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.

Is subarctic forest advance able to keep pace with climate change?

Recent climate warming and scenarios for further warming have led to expectations of rapid movement of ecological boundaries. Here we focus on the circumarctic forest-tundra ecotone (FTE), which represents an important bioclimatic zone with feedbacks from forest advance and corresponding tundra disappearance (up to 50% loss predicted this century) driving widespread ecological and climatic changes. We address FTE advance and climate history relations over the 20th century, using FTE response data from 151 sites across the circumarctic area and site-specific climate data. Specifically, we investigate spatial uniformity of FTE advance, statistical associations with 20th century climate trends, and whether advance rates match climate change velocities (CCVs). Study sites diverged into four regions (Eastern Canada; Central and Western Canada and Alaska; Siberia; and Western Eurasia) based on their climate history, although all were characterized by similar qualitative patterns of behaviour (with about half of the sites showing advancing behaviour). The main associations between climate trend variables and behaviour indicate the importance of precipitation rather than temperature for both qualitative and quantitative behaviours, and the importance of non-growing season as well as growing season months. Poleward latitudinal advance rates differed significantly among regions, being smallest in Eastern Canada (~10 m/year) and largest in Western Eurasia (~100 m/year). These rates were 1-2 orders of magnitude smaller than expected if vegetation distribution remained in equilibrium with climate. The many biotic and abiotic factors influencing FTE behaviour make poleward advance rates matching predicted 21st century CCVs (~103 -104  m/year) unlikely. The lack of empirical evidence for swift forest relocation and the discrepancy between CCV and FTE response contradict equilibrium model-based assumptions and warrant caution when assessing global-change-related biotic and abiotic implications, including land-atmosphere feedbacks and carbon sequestration.

Experiment, monitoring, and gradient methods used to infer climate change effects on plant communities yield consistent patterns.

Inference about future climate change impacts typically relies on one of three approaches: manipulative experiments, historical comparisons (broadly defined to include monitoring the response to ambient climate fluctuations using repeat sampling of plots, dendroecology, and paleoecology techniques), and space-for-time substitutions derived from sampling along environmental gradients. Potential limitations of all three approaches are recognized. Here we address the congruence among these three main approaches by comparing the degree to which tundra plant community composition changes (i) in response to in situ experimental warming, (ii) with interannual variability in summer temperature within sites, and (iii) over spatial gradients in summer temperature. We analyzed changes in plant community composition from repeat sampling (85 plant communities in 28 regions) and experimental warming studies (28 experiments in 14 regions) throughout arctic and alpine North America and Europe. Increases in the relative abundance of species with a warmer thermal niche were observed in response to warmer summer temperatures using all three methods; however, effect sizes were greater over broad-scale spatial gradients relative to either temporal variability in summer temperature within a site or summer temperature increases induced by experimental warming. The effect sizes for change over time within a site and with experimental warming were nearly identical. These results support the view that inferences based on space-for-time substitution overestimate the magnitude of responses to contemporary climate warming, because spatial gradients reflect long-term processes. In contrast, in situ experimental warming and monitoring approaches yield consistent estimates of the magnitude of response of plant communities to climate warming.

Climate warming feedback from mountain birch forest expansion: reduced albedo dominates carbon uptake.

Expanding high-elevation and high-latitude forest has contrasting climate feedbacks through carbon sequestration (cooling) and reduced surface reflectance (warming), which are yet poorly quantified. Here, we present an empirically based projection of mountain birch forest expansion in south-central Norway under climate change and absence of land use. Climate effects of carbon sequestration and albedo change are compared using four emission metrics. Forest expansion was modeled for a projected 2.6 °C increase in summer temperature in 2100, with associated reduced snow cover. We find that the current (year 2000) forest line of the region is circa 100 m lower than its climatic potential due to land-use history. In the future scenarios, forest cover increased from 12% to 27% between 2000 and 2100, resulting in a 59% increase in biomass carbon storage and an albedo change from 0.46 to 0.30. Forest expansion in 2100 was behind its climatic potential, forest migration rates being the primary limiting factor. In 2100, the warming caused by lower albedo from expanding forest was 10 to 17 times stronger than the cooling effect from carbon sequestration for all emission metrics considered. Reduced snow cover further exacerbated the net warming feedback. The warming effect is considerably stronger than previously reported for boreal forest cover, because of the typically low biomass density in mountain forests and the large changes in albedo of snow-covered tundra areas. The positive climate feedback of high-latitude and high-elevation expanding forests with seasonal snow cover exceeds those of afforestation at lower elevation, and calls for further attention of both modelers and empiricists. The inclusion and upscaling of these climate feedbacks from mountain forests into global models is warranted to assess the potential global impacts.

Global assessment of experimental climate warming on tundra vegetation: heterogeneity over space and time.

Understanding the sensitivity of tundra vegetation to climate warming is critical to forecasting future biodiversity and vegetation feedbacks to climate. In situ warming experiments accelerate climate change on a small scale to forecast responses of local plant communities. Limitations of this approach include the apparent site-specificity of results and uncertainty about the power of short-term studies to anticipate longer term change. We address these issues with a synthesis of 61 experimental warming studies, of up to 20 years duration, in tundra sites worldwide. The response of plant groups to warming often differed with ambient summer temperature, soil moisture and experimental duration. Shrubs increased with warming only where ambient temperature was high, whereas graminoids increased primarily in the coldest study sites. Linear increases in effect size over time were frequently observed. There was little indication of saturating or accelerating effects, as would be predicted if negative or positive vegetation feedbacks were common. These results indicate that tundra vegetation exhibits strong regional variation in response to warming, and that in vulnerable regions, cumulative effects of long-term warming on tundra vegetation - and associated ecosystem consequences - have the potential to be much greater than we have observed to date.

Reproduction and seedling establishment of Picea glauca across the northernmost forest-tundra region in Canada.

The northern boundary of boreal forest and the ranges of tree species are expected to shift northward in response to climate warming, which will result in a decrease in the albedo of areas currently covered by tundra vegetation, an increase in terrestrial carbon sequestration, and an alteration of biodiversity in the current Low Arctic. Central to the prediction of forest expansion is an increase in the reproductive capacity and establishment of individual trees. We assessed cone production, seed viability, and transplanted seedling success of Picea glauca (Moench.) Voss. (white spruce) in the early 1990s and again in the late 2000s at four forest stand sites and eight tree island sites (clonal populations beyond present treeline) in the Mackenzie Delta region of the Northwest Territories, Canada. Over the past 20 years, average temperatures in this region have increased by 0.9 °C. This area has the northernmost forest-tundra ecotone in North America and is one of the few circumpolar regions where the northern limit of conifer trees reaches the Arctic Ocean. We found that cone production and seed viability did not change between the two periods of examination and that both variables decreased northward across the forest-tundra ecotone. Nevertheless, white spruce individuals at the northern limit of the forest-tundra ecotone produced viable seeds. Furthermore, transplanted seedlings were able to survive in the northernmost sites for 15 years, but there were no signs of natural regeneration. These results indicate that if climatic conditions continue to ameliorate, reproductive output will likely increase, but seedling establishment and forest expansion within the forest-tundra of this region is unlikely to occur without the availability of suitable recruitment sites. Processes that affect the availability of recruitment sites are likely to be important elsewhere in the circumpolar ecotone, and should be incorporated into models and predictions of climate change and its effects on the northern forest-tundra ecotone.

Limited decadal growth of mountain birch saplings has minor impact on surrounding tundra vegetation.

Temperatures over the Arctic region are increasing at three times the rate of the global average. Consequently, Arctic vegetation is changing and trees are encroaching into the tundra. In this study, we examine the establishment and growth of mountain birch (Betula pubescens ssp. tortuosa), which forms the treeline in subarctic Europe, and its impact on community composition across the treeline ecotone nearby Abisko, Sweden. Birch advancement along elevational gradients was studied by comparing data collected in 2016 with data collected 10 and 15 years previously. Species identity, cover, and phylogenetic relatedness were used to assess the impact of birch encroachment on community composition. Our results show that birch occurrence above the treeline did not affect plant community composition, probably owing to the observed lack of significant growth due to herbivore browsing, nitrogen limitation, or a reduction in snow cover. Independent of birch performance, the tundra community structure shifted toward a novel community dissimilar from the forest plant community found below the treeline. Taken together, our findings are explained by species-specific responses to climate change, rather than by a linear forest advance. Future treeline advancements are likely more restricted than previously expected.

The missing pieces for better future predictions in subarctic ecosystems: A Torneträsk case study.

Arctic and subarctic ecosystems are experiencing substantial changes in hydrology, vegetation, permafrost conditions, and carbon cycling, in response to climatic change and other anthropogenic drivers, and these changes are likely to continue over this century. The total magnitude of these changes results from multiple interactions among these drivers. Field measurements can address the overall responses to different changing drivers, but are less capable of quantifying the interactions among them. Currently, a comprehensive assessment of the drivers of ecosystem changes, and the magnitude of their direct and indirect impacts on subarctic ecosystems, is missing. The Torneträsk area, in the Swedish subarctic, has an unrivalled history of environmental observation over 100 years, and is one of the most studied sites in the Arctic. In this study, we summarize and rank the drivers of ecosystem change in the Torneträsk area, and propose research priorities identified, by expert assessment, to improve predictions of ecosystem changes. The research priorities identified include understanding impacts on ecosystems brought on by altered frequency and intensity of winter warming events, evapotranspiration rates, rainfall, duration of snow cover and lake-ice, changed soil moisture, and droughts. This case study can help us understand the ongoing ecosystem changes occurring in the Torneträsk area, and contribute to improve predictions of future ecosystem changes at a larger scale. This understanding will provide the basis for the future mitigation and adaptation plans needed in a changing climate.

Global negative vegetation feedback to climate warming responses of leaf litter decomposition rates in cold biomes.

Whether climate change will turn cold biomes from large long-term carbon sinks into sources is hotly debated because of the great potential for ecosystem-mediated feedbacks to global climate. Critical are the direction, magnitude and generality of climate responses of plant litter decomposition. Here, we present the first quantitative analysis of the major climate-change-related drivers of litter decomposition rates in cold northern biomes worldwide. Leaf litters collected from the predominant species in 33 global change manipulation experiments in circum-arctic-alpine ecosystems were incubated simultaneously in two contrasting arctic life zones. We demonstrate that longer-term, large-scale changes to leaf litter decomposition will be driven primarily by both direct warming effects and concomitant shifts in plant growth form composition, with a much smaller role for changes in litter quality within species. Specifically, the ongoing warming-induced expansion of shrubs with recalcitrant leaf litter across cold biomes would constitute a negative feedback to global warming. Depending on the strength of other (previously reported) positive feedbacks of shrub expansion on soil carbon turnover, this may partly counteract direct warming enhancement of litter decomposition.

Stand structure, recruitment and growth dynamics in mixed subalpine spruce and Swiss stone pine forests in the Eastern Carpathians.

Natural subalpine forests are considered to be sensitive to climate change, and forest characteristics are assumed to reflect the prevalent disturbance regime. We hypothesize that stand history determines different stand structures. Based on large full inventory datasets (including tree biometric data, spatial coordinates, tree age, and basal area increment) we assessed the size structure, tree recruitment dynamics and radial growth patterns in three permanent plots along an altitudinal gradient in a mixed coniferous forest (Picea abies and Pinus cembra) in the Eastern Carpathians. Both discrete disturbances (large scale or small scale) and chronic disturbances (climate change) were identified as drivers of stand structure development in the studied plots. A stand replacing wind disturbance generated a unimodal bell-shaped size and age distribution for both species characterized by a sharp increase in post-disturbance recruitment. By contrast, small-scale wind-caused gaps led to a negative exponential diameter distribution for spruce and a left-asymmetric unimodal for pine. Climate-driven infilling processes in the upper subalpine forest were reflected as J-shaped size and age distributions for both species, but with pine predating spruce. The growth patterns for both species demonstrated an increased basal area increment since the early 1900s, with an emphasis in the last few decades, irrespective of stand history. Pine demonstrated a competitive advantage compared to spruce due to the higher growth rate and size at the same age. Recognition of combined discrete and chronic disturbances as drivers of the tree layer characteristics in a subalpine coniferous forest is essential in both stand history analyses and growth predictions.

Seed rain quantity and quality, 1984-1992, in a high altitude old-growth spruce forest, northern Sweden.

Short-term regeneration ability of a cold-marginal Picea abies forest was studied by analyzing spruce seed rain during nine years and by assessing resulting recruitment at the end of the studied period. Quantitatively large differences occurred between studied years, and seed viability was low during the whole study period. Only exceptionally did seed viability exceed 1 %. Sparse regeneration occurred in 1984, which is the year when most viable seeds/ha were recorded. In addition, the results are compared with recruitment data from the 1930s (from the same forest), a time period when regeneration peaked. It is suggested that the period 1984-1992, due to the sparse regeneration, retrospectively probably will appear as a regeneration trough or a decreasing regeneration trend. Furthermore, it is concluded that, in a long-term perspective, periodic regeneration is sufficient for maintenance of the forest because of the longevity of spruce. Additionally, climatically induced variations of regeneration success give rise to natural fluctuations in stand density.

Natural causes of the tundra-taiga boundary.

The tundra-taiga interface is characterized by a change in tree cover or density, tree size and shape, tree growth, and reproduction. Generally, trees get denser, taller, and less damaged as one moves from the tundra into the taiga proper. The environmental covariates and possible mechanisms resulting in these patterns are addressed in the paper. Low seed rain density, lack of safe sites caused by microclimatic variation, low surface substrate moisture, and low soil nutrient availability may limit the density of the tree species. Tree growth may be limited by a short growing season and further diminished, by shoot and root damage reducing carbon and nutrient stores as well as by reducing carbon and nutrient uptake capacities. Positive and negative feedbacks of tree density on tree growth exist at treeline. Increased tree density leads to increased air temperature and decreased wind damage, but also to lower soil temperature, reduced nutrient availability, and greater nutrient competition.

The dynamics of the tundra-taiga boundary: an overview and suggested coordinated and integrated approach to research.

The tundra-taiga boundary stretches for more than 13,400 km around the Northern Hemisphere and is probably the Earth's greatest vegetation transition. The trees that define the boundary have been sensitive to climate changes in the past and models of future vegetation distribution suggest a rapid and dramatic invasion of the tundra by the taiga. Such changes would generate both positive and negative feedbacks to the climate system and the balance could result in a net warming effect. However, the boundary is becoming increasingly affected by human activities that remove trees and degrade forest-tundra into tundra-like areas. Because of the vastness and remoteness of the tundra-taiga boundary, and of methodological problems such as problematic definitions and lack of standardized methods to record the location and characteristics of the ecotone, a project group has been established under the auspices of the International Arctic Science Committee (IASC). This paper summarizes the initial output of the group and focuses on our uncertainties in understanding the current processes at the tundra-taiga boundary and the conflicts between model predictions of changes in the location of the boundary and contrasting recently observed changes due to human activities. Finally, we present recommendations for a coordinated international approach to the problem and invite the international community to join us in reducing the uncertainties about the dynamics of the ecotone and their consequences.

Variable temperature effects of Open Top Chambers at polar and alpine sites explained by irradiance and snow depth.

Environmental manipulation studies are integral to determining biological consequences of climate warming. Open Top Chambers (OTCs) have been widely used to assess summer warming effects on terrestrial biota, with their effects during other seasons normally being given less attention even though chambers are often deployed year-round. In addition, their effects on temperature extremes and freeze-thaw events are poorly documented. To provide robust documentation of the microclimatic influences of OTCs throughout the year, we analysed temperature data from 20 studies distributed across polar and alpine regions. The effects of OTCs on mean temperature showed a large range (-0.9 to 2.1 °C) throughout the year, but did not differ significantly between studies. Increases in mean monthly and diurnal temperature were strongly related (R(2)  = 0.70) with irradiance, indicating that PAR can be used to predict the mean warming effect of OTCs. Deeper snow trapped in OTCs also induced higher temperatures at soil/vegetation level. OTC-induced changes in the frequency of freeze-thaw events included an increase in autumn and decreases in spring and summer. Frequency of high-temperature events in OTCs increased in spring, summer and autumn compared with non-manipulated control plots. Frequency of low-temperature events was reduced by deeper snow accumulation and higher mean temperatures. The strong interactions identified between aspects of ambient environmental conditions and effects of OTCs suggest that a detailed knowledge of snow depth, temperature and irradiance levels enables us to predict how OTCs will modify the microclimate at a particular site and season. Such predictive power allows a better mechanistic understanding of observed biotic response to experimental warming studies and for more informed design of future experiments. However, a need remains to quantify OTC effects on water availability and wind speed (affecting, for example, drying rates and water stress) in combination with microclimate measurements at organism level.