Aim We used a landscape-scale study of birch invasion onto heather moorland to determine the consistency of changes in vegetation type and soil properties and in the community composition of five soil organism groups. Our aim was to determine whether the degree to which soil organisms respond to natural changes and/or induced changes (e.g. changes in land-use type and climate) in habitat is consistent across trophic and taxonomic groups in the context of conservation policies for birch woodland and heather moorland. Location Mainland Scotland.Methods We sampled mesostigmatid mites, oribatid mites, fungi, bacteria and archaea in adjacent patches of birch woodland (dominated by Betula pubescens) and heather moorland (dominated by Calluna vulgaris) at 12 sites for which annual rainfall ranged between 713 and 2251 mm. Differences in community composition were visualized using non-metric multidimensional scaling based on Bray-Curtis dissimilarities. The factors contributing to differences between habitats within sites were explored using general linear models and those among sites using redundancy analysis. ResultsThe communities of all groups differed between habitats within sites, but only the oribatid mites and fungi differed consistently between habitats across sites. Within sites, dissimilarity in fungal communities was positively related to the difference in C. vulgaris cover between habitats, whereas dissimilarities in bacteria and archaea were positively related to differences in soil pH and C:N ratio between habitats, respectively. Main conclusionsThe influence of vegetation type and soil properties differed between groups of soil organisms, albeit in a predictable manner, across the 12 sites. Organisms directly associated with plants (fungi), and organisms with microhabitat and resource preferences (Oribatida) were strongly responsive to changes in habitat type. The response of organisms not directly associated with plants (bacteria, archaea) depended on differences in soil properties, while organisms with less clear microhabitat and resource preferences (Mesostigmata) were not strongly responsive to either vegetation type or soil properties. These results show that it is possible to predict the impact of habitat change on specific soil organisms depending on their ecology. Moreover, the community composition of all groups was related to variation in precipitation within the study area, which shows that external factors, such as those caused by climate change, can have a direct effect on belowground communities.

Increasing pressures on ecosystems through global climate and other land-use changes require predictive models of their consequences for vital processes such as soil carbon and nitrogen cycling. These environmental changes will undoubtedly affect soil fauna. There is sufficient evidence that soil fauna have significant effects on all of the pools and fluxes in these cycles, and soil fauna mineralize more N than microbes in some habitats. It is therefore essential that their role in the C and N cycle be understood. Here we introduce a new framework that attempts to reconcile our current understanding of the role of soil fauna within the C and N cycle with biogeochemical models and soil food web models. Using a simple stoichiometric approach to integrate our understanding of N mineralization and immobilization with the C:N ratio of substrates and faunal life history characteristics, as used in food web studies, we consider two mechanisms through which soil fauna can directly affect N cycling. First, fauna that are efficient assimilators of C and that have prey with similar C:N ratios as themselves, are likely to contribute directly to the mineral N pool. Second, fauna that are inefficient assimilators of C and that have prey with higher C:N ratios than themselves are likely to contribute most to the dissolved organic matter (DOM) pool. Different groups of fauna are likely to contribute to these two pathways. Protists and bacteria-feeding nematodes are more likely to be important for N mineralization through grazing on microbial biomass, while the effects of enchytraeids and fungal-feeding microarthropods are most likely to be important for DOM production. The model is consistent with experimental evidence and, despite its simplicity, provides a new framework in which the effects of soil fauna on pools and fluxes can be understood. Further, the model highlights our gaps in knowledge, not only for effects of soil fauna on processes, but also for understanding of the soil C and N cycle in general.

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Summary 1Single species can have a disproportionate effect on ecosystem function and diversity, yet our understanding of the importance of single species in driving terrestrial ecosystems during succession remains poor. 2 Utilizing a long-term experiment, where birch was planted on heather moorland 20 years ago, the cascading effects of a single tree species ( Betula pubescens ) on ecosystem characteristics (plant species richness, soil chemistry, soil fauna and decomposition rates) were tested. 3 Under the birch, plant species richness decreased and the vegetation composition changed, with lower cover of grasses and Vaccinium myrtillus . The depth of the soil organic horizon, its moisture content and percentage carbon were all smaller under the birch than under the heather. Concentrations of available phosphorus and mineralizable-N were significantly greater in the soil under birch than under the heather plots. Decomposition was faster in the birch than in the heather plots. The abundance and species richness of collembola and oribatid, mesostigmatid and prostigmatid mites were all significantly greater under the birch than under the heather. 4 The durability of the engineering effects of the birch was studied in a second experiment. Plots were established in first generation birch woodland that had developed on Calluna -dominanted moorland. The plots were cleared of birch and planted with heather. After 20 years soil chemical properties, microarthropod communities and decomposition rates were not significantly different between plots with and without the birch. However, the mass of the soil O-horizon was significantly greater in the felled birch plots than in the control birch plots, providing the first indication of a change towards soil properties more typical of a Calluna moorland. Thus for most of the birch engineering effects measured here their durability in the absence of the engineering species is at least 20 years. 5 This work has provided experimental evidence that birch acts as a top-down engineer, driving cascading effects on both above-and below-ground communities, soil chemical and physical properties and ecosystem processes. The work also shows that the role of birch in driving changes in the ecosystem is durable 20 years after the removal of the birch.

Background“The enigma of soil animal species diversity” was the title of a popular article by J. M. Anderson published in 1975. In that paper, Anderson provided insights on the great richness of species found in soils, but emphasized that the mechanisms contributing to the high species richness belowground were largely unknown. Yet, exploration of the mechanisms driving species richness has focused, almost exclusively, on above-ground plant and animal communities, and nearly 35 years later we have several new hypotheses but are not much closer to revealing why soils are so rich in species. One persistent but untested hypothesis is that species richness is promoted by small-scale environmental heterogeneity.Methodology/Principal FindingsTo test this hypothesis we manipulated small-scale heterogeneity in soil properties in a one-year field experiment and investigated the impacts on the richness of soil fauna and evenness of the microbial communities. We found that heterogeneity substantially increased the species richness of oribatid mites, collembolans and nematodes, whereas heterogeneity had no direct influence on the evenness of either the fungal, bacterial or archaeal communities or on species richness of the large and mobile mesostigmatid mites. These results suggest that the heterogeneity-species richness relationship is scale dependent.ConclusionsOur results provide direct evidence for the hypothesis that small-scale heterogeneity in soils increase species richness of intermediate-sized soil fauna. The concordance of mechanisms between above and belowground communities suggests that the relationship between environmental heterogeneity and species richness may be a general property of ecological communities.