Climatic and biotic extreme events moderate long-term responses of above- and belowground sub-Arctic heathland communities to climate change.

Climate change impacts are not uniform across the Arctic region because interacting factors causes large variations in local ecosystem change. Extreme climatic events and population cycles of herbivores occur simultaneously against a background of gradual climate warming trends and can redirect ecosystem change along routes that are difficult to predict. Here, we present the results from sub-Arctic heath vegetation and its belowground micro-arthropod community in response to the two main drivers of vegetation damage in this region: extreme winter warming events and subsequent outbreaks of the defoliating autumnal moth caterpillar (Epirrita autumnata). Evergreen dwarf shrub biomass decreased (30%) following extreme winter warming events and again by moth caterpillar grazing. Deciduous shrubs that were previously exposed to an extreme winter warming event were not affected by the moth caterpillar grazing, while those that were not exposed to warming events (control plots) showed reduced (23%) biomass from grazing. Cryptogam cover increased irrespective of grazing or winter warming events. Micro-arthropods declined (46%) following winter warming but did not respond to changes in plant community. Extreme winter warming and caterpillar grazing suppressed the CO2 fluxes of the ecosystem. Evergreen dwarf shrubs are disadvantaged in a future sub-Arctic with more stochastic climatic and biotic events. Given that summer warming may further benefit deciduous over evergreen shrubs, event and trend climate change may both act against evergreen shrubs and the ecosystem functions they provide. This is of particular concern given that Arctic heath vegetation is typically dominated by evergreen shrubs. Other components of the vegetation showed variable responses to abiotic and biotic events, and their interaction indicates that sub-Arctic vegetation response to multiple pressures is not easy to predict from single-factor responses. Therefore, while biotic and climatic events may have clear impacts, more work is needed to understand their net effect on Arctic ecosystems.

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

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

Experimental reduction of food quality is not compensated with increased food intake in high-Arctic muskoxen (Ovibos moschatus).

A total of four barren adult female muskoxen (Ovibos moschatus) were used over a period of 2 years for the purpose of the present study. During the first year, the natural changes in appetite (ad libitum intake of standard pelleted reindeer feed) and body mass were determined in two of the animals. During the second year, the effect of reduced food quality on ad libitum food intake was tested in all four animals in July when the appetite had been found to be at a high. We found that the experimentally reduced food quality was not compensated with increased food intake in these large high-Arctic herbivores.

Impacts of winter icing events on the growth, phenology and physiology of sub-arctic dwarf shrubs.

The Arctic is experiencing the greatest climate change in winter, including increases in freeze-thaw cycles that can result in ice encasement of vegetation. Ice encasement can expose plants to hypoxia and greater temperature extremes, but currently the impacts of icing on plants in the field remain little understood. With this in mind, a unique field manipulation experiment was established in heathland in northern Sweden with ice encasement simulated in early March 2008, 2009 and 2010 until natural thaw each spring. In the following summers we assessed the impacts on flowering, bud phenology, shoot growth and mortality and leaf damage (measured by chlorophyll fluorescence and electrolyte leakage) of the three dominant dwarf shrub species Empetrum nigrum, Vaccinium vitis-idaea (both evergreen) and Vaccinium myrtillus (deciduous). Two consecutive winters of icing decreased V. vitis-idaea flowering by 57%, while flowering of V. myrtillus and E. nigrum remained unaffected. Vaccinium myrtillus showed earlier budburst but shoot growth for all species was unchanged. Shoot mortality of V. myrtillus and V. vitis-idaea increased after the first year (by 70 and 165%, respectively) and again for V. myrtillus following the third year (by 67%), while E. nigrum shoot mortality remained unaffected, as were chlorophyll fluorescence and electrolyte leakage in all species. Overall, the sub-arctic heathland was relatively tolerant to icing, but the considerable shoot mortality of V. myrtillus contrasting with the general tolerance of E. nigrum suggests plant community structure in the longer term could change if winters continue to see a greater frequency of icing events.

Environmental monitoring and research in the Abisko area-an overview.

This article gives an overview of the studies on the environment surrounding the Abisko Scientific Research Station in Swedish Lapland. The long-term monitoring of the Station on processes related to the climate, and to the physical, biotic, and chemical environmental conditions is particularly addressed. Some variables are recorded since more than 100 years. The obtained data in combination with results from short-term studies and manipulation experiments are important to understand past and future conditions of the ecosystems. This has practical applications for the planning of tourism, transports, reindeer herding, and for societal purposes.

Linking fine-scale sub-arctic vegetation distribution in complex topography with surface-air-temperature modelled at 50-m resolution.

Recent studies have shown that the complexities of the surface features in mountainous terrain require a re-assessment of climate impacts at the local level. We explored the importance of surface-air-temperature based on a recently published 50-m-gridded dataset, versus soil variables for explaining vegetation distribution in Swedish Lapland using generalised linear models (GLMs). The results demonstrated that the current distribution of the birch forest and snowbed community strongly relied on the surface-air-temperature. However, temperature alone is a poor predictor of many plant communities (wetland, meadow). Because of diminishing sample representation with increasing altitude, the snowbed community was under-sampled at higher altitudes. This results in underestimation of the current distribution of the snowbed community around the mountain summits. The analysis suggests that caution is warranted when applying GLMs at the local level.

Changes versus homeostasis in alpine and sub-alpine vegetation over three decades in the sub-arctic.

Plant species distributions are expected to shift and diversity is expected to decline as a result of global climate change, particularly in the Arctic where climate warming is amplified. We have recorded the changes in richness and abundance of vascular plants at Abisko, sub-Arctic Sweden, by re-sampling five studies consisting of seven datasets; one in the mountain birch forest and six at open sites. The oldest study was initiated in 1977-1979 and the latest in 1992. Total species number increased at all sites except for the birch forest site where richness decreased. We found no general pattern in how composition of vascular plants has changed over time. Three species, Calamagrostis lapponica, Carex vaginata and Salix reticulata, showed an overall increase in cover/frequency, while two Equisetum taxa decreased. Instead, we showed that the magnitude and direction of changes in species richness and composition differ among sites.

Enhanced UV-B and elevated CO(2) impacts sub-arctic shrub berry abundance, quality and seed germination.

This study investigated the effects of long-term-enhanced UV-B, and combined UV-B with elevated CO(2) on dwarf shrub berry characteristics in a sub-arctic heath community. Germination of Vaccinium myrtillus was enhanced in seeds produced at elevated UV-B, but seed numbers and berry size were unaffected. Elevated UV-B and CO(2) stimulated the abundance of V. myrtillus berries, whilst UV-B alone stimulated the berry abundance of V. vitis-idaea and Empetrum hermaphroditum. Enhanced UV-B reduced concentrations of several polyphenolics in V. myrtillus berries, whilst elevated CO(2) increased quercetin glycosides in V. myrtillus, and syringetin glycosides and anthocyanins in E. hermaphroditum berries. UV-B × CO(2) interactions were found for total anthocyanins, delphinidin-3-hexoside and peonidin-3-pentosidein in V. myrtillus berries but not E. hermaphroditum. Results suggest positive impacts of UV-B on the germination of V. myrtillus and species-specific impacts of UV-B × elevated CO(2) on berry abundance and quality. The findings have relevance and implications for human and animal consumers plus seed dispersal and seedling establishment.

The need to understand the stability of arctic vegetation during rapid climate change: An assessment of imbalance in the literature.

In early studies, northern vegetation response to global warming recognised both increases in biomass/cover and shrinking of species' distributional ranges. Subsequent field measurements focussed on vegetation cover and biomass increases ("greening"), and more recently decreases ("browning"). However, satellite observations show that more than 50% of arctic vegetation has not changed significantly despite rapid warming. While absence of change in remote sensing data does not necessarily mean no ecological change on the ground, the significant proportion of the Arctic that appears to be stable in the face of considerable climate change points to a greater need to understand Arctic ecosystem stability. In this paper, we performed an extensive review of the available literature to seek balances or imbalances between research focussing on "greening", "browning" and "stability/no change". We find that greening studies dominate the literature though two relatively small areas of the Arctic are disproportionately represented for this main change process. Critically, there are too few studies anywhere investigating stability. We highlight the need to understand the mechanisms driving Arctic ecosystem stability, and the potential longer-term consequences of remaining stable in a rapidly changing climate.

Multi-decadal changes in tundra environments and ecosystems: the International Polar Year-Back to the Future Project (IPY-BTF).

Polar and alpine environments are changing rapidly due to increases in temperature, which are amplified in the Arctic, as well as changes in many local factors. The impacts on ecosystems and their function have potential consequences for local residents and the global community. Tundra areas are vast and diverse, and the knowledge of geographical variation in environmental and ecosystem change is limited to relatively few locations, or to remote sensing approaches that are limited mostly to the past few decades. The International Polar Year, IPY, provided a context, stimulus and timely opportunities for re-visiting old research sites and data sets to collate data on past changes, to pass knowledge from old to new generations of researchers and to document environmental characteristics of sites to facilitate detection and attribution of future changes. Consequently, the project "Retrospective and Prospective Vegetation Change in the Polar Regions: Back to the Future," BTF, was proposed and endorsed as an IPY activity (project #512). With national funding support, teams of researchers re-visited former sites and data sets throughout the Arctic and some alpine regions. These efforts have amounted to a gamut of "BTF" studies that are collectively geographically expansive and disciplinary diverse. A selection of these studies are introduced and presented in the current issue together with a brief synthesis of their findings.

Multi-decadal changes in snow characteristics in sub-Arctic Sweden.

A unique long term, 49-year record (divided into three time periods 1961-1976, 1977-1992, and 1993-2009) of snow profile stratigraphy from the Swedish sub Arctic, was analyzed with a focus on changes in snow characteristics. The data set contained grain size, snow layer hardness, grain compactness, and snow layer dryness, observed every second week during the winter season. The results showed an increase in very hard snow layers, with harder snow in early winter and more moist snow during spring. There was a striking increase in the number of observations with very hard snow at ground level over time. More than twice as many occasions with hard snow at ground level were observed between 1993 and 2009 compared to previous years, which may have a significant effect on plants and animals. The changes in snow characteristics are most likely a result of the increasing temperatures during the start and the end of the snow season.

Plant and vegetation dynamics on Disko Island, west Greenland: snapshots separated by over 40 years.

We report on a revisit in 2009 to sites where vegetation was recorded in 1967 and 1970 on Disko Island, West Greenland. Re-sampling of the same clones of the grass Phleum alpinum after 39 years showed complete stability in biometrics but dramatic earlier onset of various phenological stages that were not related to changes in population density. In a fell-field community, there was a net species loss, but in a herb-slope community, species losses balanced those that were gained. The type of species establishing and increasing in frequency and/or cover abundance at the fell-field site, particularly prostrate dwarf shrubs, indicates a possible start of a shift towards a heath, rather than a fell-field community. At the herb-slope site, those species that established or increased markedly in frequency and/or cover abundance indicate a change to drier conditions. This is confirmed both by the decrease in abundance of Alchemilla glomerulans and Epilobium hornemanii, and the drying of a nearby pond. The causes of these changes are unknown, although mean annual temperature has risen since 1984.

Past and present permafrost temperatures in the Abisko area: redrilling of boreholes.

Monitoring of permafrost has been ongoing since 1978 in the Abisko area, northernmost Sweden, when measurements of active layer thickness started. In 1980, boreholes were drilled in three mires in the area to record permafrost temperatures. Recordings were made twice per year, and the last data were obtained in 2002. During the International Polar Year (2007-2008), new boreholes were drilled within the 'Back to the Future' (BTF) and 'Thermal State of Permafrost' (TSP) projects that enabled year-round temperature monitoring. Mean annual ground temperatures (MAGT) in the mires are close to 0 degrees C, ranging from -0.16 to -0.47 degrees C at 5 m depth. Data from the boreholes show increasing ground temperatures in the upper and lower part by 0.4 to 1 degree C between 1980 and 2002. At one mire, permafrost thickness has decreased from 15 m in 1980 to ca. 9 m in 2009, with an accelerating thawing trend during the last decade.

Changes in tree growth, biomass and vegetation over a 13-year period in the Swedish sub-Arctic.

This study was conducted in the Swedish subArctic, near Abisko, in order to assess the direction and scale of possible vegetation changes in the alpine-birch forest ecotone. We have re-surveyed shrub, tree and vegetation data at 549 plots grouped into 61 clusters. The plots were originally surveyed in 1997 and re-surveyed in 2010. Our study is unique for the area as we have quantitatively estimated a 19% increase in tree biomass mainly within the existing birch forest. We also found significant increases in the cover of two vegetation types--"birch forest-heath with mosses" and "meadow with low herbs", while the cover of snowbed vegetation decreased significantly. The vegetation changes might be caused by climate, herbivory and past human impact but irrespective of the causes, the observed transition of the vegetation will have substantial effects on the mountain ecosystems.

Tree and shrub expansion over the past 34 years at the tree-line near Abisko, Sweden.

Shrubs and trees are expected to expand in the sub-Arctic due to global warming. Our study was conducted in Abisko, sub-arctic Sweden. We recorded the change in coverage of shrub and tree species over a 32- to 34-year period, in three 50 x 50 m plots; in the alpine-tree-line ecotone. The cover of shrubs and trees (<3.5 cm diameter at breast height) were estimated during 2009-2010 and compared with historical documentation from 1976 to 1977. Similarly, all tree stems (> or =3.5 cm) were noted and positions determined. There has been a substantial increase of cover of shrubs and trees, particularly dwarf birch (Betula nana), and mountain birch (Betula pubescens ssp. czerepanovii), and an establishment of aspen (Populus tremula). The other species willows (Salix spp.), juniper (Juniperus communis), and rowan (Sorbus aucuparia) revealed inconsistent changes among the plots. Although this study was unable to identify the causes for the change in shrubs and small trees, they are consistent with anticipated changes due to climate change and reduced herbivory.

Multi-decadal changes in tundra environments and ecosystems: synthesis of the International Polar Year-Back to the Future project (IPY-BTF).

Understanding the responses of tundra systems to global change has global implications. Most tundra regions lack sustained environmental monitoring and one of the only ways to document multi-decadal change is to resample historic research sites. The International Polar Year (IPY) provided a unique opportunity for such research through the Back to the Future (BTF) project (IPY project #512). This article synthesizes the results from 13 papers within this Ambio Special Issue. Abiotic changes include glacial recession in the Altai Mountains, Russia; increased snow depth and hardness, permafrost warming, and increased growing season length in sub-arctic Sweden; drying of ponds in Greenland; increased nutrient availability in Alaskan tundra ponds, and warming at most locations studied. Biotic changes ranged from relatively minor plant community change at two sites in Greenland to moderate change in the Yukon, and to dramatic increases in shrub and tree density on Herschel Island, and in subarctic Sweden. The population of geese tripled at one site in northeast Greenland where biomass in non-grazed plots doubled. A model parameterized using results from a BTF study forecasts substantial declines in all snowbeds and increases in shrub tundra on Niwot Ridge, Colorado over the next century. In general, results support and provide improved capacities for validating experimental manipulation, remote sensing, and modeling studies.

Siberian environmental change: Synthesis of recent studies and opportunities for networking.

A recent multidisciplinary compilation of studies on changes in the Siberian environment details how climate is changing faster than most places on Earth with exceptional warming in the north and increased aridity in the south. Impacts of these changes are rapid permafrost thaw and melt of glaciers, increased flooding, extreme weather events leading to sudden changes in biodiversity, increased forest fires, more insect pest outbreaks, and increased emissions of CO2 and methane. These trends interact with sociological changes leading to land-use change, globalisation of diets, impaired health of Arctic Peoples, and challenges for transport. Local mitigation and adaptation measures are likely to be limited by a range of public perceptions of climate change that vary according to personal background. However, Siberia has the possibility through land surface feedbacks to amplify or suppress climate change impacts at potentially global levels. Based on the diverse studies presented in this Ambio Special Issue, we suggest ways forward for more sustainable environmental research and management.

South-Siberian mountain mires: Perspectives on a potentially vulnerable remote source of biodiversity.

Changes in climate, land-use and pollution are having disproportionate impacts on ecosystems and biodiversity of arctic and mountain ecosystems. While these impacts are well-documented for many areas of the Arctic and alpine regions, some isolated and inaccessible mountain areas are poorly studied. Furthermore, even in well-studied regions, assessments of biodiversity and species responses to environmental change are biased towards vascular plants and cryptogams, particularly bryophytes are far less represented. This paper aims to document the environments of the remote and inaccessible Altai-Sayan mountain mires and particularly their bryofloras where threatened species exist and species new to the regional flora are still being found. As these mountain mires are relatively inaccessible, changes in drivers of change and their ecosystem and biodiversity impacts have not been monitored. However, the remoteness of the mires has so far protected them and their species. In this study, we describe the mires, their bryophyte species and the expected impacts of environmental stressors to bring attention to the urgency of documenting change and conserving these pristine ecosystems.

The dynamic land-cover of the Altai Mountains: Perspectives based on past and current environmental and biodiversity changes.

We present climate-dependent changes in the high-mountain forest ecotone, old-growth forests, alpine phytocenoses, and deglaciated forelands in the Aktru glacial basin (Altai Republic, Russia). A number of independent sources (variations in upper treeline altitude, dendrochronological data, analysis of lacustrine sediments and botanical and geographical studies linked with the dynamics of glacial-dammed lakes in the Chuya and Kurai intermountain depressions) suggest Holocene temperatures reached about 4 °C higher than today. Unlike the European Alps, glaciers in the continental Altai Mountains disappeared before forming again. Also, the upper altitudinal limit of mountain forests during the Holocene was greater than in the European Alps. The high variability of mountain ecosystems in southern Siberia suggests their potential instability in a currently changing climate. However, periglacial successions associated with the strong continental climate and glacier retreat represent an area of increasing biodiversity and plant cover. The historical and current sensitivity of the continental mountains to climate variations which exceeds that of the European Alps requires greater understanding, environmental protection, and increased social responsibility for the consequences of anthropogenic contributions to climate change: the isolated Altai areas contribute little to climate changes, but are greatly affected by them.

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

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