Biological diversity in mountain ecosystems has been increasingly studied over the last decade. This is also the case for mountain soils, but no study to date has provided an overall synthesis of the current state of knowledge. Here we fill this gap with a first global analysis of published research on cryptogams, microorganisms, and fauna in mountain soils above the treeline, and a structured synthesis of current knowledge. Based on a corpus of almost 1400 publications and the expertise of 37 mountain soil scientists worldwide, we summarise what is known about the diversity and distribution patterns of each of these organismal groups, specifically along elevation, and provide an overview of available knowledge on the drivers explaining these patterns and their changes. In particular, we document an elevation-dependent decrease in faunal diversity above the treeline, while for cryptogams there is an initial increase above the treeline, followed by a decrease towards the nival belt. Thus, our data confirm the key role that elevation plays in shaping the biodiversity and distribution of these organisms in mountain soils. The response of prokaryote diversity to elevation, in turn, was more diverse, whereas fungal diversity appeared to be substantially influenced by plants. As far as available, we describe key characteristics, adaptations, and functions of mountain soil species, and despite a lack of ecological information about the uncultivated majority of prokaryotes, fungi, and protists, we illustrate the remarkable and unique diversity of life forms and life histories encountered in alpine mountain soils. By applying rule- as well as pattern-based literature-mining approaches and semi-quantitative analyses, we identified hotspots of mountain soil research in the European Alps and Central Asia and revealed significant gaps in taxonomic coverage, particularly among biocrusts, soil protists, and soil fauna. We further report thematic priorities for research on mountain soil biodiversity above the treeline and identify unanswered research questions. Building upon the outcomes of this synthesis, we conclude with a set of research opportunities for mountain soil biodiversity research worldwide. Soils in mountain ecosystems above the treeline fulfil critical functions and make essential contributions to life on land. Accordingly, seizing these opportunities and closing knowledge gaps appears crucial to enable science-based decision making in mountain regions and formulating laws and guidelines in support of mountain soil biodiversity conservation targets.

The GBNVPD (GIVD-code EU-GB-007) is the new centralised repository for the collection and storage of standardised vegetation plot data in Great Britain and the Crown Dependencies. The initial version of the GBNVPD (v1.0) described in this long database report comprises data from 63 individual sources and contains 277,070 samples from 200,733 plots, with a total of 4,463,300 occurrences of 4,086 accepted taxa (3,464 accepted species), surveyed between 1949 and 2024. Of the 63 constituent datasets 52 are newly submitted to the EVA and 9 to ReSurveyEurope. The database is available under a discretionary mixed-access regime and will be maintained and updated continuously in response to further digitisation of historic paper records, collection of pre-existing digitised survey data, and addition of future survey data. Abbreviations: EVA = European Vegetation Archive; GB = Great Britain; GBNVPD = National Vegetation Plot Database for Great Britain and the Crown Dependencies; NVC = National Vegetation Classification; UKCEH = United Kingdom Centre for Ecology and Hydrology; UKSI = United Kingdom Species Inventory.

Aims: We introduce ReSurveyEurope - a new data source of resurveyed vegetation plots in Europe, compiled by a collaborative network of vegetation scientists. We describe the scope of this initiative, provide an overview of currently available data, governance, data contribution rules, and accessibility. In addition, we outline further steps, including potential research questions. Results: ReSurveyEurope includes resurveyed vegetation plots from all habitats. Version 1.0 of ReSurveyEurope contains 283,135 observations (i.e., individual surveys of each plot) from 79,190 plots sampled in 449 independent resurvey projects. Of these, 62,139 (78%) are permanent plots, that is, marked in situ, or located with GPS, which allow for high spatial accuracy in resurvey. The remaining 17,051 (22%) plots are from studies in which plots from the initial survey could not be exactly relocated. Four data sets, which together account for 28,470 (36%) plots, provide only presence/absence information on plant species, while the remaining 50,720 (64%) plots contain abundance information (e.g., percentage cover or cover-abundance classes such as variants of the Braun-Blanquet scale). The oldest plots were sampled in 1911 in the Swiss Alps, while most plots were sampled between 1950 and 2020. Conclusions: ReSurveyEurope is a new resource to address a wide range of research questions on fine-scale changes in European vegetation. The initiative is devoted to an inclusive and transparent governance and data usage approach, based on slightly adapted rules of the well-established European Vegetation Archive (EVA). ReSurvey:Europe data are ready for use, and proposals for analyses of the data set can be submitted at any time to the coordinators. Still, further data contributions are highly welcome.

ABSTRACT Despite the importance of healthy soils for human livelihood, wellbeing, and safety, current gaps in our knowledge and understanding of biodiversity in soil are numerous, undermining conservation efforts. These gaps are particularly wide in mountain regions where healthy soils are especially important for human safety and yet evidence is accumulating of ongoing degradation, posing significant threats to ecosystem functioning and human settlements. To analyse these gaps in detail, we synthesise current research on the global diversity of microorganisms, cryptogams, and invertebrates in mountain soils above the treeline. This synthesis is based on a semi-quantitative survey of the literature and an expert-based analysis. Our work reveals not only deficiencies in geographic cover but also significant gaps in taxonomic coverage, particularly among soil protists and invertebrates, and a lack of (functional and ecological) description of the uncultivated majority of prokaryotes, fungi, and protists. We subsequently build on this overview to highlight opportunities for research on mountain soils as systems of co-occurring species that interact in complex environmental matrices to fulfil critical functions and make essential contributions to life on land. Closing gaps in biodiversity research in mountain soil is crucial to enhance our understanding and to promote laws and guidelines advancing international soil biodiversity conservation targets in mountains. Addressing sparse and biased data, recognizing the impact of environmental changes on mountain ecosystems, and advocating dedicated policies are essential strategies to safeguard mountain soils and their biodiversity. GLOSSARY

Understanding the chemical composition of our planet's crust was one of the biggest questions of the 20th century. More than 100 years later, we are still far from understanding the global patterns in the bioavailability and spatial coupling of elements in topsoils worldwide, despite their importance for the productivity and functioning of terrestrial ecosystems. Here, we measured the bioavailability and coupling of thirteen macro‐ and micronutrients and phytotoxic elements in topsoils (3–8 cm) from a range of terrestrial ecosystems across all continents (∼10,000 observations) and in response to global change manipulations (∼5,000 observations). For this, we incubated between 1 and 4 pairs of anionic and cationic exchange membranes per site for a mean period of 53 days. The most bioavailable elements (Ca, Mg, and K) were also amongst the most abundant in the crust. Patterns of bioavailability were biome‐dependent and controlled by soil properties such as pH, organic matter content and texture, plant cover, and climate. However, global change simulations resulted in important alterations in the bioavailability of elements. Elements were highly coupled, and coupling was predictable by the atomic properties of elements, particularly mass, mass to charge ratio, and second ionization energy. Deviations from the predictable coupling‐atomic mass relationship were attributed to global change and agriculture. Our work illustrates the tight links between the bioavailability and coupling of topsoil elements and environmental context, human activities, and atomic properties of elements, thus deeply enhancing our integrated understanding of the biogeochemical connections that underlie the productivity and functioning of terrestrial ecosystems in a changing world.

Ecological communities are being impacted by global change worldwide. Experiments are a powerful tool to understand how global change will impact communities by comparing control and treatment replicates. Communities consist of multiple species, and their associated abundances make multivariate methods an effective approach to study community compositional differences between control and treated replicates. Dissimilarity metrics are a commonly employed multivariate measure of compositional differences; however, while highly informative, dissimilarity metrics do not elucidate the specific ways in which communities differ. Integrating two multivariate methods, dissimilarity metrics and rank abundance curves (RACs), have the potential to detect complex differences based on dissimilarity metrics and detail the how these differences came about through differences in richness, evenness, species ranks, or species identity. Here we use a database of 106 global change experiments located in herbaceous ecosystems and explore how patterns of ordinations based on dissimilarity metrics relate to RAC-based differences. We find that combining dissimilarity metrics alongside RAC-based measures clarifies how global change treatments are altering communities. We find that when there is no difference in community composition (no distance between centroids of control and treated replicates), there are rarely differences in species ranks or species identities and more often differences in richness or evenness alone. In contrast, when there are differences between centroids of control and treated replicates, this is most often associated with differences in ranks either alone or co-occurring with differences in richness, evenness, or species identities. We suggest that integrating these two multi-variate measures of community composition results in a deeper understanding of how global change impacts communities.

Moss-sedge heath, an important habitat of oceanic alpine regions, is under threat from elevated nitrogen (N) deposition and rising temperatures increasing decomposition rates and fragmentation of the extensive, thick moss mats of Racomitrium lanuginosum which characterise this habitat. Here, we examine the potential effects of N deposition and temperature on fungal communities associated with R. lanuginosum mats. Fungal community composition in both green moss shoots and underlying soil were significantly influenced by mat temperatures and green moss shoot N contents. Total OTU, Ascomycota, Basidiomycota and saprotrophic fungal richness in shoots were all positively related to temperature, while in soil, the richness of all groups, except Basidiomycota, was positively related to moss tissue N. The observed community changes suggest that increased N loading and increasing temperatures are ameliorating growing conditions for fungi associated with moss-sedge heath. Further study is required to determine if the observed changes in fungal communities contribute to accelerated decomposition of moss mats and degradation of the habitat.

Global change is impacting plant community composition, but the mechanisms underlying these changes are unclear. Using a dataset of 58 global change experiments, we tested the five fundamental mechanisms of community change: changes in evenness and richness, reordering, species gains and losses. We found 71% of communities were impacted by global change treatments, and 88% of communities that were exposed to two or more global change drivers were impacted. Further, all mechanisms of change were equally likely to be affected by global change treatments-species losses and changes in richness were just as common as species gains and reordering. We also found no evidence of a progression of community changes, for example, reordering and changes in evenness did not precede species gains and losses. We demonstrate that all processes underlying plant community composition changes are equally affected by treatments and often occur simultaneously, necessitating a wholistic approach to quantifying community changes.

Summary Soil carbon (C) pools and plant community composition are regulated by nitrogen (N) and phosphorus (P) availability. Atmospheric N deposition impacts ecosystem C storage, but the direction of response varies between systems. Phosphorus limitation may constrain C storage response to N, hence P application to increase plant productivity and thus C sequestration has been suggested. We revisited a 23‐yr‐old field experiment where N and P had been applied to upland heath, a widespread habitat supporting large soil C stocks. At 10 yr after the last nutrient application we quantified long‐term changes in vegetation composition and in soil and vegetation C and P stocks. Nitrogen addition, particularly when combined with P, strongly influenced vegetation composition, favouring grasses over Calluna vulgaris, and led to a reduction in vegetation C stocks. However, soil C stocks did not respond to nutrient treatments. We found 40% of the added P had accumulated in the soil. This study showed persistent effects of N and N + P on vegetation composition, whereas effects of P alone were small and showed recovery. We found no indication that P application could mitigate the effects of N on vegetation or increase C sequestration in this system.

Empirical critical loads are widely used to quantify and manage the ecological impacts of reactive nitrogen (N) deposition. Critical load values aim to identify a level of N deposition below which significant harmful effects do not occur according to present knowledge. Critical loads have been primarily based on experiments, but these are few in number and have well-known limitations, so there is a strong imperative to test and validate values with other forms of evidence. We assembled data on the spatial variability in vegetation communities in the United Kingdom and used Threshold Indicator Taxa Analyses (TITAN) to investigate linkages between species changes and modelled current and cumulative N deposition. Our analyses focused on five datasets: acid grasslands, alpine habitats, coastal fixed dunes, dune slacks and wet grasslands. In four of these habitats there was evidence for a significant decline in the cover of at least one species (a ‘species-loss change-point’) occurring below the critical load, and often at very low levels of N deposition. In all of the habitats there was evidence for clustering of many individual species-loss change-points, implying a community change-point analogous to an ecological threshold. Three of these community change-points occurred below the critical load and the remaining two overlapped with the critical load range. Studies using similar approaches are now increasingly common, with similar results. Across 19 similar analyses there has been evidence for plant species loss change-points below the critical load in 18 analyses, and community-level species loss change-points below the critical load in 13 analyses. None of these analyses has shown community change-points above the critical load. Field data increasingly suggest that many European critical loads are too high to confidently prevent loss of sensitive species.

Nitrogen (N) deposition poses a severe risk to global terrestrial ecosystems, and managing this threat is an important focus for air pollution science and policy. To understand and manage the impacts of N deposition, we need metrics which accurately reflect N deposition pressure on the environment, and are responsive to changes in both N deposition and its impacts over time. In the UK, the metric typically used is a measure of total N deposition over 1-3 years, despite evidence that N accumulates in many ecosystems and impacts from low-level exposure can take considerable time to develop. Improvements in N deposition modelling now allow the development of metrics which incorporate the long-term history of pollution, as well as current exposure. Here we test the potential of alternative N deposition metrics to explain vegetation compositional variability in British semi-natural habitats. We assembled 36 individual datasets representing 48,332 occurrence records in 5479 quadrats from 1683 sites, and used redundancy analyses to test the explanatory power of 33 alternative N metrics based on national pollutant deposition models. We find convincing evidence for N deposition impacts across datasets and habitats, even when accounting for other large-scale drivers of vegetation change. Metrics that incorporate long-term N deposition trajectories consistently explain greater compositional variance than 1-3 year N deposition. There is considerable variability in results across habitats and between similar metrics, but overall we propose that a thirty-year moving window of cumulative deposition is optimal to represent impacts on plant communities for application in science, policy and management. Crown Copyright (C) 2019 Published by Elsevier Ltd. All rights reserved.

Forests play a key role in climate change mitigation, adaptation and delivery of a range of ecosystem services. There is increasing evidence for impacts of climate and other drivers on plant community change, and fragmented habitats are predicted to be much less resilient to negative impacts on biodiversity and other services. Within Europe, Scotland's native forests are highly fragmented and now cover 4% of the land after many centuries of degradation and loss, but little is known about how their species composition has changed. We recorded long-term vegetation change (from resurvey data) and examined the relationships with climate, pollutant deposition and grazing as key drivers of change, focusing on four forest types: pine, ash, acid- and base-rich oak birch. All four forest types showed dynamic compositional change during 30-50 years between surveys, with increased species richness and decreased diversity. There was no evidence for homogenisation - the opposite was the case for all except pine (no change). Analyses indicate significant and varied climate, pollution and grazing impacts; NHy deposition showed the most frequent association with species compositional changes. Notable species changes include increases in pteridophytes and declines in forb cover, and a doubling in frequency and cover of Fagus sylvatica between surveys. Our findings suggest a possible extinction debt, with many more species declining than increasing between surveys. This trajectory of change and our other findings indicate a pressing need for mitigation management to reduce the risks of future species losses, with forest expansion planning explicitly considering spatial location in relation to existing native forest and those plant species identified as most at risk.

Understanding the full range of biodiversity patterns from local to global scales, through the study of the drivers of multiscale plant community composition and diversity, is a current goal of biogeography. A synthetic understanding of to what extent vegetation compositional patterns are produced by biotic factors, geography, or climate and how these patterns vary across scales is needed. This lack hinders prediction of the effects of climate change in global vegetation. Variation in community composition is examined in relation to climatic difference and geographic distance at hemispheric and continental scales. Vascular plants and bryophytes in thirteen mountain regions were analyzed: eight in Europe and five in North America, nine midlatitude and four oroarctic. Species composition differed between continents and between oroarctic and midlatitude regions. Patterns of paired regional similarity with distance were significant for all pairs and intercontinental pairs but not for those within Europe and North America. Climatic variables accounted for most of the variance in vegetation patterns revealed by general linear models of ordinations, but geographic variables of Moran eigenvectors and latitudinal zones were also important and significant. The effects of geography were typically twice as strong for vascular plants as for bryophytes. The importance of geography at these scales suggests that past evolutionary and ecological processes are as important as current fit to any climatic niche. Interpretation of observations of the impacts of global climate change should recognize geographic context and phylogeny, and policies to mitigate them, such as assisted migration, should be cautious. Key Words: Alpine, beta diversity, climate, distance, oroarctic.

Deposition of reactive nitrogen (N) is a major threat to terrestrial ecosystems associated with impacts on ecosystem properties and functions including carbon (C) and nutrient stocks, soil water quality and nutrient retention. In the oceanic-alpine Racomitrium heath habitat, N deposition is associated with moss mat degradation and a shift from bryophyte to graminoid dominance. To investigate the effects of moss mat decline on C and N stocks and fluxes, we collected Racomitrium heath vegetation soil cores from sites along a gradient of N deposition in the UK. Cores were maintained under controlled conditions and exposed to scenarios of current (8-40 kg N ha(-1) y(-1)), reduced (8 kg N ha(-1) y(-1)) and elevated (50 kg N ha(-1) y(-1)) N deposition. Cores from high N deposition sites had smaller aboveground C and N stocks and, under current conditions, leached large amounts of inorganic N and had low soil water pH compared with low N deposition sites. With reduced N deposition there was evidence for rapid recovery of soil water quality in terms of reduced N leaching and small increases in pH. Under high N deposition, cores from low N deposition sites retained much of the applied N while those with a history of high N deposition leached large amounts of inorganic N. Carbon fluxes in soil water and net CO2 fluxes varied according to core source site but were not affected by the N deposition scenarios. We conclude that C and N stocks and cycling in Racomitrium heath are strongly affected by long-term exposure to N deposition but that soil water quality may improve rapidly, if N deposition rates are reduced. The legacy of N deposition impacts on moss mat cover and vegetation composition however, mean that the ecosystem remains sensitive to future pulses in N input. (C) 2019 Elsevier Ltd. All rights reserved.

Significance Accurate prediction of community responses to global change drivers (GCDs) is critical given the effects of biodiversity on ecosystem services. There is consensus that human activities are driving species extinctions at the global scale, but debate remains over whether GCDs are systematically altering local communities worldwide. Across 105 experiments that included over 400 experimental manipulations, we found evidence for a lagged response of herbaceous plant communities to GCDs caused by shifts in the identities and relative abundances of species, often without a corresponding difference in species richness. These results provide evidence that community responses are pervasive across a wide variety of GCDs on long-term temporal scales and that these responses increase in strength when multiple GCDs are simultaneously imposed.

1. Assisted colonisation, one form of species translocation, has been proposed as a tool for helping species to track suitable conditions in a changing climate. There are considerable practical challenges associated with it, including predicting where to place translocated individuals. This problem may be particularly big for small and immobile species, where small-scale microenvironmental conditions de-couple them from environmental conditions as projected in large-scale climate models. 2. To investigate this problem, we developed a survey-based model to predict the occurrence of our target species, the fruticose terricolous arctic-alpine lichen, Flavocetraria nivalis, within the Cairngorm Mountains. 3. We then undertook an experimental translocation of this species. A second model, using variables that were significant in the survey-based model, was only fair at predicting the initial pattern of survival at the recipient site. 4. However, model fit of the translocation survival model improved over time as the distribution of surviving individuals more accurately reflected the distribution of suitable environmental conditions. In addition, model predictive power increased with the addition of data on microclimatic conditions at recipient plots. 5. Synthesis and applications. Our results demonstrate that, for species which respond strongly to local environmental conditions, are immobile and, to some extent, decoupled from larger scale climates, it may be difficult to build a priori accurate predictive models of habitat suitability. In these cases, a combination of modelling and expert judgement, along with the movement of substantial numbers of transplants, may be the appropriate options for maximising the success of assisted colonisation.