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International audience; Seed dispersal and local filtering interactively govern community membership and scale up to shape regional vegetation patterns, but data revealing how and why particular species are excluded from specific communities in nature are scarce. This lack of data is a missing link between our theoretical understanding of how diversity patterns can form and how they actually form in nature, and it hampers our ability to predict community responses to climate change. Here, we compare seed, seedling, and adult plant communities at 12 grassland sites with different climates in southern Norway to examine how community membership is interactively shaped by seed dispersal and local filtering, and how this process varies with climate across sites. To do this, we divide species at each site into two groups: locally transient species, which occur as seeds but are rare or absent as adults (i.e., they arrive but are filtered out), and locally persistent species, which occur consistently as adults in annual vegetation surveys. We then ask how and why locally transient species are disfavored during community assembly. Our results led to four main conclusions: (1) the total numbers of seeds and species that arrived, but failed to establish locally persistent populations, rose with temperature, indicating an increase in the realized effects of local filtering on community assembly, as well as an increase in the number of species poised to rapidly colonize those warmer sites if local conditions change in their favor, (2) locally transient species were selectively filtered out during seedling emergence, but not during seedling establishment, (3) selective filtering was partly driven by species climate preferences, exemplified by the poor performance of seeds dispersing outside of their realized climate niches into colder and drier foreign climates, and (4) locally transient species had traits that likely made them better dispersers (i.e., smaller seeds) but poorer competitors for light (i.e., shorter statures and less persistent clonal connections) than locally persistent species, potentially explaining why these species arrived to new sites but did not establish locally persistent adult populations. Our study is the first to combine seed, seedling, and adult survey data across sites to rigorously characterize how seed dispersal and local filtering govern community membership and shape climate‐associated vegetation patterns

Across environmental gradients, community‐level functional traits of plants can change due to species turnover, intraspecific variation and their covariation. Studies on vascular plants suggest that species turnover is the main driver of trait variation across gradients, although intraspecific variation can also be important. However, there is limited knowledge about whether this holds for non‐vascular primary producers such as lichens and bryophytes. We hypothesized that intraspecific variation is more important for non‐vascular than for vascular primary producers because they lack specialized structures to maintain homeostasis and should therefore be more responsive to extrinsic factors. To assess the relative importance of species turnover versus intraspecific variation for vascular plants, lichens and bryophytes, we estimated species abundance and measured chemical (tissue nitrogen (N) and phosphorous (P) content, N:P ratio and pH) and non‐chemical (specific leaf or thallus area, dry matter content and water holding capacity) functional traits along an elevational gradient in alpine southern Norway. We calculated community‐weighted mean traits and quantified the relative contribution of species turnover, intraspecific variation and their covariation to total trait variation across the gradient. We found mixed support for our hypothesis: the contribution of intraspecific variation to total trait variation for N and N:P was higher in lichens than in vascular plants and bryophytes, but in general the contribution of intraspecific variation differed among functional traits and producer groups. Nutrient variables (N, P and N:P) were significantly impacted by intraspecific variation for vascular plants and lichens but not for bryophytes. Non‐chemical traits and pH were mainly driven by species turnover effects in all primary producer groups. Our results highlight that while nearly all studies on primary producer trait variation across environments have focused on vascular plants, trait variation of other largely neglected but ecologically important producer groups, such as lichens and bryophytes, may show very different responses to the same environmental factors. In order to fully understand how future environmental changes impact on community‐ and ecosystem‐level processes, traits of primary producers other than vascular plants—and their within‐species variation—need to be considered in systems where these groups are abundant. A free Plain Language Summary can be found within the Supporting Information of this article. Published version Norges Forskningsråd, Grant/Award Number: 249902/F20

Globally accelerating trends in societal development and human environmental impacts since the mid-twentieth century 1–7 are known as the Great Acceleration and have been discussed as a key indicator of the onset of the Anthropocene epoch 6 . While reports on ecological responses (for example, changes in species range or local extinctions) to the Great Acceleration are multiplying 8, 9 , it is unknown whether such biotic responses are undergoing a similar acceleration over time. This knowledge gap stems from the limited availability of time series data on biodiversity changes across large temporal and geographical extents. Here we use a dataset of repeated plant surveys from 302 mountain summits across Europe, spanning 145 years of observation, to assess the temporal trajectory of mountain biodiversity changes as a globally coherent imprint of the Anthropocene. We find a continent-wide acceleration in the rate of increase in plant species richness, with five times as much species enrichment between 2007 and 2016 as fifty years ago, between 1957 and 1966. This acceleration is strikingly synchronized with accelerated global warming and is not linked to alternative global change drivers. The accelerating increases in species richness on mountain summits across this broad spatial extent demonstrate that acceleration in climate-induced biotic change is occurring even in remote places on Earth, with potentially far-ranging consequences not only for biodiversity, but also for ecosystem functioning and services.

Background The stress-gradient hypothesis predicts a shift from facilitative to competitive plant interactions with decreasing abiotic stress. This has been supported by studies along elevation and temperature gradients, but also challenged by the hypothesis of a facilitation collapse at extremely harsh sites. Although facilitation is known to be important in primary succession, few studies have examined these hypotheses along primary succession gradients. Aim To examine whether there is a relationship between the presence of the circumpolar cushion plant Silene acaulis and other species, and if so, whether there is a shift between positive and negative interactions along a primary succession gradient in a glacier foreland. Location Finse, southern Norway. Methods We examined the performance of the common alpine forb Bistorta vivipara, species richness of vascular plants, bryophytes and lichens, and the number of seedlings and fertile vascular plants in S. acaulis cushions, and control plots without S. acaulis, along a succession gradient with increasing distance from a glacier front, and thus decreasing abiotic stress. To examine if S. acaulis cushions modify the abiotic environment, we recorded soil temperature, moisture, organic content and pH in cushions and control plots. Results B. vivipara performed better, as shown by bigger leaves in S. acaulis cushions compared to control plots in the harshest part of the gradient close to the glacier. There were few differences in B. vivipara performance between cushion and control plots in the more benign environment further away from the glacier. This suggests a shift from facilitative to mainly neutral interactions by S. acaulis on the performance of B. vivipara with decreasing abiotic stress. A trend, although not significant, of higher vascular species richness and fertility inside S. acaulis cushions along the whole gradient, suggests that S. acaulis also facilitates community-level species richness. The causal mechanism of this facilitation is likely that the cushions buffer extreme temperatures. Conclusions Our results support the stress-gradient hypothesis for the relationship between the cushion plant S. acaulis and the performance of a single species along a primary succession gradient in a glacier foreland. S. acaulis also tended to increase vascular plant species richness and fertility regardless of stress level along the gradient, suggesting facilitation at the community level. We found no collapse of facilitation at the most stressful end of the gradient in this alpine glacier foreland.

Questions Is there a shift from positive to negative biotic interaction effects on seedling recruitment along two different stress gradients, temperature and precipitation (the stress-gradient hypothesis); do such interaction effects differ between species with different bioclimatic affinities? Location Boreal, sub-alpine and alpine grassland in southern Norway. Methods We tested the stress-gradient hypothesis by comparing seedling recruitment in bare-ground gaps where vegetation has been removed vs in extant grassland vegetation in 12 boreal, sub-alpine and alpine grassland sites along a precipitation gradient. This was tested in (1) a seed-sowing experiment and (2) in naturally occurring recruitment of alpine, generalist and boreal species. Results Emergence of the sown alpine species was higher in the cold alpine than in the warmer sub-alpine sites, with no effects of precipitation or vegetation removal. The sown generalists also decreased in emergence towards warmer sites, whereas there was no effect of temperature on the sown boreal species. Vegetation removal, interacting with precipitation, increased the emergence of the generalist and boreal species sown at intermediate precipitation levels. In contrast, interactions between temperature and vegetation removal regulated the emergence of all groups of naturally occurring seedlings. Alpine and generalist species emerged at the highest rate in alpine sites, whereas boreal species had highest emergence in the lowlands. Conclusion For all species groups, strong effects of vegetation removal show that competition from the extant vegetation dominates in controlling seedling emergence across all study sites and species. In generalist and boreal species, positive interactions between vegetation removal and temperature show that competitive interactions affect seedling emergence more strongly towards warmer climates, in line with the stress-gradient hypothesis. In contrast, alpine species show no such interactions. This suggests that species’ adaptations to climate, in combination with environmental forcing, control seedling emergence along the bioclimatic gradients. Our results have implications for nature conservation, as we propose that disturbance from grazing animals can be useful to release competition and thereby increase seedling recruitment and biodiversity in boreal and alpine grasslands in a warmer future.

International audience; Understanding the degree to which plant communities are open to seedling recruitment is key to predicting how they will be impacted by climate change. We experimentally assessed whether communities assembled under colder climates were inherently more open to recruitment than warmer‐climate communities, after controlling for differences in the current climate under which the communities were growing. We then tested whether variation in openness to recruitment could be explained by community biomass or by the plant functional traits of the community.The study was conducted in a climate grid of twelve grassland sites across southern Norway, differing systematically in temperature and precipitation. Along a 2000 mm precipitation gradient, we transplanted turfs with intact plant communities from alpine and sub‐alpine sites into 2°C warmer sites, and measured natural seedling emergence in these transplanted turfs versus locally replanted control turfs at the transplant destination sites. Mixed effect models were used to assess the effect of origin (cold versus warm climate), biomass, and functional trait composition of the communities on seedling emergence. We further assessed variation in these effects across different climatic contexts (the temperature and precipitation gradients).Communities originating from colder climates were consistently more open to recruitment, with on average 44% more seedlings emerging, than the locally replanted control communities. The higher rates of seedling emergence in colder‐climate communities were attributable to systematic differences in plant functional traits, but not in biomass. The colder‐climate communities were composed of species with smaller leaves and lower maximum plant heights; traits that may make these communities less effective at excluding new recruits. These trait‐related responses were not significant in the warmest sites and did not very across the precipitation gradient.Our results suggest that alpine species lack the competitive effect traits required to make their communities resistant to invasion by novel competitors under climate change.

Fil: Komatsu, Kimberly J. Smithsonian Environmental Research Center, Edgewater. United States. Fil: Avolio, Meghan L. Johns Hopkins University. Department of Earth and Planetary Sciences. Baltimore, United States. Fil: Lemoine, Nathan P. Marquette University. Department of Biological Sciences. Milwaukee, United States. Fil: Chaneton, Enrique José. Universidad de Buenos Aires. Facultad de Agronomía. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura (IFEVA). Buenos Aires, Argentina. Fil: Chaneton, Enrique José. CONICET – Universidad de Buenos Aires. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura (IFEVA). Buenos Aires, Argentina. Fil: Tognetti, Pedro Maximiliano.Universidad de Buenos Aires. Facultad de Agronomía. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura (IFEVA). Buenos Aires, Argentina. Fil: Tognetti, Pedro Maximiliano.CONICET – Universidad de Buenos Aires. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura (IFEVA). Buenos Aires, Argentina. Fil: Yahdjian, María Laura. Universidad de Buenos Aires. Facultad de Agronomía. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura (IFEVA). Buenos Aires, Argentina. Fil: Yahdjian, María Laura. CONICET – Universidad de Buenos Aires. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura (IFEVA). Buenos Aires, Argentina. Fil: Isbell, Forest. University of Minnesota. Department of Ecology, Evolution and Behavior. Saint Paul, United States. Fil: Grman, Emily. Eastern Michigan University. Department of Biology. Ypsilanti, United States. Global change drivers (GCDs) are expected to alter community structure and consequently, the services that ecosystems provide. Yet, few experimental investigations have examined effects of CDs on plant community structure across multiple ecosystem types, and those that do exist present conflicting patterns. In an unprecedented global synthesis of over 100 experiments that manipulated factors linked to GCDs, we show that herbaceous plant community responses depend on experimental manipulation length and number of factors manipulated. We found that plant communities are fairly resistant to experimentally manipulated GCDs in the short term ( minor to 10 y). In contrast, long-term (major or equal to 10 y) experiments show increasing community divergence of treatments from control conditions. Surprisingly, these community responses occurred with similar frequency across the GCD types manipulated in our database. However, community responses were more common when 3 or more GCDs were simultaneously manipulated, suggesting the emergence of additive or synergistic effects of multiple drivers, particularly over long time periods. In half of the cases, GCD manipulations caused a difference in community composition without a corresponding species richness difference, indicating that species reordering or replacement is an important mechanism of community responses to GCDs and should be given greater consideration when examining consequences of GCDs for the biodiversity–ecosystem function relationship. Human activities are currently driving unparalleled global changes worldwide. Our analyses provide the most comprehensive evidence to date that these human activities may have widespread impacts on plant community composition globally, which will increase in frequency over time and be greater in areas where communities face multiple GCDs simultaneously. tbls., grafs.

In the version of this Article originally published, the following sentence was missing from the Acknowledgements: “This work was supported by the Norwegian Research Council SnoEco project, grant number 230970”. This text has now been added.

Dispersal dynamics and local filtering interactively generate regional vegetation patterns, but empirical evidence of their combined influence in nature is scarce, representing a missing link between our theoretical understanding of community assembly and real-world observation. Here, we compare seed and adult plant communities at twelve grassland sites with different climates in southern Norway to explore the degree to which community membership is shaped by dispersal limitation and local niche-based filtering, and how this varies with climate. To do this, we first divide species at each site into two groups: “locally-transient” species, which occur as seeds but are rare or absent as adults (i.e., they arrive but are filtered out), or “locally-persistent” species, which occur consistently as adults in annual vegetation surveys. We then ask questions to reveal where, when, why, and how locally-transient species are systematically disfavored during community assembly. Our results led to four main conclusions: (1) the strength of local filtering on community membership increased with temperature, (2) surprisingly, local filtering was stronger during seedling emergence than during seedling establishment, (3) climate-based niche differences drove differential performance among species, especially for seeds dispersing outside of their realized climate niches into more stressful (colder and drier) climates, and (4) locally-transient species had traits that may made them better dispersers (smaller seeds) but poorer competitors for light (shorter statures, less persistent clonal connections) than locally-persistent species, providing a potential explanation for why they arrived to new sites but failed to establish persistent adult populations. Our study is one of the first to combine seed, seedling, and adult survey data across multiple sites with different climates to provide a rigorous empirical evaluation of the combined influence of dispersal limitation and local filtering on the generation and maintenance of climate-associated vegetation patterns.

The Arctic is warming twice as fast as the rest of the planet, leading to rapid changes in species composition and plant functional trait variation. Landscape-level maps of vegetation composition and trait distributions are required to expand spatially-limited plot studies, overcome sampling biases associated with the most accessible research areas, and create baselines from which to monitor environmental change. Unmanned aerial vehicles (UAVs) have emerged as a low-cost method to generate high-resolution imagery and bridge the gap between fine-scale field studies and lower resolution satellite analyses. Here we used field spectroscopy data (400–2500 nm) and UAV multispectral imagery to test spectral methods of species identification and plant water and chemistry retrieval near Longyearbyen, Svalbard. Using the field spectroscopy data and Random Forest analysis, we were able to distinguish eight common High Arctic plant tundra species with 74% accuracy. Using partial least squares regression (PLSR), we were able to predict corresponding water, nitrogen, phosphorus and C:N values (r 2 = 0.61–0.88, RMSEmean = 12%–64%). We developed analogous models using UAV imagery (five bands: Blue, Green, Red, Red Edge and Near-Infrared) and scaled up the results across a 450 m long nutrient gradient located underneath a seabird colony. At the UAV level, we were able to map three plant functional groups (mosses, graminoids and dwarf shrubs) at 72% accuracy and generate maps of plant chemistry. Our maps show a clear marine-derived fertility gradient, mediated by geomorphology. We used the UAV results to explore two methods of upscaling plant water content to the wider landscape using Sentinel-2A imagery. Our results are pertinent for high resolution, low-cost mapping of the Arctic.

Questions Vegetation mapping based on field surveys is time-consuming and expensive. Distribution modelling might be used to overcome these challenges. What is the performance of distribution modelling of vegetation compared to traditional vegetation mapping when projected locally? Does the modelling performance vary among ecosystems? Does vegetation type distribution and abundance influence the modelling performance? Location Gravfjellet, Oystre Slidre commune, southern Norway. Methods Two comparable neighbouring areas, each of 4 km2, were mapped for species-defined vegetation types. One area was used for model training, the other for model projection. Maximum entropy models were run for six vegetation types, two from each of the ecosystems present in the area: forest, wetland and mountain heath- and shrublands. For each ecosystem, one locally abundant and one locally rare vegetation type were tested. AUC, the area under the receiver operating curve, was used as the model selection criterion. Environmental variables (n = 9) were selected through a backwards selection scheme, and model complexity was kept low. The models were evaluated using independent data. Results Distribution modelling of vegetation types by local projection gave high AUC values, and the results were supported by the evaluation using independent data. The modelling ability was not affected by ecosystem differences. A negative relationship between the number of points used to train the models and the AUC value before evaluation suggests that models for locally rare vegetation types had better predictive performance than the models for abundant types. This result was not significant after evaluation. Conclusion Provided that relevant explanatory variables are available at an appropriate scale, and that field-validated training points are available, distribution modelling can be used for local projection of the six tested vegetation types from the boreal–alpine ecotone.

Biotic interactions are often ignored in assessments of climate change impacts. However, climate-related changes in species interactions, often mediated through increased dominance of certain species or functional groups, may have important implications for how species respond to climate warming and altered precipitation patterns. We examined how a dominant plant functional group affected the population dynamics of four co-occurring forb species by experimentally removing graminoids in seminatural grasslands. Specifically, we explored how the interaction between dominants and subordinates varied with climate by replicating the removal experiment across a climate grid consisting of 12 field sites spanning broad-scale temperature and precipitation gradients in southern Norway. Biotic interactions affected population growth rates of all study species, and the net outcome of interactions between dominants and subordinates switched from facilitation to competition with increasing temperature along the temperature gradient. The impacts of competitive interactions on subordinates in the warmer sites could primarily be attributed to reduced plant survival. Whereas the response to dominant removal varied with temperature, there was no overall effect of precipitation on the balance between competition and facilitation. Our findings suggest that global warming may increase the relative importance of competitive interactions in seminatural grasslands across a wide range of precipitation levels, thereby favouring highly competitive dominant species over subordinate species. As a result, seminatural grasslands may become increasingly dependent on disturbance (i.e. traditional management such as grazing and mowing) to maintain viable populations of subordinate species and thereby biodiversity under future climates. Our study highlights the importance of population-level studies replicated under different climatic conditions for understanding the underlying mechanisms of climate change impacts on plants.

Forest certification management standards aim at maintaining forest ecosystem integrity, including forest biodiversity conservation. However, studies from the Amazon and Congo basin find that forest certification may not protect forest biodiversity and ecosystems, and may therefore be unsustainable. This study evaluates the influence of forest certification on conserving biodiversity. Specifically, we (a) estimate tree (adult and seedling) species richness, diversity and density among different forest management regimes; (b) assess the relationship between environmental and human forest use variables, and species richness, diversity and density among the forest management regimes; and (c) assess the influence of forest governance of villages adjacent to the forests on tree (adult and seedling) species richness, diversity and density among the forest management regimes. This is achieved in a comparative study of Forest Stewardship Council certified community forests, non-certified open access forests, and non-certified state forest reserves in the Kilwa District in Tanzania. Our results show that forest certification standards and implementation processes are positively related to biodiversity conservation. There are significantly higher tree (adults) species richness, diversity, and density in certified community forests than in open access forests and state forest reserves. These findings suggest that forest certification may be a good policy option to conserve biodiversity. The present study is one of the first studies in tropical Africa, which contributes to the limited data on the influence of forest certification on conserving biodiversity. Our results may also serve as baseline for further research on the contribution of certified forests in conserving biodiversity at both temporal and spatial scales.

We studied the effects of different mat-forming lichens on microclimate and litter decomposition in an alpine ecosystem where lichens constitute a major part of the vegetation. We used four lichens with contrasting colour and water-holding capacity. We recorded soil temperature and moisture, and decomposition rate of plant litter under the different lichen mats and in bare soil. While soil temperature and freeze-thaw cycles were reduced under all lichen mats compared to bare soil, Cladonia rangiferina/Cladonia stygia insulated stronger than other lichens. Litter decomposition was faster under Flavocetraria nivalis than under Alectoria ochroleuca, but this was not related to lichen traits or microclimate. We conclude that insulation by lichen mats is stronger with higher water-holding capacity, and we suggest that microbial communities associated with different lichens have a larger effect on litter decomposition than soil conditions. Our findings highlight the importance of mat-forming lichens on microclimate and ecosystem processes in alpine environments.

Olea europaea L. subsp cuspidata (Wall. ex G. Don) Cif., an endangered tree species in dry Afromontane forests, has multiple uses in local communities in Ethiopia, making it susceptible to overexploitation. The study investigated the rates and causes of O. europaea harvesting in the Hugumburda National Forest Priority Area (NFPA). We measured the diameter at stump height of harvested stumps from 70 (20 &times, 20 m) plots and estimated the time since cutting to determine the biomass of O. europaea wood harvested annually in the forest. We performed a socioeconomic survey of the reasons for wood harvesting by conducting 163 stratified random individual interviews in the villages surrounding the forest. The average annual quantity of O. europaea wood illegally harvested from the forest was estimated to be 430 kg ha&minus, 1, mainly for farm implements, fuel wood, and fumigation purposes. The results of a General Linear Model (GLM) show that the extraction of O. europaea wood in the forest is higher at higher elevations than at lower, and the number of O. europaea stumps in the forest is higher at an intermediate distance to the villages. We show that O. europaea is harvested in the forest despite the fact that the forest is protected. Permanent sample plots should be established to monitor the increment.

Motivation The BioTIME database contains raw data on species identities and abundances in ecological assemblages through time. These data enable users to calculate temporal trends in biodiversity within and amongst assemblages using a broad range of metrics. BioTIME is being developed as a community-led open-source database of biodiversity time series. Our goal is to accelerate and facilitate quantitative analysis of temporal patterns of biodiversity in the Anthropocene. Main types of variables included The database contains 8,777,413 species abundance records, from assemblages consistently sampled for a minimum of 2 years, which need not necessarily be consecutive. In addition, the database contains metadata relating to sampling methodology and contextual information about each record. Spatial location and grain BioTIME is a global database of 547,161 unique sampling locations spanning the marine, freshwater and terrestrial realms. Grain size varies across datasets from 0.0000000158 km2 (158 cm2) to 100 km2 (1,000,000,000,000 cm2). Time period and grain BioTIME records span from 1874 to 2016. The minimal temporal grain across all datasets in BioTIME is a year. Major taxa and level of measurement BioTIME includes data from 44,440 species across the plant and animal kingdoms, ranging from plants, plankton and terrestrial invertebrates to small and large vertebrates. Software format .csv and .SQL., Global Ecology and Biogeography, 27 (7), ISSN:1466-822X, ISSN:1466-8238

Aim Previous research on how climatic niches vary across species ranges has focused on a limited number of species, mostly invasive, and has not, to date, been very conclusive. Here we assess the d ...

Seedling recruitment is a critical life history stage for trees, and successful recruitment is tightly linked to both abiotic factors and biotic interactions. In order to better understand how tree species’ distributions may change in response to anticipated climate change, more knowledge of the effects of complex climate and biotic interactions is needed. We conducted a seed-sowing experiment to investigate how temperature, precipitation and biotic interactions impact recruitment of Scots pine (Pinus sylvestris) and Norway spruce (Picea abies) seedlings in southern Norway. Seeds were sown into intact vegetation and experimentally created gaps. To study the combined effects of temperature and precipitation, the experiment was replicated across 12 sites, spanning a natural climate gradient from boreal to alpine and from sub-continental to oceanic. Seedling emergence and survival were assessed 12 and 16 months after sowing, respectively, and aboveground biomass and height were determined at the end of the experiment. Interestingly, very few seedlings were detected in the boreal sites, and the highest number of seedlings emerged and established in the alpine sites, indicating that low temperature did not limit seedling recruitment. Site precipitation had an overall positive effect on seedling recruitment, especially at intermediate precipitation levels. Seedling emergence, establishment and biomass were higher in gap plots compared to intact vegetation at all temperature levels. These results suggest that biotic interactions in the form of competition may be more important than temperature as a limiting factor for tree seedling recruitment in the sub- and low-alpine zone of southern Norway. acceptedVersion

Accepted manuscript version. Published version available at https://doi.org/10.1016/j.ppees.2017.09.008. Accepted manuscript version, licensed CC BY-NC-ND 4.0. In the face of climate change, populations have two survival options − they can remain in situ and tolerate the new climatic conditions (“stay”), or they can move to track their climatic niches (“go”). For sessile and small-stature organisms like alpine plants, staying requires broad climatic tolerances, realized niche shifts due to changing biotic interactions, acclimation through plasticity, or rapid genetic adaptation. Going, in contrast, requires good dispersal and colonization capacities. Neither the magnitude of climate change experienced locally nor the capacities required for staying/going in response to climate change are constant across landscapes, and both aspects may be strongly affected by local microclimatic variation associated with topographic complexity. We combine ideas from population and community ecology to discuss the effects of topographic complexity in the landscape on the immediate “stay” or “go” opportunities of local populations and communities, and on the selective pressures that may have shaped the stay or go capacities of the species occupying contrasting landscapes. We demonstrate, using example landscapes of different topographical complexity, how species’ thermal niches could be distributed across these landscapes, and how these, in turn, may affect many population and community ecological processes that are related to adaptation or dispersal. Focusing on treeless alpine or Arctic landscapes, where temperature is expected to be a strong determinant, our theorethical framework leads to the hypothesis that populations and communities of topographically complex (rough and patchy) landscapes should be both more resistant and more resilient to climate change than those of topographically simple (flat and homogeneous) landscapes. Our theorethical framework further points to how meta-community dynamics such as mass effects in topographically complex landscapes and extinction lags in simple landscapes, may mask and delay the long-term outcomes of these landscape differences under rapidly changing climates.

Summary Global change, such as climate warming and nitrogen deposition, has been predicted to induce non-reversible regime shifts in natural ecosystems. However, we lack knowledge of the potential for recovery from global change perturbations and factors influencing the recovery rate. We examined the recovery of an alpine plant community from a combined warming and nutrient addition experiment, which initially caused profound changes in plant community composition and diversity. We also examined whether the recovery process was affected by herbivory, as the presence of herbivores has been shown to modify the response of plant communities to global change. Many aspects of the alpine plant community had not recovered from experimental warming and nutrient addition 6 years after cessation of the treatments. A persistent shift in vascular plant species composition seemed to inhibit recovery and maintain a community dominated by highly competitive grasses at the expense of the previously dominating dwarf shrub Dryas octopetala, lichens and bryophytes. The exclusion of herbivores decreased the community recovery rate by maintaining unfavourable conditions for lichens and bryophytes. Synthesis. Our findings suggest that a shift in dominance hierarchies in arctic and alpine plant communities due to global change is not readily reversible. Herbivory may, however, facilitate plant community recovery.

Aim In recent decades species ranges have shifted upwards in elevation and northwards in latitude. These shifts are commonly interpreted as a response to recent climate warming. However, several alternative hypotheses have been proposed to explain the elevational shifts, including increased deposition of atmospheric nitrogen, changes in precipitation and dispersal limitation. We evaluate these hypotheses and attempt to identify the dominant drivers for the observed shifts in the upper range limits of alpine plant species. Location European mountains from Svalbard to the southern Alps. Methods We assembled data on observed shifts in the upper range limit of alpine plants over 40 to 100 years on 114 mountains. We related the observed shifts to recent changes in temperature and precipitation and to recent deposition of atmospheric nitrogen. Changes in traits and habitat preferences of species in the summit assemblages were used to evaluate the potential role of different drivers. Results Seventy per cent of the species that showed a detectable change in their upper range limits between surveys shifted their range limits upwards. The same species tend to move up on different mountains. There are, however, large differences between mountains in the proportion of species shifting upwards. This proportion is not found to be statistically related to local changes in temperature. Correspondingly, warmth-demanding species did not move upward more frequently than expected by chance. Snow-bed species have become more common on summits. Main conclusions Our data do not support the idea that climate warming is the dominant factor causing the observed range shifts of alpine plant species on European mountains: first, the amount of change in species assemblages on the summits studied is not related statistically to the amount of climate warming; second, those species that have moved upwards are not particularly warmth demanding. publishedVersion

Whether invasion of introduced plant species may be aided by certain community properties is poorly understood for species-rich ecosystems, such as tropical montane forests. In Kenya, the non-native tree Fraxinus pennsylvanica has invaded degraded montane forests. We used generalized linear mixed models to examine the relative importance of different community properties to Fraxinus invasion after agricultural abandonment and in the secondary forest. Fraxinus invasion was positively related to plant community species diversity and the abundance of tree saplings, shrubs, ferns, and herbs in the abandoned fallows, but negatively related to the same community properties in the secondary forest. The number of Fraxinus recruits declined with declining propagule pressure in the fallows, but not in the secondary forest. Although adult and saplings of Fraxinus were positively related to community diversity in the fallows, Fraxinus appeared to decrease diversity in the secondary forest. These results show that the success of non-native species invasion and the effects of an invader on the resident community may depend both on properties and degree of disturbance of the community. Plant community diversity and evenness appeared to determine the invasion success by increasing invasibility of the abandoned fallows, but decreasing invasibility of the secondary forest. Our results from a tropical degraded forest emphasize the importance of including habitat characteristics when predicting both the potential of non-native plant invasion and effects of invasives on the particular community.

Background: Alpine and arctic ecosystems at high latitudes have been shown to be particularly vulnerable to climate change, but little has been reported about plant community responses from lower latitudes, such as the vast Qinghai–Tibetan Plateau.Aim: To examine how alpine fen plant community species richness, diversity, cover, above- and below-ground biomass, and how the abundance of two sedges, the dominant Kobresia tibetica and the subordinate Carex moorcroftii, respond to experimental warming.Methods: We used open-top chambers of two heights, with identical open-top area, to simulate two levels of experimental warming at 4700 m on the Qinghai–Tibetan Plateau.Results: Five years of experimental warming reduced species richness and diversity, primarily due to a decrease in the diversity of graminoids. Above-ground biomass, vegetation height and cover increased at both levels of warming. Below-ground biomass increased at a depth of 5–20 cm, but not at 0–5 cm, indicating an allocation of resource to plan...

Th e determinants of local species richness in plant communities have been the subject of much debate. Is species richness the result of stochastic events such as dispersal processes, or do local environmental fi lters sort species into communities according to their ecological niches? Recent studies suggest that these two processes simultaneously limit species richness, although their relative importance may vary in space and time. Understanding the limiting factors for species richness is especially important in light of the ongoing global warming, as new species establish in resident plant communities as a result of climate-driven migration. We examined the relative importance of dispersal and environmental fi ltering during seedling recruitment and plant establishment in an alpine plant community subjected to seed addition and long-term experimental warming. Seed addition increased species richness during the seedling recruitment stage, but this initial increase was cancelled out by a corresponding decrease in species richness during plant establishment, suggesting that environmental fi lters limit local species richness in the long term. While initial recruitment success of the sown species was related to both abiotic and biotic factors, long-term establishment was controlled mainly by biotic factors, indicating an increase in the relative importance of biotic interactions once plants have germinated in a microhabitat with favourable abiotic conditions. Th e relative importance of biotic interactions also seemed to increase with experimental warming, suggesting that increased competition within the resident vegetation may decrease community invasibility as the climate warms.

Background: The effect of the anticipated climate change on the stability of vegetation and the factors underlying this stability are not well understood. Aims: Our objective was to quantify long-term vegetation changes in a range of habitats in northern Europe by exploring species co-occurrences and their links to diversity and productivity gradients. Methods: We re-sampled vegetation in 16 arctic, mountain and mire sites 20 to 90 years after the original inventories. A site-specific change in species assemblages (stability) was quantified using species co-occurrences. Using a randomisation test we tested whether the changes observed were significantly greater than those expected by chance. Relationships between patterns in vegetation stability and time between surveys, numbers of plots, or species diversity and proxies for productivity, were tested using regression analysis. Results: At most sites the changes in species co-occurrences of vascular plants and bryophytes were greater than those expected by...

Plant functional traits vary consistently along climate gradients and are therefore potential predictors of plant community response to climate change. We test this space-for-time assumption by combining a spatial gradient study with whole-community turf transplantation along temperature and precipitation gradients in a network of 12 grassland sites in Southern Norway. Using data on eight traits for 169 species and annual vegetation censuses of 235 turfs over 5 yr, we quantify trait-based responses to climate change by comparing observed community dynamics in transplanted turfs to field-parameterized null model simulations. Three traits related to species architecture (maximum height, number of dormant meristems, and ramet-ramet connection persistence) varied consistently along spatial temperature gradients and also correlated to changes in species abundances in turfs transplanted to warmer climates. Two traits associated with resource acquisition strategy (SLA, leaf area) increased along spatial temperature gradients but did not correlate to changes in species abundances following warming. No traits correlated consistently with precipitation. Our study supports the hypothesis that spatial associations between plant traits and broad-scale climate variables can be predictive of community response to climate change, but it also suggests that not all traits with clear patterns along climate gradients will necessarily influence community response to an equal degree.

Climate warming is predicted to affect species and trophic interactions worldwide, and alpine ecosystems are expected to be especially sensitive to changes. In this study, we used two ongoing climate warming (open‐top chambers) experiments at Finse, southern Norway, to examine whether warming had an effect on herbivory by leaf‐chewing insects in an alpine Dryas heath community. We recorded feeding marks on the most common vascular plant species in warmed and control plots at two experimental sites at different elevations and carried out a brief inventory of insect herbivores. Experimental warming increased herbivory on Dryas octopetala and Bistorta vivipara. Dryas octopetala also experienced increased herbivory at the lower and warmer site, indicating an overall positive effect of warming, whereas B. vivipara experienced an increased herbivory at the colder and higher site indicating a mixed effect of warming. The Lepidoptera Zygaena exulans and Sympistis nigrita were the two most common leaf‐chewing insects in the Dryas heath. Based on the observed patterns of herbivory, the insects life cycles and feeding preferences, we argue that Z. exulans is the most important herbivore on B. vivipara, and S. nigrita the most important herbivore on D. octopetala. We conclude that if the degree of insect herbivory increases in a warmer world, as suggested by this study and others, complex interactions between plants, insects, and site‐specific conditions make it hard to predict overall effects on plant communities.

Temperature is increasing at unprecedented rates across most of the tundra biome. Remote-sensing data indicate that contemporary climate warming has already resulted in increased productivity over much of the Arctic, but plot-based evidence for vegetation transformation is not widespread. We analysed change in tundra vegetation surveyed between 1980 and 2010 in 158 plant communities spread across 46 locations. We found biome-wide trends of increased height of the plant canopy and maximum observed plant height for most vascular growth forms; increased abundance of litter; increased abundance of evergreen, low-growing and tall shrubs; and decreased abundance of bare ground. Intersite comparisons indicated an association between the degree of summer warming and change in vascular plant abundance, with shrubs, forbs and rushes increasing with warming. However, the association was dependent on the climate zone, the moisture regime and the presence of permafrost. Our data provide plot-scale evidence linking changes in vascular plant abundance to local summer warming in widely dispersed tundra locations across the globe. © 2012 Macmillan Publishers Limited. All rights reserved.

35 Abstract 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.

Accepted version of an article published in the journal: Environmental and Experimental Botany. Published version available on Science Direct: http://dx.doi.org/10.1016/j.envexpbot.2011.04.011

Cultivation of annual crops in the initial stage of reforestation has been commonly practiced in the tropics. In recent decades, however, cultivation of such areas has been discontinued, resulting in widespread abandoned settlements. In this article we used a former forest village settlement in Kenya, which had been cleared, cultivated and then abandoned, to study how natural vegetation recovers after such disturbances. Species richness, abundance, and composition of tree seedlings, saplings, adult trees, shrubs, and herbs were recorded in different zones, from a heavily degraded zone in the center of the settlement, through less disturbed transition zones (TZs), and in the surrounding secondary forest (SF). Species richness and abundance of tree seedlings, saplings, and adult trees increased gradually from the heavily degraded zone to the SF, whereas shrub and herb richness were the same for TZs and SF and abundance was lowest in the SF. Total species richness was highest in the SF. Some pioneer tree species were highly associated with the TZs, whereas sub-canopy tree species were associated with the SF. A group of tree species were not particularly associated with any of the four zones. Thus, these species might have good potential as restoration species. The results of our study contribute to the knowledge of natural regeneration in general, and of individual species characterizing the different stages of recovery of abandoned settlements in particular. Such information is urgently needed in designing ecologically sound management strategies for restoring abandoned forest settlements in tropical areas.

Assessing the influence of warming on leaf traits, carbon, and nutrient concentrations above and below ground to understand how the dominant sedge Kobresia pygmaea (C. B. Clarke) C. B. Clarke may respond and adapt to extant and future climate in the alpine meadow of the Qinghai-Tibetan Plateau. A warming experiment was conducted in the permafrost region of the Qinghai-Tibetan Plateau from 2008 to 2009. Two 2-year warming treatments (T1, annual warming of 2.1°C; T2, annual warming of 4.4°C) were used, and responses of leaf traits and above- and belowground carbon, nitrogen, and phosphorus concentrations of K. pygmaea were examined. The results show that both moderate (T1) and more extensive (T2) warming decreased leaf mass, leaf thickness, and vascular bundle size, and increased the mass-based photosynthetic rate (Amass) and photosynthetic nitrogen use efficiency (PNUE). A moderate warming significantly decreased leaf carbon (C), nitrogen (N), and phosphorus (P), and root C and N concentrations of K. pygmaea. These decreases were even more pronounced under the more extensive warming. The decreases in leaf N and P were significantly larger than the decrease in leaf C concentration. Root P concentration increased more under the extensive than the moderate warming. The observed increase in leaf C:N ratio in the warming treatment indicates that enhanced temperature may increase the long-term nitrogen use efficiency of K. pygmaea leaves. This again suggests that K. pygmaea might adapt well to future climate warming, and that nitrogen might be a more important factor for K. pygmaea dominated alpine meadows under future climate warming.

Summary 1. It is not clear how local and regional processes influence plant species recruitment. Local abiotic factors are predicted to constrain recruitment under harsh environmental conditions and low productivity, whereas the role of seed availability may be more important at intermediate productivity. The role of establishment limitation (local interactions) is predicted to increase under high productivity due to competition. 2. This study examined the effects of local biotic interactions, such as competition and facilitation from the resident vegetation, on alpine plant species recruitment, and investigated whether the role of species interactions depended on productivity. All naturally emerging seedlings were recorded in plots with intact vegetation and where vegetation had been removed, in a natural low-productivity alpine Dryas heath and in plots where productivity had increased after fertilizing. 3. Positive effects of vegetation removal on seedling emergence suggested that competition from neighbour vegetation may have an important role for recruitment in alpine plant communities. The role of competitive release increased with productivity. Productivity increased short-term, but not long-term seedling recruitment, likely due to increased competition from re-growth over time. There was no effect of productivity on species richness recovery after vegetation removal. 4. There was a positive relationship between neighbour species richness and seedling density in the low-productivity plots, but not in the high-productivity plots. Local seed production was the major seed source in both parts of the heath, but in the high-productivity plots, the high abundance of a few species in the species pool was more important for seedling recruitment than species richness. 5. Synthesis. Competition may play a significant role for plant species recruitment in environmentally harsh habitats of low productivity, where abiotic factors or facilitation traditionally are predicted to be more important ecological filters. Climate warming and increased nitrogen deposition may result in higher productivity of tundra plant communities, which may increase the role of competition, and the identity of neighbour species may become more important for recruitment than species richness.

Slash-and-burn agriculture is an important driver of deforestation and ecosystem degradation, with large effects on biodiversity and carbon sequestration. This study was conducted in a forest in Madagascar, which consists of fragments of slash-and-burn patches, within a matrix of secondary and primary forest. By recording species richness, abundance, and composition of trees, shrubs, and herbs in fallows of various age and slash-and-burn history, and in the secondary and primary forest, we show how slash-and-burn intensity (number of cycles, duration of abandonment), years since last abandonment, and environmental factors (distance to primary forest and topography) affect the natural succession and recovery of the forest ecosystem. We used ordination analyses to examine how the species composition varied between the different successions stages, and to examine tree recruitment. Our results show shrub dominance the first years after abandonment. Thereafter, a subsequent increase in species richness and abundance of tree seedlings and saplings suggests a succession towards the diversity and composition of the secondary and primary forest, although a big gap between the oldest fallows and the secondary forest shows that this will take much more than 30 years. A high number and frequency of slash-and-burn cycles decreased tree seedling and sapling richness and abundance, suggesting that reducing slash-and-burn intensity will increase the speed of tree recruitment and fallow recovery. Trees can be planted into fallows to speed up vegetation and soil recovery, such that fallows can be usable within needed time and thus the extension of cultivated areas reduced. We recommend further testing of six potential species for restoration based on their early colonization of the fallows and their survival through vegetation succession.

The effects of environmental change on soil animal communities are poorly known. Norwegian mountains are subject to both atmospheric nitrogen deposition and increased temperature. In a nutrient poor alpine Dryas heath in south Norway, soil arthropods were studied after 4 years of simulated environmental change by warming and/or nutrient addition. Warming alone only affected three low-density Collembola species, while nutrient addition, with or without warming, greatly changed the dominance hierarchy of the microarthropod community. Certain Collembola species with a short (1 year) life cycle and predatory Gamasina mites increased markedly in density. These groups may have been favored by increased litter production, as plant biomass and litter producing graminoids and forbs increased significantly in plots with nutrient addition and nutrient addition combined with warming. Microarthropods with a longer life cycle, such as Oribatida and certain Collembola, were generally unaffected by nutrient addition and probably need more time to respond. The number of Oribatida taxa was, however, reduced in plots with nutrient addition, both with and without warming. A ground-living species of Coccoidea (Homoptera) declined in plots with nutrient addition and warming compared with only warming, probably due to reduced cover of its host plant Dryas. The density of Diptera larvae (Sciaridae and Chironomidae) was unaffected by the treatments. Our results show that increased nutrient availability in nutrient poor alpine soils may have large but different effects on different taxa of soil animals. Species with short life cycles reacted first. Nutrient addition and nutrient addition combined with warming resulted in several effects below ground on microarthropods as previously shown above ground on plants: Increased biomass, high dominance of a few rapid-growing species, contrasting responses of closely related species, and a reduction in species numbers. These short-term responses may have profound long-term effects in this alpine ecosystem.

Environmental change, caused by nitrogen deposition and temperature increase, is predicted to affect allocation to carbon-based secondary compounds (CBSCs) in plants, due to changes in their internal carbon resources. The CBSCs are considered important for plant resistance to biotic and abiotic environmental stresses, such as herbivory, pathogen attacks, and UV radiation. To determine how allocation to putative defense compounds is affected by N deposition and increased temperature, we analyzed the composition of CBSCs in leaves of three arctic-alpine plant species: Bistorta vivipara, Dryas octopetala, and Salix reticulata after 5 years of warming (by open-top chambers) and experimental nutrient addition in an alpine Dryas heath in southern Norway. The dry weight of leaves increased after nutrient addition and warming combined with nutrient addition in all three species, while the weight of D. octopetala leaves also increased with warming alone. Individual chemical compounds or compound groups reacted to ...

We examined the relationship between plant community diversity (species richness, evenness) and stability in an alpine Dryas octopetala heath exposed to four years of warming (open top chambers) and nutrient addition at Finse, southern Norway. Furthermore, we examined if different functional types responded differently to the interaction between environmental change and initial community diversity. We used the temporal change in species composition, calculated as the sum of change in sample scores in detrended correspondence analyses, as a measure of variability (opposite of stability). Under ambient conditions, a high initial species richness was associated with less stability of the total species composition. Under experimental warming, initial high species richness and evenness were associated with a more stable subsequent vascular species composition. Vascular stability decreased, however, with higher species richness under nutrient addition. When warming and nutrient addition were combined, high init...

Question: How will warming and increased nutrient availability affect individual alpine plant species abundances (non-vascular and vascular) and community composition? Location: Dryas octopetala heath at alpine Finse, southern Norway. Methods: Four years with experimental warming (open top chambers) and nutrient addition. Detrended Correspondence Analysis and Redundancy Analysis were used to examine changes in community composition. GLM-ANOVA was used to examine treatment effects on individual species. Results: Warming alone decreased the abundance of some Carex and bryophyte species, but did not affect community composition. Nutrient addition and warming combined with nutrient addition increased the abundance of high stature species, such as grasses (Festuca spp., Poa alpina) and some forbs (e.g. Cerastium alpinum, Potentilla crantzii). Low stature forbs (e.g. Tofieldia pusilla), a lycophyte (Selaginella selaginoides) and most bryophytes and lichens decreased in abundance. After four years of wa...

Most studies on factors determining diversity are conducted in temperate or warm regions, whereas studies in climatically harsh and low productivity areas, such as alpine regions, are rare. We examined the relative roles of seed availability and different biotic and abiotic factors for the diversity of an alpine plant community in southern Norway. Furthermore, because climate warming is predicted to be an important driver of alpine species diversity, we assessed how the relative impacts of dispersal and local interactions on diversity might change under experimental warming (open top chambers, OTCs). Addition of seeds from 27 regional species increased community diversity. The establishment of the species was negatively related both to the diversity of the existing system and the cover of the abundant dwarf shrub Dryas octopetala. These results show that both species dispersal limitation and local biotic interactions are important factors for alpine plant community diversity. Despite relatively harsh environmental conditions and low productivity, competition from the resident vegetation appeared to have a greater role for species establishment and diversity than facilitation and experimental warming. Higher temperature appeared to increase the negative relationship between resident species diversity and species establishment. This may suggest that climate warming can increase the role of interspecific competition for alpine plant community structure, and thus alter the long-term effects of biotic interactions on diversity.

Soil seed banks offer plants the possibility to disperse through time. This has implications for population and community dynamics, as recognised by ecological and evolutionary theory. In contrast, the conservation and restoration literature often find seed banks to be depauperate, weedy and without much conservation value or restoration potential. One explanation for these contrasting views might lie in a systematic bias in the sampling of seed banks versus established plant communities. We use the species–area relationship as a tool to assess and compare the per-area species richness and spatial structuring of the diversity of the established plant community versus soil seed banks. To allow this direct comparison we extensively survey the species–area relationship of the vegetation and underlying seed bank of a grassland community across twelve sites spanning regional bioclimatic gradients. We also compile a global dataset of established vegetation and seed banks from published sources. We find that seed banks have consistently higher intercepts and slopes of the relationship, and hence higher diversity at any given spatial scale, than the vegetation both in the field and literature study. This is consistent across habitat types, climate gradients, and biomes. Similarity indices are commonly used to compare vegetation and seed bank, and we find that sampling effort (% of the vegetation area sampled for seed bank) was the strongest predictor of vegetation–seed bank similarity for both the Sørensen (R2 = 0.70) and the Raup–Crick (R2 = 0.25) index. Our study suggests that the perception that seed banks are intrinsically less diverse than established plant communities has been based more on inadequate sampling than on biological reality. Across a range of ecosystems and climatic settings, we find high diversity in seed banks relative to the established community, suggesting potentially important roles of seed banks in population dynamics and diversity maintenance. publishedVersion

Alpine and arctic ecosystems may be particularly vulnerable to climate change. We know little about alpine plant community responses to the predicted abiotic changes, or to possible changes in the biotic environment caused by climate change. Four years of experimental warming and nutrient addition altered dominance hierarchies, community structure, and diversity of an alpine biodiversity hotspot in south Norway. The previously dominant dwarf shrub Dryas octopetala was replaced by graminoids and forbs under nutrient addition and warming with nutrients. Community diversity declined due to decreased bryophyte and lichen richness and abundances, and dwarf shrub abundances. In controls and in plots with only warming, where Dryas maintained dominance, the relationships between changes in Dryas cover and changes in community parameters were negative, suggesting that Dryas controls community processes. Under nutrient addition, bryophyte and lichen diversity decreased with decreasing Dryas cover, probably due to increased competition from graminoids and forbs. The shift in dominance hierarchies changed community structure and dynamics through increased biomass, vegetation height, and competition for light. Community diversity dropped primarily because changes in the abiotic environment modified biotic interactions, highlighting that species interaction must be considered in climate change experiments and in models predicting climate change effects.

1 Vegetation removal manipulations over four growing seasons in an alpine Dryas heath at Finse, south Norway, were used to examine the role of neighbour vegetation in the population dynamics of the forb Thalictrum alpinum and the sedge Carex vaginata. We also increased temperature (by open top chambers) and nutrient availability to assess if effects of surrounding vegetation may be modified by environmental change. 2 Removal of neighbour vegetation increased the number of leaves of Thalictrum and Carex, whereas flowering stems and leaf stalks of Thalictrum became significantly shorter. Furthermore, increased plant growth after nutrient addition and warming suggests that both abiotic environmental conditions and biotic interactions limit plant growth at Finse. 3 Our results suggest that there is competition for nutrients at Finse, but that surrounding vegetation may also have facilitative sheltering effects on other species. 4 Interactions between vegetation removal and abiotic factors on population dynamic parameters of Thalictrum suggest that plant-plant interactions may be modified if temperature and nutrient availability increase under climate change. 5 The responses of the two target species to the removal of the general neighbour vegetation were less pronounced than their responses to the removal of the dominant Dryas octopetala examined in a previous study. This suggests that one dominant species may affect the population dynamics of other species more than the net effect of all the other species in the community in concert. 6 Our results show that biotic interactions may affect alpine plant populations and thereby structure alpine plant communities. Furthermore, the effects of neighbours and abiotic environmental changes on alpine plant growth may depend on the species identity of the neighbours.

Summary 1I examined the role of species interactions on the population dynamics of the herb Thalictrum alpinum and the sedge Carex vaginata by removing the dwarf shrub Dryas octopetala over four growing seasons at Finse, in the alpine region of south Norway, Furthermore, by increasing temperature (open top chambers) and nutrient availability (nutrient addition), I assessed the effects of climate warming on biotic interactions. 2Dryas removal increased the number of Thalictrum and Carex leaves, and flowering frequency of Carex. Flower stems and leaf stalks of Thalictrum, and the leaves of Carex, became significantly shorter following Dryas removal. Warming and nutrient addition increased vegetative plant growth, whereas warming alone had positive effects on sexual reproductive effort of the target species. My results suggest that there is both competition for nutrients, and a facilitative shelter effect of Dryas on its neighbours. 3Species interactions affected population dynamics of Thalictrum and Carex. Interactions between the impacts of Dryas removal and abiotic factors on leaf production, suggested that interactions between Dryas and neighbouring species might be modified if temperature and, in particular, nutrient availability increase under global warming. 4This study shows that both biotic interactions and abiotic environmental conditions may affect alpine plant population dynamics. Furthermore, it shows that climate change may modify species interactions. 5Species interaction effects should be included in climate change experiments and in future models predicting plant community changes under global warming.

The rapidly warming temperatures in high-latitude and alpine regions have the potential to alter the phenology of Arctic and alpine plants, affecting processes ranging from food webs to ecosystem trace gas fluxes. The International Tundra Experiment (ITEX) was initiated in 1990 to evaluate the effects of expected rapid changes in temperature on tundra plant phenology, growth and community changes using experimental warming. Here, we used the ITEX control data to test the phenological responses to background temperature variation across sites spanning latitudinal and moisture gradients. The dataset overall did not show an advance in phenology; instead, temperature variability during the years sampled and an absence of warming at some sites resulted in mixed responses. Phenological transitions of high Arctic plants clearly occurred at lower heat sum thresholds than those of low Arctic and alpine plants. However, sensitivity to temperature change was similar among plants from the different climate zones. Plants of different communities and growth forms differed for some phenological responses. Heat sums associated with flowering and greening appear to have increased over time. These results point to a complex suite of changes in plant communities and ecosystem function in high latitudes and elevations as the climate warms.

Recent studies from mountainous areas of small spatial extent (2500 km(2) ) suggest that fine-grained thermal variability over tens or hundreds of metres exceeds much of the climate warming expected for the coming decades. Such variability in temperature provides buffering to mitigate climate-change impacts. Is this local spatial buffering restricted to topographically complex terrains? To answer this, we here study fine-grained thermal variability across a 2500-km wide latitudinal gradient in Northern Europe encompassing a large array of topographic complexities. We first combined plant community data, Ellenberg temperature indicator values, locally measured temperatures (LmT) and globally interpolated temperatures (GiT) in a modelling framework to infer biologically relevant temperature conditions from plant assemblages within1000-m(2) units (community-inferred temperatures: CiT). We then assessed: (1) CiT range (thermal variability) within 1-km(2) units; (2) the relationship between CiT range and topographically and geographically derived predictors at 1-km resolution; and (3) whether spatial turnover in CiT is greater than spatial turnover in GiT within 100-km(2) units. Ellenberg temperature indicator values in combination with plant assemblages explained 46-72% of variation in LmT and 92-96% of variation in GiT during the growing season (June, July, August). Growing-season CiT range within 1-km(2) units peaked at 60-65°N and increased with terrain roughness, averaging 1.97 °C (SD = 0.84 °C) and 2.68 °C (SD = 1.26 °C) within the flattest and roughest units respectively. Complex interactions between topography-related variables and latitude explained 35% of variation in growing-season CiT range when accounting for sampling effort and residual spatial autocorrelation. Spatial turnover in growing-season CiT within 100-km(2) units was, on average, 1.8 times greater (0.32 °C km(-1) ) than spatial turnover in growing-season GiT (0.18 °C km(-1) ). We conclude that thermal variability within 1-km(2) units strongly increases local spatial buffering of future climate warming across Northern Europe, even in the flattest terrains.

Opportunities for observing long-term changes in natural biota are rare. Observations on the distribution and frequency of vascular plants were performed on 23 mountains situated along a west–east gradient in Jotunhei men, central Norway, where detailed site descriptions and species lists exist from ad 1930–31. The sites were resurveyed during the summer of 1998, to examine possible changes in species richness and species distributions along the altitudinal gradient during a 68-year period. Increased species richness was found on 19 of the mountains and was most pronounced at lower altitudes and in the eastern areas. Lowland species, dwarf shrubs and species with wide altitudinal and ecological ranges showed the greatest increases in abundance and altitudinal advances since the 1930–31 study. Species with more restricted habitat demands, such as some hygrophilous snow-bed species, have declined. High-altitude species have disappeared from their lower-elevation sites and increased their abundance at the highest altitudes. Climatic warming occurring in the last 100 years might have allowed the invasion of lowland and lee-slope species. Increased competition at sites where such species have invaded may have led to a decreased abundance of the less competitive species and a concentration of high-altitude species on the highest ridges. Natural succession since the ‘Little Ice Age’, increased deposition of nitrogen during recent years and changes in grazing and tourism might have in‘ uenced some of the species turnovers, but recent climatic changes are considered to be the most likely major driving factor for the changes observed.