Climate change is driving significant changes in alpine ecosystems, where temperature-sensitive plant communities are particularly vulnerable. As plants migrate to higher elevations, understanding the mechanisms behind these changes is critical for predicting ecosystem dynamics. Morphological, physiological, and chemical functional traits may help understand species' responses to environmental change. This study examines whether functional traits can explain recent changes in presence and cover of alpine plant species in permanent plots. We analyzed vegetation data from repeated surveys of permanent plots at two sites. Morphological traits (e.g., plant height, leaf area) and physiological traits (e.g., frost and drought resistance) were measured locally for common species and supplemented with data from the TRY trait database to broaden species coverage. Linear models were applied to assess the relationships between traits and abundance changes (presence and cover), using multiple metrics. Our results reveal that plant height and leaf area are significant predictors of species abundance changes, emphasizing the role of morphological traits in shaping alpine plant communities. In contrast, physiological traits showed limited explanatory power. Notably, leaf carbon content emerged as a key predictor, suggesting that conservative strategies may provide advantages under warming conditions. Measures describing biomass dynamics (e.g., cover) differed from those describing establishment (e.g., presence), highlighting the multifaceted nature of species responses to environmental changes. Although our findings emphasize the importance of competitive interactions and resource acquisition, the study is limited by the absence of traits related to heat tolerance and prolonged warming. Future research should address these gaps to better understand the impacts of sustained temperature increases on alpine ecosystems.

Climate change increases plant species richness in alpine ecosystems. However, to what extent this diversity enrichment masks extinction dynamics of resident species remains elusive. In this study, we used floristic resurvey data from 896 permanent vegetation plots across 62 European mountain summits to show that local extinctions have increased over the past 21 years. Extinction rates rose with the magnitude of warming, and species were more likely to go extinct toward their low-elevation range margins and in communities undergoing stronger thermophilization. Moreover, local extinctions were significantly related to preceding abundance declines, which can serve as an early warning signal. These findings suggest that despite increasing plant species richness, plant assemblages above the treeline face an accelerating but so far neglected loss of their most characteristic species.

Snow cover is a crucial driver for plant species distributions in cold environments. The primary source of snow cover data used in distribution models is remotely sensed satellite imagery, which is characterized by coarser spatial resolutions than plot-scale observations of plant distributions. This scale-mismatch was hypothesized to limit model accuracy. Here, we used a common modeling framework to assess the contribution of snow melt-out dates derived from four data sources (satellite imagery, numerical snowpack modeling, webcam imagery and in-situ soil temperature measurements) at 1 m and 20 m spatial resolution to the predictive power of distribution models of 74 plant species in an alpine landscape of the Austrian Alps. We found that >80 % of the distribution models of all species were significantly improved by at least one snow melt-out data set when considering Area Under the Curve (AUC). Satellite-based melt-out led to significantly improved models for the highest number of species (>50 % for AUC) and increased True-Skill-Statistic and AUC on average by 16 % and 5 %, respectively. Surprisingly, fine-scale and in-situ measured melt-out data did not improve models more than the coarser scale (20 m) satellite-based melt-out data. Moreover, numerical snowpack modeling delivered results comparable to the other sources, which supports its use for projecting future species distributions. We conclude that the additional effort needed for producing high resolution, in-situ datasets as compared to commonly used satellite imagery might hence be worthwhile for some species but not for plant distribution modeling in cold ecosystems in general.

Recent changes in alpine vegetation are often attributed to climate warming, particularly community composition shifts towards more warmth-associated species, or thermophilization. Here we assess the link between thermophilization and warming across 53 European mountain summits. We combine long-term macroclimatic and microclimatic temperature time series with vegetation surveys in 724 permanent plots, monitored over 21 years, to evaluate a possible thermophilization signal and relate it to rates of change in 10 temperature metrics. We find evidence of both thermophilization of alpine plant communities and an increase in temperatures. However, although these two trends are related when averaged across mountain regions, their relationship is weak at the individual plot scale, especially when considering microclimatic temperature metrics. Instead, substrate conditions and particularly the availability of thermophilic colonizers in the surrounding vegetation have a major influence on plot-level thermophilization rates. We conclude that the response of plant communities to climate change strongly depends on the abiotic and biotic context, and intensified monitoring efforts are needed to reduce the resulting uncertainties.

AimUnderstanding the resilience and adaptability of alpine flora under climate change is crucial for biodiversity conservation. While functional traits are key to predicting alpine plants' responses to climate change, the role of regeneration traits remains underexplored. We hypothesised that alpine species thriving under climate change produce seeds with higher dispersal ability, longer soil persistence, lower dormancy requirements, and faster germination, while declining species would show opposite traits.LocationTwenty-three summits across six mountain ranges in Central and Southern Europe: Sierra Nevada, Northern and Central Apennines, and Northeastern, Central, and Southern Alps.MethodsWe analysed long-term data on frequency and abundance changes and eight seed traits related to dispersal, establishment, and soil persistence for 177 alpine species using linear mixed-effect models.ResultsOver two decades, alpine plant populations remained stable, with nearly 90% of species showing minimal frequency change and 70% showing minimal abundance change. However, abundance shifts varied by region: 16%-25% of species declined in Sierra Nevada, the Central Apennines, and the Southern Alps, while the Northeastern Alps and the Northern Apennines showed the largest increases (27% and 17%, respectively). Significant but limited relationships between seed traits and population dynamics were captured, primarily in the Central and Northern Apennines. Species with lower potential for epizoochory or anemochory were more likely to increase in abundance, while smaller seeds were linked to 'winners' in some regions. Germination traits, such as broader temperature requirements and slower germination, characterised species with increased abundance in the Northern Apennines.Main ConclusionsSeed traits had limited predictive power in distinguishing 'losers' and 'winners' of climate change among European alpine plants. This likely reflects the longevity of alpine plants, short observation periods, and potential mismatches between seed-level microenvironmental conditions and broader climatic trends.

Although variation in effect sizes and predicted values among studies of similar phenomena is inevitable, such variation far exceeds what might be produced by sampling error alone. One possible explanation for variation among results is differences among researchers in the decisions they make regarding statistical analyses. A growing array of studies has explored this analytical variability in different fields and has found substantial variability among results despite analysts having the same data and research question. Many of these studies have been in the social sciences, but one small “many analyst” study found similar variability in ecology. We expanded the scope of this prior work by implementing a large-scale empirical exploration of the variation in effect sizes and model predictions generated by the analytical decisions of different researchers in ecology and evolutionary biology. We used two unpublished datasets, one from evolutionary ecology (blue tit, Cyanistes caeruleus, to compare sibling number and nestling growth) and one from conservation ecology (Eucalyptus, to compare grass cover and tree seedling recruitment). The project leaders recruited 174 analyst teams, comprising 246 analysts, to investigate the answers to prespecified research questions. Analyses conducted by these teams yielded 141 usable effects (compatible with our meta-analyses and with all necessary information provided) for the blue tit dataset, and 85 usable effects for the Eucalyptus dataset. We found substantial heterogeneity among results for both datasets, although the patterns of variation differed between them. For the blue tit analyses, the average effect was convincingly negative, with less growth for nestlings living with more siblings, but there was near continuous variation in effect size from large negative effects to effects near zero, and even effects crossing the traditional threshold of statistical significance in the opposite direction. In contrast, the average relationship between grass cover and Eucalyptus seedling number was only slightly negative and not convincingly different from zero, and most effects ranged from weakly negative to weakly positive, with about a third of effects crossing the traditional threshold of significance in one direction or the other. However, there were also several striking outliers in the Eucalyptus dataset, with effects far from zero. For both datasets, we found substantial variation in the variable selection and random effects structures among analyses, as well as in the ratings of the analytical methods by peer reviewers, but we found no strong relationship between any of these and deviation from the meta-analytic mean. In other words, analyses with results that were far from the mean were no more or less likely to have dissimilar variable sets, use random effects in their models, or receive poor peer reviews than those analyses that found results that were close to the mean. The existence of substantial variability among analysis outcomes raises important questions about how ecologists and evolutionary biologists should interpret published results, and how they should conduct analyses in the future.

Boreal and tundra plant communities are expected to change in biodiversity due to increasing global change pressures such as climate warming. One long-term scenario is increasing compositional similarity, i.e. biotic homogenization, which has been relatively little studied in high-latitude plant communities. Here, we study how the composition and diversity of heathland and tundra plant communities have changed in northern Fennoscandia over several decades. In 2013-2023, we resurveyed 275 historic vegetation plots, originally surveyed in 1964-1975, with percentage covers for vascular plant, bryophyte and lichen species. We analyzed temporal changes in community composition and diversity across the study area and in different biogeographic zones, continentality-humidity classes and habitat types. We found a strong homogenization trend across the study area, with plant communities becoming more similar in composition over the decades when all taxa were treated together. The observed homogenization was driven especially by the increased similarity of vascular plant and lichen communities and was largely independent of biogeographic zones or continentality-humidity gradient. Homogenization was particularly associated with the drastic encroachment of the evergreen dwarf shrub Empetrum nigrum in habitat types originally dominated by other species, and with the decrease in lichen cover. In general, our findings suggest that Fennoscandian heathland and tundra vegetation is transforming towards a more homogeneous evergreen dwarf shrub-dominated system, which may threaten ecosystem multifunctionality. Our results highlight the importance of exploring biodiversity among different metrics and growth forms to understand the overall changes in heathland and tundra biodiversity.

The increasingly acknowledged and consequently also better understood microclimatic variability in terrestrial ecosystems has motivated a call for finer spatial resolution in species distribution modelling, especially in the case of sedentary low‐stature organisms such as plants. In contrast, less attention has so far been paid to the way climate should be represented in these models. In fact, most modelling applications rely only on a handful of so‐called bioclimatic variables (i.e. essential climatic variables designed for ecological applications), which is at odds with the hypothesised variation in the sensitivity of individual plant species to different facets of climate. We argue that the recent shift towards microclimate modelling provides a window of opportunity for re‐evaluating the predominant reliance on the small set of bioclimatic variables. We used a unique dataset of 895 1‐m 2 plots with vascular plant species observations and in situ soil temperature measurements across a high‐mountain landscape spanning 1700 elevational metres. From the hourly temperature measurements, we calculated bioclimatic variables as well as 188 additional ‘grid variables' arising from an aggregation of various summary statistics over different parts of the year. We then used those ‘grid variables', their subsets, and the bioclimatic variables to fit species distribution models for 101 plant species with combinations of one, two or three predictors. We found that bioclimatic variables consistently delivered less accurate models than many of the grid variables. Models based on subsets of grid variables only slightly decreased in accuracy and remained superior over bioclimatic variables even after the set was narrowed from the initial 188 to only six variables on the basis of a cluster analysis. These results highlight that modelling species distributions with only a few climatic variables is a viable strategy. However, the most suitable variables may often be different from those that are commonly used nowadays.

AimMicrorefugia on alpine slopes may allow species to persist in a warming climate. How plant species richness could respond to of the effectiveness of climatic buffering, defined as the difference in climate change in a microrefugium in comparison to that of an open alpine slope as mediated by competitive priority effects, is explored.LocationAlpine habitat anywhere.TaxonVascular plants.MethodsA simple spatially explicit model of plant species with adaptations across a climatic gradient simulates reproduction, dispersal, and mortality through climatic change. The effectiveness of microrefugia in buffering climate change and levels of competitive priority effects are included as factors that alter demographic rates in an experimental framework. Spatial patterns of microrefugia and differences in dispersal are also simulated. Differences in mean species richness were analysed.ResultsThe number of species conserved, relative to an instantaneous equilibrium calculation with the same climatic change, decreases with inclusion of a period of change (transient vs. instantaneous, equilibrium change) and further with competitive priority effects. In these simulations, the number of species conserved does not simply increase with buffer effectiveness, as hypothesized, but instead is bimodal.Main ConclusionsThe dip in number of species conserved in the middle range of buffer effectiveness occurs because barriers develop. These are temporal ecological traps, wherein species adapted to the middle of a climatic gradient become extinct, because their colonization of microrefugia is blocked by an extinction debt of resident species. The inertia that allows these demographic consequences is increased by competitive priorities. The highest levels of buffering and resulting inertia will not be ubiquitous but the bimodal pattern indicates that assessments of the role of microrefugia need to recognize the temporal species-environment and species-species interactions that will change the number of species conserved versus extinctions.

Complex topography regulates near-surface temperature above the treeline. It may thus sustain microrefugia for alpine plants and relax the need of shifting upward when the climate warms. The effectiveness of these microrefugia rests on the premise that plant distributions in alpine landscapes are mainly controlled by fine-scale topographic variation. We tested this assumption by relating the distribution of 79 plant species and 10 community attributes across 900 1 m² plots in a landscape spanning 1677 m of elevation to 17 topographical descriptors at resolutions between 1 and 301 m. We found that the presence of most species and most community attributes were better explained by topographic variation at coarser scales (> 20 m). Fine-scale topography is more clearly reflected in moisture than in temperature requirements of species. The elevational gradient rather than topographic variation at any scale, is the single most important driver of both species distributions and the variation in community attributes in the area studied. We hypothesise that our results reveal a hitherto underestimated influence of spatial mass effects on alpine plant distributions. These effects can override environmental filtering at fine scales and will thus impede the survival of cold-adapted plants in small and fragmented refugia under climate warming.

Difference in vegetation composition between communities from calcareous and siliceous soils might be due to high competition on siliceous soils for species from calcareous origin, and high rock-induced drought stress on calcareous soils for species from siliceous origin. We tested the hypothesis that, with increasing climate stress, competition should decrease on siliceous rock for species of calcareous origin and drought stress increase on calcareous rock for species of siliceous origin because of decreasing community biomass with increasing cold or drought stress. This question is of high interest for predicting changes in species distribution with climate change since bedrock type and climate change are both complex factors that are likely to interact with climate. We set up a transplant removal experiment in contrasting climate conditions of the south of France, warm and wet temperate, warm and dry Mediterranean, and cold temperate in the Alps and the Pyrenees. In each climate condition, three targets (two species and one population from a third species) from each origin were transplanted with and without neighbours on the two rock types and during two years with different levels of drought stress. Variation in the effects of neighbours and rock-induced drought stress on transplant survival with and without neighbours were analysed with linear modelling at each site, separately, and with ANCOVA in the whole design. Competition was the highest on siliceous rock in the more favourable climate conditions but for species of siliceous origin. The lower competition found for species of calcareous origin was likely due to the occurrence of allelopathic effects that decreased the negative effect of neighbours as measured with the removal method. The rock-induced drought stress was the highest on calcareous rock, in particular for species of siliceous origin. Thus, the decrease in competition intensity on siliceous rocks with increasing climate stress was more important for species from siliceous than calcareous origin. Additionally, the importance of the rock-induced drought stress was relatively low in the most stressful climate conditions as compared to the overwhelming importance of climate drought stress over the two years of our experiment, which likely induced a collapse of the positive effect of growing on a siliceous rock.

Complex topography regulates near‐surface temperature above the treeline. It may thus sustain microrefugia for alpine plants and relax the need of shifting upward when the climate warms. The effectiveness of these microrefugia rests on the premise that plant distributions in alpine landscapes are mainly controlled by fine‐scale topographic variation. We tested this assumption by relating the distribution of 79 plant species and 10 community attributes across 900 1 m² plots in a landscape spanning 1677 m of elevation to 17 topographical descriptors at resolutions between 1 and 301 m. We found that the presence of most species and most community attributes were better explained by topographic variation at coarser scales (> 20 m). Fine‐scale topography is more clearly reflected in moisture than in temperature requirements of species. The elevational gradient rather than topographic variation at any scale, is the single most important driver of both species distributions and the variation in community attributes in the area studied. We hypothesise that our results reveal a hitherto underestimated influence of spatial mass effects on alpine plant distributions. These effects can override environmental filtering at fine scales and will thus impede the survival of cold‐adapted plants in small and fragmented refugia under climate warming.

Boreal and tundra plant communities are expected to change in biodiversity due to increasing global change pressures such as climate warming. One long-term scenario is increasing compositional similarity, i.e., biotic homogenization, which has been relatively little studied in high-latitude plant communities. Here, we study how the composition and diversity of heathland and tundra plant communities have changed in northern Fennoscandia over several decades. In 2013–2023, we resurveyed 275 historic vegetation plots, originally surveyed in 1964–1975, with percentage covers for vascular plant, bryophyte and lichen species. We analyzed temporal changes in community composition and diversity across the study area and in different habitat types, biogeographic zones, and along the continentality-humidity gradient. We found a strong trend across the study area, with plant communities becoming more similar in composition over the decades. The observed homogenization was associated with compositional changes in vascular plant and lichen communities, and in particular with the encroachment of the evergreen dwarf shrub Empetrum nigrum. In comparison to vascular plants and lichens, the diversity of bryophytes generally remained more stable over time. Our findings suggest that Fennoscandian heathland and tundra vegetation is transforming towards a more homogenous evergreen dwarf shrub dominated system, which may threaten ecosystem multifunctionality. Our results highlight the importance of exploring biodiversity among different metrics and growth forms to understand the overall changes in heathland and tundra biodiversity. ### Competing Interest Statement The authors have declared no competing interest.

Besides environmental sorting, other processes like biotic interactions and dispersal limitation are vital for the assembly of plant communities in high mountains and their re-assembly under changing climatic conditions. Nevertheless, studies that compare the impact of these factors on plant community assembly above the tree line are largely lacking so far. We analysed occurrence changes in vascular plant communities of 492 permanent 1-m(2) plots in the alpine-nival ecotone of Mt. Schrankogel, Austrian Alps by comparing resurvey data from 2014 with data from the initial survey in 1994. We combined these data with species inventories from 899 additional plots sampled in 2021 and 2022 across a larger landscape above the tree line covering an elevational range of 1700 m, which we used for fine-scale habitat suitability modelling. We assessed the relative effects of projected habitat suitability, propagule pressure from surrounding populations and biomass density of neighbours on 1532 colonization and 372 extirpation events of 31 species observed on the permanent plots. We found that all three factors are significantly related to both colonisations and extirpations, with habitat suitability having the strongest, propagule pressure a slightly weaker, and vegetation density the weakest effect. Colonisations can be better explained by the three process proxies than extirpations. Our results indicate a crucial role of dispersal limitation besides the predominant effect of environmental filtering on the (re-)assembly of the alpine-nival plant community, while competitive/facilitative effects between plants tend to play a minor role. The strong imprint of nearby source plant populations on colonization/extirpation events suggests that recent plant migrations predominantly occur in small steps. This implies that while the topographically complex alpine terrain offers climatic microrefugia for plants, it may also pose potential barriers, hindering species from following their suitable climatic niches upwards.

Brief introduction: What are microclimates and why are they important?Microclimate science has developed into a global discipline. Microclimate science is increasingly used to understand and mitigate climate and biodiversity shifts. Here, we provide an overview of the current status of microclimate ecology and biogeography in terrestrial ecosystems, and where this field is heading next. Microclimate investigations in ecology and biogeography: We highlight the latest research on interactions between microclimates and organisms, including how microclimates influence individuals, and through them populations, communities and entire ecosystems and their processes. We also briefly discuss recent research on how organisms shape microclimates from the tropics to the poles. Microclimate applications in ecosystem management: Microclimates are also important in ecosystem management under climate change. We showcase new research in microclimate management with examples from biodiversity conservation, forestry and urban ecology. We discuss the importance of microrefugia in conservation and how to promote microclimate heterogeneity. Methods for microclimate science: We showcase the recent advances in data acquisition, such as novel field sensors and remote sensing methods. We discuss microclimate modelling, mapping and data processing, including accessibility of modelling tools, advantages of mechanistic and statistical modelling and solutions for computational challenges that have pushed the state-of-the-art of the field. What's next?We identify major knowledge gaps that need to be filled for further advancing microclimate investigations, applications and methods. These gaps include spatiotemporal scaling of microclimate data, mismatches between macroclimate and microclimate in predicting responses of organisms to climate change, and the need for more evidence on the outcomes of microclimate management.

Research in global change ecology relies heavily on global climatic grids derived from estimates of air temperature in open areas at around 2 m above the ground. These climatic grids do not reflect conditions below vegetation canopies and near the ground surface, where critical ecosystem functions occur and most terrestrial species reside. Here, we provide global maps of soil temperature and bioclimatic variables at a 1‐km 2 resolution for 0–5 and 5–15 cm soil depth. These maps were created by calculating the difference (i.e. offset) between in situ soil temperature measurements, based on time series from over 1200 1‐km 2 pixels (summarized from 8519 unique temperature sensors) across all the world's major terrestrial biomes, and coarse‐grained air temperature estimates from ERA5‐Land (an atmospheric reanalysis by the European Centre for Medium‐Range Weather Forecasts). We show that mean annual soil temperature differs markedly from the corresponding gridded air temperature, by up to 10°C (mean = 3.0 ± 2.1°C), with substantial variation across biomes and seasons. Over the year, soils in cold and/or dry biomes are substantially warmer (+3.6 ± 2.3°C) than gridded air temperature, whereas soils in warm and humid environments are on average slightly cooler (−0.7 ± 2.3°C). The observed substantial and biome‐specific offsets emphasize that the projected impacts of climate and climate change on near‐surface biodiversity and ecosystem functioning are inaccurately assessed when air rather than soil temperature is used, especially in cold environments. The global soil‐related bioclimatic variables provided here are an important step forward for any application in ecology and related disciplines. Nevertheless, we highlight the need to fill remaining geographic gaps by collecting more in situ measurements of microclimate conditions to further enhance the spatiotemporal resolution of global soil temperature products for ecological applications.

The sustainable development of mountain regions requires inter- and transdisciplinary knowledge. The Institute for Interdisciplinary Mountain Research contributes to this global endeavor as part of the Austrian Academy of Sciences and as a member of international scientific networks, together with local partners and stakeholders. As a joint effort of individual researchers covering multiple fields, this article highlights our views on mountains as research objects, the phenomena we investigate as parts of entire mountain systems, and the synergies and differences of the disciplinary frames within which we work. Alles ist Wechselwirkung [Everything is interaction] (A. von Humboldt, 18XX, 27r)

Arctic soils are an important reservoir of soil organic carbon (SOC) and their role in determining arctic ecosystem functioning in global carbon budgets requires closer attention. We investigated the coupling of soil properties and SOC stabilization mechanisms in high Arctic terrestrial habitats differing in vegetation cover and organic matter input. We focused on soil physical and chemical properties in glacier foreland, soil crust, dry tundra, wet tundra, and bird cliff meadow habitats on Svalbard (Norway). Concurrently, we performed physical fractionation to determine the amount of SOC stabilized by mineral associations or occlusion in macro and microaggregates. Initial stages of soil development (glacier foreland and soil crust habitats) exhibited characteristically high bulk density and pH, and low moisture and nutrient contents, whereas more developed soils (dry and wet tundra habitats) showed opposite trends. Contrastingly, bird cliff meadow showed low bulk density, intermediate moisture, and very high nutrient content. The amount of SOC stabilized by mineral associations and occlusion in aggregates generally increased with vegetation cover; hence, the more developed habitats supported higher contents of stabilized SOC. However, SOC was stabilized in aggregates even in initial stages of soil development. SOC content in most fractions correlated positively with contents of dissolved organic carbon and nitrogen, suggesting that both dissolved organic carbon and nitrogen might have provided some degree of SOC stabilization through increased formation of aggregates and suppression of microbial mineralization of soil organic matter, respectively. Our findings underscore the notion that models of SOC sequestration in the Arctic should account not only for total SOC content, but also SOC stabilization mechanisms, as represented by SOC content in respective soil fractions.

QuestionIn recent decades, high-latitude climate has shown regionally variable trends towards warmer and moister conditions. These changes have been predicted to cause afforestation or shrubification of open tundra, increases of warmth-demanding southern species and plant groups favoured by increased moisture, and decline of species and habitats that are dependent on snow cover. In this study, we explore temporal changes in northern tundra upland plant communities along regional gradients and in local habitats. We ask how vegetation changes are linked with long-term trends in regional climate and grazing pressure. LocationNorthern Europe. MethodsIn 2013-2014, we resurveyed a total of 108 vegetation plots on wind-exposed and snow-protected tundra habitats in three subareas along a bioclimatic gradient from the northern boreal to the arctic zone. Vegetation plots were originally sampled in 1964-1967. We related observed vegetation changes to changes in temperature, precipitation and grazing pressure, which all showed regionally variable increases over the study period. ResultsWe found a significant increase of the evergreen dwarf shrub Empetrum nigrum subsp. hermaphroditum in snow-protected communities and a prominent decrease of lichens throughout the study area. No evidence for extensive tree or larger shrub (Betula spp., Salix spp. or Juniperus communis) encroachment despite climatic warming trends was found. Among studied communities, most pronounced changes in vegetation were observed in snow-protected boreal heaths on small isolated uplands, where community composition showed low resemblance to the original composition described decades ago. Changes in plant communities correlated with changes in summer and winter temperatures, summer precipitation and reindeer grazing pressure, yet correlations varied depending on region and habitat. ConclusionsNorthern tundra uplands vary in their resistance to on-going climate change and reindeer grazing. Isolated treeless heaths of boreal forest-tundra ecotone appear least resistant to climate change and have already shifted towards new community states.