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Eisenhauer, N.

Publications and source records attributed to Eisenhauer, N..

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Global monitoring of soil animal communities using a common methodology

Here we introduce the Soil BON Foodweb Team, a cross-continental collaborative network that aims to monitor soil animal communities and food webs using consistent methodology at a global scale. Soil animals support vital soil processes via soil structure modification, direct consumption of dead organic matter, and interactions with microbial and plant communities. Soil animal effects on ecosystem functions have been demonstrated by correlative analyses as well as in laboratory and field experiments, but these studies typically focus on selected animal groups or species at one or few sites with limited variation in environmental conditions. The lack of comprehensive harmonised large-scale soil animal community data including microfauna, mesofauna, and macrofauna, in conjunction with related soil functions, limits our understanding of biological interactions in soil communities and how these interactions affect ecosystem functioning. To provide such data, the Soil BON Foodweb Team invites researchers worldwide to use a common methodology to address six long-term goals: (1) to collect globally representative harmonised data on soil micro-, meso-, and macrofauna communities; (2) to describe key environmental drivers of soil animal communities and food webs; (3) to assess the efficiency of conservation approaches for the protection of soil animal communities; (4) to describe soil food webs and their association with soil functioning globally; (5) to establish a global research network for soil biodiversity monitoring and collaborative projects in related topics; (6) to reinforce local collaboration networks and expertise and support capacity building for soil animal research around the world. In this paper, we describe the vision of the global research network and the common sampling protocol to assess soil animal communities and advocate for the use of standard methodologies across observational and experimental soil animal studies. We will use this protocol to conduct soil animal assessments and reconstruct soil food webs on the sites included in the global soil biodiversity monitoring network, Soil BON, allowing us to assess linkages among soil biodiversity, vegetation, soil physico-chemical properties, and ecosystem functions. In the present paper, we call for researchers especially from countries and ecoregions that remain underrepresented in the majority of soil biodiversity assessments to join us. Together we will be able to provide science-based evidence to support soil biodiversity conservation and functioning of terrestrial ecosystems.

ecology↗

Aboveground impacts of a belowground invader: how invasive earthworms alter aboveground arthropod communities in a northern North American forest

Declining arthropod communities have recently gained a lot of attention with climate and land-use change among the most-frequently discussed drivers. Here, we focus on a seemingly underrepresented driver of arthropod-community decline: biological invasions. For [~]12,000 years, earthworms have been absent from wide parts of northern North America, but they have been re-introduced with dramatic consequences. Most studies investigating earthworm-invasion impacts focus on the belowground world, resulting in limited knowledge on aboveground-community changes. We present observational data on earthworm, plant, and aboveground-arthropod communities in 60 plots, distributed across areas with increasing invasion status (low, medium, high) in a Canadian forest. We analyzed how earthworm-invasion status and biomass impact aboveground arthropod community abundance, biomass, and species richness, and how earthworm impacts cascade across trophic levels. We sampled [~]13,000 arthropods, dominated by Hemiptera, Diptera, Araneae, Thysanoptera, and Hymenoptera. Total arthropod abundance, biomass, and species richness declined significantly from areas of low to those with high invasion status with reductions of 61, 27, and 18%, respectively. Structural Equation Models unraveled that earthworms directly and indirectly impact arthropods across trophic levels. We show that earthworm invasion can alter aboveground multitrophic arthropod communities and suggest that belowground invasions can be important drivers of aboveground-arthropod decline.

ecology↗

Diversity-induced plant history and soil history effects modulate plant responses to global change

Global change has dramatic impacts on grassland diversity. However, little is known about how fast species can adapt to these changes and how this affects their responses to global change. To close this gap, we performed a common garden experiment testing whether plant responses to global change are influenced by the selection history of the plants and the conditioning history of soil at different levels of plant diversity. Therefore, we collected seeds and took soil samples from 14-year old plant communities of a biodiversity experiment. Offspring of plants from low- and high-diversity communities were either grown in their own soil or in soil of a different community, and were either exposed to drought, increased nitrogen input, or a combination of both. Results show that, under nitrogen addition, offspring of plants selected at high diversity produced more biomass than those selected at low diversity, while drought neutralized differences in biomass production. Moreover, under the influence of global change drivers, mainly soil, and to a lesser extent plant history, influenced the expression of plant traits. Our results show that plant diversity modulates plant-soil interactions and growth strategies of plants, which feedback on the eco-evolutionary pathways of the plants and thus their responses to global change.

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Distinct effects of host and neighbour tree identity on arbuscular and ectomycorrhizal fungi along a tree diversity gradient

Plant diversity and plant-related ecosystem functions have been in focus in biodiversity-ecosystem functioning studies. However, in this context, biotic interactions with mycorrhizal fungi have been understudied although they are crucial for plant-resource acquisition. We investigated the effects of tree species richness, tree mycorrhizal type on arbuscular (AMF) and ectomycorrhizal fungal (EMF) communities. We aimed to understand how dissimilarities in taxa composition and beta-diversity are related to target trees and neighbours of the same/different mycorrhizal type. We sampled a tree experiment with saplings ([~]7 years old), where tree species richness (monocultures, 2-species, and 4-species mixtures) and mycorrhizal type were manipulated. AMF and EMF richness significantly increased with increasing tree species richness. AMF richness of mixture plots resembled that of the sum of the respective monocultures, whereas EMF richness of mixture plots was lower compared to the sum of the respective monocultures. Specialisation scores revealed significantly more specialised AMF than EMF suggesting that, in contrast to previous studies, AMF were more specialised, whereas EMF were not. We further found that AMF communities were little driven by the surrounding trees, whereas EMF communities were. Our study revealed the drivers of mycorrhizal fungal communities and further highlights the distinct strategies of AMF and EMF.

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The archives are half-empty: a field-wide assessment of the availability of microbial community sequencing data

The sequencing revolution has resulted in the explosive growth of public genetic repositories. These repositories now hold invaluable collections of 16S rRNA gene amplicon sequences, but the extent to which the currently archived data is findable, accessible, and reusable has not been evaluated. We conducted a field-wide assessment of the availability and state of publicly archived 16S rRNA gene amplicon sequencing data. Using custom-built pattern-based text extraction algorithms, we searched 26,927 publications in 17 microbiology or microbial ecology journals, and identified 2,015 studies which performed 16S rRNA gene amplicon sequencing. We found, for example, that 7.2% of these had not been made public at the time of analysis, a trend which increased over time. Of the 635 studies targeting the V3-V4 region of the 16S rRNA gene, 40.3% contained data which was not available or not reusable, and for 25.5% of the studies, faults in data formatting or data labelling were likely to create obstacles in data reuse. Taken together, only 34% of these datasets had potentially reusable data. Our study reveals significant gaps in the availability of currently deposited community sequencing data, identifies major contributors to data loss, and offers suggestions for improving data archiving practices in the future.

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Biodiversity mediates the effects of stressors but not nutrients on litter decomposition

Understanding the consequences of ongoing biodiversity changes for ecosystems is a pressing challenge. Controlled biodiversity-ecosystem function experiments with random biodiversity loss scenarios have demonstrated that more diverse communities usually provide higher levels of ecosystem functioning. However, it is not clear if these results predict the ecosystem consequences of environmental changes that cause non-random alterations in biodiversity and community composition. We synthesized 69 independent studies reporting 660 observations of the impacts of two pervasive drivers of global change (chemical stressors and nutrient enrichment) on animal and microbial decomposer diversity and litter decomposition. Using meta-analysis and structural equation modelling, we show that declines in decomposer diversity and abundance explain reduced litter decomposition in response to stressors but not to nutrients. While chemical stressors generally reduced biodiversity and ecosystem functioning, detrimental effects of nutrients occurred only at high levels of nutrient inputs. Thus, more intense environmental change does not always result in stronger responses, illustrating the complexity of ecosystem consequences of biodiversity change. Overall, these findings provide strong empirical evidence for significant real-world biodiversity-ecosystem functioning relationships when human activities decrease biodiversity. This highlights that the consequences of biodiversity change for ecosystems are nontrivial and depend on the kind of environmental change.

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Modeling How Community Assembly Alters the Functioning of Ecosystems

Although the effects of species richness on ecosystem functioning have been extensively studied, there is increased interest in understanding how community assembly in general might alter the functioning of ecosystems. We focus on two complementary approaches for evaluating how community assembly influences ecosystem function (here, productivity). The first quantifies the relative importance of complementarity and selection by contrasting monocultures with polycultures. The second identifies the effects of species losses and/or gains relative to the baseline polyculture, as well as the indirect effects on other species productivity. We evaluate and contrast these two approaches, using simulated communities structured by different, known competition mechanisms, where species compete for different resources and experience varying levels of environmental heterogeneity. We find that the metrics provided by these approaches can jointly discriminate the mechanisms of competition driving productivity. We then apply our methods to data from a long-term biodiversity-ecosystem experiment (the Jena Experiment) and find that the data do not correspond to any of the competition scenarios we modeled. We address two additional possible complications: facilitation by nitrogen fixing plants, and non-equilibrium behavior during community assembly, and find that a combination of resource competition and facilitation by nitrogen fixing plants is the more likely explanation for the results obtained at Jena.

ecology↗

Biotic and abiotic drivers of soil microbial functions across tree diversity experiments

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

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Climate change and intensive land use reduce soil animal biomass through dissimilar pathways

Global change drivers, such as climate and land use, may profoundly influence body size, density, and biomass of organisms. It is still poorly understood how these concurrent drivers interact in affecting ecological communities. We present results of an experimental field study assessing the interactive effects of climate change and land-use intensification on body size, density, and biomass of soil microarthropods. We found that both climate change and intensive land use decreased their total biomass. Strikingly, this reduction was realized via two dissimilar pathways: climate change reduced mean body size, while intensive land use decreased population size. These findings highlight that two of the most pervasive global change drivers operate via different pathways when decreasing soil animal biomass. These shifts in soil communities may threaten essential ecosystem functions like organic matter turnover and nutrient cycling in future ecosystems. SignificanceMany important ecosystem functions are determined by the biomass of soil animal, however, how their biomass may respond to climate change and land-use intensification still remains unknown. We conducted a large field study to investigate the potential interaction between these two pervasive global change drivers, and disentangle the pathways where they contribute to the changes in soil animal biomass. Our findings are exceptionally novel by showing detrimental, but largely independent, effects of climate change and land-use intensity on soil animal biomass, and that these independent effects can be explained by two dissimilar pathways: climate change reduced mean body size, while intensive land use decreased population size. Notably, consistent climate change effects under different land-use regimes suggest that (1) the identified pathways may apply to a wide range of environmental conditions, and (2) current extensive land-use regimes do not mitigate detrimental climate change effects on ecosystems.

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Plant traits are poor predictors of long-term ecosystem functioning

Earth is home to over 350,000 vascular plant species1 that differ in their traits in innumerable ways. Yet, a handful of functional traits can help explaining major differences among species in photosynthetic rate, growth rate, reproductive output and other aspects of plant performance2-6. A key challenge, coined "the Holy Grail" in ecology, is to upscale this understanding in order to predict how natural or anthropogenically driven changes in the identity and diversity of co-occurring plant species drive the functioning of ecosystems7, 8. Here, we analyze the extent to which 42 different ecosystem functions can be predicted by 41 plant traits in 78 experimentally manipulated grassland plots over 10 years. Despite the unprecedented number of traits analyzed, the average percentage of variation in ecosystem functioning that they jointly explained was only moderate (32.6%) within individual years, and even much lower (12.7%) across years. Most other studies linking ecosystem functioning to plant traits analyzed no more than six traits, and when including either only six random or the six most frequently studied traits in our analysis, the average percentage of explained variation in across-year ecosystem functioning dropped to 4.8%. Furthermore, different ecosystem functions were driven by different traits, with on average only 12.2% overlap in significant predictors. Thus, we did not find evidence for the existence of a small set of key traits able to explain variation in multiple ecosystem functions across years. Our results therefore suggest that there are strong limits in the extent to which we can predict the long-term functional consequences of the ongoing, rapid changes in the composition and diversity of plant communities that humanity is currently facing.

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Blind spots in global soil biodiversity and ecosystem function research

Soils harbor a substantial fraction of the worlds biodiversity, contributing to many crucial ecosystem functions. It is thus essential to identify general macroecological patterns related to the distribution and functioning of soil organisms to support their conservation and governance. Here we identify and characterize the existing gaps in soil biodiversity and ecosystem function data across soil macroecological studies and >11,000 sampling sites. These include significant spatial, environmental, taxonomic, and functional gaps, and an almost complete absence of temporally explicit data. We also identify the limitations of soil macroecological studies to explore general patterns in soil biodiversity-ecosystem functioning relationships, with only 0.6% of all sampling sites having a non-systematic coverage of both biodiversity and function datasets. Based on this information, we provide clear priorities to support and expand soil macroecological research.

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The results of biodiversity-ecosystem functioning experiments are realistic

A large body of research shows that biodiversity loss can reduce ecosystem functioning, thus providing support for the conservation of biological diversity1-4. Much of the evidence for this relationship is drawn from biodiversity-ecosystem functioning experiments (hereafter: biodiversity experiments), in which biodiversity loss is simulated by randomly assembling communities of varying species diversity, and ecosystem functions are measured5-9. This random assembly has led some ecologists to question the relevance of biodiversity experiments to real-world ecosystems, where community assembly may often be non-random and influenced by external drivers, such as climate or land-use intensification10-18. Despite these repeated criticisms, there has been no comprehensive, quantitative assessment of how experimental and real-world plant communities really differ, and whether these differences invalidate the experimental results. Here, we compare data from two of the largest and longest-running grassland biodiversity experiments globally (Jena Experiment, Germany; BioDIV, USA) to related real-world grassland plant communities in terms of their taxonomic, functional, and phylogenetic diversity and functional-trait composition. We found that plant communities of biodiversity experiments have greater variance in these compositional features than their real-world counterparts, covering almost all of the variation of the real-world communities (82-96%) while also containing community types that are not currently observed in the real world. We then re-analysed a subset of experimental data that included only ecologically-realistic communities, i.e. those comparable to real-world communities. For ten out of twelve biodiversity-ecosystem functioning relationships, biodiversity effects did not differ significantly between the full dataset of biodiversity experiments and the ecologically-realistic subset of experimental communities. This demonstrates that the results of biodiversity experiments are largely insensitive to the inclusion/exclusion of unrealistic communities. By bridging the gap between experimental and real-world studies, these results demonstrate the validity of inferences from biodiversity experiments, a key step in translating their results into specific recommendations for real-world biodiversity management.

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Global distribution of earthworm diversity

Soil organisms provide crucial ecosystem services that support human life. However, little is known about their diversity, distribution, and the threats affecting them. Here, we compiled a global dataset of sampled earthworm communities from over 7000 sites in 56 countries to predict patterns in earthworm diversity, abundance, and biomass. We identify the environmental drivers shaping these patterns. Local species richness and abundance typically peaked at higher latitudes, while biomass peaked in the tropics, patterns opposite to those observed in aboveground organisms. Similar to many aboveground taxa, climate variables were more important in shaping earthworm communities than soil properties or habitat cover. These findings highlight that, while the environmental drivers are similar, conservation strategies to conserve aboveground biodiversity might not be appropriate for earthworm diversity, especially in a changing climate.\n\nOne sentence summaryGlobal patterns of earthworm diversity, abundance and biomass are driven by climate but patterns differ from many aboveground taxa.

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Diversity-dependent plant-soil feedbacks underlie long-term plant diversity effects on primary productivity

Although diversity-dependent plant-soil feedbacks (PSFs) may contribute significantly to plant diversity effects on ecosystem functioning, the influence of underlying abiotic and biotic mechanistic pathways have been little explored to date. Here, we assessed such pathways with a PSF experiment using soil conditioned for [≥]12 years from two grassland biodiversity experiments. Model plant communities differing in diversity were grown in soils conditioned by plant communities with either low- or high-diversity (soil history). Our results reveal that plant diversity can modify plant productivity through both diversity-mediated plant-plant and plant-soil interactions, with the main driver (current plant diversity or soil history) differing with experimental context. The underlying mechanisms of PSFs were explained to a significant extent by both abiotic and biotic pathways (specifically, nematode richness and soil nitrogen availability). Thus, effects of plant diversity loss on ecosystem functioning may persist or even increase over time because of biotic and abiotic soil legacy effects.

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The effects of drought and nutrient addition on soil organisms vary across taxonomic groups, but are constant across seasons

Anthropogenic global change alters the activity and functional composition of soil communities that are responsible for crucial ecosystem functions and services. Two of the most pervasive global change drivers are drought and nutrient enrichment. However, the responses of soil organisms to interacting global change drivers remain widely unknown. We tested the interactive effects of extreme drought and fertilization on soil biota ranging from microbes to invertebrates across seasons. We expected drought to reduce the activity of soil organisms and fertilization to induce positive bottom-up effects via increased plant productivity. Furthermore, we hypothesized fertilization to reinforce drought effects through enhanced plant growth, resulting in even dryer soil conditions. Our results revealed that drought had detrimental effects on soil invertebrate feeding activity and simplified nematode community structure, whereas soil microbial activity and biomass were unaffected. Microbial biomass increased in response to fertilization, whereas invertebrate feeding activity substantially declined. Notably, these effects were consistent across seasons. The dissimilar responses suggest that soil biota differ vastly in their vulnerability to global change drivers. As decomposition and nutrient cycling are driven by the interdependent concurrence of microbial and faunal activity, this may imply far-reaching consequences for crucial ecosystem processes in a changing world.

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Testing soil nematode extraction efficiency using different variations of the Baermann funnel method

Nematodes are increasingly used as powerful bioindicators of soil food web composition and functioning in ecological studies. Todays ecological research aims to investigate not only local relationships but global patterns, which requires consistent methodology across locales. Thus, a common and easy extraction protocol of soil nematodes is needed. In this study, we present a detailed protocol of the Baermann funnel method and highlight how different soil pre-treatments and equipment (soil type, amount of soil, sieving, filter type) can affect extraction efficiency and community composition by using natural nematode communities. We found that highest nematode extraction efficiency was achieved using lowest soil weight (25 g instead of 50 g or 100 g) in combination with soil sieving, and by using milk filters (instead of paper towels). PCA at the family level revealed that different pre-treatments significantly affected nematode community composition. Increasing the amount of soil increased the proportion of larger-sized nematodes being able to overcome long distances. Sieving is suggested to break up soil aggregates and, therefore, facilitate moving in general. Interestingly, sieving did not negatively affect larger nematodes that are supposed to have a higher probability of getting bruised during sieving. The present study shows that variations in the extraction protocol can alter the total density and community composition of extracted nematodes and provides recommendations for an efficient and standardized approach in future studies. Having a simple, cheap, and standardized extraction protocol can facilitate the assessment of soil biodiversity in global contexts.

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Evolution increases ecosystem temporal stability and recovery from a flood in grassland communities

Understanding factors that increase ecosystem stability is critical in the face of environmental change. Experiments simulating species loss from grassland ecosystems have shown that losing biodiversity decreases the ability of ecosystems to buffer negative effects of disturbances. However, as the originally sown experimental communities with reduced biodiversity develop, plant evolutionary processes or the assembly of interacting soil organisms may allow them to develop stability and resilience over time. We explored such effects in a long-term grassland biodiversity experiment with plant communities with either a history of co-occurrence (selected communities) or no such history (naive communities) over a four-year period in which a major flood disturbance occurred. We found selected communities had temporally more stable biomass than the same communities of naive plants, especially at low species richness. Furthermore, selected communities showed greater short-term biomass recovery after flooding, resulting in more stable post-flood productivity. In contrast to a previous study, the positive diversity-stability relationship was maintained after the flooding. Our results were consistent across three soil treatments simulating the presence or absence of co-selected microbial communities. We suggest that prolonged exposure of plant populations to a particular community context and abiotic site conditions can increase ecosystem temporal stability and resistance to disturbance. We argue that selection during the course of a biodiversity experiment is the most parsimonious explanation for these effects. A history of co-occurrence can in part compensate for species loss, as can high plant diversity in part compensate for the missing opportunity of such adaptive adjustments.

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