Search

The soil microbiome is crucial for regulating biogeochemical processes and can, thus, strongly influence tree health, especially under stress conditions. However, little is known about the effect of prolonged water deficit on soil microbial communities during the development of saplings. We assessed the response of prokaryotic and fungal communities to different levels of experimental water limitation in mesocosms with Scots pine saplings. We combined analyses of physicochemical soil properties and tree growth with DNA metabarcoding of soil microbial communities throughout four seasons. Seasonal changes in soil temperature and soil water content and a decreasing soil pH strongly influenced the composition of microbial communities but not their total abundance. Contrasting levels of soil water contents gradually altered the soil microbial community structure over the four seasons. Results indicated that prokaryotic communities were less resistant to water limitation than fungal communities. Water limitation promoted the proliferation of desiccation tolerant, oligotrophic taxa. Moreover, water limitation and an associated increase in soil C/N ratio induced a shift in the potential lifestyle of taxa from symbiotic to saprotrophic. Overall, water limitation appeared to alter soil microbial communities involved in nutrient cycling, pointing to potential consequences for forest health affected by prolonged episodes of drought., FEMS Microbiology Ecology, 99 (6), ISSN:0168-6496, ISSN:1574-6941

Background Forests perform various important ecosystem functions that contribute to ecosystem services. In many parts of the world, forest management has shifted from a focus on timber production to multi-purpose forestry, combining timber production with the supply of other forest ecosystem services. However, it is unclear which forest types provide which ecosystem services and to what extent forests primarily managed for timber already supply multiple ecosystem services. Based on a comprehensive dataset collected across 150 forest plots in three regions differing in management intensity and species composition, we develop models to predict the potential supply of 13 ecosystem services. We use those models to assess the level of multifunctionality of managed forests at the national level using national forest inventory data. Results Looking at the potential supply of ecosystem services, we found trade-offs (e.g. between both bark beetle control or dung decomposition and both productivity or soil carbon stocks) as well as synergies (e.g. for temperature regulation, carbon storage and culturally interesting plants) across the 53 most dominant forest types in Germany. No single forest type provided all ecosystem services equally. Some ecosystem services showed comparable levels across forest types (e.g. decomposition or richness of saprotrophs), while others varied strongly, depending on forest structural attributes (e.g. phosphorous availability or cover of edible plants) or tree species composition (e.g. potential nitrification activity). Variability in potential supply of ecosystem services was only to a lesser extent driven by environmental conditions. However, the geographic variation in ecosystem function supply across Germany was closely linked with the distribution of main tree species. Conclusions Our results show that forest multifunctionality is limited to subsets of ecosystem services. The importance of tree species composition highlights that a lack of multifunctionality at the stand level can be compensated by managing forests at the landscape level, when stands of complementary forest types are combined. These results imply that multi-purpose forestry should be based on a variety of forest types requiring coordinated planning across larger spatial scales.

Rising temperatures have the potential to directly affect carbon cycling in peatlands by enhancing organic matter (OM) decomposition, contributing to the release of CO2 and CH4 to the atmosphere. In turn, increasing atmospheric CO2 concentration may stimulate photosynthesis, potentially increasing plant litter inputs belowground and transferring carbon from the atmosphere into terrestrial ecosystems. Key questions remain about the magnitude and rate of these interacting and opposing environmental change drivers. Here, we assess the incorporation and degradation of plant- and microbe-derived OM in an ombrotrophic peatland after 4 years of whole-ecosystem warming (+0, +2.25, +4.5, +6.75 and +9 degrees C) and two years of elevated CO2 manipulation (500 ppm above ambient). We show that OM molecular composition was substantially altered in the aerobic acrotelm, highlighting the sensitivity of acrotelm carbon to rising temperatures and atmospheric CO2 concentration. While warming accelerated OM decomposition under ambient CO2, new carbon incorporation into peat increased in warming x elevated CO2 treatments for both plant- and microbe-derived OM. Using the isotopic signature of the applied CO2 enrichment as a label for recently photosynthesized OM, our data demonstrate that new plant inputs have been rapidly incorporated into peat carbon. Our results suggest that under current hydrological conditions, rising temperatures and atmospheric CO2 levels will likely offset each other in boreal peatlands., Global Change Biology, 28 (3), ISSN:1354-1013, ISSN:1365-2486

Rising temperatures have the potential to directly affect carbon cycling in peatlands by enhancing organic matter (OM) decomposition, contributing to the release of CO

Evidence from experimental and established grasslands indicates that plant biodiversity can modify the water cycle. One suspected mechanism behind this is a higher infiltration capacity (νB) and hydraulic conductivity (K) of the soil on species‐rich grasslands. However, in established and agriculturally managed grasslands, biodiversity effects cannot be studied independent of land‐use effects. Therefore, we investigated in established grassland systems how land‐use intensity and associated biodiversity of plants and soil animals affect νB and K at and close to saturation. On 50 grassland plots along a land‐use intensity gradient in the Biodiversity Exploratory Schwäbische Alb, Germany, we measured νB with a hood infiltrometer at several matrix potentials and calculated the saturated and unsaturated K. We statistically analysed the relationship between νB or K and land‐use information (e.g., fertilising intensity), abiotic (e.g., soil texture) and biotic data (e.g., plant species richness, earthworm abundance). Land‐use intensity decreased and plant species richness increased νB and K, while the direction of the effects of soil animals was inconsistent. The effect of land‐use intensity on νB and K was mainly attributable to its negative effect on plant species richness. Our results demonstrate that plant species richness was a better predictor of νB and K at and close to saturation than land‐use intensity or soil physical properties in the established grassland systems of the Schwäbische Alb., Deutsche Forschungsgemeinschaft http://dx.doi.org/10.13039/501100001659, Stiftung Landesbank Baden‐Württemberg, Swiss National Science Foundation http://dx.doi.org/10.13039/501100001711, http://doi.org/10.17616/R32P9Q

Acidobacteria occur in a large variety of ecosystems worldwide and are particularly abundant and highly diverse in soils. In spite of their diversity, only few species have been characterized to date which makes

In forest ecosystems, microorganisms hold key functions as nutrient cyclers, decomposers, plant symbionts or pathogens and thereby regulate biogeochemical processes and forest health. These microbial dynamics are controlled by water availability in three fundamental ways: as resource, as solvent, and as transport medium. For one of the dominant tree species in Swiss forests - Scots pine (Pinus sylvestris L.) - high mortality rates have been observed in recent decades. In the Rhone valley of Switzerland, forest dieback appears to be primarily caused by direct effects of drought and an increasing susceptibility of trees to further constraints, such as pathogen attacks. Nonetheless, water limitation does not affect soil microbes and trees separately but rather induces a series of interconnected effects between trees and the associated soil microbiome, which could strongly alter carbon and nutrient cycling in forests. We conduct a study to investigate the effects of drought on the biological interplay between Scots pine trees and soil microbial communities. We aim to estimate how shifts in microbial community composition and functional capacity under drought may affect nutrient cycling and tree vitality potentially contributing to tree mortality. In order to understand these mechanisms, we perform greenhouse experiments with tree-soil mesocosms under controlled conditions. State-of-the art molecular methods such as metabarcoding of ribosomal markers, shotgun metagenome sequencing, and qPCR of key functional genes are used to unravel alterations in the soil microbiome and in the underlying functional metabolic potential related to drought and associated tree-mortality. Furthermore, to elucidate the impact of drought on microbial carbon dynamics, stable isotope labelling techniques have been applied to trace 13C labeled plant photosynthates into the soil microbial communities by analyzing 13C signatures of phospholipid fatty acids. Investigation of soil physicochemical properties and tree-vitality is done in parallel with the microbial assessments to understand the feedbacks on nutrient-cycling and the soil-tree continuum. The overarching aim of this study is to gain new insights into the complex relationships between soil, trees and microbes under drought.

The microbial community composition in subsoils remains understudied and it is largely unknown whether subsoil microorganisms show a similar response to global warming as do microorganisms at the soil surface. Since microorganisms are key drivers of soil organic carbon decomposition, this knowledge gap causes uncertainty in predictions of future carbon cycling in the subsoil carbon pool (>50 % of the soil organic carbon stocks are below 30 cm soil depth). In the Blodgett forest field warming experiment (California, USA) we investigated how +4 °C warming the whole soil profile to 100 cm soil depth for 4.5 years has affected the abundance and community structure of microorganisms. We used proxies for bulk microbial biomass carbon (MBC) and functional microbial groups based on lipid biomarkers, such as phospholipid fatty acids (PLFAs) and branched glycerol dialkyl glycerol tetraethers (brGDGTs). Microbial biomass decreased and community composition changed with depth. Our results show that the concentration of PLFAs decreased with warming in the subsoil (below 30 cm) by 28 % but was not affected in the topsoil. Phospholipid fatty acid concentrations changed in concert with soil organic carbon. The microbial community response to warming was depth dependent. The relative abundance of actinobacteria increased in subsoil, and gram+ bacteria in subsoils adapted their cell-membrane structure to warming induced stress as indicated by the ratio of anteiso to iso PLFAs. Our results show for the first time that subsoil microorganisms can be more affected by warming as compared to topsoil microorganisms. These microbial responses could be explained by the observed decrease in subsoil organic carbon concentration in the warmed plots. A decrease in microbial abundance in warmed subsoils might reduce the magnitude of the respiration response over time. The shift in the subsoil microbial community towards more actinobacteria might disproportionately enhance degradation of previously stable subsoil carbon, as this group is able to metabolize complex carbon sources.

Soils will warm in near synchrony with the air over the whole profiles following global climate change. It is largely unknown how subsoil (below 30 cm) microbial communities will respond to this warming and how plant-derived soil organic carbon (SOC) will be affected. Predictions how climate change will affect the large subsoil carbon pool (>50 % of SOC is below 30 cm soil depth) remain uncertain. At Blodgett forest (California, USA) a field warming experiment was set up in 2013 warming whole soil profiles to 100 cm soil depth by +4°C compared to control plots. We took samples in 2018, after 4.5 years of continuous warming and investigated how warming has affected the abundance and community structure of microoganisms (using phospholipid fatty acids, PLFAs), and plant litter (using cutin and suberin). The warmed subsoil (below 30 cm) contained significantly less microbial biomass (28%) compared to control plots, whereas the topsoil remained unchanged. Additionally below 50 cm, the microbial community was different in warmed as compared to control plots. Actinobacteria were relatively more abundant and Gram+ bacteria adapted their cell-membrane structure to warming. The decrease in microbial abundance might be related to lower SOC concentrations in warmed compared to control subsoils. In contrast to smaller SOC concentrations and less fine root mass in the warmed plots, the concentrations of the plant polymers suberin and cutin did not change. Overall our results demonstrate that already four seasons of simulated whole-soil warming caused distinct depth-specific responses of soil biogeochemistry: warming altered the subsoil microbial community, but not concentrations of plant-derived soil organic carbon., EGUsphere

Worldwide tree species have been observed to be suffering from extended periods of water limitation, for example due to warmer climate that increases soil evaporation and plant transpiration. These conditions likely do not only affect the growth and vitality of trees but may also feed back on the cycling of carbon and nitrogen at the interface between roots and soils. In September 2019, we established a mesocosm experiment to mechanistically study on a seasonal basis how the interactions between plants and soil biotic and abiotic resources are altered during events of drought. The mesocosms feature young Scots pine (Pinus sylvestris L.) trees and soil collected from a drought-affected natural forest in the Rhone valley, Switzerland; and are treated with three different levels of water availability (control, sufficient water; intermediate drought, 40% reduction; severe drought, 75% reduction). One year after the start of the experiment an isotopic labelling campaign with 13CO2 was conducted to trace the natural pathway of photosynthetic assimilates into above- and belowground carbon pools and fluxes. During the first growing season of the experiment, severe drought more than doubled the growth of fine roots when compared to the control treatment. In turn, the mean diameter of the fine roots significantly decreased by 22%, and fewer ectomycorrhizal root tips were observed. These findings suggest that trees exposed to drought invest more in within-plant carbon maintenance and in the growth of root systems, rather than in the allocation of carbon to sustain the biology in the rhizosphere for nutrient acquisition. Moreover, post-label soil pore 13CO2 concentrations and total soil CO2 concentrations were lower under severe drought compared to intermediate and control treatments, indicating a generally reduced carbon metabolism. By tracking the fate of 13C assimilates into fine roots, soils and microbial communities over time we now investigate whether there is a threshold at which Scots pine trees stop investing in providing carbon to the rhizosphere and rather succumb to drought., EGUsphere

Increasing global temperatures have the potential to stimulate decomposition and alter the composition of soil organic matter (SOM). However, questions remain about the extent to which SOM quality and quantity along the soil profile may change under future warming. In this study we assessed how +4 °C whole-soil warming affected the quantity and quality of SOM down to 90 cm depth in a mixed-coniferous temperate forest using biomarker analyses. Our findings indicate that 4.5 years of soil warming led to divergent responses in subsoils (>20 cm) as compared to surface soils. Warming enhanced the accumulation of plant-derived n-alkanes over the whole soil profile. In the subsoil, this was at the expense of plant- and microorganism-derived fatty acids, and the relative abundance of SOM molecular components shifted from less microbially transformed to more transformed organic matter. Fine root mass declined by 24.0 ± 7.5% with warming over the whole soil profile, accompanied by reduced plant-derived inputs and accelerated decomposition of aromatic compounds and plant-derived fatty acids in the subsoils. Our study suggests that warming accelerated microbial decomposition of plant-derived inputs, leaving behind more degraded organic matter. The non-uniform, and depth dependent SOM composition and warming response implies that subsoil carbon cycling is as sensitive and complex as in surface soils., Soil Biology and Biochemistry, 156, ISSN:0038-0717, ISSN:1879-3428

In this case study analysis, we identified fungal traits that were associated with the responses of taxa to 4 global change factors: elevated CO2, warming and drying, increased precipitation, and nitrogen (N) enrichment. We developed a trait-based framework predicting that as global change increases limitation of a given nutrient, fungal taxa with traits that target that nutrient will represent a larger proportion of the community (and vice versa). In addition, we expected that warming and drying and N enrichment would generate environmental stress for fungi and may select for stress tolerance traits. We tested the framework by analyzing fungal community data from previously published field manipulations and linking taxa to functional gene traits from the MycoCosm Fungal Portal. Altogether, fungal genera tended to respond similarly to 3 elements of global change: increased precipitation, N enrichment, and warming and drying. The genera that proliferated under these changes also tended to possess functional genes for stress tolerance, which suggests that these global changes—even increases in precipitation—could have caused environmental stress that selected for certain taxa. In addition, these genera did not exhibit a strong capacity for C breakdown or P acquisition, so soil C turnover may slow down or remain unchanged following shifts in fungal community composition under global change. Since we did not find strong evidence that changes in nutrient limitation select for taxa with traits that target the more limiting nutrient, we revised our trait-based framework. The new framework sorts fungal taxa into Stress Tolerating versus C and P Targeting groups, with the global change elements of increased precipitation, warming and drying, and N enrichment selecting for the stress tolerators.

Land-use intensification can increase provisioning ecosystem services, such as food and timber production, but it also drives changes in ecosystem functioning and biodiversity loss, which may ultimately compromise human wellbeing. To understand how changes in land-use intensity affect the relationships between biodiversity, ecosystem functions, and services, we built networks from correlations between the species richness of 16 trophic groups, 10 ecosystem functions, and 15 ecosystem services. We evaluated how the properties of these networks varied across land-use intensity gradients for 150 forests and 150 grasslands. Land-use intensity significantly affected network structure in both habitats. Changes in connectance were larger in forests, while changes in modularity and evenness were more evident in grasslands. Our results show that increasing land-use intensity leads to more homogeneous networks with less integration within modules in both habitats, driven by the belowground compartment in grasslands, while forest responses to land management were more complex. Land-use intensity strongly altered hub identity and module composition in both habitats, showing that the positive correlations of provisioning services with biodiversity and ecosystem functions found at low land-use intensity levels, decline at higher intensity levels. Our approach provides a comprehensive view of the relationships between multiple components of biodiversity, ecosystem functions, and ecosystem services and how they respond to land use. This can be used to identify overall changes in the ecosystem, to derive mechanistic hypotheses, and it can be readily applied to further global change drivers. The work has been supported by the DFG Priority Program 1374 “Infrastructure-Biodiversity-Exploratories”. S.S. was supported by the Spanish Government under Ramón y Cajal Contract RYC-2016-20604.

Water limiting conditions for the growth and physiology of trees as well as episodes of tree mortality triggered by drought have recently been documented in several bioregions across the world. In parallel to these major vegetation alterations, the impact of water scarcity also has prominent effects on soil processes mediated by the microbiome such as the transformation of organic matter, heterotrophic respiration, microbial uptake as well as nutrient mineralization. Although currently little explored, shifts in the interplay occurring between tree functioning and soil microbial processes may be crucial during tree mortality events and may feed back on ecosystem carbon and nitrogen cycling. We will present a multidisciplinary setup to mechanistically explore how water limitation acts synergistically on the interplay between trees and soil microorganisms, with potential consequences for ecosystem biogeochemical fluxes. The experimental setup focusses on a key temperate forest species, Scots pine (Pinus sylvestris L.), which is currently facing high mortality rates in several inner-Alpine valleys of Europe due to drier climatic conditions during parts of the year. We make use of small scale mesocosms featuring young trees and soil collected from a drought-affected natural forest. The mesocosms are treated with different levels of water availability under controlled conditions. Plant growth and physiological changes related to water limitation are investigated in parallel to various soil properties. State-of-the-art isotopic labelling techniques are used to trace alterations in carbon and nitrogen transfers within the plant-soil-microbe continuum. We will specifically test whether extended periods of drought suppress the flux of carbon from plants to soil and lead plants to invest more in the maintenance of fine root systems. Moreover, we will follow the potential changes in the rates of decomposition, mineralization and incorporation of plant debris into soil organic matter over time and link them to potential alterations of the soil microbiota. These experimental observations will be validated by measurements in drought-affected Scots pine forests in inner-Alpine valleys. We expect the outcomes of this work to advance the fundamental understanding of the alterations occurring in the plant-soil-microbe system related to drought as well as to improve the detection of mechanisms leading to Scots pine mortality., EGUsphere

While forest management strongly influences biodiversity, it remains unclear how the structural and compositional changes caused by management affect different community dimensions (e.g. richness, specialisation, abundance or completeness) and how this differs between taxa. We assessed the effects of nine forest features (representing stand structure, heterogeneity and tree composition) on thirteen above‐ and belowground trophic groups of plants, animals, fungi and bacteria in 150 temperate forest plots differing in their management type. Canopy cover decreased light resources, which increased community specialisation but reduced overall diversity and abundance. Features increasing resource types and diversifying microhabitats (admixing of oaks and conifers) were important and mostly affected richness. Belowground groups responded differently to those aboveground and had weaker responses to most forest features. Our results show that we need to consider forest features rather than broad management types and highlight the importance of considering several groups and community dimensions to better inform conservation. The work was partly funded by the DFG Priority Program 1374 ‘Infrastructure-Biodiversity-Exploratories’ (DFG-Refno. Po362/18-3). SSo was supported by the Spanish Government under a Ramón y Cajal contract (RYC-2016-20604).

1.Rapid growth of the world's human population has increased pressure on landscapes to deliver high levels of multiple ecosystem services, including food and fibre production, carbon storage, biodiversity conservation and recreation. However, we currently lack general principles describing how to achieve this landscape multifunctionality. 2.We combine theoretical simulations and empirical data on 14 ecosystem services measured across 150 grasslands in three German regions. In doing so, we investigate the circumstances under which spatial heterogeneity in a driver of ecosystem functioning (an ‘ecosystem‐driver,’ e.g. the presence of keystone species, land‐use intensification or habitat types) increases landscape‐level ecosystem multifunctionality. 3.Simulations based on theoretical data demonstrated that relationships between heterogeneity and landscape multifunctionality are highly variable and can range from non‐significant to strongly positive. Despite this variability, we could identify criteria under which heterogeneity‐landscape multifunctionality relationships were most strongly positive: this happened when multiple ecosystem services responded contrastingly (both positively and negatively) to an ecosystem‐driver. 4.These findings were confirmed using empirical data, which showed that heterogeneity in land‐use intensity promoted landscape multifunctionality in cases where functions with both positive (e.g. plant biomass) and negative (e.g. flower cover) responses to land use intensification were included. For example, the simultaneous provisioning of ecosystem functions related to forage production (generally profiting from land‐use intensification), biodiversity conservation and recreation (generally decreasing with land‐use intensification) was highest in landscapes consisting of sites varying in land‐use intensity. 5.Synthesis and applications. Our findings show that there are general principles governing landscape multifunctionality. A knowledge of these principles may support land management decisions. For example, knowledge of relationships between ecosystem services and their drivers, such as land use type, can help estimate the consequences of increasing or decreasing heterogeneity for landscape‐level ecosystem service supply, although interactions between landscape units (e.g. the movement of pollinators) must also be considered.

Climate warming and elevated CO2 can modify nutrient cycling mediated by enzymes in soils, especially in cold-limited ecosystems with a low availability of nutrients and a high temperature sensitivity of decomposition and mineralization. We estimated responses of soil extracellular enzyme activities (EEAs) to 6 years of soil warming and 9 years of CO2 enrichment at an Alpine treeline site. EEAs were measured in the litter (L), fermentation (F) and humified (H) horizons under Larix decidua and Pinus uncinata trees. Soil warming indirectly affected EEAs through altered soil moisture, fine root biomass, and C:N ratio of the organic horizons. Warming increased β-glucosidase and β-xylosidase activities in the F horizon but led to reduced laccase activity in the L horizon, probably caused by drying of the litter horizon associated with the treatment. In the H horizon, previous CO2 enrichment altered the activity of leucine amino peptidase, N-acetylglucosaminidase, and phosphatase. No interactive effects between warming and CO2 enrichment were detected. Warming affected the temperature sensitivity of β-xylosidase but not of the other enzymes. Altered EEAs after six years of soil warming indicate a sustained stimulation of carbon, nitrogen and nutrient cycling under climatic warming at the alpine treeline.

The content of organic carbon (C) in soils is not stable, but depends on a number of environmental variables and biogeochemical processes that actively regulate its balance. An improved identification of the environmental variables that can be used as predictors of soil organic C (SOC) content is needed to reduce uncertainties of how the soil C reservoir will respond to environmental change. Although several simulations rely on the amount of clay to reproduce changes in the balance of SOC, recent efforts have suggested that other soil physicochemical properties may serve as better predictors. Here we tested whether the effective cation exchange capacity (CEC eff.), may be a more suitable predictor of the content and potential preservation of SOC as compared to the mere quantification of clay-size particles. We further assessed how various climatic, vegetation and edaphic variables explain the variance of SOC content across different soil depths and soil pH classes. A set of more than 1000 forest sites across Switzerland, spanning a unique gradient of mean annual precipitation (636–2484 mm), altitude (277–2207 m a.s.l), pH (2.8–8.1) and representing different geologies and soil orders was used as a case study for this linear model analysis. Our results showed that CEC eff. has the largest explanatory potential of SOC content (35 % of response variance in the complete mineral soil profile) as compared to the amount of clay (which only explained 7 % of the response variance in the complete mineral soil profile) and other environmental variables. CEC eff. is strongly linked to SOC especially in the top mineral soil (0–30 cm depth) with the larger presence of organic matter. At deeper soil depths most of the variance in SOC is instead explained by climate, which in Switzerland is related to a greater weathering activity and translocation of organic C through leaching with increasing mean annual precipitation. We further observed soil pH to have a complex influence on SOC content, with CEC eff. being a dominant variable controlling SOC content at pH > 4.5 in the upper mineral soil and pH > 6 in the subsoil. Since CEC eff. is an edaphic property which is intimately associated to both the conditions that shaped the soil and the current edaphic physicochemical conditions, these findings indicate that considering CEC eff. as an integrative proxy for the potential preservation of SOC and its alteration could improve future predictions of how the soil C reservoir will feed back to environmental change.

Many experiments have shown that loss of biodiversity reduces the capacity of ecosystems to provide the multiple services on which humans depend. However, experiments necessarily simplify the complexity of natural ecosystems and will normally control for other important drivers of ecosystem functioning, such as the environment or land use. In addition, existing studies typically focus on the diversity of single trophic groups, neglecting the fact that biodiversity loss occurs across many taxa and that the functional effects of any trophic group may depend on the abundance and diversity of others. Here we report analysis of the relationships between the species richness and abundance of nine trophic groups, including 4,600 above- and below-ground taxa, and 14 ecosystem services and functions and with their simultaneous provision (or multifunctionality) in 150 grasslands. We show that high species richness in multiple trophic groups (multitrophic richness) had stronger positive effects on ecosystem services than richness in any individual trophic group; this includes plant species richness, the most widely used measure of biodiversity. On average, three trophic groups influenced each ecosystem service, with each trophic group influencing at least one service. Multitrophic richness was particularly beneficial for 'regulating' and 'cultural' services, and for multifunctionality, whereas a change in the total abundance of species or biomass in multiple trophic groups (the multitrophic abundance) positively affected supporting services. Multitrophic richness and abundance drove ecosystem functioning as strongly as abiotic conditions and land-use intensity, extending previous experimental results to real-world ecosystems. Primary producers, herbivorous insects and microbial decomposers seem to be particularly important drivers of ecosystem functioning, as shown by the strong and frequent positive associations of their richness or abundance with multiple ecosystem services. Our results show that multitrophic richness and abundance support ecosystem functioning, and demonstrate that a focus on single groups has led to researchers to greatly underestimate the functional importance of biodiversity.

Fine roots support the water and nutrient demands of plants and supply carbon to soils. Quantifying turnover times of fine roots is crucial for modeling soil organic matter dynamics and constraining carbon cycle–climate feedbacks. Here we challenge widely used isotope-based estimates suggesting the turnover of fine roots of trees to be as slow as a decade. By recording annual growth rings of roots from woody plant species, we show that mean chronological ages of fine roots vary from, Fine-root lifetimes and carbon inputs from roots into soil impact carbon cycle-climate feedbacks yet remain poorly constrained. Here, using annual-growth rings and radiocarbon dating, the authors show that the chronological age of fine roots is substantially younger than that of the carbon used for their growth.

Trade-offs and synergies in the supply of forest ecosystem services are common but the drivers of these relationships are poorly understood. To guide management that seeks to promote multiple services, we investigated the relationships between 12 stand-level forest attributes, including structure, composition, heterogeneity and plant diversity, plus 4 environmental factors, and proxies for 14 ecosystem services in 150 temperate forest plots. Our results show that forest attributes are the best predictors of most ecosystem services and are also good predictors of several synergies and trade-offs between services. Environmental factors also play an important role, mostly in combination with forest attributes. Our study suggests that managing forests to increase structural heterogeneity, maintain large trees, and canopy gaps would promote the supply of multiple ecosystem services. These results highlight the potential for forest management to encourage multifunctional forests and suggest that a coordinated landscape-scale strategy could help to mitigate trade-offs in human-dominated landscapes., Managing forests for the supply of multiple ecosystem services (ES) is key given potential trade-offs among services. Here, the authors analyse how forest stand attributes generate trade-offs among ES and the relative contribution of forest attributes and environmental factors to predict services.

© 2015, Springer International Publishing Switzerland. Aims: Subsoil organic carbon (OC) tends to be older and is presumed to be more stable than topsoil OC, but the reasons for this are not yet resolved. One hypothesis is that decomposition rates decrease with increasing soil depth. We tested whether decomposition rates of beech fine root litter varied with depth for a range of soils using a litterbag experiment in German beech forest plots. Methods: In three study regions (Schorfheide-Chorin, Hainich-Dün and Schwäbische-Alb), we buried 432 litterbags containing 0.5 g of standardized beech root material (fine roots with a similar chemical composition collected from 2 year old Fagus sylvatica L. saplings, root diameter < 2 mm) at three different soil depths (5, 20 and 35 cm). The decomposition rates as well as the changes in the carbon (C) and nitrogen (N) concentrations of the decomposing fine root litter were determined at a 6 months interval during a 2 years field experiment. Results: The amount of root litter remaining after 2 years of field incubation differed between the study regions (76 ± 2 % in Schorfheide-Chorin, 85 ± 2 % in Schwäbische-Alb, and 88 ± 2 % in Hainich-Dün) but did not vary with soil depth. Conclusions: Our results indicate that the initial fine root decomposition rates are more influenced by regional scale differences in environmental conditions including climate and soil parent material, than by changes in microbial activities with soil depth. Moreover, they suggest that a similar potential to decompose new resources in the form of root litter exists in both surface and deep soils.

Increased CO2 emissions and global warming may alter the composition of fungal communities through the removal of temperature limitation in the plant–soil system, faster nitrogen (N) cycling and changes in the carbon (C) allocation of host plants to the rhizosphere. At a Swiss treeline featuring Larix decidua and Pinus uncinata, the effects of multiple years of CO2 enrichment and experimental soil warming on the fungal community composition in the organic horizons were analysed using 454-pyrosequencing of ITS2 amplicons. Sporocarp production and colonization of ectomycorrhizal root tips were investigated in parallel. Fungal community composition was significantly altered by soil warming, whereas CO2 enrichment had little effect. Tree species influenced fungal community composition and the magnitude of the warming responses. The abundance of ectomycorrhizal fungal taxa was positively correlated with N availability, and ectomycorrhizal taxa specialized for conditions of high N availability proliferated with warming, corresponding to considerable increases in inorganic N in warmed soils. Traits related to N utilization are important in determining the responses of ectomycorrhizal fungi to warming in N-poor cold ecosystems. Shifts in the overall fungal community composition in response to higher temperatures may alter fungal-driven processes with potential feedbacks on ecosystem N cycling and C storage at the alpine treeline.

Land-use change and intensification play a key role in the current biodiversity crisis. The resulting species loss can have severe effects on ecosystem functions and services, thereby increasing ecosystem vulnerability to climate change. We explored whether land-use intensification (i.e. fertilization intensity), plant diversity and other potentially confounding environmental factors may be significantly related to water use (i.e. drought stress) of grassland plants. Drought stress was assessed using δ13C abundances in aboveground plant biomass of 150 grassland plots across a gradient of land-use intensity. Under water shortage, plants are forced to increasingly take up the heavier 13C due to closing stomata leading to an enrichment of 13C in biomass. Plants were sampled at the community level and for single species, which belong to three different functional groups (one grass, one herb, two legumes). Results show that plant diversity was significantly related to the δ13C signal in community, grass and legume biomass indicating that drought stress was lower under higher diversity, although this relation was not significant for the herb species under study. Fertilization, in turn, mostly increased drought stress as indicated by more positive δ13C values. This effect was mostly indirect by decreasing plant diversity. In line with these results, we found similar patterns in the δ13C signal of the organic matter in the topsoil, indicating a long history of these processes. Our study provided strong indication for a positive biodiversity-ecosystem functioning relationship with reduced drought stress at higher plant diversity. However, it also underlined a negative reinforcing situation: as land-use intensification decreases plant diversity in grasslands, this might subsequently increases drought sensitivity. Vice-versa, enhancing plant diversity in species-poor agricultural grasslands may moderate negative effects of future climate change., Science of The Total Environment, 566-567, ISSN:0048-9697, ISSN:1879-1026

Background and Aims The response of forest ecosystems to continuous nitrogen (N) deposition is still uncertain. We investigated imports and exports of dissolved N from mull-type organic layers to identify the controls of N leaching in Central European beech forests under continuous N deposition. Methods Dissolved N fluxes with through fall and through mull-type organic layers (litter leachate) were measured continuously in 12 beech forests on calcareous soil in two regions in Germany over three consecutive growing seasons. Results Mean growing season net (i.e. litter leachate – throughfall flux) fluxes of total dissolved N (TDN) from the organic layer were low (2.3 ± 5.6 kg ha⁻¹) but varied widely from 12.9 kg ha⁻¹ to–8.3kgha⁻¹. The small increase of dissolved N fluxes during the water passage through mull-type organic layers suggested that high turnover rates coincided with high microbial N assimilation and plant N uptake. Stand basal area had a positive feedback on N fluxes by providing litter for soil organic matter formation. Plant diversity, especially herb diversity, reduced dissolved N fluxes. Soil fauna biomass increased NO−3 -N fluxes with litter leachate by stimulating mineralization. Microbial biomass measures were not related to dissolved N fluxes. Conclusions Our results show that dissolved N exports from organic layers contain significant amounts of throughfall-derived N (mainly NO−3 -N) that flushes through the organic layer but also highlight that N leaching from organic layers is driven by the complex interplay of plants, animals and microbes. Furthermore, diverse understories reduce N leaching from Central European beech forests.

Climate warming is shifting the elevational boundary between forests and tundra upwards, but the related belowground responses are poorly understood. In the pristine South and Polar Urals with shifts of the treeline ecotone documented by historical photographs, we investigated fine root dynamics and production of extramatrical mycorrhizal mycelia (EMM) along four elevational transects reaching from the closed forest to the treeless tundra. In addition, we analysed elevational differences in climate and vegetation structure, and excavated trees to estimate related changes in the partitioning between below- and aboveground biomass. Fine root biomass of trees (

Global change, especially land‐use intensification, affects human well‐being by impacting the delivery of multiple ecosystem services (multifunctionality). However, whether biodiversity loss is a major component of global change effects on multifunctionality in real‐world ecosystems, as in experimental ones, remains unclear. Therefore, we assessed biodiversity, functional composition and 14 ecosystem services on 150 agricultural grasslands differing in land‐use intensity. We also introduce five multifunctionality measures in which ecosystem services were weighted according to realistic land‐use objectives. We found that indirect land‐use effects, i.e. those mediated by biodiversity loss and by changes to functional composition, were as strong as direct effects on average. Their strength varied with land‐use objectives and regional context. Biodiversity loss explained indirect effects in a region of intermediate productivity and was most damaging when land‐use objectives favoured supporting and cultural services. In contrast, functional composition shifts, towards fast‐growing plant species, strongly increased provisioning services in more inherently unproductive grasslands.

Interactions occur between two or more organisms affecting each other. Interactions are decisive for the ecology of the organisms. Without direct experimental evidence the analysis of interactions is difficult. Correlation analyses that are based on co-occurrences are often used to approximate interaction. Here, we present a new mathematical model to estimate the interaction strengths between taxa, based on changes in their relative abundances across environmental gradients.

We used bomb-radiocarbon and raw minirhizotron lifetimes of fine-roots (

Background and aims: Fine root decomposition contributes significantly to element cycling in terrestrial ecosystems. However, studies on root decomposition rates and on the factors that potentially influence them are fewer than those on leaf litter decomposition. To study the effects of region and land use intensity on fine root decomposition, we established a large scale study in three German regions with different climate regimes and soil properties. Methods In 150 forest and 150 grassland sites we deployed litterbags (100 μm mesh size) with standardized litter consisting of fine roots from European beech in forests and from a lowland mesophilous hay meadow in grasslands. In the central study region, we compared decomposition rates of this standardized litter with root litter collected on-site to separate the effect of litter quality from environmental factors. Results: Standardized herbaceous roots in grassland soils decomposed on average significantly faster (24 ± 6 % mass loss after 12 months, mean ± SD) than beech roots in forest soils (12 ± 4 %; p < 0.001). Fine root decomposition varied among the three study regions. Land use intensity, in particular N addition, decreased fine root decomposition in grasslands. The initial lignin:N ratio explained 15 % of the variance in grasslands and 11 % in forests. Soil moisture, soil temperature, and C:N ratios of soils together explained 34 % of the variance of the fine root mass loss in grasslands, and 24 % in forests. Conclusions: Grasslands, which have higher fine root biomass and root turnover compared to forests, also have higher rates of root decomposition. Our results further show that at the regional scale fine root decomposition is influenced by environmental variables such as soil moisture, soil temperature and soil nutrient content. Additional variation is explained by root litter quality. © 2014 The Author(s).

Species diversity promotes the delivery of multiple ecosystem functions (multifunctionality). However, the relative functional importance of rare and common species in driving the biodiversity–multifunctionality relationship remains unknown. We studied the relationship between the diversity of rare and common species (according to their local abundances and across nine different trophic groups), and multifunctionality indices derived from 14 ecosystem functions on 150 grasslands across a land-use intensity (LUI) gradient. The diversity of above- and below-ground rare species had opposite effects, with rare above-ground species being associated with high levels of multifunctionality, probably because their effects on different functions did not trade off against each other. Conversely, common species were only related to average, not high, levels of multifunctionality, and their functional effects declined with LUI. Apart from the community-level effects of diversity, we found significant positive associations between the abundance of individual species and multifunctionality in 6% of the species tested. Species-specific functional effects were best predicted by their response to LUI: species that declined in abundance with land use intensification were those associated with higher levels of multifunctionality. Our results highlight the importance of rare species for ecosystem multifunctionality and help guiding future conservation priorities.