Among-tree variability and feedback effects result in different growth responses to climate change at the upper treeline in the Swiss Alps

Upper treeline ecotones are important life form boundaries and particularly sensitive to a warming climate. Changes in growth conditions at these ecotones have wide-ranging implications for the provision of ecosystem services in densely populated mountain regions like the European Alps. We quantify climate effects on short- and long-term tree growth responses, focusing on among-tree variability and potential feedback effects. Although among-tree variability is thought to be substantial, it has not been considered systematically yet in studies on growth-climate relationships. We compiled tree-ring data including almost 600 trees of major treeline species (Larix decidua, Picea abies, Pinus cembra, and Pinus mugo) from three climate regions of the Swiss Alps. We further acquired tree size distribution data using unmanned aerial vehicles. To account for among-tree variability, we employed information-theoretic model selections based on linear mixed-effects models (LMMs) with flexible choice of monthly temperature effects on growth. We isolated long-term trends in ring-width indices (RWI) in interaction with elevation. The LMMs revealed substantial amounts of previously unquantified among-tree variability, indicating different strategies of single trees regarding when and to what extent to invest assimilates into growth. Furthermore, the LMMs indicated strongly positive temperature effects on growth during short summer periods across all species, and significant contributions of fall (L.decidua) and current year's spring (L.decidua, P.abies). In the longer term, all species showed consistently positive RWI trends at highest elevations, but different patterns with decreasing elevation. L.decidua exhibited even negative RWI trends compared to the highest treeline sites, whereas P.abies, P.cembra, and P.mugo showed steeper or flatter trends with decreasing elevation. This does not only reflect effects of ameliorated climate conditions on tree growth over time, but also reveals first signs of long-suspected negative and positive feedback of climate change on stand dynamics at treeline.