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Penalized regression splines and distributed lag models were used to evaluate the effects of species mixing on productivity and climate-related resistance via tree-ring width measurements from sample cores. Data were collected in Lower Austria from sample plots arranged in a triplet design. Triplets were established for sessile oak [Quercus petraea (Matt.) Liebl.] and Scots pine (Pinus sylvestris L.), European beech (Fagus sylvatica L.) and Norway spruce [Picea abies (L.) H. Karst.], and European beech and European larch (Larix decidua Mill.). Mixing shortened the temporal range of time-lagged climate effects for beech, spruce, and larch, but only slightly changed the effects for oak and pine. Beech and spruce as well as beech and larch exhibited contrasting climate responses, which were consequently reversed by mixing. Single-tree productivity was reduced by between − 15% and − 28% in both the mixed oak–pine and beech–spruce stands but only slightly reduced in the mixed beech–larch stands. Measures of climate sensitivity and resistance were derived by model predictions of conditional expectations for simulated climate sequences. The relative climate sensitivity was, respectively, reduced by between − 16 and − 39 percentage points in both the beech–spruce and beech–larch mixed stands. The relative climate sensitivity of pine increased through mixing, but remained unaffected for oak. Mixing increased the resistance in both the beech–larch and the beech–spruce mixed stand. In the mixed oak–pine stand, resistance of pine was decreased and remained unchanged for oak.

15 Pág. Ecological Modelling, Tree species mixing of oak (Quercus petraea (Matt.) Liebl./Quercus robur L.) and pine (Pinus sylvestris L.) has been shown to have positive effects on ecosystem service provision. From a management perspective, however, it is still uncertain which thinning regime provides the highest possible productivity of mixed oak–pine forests in the long term. Because of a lack of empirical studies dealing with thinning and species mixing effects on oak–pine forests, we simulated forest growth in order to test which thinning type and intensity may provide the highest productivity in the long-term. To achieve this, we simulated the growth of pure and mixed stands of oak and pine for 100 years in 23 triplets located on an ecological gradient across Europe. For this purpose, we applied four different growth simulators and compared their results: the distance-independent single-tree simulator PROGNAUS, the distance-dependent single-tree simulator SILVA, the gap model ForCEEPS, and the process-based simulator 3D-CMCC-FEM. We investigated the effects of species mixing and thinning from the upper (thinning from above) and lower tail (thinning from below) of the diameter distribution by reducing the stand basal area to 50 and 80% of the maximum basal area. We compared simulated results of the relative volume productivity of mixed versus pure stands and of thinned versus unthinned stands to empirical results previously obtained on the same set of triplets. Simulated relative volume productivity ranged between 61 and 156%, although extremes of 10% and of 300% could be observed. We found the relative volume productivity to be influenced by stand age, but not by stand density, except for PROGNAUS. Relative volume productivity did not increase with the site water supply of the triplet location. Highest long-term productivity for oak, pine and oak–pine stands can be expected in consequence of thinning from above, but the effect of thinning intensity differed between simulators. Thinning effects were positively affected by stand density, but not by stand age, except for thinning from above predicted by PROGNAUS. Predicted thinning effects showed good approximation of results from thinning experiments for oak, but not for pine stands. We hypothesize the results might be caused by the insufficient simulator representation of climate and its interaction with other site variables and stand structure. Further work is needed to reduce the revealed limitations of the existing growth models, as we currently see no alternative to such kind of studies and simulators., The authors thank the European Union for funding the project “Mixed species forest management. Lowering risk, increasing resilience (REFORM)” under the framework of Sumforest ERA-NET. The authors from Austria also thank the Austrian Federal Ministry for Sustainability and Tourism for supporting the establishment of the Austrian triplet plots and for covering the work expenses of Markus Engel within the project ”Forstwirtschaft mit Mischwäldern – geringes Risiko, hohe Widerstandskraft – REFORM” under the grant number 101199. All contributors thank their national funding institutions to establish, measure and analyze data from the triplets. Maude Toïgo and Xavier Morin thank François de Coligny and Nicolas Beudez for their help in the development of the ForCEEPS model. Antonio Tomao, Carlo Trotta and Anna Barbati thank Alessio Collalti for his support and suggestions about the simulations with the 3D-CMCC-FEM model.

Mixed forests are suggested as a strategic adaptation of forest management to climate change. Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) are tree species of high economic and ecological value for European forestry. Both species coexist naturally in a large part of their distributions but there is a lack of knowledge on the ecological functioning of mixtures of these species and how to manage such stands. This paper analyses these species’ intra- and inter-specific competition, including size-symmetric vs. size-asymmetric competition, and explore the effect of weather conditions on tree growth and competition. We studied basal area growth at tree level for Scots pine and Norway spruce in mixed versus pure stands in 22 triplets of fully-stocked plots along a broad range of ecological conditions across Europe. Stand inventory and increment cores provided insights into how species mixing modifies tree growth compared with neighbouring pure stands. Five different competition indices, weather variables and their interactions were included and checked in basal area growth models using a linear mixed model approach. Interspecific size-asymmetric competition strongly influenced growth for both tree species, and was modulated by weather conditions. However, species height stratification in mixed stands resulted in a greater tree basal area growth of Scots pine (10.5 cm2 year−1) than in pure stands (9.3 cm2 year−1), as this species occupies the upper canopy layer. Scots pine growth depended on temperature and drought, whereas Norway spruce growth was influenced only by drought. Interspecific site-asymmetric competition increased in cold winters for Scots pine, and decreased after a drought year for Norway spruce. Although mixtures of these species may reduce tree size for Norway spruce, our results suggest that this could be offset by faster growth in Scots pine. How inter-specific competition and weather conditions alter tree growth may have strong implications for the management of Scots pine-Norway spruce mixtures along the rotation period into the ongoing climate change scenario.

Centro de Investigación Forestal (CIFOR), Mixed-species stands have been found to be more productive than would be expected from the performance of their component species in monocultures due to facilitation and complementarity between species, although these interactions depend on the combination of species present. Our study focuses on monospecific and mixed-species stands of Scots pine and Norway spruce using 20 triplets established in nine countries along a climatic gradient across Europe. Differences in mean tree and stand characteristics, productivity and stand structure were assessed. Basal area increment in mixed stands was 8% higher than expected while volume increment was only 2% greater. Scots pine trees growing in mixed-species stands showed 11% larger quadratic mean diameter, 7% larger dominant diameter, 17% higher basal area and 25% higher stand volume than trees growing in monospecific stands. Norway spruce showed only a non-significant tendency to lower mean values of diameters, heights, basal area, as well standing volume in mixtures than monocultures. Stand structure indices differed between mixed stands and monocultures of Scots pine showing a greater stratification in mixed-species stands. Furthermore, the studied morphological traits showed little variability for trees growing in monospecific stands, except for diameter at breast height, crown length and crown length ratio. For trees growing in mixed stands, all the morphological traits of the trees were identified as different. Some of these morphological traits were associated with relative productivity. Nevertheless, relative productivity in mixed-species stands was not related to site conditions., We thank national funders of REFORM project (Spanish Ministry of Science and Innovation: PCIN2017-026; Research Council of Lithuania: S-SUMFOREST-17-1) and those funders for the support to non-participants in the REFORM project (Denmark: Contract between Danish Ministry of Environment and Food and Department of Geosciences and Natural Resource Management of UCPH; Estonia: Estonian University of Life Sciences projects number P180024MIME and P200029MIME; Poland: EU CARE4C project (GA 778322) supported by the Polish Government MNiSW2018-2021 matching fund (W117/H2020/2018); Slovakia: APVV-15-0265) that allowed the establishment and measurement of the triplets. A grant for a research stay was received by RR-P from the CARE4C project (Carbon smart forestry under climate change, H2020-MSCA-RISE-2017)., 13 Pág.

Centro de Investigación Forestal (CIFOR), Despite growing evidence that diverse forests play an important role in ecosystem functioning, ensuring the provision of different ecosystem services, whether such diversity improves their response to drought events remains unclear. In this study, we use a large tree-ring database from thirty case studies across nine European countries and eleven species, covering from Mediterranean to hemiboreal forests, to test if the growth response to site specific drought events that occurred between 1975 and 2015 varied between mixed and monospecific stands. In particular, we quantify how stands resist those specific drought events and recover after them, thus analyzing their resilience. For each drought event and forest stand we calculated resistance, recovery, resilience and relative resilience and we related the variation in these indices between monospecific and mixed stands with type of admixture, tree species identity, site aridity gradient, stand basal area and stand age. We found a large variability among case studies, even for those that share similar species composition and have similar climates. On average, mixed stands showed higher resistance, resilience and relative resilience to drought events than monospecific stands. However, the beneficial effect of mixtures could not be generalized, being greatly modulated by the type of admixture and tree species identity, and depending on site water supply and stand characteristics, such as basal area and age. The increase in resilience in mixtures compared with monocultures was greater on the conifer-broadleaved admixtures, and to a lesser extent in the broadleaved-broadleaved combinations. The observed response patterns to drought largely varied among the eleven studied species, thus revealing the importance of functional traits for understanding a species’ response to drought across its distribution range. Along the site aridity gradient, resilience and relative resilience to drought increased in drier sites for both monospecific and mixed stands, with an observed trend towards higher resilience in mixed stands in the drier and hotter sites. Our results confirm the complexity of the relationships found of resistance, recovery, resilience and relative resilience with drought when comparing pure vs mixed stands., The study was supported by the ERA-Net SUMFOREST project REFORM “Mixed species forest management. Lowering risk, increasing resilience” (PCIN2017-026) and Marie Sklodowska Curie Actions-RISE , CARE4C . Special thankW117/H2020/2018s to Nuria Simón and Guillermo Madrigal for their assistance building up the common database. All coauthors thank their national funding institutions for supporting the establishment, measure and analysis of core data in the studied plots. Research at the Polish case studies was additionally supported by the Ministry of Science and Higher Education of the Republic of Poland (No W117/H2020/2018 )., 15 Pág.

Radial tree stem growth of Norway spruce (Picea abies (L.) H. Karst.), European beech (Fagus sylvatica L.), Scots pine (Pinus sylvestris L.), and stone pine (Pinus cembra L.) was monitored from 2012 to 2015 across sites in Austria with high-resolution dendrometers. Seasonal cumulative diameter increment was modeled using a hierarchical nonlinear mixed-effects model framework based on a logistic growth curve. In the dry and warm year 2015, the average annual diameter increment of 0.30 cm decreased by 50% on lower elevation sites and by 10% on higher elevation sites. In the cool and moist year 2014, Norway spruce achieved a higher annual diameter increment than European beech, whereas the opposite occurred in the dry and warm years 2013 and 2015. In the mixed beech–spruce stand, beech’s consumptive water-use strategy has obviously caused intensified stress for spruce in these drought periods. On higher elevation sites, Norway spruce reacted more sensitively to climate fluctuation compared with stone pine, but overall reactions were only weak. Productivity varied strongly depending on the social status of the tree, with dominant and intermediate trees suffering more from drought. As warming and drought lowers increment rates on lower elevation sites and as trees on higher elevation sites react less flexibly, productivity losses are expected for Austrian forests due to climate warming.

Tree species mixing has been widely promoted as a promising silvicultural tool for reducing drought stress. However, so far only a limited number of species combinations have been studied in detail, revealing inconsistent results. In this study, we analysed the effect of mixing Scots pine and oak (pedunculate oak and sessile oak) trees on their drought response along a comprehensive ecological gradient across Europe. The objective was to improve our knowledge of general drought response patterns of two fundamental European tree species in mixed versus monospecific stands. We focused on three null hypotheses: () tree drought response does not differ between Scots pine and oak, () tree drought response of Scots pine and oak is not affected by stand composition (mixture versus monoculture) and () tree drought response of Scots pine and oak in mixtures and monocultures is not modified by tree size or site conditions. To test the hypotheses, we analysed increment cores of Scots pine and oak, sampled in mixed and monospecific stands, covering a wide range of site conditions. We investigated resistance (the ability to maintain growth levels during drought), recovery (the ability to restore a level of growth after drought) and resilience (the capacity to recover to pre-drought growth levels), involving site-specific drought events that occurred between 1976 and 2015. In monocultures, oak showed a higher resistance and resilience than Scots pine, while recovery was lower. Scots pine in mixed stands exhibited a higher resistance, but also a lower recovery compared with Scots pine in monocultures. Mixing increased the resistance and resilience of oak. Ecological factors such as tree size, site water supply and site fertility were found to have significant effects on the drought response. In the case of Scots pine, resistance was increased by tree size, while recovery was lowered. Resistance of oak increased with site water supply. The observed mixing effect on the tree drought response of Scots pine and oak was in some cases modified by the site conditions studied. Positive mixing effects in terms of resistance and resilience of oak increased with site water supply, while the opposite was found regarding recovery. In contrast, site fertility lessened the positive mixing effect on the resistance of Scots pine. We hypothesise that the observed positive mixing effects under drought mainly result from water- and/or light-related species interactions that improve resource availability and uptake according to temporal and spatial variations in environmental conditions. This work was supported by the European Union as part of the ERA-Net SUMFOREST project REFORM – Mixed species forest management. Lowering risk, increasing resilience (2816ERA02S, PCIN2017-026) and the Horizon 2020 research and innovation programme under the Marie Skłodowska-Curie grant agreement No 778322. All contributors thank their national funding institutions for supporting the establishment, mensuration and analysis of the studied triplets. The first author wants to thank the German Federal Ministry of Food and Agriculture (BMEL) for financial support through the Federal Office for Agriculture and Food (BLE) (grant number 2816ERA02S), as well as the Bayerische Staatsforsten (BaySF) and Landesbetrieb Forst Brandenburg for providing suitable research sites. Research on the Lithuanian triplets (LT 1, LT 2) was made possible by the national funding institution Research Council of Lithuania (LMTLT) (agreement number S-SUMFOREST-17-1). The French site FR 1 belongs to the OPTMix experimental site (https://optmix.irstea.fr), which is supported annually by Ecofor, Allenvi, and the French national research infrastructure ANAEE-F. A special thank is due to Peter Biber for supporting the statistical analysis.

13 Pág. Instituto de Ciencias Forestales (ICIFOR), Mixed forests are usually associated with higher aboveground carbon storage compared to the corresponding monocultures but information on the impact of tree species mixing on soil organic carbon (SOC) is still limited. Yet, maximizing SOC storage is crucial for ecosystem C sequestration and many other ecosystem services. This study used a triplet approach (ie. two-species mixed stand and respective pure stands at the same site) to assess the impact of tree species identity and mixing on SOC storage in eight pine-oak, eight pine-beech and five beech-oak triplets in Europe. We sampled the forest floor (FF) and 0–40 cm in the mineral soil per 10 cm interval. For each triplet type, we fitted basal area (BA) proportion of one component species (for species identity) and a BA-based plot-level True Shannon Diversity index (for species mixing) as explanatory variables for SOC stocks in linear mixed effects models, which included stone content and plot BA as covariates, and site as a random intercept. Considering the total soil depth (FF + 0–40 cm), species identity effect on SOC stocks was only significant for pine-beech and pine-oak triplets but explained more variability in SOC stocks than species mixing across triplet types. Species mixing effect was not significant for any triplet type in the total soil depth. While species identity consistently drove SOC storage in the topsoil layers across triplet types, species mixing explained more variability in SOC stocks in the deeper soil layers except for pine-oak triplets. The results showed that species identity is a stronger driver of SOC storage than species mixing. While tree species identity effect was strongly related to a conifers vs broadleaves signature, the drivers behind mixing effects remained elusive. The results suggest that targeted selection of tree species could better enhance SOC storage in European forests than a mere increase in species richness., The European Union and the National Fund for Scientific Research (Fonds de la recherche scientifique) of Belgium supported this research as part the ERA-Net SUMFOREST project REFORM – Mixed species forest management. Lowering risk, increasing resilience (www.reform-mixing.eu). Some triplets were selected, established and measured with funding from the EuMIXFOR FP1206 Cost Action project. The Polish State Forests Enterprise also supported one of the Polish co-authors (Grant No: OR.271.3.15.2017). We are grateful to Karine Henin for her key role in the laboratory analyses of the samples. We appreciate the technical assistance by Benjamin Willems for soil sampling across Europe. We would like to thank Thomas Pérot for the dendrometric measurements in the Orléans sites. Finally, we are debtful to the two anonymous reviewers who contributed to improving a previous version of this manuscript.

A novel methodological framework is presented for climate-sensitive modeling of annual radial stem increments using tree-ring width time series. The approach is based on a hierarchical Bayes model together with a distributed time lag model that take into account the effects of a series of monthly temperature and precipitation values, as well as their interactions. By using a set of random walk priors, the hierarchical Bayes model allows both the detrending of the individual time series and the regression modeling to be performed simultaneously in a single model step. The approach was applied to comprehensive tree-ring width data from Austria collected on sample plots arranged in triplets representing different mixture types. Bayesian predictions revealed that European larch (Larix decidua Mill.), Norway spruce (Picea abies (L.) H. Karst.), and Scots pine (Pinus sylvestris L.) show positive climate-related growth trends throughout higher elevation sites in Tyrol, and these trends remain unchanged under a mixed-stand scenario. At the lower Austrian sites, Norway spruce was found to show a severely negative growth trend under both the pure- and mixed-stand scenario. The increment rates of European beech (Fagus sylvatica L.) were found to have a negative climate-related trend in pure stands, and the trend diminished through an admixture of spruce or larch. The trends of European larch and sessile oak (Quercus petraea (Matt.) Liebl.) showed stationary behavior, irrespective of the mixture scenario. Scots pine data showed a positive trend at the lower elevation sites under both the pure- and mixed-stand scenario. These findings indicate that species mixing does not lower the climate-related increment fluctuations of beech, oak, pine, and spruce at lower elevation sites.

Past failures of monocultures, caused by wind-throw or insect damages, and ongoing climate change currently strongly stimulate research into mixed-species stands. So far, the focus has mainly been on combinations of species with obvious complementary functional traits. However, for any generalization, a broad overview of the mixing reactions of functionally different tree species in different mixing proportions, patterns and under different site conditions is needed, including assemblages of species with rather similar demands on resources such as light. Here, we studied the growth of Scots pine and oak in mixed versus monospecific stands on 36 triplets located along a productivity gradient across Europe, reaching from Sweden to Spain and from France to Georgia. The set-up represents a wide variation in precipitation (456–1250 mm year−1), mean annual temperature (6.7–11.5 °C) and drought index by de Martonne (21–63 mm °C−1). Stand inventories and increment cores of trees stemming from 40- to 132-year-old, fully stocked stands on 0.04–0.94-ha-sized plots provided insight into how species mixing modifies stand growth and structure compared with neighbouring monospecific stands. On average, the standing stem volume was 436 and 360 m3 ha−1 in the monocultures of Scots pine and oak, respectively, and 418 m3 ha−1 in the mixed stands. The corresponding periodical annual volume increment amounted to 10.5 and 9.1 m3 ha−1 year−1 in the monocultures and 10.5 m3 ha−1 year−1 in the mixed stands. Scots pine showed a 10% larger quadratic mean diameter (p p p p p p p

The rapid re-establishment of forests following large disturbances is being seen as one option to increase the contribution of forests to climate change mitigation. The temporary inclusion of pioneer trees as nurse crops on disturbed sites can facilitate the establishment of target tree species and may additionally benefit productivity and soil fertility. In this study we compared productivity and nutrient cycling between stands of oak target species ( Quercus robur and Quercus petraea ) that were established with and without widely spaced Betula ssp. or Populus ssp. nurse crops. Simulation results for a full rotation of oaks (180 years) indicated that both types of forests, with and without nurse crops, have a comparable total productivity. However, stands with nurse crops supplied 59–96 Mg ha −1 harvestable biomass after 20 years, whereas the first harvest of biomass from stands without nurse crops would occur at least 30 years later. Nutrient element costs associated with the removal of Betula ssp. wood were low compared to Populus ssp. Also, nurse crop stands had up to 2.5 times larger pools of exchangeable base cations in top mineral soils (0–30 cm) compared to mono-specific oak stands. The high soil cation pools may have resulted from reduced leaching under nurse crops or the increased recycling of cations, also from deeper soil depth, via litter fall and fine-root turnover. Our results show that forest reestablishment with pioneer tree species may be a suitable tool for the rapid recovery of forest productivity and mitigation potential following disturbances while simultaneously helping to maintain or increase soil fertility.

An approach is presented to predict the effects climate change may have on mortality of forest trees. Mortality is modeled using long-term observations from the Pan-European Programme for Intensive and Continuous Monitoring of Forest Ecosystems plots, retrospective climate data and frailty models having a parametric baseline hazard function. The linear predictor is modeled by B-spline regression techniques to allow for nonlinear cause-and-effect curves. Spatio-temporal predictions of mortality of four major tree species in the German state of Baden-Wurttemberg were derived in terms of unconditional hazard ratios and based on climate projection data. According to the model, marginal risk of tree death for 100 year old Norway spruce trees will be doubled until 2100. Hazard rates of common beech will be halved in low elevation areas and will be reduced by 25% in higher elevations until 2100. Hazard rates of silver fir will be less affected by a changing climate and will increase by at least 25% and by a maximum of 100% in mountainous regions. Scots pines hazard rates will be halved on higher elevation sites and will increase on lower elevation sites.

A methodological framework is provided for the quantification of climate change effects on site index. Spatio-temporal predictions of site index are derived for six major tree species in the German state of Baden-Wurttemberg using simplified universal kriging (UK) based on large data sets from forest inventories and a climate sensitive site-index model. It is shown by a simulation study that, with the underlying large sample size, residual kriging using ordinary least squares (OLS) estimates of the mean function leads to an approximately unbiased spatial predictor. Moreover, the simulated coverage probabilities of resulting prediction intervals are quite close to the required level. B-spline regression techniques are applied to model nonlinear cause-and-effect curves for estimating site indexes at existing inventory plots dependent on retrospective climate covariates. The spatially structured error is modeled by exponential covariance functions. The mean model is then applied to downscaled climate projection data to spatially predict the relative changes of site index under perturbed climate conditions. Applying climate projections of an existing regional climate model based on IPCC emission scenarios A1B and A2, it is found that site index of all tree species would be decreased in lowland areas, and may increase in mountainous regions. Silver fir and common oak stands would also show increased site indexes in mountainous regions, but further extended to lower elevation levels. Site conditions in the Alpine foothills may remain highly productive for growth of Norway spruce, Baden-Wurttemberg’s most dominant tree species. Whereas site index of common beech and Douglas-fir may decrease to almost the same relative amount and on nearly the same sites as Norway spruce, site index of Scots pine may be less affected by future climate change.