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Recent climatic changes significantly affected forest ecosystems in northern Eurasia. Trees growing in Siberia are very sensitive to climate change due to strong temperature limitation of their growth. Our study covers high-latitude (northeastern Yakutia, eastern Taimyr, central Evenkia) and high-altitude (Russian Altai) zones in Eurasia, where tree-ring parameters (tree-ring width, cell-wall thickness, and maximum latewood density) mainly record summer air temperature variations. To reveal the impact of moisture changes (e.g., amount of precipitation, vapor pressure deficit, relative humidity and potential evapotranspiration) on tree growth in Siberian forest ecosystems, we evaluated δ13C in tree-ring cellulose over the past century. We found that at all the study sites mainly June-July precipitation and June-July evapotranspiration affect larch radial growth, while the strongest influence of vapor pressure deficit on the δ13C was observed in northeastern Yakutia. Further increase of vapor pressure deficit and rise of air temperature in the coming decades in Siberian regions will probably lead to drought and related forest mortality even under additional source of water due to permafrost thaw.

Although it has been recognized that rising temperatures and shifts in the hydrological cycle affect the depth of the seasonally thawing upper permafrost stratum, it remains unclear to what extent the frequency and intensity of wildfires, and subsequent changes in vegetation cover, influence the soil active layer on different spatiotemporal scales. Here, we use ring width measurements of the subterranean stem part of 15 larch trees from a Sphagnum bog site in Central Siberia to reconstruct long-term changes in the thickness of the active layer since the last wildfire occurred in 1899. Our approach reveals a three-step feedback loop between above- and belowground ecosystem components. After all vegetation is burned, direct atmospheric heat penetration over the first ~20 years caused thawing of the upper permafrost stratum. The slow recovery of the insulating ground vegetation reverses the process and initiates a gradual decrease of the active layer thickness. Due to the continuous spreading and thickening of the peat layer during the last decades, the upper permafrost horizon has increased by 0.52 cm/year. This study demonstrates the strength of annually resolved and absolutely dated tree-ring series to reconstruct the effects of historical wildfires on the functioning and productivity of boreal forest ecosystems at multi-decadal to centennial time-scale. In so doing, we show how complex interactions of above- and belowground components translate into successive changes in the active permafrost stratum. Our results are particularly relevant for improving long-term estimates of the global carbon cycle that strongly depends on the source and sink behavior of the boreal forest zone.

Though the extraction of increment cores is common practice in tree-ring research, there is no standard for the number of samples per tree, or trees per site needed to accurately describe the common growth pattern of a discrete population of trees over space and time. Tree-ring chronologies composed of living, subfossil and archaeological material often combine an uneven distribution of increment cores and disc samples. The effects of taking one or two cores per tree, or even the inclusion of multiple radii measurements from entire discs, on chronology development and quality remain unreported. Here, we present four new larch (Larix cajanderi Mayr) ring width chronologies from the same 20 trees in northeastern Siberia that have been independently developed using different combinations of core and disc samples. Our experiment reveals: i) sawing is much faster than coring, with the latter not always hitting the pith; ii) the disc-based chronology contains fewer locally absent rings, extends further back in time and exhibits more growth coherency; iii) although the sampling design has little impact on the overall chronology behaviour, lower frequency information is more robustly obtained from the disc measurements that also tend to reflect a slightly stronger temperature signal. In quantifying the influence of sampling strategy on the quality of tree-ring width chronologies, and their suitability for climate reconstructions, this study provides useful insights for optimizing fieldwork campaigns, as well as for developing composite chronologies from different wood sources.

Tracheids fulfil most wood functions in conifers (mechanical support and water transport) and earlywood tracheids account for most hydraulic conductivity within the annual tree ring. Dry conditions during the early growing season, when earlywood is formed, could lead to the formation of narrow tracheid lumens and a dense earlywood. Here, we assessed if there is a negative association between minimum wood density and early growing-season (spring) precipitation. Using dendrochronology, we studied growth and density data at nine forest stands of three Pinaceae species (Larix sibirica, Pinus nigra, and Pinus sylvestris) widely distributed in three cool–dry Eurasian regions from the forest-steppe (Russia, Mongolia) and Mediterranean (Spain) biomes. We measured for each annual tree ring and the common 1950–2002 period the following variables: earlywood and latewood width, and minimum and maximum wood density. As expected, dry early growing season (spring) conditions were associated with low earlywood production but, most importantly, to high minimum density in the three conifer species. The associations between minimum density and spring precipitation were stronger (r = -0.65) than those observed with earlywood width (r = 0.57). We interpret the relationship between spring water availability and high minimum density as a drought-induced reduction in lumen diameter, hydraulic conductivity, and growth. Consequently, forecasted growing-season drier conditions would translate into increased minimum wood density and reflect a reduction in hydraulic conductivity, radial growth, and wood formation. Given the case-study-like nature of this work, more research on other cold–dry sites with additional conifer species is needed to test if minimum wood density is a robust proxy of early season water availability. We acknowledge the support of Spanish Ministry of Economy Projects (Fundiver, CGL2015-69186-C2-1-R). Tree-ring density data were obtained and analysed under support of Russian Science Foundation (Project 14-14-00295).

Arctic driftwood represents a unique proxy archive at the interface of marine and terrestrial environments. Combined wood anatomical and dendrochronological analyses have been used to detect the origin of driftwood and may allow past timber floating activities, as well as past sea ice and ocean current dynamics to be reconstructed. However, the success of driftwood provenancing studies depends on the length, number, and quality of circumpolar boreal reference chronologies. Here, we introduce a Eurasian-wide high-latitude network of 286 ring width chronologies from the International Tree Ring Data Bank (ITRDB) and 160 additional sites comprising the three main boreal conifers Pinus, Larix, and Picea. We assess the correlation structure within the network to identify growth patterns in the catchment areas of large Eurasian rivers, the main driftwood deliverers. The occurrence of common growth patterns between and differing patterns within catchments indicates the importance of biogeographic zones for ring width formation and emphasizes the degree of spatial precision when provenancing. Reference chronologies covering millennial timescales are so far restricted to a few larch sites in Central and Eastern Siberia (eastern Taimyr, Yamal Peninsula and north-eastern Yakutia), as well as several pine sites in Scandinavia, where large rivers are missing though. The general good spatial coverage of tree-ring sites across northern Eurasia indicates the need for updating and extending existing chronologies rather than developing new sites.

It is generally assumed in dendroecological studies that annual tree-ring growth is adequately determined by a linear function of local or regional precipitation and temperature with a set of coefficients that are temporally invariant. However, various researchers have maintained that tree-ring records are the result of multivariate, often nonlinear biological and physical processes. To describe critical processes linking climate variables with tree-ring formation, the process-based tree-ring Vaganov–Shashkin model (VS-model) was successfully used. However, the VS-model is a complex tool requiring a considerable number of model parameters that should be re-estimated for each forest stand. Here we present a new visual approach of process-based tree-ring model parameterization (the so-called VS-oscilloscope) which allows the simulation of tree-ring growth and can be easily used by researchers and students. The VS-oscilloscope was tested on tree-ring data for two species (Larix gmeliniiand Picea obovata) growing in the permafrost zone of Central Siberia. The parameterization of the VS-model provided highly significant positive correlations (p < 0.0001) between simulated growth curves and original tree-ring chronologies for the period 1950–2009. The model outputs have shown differences in seasonal tree-ring growth between species that were well supported by the field observations. To better understand seasonal tree-ring growth and to verify the VS-model findings, a multi-year natural field study is needed, including seasonal observation of the thermo-hydrological regime of the soil, duration and rate of tracheid development, as well as measurements of their anatomical features.

This research is an attempt to study seasonal translocation patterns of photoassimilated carbon within trees of one of the high latitudes widespread deciduous conifer species Larix gmelinii (Rupr. Rupr). For this purpose, we applied whole-tree labeling by 13CO2, which is a powerful and effective tool for tracing newly developed assimilates translocation to tissues and organs of a tree. Experimental plot has been established in a mature 105-year-old larch stand located within the continuous permafrost area near Tura settlement (Central Siberia, 64°17′13″N, 100°11′55″E, 148 m a.s.l.). Measurements of seasonal photosynthetic activity and foliage parameters (i.e., leaf length, area, biomass, etc.), and sampling were arranged from early growing season (June 8, 2013; May 14, 2014) until yellowing and senescence of needles (September 17, 2013; September 14, 2014). Labeling by 13C of the tree branch (June 2013, for 3 branch replicates in 3 different trees) and the whole tree was conducted at early (June 2014), middle (July 2014), and late (August 2013) phase of growing season (for different trees in 3 replicates each time) by three pulses [(CO2)max = 3000–4000 ppmv, 13CO2 (30 % v/v)]. We found at least two different patterns of carbon translocation associated with larch CO2 assimilation depending on needle phenology. In early period of growing season (June), 13C appearing in newly developed needles is a result of remobilized storage material use for growth purposes. Then approximately at the end of June, growth processes is switching to storage processes lasting to the end of growing season.

Plants exhibit morphological and anatomical adaptations to cope the environmental constraints of their habitat. How can mechanisms for adapting to contrasting environmental conditions change the patterns of tree rings formation? In this study, we explored differences in climatic conditions of permafrost and non-permafrost zones and assessed their influence on radial growth and wood traits of Larix gmelinii Rupr (Rupr) and Larix sibirica L., respectively. We quantified the contribution of xylem cell anatomy to the tree-ring width variability. Comparison of the anatomical tree-ring parameters over the period 1963–2011 was tested based on non-parametric Mann-Whitney U test. The generalized linear modeling shows the common dependence between TRW and the cell structure characteristics in contrasting environments, which can be defined as non-specific to external conditions. Thus, the relationship between the tree-ring width and the cell production in early- and latewood are assessed as linear, whereas the dependence between the radial cell size in early- and latewood and the tree-ring width becomes significantly non-linear for both habitats. Moreover, contribution of earlywood (EW) and latewood (LW) cells to the variation of TRW (in average 56.8% and 24.4% respectively) was significantly higher than the effect of cell diameters (3.3% (EW) and 17.4% (LW)) for the environments. The results show that different larch species from sites with diverging climatic conditions converge towards similar xylem cell structures and relationships between xylem production and cell traits. The work makes a link between climate and tree-ring structure, and promotes a better understanding the anatomical adaptation of larch species to local environment conditions.

Tree-ring width of Larix gmelinii (Rupr.) Rupr., ratios of stable isotopes of C (delta(13)C) and O (delta(18)O) of whole wood and cellulose chronologies were obtained for the northern part of central Siberia (Tura, Russia) for the period 1864-2006. A strong decrease in the isotope ratios of O and C (after atmospheric delta(13)C corrections) and tree-ring width was observed for the period 1967-2005, while weather station data show a decrease in July precipitation, along with increasing July air temperature and vapor pressure deficit (VPD). Temperature at the end of May and the whole month of June mainly determines tree radial growth and marks the beginning of the vegetation period in this region. A positive correlation between tree-ring width and July precipitation was found for the calibration period 1929-2005. Positive significant correlations between C isotope chronologies and temperatures of June and July were found for whole wood and cellulose and negative relationships with July precipitation. These relationships are strengthened when the likely physiological response of trees to increased CO(2) is taken into account (by applying a recently developed delta(13)C correction). For the O isotope ratios, positive relationships with annual temperature, VPD of July and a negative correlation with annual precipitation were observed. The delta(18)O in tree rings may reflect annual rather than summer temperatures, due to the late melting of the winter snow and its contribution to the tree water supply in summer. We observed a clear change in the isotope and climate trends after the 1960s, resulting in a drastic change in the relationship between C and O isotope ratios from a negative to a positive correlation. According to isotope fractionation models, this indicates reduced stomatal conductance at a relatively constant photosynthetic rate, as a response of trees to water deficit for the last half century in this permafrost region.

Permafrost exerts strong controls on forest development through nutrient availability. The key question of this study was to assess the effect of site conditions on macroelement concentration and stable isotope (&delta, 13C and &delta, 15N) dynamics during the growing season, and nutrient stoichiometry and resorption efficiency in the foliage of two common larch species in Siberia. Foliar nutrient (N, P and K) concentrations of larches grown on permafrost soils were exceptionally high in juvenile needles compared to those from a permafrost-free region (+50% and 130% for P and K), but were two-fold lower at needle maturation. Within permafrost terrain trees, sites with a warmer and deeper soil active layer had 15&ndash, 60% greater nutrient concentrations and higher &delta, 15N in their needles compared to shallower, colder soils. Larch of permafrost-free sites demonstrated an enrichment of foliage in 15N (+1.4% to +2.4&permil, ) in comparison to permafrost terrain (&minus, 2.0% to &minus, 6.9&permil, ). At all sites, foliar &delta, 13C decreased from June to August, which very likely results from an increasing contribution of current photoassimilates to build foliar biomass. With senescence, nutrient concentrations in larch needles decreased significantly by 60&ndash, 90%. This strong ability of larch to retain nutrients through resorption is the essential mechanism that maintains tree growth early in the growing season when soil remains frozen. The high resorptive efficiency found for K and P for larches established on permafrost suggests nutrient limitation of tree growth within the Central Siberian Plateau not only by N, as previously reported, but also by P and K. The increasing nutrient concentrations and a 15N enrichment of foliage towards warmer sites was paralleled by an up to 50-fold increase in biomass production, strongly suggesting that accelerated nutrient cycling with permafrost degradation contributes to an increased productivity of Siberian larch forests.

Stable oxygen isotope ratios (δ(18) O) in trees from high latitude ecosystems are valuable sources of information for recent and past environmental changes, but the interpretation is hampered by the complex hydrology of forests growing under permafrost conditions, where only a shallow layer of soil thaws in summer. We investigated larch trees (Larix gmelinii) at two sites with contrasting soil conditions in Siberia and determined δ(18) O of water from different soil depths, roots, twigs, and needles as well as δ(18) O of soluble carbohydrates regularly over two growing seasons. A comparison of results from the 2 yrs revealed an unexpected 'inverse' climate-isotope relationship, as dry and warm summer conditions resulted in lower soil and root δ(18) O values. This was due to a stronger uptake of isotopically depleted water pools originating from melted permafrost or previous winter snow. We developed a conceptual framework that considers the dependence of soil water profiles on climatic conditions for explaining δ(18) O in needle water, needle soluble carbohydrates and stem cellulose. The negative feedback of drought conditions on the source isotope value could explain decreasing tree-ring δ(18) O trends in a warming climate and is likely relevant in many ecosystems, where a soil isotope gradient with depth is observed.

Key message This study provides new data and an alternative framework to the debate of tree carbon economy in a context of increasing stress. Abstract For long-living trees, the resilience in times of stress is directly linked to the amount of accessible re- serves. Despite the simplicity of this principle, the under- standing of how carbon reserves limit growth and/or induce mortality under global change is still debated. In this study, we quantify how anatomical properties of rays—one of the main container for carbon reserves in tree stems—vary among sites, individuals, and annual rings of Larix gmelinii growing in contrasting sites in Siberia to verify if (1) the ray proportion and anatomical structure is linked to the environment, and/or (2) to changes in other wood tissues. Our observations have highlighted that ray proportion mainly varies among individuals, but little among sites and consecutive annual rings. We also observed that ray size and density scale to the wood structure with a relatively constant ratio of *2.5 rays per tracheid, independent of site conditions. These results suggest that the functional connection between the anatomy of rays and tracheid is unaffected by environment and highlight the importance of considering allometric relations in ecological comparisons. Comparative studies of long-term trajectory of ray pro- portion of living and dead trees might unravel observed variability among individuals validating the link between long-term depleted reserves and mortality.

Global warming is most pronounced in high-latitude regions by altering habitat conditions and affecting permafrost degradation, which may significantly influence tree productivity and vegetation changes. In this study, by applying a “space-for-time” approach, we selected three plots of Larix gmelinii forest from a continuous permafrost zone in Siberia with different thermo-hydrological soil regimes and ground cover vegetation with the objective of assessing how tree growth and productivity will change under different stages of permafrost degradation. A tree-ring multi-proxy characterization of mature trees was used to identify shift in ecophysiological responses related to the modified plant-soil system. Variability of tree-ring width (1975–2009), stable isotope ratios (oxygen and carbon, 2000–2009) and xylem structural characteristics (2000–2009) under climatic conditions of particular years indicated that an increased depth of the soil active layer will initially lead to increase of tree productivity. However, due to an expected water use increase through transpiration, the system might progressively shift from a temperature to a moisture-limited environment.

The radial growth of Siberian larch under the impact of pollutants emitted by enterprises of Norilsk has been analyzed to reconstruct the die-off dynamics of larch stands located along the Rybnaya River, along the main direction of pollutant air transport. Dendrochronological cross dating is used to detect the year of die off of 268 trees growing in 4 sites located 22, 45, 68, and 85 km from Norilsk. The death of individual trees in the sites closest to Norilsk was recorded immediately after the first enterprises started to operate in the early 1940s. The mass mortality of the trees started in the 1960s due to the operation of new smelters and the consequent increase in pollutant emissions. The complete destruction of the stands (100% die off of larch trees) occurred in the 1970s. At the most distant site (85 km), the highest rate of larch death was observed between 1975 and 1980, and in 2004 only 23% of larch trees were alive. A comparative analysis of the tree-ring width of the studied trees testified to the decrease in tree radial growth at the period before the complete degradation of stands. Unfavorable climatic conditions became an additional factor that enhanced the rate of tree die off due to the impact of pollutants. Although the increase in tree radial growth had been found in late 1990s to early 2000s at the site 85 km from Norilsk, the status of studied trees indicated that the area of completely degraded forest ecosystems might become larger under the present conditions.

Processes of xylem formation in Dahurian larch have been studied at three sites differing in the hydrothermal regime of soils in the permafrost zone of Middle Siberia. It has been found that the start and end dates of different phases of tree ring formation may differ between the sites by up to 14 days, depending on site conditions. The data obtained contribute to knowledge of possible changes in larch forest phytomass production and provide the possibility of predicting its dynamics under conditions of climate change.

The development of research into the history of tree growth and inferred summer temperature changes in Yamalia spanning the last 2000 years is reviewed. One focus is the evolving production of tree-ring width (TRW) and tree-ring maximum-latewood density (MXD) larch (Larix sibirica) chronologies, incorporating different applications of Regional Curve Standardisation (RCS). Another focus is the comparison of independent data representing past tree growth in adjacent Yamalia areas: Yamal and Polar Urals, and the examination of the evidence for common growth behaviour at different timescales. The sample data we use are far more numerous and cover a longer time-span at Yamal compared to the Polar Urals, but Yamal has only TRW, while there are both TRW and MXD for the Polar Urals. We use more data (sub-fossil and from living trees) than in previous dendroclimatic studies in this region. We develop a new TRW chronology for Yamal, more than 2000 years long and running up to 2005. For the Polar Urals we develop new TRW and MXD chronologies that show good agreement at short (

 Premise of the study: Xylem structure determines the hydraulic and mechanical properties of a stem, and its plasticity is fundamental for maintaining tree performance under changing conditions. Unveiling the mechanism and the range of xylem adjustment is thus necessary to anticipate climate change impacts on vegetation.  Methods: To understand the mechanistic process and the functional impact of xylem responses to warming in a cold-limited environment, we investigated the relationship between temperature and tracheid anatomy along a 312-yr tree-ring chronology of Larix sibirica trees from the Altay Mountains in Russia.  Key results: Climate-growth analyses indicated that warming favors wider earlywood cell lumen, thicker latewood walls, denser maximum latewood, and wider rings. The temperature signal of the latewood was stronger ( r > 0.7) and covered a longer and more stable period (from June to August) than that of earlywood and tree-ring width. Long-term analyses indicated a diverging trend between lumen and cell wall of early- and latewood.  Conclusions: Xylem anatomy appears to respond to warming temperatures. A warmer early-growing season raises water conduction capacity by increasing the number and size of earlywood tracheids. The higher-performing earlywood tracheids promote more carbon fi xation of the latewood cells by incrementing the rate of assimilation when summer conditions are favorable for growth. The diverging long-term variation of lumen and cell wall in earlywood vs. latewood suggests that xylem adjustments in latewood increase mechanical integrity and support increasing tree size under the ameliorated growing conditions.

Understanding climate change impacts on drought-prone forests is a critical issue. We investigated ring-width and stable isotopes (Δ13C and δ18O) in two Pinus sylvestris stands of the cold-dry Siberian forest-steppe growing under contrasting climatic trends over the last 75 years. Despite regional warming, there was increasing precipitation during the growing period at the southern site (MIN) but increasing water deficit (WD) at the northern site (BER). Intrinsic water use efficiency (WUEi) increased similarly (ca. 22%) in response to warming and rising atmospheric CO2. However, the steady increase in WUEi was accompanied by divergent growth patterns since 1980: increasing basal area increment (BAI) in MIN (slope = 0.102 cm2 year−2) and decreasing BAI in BER (slope = −0.129 cm2 year−2). This suggests that increased precipitation, mediated by CO2 effects, promoted growth in MIN, whereas intensified drought stress led to decreased carbon gain and productivity in BER. When compared to warm-dry stands of eastern Spain, the WUEi dependence on WD was three-fold greater in Siberia. Conversely, BAI was more affected by the relative impact of water stress within each region. These results indicate contrasting future trajectories of P. sylvestris forests, which challenge forecasting growth and carbon sequestration in cold-dry areas. This study was funded by the Spanish Government (grant number AGL2015-68274-C3-3-R) and the Russian Science Foundation (project numbers 14-14-00295, sampling and tree-ring data obtainment and 14-14-00219-P, mathematical approach). We acknowledge P. Sopeña and M.J. Pau for technical assistance.

[1] Arctic environments, where surface temperatures increase and sea ice cover and permafrost depth decrease, are very sensitive to even slight climatic variations. Placing recent environmental change of the high-northern latitudes in a long-term context is, however, complicated by too short meteorological observations and too few proxy records. Driftwood may represent a unique cross-disciplinary archive at the interface of marine and terrestrial processes. Here, we introduce 1445 driftwood remains from coastal East Greenland and Svalbard. Macroscopy and microscopy were applied for wood anatomical classification; a multi-species subset was used for detecting fungi; and information on boreal vegetation patterns, circumpolar river systems, and ocean current dynamics was reviewed and evaluated. Four conifer (Pinus, Larix, Picea, and Abies) and three deciduous (Populus, Salix, and Betula) genera were differentiated. Species-specific identification also separated Pinus sylvestris and Pinus sibirica, which account for ~40% of all driftwood and predominantly originate from western and central Siberia. Larch and spruce from Siberia or North America represents ~26% and ~18% of all materials, respectively. Fungal colonization caused different levels of driftwood staining and/or decay. Our results demonstrate the importance of combining wood anatomical knowledge with insight on boreal forest composition for successfully tracing the origin of Arctic driftwood. To ultimately reconstruct spatiotemporal variations in ocean currents, and to better quantify postglacial uplift rates, we recommend consideration of dendrochronologically dated material from many more circumpolar sites.

While the forest-tundra zone in Siberia, Russia has been dendroclimatologically well-studied in recent decades, much less emphasis has been given to a wide belt of northern taiga larch forests located to the south. In this study, climate and local site conditions are explored to trace their influence on radial growth of Gmelin larch ( Larix gmelinii (Rupr.) Rupr.) trees developed on permafrost soils in the northern taiga. Three dendrochronological sites characterized by great differences in thermo-hydrological regime of soils were established along a short (ca. 100 m long) transect: on a river bank (RB), at riparian zone of a stream (RZ) and on a terrace (TER). Comparative analysis of the rate and year-to-year dynamics of tree radial growth among sites revealed considerable difference in both raw and standardized tree-ring width (TRW) chronologies obtained for the RZ site, characterized by shallow soil active layer depth and saturated soils. Results of dendroclimatic analysis indicated that tree-ring growth at all the sites is mostly defined by climatic conditions of a previous year and precipitation has stronger effect on TRW chronologies in comparison to the air temperatures. Remarkably, a great difference in the climatic response of TRW chronologies has been obtained for trees growing within a very short distance from each other. The positive relation of tree-ring growth with precipitation, and negative to temperature was observed in the dry site RB. In contrary, precipitation negatively and temperature positively influenced tree radial growth of larch at the water saturated RZ. Thus, a complicate response of northern Siberian larch forest productivity to the possible climate changes is expected due to great mosaic of site conditions and variability of environmental factors controlling tree-ring growth at different sites. Our study demonstrates the new possibilities for the future dendroclimatic research in the region, as various climatic parameters can be reconstructed from tree-ring chronologies obtained for different sites.

We measured the seasonal dynamics of major and trace elements concentrations in foliage of larch, main conifer species of Siberia, and we analyzed cryogenic soils collected in typical permafrost-dominated habitats in the Central Siberia. This region offers a unique opportunity to study element fractionation between the soil and the plant because of (i) the homogeneous geological substratum, (ii) the monospecific stands (Larix gmelinii) and (iii) the contrasted habitats (North-facing slope, South-facing slope, and Sphagnum peatbog) in terms of soil temperature, moisture, thickness of the active layer, tree biomass and rooting depth. The variation of these parameters from one habitat to the other allowed us to test the effects of these parameters on the element concentration in larch foliage considered with high seasonal resolution. Statistical treatment of data on larch needles collected 4 times in 3 locations during entire growing season (June–September) demonstrated that : (1) there is a high similarity of foliar chemical composition of larch trees in various habitats suggesting intrinsically similar requirements of larch tree growth for nutrients, (2) the variation of elemental concentrations in larch needles is controlled by the period (within the growing season) and not by the geographical location (South-facing slope, North-facing slope or bog zone) and (3) there are three groups of elements according to their patterns of elements concentration in needles over the growing season from June to September can be identified: (1): nutrient elements [P, Cu, Rb, K, B, Na, Zn, Ni and Cd] showing a decrease of concentration from June to September similar to the behaviour of major nutrients such as N, P and K; (2): accumulating elements [Ca, Mg, Mo, Co, Sr, Mn, Pb and Cr] showing an increase of concentration from June–July to September; (3): indifferent elements [Al, Zr, Fe, Ba, Ti, REEs (Pr, Nd, Ce, La, Gd, Er, Dy, Tb, Lu, Yb, Tm, Sm, Ho, Eu), Y, Th and U] showing a decrease of concentration from June to July and then an increase of concentration to September. A number of micronutrients (e.g., Cu, Zn) demonstrate significant resorption at the end of growing season suggesting possible limitation by these elements. Although the intrinsic requirement seems to be similar among habitats, the total amount of element stored within the different habitats is drastically different due to the differences in standing tree biomass. The partitioning coefficients between soil and larch appear to be among the lowest compared to other environments with variable plants, soils and climates. Applying the “space for time” substitution scenario, it follows that under ongoing climate warming there will be an increase of the element stock following enhanced above-ground biomass accumulation, even considering zero modification of element ratios and their relative mobility. In this sense, the habitats like south-facing slopes can serve as resultant of climate warming effect on element cycling in larch ecosystems for the larger territory of Central Siberia.

Kirdyanov, A. V., Hagedorn, F., Knorre, A. A., Fedotova, E. V., Vaganov, E. A., Naurzbaev, M. M., Moiseev, P. A. & Rigling, A. 2012 (January): 20th century tree-line advance and vegetation changes along an altitudinal transect in the Putorana Mountains, northern Siberia. Boreas, Vol. 41, pp. 56–67. 10.1111/j.1502-3885.2011.00214.x. ISSN 0300-9483. Ongoing climatic changes potentially affect tree-line ecosystems, but in many regions the observed changes are superimposed by human activities. We assessed how the forest-tundra ecotone has changed during the last century in the Putorana Mountains, northern Siberia, an extremely remote and untouched area in Eurasia. A space-for-time approach was used to determine the spatio-temporal dynamics of forest structure and biomass along an altitudinal transect. From the closed larch forest to the upper tree line, the mean age of Larix gmelinii (Rupr.) decreased considerably from 220 to 50 years ago. At the current upper species line, there is a strong and successful germination of larch, with 1500 saplings per hectare, indicating an ongoing filling-in, a densification of a formerly open forest and an upslope shift of the tree-line position (approximately 30 to 50 m in altitude during the last century). The forest expansion coincided with large increases in winter precipitation during the 20th century. In contrast, tree growth rates were significantly positively related to summer temperatures, neither of which increased markedly. The total aboveground biomass decreased from approximately 40 t ha−1 in the closed larch forest to 5 t ha−1 at the tree line. Our study demonstrates that ongoing climatic changes lead to an upslope expansion of forests in the remote Putorana Mountains, which alters the stand structure and productivity of the forest-tundra ecotone. These vegetation changes are very probably of minor importance for aboveground carbon sequestration, but soil carbon data are needed to estimate the impact of the forest expansion on the total ecosystem carbon storage.

The current paper presents a new 1929-year tree-ring chronology for the Altai-Sayan region (Western Tuva) based on analyses of wood material from living trees and trunk remains of Siberian larch (Larix sibirica Ldb) from the upper treeline (2400 m asl). The correlations between tree-ring indices and meteorological data suggest that maximum effect on radial growth is produced by temperatures in June and July. This makes it possible to use the resulting chronology for early summer temperature reconstructions in the Altai-Sayan region. In respect to archaeology, a chronology of this length provides the opportunity of dating ancient wood samples, i.e. for exact definition of the calendar dates of the archaeological objects in the relevant territory over the entire period of the chronology.

Tree rings are natural archives of climate and environmental information with a yearly resolution. Indeed, wood anatomical, chemical, and other properties of tree rings are a synthesis of several intrinsic and external factors, and their interaction during tree growth. In particular, Intra-Annual Density Fluctuations (IADFs) can be considered as tree-ring anomalies that can be used to better understand tree growth and to reconstruct past climate conditions with intra-annual resolution. However, the ecophysiological processes behind IADF formation, as well as their functional impact, remain unclear. Are IADFs resulting from a prompt adjustment to fluctuations in environmental conditions to avoid stressful conditions and/or to take advantage from favorable conditions? In this paper we discuss: (1) the influence of climatic factors on the formation of IADFs; (2) the occurrence of IADFs in different species and environments; (3) the potential of new approaches to study IADFs and identify their triggering factors. Our final aim is to underscore the advantages offered by network analyses of data and the importance of high-resolution measurements to gain insight into IADFs formation processes and their relations with climatic conditions, including extreme weather events. This research is linked to activities conducted within the COST FP1106 “STReESS” network. Collection of datasets used for this work was supported by the projects: – ELENA (CGL2012-31668) and CGL2015-69985-R funded by the Spanish Science and Innovation Ministry (MICINN) and FEDER funds. – PGIDIT06PXIB502262PR funded by Dirección Xeral de Investigación, Desenvolvemento e Innovación, Xunta de Galicia. – Slovenian Research Agency (program P4-0015). – Austrian Science Fund (FWF-TRP 122-B16). – Russian Science Foundation 14-14-00295. – POCI/CLI/58680/2004 and PTDC/AAC-AMB/111675/2009 funded by the Portuguese Foundation for Science and Technology (FCT) and the European Union (POCI 2010) SI

Significant gaps still exist in our knowledge about post-photosynthetic leaf level and downstream metabolic processes and isotopic fractionations. This includes their impact on the isotopic climate signal stored in the carbon isotope composition (δ(13)C) of leaf assimilates and tree rings. For the first time, we compared the seasonal δ(13)C variability of leaf sucrose with intra-annual, high-resolution δ(13)C signature of tree rings from larch (Larix gmelinii Rupr.). The trees were growing at two sites in the continuous permafrost zone of Siberia with different growth conditions. Our results indicate very similar low-frequency intra-seasonal trends of the sucrose and tree ring δ(13)C records with little or no indication for the use of 'old' photosynthates formed during the previous year(s). The comparison of leaf sucrose δ(13)C values with that in other leaf sugars and in tree rings elucidates the cause for the reported (13)C-enrichment of sink organs compared with leaves. We observed that while the average δ(13)C of all needle sugars was 1.2‰ more negative than δ(13)C value of wood, the δ(13)C value of the transport sugar sucrose was on an average 1.0‰ more positive than that of wood. Our study shows a high potential of the combined use of compound-specific isotope analysis of sugars (leaf and phloem) with intra-annual tree ring δ(13)C measurements for deepening our understanding about the mechanisms controlling the isotope variability in tree rings under different environmental conditions.

We propose a technique for separating the climatic signal which is contained in two tree-ring parameters widely used in dendroclimatology. The method is based on the removal of the relationship between tree-ring width (TRW) and maximum latewood density (MXD) observed for narrow tree rings from high latitudes. The new technique is tested on data from three larch stands located along the northern timberline in Eurasia. Correlations were calculated between the temperatures of pentads (five consecutive days), TRW chronologies and MXD chronologies calculated according to the standard and proposed methods. The analysis confirms the great importance of summer temperature for tree radial growth and tree-ring formation. TRW is positively correlated with the temperature of four to eight pentads (depending on the region) at the beginning of the growth season, but MXD as obtained by the standard technique is correlated with temperature over a much longer period. For maximum density series from which the relationship between MXD and TRW has been removed (MXD′), there is a clear correlation with temperatures in the second part of the growing season. These results are consistent with the known dynamics of tree-ring growth in high latitudes and mechanisms of tree-ring formation.

Productivity of the moss cover and necromass accumulation in the litter of a sphagnum larch forest have been estimated on the basis of tree age. It has been shown that the total carbon stock in the litter of a 100-year-old stand, including organic matter not destroyed by fire, exceeds the corresponding value for the tree stand itself by more than an order of magnitude. The accumulation of organic matter on the soil surface inhibits the growth of larch. In particular, this factor impairs hydrothermal conditions in the soil and causes a rise of the permafrost table; as a consequence, lower layers of the root system die off.

To investigate the variability of primary production of boreal forest ecosystems under the current climatic changes, we compared the dynamics of annual increments and productivity of the main components of plant community (trees, shrubs, mosses) at three sites in the north of Siberia (Russia). Annual radial growth of trees and shrubs was mostly defined by summer temperature regime (positive correlation), but climatic response of woody plants was species specific and depends on local conditions. Dynamics of annual increments of mosses were opposite to tree growth. The difference in climatic response of the different vegetation components of the forest ecosystems indicates that these components seem to be adapted to use climatic conditions during the short and severe northern summer, and decreasing in annual production of one component is usually combined with the increase of other component productivity. Average productivity in the northern forest ecosystems varies from 0.05 to 0.14 t ha(-1) year(-1) for trees, from 0.05 to 0.18 t ha(-1) year(-1) for shrubs and from 0.54 to 0.66 t ha(-1) year(-1) for mosses. Higher values of tree productivity combined with lower annual moss productivity were found in sites in northern taiga in comparison with forest-tundra. Different tendencies in the productivity of the dominant species from each vegetation level (trees, shrubs, mosses) were indicated for the last 10 years studied (1990-1999): while productivity of mosses is increasing, productivity of trees is decreasing, but there is no obvious trend in the productivity of shrubs. Our results show that in the long term, the main contribution to changes in annual biomass productivity in forest-tundra and northern taiga ecosystems under the predicted climatic changes will be determined by living ground cover.

Summary Mass accumulation dynamics in earlywood and latewood cell walls of larch from northern regions of Central Siberia are investigated and correlations among cell mass of different tree-ring zones, radial tracheid sizes and tree-ring width are found. A linear relationship exists between cell wall mass and radial cell size. The deviation of cell mass from the regression line (index of mass deviation) and the mean density of the respective tree-ring zone are similarly functionally dependent on the radial cell size and reflect the realization of the potential determined during the period of cell formation. There is a critical value of larch tree-ring width in relation to cell mass. For the tree-rings with width less than this critical value the difference in the mass of earlywood and latewood cells increases with decreasing tree-ring width.The cell wall mass correlates with the monthly temperatures of June and July. Under favorable growing conditions tracheids with similar mass of cell walls are formed in earlywood and latewood, while under unfavorable conditions cell wall mass accumulation in latewood is severly limited. A comparative analysis of the cell wall mass in earlywood and latewood indicates that mass accumulation is independent of the switching processes from earlywood production to the production of latewood.

Wood material for at least 12 larch trees at six sites [Larix sibirica Ldb, Larix gmelinii (Rupr.) Rupr, Larix cajanderi Mayr] near the northern timberline in Siberia was analyzed to investigate influence of climatic factor changes on tree-ring growth at high latitudes. Tree-ring cell size, maximum latewood density and ring width measured by means of image analysis and X-ray radiodensitometry and calculated latewood cell-wall thickness were used. Correlation analysis of tree-ring structure parameter chronologies with temperatures averaged over periods of 5 days (pentad) shows that early summer temperature (mean for 5–6 pentads, depending on the region, starting from the middle of June) and date of snow melt are the most important factors that define seasonal growth and tree-ring structure. Analysis of instrumental climatic data indicates that a positive trend of early summer temperature was combined with winter precipitation (October–April) increase and this combination leads to later snow melt. Based of the results of tree-ring growth modelling, it was shown that later snow melt (hence, delayed initiation of cambial activity and, as a result, decrease of wood production) explains the changes in the relationship between tree ring width and summer temperature dynamics observed after the 1960s for a large area of the Siberian Subarctic. The understanding of the role of winter precipitation in controlling ring growth, through its effect on the timing of cambial activation, suggests the possibility of using ring structure parameters to create reconstructions of past winter precipitation variations.

Wood from the boreal forest represents an important resource for paper production and sawmill processing. Due to poor infrastructure and high transportation costs on land, timbers are often transported over long distances along large river systems. Industrial river rafting activities started at the end of the 19th century and were intensified in western Russia and central Siberia from the 1920s to the 1980s. After initial single stem rafting, timber is today mostly floated in ship-guided rafts. Lost wood can be transported further to the Arctic Ocean, where it may drift within sea ice over several years and thousands of kilometers before being deposited along (sub-)Arctic coastlines. Here, we introduce dendro-dated tree-ring width series of 383 driftwood samples from logged timber that were collected along different driftwood-recipient coastlines in Greenland, Iceland and Svalbard. The majority of driftwood is Pinus sylvestris from the southern Yenisei region in central Siberia, whereas Larix sp. and Picea sp. from western Russia and eastern Siberia are rare. Although our results are based on a small sample collection, they clearly show the importance of timber rafting on species, age and origin of Arctic driftwood and indicate the immense loss of material during wood industrial river floating.

[1] Tree ring width, density, and ratio of stable isotopes (13C/12C and 18O/16O) in wood and cellulose were determined for larch (Larix sibirica Ledeb.) growing under water deficit conditions in the forest steppe zone in central Siberia (54°24′N, 89°57′E) for the period 1850–2005. Dendroclimatic analysis of the chronologies indicated precipitation to be the most important factor determining indicated parameters. Precipitation of June is significantly correlated with tree ring width and maximum density (r = 0.36 and 0.43, p < 0.05, respectively). Relations of δ13C and δ18O to precipitation are similar, but the most important month is July (r = −0.47, p < 0.05 for δ13C; r = −0.29, p < 0.05 for δ18O). Further, δ18O is positively related to the mean temperatures of July (r = 0.30, p < 0.05). Carbon and oxygen isotope ratios in wood and cellulose showed strong negative trends for the last 100 years. The reason for this decrease could be the observed shift of the beginning of the vegetation period to earlier dates (up to 1 week) in spring and the increased use of precipitation water stored in the soil from October of the previous year, although other explanations cannot be excluded (in particular the physiological effect of increasing atmospheric CO2 also responsible for lower δ13C values). Thus, an earlier start of the vegetation period could lead to tree ring formation during a period with higher water availability, resulting in stronger isotopic fractionation and 13C depletion, also reflected in a higher earlywood to latewood ratio. At the same time, highly 18O depleted water from October precipitation of the previous year is absorbed. The incorporation of this isotopically lighter water during photosynthesis is reflected in the wood and cellulose of tree rings.

Larch is the tree species most widely spread in the permafrost region of Siberia (Anonymous 1977; Dilis 1981; Koropachinskii 1983; Abaimov et al. 1997). Morphological and physiological features of the larch allowed it to adapt to severe conditions of climate, permafrost, and poor soils. The northern tree line of this species corresponds to the latitudinal timberline (Dilis 1981; Abaimov 1995). Tree-ring growth of larch at permafrost regions is characterized by features typical for the trees growing under strong influence of temperature (Vaganov et al. 1996). Dendrochronological method allows us to investigate various aspects of tree-ring formation at different locations under different conditions (Schweingruber 1996; see Chap. 17).

We present the first and longest (413 years) dataset on stable carbon isotope ratios in tree-ring cellulose (δ13C), tree-ring width (TRW), and maximum latewood density (MXD) obtained from larch trees growing on permafrost under continental climate in the Suntar Khayata mountain ridge in Eastern Siberia (Russia). With this first study we calibrate tree-ring parameters against climate quantities, and based on these results assess the potential added value of MXD and especially of δ13C complementing TRW analysis for future climate reconstruction purposes. δ13C chronologies were corrected for human induced changes in atmospheric CO2 since AD 1800. Two different approaches were compared i) a correction referring merely to the decline in atmospheric δ13C (δ13Catm) and ii) a correction additionally accounting for the increase in atmospheric partial pressure of CO2. δ13C chronologies are characterized by strong signal strength with only 4 trees representing the population signal at the site (mean inter-series correlation = 0.71 and EPS = 0.90). δ13C variation shows low similarity to TRW and MXD, while correlation between TRW and MXD is highly significant. Correlation analysis of tree-ring parameters with gridded instrumental data (Climate Research Unit, CRU TS 2.1) over the AD 1929–2000 calibration period demonstrates that TRW and MXD react as reported from other sites at cold and humid northern latitudes: precipitation plays no significant role, but strong dependencies on monthly mean, maximum and minimum temperatures, particularly of the current summer (June to August), are found (up to r = 0.60, p < 0.001). Combining instrumental data to a summer season mean (JJA) and TRW and MXD to a growth parameter mean (TRW + MXD), clearly shows the importance of the number of frost days and minimum temperatures during summer (r = 0.67, p < 0.001) to dominate tree growth and highlights the potential for climate reconstruction. Carbon isotope fixation in tree rings is obviously less controlled by temperature variables. In particular, the frost days and minimum temperature have a much smaller influence on δ13C than on tree growth. δ13C strongly reacts to current-year July precipitation (r = − 0.44, p < 0.05) and June–July maximum temperature (r = 0.46, p < 0.001). All significant (p < 0.05) correlation coefficients are higher when using the corrected δ13C chronology considering an additional plant physiological response on increasing atmospheric CO2 concentration, than using the chronology corrected for δ13Catm changes alone. Spatial distribution of correlations between tree-ring data and climate variables for Eastern Siberia indicates that the summer temperature regime in the studied region is mostly influenced by Arctic air masses, but precipitation in July seems to be brought out from the Pacific region. Both the combined TRW + MXD record and the δ13С record revealed a high reconstruction potential for summer temperature and precipitation, respectively, particularly on decadal and longer-term scales.