Fennoscandian tree-ring anatomy shows a warmer modern than medieval climate.

Earth system models and various climate proxy sources indicate global warming is unprecedented during at least the Common Era1. However, tree-ring proxies often estimate temperatures during the Medieval Climate Anomaly (950-1250 CE) that are similar to, or exceed, those recorded for the past century2,3, in contrast to simulation experiments at regional scales4. This not only calls into question the reliability of models and proxies but also contributes to uncertainty in future climate projections5. Here we show that the current climate of the Fennoscandian Peninsula is substantially warmer than that of the medieval period. This highlights the dominant role of anthropogenic forcing in climate warming even at the regional scale, thereby reconciling inconsistencies between reconstructions and model simulations. We used an annually resolved 1,170-year-long tree-ring record that relies exclusively on tracheid anatomical measurements from Pinus sylvestris trees, providing high-fidelity measurements of instrumental temperature variability during the warm season. We therefore call for the construction of more such millennia-long records to further improve our understanding and reduce uncertainties around historical and future climate change at inter-regional and eventually global scales.

Axial conduit widening, tree height, and height growth rate set the hydraulic transition of sapwood into heartwood.

The size-related xylem adjustments required to maintain a constant leaf-specific sapwood conductance (KLEAF) with increasing height (H) are still under discussion. Alternative hypotheses are that: (i) the conduit hydraulic diameter (Dh) at any position in the stem and/or (ii) the number of sapwood rings at stem base (NSWr) increase with H. In addition, (iii) reduced stem elongation (ΔH) increases the tip-to-base conductance through inner xylem rings, thus possibly the NSWr contributing to KLEAF. A detailed stem analysis showed that Dh increased with the distance from the ring apex (DCA) in all rings of a Picea abies and a Fagus sylvatica tree. Net of DCA effect, Dh did not increase with H. Using sapwood traits from a global dataset, NSWr increased with H, decreased with ΔH, and the mean sapwood ring width (SWrw) increased with ΔH. A numerical model based on anatomical patterns predicted the effects of H and ΔH on the conductance of inner xylem rings. Our results suggest that the sapwood/heartwood transition depends on both H and ΔH, and is set when the carbon allocation to maintenance respiration of living cells in inner sapwood rings produces a lower gain in total conductance than investing the same carbon in new vascular conduits.

Growth form and leaf habit drive contrasting effects of Arctic amplification in long-lived woody species.

Current global change is inducing heterogeneous warming trends worldwide, with faster rates at higher latitudes in the Northern Hemisphere. Consequently, tundra vegetation is experiencing an increase in growth rate and uneven but expanding distribution. Yet, the drivers of this heterogeneity in woody species responses are still unclear. Here, applying a retrospective approach and focusing on long-term responses, we aim to get insight into growth trends and climate sensitivity of long-lived woody species belonging to different functional types with contrasting growth forms and leaf habits (shrub vs. tree and deciduous vs. evergreen). A total of 530 samples from 7 species (common juniper, dwarf birch, woolly willow, Norway spruce, lodgepole pine, rowan, and downy birch) were collected in 10 sites across Iceland. We modelled growth trends and contrasted yearly ring-width measurements, filtering in high- and low-frequency components, with precipitation, land- and sea-surface temperature records (1967-2018). Shrubs and trees showed divergent growth trends, with shrubs closely tracking the recent warming, whereas trees, especially broadleaved, showed strong fluctuations but no long-term growth trends. Secondary growth, particularly the high-frequency component, was positively correlated with summer temperatures for most of the species. On the contrary, growth responses to sea surface temperature, especially in the low frequency, were highly diverging between growth forms, with a strong positive association for shrubs and a negative for trees. Within comparable vegetation assemblage, long-lived woody species could show contrasting responses to similar climatic conditions. Given the predominant role of oceanic masses in shaping climate patterns in the Arctic and Low Arctic, further investigations are needed to deepen the knowledge on the complex interplay between coastal tundra ecosystems and land-sea surface temperature dynamics.

Retrospective analysis of wood anatomical traits and tree-ring isotopes suggests site-specific mechanisms triggering Araucaria araucana drought-induced dieback.

In 2010-2018, Northern Patagonia featured the longest severe drought of the last millennium. This extreme dry spell triggered widespread growth decline and forest dieback. Nonetheless, the roles played by the two major mechanisms driving dieback, hydraulic failure and carbon starvation, are still not clear and understudied in this seasonally dry region. Here, for the 1800-2017 period, we apply a retrospective analysis of radial growth, wood anatomical traits (lumen area, cell-wall thickness) and δ13 C and δ18 O stable isotopes to assess dieback causes of the iconic conifer Araucaria araucana. We selected three stands where declining (defoliated) and nondeclining (not defoliated) trees coexisted along a precipitation gradient from the warm-dry Coastal Range to the cool-wet Andes. At all sites declining trees showed lower radial growth and lower theoretical hydraulic conductivity, suggesting a long-lasting process of hydraulic deterioration in their water transport system compared to nondeclining, coexisting trees. Wood anatomical traits evidenced that this divergence between declining and nondeclining trees started at least seven decades before canopy dieback. In the drier stands, declining trees showed higher water-use efficiency (WUE) throughout the whole period, which we attributed to early stomatal closure, suggesting a greater carbon starvation risk consistent with thinner cell walls. In the wettest stand, we found the opposite pattern. Here, a reduction in WUE coupled with thicker cell walls suggested increased carbon assimilation rates and exposure to drought-induced hydraulic failure. The δ18 O values indicated different strategies of gas exchange between sites, which are likely a consequence of microsite conditions and water sources. Multiproxy, retrospective quantifications of xylem anatomical traits and tree-ring isotopes provide a robust tool to identify and forecast, which stands or trees will show dieback or, on the contrary, which will likely withstand and be more resilient to future hotter droughts.

Global fading of the temperature-growth coupling at alpine and polar treelines.

Climate warming is expected to positively alter upward and poleward treelines which are controlled by low temperature and a short growing season. Despite the importance of treelines as a bioassay of climate change, a global field assessment and posterior forecasting of tree growth at annual scales is lacking. Using annually resolved tree-ring data located across Eurasia and the Americas, we quantified and modeled the relationship between temperature and radial growth at treeline during the 20th century. We then tested whether this temperature-growth association will remain stable during the 21st century using a forward model under two climate scenarios (RCP 4.5 and 8.5). During the 20th century, growth enhancements were common in most sites, and temperature and growth showed positive trends. Interestingly, the relationship between temperature and growth trends was contingent on tree age suggesting biogeographic patterns in treeline growth are contingent on local factors besides climate warming. Simulations forecast temperature-growth decoupling during the 21st century. The growing season at treeline is projected to lengthen and growth rates would increase and become less dependent on temperature rise. These forecasts illustrate how growth may decouple from climate warming in cold regions and near the margins of tree existence. Such projected temperature-growth decoupling could impact ecosystem processes in mountain and polar biomes, with feedbacks on climate warming.

Wood anatomical traits in black spruce reveal latent water constraints on the boreal forest.

The effects of climate change on high-latitude forest ecosystems are complex, making forecasts of future scenarios uncertain. The predicted lengthening of the growing season under warming conditions is expected to increase tree growth rates. However, there is evidence of an increasing sensitivity of the boreal forest to drought stress. To assess the influence of temperature and precipitation on the growth of black spruce (Picea mariana), we investigated long-term series of wood anatomical traits on 20 trees from four sites along 600 km, the latitudinal range of the closed boreal forest in Quebec, Canada. We correlated the anatomical traits resolved at intraring level with daily temperature, vapor pressure deficit (VPD), and precipitation during the 1943-2010 period. Tree-ring width, number of cells per ring and cell wall thickness were positively affected by spring and summer daily mean and maximum temperature at the northern sites. These results agree with the well-known positive effect of high temperatures on tree ring formation at high latitudes. However, we captured, for the first time in this region, the latent impact of water availability on xylem traits. Indeed, in all the four sites, cell lumen area showed positive correlations with daily precipitation (mostly at low latitude), and/or negative correlations with daily mean and maximum temperature and VPD (mostly at high latitude). We inferred that drought, due to high temperatures, low precipitations, or both, negatively affects cell enlargement across the closed boreal forest, including the northernmost sites. The production of tracheids with narrower lumen, potentially more resistant to cavitation, could increase xylem hydraulic safety under a warmer and drier climate. However, this would result in lower xylem conductivity, with consequent long-term hydraulic deterioration, growth decline, and possibly lead to tree dieback, as observed in other forest ecosystems at lower latitudes.

Moisture-driven shift in the climate sensitivity of white spruce xylem anatomical traits is coupled to large-scale oscillation patterns across northern treeline in northwest North America.

Tree growth at northern treelines is generally temperature-limited due to cold and short growing seasons. However, temperature-induced drought stress was repeatedly reported for certain regions of the boreal forest in northwestern North America, provoked by a significant increase in temperature and possibly reinforced by a regime shift of the pacific decadal oscillation (PDO). The aim of this study is to better understand physiological growth reactions of white spruce, a dominant species of the North American boreal forest, to PDO regime shifts using quantitative wood anatomy and traditional tree-ring width (TRW) analysis. We investigated white spruce growth at latitudinal treeline across a >1,000 km gradient in northwestern North America. Functionally important xylem anatomical traits (lumen area, cell-wall thickness, cell number) and TRW were correlated with the drought-sensitive standardized precipitation-evapotranspiration index of the growing season. Correlations were computed separately for complete phases of the PDO in the 20th century, representing alternating warm/dry (1925-1946), cool/wet (1947-1976) and again warm/dry (1977-1998) climate regimes. Xylem anatomical traits revealed water-limiting conditions in both warm/dry PDO regimes, while no or spatially contrasting associations were found for the cool/wet regime, indicating a moisture-driven shift in growth-limiting factors between PDO periods. TRW reflected only the last shift of 1976/1977, suggesting different climate thresholds and a higher sensitivity to moisture availability of xylem anatomical traits compared to TRW. This high sensitivity of xylem anatomical traits permits to identify first signs of moisture-driven growth in treeline white spruce at an early stage, suggesting quantitative wood anatomy being a powerful tool to study climate change effects in the northwestern North American treeline ecotone. Projected temperature increase might challenge growth performance of white spruce as a key component of the North American boreal forest biome in the future, when drier conditions are likely to occur with higher frequency and intensity.

Climate extremes and predicted warming threaten Mediterranean Holocene firs forests refugia.

Warmer and drier climatic conditions are projected for the 21st century; however, the role played by extreme climatic events on forest vulnerability is still little understood. For example, more severe droughts and heat waves could threaten quaternary relict tree refugia such as Circum-Mediterranean fir forests (CMFF). Using tree-ring data and a process-based model, we characterized the major climate constraints of recent (1950-2010) CMFF growth to project their vulnerability to 21st-century climate. Simulations predict a 30% growth reduction in some fir species with the 2050s business-as-usual emission scenario, whereas growth would increase in moist refugia due to a longer and warmer growing season. Fir populations currently subjected to warm and dry conditions will be the most vulnerable in the late 21st century when climatic conditions will be analogous to the most severe dry/heat spells causing dieback in the late 20th century. Quantification of growth trends based on climate scenarios could allow defining vulnerability thresholds in tree populations. The presented predictions call for conservation strategies to safeguard relict tree populations and anticipate how many refugia could be threatened by 21st-century dry spells.

Wood anatomical traits highlight complex temperature influence on Pinus cembra at high elevation in the Eastern Alps.

Climate sensitivity of populations at the margins of their distribution range is of key importance to understand species' responses to future warming conditions. Pinus cembra is of particular interest being a typical high-elevation taxon, spread with mostly scattered populations within its actual range, but still overlooked in traditional dendrochronological researches due to low tree-ring variability and climate sensitivity. With a different approach, we analyzed time series of xylem anatomical traits, split into intra-ring sectors, and used daily climate records over 89 years (1926-2014) aiming to improve the quality and time resolution of the climate/growth associations. From nine trees growing at their altitudinal limit and on 1.5 × 106 tracheids, we measured ring width (MRW), cell number per ring, lumen area (LA), and cell-wall thickness (CWT). We then computed correlations with monthly and fortnightly climate data. Late-spring and summer temperature emerged as the most important factors. LA and especially CWT showed a stronger temperature response than MRW, starting in mid-May and early June, respectively. CWT also evidenced the longest period of correlations with temperature and a significant difference between latewood radial and tangential walls. Analysis of xylem anatomical traits at intra-ring level and the use of daily temperature records proved to be useful for high resolution and detailed climate/growth association inferences in Pinus cembra.

Cell size and wall dimensions drive distinct variability of earlywood and latewood density in Northern Hemisphere conifers.

Interannual variability of wood density - an important plant functional trait and environmental proxy - in conifers is poorly understood. We therefore explored the anatomical basis of density. We hypothesized that earlywood density is determined by tracheid size and latewood density by wall dimensions, reflecting their different functional tasks. To determine general patterns of variability, density parameters from 27 species and 349 sites across the Northern Hemisphere were correlated to tree-ring width parameters and local climate. We performed the same analyses with density and width derived from anatomical data comprising two species and eight sites. The contributions of tracheid size and wall dimensions to density were disentangled with sensitivity analyses. Notably, correlations between density and width shifted from negative to positive moving from earlywood to latewood. Temperature responses of density varied intraseasonally in strength and sign. The sensitivity analyses revealed tracheid size as the main determinant of earlywood density, while wall dimensions become more influential for latewood density. Our novel approach of integrating detailed anatomical data with large-scale tree-ring data allowed us to contribute to an improved understanding of interannual variations of conifer growth and to illustrate how conifers balance investments in the competing xylem functions of hydraulics and mechanical support.

Diverging shrub and tree growth from the Polar to the Mediterranean biomes across the European continent.

Climate warming is expected to enhance productivity and growth of woody plants, particularly in temperature-limited environments at the northernmost or uppermost limits of their distribution. However, this warming is spatially uneven and temporally variable, and the rise in temperatures differently affects biomes and growth forms. Here, applying a dendroecological approach with generalized additive mixed models, we analysed how the growth of shrubby junipers and coexisting trees (larch and pine species) responds to rising temperatures along a 5000-km latitudinal range including sites from the Polar, Alpine to the Mediterranean biomes. We hypothesize that, being more coupled to ground microclimate, junipers will be less influenced by atmospheric conditions and will less respond to the post-1950 climate warming than coexisting standing trees. Unexpectedly, shrub and tree growth forms revealed divergent growth trends in all the three biomes, with juniper performing better than trees at Mediterranean than at Polar and Alpine sites. The post-1980s decline of tree growth in Mediterranean sites might be induced by drought stress amplified by climate warming and did not affect junipers. We conclude that different but coexisting long-living growth forms can respond differently to the same climate factor and that, even in temperature-limited area, other drivers like the duration of snow cover might locally play a fundamental role on woody plants growth across Europe.

How does climate influence xylem morphogenesis over the growing season? Insights from long-term intra-ring anatomy in Picea abies.

Background and Aims: During the growing season, the cambium of conifer trees produces successive rows of xylem cells, the tracheids, that sequentially pass through the phases of enlargement and secondary wall thickening before dying and becoming functional. Climate variability can strongly influence the kinetics of morphogenetic processes, eventually affecting tracheid shape and size. This study investigates xylem anatomical structure in the stem of Picea abies to retrospectively infer how, in the long term, climate affects the processes of cell enlargement and wall thickening. Methods: Tracheid anatomical traits related to the phases of enlargement (diameter) and wall thickening (wall thickness) were innovatively inspected at the intra-ring level on 87-year-long tree-ring series in Picea abies trees along a 900 m elevation gradient in the Italian Alps. Anatomical traits in ten successive tree-ring sectors were related to daily temperature and precipitation data using running correlations. Key Results: Close to the altitudinal tree limit, low early-summer temperature negatively affected cell enlargement. At lower elevation, water availability in early summer was positively related to cell diameter. The timing of these relationships shifted forward by about 20 (high elevation) to 40 (low elevation) d from the first to the last tracheids in the ring. Cell wall thickening was affected by climate in a different period in the season. In particular, wall thickness of late-formed tracheids was strongly positively related to August-September temperature at high elevation. Conclusions: Morphogenesis of tracheids sequentially formed in the growing season is influenced by climate conditions in successive periods. The distinct climate impacts on cell enlargement and wall thickening indicate that different morphogenetic mechanisms are responsible for different tracheid traits. Our approach of long-term and high-resolution analysis of xylem anatomy can support and extend short-term xylogenesis observations, and increase our understanding of climate control of tree growth and functioning under different environmental conditions.

Wood anatomy and carbon-isotope discrimination support long-term hydraulic deterioration as a major cause of drought-induced dieback.

Hydraulic impairment due to xylem embolism and carbon starvation are the two proposed mechanisms explaining drought-induced forest dieback and tree death. Here, we evaluate the relative role played by these two mechanisms in the long-term by quantifying wood-anatomical traits (tracheid size and area of parenchyma rays) and estimating the intrinsic water-use efficiency (iWUE) from carbon isotopic discrimination. We selected silver fir and Scots pine stands in NE Spain with ongoing dieback processes and compared trees showing contrasting vigour (declining vs nondeclining trees). In both species earlywood tracheids in declining trees showed smaller lumen area with thicker cell wall, inducing a lower theoretical hydraulic conductivity. Parenchyma ray area was similar between the two vigour classes. Wet spring and summer conditions promoted the formation of larger lumen areas, particularly in the case of nondeclining trees. Declining silver firs presented a lower iWUE than conspecific nondeclining trees, but the reverse pattern was observed in Scots pine. The described patterns in wood anatomical traits and iWUE are coherent with a long-lasting deterioration of the hydraulic system in declining trees prior to their dieback. Retrospective quantifications of lumen area permit to forecast dieback in declining trees 2-5 decades before growth decline started. Wood anatomical traits provide a robust tool to reconstruct the long-term capacity of trees to withstand drought-induced dieback.

An allometry-based approach for understanding forest structure, predicting tree-size distribution and assessing the degree of disturbance.

Tree-size distribution is one of the most investigated subjects in plant population biology. The forestry literature reports that tree-size distribution trajectories vary across different stands and/or species, whereas the metabolic scaling theory suggests that the tree number scales universally as -2 power of diameter. Here, we propose a simple functional scaling model in which these two opposing results are reconciled. Basic principles related to crown shape, energy optimization and the finite-size scaling approach were used to define a set of relationships based on a single parameter that allows us to predict the slope of the tree-size distributions in a steady-state condition. We tested the model predictions on four temperate mountain forests. Plots (4 ha each, fully mapped) were selected with different degrees of human disturbance (semi-natural stands versus formerly managed). Results showed that the size distribution range successfully fitted by the model is related to the degree of forest disturbance: in semi-natural forests the range is wide, whereas in formerly managed forests, the agreement with the model is confined to a very restricted range. We argue that simple allometric relationships, at an individual level, shape the structure of the whole forest community.

Self-similarity and scaling in forest communities.

Ecological communities exhibit pervasive patterns and interrelationships between size, abundance, and the availability of resources. We use scaling ideas to develop a unified, model-independent framework for understanding the distribution of tree sizes, their energy use, and spatial distribution in tropical forests. We demonstrate that the scaling of the tree crown at the individual level drives the forest structure when resources are fully used. Our predictions match perfectly with the scaling behavior of an exactly solvable self-similar model of a forest and are in good accord with empirical data. The range, over which pure power law behavior is observed, depends on the available amount of resources. The scaling framework can be used for assessing the effects of natural and anthropogenic disturbances on ecosystem structure and functionality.

Tree-ring and remote sensing analyses uncover the role played by elevation on European beech sensitivity to late spring frost.

Extreme climate events such as late spring frosts (LSFs) negatively affect productivity and tree growth in temperate beech forests. However, detailed information on how these forests recover after such events are still missing. We investigated how LSFs affected forest cover and radial growth in European beech (Fagus sylvatica L.) populations located at different elevations at four sites in the Italian Apennines, where LSFs have been recorded. We combined tree-ring and remote-sensing data to analyse the sensitivity and recovery capacity of beech populations to LSFs. Using daily temperature records, we reconstructed LSF events and assessed legacy effects on growth. We also evaluated the role played by elevation and stand structure as modulators of LSFs impacts. Finally, using satellite images we computed Normalized Difference Vegetation Index (NDVI), Enhanced Vegetation Index (EVI) and LAI (Leaf Area Index) to evaluate the post-LSF canopy recovery. The growth reduction in LSF-affected trees ranged from 36 % to 84 %. We detected a negative impact of LSF on growth only during the LSF year, with growth recovery occurring within 1-2 years after the event. LSF-affected stands featured low vegetation indices until late June, i.e. on average 75 days after the frost events. We did not find a clear relationship between beech forest elevation and occurrence of LSFs defoliations. Our results indicate a high recovery capacity of common beech and no legacy effects of LSFs.

The admixture of Quercus sp. in Pinus sylvestris stands influences wood anatomical trait responses to climatic variability and drought events.

Introduction: Forests are threatened by increasingly severe and more frequent drought events worldwide. Mono-specific forests, developed as a consequence of widespread management practices established early last century, seem particularly susceptible to global warming and drought compared with mixed-species forests. Although, in several contexts, mixed-species forests display higher species diversity, higher productivity, and higher resilience, previous studies highlighted contrasting findings, with not only many positive but also neutral or negative effects on tree performance that could be related to tree species diversity. Processes underlying this relationship need to be investigated. Wood anatomical traits are informative proxies of tree functioning, and they can potentially provide novel long-term insights in this regard. However, wood anatomical traits are critically understudied in such a context. Here, we assess the role of tree admixture on Pinus sylvestris L. xylem traits such as mean hydraulic diameter, cell wall thickness, and anatomical wood density, and we test the variability of these traits in response to climatic parameters such as temperature, precipitation, and drought event frequency and intensity. Methods: Three monocultural plots of P. sylvestris and three mixed-stand plots of P. sylvestris and Quercus sp. were identified in Poland and Spain, representing Continental and Mediterranean climate types, respectively. In each plot, we analyzed xylem traits from three P. sylvestris trees, for a total of nine trees in monocultures and nine in mixed stands per study location. Results: The results highlighted that anatomical wood density was one of the most sensitive traits to detect tree responses to climatic conditions and drought under different climate and forest types. Inter-specific facilitation mechanisms were detected in the admixture between P. sylvestris and Quercus sp., especially during the early growing season and during stressful events such as spring droughts, although they had negligible effects in the late growing season. Discussion: Our findings suggest that the admixture between P. sylvestris and Quercus sp. increases the resilience of P. sylvestris to extreme droughts. In a global warming scenario, this admixture could represent a useful adaptive management option.

Contrasting Climate Sensitivity of Pinus cembra Tree-Ring Traits in the Carpathians.

High-elevation ecosystems are one of the most sensitive to climate change. The analysis of growth and xylem structure of trees from marginal populations, especially the ones growing at the treeline, could provide early-warning signs to better understand species-specific responses to future climate conditions. In this study, we combined classical dendrochronology with wood density and anatomical measurements to investigate the climate sensitivity of Pinus cembra L., a typical European high-elevation tree species distributed in isolated patches in the Carpathians. Samples were collected from the Retezat Mountains, South-Western Romania. We analyzed ring width (TRW), maximum density (MXD), xylem anatomical traits [cell number per ring (CNo), cell density (CD), conduit area (CA), and cell wall thickness (CWT)] time series, split into ring sectors and assessed the relationships with monthly and daily climate records over the last century (1901-2015). The analysis showed a strong dependency of TRW on CNo and MXD on CWT. Summer temperature positively correlated with MXD and CWT [monthly correlation (r) were 0.65 and 0.48 respectively] from the early to late wood but not TRW (r = 0.22). CA positively correlated with water availability (r = 0.37) and negatively correlated with temperature (r = -0.39). This study improves our general understanding of the climate-growth relationships of a European high-elevation tree species and the results could be considered for forecasting population dynamics on projected changes in climate.

Transient Effects of Snow Cover Duration on Primary Growth and Leaf Traits in a Tundra Shrub.

With the recent climate warming, tundra ecotones are facing a progressive acceleration of spring snowpack melting and extension of the growing season, with evident consequences to vegetation. Along with summer temperature, winter precipitation has been recently recognised as a crucial factor for tundra shrub growth and physiology. However, gaps of knowledge still exist on long-living plant responses to different snowpack duration, especially on how intra-specific and year-to-year variability together with multiple functional trait adjustments could influence the long-term responses. To fill this gap, we conducted a 3 years snow manipulation experiment above the Alpine treeline on the typical tundra species Juniperus communis, the conifer with the widest distributional range in the north emisphere. We tested shoot elongation, leaf area, stomatal density, leaf dry weight and leaf non-structural carbohydrate content of plants subjected to anticipated, natural and postponed snowpack duration. Anticipated snowpack melting enhanced new shoot elongation and increased stomatal density. However, plants under prolonged snow cover seemed to compensate for the shorter growing period, likely increasing carbon allocation to growth. In fact, these latter showed larger needles and low starch content at the beginning of the growing season. Variability between treatments slightly decreased over time, suggesting a progressive acclimation of juniper to new conditions. In the context of future warming scenarios, our results support the hypothesis of shrub biomass increase within the tundra biome. Yet, the picture is still far from being complete and further research should focus on transient and fading effects of changing conditions in the long term.

Influences of summer warming and nutrient availability on Salix glauca L. growth in Greenland along an ice to sea gradient.

The combined effects of climate change and nutrient availability on Arctic vegetation growth are poorly understood. Archaeological sites in the Arctic could represent unique nutrient hotspots for studying the long-term effect of nutrient enrichment. In this study, we analysed a time-series of ring widths of Salix glauca L. collected at nine archaeological sites and in their natural surroundings along a climate gradient in the Nuuk fjord region, Southwest Greenland, stretching from the edge of the Greenlandic Ice Sheet in the east to the open sea in the west. We assessed the temperature-growth relationship for the last four decades distinguishing between soils with past anthropogenic nutrient enrichment (PANE) and without (controls). Along the East-West gradient, the inner fjord sites showed a stronger temperature signal compared to the outermost ones. Individuals growing in PANE soils had wider ring widths than individuals growing in the control soils and a stronger climate-growth relation, especially in the inner fjord sites. Thereby, the individuals growing on the archaeological sites seem to have benefited more from the climate warming in recent decades. Our results suggest that higher nutrient availability due to past human activities plays a role in Arctic vegetation growth and should be considered when assessing both the future impact of plants on archaeological sites and the general greening in landscapes with contrasting nutrient availability.