to track developmental changes in forest structural components over this time. We assessed changes in structural components that play an important role in biodiversity (dead wood, large living trees, tree species composition, and understory vegetation) in four forest types with descending tiers of biodiversity protection: protected areas, woodland key habitats, low-productivity forests and production forests. Overall, we found a positive trend in the volumes of dead wood and large living trees, as well as in tree species diversity, while there was a general decline in understory vegetation coverage. Most observed changes were consistent with the intended outcomes of the current forest policy, adapted in the early 1990s. The implementation of retention forestry is likely driving some of the observed changes in forest structural components in the south. In contrast, we observed no changes in any of the focal structural components in the north, which could be attributed to the ongoing clear-cutting of forests previously managed less intensively. Dead wood and large living trees increased not only in managed, but also in unmanaged forests, likely reflecting historical management. The increased tree species diversity can be explained through current forest management practices that encourages maintenance of additional tree species. Decreasing understory vegetation coverage in both dense managed and unmanaged forests suggests that factors other than forestry contribute to the ongoing changes in understory vegetation in Swedish forests. Overall, the observed increase in structural components has not yet been reflected in documented improvements for red-listed forest species, which may be due to delays in species responses to small improvements, as well as a lack of detailed monitoring. Similarly, the increased availability of forest structural components might still be insufficient to meet the specific habitat requirements of red-listed species

Effects of local forest continuity on the diversity of fungi on standing dead pines

Human-induced fragmentation affects forest continuity, i.e. availability of a suitable habitat for the target species over a time period. The dependence of wood-inhabiting fungi on landscape level continuity has been well demonstrated, but the importance of local continuity has remained controversial. In this study, we explored the effects of local forest continuity (microhabitat and stand level) on the diversity of wood-inhabiting fungi on standing dead trunks of Scots pine (Paws sylvestris L.). We studied species richness and community composition of decomposers and Micarea lichens on 70 trunks in 14 forests in central Finland that differed in their state of continuity. We used dendrochronological methods to assess the detailed history of each study trunk, i.e. the microhabitat continuity. The stand continuity was estimated as dead wood diversity and past management intensity (number of stumps). We recorded 107 species (91 decomposers, 16 Micarea lichens), with a total of 510 occurrences. Using generalized linear mixed models, we found that none of the variables explained decomposer species richness, but that Micarea species richness was positively dependent on the time since tree death. Dead wood diversity was the most important variable determining the composition of decomposer communities. For Micarea lichens, the community composition was best explained by the combined effect of years from death, site and dead wood diversity. However, these effects were rather tentative. The results are in line with those of previous studies suggesting the restricted significance of local forest continuity for wood-inhabiting fungi. However, standing dead pines that have been available continuously over long periods seem to be important for species-rich communities of Micarea lichens. Rare specialists (e.g. on veteran trees) may be more sensitive to local continuity, and should be at the center of future research.Peer reviewe

Spatial tree community structure in three stands across a forest succession gradient in northern boreal Fennoscandia

Development of species composition during succession is well studied in natural boreal forests, but empirical assessments of how within-stand spatial structure develops in late-successional stages are few. Here, we quantified spatial patterns in three unmanaged stands consisting of Picea abies (L.) Karst. and Betula pubescens Ehrh. and Betula pendula Roth (hereafter Betula spp.) in northern boreal Fennoscandia. We conducted a comprehensive analysis of small-scale spatial point patterns in three fully mapped 1.2-ha sample plots, representing different forest developmental stages: mid-successional, late-successional and old-growth forest. We used several variants of Ripley’s K-function to analyze the spatial point patterns along the successional gradient. Univariate analyses showed that mature trees of both species were either randomly distributed or clumped. P. abies saplings were clumped, and Betula spp. saplings occurred in a random or clumped manner. In the bivariate analyses, saplings were more likely to be found in the surroundings of mature trees of the same species, but occurred independent of the individuals of other tree species. Mature trees showed interspecific repulsion. Only modest differences occurred in the univariate patterns between the three successional stages, but in the bivariate analyses the most evident patterns, i.e. intraspecific attraction and interspecific repulsion, were stronger in the older successional stages. Overall, the studied stands appear structured as species-specific mosaics. These mosaics, along with mixed species composition, seem to be maintained by species self-replacement, which contrasts with findings from earlier studies.Peer reviewe

At What Scales and Why Does Forest Structure Vary in Naturally Dynamic Boreal Forests? An Analysis of Forest Landscapes on Two Continents

Identifying the scales of variation in forest structures and the underlying processes are fundamental for understanding forest dynamics. Here, we studied these scale-dependencies in forest structure in naturally dynamic boreal forests on two continents. We identified the spatial scales at which forest structures varied, and analyzed how the scales of variation and the underlying drivers differed among the regions and at particular scales. We studied three 2kmx2km landscapes in northeastern Finland and two in eastern Canada. We estimated canopy cover in contiguous 0.1-ha cells from aerial photographs and used scale-derivative analysis to identify characteristic scales of variation in the canopy cover data. We analyzed the patterns of variation at these scales using Bayesian scale space analysis. We identified structural variation at three spatial scales in each landscape. Among landscapes, the largest scale of variation showed the greatest variability (20.1-321.4ha), related to topography, soil variability, and long-term disturbance history. Superimposed on this large-scale variation, forest structure varied at similar scales (1.3-2.8ha) in all landscapes. This variation correlated with recent disturbances, soil variability, and topographic position. We also detected intense variation at the smallest scale analyzed (0.1ha, grain of our data), partly driven by recent disturbances. The distinct scales of variation indicated hierarchical structure in the landscapes studied. Except for the large-scale variation, these scales were remarkably similar among the landscapes. This suggests that boreal forests may display characteristic scales of variation that occur somewhat independent of the tree species characteristics or the disturbance regime.Peer reviewe