Upward migration of plants to barren subnival areas is occurring worldwide due to raising ambient temperatures and glacial recession. In summer 2012, the presence of six vascular plants, growing in a single patch, was recorded at an unprecedented elevation of 6150 m.a.s.l. close to the summit of Mount Shukule II in the Western Himalayas (Ladakh, India). Whilst showing multiple signs of stress, all plants have managed to establish stable growth and persist for several years. To learn about the role of microbes in the process of plant upward migration, we analysed the root-associated microbial community of the plants (three individuals from each) using microscopy and tagged amplicon sequencing. No mycorrhizae were found on the roots, implying they are of little importance to the establishment and early growth of the plants. However, all roots were associated with a complex bacterial community, with richness and diversity estimates similar or even higher than the surrounding bare soil. Both soil and root-associated communities were dominated by members of the orders Sphingomonadales and Sphingobacteriales, which are typical for hot desert soils, but were different from communities of temperate subnival soils and typical rhizosphere communities. Despite taxonomic similarity on the order level, the plants harboured a unique set of highly dominant operational taxonomic units which were not found in the bare soil. These bacteria have been likely transported with the dispersing seeds and became part of the root-associated community following germination. The results indicate that developing soils act not only as a source of inoculation to plant roots but also possibly as a sink for plant-associated bacteria.

Department of Botany Faculty of Science University of South Bohemia Na Zlate stoce 1 37005 Ceske Budejovice Czech Republic

Division of Microbial Ecology Department of Microbiology and Ecosystem Science University of Vienna Althanstrasse 14 Vienna Austria

Institute of Botany The Czech Academy of Sciences Zámek 1 25243 Průhonice Czech Republic

Max Planck Institute for Terrestrial Microbiology Karl von Frisch Str 10 Marburg Germany

Swiss Federal Research Institute WSL Zuercherstrasse 111 8903 Birmensdorf Switzerland

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Highest Occurring Vascular Plants from Ladakh Provide Wood Anatomical Evidence for a Thermal Limitation of Cell Wall Lignification

Functionally distinct assembly of vascular plants colonizing alpine cushions suggests their vulnerability to climate change

Fungal root symbionts of high-altitude vascular plants in the Himalayas