Salix shrub encroachment along a 1000 m elevation gradient triggers a major ecosystem change in the European Alps

Shrub encroachment, a globally recognized response to climate warming, usually involves late successional species in mountain environments, without alteration to climax communities. We show that a major ecosystem change is occurring in the European Alps across a 1000 m elevation gradient, with pioneer hygrophilous Salix shrubs, previously typical of riparian forests, wetlands and avalanche ravines, encroaching into the climax communities of subalpine and alpine belts shrublands and grasslands, as well as snowbeds, pioneer vegetation and barren grounds in the nival belt. We analyzed Salix recruitment through dendrochronological methods, and assessed its relationships with climate and atmospheric CO2 concentration. The dendrochronological data indicated that Salix encroachment commenced in the 1950s (based on the age of the oldest Salix individuals, recruited in 1957), and that it was correlated with increasing atmospheric CO2 concentration, spring warming and snow cover decrease. Hygrophilous Salix shrubs are expanding their distribution both through range filling and upwards migration, likely achieving competitive replacement of species of subalpine and alpine climax communities. They benefit from climate warming and CO2 fertilization and are not sensitive to spring frost damage and soil limitations, being observed across a gradient of soil conditions from loose glacial sediments in recently deglaciated areas (where soils had not had sufficient time to develop) to mature soils such as podzols (when colonizing late successional subalpine shrublands). Salix encroachment may trigger ecosystem and landscape transformations, promoting the development of forests that replace pre-existing subalpine shrublands, and of open woodlands invading alpine grasslands and snowbeds, making the alpine environment similar to sub-Arctic and Arctic areas. This results in a new threat to the conservation of the plant species, communities and landscapes typical of the alpine biota, as mountain ranges such as the Alps provide limited opportunities for upward migration and range-shift.