Global evidence that cold rocky landforms support icy springs in warming mountains

Climate change is reducing the extent of cold aquatic habitats and their unique biodiversity in mountain areas. However, a variety of cold rocky landforms (CRLs) are thermally buffered and feed cold springs (<2 ◦C) that may represent climate refugia for cold-adapted organisms. These landforms, hitherto overlooked by freshwater research, include rock glaciers, debris-covered glaciers, talus slopes, protalus ramparts, and young moraines. Here, we investigated the warm-season water temperature of 228 springs from clean (ice) glaciers, CRLs, and reference slopes (not sourced by any of these features) in 13 mountain ranges of Europe, South America, and North America. Only springs from glaciers (90%) and CRLs (45%) had average stream temperatures below the thermal optimum for coldwater organisms of 2 ◦C. Springs fed by CRLs were 3 ◦C–5 ◦C (up to 9 ◦C) colder than those from nearby reference slopes. In general, cold springs were rarer in Mediterranean/semi-arid climates than in temperate and sub-polar climates. Landforms comprising barren and coarse rocky surfaces or ice/rock mix, having a simple or absent soil/vegetation structure, and higher likelihood of permafrost more often supported cold springs. When water temperatures were compared to air temperature, most CRL springs were thermally buffered against warm periods, cumulative heat, and daily temperature fluctuations. With cold conditions maintained in a variety of climates and mountain landscapes,