Expanding forests and changing growth forms of Siberian larch at the Polar Urals treeline during the 20th century

The ongoing climatic changes potentially affect plant growth and the functioning of temperature-limited high-altitude and high-latitude ecosystems; the rate and magnitude of these biotic changes are, however, uncertain. The aim of this study was to reconstruct stand structure and growth forms of Larix sibirica (Ledeb.) in undisturbed forest-tundra ecotones of the remote Polar Urals on a centennial time scale. Comparisons of the current ecotone with historic photographs from the 1960s clearly document that forests have significantly expanded since then. Similarly, the analysis of forest age structure based on more than 300 trees sampled along three altitudinal gradients reaching from forests in the valleys to the tundra indicate that more than 70% of the currently upright-growing trees are < 80 years old. Because thousands of more than 500-year-old subfossil trees occur in the same area but tree remnants of the 15-19th century are lacking almost entirely, we conclude that the forest has been expanding upwards into the formerly tree-free tundra during the last century by about 20-60 m in altitude. This upward shift of forests was accompanied by significant changes in tree growth forms: while 36% of the few trees that are more than 100 years old were multi-stem tree clusters, 90% of the trees emerging after 1950 were single-stemmed. Tree-ring analysis of horizontal and vertical stems of multi-stemmed larch trees showed that these trees had been growing in a creeping form since the 15th century. In the early 20th century, they started to grow upright with 5-20 stems per tree individual. The incipient vertical growth led to an abrupt tripling in radial growth and thus, in biomass production. Based on above- and belowground biomass measurements of 33 trees that were dug out and the mapping of tree height and diameter, we estimated that forest expansion led to a biomass increase by 40-75 t ha(-1) and a carbon accumulation of approximately 20-40 g C m(-2) yr(-1) during the last century. The forest expansion and change in growth forms coincided with significant summer warming by 0.9 degrees C and a doubling of winter precipitation during the 20th century. In summary, our results indicate that the ongoing climatic changes are already leaving a fingerprint on the appearance, structure, and productivity of the treeline ecotone in the Polar Urals.