Rising utilization of stable isotopes in tree rings for climate change and forest ecology

Analyses of stable isotopes(C,O,H)in tree rings are increasingly important cross-disciplinary programs.The rapid development in this field documented in an increasing number of publications requires a comprehensive review.This study includes a bibliometric analysis-based review to better understand research trends in tree ring stable isotope research.Overall,1475 publications were selected from the Web of Science Core Collection for 1974-2023.The findings are that:(1)numbers of annual publications and citations increased since 1974.From 1974 to 1980,there were around two relevant publications per year.However,from 2020 to 2022,this rose sharply to 109 publications per year.Likewise,average article citations were less than four per year before 1990,but were around four per article per year after 2000;(2)the major subjects using tree ring stable isotopes include forestry,geosciences,and environmental sciences,contributing to 42.5%of the total during 1974-2023;(3)the top three most productive institutions are the Chinese Academy of Sciences(423),the Swiss Federal Institute for Forest,Snow and Landscape Research(227),and the University of Arizona(204).These achievements result from strong collaborations;(4)review papers,for example,(Dawson et al.,Annu Rev Ecol Syst 33:507-559,2002)and(McCarroll and Loader,Quat Sci Rev 23:771-801,2004),are among the most cited,with more than 1000 citations;(5)tree ring stable isotope studies mainly focus on climatology and ecology,with atmospheric CO_(2) one of the most popular topics.Since 2010,precipitation and drought have received increasing attention.Based on this analysis,the research stages,key findings,debated issues,limitations and direc-tions for future research are summarized.This study serves as an important attempt to understand the progress on the use of stable isotopes in tree rings,providing scientific guid-ance for young researchers in this field.

Aging Mongolian pine plantations face high risks of drought-induced growth decline:evidence from both individual tree and forest stand measurements

Discerning vulnerability differences among different aged trees to drought-driven growth decline or to mortality is critical to implement age-specific countermeasures for forest management in water-limited areas.An important species for afforestation in dry environments of northern China,Mongolian pine(Pinus sylvestris var.mongolica Litv.)has recently exhibited growth decline and dieback on many sites,particularly pronounced in old-growth plantations.However,changes in response to drought stress by this species with age as well as the underlying mechanisms are poorly understood.In this study,tree-ring data and remotely sensed vegetation data were combined to investigate variations in growth at individual tree and stand scales for young(9-13 years)and aging(35-52 years)plantations of Mongolian pine in a water-limited area of northern China.A recent decline in tree-ring width in the older plantation also had lower values in satellited-derived normalized difference vegetation indices and normalized difference water indices relative to the younger plantations.In addition,all measured growth-related metrics were strongly correlated with the self-calibrating Palmer drought severity index during the growing season in the older plantation.Sensitivity of growth to drought of the older plantation might be attributed to more severe hydraulic limitations,as reflected by their lower sapwood-and leaf-specific hydraulic conductivities.Our study presents a comprehensive view on changes of growth with age by integrating multiple methods and provides an explanation from the perspective of plant hydraulics for growth decline with age.The results indicate that old-growth Mongolian pine plantations in water-limited environments may face increased growth declines under the context of climate warming and drying.

More tree growth reduction due to consecutive drought and its legacy effect for a semiarid larch plantation in Northwest China

Extreme climate has increasingly led to negative impacts on forest ecosystems globally,especially in semiarid areas where forest ecosystems are more vulnerable.However,it is poorly understood how tree growth is affected by different drought events.In 2006–2009,the larch plantations in the semiarid areas of Northwest China were negatively affected by four consecutive dry years,which was a very rare phenomenon that may occur frequently under future climate warming.In this study,we analyzed the effect of these consecutive dry years on tree growth based on the data of the tree rings in the dominant layer of the forest canopy on a larch plantation.We found that the tree-ring width index(RWI)in dry years was lower than that in normal years,and it experienced a rapidly decreasing trend from 2006 to 2009(slope=-0.139 year^(-1),r=-0.94)due to water supply deficits in those dry years.Drought induced legacy effects of tree growth reduction,and consecutive dry years corresponded with greater growth reductions and legacy effects.Growth reductions and legacy effects were significantly stronger in the third and fourth consecutive dry years than that of single dry year(p<0.05),which might have been due to the cumulative stress caused by consecutive dry years.Our results showed that larch trees experienced greater tree growth reduction due to consecutive dry years and their legacy effect,and the trees had lower recovery rates after consecutive dry years.Our results highlight that consecutive dry years pose a new threat to plantations under climate warming,and thus,the effect of climate extremes on tree growth should be considered in growth models in semiarid areas.

Winter-spring minimum temperature variations inferred from tree-ring δ^(13)C in southeastern China

Long-term temperature variations inferred from high-resolution proxies provide an important context to evaluate the intensity of current warming.However,tem-perature reconstructions in humid southeastern China are scarce and particularly lack long-term data,limiting us to obtain a complete picture of regional temperature evolution.In this study,we present a well-verified reconstruction of winter-spring(January–April)minimum temperatures over southeastern China based on stable carbon isotopic(δ^(13)C)records of tree rings from Taxus wallichiana var.mairei from 1860 to 2014.This reconstruction accounted for 56.4%of the total observed variance.Cold periods occurred during the 1860s–1910s and 1960s–1970s.Although temperatures have had an upward trend since the 1920s,most of the cold extremes were in recent decades.The El Niño-Southern Oscillation(ENSO)variance acted as a key modulator of regional winter-spring minimum temperature variability.However,teleconnections between them were a nonlinear process,i.e.,a reduced or enhanced ENSO variance may result in a weakened or intensified temperature-ENSO relationship.

Stability assessment of tree ring growth of Pinus armandii Franch in response to climate change based on slope directions at the Lubanling in the Funiu Mountains,China

Global warming will affect growth strategies and how trees will adapt.To compare the response of tree radial growth to climate warming in different slope directions,samples of Pinus armandii Franch were collected and tree-ring chronologies developed on northern and western slopes from the Lubanling in the Funiu Mountains.Correlation analyses showed that two chronologies were mainly limited by temperatures in the previous June-August and the com-bination of temperatures and moisture in the current May-July.The difference of the climate response to slopes was small but not negligible.Radial growth of the LBLO1 site on the northern slope was affected by the combined maximum and minimum temperatures,while that of the LBLO2 site was affected by maximum temperatures.With regards to moisture,radial growth of the trees on the north slope was influenced by the relative humidity in the current May-July,while on the western slope,it was affected by the relative humidity in the previous June-August,the current May-July and the precipitation in the current May-July.With the change in climate,the effects of the main limiting factors on growth on different slopes were visible to a certain extent,but the differences in response of trees on different slopes gradually decreased,which might be caused by factors such as different slope directions and the change in diurnal temperature range.These results may provide information for forest protection and ecological construction in this region,and a scientific reference for future climate reconstruction.

Stability assessment of tree ring growth of Pinus armandii Franch in response to climate change based on slope directions at the Lubanling in the Funiu Mountains,China

Global warming will affect growth strategies and how trees will adapt.To compare the response of tree radial growth to climate warming in different slope directions,samples of Pinus armandii Franch were collected and treering chronologies developed on northern and western slopes from the Lubanling in the Funiu Mountains.Correlation analyses showed that two chronologies were mainly limited by temperatures in the previous June–August and the combination of temperatures and moisture in the current May–July.The difference of the climate response to slopes was small but not negligible.Radial growth of the LBL01 site on the northern slope was affected by the combined maximum and minimum temperatures,while that of the LBL02 site was affected by maximum temperatures.With regards to moisture,radial growth of the trees on the north slope was influenced by the relative humidity in the current May–July,while on the western slope,it was affected by the relative humidity in the previous June–August,the current May–July and the precipitation in the current May–July.With the change in climate,the effects of the main limiting factors on growth on different slopes were visible to a certain extent,but the differences in response of trees on different slopes gradually decreased,which might be caused by factors such as different slope directions and the change in diurnal temperature range.These results may provide information for forest protection and ecological construction in this region,and a scientific reference for future climate reconstruction.

Climate warming is significantly influenced by rising summer maximum temperatures:insights from tree-ring evidence of the Western Tianshan Mountains,China

As one of the regions most affected by global climate warming,the Tianshan mountains has experienced several ecological crises,including retreating glaciers and water deficits.Climate warming in these mountains is considered mainly to be caused by increases in minimum temperatures and winter temperatures,while the influence of maximum temperatures is unclear.In this study,a 300-year tree-ring chronology developed from the Western Tianshan Mountains was used to reconstruct the summer(June-August)maximum temperature(Tmax6-8) variations from 1718 to2017.The reconstruction explained 53.1% of the variance in the observed Tmax6-8.Over the past 300 years,the Tmax6-8reconstruction showed clear interannual and decadal variabilities.There was a significant warming trend(0.18 ℃/decade) after the 1950s,which was close to the increasing rates of the minimum and mean temperatures.The increase in maximum temperature was also present over the whole Tianshan mountains and its impact on climate warming has increased.The Tmax6-8variations in the Western Tianshan mountains were influenced by frequent volcanic eruptions combined with the influence of solar activity and the summer North Atlantic Oscillation.This study reveals that climate warming is significantly influenced by the increase in maximum temperatures and clarifies possible driving mechanisms of temperature variations in the Western Tianshan mountains which should aid climate predictions.

Climate warming is significantly influenced by rising summer maximum temperatures: insights from tree-ring evidence of the Western Tianshan Mountains, China

As one of the regions most affected by global cli-mate warming,the Tianshan mountains has experienced sev-eral ecological crises,including retreating glaciers and water deficits.Climate warming in these mountains is considered mainly to be caused by increases in minimum temperatures and winter temperatures,while the influence of maximum temperatures is unclear.In this study,a 300-year tree-ring chronology developed from the Western Tianshan Moun-tains was used to reconstruct the summer(June-August)maximum temperature(T_(max6-8))variations from 1718 to 2017.The reconstruction explained 53.1% of the variance in the observed T_(max6-8).Over the past 300 years,the T_(max6-8)reconstruction showed clear interannual and decadal vari-abilities.There was a significant warming trend(0.18°C/decade)after the 1950s,which was close to the increasing rates of the minimum and mean temperatures.The increase in maximum temperature was also present over the whole Tianshan mountains and its impact on climate warming has increased.The T_(max6-8) variations in the Western Tianshan mountains were influenced by frequent volcanic eruptions combined with the influence of solar activity and the sum-mer North Atlantic Oscillation.This study reveals that cli-mate warming is significantly influenced by the increase in maximum temperatures and clarifies possible driving mech-anisms of temperature variations in the Western Tianshan mountains which should aid climate predictions.

Long-term changes in radial growth of seven tree species in the mixed broadleaf-Korean pine forest in Northeast China:Are deciduous trees favored by climate change?

The role of the temperate mixed broadleaf-Korean pine forest(BKF)in global biogeochemical cycles will depend on how the tree species community responds to climate;however,species-specific responses and vulner-abilities of common trees in BKF to extreme climates are poorly understood.Here we used dendrochronological meth-ods to assess radial growth of seven main tree species(Pinus koraiensis,Picea jezoensis,Abies nephrolepis,Fraxinus mandshurica,Phellodendron amurense,Quercus mongolica,and Ulmus davidiana)in an old-growth BKF in response to climate changes in the Xiaoxing’an Mountains and to improve predictions of changes in the tree species compo-sition.Temperature in most months and winter precipita-tion significantly negatively affected growth of P.jezoensis and A.nephrolepis,but positively impacted growth of P.koraiensis and the broadleaf species,especially F.mandshu-rica and U.davidiana.Precipitation and relative humidity in June significantly positively impacted the growth of most tree species.The positive effect of the temperature during the previous non-growing season(PNG)on growth of F.mandshurica and Q.mongolica strengthened significantly with rapid warming around 1981,while the impact of PNG temperature on the growth of P.jezoensis and A.nephrolepis changed from significantly negative to weakly negative or positive at this time.The negative response of radial growth of P.jezoensis and A.nephrolepis to precipitation during the growing season gradually weakened,and the negative response to PNG precipitation was enhanced.Among the studied species,P.koraiensis was the most resistant to drought,and U.davidiana recovered the best after extreme drought.Ulmus davidiana,P.jezoensis and A.nephrolepis were more resistant to extreme cold than the other species.Climate warming generally exacerbated the opposite growth patterns of conifer(decline)and broadleaf(increase)spe-cies.Deciduous broadleaf tree species in the old-growth BKF probably will gradually become dominant as warming continues.Species-specific growth-climate relationships should be considered in future models of biogeochemical cycles and in forestry management practices.

Correlation Between Chemical Element Contents in Tree Rings and Soils

The annual growth rings of ten trees and the soils near the tree roots were sampled from the mining area of lead-and zinc-dominant metals in the Xixia Mountain, Nanjing, for the determination of chemical element contents. The study results showed that the elemental contents in the tree rings were correlated with those in the soils, i. e., the elemental contents in the tree rings increased with those in the soils, even in the cases of different environments and different tree species. Therefore, a time-concentration sequence could be set up on the basis of determining the elemental contents in the successive annual growth rings of trees to qualitatively reflect the annual variations of relevant elements in the soils, and a time-concentration sequence of elemental contents in soils could also be established in terms of related model to reproduce the dynamic changes of the surroundings.

Chrono-Sequences of Elemental Contents in Tree Ringsand Soils

There exists a logarithmic linear correlation, i. e., In C'(Z, t) = a(Z) + b(Z) ln(Z, t) where Z is the atomicnumber of element and t the year when tree ring grows between the chemical element contents in tree ringsC(Z, t) and those in the soils near the tree roots C'(Z, t).By determining the elemental contents of the annual growth rings of trees, we could establish the chrono-sequences of elemental contents in the tree rings, thus calculating that of the soil, that is, reproducing thedynamic changes of contents of elements in the soil C'(Z, t). The background values of elements in the soilunder site conditions of the tree could be estimated from the minimum C(Z, t_0) in the chrono-sequences ofelemental contents in the tree rings.

Study on Model of Correlation Between Chemical Ele－ment Contents in Tree Rings and Soils near Tree Roots

The chemical element contents in tree rings are correlated with those in the soils near the tree roots. Theresults in the present study showed that the correlation between them could be described using the followinglogarithmic linear correlation model:lgC'(Z) = α(Z) + b(Z)lgC(Z).Therefore, by determining the chrono-sequence C(Z, t), where Z is the atomic number and t the year ofelemental contents in the annual growth rings of trees, we could get the chrono-sequence C'(Z, t) of elementalcontents in the soil, thus inferring the dynamic variations of relevant elemental contents in the soil.

Angular Distribution of Element Contents in Tree Rings and the Environmental Information

Element contents of tree rings and soils near tree roots collected from Deodar cedar (Cedrus deodara (Roxb.) G. Don) and Masson pine (Picks massoniana lamb.) were determined to study the relationship between the angular distribution of element contents in tree rings and the environmental information. The chemical composition and properties of soils are very much complicated, which leads to the non-uniform distribution of the element contents in tree rings. The statistical multi-variable regression method was used to got the information of the tree-centered distribution of element contents in the environment (soil) (C’), C’(z, θj ), from the distribution of element contents in tree rings (C), C(Z, θi), which depends on the plane azimuth angle (θi), i. e., C=C(Z,θi), where Z is the atomic number of the element, with a satisfactory result,though this study is only a primary one.

August–September Runoff Variation in the Kara Darya River Determined from Juniper(Juniperus turkestanica) Tree Rings in the Pamirs-Alai Mountains, Kyrgyzstan, Back to 1411 CE

A 606 year runoff reconstruction of the Kara Darya River was developed, based on the tree-ring width chronology of Turkestan juniper(Juniperus turkestanica) from the Pamir-Alai Mountains of Kyrgyzstan. Preliminary comparison between the snow cover variation and these climate/runoff reconstructions found that Central Asian snow cover may have strong associations with large-scale ocean-atmosphere-land circulations. The runoff reconstruction demonstrated that instrumental runoff was not representative of runoff over the past 606 years. The drought of the 1960 s-1990 s resulted in low runoff levels during the past 50 years;however, this probably does not represent a worst-case scenario for the Kara Darya because the runoff reconstruction showed additional extremely low runoff prior to the 20 th century. The reconstruction will provide a long-term perspective on runoff variation in the Kara Darya River basin, aid sustainable water resource management and be useful in guiding expectations of future variations and water resource planning.

Application and verification of simultaneous determination of cellulose δ^(13)C and δ^(18)O in Picea shrenkiana tree rings from northwestern China using the high-temperature pyrolysis method

Stable isotopes in tree-ring cellulose provide important data in ecological,archaeological,and paleoenvironmental researches,thereby,the demand for stable isotope analyses is increasing rapidly.Simultaneous measurement of cellulose δC and δO values from tree rings would reduce the cost of isotopic commodities and improve the analytical efficiency compared with conventional separate measurement.In this study,we compared the δC and δO values of tree-ringα-cellulose from Tianshan spruce(Picea schrenkiana)in an arid site in the drainage basin of the Urumqi River in Xinjiang of northwestern China based on separate and simultaneous measurements,using the combustion method(at1050°C)and the high-temperature pyrolysis method(at 1350°C and 1400°C).We verified the results of simultaneous measurement using the outputs from separate measurement and found that both methods(separate and simultaneous)produced similar δC values.The two-point calibrated method improved the results(range and variation)of δC and δO values.The mean values,standard deviations,and trends of the tree-ring δC obtained by the combustion method were similar to those by the pyrolysis method followed by two-point calibration.The simultaneously measured δO from the pyrolysis method at 1400°C had a nearly constant offset with that the pyrolysis method at 1350°C due to isotopic-dependence on the reaction temperature.However,they showed similar variations in the time series.The climate responses inferred from simultaneously and separately measured δC and δO did not differ between the two methods.The tree-ring δC and δO values were negatively correlated with standardized precipitation evapotranspiration index from May to August.In addition,the δO was significantly correlated with temperature(positive),precipitation(negative),and relative humidity(negative)from May to August.The tree-ring δC and δO values determined simultaneously through the high-temperature pyrolysis method could produce acceptable and reliable stable isotope series.The simultaneous isotopic measurement can greatly reduce the cost and time requirement compared with the separate isotopic measurement.These results are consistent with the previous studies at humid sites,suggesting that the simultaneous determination of δC and δO in tree-ringα-cellulose can be used in wide regions.

Application and Verification of Logarithmic LinearCorrelation Model of Element Contents Between Tree Ringsand Soils near the Tree Roots

Nine Platanus acerifolia (Ait.) Willd. trees growing in the Nanjing Children Teacher’s School, Nanjing,China, were selected to determine the contents of 13 chemical elements both in the 1994’s growth rings,C(Z, t), and in the soils near the roots, C’(Z, t), of the trees. The results showed that the relationship between C(Z, t) and C’(Z, t) followed the logarithmic linear correlation model, lgC’(Z, t)=a(Z)+b(Z) ig C (Z, t).Based on this model the chrono-sequences of chemical element contents in the soils were reproduced from those in the tree rings; i.e., the dynamic variations in the chemical element contents of the soils at the sites were traced. In this study the chrono-sequences of the chemical elements including Cd, Ph, Mn, Co and Zn in the soils near the roots of a Platanus acerifolia tree from 1957 to 1994 were established, and the background values of Cd, Pb, Mn, Co and Zn in the soil were calculated by taking the lowest values of the chrono-sequences of the element contents as upper limits of the background values.

Changes in Ecological Environment of the Tarim River Confirmed by Tree Rings

The tree ring is an important carrier of the evolution of climate in the study of global change.It enjoys advantages of accurate dating,strong continuity,high resolution ratio and high reconstruction precision.This paper tested the changes of Populus euphratica tree ring and the runoff in the upper and middle reaches of the Tarim River through the monotonic trend and analyzed the response of the tree ring in the lower reaches of the Tarim River to the water supply through the combined tree ring data and the historical literature,so as to confirm the accuracy of the response of Populus euphratica tree ring to river diversion.The results were included as follows.(1)From 1957 to 2010,the radial growth of Populus euphratica in the upper and middle reaches of the Tarim River decreased gradually in the time series,which was the same as the runoff.The downtrend was significant,and|Zc|was more than 0.96.(2)From1913 to 2000,the radial growth of Populus euphratica as a whole in the lower reaches of the Tarim River decreased.The downtrend was significant between 1973 and 2001,reaching the lowest level in the past 100years.The formation and diversion of the Layi River was as described in the historical literature.(3)The development of the ecological water delivery project has improved the ecological environment of the lower reaches,which has been verified from the Populus euphratica tree ring in the lower reaches.However,despite the ecological water delivery project in recent years,the growth of Populus euphratica tree ring has not reached to the level before 1920,indicating that the existing ecological water delivery is of limited value in recovering desert riparian forest.(4)In the analysis of the response of the Populus euphratica to the water resources,the wavelet analysis method and the historical literature review method have drawn the same conclusion.

Long-Term H2O and CO2 Trends in Conifer Disc Tree Rings and Meteorological Parameters

Results of investigations into the CO2 content in tree disc rings by the method proposed here have shown that a considerable part of CO2 generated under cell respiration is found in tree stems. Besides, annual CO2 distribution in tree rings exhibits a well-defined cyclicity. Results obtained from investigations into long-term CO2 and H2O variations in tree discs performed by different methods are presented. Wavelet and spectral analyses of the relationship between CO2 and H2O variations in a Siberian stone pine disc and meteorological parameters were made. The CO2 annual distributions of seven spruce tree discs were examined by the Caterpiller-SSA method. Wood samples of Siberian stone pine and spruce trees were taken from the same site in Tomsk region （Siberia, Russia） that characterized by an optimum growth habitat. Conclusions are made regarding the response of the annual CO2 and H2O conifer disc tree ring distributions to different climatic factors.

Climate-growth relationships of Pinus tabuliformis along an altitudinal gradient on Baiyunshan Mountain,Central China

A set of standard chronologies for tree-ring width(TRW),earlywood width(EWW)and latewood width(LWW)in Pinus tabuliformis Carr.along an altitudi-nal gradient(1450,1400,and 1350 m a.s.l.)on Baiyunshan Mountain,Central China to analyze the effect of varying temperature and precipitation on growth along the gradi-ent.Correlation analyses showed that at all three altitudes and the TRW and EWW chronologies generally had signifi-cant negative correlations with mean and maximum tem-peratures in the current April and May and with minimum temperatures in the prior July and August,but significant positive correlations with precipitation in the current May.Correlations were generally significantly negative between LWW chronologies and all temperatures in the prior July and August,indicating that the prior summer temperature had a strong lag effect on the growth of P.tabuliformis that increased with altitude.The correlation with the standard-ized precipitation evapotranspiration index(SPEI)confirmed that wet conditions in the current May promoted growth of TR and EW at all altitudes.Significant altitudinal differences were also found;at 1400 m,there were significant positive correlations between EWW chronologies and SPEI in the current April and significant negative correlations between LWW chronologies and SPEI in the current September,but these correlations were not significant at 1450 m.At 1350 m,there were also significant negative correlations between the TRW and the EWW chronologies and SPEI in the prior October and the current July and between LWW chronology and SPEI in the current August,but these cor-relations were not significant at 1400 m.Moving correlation results showed a stable response of EWW in relation to the SPEI in the current May at all three altitudes and of LWW to maximum temperature in the prior July-August at 1400 m from 2002 to 2018.The EWW chronology at 1400 m and the LWW chronology at 1450 m were identified as more suitable for climate reconstruction.These results provide a strong scientific basis for forest management decisions and climate reconstructions in Central China.

Elevation response of above-ground net primary productivity for Picea crassifolia to climate change in Qilian Mountains of Northwest China based on tree rings

Current ecosystem models used to simulate global terrestrial carbon balance generally suggest that terrestrial landscapes are stable and mature,but terrestrial net primary productivity(NPP)data estimated without accounting for disturbances in species composition,environment,structure,and ecological characteristics will reduce the accuracy of the global carbon budget.Therefore,the steady-state assumption and neglect of elevation-related changes in forest NPP is a concern.The Qilian Mountains are located in continental climate zone,and vegetation is highly sensitive to climate change.We quantified aboveground biomass(AGB)and aboveground net primary productivity(ANPP)sequences at three elevations using field-collected tree rings of Picea crassifolia in Qilian Mountains of Northwest China.The results showed that(1)There were significant differences between AGB and ANPP at the three elevations,and the growth rate of AGB was the highest at the low elevation(55.99 t ha^(–1)10a^(–1)).(2)There are differences in the response relationship between the ANPP and climate factors at the three elevations,and drought stress is the main climate signal affecting the change of ANPP.(3)Under the future climate scenario,drought stress intensifies,and the predicted decline trend of ANPP at the three elevations from mid-century to the end of this century is–0.025 t ha^(–1)10a^(–1),respectively;–0.022 t ha^(–1)10a^(–1);At–0.246 t ha^(–1)10a^(–1),the level of forest productivity was significantly degraded.The results reveal the elevation gradient differences in forest productivity levels and provide key information for studying the carbon sink potential of boreal forests.