Contrasting physiological responses to future ocean acidification among Arctic copepod populations

Widespread ocean acidification (OA) is modifying the chemistry of the global ocean, and the Arctic is recognized as the region where the changes will progress at the fastest rate. Moreover, Arctic species show lower capacity for cellular homeostasis and acid‐base regulation rendering them particularly vulnerable to OA. In the present study, we found physiological differences in OA response across geographically separated populations of the keystone Arctic copepod Calanus glacialis. In copepodites stage CIV, measured reaction norms of ingestion rate and metabolic rate showed severe reductions in ingestion and increased metabolic expenses in two populations from Svalbard (Kongsfjord and Billefjord) whereas no effects were observed in a population from the Disko Bay, West Greenland. At pH_T 7.87, which has been predicted for the Svalbard west coast by year 2100, these changes resulted in reductions in scope for growth of 19% in the Kongsfjord and a staggering 50% in the Billefjord. Interestingly, these effects were not observed in stage CV copepodites from any of the three locations. It seems that CVs may be more tolerant to OA perhaps due to a general physiological reorganization to meet low intracellular pH during hibernation. Needless to say, the observed changes in the CIV stage will have serious implications for the C. glacialis population health status and growth around Svalbard. However, OA tolerant populations such as the one in the Disko Bay could help to alleviate severe effects in C. glacialis as a species.     

Timing of blooms,algal food quality and Calanus glacialis reproduction and growth in a changing Arctic

The Arctic bloom consists of two distinct categories of primary producers, ice algae growing within and on the underside of the sea ice, and phytoplankton growing in open waters. Long chain omega‐3 fatty acids, a subgroup of polyunsaturated fatty acids (PUFAs) produced exclusively by these algae, are essential to all marine organisms for successful reproduction, growth, and development. During an extensive field study in the Arctic shelf seas, we followed the seasonal biomass development of ice algae and phytoplankton and their food quality in terms of their relative PUFA content. The first PUFA‐peak occurred in late April during solid ice cover at the onset of the ice algal bloom, and the second PUFA‐peak occurred in early July just after the ice break‐up at the onset of the phytoplankton bloom. The reproduction and growth of the key Arctic grazer Calanus glacialis perfectly coincided with these two bloom events. Females of C. glacialis utilized the high‐quality ice algal bloom to fuel early maturation and reproduction, whereas the resulting offspring had access to ample high‐quality food during the phytoplankton bloom 2 months later. Reduction in sea ice thickness and coverage area will alter the current primary production regime due to earlier ice break‐up and onset of the phytoplankton bloom. A potential mismatch between the two primary production peaks of high‐quality food and the reproductive cycle of key Arctic grazers may have negative consequences for the entire lipid‐driven Arctic marine ecosystem.     

Arctic sea ice as feeding ground for amphipods – food sources and strategies

The amphipod species Gammarus wilkitzkii, Apherusa glacialis, Onisimus nanseni and O. glacialis live permanently associated with the Arctic sea ice. Qualitative and semi-quantitative investigations of gut contents and faeces showed that all four species use detritus as the main food source. Detrital lumps from the underside of sea ice had the same item composition as amphipod gut contents and faeces. Crustacean remains and ice algae were additional food items, but overall they were quantitatively less important. All species are omnivorous; however, differences in gut contents, behavioural observations and functional–morphological studies of the mandibles suggest a differentiation within this feeding strategy. G. wilkitzkii is a detritivorous-carnivorous-necrophagous-suspension-feeding species and shows the most complex feeding strategy. O. nanseni and O. glacialis are predominantly detritivorous-necrophagous, whereas A. glacialis is characterised as a more herbivorous-detritivorous species. By using a variety of the available food sources under Arctic sea ice, the amphipods are well adapted to the under-ice habitat and are less influenced by temporal and spatial variations. Furthermore, the wide food spectrum of all four species reduces the intra- and interspecific competition in a habitat where certain food sources are limited or only seasonally available. Accepted: 30 June 2000     

Reproduction of Calanus glacialis in the Laptev Sea, Arctic Ocean

Abundance and reproductive biology (gonad maturation and egg production) of the Arctic copepod Calanus glacialis were studied in the Laptev Sea and adjacent Arctic Ocean in September 1993 and from July to September 1995. Both abundance and reproductive activity were subject to strong spatial and seasonal variability, which was related to the ice cover, feeding conditions and circulation pattern. Maximum abundance of the C. glacialis population was generally confined to the outer shelf and slope with depths between 50 and 1000 m. During both cruises, highest egg production rates and largest number of young copepodite stages were observed in the eastern Laptev Sea, where the development of the C. glacialis population seems to follow the opening of the “Siberian Polynya”. In the western part, which is usually covered by pack ice, females were all immature, and no young stages were found. However, females responded quickly to a temporary opening of the ice there in 1995 and spawned. Starvation experiments showed that food-independent reproduction fuelled by internal energy resources was at least partly responsible for relatively high egg production rates at low ambient food concentrations. Egg production rates in starved females were considerably higher than those previously reported. Accepted: 14 June 2000     

Life strategy and diet of Calanus glacialis during the winter�Cspring transition in Amundsen Gulf, south-eastern Beaufort Sea

The copepod Calanus glacialis plays a key role in the lipid-based energy flux in Arctic shelf seas. By utilizing both ice algae and phytoplankton, this species is able to extend its growth season considerably in these seasonally ice-covered seas. This study investigated the impacts of the variability in timing and extent of the ice algal bloom on the reproduction and population success of C. glacialis. The vertical distribution, reproduction, amount of storage lipids, stable isotopes, fatty acid and fatty alcohol composition of C. glacialis were assessed during the Circumpolar Flaw Lead System Study. Data were collected in the Amundsen Gulf, south-eastern Beaufort Sea, from January to July 2008 with the core-sampling from March to April. The reduction in sea ice thickness and coverage observed in the Amundsen Gulf in 2007 and 2008 affected the life strategy and reproduction of C. glacialis. Developmental stages CIII and CIV dominated the overwintering population, which resulted in the presence of very few CV and females during spring 2008. Spawning began at the peak of the ice algal bloom that preceded the precocious May ice break-up. Although the main recruitment may have occurred later in the season, low abundance of females combined with a potential mismatch between egg production/development to the first feeding stage and phytoplankton bloom resulted in low recruitment of C. glacialis in the early summer of 2008.     

Zooplankton boom and ice amphipod bust below melting sea ice in the Amundsen Gulf, Arctic Canada

Early summer in the Arctic with extensive ice melt and break-up represents a dramatic change for sympagic–pelagic fauna below seasonal sea ice. As part of the International Polar Year-Circumpolar Flaw Lead system study (IPY-CFL), this investigation quantified zooplankton in the meltwater layer below landfast ice and remaining ice fauna below melting ice during June (2008) in Franklin Bay and Darnley Bay, Amundsen Gulf, Canada. The ice was in a state of advanced melt, with fully developed melt ponds. Intense melting resulted in a 0.3- to 0.5-m-thick meltwater layer below the ice, with a strong halocline to the Arctic water below. Zooplankton under the ice, in and below the meltwater layer, was sampled by SCUBA divers. Dense concentrations (max. 1,400 ind. m⁻³) of Calanus glacialis were associated with the meltwater layer, with dominant copepodid stages CIV and CV and high abundance of nauplii. Less abundant species included Pseudocalanus spp., Oithona similis and C. hyperboreus. The copepods were likely feeding on phytoplankton (0.5–2.3 mg Chl-a m⁻³) in the meltwater layer. Ice amphipods were present at low abundance (<10 ind. m⁻²) and wet biomass (<0.2 g m⁻²). Onisimus glacialis and Apherusa glacialis made up 64 and 51% of the total ice faunal abundance in Darnley Bay and Franklin Bay, respectively. During early summer, the autochthonous ice fauna becomes gradually replaced by allochthonous zooplankton, with an abundance boom near the meltwater layer. The ice amphipod bust occurs during late stages of melting and break-up, when their sympagic habitat is diminished then lost.     

Ice amphipod distribution relative to ice density and under-ice topography in the northern Barents Sea

Arctic ice amphipods are part of the sympagic macrofauna in the Marginal Ice Zone of the northern Barents Sea and represent an important link from lower to higher trophic levels in some Arctic marine food chains. The species diversity in this area (1995/1996) consisted of four species: Gammarus wilkitzkii, Apherusa glacialis, Onisimus nanseni and Onisimus glacialis. The larger ice amphipod, G. wilkitzkii, was the most abundant with the highest biomass (>90%), whereas A. glacialis was abundant, but contributed little to the total biomass (<4%). The other two species were found only in small numbers. Both abundance and biomass of ice amphipods decreased along a latitudinal gradient from north to south across the Marginal Ice Zone. Their distribution was also related to the under-ice topography with regard to mesoscale structures (edge, flat area, dome and ridge). Overall, the abundance and biomass on ridges were much higher in comparison to other mesoscale structures, although edges also showed high abundance, but low biomass. The large G. wilkitzkii was consistently abundant on ridges. The small A. glacialis was predominately associated with edges, but also showed high numbers in dome-shaped areas. The Onisimus species were present in low numbers at all structures, and their biomass contributed <10% on any one structure. The reasons for different distribution patterns of the dominant amphipod species under Arctic sea ice are probably related to different requirements of the species, especially for food, shelter and physiological conditions. Accepted: 27 November 1999     

Do marine phytoplankton follow Bergmann's rule sensu lato?

Global warming has revitalized interest in the relationship between body size and temperature, proposed by Bergmann's rule 150 years ago, one of the oldest manifestations of a ‘biogeography of traits’. We review biogeographic evidence, results from clonal cultures and recent micro‐ and mesocosm experiments with naturally mixed phytoplankton communities regarding the response of phytoplankton body size to temperature, either as a single factor or in combination with other factors such as grazing, nutrient limitation, and ocean acidification. Where possible, we also focus on the comparison between intraspecific size shifts and size shifts resulting from changes in species composition. Taken together, biogeographic evidence, community‐level experiments and single‐species experiments indicate that phytoplankton average cell sizes tend to become smaller in warmer waters, although temperature is not necessarily the proximate environmental factor driving size shifts. Indirect effects via nutrient supply and grazing are important and often dominate. In a substantial proportion of field studies, resource availability is seen as the only factor of relevance. Interspecific size effects are greater than intraspecific effects. Direct temperature effects tend to be exacerbated by indirect ones, if warming leads to intensified nutrient limitation or copepod grazing while ocean acidification tends to counteract the temperature effect on cell size in non‐calcifying phytoplankton. We discuss the implications of the temperature‐related size trends in a global‐warming context, based on known functional traits associated with phytoplankton size. These are a higher affinity for nutrients of smaller cells, highest maximal growth rates of moderately small phytoplankton (ca. 10² µm³), size‐related sensitivities for different types of grazers, and impacts on sinking rates. For a phytoplankton community increasingly dominated by smaller algae we predict that: (i) a higher proportion of primary production will be respired within the microbial food web; (ii) a smaller share of primary production will be channeled to the classic phytoplankton – crustacean zooplankton – fish food chain, thus leading to decreased ecological efficiency from a fish‐production point of view; (iii) a smaller share of primary production will be exported through sedimentation, thus leading to decreased efficiency of the biological carbon pump.     

Abundance, biomass and small-scale distribution of cryopelagic amphipods in the Franz Josef Land area (Arctic)

Arctic sea ice is inhabited by several amphipod species. Abundance, biomass and small-scale distribution of these cryopelagic (=ice associated) amphipods were investigated near Franz Josef Land in summer 1994. The mean abundance of all species was 420 ind./m²; the mean biomass was 10.61 g ww/m². Gammarus wilkitzkii was the dominant species, whereas Apherusa glacialis, Onisimus nanseni and O. glacialis were only scarcely found. Amphipods were concentrated at the edges of ice floes and were less frequent in areas further away under the ice. The relationship between the distribution and ecological/physiological requirements of cryopelagic amphipods, as well as the small-scale morphology of Arctic sea ice, are discussed. Received: 14 January 1998 / Accepted 14 April 1998     

Identification of the sea ice diatom biomarker IP25 in Arctic benthic macrofauna: direct evidence for a sea ice diatom diet in Arctic heterotrophs

Currently, the impact of declining seasonal sea ice extent in the Arctic on polar food webs remains uncertain. Previously, a range of proxy techniques has been employed to determine links between sea ice or phytoplankton primary production and the Arctic marine food web, although it is accepted that such approaches have their limitations. Here, we propose a novel approach to tracing sea ice primary production through Arctic food webs using the sea ice diatom biomarker, IP₂₅. Various benthic macrofaunal specimens were collected between March and May 2008 from Franklin Bay in the Amundsen Gulf, Arctic Canada, as part of the International Polar Year–Circumpolar Flaw Lead system study. Each specimen was analysed for the presence of the sea ice diatom biomarker IP₂₅ in order to provide evidence for feeding by benthic organisms on sea ice algae. IP₂₅ was found in nineteen out of the twenty-one specimens analysed, often as the most abundant of the highly branched isoprenoid biomarkers detected. The stable isotope composition of IP₂₅ (δ¹³C = −17.1 ± 0.5‰) in the sea urchin (Strongylocentrotus sp.) specimens was similar to that reported previously for this biomarker in Arctic sea ice, sedimenting particles and sediments. It is concluded that detection of IP₂₅ in Arctic benthic macrofauna represents a novel approach to providing convincing evidence for feeding on sea ice algae. It is also proposed that analysis of IP₂₅ may be used to trace trophic transfer of sea ice algal-derived organic matter through Arctic food webs in the future.     

Temporal and spatial trends in marine carbon isotopes in the Arctic Ocean and implications for food web studies

The Arctic is undergoing unprecedented environmental change. Rapid warming, decline in sea ice extent, increase in riverine input, ocean acidification and changes in primary productivity are creating a crucible for multiple concurrent environmental stressors, with unknown consequences for the entire arctic ecosystem. Here, we synthesized 30 years of data on the stable carbon isotope (δ¹³C) signatures in dissolved inorganic carbon (δ¹³C‐DIC; 1977–2014), marine and riverine particulate organic carbon (δ¹³C‐POC; 1986–2013) and tissues of marine mammals in the Arctic. δ¹³C values in consumers can change as a result of environmentally driven variation in the δ¹³C values at the base of the food web or alteration in the trophic structure, thus providing a method to assess the sensitivity of food webs to environmental change. Our synthesis reveals a spatially heterogeneous and temporally evolving δ¹³C baseline, with spatial gradients in the δ¹³C‐POC values between arctic shelves and arctic basins likely driven by differences in productivity and riverine and coastal influence. We report a decline in δ¹³C‐DIC values (?0.011‰ per year) in the Arctic, reflecting increasing anthropogenic carbon dioxide (CO₂) in the Arctic Ocean (i.e. Suess effect), which is larger than predicted. The larger decline in δ¹³C‐POC values and δ¹³C in arctic marine mammals reflects the anthropogenic CO₂ signal as well as the influence of a changing arctic environment. Combining the influence of changing sea ice conditions and isotopic fractionation by phytoplankton, we explain the decadal decline in δ¹³C‐POC values in the Arctic Ocean and partially explain the δ¹³C values in marine mammals with consideration of time‐varying integration of δ¹³C values. The response of the arctic ecosystem to ongoing environmental change is stronger than we would predict theoretically, which has tremendous implications for the study of food webs in the rapidly changing Arctic Ocean.     

Regulation of gene expression is associated with tolerance of the Arctic copepod Calanus glacialis to CO2‐acidified sea water

Ocean acidification is the increase in seawater pCO₂ due to the uptake of atmospheric anthropogenic CO₂, with the largest changes predicted to occur in the Arctic seas. For some marine organisms, this change in pCO₂, and associated decrease in pH, represents a climate change‐related stressor. In this study, we investigated the gene expression patterns of nauplii of the Arctic copepod Calanus glacialis cultured at low pH levels. We have previously shown that organismal‐level performance (development, growth, respiration) of C. glacialis nauplii is unaffected by low pH. Here, we investigated the molecular‐level response to lowered pH in order to elucidate the physiological processes involved in this tolerance. Nauplii from wild‐caught C. glacialis were cultured at four pH levels (8.05, 7.9, 7.7, 7.5). At stage N6, mRNA was extracted and sequenced using RNA‐seq. The physiological functionality of the proteins identified was categorized using Gene Ontology and KEGG pathways. We found that the expression of 151 contigs varied significantly with pH on a continuous scale (93% downregulated with decreasing pH). Gene set enrichment analysis revealed that, of the processes downregulated, many were components of the universal cellular stress response, including DNA repair, redox regulation, protein folding, and proteolysis. Sodium:proton antiporters were among the processes significantly upregulated, indicating that these ion pumps were involved in maintaining cellular pH homeostasis. C. glacialis significantly alters its gene expression at low pH, although they maintain normal larval development. Understanding what confers tolerance to some species will support our ability to predict the effects of future ocean acidification on marine organisms.     

Unusual forest growth decline in boreal North America covaries with the retreat of Arctic sea ice

The 20th century was a pivotal period at high northern latitudes as it marked the onset of rapid climatic warming brought on by major anthropogenic changes in global atmospheric composition. In parallel, Arctic sea ice extent has been decreasing over the period of available satellite data records. Here, we document how these changes influenced vegetation productivity in adjacent eastern boreal North America. To do this, we used normalized difference vegetation index (NDVI) data, model simulations of net primary productivity (NPP) and tree‐ring width measurements covering the last 300 years. Climatic and proxy‐climatic data sets were used to explore the relationships between vegetation productivity and Arctic sea ice concentration and extent, and temperatures. Results indicate that an unusually large number of black spruce (Picea mariana) trees entered into a period of growth decline during the late‐20th century (62% of sampled trees; n = 724 cross sections of age >70 years). This finding is coherent with evidence encoded in NDVI and simulated NPP data. Analyses of climatic and vegetation productivity relationships indicate that the influence of recent climatic changes in the studied forests has been via the enhanced moisture stress (i.e. greater water demands) and autotrophic respiration amplified by the declining sea ice concentration in Hudson Bay and Hudson Strait. The recent decline strongly contrasts with other growth reduction events that occurred during the 19th century, which were associated with cooling and high sea ice severity. The recent decline of vegetation productivity is the first one to occur under circumstances related to excess heat in a 300‐year period, and further culminates with an intensifying wildfire regime in the region. Our results concur with observations from other forest ecosystems about intensifying temperature‐driven drought stress and tree mortality with ongoing climatic changes.     

Observations on the relationship between the Antarctic coastal diatoms Thalassiosira antarctica Comber and Porosira glacialis (Grunow) Jørgensen and sea ice concentrations during the late Quaternary

The available ecological and palaeoecological information for two sea ice-related marine diatoms (Bacillariophyceae), Thalassiosira antarctica Comber and Porosira glacialis (Grunow) Jørgensen, suggests that these two species have similar sea surface temperature (SST), sea surface salinity (SSS) and sea ice proximity preferences. From phytoplankton observations, both are described as summer or autumn bloom species, commonly found in low SST waters associated with sea ice, although rarely within the ice. Both species form resting spores (RS) as irradiance decreases, SST falls and SSS increases in response to freezing ice in autumn. Recent work analysing late Quaternary seasonally laminated diatom ooze from coastal Antarctic sites has revealed that sub-laminae dominated either by T. antarctica RS, or by P. glacialis RS, are nearly always deposited as the last sediment increment of the year, interpreted as representing autumn flux. In this study, we focus on sites from the East Antarctic margin and show that there is a spatial and temporal separation in whether T. antarctica RS or P. glacialis RS form the autumnal sub-laminae. For instance, in deglacial sediments from the Mertz Ninnis Trough (George V Coast) P. glacialis RS form the sub-laminae whereas in similar age sediments from Iceberg Alley (Mac.Robertson Shelf) T. antarctica RS dominate the autumn sub-lamina. In the Dumont d'Urville Trough (Adélie Land), mid-Holocene (Hypsithermal warm period) autumnal sub-laminae are dominated by T. antarctica RS whereas late Holocene (Neoglacial cool period) sub-laminae are dominated by P. glacialis RS. These observations from late Quaternary seasonally laminated sediments would appear to indicate that P. glacialis prefers slightly cooler ocean–climate conditions than T. antarctica. We test this relationship against two down-core Holocene quantitative diatom abundance records from Dumont d'Urville Trough and Svenner Channel (Princess Elizabeth Land) and compare the results with SST and sea ice concentration results of an Antarctic and Southern Ocean Holocene climate simulation that used a coupled atmosphere–sea ice–vegation model forced with orbital parameters and greenhouse gas concentrations. We find that abundance of P. glacialis RS is favoured by higher winter and spring sea ice concentrations and that a climatically-sensitive threshold exists between the abundance of P. glacialis RS and T. antarctica RS in the sediments. An increase to > 0.1 for the ratio of P. glacialis RS:T. antarctica RS indicates a change to increased winter sea ice concentration (to >80% concentration), cooler spring seasons with increased sea ice, slightly warmer autumn seasons with less sea ice and a change from ~ 7.5 months annual sea ice cover at a site to much greater than 7.5 months. In the East Antarctic sediment record, an increase in the ratio from <0.1 to above 0.1 occurs at the transition from the warmer Hypsithermal climate into the cooler Neoglacial climate (~ 4 cal kyr) indicating that the ratio between these two diatoms has the potential to be used as a semi-quantitative climate proxy.     

Sea ice phenology and timing of primary production pulses in the Arctic Ocean

Arctic organisms are adapted to the strong seasonality of environmental forcing. A small timing mismatch between biological processes and the environment could potentially have significant consequences for the entire food web. Climate warming causes shrinking ice coverage and earlier ice retreat in the Arctic, which is likely to change the timing of primary production. In this study, we test predictions on the interactions among sea ice phenology and production timing of ice algae and pelagic phytoplankton. We do so using the following (1) a synthesis of available satellite observation data; and (2) the application of a coupled ice‐ocean ecosystem model. The data and model results suggest that, over a large portion of the Arctic marginal seas, the timing variability in ice retreat at a specific location has a strong impact on the timing variability in pelagic phytoplankton peaks, but weak or no impact on the timing of ice‐algae peaks in those regions. The model predicts latitudinal and regional differences in the timing of ice algae biomass peak (varying from April to May) and the time lags between ice algae and pelagic phytoplankton peaks (varying from 45 to 90 days). The correlation between the time lag and ice retreat is significant in areas where ice retreat has no significant impact on ice‐algae peak timing, suggesting that changes in pelagic phytoplankton peak timing control the variability in time lags. Phenological variability in primary production is likely to have consequences for higher trophic levels, particularly for the zooplankton grazers, whose main food source is composed of the dually pulsed algae production of the Arctic.     

An r-strategist in Antarctic pack ice

Summary The Antarctic copepod Drescheriella glacialis, an inhabitant of sea ice, is the first polar invertebrate metazoan to have been cultured throughout its life cycle. We describe its demographic characteristics on the basis of a laboratory cohort study and correlative field data. When compared to its closest temperate-zone relatives, D. glacialis shows temperature compensation of developmental and reproductive rates. A genuine r-strategist in every respect, it does not fit established trends for Antarctic invertebrates but appears well adapted to the peculiar spatio-temporal variability of the sea ice habitat.     

Variable sea‐ice conditions influence trophic dynamics in an Arctic community of marine top predators

Sea‐ice coverage is a key abiotic driver of annual environmental conditions in Arctic marine ecosystems and could be a major factor affecting seabird trophic dynamics. Using stable isotope ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) in eggs of thick‐billed murres (Uria lomvia), northern fulmars (Fulmarus glacialis), glaucous gulls (Larus hyperboreus), and black‐legged kittiwakes (Rissa tridactyla), we investigated the trophic ecology of prebreeding seabirds nesting at Prince Leopold Island, Nunavut, and its relationship with sea‐ice conditions. The seabird community of Prince Leopold Island had a broader isotopic niche during lower sea‐ice conditions, thus having a more divergent diet, while the opposite was observed during years with more extensive sea‐ice conditions. Species' trophic position was influenced by sea ice; in years of lower sea‐ice concentration, gulls and kittiwakes foraged at higher trophic levels while the opposite was observed for murres and fulmars. For murres and fulmars over a longer time series, there was no evidence of the effect of sea‐ice concentration on species' isotopic niche. Results suggest a high degree of adaptation in populations of high Arctic species that cope with harsh and unpredictable conditions. Such different responses of the community isotopic niche also show that the effect of variable sea‐ice conditions, despite being subtle at the species level, might have larger implications when considering the trophic ecology of the larger seabird community. Species‐specific responses in foraging patterns, in particular trophic position in relation to sea ice, are critical to understanding effects of ecosystem change predicted for a changing climate.     

Osmotic and ionic regulation in response to salinity variations and cold resistance in the Arctic under-ice amphipod <Emphasis Type="Italic">Apherusa glacialis</Emphasis>

Amphipods living at the underside of Arctic sea ice are exposed to varying salinities due to freezing and melting, and have to cope with the resulting osmotic stress. Extracellular osmotic and ionic regulation at different salinities, thermal hysteresis, and supercooling points (SCPs) were studied in the under-ice amphipod Apherusa glacialis. The species is euryhaline, capable to regulate hyperosmotically at salinities S _R < 30 g/kg, and osmoconforms at salinities S _R ≥ 30 g/kg. Hyperosmotic regulation is an adaptation to thrive in low-salinity meltwater below the ice. Conforming to the ambient salinity during freezing reduces the risk of internal ice formation. Thermal hysteresis was not observed in the haemolymph of A. glacialis. The SCP of the species was −7.8 ± 1.9°C. Several ions were specifically downregulated ([Mg²⁺], [SO₄ ²⁻]), or upregulated ([K⁺], [Ca²⁺]) in comparison to the medium. Strong downregulation of [Mg²⁺], is probably necessary to avoid an anaesthetic effect at low temperatures.     

High Arctic summer warming tracked by increased Cassiope tetragona growth in the world's northernmost polar desert

Rapid climate warming has resulted in shrub expansion, mainly of erect deciduous shrubs in the Low Arctic, but the more extreme, sparsely vegetated, cold and dry High Arctic is generally considered to remain resistant to such shrub expansion in the next decades. Dwarf shrub dendrochronology may reveal climatological causes of past changes in growth, but is hindered at many High Arctic sites by short and fragmented instrumental climate records. Moreover, only few High Arctic shrub chronologies cover the recent decade of substantial warming. This study investigated the climatic causes of growth variability of the evergreen dwarf shrub Cassiope tetragona between 1927 and 2012 in the northernmost polar desert at 83°N in North Greenland. We analysed climate–growth relationships over the period with available instrumental data (1950–2012) between a 102‐year‐long C. tetragona shoot length chronology and instrumental climate records from the three nearest meteorological stations, gridded climate data, and North Atlantic Oscillation (NAO) and Arctic Oscillation (AO) indices. July extreme maximum temperatures (JulT_emx), as measured at Alert, Canada, June NAO, and previous October AO, together explained 41% of the observed variance in annual C. tetragona growth and likely represent in situ summer temperatures. JulT_emx explained 27% and was reconstructed back to 1927. The reconstruction showed relatively high growing season temperatures in the early to mid‐twentieth century, as well as warming in recent decades. The rapid growth increase in C. tetragona shrubs in response to recent High Arctic summer warming shows that recent and future warming might promote an expansion of this evergreen dwarf shrub, mainly through densification of existing shrub patches, at High Arctic sites with sufficient winter snow cover and ample water supply during summer from melting snow and ice as well as thawing permafrost, contrasting earlier notions of limited shrub growth sensitivity to summer warming in the High Arctic.