Plant functional traits have globally consistent effects on competition.

Phenotypic traits and their associated trade-offs have been shown to have globally consistent effects on individual plant physiological functions, but how these effects scale up to influence competition, a key driver of community assembly in terrestrial vegetation, has remained unclear. Here we use growth data from more than 3 million trees in over 140,000 plots across the world to show how three key functional traits--wood density, specific leaf area and maximum height--consistently influence competitive interactions. Fast maximum growth of a species was correlated negatively with its wood density in all biomes, and positively with its specific leaf area in most biomes. Low wood density was also correlated with a low ability to tolerate competition and a low competitive effect on neighbours, while high specific leaf area was correlated with a low competitive effect. Thus, traits generate trade-offs between performance with competition versus performance without competition, a fundamental ingredient in the classical hypothesis that the coexistence of plant species is enabled via differentiation in their successional strategies. Competition within species was stronger than between species, but an increase in trait dissimilarity between species had little influence in weakening competition. No benefit of dissimilarity was detected for specific leaf area or wood density, and only a weak benefit for maximum height. Our trait-based approach to modelling competition makes generalization possible across the forest ecosystems of the world and their highly diverse species composition.

Growth resistance and resilience of mixed silver fir and Norway spruce forests in central Europe: Contrasting responses to mild and severe droughts.

Extreme droughts are expected to increase in frequency and severity in many regions of the world, threatening multiple ecosystem services provided by forests. Effective strategies to adapt forests to such droughts require comprehensive information on the effects and importance of the factors influencing forest resistance and resilience. We used a unique combination of inventory and dendrochronological data from a long-term (>30 years) silvicultural experiment in mixed silver fir and Norway spruce mountain forests along a temperature and precipitation gradient in southwestern Germany. We aimed at examining the mechanisms and forest stand characteristics underpinning the resistance and resilience to past mild and severe droughts. We found that (i) fir benefited from mild droughts and showed higher resistance (i.e., lower growth loss during drought) and resilience (i.e., faster return to pre-drought growth levels) than spruce to all droughts; (ii) species identity determined mild drought responses while species interactions and management-related factors strongly influenced the responses to severe droughts; (iii) intraspecific and interspecific interactions had contrasting effects on the two species, with spruce being less resistant to severe droughts when exposed to interaction with fir and beech; (iv) higher values of residual stand basal area following thinning were associated with lower resistance and resilience to severe droughts; and (v) larger trees were resilient to mild drought events but highly vulnerable to severe droughts. Our study provides an analytical approach for examining the effects of different factors on individual tree- and stand-level drought response. The forests investigated here were to a certain extent resilient to mild droughts, and even benefited from such conditions, but were strongly affected by severe droughts. Lastly, negative effects of severe droughts can be reduced through modifying species composition, tree size distribution and stand density in mixed silver fir-Norway spruce forests.

Climate change and the provision of biodiversity in public temperate forests - A mechanism design approach for the implementation of biodiversity conservation policies.

The provision of forest biodiversity remains a major challenge in the management of forest resources. Biodiversity is mostly considered a public good and the fact that societal benefits from biodiversity are private information, hinders its supply at adequate levels. Here we investigate how the government, as a forest owner, may increase the biodiversity supply in publicly-owned forests. We employ a mechanism design approach to find the biodiversity provision choices, which take into account agents' strategic behavior and values towards biodiversity. We applied our framework to a forest landscape in Southwestern Germany, using forest birds as biodiversity indicators and evaluating the impacts of climate change on forest dynamics and on the costs of biodiversity provision. Our results show that climate change has important implications to the opportunity cost of biodiversity and the provision levels (ranging from 10 to 12.5% increase of the bird indicator abundance). In general, biodiversity valuations needed to surpass the opportunity cost by more than 18% to cope with the private information held by the agents. Moreover, higher costs under more intense climate change (e.g. Representative Concentration Pathway 8.5) reduced the attainable bird abundance increase from 12.5 to 10%. We conclude that mechanism design may provide key information for planning conservation policies and identify conditions for a successful implementation of biodiversity-oriented forest management.

Survival of Norway spruce remains higher in mixed stands under a dryer and warmer climate.

Shifts in tree species distributions caused by climatic change are expected to cause severe losses in the economic value of European forestland. However, this projection disregards potential adaptation options such as tree species conversion, shorter production periods, or establishment of mixed species forests. The effect of tree species mixture has, as yet, not been quantitatively investigated for its potential to mitigate future increases in production risks. For the first time, we use survival time analysis to assess the effects of climate, species mixture and soil condition on survival probabilities for Norway spruce and European beech. Accelerated Failure Time (AFT) models based on an extensive dataset of almost 65,000 trees from the European Forest Damage Survey (FDS)--part of the European-wide Level I monitoring network--predicted a 24% decrease in survival probability for Norway spruce in pure stands at age 120 when unfavorable changes in climate conditions were assumed. Increasing species admixture greatly reduced the negative effects of unfavorable climate conditions, resulting in a decline in survival probabilities of only 7%. We conclude that future studies of forest management under climate change as well as forest policy measures need to take this, as yet unconsidered, strongly advantageous effect of tree species mixture into account.

Converting probabilistic tree species range shift projections into meaningful classes for management.

The paper deals with the management problem how to decide on tree species suitability under changing environmental conditions. It presents an algorithm that classifies the output of a range shift model for major tree species in Europe into multiple classes that can be linked to qualities characterizing the ecological niche of the species. The classes: i) Core distribution area, ii) Extended distribution area, iii) Occasional occurrence area, and iv) No occurrence area are first theoretically developed and then statistically described. The classes are interpreted from an ecological point of view using criteria like population structure, competitive strength, site spectrum and vulnerability to biotic hazards. The functioning of the algorithm is demonstrated using the example of a generalized linear model that was fitted to a pan-European dataset of presence/absence of major tree species with downscaled climate data from a General Circulation Model (GCM). Applications of the algorithm to tree species suitability classification on a European and regional level are shown. The thresholds that are used by the algorithm are precision-based and include Cohen's Kappa. A validation of the algorithm using an independent dataset of the German National Forest Inventory shows good accordance of the statistically derived classes with ecological traits for Norway spruce, while the differentiation especially between core and extended distribution for European beech that is in the centre of its natural range in this area is less accurate. We hypothesize that for species in the core of their range regional factors like forest history superimpose climatic factors. Problems of uncertainty issued from potentially applying a multitude of modelling approaches and/or climate realizations within the range shift model are discussed and a way to deal with the uncertainty by revealing the underlying attitude towards risk of the decision maker is proposed.

Climate change and European forests: what do we know, what are the uncertainties, and what are the implications for forest management?

The knowledge about potential climate change impacts on forests is continuously expanding and some changes in growth, drought induced mortality and species distribution have been observed. However despite a significant body of research, a knowledge and communication gap exists between scientists and non-scientists as to how climate change impact scenarios can be interpreted and what they imply for European forests. It is still challenging to advise forest decision makers on how best to plan for climate change as many uncertainties and unknowns remain and it is difficult to communicate these to practitioners and other decision makers while retaining emphasis on the importance of planning for adaptation. In this paper, recent developments in climate change observations and projections, observed and projected impacts on European forests and the associated uncertainties are reviewed and synthesised with a view to understanding the implications for forest management. Current impact assessments with simulation models contain several simplifications, which explain the discrepancy between results of many simulation studies and the rapidly increasing body of evidence about already observed changes in forest productivity and species distribution. In simulation models uncertainties tend to cascade onto one another; from estimating what future societies will be like and general circulation models (GCMs) at the global level, down to forest models and forest management at the local level. Individual climate change impact studies should not be uncritically used for decision-making without reflection on possible shortcomings in system understanding, model accuracy and other assumptions made. It is important for decision makers in forest management to realise that they have to take long-lasting management decisions while uncertainty about climate change impacts are still large. We discuss how to communicate about uncertainty - which is imperative for decision making - without diluting the overall message. Considering the range of possible trends and uncertainties in adaptive forest management requires expert knowledge and enhanced efforts for providing science-based decision support.

Updating beliefs and combining evidence in adaptive forest management under climate change: a case study of Norway spruce (Picea abies L. Karst) in the Black Forest, Germany.

We study climate uncertainty and how managers' beliefs about climate change develop and influence their decisions. We develop an approach for updating knowledge and beliefs based on the observation of forest and climate variables and illustrate its application for the adaptive management of an even-aged Norway spruce (Picea abies L. Karst) forest in the Black Forest, Germany. We simulated forest development under a range of climate change scenarios and forest management alternatives. Our analysis used Bayesian updating and Dempster's rule of combination to simulate how observations of climate and forest variables may influence a decision maker's beliefs about climate development and thereby management decisions. While forest managers may be inclined to rely on observed forest variables to infer climate change and impacts, we found that observation of climate state, e.g. temperature or precipitation is superior for updating beliefs and supporting decision-making. However, with little conflict among information sources, the strongest evidence would be offered by a combination of at least two informative variables, e.g., temperature and precipitation. The success of adaptive forest management depends on when managers switch to forward-looking management schemes. Thus, robust climate adaptation policies may depend crucially on a better understanding of what factors influence managers' belief in climate change.

High economic costs of reduced carbon sinks and declining biome stability in Central American forests.

Tropical forests represent important supporting pillars for society, supplying global ecosystem services (ES), e.g., as carbon sinks for climate regulation and as crucial habitats for unique biodiversity. However, climate change impacts including implications for the economic value of these services have been rarely explored before. Here, we derive monetary estimates for the effect of climate change on climate regulation and habitat services for the forests of Central America. Our results projected ES declines in 24-62% of the study region with associated economic costs of $51-314 billion/year until 2100. These declines particularly affected montane and dry forests and had strong economic implications for Central America's lower-middle income countries (losses of up to 335% gross domestic product). In addition, economic losses were mostly higher for habitat services than for climate regulation. This highlights the need to expand the focus from mere maximization of CO2 sequestration and avoid false incentives from carbon markets.

Sharp decline in future productivity of tropical reforestation above 29°C mean annual temperature.

Tropical reforestation is among the most powerful tools for carbon sequestration. Yet, climate change impacts on productivity are often not accounted for when estimating its mitigation potential. Using the process-based forest growth model 3-PGmix, we analyzed future productivity of tropical reforestation in Central America. Around 29°C mean annual temperature, productivity sharply and consistently declined (-11% per 1°C of warming) across all tropical lowland climate zones and five tree species spanning a wide range of ecological characteristics. Under a high-emission scenario (SSP3-7.0), productivity of dry tropical reforestation nearly halved and tropical moist and rain forest sites showed moderate losses around 10% by the end of the century. Under SSP2-4.5, tropical moist and rain forest sites were resilient and tropical dry forest sites showed moderate losses (-17%). Increased vapor pressure deficit, caused by increasing temperatures, was the main driver of growth decline. Thus, to continue following high-emission pathways could reduce the effectiveness of reforestation as climate action tool.

Climate change may induce connectivity loss and mountaintop extinction in Central American forests.

The tropical forests of Central America serve a pivotal role as biodiversity hotspots and provide ecosystem services securing human livelihood. However, climate change is expected to affect the species composition of forest ecosystems, lead to forest type transitions and trigger irrecoverable losses of habitat and biodiversity. Here, we investigate potential impacts of climate change on the environmental suitability of main plant functional types (PFTs) across Central America. Using a large database of occurrence records and physiological data, we classify tree species into trait-based groups and project their suitability under three representative concentration pathways (RCPs 2.6, 4.5 and 8.5) with an ensemble of state-of-the-art correlative modelling methods. Our results forecast transitions from wet towards generalist or dry forest PFTs for large parts of the study region. Moreover, suitable area for wet-adapted PFTs is projected to latitudinally diverge and lose connectivity, while expected upslope shifts of montane species point to high risks of mountaintop extinction. These findings underline the urgent need to safeguard the connectivity of habitats through biological corridors and extend protected areas in the identified transition hotspots.

Evaluating the suitability of management strategies of pure Norway spruce forests in the Black Forest area of southwest Germany for adaptation to or mitigation of climate change.

The study deals with the problem of evaluating management strategies for pure stands of Norway spruce (Picea abies Karst) to balance adaptation to and mitigation of climate change, taking into account multiple objectives of a forest owner. A simulation and optimization approach was used to evaluate the management of a 1000 ha model Age-Class forest, representing the age-class distribution of an area of 66,000 ha of pure Norway spruce forests in the Black Forest region of Southwest Germany. Eight silvicultural scenarios comprising five forest conversion schemes which were interpreted as "adaptation" strategies which aims at increasing the proportion of Beech, that is expected to better cope with climate change than the existing Norway spruce, and three conventional strategies including a "Do-nothing" alternative classified as "mitigation", trying to keep rather higher levels of growing stock of spruce, were simulated using the empirical growth simulator BWINPro-S. A linear programming approach was adapted to simultaneously maximize the net present values of carbon sequestration and timber production subject to the two constraints of wood even flow and partial protection of the oldest (nature protection). The optimized plan, with the global utility of 11,687 /ha in forty years, allocated a combination of silvicultural scenarios to the entire forest area. Overall, strategies classified as "mitigation" were favored, while strategies falling into the "adaptation"-category were limited to the youngest age-classes in the optimal solution. Carbon sequestration of the "Do-nothing" alternative was between 1.72 and 1.85 million tons higher than the other alternatives for the entire forest area while the differences between the adaptation and mitigation approaches were approximately 133,000 tons. Sensitivity analysis showed that a carbon price of 21 /t is the threshold at which carbon sequestration is promoted, while an interest rate of above 2% would decrease the amount of carbon.

Retention as an integrated biodiversity conservation approach for continuous-cover forestry in Europe.

Retention forestry implies that biological legacies like dead and living trees are deliberately selected and retained beyond harvesting cycles to benefit biodiversity and ecosystem functioning. This model has been applied for several decades in even-aged, clearcutting (CC) systems but less so in uneven-aged, continuous-cover forestry (CCF). We provide an overview of retention in CCF in temperate regions of Europe, currently largely focused on habitat trees and dead wood. The relevance of current meta-analyses and many other studies on retention in CC is limited since they emphasize larger patches in open surroundings. Therefore, we reflect here on the ecological foundations and socio-economic frameworks of retention approaches in CCF, and highlight several areas with development potential for the future. Conclusions from this perspective paper, based on both research and current practice on several continents, although highlighting Europe, are also relevant to other temperate regions of the world using continuous-cover forest management approaches.

Evaluating the effectiveness of retention forestry to enhance biodiversity in production forests of Central Europe using an interdisciplinary, multi-scale approach.

Retention forestry, which retains a portion of the original stand at the time of harvesting to maintain continuity of structural and compositional diversity, has been originally developed to mitigate the impacts of clear-cutting. Retention of habitat trees and deadwood has since become common practice also in continuous-cover forests of Central Europe. While the use of retention in these forests is plausible, the evidence base for its application is lacking, trade-offs have not been quantified, it is not clear what support it receives from forest owners and other stakeholders and how it is best integrated into forest management practices. The Research Training Group ConFoBi (Conservation of Forest Biodiversity in Multiple-use Landscapes of Central Europe) focusses on the effectiveness of retention forestry, combining ecological studies on forest biodiversity with social and economic studies of biodiversity conservation across multiple spatial scales. The aim of ConFoBi is to assess whether and how structural retention measures are appropriate for the conservation of forest biodiversity in uneven-aged and selectively harvested continuous-cover forests of temperate Europe. The study design is based on a pool of 135 plots (1 ha) distributed along gradients of forest connectivity and structure. The main objectives are (a) to investigate the effects of structural elements and landscape context on multiple taxa, including different trophic and functional groups, to evaluate the effectiveness of retention practices for biodiversity conservation; (b) to analyze how forest biodiversity conservation is perceived and practiced, and what costs and benefits it creates; and (c) to identify how biodiversity conservation can be effectively integrated in multi-functional forest management. ConFoBi will quantify retention levels required across the landscape, as well as the socio-economic prerequisites for their implementation by forest owners and managers. ConFoBi's research results will provide an evidence base for integrating biodiversity conservation into forest management in temperate forests.

Diversification of forest management regimes secures tree microhabitats and bird abundance under climate change.

The loss of biodiversity in temperate forests due to combined effect of climate change and forest management poses a major threat to the functioning of these ecosystems in the future. Climate change is expected to modify ecological processes and amplify disturbances, compromising the provisioning of multiple ecosystem services. Here we investigate the impacts of climate change and forest management on the abundance of tree microhabitats and forest birds as biodiversity proxies, using an integrated modelling approach. To perform our analysis, we calibrated tree microhabitat and bird abundance in a forest landscape in Southwestern Germany, and coupled them with a climate sensitive forest growth model. Our results show generally positive impacts of climate warming and higher harvesting intensity on bird abundance, with up to 30% increase. Conversely, climate change and wood removals above 5% of the standing volume led to a loss of tree microhabitats. A diversified set of management regimes with different harvesting intensities applied in a landscape scale was required to balance this trade-off. For example, to maximize the expected bird abundance (up to 11%) and to avoid tree microhabitat abundance loss of >20% necessitates setting aside 10.2% of the forest area aside and application of harvesting intensities < 10.4% of the standing volume. We conclude that promoting forest structural complexity by diversifying management regimes across the landscape will be key to maintain forest biodiversity in temperate forests under climate change.

Multiple uncertainties require a change of conservation practices for saproxylic beetles in managed temperate forests.

In Europe, intensive forest management has severely compromised the habitat of forest insects, especially saproxylic beetles, due to the removal of deadwood and veteran trees. The loss of insect diversity may disrupt ecosystem functioning and affect the provision of important ecosystem goods and services in the future. Here we propose a novel approach for the implementation of conservation policies, by optimally allocating forest reserves and deadwood islands under multiple sources of uncertainty and minimizing economic risk. We use the saproxylic beetle Lucanus cervus as umbrella species, requiring that deadwood islands were spaced within its dispersal capacity. We show that current management and conservation practices are increasingly inefficient under changing environmental conditions and that the consideration of uncertainty requires a major expansion of conservation areas. Moreover, our results indicate that a strong diversification of management regimes, with a focus on selection forest systems, is required to reduce economic risk of forest management. We conclude that the integration of uncertainty into conservation planning may reduce the trade-off between production and conservation objectives in forest landscapes and is key to increase the efficiency of forest management in the future.

Realizing Mitigation Efficiency of European Commercial Forests by Climate Smart Forestry.

European temperate and boreal forests sequester up to 12% of Europe's annual carbon emissions. Forest carbon density can be manipulated through management to maximize its climate mitigation potential, and fast-growing tree species may contribute the most to Climate Smart Forestry (CSF) compared to slow-growing hardwoods. This type of CSF takes into account not only forest resource potentials in sequestering carbon, but also the economic impact of regional forest products and discounts both variables over time. We used the process-based forest model 4 C to simulate European commercial forests' growth conditions and coupled it with an optimization algorithm to simulate the implementation of CSF for 18 European countries encompassing 68.3 million ha of forest (42.4% of total EU-28 forest area). We found a European CSF policy that could sequester 7.3-11.1 billion tons of carbon, projected to be worth 103 to 141 billion euros in the 21st century. An efficient CSF policy would allocate carbon sequestration to European countries with a lower wood price, lower labor costs, high harvest costs, or a mixture thereof to increase its economic efficiency. This policy prioritized the allocation of mitigation efforts to northern, eastern and central European countries and favored fast growing conifers Picea abies and Pinus sylvestris to broadleaves Fagus sylvatica and Quercus species.

Are forest disturbances amplifying or canceling out climate change-induced productivity changes in European forests?

Recent studies projecting future climate change impacts on forests mainly consider either the effects of climate change on productivity or on disturbances. However, productivity and disturbances are intrinsically linked because 1) disturbances directly affect forest productivity (e.g. via a reduction in leaf area, growing stock or resource-use efficiency), and 2) disturbance susceptibility is often coupled to a certain development phase of the forest with productivity determining the time a forest is in this specific phase of susceptibility. The objective of this paper is to provide an overview of forest productivity changes in different forest regions in Europe under climate change, and partition these changes into effects induced by climate change alone and by climate change and disturbances. We present projections of climate change impacts on forest productivity from state-of-the-art forest models that dynamically simulate forest productivity and the effects of the main European disturbance agents (fire, storm, insects), driven by the same climate scenario in seven forest case studies along a large climatic gradient throughout Europe. Our study shows that, in most cases, including disturbances in the simulations exaggerate ongoing productivity declines or cancel out productivity gains in response to climate change. In fewer cases, disturbances also increase productivity or buffer climate-change induced productivity losses, e.g. because low severity fires can alleviate resource competition and increase fertilization. Even though our results cannot simply be extrapolated to other types of forests and disturbances, we argue that it is necessary to interpret climate change-induced productivity and disturbance changes jointly to capture the full range of climate change impacts on forests and to plan adaptation measures.

Forest Owners' Response to Climate Change: University Education Trumps Value Profile.

Do forest owners' levels of education or value profiles explain their responses to climate change? The cultural cognition thesis (CCT) has cast serious doubt on the familiar and often criticized "knowledge deficit" model, which says that laypeople are less concerned about climate change because they lack scientific knowledge. Advocates of CCT maintain that citizens with the highest degrees of scientific literacy and numeracy are not the most concerned about climate change. Rather, this is the group in which cultural polarization is greatest, and thus individuals with more limited scientific literacy and numeracy are more concerned about climate change under certain circumstances than those with higher scientific literacy and numeracy. The CCT predicts that cultural and other values will trump the positive effects of education on some forest owners' attitudes to climate change. Here, using survey data collected in 2010 from 766 private forest owners in Sweden and Germany, we provide the first evidence that perceptions of climate change risk are uncorrelated with, or sometimes positively correlated with, education level and can be explained without reference to cultural or other values. We conclude that the recent claim that advanced scientific literacy and numeracy polarizes perceptions of climate change risk is unsupported by the forest owner data. In neither of the two countries was university education found to reduce the perception of risk from climate change. Indeed in most cases university education increased the perception of risk. Even more importantly, the effect of university education was not dependent on the individuals' value profile.

Climate Change Impairs Nitrogen Cycling in European Beech Forests.

European beech forests growing on marginal calcareous soils have been proposed to be vulnerable to decreased soil water availability. This could result in a large-scale loss of ecological services and economical value in a changing climate. In order to evaluate the potential consequences of this drought-sensitivity, we investigated potential species range shifts for European beech forests on calcareous soil in the 21st century by statistical species range distribution modelling for present day and projected future climate conditions. We found a dramatic decline by 78% until 2080. Still the physiological or biogeochemical mechanisms underlying the drought sensitivity of European beech are largely unknown. Drought sensitivity of beech is commonly attributed to plant physiological constraints. Furthermore, it has also been proposed that reduced soil water availability could promote nitrogen (N) limitation of European beech due to impaired microbial N cycling in soil, but this hypothesis has not yet been tested. Hence we investigated the influence of simulated climate change (increased temperatures, reduced soil water availability) on soil gross microbial N turnover and plant N uptake in the beech-soil interface of a typical mountainous beech forest stocking on calcareous soil in SW Germany. For this purpose, triple 15N isotope labelling of intact beech seedling-soil-microbe systems was combined with a space-for-time climate change experiment. We found that nitrate was the dominant N source for beech natural regeneration. Reduced soil water content caused a persistent decline of ammonia oxidizing bacteria and therefore, a massive attenuation of gross nitrification rates and nitrate availability in the soil. Consequently, nitrate and total N uptake of beech seedlings were strongly reduced so that impaired growth of beech seedlings was observed already after one year of exposure to simulated climatic change. We conclude that the N cycle in this ecosystem and here specifically nitrification is vulnerable to reduced water availability, which can directly lead to nutritional limitations of beech seedlings. This tight link between reduced water availability, drought stress for nitrifiers, decreased gross nitrification rates and nitrate availability and finally nitrate uptake by beech seedlings could represent the Achilles' heel for beech under climate change stresses.