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Functional diversity informs about biodiversity-ecosystem functioning relationships. The intraspecific component of functional diversity (i.e. the phenotypic space of each species) depicts individual differences in the resource use and fitness among conspecifics, and gives valuable information about the functional similarity (competition) or dissimilarity (complementarity) of coexisting species. Here, we quantified trait differences within tree species along local diversity gradients to shed light on the role that this intraspecific variability exerts on functional complementarity of tree species. We measured architectural traits in 5,036 individuals and leaf traits in 1,403 individuals from nine dominant tree species, surveyed in 92 plots located in three major European forest types (Mediterranean, temperate and boreal forests). In each forest type, plots were positioned along a canopy richness gradient, with every study species present in different species richness levels, including monocultures. Our results showed that the relative magnitude of intraspecific trait variability to community-level variability is high in these forests. At the species level, we found adjustments of species leaf traits (mean shifts) in response to neighbouring trees, suggesting the existence of processes that limit niche overlap. We also found higher variability in architectural traits of conspecific individuals in more diverse canopies, suggesting greater niche packing and a more efficient use of available space as the number of species in the canopy increases. Altogether, our results support the hypothesis that differential responses of individuals within a species promote species complementarity, suggesting that biodiversity-ecosystem functioning relationships cannot be properly estimated without accounting for the intraspecific level of functional variation. ; All authors acknowledge support from the European Union FunDivEUROPE project (FP7-ENV-2010. Grant agreement No. 265171). RB was funded by a Marie Curie IEF fellowship (DIVEFOR. FP7-PEOPLE-2011-IEF. Grant Agreement No. 302445), together with the European Union's Horizon 2020 Research and Innovation Programme Project GenTree (Grant Agreement No. 676876), and REMEDINAL3-CM (Autonomous Community of Madrid, S2013/MAE-2719), LINCGlobal (4540-143AP), COMEDIAS (CGL2017-83170-R, Spanish Ministry of Science, Innovation and Universities) projects. ; Peer reviewed

Functional diversity informs about biodiversity-ecosystem functioning relationships. The intraspecific component of functional diversity (i.e. the phenotypic space of each species) depicts individual differences in the resource use and fitness among conspecifics, and gives valuable information about the functional similarity (competition) or dissimilarity (complementarity) of coexisting species. Here, we quantified trait differences within tree species along local diversity gradients to shed light on the role that this intraspecific variability exerts on functional complementarity of tree species. We measured architectural traits in 5,036 individuals and leaf traits in 1,403 individuals from nine dominant tree species, surveyed in 92 plots located in three major European forest types (Mediterranean, temperate and boreal forests). In each forest type, plots were positioned along a canopy richness gradient, with every study species present in different species richness levels, including monocultures. Our results showed that the relative magnitude of intraspecific trait variability to community-level variability is high in these forests. At the species level, we found adjustments of species leaf traits (mean shifts) in response to neighbouring trees, suggesting the existence of processes that limit niche overlap. We also found higher variability in architectural traits of conspecific individuals in more diverse canopies, suggesting greater niche packing and a more efficient use of available space as the number of species in the canopy increases. Altogether, our results support the hypothesis that differential responses of individuals within a species promote species complementarity, suggesting that biodiversity-ecosystem functioning relationships cannot be properly estimated without accounting for the intraspecific level of functional variation. ; All authors acknowledge support from the European Union FunDivEUROPE project (FP7-ENV-2010. Grant agreement No. 265171). RB was funded by a Marie Curie IEF fellowship (DIVEFOR. FP7-PEOPLE-2011-IEF. Grant Agreement No. 302445), together with the European Union's Horizon 2020 Research and Innovation Programme Project GenTree (Grant Agreement No. 676876), and REMEDINAL3-CM (Autonomous Community of Madrid, S2013/MAE-2719), LINCGlobal (4540-143AP), COMEDIAS (CGL2017-83170-R, Spanish Ministry of Science, Innovation and Universities) projects. ; No

[Motivation] Trait variation within species can reveal plastic and/or genetic responses to environmental gradients, and may indicate where local adaptation has occurred. Here, we present a dataset of rangewide variation in leaf traits from seven of the most ecologically and economically important tree species in Europe. Sample collection and trait assessment are embedded in the GenTree project (EU-Horizon 2020), which aims at characterizing the genetic and phenotypic variability of forest tree species to optimize the management and sustainable use of forest genetic resources. Our dataset captures substantial intra- and interspecific leaf phenotypic variability, and provides valuable information for studying the relationship between ecosystem functioning and trait variability of individuals, and the response and resilience of species to environmental changes. ; [Main types of variable contained] We chose morphological and chemical characters linked to trade-offs between acquisition and conservation of resources and water use, namely specific leaf area, leaf size, carbon and nitrogen content and their ratio, and the isotopic signature of stable isotope 13C and 15N in leaves. ; [Spatial location and grain] We surveyed between 18 and 22 populations per species, 141 in total, across Europe. ; [Time period] Leaf sampling took place between 2016 and 2017. ; [Major taxa and level of measurement] We sampled at least 25 individuals in each population, 3,569 trees in total, and measured traits in 35,755 leaves from seven European tree species, i.e. the conifers Picea abies, Pinus pinaster and Pinus sylvestris, and the broadleaves Betula pendula, Fagus sylvatica, Populus nigra and Quercus petraea. ; [Software format] The data files are in ASCII text, tab delimited, not compressed. ; All authors acknowledge support from the European Union's Horizon 2020 Research and Innovation Programme under grant agreement no. 676876 (Project GenTree). Laboratory work was also supported by other projects: COMEDIAS (CGL2017- 83170-R, Spanish Ministry of Science, Innovation and Universities), REMEDINAL IV (TE-CM S2018/EMT-4338, Project of the Autonomous Community of Madrid, Spain), SPONFOREST (BIODIVERSA-3, PCIN-2016-055), and International Laboratory of Global Change (LINCGlobal 4540-143AP). The Swiss contribution was supported by the Swiss Secretariat for Education, Research and Innovation (SERI) under contract no. 6.0032. BC was funded by a Scholarship from Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - CAPES (Brazil). ; Peer reviewed

Received 30 April 2015, Revised 26 October 2015, Accepted 26 October 2015, Available online 2 November 2015 ; Evidence of tree regeneration failure of some species in the Iberian Peninsula forests warns us about the impact that the global change may exert on the preservation of Mediterranean forests, such as we know them. Predictions agree about an exacerbation of the summer drought there, acknowledged as the main limiting factor for the recruits' survival. On the other hand, many studies have also proved the relevant role that local heterogeneity has over the spatial distribution of forest species recruitment by providing safe sites. Therefore, to unravel how climate interacts with local factors over juveniles' performance seems crucial for the design of successful management strategies that allow facing the global warming. Here, we surveyed the natural recruitment of four dominant tree species in seven mountainous regions in the Iberian Peninsula, along entire elevational ranges as surrogates of their climatic ranges. Two of them have alpine and temperate distributions with populations at their rear edge in the Spanish mountains: Fagus sylvatica and Pinus uncinata; and the other two have a genuine Mediterranean distribution: Quercus ilex and Pinus nigra. Our main goal was to analyze for each species the effect of climate, local factors (i.e. light availability, stand structure and ground cover) and the interactions among them to identify the main drivers leading the regeneration process, assessed in terms of presence, abundance and mean annual growth of juveniles. The results showed different environmental factors determining the recruitment patterns of each species. Nevertheless, they highlighted the pervasive role exerted by both climate and fine scale factors, particularly the co-occurring vegetation on recruits' abundance, and the light availability on their growth. Moreover, we found some interactions among annual mean temperature and local factors, suggesting that climate and local heterogeneity act hierarchically, i.e. the local conditions may mitigate or exacerbate the impact of climate on juveniles. These results advocate for further research to increase our knowledge on the complex net of interactions among factors involved in recruitment at different scales, which in turn should be taken into account and incorporated in forthcoming management strategies. ; Funding was also provided by the Spanish Ministry for Innovation and Science with the Grant Consolider-Montes (CSD2008_00040), and the European Union with the projects BACCARA (CE: FP7-226299, 7FP), FunDivEUROPE (CE: FP7-ENV-2010. 265171), and IMBALANCE-P (ERC Synergy project SyG-2013-610028). ; RB was funded by a Marie Curie IEF fellowship (FP7-PEOPLE-2011-IEF). ; Peer reviewed

There is growing concern that the implementation of political agreements on climate change and biodiversity will not be enough to protect forests in the short run and up to the end of the 21st century. As mitigation efforts are lagging behind self-imposed, reasonable targets, genetic diversity will have a large and significant part to play in the process of adapting forests to climate change. Genetic diversity, the raw material of evolution, can be used for adaptation by natural selection and artificial breeding, in naturally regenerated and plantation forests alike. The two-day scientific conference: "#rescueforests: Genetics to the rescue - Managing forests sustainably in a changing world", addressed the genetic diversity of forests. More specifically, the conference was about natural as well as assisted adaptive processes, their spatial scale, from fine grain to landscape and ecoregions, and how much of the genome it involves. It also dealt with phenotypes and how much of their variation is determined by underlying genetic diversity. And finally, and perhaps most importantly, the conference emphasized the importance of conservation and sustainable use of this genetic diversity as a nature-based solution to adapt under the fast pace of climate change. The conference demonstrated how improved knowledge on genomic diversity and evolutionary mechanisms can help to rescue forests, either naturally or by means of management.

[Background]: In contrast with the negligible contribution of the forest understorey to the total aboveground phytobiomass of a forest, its share in annual litter production and nutrient cycling may be more important. Whether and how this functional role of the understorey differs across forest types and depends upon overstorey characteristics remains to be investigated. ; [Methods]: We sampled 209 plots of the FunDivEUROPE Exploratory Platform, a network of study plots covering local gradients of tree diversity spread over six contrasting forest types in Europe. To estimate the relative contribution of the understorey to carbon and nutrient cycling, we sampled non-lignified aboveground understorey biomass and overstorey leaf litterfall in all plots. Understorey samples were analysed for C, N and P concentrations, overstorey leaf litterfall for C and N concentrations. We additionally quantified a set of overstorey attributes, including species richness, proportion of evergreen species, light availability (representing crown density) and litter quality, and investigated whether they drive the understorey's contribution to carbon and nutrient cycling. [Results and conclusions]: Overstorey litter production and nutrient stocks in litterfall clearly exceeded the contribution of the understorey for all forest types, and the share of the understorey was higher in forests at the extremes of the climatic gradient. In most of the investigated forest types, it was mainly light availability that determined the contribution of the understorey to yearly carbon and nutrient cycling. Overstorey species richness did not affect the contribution of the understorey to carbon and nutrient cycling in any of the investigated forest types. ; [Results and conclusions]: Overstorey litter production and nutrient stocks in litterfall clearly exceeded the contribution of the understorey for all forest types, and the share of the understorey was higher in forests at the extremes of the climatic gradient. In most of the investigated forest types, it was mainly light availability that determined the contribution of the understorey to yearly carbon and nutrient cycling. Overstorey species richness did not affect the contribution of the understorey to carbon and nutrient cycling in any of the investigated forest types. ; This study was performed within the framework of the FunDivEUROPE project and has received funding from the European Union Seventh Framework Programme (FP7/2007–2013) under grant agreement n° 265171. Dries Landuyt was supported by a postdoctoral fellowship of the Research Foundation-Flanders (FWO). Kris Verheyen was supported by the ERC Consolidator Grant 614839 that is linked with the project PASTFORWARD.

Global change triggers shifts in forest composition, with warming and aridification being particularly threatening for the populations located at the rear edge of the species distributions. This is the case of Scots pine (Pinus sylvestris) in the Mediterranean Basin where uncertainties in relation to its dynamics under these changing scenarios are still high. We analysed the relative effect of climate on the recruitment patterns of Scots pine and its interactions with local biotic and abiotic variables at different spatial scales. Number of seedlings and saplings was surveyed, and their annual shoot growth measured in 96 plots located across altitudinal gradients in three different regions in the Iberian Peninsula. We found a significant influence of climate on demography and performance of recruits, with a non-linear effect of temperature on the presence of juveniles, and a positive effect of precipitation on their survival. Abundance of juveniles of P. sylvestris that underwent their first summer drought was skewed towards higher altitudes than the altitudinal mean range of the conspecific adults and the optimum elevation for seedlings¿ emergence. At local level, light availability did not influence juveniles¿ density, but it enhanced their growth. Biotic interactions were found between juveniles and the herb cover (competition) and between the number of newly emerged seedlings and shrubs (facilitation). Results also highlighted the indirect effect that climate exerts over the local factors, modulating the interactions with the pre-existing vegetation that were more evident at more stressful sites. This multiscale approach improves our understanding of the dynamics of these marginal populations and some management criteria can be inferred to boost their conservation under the current global warming. ; Funding was provided by the Spanish Ministry for Innovation and Science with the grant Consolider-Montes (CSD2008_00040), and the European Union with the projects BACCARA (CE: FP7-226299, 7FP) and FunDivEUROPE (CE: FP7-ENV-2010. 265171). ; Peer Reviewed

Global change triggers shifts in forest composition, with warming and aridification being particularly threatening for the populations located at the rear edge of the species distributions. This is the case of Scots pine (Pinus sylvestris) in the Mediterranean Basin where uncertainties in relation to its dynamics under these changing scenarios are still high. We analysed the relative effect of climate on the recruitment patterns of Scots pine and its interactions with local biotic and abiotic variables at different spatial scales. Number of seedlings and saplings was surveyed, and their annual shoot growth measured in 96 plots located across altitudinal gradients in three different regions in the Iberian Peninsula. We found a significant influence of climate on demography and performance of recruits, with a non-linear effect of temperature on the presence of juveniles, and a positive effect of precipitation on their survival. Abundance of juveniles of P. sylvestris that underwent their first summer drought was skewed towards higher altitudes than the altitudinal mean range of the conspecific adults and the optimum elevation for seedlings' emergence. At local level, light availability did not influence juveniles' density, but it enhanced their growth. Biotic interactions were found between juveniles and the herb cover (competition) and between the number of newly emerged seedlings and shrubs (facilitation). Results also highlighted the indirect effect that climate exerts over the local factors, modulating the interactions with the pre-existing vegetation that were more evident at more stressful sites. This multiscale approach improves our understanding of the dynamics of these marginal populations and some management criteria can be inferred to boost their conservation under the current global warming. ; Funding was provided by the Spanish Ministry for Innovation and Science with the grant Consolider-Montes (CSD2008_00040), and the European Union with the projects BACCARA (CE: FP7-226299, 7FP) and FunDivEUROPE (CE: FP7-ENV-2010. 265171).

Disentangling the mechanisms that shape community assembly across diversity gradients is a central matter in ecology. While many studies have explored community assembly through species average trait values, there is a growing understanding that intraspecific trait variation (ITV) can also play a critical role in species coexistence. Classic biodiversity theory hypothesizes that higher diversity at species-rich sites can arise from narrower niches relative to species-poor sites, which would be reflected in reduced ITV as species richness increases. To explore how ITV in woody plant communities changes with species richness, we compiled leaf trait data (leaf size and specific leaf area) in a total of 521 woody plant species from 21 forest communities that differed dramatically in species richness, ranging from boreal to tropical rainforests. At each forest, we assessed ITV as an estimate of species niche breadth and we quantified the degree of trait overlap among co-occurring species as a measure of species functional similarity. We found ITV was relatively invariant across the species richness gradient. In addition, we found that species functional similarity increased with diversity. Contrary to the expectation from classic biodiversity theory, our results rather suggest that neutral processes or equalizing mechanisms can be acting as potential drivers shaping community assembly in hyperdiverse forests. ; Leaf data used to come to this manuscript resulted from many different studies involving different funding sources: European Union Seventh Framework Programme (FP7/2007-2013) under grant agreement no. 265171; Spanish-funded project REMEDINAL3eCM (S2013/MAE-2719); Ecometas excellence network (CGL2014-53840-REDT); a FPU grant by the Spanish Government (AP2010-5600); a Marie Curie IEF fellowship (FP7-PEOPLE-2011-IEF. no. 302445); ; Peer Reviewed

[Background]: Progress in the field of evolutionary forest ecology has been hampered by the huge challenge of phenotyping trees across their ranges in their natural environments, and the limitation in high-resolution environmental information. ; [Findings]: The GenTree Platform contains phenotypic and environmental data from 4,959 trees from 12 ecologically and economically important European forest tree species: Abies alba Mill. (silver fir), Betula pendula Roth. (silver birch), Fagus sylvatica L. (European beech), Picea abies (L.) H. Karst (Norway spruce), Pinus cembra L. (Swiss stone pine), Pinus halepensis Mill. (Aleppo pine), Pinus nigra Arnold (European black pine), Pinus pinaster Aiton (maritime pine), Pinus sylvestris L. (Scots pine), Populus nigra L. (European black poplar), Taxus baccata L. (English yew), and Quercus petraea (Matt.) Liebl. (sessile oak). Phenotypic (height, diameter at breast height, crown size, bark thickness, biomass, straightness, forking, branch angle, fructification), regeneration, environmental in situ measurements (soil depth, vegetation cover, competition indices), and environmental modeling data extracted by using bilinear interpolation accounting for surrounding conditions of each tree (precipitation, temperature, insolation, drought indices) were obtained from trees in 194 sites covering the species' geographic ranges and reflecting local environmental gradients. ; [Conclusion]: The GenTree Platform is a new resource for investigating ecological and evolutionary processes in forest trees. The coherent phenotyping and environmental characterization across 12 species in their European ranges allow for a wide range of analyses from forest ecologists, conservationists, and macro-ecologists. Also, the data here presented can be linked to the GenTree Dendroecological collection, the GenTree Leaf Trait collection, and the GenTree Genomic collection presented elsewhere, which together build the largest evolutionary forest ecology data collection available. ; This publication is part of the GenTree project, which was funded by the European Union's Horizon 2020 research and innovation program under grant agreement No. 676876 (GenTree). This work was also supported by the Swiss Secretariat for Education, Research and Innovation (SERI) under contract No. 6.0032. ; Peer reviewed

© 2015 Geobotanisches Institut ETH, Stiftung Ruebel. Climate change is expected to alter the geographic distribution of many plant species worldwide. However, there is still no clear evidence showing a generalised direction and magnitude of these distribution shifts. Here, we have surveyed, in nine mountainous regions in Spain, an array of tree species along entire elevational ranges, as surrogates of their global climatic ranges, to test for elevational shifts towards cooler locations. We analysed the distribution recruitment patterns of five dominant tree species, recording the abundance and measuring the primary growth of juveniles in 306 plots. Three of the species have a temperate-boreal distribution with populations at their southern edge in the Mediterranean mountain ranges: Pinus sylvestris, Pinus uncinata and Fagus sylvatica; and the other two species have a Mediterranean distribution: Quercus ilex and Pinus nigra. Despite the contrasting phylogenies and biogeographies, we identified a similar pattern in recruitment abundance across species, with an asymmetric distribution of juveniles (more recruits in the middle-upper elevation of their range), but higher annual growths at lower elevations. This survival-growth trade-off at the early recruitment stage may potentially counterbalance at population level the negative effect of global warming on recruit survival at the lower edge of species ranges. These findings suggest a demographic stabilisation process at the early recruitment stage of these tree species, and highlight the importance of considering the different demographic stages across the whole climatic range to understand the effects that climate change may exert on species distributions and population dynamics. ; Funding also was provided by the Spanish Ministry for Innovation and Science with the grant Consolider-Montes (CSD2008 00040), VULGLO (CGL201022180 C03 03), MOUNTAINS (CGL-2012-38427), the Community of Madrid grant REMEDINAL 2 (CM S2009 AMB 1783) and the European Union with the projects BACCARA (CE: FP7-226299, 7FP) and FunDivEUROPE (CE: FP7-ENV-2010. 265171). ; RB was funded by a MarieCurie IEF fellowship (FP7-PEOPLE-2011-IEF). ; Peer Reviewed

Many experiments have shown that local biodiversity loss impairs the ability of ecosystems to maintain multiple ecosystem functions at high levels (multifunctionality). In contrast, the role of biodiversity in driving ecosystem multifunctionality at landscape scales remains unresolved. We used a comprehensive pan-European dataset, including 16 ecosystem functions measured in 209 forest plots across six European countries, and performed simulations to investigate how local plot-scale richness of tree species (α-diversity) and their turnover between plots (β-diversity) are related to landscape-scale multifunctionality. After accounting for variation in environmental conditions, we found that relationships between α-diversity and landscape-scale multifunctionality varied from positive to negative depending on the multifunctionality metric used. In contrast, when significant, relationships between β-diversity and landscape-scale multifunctionality were always positive, because a high spatial turnover in species composition was closely related to a high spatial turnover in functions that were supported at high levels. Our findings have major implications for forest management and indicate that biotic homogenization can have previously unrecognized and negative consequences for large-scale ecosystem multifunctionality. ; The research leading to these results received funding from the European Union Seventh Framework Programme (FP7/2007-2013) under Grant Agreement 265171. ; Peer Reviewed

Many experiments have shown that local biodiversity loss impairs the ability of ecosystems to maintain multiple ecosystem functions at high levels (multifunctionality). In contrast, the role of biodiversity in driving ecosystem multifunctionality at landscape scales remains unresolved. We used a comprehensive pan-European dataset, including 16 ecosystem functions measured in 209 forest plots across six European countries, and performed simulations to investigate how local plot-scale richness of tree species (α-diversity) and their turnover between plots (β-diversity) are related to landscape-scale multifunctionality. After accounting for variation in environmental conditions, we found that relationships between α-diversity and landscape-scale multifunctionality varied from positive to negative depending on the multifunctionality metric used. In contrast, when significant, relationships between β-diversity and landscape-scale multifunctionality were always positive, because a high spatial turnover in species composition was closely related to a high spatial turnover in functions that were supported at high levels. Our findings have major implications for forest management and indicate that biotic homogenization can have previously unrecognized and negative consequences for large-scale ecosystem multifunctionality. ; We thank the Hainich National Park administration as well as Felix Berthold and Carsten Beinhoff for support of this study and Gerald Kaendler and the Johann Heinrich von Thünen-Institut for providing access to the German National Forest Inventory data. The research leading to these results received funding from the European Union Seventh Framework Programme (FP7/2007-2013) under Grant Agreement 265171. ; This is the final version of the article. It first appeared from the National Academy of Sciences via https://doi.org//10.1073/pnas.1517903113

There is considerable evidence that biodiversity promotes multiple ecosystem functions (multifunctionality), thus ensuring the delivery of ecosystem services important for human well-being. However, the mechanisms underlying this relationship are poorly understood, especially in natural ecosystems. We develop a novel approach to partition biodiversity effects on multifunctionality into three mechanisms and apply this to European forest data. We show that throughout Europe, tree diversity is positively related with multifunctionality when moderate levels of functioning are required, but negatively when very high function levels are desired. For two well-known mechanisms, 'complementarity' and 'selection', we detect only minor effects on multifunctionality. Instead a third, so far overlooked mechanism, the 'jack-of-all-trades' effect, caused by the averaging of individual species effects on function, drives observed patterns. Simulations demonstrate that jack-of-all-trades effects occur whenever species effects on different functions are not perfectly correlated, meaning they may contribute to diversity-multifunctionality relationships in many of the world's ecosystems. ; The research leading to these results received funding from the European Union Seventh Framework Programme (FP7/2007-2013) under grant agreement no. 265171. ; This is the final version of the article. It first appeared from Nature Publishing Group via http://dx.doi.org/10.1038/ncomms11109