Mapping carbon accumulation potential from global natural forest regrowth.

To constrain global warming, we must strongly curtail greenhouse gas emissions and capture excess atmospheric carbon dioxide1,2. Regrowing natural forests is a prominent strategy for capturing additional carbon3, but accurate assessments of its potential are limited by uncertainty and variability in carbon accumulation rates2,3. To assess why and where rates differ, here we compile 13,112 georeferenced measurements of carbon accumulation. Climatic factors explain variation in rates better than land-use history, so we combine the field measurements with 66 environmental covariate layers to create a global, one-kilometre-resolution map of potential aboveground carbon accumulation rates for the first 30 years of natural forest regrowth. This map shows over 100-fold variation in rates across the globe, and indicates that default rates from the Intergovernmental Panel on Climate Change (IPCC)4,5 may underestimate aboveground carbon accumulation rates by 32 per cent on average and do not capture eight-fold variation within ecozones. Conversely, we conclude that maximum climate mitigation potential from natural forest regrowth is 11 per cent lower than previously reported3 owing to the use of overly high rates for the location of potential new forest. Although our data compilation includes more studies and sites than previous efforts, our results depend on data availability, which is concentrated in ten countries, and data quality, which varies across studies. However, the plots cover most of the environmental conditions across the areas for which we predicted carbon accumulation rates (except for northern Africa and northeast Asia). We therefore provide a robust and globally consistent tool for assessing natural forest regrowth as a climate mitigation strategy.

Higher tree diversity is linked to higher tree mortality.

Examining the relationship between tree diversity and ecosystem functioning has been a recent focus of forest ecology. Particular emphasis has been given to the impact of tree diversity on productivity and to its potential to mitigate negative global change effects; however, little attention has been paid to tree mortality. This is critical because both tree mortality and productivity underpin forest ecosystem dynamics and therefore forest carbon sequestration. Neglecting tree mortality leaves a large part of the picture undocumented. Here we show that increasingly diverse forest stands have increasingly high mortality probabilities. We found that the most species-rich stands in temperate biomes had mortality probabilities more than sevenfold higher than monospecific stands (∼0.6% year−1 in monospecific stands to 4.0% year−1 in the most species-rich stands) while in boreal stands increases were less pronounced but still significant (∼1.1% year−1 in monospecific stands to 1.8% year−1 in the most species-rich stands). Tree species richness was the third-most-important predictor of mortality in our models in temperate forests and the fifth-most-important predictor in boreal forests. Our results highlight that while the promotion of tree diversity undoubtedly has many positive effects on ecosystem functioning and the services that trees provide to humanity, it remains important to consider all aspects of forest dynamics in order to properly predict the implications of maintaining and promoting tree diversity.

Tree diversity enhances predation by birds but not by arthropods across climate gradients.

Tree diversity can promote both predator abundance and diversity. However, whether this translates into increased predation and top-down control of herbivores across predator taxonomic groups and contrasting environmental conditions remains unresolved. We used a global network of tree diversity experiments (TreeDivNet) spread across three continents and three biomes to test the effects of tree species richness on predation across varying climatic conditions of temperature and precipitation. We recorded bird and arthropod predation attempts on plasticine caterpillars in monocultures and tree species mixtures. Both tree species richness and temperature increased predation by birds but not by arthropods. Furthermore, the effects of tree species richness on predation were consistent across the studied climatic gradient. Our findings provide evidence that tree diversity strengthens top-down control of insect herbivores by birds, underscoring the need to implement conservation strategies that safeguard tree diversity to sustain ecosystem services provided by natural enemies in forests.

Mycorrhizal associations modify tree diversity-productivity relationships across experimental tree plantations.

Decades of studies have demonstrated links between biodiversity and ecosystem functioning, yet the generality of the relationships and the underlying mechanisms remain unclear, especially for forest ecosystems. Using 11 tree-diversity experiments, we tested tree species richness-community productivity relationships and the role of arbuscular (AM) or ectomycorrhizal (ECM) fungal-associated tree species in these relationships. Tree species richness had a positive effect on community productivity across experiments, modified by the diversity of tree mycorrhizal associations. In communities with both AM and ECM trees, species richness showed positive effects on community productivity, which could have resulted from complementarity between AM and ECM trees. Moreover, both AM and ECM trees were more productive in mixed communities with both AM and ECM trees than in communities assembled by their own mycorrhizal type of trees. In communities containing only ECM trees, species richness had a significant positive effect on productivity, whereas species richness did not show any significant effects on productivity in communities containing only AM trees. Our study provides novel explanations for variations in diversity-productivity relationships by suggesting that tree-mycorrhiza interactions can shape productivity in mixed-species forest ecosystems.

Leaf bacterial diversity mediates plant diversity and ecosystem function relationships.

Research on biodiversity and ecosystem functioning has demonstrated links between plant diversity and ecosystem functions such as productivity. At other trophic levels, the plant microbiome has been shown to influence host plant fitness and function, and host-associated microbes have been proposed to influence ecosystem function through their role in defining the extended phenotype of host organisms However, the importance of the plant microbiome for ecosystem function has not been quantified in the context of the known importance of plant diversity and traits. Here, using a tree biodiversity-ecosystem functioning experiment, we provide strong support for the hypothesis that leaf bacterial diversity is positively linked to ecosystem productivity, even after accounting for the role of plant diversity. Our results also show that host species identity, functional identity and functional diversity are the main determinants of leaf bacterial community structure and diversity. Our study provides evidence of a positive correlation between plant-associated microbial diversity and terrestrial ecosystem productivity, and a new mechanism by which models of biodiversity-ecosystem functioning relationships can be improved.

Late-spring frost risk between 1959 and 2017 decreased in North America but increased in Europe and Asia.

Late-spring frosts (LSFs) affect the performance of plants and animals across the world's temperate and boreal zones, but despite their ecological and economic impact on agriculture and forestry, the geographic distribution and evolutionary impact of these frost events are poorly understood. Here, we analyze LSFs between 1959 and 2017 and the resistance strategies of Northern Hemisphere woody species to infer trees' adaptations for minimizing frost damage to their leaves and to forecast forest vulnerability under the ongoing changes in frost frequencies. Trait values on leaf-out and leaf-freezing resistance come from up to 1,500 temperate and boreal woody species cultivated in common gardens. We find that areas in which LSFs are common, such as eastern North America, harbor tree species with cautious (late-leafing) leaf-out strategies. Areas in which LSFs used to be unlikely, such as broad-leaved forests and shrublands in Europe and Asia, instead harbor opportunistic tree species (quickly reacting to warming air temperatures). LSFs in the latter regions are currently increasing, and given species' innate resistance strategies, we estimate that ∼35% of the European and ∼26% of the Asian temperate forest area, but only ∼10% of the North American, will experience increasing late-frost damage in the future. Our findings reveal region-specific changes in the spring-frost risk that can inform decision-making in land management, forestry, agriculture, and insurance policy.

No complementarity no gain-Net diversity effects on tree productivity occur once complementarity emerges during early stand development.

Although there is compelling evidence that tree diversity has an overall positive effect on forest productivity, there are important divergences among studies on the nature and strength of these diversity effects and their timing during forest stand development. To clarify conflicting results related to stand developmental stage, we explored how diversity effects on productivity change through time in a diversity experiment spanning 11 years. We show that the strength of diversity effects on productivity progressively increases through time, becoming significantly positive after 9 years. Moreover, we demonstrate that the strengthening of diversity effects is driven primarily by gradual increases in complementarity. We also show that mixing species with contrasting resource-acquisition strategies, and the dominance of deciduous, fast-developing species, promote positive diversity effects on productivity. Our results suggest that the canopy closure and subsequent stem exclusion phase are key for promoting niche complementarity in diverse tree communities.

Mutually inclusive mechanisms of drought-induced tree mortality.

Unprecedented tree dieback across Central Europe caused by recent global change-type drought events highlights the need for a better mechanistic understanding of drought-induced tree mortality. Although numerous physiological risk factors have been identified, the importance of two principal mechanisms, hydraulic failure and carbon starvation, is still debated. It further remains largely unresolved how the local neighborhood composition affects individual mortality risk. We studied 9435 young trees of 12 temperate species planted in a diversity experiment in 2013 to assess how hydraulic traits, carbon dynamics, pest infestation, tree height and neighborhood competition influence individual mortality risk. Following the most extreme global change-type drought since record in 2018, one third of these trees died. Across species, hydraulic safety margins (HSMs) were negatively and a shift towards a higher sugar fraction in the non-structural carbohydrate (NSC) pool positively associated with mortality risk. Moreover, trees infested by bark beetles had a higher mortality risk, and taller trees a lower mortality risk. Most neighborhood interactions were beneficial, although neighborhood effects were highly species-specific. Species that suffered more from drought, especially Larix spp. and Betula spp., tended to increase the survival probability of their neighbors and vice versa. While severe tissue dehydration marks the final stage of drought-induced tree mortality, we show that hydraulic failure is interrelated with a series of other, mutually inclusive processes. These include shifts in NSC pools driven by osmotic adjustment and/or starch depletion as well as pest infestation and are modulated by the size and species identity of a tree and its neighbors. A more holistic view that accounts for multiple causes of drought-induced tree mortality is required to improve predictions of trends in global forest dynamics and to identify mutually beneficial species combinations.

Enhanced light interception and light use efficiency explain overyielding in young tree communities.

Diverse plant communities are often more productive than mono-specific ones. Several possible mechanisms underlie this phenomenon but their relative importance remains unknown. Here we investigated whether light interception alone or in combination with light use efficiency (LUE) of dominant and subordinate species explained greater productivity of mixtures relative to monocultures (i.e. overyielding) in 108 young experimental tree communities. We found mixed-species communities that intercepted more light than their corresponding monocultures had 84% probability of overyielding. Enhanced LUE, which arose via several pathways, also mattered: the probability of overyielding was 71% when, in a mixture, species with higher 'inherent' LUE (i.e. LUE in monoculture) intercepted more light than species with lower LUE; 94% when dominant species increased their LUE in mixture; and 79% when subordinate species increased their LUE. Our results suggest that greater light interception and greater LUE, generated by inter and intraspecific variation, together drive overyielding in mixed-species forests.

Tree characteristics and environmental noise in complex urban settings - A case study from Montreal, Canada.

Field studies have shown that dense tree canopies and regular tree arrangements reduce noise from a point source. In urban areas, noise sources are multiple and tree arrangements are rarely dense. There is a lack of data on the association between the urban tree canopy characteristics and noise in complex urban settings. Our aim was to investigate the spatial variation of urban tree canopy characteristics, indices of vegetation abundance, and environmental noise levels. Using Light Detection and Ranging point cloud data for 2015, we extracted the characteristics of 1,272,069 public and private trees across the island of Montreal, Canada. We distinguished needle-leaf from broadleaf trees, and calculated the percentage of broadleaf trees, the total area of the crown footprint, the mean crown centroid height, and the mean volume of crowns of trees that were located within 100m, 250m, 500m, and 1000m buffers around 87 in situ noise measurement sites. A random forest model incorporating tree characteristics, the normalized difference vegetation index (NDVI) values, and the distances to major urban noise sources (highways, railways and roads) was employed to estimate variation in noise among measurement locations. We found decreasing trends in noise levels with increases in total area of the crown footprint and mean crown centroid height. The percentages of increased mean squared error of the regression models indicated that in 500m buffers the total area of the crown footprint (29.2%) and the mean crown centroid height (12.6%) had a stronger influence than NDVI (3.2%) in modeling noise levels; similar patterns of influence were observed using other buffers. Our findings suggest that municipal initiatives designed to reduce urban noise should account for tree features, and not just the number of trees or the overall amount of vegetation.

Evergreenness influences fine root growth more than tree diversity in a common garden experiment.

Recent studies have reported positive net diversity effects on aboveground tree growth. However, whether similar effects occur belowground through root investment, and whether such effects are related to evergreenness of tree communities, is less clear. Here we studied vertical distribution of standing fine root biomass of twelve North American temperate tree species planted in a common garden tree diversity experiment of varying species richness and evergreenness to test whether belowground niche complementarity of trees could explain positive diversity effects reported aboveground. We tested two alternative hypotheses: trees in mixtures increase uptake of soil resources (1) by increasing vertical root stratification and/or producing a greater fine root density (mg cm-3) or (2) by producing similar or fewer fine roots that are potentially more efficient. Additionally, we hypothesized that proportional allocation to belowground biomass increases with evergreenness of tree communities. Fine roots were sampled in six layers of 5-10 cm, from 0 to 40 cm depth in single-, two- and four-species mixtures. We did not observe an effect of species richness on rooting depth or root density, refuting the hypothesis that aboveground overyielding in tree mixtures is linked to fine root overyielding. Rather, we observed a significant negative diversity effect (- 7.6%) on total fine root density, suggesting overall less investment to fine roots with increasing diversity. The strong positive effect of evergreeness on proportional allocation to fine roots over aboveground parts suggests that deciduous tree roots may be generally more efficient at absorbing soil resources, at least in the early years after tree establishment.

Ecosystem context illuminates conflicting roles of plant diversity in carbon storage.

Plant diversity can increase biomass production in plot-scale studies, but applying these results to ecosystem carbon (C) storage at larger spatial and temporal scales remains problematic. Other ecosystem controls interact with diversity and plant production, and may influence soil pools differently from plant pools. We integrated diversity with the state-factor framework, which identifies key controls, or 'state factors', over ecosystem properties and services such as C storage. We used this framework to assess the effects of diversity, plant traits and state factors (climate, topography, time) on live tree, standing dead, organic horizon and total C in Québec forests. Four patterns emerged: (1) while state factors were usually the most important model predictors, models with both state and biotic factors (mean plant traits and diversity) better predicted C pools; (2) mean plant traits were better predictors than diversity; (3) diversity increased live tree C but reduced organic horizon C; (4) different C pools responded to different traits and diversity metrics. These results suggest that, where ecosystem properties result from multiple processes, no simple relationship may exist with any one organismal factor. Integrating biodiversity into ecosystem ecology and assessing both traits and diversity improves our mechanistic understanding of biotic effects on ecosystems.

The positive carbon stocks-biodiversity relationship in forests: co-occurrence and drivers across five subclimates.

Carbon storage in forests and its ability to offset global greenhouse gas emissions, as well as biodiversity and its capacity to support ecosystem functions and services, are often considered separately in landscape planning. However, the potential synergies between them are currently poorly understood. Identifying the spatial patterns and factors driving their co-occurrence across different climatic zones is critical to more effectively conserve forest ecosystems at the regional level. Here, we integrated information of National Forest Inventories and Breeding Bird Atlases across Europe and North America (Spain and Quebec, respectively), covering five subclimates (steppe, dry Mediterranean, humid Mediterranean, boreal, and temperate). In particular, this study aimed to (1) determine the spatial patterns of both forest carbon stocks and biodiversity (bird richness, tree richness, and overall biodiversity) and the factors that influence them; (2) establish the relationships between forest carbon stocks and biodiversity; and (3) define and characterize the areas of high (hotspots) and low (coldspots) values of carbon and biodiversity, and ultimately quantify their spatial overlap. Our results show that the factors affecting carbon and biodiversity vary between regions and subclimates. The highest values of carbon and biodiversity were found in northern Spain (humid Mediterranean subclimate) and southern Quebec (temperate subclimate) where there was more carbon as climate conditions were less limiting. High density and structural diversity simultaneously favored carbon stocks, tree, and overall biodiversity, especially in isolated and mountainous areas, often associated with steeper slopes and low accessibility. In addition, the relationship between carbon stocks and biodiversity was positive in both regions and all subclimates, being stronger where climate is a limiting factor for forest growth. The spatial overlap between hotspots of carbon and biodiversity provides an excellent opportunity for landscape planning to maintain carbon stocks and conserve biodiversity. The variables positively affecting carbon and biodiversity were also driving the hotspots of both carbon and biodiversity, emphasizing the viability of "win-win" solutions. Our results highlight the need to jointly determine the spatial patterns of ecosystem services and biodiversity for an effective and sustainable planning of forest landscapes that simultaneously support conservation and mitigate climate change.

Biodiversity and ecosystem functioning relations in European forests depend on environmental context.

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.

Tree range expansion in eastern North America fails to keep pace with climate warming at northern range limits.

Rising global temperatures are suggested to be drivers of shifts in tree species ranges. The resulting changes in community composition may negatively impact forest ecosystem function. However, long-term shifts in tree species ranges remain poorly documented. We test for shifts in the northern range limits of 16 temperate tree species in Quebec, Canada, using forest inventory data spanning three decades, 15° of longitude and 7° of latitude. Range shifts were correlated with climate warming and dispersal traits to understand potential mechanisms underlying changes. Shifts were calculated as the change in the 95th percentile of latitudinal occurrence between two inventory periods (1970-1978, 2000-2012) and for two life stages: saplings and adults. We also examined sapling and adult range offsets within each inventory, and changes in the offset through time. Tree species ranges shifted predominantly northward, although species responses varied. As expected shifts were greater for tree saplings, 0.34 km yr-1 , than for adults, 0.13 km yr-1 . Range limits were generally further north for adults compared to saplings, but the difference diminished through time, consistent with patterns observed for range shifts within each life stage. This suggests caution should be exercised when interpreting geographic range offsets between life stages as evidence of range shifts in the absence of temporal data. Species latitudinal velocities were on average <50% of the velocity required to equal the spatial velocity of climate change and were mostly unrelated to dispersal traits. Finally, our results add to the body of evidence suggesting tree species are mostly limited in their capacity to track climate warming, supporting concerns that warming will negatively impact the functioning of forest ecosystems.

Functional identity is the main driver of diversity effects in young tree communities.

Two main effects are proposed to explain biodiversity-ecosystem functioning relationships: niche complementarity and selection effects. Both can be functionally defined using the functional diversity (FD) and functional identity (FI) of the community respectively. Herein, we present results from the first tree diversity experiment that separated the effect of selection from that of complementarity by varying community composition in high-density plots along a gradient of FD, independent of species richness and testing for the effects of FD and community weighted means of traits (a proxy for FI) on stem biomass increment (a proxy for productivity). After 4 years of growth, most mixtures did not differ in productivity from the averages of their respective monocultures, but some did overyield significantly. Those positive diversity effects resulted mostly from selection effects, primarily driven by fast-growing deciduous species and associated traits. Net diversity effect did not increase with time over 4 years.

Ectomycorrhizal fungal diversity and saprotrophic fungal diversity are linked to different tree community attributes in a field-based tree experiment.

Exploring the link between above- and belowground biodiversity has been a major theme of recent ecological research, due in large part to the increasingly well-recognized role that soil microorganisms play in driving plant community processes. In this study, we utilized a field-based tree experiment in Minnesota, USA, to assess the effect of changes in plant species richness and phylogenetic diversity on the richness and composition of both ectomycorrhizal and saprotrophic fungal communities. We found that ectomycorrhizal fungal species richness was significantly positively influenced by increasing plant phylogenetic diversity, while saprotrophic fungal species richness was significantly affected by plant leaf nitrogen content, specific root length and standing biomass. The increasing ectomycorrhizal fungal richness associated with increasing plant phylogenetic diversity was driven by the combined presence of ectomycorrhizal fungal specialists in plots with both gymnosperm and angiosperm hosts. Although the species composition of both the ectomycorrhizal and saprotrophic fungal communities changed significantly in response to changes in plant species composition, the effect was much greater for ectomycorrhizal fungi. In addition, ectomycorrhizal but not saprotrophic fungal species composition was significantly influenced by both plant phylum (angiosperm, gymnosperm, both) and origin (Europe, America, both). The phylum effect was caused by differences in ectomycorrhizal fungal community composition, while the origin effect was attributable to differences in community heterogeneity. Taken together, this study emphasizes that plant-associated effects on soil fungal communities are largely guild-specific and provides a mechanistic basis for the positive link between plant phylogenetic diversity and ectomycorrhizal fungal richness.

A traits-based test of the home-field advantage in mixed-species tree litter decomposition.

BACKGROUND AND AIMS: Litter often decomposes faster in its environment of origin (at 'home') than in a foreign environment ('away'), which has become known as the home-field advantage (HFA). However, many studies have highlighted the conditional nature of the HFA, suggesting that current understanding of this phenomenon is not yet sufficient to generalize across systems. METHODS: The HFA hypothesis was tested for mono-specific and mixed-species litter using a tree-based experiment that manipulated the functional identity and diversity of the host tree community. Litter types of varying quality were transplanted between several host tree communities and decomposition rates were measured using litterbags. Since the decomposer community should respond to traits of the litter input and not their taxonomic identity, a traits-based index of litter-tree similarity was developed. KEY RESULTS: Mono-specific litter exhibited HFA, but when the same litter was decomposed in mixture, this trend was not observed. Mixed-species litter decomposed on average no faster or slower than monoculture litter and exhibited both positive and negative species interactions. These non-additive interactions of decomposition rates in mixture were influenced by the degree of similarity between litter and tree traits. Both synergistic and antagonistic interactions decreased in magnitude with increasing litter-tree similarity such that mixture rates were predictable from monocultures. CONCLUSIONS: The HFA occurred more strongly for mono-specific litter than for the litter types mixed together because interactions between species may have masked this effect. However, when expressed as a function of trait similarity between litters and tree communities, the HFA was not detected.

Advancing biodiversity-ecosystem functioning science using high-density tree-based experiments over functional diversity gradients.

Increasing concern about loss of biodiversity and its effects on ecosystem functioning has triggered a series of manipulative experiments worldwide, which have demonstrated a general trend for ecosystem functioning to increase with diversity. General mechanisms proposed to explain diversity effects include complementary resource use and invoke a key role for species' functional traits. The actual mechanisms by which complementary resource use occurs remain, however, poorly understood, as well as whether they apply to tree-dominated ecosystems. Here we present an experimental approach offering multiple innovative aspects to the field of biodiversity-ecosystem functioning (BEF) research. The International Diversity Experiment Network with Trees (IDENT) allows research to be conducted at several hierarchical levels within individuals, neighborhoods, and communities. The network investigates questions related to intraspecific trait variation, complementarity, and environmental stress. The goal of IDENT is to identify some of the mechanisms through which individuals and species interact to promote coexistence and the complementary use of resources. IDENT includes several implemented and planned sites in North America and Europe, and uses a replicated design of high-density tree plots of fixed species-richness levels varying in functional diversity (FD). The design reduces the space and time needed for trees to interact allowing a thorough set of mixtures varying over different diversity gradients (specific, functional, phylogenetic) and environmental conditions (e.g., water stress) to be tested in the field. The intention of this paper is to share the experience in designing FD-focused BEF experiments with trees, to favor collaborations and expand the network to different conditions.