Non-linear response of Norway spruce to climate variation along elevational and age gradients in the Carpathians.

Climate change, namely increased warming coupled with a rise in extreme events (e.g., droughts, storms, heatwaves), is negatively affecting forest ecosystems worldwide. In these ecosystems, growth dynamics and biomass accumulation are driven mainly by environmental constraints, inter-tree competition, and disturbance regimes. Usually, climate-growth relationships are assessed by linear correlation due to the simplicity and straightforwardness of modeling. However, applying this method may bias results, since the ecological and physiological responses of trees to environmental factors are non-linear, and usually bell-shaped. In the Eastern Carpathian, Norway spruce is at the southeasternmost edge of its natural occurrence; this region is thus potentially vulnerable to climate change. A non-linear assessment of climate-growth relationships using machine-learning techniques for Norway spruce in this area had not been conducted prior to this study. To address this knowledge gap, we analyzed a large tree-ring network from 158 stands, with over 3000 trees of varying age distributed along an elevational gradient. Our results showed that non-linearity in the growth-climate response of spruce was season-specific: temperatures from the previous autumn and current growing season, along with water availability during winter, induced a bell-shaped response. Moreover, we found that at low elevations, spruce growth was mainly limited by water availability in the growing season, while winter temperatures are likely to have had a slight influence along the entire elevational gradient. Furthermore, at elevations lower than 1400 m, spruce trees were also found to be sensitive to previous autumn water availability. Overall, our results shed new light on the response of Norway spruce to climate in the Carpathians, which may aid in management decisions.