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1. There is increasing evidence that species diversity enhances the temporal stability (TS) of community productivity in different ecosystems, although its effect at the population and tree levels seems to be negative or neutral. Asynchrony in species responses to environmental conditions was found to be one of the main drivers of this stabilizing process. However, the effect of species mixing on the stability of productivity, and the relative importance of the associated mechanisms, remain poorly understood in forest communities. 2. We investigated the way mixing species influenced the TS of productivity in Pinus sylvestris L. and Fagus sylvatica L. forests, and attempted to determine the main drivers among overyielding, asynchrony between species annual growth responses to environmental conditions, and temporal shifts in species interactions. We used a network of 93 experimental plots distributed across Europe to compare the TS of basal area growth over a 15-year period (1999-2013) in mixed and monospecific forest stands at different organizational levels, namely the community, population and individual tree levels. 3. Mixed stands showed a higher TS of basal area growth than monospecific stands at the community level, but not at the population or individual tree levels. The TS at the community level was related to asynchrony between species growth in mixtures, but not to overyielding nor to asynchrony between species growth in monospecific stands. Temporal shifts in species interactions were also related to asynchrony and to the mixing effect on the TS. 4. Synthesis. Our findings confirm that species mixing can stabilize productivity at the community level, whereas there is a neutral or negative effect on stability at the population and individual tree levels. The contrasting findings regarding the relationships between the temporal stability and asynchrony in species growth in mixed and monospecific stands suggest that the main driver in the stabilizing process may be the temporal niche complementarity between species rather than differences in species' intrinsic responses to environmental conditions.

Drought frequency and intensity are predicted to increase in many parts of the Northern Hemisphere and the effects of such changes on forest growth and tree mortality are already evident in many regions around the world. Mixed-species forests and increasing tree species diversity have been put forward as important risk reduction and adaptation strategies in the face of climate change. However, little is known about whether the species interactions that occur in diverse forests will reduce drought susceptibility or water stress. In this study, we focused on the effect of drought on individual tree species (n=16) within six regions of Europe and assessed whether this response was related to tree species diversity and stand density, and whether community-level responses resulted from many similar or contrasting species-level responses. For each species in each plot, we calculated the increase in carbon isotope composition of latewood from a wet to a dry year (C-13) as an estimate of its drought stress level. When significant community-level relationships occurred (three of six regions), there was only one species within the given community that showed a significant relationship (three of 25 species-region combinations), showing that information about a single species can be a poor indicator of the response of other species or the whole community. There were many two-species mixtures in which both species were less water-stressed compared with their monocultures, but also many mixtures where both species were more stressed compared with their monocultures. Furthermore, a given species combination responded differently in different regions.Synthesis and applications. Our study shows that drought stress may sometimes be reduced in mixed-species forests, but this is not a general pattern, and even varies between sites for a given combination. The management or prediction of drought stress requires consideration of the physiological characteristics of the mixed species, and how this complements the water-related climatic and edaphic features of the site, rather than species richness.Our study shows that drought stress may sometimes be reduced in mixed-species forests, but this is not a general pattern, and even varies between sites for a given combination. The management or prediction of drought stress requires consideration of the physiological characteristics of the mixed species, and how this complements the water-related climatic and edaphic features of the site, rather than species richness.

Questions: We aim for a better understanding of the different modes of intra- and inter-specific competition in two- and three-species mixed-forests. How can the effect of different modes of competitive interactions be detected and integrated into individual tree growth models? Are species interactions in spruce–fir–beech forests more associated with size-symmetric or size-asymmetric competition? Do competitive interactions between two of these species change from two- to three-species mixtures? Location: Temperate mixed-species forests in Central Europe (Switzerland). Methods: We used data from the Swiss National Forest Inventory to fit basal area increment models at the individual tree level, including the effect of ecological site conditions and indices of size-symmetric and size-asymmetric competition. Interaction terms between species-specific competition indices were used to disentangle significant differences in species interactions from two- to three-species mixtures. Results: The growth of spruce and fir was positively affected by increasing proportions of the other species in spruce–fir mixtures, but negative effects were detected with increasing presence of beech. We found that competitive interactions for spruce and fir were more related to size-symmetric competition, indicating that species interactions might be more associated with competition for below-ground resources. Under constant amounts of stand basal area, the growth of beech clearly benefited from the increasing admixture of spruce and fir. For this species, patterns of size-symmetric and size-asymmetric competitive interactions were similar, indicating that beech is a strong self-competitor for both above-ground and below-ground resources. Only for silver fir and beech, we found significant changes in species interactions from two- to three-species mixtures, but these were not as prominent as the effects due to differences between intra- and inter-specific competition. Conclusions: Species interactions in spruce–fir–beech, or other mixed forests, can be characterized depending on the mode of competition, allowing interpretations of whether they occur mainly above or below ground level. Our outcomes illustrate that species-specific competition indices can be integrated in individual tree growth functions to express the different modes of competition between species, and highlight the importance of considering the symmetry of competition alongside competitive interactions in models aimed at depicting growth in mixed-species forests.

1. Mixed species forests can often be more productive and deliver higher levels of ecosystem services and functions than monocultures. However, complementarity effects for any given tree species are difficult to generalize because they can vary greatly along gradients of climatic conditions and resource availability. Identifying the conditions where species diversity can positively influence productivity is crucial. To date, few studies have examined how growth complementarity across species and mixture types is modulated by stand and environmental factors, and fewer have considered more than one or two factors. 2. We investigated how complementarity effects for several major Central European tree species change with climatic and edaphic conditions, and with stand structural characteristics, including species composition. We used data from the Swiss National Forest Inventory, which is based on 3,231 plots of pure and mixed stands (19 mixture types) across a broad environmental gradient, to test (i) how mixing effects change depending on the identity of the admixed species and (ii) if complementarity consistently increases when environmental conditions become harsher. 3. The magnitude, whether positive or negative, of complementarity increased with increasing stand density and stand developmental stage, but no general pattern could be identified across mixture types. Complementarity for many species increased as drought intensity and temperature increased, but not for all species and mixture types. While soil conditions, nitrogen and site topography influenced complementarity for many species, there was no general pattern (increases and decreases were observed). 4. Synthesis. Our study indicates that complementarity varies strongly with stand density and stand development as well as with topographic, climatic and soil conditions. This emphasizes the need to account for site-dependent conditions when exploring mixture effects in relation to forest productivity. We found that under certain conditions (i.e. increasing drought, higher temperature), mixed forests can promote individual tree growth in Central European temperate forests. However, careful assessments depending on the species composing the stands are required under changing resource availability as well as under different levels of stand density and development.

Aim We assessed how avian biodiversity and above-ground carbon storage were related in different forest age-classes, including mature stands (> 100 years), in a managed, mixed-species eucalypt forest. Location Gippsland, south-eastern Australia. Methods In 50 2-ha stands ranging in age from ≤ 5 years to mature stands > 100 years, we undertook repeated avian surveys, performed detailed habitat measurements and estimated amounts of above-ground carbon. Extensive wildfire reduced the number of sites to 28 (seven in each of four age classes) upon which analyses and inferences were made. We also analysed data on carbon storage and some bird responses from previously published studies. Results Mature vegetation (> 100 years) had the greatest richness, abundance and biomass of birds. Key ecological resources, such as tree-hollows for nesting, generally occurred mostly in stands > 60 years. Avian richness per unit of above-ground carbon storage was relatively low for stands of 20-60 years. While above-ground carbon storage appeared to increase in a monotonic fashion as stands age and mature, there were quantum increases in all measures of avian biodiversity in mature stands (> 100 years). Main conclusions Our results suggest that carbon is organized in a different way, with substantially greater biodiversity benefits, in very old stands. Mature vegetation simultaneously maximizes both avian biodiversity and above-ground carbon storage. These results bolster arguments for allocating highest priorities to the preservation of old-growth forest stands rather than alternative investments (e.g. reafforestation for carbon sequestration).

Process-based forest models are important tools for predicting forest growth and their vulnerability to factors such as climate change or responses to management. One of the most widely used stand-level process-based models is the 3-PG model (Physiological Processes Predicting Growth), which is used for applications including estimating wood production, carbon budgets, water balance and susceptibility to climate change. Few 3-PG parameter sets are available for central European species and even fewer are appropriate for mixed-species forests. Here we estimated 3-PG parameters for twelve major central European tree species using 1418 long-term permanent forest monitoring plots from managed forests, 297 from un-managed forest reserves and 784 Swiss National Forest Inventory plots. A literature review of tree physiological characteristics, as well as regression analyses and Bayesian inference, were used to calculate the 3-PG parameters.The Swiss-wide calibration, based on monospecific plots, showed a robust performance in predicting forest stocks such as stem, foliage and root biomass. The plots used to inform the Bayesian calibration resulted in posterior ranges of the calibrated parameters that were, on average, 69% of the prior range. The bias of stem, foliage and root biomass predictions was generally less than 20%, and less than 10% for several species. The parameter sets also provided reliable predictions of biomass and mean tree sizes in mixed-species forests. Given that the information sources used to develop the parameters included a wide range of climatic, edaphic and management conditions and long time spans (from 1930 to present), these species parameters for 3-PG are likely to be appropriate for most central European forests and conditions.

Mixed-species forests can provide higher levels of ecosystem functions and services and can be more resistant and resilient in the face of global change. While many studies focus on the growth and yield of mixed forests, fewer have examined the underlying processes. Inter- or intra-specific differences and interactions influence tree- and stand-level light absorption by determining the vertical structure of stratified canopies, stand density, leaf area index, and the size or allometry of trees. While canopy light absorption is a very important process, it is difficult to quantify it for individual species within a mixture and is rarely examined. A detailed tree-level model (MAESTRA) was used in combination with measurements of tree sizes and stand structures to examine effects of mixing on absorbed photosynthetically active radiation (APAR) in 41–63-year-old stands of Pseudotsuga menziesii and Fagus sylvatica at three sites in Bavaria, Germany. The effects of initial stand density on APAR were analysed in 46-year-old P. menziesii stands of a spacing experiment at two sites. At the tree level, mixing increased mean height and leaf area, growth (185% higher) and APAR (85% higher) of P. menziesii at all sites. Mean tree heights and crown sizes of F. sylvatica were larger in mixtures, while recent growth rates and APAR were not significantly different to monocultures. Planting density did not influence mean tree variables (e.g. height, leaf area, crown volume), because any initial spacing effects had been gradually removed by thinning across all treatments. At the stand level, there were no differences in growth, basal area or in the annual growth per annual APAR (light use efficiency, LUE) between monocultures and mixtures. The highest APAR values were observed in P. menziesii monocultures, while the lowest APAR values were observed in F. sylvatica monocultures. While mixing these species may not increase stand-level growth during later phases of development, mixing accelerated initial growth of individual trees and reduced the time to reach target diameters, which are both important aspects in adapting forests to global change.

The role of trees, in addition to that of the soil, must be considered in CH4 budget for forests. Although trees can emit CH4 through their stems, there are uncertainties about the main factors that explain inter- and intraspecific variations, which impedes upscaling of measurements from the stem to the ecosystem level. This study aimed to characterize the variability in CH4 emissions (FCH4) from stems between species and individuals, and within individuals. We measured FCH4 in situ during the snow-free period in five species in a temperate mountain forest, using individuals of different sizes and chambers at different heights along the stems. One coniferous species emitted almost no CH4, whereas four broadleaved species exhibited high intraspecific variability in FCH4 (0–3.7 nmol m−2 s−1). Increasing trends in FCH4 with tree diameter were observed for four species. The vertical patterns in FCH4 were complex. Seasonal variations in FCH4, measured on two trees per species, were well explained by air temperature with apparent temperature sensitivity coefficients (Q10) between 1.2 and 2, which were not related to the antecedent precipitation indices, whether calculated over 7 or 30 days. Potential CH4 production was detected in wood core segments incubated under anoxic conditions in the majority of individual trees of all species. Our results suggest that the CH4 emitted by trunks can originate either from soil or internal sources. Scaling FCH4 from trees at the stand level and developing process-based models of FCH4 will remain challenging until the sources of variation are better explained.

Disturbance to spruce forests from wind and bark beetles is projected to worsen. It has been suggested that mixed-species forests could provide a more disturbance-resilient option than spruce monocultures. We used dynamic optimization to study how profitable mixed forests are compared to pure spruce forests by examining two similar neighboring stands facing a windthrow risk. We found that under high current windthrow risk levels in northeastern Finland, Norway spruce (Picea abies)-only forests are more profitable than mixed forests consisting of spruce and silver birch (Betula pendula). However, if the windthrow risk to spruce trees increases by 35% compared to its current level, and the risk level of birch remains at its current level, mixed forests become more profitable. When carbon is priced at €50 per ton of CO2, the additional income from carbon storage in mixed forests outweighs the economic advantage of spruce timber, making mixed forests more profitable—even if the risk to spruce increases by only 25%. Hence, mixed-species forests become increasingly profitable as carbon prices rise. Therefore, mixed-species forestry represents a cost-efficient strategy for adapting to increasing environmental risks and mitigating climate change.

Functional trait-based approaches are extensively applied to the study of mechanisms governing community assembly along environmental gradients. These approaches have been classically based on studying differences in mean values among species, but there is increasing recognition that alternative metrics of trait distributions should be considered to decipher the mechanisms determining community assembly and species coexistence. Under this framework, the main aim of this study is to unravel the effects of environmental conditions as drivers of plant community assembly in sub-Mediterranean ecotones. We set 60 plots in six plant communities of a sub-Mediterranean forest in Central Spain, and measured key above- and belowground functional traits in 411 individuals belonging to 19 species, along with abiotic variables. We calculated community-weighted mean (CWM), skewness (CWS) and kurtosis (CWK) of three plant dimensions, and used maximum likelihood techniques to analyze how variation in these functional community traits was driven by abiotic factors. Additionally, we estimated the relative contribution of intraspecific trait variability and species turnover to variation in CWM. The first three axes of variation of the principal component analyses were related to three main plant ecological dimensions: Leaf Economics Spectrum, Root Economics Spectrum and plant hydraulic architecture, respectively. Type of community was the most important factor determining differences in the functional structure among communities, as compared to the role of abiotic variables. We found strong differences among communities in their CWMs in line with their biogeographic origin (Eurosiberian vs Mediterranean), while differences in CWS and CWK indicate different trends in the functional structure among communities and the coexistence of different functional strategies, respectively. Moreover, changes in functional composition were primarily due to intraspecific variability. We observed a high number of strategies in the forest with the different communities spreading along the acquisitive-conservative axis of resource-use, partly matching their Eurosiberian-Mediterranean nature, respectively. Intraspecific trait variability, rather than species turnover, stood as the most relevant factor when analyzing functional changes and assembly patterns among communities. Altogether, our data support the notion that ecotones are ecosystems where relatively minor environmental shifts may result in changes in plant and functional composition.