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.

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.

Effects of plant diversity on productivity strengthen over time due to trait-dependent shifts in species overyielding.

Plant diversity effects on community productivity often increase over time. Whether the strengthening of diversity effects is caused by temporal shifts in species-level overyielding (i.e., higher species-level productivity in diverse communities compared with monocultures) remains unclear. Here, using data from 65 grassland and forest biodiversity experiments, we show that the temporal strength of diversity effects at the community scale is underpinned by temporal changes in the species that yield. These temporal trends of species-level overyielding are shaped by plant ecological strategies, which can be quantitatively delimited by functional traits. In grasslands, the temporal strengthening of biodiversity effects on community productivity was associated with increasing biomass overyielding of resource-conservative species increasing over time, and with overyielding of species characterized by fast resource acquisition either decreasing or increasing. In forests, temporal trends in species overyielding differ when considering above- versus belowground resource acquisition strategies. Overyielding in stem growth decreased for species with high light capture capacity but increased for those with high soil resource acquisition capacity. Our results imply that a diversity of species with different, and potentially complementary, ecological strategies is beneficial for maintaining community productivity over time in both grassland and forest ecosystems.

Young temperate tree species show different fine root acclimation capacity to growing season water availability.

Background and aims: Changes in water availability during the growing season are becoming more frequent due to climate change. Our study aimed to compare the fine-root acclimation capacity (plasticity) of six temperate tree species aged six years and exposed to high or low growing season soil water availability over five years. Methods: Root samples were collected from the five upper strata of mineral soil to a total soil depth of 30 cm in monoculture plots of Acer saccharum Marsh., Betula papyrifera Marsh., Larix laricina K. Koch, Pinus strobus L., Picea glauca (Moench) Voss and Quercus rubra L. established at the International Diversity Experiment Network with Trees (IDENT) field experiment in Sault Ste. Marie, Ontario, Canada. Four replicates of each monoculture were subjected to high or low water availability treatments. Results: Absorptive fine root density increased by 67% for Larix laricina, and 90% for Picea glauca, under the high-water availability treatment at 0-5 cm soil depth. The two late successional, slower growing tree species, Acer saccharum and Picea glauca, showed higher plasticity in absorptive fine root biomass in the upper 5 cm of soil (PIv = 0.36 & 0.54 respectively), and lower plasticity in fine root depth over the entire 30 cm soil profile compared to the early successional, faster growing tree species Betula papyrifera and Larix laricina. Conclusion: Temperate tree species show contrasting acclimation responses in absorptive fine root biomass and rooting depth to differences in water availability. Some of these responses vary with tree species successional status and seem to benefit both early and late successional tree species. Supplementary Information: The online version contains supplementary material available at 10.1007/s11104-023-06377-w.

It takes a village to grow a tree: Most tree species benefit from dissimilar neighbors.

Scientific consensus is that diverse tree species positively impact forest productivity, especially when species are functionally dissimilar. Under the complementarity hypothesis, differences in species traits reduce competition among neighboring tree species. However, while this relationship has been extensively studied at the community level, there is a lack of understanding regarding how individuals of different species specifically respond to a functionally dissimilar neighborhood. In this study, we used permanent plots from Quebec, Canada, and 19 focal tree species to test whether: (1) tree growth response to neighborhood dissimilarity varies with their identity and competition intensity, and (2) focal tree species' traits explain their response to neighborhood dissimilarity. We demonstrate that: tree growth is primarily influenced by competition, species identity, and their interactions, but that dissimilarity, alone and in interaction with the main drivers of tree growth, explains an additional 1.8% of the variation in species growth. Within this context, (1) most species' respond positively to neighborhood dissimilarity, with magnitude being species and competition dependent, and (2) focal tree traits partly explain these dependencies, with shade-intolerant species benefiting most from dissimilar neighbors under high competition. Our study provides empirical support for the complementarity hypothesis, emphasizing the small but consistent positive effect of functional dissimilarity on tree growth in local neighborhoods. Our findings identify the species with the highest potential of benefiting from dissimilar neighbors but also demonstrate that the positive effect of neighborhood dissimilarity is not limited to a select few species with specific traits; rather, it is observed across a diverse range of species. The cumulative growth responses of individuals to functionally dissimilar neighbors may help explain the commonly observed higher productivity in more diverse communities.

Exotic tree species have consistently lower herbivore load in a cross-Atlantic tree biodiversity experiment.

It is commonly expected that exotic plants experience reduced herbivory, but experimental evidence for such enemy release is still controversial. One reason for conflicting results might be that community context has rarely been accounted for, although the surrounding plant diversity may moderate enemy release. Here, we tested the effects of focal tree origin and surrounding tree diversity on herbivore abundance and leaf damage in a cross-Atlantic tree-diversity experiment in Canada and Germany. We evaluated six European tree species paired with six North American congeners in both their native and exotic range, expecting lower herbivory for the exotic tree species in each pair at each site. Such reciprocal experiments have long been called for, but have not been realized thus far. In addition to a thorough evaluation of overall enemy release effects, we tested whether enemy release effects changed with the surrounding tree diversity. Herbivore abundance was indeed consistently lower on exotics across all six tree genera (12 comparisons). This effect of exotic status was independent of the continent, phylogenetic relatedness, and surrounding tree diversity. In contrast, leaf damage associated with generalist leaf chewers was consistently higher on North American tree species. Interestingly, several species of European weevils were the most abundant leaf chewers on both continents and the dominant herbivores at the Canadian site. Thus, most observed leaf damage is likely to reflect the effect of generalist herbivores that feed heavily on plant species with which they have not evolved. At the German site, sap suckers were the dominant herbivores and showed a pattern consistent with enemy release. Taken together, the consistently lower herbivory on exotics on both continents is not purely a pattern of enemy release in the strictest sense, but to some degree additionally reflects the susceptibility of native plants to invasive herbivores. In conclusion, our cross-Atlantic study is consistent with the idea that nonnative trees have generally reduced herbivory, regardless of tree community diversity and species identity, but for different reasons depending on the dominant herbivore guild.

Disentangling drivers of litter decomposition in a multi-continent network of tree diversity experiments.

Litter decomposition is a key ecosystem function in forests and varies in response to a range of climatic, edaphic, and local stand characteristics. Disentangling the relative contribution of these factors is challenging, especially along large environmental gradients. In particular, knowledge of the effect of management options, such as tree planting density and species composition, on litter decomposition would be highly valuable in forestry. In this study, we made use of 15 tree diversity experiments spread over eight countries and three continents within the global TreeDivNet network. We evaluated the effects of overstory composition (tree identity, species/mixture composition and species richness), plantation conditions (density and age), and climate (temperature and precipitation) on mass loss (after 3 months and 1 year) of two standardized litters: high-quality green tea and low-quality rooibos tea. Across continents, we found that early-stage decomposition of the low-quality rooibos tea was influenced locally by overstory tree identity. Mass loss of rooibos litter was higher under young gymnosperm overstories compared to angiosperm overstories, but this trend reversed with age of the experiment. Tree species richness did not influence decomposition and explained almost no variation in our multi-continent dataset. Hence, in the young plantations of our study, overstory composition effects on decomposition were mainly driven by tree species identity on decomposer communities and forest microclimates. After 12 months of incubation, mass loss of the high-quality green tea litter was mainly influenced by temperature whereas the low-quality rooibos tea litter decomposition showed stronger relationships with overstory composition and stand age. Our findings highlight that decomposition dynamics are not only affected by climate but also by management options, via litter quality of the identity of planted trees but also by overstory composition and structure.

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.

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.

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.

Pathways for cross-boundary effects of biodiversity on ecosystem functioning.

The biodiversity-ecosystem functioning concept asserts that processes in ecosystems are markedly influenced by species richness and other facets of biodiversity. However, biodiversity-ecosystem functioning studies have been largely restricted to single ecosystems, ignoring the importance of functional links - such as the exchange of matter, energy, and organisms - between coupled ecosystems. Here we present a basic concept and outline three pathways of cross-boundary biodiversity effects on ecosystem processes and propose an agenda to assess such effects, focusing on terrestrial-aquatic linkages to illustrate the case. This cross-boundary perspective of biodiversity-ecosystem functioning relationships presents a promising frontier for biodiversity and ecosystem science with repercussions for the conservation, restoration, and management of biodiversity and ecosystems from local to landscape scales.

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.

Strong variations in urban allergenicity riskscapes due to poor knowledge of tree pollen allergenic potential.

Exposure to allergenic tree pollen is an increasing environmental health issue in urban areas. However, reliable, well-documented, peer-reviewed data on the allergenicity of pollen from common tree species in urban environments are lacking. Using the concept of 'riskscape', we present and discuss evidence on how different tree pollen allergenicity datasets shape the risk for pollen-allergy sufferers in five cities with different urban forests and population densities: Barcelona, Montreal, New York City, Paris, and Vancouver. We also evaluate how tree diversity can modify the allergenic risk of urban forests. We show that estimates of pollen exposure risk range from 1 to 74% for trees considered to be highly allergenic in the same city. This variation results from differences in the pollen allergenicity datasets, which become more pronounced when a city's canopy is dominated by only a few species and genera. In an increasingly urbanized world, diverse urban forests offer a potentially safer strategy aimed at diluting sources of allergenic pollen until better allergenicity data is developed. Our findings highlight an urgent need for a science-based approach to guide public health and urban forest planning.

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.

Climate affects neighbour-induced changes in leaf chemical defences and tree diversity-herbivory relationships.

Associational resistance theory predicts that insect herbivory decreases with increasing tree diversity in forest ecosystems. However, the generality of this effect and its underlying mechanisms are still debated, particularly since evidence has accumulated that climate may influence the direction and strength of the relationship between diversity and herbivory.We quantified insect leaf herbivory and leaf chemical defences (phenolic compounds) of silver birch Betula pendula in pure and mixed plots with different tree species composition across 12 tree diversity experiments in different climates. We investigated whether the effects of neighbouring tree species diversity on insect herbivory in birch, that is, associational effects, were dependent on the climatic context, and whether neighbour-induced changes in birch chemical defences were involved in associational resistance to insect herbivory.We showed that herbivory on birch decreased with tree species richness (i.e. associational resistance) in colder environments but that this relationship faded as mean annual temperature increased.Birch leaf chemical defences increased with tree species richness but decreased with the phylogenetic distinctiveness of birch from its neighbours, particularly in warmer and more humid environments.Herbivory was negatively correlated with leaf chemical defences, particularly when birch was associated with closely related species. The interactive effect of tree diversity and climate on herbivory was partially mediated by changes in leaf chemical defences.Our findings confirm that tree species diversity can modify the leaf chemistry of a focal species, hence its quality for herbivores. They further stress that such neighbour-induced changes are dependent on climate and that tree diversity effects on insect herbivory are partially mediated by these neighbour-induced changes in chemical defences.

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.

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.

Complex Above- and Below-Ground Growth Responses of Two Urban Tree Species Following Root, Stem, and Foliage Damage-An Experimental Approach.

Urban trees are subjected to numerous biotic and mechanical damages, which can affect their growth rates and health. However, for most species, a systematic analysis of tree above- and below-ground growth reactions to a variety of damages is still lacking. Under a fully factorial experimental setup, using two common urban trees (Celtis occidentalis, Fraxinus pennsylvanica), we tested the effects of various degrees of frequently occurring damage as defoliation, root reduction, and stem injuries for a total of 18 treatments. We hypothesized that (i) an increasing amount of damage would proportionally negatively affect both root and stem growth; (ii) there would be a lag or lasting effect on growth; and (iii) both species would react similarly to the treatments. Contrary to our expectation, increasing levels of single or combined damage did not have an incremental effect on either stem or root growth. Although Celtis was significantly less vigorous than Fraxinus, it did not react strongly to damage treatments compared to the control. Interestingly, Celtis that experienced stem damage alone or in combination with other damages showed higher growth rates than the control. For Celtis, root injury was the treatment having the most impact, decreasing both root and stem growth consistently throughout the 5 years following treatments, whereas defoliation decreased growth only in the first 2 years. All damage treatments negatively affected stem and root growth of Fraxinus trees. Stem growth was affected the most by defoliation in the first year following the treatment, while root injury became the driving factor in subsequent years. For both species, stem injury showed the least influence on growth rates. The control and low-level damage treatments often affected growth rates in a similar way, suggesting that low-intensity stress triggers compensatory reactions stimulating photosynthetic rates and nutrient utilization. The slower-growing tree species, Celtis, showed a less negative reaction to all damage treatments compared to Fraxinus. This study illustrates that various types of above- and below-ground injuries do not have a simple additive effect on tree growth and that trees are capable of compensating for the loss of foliage, roots, or phloem to meet their metabolic demand.