Water bears dominated cryoconite hole ecosystems: densities, habitat preferences and physiological adaptations of Tardigrada on an alpine glacier

We investigated the Forni Glacier and the surrounding area in the Alps in terms of habitat preferences, densities, dispersal and desiccation tolerance of glacier tardigrades, which are one of the most common faunal representatives and top consumers in supraglacial ecosystems. To do so, we sampled supraglacial environments (cryoconite holes, debris from ice surface, dirt cones and moraine, mosses from supraglacial stones) and non-glacial habitats (mosses, freshwater sediments and algae), and we installed air traps on the glacier and the nearby area. We found that cryoconite holes on the Forni Glacier are exclusively dominated by one metazoan group of tardigrades, representing one species, Hypsibius klebelsbergi (identified by morphological and molecular approaches). Tardigrades were found in 100% of cryoconite holes and wet supraglacial sediment samples and reached up to 172 ind./ml. Additionally, we found glacier tardigrades in debris from dirt cones and sparsely in supraglacial mosses. Glacier tardigrades were absent from freshwater and terrestrial samples collected from non-glacial habitats. Despite the fact that H. klebelsbergi is a typical aquatic species, we showed it withstands desiccation in sediments, but in low temperatures only. Treatments conducted in higher temperatures and water only showed low or no recovery. We suspect successful dispersal with wind might have taken place only when tardigrades desiccated in sediments and were passively transported by cold wind. Limited ability to withstand high temperatures and desiccation may be potential barriers preventing glacier tardigrades inhabiting new, even apparently suitable high mountain water bodies like temporary rock pools.