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The role of conspecific density dependence (CDD) in the maintenance of species richness is a central focus of tropical forest ecology. However, tests of CDD often ignore the integrated effects of CDD over multiple life stages and their long-term impacts on population demography. We combined a 10-year time series of seed production, seedling recruitment and sapling and tree demography of three dominant Southeast Asian tree species that adopt a mast-fruiting phenology. We used these data to construct individual-based models that examine the effects of CDD on population growth rates ( λ ) across life-history stages. Recruitment was driven by positive CDD for all species, supporting the predator satiation hypothesis, while negative CDD affected seedling and sapling growth of two species, significantly reducing λ . This negative CDD on juvenile growth overshadowed the positive CDD of recruitment, suggesting the cumulative effects of CDD during seedling and sapling development has greater importance than the positive CDD during infrequent masting events. Overall, CDD varied among positive, neutral and negative effects across life-history stages for all species, suggesting that assessments of CDD on transitions between just two stages (e.g. seeds seedlings or juveniles mature trees) probably misrepresent the importance of CDD on population growth and stability.

Current policy is driving renewed impetus to restore forests to return ecological function, protect species, sequester carbon and secure livelihoods. Here we assess the contribution of tree planting to ecosystem restoration in tropical and sub-tropical Asia; we synthesize evidence on mortality and growth of planted trees at 176 sites and assess structural and biodiversity recovery of co-located actively restored and naturally regenerating forest plots. Mean mortality of planted trees was 18% 1 year after planting, increasing to 44% after 5 years. Mortality varied strongly by site and was typically ca 20% higher in open areas than degraded forest, with height at planting positively affecting survival. Size-standardized growth rates were negatively related to species-level wood density in degraded forest and plantations enrichment settings. Based on community-level data from 11 landscapes, active restoration resulted in faster accumulation of tree basal area and structural properties were closer to old-growth reference sites, relative to natural regeneration, but tree species richness did not differ. High variability in outcomes across sites indicates that planting for restoration is potentially rewarding but risky and context-dependent. Restoration projects must prepare for and manage commonly occurring challenges and align with efforts to protect and reconnect remaining forest areas. The abstract of this article is available in Bahasa Indonesia in the electronic supplementary material. This article is part of the theme issue 'Understanding forest landscape restoration: reinforcing scientific foundations for the UN Decade on Ecosystem Restoration'., Philosophical Transactions of the Royal Society B: Biological Sciences, 378 (1867), ISSN:0962-8436, ISSN:1471-2970, ISSN:0080-4622

Natural forest is declining globally as the area of planted forest increases. Planted forests are often monocultures, despite results suggesting that higher species richness improves ecosystem functioning and stability. To test if this is generally the case, we performed a meta-analysis of available results. We assessed aboveground carbon stocks in mixed-species planted forests vs (a) the average of constituent species monocultures, (b) the best constituent species monoculture, and (c) commercial species monocultures. We investigated whether any advantage of mixtures over monocultures was positively related to species richness, as well as potential mechanisms driving differences in carbon stocks between mixtures and monocultures. The meta-analysis dataset included 79 comparisons from 21 sites. Carbon stocks in mixed planted forests were higher than the average of stocks in monocultures of their constituent species, containing on average 70% more carbon. Mixed planted forests also out-performed commercial monocultures, containing on average 77% more carbon. There was c.25% more carbon in mixed planted forests relative to the best performing monocultures, although this difference was not statistically significant. Overyielding was highest in four-species mixtures (richness range 2-6 species). More data providing better coverage of richness and age gradients (study sites aged 3.5-28 years) is needed to increase confidence in these results. None of the potential mechanisms we examined (nitrogen-fixer present vs absent; native vs non-native/mixed origin; tree diversity experiment vs forestry plantation) consistently explained variation in the diversity effects. This suggests that our findings are driven by a combination of small (statistically insignificant) effects from these sources or further unidentified mechanisms or some combination of the two. We conclude that increasing tree species richness in planted forests can increase carbon stocks while bringing other potential benefits associated with diversification. However, implementation will depend on the balance of these benefits relative to the operational challenges and costs of diversification.

Given the worldwide plans for extensive tree planting, we urgently need to understand how and where implementation will contribute to goals such as those for carbon sequestration. We used a long-term, large-scale native reforestation project in the Scottish Highlands to assess the response of carbon storage and other ecosystem functions to reforestation and grazing exclusion. We measured aboveground carbon, topsoil carbon, topsoil nitrogen, decomposition rates, soil invertebrate feeding activity, tree regeneration, and ground-layer and moss height at 14 sites that are in the early stages of reforestation and fenced to exclude grazing. Reforestation areas were compared to unforested and mature forest areas that are both grazed and ungrazed, using 10 × 10 m plots. Aboveground carbon in the reforestation plots (1.4 kg/m 2 [95% CI: 0.6, 2.6], average age 20 years since reforestation) was c. 8% of the mature forest plots (17.1 kg/m 2 [13.1, 21.8]). Topsoil carbon was lower in the reforestation plots (18.78 kg/m 2 [11.79, 25.78]) than in the unforested (29.82 kg/m 2 [24.34, 35.29]) or mature forest (31.39 kg/m 2 [22.91, 39.88]) plots. Responses of other functions to the reforestation and grazing interventions varied. Our results suggest that reforestation may trigger carbon loss from areas with high initial soil carbon even with low disturbance establishment, at least in the short term. Our work emphasizes where we lack knowledge: on the potential for long-term re-accumulation of soil carbon under semi-natural native reforestation, soil carbon sequestration in the deeper soil layers, and the response of soil carbon to natural regeneration.

Diversifying planted forests by increasing genetic and species diversity is often promoted as a method to improve forest resilience to climate change and reduce pest and pathogen damage. In this study, we used a young tree diversity experiment replicated at two sites in the UK to study the impacts of tree diversity and tree provenance (geographic origin) on the oak (Quercus robur) insect herbivore community and a specialist biotrophic pathogen, oak powdery mildew. Local UK, French, and Italian provenances were planted in monocultures, provenance mixtures, and species mixes, allowing us to test whether: (a) local and nonlocal provenances differ in their insect herbivore and pathogen communities, and (b) admixing trees leads to associational effects on insect herbivore and pathogen damage. Tree diversity had variable impacts on foliar organisms across sites and years, suggesting that diversity effects can be highly dependent on environmental context. Provenance identity impacted upon both herbivores and powdery mildew, but we did not find consistent support for the local adaptation hypothesis for any group of organisms studied. Independent of provenance, we found tree vigor traits (shoot length, tree height) and tree apparency (the height of focal trees relative to their surroundings) were consistent positive predictors of powdery mildew and insect herbivory. Synthesis. Our results have implications for understanding the complex interplay between tree identity and diversity in determining pest damage, and show that tree traits, partially influenced by tree genotype, can be important drivers of tree pest and pathogen loads.

Statistics is a fundamental component of the scientific toolbox, but learning the basics of this area of mathematics is one of the most challenging parts of a research training. This book gives an up-to-date introduction to the classical techniques and modern extensions of linear-model analysis—one of the most useful approaches in the analysis of scientific data in the life and environmental sciences. The book emphasizes an estimation-based approach that takes account of recent criticisms of overuse of probability values and introduces the alternative approach using information criteria. The book is based on the use of the open-source R programming language for statistics and graphics, which is rapidly becoming the lingua franca in many areas of science. This second edition adds new chapters, including one discussing some of the complexities of linear-model analysis and another introducing reproducible research documents using the R Markdown package. Statistics is introduced through worked analyses performed in R using interesting data sets from ecology, evolutionary biology, and environmental science. The data sets and R scripts are available as supporting material.

Arbuscular mycorrhizal (AM) and ectomycorrhizal (EcM) associations are critical for host-tree performance. However, how mycorrhizal associations correlate with the latitudinal tree beta-diversity remains untested. Using a global dataset of 45 forest plots representing 2,804,270 trees across 3840 species, we test how AM and EcM trees contribute to total beta-diversity and its components (turnover and nestedness) of all trees. We find AM rather than EcM trees predominantly contribute to decreasing total beta-diversity and turnover and increasing nestedness with increasing latitude, probably because wide distributions of EcM trees do not generate strong compositional differences among localities. Environmental variables, especially temperature and precipitation, are strongly correlated with beta-diversity patterns for both AM trees and all trees rather than EcM trees. Results support our hypotheses that latitudinal beta-diversity patterns and environmental effects on these patterns are highly dependent on mycorrhizal types. Our findings highlight the importance of AM-dominated forests for conserving global forest biodiversity., The relationship of mycorrhizal associations with latitudinal gradients in tree beta-diversity is unexplored. Using a global dataset approach, this study examines how trees with arbuscular mycorrhizal and ectomycorrhizal associations contribute to latitudinal beta-diversity patterns and the environmental controls of these patterns.

Globally, there is increasing interest in tree planting, leading to many country-level commitments to reforestation. In the UK, current commitments would achieve 17% forest cover by 2050, with the highest rates of forest expansion expected in Scotland. Forest expansion with native trees is expected to increase biodiversity, particularly woodland specialist species, and associated ecosystem services. Despite this, data on biodiversity changes over the early stages of reforestation are sparse, particularly for upland areas in Scotland where opportunities for forest expansion are greatest. We collected data on the response of plants, carabid beetles and birds to native reforestation and grazing exclusion, using sites reforested over the last 30 years in the Scottish Highlands. Biodiversity in ungrazed, reforested sites was compared to unforested controls and mature native forest, both grazed and ungrazed. Mean bird species richness in reforested plots (4.4 [95% CI: 3.2, 5.9]) was higher than in unforested plots (0.8 [0.5, 1.3]), but lower than in mature forest plots (7.0 [5.4, 8.3]). In contrast, there was no systematic difference in plant or carabid beetle species richness in reforested, unforested or mature forest plots, or between grazed and ungrazed plots for the species richness of any groups. Woodland specialist bird and plant species were found in the reforested plots, and richness of woodland specialist bird species was predicted to reach levels in mature forest c. 36 years after reforestation. Species assemblages differed across habitat categories. For birds and plants, species assemblages in reforested sites were intermediate to unforested and mature sites. For carabid beetles, the assemblages in mature and reforested sites were comparable and differed from unforested sites. Grazing did not strongly influence species assemblages. Policy implications. We show that woodland specialists colonise reforested sites and species assemblages transition towards those found in the target habitat within the first 30 years of reforestation with native species. Native forest should be prioritised in Scotland's future forest expansion targets, given that mature native forest is scarce and fragmented in the Scottish Highlands and that the ultimate gain from native forest expansion may accrue over long time-scales.

International audience; Our planet is facing significant changes of biodiversity across spatial scales. Although the negative effects of local biodiversity (α diversity) loss on ecosystem stability are well documented, the consequences of biodiversity changes at larger spatial scales, in particular biotic homogenization, that is, reduced species turnover across space (β diversity), remain poorly known. Using data from 39 grassland biodiversity experiments, we examine the effects of β diversity on the stability of simulated landscapes while controlling for potentially confounding biotic and abiotic factors. Our results show that higher β diversity generates more asynchronous dynamics among local communities and thereby contributes to the stability of ecosystem productivity at larger spatial scales. We further quantify the relative contributions of α and β diversity to ecosystem stability and find a relatively stronger effect of α diversity, possibly due to the limited spatial scale of our experiments. The stabilizing effects of both α and β diversity lead to a positive diversity-stability relationship at the landscape scale. Our findings demonstrate the destabilizing effect of biotic homogenization and suggest that biodiversity should be conserved at multiple spatial scales to maintain the stability of ecosystem functions and services.

Eutrophication is a widespread environmental change that usually reduces the stabilizing effect of plant diversity on productivity in local communities. Whether this effect is scale dependent remains to be elucidated. Here, we determine the relationship between plant diversity and temporal stability of productivity for 243 plant communities from 42 grasslands across the globe and quantify the effect of chronic fertilization on these relationships. Unfertilized local communities with more plant species exhibit greater asynchronous dynamics among species in response to natural environmental fluctuations, resulting in greater local stability (alpha stability). Moreover, neighborhood communities that have greater spatial variation in plant species composition within sites (higher beta diversity) have greater spatial asynchrony of productivity among communities, resulting in greater stability at the larger scale (gamma stability). Importantly, fertilization consistently weakens the contribution of plant diversity to both of these stabilizing mechanisms, thus diminishing the positive effect of biodiversity on stability at differing spatial scales. Our findings suggest that preserving grassland functional stability requires conservation of plant diversity within and among ecological communities., Eutrophication has been shown to weaken diversity-stability relationships in grasslands, but it is unclear whether the effect depends on scale. Analysing a globally distributed network of grassland sites, the authors show a positive role of beta diversity and spatial asynchrony as drivers of stability but find that nitrogen enrichment weakens the diversity-stability relationships at different spatial scales.

Significance Identifying and explaining regional differences in tropical forest dynamics, structure, diversity, and composition are critical for anticipating region-specific responses to global environmental change. Floristic classifications are of fundamental importance for these efforts. Here we provide a global tropical forest classification that is explicitly based on community evolutionary similarity, resulting in identification of five major tropical forest regions and their relationships: (i) Indo-Pacific, (ii) Subtropical, (iii) African, (iv) American, and (v) Dry forests. African and American forests are grouped, reflecting their former western Gondwanan connection, while Indo-Pacific forests range from eastern Africa and Madagascar to Australia and the Pacific. The connection between northern-hemisphere Asian and American forests is confirmed, while Dry forests are identified as a single tropical biome., Knowledge about the biogeographic affinities of the world’s tropical forests helps to better understand regional differences in forest structure, diversity, composition, and dynamics. Such understanding will enable anticipation of region-specific responses to global environmental change. Modern phylogenies, in combination with broad coverage of species inventory data, now allow for global biogeographic analyses that take species evolutionary distance into account. Here we present a classification of the world’s tropical forests based on their phylogenetic similarity. We identify five principal floristic regions and their floristic relationships: (i) Indo-Pacific, (ii) Subtropical, (iii) African, (iv) American, and (v) Dry forests. Our results do not support the traditional neo- versus paleotropical forest division but instead separate the combined American and African forests from their Indo-Pacific counterparts. We also find indications for the existence of a global dry forest region, with representatives in America, Africa, Madagascar, and India. Additionally, a northern-hemisphere Subtropical forest region was identified with representatives in Asia and America, providing support for a link between Asian and American northern-hemisphere forests.

Occasional periods of drought are typical of most tropical forests, but climate change is increasing drought frequency and intensity in many areas across the globe, threatening the structure and function of these ecosystems. The effects of intermittent drought on tropical tree communities remain poorly understood and the potential impacts of intensified drought under future climatic conditions are even less well known. The response of forests to altered precipitation will be determined by the tolerances of different species to reduced water availability and the interactions among plants that alleviate or exacerbate the effects of drought. Here, we report the response of experimental monocultures and mixtures of tropical trees to simulated drought, which reveals a fundamental shift in the nature of interactions among species. Weaker competition for water in diverse communities allowed seedlings to maintain growth under drought while more intense competition among conspecifics inhibited growth under the same conditions. These results show that reduced competition for water among species in mixtures mediates community resistance to drought. The delayed onset of competition for water among species in more diverse neighbourhoods during drought has potential implications for the coexistence of species in tropical forests and the resilience of these systems to climate change. Reduced competition for water among species in mixed tropical plant communities mediates community resistance to drought: weaker competition permits growth maintenance in drought, whereas stronger competition inhibits it.

Aim Palms are an iconic, diverse and often abundant component of tropical ecosystems that provide many ecosystem services. Being monocots, tree palms are evolutionarily, morphologically and physiologically distinct from other trees, and these differences have important consequences for ecosystem services (e.g., carbon sequestration and storage) and in terms of responses to climate change. We quantified global patterns of tree palm relative abundance to help improve understanding of tropical forests and reduce uncertainty about these ecosystems under climate change. Location Tropical and subtropical moist forests. Time period Current. Major taxa studied Palms (Arecaceae). Methods We assembled a pantropical dataset of 2,548 forest plots (covering 1,191ha) and quantified tree palm (i.e., ≥10cm diameter at breast height) abundance relative to co‐occurring non‐palm trees. We compared the relative abundance of tree palms across biogeographical realms and tested for associations with palaeoclimate stability, current climate, edaphic conditions and metrics of forest structure. Results On average, the relative abundance of tree palms was more than five times larger between Neotropical locations and other biogeographical realms. Tree palms were absent in most locations outside the Neotropics but present in >80% of Neotropical locations. The relative abundance of tree palms was more strongly associated with local conditions (e.g., higher mean annual precipitation, lower soil fertility, shallower water table and lower plot mean wood density) than metrics of long‐term climate stability. Life‐form diversity also influenced the patterns; palm assemblages outside the Neotropics comprise many non‐tree (e.g., climbing) palms. Finally, we show that tree palms can influence estimates of above‐ground biomass, but the magnitude and direction of the effect require additional work. Conclusions Tree palms are not only quintessentially tropical, but they are also overwhelmingly Neotropical. Future work to understand the contributions of tree palms to biomass estimates and carbon cycling will be particularly crucial in Neotropical forests.

AimSoil microorganisms are essential for the functioning of terrestrial ecosystems. Although soil microbial communities and functions may be linked to the tree species composition and diversity of forests, there has been no comprehensive study of how general potential relationships are and if these are context-dependent. A global network of tree diversity experiments (TreeDivNet) allows for a first examination of tree diversity-soil microbial function relationships across environmental gradients.LocationGlobalMajor Taxa StudiedSoil microorganismsMethodsSoil samples collected from eleven tree diversity experiments in four biomes across four continents were used to measure soil basal respiration, microbial biomass, and carbon use efficiency using the substrate-induced respiration method. All samples were measured using the same analytical device in the same laboratory to prevent measurement bias. We used linear mixed-effects models to examine the effects of tree species diversity, environmental conditions, and their interactions on soil microbial functions.ResultsAcross biodiversity experiments, abiotic drivers, mainly soil water content, significantly increased soil microbial functions. Potential evapotranspiration (PET) increased, whereas soil C-to-N ratio (CN) decreased soil microbial functions under dry soil conditions, but high soil water content reduced the importance of other abiotic drivers. Tree species richness and phylogenetic diversity had overall similar, but weak and context-dependent (climate, soil abiotic variables) effects on soil microbial respiration. Positive tree diversity effects on soil microbial respiration were most pronounced at low PET, low soil CN, and high tree density. Soil microbial functions increased with the age of the experiment.Main conclusionsOur results point at the importance of soil water content for maintaining high levels of soil microbial functions and modulating effects of other environmental drivers. Moreover, overall tree diversity effects on soil microbial functions seem to be negligible in the short term (experiments were 1-18 years old). However, context-dependent tree diversity effects (climate, soil abiotic variables) have greater importance at high tree density, and significant effects of experimental age call for longer-term studies. Such systematic insights are key to better integrate soil carbon dynamics into the management of afforestation projects across environmental contexts, as today’s reforestation efforts remain focused largely on aboveground carbon storage and are still dominated by less diverse forests stands of commercial species.

1. Tree health declines can be caused by interactions between pests and pathogens and many studies have shown a reduction in their damage in mixed species forests compared to monocultures. Yet few authors have considered tree diversity effects on both groups simultaneously. Moreover, it is unclear whether diversity effects on tree pests and pathogens are robust to changes in abiotic conditions, such as drought. 2. We addressed tree diversity effects on foliar insect herbivory, oak powdery mildew and their interaction under contrasting water regimes in a large-scale tree diversity experiment in SW France. Using an irrigation treatment that alleviated drought conditions, we were able to experimentally assess the effects of tree diversity under contrasting abiotic environments. We surveyed plots along a richness gradient from one to four tree species, in which a focal study species of oak (Quercus robur) was mixed with other oak species (Q. pyrenaica and Q. ilex) and a taller, broadleaved species (Betula pendula). 3. Increasing tree species richness lowered leaf miner abundance, leaf chewer damage and oak powdery mildew infection, consistent with a protective effect of resource dilution. However, richness effects on leaf miners were stronger in irrigated compared to non-irrigated blocks, indicating that environmental conditions can modulate diversity effects. Separate from the effect of tree species richness, the presence of birch in a plot increased damage by leaf chewers and powdery mildew, but lowered leaf miner damage, suggesting additional tree neighbour identity effects potentially linked to modulation of microclimate. We found a negative association between leaf miner abundance and oak powdery mildew, consistent with antagonism between oak damage agents. 4. Synthesis . Overall, our study illustrates the importance of considering both tree diversity and composition (neighbour identity) in designing forests more resistant to pest and pathogen damage.

International audience; The Loreau and Hector (2019) Comment on our paper (Pillai and Gouhier, 2019) failed to address the two core elements of our critique, both the circularity of the BEF research program, in general, and the mathematical flaws of the Loreau-Hector partitioning scheme, in particular. Loreau and Hector avoided dealing with the first part of our critique by arguing against a non-existent claim that all biodiversity effects could be reduced to coexistence, while the mathematical flaws in the Loreau-Hector partitioning method that we described in the second part of our critique were ignored altogether. Here, we address these misconceptions and demonstrate that all of the claims that were made in our original paper hold. We conclude that (i) BEF studies need to adopt baselines that account for coexistence in order to avoid overestimating the effects of biodiversity and (ii) the LH partitioning method should not be used unless the linearity of the abundance-ecosystem functioning relationship in monocultures can be verified for all species.

Summary Invasive tree pests and diseases present some of the greatest global threats to forests, and the recent global acceleration in invasions has caused massive ecological damage 1 , 2 . Calls to improve biosecurity have, however, often lost out to economic arguments in favour of trade [3] . Human activities, such as trade, move organisms between continents, and interventions to reduce risk of introductions inevitably incur financial costs. No previous studies have attempted to estimate the full economic cost of a tree disease, and the economic imperative to improve biosecurity may have been underappreciated. We set out to estimate the cost of the dieback of ash, Fraxinus excelsior, caused by Hymenoscyphus fraxineus, in Great Britain, and investigate whether this may be the case [4] .

Logged tropical forests represent a major opportunity for preserving biodiversity and sequestering carbon, playing a large role in meeting global forest restoration targets. Left alone, these ecosystems have been expected to undergo natural regeneration and succession towards old growth forests, but extreme drought events may challenge this process. While old growth forests possess a certain level of resilience, we lack understanding as to how logging may affect forest responses to drought. This study examines the drought–logging interaction in seedling dynamics within a landscape of logged and unlogged forests in Sabah Malaysia, based on 73 plots monitored before and after the 2015-16 El Niño drought. Drought increased seedling mortality in all forests, but the magnitude of this impact was modulated by logging intensity, with forests with lower canopy leaf area index (LAI) and above ground biomass (AGB) experiencing greater drought induced mortality. Moreover, community traits in more heavily logged forests shifted towards being more ruderal after drought, suggesting that the trajectory of forest succession had been reversed. These results indicate that with reoccurring strong droughts under a changing climate, logged forests that have had over half of their biomass removed may suffer permanently arrested succession. Targeted management interventions may therefore be necessary to lift the vulnerable forests above the biomass threshold.

1. High-quality abundance data is expensive and time consuming to collect and often highly limited in availability. Nonetheless, accurate, high-resolution abundance distributions are essential for many ecological applications ranging from species conservation to epidemiology. Producing models that can predict abundance well, with good resolution over large areas, has therefore been an important aim in ecology, but poses considerable challenges. 2. We present a two-stage approach to modelling abundance, combining two established techniques. First, we produce ensemble species distribution models (SDMs) of trees in Great Britain at a fine resolution, using much more common presence-absence data and key environmental variables. We then use Random Forest regression to predict abundance by linking the results of the SDMs to a much smaller amount of abundance data. 3. We show that this method performs well in predicting the abundance of 20 out of 25 tested British tree species, a group that is generally considered challenging for modelling distributions due to the strong influence of human activities. 4. Maps of predicted tree abundance for the whole of Great Britain are provided at 1 km resolution. Abundance maps have a far wider variety of applications than presence only maps, and these maps should allow improvements to aspects of woodland management and conservation including analysis of habitats and ecosystem functioning, epidemiology and disease management, providing a useful contribution to the protection of British trees. We also provide complete R scripts to facilitate application of the approach to other scenarios.

Complementary resource use has often been claimed to explain positive effects of plant diversity on ecosystem functioning, but the underlying mechanisms of complementarity have rarely been directly quantified in forest systems. The aim of this study was to characterize spatial and temporal water uptake of subtropical tree saplings, to assess the role of interspecific interaction on water uptake patterns and to quantify species niche breadth and overlap. Our experiment comprised two deciduous ( Castanea henryi , Quercus serrata ) and two evergreen tree species ( Elaeocarpus decipiens , Schima superba ) that were each planted in monoculture and 4-species mixture. We used deuterium-enriched water to trace seasonal water uptake from 5 cm and 20 cm soil depth. Castanea exploited predominantly the upper soil (74% of overall tracer uptake across treatments) whereas Quercus (50%), Elaeocarpus (57%) and Schima (62%) tended to use both soil layers more equally. Species identity had an overall significant effect on isotopic enrichment in stem water. There was no effect of species richness: niche breadth and overlap of single species was not affected by interspecific interactions in mixtures. Niche overlap between deciduous species was lowest (74%) whereas the two evergreen species had similar water uptake patterns (91%). According to our results, interspecific competition did not alter water uptake patterns of the studied species during the early phase of forest establishment. Thus, soil water uptake complementarity could only occur through inherent (fundamental) specific differences in water uptake niches based on sapling specialization, while phenotypic adjustments to interspecific interaction or neighbor diversity are less important.

A Correction to this paper has been published: https://doi.org/10.1038/s41467-021-20997-9.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

1.Acute outbreaks of pests and disease are increasingly affecting tree populations around the world, causing widespread ecological effects. In Britain, ash dieback Hymenoscyphus fraxineus (Baral et al.) has severe impacts on common ash (Fraxinus excelsior L.) populations, and the emerald ash borer (Agrilus planipennis Fairmaire) is likely to add to the impact in future. This will cause significant changes to the character and functioning of many ecosystems. However, the nature of these changes and the best approach for conserving ecosystems after ash loss are not clear. 2.We present a method to locate those areas most ecologically vulnerable to loss of a major tree species (common ash) and identify the resultant damage to distinctive ecosystem properties. This method uses the functional traits of species and their distributions to map the potential degree of change in traits across space and recommend management approaches to reduce the change. An analytic hierarchy process is used to score traits according to ecological importance. 3.Our results indicate that in some areas of Britain, provision of ash‐associated traits could be reduced by over 50% if all ash is lost. Certain woodland types, and trees outside woodlands, may be especially vulnerable to ash loss. However, compensatory growth by other species could halve this impact in the longer term. 4.We offer management guidance for reducing ecosystem vulnerability to ash loss, including recommending appropriate alternative tree species to encourage through planting or management in particular areas and woodland types. 5.Synthesis and applications. The method described in this paper allows spatially explicit assessment of species traits to be used in the restoration of ecosystems for the first time. We offer practical recommendations for the ash dieback outbreak in Britain to help conserve functional traits in ecosystems affected by the loss of ash. This technique is widely applicable to a range of restoration and conservation scenarios and represents a step forward in the use of functional traits in conservation.

International audience; Despite considerable research demonstrating that biodiversity increases productivity in forests and regulates herbivory and pathogen damage, there remain gaps in our understanding of the shape, magnitude, and generality of these biodiversity-ecosystem functioning (BEF) relationships. Here, we review findings from TreeDivNet, a global network of 25 tree diversity experiments, on relationships between levels of biodiversity and (a) tree growth and survival and (b) damage to trees from pests and pathogens. Tree diversity often improved the survival and above- and belowground growth of young trees. The mechanistic bases of the diversity effects on tree growth and survival include both selection effects (i.e., an increasing impact of particular species in more species-rich communities) and complementary effects (e.g. related to resource differentiation and facilitation). Plant traits and abiotic stressors may mediate these relationships. Studies of the responses of invertebrate and vertebrate herbivory and pathogen damage have demonstrated that trees in more diverse experimental plots may experience more, less, or similar damage compared to conspecific trees in less diverse plots. Documented mechanisms producing these patterns include changes in concentration, frequency, and apparency of hosts; herbivore and pathogen diet breadth; the spatial scale of interactions; and herbivore and pathogen regulation by natural enemies. Our review of findings from TreeDivNet indicates that tree diversity experiments are extending BEF research across systems and scales, complementing previous BEF work in grasslands by providing opportunities to use remote sensing and spectral approaches to study BEF dynamics, integrate belowground and aboveground approaches, and trace the consequences of tree physiology for ecosystem functioning. This extension of BEF research into tree-dominated systems is improving ecologists’ capacity to understand the mechanistic bases behind BEF relationships. Tree diversity experiments also present opportunities for novel research. Since experimental tree diversity plantations enable measurements at tree, neighbourhood and plot level, they allow for explicit consideration of temporal and spatial scales in BEF dynamics. Presently, most TreeDivNet experiments have run for less than ten years. Given the longevity of trees, exciting results on BEF relationships are expected in the future.

International audience; Biodiversity loss decreases ecosystem functioning at the local scales at which species interact, but it remains unclear how biodiversity loss affects ecosystem functioning at the larger scales of space and time that are most relevant to biodiversity conservation and policy. Theory predicts that additional insurance effects of biodiversity on ecosystem functioning could emerge across time and space if species respond asynchronously to environmental variation and if species become increasingly dominant when and where they are most productive. Even if only a few dominant species maintain ecosystem functioning within a particular time and place, ecosystem functioning may be enhanced by many different species across many times and places (b-diversity). Here, we develop and apply a new approach to estimate these previously unquantified insurance effects of biodiversity on ecosystem functioning that arise due to species turnover across times and places. In a long-term (18-year) grassland plant diversity experiment, we find that total insurance effects are positive in sign and substantial in magnitude, amounting to 19% of the net biodiversity effect, mostly due to temporal insurance effects. Species loss can therefore reduce ecosystem functioning both locally and by eliminating species that would otherwise enhance ecosystem functioning across temporally fluctuating and spatially heterogeneous environments.

Tree diversity improves forest productivity Experimental studies in grasslands have shown that the loss of species has negative consequences for ecosystem functioning. Is the same true for forests? Huang et al. report the first results from a large biodiversity experiment in a subtropical forest in China. The study combines many replicates, realistic tree densities, and large plot sizes with a wide range of species richness levels. After 8 years of the experiment, the findings suggest strong positive effects of tree diversity on forest productivity and carbon accumulation. Thus, changing from monocultures to more mixed forests could benefit both restoration of biodiversity and mitigation of climate change. Science , this issue p. 80

Climate is widely recognised as an important determinant of the latitudinal diversity gradient. However, most existing studies make no distinction between direct and indirect effects of climate, which substantially hinders our understanding of how climate constrains biodiversity globally. Using data from 35 large forest plots, we test hypothesised relationships amongst climate, topography, forest structural attributes (stem abundance, tree size variation and stand basal area) and tree species richness to better understand drivers of latitudinal tree diversity patterns. Climate influences tree richness both directly, with more species in warm, moist, aseasonal climates and indirectly, with more species at higher stem abundance. These results imply direct limitation of species diversity by climatic stress and more rapid (co-)evolution and narrower niche partitioning in warm climates. They also support the idea that increased numbers of individuals associated with high primary productivity are partitioned to support a greater number of species.

Biodiversity is declining in many local communities while also becoming increasingly homogenized across space. Experimentsshow that local plant species loss reduces ecosystem functioning and services, but the role of spatial homogenization of communitycomposition and the potential interaction between diversity at different scales in maintaining ecosystem functioningremains unclear, especially when many functions are considered (ecosystem multifunctionality). We present an analysis ofeight ecosystem functions measured in 65 grasslands worldwide. We find that more diverse grasslands?those with both species-rich local communities (α-diversity) and large compositional differences among localities (β-diversity)?had higher levelsof multifunctionality. Moreover, α- and β-diversity synergistically affected multifunctionality, with higher levels of diversity at one scale amplifying the contribution to ecological functions at the other scale. The identity of species influencing ecosystem functioning differed among functions and across local communities, explaining why more diverse grasslands maintained greater functionality when more functions and localities were considered. These results were robust to variation in environmental drivers. Our findings reveal that plant diversity, at both local and landscape scales, contributes to the maintenance of multiple ecosystem services provided by grasslands. Preserving ecosystem functioning therefore requires conservation of biodiversity both within and among ecological communities. Fil: Hautier, Yann. University of Utrecht; Países Bajos Fil: Isbell, Forest. University of Minnesota; Estados Unidos Fil: Borer, Elizabeth T.. University of Minnesota; Estados Unidos Fil: Seabloom, Eric W.. University of Minnesota; Estados Unidos Fil: Harpole, Stanley. Martin Luther University Halle-Wittenberg; Alemania Fil: Lind, Eric M.. University of Minnesota; Estados Unidos Fil: MacDougall, Andrew. University of Guelph; Canadá Fil: Stevens, Carly. Lancaster University, ; Reino Unido Fil: Adler, Peter. Utah State University; Estados Unidos Fil: Alberti, Juan. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Bakker, Jonathan D. University of Washington, ; Estados Unidos Fil: Brudvig, Lars A. Michigan State University; Estados Unidos Fil: Buckley, Yvonne M. Trinity College Dublin; Irlanda Fil: Cadotte, Marc. University of Toronto Scarborough,; Canadá Fil: Caldeira, María C. University of Lisbon; Portugal Fil: Chaneton, Enrique Jose. Universidad de Buenos Aires; Argentina Fil: Chu, Chengjin. Sun Yat-sen University; China Fil: Daleo, Pedro. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Dickman, Christopher R. The University of Sydney; Australia Fil: Dwyer, John M. The University of Queensland; Australia Fil: Eskelinen, Anu. University of Oulu; Finlandia Fil: Fay, Philip. USDA-ARS, Grassland Soil and Water Lab; Estados Unidos Fil: Firn, Jennifer. Queensland University of Technology; Australia Fil: Hagenah, Nicole. University of KwaZulu-Natal; Sudáfrica Fil: Hillebrand, Helmut. University Oldenburg; Alemania Fil: Iribarne, Oscar Osvaldo. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Kirkman, Kevin P.. University of KwaZulu-Natal; Sudáfrica Fil: Knops, Johannes M. H. University of Nebraska, Lincoln; Estados Unidos Fil: La Pierre, Kimberly J. Smithsonian Environmental Research Center; Estados Unidos Fil: McCulley, Rebecca L. University of Kentucky; Estados Unidos Fil: Morgan, John W.. La Trobe University; Australia Fil: Pärtel, Meelis. University of Tartu; Estonia Fil: Pascual, Jesus Maria. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Price, Jodi N. Charles Sturt University; Australia Fil: Prober, Suzanne M. CSIRO Land and Water; Australia Fil: Risch, Anita C. Swiss Federal Institute of Forest, Snow and Landscape Research; Suiza Fil: Sankaran, Mahesh. National Centre for Biological Sciences; India Fil: Schuetz, Martin. Swiss Federal Institute of Forest, Snow and Landscape Research; Suiza Fil: Standish, Rachel J. Murdoch University; Australia Fil: Virtanen, Risto. University of Oulu; Finlandia Fil: Wardle, Glenda M. The University of Sydney; Australia Fil: Yahdjian, María Laura. UBA; Argentina Fil: Hector, Andy. University of Oxford; Reino Unido

Biodiversity-ecosystem functioning (BEF) research has extended its scope from communities that are short-lived or reshape their structure annually to structurally complex forest ecosystems. The establishment of tree diversity experiments poses specific methodological challenges for assessing the multiple functions provided by forest ecosystems. In particular, methodological inconsistencies and nonstandardized protocols impede the analysis of multifunctionality within, and comparability across the increasing number of tree diversity experiments. By providing an overview on key methods currently applied in one of the largest forest biodiversity experiments, we show how methods differing in scale and simplicity can be combined to retrieve consistent data allowing novel insights into forest ecosystem functioning. Furthermore, we discuss and develop recommendations for the integration and transferability of diverse methodical approaches to present and future forest biodiversity experiments. We identified four principles that should guide basic decisions concerning method selection for tree diversity experiments and forest BEF research: (1) method selection should be directed toward maximizing data density to increase the number of measured variables in each plot. (2) Methods should cover all relevant scales of the experiment to consider scale dependencies of biodiversity effects. (3) The same variable should be evaluated with the same method across space and time for adequate larger-scale and longer-time data analysis and to reduce errors due to changing measurement protocols. (4) Standardized, practical and rapid methods for assessing biodiversity and ecosystem functions should be promoted to increase comparability among forest BEF experiments. We demonstrate that currently available methods provide us with a sophisticated toolbox to improve a synergistic understanding of forest multifunctionality. However, these methods require further adjustment to the specific requirements of structurally complex and long-lived forest ecosystems. By applying methods connecting relevant scales, trophic levels, and above- and belowground ecosystem compartments, knowledge gain from large tree diversity experiments can be optimized. Biodiversity–ecosystem functioning (BEF) research has extended its scope from communities that are short‐lived or reshape their structure annually to structurally complex forest ecosystems. The establishment of tree diversity experiments poses specific methodological challenges for assessing the multiple functions provided by forest ecosystems. In particular, methodological inconsistencies and nonstandardized protocols impede the analysis of multifunctionality within, and comparability across the increasing number of tree diversity experiments. By providing an overview on key methods currently applied in one of the largest forest biodiversity experiments, we show how methods differing in scale and simplicity can be combined to retrieve consistent data allowing novel insights into forest ecosystem functioning. Furthermore, we discuss and develop recommendations for the integration and transferability of diverse methodical approaches to present and future forest biodiversity experiments. We identified four principles that should guide basic decisions concerning method selection for tree diversity experiments and forest BEF research: (1) method selection should be directed toward maximizing data density to increase the number of measured variables in each plot. (2) Methods should cover all relevant scales of the experiment to consider scale dependencies of biodiversity effects. (3) The same variable should be evaluated with the same method across space and time for adequate larger‐scale and longer‐time data analysis and to reduce errors due to changing measurement protocols. (4) Standardized, practical and rapid methods for assessing biodiversity and ecosystem functions should be promoted to increase comparability among forest BEF experiments. We demonstrate that currently available methods provide us with a sophisticated toolbox to improve a synergistic understanding of forest multifunctionality. However, these methods require further adjustment to the specific requirements of structurally complex and long‐lived forest ecosystems. By applying methods connecting relevant scales, trophic levels, and above‐ and belowground ecosystem compartments, knowledge gain from large tree diversity experiments can be optimized.

Forest ecosystems contribute substantially to global terrestrial primary productivity and climate regulation, but, in contrast to grasslands, experimental evidence for a positive biodiversity-productivity relationship in highly diverse forests is still lacking1. Here, we provide such evidence from a large forest biodiversity experiment with a novel design2 in subtropical China. Productivity (stand-level tree basal area, aboveground volume and carbon and their annual increment) increased linearly with the logarithm of tree species richness. Additive partitioning3 showed that increasing positive complementarity effects combined with weakening negative selection effects caused a strengthening of the relationship over time. In 2-species mixed stands, complementary effects increased with functional distance and selection effects with vertical crown dissimilarity between species. Understorey shrubs reduced stand-level tree productivity, but this effect of competition was attenuated by shrub species richness, indicating that a diverse understorey may facilitate overall ecosystem functioning. Identical biodiversity-productivity relationships were found in plots of different size, suggesting that extrapolation to larger scales is possible. Our results highlight the potential of multi-species afforestation strategies to simultaneously contribute to mitigation of climate change and biodiversity restoration.

The effects of biodiversity on ecosystem functioning generally increase over time, but the underlying processes remain unclear. Using 26 long-term grassland and forest experimental ecosystems, we demonstrate that biodiversity–ecosystem functioning relationships strengthen mainly by greater increases in functioning in high-diversity communities in grasslands and forests. In grasslands, biodiversity effects also strengthen due to decreases in functioning in low-diversity communities. Contrasting trends across grasslands are associated with differences in soil characteristics.

[Departement_IRSTEA]Territoires [TR1_IRSTEA]SEDYVIN; International audience; Widespread tree mortality associated with drought has been observed on all forested continents and global change is expected to exacerbate vegetation vulnerability. Forest mortality has implications for future biosphere–atmosphere interactions of carbon, water and energy balance, and is poorly represented in dynamic vegetation models. Reducing uncertainty requires improved mortality projections founded on robust physiological processes. However, the proposed mechanisms of drought-induced mortality, including hydraulic failure and carbon starvation, are unresolved. A growing number of empirical studies have investigated these mechanisms, but data have not been consistently analysed across species and biomes using a standardized physiological framework. Here, we show that xylem hydraulic failure was ubiquitous across multiple tree taxa at drought-induced mortality. All species assessed had 60% or higher loss of xylem hydraulic conductivity, consistent with proposed theoretical and modelled survival thresholds. We found diverse responses in non-structural carbohydrate reserves at mortality, indicating that evidence supporting carbon starvation was not universal. Reduced non-structural carbohydrates were more common for gymnosperms than angiosperms, associated with xylem hydraulic vulnerability, and may have a role in reducing hydraulic function. Our finding that hydraulic failure at drought-induced mortality was persistent across species indicates that substantial improvement in vegetation modelling can be achieved using thresholds in hydraulic function

Biodiversity enhances many of nature?s benefits to people, including the regulation of climate and the production of woodin forests, livestock forage in grasslands and fish in aquatic ecosystems. Yet people are now driving the sixth mass extinctionevent in Earth?s history. Human dependence and influence on biodiversity have mainly been studied separately and atcontrasting scales of space and time, but new multiscale knowledge is beginning to link these relationships. Biodiversityloss substantially diminishes several ecosystem services by altering ecosystem functioning and stability, especially at thelarge temporal and spatial scales that are most relevant for policy and conservation. Fil: Isbell, Forest. University of Minnesota; Estados Unidos Fil: Gonzalez, Andrew. Mcgill University; Canadá Fil: Loreau, Michael. Paul Sabatier University. Centre for Biodiversity Theory and Modelling, Theoretical and Experimental Ecology Station; Francia Fil: Cowles, Jane. University of Minnesota; Estados Unidos Fil: Díaz, Sandra Myrna. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto Multidisciplinario de Biología Vegetal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto Multidisciplinario de Biología Vegetal; Argentina Fil: Hector, Andy. University of Oxford; Reino Unido Fil: Mace, Georgina M.. University College London; Reino Unido Fil: Wardle, David A.. Swedish University of Agricultural Sciences; Suecia. Nanyang Technological University; Singapur Fil: O'Connor, Mary I.. University of British Columbia; Canadá Fil: Duffy, J. Emmett. Smithsonian Institution. Tennenbaum Marine Observatories Network; Estados Unidos Fil: Turnbull, Lindsay A.. University of Oxford. Department of Plant Sciences; Reino Unido Fil: Thompson, Patrick L.. University of British Columbia; Canadá Fil: Larigauderie, Anne. Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services; Alemania

Drought regimes can be characterized by the variability in the quantity of rainfall and the duration of rainless periods. However, most research on plant response to drought has ignored the impacts of rainfall variation, especially with regard to the influence of nonstructural carbohydrates (NSCs) in promoting drought resistance. To test the hypothesis that these components of drought differentially affect NSC dynamics and seedling resistance, we tracked NSC in plant tissues of tropical tree seedlings in response to manipulations of the volume and frequency of water applied. NSC concentrations decreased in woody tissues under infrequent-high watering but increased under no watering. A faster decline of growth relative to stomatal conductance in the no watering treatment was consistent with NSC accumulation as a result of an uncoupling of growth and photosynthesis, while usage of stored NSCs in woody tissues to maintain function may account for the NSC decline under infrequent-high watering. NSCs, and specifically stem NSCs, contributed to drought resistance under severe water deficits, while NSCs had a less clear role in drought resistance to variability in water availability. The contrasting response of NSCs to water variability and deficit indicates that unique processes support seedling resistance to these components of drought.

Background: Connections between mature trees and seedlings via ectomycorrhizal (EcM) hyphal networks existing in dipterocarp-dominated tropical rain forests of South-east Asia could have strong implications for seedling growth and survival and the maintenance of high diversity in such forests. Aim: To test whether EcM hyphal network connections are important for the growth and survival of dipterocarp seedlings. Methods: We conducted four independent experiments that prevented contact of experimental seedlings with an EcM network by using a series of fine meshes and/or plastic barriers. We measured the growth and survival (and foliar δ13C in one experiment) of seedlings of six dipterocarp species over intervals ranging from 11 to 29 months. Results: Seedling growth (diameter, height or leaf number) was unaffected by exclusion from the EcM network in three experiments and there were no differences in foliar δ13C values in the fourth. Seedling survival was reduced following exclusion from the EcM network in one experiment. Our results give little support to the hypothesis that dipterocarp seedlings growing in the shaded forest understorey benefit from being connected, through a common EcM network, to surrounding trees. Conclusions: We suggest that our negative results, in contrast to studies conducted in low diversity boreo-temperate or tropical forests, are due to these high diversity forests lacking host species-specific EcM fungi and therefore providing little opportunity for adaptive support of seedlings via hyphal networks.