20th century climate warming and tree-limit rise in the southern Scandes of Sweden

Climate warming by ca. 0.8 degree C between the late-19th and late-20th century, although with some fluctuations, has forced multispecies elevational tree-limit advance by > 100 m for the principal tree species in the Swedish part of the Scandinavian mountain range. Predominantly, these processes imply growth in height of old-established individuals and less frequently upslope migration of new individuals. After a slight retardation during some cooler decades after 1940, a new active phase of tree-limit advance has occurred with a series of exceptionally mild winters and some warm summers during the 1990s. The magnitude of total 20th century tree-limit rise varies with topoclimate and is mainly confined to wind-sheltered and snow-rich segments of the landscape. Thickening of birch tree stands in the "advance belt" has profoundly altered the general character of the subalpine/low alpine landscape and provides a positive feedback loop for further progressive change and resilience to short-term cooling episodes. All upslope tree-limit shifts and associated landscape transformations during the 20th century have occurred without appreciable time lags, which constitutes knowledge fundamental to the generation of realistic models concerning vegetation responses to potential future warming. The new and elevated pine tree-limit may be the highest during the past 4000 14C years. Thus, it is tentatively inferred that the 20th century climate is unusually warm in a late-Holocene perspective.     

A boreal invasion in response to climate change? Range shifts and community effects in the borderland between forest and tundra

It has been hypothesized that climate warming will allow southern species to advance north and invade northern ecosystems. We review the changes in the Swedish mammal and bird community in boreal forest and alpine tundra since the nineteenth century, as well as suggested drivers of change. Observed changes include (1) range expansion and increased abundance in southern birds, ungulates, and carnivores; (2) range contraction and decline in northern birds and carnivores; and (3) abundance decline or periodically disrupted dynamics in cyclic populations of small and medium-sized mammals and birds. The first warm spell, 1930–1960, stands out as a period of substantial faunal change. However, in addition to climate warming, suggested drivers of change include land use and other anthropogenic factors. We hypothesize all these drivers interacted, primarily favoring southern generalists. Future research should aim to distinguish between effects of climate and land-use change in boreal and tundra ecosystems.     

Flora and Vegetation of Tasiilaq,Formerly Angmagssalik,Southeast Greenland: A Comparison of Data Between Around 1900 and 2007

The changes in the vascular plant flora of Tasiilaq, low arctic Southeast Greenland, between around 1900 and 2007 were studied by comparing the data from historical literature with those of the field observations performed between the late 1960s and 2007. Since 1900, the percentage of widely distributed arctic species distinctly decreased, whereas that of the low arctic species somewhat increased, and boreal species hardly increased. Vegetation monitoring revealed minor changes and showed that several thermophilous and xerophilous species increased between 1968/1969 and 2007, whereas some hygrophilous species decreased. Repeated vegetation mapping of a shallow pond revealed conspicuous changes suggesting increased evaporation/precipitation ratios associated with environmental warming up and decreasing snow accumulation in winter, in line with results of previous investigations. In spite of climate warming, expansion of the town and increasing human impact, flora and vegetation on the whole appeared rather stable during the last 40 years without invading species or introductions.     

Status and trends in Arctic vegetation: Evidence from experimental warming and long-term monitoring

Changes in Arctic vegetation can have important implications for trophic interactions and ecosystem functioning leading to climate feedbacks. Plot-based vegetation surveys provide detailed insight into vegetation changes at sites around the Arctic and improve our ability to predict the impacts of environmental change on tundra ecosystems. Here, we review studies of changes in plant community composition and phenology from both long-term monitoring and warming experiments in Arctic environments. We find that Arctic plant communities and species are generally sensitive to warming, but trends over a period of time are heterogeneous and complex and do not always mirror expectations based on responses to experimental manipulations. Our findings highlight the need for more geographically widespread, integrated, and comprehensive monitoring efforts that can better resolve the interacting effects of warming and other local and regional ecological factors.     

Spitsbergen landscape under 20th century climate change: Sørkapp Land

A reaction of the European Arctic landscape to a climate change on the scale of a typical middle-sized region is outlined. A wide scope of the methods was used, first of all field mapping and observations. Glaciers are important in the S?rkapp Land landscape because they cover the majority of its territory and undergo quick recessions as a result of the 20th century warming. Glacial recession influence intensively: relief with Quaternary deposits, waters, animals, vegetation and soils. The most important landscape changes in the 20th century are: uplift of the equilibrium line altitude on glaciers by 100-200 m; large glacial recession in both surface and volume; significant decrease of the land area due to recession of tidewater glaciers; lengthening of the coastline, and especially of glacial cliffs; development of the land water network; start of the plant succession in areas abandoned by glaciers. No isostatic uplift has taken place in S?rkapp Land since the Little Ice Age.     

Synthesis of effects in four Arctic subregions

An assessment of impacts on Arctic terrestrial ecosystems has emphasized geographical variability in responses of species and ecosystems to environmental change. This variability is usually associated with north-south gradients in climate, biodiversity, vegetation zones, and ecosystem structure and function. It is clear, however, that significant east-west variability in environment, ecosystem structure and function, environmental history, and recent climate variability is also important. Some areas have cooled while others have become warmer. Also, east-west differences between geographical barriers of oceans, archipelagos and mountains have contributed significantly in the past to the ability of species and vegetation zones to relocate in response to climate changes, and they have created the isolation necessary for genetic differentiation of populations and biodiversity hot-spots to occur. These barriers will also affect the ability of species to relocate during projected future warming. To include this east-west variability and also to strike a balance between overgeneralization and overspecialization, the ACIA identified four major sub regions based on large-scale differences in weather and climate-shaping factors. Drawing on information, mostly model output that can be related to the four ACIA subregions, it is evident that geographical barriers to species re-location, particularly the distribution of landmasses and separation by seas, will affect the northwards shift in vegetation zones. The geographical constraints--or facilitation--of northward movement of vegetation zones will affect the future storage and release of carbon, and the exchange of energy and water between biosphere and atmosphere. In addition, differences in the ability of vegetation zones to re-locate will affect the biodiversity associated with each zone while the number of species threatened by climate change varies greatly between subregions with a significant hot-spot in Beringia. Overall, the subregional synthesis demonstrates the difficulty of generalizing projections of responses of ecosystem structure and function, species loss, and biospheric feedbacks to the climate system for the whole Arctic region and implies a need for a far greater understanding of the spatial variability in the responses of terrestrial arctic ecosystems to climate change.     

A reflection on four impactful Ambio papers: The biotic perspective: This article belongs to Ambio’s 50th Anniversary Collection. Theme: Climate change impacts

Climate change represents one of the most pressing societal and scientific challenges of our time. While much of the current research on climate change focuses on future prediction, some of the strongest signals of warming can already be seen in Arctic and alpine areas, where temperatures are rising faster than the global average, and in the oceans, where the combination of rising temperatures and acidification due to increased CO₂ concentrations has had catastrophic consequences for sensitive marine organisms inhabiting coral reefs. The scientific papers highlighted as part of this anniversary issue represent some of the most impactful advances in our understanding of the consequences of anthropogenic climate change. Here, we reflect on the legacy of these papers from the biotic perspective.     

Using Available Information to Assess the Potential Effects of Climate Change on Vegetation in the High Arctic: North Billjefjorden, Central Spitsbergen (Svalbard)

We review the available data that can be used to assess the potential impact of climate change on vegetation, and we use central Spitsbergen, Svalbard, as a model location for the High Arctic. We used two sources of information: recent and short-term historical records, which enable assessment on scales of particular plant communities and the landscape over a period of decades, and palynological and macrofossil analyses, which enable assessment on time scales of hundreds and thousands of years and on the spatial scale of the landscape. Both of these substitutes for standardized monitoring revealed stability of vegetation, which is probably attributable to the harsh conditions and the distance of the area from sources of diaspores of potential new incomers. The only evident recent vegetation changes related to climate change are associated with succession after glacial retreats. By establishing a network of permanent plots, researchers will be able to monitor immigration of new species from diversity 'hot spots' and from an abandoned settlement nearby. This will greatly enhance our ability to understand the effects of climate change on vegetation in the High Arctic.     

Ecosystem Response to Climatic Change: The Importance of the Cold Season

Winter climate and snow cover are the important drivers of plant community development in polar regions. However, the impacts of changing winter climate and associated changes in snow regime have received much less attention than changes during summer. Here, we synthesize the results from studies on the impacts of extreme winter weather events on polar heathland and lichen communities. Dwarf shrubs, mosses and soil arthropods were negatively impacted by extreme warming events while lichens showed variable responses to changes in extreme winter weather events. Snow mould formation underneath the snow may contribute to spatial heterogeneity in plant growth, arthropod communities and carbon cycling. Winter snow cover and depth will drive the reported impacts of winter climate change and add to spatial patterns in vegetation heterogeneity. The challenges ahead lie in obtaining better predictions on the snow patterns across the landscape and how these will be altered due to winter climate change.     

Climate change,phenology, and butterfly host plant utilization

Knowledge of how species interactions are influenced by climate warming is paramount to understand current biodiversity changes. We review phenological changes of Swedish butterflies during the latest decades and explore potential climate effects on butterfly–host plant interactions using the Orange tip butterfly Anthocharis cardamines and its host plants as a model system. This butterfly has advanced its appearance dates substantially, and its mean flight date shows a positive correlation with latitude. We show that there is a large latitudinal variation in host use and that butterfly populations select plant individuals based on their flowering phenology. We conclude that A. cardamines is a phenological specialist but a host species generalist. This implies that thermal plasticity for spring development influences host utilization of the butterfly through effects on the phenological matching with its host plants. However, the host utilization strategy of A. cardamines appears to render it resilient to relatively large variation in climate.     

Ecosystem response to climatic change: the importance of the cold season

Winter climate and snow cover are the important drivers of plant community development in polar regions. However, the impacts of changing winter climate and associated changes in snow regime have received much less attention than changes during summer. Here, we synthesize the results from studies on the impacts of extreme winter weather events on polar heathland and lichen communities. Dwarf shrubs, mosses and soil arthropods were negatively impacted by extreme warming events while lichens showed variable responses to changes in extreme winter weather events. Snow mould formation underneath the snow may contribute to spatial heterogeneity in plant growth, arthropod communities and carbon cycling. Winter snow cover and depth will drive the reported impacts of winter climate change and add to spatial patterns in vegetation heterogeneity. The challenges ahead lie in obtaining better predictions on the snow patterns across the landscape and how these will be altered due to winter climate change.     

Four Decades of Plant Community Change in the Alpine Tundra of Southwest Yukon,Canada

Repeat measurements from long-term plots provide precise data for studying plant community change. In 2010, we visited a remote location in Yukon, Canada, where a detailed survey of alpine tundra communities was conducted in 1968. Plant community composition was resurveyed on the same four slopes using the same methods as the original study. Species richness and diversity increased significantly over the 42 years and non-metric multidimensional scaling indicated that community composition had also changed significantly. However, the direction and magnitude of change varied with aspect. Dominant species were not replaced or eliminated but, instead, declined in relative importance. Fine-scale changes in vegetation were evident from repeat photography and dendro-ecological analysis of erect shrubs, supporting the community-level analysis. The period of study corresponds to a mean annual temperature increase of 2°C, suggesting that climate warming has influenced these changes.     

High Arctic ecosystem states: Conceptual models of vegetation change to guide long-term monitoring and research

Vegetation change has consequences for terrestrial ecosystem structure and functioning and may involve climate feedbacks. Hence, when monitoring ecosystem states and changes thereof, the vegetation is often a primary monitoring target. Here, we summarize current understanding of vegetation change in the High Arctic—the World’s most rapidly warming region—in the context of ecosystem monitoring. To foster development of deployable monitoring strategies, we categorize different kinds of drivers (disturbances or stresses) of vegetation change either as pulse (i.e. drivers that occur as sudden and short events, though their effects may be long lasting) or press (i.e. drivers where change in conditions remains in place for a prolonged period, or slowly increases in pressure). To account for the great heterogeneity in vegetation responses to climate change and other drivers, we stress the need for increased use of ecosystem-specific conceptual models to guide monitoring and ecological studies in the Arctic. We discuss a conceptual model with three hypothesized alternative vegetation states characterized by mosses, herbaceous plants, and bare ground patches, respectively. We use moss-graminoid tundra of Svalbard as a case study to discuss the documented and potential impacts of different drivers on the possible transitions between those states. Our current understanding points to likely additive effects of herbivores and a warming climate, driving this ecosystem from a moss-dominated state with cool soils, shallow active layer and slow nutrient cycling to an ecosystem with warmer soil, deeper permafrost thaw, and faster nutrient cycling. Herbaceous-dominated vegetation and (patchy) bare ground would present two states in response to those drivers. Conceptual models are an operational tool to focus monitoring efforts towards management needs and identify the most pressing scientific questions. We promote greater use of conceptual models in conjunction with a state-and-transition framework in monitoring to ensure fit for purpose approaches. Defined expectations of the focal systems’ responses to different drivers also facilitate linking local and regional monitoring efforts to international initiatives, such as the Circumpolar Biodiversity Monitoring Program.     

Implications of climate and land-use change for landscape processes,biodiversity, ecosystem services,and governance

This introduction to the Special Issue summarizes the results of 14 scientific articles from the interdisciplinary research program Ekoklim at Stockholm University, Sweden. In this program, we investigate effects of changing climate and land use on landscape processes, biodiversity, and ecosystem services, and analyze issues related to adaptive governance in the face of climate and land-use change. We not only have a research focus on the 22 650 km² Norrström catchment surrounding lake Mälaren in south-central Sweden, but we also conduct research in other Swedish regions. The articles presented here show complex interactions between multiple drivers of change, as well as feedback processes at different spatiotemporal scales. Thus, the Ekoklim program highlights and deals with issues relevant for the future challenges society will face when land-use change interacts with climate change.     

Responses to projected changes in climate and UV-B at the species level

Environmental manipulation experiments showed that species respond individualistically to each environmental-change variable. The greatest responses of plants were generally to nutrient, particularly nitrogen, addition. Summer warming experiments showed that woody plant responses were dominant and that mosses and lichens became less abundant. Responses to warming were controlled by moisture availability and snow cover. Many invertebrates increased population growth in response to summer warming, as long as desiccation was not induced. CO2 and UV-B enrichment experiments showed that plant and animal responses were small. However, some microorganisms and species of fungi were sensitive to increased UV-B and some intensive mutagenic actions could, perhaps, lead to unexpected epidemic outbreaks. Tundra soil heating, CO2 enrichment and amendment with mineral nutrients generally accelerated microbial activity. Algae are likely to dominate cyanobacteria in milder climates. Expected increases in winter freeze-thaw cycles leading to ice-crust formation are likely to severely reduce winter survival rate and disrupt the population dynamics of many terrestrial animals. A deeper snow cover is likely to restrict access to winter pastures by reindeer/caribou and their ability to flee from predators while any earlier onset of the snow-free period is likely to stimulate increased plant growth. Initial species responses to climate change might occur at the sub-species level: an Arctic plant or animal species with high genetic/racial diversity has proved an ability to adapt to different environmental conditions in the past and is likely to do so also in the future. Indigenous knowledge, air photographs, satellite images and monitoring show that changes in the distributions of some species are already occurring: Arctic vegetation is becoming more shrubby and more productive, there have been recent changes in the ranges of caribou, and "new" species of insects and birds previously associated with areas south of the treeline have been recorded. In contrast, almost all Arctic breeding bird species are declining and models predict further quite dramatic reductions of the populations of tundra birds due to warming. Species-climate response surface models predict potential future ranges of current Arctic species that are often markedly reduced and displaced northwards in response to warming. In contrast, invertebrates and microorganisms are very likely to quickly expand their ranges northwards into the Arctic.     

Effects of changes in climate on landscape and regional processes, and feedbacks to the climate system

Biological and physical processes in the Arctic system operate at various temporal and spatial scales to impact large-scale feedbacks and interactions with the earth system. There are four main potential feedback mechanisms between the impacts of climate change on the Arctic and the global climate system: albedo, greenhouse gas emissions or uptake by ecosystems, greenhouse gas emissions from methane hydrates, and increased freshwater fluxes that could affect the thermohaline circulation. All these feedbacks are controlled to some extent by changes in ecosystem distribution and character and particularly by large-scale movement of vegetation zones. Indications from a few, full annual measurements of CO2 fluxes are that currently the source areas exceed sink areas in geographical distribution. The little available information on CH4 sources indicates that emissions at the landscape level are of great importance for the total greenhouse balance of the circumpolar North. Energy and water balances of Arctic landscapes are also important feedback mechanisms in a changing climate. Increasing density and spatial expansion of vegetation will cause a lowering of the albedo and more energy to be absorbed on the ground. This effect is likely to exceed the negative feedback of increased C sequestration in greater primary productivity resulting from the displacements of areas of polar desert by tundra, and areas of tundra by forest. The degradation of permafrost has complex consequences for trace gas dynamics. In areas of discontinuous permafrost, warming, will lead to a complete loss of the permafrost. Depending on local hydrological conditions this may in turn lead to a wetting or drying of the environment with subsequent implications for greenhouse gas fluxes. Overall, the complex interactions between processes contributing to feedbacks, variability over time and space in these processes, and insufficient data have generated considerable uncertainties in estimating the net effects of climate change on terrestrial feedbacks to the climate system. This uncertainty applies to magnitude, and even direction of some of the feedbacks.     

Long-Term Effects of Grazing and Global Warming on the Composition and Carrying Capacity of Graminoid Marshes for Moulting Geese in East Greenland

Greening of the Arctic due to climate warming may provide herbivores with richer food supplies, resulting in higher herbivore densities. In turn, this may cause changes in vegetation composition and ecosystem function. In 1982-1984, we studied the ecology of non-breeding moulting geese in Jameson Land, low Arctic East Greenland. By then, geese consumed most of the graminoid production in available moss fens, and it appeared that the geese had filled up the available habitat. In 2008, we revisited the area and found that the number of moulting geese and the temperature sum for June-July had tripled, while the above-ground biomass in a moss fen ungrazed by geese had more than doubled. In a goose-grazed fen, the overall plant composition was unchanged, but the frequency of graminoids had decreased and the area with dead vegetation and open spots had increased. We suggest that climate warming has lead to increased productivity, allowing for higher numbers of moulting geese. However, the reduction of vegetation cover by grazing may have longer term negative consequences for the number of geese the habitat can sustain.     

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.     

Effects on the function of Arctic ecosystems in the short- and long-term perspectives

Historically, the function of Arctic ecosystems in terms of cycles of nutrients and carbon has led to low levels of primary production and exchanges of energy, water and greenhouse gases have led to low local and regional cooling. Sequestration of carbon from atmospheric CO2, in extensive, cold organic soils and the high albedo from low, snow-covered vegetation have had impacts on regional climate. However, many aspects of the functioning of Arctic ecosystems are sensitive to changes in climate and its impacts on biodiversity. The current Arctic climate results in slow rates of organic matter decomposition. Arctic ecosystems therefore tend to accumulate organic matter and elements despite low inputs. As a result, soil-available elements like nitrogen and phosphorus are key limitations to increases in carbon fixation and further biomass and organic matter accumulation. Climate warming is expected to increase carbon and element turnover, particularly in soils, which may lead to initial losses of elements but eventual, slow recovery. Individual species and species diversity have clear impacts on element inputs and retention in Arctic ecosystems. Effects of increased CO2 and UV-B on whole ecosystems, on the other hand, are likely to be small although effects on plant tissue chemisty, decomposition and nitrogen fixation may become important in the long-term. Cycling of carbon in trace gas form is mainly as CO2 and CH4. Most carbon loss is in the form of CO2, produced by both plants and soil biota. Carbon emissions as methane from wet and moist tundra ecosystems are about 5% of emissions as CO2 and are responsive to warming in the absence of any other changes. Winter processes and vegetation type also affect CH4 emissions as well as exchanges of energy between biosphere and atmosphere. Arctic ecosystems exhibit the largest seasonal changes in energy exchange of any terrestrial ecosystem because of the large changes in albedo from late winter, when snow reflects most incoming radiation, to summer when the ecosystem absorbs most incoming radiation. Vegetation profoundly influences the water and energy exchange of Arctic ecosystems. Albedo during the period of snow cover declines from tundra to forest tundra to deciduous forest to evergreen forest. Shrubs and trees increase snow depth which in turn increases winter soil temperatures. Future changes in vegetation driven by climate change are therefore, very likely to profoundly alter regional climate.     

Ecosystem responses to climate change at a Low Arctic and a High Arctic long-term research site

Long-term measurements of ecological effects of warming are often not statistically significant because of annual variability or signal noise. These are reduced in indicators that filter or reduce the noise around the signal and allow effects of climate warming to emerge. In this way, certain indicators act as medium pass filters integrating the signal over years-to-decades. In the Alaskan Arctic, the 25-year record of warming of air temperature revealed no significant trend, yet environmental and ecological changes prove that warming is affecting the ecosystem. The useful indicators are deep permafrost temperatures, vegetation and shrub biomass, satellite measures of canopy reflectance (NDVI), and chemical measures of soil weathering. In contrast, the 18-year record in the Greenland Arctic revealed an extremely high summer air-warming of 1.3 °C/decade; the cover of some plant species increased while the cover of others decreased. Useful indicators of change are NDVI and the active layer thickness.