Snow evens fragmentation effects and food determines overwintering success in ground-dwelling voles

Climate instability strongly affects overwintering conditions in organisms living in a strongly seasonal environment and consequently their survival and population dynamics. Food, predation and density effects are also strong during winter, but the effect of fragmentation of ground vegetation on ground-dwelling small mammals is unknown. Here, we report the results of a winter experiment on the effects of habitat fragmentation and food on experimental overwintering populations of bank voles Myodes glareolus. The study was conducted in large outdoor enclosures containing one large, two medium-sized or four small habitat patches or the total enclosure area covered with protective tall-grass habitat. During the stable snow cover of midwinter, only food affected the overwintering success, body condition, trappability and earlier onset of breeding in bank voles. However, after the snow thaw in spring, habitat fragmentation gained importance again, and breeding activities increased the movements of voles in the most fragmented habitat. The use of an open, risky matrix area increased along the habitat fragmentation. Our experiment showed that long-lasting stable snow cover protects overwintering individuals in otherwise exposed and risky ground habitats. We conclude that a stable winter climate and snow cover should even out habitat fragmentation effects on small mammals. However, along prolonged snow-free early winter and in an earlier spring thaw, this means loss of protection by snow cover both in terms of thermoregulation and predation. Thus, habitat cover is important for the survival of small ground-dwelling boreal mammals also during the non-breeding season.     

Demographic responses to a mild winter in enclosed vole populations

Mild winter weather causing snow to melt and ice to accumulate on the ground has been proposed to cause the decreased survival of individuals, and less pronounced cyclicity, of small rodent populations in Fennoscandia. However, detailed data linking ice accumulation to decreased winter survival is lacking. We live-trapped and monitored with passive integrated transponders enclosed populations of root voles (Microtus oeconomus) exposed to different amounts of ice accumulation through a mild winter. We studied how social behaviour and survival responded to snow melt and ice accumulation. Voles avoided ground ice by moving their home ranges, thus increasing home range overlap in enclosed populations experiencing more extensive ice cover. Winter survival was not affected by the amount of ice accumulation, and was only slightly reduced during ice formation in early winter. The lowest survival rates were found at the onset of snow melt in early spring. These results suggest that ice accumulation does not cause lower survival during mild winters, probably because plastic social behaviour enables root voles to reduce the negative effects of varying winter weather on survival. The mechanisms for lower survival during mild winters may operate during spring and be related to spring floods or increased susceptibility to predators. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Winter distribution of small mammals in relation to snow cover in the subalpine zone, Australia

We present the results of a study that is the first to investigate the landscape-scale distribution of small mammals in direct relation to the spatial and temporal variation of snow cover. We investigated the distribution of the dusky antechinus Antechinus swainsonii and bush rat Rattus fuscipes in the subalpine zone of Kosciuszko National Park in the Snowy Mountains of south-eastern Australia. A new hair tube technique was used to detect the presence of small mammals active in the subnivean space. In 2002, we sampled 72 sites stratified by elevation (1501–1600, 1601–1700, 1701–1800 m), aspect (accumulating, ablating) and habitat type (woodland, wet heath, dry heath, grassland). These factors were considered important in influencing snow accumulation and generally reflected the types of habitats available to small mammals. In 2003, the presence of small mammals was investigated at 30 high-elevation sites including six boulderfields. The development of the subnivean space in the Snowy Mountains is dependent on the presence of structures such as shrubs, boulders and microtopographic features that are capable of supporting a snow layer above ground level. When snow was present, small mammal detections were negatively correlated with snow depth and duration, and positively correlated with the complexity of structures and microtopography. At high elevations detections were largely confined to boulderfields, and at mid and low elevations small mammals were detected primarily in habitats where the subnivean space was most extensive. Antechinus swainsonii and R. fuscipes responded differently to snow cover, with the latter seeming better able to overwinter where snow cover was shallow and patchy, in contrast to A. swainsonii whose occurrence was correlated to the size of the subnivean space.     

Measuring the effects of reduced snow cover on Australia's alpine arthropods

Snow is one of the most important factors in the ecology of alpine ecosystems. In Australia, both the depth and duration of snow cover have declined significantly in recent decades and this trend is projected to continue with global warming. Many small arthropods remain active throughout the winter, within a space beneath the snowpack (subnivean) where the snow's insulation creates a thermally stable environment. Using field surveys and experimental manipulation of snow depth at two locations in the Australian alpine region, we explored the diversity of winter‐active arthropods and their response to reduced snow. Individuals from 18 arthropod Orders were detected beneath the snow during winter, with Collembola, Araneae, Acari and Coleoptera accounting for 95–98% of the individuals collected. The subnivean taxa represented a distinct subset of those active outside the winter months. Removal of the snow layer increased daily temperature fluctuations, increased the number of days below freezing and raised the mean surface temperatures. Community composition was altered by snow removal, driven by changes in the numbers of two abundant springtail taxa at each location. We found a strong reduction in the abundances of both taxa at one study site, and contrasting responses (one strong positive and one strong negative) to snow removal at the second study site. Subnivean arthropod communities in Australia thus appear sensitive to snow conditions at small spatial scales.     

Habitat selection,reproduction and predation of wintering lemmings in the Arctic

Snow cover has dramatic effects on the structure and functioning of Arctic ecosystems in winter. In the tundra, the subnivean space is the primary habitat of wintering small mammals and may be critical for their survival and reproduction. We have investigated the effects of snow cover and habitat features on the distributions of collared lemming (Dicrostonyx groenlandicus) and brown lemming (Lemmus trimucronatus) winter nests, as well as on their probabilities of reproduction and predation by stoats (Mustela erminea) and arctic foxes (Vulpes lagopus). We sampled 193 lemming winter nests and measured habitat features at all of these nests and at random sites at two spatial scales. We also monitored overwinter ground temperature at a subsample of nest and random sites. Our results demonstrate that nests were primarily located in areas with high micro-topography heterogeneity, steep slopes, deep snow cover providing thermal protection (reduced daily temperature fluctuations) and a high abundance of mosses. The probability of reproduction increased in collared lemming nests at low elevation and in brown lemming nests with high availability of some graminoid species. The probability of predation by stoats was density dependent and was higher in nests used by collared lemmings. Snow cover did not affect the probability of predation of lemming nests by stoats, but deep snow cover limited predation attempts by arctic foxes. We conclude that snow cover plays a key role in the spatial structure of wintering lemming populations and potentially in their population dynamics in the Arctic.     

The role of ice dynamics in shaping vegetation in flowing waters

Ice dynamics is an important factor affecting vegetation in high‐altitude and high‐latitude streams and rivers. During the last few decades, knowledge about ice in streams and rivers has increased significantly and a respectable body of literature is now available. Here we review the literature on how ice dynamics influence riparian and aquatic vegetation. Traditionally, plant ecologists have focused their studies on the summer period, largely ignoring the fact that processes during winter also impact vegetation dynamics. For example, the freeze‐up period in early winter may result in extensive formation of underwater ice that can restructure the channel, obstruct flow, and cause flooding and thus formation of more ice. In midwinter, slow‐flowing reaches develop a surface‐ice cover that accumulates snow, protecting habitats under the ice from formation of underwater ice but also reducing underwater light, thus suppressing photosynthesis. Towards the end of winter, ice breaks up and moves downstream. During this transport, ice floes can jam up and cause floods and major erosion. The magnitudes of the floods and their erosive power mainly depend on the size of the watercourse, also resulting in different degrees of disturbance to the vegetation. Vegetation responds both physically and physiologically to ice dynamics. Physical action involves the erosive force of moving ice and damage caused by ground frost, whereas physiological effects – mostly cell damage – happen as a result of plants freezing into the ice. On a community level, large magnitudes of ice dynamics seem to favour species richness, but can be detrimental for individual plants. Human impacts, such as flow regulation, channelisation, agriculturalisation and water pollution have modified ice dynamics; further changes are expected as a result of current and predicted future climate change. Human impacts and climate change can both favour and disfavour riverine vegetation dynamics. Restoration of streams and rivers may mitigate some effects of anticipated climate change on ice and vegetation dynamics by, for example, slowing down flows and increasing water depth, thus reducing the potential for massive formation of underwater ice.     

Snow cover manipulations alter survival of early life stages of cold‐temperate tree species

Projections of future climate suggest increases in global temperatures that are especially pronounced in winter in cold‐temperate regions. Thermal insulation provided by snow cover to litter, soil, and overwintering plants will likely be affected by changing winter temperatures and might influence future species composition and ranges. We investigated effects of changing snow cover on seed germination and sapling survival of several cold‐temperate tree species using a snow manipulation approach. Post‐winter seed germination increased or decreased with increasing snow cover, depending on species; decreased seed germination was found in species that characteristically disperse seed in summer or fall months prior to snowfall. Post‐winter sapling survival increased with increasing snow cover for all species, though some species benefitted more from increased snow cover than others. Sapling mortality was associated with root exposure, suggesting the possibility that soil frost heaving could be an important mechanism for observed effects. Our results suggest that altered snow regimes may cause re‐assembly of current species habitat relationships and may drive changes in species’ biogeographic range. However, local snow regimes also vary with associated vegetation cover and topography, suggesting that species distribution patterns may be strongly influenced by spatial heterogeneity in snow regimes and complicating future projections.     

Activity of temperate grassland plants and symbiotic fungi during the winter – implications for community structure and carbon cycling in a changing climate

Several investigations have revealed surprisingly high activities during the winter in vegetation and soil in temperate and subarctic areas. Plants have been found to photosynthesize even under snow cover and at temperatures below freezing, and decomposer microorganisms can function, at low rates, all year around. In temperate grasslands, the vegetation includes winter annual herbs as well as bryophytes, which have the potential to be active and are thus susceptible to changing temperatures during winter. If temperatures stay below freezing and there is a snow cover, an increase in temperatures could in fact decrease the soil temperature due to reduced insulation by snow cover. On the other hand, if winter temperatures initially fluctuate around the freezing point, an increase by a few degrees might produce frost‐free conditions. Based on available data, the composition of plant communities are strongly influenced by temperature conditions in the preceding winter. We conclude that the winter season in grasslands needs more research attention, to start to resolve which species are active and how they respond to a changing climate.     

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.     

Ice‐dependent winter survival of juvenile Atlantic salmon

Changes in snow and ice conditions are some of the most distinctive impacts of global warming in cold temperate and Arctic regions, altering the environment during a critical period for survival for most animals. Laboratories studies have suggested that reduced ice cover may reduce the survival of stream dwelling fishes in Northern environments. This, however, has not been empirically investigated in natural populations in large rivers. Here, we examine how the winter survival of juvenile Atlantic salmon in a large natural river, the River Alta (Norway, 70°N), is affected by the presence or absence of surface ice. Apparent survival rates for size classes corresponding to parr and presmolts were estimated using capture‐mark‐recapture and Cormack‐Jolly‐Seber models for an ice‐covered and an ice‐free site. Apparent survival (Φ) in the ice‐covered site was greater than in the ice‐free site, but did not depend on size class (0.64 for both parr and presmolt). In contrast, apparent survival in the ice‐free site was lower for larger individuals (0.33) than smaller individuals (0.45). The over‐winter decline in storage energy was greater for the ice‐free site than the ice‐covered site, suggesting that environmental conditions in the ice‐free site caused a strong depletion in energy reserves likely affecting survival. Our findings highlight the importance of surface ice for the winter survival of juvenile fish, thus, underpinning that climate change, by reducing ice cover, may have a negative effect on the survival of fish adapted to ice‐covered habitats during winter.     

Effects of ice and floods on vegetation in streams in cold regions: implications for climate change

Riparian zones support some of the most dynamic and species‐rich plant communities in cold regions. A common conception among plant ecologists is that flooding during the season when plants are dormant generally has little effect on the survival and production of riparian vegetation. We show that winter floods may also be of fundamental importance for the composition of riverine vegetation. We investigated the effects of ice formation on riparian and in‐stream vegetation in northern Sweden using a combination of experiments and observations in 25 reaches, spanning a gradient from ice‐free to ice‐rich reaches. The ice‐rich reaches were characterized by high production of frazil and anchor ice. In a couple of experiments, we exposed riparian vegetation to experimentally induced winter flooding, which reduced the dominant dwarf‐shrub cover and led to colonization of a species‐rich forb‐dominated vegetation. In another experiment, natural winter floods caused by anchor‐ice formation removed plant mimics both in the in‐stream and in the riparian zone, further supporting the result that anchor ice maintains dynamic plant communities. With a warmer winter climate, ice‐induced winter floods may first increase in frequency because of more frequent shifts between freezing and thawing during winter, but further warming and shortening of the winter might make them less common than today. If ice‐induced winter floods become reduced in number because of a warming climate, an important disturbance agent for riparian and in‐stream vegetation will be removed, leading to reduced species richness in streams and rivers in cold regions. Given that such regions are expected to have more plant species in the future because of immigration from the south, the distribution of species richness among habitats can be expected to show novel patterns.     

Vegetation recovery following extreme winter warming events in the sub-Arctic estimated using NDVI from remote sensing and handheld passive proximal sensors

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.     

Winter energy requirements of soricine shrews

The energy requirements of shrews under snow cover have not been determined. This is an expository paper attempting to ascertain the daily energy budget of a soricine in subnivean conditions. By means of extrapolation from past work (Morrison, Ryser & Dawe, 1959; Gebczynski, 1965, 1971), a winter individual of Sorex cinereus (mean weight of 2–6 g) in southern Manitoba has a rather high energy budget at 2^oC of about 38-6 kj or 9-2kcal per day. It is assumed that winter prey include invertebrates and small mammals, and their biomass and energy content are estimated. Compensatory factors reducing energy needs are considered, i.e. Dehnel's Phenomenon, extended periods in the nest, low population densities and adequate snow cover. We can probably reduce the above figure for a daily energy budget when considering all the above factors, so that about 30 kj are required daily at the mean subnivean temperature of – 4–5^oC.     

The ice cover on small and large lakes: scaling analysis and mathematical modelling

Lake ice cover is described by its thickness, temperature, stratigraphy and overlying snow layer. When the ratio of ice thickness to lake size is above ～10^?5, the ice cover is stable; otherwise, mechanical forcing breaks the ice cover, and ice drifting takes place with lead-opening and ridging. This transition enables a convenient distinction to be made between small and large lakes. The evolution of the ice cover on small lakes is solved by a wholly thermodynamic model, but a coupled mechanical–thermodynamic model is needed for large lakes. The latter indicates a wide distribution of ice thickness, and frazil ice may be formed in openings. Ecological conditions in large lakes differ markedly from those in small lakes because vertical mixing and oxygen renewal may take place during the ice season, and the euphotic zone penetrates well into the water column in thin ice regions. Mesoscale sea ice models are applicable to large lakes with only minor tuning of the key parameters. These model systems are presented and analysed using Lake Peipsi as an example. As the climate changes, the transition size between small and large lake ice cover will change.     

Experimental icing affects growth,mortality, and flowering in a high Arctic dwarf shrub

Effects of climate change are predicted to be greatest at high latitudes, with more pronounced warming in winter than summer. Extreme mid‐winter warm spells and heavy rain‐on‐snow events are already increasing in frequency in the Arctic, with implications for snow‐pack and ground‐ice formation. These may in turn affect key components of Arctic ecosystems. However, the fitness consequences of extreme winter weather events for tundra plants are not well understood, especially in the high Arctic. We simulated an extreme mid‐winter rain‐on‐snow event at a field site in high Arctic Svalbard (78°N) by experimentally encasing tundra vegetation in ice. After the subsequent growing season, we measured the effects of icing on growth and fitness indices in the common tundra plant, Arctic bell‐heather (Cassiope tetragona). The suitability of this species for retrospective growth analysis enabled us to compare shoot growth in pre and postmanipulation years in icing treatment and control plants, as well as shoot survival and flowering. Plants from icing treatment plots had higher shoot mortality and lower flowering success than controls. At the individual sample level, heavily flowering plants invested less in shoot growth than nonflowering plants, while shoot growth was positively related to the degree of shoot mortality. Therefore, contrary to expectation, undamaged shoots showed enhanced growth in ice treatment plants. This suggests that following damage, aboveground resources were allocated to the few remaining undamaged meristems. The enhanced shoot growth measured in our icing treatment plants has implications for climate studies based on retrospective analyses of Cassiope. As shoot growth in this species responds positively to summer warming, it also highlights a potentially complex interaction between summer and winter conditions. By documenting strong effects of icing on growth and reproduction of a widespread tundra plant, our study contributes to an understanding of Arctic plant responses to projected changes in winter climatic conditions.     

Vegetation controls on northern high latitude snow‐albedo feedback: observations and CMIP5 model simulations

The snow‐masking effect of vegetation exerts strong control on albedo in northern high latitude ecosystems. Large‐scale changes in the distribution and stature of vegetation in this region will thus have important feedbacks to climate. The snow‐albedo feedback is controlled largely by the contrast between snow‐covered and snow‐free albedo (Δα), which influences predictions of future warming in coupled climate models, despite being poorly constrained at seasonal and century time scales. Here, we compare satellite observations and coupled climate model representations of albedo and tree cover for the boreal and Arctic region. Our analyses reveal consistent declines in albedo with increasing tree cover, occurring south of latitudinal tree line, that are poorly represented in coupled climate models. Observed relationships between albedo and tree cover differ substantially between snow‐covered and snow‐free periods, and among plant functional type. Tree cover in models varies widely but surprisingly does not correlate well with model albedo. Furthermore, our results demonstrate a relationship between tree cover and snow‐albedo feedback that may be used to accurately constrain high latitude albedo feedbacks in coupled climate models under current and future vegetation distributions.     

Rapid photosynthetic recovery of a snow-covered feather moss and Peltigera lichen during sub-Arctic midwinter warming

Background: Arctic lichens and mosses are covered by snow for more than half the year and are generally considered as being dormant for most of this period. However, enhanced frequency of winter warming events due to climate change can cause increased disturbance of their protective subnivean environment.

Aim: To further understand cryptogamic responses to midwinter warming we compared the ecophysiological performance of one lichen and one moss species during a simulated warming event.

Methods: We measured photosynthesis and dark respiration in samples of the moss Hylocomium splendens and the lichen Peltigera aphthosa removed from under snow, and on natural refreezing after the warming event, which was simulated by using infrared heaters suspended above the ground.

Results: The moss exposed to light at +5 °C immediately after removal from their subnivean environment and from warmed plots showed positive net gas exchange within 332 s; the lichen required 1238 s. Photosynthesis and nitrogen fixation rates were equal to that, or higher than, during the preceding growing season. Upon refreezing after the event, moss photosynthesis declined considerably.

Conclusions: The moss, and to a lesser extent the lichen, may contribute to subnivean midwinter ecosystem respiration, and both are opportunistic, and can take advantage of warmer winter phases for photosynthesis and growth. This ought to be taken into account in vegetation change projections of cryptogam-rich ecosystems.     

Effects of snow accumulation on survival of beech (Fagus crenata) seed

The escape of beech seeds from seed predators and winter desiccation due to snow accumulation was studied by comparing two sites in Japan: one site that experiences much snow and another site that experiences less snow cover. At the site with greater snow cover, about 70% of the beech seeds escaped seed predation by rodents during winter and about 70% of surviving seeds germinated successfully in spring. At the site with less snow cover, however, all of the beech seeds were eaten by rodents, and all seeds that were protected from feeding were killed by winter desiccation. We confirmed that snow prevents beech seeds from predation by rodents because it conceals their sight and scent. These effects are one of the main reasons why beeches in snowy areas regenerate constantly and those in less snowy areas do not.     

Review of winter trophic relations of soricine shrews

This paper attempts to review all those factors that affect the winter trophic relations of soricines in northern, Holarctic areas, from physiological and ecological aspects to prey and predators. Winter is generally a time of food scarcity and low temperatures, often accompanied by snow cover. Winter shrews are characterized by decreased metabolic rates, longer periods in the nest, short foraging periods and the subsequent decreased need to forage. In smaller species the diet generally consists of epigean fauna, while in larger species hypogean fauna is also included, the proportions of which vary seasonally. In areas of snow cover their diets are derived from the subnivean fauna, i.e. soil and litter fauna, overwintering on the soil surface, and consist of many insects, especially beetles, arachnids, small vertebrates and carrion. Their predators include owls and, to a lesser extent, weasels and foxes.     

Spatio-temporal variability of snow over sea ice in western Hudson Bay,with reference to ringed seal pup survival

Snow-covered sea ice plays a significant role in the ecology of the Arctic marine system and is a critical habitat for ice-adapted ringed seals; however, limited research has focused on the role of snow. The first two objectives of this study characterize the spatial and temporal variability in snow over the sea ice of western Hudson Bay measured from satellite (2002–2010) and how this variability relates to ringed seal pup demographic parameters. The final objective uses a regional circulation model (RCM) to estimate the future snow cover within the study area. Results indicated that the snow cover experienced interannual and interseasonal variability, however, typically increased in late spring. In addition, the Moran’s I statistics indicated clustering at small spatial lags for both seasons, suggesting similar snow depths (i.e., large-scale drifting) occurring at distances of 125 km. There was greater interannual variability in the clustering of snow during spring, compared to the winter season. These trends in snow depth were related to variability in seal pup survival; however, seal pup growth and body condition were not related to winter/spring snow conditions. The results from the RCM member runs suggest that snow will decrease by the end of this century, with a larger decrease occurring in the spring period. In addition, there will also be an increase in interannual and spatial variability during both seasons, which may have significant consequences to ringed seal population abundance through reduced pup survival within the study area.