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Many insect species live in the arctic and show a wide range of adaptations to its extreme severity and seasonality. Long, cold winters are met, for example, by cold hardiness and choice of protected sites. Cold hardiness includes both widespread tolerance to freezing and extreme supercooling ability, as well as unusual responses in a few species, such as lack of typical cryoprotectants. Adaptations to short, cool summers include activity at low temperatures, selection of warm habitats and microhabitats, melanism and hairiness coupled with basking behaviour, and prolonged or abbreviated life cycles. Diapause ensures that many species emerge early in summer, with brief synchronized reproduction that maximizes the time for offspring development before winter returns. Some species overwinter in sites that thaw earliest in spring, even if they are relatively exposed in winter. Other adaptations respond to year-to-year variability: for example, prolonged diapause can provide insurance against unsuitable summers. All of these adaptations are co-ordinated. For example, cold hardiness relies on physiological and biochemical adaptations but also on habitat choice and timing. Because the adaptations are complex, predicted climatic warming probably will have unexpected effects. In particular, an increase in temperature that increases summer cloud when sea ice melts would likely reduce temperatures for insect development and activity, because sunshine provides critical warmth to insects and their microhabitats. Changes in moisture will also be important. Moreover, responses differ among species, depending especially on their microhabitats. The complexity of the responses of insects to arctic conditions reinforces the need for research that is sufficiently detailed.
Northern environments present ecological and physiological problems for homeotherms that require adaptations to cope with severe and less predictable physical factors while at the same time continuing to have to cope with the biological ones, such as competition and predation. The stress axis plays a central role in these adaptations and I discuss the range of solutions that birds and mammals have evolved. The stress response in these animals is not static when a challenge occurs, but may be modulated depending on the biological function during the annual cycle (breeding versus nonbreeding), either under-responding to permit reproduction (some song birds) or responding vigorously, yet not having this compromise reproduction (Arctic ground squirrels). Both may trade off survival for reproduction. In contrast, the snowshoe hare shows the expected stress response to chronic high predation risk over 2–3 years: body resources are geared to survival and reproduction is inhibited. Two long term, persistent, and pervasive changes will confront northern birds and mammals in the 21st century: global change and persistent organochlorine pollutants (POPs). These may result in either adaptations or shifts in distribution and abundance. For the former, latitudinal variation in the stress axis may help song birds respond rapidly; population variation in the stress axis response is unknown in northern mammals and relatively sedentary mammals may be unable to shift their distribution rapidly to adjust major climate shifts. For the latter, the few POPs studies that have examined the stress axis indicate marked negative effects.
Important drivers for emergence of infectious disease in wildlife include changes in the environment, shrinking habitats or concentration of wildlife, and movement of people, animals, pathogens, or vectors. In this paper we present three case-studies of emerging parasitic infections and diseases in ungulates in the Canadian north. First we discuss climate warming as an important driver for the emergence of disease associated with Umingmakstrongylus pallikuukensis, a nematode lungworm of muskoxen. Then we examine how Protostrongylus stilesi, the sheep lungworm, emerged (or re-emerged) in muskoxen after re-introduction of this host into its historical range made it sympatric with Dall's sheep. Finally, we consider Teladorsagia boreoarcticus, a newly described and common abomasal nematode of muskoxen that is emerging as a disease-causing parasite and may be an important regulator for muskox populations on Banks Island, Northwest Territories. These and other arctic host-parasite systems are exquisitely tuned and constrained by a harsh and highly seasonal environment. The dynamics of these systems will be impacted by climate change and other ecological disruptions. Baseline knowledge of parasite biodiversity and parasite and host ecology, together with predictive models and long-term monitoring programs, are essential for anticipating and detecting altered patterns of host range, geographic distribution, and the emergence of parasitic infections and diseases.
We examined the role of trophic interactions in structuring a high arctic tundra community characterized by a large breeding colony of greater snow geese (Chen caerulescens atlantica). According to the exploitation ecosystem hypothesis of Oksanen et al. (1981), food chains are controlled by top-down interactions. However, because the arctic primary productivity is low, herbivore populations are too small to support functional predator populations and these communities should thus be dominated by the plant/ herbivore trophic-level interaction. Since 1990, we have been monitoring annual abundance and productivity of geese, the impact of goose grazing, predator abundance (mostly arctic foxes, Alopex lagopus) and the abundance of lemmings, the other significant herbivore in this community, on Bylot Island, Nunavut, Canada. Goose grazing consistently removed a significant proportion of the standing crop (∼40%) in tundra wetlands every year. Grazing changed plant community composition and reduced the production of grasses and sedges to a low-level equilibrium compared to the situation where the presence of geese had been removed. Lemming cyclic fluctuations were strong and affected fox reproduction. Fox predation on goose eggs was severe and generated marked annual variation in goose productivity. Predation intensity on geese was closely related to the lemming cycle, a consequence of an indirect interaction between lemming and geese via shared predators. We conclude that, contrary to the exploitation ecosystem hypothesis, both the plant/herbivore and predator/prey interactions are significant in this arctic community.
An allochthonous input can modify trophic relationships, by providing an external resource that is normally limiting within a system. The subsidy may not only elicit a growth response of the primary producers via a bottom-up effect, but it also may lead to runaway herbivore growth in the absence of increased predation. If the consumer is migratory and predation is similarly dampened in the alternative system, the increased numbers may produce a top-down cascade of direct and indirect effects on an ecosystem that may be a great distance from the source of the subsidy. In an extreme case, it can lead to a catastrophic shift in ecosystem functioning as a result of biotic exploitation that produces an alternative stable state. The loss of resilience is particularly sensitive to herbivore density which can result in two different outcomes to the vegetation on which the consumer feeds. Over-compensatory growth of above-ground biomass gives way to sward destruction and near irreversible changes in soil properties as density of a herbivore increases. A striking temporal asymmetry exists between a reduction in the consumer population and recovery of damaged vegetation and degraded soils.
Adaptations to the cold and to short growing seasons characterize arctic life, but climate in the Arctic is warming at an unprecedented rate. Will plant and animal populations of the Arctic be able to cope with these drastic changes in environmental conditions? Here we explore the potential contribution of evolution by natural selection to the current response of populations to climate change. We focus on the spring phenology of populations because it is highly responsive to climate change and easy to document across a wide range of species. We show that evolution can be fast and can occur at the time scale of a few decades. We present an example of reproductive phenological change associated with climate change (North American red squirrels in the Yukon), where a detailed analysis of quantitative genetic parameters demonstrates contemporary evolution. We answer a series of frequently asked questions that should help biologists less familiar with evolutionary theory and quantitative genetic methods to think about the role of evolution in current responses of ecological systems to climate change. Our conclusion is that evolution by natural selection is a pertinent force to consider even at the time scale of contemporary climate changes. However, all species may not be equal in their capacity to benefit from contemporary evolution.
Climate change will likely alter the distribution and abundance of northern mammals through a combination of direct, abiotic effects (e.g., changes in temperature and precipitation) and indirect, biotic effects (e.g., changes in the abundance of resources, competitors, and predators). Bioenergetic approaches are ideally suited to predicting the impacts of climate change because individual energy budgets integrate biotic and abiotic influences, and translate individual function into population and community outcomes. In this review, we illustrate how bioenergetics can be used to predict the regional biodiversity, species range limits, and community trophic organization of mammals under future climate scenarios. Although reliable prediction of climate change impacts for particular species requires better data and theory on the physiological ecology of northern mammals, two robust hypotheses emerge from the bioenergetic approaches presented here. First, the impacts of climate change in northern regions will be shaped by the appearance of new species at least as much as by the disappearance of current species. Second, seasonally inactive mammal species (e.g., hibernators), which are largely absent from the Canadian arctic at present, should undergo substantial increases in abundance and distribution in response to climate change, probably at the expense of continuously active mammals already present in the arctic.
Polar bears (Ursus maritimus) live throughout the ice-covered waters of the circumpolar Arctic, particularly in near shore annual ice over the continental shelf where biological productivity is highest. However, to a large degree under scenarios predicted by climate change models, these preferred sea ice habitats will be substantially altered. Spatial and temporal sea ice changes will lead to shifts in trophic interactions involving polar bears through reduced availability and abundance of their main prey: seals. In the short term, climatic warming may improve bear and seal habitats in higher latitudes over continental shelves if currently thick multiyear ice is replaced by annual ice with more leads, making it more suitable for seals. A cascade of impacts beginning with reduced sea ice will be manifested in reduced adipose stores leading to lowered reproductive rates because females will have less fat to invest in cubs during the winter fast. Non-pregnant bears may have to fast on land or offshore on the remaining multiyear ice through progressively longer periods of open water while they await freeze-up and a return to hunting seals. As sea ice thins, and becomes more fractured and labile, it is likely to move more in response to winds and currents so that polar bears will need to walk or swim more and thus use greater amounts of energy to maintain contact with the remaining preferred habitats. The effects of climate change are likely to show large geographic, temporal and even individual differences and be highly variable, making it difficult to develop adequate monitoring and research programs. All ursids show behavioural plasticity but given the rapid pace of ecological change in the Arctic, the long generation time, and the highly specialised nature of polar bears, it is unlikely that polar bears will survive as a species if the sea ice disappears completely as has been predicted by some.
As ground nesting homeotherms, alpine and arctic birds must meet similar physiological requirements for breeding as other birds, but must do so in more extreme conditions. Annual spring snowfall and timing of snow melt can vary by up to 1 month and daily temperatures near the ground surface vary from below freezing to over 45°C in alpine and arctic habitats. Species breeding in these environments have various behavioral, physiological, and morphological adaptations to cope with energetically demanding conditions. We review the ways birds cope with harsh and variable weather, and present data from long term field studies of ptarmigan to examine effects of spring weather on reproduction. In variable but normal spring conditions, timing of breeding was not influenced by snow melt, snow depth or daily temperatures in the alpine, as breeding did not commence until conditions were generally favorable. Arctic ptarmigan tended to vary breeding onset in response to spring conditions. Generally, birds breeding in alpine and arctic habitats suffer a seasonal reproductive disadvantage compared to birds at lower latitudes or elevations because the breeding window is short and in late years, nest failure may be high with little opportunity for renesting. Coping mechanisms may only be effective below a threshold of climactic extremes. Despite strong resilience in fecundity parameters, when snowmelt is extremely delayed breeding success is greatly reduced. Alpine and arctic birds will be further challenged as they attempt to cope with anticipated increases in the frequency and severity of weather events (climate variability), as well as general climate warming.
The length of the snow-free season has a significant influence on reproduction and growth in northern alpine environments, and these life history traits may provide sensitive indicators of the responses of organisms to climate change. We examined growth rates and timing of parturition of collared pikas (Ochotona collaris) from 1995–2002 in the Ruby Range, Yukon Territory, Canada. Growth rates were best described using a Gompertz model, in which the asymptotic mass, determined from the average male and female weights, was 157 g, the growth rate constant (K) was 0.0557, and the age at inflection (I) was 18.12 days, for a birth weight of 10 g. The maximum growth rate for North American pikas (O. collaris and O. princeps) increased with latitude, with maximum growth rates being approximately one-third greater in northern populations where the snow-free season is less than three months long. The mean parturition date varied significantly among years from 3 June to 3 July, and delayed parturition was correlated with indices of high snow accumulation and, to a lesser extent, late spring snowmelt. However, parturition date did not significantly affect the subsequent over-winter survival of juveniles in this population, suggesting that pikas are able to adjust to seasonal uncertainty associated with highly variable spring conditions.