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Motivation and aim: Soil biodiversity is central to ecosystem function and services. It represents most of terrestrial biodiversity and at least a quarter of all biodiversity on Earth. Yet, research into broad, generalizable spatial and temporal patterns of soil biota has been limited compared to aboveground systems due to complexities of the soil system. We review the literature and identify key considerations necessary to expand soil macroecology beyond the recent surge of global maps of soil taxa, so that we can gain greater insight into the mechanisms and processes shaping soil biodiversity. We focus primarily on three groups of soil taxa (earthworms, mycorrhizal fungi and soil bacteria) that represent a range of body sizes and ecologies, and, therefore, interact with their environment at different spatial scales. Results: The complexities of soil, including fine‐scale heterogeneity, 3‐D habitat structure, difficulties with taxonomic delimitation, and the wide‐ranging ecologies of its inhabitants, require the classical macroecological toolbox to be expanded to consider novel sampling, molecular identification, functional approaches, environmental variables, and modelling techniques. Main conclusions: Soil provides a complex system within which to apply macroecological research, yet, it is this property that itself makes soil macroecology a field ripe for innovative methodologies and approaches. To achieve this, soil‐specific data, spatio‐temporal, biotic, and abiotic considerations are necessary at all stages of research, from sampling design to statistical analyses. Insights into whole ecosystems and new approaches to link genes, functions and diversity across spatial and temporal scales, alongside methodologies already applied in aboveground macroecology, invasion ecology and aquatic ecology, will facilitate the investigation of macroecological processes in soil biota, which is key to understanding the link between biodiversity and ecosystem functioning in terrestrial ecosystems. [ABSTRACT FROM AUTHOR]

Theory suggests that more complex food webs promote stability and can buffer the effects of perturbations, such as drought, on soil organisms and ecosystem functions. Here, we tested experimentally how soil food web trophic complexity modulates the response to drought of soil functions related to carbon cycling and the capture and transfer below‐ground of recent photosynthate by plants. We constructed experimental systems comprising soil communities with one, two or three trophic levels (microorganisms, detritivores and predators) and subjected them to drought. We investigated how food web trophic complexity in interaction with drought influenced litter decomposition, soil CO2 efflux, mycorrhizal colonization, fungal production, microbial communities and soil fauna biomass. Plants were pulse‐labelled after the drought with 13C‐CO2 to quantify the capture of recent photosynthate and its transfer below‐ground. Overall, our results show that drought and soil food web trophic complexity do not interact to affect soil functions and microbial community composition, but act independently, with an overall stronger effect of drought. After drought, the net uptake of 13C by plants was reduced and its retention in plant biomass was greater, leading to a strong decrease in carbon transfer below‐ground. Although food web trophic complexity influenced the biomass of Collembola and fungal hyphal length, 13C enrichment and the net transfer of carbon from plant shoots to microbes and soil CO2 efflux were not affected significantly by varying the number of trophic groups. Our results indicate that drought has a strong effect on above‐ground–below‐ground linkages by reducing the flow of recent photosynthate. Our results emphasize the sensitivity of the critical pathway of recent photosynthate transfer from plants to soil organisms to a drought perturbation, and show that these effects may not be mitigated by the trophic complexity of soil communities, at least at the level manipulated in this experiment. [ABSTRACT FROM AUTHOR]

Urban soil biota can be surprisingly diverse and recent studies hypothesize that such biodiversity is partly due to stochastic community dynamics caused by fragmentation and high environmental variability. We aimed to quantify the relative effects of these factors on the community structure of soil oribatid mites inhabiting holm oak woodlands in two Mediterranean cities. We partitioned the community variation into fractions uniquely attributable to gradients in soil abiotic properties, pollution, microbiological properties, and spatial and temporal variation that could not be related to measured soil factors. As we found strong spatial structure at the local scales, a neutral model was fitted to test whether beta diversity patterns of relatively isolated communities were consistent with a purely stochastic assembly process. The overall diversity of the assemblage was remarkable: a total of 124 species were identified and rarefaction curves demonstrated that urban parks are as rich in species as their counterpart woodlands in suburban areas. The fraction of variation explainable in terms of soil properties, microbiology and pollution was statistically significant but surprisingly low. At the site scale, patterns of beta diversity (i.e. spatial turnover in species composition and relative abundances) were not significantly different from those predicted by neutral models. Stochastic models can parsimoniously predict background levels of urban soil biodiversity at local scales, while disturbance and environmental variation still play some significant but not major role at broader scales. [ABSTRACT FROM AUTHOR]

Beta diversity describes how local communities within an area or region differ in species composition/abundance. There have been attempts to use changes in beta diversity as a biotic indicator of disturbance, but lack of theory and methodological caveats have hampered progress. We here propose that the neutral theory of biodiversity plus the definition of beta diversity as the total variance of a community matrix provide a suitable, novel, starting point for ecological applications. Observed levels of beta diversity (BD) can be compared to neutral predictions with three possible outcomes: Observed BD equals neutral prediction or is larger (divergence) or smaller (convergence) than the neutral prediction. Disturbance might lead to either divergence or convergence, depending on type and strength. We here apply these ideas to datasets collected on oribatid mites (a key, very diverse soil taxon) under several regimes of disturbances. When disturbance is expected to increase the heterogeneity of soil spatial properties or the sampling strategy encompassed a range of diverging environmental conditions, we observed diverging assemblages. On the contrary, we observed patterns consistent with neutrality when disturbance could determine homogenization of soil properties in space or the sampling strategy encompassed fairly homogeneous areas. With our method, spatial and temporal changes in beta diversity can be directly and easily monitored to detect significant changes in community dynamics, although the method itself cannot inform on underlying mechanisms. However, human-driven disturbances and the spatial scales at which they operate are usually known. In this case, our approach allows the formulation of testable predictions in terms of expected changes in beta diversity, thereby offering a promising monitoring tool. [ABSTRACT FROM AUTHOR]

Abstract: The study of interrelationships between soil structure and its functional properties is complicated by the fact that the quantitative description of soil structure is challenging. Soil scientists have tackled this challenge by taking advantage of approaches such as fractal geometry, which describes soil architectural complexity through a scaling exponent (D) relating mass and numbers of particles/aggregates to particle/aggregate size. Typically, soil biologists use empirical indices such as mean weight diameters (MWD) and percent of water stable aggregates (WSA), or the entire size distribution, and they have successfully related these indices to key soil features such as C and N dynamics and biological promoters of soil structure. Here, we focused on D, WSA and MWD and we tested whether: D estimated by the exponent of the power law of number-size distributions is a good and consistent correlate of MWD and WSA; D carries information that differs from MWD and WSA; the fraction of variation in D that is uncorrelated with MWD and WSA is related to soil chemical and biological properties that are thought to establish interdependence with soil structure (e.g., organic C, N, arbuscular mycorrhizal fungi). We analysed observational data from a broad scale field study and results from a greenhouse experiment where arbuscular mycorrhizal fungi (AMF) and collembola altered soil structure. We were able to develop empirical models that account for a highly significant and large portion of the correlation observed between WSA and MWD but we did not uncover the mechanisms that underlie this correlation. We conclude that most of the covariance between D and soil biotic (AMF, plant roots) and abiotic (C, N) properties can be accounted for by WSA and MWD. This result implies that the ecological effects of the fragmentation properties described by D and generally discussed under the framework of fractal models can be interpreted under the intuitive perspective of simpler indices and we suggest that the biotic components mostly impacted the largest size fractions, which dominate MWD, WSA and the scaling exponent ruling number-size distributions. [Copyright &y& Elsevier]

Abstract: The effects of spatial scale on the assessment of soil biodiversity were investigated through a field study conducted in the Pelagian Islands, Sicilian Channel, southern Mediterranean. The oribatid mite community in this archipelago was investigated from both an ecological and a biogeographical point of view. The following hierarchically nested spatial scales were considered: 1—archipelago (γ-diversity); 2—island (α- and β-diversity between the islands); 3—sites (α- and β-diversity within each island). Quantitative and semi-quantitative soil sample replicates were collected during the wet season (autumn, 1999) in Lampedusa and Linosa, the two major islands of the archipelago. Data revealed that the sample α-diversity of the two islands is similar. The density of species followed a geometrical trend (a few dominant species with the remainder fairly uncommon) typical of communities in which a single environmental factor dominates species ecology. Community structure differed significantly between the two islands. Species turnover (β-diversity) was very high at every spatial scale, from the sites to the archipelago. Biogeographical results highlighted great differences between faunas in the two islands. The formulation of hypotheses for biodiversity patterns is strongly scale dependent: the heuristic and conservation value of the biogeographical approach increases from local to regional scales, because the importance of historical factors increases while that of ecological factors decreases. [Copyright &y& Elsevier]

Hedgerows structure agricultural landscape worldwide but little is known on their soil communities. In the intensively managed agricultural landscapes of Ireland and the UK, hedgerows protect soil from degradation and are thought to provide a reservoir of soil biodiversity for the farmed fields. But hedgerows are currently threatened by emergent tree diseases spread with rising volumes of trade in plants. This study analysed the effect of the most dominant hedgerow tree species on the composition and diversity of soil communities, which we estimated with oribatid mites in the context of an imminent ash dieback outbreak in Northern Ireland. We hypothesised that the environmental heterogeneity created by different tree species and good tree health are beneficial to the abundance and diversity of these soil communities. We found that communities differ, both in terms of species composition and relative abundances, between different trees in the trimmed hedgerows, and between untrimmed and trimmed hedgerows. The presence of trees was associated with more variable species composition. The analysis of phospholipid fatty acids (PLFA) showed that pasture and adjacent hedgerows were dominated by bacteria but there was no clear relationship between bacterial biomass and oribatid abundance or richness. However, changes in bacterial PLFAs were correlated positively with changes in oribatid community composition. Soil under ash trees which were diseased supported a higher species richness and greater abundance of oribatids than healthy trees. We conclude that the presence of different tree species in hedgerows adds to the overall diversity of soil hedgerow communities and hypothesise that the structural heterogeneity created by tree canopy drives increased diversity. Ill health in ash trees is currently positively affecting oribatid communities, possibly due to the temporarily increase in organic matter in soil. The epidemic of ash dieback, however, is expected to kill and remove trees in the medium term and so reduce soil community diversity in the long term. • Hedgerows host significant soil oribatid mite biodiversity. • Trimmed hedgerows and tree standards have distinct species composition. • The presence of trees in trimmed hedgerows increases biodiversity. • The positive short term effects of diseased tree may reverse in the long term. [ABSTRACT FROM AUTHOR]

• Diverse traits of Oribatida are maintained across native and non-native forest types. • Oribatida were trophically reorganized under non-native Douglas fir. • Litter-based resources of Douglas fir support more primary decomposers and surface-living Oribatida. • Tree mixtures can sustain Oribatida functions in forests with non-native tree species. 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. [ABSTRACT FROM AUTHOR]

The ecological interactions that occur in and with soil are of consequence in many ecosystems on the planet. These interactions provide numerous essential ecosystem services, and the sustainable management of soils has attracted increasing scientific and public attention. Although soil ecology emerged as an independent field of research many decades ago, and we have gained important insights into the functioning of soils, there still are fundamental aspects that need to be better understood to ensure that the ecosystem services that soils provide are not lost and that soils can be used in a sustainable way. In this perspectives paper, we highlight some of the major knowledge gaps that should be prioritized in soil ecological research. These research priorities were compiled based on an online survey of 32 editors of Pedobiologia – Journal of Soil Ecology. These editors work at universities and research centers in Europe, North America, Asia, and Australia. The questions were categorized into four themes: (1) soil biodiversity and biogeography, (2) interactions and the functioning of ecosystems, (3) global change and soil management, and (4) new directions. The respondents identified priorities that may be achievable in the near future, as well as several that are currently achievable but remain open. While some of the identified barriers to progress were technological in nature, many respondents cited a need for substantial leadership and goodwill among members of the soil ecology research community, including the need for multi-institutional partnerships, and had substantial concerns regarding the loss of taxonomic expertise. [ABSTRACT FROM AUTHOR]

Community coalescence is a recently introduced term describing the interaction of entire communities and their environments. We here explicitly place the concept of community coalescence in a soil microbial context, exploring intrinsic and extrinsic drivers of such coalescence events. Examples of intrinsic events include the action of earthworms and the dynamics of soil aggregates, while extrinsic events are exemplified by tillage, flooding, litter-fall, outplanting, and the addition of materials containing microbial communities. Aspects of global change may alter the frequency or severity of coalescence events. We highlight functional consequences of community coalescence in soil, and suggest ways to experimentally tackle this phenomenon. Soil ecology as a whole stands to benefit from conceptualizing soil biodiversity in terms of dynamic coalescent microbial assemblages. [ABSTRACT FROM AUTHOR]