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Enhancing tree diversity may be important to fostering resilience to drought-related climate extremes. So far, little attention has been given to whether tree diversity can increase the survival of trees and reduce its variability in young forest plantations.We conducted an analysis of seedling and sapling survival from 34 globally distributed tree diversity experiments (363,167 trees, 168 species, 3744 plots, 7 biomes) to answer two questions: (1) Do drought and tree diversity alter the mean and variability in plot-level tree survival, with higher and less variable survival as diversity increases? and (2) Do species that survive poorly in monocultures survive better in mixtures and do specific functional traits explain monoculture survival?Tree species richness reduced variability in plot-level survival, while functional diversity (Rao's Q entropy) increased survival and also reduced its variability. Importantly, the reduction in survival variability became stronger as drought severity increased. We found that species with low survival in monocultures survived comparatively better in mixtures when under drought. Species survival in monoculture was positively associated with drought resistance (indicated by hydraulic traits such as turgor loss point), plant height and conservative resource-acquisition traits (e.g. low leaf nitrogen concentration and small leaf size).Synthesis. The findings highlight: (1) The effectiveness of tree diversity for decreasing the variability in seedling and sapling survival under drought; and (2) the importance of drought resistance and associated traits to explain altered tree species survival in response to tree diversity and drought. From an ecological perspective, we recommend mixing be considered to stabilize tree survival, particularly when functionally diverse forests with drought-resistant species also promote high survival of drought-sensitive species.

Water deficit in the atmosphere and soil are two key interactive factors that constrain transpiration and vegetation productivity. It is not clear which of these two factors is more important for the water and carbon flux response to drought stress in ecosystems. In this study, field data and numerical modeling were used to isolate their impact on evapotranspiration (ET) and gross primary productivity (GPP) at a tropical forest site in Barro Colorado Island (BCI), Panama, focusing on their response to the drought induced by the El Niño event of 2015–2016. Numerical simulations were performed using a plant hydrodynamic scheme (HYDRO) and a heuristic approach that ignores stomatal sensitivity to leaf water potential in the Energy Exascale Earth System Model (E3SM) Land Model (ELM). The sensitivity of canopy conductance (Gs) to vapor pressure deficit (VPD) obtained from eddy-covariance fluxes and measured sap flux shows that, at both ecosystem and plant scale, soil water stress is more important in limiting Gs than VPD at BCI during the El Niño event. The model simulations confirmed the importance of water stress limitation on Gs, but overestimated the VPD impact on Gs compared to that estimated from the observations. We also found that the predicted soil moisture is less sensitive to the diversity of plant hydraulic traits than ET and GPP. During the dry season at BCI, seasonal ET, especially soil evaporation at VPD > 0.42 kPa, simulated using HYDRO and ELM, were too strong and will require alternative parameterizations.

Water deficit in the atmosphere and soil are two key interactive factors that constrain transpiration and vegetation productivity. It is not clear which of these two factors is more important for the water and carbon flux response to drought stress in ecosystems. In this study, field data and numerical modeling were used to isolate their impact on evapotranspiration (ET) and gross primary productivity (GPP) at a tropical forest site in Barro Colorado Island (BCI), Panama, focusing on their response to the drought induced by the El Niño event of 2015–2016. Numerical simulations were performed using a plant hydrodynamic scheme (HYDRO) and a heuristic approach that ignores stomatal sensitivity to leaf water potential in the Energy Exascale Earth System Model (E3SM) Land Model (ELM). The sensitivity of canopy conductance (Gs) to vapor pressure deficit (VPD) obtained from eddy-covariance fluxes and measured sap flux shows that, at both ecosystem and plant scale, soil water stress is more important in limiting Gs than VPD at BCI during the El Niño event. The model simulations confirmed the importance of water stress limitation on Gs, but overestimated the VPD impact on Gs compared to that estimated from the observations. We also found that the predicted soil moisture is less sensitive to the diversity of plant hydraulic traits than ET and GPP. During the dry season at BCI, seasonal ET, especially soil evaporation at VPD > 0.42 kPa, simulated using HYDRO and ELM, were too strong and will require alternative parameterizations.

1. Many experiments have shown that biodiversity promotes ecosystem functioning and stability and that this relationship varies with resource availability. However, we still have a poor understanding of the underlying physiological and ecological mechanisms driving diversity effects and how they may interact with soil nutrient availability. 2. We collected data in a grassland experiment factorially manipulating fertilization, species richness (SR), functional composition (slow-growing vs. fast-growing species) and functional diversity in resource economic traits. We measured above-ground productivity (AP), nitrogen (N) uptake, photosynthesis and water use efficiency by combining a ¹⁵N labelling approach with productivity, gas exchange and stable isotope measurements in 3 years differing in rainfall. 3. We found that sown SR increased AP, N uptake and photosynthesis, suggesting that SR is the most important driver of ecosystem productivity and nutrient cycling. Similarly, photosynthesis was affected by functional composition but not by functional diversity. Water use efficiency was reduced by sown SR for communities dominated by slow-growing species but not for communities dominated by fast-growing species. Fertilization increased productivity, N uptake and water use efficiency. The positive effects of high SR on ecosystem functions were independent of fertility levels. 4. Synthesis. Our results provide evidence that high species richness in temperate grasslands could enhance productivity and reduce the negative impacts of drought events. Multiple factors and community characteristics are important in driving enhanced ecosystem functioning in biodiverse grasslands and seem to affect functioning and stability through different mechanisms.

Forest ecosystem functioning generally benefits from higher tree species richness, but variation within richness levels is typically large. This is mostly due to the contrasting performances of communities with different compositions. Evidence-based understanding of composition effects on forest productivity, as well as on multiple other functions will enable forest managers to focus on the selection of species that maximize functioning, rather than on diversity per se. We used a dataset of 30 ecosystem functions measured in stands with different species richness and composition in six European forest types. First, we quantified whether the compositions that maximize annual above-ground wood production (productivity) generally also fulfil the multiple other ecosystem functions (multifunctionality). Then, we quantified the species identity effects and strength of interspecific interactions to identify the “best” and “worst” species composition for multifunctionality. Finally, we evaluated the real-world frequency of occurrence of best and worst mixtures, using harmonized data from multiple national forest inventories. The most productive tree species combinations also tended to express relatively high multifunctionality, although we found a relatively wide range of compositions with high- or low-average multifunctionality for the same level of productivity. Monocultures were distributed among the highest as well as the lowest performing compositions. The variation in functioning between compositions was generally driven by differences in the performance of the component species and, to a lesser extent, by particular interspecific interactions. Finally, we found that the most frequent species compositions in inventory data were monospecific stands and that the most common compositions showed below-average multifunctionality and productivity. Synthesis and applications. Species identity and composition effects are essential to the development of high-performing product

It has been known for a long time that drought intensity is a critical variable in determining water stress of Mediterranean tree species. However, not as much attention has been paid to other drought characteristics, for example the timing of the dry periods. We investigated the impact of the timing and intensity of extreme droughts on growing season length, growth and water-use efficiency of three tree species, Pinus nigra ssp. Salzmannii J.F. Arnold, Quercus ilex ssp. ballota (Desf.) Samp. and Quercus faginea Lam. coexisting in a continental Mediterranean ecosystem. Over the study period (2009–13), intense droughts were observed at annual and seasonal scales, particularly during 2011 and 2012. In 2012, an atypically dry winter and spring was followed by an intense summer drought. Quercus faginea growth was affected more by drought timing than by drought intensity, probably because of its winter-deciduous leaf habit. Pinus nigra showed a lower decrease in secondary growth than observed in the two Quercus species in extremely dry years. Resilience to extreme droughts was different among species, with Q. faginea showing poorer recovery of growth after very dry years. The highest intra- and inter-annual plasticity in water-use efficiency was observed in P. nigra, which maintained a more water-saving strategy. Our results revealed that the timing of extreme drought events can affect tree function to a larger extent than drought intensity, especially in deciduous species. Legacy effects of drought over months and years significantly strengthened the impact of drought timing and intensity on tree function.It has been known for a long time that drought intensity is a critical variable in determining water stress of Mediterranean tree species. However, not as much attention has been paid to other drought characteristics, for example the timing of the dry periods. We investigated the impact of the timing and intensity of extreme droughts on growing season length, growth an

Humans require multiple services from ecosystems, but it is largely unknown whether trade-offs between ecosystem functions prevent the realisation of high ecosystem multifunctionality across spatial scales. Here, we combined a comprehensive dataset (28 ecosystem functions measured on 209 forest plots) with a forest inventory dataset (105,316 plots) to extrapolate and map relationships between various ecosystem multifunctionality measures across Europe. These multifunctionality measures reflected different management objectives, related to timber production, climate regulation and biodiversity conservation/recreation. We found that trade-offs among them were rare across Europe, at both local and continental scales. This suggests a high potential for ‘win-win’ forest management strategies, where overall multifunctionality is maximised. However, across sites, multifunctionality was on average 45.8-49.8% below maximum levels and not necessarily highest in protected areas. Therefore, using one of the most comprehensive assessments so far, our study suggests a high but largely unrealised potential for management to promote multifunctional forests.

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.

Interactions between tree species in forests can be beneficial to ecosystem functions and services related to the carbon and water cycles by improving for example transpiration and productivity. However, little is known on below- and above-ground processes leading to these positive effects. We tested whether stratification in soil water uptake depth occurred between four tree species in a 10-year-old temperate mixed species plantation during a dry summer. We selected dominant and co-dominant trees of European beech, Sessile oak, Douglas fir and Norway spruce in areas with varying species diversity, competition intensity, and where different plant functional types (broadleaf vs. conifer) were present. We applied a deuterium labelling approach that consisted of spraying labelled water to the soil surface to create a strong vertical gradient of the deuterium isotope composition in the soil water. The deuterium isotope composition of both the xylem sap and the soil water was measured before labelling, and then again three days after labelling, to estimate the soil water uptake depth using a simple modelling approach. We also sampled leaves and needles from selected trees to measure their carbon isotope composition (a proxy for water use efficiency) and total nitrogen content. At the end of the summer, we found differences in the soil water uptake depth between plant functional types but not within types: on average, coniferous species extracted water from deeper layers than did broadleaved species. Neither species diversity nor competition intensity had a detectable influence on soil water uptake depth, foliar water use efficiency or foliar nitrogen concentration in the species studied. However, when coexisting with an increasing proportion of conifers, beech extracted water from progressively deeper soil layers. We conclude that complementarity for water uptake could occur in this 10-year-old plantation because of inherent differences among functional groups (conifers

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.