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Changes in the composition, structure, and altitudinal distribution of low forests at the upper limit of their growth over the past 50 years have been studied in the Tylaisko-Konzhakovsko-Serebryanskii Massif (the Northern Urals). The qualitative and quantitative assessment of these changes has been made on the basis of descriptions, photographs, and maps made in 1956 and 2005. The results show that the upper boundary of low forests on the majority of slopes has ascended. Considerable changes have occurred in the composition, density, and height of the tree layer in the communities that formed the upper low-forest boundary in 1956. Among a fairly large number of tree species growing in the subgoltzy belt (Picea obovata, Larix sibirica, Pinus sibirica, and Abies sibirica), the birch Betula tortuosa has expanded most actively, whereas the proportions of P. obovata and especially L. sibirica in low forests have decreased. These changes are explained by climate warming and increasing humidity.

Climate warming is shifting the elevational boundary between forests and tundra upwards, but the related belowground responses are poorly understood. In the pristine South and Polar Urals with shifts of the treeline ecotone documented by historical photographs, we investigated fine root dynamics and production of extramatrical mycorrhizal mycelia (EMM) along four elevational transects reaching from the closed forest to the treeless tundra. In addition, we analysed elevational differences in climate and vegetation structure, and excavated trees to estimate related changes in the partitioning between below- and aboveground biomass. Fine root biomass of trees (<2 mm) increased by 13-79% with elevation, paralleled by a 35-72% increase in ground vegetation fine roots from the closed forest to the tundra. During the first year of decomposition, mass loss of fine root litter from different vegetation types was greater at lower elevations in the forest-tundra ecotone. The ratio between fine roots of trees and stem biomass largely increased with elevation in both regions, but these increases were not accompanied by a distinct production of EMM. Production of EMM, however, increased with the presence of ectomycorrhizal trees at the transition from the tundra to the forest. Our results imply that the recorded upward expansion of forest into former tundra in the Ural Mountains by 4-8 m per decade is decreasing the partitioning of plant biomass to fine roots. They further suggest that climate-driven forest advances will alter EMM production rates with potential feedbacks on soil carbon and nutrient cycling in these ecosystems.

The composition and structure of tree stands near the timberline have been studied on different slopes and at different elevations in the Tylaisko-Konzhakovsko-Serebryanskii Massif, the North Urals. It has been found that the upper limits of tree stands with different degrees of canopy closure have risen considerably (by about 100 m of elevation) since the mid-19th century, although the formation of these stands started as early as the late 18th century. Woodless areas in the eastern part of the massif started to be colonized by Larix sibirica in the late 18th to early 19th centuries; those in the western part, by Picea obovata in the mid-19th century; and in the southern part, by Betula tortuosa in the late 19th century. Analysis of meteorological data provides evidence for warming and increasing humidity of the climate since the late 19th century. Favorable climatic changes that facilitated the expansion of the forest have taken place both in the summer (prolongation of the growing period) and in winter seasons (increase of air temperature and precipitation). The observed differences in the composition and dynamics of tree stands between the studied areas of the mountain range are most probably explained by different requirements of tree species for the depth of snow cover and the degree of soil freezing.

AbstractHigh‐altitude treelines are temperature‐limited vegetation boundaries, but little quantitative evidence exists about the impact of climate change on treelines in untouched areas of Russia. Here, we estimated how forest‐tundra ecotones have changed during the last century along the Ural mountains. In the South, North, Sub‐Polar, and Polar Urals, we compared 450 historical and recent photographs and determined the ages of 11 100 trees along 16 altitudinal gradients. In these four regions, boundaries of open and closed forests (crown covers above 20% and 40%) expanded upwards by 4 to 8 m in altitude per decade. Results strongly suggest that snow was an important driver for these forest advances: (i) Winter precipitation has increased substantially throughout the Urals (~7 mm decade−1), which corresponds to almost a doubling in the Polar Urals, while summer temperatures have only changed slightly (~0.05 °C decade−1). (ii) There was a positive correlation between canopy cover, snow height and soil temperatures, suggesting that an increasing canopy cover promotes snow accumulation and, hence, a more favorable microclimate. (iii) Tree age analysis showed that forest expansion mainly began around the year 1900 on concave wind‐sheltered slopes with thick snow covers, while it started in the 1950s and 1970s on slopes with shallower snow covers. (iv) During the 20th century, dominant growth forms of trees have changed from multistemmed trees, resulting from harsh winter conditions, to single‐stemmed trees. While 87%, 31%, and 93% of stems appearing before 1950 were from multistemmed trees in the South, North and Polar Urals, more than 95% of the younger trees had a single stem. Currently, there is a high density of seedlings and saplings in the forest‐tundra ecotone, indicating that forest expansion is ongoing and that alpine tundra vegetation will disappear from most mountains of the South and North Urals where treeline is already close to the highest peaks.

AbstractAimIn the alpine life zone, plant diversity is strongly determined by local topography and microclimate. We assessed the extent to which aspect and its relatedness to temperature affect plant species diversity, and the colonization and disappearance of species on alpine summits on a pan‐European scale.LocationMountain summits in Europe's alpine life zone.MethodsVascular plant species and their percentage cover were recorded in permanent plots in each cardinal direction on 123 summits in 32 regions across Europe. For a subset from 17 regions, resurvey data and 6‐year soil temperature series were available. Differences in temperature sum and Shannon index as well as species richness, colonization and disappearance of species among cardinal directions were analysed using linear mixed‐effects and generalised mixed‐effects models, respectively.ResultsTemperature sums were higher in east‐ and south‐facing aspects than in the north‐facing ones, while the west‐facing ones were intermediate; differences were smallest in northern Europe. The patterns of temperature sums among aspects were consistent among years. In temperate regions, thermal differences were reflected by plant diversity, whereas this relationship was weaker or absent on Mediterranean and boreal mountains. Colonization of species was positively related to temperature on Mediterranean and temperate mountains, whereas disappearance of species was not related to temperature.Main conclusionsThermal differences caused by solar radiation determine plant species diversity on temperate mountains. Advantages for plants on eastern slopes may result from the combined effects of a longer diurnal period of radiation due to convection cloud effects in the afternoon and the sheltered position against the prevailing westerly winds. In northern Europe, long summer days and low sun angles can even out differences among aspects. On Mediterranean summits, summer drought may limit species numbers on the warmer slopes. Warmer aspects support a higher number of colonization events. Hence, aspect can be a principal determinant of the pace of climate‐induced migration processes.

Changes in the altitudinal position of the timberline in high mountain areas of the Nether-Polar Urals and basic factors that influence such changes have been revealed on the basis of comparison of the age structure of Siberian larch (Larix sibirica) and arctic birch (Betula tortuosa) tree stands and photographs made in different years. On the mountain slopes studied, an upward shift of the timberline took place in areas covered in winter with thick snow (in the late 18th century), with Siberian larch being the pioneer species. Larch began colonizing areas with a thin snow cover in the 20th century. Birch appeared later and has since strengthened its positions. The increase in winter temperatures and precipitation facilitated the expansion of the forest.

The correlation between climatic changes and the numbers of trees in the generations of Siberian spruce (Picea obovata Ledeb.) in subgoltsy tree stands of the Southern Urals was analyzed over the past 200 years. The results showed that the most abundant generations appeared in 1809–1816, 1821–1827, 1839–1845, 1850–1867, 1875–1887, 1891–1896, 1900–1904, 1911–1918, 1923–1932, 1944–1952, 1958–1966, and 1975–1995. Their formation proved to be related to the improvement of thermal conditions of the warm season during the five to seven years before and after the emergence of trees and conditions of the cold season in the year of their emergence, as well as to the periods of increased snow depth in late April in the years 27–32 of tree growth.

Within the ecotone of the upper limit of woody vegetation on the southeastern macroslope of the Khibiny Mountain ridge (Kola Peninsula), the spatial and age structure, as well as features of the phytomass accumulation of spruce–birch stands, were studied. Analysis revealed that there was a manifold increase in the density and productivity of forest stands in the last century, and the upper border of the woodlands and dense forests has moved considerably higher into the mountains. All of this happened against the background of an increase in early summer temperatures and a longer growing season in the area in the 20th century. Our data will help simulate the response of mountain ecosystems in the region to future climate change.