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.

Factors affecting composition of fatty acids in wild-growing forest mushrooms.

The importance of mushrooms as a food source is continually increasing. To investigate how environmental factors affect the nutritional value of mushrooms, we harvested them in eastern Poland, south-central Germany, and northwestern Belgium in plots with similar environmental conditions but varying in tree species composition and richness. We used gas chromatography-mass spectrometry (GC-MS) to analyze the fatty acid (FA) content of the mushrooms. Fungal species identity explained the largest part (40%) of the total variation in FA concentration and composition. Environmental factors accounted for 1-12% of variation. The concentration of FA, especially saturated fatty acids, decreased with increasing understory cover and increasing nitrogen concentration in the topsoil. The effect of tree species richness or tree species identity was negligible. Our results suggest that the nutritional value of mushrooms depends mainly on the species identity of fungi, but that their FA content is slightly higher in forests with less undergrowth and in nitrogen-poor soils.

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.

Hill-Chao numbers allow decomposing gamma multifunctionality into alpha and beta components.

Biodiversity-ecosystem functioning (BEF) research has provided strong evidence and mechanistic underpinnings to support positive effects of biodiversity on ecosystem functioning, from single to multiple functions. This research has provided knowledge gained mainly at the local alpha scale (i.e. within ecosystems), but the increasing homogenization of landscapes in the Anthropocene has raised the potential that declining biodiversity at the beta (across ecosystems) and gamma scales is likely to also impact ecosystem functioning. Drawing on biodiversity theory, we propose a new statistical framework based on Hill-Chao numbers. The framework allows decomposition of multifunctionality at gamma scales into alpha and beta components, a critical but hitherto missing tool in BEF research; it also allows weighting of individual ecosystem functions. Through the proposed decomposition, new BEF results for beta and gamma scales are discovered. Our novel approach is applicable across ecosystems and connects local- and landscape-scale BEF assessments from experiments to natural settings.

Impact of patch age and size on forest soil characteristics in European agricultural landscapes.

Many landscapes worldwide are characterized by the presence of a mosaic of forest patches with contrasting age and size embedded in a matrix of agricultural land. However, our understanding of the effects of these key forest patch features on the soil nutrient status (in terms of nitrogen, carbon, and phosphorus) and soil pH is still limited due to a lack of large-scale data. To address this research gap, we analyzed 830 soil samples from nearly 200 forest patches varying in age (recent versus ancient forests) and size (small versus larger patches) along a 2500-km latitudinal gradient across Europe. We also considered environmental covariates at multiple scales to increase the generality of our research, including variation in macroclimate, nitrogen deposition rates, forest cover in a buffer zone, basal area and soil type. Multiple linear mixed-effects models were performed to test the combined effects of patch features and environmental covariates on soil nutrients and pH. Recent patches had higher total soil phosphorus concentrations and stocks in the mineral soil layer, along with a lower nitrogen to phosphorus ratio within that layer. Small patches generally had a higher mineral soil pH. Mineral soil nitrogen stocks were lower in forest patches with older age and larger size, as a result of a significant interactive effect. Additionally, environmental covariates had significant effects on soil nutrients, including carbon, nitrogen, phosphorus, and their stoichiometry, depending on the specific covariates. In some cases, the effect of patch age on mineral soil phosphorus stocks was greater than that of environmental covariates. Our findings underpin the important roles of forest patch age and size for the forest soil nutrient status. Long-term studies assessing edge effects and soil development in post-agricultural forests are needed, especially in a context of changing land use and climate.

Carbon-biodiversity relationships in a highly diverse subtropical forest.

Carbon-focused climate mitigation strategies are becoming increasingly important in forests. However, with ongoing biodiversity declines we require better knowledge of how much such strategies account for biodiversity. We particularly lack information across multiple trophic levels and on established forests, where the interplay between carbon stocks, stand age, and tree diversity might influence carbon-biodiversity relationships. Using a large dataset (>4600 heterotrophic species of 23 taxonomic groups) from secondary, subtropical forests, we tested how multitrophic diversity and diversity within trophic groups relate to aboveground, belowground, and total carbon stocks at different levels of tree species richness and stand age. Our study revealed that aboveground carbon, the key component of climate-based management, was largely unrelated to multitrophic diversity. By contrast, total carbon stocks-that is, including belowground carbon-emerged as a significant predictor of multitrophic diversity. Relationships were nonlinear and strongest for lower trophic levels, but nonsignificant for higher trophic level diversity. Tree species richness and stand age moderated these relationships, suggesting long-term regeneration of forests may be particularly effective in reconciling carbon and biodiversity targets. Our findings highlight that biodiversity benefits of climate-oriented management need to be evaluated carefully, and only maximizing aboveground carbon may fail to account for biodiversity conservation requirements.

Resolving the intricate role of climate in litter decomposition.

With approximately 60 Pg of carbon (C) released as CO2 annually, the decomposition of dead organic matter feeds the major terrestrial global CO2 flux to the atmosphere. Macroclimate control over this critical C flux facilitates the parametrization of the C cycle in Earth system models and the understanding of climate change effects on the global C balance. Yet, the long-standing paradigm of climate control was recently challenged by the so far underestimated environmental heterogeneity at local scales, questioning the conceptual framework of thousands of decomposition studies and accuracy of current predictive models. Using three complementary decomposition experiments at a European scale, we showed that macroclimate and litter characteristics largely control plant litter decomposition, reaffirming the role of macroclimate as an integrative decomposition driver through direct environmental control and by influencing co-evolving local plant and decomposer communities. Neglecting this latter indirect effect, commonly used standard litter types overrated micro-environmental control and failed to predict local decomposition of plot-specific litter. Our data help clarify a key question on the regulation of the global C cycle by identifying the relative role of control factors over decomposition and the scales at which they matter and by highlighting sources of confusion in the literature.

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.

The supply of multiple ecosystem services requires biodiversity across spatial scales.

The impact of local biodiversity loss on ecosystem functioning is well established, but the role of larger-scale biodiversity dynamics in the delivery of ecosystem services remains poorly understood. Here we address this gap using a comprehensive dataset describing the supply of 16 cultural, regulating and provisioning ecosystem services in 150 European agricultural grassland plots, and detailed multi-scale data on land use and plant diversity. After controlling for land-use and abiotic factors, we show that both plot-level and surrounding plant diversity play an important role in the supply of cultural and aboveground regulating ecosystem services. In contrast, provisioning and belowground regulating ecosystem services are more strongly driven by field-level management and abiotic factors. Structural equation models revealed that surrounding plant diversity promotes ecosystem services both directly, probably by fostering the spill-over of ecosystem service providers from surrounding areas, and indirectly, by maintaining plot-level diversity. By influencing the ecosystem services that local stakeholders prioritized, biodiversity at different scales was also shown to positively influence a wide range of stakeholder groups. These results provide a comprehensive picture of which ecosystem services rely most strongly on biodiversity, and the respective scales of biodiversity that drive these services. This key information is required for the upscaling of biodiversity-ecosystem service relationships, and the informed management of biodiversity within agricultural landscapes.

Enhancing the structural diversity between forest patches-A concept and real-world experiment to study biodiversity, multifunctionality and forest resilience across spatial scales.

Intensification of land use by humans has led to a homogenization of landscapes and decreasing resilience of ecosystems globally due to a loss of biodiversity, including the majority of forests. Biodiversity-ecosystem functioning (BEF) research has provided compelling evidence for a positive effect of biodiversity on ecosystem functions and services at the local (α-diversity) scale, but we largely lack empirical evidence on how the loss of between-patch ß-diversity affects biodiversity and multifunctionality at the landscape scale (γ-diversity). Here, we present a novel concept and experimental framework for elucidating BEF patterns at α-, ß-, and γ-scales in real landscapes at a forest management-relevant scale. We examine this framework using 22 temperate broadleaf production forests, dominated by Fagus sylvatica. In 11 of these forests, we manipulated the structure between forest patches by increasing variation in canopy cover and deadwood. We hypothesized that an increase in landscape heterogeneity would enhance the ß-diversity of different trophic levels, as well as the ß-functionality of various ecosystem functions. We will develop a new statistical framework for BEF studies extending across scales and incorporating biodiversity measures from taxonomic to functional to phylogenetic diversity using Hill numbers. We will further expand the Hill number concept to multifunctionality allowing the decomposition of γ-multifunctionality into α- and ß-components. Combining this analytic framework with our experimental data will allow us to test how an increase in between patch heterogeneity affects biodiversity and multifunctionality across spatial scales and trophic levels to help inform and improve forest resilience under climate change. Such an integrative concept for biodiversity and functionality, including spatial scales and multiple aspects of diversity and multifunctionality as well as physical and environmental structure in forests, will go far beyond the current widely applied approach in forestry to increase resilience of future forests through the manipulation of tree species composition.

Biodiversity-stability relationships strengthen over time in a long-term grassland experiment.

Numerous studies have demonstrated that biodiversity drives ecosystem functioning, yet how biodiversity loss alters ecosystems functioning and stability in the long-term lacks experimental evidence. We report temporal effects of species richness on community productivity, stability, species asynchrony, and complementarity, and how the relationships among them change over 17 years in a grassland biodiversity experiment. Productivity declined more rapidly in less diverse communities resulting in temporally strengthening positive effects of richness on productivity, complementarity, and stability. In later years asynchrony played a more important role in increasing community stability as the negative effect of richness on population stability diminished. Only during later years did species complementarity relate to species asynchrony. These results show that species complementarity and asynchrony can take more than a decade to develop strong stabilizing effects on ecosystem functioning in diverse plant communities. Thus, the mechanisms stabilizing ecosystem functioning change with community age.

Forest structure and composition alleviate human thermal stress.

Current climate change aggravates human health hazards posed by heat stress. Forests can locally mitigate this by acting as strong thermal buffers, yet potential mediation by forest ecological characteristics remains underexplored. We report over 14 months of hourly microclimate data from 131 forest plots across four European countries and compare these to open-field controls using physiologically equivalent temperature (PET) to reflect human thermal perception. Forests slightly tempered cold extremes, but the strongest buffering occurred under very hot conditions (PET >35°C), where forests reduced strong to extreme heat stress day occurrence by 84.1%. Mature forests cooled the microclimate by 12.1 to 14.5°C PET under, respectively, strong and extreme heat stress conditions. Even young plantations reduced those conditions by 10°C PET. Forest structure strongly modulated the buffering capacity, which was enhanced by increasing stand density, canopy height and canopy closure. Tree species composition had a more modest yet significant influence: that is, strongly shade-casting, small-leaved evergreen species amplified cooling. Tree diversity had little direct influences, though indirect effects through stand structure remain possible. Forests in general, both young and mature, are thus strong thermal stress reducers, but their cooling potential can be even further amplified, given targeted (urban) forest management that considers these new insights.

Drought-exposure history increases complementarity between plant species in response to a subsequent drought.

Growing threats from extreme climatic events and biodiversity loss have raised concerns about their interactive consequences for ecosystem functioning. Evidence suggests biodiversity can buffer ecosystem functioning during such climatic events. However, whether exposure to extreme climatic events will strengthen the biodiversity-dependent buffering effects for future generations remains elusive. We assess such transgenerational effects by exposing experimental grassland communities to eight recurrent summer droughts versus ambient conditions in the field. Seed offspring of 12 species are then subjected to a subsequent drought event in the glasshouse, grown individually, in monocultures or in 2-species mixtures. Comparing productivity between mixtures and monocultures, drought-selected plants show greater between-species complementarity than ambient-selected plants when recovering from the subsequent drought, causing stronger biodiversity effects on productivity and better recovery of drought-selected mixtures after the drought. These findings suggest exposure to recurrent climatic events can improve ecosystem responses to future events through transgenerational reinforcement of species complementarity.

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.

Light heterogeneity affects understory plant species richness in temperate forests supporting the heterogeneity-diversity hypothesis.

One of the most important drivers for the coexistence of plant species is the resource heterogeneity of a certain environment, and several studies in different ecosystems have supported this resource heterogeneity-diversity hypothesis. However, to date, only a few studies have measured heterogeneity of light and soil resources below forest canopies to investigate their influence on understory plant species richness. Here, we aim to determine (1) the influence of forest stand structural complexity on the heterogeneity of light and soil resources below the forest canopy and (2) whether heterogeneity of resources increases understory plant species richness. Measures of stand structural complexity were obtained through inventories and remote sensing techniques in 135 1-ha study plots of temperate forests, established along a gradient of forest structural complexity. We measured light intensity and soil chemical properties on six 25 m² subplots on each of these 135 plots and surveyed understory vegetation. We calculated the coefficient of variation of light and soil parameters to obtain measures of resource heterogeneity and determined understory plant species richness at plot level. Spatial heterogeneity of light and of soil pH increased with higher stand structural complexity, although heterogeneity of soil pH did not increase in conditions of generally high levels of light availability. Increasing light heterogeneity was also associated with increasing understory plant species richness. However, light heterogeneity had no such effects in conditions where soil resource heterogeneity (variation in soil C:N ratios) was low. Our results support the resource heterogeneity-diversity hypothesis for temperate forest understory at the stand scale. Our results also highlight the importance of interaction effects between the heterogeneity of both light and soil resources in determining plant species richness.

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.

Intra-specific leaf trait variability of F. sylvatica, Q. petraea and P. abies in response to inter-specific competition and implications for forest functioning.

Variability in functional traits (FT) is increasingly used to understand the mechanisms behind tree species interactions and ecosystem functioning. In order to explore how FT differ due to interactions between tree species and its influence on stand productivity and other ecological processes, we examined the effects of tree species composition on the intra-specific variability of four widely measured FT: specific leaf area, leaf nitrogen content, leaf angle and stomatal conductance response to vapor pressure deficit. This study focused on three major central European tree species: European beech (Fagus sylvatica L.), Sessile oak (Quercus petraea Liebl.) and Norway spruce (Picea abies [L.] H. Karst.). Each species was examined in monoculture and two-species mixtures in the 13-year-old tree biodiversity experiment BIOTREE-Kaltenborn. Trait distributions and linear mixed models were used to analyze the effect of species mixing, tree size and stand variables on the intra-specific FT variability. A significant effect of branch height on most traits and species indicated a vertical gradient of foliar trait frequently related to light availability. Beech and oak showed a high overall trait variability and sensitivity to species mixing and stand basal area, while the trait variability of spruce was limited. Greater shifts in trait distributions due to mixing were found in specific leaf area for oak and leaf nitrogen content for beech. Thus intra-specific variability of key leaf traits was already influenced at this young development stage by inter-specific interactions. Finally, we used the 3-PG (Physiological Processes Predicting Growth) process-based forest growth model to show that the measured intra-specific variability on single FT values could influence stand productivity, light absorption and transpiration, although the net effect depends on the considered trait and the species composition of the mixture. The results of this study will aid better understanding of the effects of inter-specific competition on intra-specific FT variability, which has implications for the parameterization of process-based forest growth models and our understanding of ecosystem functioning.

Climatic conditions, not above- and belowground resource availability and uptake capacity, mediate tree diversity effects on productivity and stability.

Tree species diversity promotes multiple ecosystem functions and services. However, little is known about how above- and belowground resource availability (light, nutrients, and water) and resource uptake capacity mediate tree species diversity effects on aboveground wood productivity and temporal stability of productivity in European forests and whether the effects differ between humid and arid regions. We used the data from six major European forest types along a latitudinal gradient to address those two questions. We found that neither leaf area index (a proxy for light uptake capacity), nor fine root biomass (a proxy for soil nutrient and water uptake capacity) was related to tree species richness. Leaf area index did, however, enhance productivity, but negatively affected stability. Productivity was further promoted by soil nutrient availability, while stability was enhanced by fine root biomass. We only found a positive effect of tree species richness on productivity in arid regions and a positive effect on stability in humid regions. This indicates a possible disconnection between productivity and stability regarding tree species richness effects. In other words, the mechanisms that drive the positive effects of tree species richness on productivity do not per se benefit stability simultaneously. Our findings therefore suggest that tree species richness effects are largely mediated by differences in climatic conditions rather than by differences in above- and belowground resource availability and uptake capacity at the regional scales.