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Abstract Biodiversity experiments revealed that plant diversity loss can decrease ecosystem functions across trophic levels. To address why such biodiversity-function relationships strengthen over time, we established experimental mesocosms replicating a gradient in plant species richness across treatments of shared versus non-shared history of (1) the plant community and (2) the soil fauna community. After 4 months, we assessed the multitrophic functioning of soil fauna via biomass stocks and energy fluxes across the food webs. We find that soil community history significantly enhanced belowground multitrophic function via changes in biomass stocks and community-average body masses across the food webs. However, variation in plant diversity and plant community history had unclear effects. Our findings underscore the importance of long-term community assembly processes for soil fauna-driven ecosystem function, with species richness and short-term plant adaptations playing a minimal role. Disturbances that disrupt soil community stability may hinder fauna-driven ecosystem functions, while recovery may require several years.

Long-term research in grassland biodiversity experiments has provided empirical evidence that ecological and evolutionary processes are intertwined in determining both biodiversity–ecosystem functioning (BEF) and biodiversity–stability relationships. Focusing on plant diversity, we hypothesize that multifunctional stability is highest in high-diversity plant communities and that biodiversity–stability relationships increase over time due to a variety of forms of ecological complementarity including the interaction with other biota above and below ground. We introduce the multiple-mechanisms hypothesis of biodiversity–stability relationships suggesting that it is not an individual mechanism that drives long-term biodiversity effects on ecosystem functioning and stability but that several intertwined processes produce increasingly positive ecosystem effects. The following six mechanisms are important. Low-diversity plant communities accumulate more plant antagonists over time (1), and use resources less efficiently and have more open, leaky nutrient cycles (2). Conversely, high-diversity plant communities support a greater diversity and activity of beneficial interaction partners across trophic levels (3); diversify in their traits over time and space, within and across species, to optimize temporal (intra- and interannual) and spatial complementarity (4), create a more stable microclimate (5), and foster higher top-down control of aboveground and belowground herbivores by predators (6). In line with the observation that different species play unique roles in ecosystems that are dynamic and multifaceted, the particular mechanism contributing most to the higher performance and stability of diverse plant communities might differ across ecosystem functions, years, locations, and environmental change scenarios. This indicates “between-context insurance” or “across-context complementarity” of different mechanisms. We introduce examples of experiments that will be conducted to test our hypotheses and which might inspire additional work.

Soil biodiversity and the structure of soil animal communities are important foundations for forest ecosystem functions. Forest gap formation is an important forest management practice used to transform monocultures into mixed forests. However, whether and how gap size and age affect soil biodiversity and modify nematode communities remains limited. We manipulated gap size (100, 200, and 400 m2) in Pinus massoniana plantations and studied the communities of soil nematodes, bacteria, fungi, and understory plants two and four years after gap formation. Compared to the no-gap treatment, soil nematode abundance across forest gaps increased by a factor of 1.40, which was largely attributed to the increase in herbivorous nematodes as the abundance and diversity of understory plants increased. The increased abundance of soil nematodes in forest gaps was also associated with increased soil pH presumably related to reduced input of pine needles. Furthermore, the abundance (−5.3 %) and diversity (−25.1 %) of soil nematodes decreased with gap age, presumably because of increased soil temperature and decreased soil moisture in the four- compared to the two-year-old gaps. In contrast to nematodes, the abundance and diversity of soil bacteria (21.8 % and 7.1 %) and fungi (10.5 % and 10.0 %) increased significantly with gap age. Overall, forest gaps increased the diversity of understory plants and soil biota, and changed the community and functional group structure of soil nematodes. These results provide guidelines for fostering soil biodiversity and maintaining soil functioning when transforming coniferous forests into mixed forests.

Abstract The ongoing climate change calls for managing forest ecosystems in temperate regions toward more drought‐resistant and climate‐resilient stands. Yet ecological consequences of management options such as planting non‐native tree species and mixing coniferous and deciduous tree species have been little studied, especially on soil animal communities, key in litter decomposition and pest control. Here, we investigated the taxonomic and trophic structure of soil macrofauna communities in five forest types including native European beech (Fagus sylvatica), range‐expanding Norway spruce (Picea abies) and non‐native Douglas fir (Pseudotsuga menziesii) as well as conifer‐beech mixtures across loamy and sandy sites in northern Germany. Abundance of primary decomposers (feeding predominantly on litter) was high in Douglas fir and beech forests, benefiting from less acidic soil and more favorable litter resources compared to spruce forests, while secondary decomposers (feeding predominantly on microorganisms and microbial residues) reached highest densities in spruce forests. Differences in abundance and species richness among forest types generally varied between regions and were most pronounced in Douglas fir of the sandy region. However, trophic guilds differed more between regions than between forest types, indicating that environmental factors outweigh the importance of forest type on soil macrofauna communities. The analysis of stable isotopes (δ15N and δ13C values) supported the general robustness in trophic position of macrofauna trophic guilds against variations in forest types and regions, but indicated reduced detrital shifts and food‐chain lengths in coniferous compared to European beech forests with mixtures mitigating these effects. Overall, for evaluating consequences of future forest management practices on the structure and functioning of soil animal communities, regional factors need to be considered, but in particular at loamy sites the taxonomic and trophic structure of soil macrofauna communities are resistant against changes in forest types.

Abstract Soils, just like all other ecosystem compartments, change over time and, consequently, conditions for soil‐inhabiting organisms are also changing, affecting their composition and diversity. Soil biodiversity is a critical component of ecosystems that supports many essential ecosystem functions and services, such as nutrient cycling, carbon sequestration, water regulation and biomass production for food, fodder, fibre and energy. However, and despite the importance of soil biodiversity for ecosystem health and human well‐being, neither current state, drivers, potential consequences for ecosystem services nor options for sustainable governance of soil biodiversity are well understood. Here, we provide a framework for and argue that conducting a national assessment of soil biodiversity, albeit being a complex endeavour, is fundamental to building a baseline to understand the current state and trends of soil biodiversity, but also to identify the main drivers of change, the impacts of soil biodiversity loss and the potential pathways for conservation and sustainable governance of soil biodiversity.

Large tropical trees are rightly perceived as supporting a plethora of organisms. However, baseline data about the variety of taxa coexisting on single large tropical trees are lacking and prevent a full understanding of both the magnitude of biodiversity and the complexity of interactions among organisms in tropical rainforests. The two main aims of the research program “Life on Trees” (LOT) are (1) to establish baseline knowledge on the number of eukaryote species supported/hosted by the above-ground part of a single tropical tree and (2) to understand how these communities of organisms are assembled and distributed on or inside the tree. To achieve the first goal, we integrated a set of 36 methods for comprehensively sampling eukaryotes (plants, fungi, animals, protists) present on a tropical tree. The resulting LOT protocol was conceived and implemented during projects in the Andean Amazon region and is proposed here as a guideline for future projects of a similar nature. To address the second objective, we evaluated the microclimatic differences between tree zones and tested state-of-the-art terrestrial laser scanning (TLS) and positioning technologies incorporating satellite and fixed base station signals (dGNSS). A marked variation in temperature and relative humidity was detected along a 6-zones Johansson scheme, a tree structure subdivision system commonly used to study the stratification of epiphytic plants. Samples were collected from these six zones, including three along the trunk and three in the canopy. To better understand how different tree components (e.g., bark, leaves, fruits, flowers, dead wood) contribute to overall tree biodiversity, we categorized observations into communities based on Johansson zones and microhabitats. TLS was an essential aid in understanding the complex tree architecture. By contrast, the accuracy of positioning samples in the tree with dGNSS was low. Comprehensively sampling the biota of individual trees offers an alternative to assessing the biodiversity of fewer groups of organisms at the forest scale. Large old tropical trees provide a wealth of microhabitats that encompass a wide range of ecological conditions, thereby capturing a broad spectrum of biodiversity.

Biodiversity loss and its potential threat on ecosystem functions call for a critical evaluation of human impacts on forest ecosystems. Management practices based on stand diversification offer a possible solution to biodiversity loss due to monoculture plantations, and these practices often involve planting introduced tree species. Although introduced non-native tree species may provide high economic returns, they may also form novel ecosystems and threaten local biodiversity, but this has been little studied. Here, we combined a taxonomic and trait-based approach and investigated communities of oribatid mites (Oribatida, Acari) across forest types of both native and introduced tree species in Northern Germany. Both trophic and life-history traits of oribatid mites were evaluated using native European beech (Fagus sylvatica) as reference, compared to native Norway spruce (Picea abies), introduced Douglas fir (Pseudotsuga menziesii) and beech-conifer mixtures. The abundance and diversity of oribatid mites were generally similar among monocultures of European beech, Norway spruce and Douglas fir. By contrast, species and trait compositions of oribatid mite communities were shifted to include more primary decomposers and more surface-living oribatid mites in Douglas fir, resulting in a trophic reorganization with less predators than in European beech forests. These results suggest that oribatid mites maintain a similar level of trait diversity regardless of forest type, but the changes in trophic guild composition and vertical distribution indicate greater availability of litter-based resources in Douglas fir than in European beech forests. The similar abundance and diversity of oribatid mite communities in Douglas fir mixed stands as in native European beech forests points to mixed forests as a promising management option for future forestry. Overall, our trait-based analyses provided insights into the changes of soil biota composition, revealing the impact of introduced tree species on the structure and functions of soil animal communities.

Belowground invertebrate communities are dominated by species-rich and very small microarthropods that require long handling times and high taxonomic expertise for species determination. Molecular based methods like metabarcoding circumvent the morphological determination process by assigning taxa bioinformatically based on sequence information. The potential to analyse diverse and cryptic communities in short time at high taxonomic resolution is promising. However, metabarcoding studies revealed that taxonomic assignment below family-level in Collembola (Hexapoda) and Oribatida (Acariformes) is difficult and often fails. These are the most abundant and species-rich soil-living microarthropods, and the application of molecular-based, automated species determination would be most beneficial in these taxa. In this study, we analysed the presence of a barcoding gap in the standard barcoding gene cytochrome oxidase I (COI) in Collembola and Oribatida. The barcoding gap describes a significant difference between intra- and interspecific genetic distances among taxa and is essential for bioinformatic taxa assignment. We collected COI sequences of Collembola and Oribatida from BOLD and NCBI and focused on species with a wide geographic sampling to capture the range of their intraspecific variance. Our results show that intra- and interspecific genetic distances in COI overlapped in most species, impeding accurate assignment. When a barcoding gap was present, it exceeded the standard threshold of 3% intraspecific distances and also differed between species. Automatic specimen assignments also showed that most species comprised of multiple genetic lineages that caused ambiguous taxon assignments in distance-based methods. Character-based taxonomic assignment using phylogenetic trees and monophyletic clades as criteria worked for some species of Oribatida but failed completely for Collembola. Notably, parthenogenetic species showed lower genetic variance in COI and more accurate species assignment than sexual species. The different patterns in genetic diversity among species suggest that the different degrees of genetic variance result from deep evolutionary distances. This indicates that a single genetic threshold, or a single standard gene, will probably not be sufficient for the molecular species identification of many Collembola and Oribatida taxa. Our results also show that haplotype diversity in some of the investigated taxa was not even nearly covered, but coverage was better for Collembola than for Oribatida. Additional use of secondary barcoding genes and long-read sequencing of marker genes can improve metabarcoding studies. We also recommend the construction of pan-genomes and pan-barcodes of species lacking a barcoding gap. This will allow both to identify species boundaries, and to cover the full range of variability in the marker genes, making molecular identification also possible for species with highly diverse barcode sequences.

Abstract Springtails (Collembola) inhabit soils from the Arctic to the Antarctic and comprise an estimated ~32% of all terrestrial arthropods on Earth. Here, we present a global, spatially-explicit database on springtail communities that includes 249,912 occurrences from 44,999 samples and 2,990 sites. These data are mainly raw sample-level records at the species level collected predominantly from private archives of the authors that were quality-controlled and taxonomically-standardised. Despite covering all continents, most of the sample-level data come from the European continent (82.5% of all samples) and represent four habitats: woodlands (57.4%), grasslands (14.0%), agrosystems (13.7%) and scrublands (9.0%). We included sampling by soil layers, and across seasons and years, representing temporal and spatial within-site variation in springtail communities. We also provided data use and sharing guidelines and R code to facilitate the use of the database by other researchers. This data paper describes a static version of the database at the publication date, but the database will be further expanded to include underrepresented regions and linked with trait data.

Maintaining a balance between growth needs and available food resources is critical to the development of any organism. Ecological stoichiometry provides a theoretical basis for studying stoichiometric mismatches between organisms and their food resources. Recent studies have shown that detritivore taxa occupying different multidimensional stoichiometric niches (MSNs) exhibit different stoichiometric mismatch patterns. However, changes in stoichiometric mismatch patterns of different functional groups within the same taxon in response to changes in food resources are poorly understood. Here, we investigated changes in the stoichiometric mismatch between litter as major food resource and five functional groups of soil detritivores (millipedes; Diplopoda) for eleven elements along an altitudinal gradient from 800 to 1850 m on Changbai Mountain, Northeast China. We quantified MSNs and multidimensional stoichiometric mismatch patterns, and investigated changes with altitude and relationships with body size and population density. The concentration of chemical elements of litter varied significantly with altitude and this also was true for the trophic stoichiometric mismatch in millipedes. Furthermore, millipede stoichiometry homeostasis resulted in changing stoichiometric mismatches with changing litter element ratios. Element concentrations in litter, specifically N, Ca, P, Na and Cu, impacted the body size and population density of millipedes, with different elements correlating with individual functional groups. The results indicate that consumer-resource stoichiometric mismatches significantly affect detritivore populations, highlighting the predictive power of the multidimensional stoichiometry framework for detritivore community composition in context of environmental change.

Abstract Forest soils are a critical component of terrestrial ecosystems and host a large number of animal decomposer species. One diverse and abundant decomposer taxon is oribatid mites (Acari: Oribatida), whose species composition varies with forest type and tree species composition. We used functional traits that indicate different niche dimensions, to infer assembly processes of oribatid mite communities in monocultures and mixed forests of native and introduced tree species. We found that coexisting species differed more in the resource‐related niche dimension, i.e., reproductive mode and trophic guild, than in the morphological dimension, e.g., body length and width, sclerotization and concealability. These results suggest that both filtering and partitioning processes structure oribatid mite communities. In native European beech forests, but not in non‐native Douglas fir forests, oribatid mites were mainly structured by filtering processes acting via traits related both to environmental tolerance and to resources. Furthermore, oribatid mite trait diversity, but not phylogenetic diversity, differed significantly between monocultures and mixed forests, demonstrating that multidimensional diversity indices provide additional information on soil biodiversity. Overall, the study provides evidence that traits representing different niche dimensions need to be considered for understanding assembly processes in soil animal communities and thereby soil biodiversity.

Rainforest conversion and expansion of plantations in tropical regions change local microclimate and are associated with biodiversity decline. Tropical soils are a hotspot of animal biodiversity and may sensitively respond to microclimate changes, but these responses remain unexplored. To address this knowledge gap, here we investigated seasonal fluctuations in density and community composition of Collembola, a dominant group of soil invertebrates, in rainforest, and in rubber and oil palm plantations in Jambi province (Sumatra, Indonesia). Across land-use systems, the density of Collembola in the litter was at a maximum at the beginning of the wet season, whereas in soil it generally varied little. The community composition of Collembola changed with season and the differences between land-use systems were most pronounced at the beginning of the dry season. Water content, pH, fungal and bacterial biomarkers, C/N ratio and root biomass were identified as factors related to seasonal variations in species composition of Collembola across different land-use systems. We conclude that (1) conversion of rainforest into plantation systems aggravates detrimental effects of low moisture during the dry season on soil invertebrate communities; (2) Collembola communities are driven by common environmental factors across land-use systems, with water content, pH and food availability being most important; (3) Collembola in litter are more sensitive to climatic variations than those in soil. Overall, the results document the sensitivity of tropical soil invertebrate communities to seasonal climatic variations, which intensifies the effects of the conversion of rainforest into plantation systems on soil biodiversity.

Abstract Carbon allocation of trees to ectomycorrhizas is thought to shape forest nutrient cycling, but the sink activities of different fungal taxa for host resources are unknown. Here, we investigate fungal taxon-specific differences in naturally composed ectomycorrhizal (EM) communities for plant-derived carbon and nitrogen. After aboveground dual labeling of young beech with 15N and 13C, ectomycorrhizas formed with different fungal taxa exhibit strong differences in label enrichment. Secondary Ion Mass Spectrometry (SIMS) imaging of nitrogen in cross sections of ectomycorrhizas demonstrates plant-derived 15N in both root and fungal structures. Isotope enrichment in ectomycorrhizas correlates with that in the corresponding ectomycorrhiza-attached lateral root, supporting fungal taxon-specific N and C fluxes in ectomycorrhizas. The enrichments with 13C and 15N in the symbiosis decrease with increasing C/N ratio of ectomycorrhizas, converging to zero at high C/N. The relative abundances of EM fungal species on roots are positively correlated with 13C enrichment, demonstrating higher fitness of stronger than of less C-demanding symbioses. Overall, our results support that differences among the C/N ratios in ectomycorrhizas formed with different fungal species regulate the supply of the symbioses with host-derived carbon and provide insights on functional traits of ectomycorrhizas, which are important for major ecosystem processes.

Global change is transforming Earth’s ecological communities with severe consequences for the functions and services they provide. In temperate grasslands, home to a mesmerising diversity of invertebrates controlling multiple ecosystem processes and services, land-use intensification and climate change are two of the most important global-change drivers. While we know a lot about their independent effects on grassland biodiversity and ecosystem functioning, little is known about how these stressors interact. Moreover, most research on biodiversity change focuses on decreasing biomass or species richness, while a major aspect is commonly ignored – altered ecological interactions. This is problematic because these interactions represent and control many important ecosystem processes, such as predation, herbivory or decomposition. Networks of trophic interactions, so-called food webs, link the structure and functioning of ecological communities and unravel mechanistic relationships between environmental change, ecological communities and ecosystem multifunctionality – the ability of a system to simultaneously support multiple processes. Consequently, we need to study how ecological interactions and the food webs they comprise respond to environmental change and to multiple interacting global-change drivers. Fortunately, novel tools offer unprecedented opportunities in studying trophic interactions and their impact on ecosystem processes. In addition, we know far more about how global change impacts the aboveground world than its belowground counterpart. However, belowground communities are just as important for the overall functioning of terrestrial ecosystems. Thus, to comprehensively understand global-change impacts on temperate grasslands, we need to study above- and belowground multitrophic interactions and ecosystem processes together, also accounting for their interdependencies. Here, we propose to use the Global Change Experimental Facility (GCEF, Bad Lauchstädt, Germany) to study joint impacts of land-use intensity and climate change on above-belowground multitrophic interactions and ecosystem multifunctionality in a temperate grassland global-change experiment. We will combine novel approaches to assessing trophic interactions and basal-resource dependency with an innovative method to quantify energy flux through ecological interaction networks. We will disentangle separate and interactive effects of land use and climate change and unravel how global-change driven modifications in multitrophic interactions mechanistically translate into altered ecosystem processes and multifunctionality – above and below the ground. Combining a field-experimental approach with novel molecular and quantitative techniques will allow for a leap forward in our understanding of global-change impacts on temperate grasslands, which will be crucial to manage and conserve these important ecosystems.

Biodiversity is declining on a global scale with detrimental effects on ecosystem functioning. Effects of reduced tree diversity on the diversity of aboveground animals have been studied in detail, whereas the response of soil animals remains poorly understood. We analyzed seasonal variations of soil oribatid mite communities as major soil detritivores along a tree diversity gradient as implemented in the Biodiversity - Ecosystem Functioning Experiment in China. A total of 24,898 oribatid mites were collected over two years and eight sampling periods. Generally, density and species richness of oribatid mites significantly differed among the four seasons and were highest in winter. By contrast, tree diversity did not significantly affect the density and species richness of oribatid mites as main factor, but its effect varied with season, however, the variations were generally small. Also, oribatid mite community composition varied with seasons but not significantly with tree diversity. Further, functional traits and functional diversity of oribatid mites significantly changed with seasons, but this varied with tree diversity although these variations were small. Overall, the effect of season on soil oribatid mite communities considerably exceeded that of tree diversity, indicating that oribatid mite communities in soil are mainly structured by seasonal variations in abiotic factors and resources outweighing the role of tree diversity. The results highlight the necessity of considering temporal variations when analyzing relationships between plant diversity and soil animal communities.

IntroductionHigh-throughput sequencing (HTS) provides an efficient and cost-effective way to generate large amounts of sequence data, providing a very powerful tool to analyze biodiversity of soil organisms. However, marker-based methods and the resulting datasets come with a range of challenges and disputes, including incomplete reference databases, controversial sequence similarity thresholds for delimitating taxa, and downstream compositional data analysis. MethodsHere, we use HTS data from a soil nematode biodiversity experiment to explore standardized HTS data processing procedures. We compared the taxonomic assignment performance of two main rDNA reference databases (SILVA and PR2). We tested whether the same ecological patterns are detected with Amplicon Sequence Variants (ASV; 100% similarity) versus classical Operational Taxonomic Units (OTU; 97% similarity). Further, we tested how different HTS data normalization methods affect the recovery of beta diversity patterns and the identification of differentially abundant taxa.ResultsAt this time, the SILVA 138 eukaryotic database performed better than the PR2 4.12 database, assigning more reads to family level and providing higher phylogenetic resolution. ASV- and OTU-based alpha and beta diversity of nematodes correlated closely, indicating that OTU-based studies represent useful reference points. For downstream data analyses, our results indicate that loss of data during subsampling under rarefaction-based methods might reduce the sensitivity of the method, e.g. underestimate the differences between nematode communities under different treatments, while the clr-transformation-based methods may overestimate effects. The Analysis of Compositions of Microbiome with Bias Correction approach (ANCOM-BC) retains all data and accounts for uneven sampling fractions for each sample, suggesting that this is currently the optimal method to analyze compositional data.DiscussionOverall, our study highlights the importance of comparing and selecting taxonomic reference databases before data analyses, and provides solid evidence for the similarity and comparability between OTU- and ASV-based nematode studies. Further, the results highlight the potential weakness of rarefaction-based and clr-transformation-based methods. We recommend future studies use ASV and that both the taxonomic reference databases and normalization strategies are carefully tested and selected before analyzing the data.

Investigating functional facets of biodiversity across elevation gradients provides the perspective to deepen understanding of the response of communities to global changes. Functional ecology approaches typically assume that filtering of traits across broad environmental gradients is largely due to species turnover rather than intraspecific trait variability. However, few studies quantified the relative importance of species turnover versus intraspecific variability for the composition of soil animal communities across environmental gradients. Here, for the first time, we investigated the relative contribution of species turnover, intraspecific trait variability and their covariation to changes in Collembola body size based on measurements at the individual level along a temperate-boreal elevation gradient in northeastern China. The results indicated that community weighted mean (CWM) body size of Collembola varied significantly with elevation as well as with life form, with the variations in CWM body size in total, epedaphic and hemiedaphic species being similar and more pronounced than in euedaphic Collembola. These variations in body size were driven to a similar extent by intraspecific trait variability and species turnover, with the former being somewhat higher in total, epedaphic and euedaphic but not in hemiedaphic Collembola. Further, variations in body size were better explained by local habitat-related soil factors than by temperature, but the relative importance of these factors differed among ecological groups of Collembola. Overall, these findings show that intraspecific variations in traits may be of similar importance than the turnover of species and needs closer consideration when using trait approaches to understand biodiversity – ecosystem functioning relationships in face of global change.

Soil invertebrates are key to decomposition, a central ecosystem process related to soil health. In many temperate areas climate change will decrease soil water content, which strongly modulates biological activity. However, data are lacking on how shifts in rainfall patterns affect soil biota and the ecosystem processes they provide. Here, we used the bait-lamina test to experimentally assess how a severe drought event influenced detritivore feeding activity, during a wheat growing season, in soils under long-term organic or conventional farming. Additionally, biotic and abiotic soil parameters were measured. Feeding activity was reduced under extreme drought and conventional management, although no climate-management synergies were found. Vertical migrations of Collembola and Oribatida partially explained the unexpectedly higher bait consumption at shallower depths in response to drought. Exploratory mixed-effects longitudinal random forests (a novel machine learning technique) were used to explore whether the relative abundances of meso‑, microfauna and microbes of the decomposer food web, or abiotic soil parameters, affected the feeding activity of detritivores. The model including meso‑ and microfauna selected four Nematoda taxa and explained higher variance than the model with only microbiota, indicating that detritivore feeding is closely associated with nematodes but not with microbes. Additionally, the model combining fauna and microbiota explained less variance than the faunal model, suggesting that microbe-fauna synergies barely affected detritivore feeding. Moreover, soil water and mineral nitrogen contents were found to strongly determine detritivore feeding, in a positive and negative way, respectively. Hence, our results suggest that severe drought and conventional farming impair the feeding activity of soil detritivores and thus, probably, decomposition and nutrient mineralization in soils. Furthermore, machine learning algorithms arise as a powerful technique to explore the identity of potential key drivers relating biodiversity to ecosystem functioning.

Abstract Enhancing biodiversity in monoculture-dominated landscapes is a key sustainability challenge that requires considering the spatial organization of ecological communities (beta diversity). Here, we tested whether increasing landscape heterogeneity, through establishing 52 tree islands in an oil-palm landscape, is a suitable restoration strategy to enhance the diversity of six taxa (multi-taxa diversity). Further, we elucidated whether patterns in the spatial distribution of above- and below-ground taxa are related, and their role in shaping multi-taxa beta diversity. After five years, islands enhanced diversity at the landscape scale by fostering unique species (turnover). Partial correlation networks revealed that dissimilarity, in vegetation structural complexity and soil conditions, impacts multi-taxa beta diversity and turnover. In addition, soil fauna, bacteria, and fungi were more strongly associated with the overall community than aboveground taxa. Thus, strategies aiming to enhance multi-taxa diversity should consider the central role of landscape heterogeneity and soil biota.

Molecular gut content analysis via diagnostic PCR or high-throughput sequencing (metabarcoding) of consumers allows unravelling of feeding interactions in a wide range of animals. This is of particular advantage for analyzing the diet of small invertebrates living in opaque habitats such as the soil. Due to their small body size, which complicates dissection, microarthropods are subjected to whole-body DNA extraction-step before their gut content is screened for DNA of their food. This poses the problem that body surface contaminants, such as fungal spores may be incorrectly identified as ingested food particles for fungivorous species. We investigated the effectiveness of ten methods for body surface decontamination in litter-dwelling oribatid mites using Steganacarus magnus as model species. Furthermore, we tested for potential adverse effects of the decontamination techniques on the molecular detection of ingested prey organisms. Prior to decontamination, oribatid mites were fed with an oversupply of nematodes (Plectus sp.) and postmortem contaminated with fungal spores (Chaetomium globosum). We used diagnostic PCR with primers specific for C. globosum and Plectus sp. to detect contaminants and prey, respectively. The results suggest that chlorine bleach (sodium hypochloride, NaClO, 5%) is most efficient in removing fungal surface contamination without significantly affecting the detection of prey DNA in the gut. Based on these results, we provide a standard protocol for efficient body surface decontamination allowing to trace the prey spectrum of microarthropods using molecular gut content analysis.

Abstract Biodiversity is widely linked to human health, however, connections between human health and soil biodiversity in urban environments remain poorly understood. Here, we stress that reductions in urban soil biodiversity elevate risks to human health, but soil biodiversity can improve human health through pathways including suppressing pathogens, remediating soil, shaping a beneficial human microbiome and promoting immune fitness. We argue that targeted enhancement of urban soil biodiversity could support human health, in both outdoor and indoor settings. The potential of enhanced urban soil biodiversity to benefit human health reflects an important yet understudied field of fundamental and applied research.

Springtails are omnipresent soil arthropods, vital for ecosystems. In the first global assessment of springtails, this study shows a 20-fold biomass difference between the tundra and the tropics, with distinct temperature-related patterns for diversity and metabolism that suggest climate change may restructure the functioning of soil biodiversity.

Abstract Background Functional diversity is vital for forest ecosystem resilience in times of climate-induced forest diebacks. Admixing drought resistant non-native Douglas fir, as a partial replacement of climate-sensitive Norway spruce, to native beech forests in Europe appears promising for forest management, but possible consequences for associated biota and ecosystem functioning are poorly understood. To better link forest management and functional diversity of associated biota, we investigated the trophic niches (∆13C, ∆15N) of epigeic generalist predators (spiders and ground beetles) in mixed and pure stands of European beech, Norway spruce and non-native Douglas fir in north-west Germany. We assessed the multidimensional niche structure of arthropod predator communities using community-based isotopic metrics. Results Whilst arthropod ∆13C differed most between beech (high ∆13C) and coniferous stands (low ∆13C), ∆15N was lowest in non-native Douglas fir. Tree mixtures mitigated these effects. Further, conifers increased isotopic ranges and isotopic richness, which is linked to higher canopy openness and herb complexity. Isotopic divergence of ground beetles decreased with Douglas fir presence, and isotopic evenness of spiders in Douglas fir stands was lower in loamy sites with higher precipitation than in sandy, drier sites. Conclusions We conclude that tree species and particularly non-native trees alter the trophic niche structure of generalist arthropod predators. Resource use and feeding niche breadth in non-native Douglas fir and native spruce differed significantly from native beech, with more decomposer-fueled and narrower feeding niches in beech stands (∆13C, isotopic ranges and richness). Arthropod predators in non-native Douglas fir, however, had shorter (∆15N) and simplified (isotopic divergence) food chains compared to native forest stands; especially under beneficial abiotic conditions (isotopic evenness). These findings indicate potential adverse effects of Douglas fir on functional diversity of generalist arthropod predators. As tree mixtures mitigated differences between beech and conifers, mixed stands including (non-native) conifers constitute a promising compromise between economic and conservational interests.

Abstract Background In the last decades, Southeast Asia has experienced massive conversion of rainforest into rubber and oil palm monoculture plantations. The effects of this land-use change on canopy arthropods are still largely unknown. Arboreal Collembola are among the most abundant canopy arthropods in tropical forests, potentially forming a major component of the canopy food web by contributing to the decomposition of arboreal litter and being an important prey for canopy arthropod predators. We investigated abundance, richness, and community composition of, as well as the influence of a series of environmental factors on, canopy Collembola communities in four land-use systems in Jambi Province, Sumatra, Indonesia: (1) lowland rainforest, (2) jungle rubber (rubber agroforest), and monoculture plantations of (3) rubber and (4) oil palm. Results Using canopy fogging in 32 research plots in both the dry and rainy seasons in 2013, we collected 77,104 specimens belonging to 68 (morpho) species. Generally, Collembola communities were dominated by few species including two species of the genus Salina (Paronellidae; 34% of total individuals) and two species of Lepidocyrtinae (Entomobryidae; 20%). The abundance of Collembola in lowland rainforest (53.4 ± 30.7 ind. m−2) was more than five times higher than in rubber plantations, and more than ten times higher than in oil palm plantations; abundances in jungle rubber were intermediate. Collembola species richness was highest in rainforest (18.06 ± 3.60 species) and jungle rubber (16.88 ± 2.33 species), more than twice that in rubber or oil palm. Collembola community composition was similar in rainforest and jungle rubber, but different from monoculture plantations which had similar Collembola community composition to each other. The environmental factors governing community composition differed between the land-use systems and varied between seasons. Conclusions Overall, this is the first in-depth report on the structure of arboreal Collembola communities in lowland rainforest and agricultural replacement systems in Southeast Asia. The results highlight the potentially major consequences of land-use change for the functioning of arboreal arthropod food webs.

Human activities are increasing the input of atmospheric particulate pollutants to forests. The components of particulate pollutants include inorganic anions, base cations and hydrocarbons. Continuous input of particulate pollutants may affect soil functioning in forests, but their effects may be modified by soil fauna. However, studies investigating how soil fauna affects the effects of particulate pollutants on soil functioning are lacking. Here, we investigated how earthworms and the particulate components interact in affecting soil enzymatic functions in a deciduous (Quercus variabilis) and a coniferous (Pinus massoniana) forest in southeast China. We manipulated the addition of nitrogen (N, ammonium nitrate), sodium (Na, sodium chloride) and polycyclic aromatic hydrocarbons (PAHs, five mixed PAHs) in field mesocosms with and without Eisenia fetida, an earthworm species colonizing forests in eastern China. After one year, N and Na addition increased, whereas PAHs decreased soil enzymatic functions, based on average Z scores of extracellular enzyme activities. Earthworms generally stabilized soil enzymatic functions via neutralizing the effects of N, Na and PAHs addition in the deciduous but not in the coniferous forest. Specifically, earthworms neutralized the effects of N and Na addition on soil pH and the effects of the addition of PAHs on soil microbial biomass. Further, both particulate components and earthworms changed the correlations among soil enzymatic and other ecosystem functions in the deciduous forest, but the effects depended on the type of particulate components. Generally, the effects of particulate components and earthworms on soil enzymatic functions were weaker in the coniferous than the deciduous forest. Overall, the results indicate that earthworms stabilize soil enzymatic functions in the deciduous but not the coniferous forest irrespective of the type of particulate components. This suggests that earthworms may neutralize the influence of atmospheric particulate pollutants on ecosystem functions, but the neutralization may be restricted to deciduous forests.

Abstract Cultivation of non‐native tree species is a promising option to adapt managed forests to climate change. However, consequences of non‐native tree species on flora, fauna, and microorganisms that occur in forest ecosystems (forest associates) need to be considered when managing forests. We lack a solid understanding of how cultivating non‐native tree species in pure stands and in mixture with native tree species impacts abundance and species diversity of forest associates. We compared abundance, alpha‐, beta‐ and gamma‐diversity of eight forest‐floor‐associated taxa that are relevant for ecosystem functioning (including fungi, plants, arthropods, and small mammals) between different forest ecosystems. We addressed pure stands of non‐native coniferous Douglas‐fir (Pseudotsuga menziesii) and two native species, broad‐leaved European beech (Fagus sylvatica) and coniferous Norway spruce (Picea abies), as well as mixed stands of European beech with each conifer in two regions with differing site conditions in temperate Northwest Germany. Diversity indices revealed differences among species groups. Gamma‐diversity and alpha‐diversity of forest associates in Douglas‐fir and spruce stands were usually higher than in beech stands. Estimates of species diversity of mixed stands are intermediate between estimates for the respective pure stands. Differences in the diversity between the two study regions were highly taxon specific with no clear support for a general trend toward a higher or lower species diversity of forest associates depending on site quality. Abundance values show a pattern similar to our diversity estimates, but with a higher statistical uncertainty. Non‐native Douglas‐fir stands provided habitats to support associated species communities of equally high or higher diversity than stands of native beech or spruce. Mixed stands of non‐native and native tree species may be a management option to achieve different goals, that is, to provide habitats for species that are strictly associated with beech and to increase resilience to climate change. However, the overall diversity of forest‐floor‐associated biota is not improved by cultivating Douglas‐fir or spruce in mixture with beech.

Abstract Earthworms modulate carbon and nitrogen cycling in terrestrial ecosystems, but their effect may be compromised by the deposition of pollutants from industrial emissions. However, studies investigating how deposited compounds affect the role of earthworms in carbon cycling such as litter decomposition are lacking, although the interactions of earthworms and deposited compounds are important for understanding the impact of pollutants on ecosystems and the potential of earthworms in bioremediation. We performed a 365‐day in situ litterbag decomposition experiment in a deciduous (Quercus variabilis) and coniferous (Pinus massoniana) forest in southeast China. We manipulated nitrogen (N), sodium (Na), and polycyclic aromatic hydrocarbons (PAHs) as model compounds during litter decomposition with and without earthworms (Eisenia fetida). After one year, N, Na, and PAH all slowed down litter mass loss, with the effects of Na being the strongest. By contrast, E. fetida generally increased litter mass loss, and the positive effects were uniformly maintained irrespective of the type of compounds added. However, the pathways to how earthworms increased litter mass loss varied among the compounds added and the two forests studied. As indicated by structural equation modeling, earthworms mitigated the negative effects of deposited compounds by directly increasing litter mass loss and indirectly increasing soil pH and microbial biomass. Overall, the results indicate that the acceleration of litter mass loss by earthworms is little affected by deposited compounds, and that earthworms have the potential to mitigate negative impacts of pollutants on litter decomposition and ecosystem processes.

Abstract Niche theory fundamentally contributed to the understanding of animal diversity. However, in soil, the diversity of animals seems enigmatic since the soil is a rather homogeneous habitat, and soil animals are often generalist feeders. A new approach to understand soil animal diversity is the use of ecological stoichiometry. The elemental composition of animals may explain their occurrence, distribution, and density. This approach has been used before in soil macrofauna, but this study is the first to investigate soil mesofauna. Using inductively coupled plasma optic emission spectrometry (ICP‐OES), we analyzed the concentration of a wide range of elements (Al, Ca, Cu, Fe, K, Mg, Mn, Na, P, S, Zn) in 15 soil mite taxa (Oribatida, Mesostigmata) from the litter of two different forest types (beech, spruce) in Central Europe (Germany). Additionally, the concentration of carbon and nitrogen, and their stable isotope ratios (15N/14N, 13C/12C), reflecting their trophic niche, were measured. We hypothesized that (1) stoichiometry differs between mite taxa, (2) stoichiometry of mite taxa occurring in both forest types is not different, and (3) element composition is correlated to trophic level as indicated by 15N/14N ratios. The results showed that stoichiometric niches of soil mite taxa differed considerably indicating that elemental composition is an important niche dimension of soil animal taxa. Further, stoichiometric niches of the studied taxa did not differ significantly between the two forest types. Calcium was negatively correlated with trophic level indicating that taxa incorporating calcium carbonate in their cuticle for defense occupy lower trophic positions in the food web. Furthermore, a positive correlation of phosphorus with trophic level indicated that taxa higher in the food web have higher energetic demand. Overall, the results indicate that ecological stoichiometry of soil animals is a promising tool for understanding their diversity and functioning.

Abstract Salt marshes exist along the gradient of the marine mudflat to the terrestrial dunes, with a gradient of shore height and associated plant zonation. The lower salt marsh (LSM) extends from the mean high tidal level to 35 cm above that level and is followed by the upper salt marsh (USM). Despite changes in the amount of allochthonous marine input and in abiotic conditions, little is known about changes in the trophic structure and used of basal resources by the soil macrofauna along marine—terrestrial boundaries. Natural variations in carbon stable isotope ratios (δ13C signatures) allow insight into basal resources of consumers such as marine algae, terrestrial C3 and C4 photosynthesising plants. Furthermore, variations in nitrogen stable isotope ratios (δ15N signatures) allow insight into the trophic position of consumers. We investigated spatial and temporal changes in stable isotope signatures in salt marsh soil macrofauna of the island of Spiekeroog, German Wadden Sea. The range of δ15N signatures indicated no changes in food chain length across salt marsh zones with consumers in both zones comprising primary decomposers, secondary decomposers and first order predators. However, the trophic position of individual species changed between zones, but in particular with season. Contrasting δ15N signatures, the range in δ13C signatures in the LSM was twice that in the USM indicating a wider range of resources consumed. Bayesian mixing models indicated predominant autochthonous resource use in both the LSM and USM, with the use of marine allochthonous resources never exceeding 29.6%. However, the models also indicate an increase in the use of marine resources in certain species in the LSM with no use in the USM. Overall, the results indicate that the resource use of salt marsh macrofauna varies more in space than in time, with the food web being generally based on autochthonous rather than allochthonous resources. However, there also is trophic plasticity in certain species across both temporal and spatial scales including variations in the use of allochthonous resources. Generally, however, marine input contributes little to the nutrition of salt marsh soil macroinvertebrates.

Abstract Endogenous rhythmic growth (ERG) is displayed by many tropical and some major temperate tree species and characterized by alternating root and shoot flushes (RF and SF). These flushes occur parallel to changes in biomass partitioning and in allocation of recently assimilated carbon and nitrogen. To address how biotic interactions interplay with ERG, we cross-compared the RF/SF shifts in oak microcuttings in the presence of pathogens, consumers and a mycorrhiza helper bacterium, without and with an ectomycorrhizal fungus (EMF), and present a synthesis of the observations. The typical increase in carbon allocation to sink leaves during SF did not occur in the presence of root or leaf pathogens, and the increase in nitrogen allocation to lateral roots during RF did not occur with the pathogens. The RF/SF shifts in resource allocation were mostly restored upon additional interaction with the EMF. Its presence led to increased resource allocation to principal roots during RF, also when the oaks were inoculated additionally with other interactors. The interactors affected the alternating, rhythmic growth and resource allocation shifts between shoots and roots. The restoring role of the EMF on RF/SF changes in parallel to the corresponding enhanced carbon and nitrogen allocation to sink tissues suggests that the EMF is supporting plants in maintaining the ERG.

Abstract Soil biodiversity constitutes the biological pillars of ecosystem services provided by soils worldwide. Soil life is threatened by intense agricultural management and shifts in climatic conditions as two important global change drivers which are not often jointly studied under field conditions. We addressed the effects of experimental short-term drought over the wheat growing season on soil organisms and ecosystem functions under organic and conventional farming in a Swiss long term trial. Our results suggest that activity and community metrics are suitable indicators for drought stress while microbial communities primarily responded to agricultural practices. Importantly, we found a significant loss of multiple pairwise positive and negative relationships between soil biota and process-related variables in response to conventional farming, but not in response to experimental drought. These results suggest a considerable weakening of the contribution of soil biota to ecosystem functions under long-term conventional agriculture. Independent of the farming system, experimental and seasonal (ambient) drought conditions directly affected soil biota and activity. A higher soil water content during early and intermediate stages of the growing season and a high number of significant relationships between soil biota to ecosystem functions suggest that organic farming provides a buffer against drought effects.

Abstract Biodiversity and ecosystem functions are highly threatened by global change. It has been proposed that geodiversity can be used as an easy-to-measure surrogate of biodiversity to guide conservation management. However, so far, there is mixed evidence to what extent geodiversity can predict biodiversity and ecosystem functions at the regional scale relevant for conservation planning. Here, we analyse how geodiversity computed as a compound index is suited to predict the diversity of four taxa and associated ecosystem functions in a tropical mountain hotspot of biodiversity and compare the results with the predictive power of environmental conditions and resources (climate, habitat, soil). We show that combinations of these environmental variables better explain species diversity and ecosystem functions than a geodiversity index and identified climate variables as more important predictors than habitat and soil variables, although the best predictors differ between taxa and functions. We conclude that a compound geodiversity index cannot be used as a single surrogate predictor for species diversity and ecosystem functions in tropical mountain rain forest ecosystems and is thus little suited to facilitate conservation management at the regional scale. Instead, both the selection and the combination of environmental variables are essential to guide conservation efforts to safeguard biodiversity and ecosystem functions.

Background Plant litter decomposition is a key process in carbon and nutrient cycling. Among the factors determining litter decomposition rates, the role of soil biota in the decomposition of different plant litter types and its modification by variations in climatic conditions is not well understood. Methods In this study, we used litterbags with different mesh sizes (45 µm, 1 mm and 4 mm) to investigate the effect of microorganisms and decomposer microarthropods on leaf and root litter decomposition along an altitudinal gradient of tropical montane rainforests in Ecuador. We examined decomposition rates, litter C and N concentrations, microbial biomass and activity, as well as decomposer microarthropod abundance over one year of exposure at three different altitudes (1,000, 2,000 and 3,000 m). Results Leaf litter mass loss did not differ between the 1,000 and 2,000 m sites, while root litter mass loss decreased with increasing altitude. Changes in microbial biomass and activity paralleled the changes in litter decomposition rates. Access of microarthropods to litterbags only increased root litter mass loss significantly at 3,000 m. The results suggest that the impacts of climatic conditions differentially affect the decomposition of leaf and root litter, and these modifications are modulated by the quality of the local litter material. The findings also highlight litter quality as the dominant force structuring detritivore communities. Overall, the results support the view that microorganisms mostly drive decomposition processes in tropical montane rainforests with soil microarthropods playing a more important role in decomposing low-quality litter material.

Abstract In Central Europe, summer droughts are increasing in frequency which threatens production and biodiversity in agroecosystems. The potential of different farming systems to mitigate detrimental drought effects on soil animals is largely unknown. We investigated the effects of simulated drought on the abundance and community composition of soil microarthropods (Collembola and Oribatida and Meso‐, Pro‐, and Astigmata) in winter wheat fields under long‐term conventional and organic farming in the DOK trial, Switzerland. We simulated drought by excluding 65% of the ambient precipitation during the wheat‐growing season from March to June 2017. The abundance of Collembola and Oribatida declined more consistently in conventionally managed fields compared to organically managed fields under simulated drought. The abundance of Collembola as well as Meso‐, Pro‐ and Astigmata, but not the abundance of Oribatida, increased in deeper soil layers due to simulated drought, suggesting vertical migration as a drought avoidance strategy. The species composition of Oribatida communities, but not of Collembola communities, differed significantly between drought treatments and between farming systems. Soil carbon content was a major factor structuring Oribatida communities. Our results suggest that organic farming buffers negative effects of drought on soil microarthropods, presumably due to higher soil carbon content and associated higher soil moisture and improved soil structure. This potential of organic farming systems to mitigate consequences of future droughts on soil biodiversity is promising and needs further exploration across larger climatic and spatial scales and should be extended to other groups of soil biota.

Abstract Rainforest conversion and expansion of plantations in tropical regions are associated with changes in animal communities and biodiversity decline. In soil, Collembola are one of the most numerous invertebrate groups that affect the functioning of microbial communities and support arthropod predators. Despite that, information on the impact of changes in land use in the tropics on species and trait composition of Collembola communities is very limited. We investigated the response of Collembola to the conversion of rainforest into rubber agroforestry (“jungle rubber”), rubber, and oil palm plantations in Jambi Province (Sumatra, Indonesia), a region which experienced one of the strongest recent deforestation globally. Collembola were sampled in 2013 and 2016 from the litter and soil layer using heat extraction, and environmental factors were measured (litter C/N ratio, pH, water content, composition of microbial community and predator abundance). In the litter layer, density and species richness in plantation systems were 25%–38% and 30%–40% lower, respectively, than in rainforest. However, in the soil layer, density, species richness, and trait diversity of Collembola were only slightly affected by land‐use change, contrasting the response of many other animal groups. Species and trait composition of Collembola communities in litter and soil differed between each of the land‐use systems. Water content and pH were identified as main factors related to the differences in species and trait composition in both litter and soil, followed by the density of micro‐ and macropredators. Dominant species of Collembola in rainforest and jungle rubber were characterized by small body size, absence of furca, and absence of intense pigmentation, while in plantations, larger species with long furca and diffuse or patterned pigmentation were more abundant. Overall, land‐use change negatively affected Collembola communities in the litter layer, but its impact was lower in the soil layer. Several pantropical genera of Collembola (i.e., Isotomiella, Pseudosinella, and Folsomides) dominated across land‐use systems, reflecting their high environmental adaptability and/or efficient dispersal, calling for studies on their ecology and genetic diversity. The decline in species richness and density of litter‐dwelling Collembola with the conversion of rainforest into plantation systems calls for management practices mitigating negative effects of the deterioration of the litter layer in rubber plantations, but even more in oil palm plantations.

Rainforest canopies, home to one of the most complex and diverse terrestrial arthropod communities, are threatened by conversion of rainforest into agricultural production systems. However, little is known about how predatory arthropod communities respond to such conversion. To address this, we compared canopy spider (Araneae) communities from lowland rainforest with those from three agricultural systems in Jambi Province, Sumatra, Indonesia, i.e., jungle rubber (rubber agroforest) and monoculture plantations of rubber and oil palm. Using canopy fogging, we collected 10,676 spider specimens belonging to 36 families and 445 morphospecies. The four most abundant families (Salticidae N = 2,043, Oonopidae N = 1,878, Theridiidae N = 1,533 and Clubionidae N = 1,188) together comprised 62.2% of total individuals, while the four most speciose families, Salticidae (S = 87), Theridiidae (S = 83), Araneidae (S = 48) and Thomisidae (S = 39), contained 57.8% of all morphospecies identified. In lowland rainforest, average abundance, biomass and species richness of canopy spiders was at least twice as high as in rubber or oil palm plantations, with jungle rubber showing similar abundances as rainforest, and intermediate biomass and richness. Community composition of spiders was similar in rainforest and jungle rubber, but differed from rubber and oil palm, which also differed from each other. Canonical Correspondence Analysis showed that canopy openness, aboveground tree biomass and tree density together explained 18.2% of the variation in spider communities at family level. On a morphospecies level, vascular plant species richness and tree density significantly affected the community composition but explained only 6.8% of the variance. While abundance, biomass and diversity of spiders declined strongly with the conversion of rainforest into monoculture plantations of rubber and oil palm, we also found that a large proportion of the rainforest spider community can thrive in extensive agroforestry systems such as jungle rubber. Despite being very different from rainforest, the canopy spider communities in rubber and oil palm plantations may still play a vital role in the biological control of canopy herbivore species, thus contributing important ecosystem services. The components of tree and palm canopy structure identified as major determinants of canopy spider communities may aid in decision-making processes toward establishing cash-crop plantation management systems which foster herbivore control by spiders.

Information about soil scale insect (Hemiptera: Coccomorpha) diversity, although fragmented, had been progressively improved for Asia, especially in the southern countries such as Indonesia. Recent works recorded 365 species in the country, 21 of which in below ground habit. In this paper, we contribute to the knowledge of Indonesian scale insect diversity by analyzing specimens collected from rainforest, rubber and oil palm plantations in the Province of Jambi and Sumatra, Indonesia. The results include the new records of Rhizoecus americanus, Rhizoecus omphalius, Ripersiella bacorum, Ripersiella cryphia and Ripersiella sabahica (Rhizoecidae) with description of new intraspecific variations. Pseudococcus saccharicola (Pseudococcidae), Geococcus coffeae, and Rhizoecus pignerator (Rhizoecidae) were also collected in this study. With these species, the known Indonesian scale insect fauna increases to 370 species.

Abstract Although nitrogen (N) deposition is increasing globally, N availability still limits many organisms, such as microorganisms and mesofauna. However, little is known to which extent soil organisms rely on mineral‐derived N and whether plant community composition modifies its incorporation into soil food webs. More diverse plant communities more effectively compete with microorganisms for mineral N likely reducing the incorporation of mineral‐derived N into soil food webs. We set up a field experiment in experimental grasslands with different levels of plant species and functional group richness. We labeled soil with 15NH415NO3 and analyzed the incorporation of mineral‐derived 15N into soil microorganisms and mesofauna over 3 months. Mineral‐derived N incorporation decreased over time in all investigated organisms. Plant species richness and presence of legumes reduced the uptake of mineral‐derived N into microorganisms. In parallel, the incorporation of mineral‐derived 15N into mesofauna species declined with time and decreased with increasing plant species richness in the secondary decomposer springtail Ceratophysella sp. Effects of both plant species richness and functional group richness on other mesofauna species varied with time. The presence of grasses increased the 15N incorporation into Ceratophysella sp., but decreased it in the primary decomposer oribatid mite Tectocepheus velatus sarekensis. The results highlight that mineral N is quickly channeled into soil animal food webs via microorganisms irrespective of plant diversity. The amount of mineral‐derived N incorporated into soil animals, and the plant community properties affecting this incorporation, differed markedly between soil animal taxa, reflecting species‐specific use of food resources. Our results highlight that plant diversity and community composition alter the competition for N in soil and change the transfer of N across trophic levels in soil food webs, potentially leading to changes in soil animal population dynamics and community composition. Sustaining high plant diversity may buffer detrimental effects of elevated N deposition on soil biota.

Abstract In tropical forest ecosystems leaf litter from a large variety of species enters the decomposer system, however, the impact of leaf litter diversity on the abundance and activity of soil organisms during decomposition is little known. We investigated the effect of leaf litter diversity and identity on microbial functions and the abundance of microarthropods in Ecuadorian tropical montane rainforests. We used litterbags filled with leaves of six native tree species (Cecropia andina, Dictyocaryum lamarckianum, Myrcia pubescens, Cavendishia zamorensis, Graffenrieda emarginata, and Clusia spp.) and incubated monocultures and all possible two‐ and four‐species combinations in the field for 6 and 12 months. Mass loss, microbial biomass, basal respiration, metabolic quotient, and the slope of microbial growth after glucose addition, as well as the abundance of microarthropods (Acari and Collembola), were measured at both sampling dates. Leaf litter diversity significantly increased mass loss after 6 months of exposure, but reduced microbial biomass after 12 months of exposure. Leaf litter species identity significantly changed both microbial activity and microarthropod abundance with species of high quality (low C‐to‐N ratio), such as C. andina, improving resource quality as indicated by lower metabolic quotient and higher abundance of microarthropods. Nonetheless, species of low quality, such as Clusia spp., also increased the abundance of Oribatida suggesting that leaf litter chemical composition alone is insufficient to explain variation in the abundances of soil microarthropods. Overall, the results provide evidence that decomposition and microbial biomass in litter respond to leaf litter diversity as well as litter identity (chemical and physical characteristics), while microarthropods respond only to litter identity but not litter diversity.

Abstract The existence of old species‐rich parthenogenetic taxa is a conundrum in evolutionary biology. Such taxa point to ancient parthenogenetic radiations resulting in morphologically distinct species. Ancient parthenogenetic taxa have been proposed to exist in bdelloid rotifers, darwinulid ostracods, and in several taxa of acariform mites (Acariformes, Acari), especially in oribatid mites (Oribatida, Acari). Here, we investigate the diversification of Acariformes and their ancestral mode of reproduction using 18S rRNA. Because parthenogenetic taxa tend to be more frequent in phylogenetically old taxa of Acariformes, we sequenced a wide range of members of this taxon, including early‐derivative taxa of Prostigmata, Astigmata, Endeostigmata, and Oribatida. Ancestral character state reconstruction indicated that (a) Acariformes as well as Oribatida evolved from a sexual ancestor, (b) the primary mode of reproduction during evolution of Acariformes was sexual; however, species‐rich parthenogenetic taxa radiated independently at least four times (in Brachychthonioidea (Oribatida), Enarthronota (Oribatida), and twice in Nothrina (Oribatida), (c) parthenogenesis additionally evolved frequently in species‐poor taxa, for example, Tectocepheus, Oppiella, Rostrozetes, Limnozetes, and Atropacarus, and (d) sexual reproduction likely re‐evolved at least three times from species‐rich parthenogenetic clusters, in Crotonia (Nothrina), in Mesoplophora/Apoplophora (Mesoplophoridae, Enarthronota), and in Sphaerochthonius/Prototritia (Protoplophoridae, Enarthronota). We discuss possible reasons that favored the frequent diversification of parthenogenetic taxa including the continuous long‐term availability of dead organic matter resources as well as generalist feeding of species as indicated by natural variations in stable isotope ratios.

Abstract Sex is evolutionarily more costly than parthenogenesis, evolutionary ecologists therefore wonder why sex is much more frequent than parthenogenesis in the majority of animal lineages. Intriguingly, parthenogenetic individuals and species are as common as or even more common than sexuals in some major and putative ancient animal lineages such as oribatid mites and rotifers. Here, we analyzed oribatid mites (Acari: Oribatida) as a model group because these mites are ancient (early Paleozoic), widely distributed around the globe, and include a high number of parthenogenetic species, which often co‐exist with sexual oribatid mite species. There is evidence that the reproductive mode is phylogenetically conserved in oribatid mites, which makes them an ideal model to test hypotheses on the relationship between reproductive mode and species' ecological strategies. We used oribatid mites to test the frozen niche variation hypothesis; we hypothesized that parthenogenetic oribatid mites occupy narrow specialized ecological niches. We used the geographic range of species as a proxy for specialization as specialized species typically do have narrower geographic ranges than generalistic species. After correcting for phylogenetic signal in reproductive mode and demonstrating that geographic range size has no phylogenetic signal, we found that parthenogenetic lineages have a higher probability to have broader geographic ranges than sexual species arguing against the frozen niche variation hypothesis. Rather, the results suggest that parthenogenetic oribatid mite species are more generalistic than sexual species supporting the general‐purpose genotype hypothesis. The reason why parthenogenetic oribatid mite species are generalists with wide geographic range sizes might be that they are of ancient origin reflecting that they adapted to varying environmental conditions during evolutionary history. Overall, our findings indicate that parthenogenetic oribatid mite species possess a widely adapted general‐purpose genotype and therefore might be viewed as “Jack‐of‐all‐trades.”

Abstract Deforestation and agricultural expansion in the tropics affect local and regional climatic conditions, leading to synergistic negative impacts on land ecosystems. Climatic changes manifest in increased inter‐ and intraseasonal variations and frequency of extreme climatic events (i.e., droughts and floods), which have evident consequences for aboveground biodiversity. However, until today, there have been no studies on how land use affects seasonal variations below ground in tropical ecosystems, which may be more buffered against climatic variation. Here, we analyzed seasonal variations in soil parameters, basal respiration, microbial communities, and abundances of soil invertebrates along with microclimatic conditions in rainforest and monocultures of oil palm and rubber in Sumatra, Indonesia. About 75% (20 out of 26) of the measured litter and soil, microbial, and animal parameters varied with season, with seasonal changes in 50% of the parameters depending on land use. Land use affected seasonal variations in microbial indicators associated with carbon availability and cycling rate. The magnitude of seasonal variations in microbial parameters in the soil of monocultures was almost 40% higher than in the soil of rainforest. Measured parameters were associated with short‐term climatic conditions (3‐day period air humidity) in plantations, but not in rainforest, confirming a reduced soil buffering ability in plantations. Overall, our findings suggest that land use temporally shifts and increases the magnitude of seasonal variations of the belowground ecosystem compartment, with microbial communities responding most strongly. The increased seasonal variations in soil biota in plantations likely translate into more pronounced fluctuations in essential ecosystem functions such as nutrient cycling and carbon sequestration, and these ramifications ultimately may compromise the stability of tropical ecosystems in the long term. As the observed seasonal dynamics is likely to increase with both local and global climate change, these shifts need closer attention for the long‐term sustainable management of plantation systems in the tropics.

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, 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, microbial communities, and vegetation, limits our understanding of biological interactions in soil systems 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 at sites associated with the global soil biodiversity monitoring network, Soil BON, allowing us to assess linkages among soil biodiversity, vegetation, soil physico-chemical properties, climate, 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.

Human-induced global environmental change is predicted to alter the stability and functioning of ecosystems worldwide. Most research in recent decades has focused on studying climate-change effects on aboveground systems, causing a poor understanding of belowground responses. However, gaining knowledge on environmental-change effects on soil biota is of crucial importance, as soil-ecosystem services are indispensable for human well-being and contribute fundamentally to the functioning of terrestrial ecosystems. Nematode communities play a central role in various soil ecosystem processes and are therefore commonly used as biological indicators to assess soil conditions and soil health. While causing overall shifts in community composition patterns, which are most often examined, climate change might also alter nematode population dynamics and the sex ratio (number of males per female). Previous studies on plant-parasitic nematode species suggest that changes to unfavorable environmental conditions trigger reduced development of females and favor sexual rather than parthenogenetic reproduction. Therefore, we are presenting the working hypothesis that predicted climate change causing reduced resource availability and enhanced environmental stress will lead to an increased proportion of males in soil nematode communities. Our systematic literature review revealed that climate- and environmental change effects on the sex ratio of free-living soil nematode populations are inconsistent, but heavily understudied. Data on sex ratios have been treated mostly as additional information, presented without any underlying theory and hypotheses, as well as limited discussion. In this perspective paper, we thus propose that future studies should include clear hypotheses and test if the sex ratio of free-living nematodes increases with climate change due to more stressful environmental conditions and low resource availability. Furthermore, we conclude that experimental studies investigating the specific roles of male and female nematodes are needed to better predict the implications of a changing climate on soil ecosystem functioning.

Herein, we update the diagnosis and description of the genus Krantzolaspina Datta & Bhattacharjee and provide a list of the three valid species including new combinations and synonyms, as follows: 1) Krantzolaspina angustatus (Ishikawa, 1987) comb. nov. (= Indutolaelaps jiroftensis Hajizadeh et al., 2017 syn. nov.), 2) K. rebatii Datta & Bhattacharjee, 1989 and 3) K. solimani (Metwali, 1983) comb. nov. Finally, we re-describe K. angustatus (Ishikawa, 1987) comb. nov. based on the holotype from Japan, voucher specimens from Iran and additional females that we found in soil samples from oil palm plantations in Sumatra, Indonesia.

Abstract Plants provide resources and shape the habitat of soil organisms thereby affecting the composition and functioning of soil communities. Effects of plants on soil communities are largely taxon‐dependent, but how different functional groups of herbaceous plants affect trophic niches of individual animal species in soil needs further investigation. Here, we studied the use of basal resources and trophic levels of dominating soil meso‐ and macrofauna using stable isotope ratios of carbon and nitrogen in arable fallow systems 3 and 14–16 years after abandonment. Animals were sampled from the rhizosphere of three plant species of different functional groups: a legume (Medicaco sativa), a nonlegume herb (Taraxacum officinale), and a grass (Bromus sterilis). We found virtually no consistent effects of plant identity on stable isotope composition of soil animals and on thirteen isotopic metrics that reflect general food‐web structure. However, in old fallows, the carbon isotope composition of some predatory macrofauna taxa had shifted closer to that of co‐occurring plants, which was particularly evident for Lasius, an aphid‐associated ant genus. Trophic levels and trophic‐chain lengths in food webs were similar across plant species and fallow ages. Overall, the results suggest that variations in local plant diversity of grassland communities may little affect the basal resources and the trophic level of prey consumed by individual species of meso‐ and macrofauna belowground. By contrast, successional changes in grassland communities are associated with shifts in the trophic niches of certain species, reflecting establishment of trophic interactions with time, which shapes the functioning and stability of soil food webs.

Identifying economic and ecological trade-offs of land-use transitions is important to ensure sustainability. Here, Grass et al. find biodiversity-profit trade-offs in tropical land-use transitions in Sumatra, and show that targeted landscape planning is needed to increase land-use efficiency while ensuring socio-ecological sustainability.

Agricultural expansion is among the main threats to biodiversity and functions of tropical ecosystems. It has been shown that conversion of rainforest into plantations erodes biodiversity, but further consequences for food-web structure and energetics of belowground communities remains little explored. We used a unique combination of stable isotope analysis and food-web energetics to analyze in a comprehensive way consequences of the conversion of rainforest into oil palm and rubber plantations on the structure of and channeling of energy through soil animal food webs in Sumatra, Indonesia. Across the animal groups studied, most of the taxa had lower litter-calibrated Δ13C values in plantations than in rainforests, suggesting that they switched to freshly-fixed plant carbon ('fast' energy channeling) in plantations from the detrital C pathway ('slow' energy channeling) in rainforests. These shifts led to changes in isotopic divergence, dispersion, evenness, and uniqueness. However, earthworms as major detritivores stayed unchanged in their trophic niche and monopolized the detrital pathway in plantations, resulting in similar energetic metrics across land-use systems. Functional diversity metrics of soil food webs were associated with reduced amount of litter, tree density, and species richness in plantations, providing guidelines on how to improve the complexity of the structure of and channeling of energy through soil food webs. Our results highlight the strong restructuring of soil food webs with the conversion of rainforest into plantations threatening soil functioning and ecosystem stability in the long term.

Abstract Altitudinal changes in the diversity of plants and animals have been well documented; however, soil animals received little attention in this context and it is unclear whether their diversity follows general altitudinal distribution patterns. Changbai Mountain is one of few well‐conserved mountain regions comprising natural ecosystems on the Eurasian continent. Here, we present a comprehensive analysis of the diversity and community composition of Collembola along ten altitudinal sites representing five vegetation types from forest to alpine tundra. Among 7834 Collembola individuals, 84 morphospecies were identified. Species richness varied marginally significant with altitude and generally followed a unimodal relationship with altitude. By contrast, the density of Collembola did not change in a consistent way with altitude. Collembola communities changed gradually with altitude, with local habitat‐related factors (soil and litter carbon‐to‐nitrogen ratio, litter carbon content, and soil pH) and climatic variables (precipitation seasonality) identified as major drivers of changes in Collembola community composition. Notably, local habitat‐related factors explained more variation in Collembola assemblages than climatic variables. The results suggest that local habitat‐related factors including precipitation and temperature are the main drivers of changes in Collembola communities with altitude. Specifically, soil and litter carbon‐to‐nitrogen ratio correlated positively with Collembola communities at high altitudes, whereas soil pH correlated positively at low altitudes. This documents that altitudinal gradients provide unique opportunities for identifying factors driving the community composition of not only above‐ but also belowground invertebrates.