Winter warming as an important co‐driver for Betula nana growth in western Greenland during the past century

Growing season conditions are widely recognized as the main driver for tundra shrub radial growth, but the effects of winter warming and snow remain an open question. Here, we present a more than 100 years long Betula nana ring‐width chronology from Disko Island in western Greenland that demonstrates a highly significant and positive growth response to both summer and winter air temperatures during the past century. The importance of winter temperatures for Betula nana growth is especially pronounced during the periods from 1910–1930 to 1990–2011 that were dominated by significant winter warming. To explain the strong winter importance on growth, we assessed the importance of different environmental factors using site‐specific measurements from 1991 to 2011 of soil temperatures, sea ice coverage, precipitation and snow depths. The results show a strong positive growth response to the amount of thawing and growing degree‐days as well as to winter and spring soil temperatures. In addition to these direct effects, a strong negative growth response to sea ice extent was identified, indicating a possible link between local sea ice conditions, local climate variations and Betula nana growth rates. Data also reveal a clear shift within the last 20 years from a period with thick snow depths (1991–1996) and a positive effect on Betula nana radial growth, to a period (1997–2011) with generally very shallow snow depths and no significant growth response towards snow. During this period, winter and spring soil temperatures have increased significantly suggesting that the most recent increase in Betula nana radial growth is primarily triggered by warmer winter and spring air temperatures causing earlier snowmelt that allows the soils to drain and warm quicker. The presented results may help to explain the recently observed ‘greening of the Arctic’ which may further accelerate in future years due to both direct and indirect effects of winter warming.     

The interaction between island geomorphology and environmental parameters drives Adélie penguin breeding phenology on neighboring islands near Palmer Station,Antarctica

Despite many studies on Adélie penguin breeding phenology, understanding the drivers of clutch initiation dates (CIDs, egg 1 lay date) is limited or lacks consensus. Here, we investigated Adélie penguin CIDs over 25 years (1991–2016) on two neighboring islands, Torgersen and Humble (<1 km apart), in a rapidly warming region near Palmer Station, Antarctica. We found that sea ice was the primary large‐scale driver of CIDs and precipitation was a secondary small‐scale driver that fine‐tunes CID to island‐specific nesting habitat geomorphology. In general, CIDs were earlier (later) when the spring sea ice retreat was earlier (later) and when the preceding annual ice season was shorter (longer). Island‐specific effects related to precipitation and island geomorphology caused greater snow accumulation and delayed CIDs by ~2 days on Torgersen compared to Humble Island. When CIDs on the islands were similar, conditions were mild with less snow across breeding sites. At Torgersen Island, the negative relationship between CID and breeding success highlights detrimental effects of delayed breeding and/or snow on penguin fitness. Past phenological studies reported a relationship between air temperature and CID, assumed to be related to precipitation, but we found air temperature was more highly correlated to sea ice, revealing a misinterpretation of temperature effects. Finally, contrasting trends in CIDs based on temporal shifts in regional sea ice patterns revealed trends toward earlier CIDs (4–6 day advance) from 1979 to 2009 as the annual ice season shortened, and later CIDs (7–10 day delay) from 2010 to 2016 as the annual ice season lengthened. Adélie penguins tracked environmental conditions with flexible breeding phenology, but their life history remains vulnerable to subpolar weather conditions that can delay CIDs and decrease breeding success, especially on landscapes where geomorphology facilitates snow accumulation.     

The impact of snow accumulation on a heath spider community in a sub-Arctic landscape

Patterns of snow cover across the Arctic are expected to change as a result of shrub encroachment and climate change. As snow cover impacts both the subnivean environment and the date of spring melt, these changes could impact Arctic food webs by altering the phenology and survival of overwintering arthropods, such as spiders (Araneae). In this field study, we used snow fences to increase snow cover across a series of large (375 m²) heath tundra plots and examined the effects on the local spider community during the following growing season. Fences increased snow cover and delayed melt on the treatment plots, paralleling the conditions of nearby shrub sites. Frequent sampling over the season revealed that increased snow cover did not affect spider abundance across different genera nor did it affect overall community composition. Further, our snow treatment did not affect the dates when plots achieved seasonal catch milestones (25, 50, 75 % of total seasonal catch). Increased winter snow cover did, however, produce higher body masses in adults and juveniles of the dominant species Pardosa lapponica (Lycosidae), beginning immediately after snow melt until midway through the growing season. In addition, ovary/oocyte mass of mature P. lapponica females was significantly higher on treatment plots during the peak reproductive period. This is the first experimental manipulation study to report a significant effect of landscape-level changes to winter snow cover on the biomass of an Arctic macroarthropod.     

Phenology and species determine growing‐season albedo increase at the altitudinal limit of shrub growth in the sub‐Arctic

Arctic warming is resulting in reduced snow cover and increased shrub growth, both of which have been associated with altered land surface–atmospheric feedback processes involving sensible heat flux, ground heat flux and biogeochemical cycling. Using field measurements, we show that two common Arctic shrub species (Betula glandulosa and Salix pulchra), which are largely responsible for shrub encroachment in tundra, differed markedly in albedo and that albedo of both species increased as growing season progressed when measured at their altitudinal limit. A moveable apparatus was used to repeatedly measure albedo at six precise spots during the summer of 2012, and resampled in 2013. Contrary to the generally accepted view of shrub‐covered areas having low albedo in tundra, full‐canopy prostrate B. glandulosa had almost the highest albedo of all surfaces measured during the peak of the growing season. The higher midsummer albedo is also evident in localized MODIS albedo aggregated from 2000 to 2013, which displays a similar increase in growing‐season albedo. Using our field measurements, we show the ensemble summer increase in tundra albedo counteracts the generalized effect of earlier spring snow melt on surface energy balance by approximately 40%. This summer increase in albedo, when viewed in absolute values, is as large as the difference between the forest and tundra transition. These results indicate that near future (<50 years) changes in growing‐season albedo related to Arctic vegetation change are unlikely to be particularly large and might constitute a negative feedback to climate warming in certain circumstances. Future efforts to calculate energy budgets and a sensible heating feedback in the Arctic will require more detailed information about the relative abundance of different ground cover types, particularly shrub species and their respective growth forms and phenology.     

Pan‐Arctic sea ice‐algal chl a biomass and suitable habitat are largely underestimated for multiyear ice

There is mounting evidence that multiyear ice (MYI) is a unique component of the Arctic Ocean and may play a more important ecological role than previously assumed. This study improves our understanding of the potential of MYI as a suitable habitat for sea ice algae on a pan‐Arctic scale. We sampled sea ice cores from MYI and first‐year sea ice (FYI) within the Lincoln Sea during four consecutive spring seasons. This included four MYI hummocks with a mean chl a biomass of 2.0 mg/m², a value significantly higher than FYI and MYI refrozen ponds. Our results support the hypothesis that MYI hummocks can host substantial ice‐algal biomass and represent a reliable ice‐algal habitat due to the (quasi‐) permanent low‐snow surface of these features. We identified an ice‐algal habitat threshold value for calculated light transmittance of 0.014%. Ice classes and coverage of suitable ice‐algal habitat were determined from snow and ice surveys. These ice classes and associated coverage of suitable habitat were applied to pan‐Arctic CryoSat‐2 snow and ice thickness data products. This habitat classification accounted for the variability of the snow and ice properties and showed an areal coverage of suitable ice‐algal habitat within the MYI‐covered region of 0.54 million km² (8.5% of total ice area). This is 27 times greater than the areal coverage of 0.02 million km² (0.3% of total ice area) determined using the conventional block‐model classification, which assigns single‐parameter values to each grid cell and does not account for subgrid cell variability. This emphasizes the importance of accounting for variable snow and ice conditions in all sea ice studies. Furthermore, our results indicate the loss of MYI will also mean the loss of reliable ice‐algal habitat during spring when food is sparse and many organisms depend on ice‐algae.     

Predicted responses of arctic and alpine ecosystems to altered seasonality under climate change

Global climate change is already having significant impacts on arctic and alpine ecosystems, and ongoing increases in temperature and altered precipitation patterns will affect the strong seasonal patterns that characterize these temperature‐limited systems. The length of the potential growing season in these tundra environments is increasing due to warmer temperatures and earlier spring snow melt. Here, we compare current and projected climate and ecological data from 20 Northern Hemisphere sites to identify how seasonal changes in the physical environment due to climate change will alter the seasonality of arctic and alpine ecosystems. We find that although arctic and alpine ecosystems appear similar under historical climate conditions, climate change will lead to divergent responses, particularly in the spring and fall shoulder seasons. As seasonality changes in the Arctic, plants will advance the timing of spring phenological events, which could increase plant nutrient uptake, production, and ecosystem carbon (C) gain. In alpine regions, photoperiod will constrain spring plant phenology, limiting the extent to which the growing season can lengthen, especially if decreased water availability from earlier snow melt and warmer summer temperatures lead to earlier senescence. The result could be a shorter growing season with decreased production and increased nutrient loss. These contrasting alpine and arctic ecosystem responses will have cascading effects on ecosystems, affecting community structure, biotic interactions, and biogeochemistry.     

Climate-associated changes in spring plant phenology in China

The timing of phenological events is highly responsive to global environmental change, and shifts in a phenological phase can affect terrestrial ecosystems, agriculture and economics. We analyzed changes in phenology for the spring season in China that occurred between the 1960’s and the 2000’s using four methods: species-level observations, meta-analysis, satellite measurements and phenology modeling. Previous analyses have rarely been reported due to sparse observations. Our results suggest that spring in China has started on average 2.88 days earlier per decade in response to spring warming by −4.93 days per degree Celsius over the last three decades. The shift towards an earlier start of spring was faster in two forest biomes (spring started on average 3.90 days earlier per decade) than in three grassland biomes (spring started on average 0.95 day earlier per decade). This difference was probably due to increased precipitation impacts in the grassland biomes. Interannual variations in the start of spring were most likely attributed to annual fluctuations in spring temperature (∼40%) and in large-scale circulation anomalies (∼20%).     

Changes in satellite‐derived spring vegetation green‐up date and its linkage to climate in China from 1982 to 2010: a multimethod analysis

The change in spring phenology is recognized to exert a major influence on carbon balance dynamics in temperate ecosystems. Over the past several decades, several studies focused on shifts in spring phenology; however, large uncertainties still exist, and one understudied source could be the method implemented in retrieving satellite‐derived spring phenology. To account for this potential uncertainty, we conducted a multimethod investigation to quantify changes in vegetation green‐up date from 1982 to 2010 over temperate China, and to characterize climatic controls on spring phenology. Over temperate China, the five methods estimated that the vegetation green‐up onset date advanced, on average, at a rate of 1.3 ± 0.6 days per decade (ranging from 0.4 to 1.9 days per decade) over the last 29 years. Moreover, the sign of the trends in vegetation green‐up date derived from the five methods were broadly consistent spatially and for different vegetation types, but with large differences in the magnitude of the trend. The large intermethod variance was notably observed in arid and semiarid vegetation types. Our results also showed that change in vegetation green‐up date is more closely correlated with temperature than with precipitation. However, the temperature sensitivity of spring vegetation green‐up date became higher as precipitation increased, implying that precipitation is an important regulator of the response of vegetation spring phenology to change in temperature. This intricate linkage between spring phenology and precipitation must be taken into account in current phenological models which are mostly driven by temperature.     

Greater deciduous shrub abundance extends tundra peak season and increases modeled net CO2 uptake

Satellite studies of the terrestrial Arctic report increased summer greening and longer overall growing and peak seasons since the 1980s, which increases productivity and the period of carbon uptake. These trends are attributed to increasing air temperatures and reduced snow cover duration in spring and fall. Concurrently, deciduous shrubs are becoming increasingly abundant in tundra landscapes, which may also impact canopy phenology and productivity. Our aim was to determine the influence of greater deciduous shrub abundance on tundra canopy phenology and subsequent impacts on net ecosystem carbon exchange (NEE) during the growing and peak seasons in the arctic foothills region of Alaska. We compared deciduous shrub‐dominated and evergreen/graminoid‐dominated community‐level canopy phenology throughout the growing season using the normalized difference vegetation index (NDVI). We used a tundra plant‐community‐specific leaf area index (LAI) model to estimate LAI throughout the green season and a tundra‐specific NEE model to estimate the impact of greater deciduous shrub abundance and associated shifts in both leaf area and canopy phenology on tundra carbon flux. We found that deciduous shrub canopies reached the onset of peak greenness 13 days earlier and the onset of senescence 3 days earlier compared to evergreen/graminoid canopies, resulting in a 10‐day extension of the peak season. The combined effect of the longer peak season and greater leaf area of deciduous shrub canopies almost tripled the modeled net carbon uptake of deciduous shrub communities compared to evergreen/graminoid communities, while the longer peak season alone resulted in 84% greater carbon uptake in deciduous shrub communities. These results suggest that greater deciduous shrub abundance increases carbon uptake not only due to greater leaf area, but also due to an extension of the period of peak greenness, which extends the period of maximum carbon uptake.     

Trophic mismatch and its effects on the growth of young in an Arctic herbivore

In highly seasonal environments, timing of breeding of organisms is typically set to coincide with the period of highest resource availability. However, breeding phenology may not change at a rate sufficient to keep up with rapid changes in the environment in the wake of climate change. The lack of synchrony between the phenology of consumers and that of their resources can lead to a phenomenon called trophic mismatch, which may have important consequences on the reproductive success of herbivores. We analyzed long‐term data (1991–2010) on climate, plant phenology and the reproduction of a long‐distance Arctic migrant, the greater snow goose (Chen caerulescens atlantica), in order to examine the effects of mismatched reproduction on the growth of young. We found that geese are only partially able to adjust their breeding phenology to compensate for annual changes in the timing of high‐quality food plants, leading to mismatches of up to 20 days between the two. The peak of nitrogen concentration in plants, an index of their nutritive quality for goslings, occurred earlier in warm springs with an early snow melt. Likewise, mismatch between hatch dates of young and date of peak nitrogen was more important in years with early snow melt. Gosling body mass and structural size at fledging was reduced when trophic mismatch was high, particularly when the difference between date of peak nitrogen concentration and hatching was >9 days. Our results support the hypothesis that trophic mismatch can negatively affect the fitness of Arctic herbivores and that this is likely to be exacerbated by rising global temperatures.     

Effects of snow removal and algal photoacclimation on growth and export of ice algae

Net growth of ice algae in response to changes in overlying snow cover was studied after manipulating snow thickness on land-fast, Arctic sea ice. Parallel laboratory experiments measured the effect of changing irradiance on growth rate of the ice diatom, Nitzschia frigida. After complete removal of thick snow (≥9 cm), in situ ice algae biomass declined (over 7–12 days), while removal of thin snow layers (4–5 cm), or partial snow removal, increased net algal growth. Ice bottom ablation sometimes followed snow removal, but did not always result in net loss of algae. Similarly, in laboratory experiments, small increases in irradiance increased algal growth rate, while greater light shifts suppressed growth for 3–6 days. However, N. frigida could acclimate to relatively high irradiance (110 μmol photons m² s⁻¹). The results suggest that algal loss following removal of a thick snow layer was due to the combination of photoinhibition and bottom ablation. The smaller relative increase in irradiance after removal of thin or partial snow layers allowed algae to maintain high specific-growth rates that compensated for loss from physical mechanisms. Thus, the response of ice algae to snow loss depends both on the amount of change in snow depth and algal photophysiology. The complex response of ice algae growth and export loss to frequently changing snow fields may contribute to horizontal and temporal patchiness of ecologically and biogeochemically important variables in sea ice and should be considered in predictions of how climate change will affect Arctic marine ecosystems.     

Arctic warming: nonlinear impacts of sea‐ice and glacier melt on seabird foraging

Arctic climate change has profound impacts on the cryosphere, notably via shrinking sea‐ice cover and retreating glaciers, and it is essential to evaluate and forecast the ecological consequences of such changes. We studied zooplankton‐feeding little auks (Alle alle), a key sentinel species of the Arctic, at their northernmost breeding site in Franz‐Josef Land (80°N), Russian Arctic. We tested the hypothesis that little auks still benefit from pristine arctic environmental conditions in this remote area. To this end, we analysed remote sensing data on sea‐ice and coastal glacier dynamics collected in our study area across 1979–2013. Further, we recorded little auk foraging behaviour using miniature electronic tags attached to the birds in the summer of 2013, and compared it with similar data collected at three localities across the Atlantic Arctic. We also compared current and historical data on Franz‐Josef Land little auk diet, morphometrics and chick growth curves. Our analyses reveal that summer sea‐ice retreated markedly during the last decade, leaving the Franz‐Josef Land archipelago virtually sea‐ice free each summer since 2005. This had a profound impact on little auk foraging, which lost their sea‐ice‐associated prey. Concomitantly, large coastal glaciers retreated rapidly, releasing large volumes of melt water. Zooplankton is stunned by cold and osmotic shock at the boundary between glacier melt and coastal waters, creating new foraging hotspots for little auks. Birds therefore switched from foraging at distant ice‐edge localities, to highly profitable feeding at glacier melt‐water fronts within <5 km of their breeding site. Through this behavioural plasticity, little auks maintained their chick growth rates, but showed a 4% decrease in adult body mass. Our study demonstrates that arctic cryosphere changes may have antagonistic ecological consequences on coastal trophic flow. Such nonlinear responses complicate modelling exercises of current and future polar ecosystem dynamics.     

Combined Genetic and Telemetry Data Reveal High Rates of Gene Flow,Migration, and Long-Distance Dispersal Potential in Arctic Ringed Seals (Pusa hispida)

Ringed seals (Pusa hispida) are broadly distributed in seasonally ice covered seas, and their survival and reproductive success is intricately linked to sea ice and snow. Climatic warming is diminishing Arctic snow and sea ice and threatens to endanger ringed seals in the foreseeable future. We investigated the population structure and connectedness within and among three subspecies: Arctic (P. hispida hispida), Baltic (P. hispida botnica), and Lake Saimaa (P. hispida saimensis) ringed seals to assess their capacity to respond to rapid environmental changes. We consider (a) the geographical scale of migration, (b) use of sea ice, and (c) the amount of gene flow between subspecies. Seasonal movements and use of sea ice were determined for 27 seals tracked via satellite telemetry. Additionally, population genetic analyses were conducted using 354 seals representative of each subspecies and 11 breeding sites. Genetic analyses included sequences from two mitochondrial regions and genotypes of 9 microsatellite loci. We found that ringed seals disperse on a pan-Arctic scale and both males and females may migrate long distances during the summer months when sea ice extent is minimal. Gene flow among Arctic breeding sites and between the Arctic and the Baltic Sea subspecies was high; these two subspecies are interconnected as are breeding sites within the Arctic subspecies.     

Denitrification activity and oxygen dynamics in Arctic sea ice

Denitrification and oxygen dynamics were investigated in the sea ice of Franklin Bay (70°N), Canada. These investigations were complemented with measurements of denitrification rates in sea ice from different parts of the Arctic (69°N–85°N). Potential for bacterial denitrification activity (5–194 μmol N m⁻² day⁻¹) and anammox activity (3–5 μmol N m⁻² day⁻¹) in melt water from both first-year and multi-year sea ice was found. These values correspond to 27 and 7%, respectively, of the benthic denitrification and anammox activities in Arctic sediments. Although we report only potential denitrification and anammox rates, we present several indications that active denitrification in sea ice may occur in Franklin Bay (and elsewhere): (1) despite sea ice-algal primary production in the lower sea ice layers, heterotrophic activity resulted in net oxygen consumption in the sea ice of 1–3 μmol l⁻¹ sea ice per day at in situ light conditions, suggesting that O₂ depletion may occur prior to the spring bloom. (2) The ample organic carbon (DOC) and NO₃ ⁻ present in sea ice may support an active denitrification population. (3) Measurements of O₂ conditions in melted sea ice cores showed very low bulk concentrations, and in some cases anoxic conditions prevailed. (4) Laboratory studies using planar optodes for measuring the high-resolution two-dimensional O₂ distributions in sea ice confirmed the very dynamic and heterogeneous O₂ distribution in sea ice, displaying a mosaic of microsites of high and low O₂ concentrations. Brine enclosures and channels were strongly O₂ depleted in actively melting sea ice, and anoxic conditions in parts of the brine system would favour anaerobic processes.     

Linking ice structure and microscale variability of algal biomass in Arctic first-year sea ice using an in situ photographic technique

Microscale photographs were taken of the ice bottom to examine linkages of algal chlorophyll a (chl a) biomass distribution with bottom ice features in thick Arctic first-year sea ice during a spring field program which took place from May 5 to 21, 2003. The photographic technique developed in this paper has resulted in the first in situ observations of microscale variability in bottom ice algae distribution in Arctic first-year sea ice in relation to ice morphology. Observations of brine channel diameter (1.65–2.68 mm) and number density (5.33–10.35 per 100 cm²) showed that the number of these channels at the bottom of thick first-year sea ice may be greater than previously measured on extracted ice samples. A variogram analysis showed that over areas of low chl a biomass (≤20.7 mg chl a m⁻²), patchiness in bottom ice chl a biomass was at the scale of brine layer spacing and small brine channels (∼1–3 mm). Over areas of high chl a biomass (≥34.6 mg chl a m⁻²), patchiness in biomass was related to the spacing of larger brine channels on the ice bottom (∼10–26 mm). Brine layers and channels are thought to provide microscale maxima of light, nutrient replenishment and space availability which would explain the small scale patchiness over areas of low algal biomass. However, ice melt and erosion near brine channels may play a more important role in areas with high algal biomass and low snow cover.     

Ice algal assemblages and vertical export of organic matter from sea ice in the Barents Sea and Nansen Basin (Arctic Ocean)

Algal communities and export of organic matter from sea ice were studied in the offshore marginal ice zone (MIZ) of the northern Barents Sea and Nansen Basin of the Arctic Ocean north of Svalbard by means of ice cores and short-term deployed sediment traps. The observations cover a total of ten stations within the drifting pack ice, visited over a period of 3 years during the period of ice melt in May and July. Maximum flux of particulate organic carbon and chlorophyll a from the ice at 1 m depth (1,537 mg C m⁻² per day and 20 mg Chl a m⁻² per day) exceeded the flux at 30 m by a factor of 2 during spring, a pattern that was reversed later in the season. Although diatoms dominated the ice-associated algal biomass, flagellates at times revealed similarly high biomass and typically dominated the exported algal carbon. Importance of flagellates to the vertical flux increased as melting progressed, whereas diatoms made the highest contribution during the early melting stage. High export of ice-derived organic matter and phytoplankton took place simultaneously in the offshore MIZ, likely as a consequence of ice drift dynamics and the mosaic structure of ice-covered and open water characteristic of this region.     

Demographic response of tundra small mammals to a snow fencing experiment

Snow cover is a key environmental component for tundra wildlife that will be affected by climate change. Change to the snow cover may affect the population dynamics of high‐latitude small mammals, which are active throughout the winter and reproduce under the snow. We experimentally tested the hypotheses that a deeper snow cover would enhance the densities and winter reproductive rates of small mammals, but that predation by mustelids could be higher in areas of increased small mammal density. We enhanced snow cover by setting out snow fences at three sites in the Canadian Arctic (Bylot Island, Nunavut, and Herschel Island and Komakuk Beach, Yukon) over periods ranging from one to four years. Densities of winter nests were higher where snow depth was increased but spring lemming densities did not increase on the experimental areas. Lemmings probably moved from areas of deep snow, their preferred winter habitat, to summer habitat during snow melt once the advantages associated with deep snow were gone. Our treatment had no effect on signs of reproduction in winter nests, proportion of lactating females in spring, or the proportion of juveniles caught in spring, which suggests that deep snow did not enhance reproduction. Results on predation were inconsistent across sites as predation by weasels was higher on the experimental area at one site but lower at two others and was not higher in areas of winter nest aggregations. Although this experiment provided us with several new insights about the impact of snow cover on the population dynamics of tundra small mammals, it also illustrates the challenges and difficulties associated with large‐scale experiments aimed at manipulating a critical climatic factor.     

Novel wildlife in the Arctic: the influence of changing riparian ecosystems and shrub habitat expansion on snowshoe hares

Warming during the 20th century has changed the arctic landscape, including aspects of the hydrology, vegetation, permafrost, and glaciers, but effects on wildlife have been difficult to detect. The primary aim of this study is to examine the physical and biological processes contributing to the expanded riparian habitat and range of snowshoe hares (Lepus americanus) in northern Alaska. We explore linkages between components of the riparian ecosystem in Arctic Alaska since the 1960s, including seasonality of stream flow, air temperature, floodplain shrub habitat, and snowshoe hare distributions. Our analyses show that the peak discharge during spring snowmelt has occurred on average 3.4 days per decade earlier over the last 30 years and has contributed to a longer growing season in floodplain ecosystems. We use empirical correlations between cumulative summer warmth and riparian shrub height to reconstruct annual changes in shrub height from the 1960s to the present. The effects of longer and warmer growing seasons are estimated to have stimulated a 78% increase in the height of riparian shrubs. Earlier spring discharge and the estimated increase in riparian shrub height are consistent with observed riparian shrub expansion in the region. Our browsing measurements show that snowshoe hares require a mean riparian shrub height of at least 1.24–1.36 m, a threshold which our hindcasting indicates was met between 1964 and 1989. This generally coincides with observational evidence we present suggesting that snowshoe hares became established in 1977 or 1978. Warming and expanded shrub habitat is the most plausible reason for recent snowshoe hare establishment in Arctic Alaska. The establishment of snowshoe hares and other shrub herbivores in the Arctic in response to increasing shrub habitat is a contrasting terrestrial counterpart to the decline in marine mammals reliant on decreasing sea ice.     

Notes on the biology of sea ice in the Arctic and Antarctic

The sea ice which covers large areas of the polar regions plays a major role in the marine ecosystem of both the Arctic and Southern Oceans. Not only do warmblooded animals depend on sea ice as a platform, but the sympagic organisms living internally within the sea ice or at the interfaces ice/snow and ice/water provide a substantial part of the total primary production of the ice covered regions. In addition sea ice organisms are an important food source for a variety of pelagic animals and may initiate phytoplankton spring blooms after ice melt by seeding effects.Sea ice organisms often are enriched by some orders of magnitude if the same volume of melted ice is compared to that of the underlying water column. Three hypotheses try to explain this discrepancy and are discussed. Investigations on the nutrient chemistry within the sea ice system and in-situ observations still are rare. Intense growth of sympagic organisms can result in nutrient deficiencies, at least in selected habitats. Advances in endoscopie methods may lead to a better understanding of the life within the sea ice.Paper presented at the Symposium on Polar regions: the challenge for biological and ecological research organised by the Swiss Committee for Polar Research, Basel on 2 October 1992     

Climatic effects on the breeding phenology and reproductive success of an arctic-nesting goose species

Climate warming is pronounced in the Arctic and migratory birds are expected to be among the most affected species. We examined the effects of local and regional climatic variations on the breeding phenology and reproductive success of greater snow geese ( Chen caerulescens atlantica ), a migratory species nesting in the Canadian Arctic. We used a long-term dataset based on the monitoring of 5447 nests and the measurements of 19 234 goslings over 16 years (1989–2004) on Bylot Island. About 50% of variation in the reproductive phenology of individuals was explained by spring climatic factors. High mean temperatures and, to a lesser extent, low snow cover in spring were associated with an increase in nest density and early egg-laying and hatching dates. High temperature in spring and high early summer rainfall were positively related to nesting success. These effects may result from a reduction in egg predation rate when the density of nesting geese is high and when increased water availability allows females to stay close to their nest during incubation recesses. Summer brood loss and production of young at the end of the summer increased when values of the summer Arctic Oscillation (AO) index were either very positive (low temperatures) or very negative (high temperatures), indicating that these components of the breeding success were most influenced by the regional summer climate. Gosling mass and size near fledging were reduced in years with high spring temperatures. This effect is likely due to a reduced availability of high quality food in years with early spring, either due to food depletion resulting from high brood density or a mismatch between hatching date of goslings and the timing of the peak of plant quality. Our analysis suggests that climate warming should advance the reproductive phenology of geese, but that high spring temperatures and extreme values of the summer AO index may decrease their reproductive success up to fledging.