Stand Biomass at Treeline Ecotone in Russian Subarctic Mountains Is Primarily Related to Species Composition but Its Dynamics Driven by Improvement of Climatic Conditions

Abstract

Climate change effects are strongest in forest ecosystems at the limit of their distributions. Despite the evidence that treelines have shifted upwards by hundreds of meters, knowledge of the associated changes in the stand biomass is limited. In this study, stand biomass and changes to it during the last centuries were estimated along 20 altitudinal transects reaching from the historical (located in the 1950s–1960s) closed forest line up to the current treelines on mountain slopes of three subarctic regions of Russia (Kola Peninsula, Polar Urals, and Putorana Plateau) along a 2200 km long longitudinal gradient. The estimates were based on allometric measurements of 139 trees of five species (Betula pubescens Ehrh. ssp. tortuosa, Pinus sylvestris L., Picea abies Ledeb. ssp. obovata, Larix sibirica Ledeb., and Larix gmelinii Rupr.), stand structure assessments, and the demographic patterns of 9300 trees. During the 20th century, the growth and establishment of trees at the forest–mountain tundra transition (340–500 m width) increased exponentially. Since 1910 forest expansion and densification led to an accumulation of 621–748 tons of aboveground stand biomass per km of treeline length. The accumulation was two times higher below than above the contemporary closed forest line. Data analysis of weather stations showed that the 20th century’s climate had changed in a similar manner in the three study regions, namely vegetation periods became longer (8–10 days) and warmer (0.6–0.9 °C) and more snow fell in the cold period (+10–30%). Our results indicate that regional patterns in stand biomass at the treeline ecotone are primarily related to tree species composition as determined by macroclimatic conditions (e.g., continentality, sunshine hours), snowpack depth, and growing season duration. However, the stand biomass accumulation was driven by increases of early summer temperatures and early winter precipitation during the last century.