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.     

Cloud Albedo,Greenhouse Effects,Atmospheric Chemistry,and Climate Change

Changes in global atmospheric chemistry and climate are taking place as a result of observed trends in long-lived species such as CO₂, CH₄, N₂O, and the CFCs. The continuation of these trends is expected to eventually lead to a major atmospheric warming that might profoundly affect the society we live in. Trends in short-lived species such as NOx and SOx are also suspected. These trends are not as well established, because the shorter-lived species vary spatially and temporally. Trends in NOx would be expected to lead to increases in tropospheric ozone that would add to the warming created by the other greenhouse gases. Trends in NOx could also alter tropospheric OH concentrations that could lead to changes in CH₄ and some of the CFCs. On the other hand, increases in sulfur emissions may alter cloud optical properties. The changes in cloud optical properties could possibly offset the warming expected from increases in greenhouse gases, depending on the role of natural oceanic sulfur emissions. This paper summarizes recent research in these areas and the interactions of climate and atmospheric chemistry.     

Change in spring arrival of migratory birds under an era of climate change,Swedish data from the last 140 years

Many migratory bird species have advanced their spring arrival during the latest decades, most probably due to climate change. However, studies on migratory phenology in the period before recent global warming are scarce. We have analyzed a historical dataset (1873–1917) of spring arrival to southern and central Sweden of 14 migratory bird species. In addition, we have used relative differences between historical and present-day observations (1984–2013) to evaluate the effect of latitude and migratory strategy on day of arrival over time. There was a larger change in spring phenology in short-distance migrants than in long-distance migrants. Interestingly, the results further suggest that climate change has affected the phenology of short-distance migrants more in southern than in central Sweden. The results suggest that the much earlier calculated arrival to southern Sweden among short-distance migrants mirrors a change in location of wintering areas, hence, connecting migration phenology and wintering range shifts.     

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.     

Modeling the effect of climate change on the distribution of threatened medicinal orchid Satyrium nepalense D. Don in India

It is vital to understand the distribution area of a threatened plant species for its better conservation and management planning. Satyrium nepalense (family: Orchidaceae) is a threatened terrestrial orchid species with valuable medicinal and nutritional properties. The survival of S. nepalense in wild conditions has been challenged by increasing global surface temperature. Hence, understanding the impact of climate change on its potential distribution is crucial to conserve and restore this species. In present study, Maxent species distribution modeling algorithm was used to simulate the current distribution of S. nepalense in India and predict the possible range shift in projected future climate scenarios. A set of 19 bioclimatic variables from WorldClim database were used to predict the potential suitable habitats in current climatic condition and four Representative Concentration Pathway (RCP 2.6, 4.5, 6.0, and 8.5) scenarios by integrating five General Circulation Models (GCMs) for future distribution modeling of species for the years 2050 and 2070. Furthermore, change analysis was performed to identify the suitable habitat in current and future climate for delineating range expansion (gain), contraction (loss), and stable (no change) habitats of species. The Maxent model predicted that?~?2.38% of the geographical area in India is presently climatically suitable for S. nepalense. The key bioclimatic variables affecting the distribution of studied species were the mean temperature of warmest quarter, mean temperature of wettest quarter, precipitation of warmest quarter, and temperature seasonality. Under future climate change scenarios, the total suitable habitat of S. nepalense will increase slightly in the Himalayan region and likely to migrate towards northward, but in the Western Ghats region, the suitable areas will be lost severely. The net habitat loss under four RCP scenarios was estimated from 26 to 39% for the year 2050, which could further increase from 47 to 60% by the year 2070. The finding of the predictive Maxent modeling approach indicates that warming climates could significantly affect the potential habitats of S. nepalense and hence suitable conservation measures need to be taken to protect this threatened orchid species in wild conditions.

     

Further evidence of the effects of global warming on lichens, particularly those with Trentepohlia phycobionts

Increasing evidence suggests that lichens are responding to climate change in Western Europe. More epiphytic species appear to be increasing, rather than declining, as a result of global warming. Many terricolous species, in contrast, are declining. Changes to epiphytic floras are markedly more rapid in formerly heavily polluted, generally built-up or open rural areas, as compared to forested regions. Both the distribution (southern) and ecology (warmth-loving) of the newly established or increasing species seem to be determined by global warming. Epiphytic temperate to boreo-montane species appear to be relatively unaffected. Vacant niches caused by other environmental changes are showing the most pronounced effects of global warming. Species most rapidly increasing in forests, although taxonomically unrelated, all contain Trentepohlia as phycobiont in addition to having a southern distribution. This suggests that in this habitat, Trentepohlia algae, rather than the different lichen symbioses, are affected by global warming.     

What determines the current presence or absence of permafrost in the Torneträsk region, a sub-arctic landscape in northern Sweden?

In a warming climate, permafrost is likely to be significantly reduced and eventually disappear from the sub-Arctic region. This will affect people at a range of scales, from locally by slumping of buildings and roads, to globally as melting of permafrost will most likely increase the emissions of the powerful greenhouse gas methane, which will further enhance global warming. In order to predict future changes in permafrost, it is crucial to understand what determines the presence or absence of permafrost under current climate conditions, to assess where permafrost is particularly vulnerable to climate change, and to identify where changes are already occurring. The Tornetr?sk region of northern sub-Arctic Sweden is one area where changes in permafrost have been recorded and where permafrost could be particularly vulnerable to any future climate changes. This paper therefore reviews the various physical, biological, and anthropogenic parameters that determine the presence or absence of permafrost in the Tornetr?sk region under current climate conditions, so that we can gain an understanding of its current vulnerability and potential future responses to climate change. A patchy permafrost distribution as found in the Tornetr?sk region is not random, but a consequence of site-specific factors that control the microclimate and hence the surface and subsurface temperature. It is also a product of past as well as current processes. In sub-Arctic areas such as northern Sweden, it is mainly the physical parameters, e.g., topography, soil type, and climate (in particular snow depth), that determine permafrost distribution. Even though humans have been present in the study area for centuries, their impacts on permafrost distribution can more or less be neglected at the catchment level. Because ongoing climate warming is projected to continue and lead to an increased snow cover, the permafrost in the region will most likely disappear within decades, at least at lower elevations.     

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.     

Does the stress tolerance of mixed grassland communities change in a future climate? A test with heavy metal stress (zinc pollution)

Will species that are sensitive/tolerant to Zn pollution still have the same sensitivity/tolerance in a future climate? To answer this question we analysed the response of constructed grassland communities to five levels of zinc (Zn) supply, ranging from 0 to 354 mg Zn kg⁻¹ dry soil, under a current climate and a future climate (elevated CO₂ and warming). Zn concentrations increased in roots and shoots with Zn addition but this increase did not differ between climates. Light-saturated net CO₂ assimilation rate (A_sat) of the species, on the other hand, responded differently to Zn addition depending on climate. Still, current and future climate communities have comparable biomass responses to Zn, i.e., no change in root biomass and a 13% decrease of above-ground biomass. Provided that the different response of A_sat in a future climate will not compromise productivity and survival on the long term, sensitivity is not altered by climate change.     

Forest fire emissions in Portugal: a contribution to global warming?

A forecast of expected evolution of carbon dioxide (CO(2)) emissions in Portugal between 1988 and 2010 is presented. Predictions show that CO(2) emissions will almost double in the next twenty years. The equivalent potential CO(2) emissions from nitrogen oxides (NO(x)) and volatile organic compounds (VOC), for a time horizon of 20 years, is also presented. NO(x) and VOC emissions seem to make a significant contribution to the global warming potential of Portuguese emissions. Estimates of CO(2) emissions due to forest fires have been made, oriented towards the study of the Portuguese contribution to the global warming. If the burned area exceeds 100 000 ha this contribution could reach 7% of the total Portuguese CO(2) emissions. The global warming potential of Portuguese forest emissions were also calculated. The climate change predicted to Portugal could be responsible for an increase in the forest fires and consequently for a greater contribution of its emissions to the total values. It was concluded that it is important to quantify emissions of the greenhouse gases, including the contribution of forest fire emissions, not only in Portugal, but in all the Southern European countries.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

Effect of climate change on air quality

Air quality is strongly dependent on weather and is therefore sensitive to climate change. Recent studies have provided estimates of this climate effect through correlations of air quality with meteorological variables, perturbation analyses in chemical transport models (CTMs), and CTM simulations driven by general circulation model (GCM) simulations of 21st-century climate change. We review these different approaches and their results. The future climate is expected to be more stagnant, due to a weaker global circulation and a decreasing frequency of mid-latitude cyclones. The observed correlation between surface ozone and temperature in polluted regions points to a detrimental effect of warming. Coupled GCM–CTM studies find that climate change alone will increase summertime surface ozone in polluted regions by 1–10 ppb over the coming decades, with the largest effects in urban areas and during pollution episodes. This climate penalty means that stronger emission controls will be needed to meet a given air quality standard. Higher water vapor in the future climate is expected to decrease the ozone background, so that pollution and background ozone have opposite sensitivities to climate change. The effect of climate change on particulate matter (PM) is more complicated and uncertain than for ozone. Precipitation frequency and mixing depth are important driving factors but projections for these variables are often unreliable. GCM–CTM studies find that climate change will affect PM concentrations in polluted environments by ±0.1–1 μg m^?3 over the coming decades. Wildfires fueled by climate change could become an increasingly important PM source. Major issues that should be addressed in future research include the ability of GCMs to simulate regional air pollution meteorology and its sensitivity to climate change, the response of natural emissions to climate change, and the atmospheric chemistry of isoprene. Research needs to be undertaken on the effect of climate change on mercury, particularly in view of the potential for a large increase in mercury soil emissions driven by increased respiration in boreal ecosystems.     

A Richer,Greener and Smaller Alpine World: Review and Projection of Warming-Induced Plant Cover Change in the Swedish Scandes

Alpine plant life is proliferating, biodiversity is on the rise and the mountain world appears more productive and inviting than ever. Upper range margin rise of trees and low-altitude (boreal) plant species, expansion of alpine grasslands and dwarf-shrub heaths are the modal biotic adjustments during the past few decades, after a century of substantial climate warming in the Swedish Scandes. This course of biotic landscape evolution has reached historical dimensions and broken a multi-millennial trend of plant cover retrogression, alpine tundra expansion, floristic and faunal impoverishment, all imposed by progressive and deterministic neoglacial climate cooling. Continued modest warming over the present century will likely be beneficial to alpine biodiversity, geoecological stability, resilience, sustainable reindeer husbandry and aesthetic landscape qualities. These aspects are highlighted by an integrative review of results from long-term monitoring of subalpine/alpine vegetation in the Swedish Scandes. This forms the basis for some tentative projections of landscape transformations in a potentially warmer future. Notably, these results and projections are not necessarily valid in other regions and differ in some respects from model predictions. Continued monitoring is mandatory as a basis for generation of more realistic vegetation and ecosystem models.