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Plant species that grow across environmental gradients show a range of trait expression, but traits do not vary independently. In fact, phenotypes are integrated expressions of multiple traits that covary due to trade‐offs among functions and processes. Understanding trait covariation structures will ultimately help with predicting species' responses to change and developing management actions.We measured variation and covariation (a proxy of phenotypic integration) among functional traits of Pinus sylvestris from paired populations across its European distribution. Populations within a pair were close enough to be in gene flow contact but located in contrasting environmental conditions across a local gradient. Measured traits represented three axes of variation (groups of traits) related to a tree's competitive ability and the trade‐off between resource acquisition and conservation, namely plant size measures and stem and foliar traits.Results revealed important intra‐ and inter‐population trait variability. In particular, at the population level, trait means shifted across the climatic gradient mainly described by mean annual temperature. Moreover, we found a higher degree of trait covariation in populations under harsher environments (i.e. lower environmental suitability for the species). This pattern was consistent within population pairs, suggesting that higher trait covariation may be adaptive, being more coordinated in sites with harsher conditions. At larger spatial scales, we found a less conclusive pattern with a trend of increasing covariation at the northern edge of the species distribution. This result suggests that at larger scales different processes may be involved in the trade‐off between the adaptive value of phenotypic covariation versus its constraints on trait combinations that may limit plant's response capability.Synthesis. Trait covariation varies at different spatial scales, increasing under harsher conditions, and the robustness and repeatability of this pattern suggests its adaptive role for the species' responses to different environments. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Summary: Understanding the dynamics of selection is key to predicting the response of tree species to new environmental conditions in the current context of climate change. However, selection patterns acting on early recruitment stages and their climatic drivers remain largely unknown in most tree species, despite being a critical period of their life cycle.We measured phenotypic selection on Pinus sylvestris seed mass, emergence time and early growth rate over 2 yr in four common garden experiments established along the latitudinal gradient of the species in Europe.Significant phenotypic plasticity and among‐population genetic variation were found for all measured phenotypic traits. Heat and drought negatively affected fitness in the southern sites, but heavy rainfalls also decreased early survival in middle latitudes. Climate‐driven directional selection was found for higher seed mass and earlier emergence time, while the form of selection on seedling growth rates differed among sites and populations. Evidence of adaptive and maladaptive phenotypic plasticity was found for emergence time and early growth rate, respectively.Seed mass, emergence time and early growth rate have an adaptive role in the early stages of P. sylvestris and climate strongly influences the patterns of selection on these fitness‐related traits. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The functioning of plant communities is strongly influenced by the number of species in the community and their spatial arrangement. This is because plants interact with their nearest neighbors and this interaction is expected to be stronger when the interacting individuals are ecologically similar in terms of resource use. Recent evidence shows that species richness alters the balance of intra‐ versus interspecific competition, but the effect of species richness, and phylogenetic and functional diversity on the spatial pattern of the plant communities remain less studied. Even far, how forest stand structure derived from past management practices can influence the relationship between species richness and spatial pattern is still unknown. Here, we evaluate the spatial distribution of woody individuals (DBH >7.5 cm) in 209 forest stands (i.e. plots) with an increasing level of species richness (from 1 up to 10 species) in six forest types along a latitudinal gradient in Europe. We used completely mapped plots to investigate the spatial pattern in each forest stand with point pattern techniques. We fitted linear models to analyze the effect of species richness (positively correlated with phylogenetic diversity) and functional diversity on tree spatial arrangements. We also controled this relationship by forest type and stand structure as a proxy of the management legacy. Our results showed a generalized positive effect of species richness and functional diversity on the degree of spatial clustering of trees, and on the spatial independence of tree sizes regardless of the forest type. Moreover, current tree spatial arrangements were still conditioned by its history of management; however its effect was independent of the number of species in the community. Our study showed that species richness and functional diversity are relevant attributes of forests influencing the spatial pattern of plant communities, and consequently forest functioning. [ABSTRACT FROM AUTHOR]

The importance of biodiversity in supporting ecosystem functioning is generally well accepted. However, most evidence comes from small-scale studies, and scaling-up patterns of biodiversity-ecosystem functioning (B-EF) remains challenging, in part because the importance of environmental factors in shaping B-EF relations is poorly understood. Using a forest research platform in which 26 ecosystem functions were measured along gradients of tree species richness in six regions across Europe, we investigated the extent and the potential drivers of context dependency of B-EF relations. Despite considerable variation in species richness effects across the continent, we found a tendency for stronger B-EF relations in drier climates as well as in areas with longer growing seasons and more functionally diverse tree species. The importance of water availability in driving context dependency suggests that as water limitation increases under climate change, biodiversity may become even more important to support high levels of functioning in European forests. [ABSTRACT FROM AUTHOR]

Predicting climate-driven changes in plant distribution is crucial for biodiversity conservation and management under recent climate change. Climate warming is expected to induce movement of species upslope and towards higher latitudes. However, the mechanisms and physiological processes behind the altitudinal and latitudinal distribution range of a tree species are complex and depend on each tree species features and vary over ontogenetic stages. We investigated the altitudinal distribution differences between juvenile and adult individuals of seven major European tree species along elevational transects covering a wide latitudinal range from southern Spain (37°N) to northern Sweden (67°N). By comparing juvenile and adult distributions (shifts on the optimum position and the range limits) we assessed the response of species to present climate conditions in relation to previous conditions that prevailed when adults were established. Mean temperature increased by 0.86 °C on average at our sites during the last decade compared with previous 30-year period. Only one of the species studied, Abies alba, matched the expected predictions under the observed warming, with a maximum abundance of juveniles at higher altitudes than adults. Three species, Fagus sylvatica, Picea abies and Pinus sylvestris, showed an opposite pattern while for other three species, such as Quercus ilex, Acer pseudoplatanus and Q. petraea, we were no able to detect changes in distribution. These findings are in contrast with theoretical predictions and show that tree responses to climate change are complex and are obscured not only by other environmental factors but also by internal processes related to ontogeny and demography. [ABSTRACT FROM AUTHOR]

• Differences in tree recruitment for forests in extreme and middle latitudes in Europe. • Potential shift in species diversity and composition of European forests at long-term. • Post-dispersal filters blurred the benign effect of canopy diversity on recruitment. • Recruitment patterns were mediated by seed size along a European gradient. Increasing evidence supports that canopy richness favors multifunctionality in European forests. Given that recruitment represents one of the main demographic bottlenecks in plant communities, the provision of similar levels of forest functioning in the future will depend on successful natural regeneration in the present. Here, we evaluated whether canopy diversity enhances recruitment (abundance and richness) by providing a more diverse seed supply and a more heterogeneous environmental context; or, alternatively, if post-dispersal environmental filters outweigh these expected positive effects. For this purpose, we used a large-scale European platform comprising 173 forest plots that follow a canopy species richness gradient (from monoculture to mixed plots up to 5 tree species) in five forest types from Mediterranean to boreal forests. Despite that canopy diversity promoted a more diverse seed supply and a more heterogeneous light environment, this did not translate into a richer or more abundant community of recruits. In fact, we found an important uncoupling in species composition between recruitment and canopy layers. Our results suggest that strong post-dispersal filters, mediated by an interaction between seed size and latitude, drive this mismatch. Large-seeded species were over-represented in the recruitment at both extremes of the European gradient (Mediterranean and boreal forests), while the opposite trend occurred at intermediate latitudes. Altogether, our work shows that preserving a more diverse canopy may not ensure successful recruitment, and advocates future studies analyzing the role of post-dispersal filters on the assembly of European forests. [ABSTRACT FROM AUTHOR]

In a context of accelerated human-induced biodiversity loss, remote sensing (RS) is emerging as a promising tool to map plant biodiversity from space. Proposed approaches often rely on the Spectral Variation Hypothesis (SVH), linking the heterogeneity of terrestrial vegetation to the variability of the spectroradiometric signals. Yet, due to observational limitations, the SVH has been insufficiently tested, remaining unclear which metrics, methods, and sensors could provide the most reliable estimates of plant biodiversity. Here we assessed the potential of RS to infer plant biodiversity using radiative transfer simulations and inversion. We focused specifically on "functional diversity," which represents the spatial variability in plant functional traits. First, we simulated vegetation communities and evaluated the information content of different functional diversity metrics (FDMs) derived from their optical reflectance factors (R) or the corresponding vegetation "optical traits," estimated via radiative transfer model inversion. Second, we assessed the effect of the spatial resolution, the spectral characteristics of the sensor, and signal noise on the relationships between FDMs derived from field and remote sensing datasets. Finally, we evaluated the plausibility of the simulations using Sentinel-2 (multispectral, 10 m pixel) and DESIS (hyperspectral, 30 m pixel) imagery acquired over sites of the Functional Significance of Forest Biodiversity in Europe (FunDivEUROPE) network. We demonstrate that functional diversity can be inferred both by reflectance and optical traits. However, not all the FDMs tested were suited for assessing plant functional diversity from RS. Rao's Q index, functional dispersion, and functional richness were the best-performing metrics. Furthermore, we demonstrated that spatial resolution is the most limiting RS feature. In agreement with simulations, Sentinel-2 imagery provided better estimates of plant diversity than DESIS, despite the coarser spectral resolution. However, Sentinel-2 offered inaccurate results at DESIS spatial resolution. Overall, our results identify the strengths and weaknesses of optical RS to monitor plant functional diversity. Future missions and biodiversity products should consider and benefit from the identified potentials and limitations of the SVH. [Display omitted] • Systematic evaluation of the Spectral Variation Hypothesis delimits when it works. • Rao's Q , functional dispersion & richness get plant functional diversity from space. • Reflectance and vegetation radiative transfer parameter estimates can be used. • Images individually analyzed can be compared in space and time in global products. • Spatial is the most limiting resolution, spectral only for parameter estimation. [ABSTRACT FROM AUTHOR]

Abstract: One of the current advances in functional biodiversity research is the move away from short-lived test systems towards the exploration of diversity-ecosystem functioning relationships in structurally more complex ecosystems. In forests, assumptions about the functional significance of tree species diversity have only recently produced a new generation of research on ecosystem processes and services. Novel experimental designs have now replaced traditional forestry trials, but these comparatively young experimental plots suffer from specific difficulties that are mainly related to the tree size and longevity. Tree species diversity experiments therefore need to be complemented with comparative observational studies in existing forests. Here we present the design and implementation of a new network of forest plots along tree species diversity gradients in six major European forest types: the FunDivEUROPE Exploratory Platform. Based on a review of the deficiencies of existing observational approaches and of unresolved research questions and hypotheses, we discuss the fundamental criteria that shaped the design of our platform. Key features include the extent of the species diversity gradient with mixtures up to five species, strict avoidance of a dilution gradient, special attention to community evenness and minimal covariation with other environmental factors. The new European research platform permits the most comprehensive assessment of tree species diversity effects on forest ecosystem functioning to date since it offers a common set of research plots to groups of researchers from very different disciplines and uses the same methodological approach in contrasting forest types along an extensive environmental gradient. [Copyright &y& Elsevier]

The dataset presented here was collected by the GenTree project (EU-Horizon 2020), which aims to improve the use of forest genetic resources across Europe by better understanding how trees adapt to their local environment. This dataset of individual tree-core characteristics including ring-width series and whole-core wood density was collected for seven ecologically and economically important European tree species: silver birch (Betula pendula), European beech (Fagus sylvatica), Norway spruce (Picea abies), European black poplar (Populus nigra), maritime pine (Pinus pinaster), Scots pine (Pinus sylvestris), and sessile oak (Quercus petraea). Tree-ring width measurements were obtained from 3600 trees in 142 populations and whole-core wood density was measured for 3098 trees in 125 populations. This dataset covers most of the geographical and climatic range occupied by the selected species. The potential use of it will be highly valuable for assessing ecological and evolutionary responses to environmental conditions as well as for model development and parameterization, to predict adaptability under climate change scenarios.

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.

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.