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Climate change is rapidly warming high latitude and high elevation regions influencing plant community composition. Changes in vegetation composition have motivated the coordination of ecological monitoring networks across the Arctic, including the International Tundra Experiment. We have established a long-term passive warming experiment using open-top chambers, which includes five distinct plant communities (Dry Heath; Tussock Tundra; and Dry, Mesic, and Wet Meadow). We measured changes in plant community composition based on relative abundance differences over 26 years. In addition, relative abundance changes in response to fertilization and warming treatments were analyzed based on a seven-year Community-Level Interaction Program experiment. The communities had distinct soil moisture conditions, leading to community-specific responses of the plant growth forms (deciduous shrubs, evergreen shrubs, forbs, and graminoids). Warming significantly affected growth forms, but the direction of the response was not consistent across the communities. Evidence of shrub expansion was found in nearly all communities, with soil moisture determining whether it was driven by deciduous or evergreen shrubs. Graminoids increased in relative abundance in the Dry Meadow due to warming. Growth form responses to warming are likely mediated by edaphic characteristics of the communities and their interactions with climate.

The soil nitrogen (N) cycle in cold terrestrial ecosystems is slow and organically bound N is an important source of N for plants in these ecosystems. Many plant species can take up free amino acids from these infertile soils, either directly or indirectly via their mycorrhizal fungi. We hypothesized that plant community changes and local plant community differences will alter the soil free amino acid pool and composition; and that long-term warming could enhance this effect. To test this, we studied the composition of extractable free amino acids at five separate heath, meadow, and bog locations in subarctic and alpine Scandinavia, with long-term (13 to 24 years) warming manipulations. The plant communities all included a mixture of ecto-, ericoid-, and arbuscular mycorrhizal plant species. Vegetation dominated by grasses and forbs with arbuscular and non-mycorrhizal associations showed highest soil free amino acid content, distinguishing them from the sites dominated by shrubs with ecto- and ericoid-mycorrhizal associations. Warming increased shrub and decreased moss cover at two sites, and by using redundancy analysis, we found that altered soil free amino acid composition was related to this plant cover change. From this, we conclude that the mycorrhizal type is important in controlling soil N cycling and that expansion of shrubs with ectomycorrhiza (and to some extent ericoid mycorrhiza) can help retain N within the ecosystems by tightening the N cycle.

Arctic ecosystems are warming nearly four times faster than the global average which is resulting in plant community shifts and subsequent changes in biogeochemical processes such as gaseous fluxes. Additionally, herbivores shape plant communities and thereby alter the magnitude and composition of ecosystem respiration and BVOC emissions. Here we determine the effect of large mammalian herbivores on ecosystem respiration and BVOC emissions in two southern and two northern sites in Sweden, encompassing mountain birch (LOMB) and shrub heath (LORI) communities in the south and low-herb meadow (RIGA) and shrub heath (RIRI) communities in the north. Herbivory significantly decreased ecosystem respiration at RIGA and altered the BVOC composition between sites. However, plant community composition had a larger effect on ecosystem respiration as RIGA had 35 % higher emissions than the next highest emitting site (LOMB). Additionally, LOMB had the highest emissions of terpenes with the northern sites having significantly lower emissions. Differences between sites were primarily due to differences in exclosure effects, soil temperature and prevalence of different shrub growth forms. Our results suggest that herbivory has a significant effect on trace gas fluxes in a productive meadow community and differences between communities may be driven by differences in shrub composition.

Observations of changes in phenology have provided some of the strongest signals of the effects of climate change on terrestrial ecosystems. The International Tundra Experiment (ITEX), initiated in the early 1990s, established a common protocol to measure plant phenology in tundra study areas across the globe. Today, this valuable collection of phenology measurements depicts the responses of plants at the colder extremes of our planet to experimental and ambient changes in temperature over the past decades. The database contains 150,434 phenology observations of 278 plant species taken at 28 study areas for periods of 1 to 26 years. Here we describe the full dataset to increase the visibility and use of these data in global analyses, and to invite phenology data contributions from underrepresented tundra locations. Portions of this tundra phenology database have been used in three recent syntheses, some datasets are expanded, others are from entirely new study areas, and the entirety of these data are now available at the Polar Data Catalogue., Arctic Science, 8 (3), ISSN:2368-7460

In Arctic regions, thawing permafrost soils are projected to release 50 to 250 Gt of carbon by 2100. This data is mostly derived from carbon-rich wetlands, although 71% of this carbon pool is stored in faster-thawing mineral soils, where ecosystems close to the outer boundaries of permafrost regions are especially vulnerable. Although extensive data exists from currently thawing sites and short-term thawing experiments, investigations of the long-term changes following final thaw and co-occurring drainage are scarce. Here we show ecosystem changes at two comparable tussock tundra sites with distinct permafrost thaw histories, representing 15 and 25 years of natural drainage, that resulted in a 10-fold decrease in CH4 emissions (3.2 ± 2.2 vs. 0.3 ± 0.4 mg C-CH4 m−2 day−1), while CO2 emissions were comparable. These data extend the time perspective from earlier studies based on short-term experimental drainage. The overall microbial community structures did not differ significantly between sites, although the drier top soils at the most advanced site led to a loss of methanogens and their syntrophic partners in surface layers while the abundance of methanotrophs remained unchanged. The resulting deeper aeration zones likely increased CH4 oxidation due to the longer residence time of CH4 in the oxidation zone, while the observed loss of aerenchyma plants reduced CH4 diffusion from deeper soil layers directly to the atmosphere. Our findings highlight the importance of including hydrological, vegetation and microbial specific responses when studying long-term effects of climate change on CH4 emissions and underscores the need for data from different soil types and thaw histories.

Centro de Biotecnología y Genómica de Plantas (CBGP), The evolutionary success of plants relies to a large extent on their extraordinary ability to adapt to changes in their environment. These adaptations require that plants balance their growth with their stress responses. Plant hormones are crucial mediators orchestrating the underlying adaptive processes. However, whether and how the growth-related hormone auxin and the stress-related hormones jasmonic acid, salicylic acid, and abscisic acid (ABA) are coordinated remains largely elusive. Here, we analyse the physiological role of AMIDASE 1 (AMI1) in Arabidopsis plant growth and its possible connection to plant adaptations to abiotic stresses. AMI1 contributes to cellular auxin homeostasis by catalysing the conversion of indole-acetamide into the major plant auxin indole-3-acetic acid. Functional impairment of AMI1 increases the plant's stress status rendering mutant plants more susceptible to abiotic stresses. Transcriptomic analysis of ami1 mutants disclosed the reprogramming of a considerable number of stress-related genes, including jasmonic acid and ABA biosynthesis genes. The ami1 mutants exhibit only moderately repressed growth but an enhanced ABA accumulation, which suggests a role for AMI1 in the crosstalk between auxin and ABA. Altogether, our results suggest that AMI1 is involved in coordinating the trade-off between plant growth and stress responses, balancing auxin and ABA homeostasis., This research was supported by grants from the German Research Foundation (DFG, SFB480/A10) and the Spanish Ministry of Economy, Industry and Competitiveness (MINECO, BFU2017-82826-R to SP and a grant from the Swedish Research Council (VR) to HA. JM was supported by the ‘Severo Ochoa Program for Centers of Excellence in R&D’ from the Agencia Estatal de Investigación of Spain, grant SEV-2016-0672 (2017-2021) to the CBGP., 17 Pág.

Open top chambers (OTCs) were adopted as the recommended warming mechanism by the International Tundra Experiment network in the early 1990s. Since then, OTCs have been deployed across the globe. Hundreds of papers have reported the impacts of OTCs on the abiotic environment and the biota. Here, we review the impacts of the OTC on the physical environment, with comments on the appropriateness of using OTCs to characterize the response of biota to warming. The purpose of this review is to guide readers to previously published work and to provide recommendations for continued use of OTCs to understand the implications of warming on low stature ecosystems. In short, the OTC is a useful tool to experimentally manipulate temperature; however, the characteristics and magnitude of warming varies greatly in different environments; therefore, it is important to document chamber performance to maximize the interpretation of biotic response. When coupled with long-term monitoring, warming experiments are a valuable means to understand the impacts of climate change on natural ecosystems. Open top chambers (OTCs) were adopted as the recommended warming mechanism by the International Tundra Experiment network in the early 1990s. Since then, OTCs have been deployed across the globe. Hundreds of papers have reported the im-pacts of OTCs on the abiotic environment and the biota. Here, we review the impacts of the OTC on the physical environment, with comments on the appropriateness of using OTCs to characterize the response of biota to warming. The purpose of this review is to guide readers to previously published work and to provide recommendations for continued use of OTCs to under -stand the implications of warming on low stature ecosystems. In short, the OTC is a useful tool to experimentally manipulate temperature; however, the characteristics and magnitude of warming varies greatly in different environments; therefore, it is important to document chamber performance to maximize the interpretation of biotic response. When coupled with long-term monitoring, warming experiments are a valuable means to understand the impacts of climate change on natural ecosystems.

Global warming has pronounced effects on tundra vegetation, and rising mean temperatures increase plant growth potential across the Arctic biome. Herbivores may counteract the warming impacts by reducing plant growth, but the strength of this effect may depend on prevailing regional climatic conditions. To study how ungulates interact with temperature to influence growth of tundra shrubs across the Arctic tundra biome, we assembled dendroecological data from 20 sites, comprising 1153 individual shrubs and 223 63 annual growth rings. Evidence for ungulates suppressing shrub radial growth was only observed at intermediate summer temperatures (6.5 ◦C–9 ◦C), and even at these temperatures the effect was not strong. Multiple factors, including forage preferences and landscape use by the ungulates, and favourable climatic conditions enabling effective compensatory growth of shrubs, may weaken the effects of ungulates on shrubs, possibly explaining the weakness of observed ungulate effects. Earlier local studies have shown that ungulates may counteract the impacts of warming on tundra shrub growth, but we demonstrate that ungulates’ potential to suppress shrub radial growth is not always evident, and may be limited to certain climatic conditions. Arctic, browsing, climate change, dendroecology, herbivory, shrub, tundra

Alpine ecosystems harbor remarkably diverse and distinct plant communities that are characteristically limited to harsh, and cold climatic conditions. As a result of thermal limitation to species occurrence, mountainous ecosystems are considered to be particularly sensitive to climate change. Our understanding of the impact of climate change is mainly based on vascular plants however, whereas cryptogams (i.e., lichens and bryophytes) are generally neglected or simply considered as one functional group. Here we aimed to improve our understanding of the drivers underlying temporal changes in vegetation of alpine ecosystems. To this end, we repeatedly surveyed the vegetation on four mountain summits along an elevational gradient in northern Sweden spanning a 19-year period. Our results show that the vascular plant communities remained relatively stable throughout the study period, despite fluctuations in terms of ground cover and species richness of shrubs and graminoids. In contrast, both lichens and bryophytes substantially decreased in cover and diversity, leading to alterations in community composition that were unrelated to vascular plant cover. Thermophilization of the vascular plant community was found only on the two intermediate summits. Our findings are only partially consistent with (long-term) climate-change impacts, and we argue that local non-climatic drivers such as herbivory might offset vegetation responses to warming. Hence, we underline the importance of considering local non-climatic drivers when evaluating temporal vegetation change in biologically complex systems.

1. Large herbivores influence plant community structure and ecosystem processes in many ecosystems. In large parts of the Arctic, reindeer (or caribou) are the only large herbivores present. Recent studies show that reindeer have the potential to mitigate recent warming-induced shrub encroachment in the Arctic and the associated greening of high-latitude ecosystems. This will potentially have large scale consequences for ecosystem productivity and carbon cycling. 2. To date, information on variation in the interactions between reindeer and plants across Arctic landscapes has been scarce. We utilized a network of experimental sites across a latitudinal gradient in the Scandinavian mountains where reindeer have been excluded from 59 study plots for at least 15 years. We used this study system to test the effect of long-term exclusion of reindeer on the abundance of major plant functional groups, the greenness indexes Leaf Area Index (LAI) and Normalized Difference Vegetation Index (NDVI), soil mineral nitrogen (N) and phosphorous (P), and species richness, and to determine whether the effect of reindeer exclusion is dependent on reindeer density, productivity, soil fertility or climate. 3. We found that NDVI and LAI, lichen and deciduous shrub abundances were largely reduced while soil mineral N was enhanced by reindeer. The direction and amplitude of other plant functional group responses to reindeer exclusion differed between forest and tundra as well as shrub- and herbaceous-dominated vegetation. Higher reindeer densities were related to decreased plant species richness in low -productive sites and to increased species richness in productive sites. 4. The relative reduction in LAI and associated absolute reductions of deciduous shrubs in response to reindeer were positively related to reindeer density, while the relative reduction in NDVI was not. Further, relative reductions in LAI and NDVI in response to reindeer were unrelated to climate and soil fertility. 5. Synthesis. Our results provide long-term experimental evidence highlighting the role of reindeer density in regulating plant species richness, global climate change induced greenness patterns and shrub encroachment at regional scales in the Arctic. These findings emphasize the need to consider reindeer in models predicting vegetation patterns and changes in high-latitude ecosystems. peerReviewed

Arctic environmental change has induced shifts in high latitude plant community composition and stature with impli-cations for Arctic carbon cycling and energy exchange. Two major components of high latitude ecosystems undergoing change is the advancement of trees into treeless tundra and the increased abundance and size of shrubs. How future changes in key climatic and environmental drivers will affect distributions of major ecosystem types is an active area of research. Dynamic Vegetation Models (DVMs) offer a way to investigate multiple and interacting drivers of vegeta-tion distribution and ecosystem function. We employed the LPJ-GUESS DVM over a subarctic landscape in northern Sweden, Torneträsk. Using a highly resolved climate dataset we downscaled CMIP5 climate data from three Global Climate Models and two 21st century future scenarios (RCP2.6 and RCP8.5) to investigate future impacts of climate change on these ecosystems. We also performed three model experiments where we factorially varied drivers (climate, nitrogen deposition and [CO2]) to disentangle the effects of each on ecosystem properties and functions. We found that treelines could advance by between 45 and 195 elevational meters in the landscape until the year 2100, depending on the scenario. Temperature was a strong, but not the only, driver of vegetation change. Nitrogen availability was identi-fied as an important modulator of treeline advance. While increased CO2 fertilisation drove productivity increases it did not result in any range shifts of trees. Treeline advance was realistically simulated without any temperature depend-ence on growth, but biomass was overestimated. As nitrogen was identified as an important modulator of treeline ad-vance, we support the idea that accurately representing plant-soil interactions in models will be key to future predic-tions Arctic vegetation change.

Current analyses and predictions of spatially explicit patterns and processes in ecology most often rely on climate data interpolated from standardized weather stations. This interpolated climate data represents long-term average thermal conditions at coarse spatial resolutions only. Hence, many climate-forcing factors that operate at fine spatiotemporal resolutions are overlooked. This is particularly important in relation to effects of observation height (e.g. vegetation, snow and soil characteristics) and in habitats varying in their exposure to radiation, moisture and wind (e.g. topography, radiative forcing or cold-air pooling). Since organisms living close to the ground relate more strongly to these microclimatic conditions than to free-air temperatures, microclimatic ground and near-surface data are needed to provide realistic forecasts of the fate of such organisms under anthropogenic climate change, as well as of the functioning of the ecosystems they live in. To fill this critical gap, we highlight a call for temperature time series submissions to SoilTemp, a geospatial database initiative compiling soil and near-surface temperature data from all over the world. Currently, this database contains time series from 7,538 temperature sensors from 51 countries across all key biomes. The database will pave the way toward an improved global understanding of microclimate and bridge the gap between the available climate data and the climate at fine spatiotemporal resolutions relevant to most organisms and ecosystem processes. ispartof: GLOBAL CHANGE BIOLOGY vol:26 issue:11 pages:6616-6629 ispartof: location:England status: published

The evolutionary success of plants relies to a large extent on their extraordinary ability to adapt to changes in their environment. These adaptations require that plants balance their growth with their stress responses. Plant hormones are crucial mediators orchestrating the underlying adaptive processes. However, whether and how the growth-related hormone auxin and the stress-related hormones jasmonic acid (JA), salicylic acid, and abscisic acid (ABA) are coordinated remains largely elusive. Here, we analyze the physiological role of AMIDASE 1 (AMI1) in plant growth and its possible connection to plant adaptations to abiotic stresses. AMI1 contributes to cellular auxin homeostasis by catalyzing the conversion of indole-acetamide into the major plant auxin indole-3-acetic acid. Functional impairment of AMI1 increases the plants’ stress status rendering mutant plants more susceptible to abiotic stresses. Transcriptomic analysis of ami1 mutants disclosed the reprogramming of a considerable number of stress-related genes, including JA and ABA biosynthesis genes. The ami1 mutants exhibit only moderately repressed growth, but an enhanced ABA accumulation, which suggests a role for AMI1 in the crosstalk between auxin and ABA. Altogether, our results suggest that AMI1 is involved in coordinating the trade-off between plant growth and stress responses, balancing auxin with ABA homeostasis.HIGHLIGHTThe IAM amidohydrolase AMI1 catalyzes the conversion of IAM into IAA in vivo. Expression of AMI1 is specifically repressed by osmotic stress conditions, which triggers ABA biosynthesis through the induction of NCED3, thereby linking auxin homeostasis with plant stress responses.

The majority of variation in six traits critical to the growth, survival and reproduction of plant species is thought to be organised along just two dimensions, corresponding to strategies of plant size and resource acquisition. However, it is unknown whether global plant trait relationships extend to climatic extremes, and if these interspecific relationships are confounded by trait variation within species. We test whether trait relationships extend to the cold extremes of life on Earth using the largest database of tundra plant traits yet compiled. We show that tundra plants demonstrate remarkably similar resource economic traits, but not size traits, compared to global distributions, and exhibit the same two dimensions of trait variation. Three quarters of trait variation occurs among species, mirroring global estimates of interspecific trait variation. Plant trait relationships are thus generalizable to the edge of global trait-space, informing prediction of plant community change in a warming world., It is unclear whether plant trait relationships found at the global scale extend to climatic extremes. Here the authors analyse six major aboveground traits to show that known plant trait relationships extend to the tundra biomes and exhibit the same two dimensions of variation detected at the global scale.

Vast amounts of carbon are bound in both active layer and permafrost soils in the Arctic. As a consequence of climate warming, the depth of the active layer is increasing in size and permafrost soils are thawing. We hypothesize that pulses of biogenic volatile organic compounds are released from the near-surface active layer during spring, and during late summer season from thawing permafrost, while the subsequent biogeochemical processes occurring in thawed soils also lead to emissions. Biogenic volatile organic compounds are reactive gases that have both negative and positive climate forcing impacts when introduced to the Arctic atmosphere, and the knowledge of their emission magnitude and pattern is necessary to construct reliable climate models. However, it is unclear how different ecosystems and environmental factors such as drainage conditions upon permafrost thaw affect the emission and compound composition. Here we show that incubations of frozen B horizon of the active layer and permafrost soils collected from a High Arctic heath and fen release a range of biogenic volatile organic compounds upon thaw and during subsequent incubation experiments at temperatures of 10°C and 20°C. Meltwater drainage in the fen soils increased emission rates nine times, while having no effect in the drier heath soils. Emissions generally increased with temperature, and emission profiles for the fen soils were dominated by benzenoids and alkanes, while benzenoids, ketones, and alcohols dominated in heath soils. Our results emphasize that future changes affecting the drainage conditions of the Arctic tundra will have a large influence on volatile emissions from thawing permafrost soils – particularly in wetland/fen areas.

Global warming leads to drastic changes in the diversity and structure of Arctic plant communities. Studies of functional diversity within the Arctic tundra biome have improved our understanding of plant responses to warming. However, these studies still show substantial unexplained variation in diversity responses. Complementary to functional diversity, phylogenetic diversity has been useful in climate change studies, but has so far been understudied in the Arctic. Here, we use a 25 year warming experiment to disentangle community responses in Arctic plant phylogenetic β diversity across a soil moisture gradient. We found that responses varied over the soil moisture gradient, where meadow communities with intermediate to high soil moisture had a higher magnitude of response. Warming had a negative effect on soil moisture levels in all meadow communities, however meadows with intermediate moisture levels were more sensitive. In these communities, soil moisture loss was associated with earlier snowmelt, resulting in community turnover towards a more heath-like community. This process of ‘heathification’ in the intermediate moisture meadows was driven by the expansion of ericoid and Betula shrubs. In contrast, under a more consistent water supply Salix shrub abundance increased in wet meadows. Due to its lower stature, palatability and decomposability, the increase in heath relative to meadow vegetation can have several large scale effects on the local food web as well as climate. Our study highlights the importance of the hydrological cycle as a driver of vegetation turnover in response to Arctic climate change. The observed patterns in phylogenetic β diversity were often driven by contrasting responses of species of the same functional growth form, and could thus provide important complementary information. Thus, phylogenetic diversity is an important tool in disentangling tundra response to environmental change.

Vascular and nonvascular plants are affected by environmental factors determining their distribution and shaping their diversity and cover. Despite the cryptogam commonness in Arctic communities, previous studies have often focused on limited number of factors and their impact on only selected species of vascular plants or cryptogams. Our study aimed to investigate in detail the differences in species diversity and cover of cryptogams and vascular plants in the glacier forelands and mature tundra on Svalbard. Furthermore, we determined the biotic and abiotic factors that affected diversity, cover and distribution of cryptogam and vascular plant species. In 2017, we established 201 plots in eight locations (each including habitat type of foreland and mature tundra) and surveyed species abundance, sampled soils and environmental data. Results revealed that diversity and cover of analysed groups differed significantly between locations and habitat types, except for cryptogam cover in mature tundra in terms of location. Distance to the glacier terminus, slope, soil conductivity, nutrient content, and clay content impacted both plant groups' diversity. In contrast, distance to the glacier terminus, nutrient content and soil pH affected their cover. In addition, for cryptogam diversity and cover, foreland location and vascular plant cover were also important, while for vascular plant cover time elapsed after glacier retreat was significant. Distribution of both groups' species in forelands was associated with time elapsed after glacier retreat, soil pH, and nutrient contents. Soil texture and distance to the glacier terminus additionally influenced cryptogam distribution. The positive impact of vascular plants on cryptogam diversity and cover indicates complex relationships between these groups, even in forelands' relatively simple communities. As the cryptogam diversity in the polar areas is high but still largely unknown, future studies on species ecology and climate change impact on vegetation should consider both vascular plants and cryptogams and interactions between these groups.

Altres ajuts europeus: P.A.W. was additionally supported by the European Union Fourth Environment and Climate Framework Programme (Project Number ENV4-CT970586)P.A.W. was additionally supported by the European Union Fourth Environment and Climate Framework Programme (Project Number ENV4-CT970586). The tundra is warming more rapidly than any other biome on Earth, and the potential ramifications are far-reaching because of global feedback effects between vegetation and climate. A better understanding of how environmental factors shape plant structure and function is crucial for predicting the consequences of environmental change for ecosystem functioning. Here we explore the biome-wide relationships between temperature, moisture and seven key plant functional traits both across space and over three decades of warming at 117 tundra locations. Spatial temperature-trait relationships were generally strong but soil moisture had a marked influence on the strength and direction of these relationships, highlighting the potentially important influence of changes in water availability on future trait shifts in tundra plant communities. Community height increased with warming across all sites over the past three decades, but other traits lagged far behind predicted rates of change. Our findings highlight the challenge of using space-for-time substitution to predict the functional consequences of future warming and suggest that functions that are tied closely to plant height will experience the most rapid change. They also reveal the strength with which environmental factors shape biotic communities at the coldest extremes of the planet and will help to improve projections of functional changes in tundra ecosystems with climate warming.

Effective societal responses to rapid climate change in the Arctic rely on an accurate representation of region-specific ecosystem properties and processes. However, this is limited by the scarcity and patchy distribution of field measurements. Here, we use a comprehensive, geo-referenced database of primary field measurements in 1,840 published studies across the Arctic to identify statistically significant spatial biases in field sampling and study citation across this globally important region. We find that 31% of all study citations are derived from sites located within 50 km of just two research sites: Toolik Lake in the USA and Abisko in Sweden. Furthermore, relatively colder, more rapidly warming and sparsely vegetated sites are under-sampled and under-recognized in terms of citations, particularly among microbiology-related studies. The poorly sampled and cited areas, mainly in the Canadian high-Arctic archipelago and the Arctic coastline of Russia, constitute a large fraction of the Arctic ice-free land area. Our results suggest that the current pattern of sampling and citation may bias the scientific consensuses that underpin attempts to accurately predict and effectively mitigate climate change in the region. Further work is required to increase both the quality and quantity of sampling, and incorporate existing literature from poorly cited areas to generate a more representative picture of Arctic climate change and its environmental impacts.

Afforestation has been proposed as a strategy to mitigate the often high greenhouse gas (GHG) emissions from agricultural soils with high organic matter content. However, the carbon dioxide (CO2) and nitrous oxide (N2O) fluxes after afforestation can be considerable, depending predominantly on site drainage and nutrient availability. Studies on the full GHG budget of afforested organic soils are scarce and hampered by the uncertainties associated with methodology. In this study we determined the GHG budget of a spruce-dominated forest on a drained organic soil with an agricultural history. Two different approaches for determining the net ecosystem CO2 exchange (NEE) were applied, for the year 2008, one direct (eddy covariance) and the other indirect (analyzing the different components of the GHG budget), so that uncertainties in each method could be evaluated. The annual tree production in 2008 was 8.3 ± 3.9 t C ha−1 yr−1 due to the high levels of soil nutrients, the favorable climatic conditions and the fact that the forest was probably in its phase of maximum C assimilation or shortly past it. The N2O fluxes were determined by the closed-chamber technique and amounted to 0.9 ± 0.8 t Ceq ha−1 yr−1. According to the direct measurements from the eddy covariance technique, the site acts as a minor GHG sink of −1.2 ± 0.8 t Ceq ha−1 yr−1. This contrasts with the NEE estimate derived from the indirect approach which suggests that the site is a net GHG emitter of 0.6 ± 4.5 t Ceq ha−1 yr−1. Irrespective of the approach applied, the soil CO2 effluxes counter large amounts of the C sequestration by trees. Due to accumulated uncertainties involved in the indirect approach, the direct approach is considered the more reliable tool. As the rate of C sequestration will likely decrease with forest age, the site will probably become a GHG source once again as the trees do not compensate for the soil C and N losses. Also forests in younger age stages have been shown to have lower C assimilation rates; thus, the overall GHG sink potential of this afforested nutrient-rich organic soil is probably limited to the short period of maximum C assimilation.

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.

Wood ash additions of 3.3 and 6.6 t ha(-1) reduced greenhouse gas (GHG) emissions from a spruce forest (Picea abies) on a minerotrophic drained organic soil. Emissions of carbon dioxide (CO(2)), methane (CH(4)) and nitrous oxide (N(2)O) were measured using static dark chambers for two years following the ash treatment. The CO(2) emission from the soil was significantly reduced by 17-23% by both doses during 2006-2008. The mechanism behind the reduction could not be related to a direct inhibition of soil C mineralization by the ash. The emission of N(2)O was also significantly reduced by 44 and 46% during the first year, mainly due to reductions in the winter emissions. Similar reductions of 34 and 50% were found in the second year for the low and the high wood ash, respectively. Increased pH of the soil due to the ash additions may have caused the effect. The control and amended soils consumed ambient CH(4). The low wood ash dose increased the annual net CH(4) uptake rate by 9%, due to an increased winter uptake. No changes in tree growth could be detected over the short 2-year measurement period. The net effect of wood ash application was a reduction in the total GHG emissions during the first two years after the treatment.

Recent climate change predictions suggest altered patterns of winter precipitation across the Arctic. It has been suggested that the presence, timing and quantity of snow all affect microbial activity, thus influencing CO2 production in soil. In this study annual and seasonal emissions of CO2 were estimated in High-Arctic Adventdalen, Svalbard, and sub-Arctic Latnjajaure, Sweden, using a new trace gas-based method to track real-time diffusion rates through the snow. Summer measurements from snow-free soils were made using a chamber-based method. Measurements were obtained from different snow regimes in order to evaluate the effect of snow depth on winter CO2 effluxes. Total annual emissions of CO2 from the sub-Arctic site (0.662–1.487 kg CO2 m–2 yr–1) were found to be more than double the emissions from the High-Arctic site (0.369–0.591 kg CO2 m–2 yr–1). There were no significant differences in winter effluxes between snow regimes or vegetation types, indicating that spatial variability in winter soil CO2 effluxes are not directly linked to snow cover thickness or soil temperatures. Total winter emissions (0.004– 0.248 kg CO2 m–2) were found to be in the lower range of those previously described in the literature. Winter emissions varied in their contribution to total annual production between 1 and 18%. Artificial snow drifts shortened the snow-free period by 2 weeks and decreased the annual CO2 emission by up to 20%. This study suggests that future shifts in vegetation zones may increase soil respiration from Arctic tundra regions.

Background: Facilitation plays important roles in the structuring of plant communities and several studies have found that it tends to increase with environmental severity in alpine plant communiti ...

Background: Saplings of mountain birch ( Betula pubescens ssp. czerepanovii) have established in pockets of dwarf-shrub heath approximately 250 m above the treeline in the Latnjavagge Valley, north ...

Effects of warming on root morphology, root mass distribution and microbial activity were studied in organic and mineral soil layers in two alpine ecosystems over>10 yr, using open-top chambers, in Swedish Lapland. Root mass was estimated using soil cores. Washed roots were scanned and sorted into four diameter classes, for which variables including root mass (g dry matter (g DM) m(-2)), root length density (RLD; cm cm(-3) soil), specific root length (SRL; m g DM(-1)), specific root area (SRA; m2 kg DM(-1)), and number of root tips m(-2) were determined. Nitrification (NEA) and denitrification enzyme activity (DEA) in the top 10 cm of soil were measured. Soil warming shifted the rooting zone towards the upper soil organic layer in both plant communities. In the dry heath, warming increased SRL and SRA of the finest roots in both soil layers, whereas the dry meadow was unaffected. Neither NEA nor DEA exhibited differences attributable to warming. Tundra plants may respond to climate change by altering their root morphology and mass while microbial activity may be unaffected. This suggests that carbon may be incorporated in tundra soils partly as a result of increases in the mass of the finer roots if temperatures rise.

The spatial distribution of organic soil nitrogen (N) in alpine tundra was studied along a natural environmental gradient, covering five plant communities, at the Latnjajaure Field Station, north- ern Swedish Lapland. The five communities (mesic meadow, meadow snowbed, dry heath, mesic heath, and heath snowbed) are the dominant types in this region and are differentiated by soil pH. Net N mineralization, net ammonification, and net nitrifica- tion were measured using 40-day laboratory incuba- tions based on extractable NH4 and NO3 . Nitrification enzyme activity (NEA), denitrification enzyme activity (DEA), amino acid concentrations, and microbial respiration were measured for soils from each plant community. The results show that net N mineralization rates were more than three times higher in the meadow ecosystems (mesic meadow 0.7 m gNg � 1 OM day � 1 and meadow snowbed 0.6 m gNg � 1 OM day � 1 ) than the heath ecosystems (dry heath 0.2 m gNg � 1 OM day � 1 , mesic heath 0.1 m gNg � 1 OM day � 1 and heath snowbed 0.2 m gNg � 1 OM day � 1 ). The net N mineralization rates were negatively correlated to organic soil C/N ratio (r = � 0.652, P < 0.001) and positively correlated to soil pH (r = 0.701, P < 0.001). Net nitrification, inorganic N concentrations, and NEA rates also differed between plant communities; the values for the mesic meadow were at least four times higher than the other plant communities, and the snowbeds formed an intermediate group. Moreover, the results show a different pattern of distribution for individual amino acids across the plant communities, with snowbeds tending to have the highest amino acid N concentrations. The differences between plant communities along this natural gradient also illustrate variations between the dominant mycorrhizal associ- ations in facilitating N capture by the characteristic functional groups of plants.

The ecosystems of alpine snowbed habitats are reviewed with emphasis on ecosystem functioning and capability to adapt to current and predicted global change. Snowbeds form in topographic depressions that accumulate large amounts of snow during the winter months, and the final snowmelt does not occur until late in the growing season. Many species preferentially grow in snowbed habitats and some of these are even restricted to these habitats. In this review we identify several ecosystem services which snowbeds provide to the alpine landscape. For instance, snowbeds provide a steady water and nutrient supply to adjacent plant communities and offer newly emerged high-quality food for herbivores late in the growing season. We also propose that alpine snowbeds are much more productive than earlier thought, especially when the very short growing season and often high grazing pressure are taken fully into account. Furthermore, we propose that bryophytes and graminoids (grasses, sedges, and rushes) probab...

Effects of simulated environmental change on bryophyte and lichen species richness and diversity in alpine tundra were investigated in a 5-year experiment at Latnjajaure, northern Sweden. The experiment had a factorial design including fertilisation and temperature enhancement in one meadow and one heath plant community. Responses in species richness, biodiversity, and species composition of bryophytes and lichens to experimental treatments were compared to the observed variation in six naturally occurring plant communities. The combination of fertilisation and enhanced temperature resulted in a species impoverishment, for bryophytes in the bryophyte-dominated community, and for lichens in the lichen-dominated communities, but the species composition stayed within the observed natural variation. During the course of the study, no species new to the investigated mid-alpine landscape were recorded, but that scenario is realistic within a decade when comparing with the processes seen in vascular plants.

Plant communities in Arctic and alpine areas are changing due to higher temperatures and longer vegetation periods and it is uncertain how this will affect plant-herbivore dynamics. For instance, relatively fast-growing, deciduous shrub species that are the most responsive to warming may also be the most targeted by herbivores such as reindeer, giving less palatable evergreen shrubs the chance to expand. Using herbivore exclosures, we have studied how two grasslands with contrasting nutrient and moisture regimes, a dry, nutrient-poor alpine grass heath and a wet, productive low herb meadow, changed between 1995 and 2012, in grazed and ungrazed conditions. At the grass heath, evergreen low shrub abundance had more than doubled, regardless of grazer treatment, whereas at the low herb meadow, evergreen shrubs had increased only outside exclosures while deciduous tall shrubs and forbs were significantly more abundant inside exclosures. Deciduous tall shrubs were also significantly taller in exclosures. These contrasting findings suggest that the impact of herbivores is to a great deal determined by their influence on competitive interactions between plant species, and therefore depends on the underlying composition of the plant community. Consequently, as the balance in these competitive interactions is shifting due to climate warming, we conclude that the potential of herbivory to influence this balance is considerable yet highly site dependent.

International audience; Interactions among species determine local-scale diversity, but local interactions are thought to have minor effects at larger scales. However, quantitative comparisons of the importance of biotic interactions relative to other drivers are rarely made at larger scales. Using a data set spanning 78 sites and five continents, we assessed the relative importance of biotic interactions and climate in determining plant diversity in alpine ecosystems dominated by nurse-plant cushion species. Climate variables related with water balance showed the highest correlation with richness at the global scale. Strikingly, although the effect of cushion species on diversity was lower than that of climate, its contribution was still substantial. In particular, cushion species enhanced species richness more in systems with inherently impoverished local diversity. Nurse species appear to act as a ‘safety net’ sustaining diversity under harsh conditions, demonstrating that climate and species interactions should be integrated when predicting future biodiversity effects of climate change.

Mycorrhizal fungi constitute a considerable sink for carbon in most ecosystems. This carbon is used for building extensive mycelial networks in the soil as well as for metabolic activity related to nutrient uptake. A number of methods have been developed recently to quantify production, standing biomass and turnover of extramatrical mycorrhizal mycelia (EMM) in the field. These methods include minirhizotrons, in-growth mesh bags and cores, and indirect measurements of EMM based on classification of ectomycorrhizal fungi into exploration types. Here we review the state of the art of this methodology and discuss how it can be developed and applied most effectively in the field, Furthermore, we also discuss different ways to quantify fungal biomass based on biomarkers such as chitin, ergosterol and PLFAs, as well as molecular methods, such as qPCR. The evidence thus far indicates that mycorrhizal fungi are key components of microbial biomass in many ecosystems. We highlight the need to extend the application of current methods to focus on a greater range of habitats and mycorrhizal types enabling incorporation of mycorrhizal fungal biomass and turnover into biogeochemical cycling models. (C) 2012 Elsevier Ltd. All rights reserved.

[b][u]Background and Aims[/u][/b][br/] [br/] Forest trees directly contribute to carbon cycling in forest soils through the turnover of their fine roots. In this study we aimed to calculate root turnover rates of common European forest tree species and to compare them with most frequently published values.[br/] [br/] [b][u]Methods[/u][/b][br/] [br/] We compiled available European data and applied various turnover rate calculation methods to the resulting database. We used Decision Matrix and Maximum-Minimum formula as suggested in the literature.[br/] [br/] [b][u]Results[/u][/b][br/] [br/] Mean turnover rates obtained by the combination of sequential coring and Decision Matrix were 0.86 yr−1 for Fagus sylvatica and 0.88 yr−1 for Picea abies when maximum biomass data were used for the calculation, and 1.11 yr−1 for both species when mean biomass data were used. Using mean biomass rather than maximum resulted in about 30 % higher values of root turnover. Using the Decision Matrix to calculate turnover rate doubled the rates when compared to the Maximum-Minimum formula. The Decision Matrix, however, makes use of more input information than the Maximum-Minimum formula.[br/] [br/] [b][u]Conclusions[/u][/b][br/] [br/] We propose that calculations using the Decision Matrix with mean biomass give the most reliable estimates of root turnover rates in European forests and should preferentially be used in models and C reporting.

There is growing evidence of the importance of extramatrical mycelium (EMM) of mycorrhizal fungi in carbon (C) cycling in ecosystems. However, our understanding has until recently been mainly based on laboratory experiments, and knowledge of such basic parameters as variations in mycelial production, standing biomass and turnover as well as the regulatory mechanisms behind such variations in forest soils is limited. Presently, the production of EMM by ectomycorrhizal (EM) fungi has been estimated at ~140 different forest sites to be up to several hundreds of kg per ha per year, but the published data are biased towards Picea abies in Scandinavia. Little is known about the standing biomass and turnover of EMM in other systems, and its influence on the C stored or lost from soils. Here, focussing on ectomycorrhizas, we discuss the factors that regulate the production and turnover of EMM and its role in soil C dynamics, identifying important gaps in this knowledge. C availability seems to be the key factor determining EMM production and possibly its standing biomass in forests but direct effects of mineral nutrient availability on the EMM can be important. There is great uncertainty about the rate of turnover of EMM. There is increasing evidence that residues of EM fungi play a major role in the formation of stable N and C in SOM, which highlights the need to include mycorrhizal effects in models of global soil C stores. Skupno št. avtorjev: 14. Bibliografija: str. 21-27.

The winter CO(2) efflux from subnivean environments is an important component of annual C budgets in Arctic ecosystems and consequently makes prediction and estimations of winter processes as well ...

The effects of wood ash application on soil microbial processes were investigated in three drained forested peatlands, which differed in nutrient status and time since application. Measured variables included the concentrations of soil elements and phospholipid fatty acids (PLFAs), net nitrogen (N) mineralization, nitrification and denitrification enzyme activity, potential methane (CH(4)) oxidation, CH(4) production and microbial respiration kinetics. Wood ash application had a considerable influence on soil element concentrations. This mirrored a decrease in the majority of the microbial biomarkers by more than one-third in the two oligotrophic peatlands, although the microbial community composition was not altered. The decreases in PLFAs coincided with reduced net ammonification and net N mineralization. Other measured variables did not change systematically as a result of wood ash application. No significant changes in microbial biomass or processes were found in the mesotrophic peatland, possibly because too little time (1 year) had elapsed since the wood ash application. This study suggests that oligotrophic peatlands can be substantially affected by wood ash for a period of at least 4 years after application. However, within 25 years of the wood ash application, the microbial biomass seemed to have recovered or adapted to enhanced element concentrations in the soil.

The effects of wood ash application on soil microbial processes were investigated in three drained forested peatlands, which differed in nutrient status and time since application. Measured variables included the concentrations of soil elements and phospholipid fatty acids (PLFAs), net nitrogen (N) mineralization, nitrification and denitrification enzyme activity, potential methane (CH(4)) oxidation, CH(4) production and microbial respiration kinetics. Wood ash application had a considerable influence on soil element concentrations. This mirrored a decrease in the majority of the microbial biomarkers by more than one-third in the two oligotrophic peatlands, although the microbial community composition was not altered. The decreases in PLFAs coincided with reduced net ammonification and net N mineralization. Other measured variables did not change systematically as a result of wood ash application. No significant changes in microbial biomass or processes were found in the mesotrophic peatland, possibly because too little time (1 year) had elapsed since the wood ash application. This study suggests that oligotrophic peatlands can be substantially affected by wood ash for a period of at least 4 years after application. However, within 25 years of the wood ash application, the microbial biomass seemed to have recovered or adapted to enhanced element concentrations in the soil.

Biodiversity is a complex concept that covers diversity in a continuum of organisational levels from genetic diversity within populations to functional diversity at a global scale. The scale problem is central, and organism groups reach their maximum diversity (taxonomic richness) per unit area at very different geographical scales. Cryptogams and soil invertebrates may have high genetic and taxonomic diversity at a patch size of

That reindeer grazing can have large effects on plant communities is well known, but how reindeer grazing affects plant traits and plant carbon (C) and nitrogen (N) allocation has not been studied to the same extent. This study was conducted in a sub-arctic dry heath in northern Sweden. 17-year-old reindeer exclosures were used to test whether reindeer grazing affects the C:N ratio (a plant quality index), and the δ13C and δ15N (indicators of changes in C and N dynamics) as well as the C and N content of above- and below ground parts of the evergreen dwarf shrub Vaccinium vitis-idaea L. and the perennial forb Bistorta vivipara (L.) Gray. A lower C:N ratio was found in B. vivipara compared to V. vitis-idaea suggesting a higher grazing pressure on that species. We found that grazing reduced the total C content, by 26%, and increased the δ15N, by 1‰, in the leaves of B. vivipara, while no changes were observed in V. vitis-idaea. Fine roots of B. vivipara had higher δ13C (1‰) and δ15N (2.5‰) than the leaves, while such differences were not found in V. vitis-idaea. The results also highlight the importance of analysing both above- and belowground plant parts when interpreting natural variations in δ13C and δ15N.

International audience; Foundation species can change plant community structure by modulating important ecological processes such as community assembly, yet this topic is poorly understood. In alpine systems, cushion plants commonly act as foundation species by ameliorating local conditions. Here, we analyze diversity patterns of species' assembly within cushions and in adjacent surrounding open substrates (83 sites across five continents) calculating floristic dissimilarity between replicate plots, and using linear models to analyze relationships between microhabitats and species diversity. Floristic dissimilarity did not change across biogeographic regions, but was consistently lower in the cushions than in the open microhabitat. Cushion plants appear to enable recruitment of many relatively stress-intolerant species that otherwise would not establish in these communities, yet the niche space constructed by cushion plants supports a more homogeneous composition of species than the niche space beyond the cushion's influence. As a result, cushion plants support higher α-diversity and a larger species pool, but harbor assemblies with lower ?-diversity than open microhabitats. We conclude that habitats with and without dominant foundation species can strongly differ in the processes that drive species recruitment, and thus the relationship between local and regional species diversity.