Global climate and land use change are causing woody plant encroachment in arctic, alpine, and arid/semi-arid ecosystems around the world, yet our understanding of the belowground impacts of this phenomenon is limited. We conducted a globally distributed field study of 13 alpine sites across four continents undergoing woody plant encroachment and sampled soils from both woody encroached and nearby herbaceous plant community types. We found that woody plant encroachment influenced soil microbial richness and community composition across sites based on multiple factors including woody plant traits, site level climate, and abiotic soil conditions. In particular, root symbiont type was a key determinant of belowground effects, as Nitrogen-fixing woody plants had higher soil fungal richness, while Ecto/Ericoid mycorrhizal species had higher soil bacterial richness and symbiont types had distinct soil microbial community composition. Woody plant leaf traits indirectly influenced soil microbes through their impact on soil abiotic conditions, primarily soil pH and C:N ratios. Finally, site-level climate affected the overall magnitude and direction of woody plant influence, as soil fungal and bacterial richness were either higher or lower in woody encroached versus herbaceous soils depending on mean annual temperature and precipitation. All together, these results document global impacts of woody plant encroachment on soil microbial communities, but highlight that multiple biotic and abiotic pathways must be considered to scale up globally from site- and species-level patterns. Considering both the aboveground and belowground effects of woody encroachment will be critical to predict future changes in alpine ecosystem structure and function and subsequent feedbacks to the global climate system.

Cirad UMR EcoFoG Kourou French Guiana

College of Resources and Environmental Sciences Nanjing Agricultural University Nanjing China

Department of Botany and Plant Sciences University of California Riverside Riverside CA USA

Department of Environmental Sciences Physical Geography and Environmental Change University of Basel Basel Switzerland

Department of Evolution Ecology and Organismal Biology University of California Riverside Riverside CA USA

Department of Microbiology and Plant Pathology University of California Riverside Riverside CA USA

Environmental Earth Science Hokkaido University Sapporo Japan

Global Ecology Unit Campus de Bellaterra CREAF Barcelona Spain

Institut für Evolutionsbiologie und Umweltwissenschaften Universität Zürich Zürich Switzerland

Institute of Arctic and Alpine Research University of Colorado Boulder Boulder CO USA

Institute of Botany of the Czech Academy of Sciences Průhonice Czech Republic

Instituto Pirenaico de Ecología Zaragoza Spain

Pontificia Universidad Javeriana Bogotá Colombia

Red de Estudios Moleculares Avanzados Instituto de Ecología Pátzcuaro Mexico

Università degli Studi dell'Insubria Como Italy

University Montpellier AMAP INRAE CIRAD IRD CNRS Montpellier France

US Geological Survey Southwest Biological Science Center Moab UT USA

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