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.     

General effects of climate change on Arctic fishes and fish populations

Projected shifts in climate forcing variables such as temperature and precipitation are of great relevance to arctic freshwater ecosystems and biota. These will result in many direct and indirect effects upon the ecosystems and fish present therein. Shifts projected for fish populations will range from positive to negative in overall effect, differ among species and also among populations within species depending upon their biology and tolerances, and will be integrated by the fish within their local aquascapes. This results in a wide range of future possibilities for arctic freshwater and diadromous fishes. Owing to a dearth of basic knowledge regarding fish biology and habitat interactions in the north, complicated by scaling issues and uncertainty in future climate projections, only qualitative scenarios can be developed in most cases. This limits preparedness to meet challenges of climate change in the Arctic with respect to fish and fisheries.     

Uncertainties and recommendations

An assessment of the impacts of changes in climate and UV-B radiation on Arctic terrestrial ecosystems, made within the Arctic Climate Impacts Assessment (ACIA), highlighted the profound implications of projected warming in particular for future ecosystem services, biodiversity and feedbacks to climate. However, although our current understanding of ecological processes and changes driven by climate and UV-B is strong in some geographical areas and in some disciplines, it is weak in others. Even though recently the strength of our predictions has increased dramatically with increased research effort in the Arctic and the introduction of new technologies, our current understanding is still constrained by various uncertainties. The assessment is based on a range of approaches that each have uncertainties, and on data sets that are often far from complete. Uncertainties arise from methodologies and conceptual frameworks, from unpredictable surprises, from lack of validation of models, and from the use of particular scenarios, rather than predictions, of future greenhouse gas emissions and climates. Recommendations to reduce the uncertainties are wide-ranging and relate to all disciplines within the assessment. However, a repeated theme is the critical importance of achieving an adequate spatial and long-term coverage of experiments, observations and monitoring of environmental changes and their impacts throughout the sparsely populated and remote region that is the Arctic.     

Cumulative effects of climate warming and other human activities on freshwaters of Arctic and subarctic North America

Despite their generally isolated geographic locations, the freshwaters of the north are subjected to a wide spectrum of environmental stressors. High-latitude regions are especially sensitive to the effects of recent climatic warming, which have already resulted in marked regime shifts in the biological communities of many Arctic lakes and ponds. Important drivers of these limnological changes have included changes in the amount and duration of snow and ice cover, and, for rivers and lakes in their deltas, the frequency and extent of spring floods. Other important climate-related shifts include alterations in evaporation and precipitation ratios, marked changes in the quality and quantity of lake and river water inflows due to accelerated glacier and permafrost melting, and declining percentages of precipitation that falls as snow. The depletion of stratospheric ozone over the north, together with the clarity of many Arctic lakes, renders them especially susceptible to damage from ultraviolet radiation. In addition, the long-range atmospheric transport of pollutants, coupled with the focusing effects of contaminant transport from biological vectors to some local ecosystems (e.g., salmon nursery lakes, ponds draining seabird colonies) and biomagnification in long food chains, have led to elevated concentrations of many persistent organic pollutants (e.g., insecticides, which have never been used in Arctic regions) and other pollutants (e.g., mercury). Rapid development of gas and oil pipelines, mining for diamonds and metals, increases in human populations, and the development of all-season roads, seaports, and hydroelectric dams will stress northern aquatic ecosystems. The cumulative effects of these stresses will be far more serious than those caused by changing climate alone.     

Biodiversity, distributions and adaptations of Arctic species in the context of environmental change

The individual of a species is the basic unit which responds to climate and UV-B changes, and it responds over a wide range of time scales. The diversity of animal, plant and microbial species appears to be low in the Arctic, and decreases from the boreal forests to the polar deserts of the extreme North but primitive species are particularly abundant. This latitudinal decline is associated with an increase in super-dominant species that occupy a wide range of habitats. Climate warming is expected to reduce the abundance and restrict the ranges of such species and to affect species at their northern range boundaries more than in the South: some Arctic animal and plant specialists could face extinction. Species most likely to expand into tundra are boreal species that currently exist as outlier populations in the Arctic. Many plant species have characteristics that allow them to survive short snow-free growing seasons, low solar angles, permafrost and low soil temperatures, low nutrient availability and physical disturbance. Many of these characteristics are likely to limit species' responses to climate warming, but mainly because of poor competitive ability compared with potential immigrant species. Terrestrial Arctic animals possess many adaptations that enable them to persist under a wide range of temperatures in the Arctic. Many escape unfavorable weather and resource shortage by winter dormancy or by migration. The biotic environment of Arctic animal species is relatively simple with few enemies, competitors, diseases, parasites and available food resources. Terrestrial Arctic animals are likely to be most vulnerable to warmer and drier summers, climatic changes that interfere with migration routes and staging areas, altered snow conditions and freeze-thaw cycles in winter, climate-induced disruption of the seasonal timing of reproduction and development, and influx of new competitors, predators, parasites and diseases. Arctic microorganisms are also well adapted to the Arctic's climate: some can metabolize at temperatures down to -39 degrees C. Cyanobacteria and algae have a wide range of adaptive strategies that allow them to avoid, or at least minimize UV injury. Microorganisms can tolerate most environmental conditions and they have short generation times which can facilitate rapid adaptation to new environments. In contrast, Arctic plant and animal species are very likely to change their distributions rather than evolve significantly in response to warming.     

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.Supplementary InformationThe online version contains supplementary material available at 10.1007/s13280-021-01596-w.     

Seafood from a changing Arctic

We review current knowledge about climate change impacts on Arctic seafood production. Large-scale changes in the Arctic marine food web can be expected for the next 40–100 years. Possible future trajectories under climate change for Arctic capture fisheries anticipate the movement of aquatic species into new waters and changed the dynamics of existing species. Negative consequences are expected for some fish stocks but others like the Barents Sea cod (Gadus morhua) may instead increase. Arctic aquaculture that constitutes about 2% of global farming is mainly made up of Norwegian salmon (Salmo salar) farming. The sector will face many challenges in a warmer future and some of these are already a reality impacting negatively on salmon growth. Other more indirect effects from climate change are more uncertain with respect to impacts on the economic conditions of Arctic aquaculture.     

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.     

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.     

Future distribution of Arctic char Salvelinus alpinus in Sweden under climate change: effects of temperature, lake size and species interactions

Novel communities will be formed as species with a variety of dispersal abilities and environmental tolerances respond individually to climate change. Thus, models projecting future species distributions must account for species interactions and differential dispersal abilities. We developed a species distribution model for Arctic char Salvelinus alpinus, a freshwater fish that is sensitive both to warm temperatures and to species interactions. A logistic regression model using lake area, mean annual air temperature (1961-1990), pike Esox lucius and brown trout Salmo trutta occurrence correctly classified 95?% of 467 Swedish lakes. We predicted that Arctic char will lose 73?% of its range in Sweden by 2100. Predicted extinctions could be attributed both to simulated temperature increases and to projected pike invasions. The Swedish mountains will continue to provide refugia for Arctic char in the future and should be the focus of conservation efforts for this highly valued fish.     

Arctic Climate Tipping Points

There is widespread concern that anthropogenic global warming will trigger Arctic climate tipping points. The Arctic has a long history of natural, abrupt climate changes, which together with current observations and model projections, can help us to identify which parts of the Arctic climate system might pass future tipping points. Here the climate tipping points are defined, noting that not all of them involve bifurcations leading to irreversible change. Past abrupt climate changes in the Arctic are briefly reviewed. Then, the current behaviour of a range of Arctic systems is summarised. Looking ahead, a range of potential tipping phenomena are described. This leads to a revised and expanded list of potential Arctic climate tipping elements, whose likelihood is assessed, in terms of how much warming will be required to tip them. Finally, the available responses are considered, especially the prospects for avoiding Arctic climate tipping points.     

Status and trends of tundra birds across the circumpolar Arctic

Tundra-breeding birds face diverse conservation challenges, from accelerated rates of Arctic climate change to threats associated with highly migratory life histories. Here we summarise the status and trends of Arctic terrestrial birds (88 species, 228 subspecies or distinct flyway populations) across guilds/regions, derived from published sources, raw data or, in rare cases, expert opinion. We report long-term trends in vital rates (survival, reproduction) for the handful of species and regions for which these are available. Over half of all circumpolar Arctic wader taxa are declining (51% of 91 taxa with known trends) and almost half of all waterfowl are increasing (49% of 61 taxa); these opposing trends have fostered a shift in community composition in some locations. Declines were least prevalent in the African-Eurasian Flyway (29%), but similarly prevalent in the remaining three global flyways (44–54%). Widespread, and in some cases accelerating, declines underscore the urgent conservation needs faced by many Arctic terrestrial bird species.     

Future Distribution of Arctic Char <Emphasis Type="Italic">Salvelinus alpinus</Emphasis> in Sweden under Climate Change: Effects of Temperature,Lake Size and Species Interactions

Novel communities will be formed as species with a variety of dispersal abilities and environmental tolerances respond individually to climate change. Thus, models projecting future species distributions must account for species interactions and differential dispersal abilities. We developed a species distribution model for Arctic char Salvelinus alpinus, a freshwater fish that is sensitive both to warm temperatures and to species interactions. A logistic regression model using lake area, mean annual air temperature (1961–1990), pike Esox lucius and brown trout Salmo trutta occurrence correctly classified 95 % of 467 Swedish lakes. We predicted that Arctic char will lose 73 % of its range in Sweden by 2100. Predicted extinctions could be attributed both to simulated temperature increases and to projected pike invasions. The Swedish mountains will continue to provide refugia for Arctic char in the future and should be the focus of conservation efforts for this highly valued fish.     

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.     

Tipping Points in the Arctic: Eyeballing or Statistical Significance?

Arctic ecosystems have experienced and are projected to experience continued large increases in temperature and declines in sea ice cover. It has been hypothesized that small changes in ecosystem drivers can fundamentally alter ecosystem functioning, and that this might be particularly pronounced for Arctic ecosystems. We present a suite of simple statistical analyses to identify changes in the statistical properties of data, emphasizing that changes in the standard error should be considered in addition to changes in mean properties. The methods are exemplified using sea ice extent, and suggest that the loss rate of sea ice accelerated by factor of ~5 in 1996, as reported in other studies, but increases in random fluctuations, as an early warning signal, were observed already in 1990. We recommend to employ the proposed methods more systematically for analyzing tipping points to document effects of climate change in the Arctic.     

Societal implications of a changing Arctic Ocean

The Arctic Ocean is undergoing rapid change: sea ice is being lost, waters are warming, coastlines are eroding, species are moving into new areas, and more. This paper explores the many ways that a changing Arctic Ocean affects societies in the Arctic and around the world. In the Arctic, Indigenous Peoples are again seeing their food security threatened and cultural continuity in danger of disruption. Resource development is increasing as is interest in tourism and possibilities for trans-Arctic maritime trade, creating new opportunities and also new stresses. Beyond the Arctic, changes in sea ice affect mid-latitude weather, and Arctic economic opportunities may re-shape commodities and transportation markets. Rising interest in the Arctic is also raising geopolitical tensions about the region. What happens next depends in large part on the choices made within and beyond the Arctic concerning global climate change and industrial policies and Arctic ecosystems and cultures.     

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.     

Impacts of atmospheric pollution on the plant communities of British acid grasslands

Air pollutants are recognised as important agents of ecosystem change but few studies consider the effects of multiple pollutants and their interactions. Here we use ordination, constrained cluster analysis and indicator value analyses to identify potential environmental controls on species composition, ecological groupings and indicator species in a gradient study of UK acid grasslands. The community composition of these grasslands is related to climate, grazing, ozone exposure and nitrogen deposition, with evidence for an interaction between the ecological impacts of base cation and nitrogen deposition. Ozone is a key agent in species compositional change but is not associated with a reduction in species richness or diversity indices, showing the subtly different drivers on these two aspects of ecosystem degradation. Our results demonstrate the effects of multiple interacting pollutants, which may collectively have a greater impact than any individual agent.     

Effects of environmental methylmercury on the health of wild birds, mammals, and fish

Wild piscivorous fish, mammals, and birds may be at risk for elevated dietary methylmercury intake and toxicity. In controlled feeding studies, the consumption of diets that contained Hg (as methylmercury) at environmentally realistic concentrations resulted in a range of toxic effects in fish, birds, and mammals, including behavioral, neurochemical, hormonal, and reproductive changes. Limited field-based studies, especially with certain wild piscivorous bird species, e.g., the common loon, corroborated laboratory-based results, demonstrating significant relations between methylmercury exposure and various indicators of methylmercury toxicity, including reproductive impairment. Potential population effects in fish and wildlife resulting from dietary methylmercury exposure are expected to vary as a function of species life history, as well as regional differences in fish-Hg concentrations, which, in turn, are influenced by differences in Hg deposition and environmental methylation rates. However, population modeling suggests that reductions in Hg emissions could have substantial benefits for some common loon populations that are currently experiencing elevated methylmercury exposure. Predicted benefits would be mediated primarily through improved hatching success and development of hatchlings to maturity as Hg concentrations in prey fish decline. Other piscivorous species may also benefit from decreased Hg exposure but have not been as extensively studied as the common loon.