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Tree species diversity plays a central role for forest productivity, but factors driving positive biodiversity-productivity relationships remain poorly understood. In a biodiversity experiment manipulating tree diversity and mycorrhizal associations, we examined the roles of above- and belowground processes in modulating wood productivity in young temperate tree communities, as well as potential underlying mechanisms. We found that tree species richness increased forest productivity indirectly by enhancing structural complexity within communities. After six years, structurally complex communities were twice as productive as structurally simple stands. This pattern was consistent across stands with different mycorrhizal associations. Our results also demonstrate that taxonomic diversity and functional variation in shade tolerance, but not phenotypic plasticity, are key drivers of structural complexity in mixtures, which in turn lead to overyielding. Consideration of stand structural complexity appears to be a crucial element in predicting carbon sequestration in the early successional stages of mixed-species forests.

Earthworms are considered ecosystem engineers due to their fundamental impact on soil structure, soil processes and on other soil biota. An invasion of non-native earthworm species has altered soils of North America since European settlement, a process currently expanding into still earthworm-free forest ecosystems due to continuous spread and increasing soil temperatures owing to climate change. Although earthworms are known to modify soil microbial diversity and activity, it is as yet unclear how eukaryote consumers in soil microbial food webs will be affected. Here, we investigated how earthworm invasion affects the diversity of Cercozoa, one of the most dominant protist taxa in soils. Although the composition of the native cercozoan community clearly shifted in response to earthworm invasion, the communities of the different forests showed distinct responses. We identified 39 operational taxonomic units (OTUs) exclusively indicating earthworm invasion, hinting at an earthworm-associated community of Cercozoa. In particular, Woronina pythii, a hyper-parasite of plant-parasitic Oomycota in American forests, increased strongly in the presence of invasive earthworms, indicating an influence of invasive earthworms on oomycete communities and potentially on forest health, which requires further research.

Anthropogenic global changes are impacting biodiversity, however, many previous meta-analyses investigating the impact of different global changes on biodiversity have omitted soil fauna, or are limited in the scope of the global changes studied. Threats to soil biodiversity by global changes need to be understood to mitigate effects on ecosystem services provided by soils. We conducted a meta-analysis using 3,173 effect sizes from 627 publications focused on six global changes (climate change, land-use intensification, pollution, nutrient enrichment, invasive species, and habitat fragmentation) and their associated environmental stressors on soil fauna. We classified stressors as either pulse (short-term, acute) or press (long-term, chronic) stressors, and expected pulse stressors to have less impact on soil biodiversity due to buffering effects of the soil. Unexpectedly, pollution caused the largest loss in soil fauna communities, which is worrying due to continually increasing levels of pollution, as well as the poor mechanistic understanding of pollution impacts. There was no clear pattern of pulse stressors having a smaller impact on soil biodiversity than press stressors. Overall, this work shows the importance of including soil biodiversity in large-scale global change analyses, as soil organisms often do not show the same responses as organisms above-ground.

Climate extremes in tandem with biodiversity change affect emissions of biogenic volatile organic compounds (BVOCs) from plants and, as a result, the formation of biogenic secondary organic aerosols (BSOA). The resulting BSOA can have a wide variety of impacts, such as on Earth’s radiative balance and cloud formation. However, it is unclear to what extent changes in BVOC emissions and BSOA formation are related to biodiversity. Here we present a conceptual framework of the relationships between biodiversity and BVOC emissions based on our current mechanistic understanding and existing knowledge. We tested parts of this framework using a tree diversity experiment as a case-study. We find that the amount of BVOCs in most cases decreases with biodiversity, implying that tree mixtures produce less than expected BVOC compared to tree monocultures. However, some BSOA compounds increased and some decreased relative to what is expected from monocultures. Based on these mixed results, we recommend further field measurements on the patterns of BVOC emission and BSOA formation across biodiversity gradients to improve our understanding and accuracy of the amounts of the compounds emitted. Future studies need a multidisciplinary approach to open a new research nexus where the fields of climate science, biology and atmospheric chemistry interact.

Biological invasions have major impacts on a variety of ecosystems and threaten native biodiversity. Earthworms have been absent from northern parts of North America since the last ice age, but non-native earthworms were recently introduced there and are now being spread by human activities. While past work has shown that plant communities in earthworm-invaded areas change towards a lower diversity mainly dominated by grasses, the underlying mechanisms related to changes in the biotic interactions of the plants are not well understood. Here, we used a trait-based approach to study the effect of earthworms on interspecific plant competition and aboveground herbivory. We conducted a microcosm experiment in a growth chamber with a full-factorial design using three plant species native to northern North American deciduous forests, Poa palustris (grass), Symphyotrichum laeve (herb), and Vicia americana (legume), either growing in monoculture or in a mixture of three. These plant community treatments were crossed with earthworm (presence or absence) and herbivore (presence or absence) treatments. Eight out of the eleven above- and belowground plant functional traits studied were significantly affected by earthworms, either by a general effect or in interaction with plant species identity, plant diversity level, and/or herbivore. Earthworms increased the aboveground productivity and the number of inflorescences of the grass P. palustris. Further, earthworms countervailed the increasing effect of herbivores on root tissue density of all species, and earthworms and herbivores individually increased the average root diameter of S. laeve in monoculture, but decreased it in mixture. In this study, earthworm presence gave a competitive advantage to the grass species P. palustris by inducing changes in plant functional traits. Our results suggest that invasive earthworms can alter competitive and multitrophic interactions of plants, shedding light on some of the mechanisms behind invasive earthworm-induced plant community changes in northern North America forests.

While mycorrhization rates have been studied in different contexts, not much is known about their temporal patterns across seasons. Here, we asked how mycorrhization rates of 10 deciduous trees assessed by microscopy changed from winter to spring to early summer. We made use of a tree diversity experiment on nutrient-rich soil (former farmland) in Central Germany. In the experiment, saplings of host species with a preference for either arbuscular mycorrhiza (AM) or ectomycorrhiza (EM) were planted in monocultures, two-species, and four-species mixtures. In addition, mixtures were composed of tree species of only one mycorrhizal type or by AM/EM trees. For almost all species, with the exception of Aesculus hippocastanum and Acer pseudoplatanus (only AM), dual mycorrhization with both types (AM and EM) was found at every sampling time (December, March, and May), although the expected preferences for certain mycorrhizal types were confirmed. The sampling date had a significant influence on mycorrhization rates of both EM and AM tree species. Frequencies of EM and AM were lowest in May, but there were no differences between December and March. The causes of this seasonal variation may be associated with climate-induced differences in carbon allocation to mycorrhizal tree roots in the temperate climate. Within individual trees, mycorrhization rates by AM and EM fungi were not correlated over time, pointing to asynchronous variation between both types and to independent drivers for AM and EM mycorrhization. At the community level, mycorrhiza frequency of either of the two types became more asynchronous from two-species to four-species mixtures. Thus, increased community asynchrony in mycorrhization could be another important mechanism in biodiversity–ecosystem functioning relationships.

14 páginas, 3 figuras, 2 tablas, Research aimed at understanding the mechanisms underlying the relationship between tree diversity and antagonist infestation is often neglecting resource-use complementarity among plant species. We investigated the effects of tree species identity, species richness, and mycorrhizal type on leaf herbivory and pathogen infestation. We used a tree sapling experiment manipulating the two most common mycorrhizal types, arbuscular mycorrhiza and ectomycorrhiza, via respective tree species in monocultures and two-species mixtures. We visually assessed leaf herbivory and pathogen infestation rates, and measured concentrations of a suite of plant metabolites (amino acids, sugars, and phenolics), leaf elemental concentrations (carbon, nitrogen, and phosphorus), and tree biomass. Tree species and mycorrhizal richness had no significant effect on herbivory and pathogen infestation, whereas species identity and mycorrhizal type had. Damage rates were higher in arbuscular mycorrhizal (AM) than in ectomycorrhizal (EM) trees. Our structural equation model (SEM) indicated that elemental, but not metabolite concentrations, determined herbivory and pathogen infestation, suggesting that the investigated chemical defence strategies may not have been involved in the effects found in our study with tree saplings. Other chemical and physical defence strategies as well as species identity as its determinant may have played a more crucial role in the studied saplings. Furthermore, the SEM indicated a direct positive effect of AM trees on herbivory rates, suggesting that other dominant mechanisms, not considered here, were involved as well. We found differences in the attribution of elemental concentrations between the two rates. This points to the fact that herbivory and pathogen infestation are driven by distinct mechanisms. Our study highlights the importance of biotic contexts for understanding the mechanisms underlying the effects of biodiversity on tree-antagonist interactions, We thank Nicole M. van Dam, Henriette Uthe, Fredd Vergara, Martin Volf, and Alexander Weinhold for their valuable advice on metabolite analyses as well as Beate Rothe and Michael Reichelt for their help with chemical analyses. Moreover, comments by two anonymous reviewers helped to improve the paper. This work was supported by the European Research Council (ERC) under the European Union's Horizon 2020 research and innovation program (grant agreement no. 677232). Further support came from the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig and the EcoMetEoR platform, funded by the German Research Foundation (FZT 118). AMM acknowledges support from the program for attracting talent to Salamanca from Fundación Salamanca Ciudad de Cultura y Saberes and Ayuntamiento de Salamanca.

Elevated atmospheric CO

Tree species are known to predominantly interact either with arbuscular mycorrhizal (AM) or ectomycorrhizal (EM) fungi. However, there is a knowledge gap whether these mycorrhizae differently influence biodiversity-ecosystem functioning (BEF) relationships and whether a combination of both can increase community productivity. In 2015, we established a tree-diversity experiment by growing tree communities with varying species-richness levels (1, 2, or 4 species), and either with AM or EM tree species, or a combination of both. We investigated basal area and annual basal area increment from 2015 to 2020 as proxy for community productivity. We found significant positive relationships between tree species richness and community productivity, which strengthened over time. Further, AM and EM tree species differently influenced productivity; however, there was no overyielding when AM and EM trees grew together. EM tree communities were characterized by low productivity in the beginning, but an increase of increment over time, and showed overall strong biodiversity effects. For AM tree communities the opposite was true. While young trees did not benefit from the presence of the other mycorrhizal type, dissimilar mechanisms underlying BEF relationships in AM and EM trees indicate that maximizing tree and mycorrhizal diversity may increase ecosystem functioning in the long run.

The relationship between species’ body masses and densities is strongly conserved around a three-quarter power law when pooling data across communities. However, studies of local within-community relationships have revealed major deviations from this general pattern, which has profound implications for their stability and functioning. Despite multiple contributions of soil communities to people, there is limited knowledge on the drivers of body mass-abundance relationship in these communities. We compiled a dataset comprising 155 soil-animal communities across four countries (Canada, Germany, Indonesia, USA), all sampled using the same methodology. We tested if variation in local climatic and edaphic conditions drives differences in local body mass-abundance scaling relationships. We found substantial variation in the slopes of this power-law relationship across local communities. Structural equation modeling showed that soil temperature and water content have a positive and negative net effect, respectively, on soil communities. These effects are mediated by changes in local edaphic conditions (soil pH and carbon content) and the body-mass range of the communities. These results highlight ways in which alterations of soil climatic and edaphic conditions interactively impact the distribution of abundance, and thus energy, between populations of small and large animals. These quantitative mechanistic relationships facilitate our understanding of how global changes in environmental conditions, such as temperature and precipitation, will affect community-abundance distributions and thus the stability and functioning of soil-animal communities.

Mycorrhizal fungi play an important role for the nitrogen (N) supply of trees. The influence of different mycorrhizal types on N acquisition in tree–tree interactions is, however, not well understood, particularly with regard to the competition for growth-limiting N. We studied the effect of competition between temperate forest tree species on their inorganic and organic N acquisition in relation to their mycorrhizal type (i.e., arbuscular mycorrhiza or ectomycorrhiza). In a field experiment, we quantified net N uptake capacity from inorganic and organic N sources using 15N/13C stable isotopes for arbuscular mycorrhizal tree species (i.e., Acer pseudoplatanus L., Fraxinus excelsior L., and Prunus avium L.) as well as ectomycorrhizal tree species (i.e., Carpinus betulus L., Fagus sylvatica L., and Tilia platyphyllos Scop.). All species were grown in intra- and interspecific competition (i.e., monoculture or mixture). Our results showed that N sources were not used complementarily depending on a species’ mycorrhizal association, but their uptake rather depended on the competitor, indicating species-specific effects. Generally, ammonium was preferred over glutamine and glutamine over nitrate. In conclusion, our findings suggest that the inorganic and organic N acquisition of the studied temperate tree species is less regulated by mycorrhizal association but rather by the availability of specific N sources in the soil as well as the competitive environment of different tree species.

Anthropogenic pressures alter the biodiversity, structure and organization of biological communities with severe consequences for ecosystem processes. Species invasion is such a human-induced ecosystem change with pronounced impacts on recipient ecosystems. Around the globe, earthworms invade habitats and impact abiotic soil conditions and a wide range of above- and belowground organisms. In northern North America, where earthworms have been largely absent since the last glaciation period and most earthworm species present today have only been (re-)introduced a few hundred years ago, invasion impacts have been intensively studied. However, despite several studies assessing impacts of invasive earthworms on soil fauna, studies have rarely investigated the simultaneous responses of different soil-fauna size groups and biodiversity facets which might respond differently to earthworm invasion and independently affect ecosystem processes. Our study goes beyond previously-established knowledge on earthworm-invasion effects by simultaneously assessing differences in four biodiversity facets, namely the abundance, biomass, richness and Shannon index of soil invertebrate macro-, meso- and microfauna communities between high- and low-invasion status plots (n = 80) and in relation to invasion intensity measured as earthworm biomass across four northern North American forests sampled between 2016 and 2017. Across forests and soil-fauna groups, we found reduced abundance (−33 to −45%) and richness (−18 to −25%) in high compared to low-invasion status areas. Additionally, meso- (−14%) and microfauna biomass (−38%) and macro- (−7%) and microfauna Shannon index (−8%) were reduced. Higher invasion intensity (earthworm biomass) was additionally related to reduced soil-fauna biodiversity. While the studied biodiversity facet was important for the soil fauna response, soil-fauna size group was comparably unimportant. Given the global ubiquity of earthworm invasion and the importance of soil fauna for key ecosystem processes, our observational results help to assess future impacts of this invasion and the consequences for anthropogenically-altered ecosystem functioning.

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 approximately 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 and high) in a Canadian forest. We analysed 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 approximately 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 suggest that earthworms directly and indirectly impact arthropods across trophic levels. We show that earthworm invasion can alter aboveground multi-trophic arthropod communities and suggest that belowground invasions might be underappreciated drivers of aboveground arthropod decline.

Natural forest is declining globally as the area of planted forest increases. Planted forests are often monocultures, despite results suggesting that higher species richness improves ecosystem functioning and stability. To test if this is generally the case, we performed a meta-analysis of available results. We assessed aboveground carbon stocks in mixed-species planted forests vs (a) the average of constituent species monocultures, (b) the best constituent species monoculture, and (c) commercial species monocultures. We investigated whether any advantage of mixtures over monocultures was positively related to species richness, as well as potential mechanisms driving differences in carbon stocks between mixtures and monocultures. The meta-analysis dataset included 79 comparisons from 21 sites. Carbon stocks in mixed planted forests were higher than the average of stocks in monocultures of their constituent species, containing on average 70% more carbon. Mixed planted forests also out-performed commercial monocultures, containing on average 77% more carbon. There was c.25% more carbon in mixed planted forests relative to the best performing monocultures, although this difference was not statistically significant. Overyielding was highest in four-species mixtures (richness range 2-6 species). More data providing better coverage of richness and age gradients (study sites aged 3.5-28 years) is needed to increase confidence in these results. None of the potential mechanisms we examined (nitrogen-fixer present vs absent; native vs non-native/mixed origin; tree diversity experiment vs forestry plantation) consistently explained variation in the diversity effects. This suggests that our findings are driven by a combination of small (statistically insignificant) effects from these sources or further unidentified mechanisms or some combination of the two. We conclude that increasing tree species richness in planted forests can increase carbon stocks while bringing other potential benefits associated with diversification. However, implementation will depend on the balance of these benefits relative to the operational challenges and costs of diversification.

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.

Plant diversity and plant-related ecosystem functions have been important in biodiversity-ecosystem functioning studies. However, biotic interactions with mycorrhizal fungi have been understudied although they are crucial for plant-resource acquisition. Here, we investigated the effects of tree species richness and 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 or different mycorrhizal type. We sampled a tree diversity 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 drivers of mycorrhizal fungal communities and further highlights the distinct strategies of AMF and EMF.

Bottom-up effects of plants on soil communities can be modified by the activity of exotic earthworms, by altering resource availability for soil food webs through feeding, burrowing, and casting activities. The present study explored effects of plants (planting of shrubs) on soil micro-food webs (composition of soil microbial and nematode communities), and whether these effects were altered by the activity of exotic earthworms (exotic earthworms addition). Planted shrubs resulted in a non-significant increase of bacterial biomass and significantly increased the abundance of different nematode trophic groups and total nematode biomass, indicating that planted shrubs had significant bottom-up effects on soil bacteria and nematodes. Planted shrubs decreased nematode diversity, evenness, and richness, but increased nematode dominance in the plots where the abundance of exotic earthworms was not amended. By contrast, these effects of shrub presence on soil biodiversity were not found in the plots that received exotic earthworms. In addition, planted shrubs increased the total energy flux to the nematode community. By contrast, the elevated activity of exotic earthworms mitigated the increase in total energy flux to nematodes in the presence of shrubs, and increased the ratio of fungal to bacterial PLFAs. Both of these changes indicate reduced energy flux in the plots with added exotic earthworms. Nematode diversity decreased, while nematode dominance increased with increasing total energy flux to nematodes, probably because few species benefited from high energy flux. Our study indicates that exotic earthworms can maintain soil biodiversity by reducing the energy flux through soil food webs.

Recent research shows that earthworms can alter defense traits of plants against herbivores and pathogens by affecting soil biochemistry. Yet, the effects of invasive earthworms on defense traits of native plants from previously earthworm‐free ecosystems as well as the consequences for multitrophic interactions are virtually unknown.Here we use a combination of an observational study and a complementary experimental study to investigate the effects of invasive earthworms on leaf defense traits, herbivore damage and pathogen infection in two poplar tree species (Populus balsamifera and Populus tremuloides) native to North American boreal forests.Our observational study showed that earthworm invasion was associated with enhanced leaf herbivory (by leaf‐chewing insects) in saplings of both tree species. However, we only detected significant shifts in the concentration of chemical defense compounds in response to earthworm invasion for P. balsamifera. Specifically, leaf phenolic concentrations, including salicinoids and catechin, were lower in P. balsamifera from earthworm‐invaded sites.Our experimental study confirmed an earthworm‐induced reduction in leaf defense levels in P. balsamifera for one of the defense compounds, tremulacin. The experimental study additionally showed that invasive earthworms reduced leaf dry matter content, potentially increasing leaf palatability, and enhanced susceptibility of trees to infection by a fungal pathogen, but not to aphid infestation, in the same tree species. Synthesis. Our results show that invasive earthworms can decrease the concentrations of some chemical defense compounds in P. balsamifera, which could make them susceptible to leaf‐chewing insects. Such potential impacts of invasive earthworms are likely to have implications for tree survival and competition, native tree biodiversity and ecosystem functioning., Invasive earthworms can dramatically alter native biodiversity and ecosystem functioning. Yet, we know little how invasive earthworms affect the chemical defense of native trees. Here using a combination of observational and experimental studies, we show that invasive earthworms from Europe can decrease the concentrations of some chemical defense compounds in a native North American deciduous tree, which could make them susceptible to leaf‐chewing insects.

Plant diversity has a strong impact on a plethora of ecosystem functions and services, especially ecosystem carbon (C) storage. However, the potential context-dependency of biodiversity effects across ecosystem types, environmental conditions and carbon pools remains largely unknown. In this study, we performed a meta-analysis by collecting data from 95 biodiversity-ecosystem functioning (BEF) studies across 60 sites to explore the effects of plant diversity on different C pools, including aboveground and belowground plant biomass, soil microbial biomass C and soil C content across different ecosystem types. The results showed that ecosystem C storage was significantly enhanced by plant diversity, with stronger effects on aboveground biomass than on soil C content. Moreover, the response magnitudes of ecosystem C storage increased with the level of species richness and experimental duration across all ecosystems. The effects of plant diversity were more pronounced in grasslands than in forests. Furthermore, the effects of plant diversity on belowground plant biomass increased with aridity index in grasslands and forests, suggesting that climate change might modulate biodiversity effects, which are stronger under wetter conditions but weaker under more arid conditions. Taken together, these results provide novel insights into the important role of plant diversity in ecosystem C storage across critical C pools, ecosystem types and environmental contexts.

Recent studies found that the majority of shrub and tree species are associated with both arbuscular mycorrhizal (AM) and ectomycorrhizal (EM) fungi. However, our knowledge on how different mycorrhizal types interact with each other is still limited. We asked whether the combination of hosts with a preferred association with either AM or EM fungi increases the host tree roots’ mycorrhization rate and affects AM and EM fungal richness and community composition.We established a tree diversity experiment, where five tree species of each of the two mycorrhiza types were planted in monocultures, two‐species and four‐species mixtures. We applied morphological assessment to estimate mycorrhization rates and next‐generation molecular sequencing to quantify mycobiont richness.Both the morphological and molecular assessment revealed dual‐mycorrhizal colonization in 79% and 100% of the samples, respectively. OTU community composition strongly differed between AM and EM trees. While host tree species richness did not affect mycorrhization rates, we observed significant effects of mixing AM‐ and EM‐associated hosts in AM mycorrhization rate. Glomeromycota richness was larger in monotypic AM tree combinations than in AM‐EM mixtures, pointing to a dilution or suppression effect of AM by EM trees. We found a strong match between morphological quantification of AM mycorrhization rate and Glomeromycota richness. Synthesis. We provide evidence that the combination of hosts differing in their preferred mycorrhiza association affects the host's fungal community composition, thus revealing important biotic interactions among trees and their associated fungi., In a tree diversity experiment where different mycorrhizal host species were planted in monospecific stands and in mixtures, we found that combining arbuscular or ectotrophic mycorrhizae affect mycorrhization rates and richness of AM and EM fungi. Glomeromycota richness was larger in monotypic AM tree combinations than in mixtures of AM‐EM host trees, pointing to a suppression effect of AM fungi by the presence of EM trees.

12 páginas. 4 figuras.- 2 tablas.- 70 referencias.- Additional supporting information may be found in the online version of this article at http://onlinelibrary.wiley.com/doi/ 10.1002/ecy.2936/suppinfo .- Data are available from the Dryad Digital Repository: https://doi.org/10.5061/dryad.59zw3r23d, Recent studies have shown that invasive earthworms can dramatically reduce native biodiversity, both above and below the ground. However, we still lack a synthetic understanding of the underlying mechanisms behind these changes, such as whether earthworm effects on soil chemical properties drive such relationships. Here, we investigated the effects of invasive earthworms on soil chemical properties (pH, water content, and the stocks and fluxes of carbon, nitrogen, and phosphorus) by conducting a meta‐analysis. Invasive earthworms generally increased soil pH, indicating that the removal of organic layers and the upward transport of more base‐rich mineral soil caused a shift in soil pH. Moreover, earthworms significantly decreased soil water content, suggesting that the burrowing activities of earthworms may have increased water infiltration of and/or increased evapotranspiration from soil. Notably, invasive earthworms had opposing effects on organic and mineral soil for carbon and nitrogen stocks, with decreases in organic, and increases in mineral soil. Nitrogen fluxes were higher in mineral soil, whereas fluxes in organic soil were not significantly affected by the presence of invasive earthworms, indicating that earthworms mobilize and redistribute nutrients among soil layers and increase overall nitrogen loss from the soil. Invasive earthworm effects on element stocks increased with ecological group richness only in organic soil. Earthworms further decreased ammonium stocks with negligible effects on nitrate stocks in organic soil, whereas they increased nitrate stocks but not ammonium stocks in mineral soil. Notably, all of these results were consistent across forest and grassland ecosystems underlining the generality of our findings. However, we found some significant differences between studies that were conducted in the field (observational and experimental settings) and in the lab, such as that the effects on soil pH decreased from field to lab settings, calling for a careful interpretation of lab findings. Our meta‐analysis provides strong empirical evidence that earthworm invasion may lead to substantial changes in soil chemical properties and element cycling in soil. Furthermore, our results can help explain the dramatic effects of invasive earthworms on native biodiversity, for example, shifts towards the dominance of grass species over herbaceous ones, as shown by recent meta‐analyses., This project received support from the European Research Council (ERC) under the European Union’s Horizon 2020 research and innovation program (grant agreement 677232). Further support came from the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, funded by the German Research Foundation (FZT 118).

Associational resistance theory predicts that insect herbivory decreases with increasing tree diversity in forest ecosystems. However, the generality of this effect and its underlying mechanisms are still debated, particularly since evidence has accumulated that climate may influence the direction and strength of the relationship between diversity and herbivory.We quantified insect leaf herbivory and leaf chemical defences (phenolic compounds) of silver birch Betula pendula in pure and mixed plots with different tree species composition across 12 tree diversity experiments in different climates. We investigated whether the effects of neighbouring tree species diversity on insect herbivory in birch, that is, associational effects, were dependent on the climatic context, and whether neighbour-induced changes in birch chemical defences were involved in associational resistance to insect herbivory.We showed that herbivory on birch decreased with tree species richness (i.e. associational resistance) in colder environments but that this relationship faded as mean annual temperature increased.Birch leaf chemical defences increased with tree species richness but decreased with the phylogenetic distinctiveness of birch from its neighbours, particularly in warmer and more humid environments.Herbivory was negatively correlated with leaf chemical defences, particularly when birch was associated with closely related species. The interactive effect of tree diversity and climate on herbivory was partially mediated by changes in leaf chemical defences.Our findings confirm that tree species diversity can modify the leaf chemistry of a focal species, hence its quality for herbivores. They further stress that such neighbour-induced changes are dependent on climate and that tree diversity effects on insect herbivory are partially mediated by these neighbour-induced changes in chemical defences. ispartof: FUNCTIONAL ECOLOGY vol:35 issue:1 pages:67-81 ispartof: location:England status: published

Research on mycorrhizal interactions has traditionally developed into separate disciplines addressing different organizational levels. This separation has led to an incomplete understanding of mycorrhizal functioning. Integration of mycorrhiza research at different scales is needed to understand the mechanisms underlying the context dependency of mycorrhizal associations, and to use mycorrhizae for solving environmental issues. Here, we provide a road map for the integration of mycorrhiza research into a unique framework that spans genes to ecosystems. Using two key topics, we identify parallels in mycorrhiza research at different organizational levels. Based on two current projects, we show how scientific integration creates synergies, and discuss future directions. Only by overcoming disciplinary boundaries, we will achieve a more comprehensive understanding of the functioning of mycorrhizal associations., All authors acknowledge funding from the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig funded by the German Research Foundation (DFG;FZT118) .A.M.M.,A.B.,M.P.,S.R., and N.M.v.D.acknowledge stimulating discussions within COSTActionFA1405 and funding from EUMC-ITNMiRA(grantno.765290).I.C.M. acknowledges financial supportfromtheDFG(grantno.ME4156/2-1)and Volkswagen Foundation [grantno.11-76251- 99-34/13 (ZN2928)].M.C.R.acknowledgesfundingfromtheEuropeanResearchCouncil(ERC,AdvancedGrant ‘Gradual Change’) andBMBFfundingfortheproject ‘Bridging inBiodiversityScience(BIBS)’. M.P.acknowledgessupportfrom MINECO (grantno.AGL2015-64990-C2-1-R).I.F.acknowledgessupportfromiDivFlexpoolProgram(grantno.RA-185/ 17). O.F.andN.E.acknowledgesupportfromtheERC(EuropeanUnion’s Horizon2020researchandinnovationprogram, grant no.677232).S.R.acknowledgestheSwissScienceFoundation(grantno.159869).F.B.,S.H.,andM.T.acknowl- edge DFGfor financial support(GrantsBU941/20-1andTA290/4-1).AlltheauthorsacknowledgesupportfromtheiDiv Open SciencePublicationFund.

Along with the global decline of species richness goes a loss of ecological traits. Associated biotic homogenization of animal communities and narrowing of trait diversity threaten ecosystem functioning and human well-being. High management intensity is regarded as an important ecological filter, eliminating species that lack suitable adaptations. Below-ground arthropods are assumed to be less sensitive to such effects than above-ground arthropods. Here, we compared the impact of management intensity between (grassland vs. forest) and within land-use types (local management intensity) on the trait diversity and composition in below- and above-ground arthropod communities. We used data on 722 arthropod species living above-ground (Auchenorrhyncha and Heteroptera), primarily in soil (Chilopoda and Oribatida) or at the interface (Araneae and Carabidae). Our results show that trait diversity of arthropod communities is not primarily reduced by intense local land use, but is rather affected by differences between land-use types. Communities of Auchenorrhyncha and Chilopoda had significantly lower trait diversity in grassland habitats as compared to forests. Carabidae showed the opposite pattern with higher trait diversity in grasslands. Grasslands had a lower proportion of large Auchenorrhyncha and Carabidae individuals, whereas Chilopoda and Heteroptera individuals were larger in grasslands. Body size decreased with land-use intensity across taxa, but only in grasslands. The proportion of individuals with low mobility declined with land-use intensity in Araneae and Auchenorrhyncha, but increased in Chilopoda and grassland Heteroptera. The proportion of carnivorous individuals increased with land-use intensity in Heteroptera in forests and in Oribatida and Carabidae in grasslands. Our results suggest that gradients in management intensity across land-use types will not generally reduce trait diversity in multiple taxa, but will exert strong trait filtering within individual taxa. The observed patterns for trait filtering in individual taxa are not related to major classifications into above- and below-ground species. Instead, ecologically different taxa resembled each other in their trait diversity and compositional responses to land-use differences. These previously undescribed patterns offer an opportunity to develop management strategies for the conservation of trait diversity across taxonomic groups in permanent grassland and forest habitats.

Soil organisms, including earthworms, are a key component of terrestrial ecosystems. However, little is known about their diversity, their distribution, and the threats affecting them. We compiled a global dataset of sampled earthworm communities from 6928 sites in 57 countries as a basis for predicting patterns in earthworm diversity, abundance, and biomass. We found that local species richness and abundance typically peaked at higher latitudes, displaying patterns opposite to those observed in aboveground organisms. However, high species dissimilarity across tropical locations may cause diversity across the entirety of the tropics to be higher than elsewhere. Climate variables were found to be more important in shaping earthworm communities than soil properties or habitat cover. These findings suggest that climate change may have serious implications for earthworm communities and for the functions they provide., 677232 to N.E.). K.S.R. and W.H.v.d.P. were supported by ERC-ADV grant 323020 to W.H.v.d.P. Also supported by iDiv (DFG FZT118) Flexpool proposal 34600850 (C.A.G. and N.E.); the Academy of Finland (285882) and the Natural Sciences and Engineering Research Council of Canada (postdoctoral fellowship and RGPIN-2019-05758) (E.K.C.); DOB Ecology (T.W.C., J.v.d.H., and D.R.); ERC-AdG 694368 (M.R.); and the TULIP Laboratory of Excellence (ANR-10-LABX-41) (M.L.). In addition, data collection was funded by the Russian Foundation for Basic Research (12-04-01538-a, 12-04-01734-a, 14-44-03666-r_center_a, 15-29-02724-ofi_m, 16-04-01878-a 19-05-00245); Tarbiat Modares University; Aurora Organic Dairy; UGC(NERO) (F. 1-6/Acctt./NERO/2007-08/1485); Natural Sciences and Engineering Research Council (RGPIN-2017-05391); Slovak Research and Development Agency (APVV-0098-12); Science for Global Development through Wageningen University; Norman Borlaug LEAP Programme and International Atomic Energy Agency (IAEA); São Paulo Research Foundation - FAPESP (12/22510-8); Oklahoma Agricultural Experiment Station; INIA - Spanish Agency (SUM 2006-00012-00-0); Royal Canadian Geographical Society; Environmental Protection Agency (Ireland) (2005-S-LS-8); University of Hawai‘i at Mānoa (HAW01127H; HAW01123M); European Union FP7 (FunDivEurope, 265171); U.S. Department of the Navy, Commander Pacific Fleet (W9126G-13-2-0047); Science and Engineering Research Board (SB/SO/AS-030/2013) Department of Science and Technology, New Delhi, India; Strategic Environmental Research and Development Program (SERDP) of the U.S. Department of Defense (RC-1542); Maranhão State Research Foundation (FAPEMA); Coordination for the Improvement of Higher Education Personnel (CAPES); Ministry of Education, Youth and Sports of the Czech Republic (LTT17033); Colorado Wheat Research Foundation; Zone Atelier Alpes, French National Research Agency (ANR-11-BSV7-020-01, ANR-09-STRA-02-01, ANR 06 BIODIV 009-01); Austrian Science Fund (P16027, T441); Landwirtschaftliche Rentenbank Frankfurt am Main; Welsh Government and the European Agricultural Fund for Rural Development (Project Ref. A AAB 62 03 qA731606); SÉPAQ; Ministry of Agriculture and Forestry of Finland; Science Foundation Ireland (EEB0061); University of Toronto (Faculty of Forestry); National Science and Engineering Research Council of Canada; Haliburton Forest and Wildlife Reserve; NKU College of Arts and Sciences Grant; Österreichische Forschungsförderungsgesellschaft (837393 and 837426); Mountain Agriculture Research Unit of the University of Innsbruck; Higher Education Commission of Pakistan; Kerala Forest Research Institute, Peechi, Kerala; UNEP/GEF/TSBF-CIAT Project on Conservation and Sustainable Management of Belowground Biodiversity; Ministry of Agriculture and Forestry of Finland; Complutense University of Madrid/European Union FP7 project BioBio (FPU UCM 613520); GRDC; AWI; LWRRDC; DRDC; CONICET (National Scientific and Technical Research Council) and FONCyT (National Agency of Scientific and Technological Promotion) (PICT, PAE, PIP), Universidad Nacional de Luján y FONCyT [PICT 2293 (2006)], Fonds de recherche sur la nature et les technologies du Québec (131894), Deutsche Forschungsgemeinschaft [SCHR1000/3-1, SCHR1000/6-1, 6-2 (FOR 1598), WO 670/7-1, WO 670/7-2, and SCHA 1719/1-2], CONACYT (FONDOS MIXTOS TABASCO/PROYECTO11316); NSF (DGE-0549245, DGE-0549245, DEB-BE-0909452, NSF1241932); Institute for Environmental Science and Policy at the University of Illinois at Chicago; Dean’s Scholar Program at UIC; Garden Club of America Zone VI Fellowship in Urban Forestry from the Casey Tree Endowment Fund; J. E. Weaver Competitive Grant from the Nebraska Chapter of The Nature Conservancy; the College of Liberal Arts and Sciences at DePaul University; Elmore Hadley Award for Research in Ecology and Evolution from the UIC Dept. of Biological Sciences; Spanish CICYT (AMB96-1161; REN2000-0783/GLO; REN2003-05553/GLO; REN2003-03989/GLO; CGL2007-60661/BOS); Yokohama National University; MEXT KAKENHI (25220104); Japan Society for the Promotion of Science KAKENHI (25281053, 17KT0074, 25252026); ADEME (0775C0035); Ministry of Science, Innovation and Universities of Spain (CGL2017-86926-P); Syngenta Philippines; UPSTREAM; LTSER (Val Mazia/Matschertal); Marie Sklodowska Curie Postdoctoral Fellowship (747607); National Science and Technology Base Resource Survey Project of China (2018FY100306); McKnight Foundation (14-168); Program of Fundamental Researches of Presidium of Russian Academy of Sciences (AAAA-A18-118021490070-5); Brazilian National Council of Research CNPq; and French Ministry of Foreign and European Affairs. Author contributions: H.R.P.P. led the analysis, data curation, and writing of the original manuscript draft. C.A.G. assisted in analyses and writing of the original manuscript draft. E.K.C. and N.E. revised subsequent manuscript drafts. J.v.d.H., D.R., and T.W.C. provided additional analyses. E.K.C., N.E., and M.P.T. acquired funding for the project. J.K., K.B.G., B.S., M.L.C.B., M.J.I.B., and G.B. contributed to data curation. H.R.P.P., C.A.G., M.L.C.B., M.J.I.B., G.B., O.F., A.O., E.M.B., J.B., U.B., T.D., F.T.d.V., B.K.-R., M.L., J.M., C.M., W.H.v.d.P., K.S.R., M.C.R., D.R., M.R., M.P.T., D.H.W., D.A.W., E.K.C., and N.E. contributed to the project conceptualization. All authors reviewed and edited the final draft manuscript. The majority of the authors provided data for the analyses. Competing interests: The authors declare no competing interests. Data and materials availability: Data and analysis code are available on the iDiv Data repository (DOI: 10.25829/idiv.1804-5-2593) and GitHub (https://github.com/helenphillips/GlobalEWDiversity; DOI: 10.5281/zenodo.3386456).