For centuries ecologists have studied how the diversity and functional traits of plant and animal communities vary across biomes. In contrast, we have only just begun exploring similar questions for soil microbial communities despite soil microbes being the dominant engines of biogeochemical cycles and a major pool of living biomass in terrestrial ecosystems. We used metagenomic sequencing to compare the composition and functional attributes of 16 soil microbial communities collected from cold deserts, hot deserts, forests, grasslands, and tundra. Those communities found in plant-free cold desert soils typically had the lowest levels of functional diversity (diversity of protein-coding gene categories) and the lowest levels of phylogenetic and taxonomic diversity. Across all soils, functional beta diversity was strongly correlated with taxonomic and phylogenetic beta diversity; the desert microbial communities were clearly distinct from the nondesert communities regardless of the metric used. The desert communities had higher relative abundances of genes associated with osmoregulation and dormancy, but lower relative abundances of genes associated with nutrient cycling and the catabolism of plant-derived organic compounds. Antibiotic resistance genes were consistently threefold less abundant in the desert soils than in the nondesert soils, suggesting that abiotic conditions, not competitive interactions, are more important in shaping the desert microbial communities. As the most comprehensive survey of soil taxonomic, phylogenetic, and functional diversity to date, this study demonstrates that metagenomic approaches can be used to build a predictive understanding of how microbial diversity and function vary across terrestrial biomes.shotgun metagenomics | soil microbial ecology | 16S rRNA gene sequencing | biogeography

Soil biodiversity is increasingly recognized as providing benefits to human health because it can suppress disease-causing soil organisms and provide clean air, water and food. Poor land-management practices and environmental change are, however, affecting belowground communities globally, and the resulting declines in soil biodiversity reduce and impair these benefits. Importantly, current research indicates that soil biodiversity can be maintained and partially restored if managed sustainably. Promoting the ecological complexity and robustness of soil biodiversity through improved management practices represents an underutilized resource with the ability to improve human health.

Biodiversity and carbon (C) cycling have been the focus of much research in recent decades, partly because both change as a result of anthropogenic activities that are likely to continue. Soils are extremely speciesrich and store approximately 80% of global terrestrial C. Soil organisms play a key role in C dynamics and a loss of species through global changes could influence global C dynamics. Here, we synthesize findings from published studies that have manipulated soil species richness and measured the response in terms of ecosystem functions related to C cycling (such as decomposition, respiration and the abundance or biomass of decomposer biota) to evaluate the impact of biodiversity loss on C dynamics. We grouped studies where one or more biotic groups had been manipulated to include a richness of ≤10 species or >10 species in order to reflect 'low' and 'high' extents of diversity manipulations. There was a positive relationship between species richness and C cycling in 77-100% of low-diversity experiments, even when the richness of just one biotic group was manipulated, whereas positive relationships occurred less frequently in studies with greater richness (35-64%). Moreover, when positive relationships were observed, these often indicated functional redundancy at low extents of diversity or that community composition had a stronger influence on C cycling than did species richness. Initial reductions in soil species richness resulting from global changes are unlikely to alter C dynamics significantly unless particularly influential species are lost. However, changes in community composition, and the loss of species with an ability to facilitate specialized soil processes related to C cycling, as a result of global changes, may have larger impacts on C dynamics.

Altered precipitation patterns resulting from climate change will have particularly significant consequences in water-limited ecosystems, such as arid to semi-arid ecosystems, where discontinuous inputs of water control biological processes. Given that these ecosystems cover more than a third of Earth's terrestrial surface, it is important to understand how they respond to such alterations. Altered water availability may impact both aboveground and belowground communities and the interactions between these, with potential impacts on ecosystem functioning; however, most studies to date have focused exclusively on vegetation responses to altered precipitation regimes. To synthesize our understanding of potential climate change impacts on dryland ecosystems, we present here a review of current literature that reports the effects of precipitation events and altered precipitation regimes on belowground biota and biogeochemical cycling. Increased precipitation generally increases microbial biomass and fungal:bacterial ratio. Few studies report responses to reduced precipitation but the effects likely counter those of increased precipitation. Altered precipitation regimes have also been found to alter microbial community composition but broader generalizations are difficult to make. Changes in event size and frequency influences invertebrate activity and density with cascading impacts on the soil food web, which will likely impact carbon and nutrient pools. The long-term implications for biogeochemical cycling are inconclusive but several studies suggest that increased aridity may cause decoupling of carbon and nutrient cycling. We propose a new conceptual framework that incorporates hierarchical biotic responses to individual precipitation events more explicitly, including moderation of microbial activity and biomass by invertebrate grazing, and use this framework to make some predictions on impacts of altered precipitation regimes in terms of event size and frequency as well as mean annual precipitation. While our understanding of dryland ecosystems is improving, there is still a great need for longer term in situ manipulations of precipitation regime to test our model.

Aim To conduct the first global‐scale investigation of soil nematode assemblages using a standardized approach to quantify how environmental and climatic variables influence family assemblage structure in nematodes and determine whether nematode families have restricted distributions. Location Global. Methods We collected soil nematodes within four 10 m × 10 m plots distributed evenly along a 900‐m transect at each of 12 sites representing multiple ecosystem types across a latitudinal gradient (68° N to 77° S) on six continents. We assigned > 28,000 individuals to family level and trophic group morphologically. Results We recorded a total of 43 nematode families, but sites varied considerably in family richness (1–30). Families differed in their ranges with 12 families occurring at 10 or more sites, while 14 families occurred at three or fewer sites. Total nematode and trophic group abundances were generally related to soil characteristics, including bulk density and soil moisture, but we found no good predictor of family richness, diversity or evenness at the plot level. Family richness, diversity and evenness were considerably lower in the high‐latitude polar desert than elsewhere, but only family diversity showed a significant, albeit weak, latitudinal gradient. Nematode assemblage composition was quite strongly related to climate: 65% and 58% of the variation in assemblage composition across sites could be accounted for by mean annual rainfall and temperature, respectively. Main conclusions Nematode families display macroecological patterns similar to other organisms, such as a positive abundance–range size relationship and restricted distribution of some families. Local nematode abundances were related to soil characteristics, but we found no relationships between family richness and environmental or climatic variables. Family composition was related to mean annual rainfall and temperature, suggesting that climate is a good predictor of local assemblage structure. As a result, climate change may have a significant impact on nematode assemblages, with potential implications for ecosystem functioning.

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.

Soils represent a significant reservoir of biological diversity that underpins a broad range of key processes and moderate ecosystem service provision. Our understanding of the role that soil organisms play in ecosystems is still developing, but the increased investigation into biodiversity-ecosystem functioning relationships in soils over the past couple of decades has provided insights that have greatly enhanced our ability to sustainably manage soil biodiversity. In this review, we synthesize emerging knowledge of soil biodiversity as a natural resource that supports the functioning of terrestrial ecosystems and their delivery of ecosystem services. We explore how environmental changes alter soil biodiversity and how this in turn can affect ecosystem processes as well as resistance and resilience to environmental changes. We then discuss ways to include soil biodiversity in management strategies for sustainable production and biodiversity conservation. We conclude by highlighting key research challenges to further improve our knowledge of soil biodiversity and its management. 4.1

The polar regions are experiencing rapid climate change with implications for terrestrial ecosystems. Here, despite limited knowledge, we make some early predictions on soil invertebrate community responses to predicted twenty-first century climate change. Geographic and environmental differences suggest that climate change responses will differ between the Arctic and Antarctic. We predict significant, but different, belowground community changes in both regions. This change will be driven mainly by vegetation type changes in the Arctic, while communities in Antarctica will respond to climate amelioration directly and indirectly through changes in microbial community composition and activity, and the development of, and/ or changes in, plant communities. Climate amelioration is likely to allow a greater influx of non-native species into both the Arctic and Antarctic promoting landscape scale biodiversity change. Non-native competitive species could, however, have negative effects on local biodiversity particularly in the Arctic where the communities are already species rich. Species ranges will shift in both areas as the climate changes potentially posing a problem for endemic species in the Arctic where options for northward migration are limited. Greater soil biotic activity may move the Arctic towards a trajectory of being a substantial carbon source, while Antarctica could become a carbon sink.