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.

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.

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.

Forest and woodland replacement patterns following drought-related mortality.

Forest vulnerability to drought is expected to increase under anthropogenic climate change, and drought-induced mortality and community dynamics following drought have major ecological and societal impacts. Here, we show that tree mortality concomitant with drought has led to short-term (mean 5 y, range 1 to 23 y after mortality) vegetation-type conversion in multiple biomes across the world (131 sites). Self-replacement of the dominant tree species was only prevalent in 21% of the examined cases and forests and woodlands shifted to nonwoody vegetation in 10% of them. The ultimate temporal persistence of such changes remains unknown but, given the key role of biological legacies in long-term ecological succession, this emerging picture of postdrought ecological trajectories highlights the potential for major ecosystem reorganization in the coming decades. Community changes were less pronounced under wetter postmortality conditions. Replacement was also influenced by management intensity, and postdrought shrub dominance was higher when pathogens acted as codrivers of tree mortality. Early change in community composition indicates that forests dominated by mesic species generally shifted toward more xeric communities, with replacing tree and shrub species exhibiting drier bioclimatic optima and distribution ranges. However, shifts toward more mesic communities also occurred and multiple pathways of forest replacement were observed for some species. Drought characteristics, species-specific environmental preferences, plant traits, and ecosystem legacies govern postdrought species turnover and subsequent ecological trajectories, with potential far-reaching implications for forest biodiversity and ecosystem services.

Low growth resilience to drought is related to future mortality risk in trees.

Severe droughts have the potential to reduce forest productivity and trigger tree mortality. Most trees face several drought events during their life and therefore resilience to dry conditions may be crucial to long-term survival. We assessed how growth resilience to severe droughts, including its components resistance and recovery, is related to the ability to survive future droughts by using a tree-ring database of surviving and now-dead trees from 118 sites (22 species, >3,500 trees). We found that, across the variety of regions and species sampled, trees that died during water shortages were less resilient to previous non-lethal droughts, relative to coexisting surviving trees of the same species. In angiosperms, drought-related mortality risk is associated with lower resistance (low capacity to reduce impact of the initial drought), while it is related to reduced recovery (low capacity to attain pre-drought growth rates) in gymnosperms. The different resilience strategies in these two taxonomic groups open new avenues to improve our understanding and prediction of drought-induced mortality.

TRY plant trait database - enhanced coverage and open access.

Plant traits-the morphological, anatomical, physiological, biochemical and phenological characteristics of plants-determine how plants respond to environmental factors, affect other trophic levels, and influence ecosystem properties and their benefits and detriments to people. Plant trait data thus represent the basis for a vast area of research spanning from evolutionary biology, community and functional ecology, to biodiversity conservation, ecosystem and landscape management, restoration, biogeography and earth system modelling. Since its foundation in 2007, the TRY database of plant traits has grown continuously. It now provides unprecedented data coverage under an open access data policy and is the main plant trait database used by the research community worldwide. Increasingly, the TRY database also supports new frontiers of trait-based plant research, including the identification of data gaps and the subsequent mobilization or measurement of new data. To support this development, in this article we evaluate the extent of the trait data compiled in TRY and analyse emerging patterns of data coverage and representativeness. Best species coverage is achieved for categorical traits-almost complete coverage for 'plant growth form'. However, most traits relevant for ecology and vegetation modelling are characterized by continuous intraspecific variation and trait-environmental relationships. These traits have to be measured on individual plants in their respective environment. Despite unprecedented data coverage, we observe a humbling lack of completeness and representativeness of these continuous traits in many aspects. We, therefore, conclude that reducing data gaps and biases in the TRY database remains a key challenge and requires a coordinated approach to data mobilization and trait measurements. This can only be achieved in collaboration with other initiatives.

Early-Warning Signals of Individual Tree Mortality Based on Annual Radial Growth.

Tree mortality is a key driver of forest dynamics and its occurrence is projected to increase in the future due to climate change. Despite recent advances in our understanding of the physiological mechanisms leading to death, we still lack robust indicators of mortality risk that could be applied at the individual tree scale. Here, we build on a previous contribution exploring the differences in growth level between trees that died and survived a given mortality event to assess whether changes in temporal autocorrelation, variance, and synchrony in time-series of annual radial growth data can be used as early warning signals of mortality risk. Taking advantage of a unique global ring-width database of 3065 dead trees and 4389 living trees growing together at 198 sites (belonging to 36 gymnosperm and angiosperm species), we analyzed temporal changes in autocorrelation, variance, and synchrony before tree death (diachronic analysis), and also compared these metrics between trees that died and trees that survived a given mortality event (synchronic analysis). Changes in autocorrelation were a poor indicator of mortality risk. However, we found a gradual increase in inter-annual growth variability and a decrease in growth synchrony in the last ∼20 years before mortality of gymnosperms, irrespective of the cause of mortality. These changes could be associated with drought-induced alterations in carbon economy and allocation patterns. In angiosperms, we did not find any consistent changes in any metric. Such lack of any signal might be explained by the relatively high capacity of angiosperms to recover after a stress-induced growth decline. Our analysis provides a robust method for estimating early-warning signals of tree mortality based on annual growth data. In addition to the frequently reported decrease in growth rates, an increase in inter-annual growth variability and a decrease in growth synchrony may be powerful predictors of gymnosperm mortality risk, but not necessarily so for angiosperms.

Effects of Competition, Drought Stress and Photosynthetic Productivity on the Radial Growth of White Spruce in Western Canada.

Understanding the complex interactions of competition, climate warming-induced drought stress, and photosynthetic productivity on the radial growth of trees is central to linking climate change impacts on tree growth, stand structure and in general, forest productivity. Using a mixed modeling approach, a stand-level photosynthetic production model, climate, stand competition and tree-ring data from mixedwood stands in western Canada, we investigated the radial growth response of white spruce [Picea glauca (Moench.) Voss] to simulated annual photosynthetic production, simulated drought stress, and tree and stand level competition. The long-term (~80-year) radial growth of white spruce was constrained mostly by competition, as measured by total basal area, with minor effects from drought. There was no relation of competition and drought on tree growth but dominant trees increased their growth more strongly to increases in modeled photosynthetic productivity, indicating asymmetric competition. Our results indicate a co-limitation of drought and climatic factors inhibiting photosynthetic productivity for radial growth of white spruce in western Canada. These results illustrate how a modeling approach can separate the complex factors regulating both multi-decadal average radial growth and interannual radial growth variations of white spruce, and contribute to advance our understanding on sustainable management of mixedwood boreal forests in western Canada.

Newtonian boreal forest ecology: The Scots pine ecosystem as an example.

Isaac Newton's approach to developing theories in his book Principia Mathematica proceeds in four steps. First, he defines various concepts, second, he formulates axioms utilising the concepts, third, he mathematically analyses the behaviour of the system defined by the concepts and axioms obtaining predictions and fourth, he tests the predictions with measurements. In this study, we formulated our theory of boreal forest ecosystems, called NewtonForest, following the four steps introduced by Newton. The forest ecosystem is a complicated entity and hence we needed altogether 27 concepts to describe the material and energy flows in the metabolism of trees, ground vegetation and microbes in the soil, and to describe the regularities in tree structure. Thirtyfour axioms described the most important features in the behaviour of the forest ecosystem. We utilised numerical simulations in the analysis of the behaviour of the system resulting in clear predictions that could be tested with field data. We collected retrospective time series of diameters and heights for test material from 6 stands in southern Finland and five stands in Estonia. The numerical simulations succeeded to predict the measured diameters and heights, providing clear corroboration with our theory.

Reliability of temperature signal in various climate indicators from northern Europe.

We collected relevant observational and measured annual-resolution time series dealing with climate in northern Europe, focusing in Finland. We analysed these series for the reliability of their temperature signal at annual and seasonal resolutions. Importantly, we analysed all of the indicators within the same statistical framework, which allows for their meaningful comparison. In this framework, we employed a cross-validation procedure designed to reduce the adverse effects of estimation bias that may inflate the reliability of various temperature indicators, especially when several indicators are used in a multiple regression model. In our data sets, timing of phenological observations and ice break-up were connected with spring, tree ring characteristics (width, density, carbon isotopic composition) with summer and ice formation with autumn temperatures. Baltic Sea ice extent and the duration of ice cover in different watercourses were good indicators of winter temperatures. Using combinations of various temperature indicator series resulted in reliable temperature signals for each of the four seasons, as well as a reliable annual temperature signal. The results hence demonstrated that we can obtain reliable temperature information over different seasons, using a careful selection of indicators, combining the results with regression analysis, and by determining the reliability of the obtained indicator.

A synthesis of radial growth patterns preceding tree mortality.

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.

Even-aged and uneven-aged forest management in boreal Fennoscandia: a review.

Since WWII, forest management in Fennoscandia has primarily been based on even-aged stand management, clear cut harvesting and thinning from below. As an alternative, uneven-aged management, based on selection cutting of individual trees or small groups of trees, has been proposed. In this review we discuss the theoretical aspects of ecology and economics of the two management approaches. We also review peer-reviewed studies from boreal Fennoscandia, which have aimed at comparing the outcomes of uneven-aged and the conventional even-aged forest management. According to a common view the main obstacle of practicing uneven-aged forestry is its low economic performance. However, the reviewed studies did not offer any straightforward support for this view and several studies have found uneven-aged management to be fully competitive with existing even-aged management. Studies on the ecological aspects indicated that selection cuttings maintain mature or late-successional forest characteristics and species assemblages better than even-aged management, at least at the stand scale and in the short term. We conclude that although the number of relevant studies has increased in recent years, the ecological and economic performance of alternative management methods still remains poorly examined, especially for those stands with multiple tree species and also at wider spatial and temporal scales. For future research we advocate a strategy that fully takes into consideration the interdisciplinary nature of forest management and is better connected to social goals and latest theoretical and methodological developments in ecology and economics.