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.

Biology Centre of the Czech Academy of Sciences Institute of Soil Biology and Biogeochemistry Na Sádkách 702 7 České Budějovice 37005 Czech Republic

CEFE Univ Montpellier CNRS EPHE IRD Univ Paul Valéry Montpellier 3 Route de Mende 34199 Montpellier Cedex 5 France

Centre Interdisciplinaire de Recherche sur la Montagne University of Lausanne Ch de l'Institut 18 Bramois Sion 1967 Switzerland

Department of Agricultural Forest and Food Sciences University of Turin Largo Paolo Braccini 2 Grugliasco 10095 Italy

Department of Biological Sciences Montana Technological University Butte 59701 MT USA

Department of Ecology and Evolution University of Lausanne Biophore Lausanne 1015 Switzerland

Department of Ecology Universität Innsbruck Technikerstrasse 25 Innsbruck 6020 Austria

Department of Microbiology Universität Innsbruck Technikerstrasse 25d Innsbruck 6020 Austria

Department of Soil Crop and Climate Sciences University of the Free State 205 Nelson Mandela Drive Bloemfontein 9300 South Africa

Dipartimento di Biologia Ecologia e Scienze della Terra University of Calabria Ponte Pietro Bucci 4b Rende 87036 Italy

Ecological Sciences The James Hutton Institute Craigiebuckler Aberdeen AB15 8QH Scotland UK

Forest Soils and Biogeochemistry Swiss Federal Research Institute WSL Birmensdorf 8903 Switzerland

Global Mountain Biodiversity Assessment University of Bern Altenbergrain 21 Bern 3013 Switzerland

Institute for Alpine Environment Eurac Research Viale Druso 1 Bozen Bolzano 39100 Italy

Institute of Biology Division of Plant Sciences University of Graz Holteigasse 6 Graz 8010 Austria

Institute of Earth Surface Dynamics University of Lausanne Géopolis Lausanne 1015 Switzerland

Institute of Plant Sciences University of Bern Altenbergrain 21 Bern 3013 Switzerland

Instituto Pirenaico de Ecología Avda Ntra Sra de la Victoria 16 Jaca 22700 Huesca Spain

J F Blumenbach Institute of Zoology and Anthropology University of Göttingen Untere Karspüle 2 Göttingen 37073 Germany

Key Laboratory of Vegetation Ecology Ministry of Education Northeast Normal University 2555 Jingyue Street Changchun 130117 China

Laboratory of Nematology Wageningen University and Research Droevendaalsesteeg 1 Wageningen 6708PB The Netherlands

Laboratory of Soil Biodiversity University of Neuchâtel Rue Emile Argand 11 Neuchâtel 2000 Switzerland

Northeast Institute of Geography and Agroecology Chinese Academy of Sciences 4888 Shengbei Street Changchun 130102 China

Research Center on Natural Risks in Mountain and Hilly Environments University of Turin Largo Paolo Braccini 2 Grugliasco 10095 Italy

School of Food Technology and Natural Sciences Massey University Riddett Road Palmerston North 4410 New Zealand

Terrestrial Ecology Cologne Biocenter University of Cologne Zülpicher Strasse 47b Cologne 50674 Germany

UMR 6553 EcoBio CNRS University of Rennes Biological Station Paimpont 35380 France

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