(Table 1) Normalized difference vegetation index in control and warming plots, Abisco Research Station

Extreme winter warming events in the sub-Arctic have caused considerable vegetation damage due to rapid changes in temperature and loss of snow cover. The frequency of extreme weather is expected to increase due to climate change thereby increasing the potential for recurring vegetation damage in Arctic regions. Here we present data on vegetation recovery from one such natural event and multiple experimental simulations in the sub-Arctic using remote sensing, handheld passive proximal sensors and ground surveys. Normalized difference vegetation index (NDVI) recovered fast (2 years), from the 26% decline following one natural extreme winter warming event. Recovery was associated with declines in dead Empetrum nigrum (dominant dwarf shrub) from ground surveys. However, E. nigrum healthy leaf NDVI was also reduced (16%) following this winter warming event in experimental plots (both control and treatments), suggesting that non-obvious plant damage (i.e., physiological stress) had occurred in addition to the dead E. nigrum shoots that was considered responsible for the regional 26% NDVI decline. Plot and leaf level NDVI provided useful additional information that could not be obtained from vegetation surveys and regional remote sensing (MODIS) alone. The major damage of an extreme winter warming event appears to be relatively transitory. However, potential knock-on effects on higher trophic levels (e.g., rodents, reindeer, and bear) could be unpredictable and large. Repeated warming events year after year, which can be expected under winter climate warming, could result in damage that may take much longer to recover.

Plant distribution and climate data across western North America

Using an extensive network of occurrence records for 293 plant species collected over the past 40 years across a climatically diverse geographic section of western North America, we find that plant species distributions were just as likely to shift upwards (i.e., towards higher elevations) as downward (i.e., towards lower elevations) - despite consistent warming across the study area. Although there was no clear directional response to climate warming across the entire study area, there was significant region-to region- variation in responses (i.e. from as many as 73% to as few as32% of species shifting upward or downward). To understand the factors that might be controlling region-specific distributional shifts, we explored the relationship between the direction of change in distribution limits and the nature of recent climate change. We found that the direction of distribution limit shifts was explained by an interaction between the rate of change in local summer temperatures and seasonal precipitation. Specifically, species shifted upward at their upper elevational limit when snowfall declined at slower rates and minimum temperatures increased. By contrast, species shifted upwards at their lower elevation limit when maximum temperatures increased or both temperature and precipitation decreased. Our results suggest that future species' elevational distribution shifts will be complex, depending on the interaction between seasonal temperature and precipitation change.

Size matters: plasticity in metabolic scaling shows body-size may modulate responses to climate change

Variability in metabolic scaling in animals, the relationship between metabolic rate ( R) and body mass ( M), has been a source of debate and controversy for decades. R is proportional to Mb, the precise value of b much debated, but historically considered equal in all organisms. Recent metabolic theory, however, predicts b to vary among species with ecology and metabolic level, and may also vary within species under different abiotic conditions. Under climate change, most species will experience increased temperatures, and marine organisms will experience the additional stressor of decreased seawater pH ('ocean acidification'). Responses to these environmental changes are modulated by myriad species-specific factors. Body-size is a fundamental biological parameter, but its modulating role is relatively unexplored. Here, we show that changes to metabolic scaling reveal asymmetric responses to stressors across body-size ranges; b is systematically decreased under increasing temperature in three grazing molluscs, indicating smaller individuals were more responsive to warming. Larger individuals were, however, more responsive to reduced seawater pH in low temperatures. These alterations to the allometry of metabolism highlight abiotic control of metabolic scaling, and indicate that responses to climate warming and ocean acidification may be modulated by body-size.

(Table 1) Climate characteristics of the four North American Tundra Experiment (ITEX) sites

Climate warming is expected to differentially affect CO2 exchange of the diverse ecosystems in the Arctic. Quantifying responses of CO2 exchange to warming in these ecosystems will require coordinated experimentation using standard temperature manipulations and measurements. Here, we used the International Tundra Experiment (ITEX) standard warming treatment to determine CO2 flux responses to growing-season warming for ecosystems spanning natural temperature and moisture ranges across the Arctic biome. We used the four North American Arctic ITEX sites (Toolik Lake, Atqasuk, and Barrow [USA] and Alexandra Fiord [Canada]) that span 10° of latitude. At each site, we investigated the CO2 responses to warming in both dry and wet or moist ecosystems. Net ecosystem CO2 exchange (NEE), ecosystem respiration (ER), and gross ecosystem photosynthesis (GEP) were assessed using chamber techniques conducted over 24-h periods sampled regularly throughout the summers of two years at all sites. At Toolik Lake, warming increased net CO2 losses in both moist and dry ecosystems. In contrast, at Atqasuk and Barrow, warming increased net CO2 uptake in wet ecosystems but increased losses from dry ecosystems. At Alexandra Fiord, warming improved net carbon uptake in the moist ecosystem in both years, but in the wet and dry ecosystems uptake increased in one year and decreased the other. Warming generally increased ER, with the largest increases in dry ecosystems. In wet ecosystems, high soil moisture limited increases in respiration relative to increases in photosynthesis. Warming generally increased GEP, with the notable exception of the Toolik Lake moist ecosystem, where warming unexpectedly decreased GEP >25%. Overall, the respiration response determined the effect of warming on ecosystem CO2 balance. Our results provide the first multiple-site comparison of arctic tundra CO2 flux responses to standard warming treatments across a large climate gradient. These results indicate that (1) dry tundra may be initially the most responsive ecosystems to climate warming by virtue of strong increases in ER, (2) moist and wet tundra responses are dampened by higher water tables and soil water contents, and (3) both GEP and ER are responsive to climate warming, but the magnitudes and directions are ecosystem-dependent.

Initial soil properties and biomass after experimental grazing and warming in mesic and wet sites, Adventdalen valley, Spitzbergen

High-latitude ecosystems store large amounts of carbon (C); however, the C storage of these ecosystems is under threat from both climate warming and increased levels of herbivory. In this study we examined the combined role of herbivores and climate warming as. drivers of CO2 fluxes in two typical high-latitude habitats (mesic heath and wet meadow). We hypothesized that both herbivory and climate warming would reduce the C sink strength of Arctic tundra through their combined effects on plant biomass and gross ecosystem photosynthesis and on decomposition rates and the abiotic environment. To test this hypothesis we employed experimental warming (via International Tundra Experiment [ITEX] chambers) and grazing (via captive Barnacle Geese) in a three-year factorial field experiment. Ecosystem CO2 fluxes (net ecosystem exchange of CO2, ecosystem respiration, and gross ecosystem photosynthesis) were measured in all treatments at varying intensity over the three growing seasons to capture the impact of the treatments on a range of temporal scales (diurnal, seasonal, and interannual). Grazing and warming treatments had markedly different effects on CO2 fluxes in the two tundra habitats. Grazing caused a strong reduction in CO2 assimilation in the wet meadow, while warming reduced CO2 efflux from the mesic heath. Treatment effects on net ecosystem exchange largely derived from the modification of gross ecosystem photosynthesis rather than ecosystem respiration. In this study we have demonstrated that on the habitat scale, grazing by geese is a strong driver of net ecosystem exchange of CO2, with the potential to reduce the CO2 sink strength of Arctic ecosystems. Our results highlight that the large reduction in plant biomass due to goose grazing in the Arctic noted in several studies can alter the C balance of wet tundra ecosystems. We conclude that herbivory will modulate direct climate warming responses of Arctic tundra with implications for the ecosystem C balance; however, the magnitude and direction of the response will be habitat-specific.

Mesocosm experiment on warming and acidification effects on phytoplankton composition and cell size

While the isolated responses of marine phytoplankton to climate warming and to ocean acidification have been studies intensively, studies on the combined effect of both aspects of Global Change are still scarce. Therefore, we performed a mesocosm experiment with a factorial combination of temperature (9 and 15°C) and pCO2 (560 ppm and 1400 ppm) with a natural autumn plankton community from the western Baltic Sea. Temporal trajectories of total biomass and of the biomass of the most important higher taxa followed similar patterns in all treatments. When averaging over the entire time course, phytoplankton biomass decreased with warming and increased with CO2 under warm conditions. The contribution of the two dominant higher phytoplankton taxa (diatoms and cryptophytes) and of the 4 most important species (3 diatoms, 1 cryptophyte) did not respond to the experimental treatments. Taxonomic composition of phytoplankton showed only responses at the level of subdominant and rare species. Phytoplankton cell sizes increased with CO2 addition and decreased with warming. Both effects were stronger for larger species. Warming effects were stronger than CO2 effects and tended to counteract each other. Phytoplankton communities without calcifying species and exposed to short-term variation of COO2 seem to be rather resistant to ocean acidification.

Planktonic foraminifera in the latest Maastrichtian of Walvis Ridge, South Atlantic

An abrupt global warming of 3-4°C occurred near the end of the Maastrichtian at 65.45-65.10 Ma. The environmental effects of this warm event are here documented based on stable isotopes and quantitative analysis of planktonic foraminifera at the South Atlantic DSDP Site 525A. Stable isotopes of individual species mark a rapid increase in temperature and a reduction in the vertical water mass stratification that is accompanied by a decrease in niche habitats, reduced species diversity and/or abundance, smaller species morphologies or dwarfing, and reduced photosymbiotic activity. During the warm event, the relative abundance of a large number of species decreased, including tropical-subtropical affiliated species, whereas typical mid-latitude species retained high abundances. This indicates that climate warming did not create favorable conditions for all tropical-subtropical species at mid-latitudes and did not cause a massive retreat in the local mid-latitude population. A noticeable exception is the ecological generalist Heterohelix dentata Stenestad that dominated during the cool intervals, but significantly decreased during the warm event. However, dwarfing is the most striking response to the abrupt warming and occurred in various species of different morphologies and lineages (e.g. biserial, trochospiral, keeled globotruncanids). Dwarfing is a typical reaction to environmental stress conditions and was likely the result of increased reproduction rates. Similarly, photosymbiotic activity appears to have been reduced significantly during the maximum warming, as indicated by decreased delta13C values. The foraminiferal response to climate change is thus multifaceted resulting in decreased species diversity, decreased species populations, increased competition due to reduced niche habitats, dwarfing and reduced photosymbiotic activity.

Numer and type of seeds introduced by footwear of travelers to Svalbard and germination success in 2008

Expanding visitation to Polar regions combined with climate warming increases the potential for alien species introduction and establishment. We quantified vascular plant propagule pressure associated with different groups of travelers to the high-Arctic archipelago of Svalbard, and evaluated the potential of introduced seeds to germinate under the most favorable average Svalbard soil temperature (10°C). We sampled the footwear of 259 travelers arriving by air to Svalbard during the summer of 2008, recording 1,019 seeds: a mean of 3.9 (±0.8) seeds per traveler. Assuming the seed influx is representative for the whole year, we estimate a yearly seed load of around 270,000 by this vector alone. Seeds of 53 species were identified from 17 families, with Poaceae having both highest diversity and number of seeds. Eight of the families identified are among those most invasive worldwide, while the majority of the species identified were non-native to Svalbard. The number of seeds was highest on footwear that had been used in forested and alpine areas in the 3 months prior to traveling to Svalbard, and increased with the amount of soil affixed to footwear. In total, 26% of the collected seeds germinated under simulated Svalbard conditions. Our results demonstrate high propagule transport through aviation to highly visited cold-climate regions and isolated islands is occurring. Alien species establishment is expected to increase with climate change, particularly in high latitude regions, making the need for regional management considerations a priority.

Soil characteristics and decomposition of organic matter on Anchorage, Signy and Falkland Islands

Antarctic terrestrial ecosystems have poorly developed soils and currently experience one of the greatest rates of climate warming on the globe. We investigated the responsiveness of organic matter decomposition in Maritime Antarctic terrestrial ecosystems to climate change, using two study sites in the Antarctic Peninsula region (Anchorage Island, 67°S; Signy Island, 61°S), and contrasted the responses found with those at the cool temperate Falkland Islands (52°S). Our approach consisted of two complementary methods: (1) Laboratory measurements of decomposition at different temperatures (2, 6 and 10 °C) of plant material and soil organic matter from all three locations. (2) Field measurements at all three locations on the decomposition of soil organic matter, plant material and cellulose, both under natural conditions and under experimental warming (about 0.8 °C) achieved using open top chambers. Higher temperatures led to higher organic matter breakdown in the laboratory studies, indicating that decomposition in Maritime Antarctic terrestrial ecosystems is likely to increase with increasing soil temperatures. However, both laboratory and field studies showed that decomposition was more strongly influenced by local substratum characteristics (especially soil N availability) and plant functional type composition than by large-scale temperature differences. The very small responsiveness of organic matter decomposition in the field (experimental temperature increase <1 °C) compared with the laboratory (experimental increases of 4 or 8 °C) shows that substantial warming is required before significant effects can be detected.

(Table 2) Survivorship of transplanted seedlings of Picea glauca inside and outside three tree islands in 1994, 1995, 2009 and after 15 years

The northern boundary of boreal forest and the ranges of tree species are expected to shift northward in response to climate warming, which will result in a decrease in the albedo of areas currently covered by tundra vegetation, an increase in terrestrial carbon sequestration, and an alteration of biodiversity in the current Low Arctic. Central to the prediction of forest expansion is an increase in the reproductive capacity and establishment of individual trees. We assessed cone production, seed viability, and transplanted seedling success of Picea glauca (Moench.) Voss. (white spruce) in the early 1990s and again in the late 2000s at four forest stand sites and eight tree island sites (clonal populations beyond present treeline) in the Mackenzie Delta region of the Northwest Territories, Canada. Over the past 20 years, average temperatures in this region have increased by 0.9 °C. This area has the northernmost forest-tundra ecotone in North America and is one of the few circumpolar regions where the northern limit of conifer trees reaches the Arctic Ocean. We found that cone production and seed viability did not change between the two periods of examination and that both variables decreased northward across the forest-tundra ecotone. Nevertheless, white spruce individuals at the northern limit of the forest-tundra ecotone produced viable seeds. Furthermore, transplanted seedlings were able to survive in the northernmost sites for 15 years, but there were no signs of natural regeneration. These results indicate that if climatic conditions continue to ameliorate, reproductive output will likely increase, but seedling establishment and forest expansion within the forest-tundra of this region is unlikely to occur without the availability of suitable recruitment sites. Processes that affect the availability of recruitment sites are likely to be important elsewhere in the circumpolar ecotone, and should be incorporated into models and predictions of climate change and its effects on the northern forest-tundra ecotone.

Mean cover and frequencies of vascular plant species during three decades around Abisko Scientific Research Station

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.

Pollen record and age determination of a sediment profile from Lake Teletskoye, Russia

A high-resolution pollen record from Lake Teletskoye documents the climate-related vegetation history of the northern Altai Mountain region during the last millennium. Siberian pine taiga with Scots pine, fir, spruce, and birch dominated the vegetation between ca. AD 1050 and 1100. The climate was similar to modern. In the beginning of the 12th century, birch and shrub alder increased. Lowered pollen concentrations and simultaneous peaks in herbs (especially Artemisia and Poaceae), ferns, and charcoal fragments point to colder and more arid climate conditions than before, with frequent fire events. Around AD 1200, regional climate became warmer and more humid than present, as revealed by an increase of Siberian pine and decreases of dry herb taxa and charcoal contents. Climatic conditions were rather stable until ca. AD 1410. An increase of Artemisia pollen may reflect slightly drier climate conditions between AD 1410 and 1560. Increases in Alnus, Betula, Artemisia, and Chenopodiaceae pollen and in charcoal particle contents may reflect further deterioration of climate conditions between AD 1560 and 1810, consistent with the Little Ice Age. After AD 1850 the vegetation gradually approached the modern one, in conjunction with ongoing climate warming.

(Table 1) Bioclimatic subzones and soil characteristics along the Yamal Arctic Transect, northwestern Siberia

Sustainability of tundra vegetation under changing climate on the Yamal Peninsula, northwestern Siberia, home to the world's largest area of reindeer husbandry, is of crucial importance to the local native community. An integrated investigation is needed for better understanding of the effects of soils, climate change and grazing on tundra vegetation in the Yamal region. In this study we applied a nutrient-based plant community model - ArcVeg - to evaluate how two factors (soil organic nitrogen (SON) levels and grazing) interact to affect tundra responses to climate warming across a latitudinal climatic gradient on the Yamal Peninsula. Model simulations were driven by field-collected soil data and expected grazing patterns along the Yamal Arctic Transect (YAT), within bioclimate subzones C (high arctic), D (northern low arctic) and E (southern low arctic). Plant biomass and NPP (net primary productivity) were significantly increased with warmer bioclimate subzones, greater soil nutrient levels and temporal climate warming, while they declined with higher grazing frequency. Temporal climate warming of 2 °C caused an increase of 665 g/m**2 in total biomass at the high SON site in subzone E, but only 298 g/m**2 at the low SON site. When grazing frequency was also increased, total biomass increased by only 369 g/m**2 at the high SON site in contrast to 184 g/m**2 at the low SON site in subzone E. Our results suggest that high SON can support greater plant biomass and plant responses to climate warming, while low SON and grazing may limit plant response to climate change. In addition to the first order factors (SON, bioclimate subzones, grazing and temporal climate warming), interactions among these significantly affect plant biomass and productivity in the arctic tundra and should not be ignored in regional scale studies.