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Baseline studies of small rodent populations in undisturbed ecosystems are rare. We report here 50 years of monitoring and experimentation in Yukon of a dominant rodent species in the North American boreal forest, the red‐backed vole Clethrionomys rutilus. These voles breed in summer, weigh 20–25 g, and reach a maximum density of 20 to 25 per ha. Their populations have shown consistent 3–4‐year cycles for the last 50 years with the only change being that peak densities averaged 8/ha until 2000 and 18/ha since that year. During the last 25 years, we have measured food resources, predator numbers, and winter weather, and for 1‐year social interactions, to estimate their contribution to changes in the rate of summer increase and the rate of overwinter decline. All these potential limiting factors could contribute to changes in density, and we measured their relative contributions statistically with multiple regressions. The rate of winter decline in density was related to both food supply and winter severity. The rate of summer increase was related to summer berry crops and white spruce cone production. No measure of predator numbers was related to winter or summer changes in vole abundance. There was a large signal of climate change effects in these populations. There is no density dependence in summer population growth and only a weak one in winter population declines. None of our results provide a clear understanding of what generates 3–4‐year cycles in these voles, and the major missing piece may be an understanding of social interactions at high density. [ABSTRACT FROM AUTHOR]

Wild mammal populations exhibit a variety of dynamics, ranging from fairly stable with little change in population size over time to high‐amplitude cyclic or erratic fluctuations. A persistent question in population ecology is why populations fluctuate as they do. Answering this seemingly simple question has proven to be challenging. Broadly, density‐dependent feedback mechanisms should allow populations to grow at low density and slow or halt growth at high density. However, experimental tests of what demographic processes result in density‐dependent feedback and on what timescale have proven elusive. Here, we used replicated density perturbation experiments and capture‐mark‐recapture analyses to test density‐dependent population growth in populations of meadow voles (Microtus pennsylvanicus) during the summer breeding season by manipulating founding population density and observing the pattern of survival, reproduction, and population growth. High population density had no consistent effect on survival rates but generally negatively influenced recruitment and population growth rates. However, these density‐dependent effects varied within the breeding season and across years. Our study provides evidence that density‐dependent feedback mechanisms operate at finer time scales than previously believed and that process, additively with delayed year effects, is key to understanding multiyear population demography. [ABSTRACT FROM AUTHOR]

Climate warming is causing asynchronies between animal phenology and environments. Mismatched traits, such as coat color change mismatched with snow, can decrease survival. However, coat change does not serve a singular adaptive benefit of camouflage, and alternate coat change functions may confer advantages that supersede mismatch costs. We found that mismatch reduced, rather than increased, autumn mortality risk of snowshoe hares in Yukon by 86.5% when mismatch occurred. We suggest that the increased coat insulation and lower metabolic rates of winter‐acclimatized hares confer energetic advantages to white mismatched hares that reduce their mortality risk. We found that white mismatched hares forage 17–77 min less per day than matched brown hares between 0°C and −10°C, thus lowering their predation risk and increasing survival. We found no effect of mismatch on spring mortality risk, during which mismatch occurred at warmer temperatures, suggesting a potential temperature limit at which the costs of conspicuousness outweigh energetic benefits. [ABSTRACT FROM AUTHOR]

Snowshoe hare cycles are one of the most prominent phenomena in ecology. Experimental studies point to predation as the dominant driving factor, but previous experiments combining food supplementation and predator removal produced unexplained multiplicative effects on density. We examined the potential interactive effects of food limitation and predation in causing hare cycles using an individual‐based food‐supplementation experiment over‐winter across three cycle phases that naturally varied in predation risk. Supplementation doubled over‐winter survival with the largest effects occurring in the late increase phase. Although the proximate cause of mortality was predation, supplemented hares significantly decreased foraging time and selected for conifer habitat, potentially reducing their predation risk. Supplemented hares also lost less body mass which resulted in the production of larger leverets. Our results establish a mechanistic link between how foraging time, mass loss and predation risk affect survival and reproduction, potentially driving demographic changes associated with hare cycles. [ABSTRACT FROM AUTHOR]

A large body of research shows that maternal stress during an offspring's early life can impact its phenotype in both the short and long term. In the Vertebrata, most research has been focused on maternal stress during the prenatal period. However, the postnatal period is particularly important in mammals because maternal milk provides a conduit by which maternal hormones secreted in response to stressors (glucocorticoids, GCs) can reach offspring. Moreover, lactation outlasts gestation in many species.Though GCs were first detected in milk over 40 years ago, few studies have explored how they affect nursing offspring, and no reviews have been written on how maternal stress affects nursing offspring in the natural world.We discuss the evolution of milk and highlight its importance in each of the three mammalian lineages: monotremes (subclass Monotremata), marsupials (infraclass Marsupialia), and eutherians (infraclass Placentalia). Most research on the effects of milk GCs on offspring has been focused on eutherians, but monotremes and marsupials rely on their mothers' milk for a proportionally longer period of time, and so research on these taxa may yield more insight.We show that GCs are important for milk production, both during an individual nursing bout and over the entire lactation period, and review evidence of GCs moving from maternal blood to milk, and eventually to nursing offspring. We examine evidence from rodents and primates of associations between GC levels in lactating females (either blood or milk) and offspring behaviour and growth rates. We discuss ways that maternal stress may impact these offspring phenotypes outside of milk GCs, such as changes to: (1) milk output, (2) other milk constituents (e.g. macronutrients, growth factors, cytokines), and (3) maternal care behaviour.Critical to understanding the fitness impacts of elevated maternal GC levels during lactation is to place this within the context of the natural environment. Species‐specific traits and natural histories will help us to understand why such maternal stress produces different offspring phenotypes that equip them to cope with and succeed in the environment they are about to enter. [ABSTRACT FROM AUTHOR]

Vertebrate scavenging can impact food web dynamics, but our understanding of this process stems predominantly from monitoring herbivore carrion and extrapolating results across carcass types. Recent evidence suggests carnivores may avoid intraguild scavenging to reduce parasite transmission. If this behavior is widespread across diverse ecosystems, estimation of nutrient cycling and community scavenging rates are likely biased to a currently unknown degree. We examined whether the time to initiate scavenging, carcass persistence, or the richness of species scavenging in the boreal forest of Yukon, Canada, differed between carnivore and herbivore carcasses. Vertebrates took longer to initiate scavenging on carnivore carcasses (3.2 d) relative to herbivore carcasses (1.1 d), and carnivore carcasses persisted on the landscape for over a month longer (48.4 d and 5.5 d, respectively). The longer persistence times were due to the reduction in scavenging by carnivores such as Canada lynx (Lynx canadensis). Decreased scavenging was caused by changes in the propensity to consume carnivore carrion, as the number of species detecting a carcass within the first week did not differ between carnivore and herbivore carcasses. These results have ramifications for our understanding of nutrient cycling and food web dynamics in the boreal forest and provide further support that carcass type should be included in future studies. [ABSTRACT FROM AUTHOR]

Population and community ecology as a science are about 100 years old, and we discuss here our opinion of what approaches have progressed well and which point to possible future directions. The three major threads within population and community ecology are theoretical ecology, statistical tests and models, and experimental ecology. We suggest that our major objective is to understand what factors determine the distribution and abundance of organisms within populations and communities, and we evaluate these threads against this major objective. Theoretical ecology is elegant and compelling and has laid the groundwork for achieving our overall objectives with useful simple models. Statistics and statistical models have contributed informative methods to analyze quantitatively our understanding of distribution and abundance for future research. Population ecology is difficult to carry out in the field, even though we may have all the statistical methods and models needed to achieve results. Community ecology is growing rapidly with much description but less understanding of why changes occur. Biodiversity science cuts across all these subdivisions but rarely digs into the necessary population and community science that might solve conservation problems. Climate change affects all aspects of ecology but to assume that everything in population and community ecology is driven by climate change is oversimplified. We make recommendations on how to advance the field with advice for present and future generations of population and community ecologists. [ABSTRACT FROM AUTHOR]

Nearly 100 years ago, Charles Elton described lemming and vole population cycles as ecological models for understanding population regulation in nature. Yet, the mechanisms driving these cycles are still not fully understood. These rodent populations can continue to cycle in the absence of predation and with food supplementation, and represent a major unsolved problem in population ecology.It has been hypothesized that the social environment at high population density can drive selection for a low‐reproduction phenotype, resulting in population self‐regulation as an intrinsic mechanism driving the cycles. However, a physiological mechanism for this self‐regulation has not been demonstrated. We manipulated population density in wild meadow voles Microtus pennsylvanicus using large‐scale field enclosures over 3 years and examined reproductive performance and physiology.Within the field enclosures, we assessed the proportion of breeding animals, mass at sexual maturation, and faecal androgen and oestrogen metabolites. We then collected brain tissue from juvenile voles born at high or low density, quantified mRNA expression of gonadotropin‐releasing hormone (GnRH) and oestrogen receptor alpha (ERα) and measured DNA methylation at six CpG sites in a region that was highly conserved with the mouse GnRH promoter.At high density, there was a lower proportion of reproductive animals. Juvenile voles born at high densities had reduced expression of GnRH in the hypothalamus, accompanied by marginally lower faecal sex hormone metabolites. Female juvenile voles born at high density also had higher methylation levels at two CpG sites while males did not, aligning with prior observations that females (but not males) from high‐density environments retain reduced reproduction long term.Our results support a physiological basis for population self‐regulation in vole cycles, as altering population density alone induced reproductive downregulation at the hypothalamic level. Our results demonstrate that altering the early‐life social environment can fundamentally impact reproductive function in the brain. This, in turn, can drive population demography changes in wild animals. [ABSTRACT FROM AUTHOR]

Vertebrates have high species‐level variation in circulating hormone concentrations, and the functional significance of this variation is largely unknown. We tested the hypothesis that interspecific differences in hormone concentrations are partially driven by plant consumption, based on the prediction that herbivores should have higher basal hormone levels to 'outcompete' plant endocrine disruptors. We compared levels of glucocorticoids (GCs), the hormones with the most available data, across 166 species. Using phylogenetically informed comparisons, we found that herbivores had higher GC levels than carnivores. Furthermore, we found that the previously described negative relationship between GC levels and body mass only held in herbivores, not carnivores, and that the effect of diet was greatest at extreme body sizes. These findings demonstrate the far‐reaching effects of diet on animal physiology, and provide evidence that herbivory influences circulating hormone concentrations. We urge future direct testing of the relationship between phytochemical load and GC levels. [ABSTRACT FROM AUTHOR]

Scavenging by vertebrates can have important impacts on food web stability and persistence, and can alter the distribution of nutrients throughout the landscape. However, scavenging communities have been understudied in most regions around the globe, and we lack understanding of the biotic drivers of vertebrate scavenging dynamics.In this paper, we examined how changes in prey density and carrion biomass caused by population cycles of a primary prey species, the snowshoe hare Lepus americanus, influence scavenging communities in the northern boreal forest. We further examined the impact of habitat and temperature on scavenging dynamics.We monitored the persistence time, time until first scavenger, and number of species scavenging experimentally‐placed hare carcasses over four consecutive years in the southwestern Yukon. We simultaneously monitored hare density and carrion biomass to examine their influence relative to temperature, habitat, and seasonal effects. For the primary scavengers, we developed species‐specific scavenging models to determine variation on the effects of these factors across species, and determine which species may be driving temporal patterns in the entire community.We found that the efficiency of the scavenging community was affected by hare density, with carcass persistence decreasing when snowshoe hare densities declined, mainly due to increased scavenging rates by Canada lynx Lynx canadensis. However, prey density did not influence the number of species scavenging a given carcass, suggesting prey abundance affects carrion recycling but not necessarily the number of connections in the food web. In addition, scavenging rates increased in warmer temperatures, and there were strong seasonal effects on the richness of the vertebrate scavenging community.Our results demonstrate that vertebrate scavenging communities are sensitive to changes in species' demography and environmental change, and that future assessments of food web dynamics should consider links established through scavenging. [ABSTRACT FROM AUTHOR]

Arctic ecosystems are changing rapidly in response to climate warming. While Arctic mammals are highly evolved to these extreme environments, particularly with respect to their stress axis, some species may have limited capacity to adapt to this change. We examined changes in key components of the stress axis (cortisol and its carrier protein—corticosteroid binding globulin [CBG]) in polar bears (Ursus maritimus) from western Hudson Bay (N = 300) over a 33 year period (1983–2015) during which time the ice‐free period was increasing. Changing sea ice phenology limits spring hunting opportunities and extends the period of onshore fasting. We assessed the response of polar bears to a standardized stressor (helicopter pursuit, darting, and immobilization) during their onshore fasting period (late summer–autumn) and quantified the serum levels of the maximum corticosteroid binding capacity (MCBC) of CBG, the serum protein that binds cortisol strongly, and free cortisol (FC). We quantified bear condition (age, sex, female with cubs or not, fat condition), sea ice (breakup in spring–summer, 1 year lagged freeze‐up in autumn), and duration of fasting until sample collection as well as cumulative impacts of the latter environmental traits from the previous year. Data were separated into "good" years (1983–1990) when conditions were thought to be optimal and "poor" years (1991–2015) when sea ice conditions deteriorated and fasting on land was extended. MCBC explained 39.4% of the variation in the good years, but only 28.1% in the poor ones, using both biological and environmental variables. MCBC levels decreased with age. Changes in FC were complex, but more poorly explained. Counterintuitively, MCBC levels increased with increased time onshore, 1 year lag effects, and in poor ice years. We conclude that MCBC is a biomarker of stress in polar bears and that the changes we document are a consequence of climate warming. [ABSTRACT FROM AUTHOR]

Coyotes are one of the major predators of snowshoe hares in the Yukon boreal forest. Lynx are the classic major predator of snowshoe hares because they are well adapted to cold and their large feet are effective snowshoes. Photo credit: Alice J. Kenney. gl GLO:69VH/01apr20:bes21678-fig-0003.jpg PHOTO (COLOR): Snowshoe hare (Lepus americanus). [Extracted from the article]

Anogenital distance (AGD) is positively correlated to fetal androgen exposure and developmental masculinization in mammals. Independent of overall body size, AGD shows a strong positive correlation with male fertility and in rodents, AGD is a good indicator of male competitive ability and is associated with female choice. We hypothesized that AGD will also predict male competitive ability in non‐human primates. To test this, we measured AGD noninvasively with a parallel laser in a wild population of Angolan colobus monkeys (Colobus angolensis ruwenzorii) in Uganda and correlated to it to their social structure. C. angolensis ruwenzorii form a multilevel society with both one‐male/multifemale units (OMUs) and multimale/multifemale units (MMUs). We compared AGD in males from five OMUs and six MMUs and related it to their fecal androgen metabolite concentrations, dominance rank and body size, and to the number of females in their unit. Males in OMUs had greater access to females, so were predicted to have longer AGDs, but this was not found. AGD also did not correlate overall with mean fecal androgen metabolites in MMUs. However, AGD was correlated with dominance rank in MMUs, demonstrating that higher‐ranking males in these multimale units had longer AGDs. Body size did not show the same relationship with dominance rank, suggesting that male rank was not just a reflection of absolute male size. Our findings indicate that AGD predicts male competitive ability in this species and that it may be a useful correlate throughout the non‐human primates. These results also support the idea that prenatal androgen exposure increases the likelihood of the expression of behaviors that maintain high dominance rank. Research Highlights: Anogenital distance (AGD) is an accurate measure of androgen exposure in utero.In multimale colobus core units, male AGD was correlated with dominance rank.This suggests that AGD is a useful measure of competitive ability in non‐human primates. [ABSTRACT FROM AUTHOR]

Cyclic fluctuations in abundance exhibited by some mammalian populations in northern habitats ("population cycles") are key processes in the functioning of many boreal and tundra ecosystems. Understanding population cycles, essentially demographic processes, necessitates discerning the demographic mechanisms that underlie numerical changes. Using mark–recapture data spanning five population cycles (1977–2017), we examined demographic mechanisms underlying the 9–10‐yr cycles exhibited by snowshoe hares (Lepus americanus Erxleben) in southwestern Yukon, Canada. Snowshoe hare populations always decreased during winter and increased during summer; the balance between winter declines and summer increases characterized the four, multiyear cyclic phases: increase, peak, decline, and low. Little or no recruitment occurred during winter, but summer recruitment varied markedly across the four phases with the highest and lowest recruitment observed during the increase and decline phase, respectively. Population crashes during the decline were triggered by a substantial decline in winter survival and by a lack of subsequent summer recruitment. In contrast, initiation of the increase phase was triggered by a twofold increase in summer recruitment abetted secondarily by improvements in subsequent winter survival. We show that differences in peak density across cycles are explained by differences in overall population growth rate, amount of time available for population growth to occur, and starting population density. Demographic mechanisms underlying snowshoe hare population cycles were consistent across cycles in our study site but we do not yet know if similar demographic processes underlie population cycles in other northern snowshoe hare populations. [ABSTRACT FROM AUTHOR]

Long‐term monitoring is critical to determine the stability and sustainability of wildlife populations, and if change has occurred, why. We have followed population density changes in the small mammal community in the boreal forest of the southern Yukon for 46 years with density estimates by live trapping on 3–5 unmanipulated grids in spring and autumn. This community consists of 10 species and was responsible for 9% of the energy flow in the herbivore component of this ecosystem from 1986 to 1996, but this increased to 38% from 2003 to 2014. Small mammals, although small in size, are large in the transfer of energy from plants to predators and decomposers. Four species form the bulk of the biomass. There was a shift in the dominant species from the 1970s to the 2000s, with Myodes rutilus increasing in relative abundance by 22% and Peromyscus maniculatus decreasing by 22%. From 2007 to 2018, Myodes comprised 63% of the catch, Peromyscus 20%, and Microtus species 17%. Possible causes of these changes involve climate change, which is increasing primary production in this boreal forest, and an associated increase in the abundance of 3 rodent predators, marten (Martes americana), ermine (Mustela ermine) and coyotes (Canis latrans). Following and understanding these and potential future changes will require long‐term monitoring studies on a large scale to measure metapopulation dynamics. The small mammal community in northern Canada is being affected by climate change and cannot remain stable. Changes will be critically dependent on food–web interactions that are species‐specific. [ABSTRACT FROM AUTHOR]

In contrast, binding by CBG is termed "specific binding" because CBG tightly binds GC with high specificity but just one GC molecule per CBG molecule (Hammond I et al i . For example, when GC concentrations increase in response to a stressor, a species with high affinity CBG will initially bind more of the released GC than a species with lower affinity CBG, all other things being equal. However, if a species with lower affinity CBG has more CBG relative to their GC concentrations than does a high affinity species, then both would experience the more gradual increase in free GC. [Extracted from the article]

Abstract: In recent years, it has been argued that the effect of predator fear exacts a greater demographic toll on prey populations than the direct killing of prey. However, efforts to quantify the effects of fear have primarily relied on experiments that replace predators with predator cues. Interpretation of these experiments must consider two important caveats: (1) the magnitude of experimenter‐induced predator cues may not be realistically comparable to those of the prey's natural sensory environment, and (2) given functional predators are removed from the treatments, the fear effect is measured in the absence of any consumptive effects, a situation which never occurs in nature. We contend that demographic consequences of fear in natural populations may have been overestimated because the intensity of predator cues applied by experimenters in the majority of studies has been unnaturally high, in some instances rarely occurring in nature without consumption. Furthermore, the removal of consumption from the treatments creates the potential situation that individual prey in poor condition (those most likely to contribute strongly to the observed fear effects via starvation or reduced reproductive output) may have been consumed by predators in nature prior to the expression of fear effects, thus confounding consumptive and fear effects. Here, we describe an alternative treatment design that does not utilize predator cues, and in so doing, better quantifies the demographic effect of fear on wild populations. This treatment substitutes the traditional cue experiment where consumptive effects are eliminated and fear is simulated with a design where fear is removed and consumptive effects are simulated through the experimental removal of prey. Comparison to a natural population would give a more robust estimate of the effect of fear in the presence of consumption on the demographic variable of interest. This approach represents a critical advance in quantifying the mechanistic pathways through which predation structures ecological communities. Discussing the merits of both treatments will motivate researchers to go beyond simply describing the existence of fear effects and focus on testing their true magnitude in wild populations and natural communities. [ABSTRACT FROM AUTHOR]

Abstract: Community and ecosystem changes are happening in the pristine boreal forest ecosystem of the Yukon for 2 reasons. First, climate change is affecting the abiotic environment (temperature, rainfall and growing season) and driving changes in plant productivity and predator–prey interactions. Second, simultaneously change is occurring because of mammal species reintroductions and rewilding. The key ecological question is the impact these faunal changes will have on trophic dynamics. Primary productivity in the boreal forest is increasing because of climatic warming, but plant species composition is unlikely to change significantly during the next 50–100 years. The 9–10‐year population cycle of snowshoe hares will persist but could be reduced in amplitude if winter weather increases predator hunting efficiency. Small rodents have increased in abundance because of increased vegetation growth. Arctic ground squirrels have disappeared from the forest because of increased predator hunting efficiency associated with shrub growth. Reintroductions have occurred for 2 reasons: human reintroductions of large ungulates and natural recolonization of mammals and birds extending their geographic ranges. The deliberate rewilding of wood bison (Bison bison) and elk (Cervus canadensis) has changed the trophic structure of this boreal ecosystem very little. The natural range expansion of mountain lions (Puma concolor), mule deer (Odocoileus hemionus) and American marten (Martes americana) should have few ecosystem effects. Understanding potential changes will require long‐term monitoring studies and experiments on a scale we rarely deem possible. Ecosystems affected by climate change, species reintroductions and human alteration of habitats cannot remain stable and changes will be critically dependent on food web interactions. [ABSTRACT FROM AUTHOR]

Predators play a critical, top–down role in shaping ecosystems, driving prey population and community dynamics. Traditionally, studies of predator‐prey interactions have focused on direct effects of predators, namely the killing of prey. More recently, the non‐consumptive effects of predation risk are being appreciated; e.g. the ‘ecology of fear’. Prey responses to predation risk can be morphological, behavioural, and physiological, and are assumed to come at a cost to prey fitness. However, few studies have examined the relationship between predation risk and survival in wild animals. We tested the hypothesis that predation risk itself could reduce survival in wild‐caught snowshoe hares. We exposed female snowshoe hares to a simulated predator (a trained dog) during gestation only, and measured adult survival and, in surviving females, their ability to successfully wean offspring. We show for the first time in a wild mammal that the risk of predation can itself be lethal. Predation risk reduced adult female survival by 30%, and had trans‐generational effects, reducing offspring survival to weaning by over 85% – even though the period of risk ended at birth. As a consequence of these effects the predator‐exposed group experienced a decrease in number, while the control group substantially increased. Challenges remain in determining the importance of risk‐induced mortality in natural field settings; however, our findings show that non‐lethal predator encounters can influence survival of both adults and offspring. Future work is needed to test these effects in free‐living animals. [ABSTRACT FROM AUTHOR]

Population cycles have long fascinated ecologists from the time of Charles Elton in the 1920s. The discovery of large population fluctuations in undisturbed ecosystems challenged the idea that pristine nature was in a state of balance. The 10-year cycle of snowshoe hares ( Lepus americanus Erxleben) across the boreal forests of Canada and Alaska is a classic cycle, recognized by fur traders for more than 300 years., Since the 1930s, ecologists have investigated the mechanisms that might cause these cycles. Proposed causal mechanisms have varied from sunspots to food supplies, parasites, diseases, predation and social behaviour. Both the birth rate and the death rate change dramatically over the cycle. Social behaviour was eliminated as a possible cause because snowshoe hares are not territorial and do not commit infanticide., Since the 1960s, large-scale manipulative experiments have been used to discover the major limiting factors. Food supply and predation quickly became recognized as potential key factors causing the cycle. Experiments adding food and restricting predator access to field populations have been decisive in pinpointing predation as the key mechanism causing these fluctuations., The immediate cause of death of most snowshoe hares is predation by a variety of predators, including the Canada lynx ( Lynx canadensis Kerr). The collapse in the reproductive rate is not due to food shortage as was originally thought, but is a result of chronic stress from predator chases., Five major issues remain unresolved. First, what is the nature of the predator-induced memory that results in the prolonged low phase of the cycle? Second, why do hare cycles form a travelling wave, starting in the centre of the boreal forest in Saskatchewan and travelling across western Canada and Alaska? Third, why does the amplitude of the cycle vary greatly from one cycle to the next in the same area? Fourth, do the same mechanisms of population limitation apply to snowshoe hares in eastern North American or in similar ecosystems across Siberia? Finally, what effect will climatic warming have on all the above issues? The answers to these questions remain for future generations of biologists to determine. [ABSTRACT FROM AUTHOR]

Vole population cycles are a major force driving boreal ecosystem dynamics in northwestern Eurasia. However, our understanding of the impact of winter on these cycles is increasingly uncertain, especially because climate change is affecting snow predictability, quality, and abundance. We examined the role of winter weather and snow conditions, the lack of suitable habitat structure during freeze-thaw periods, and the lack of sufficient food as potential causes for winter population crashes. We live-trapped bank voles Myodes glareolus on 26 plots (0.36 ha each) at two different elevations (representing different winter conditions) in southeast Norway in the winters 2013/2014 and 2014/2015. We carried out two manipulations: supplementing six plots with food to eliminate food limitation and six plots with straw to improve habitat structure and limit the effect of icing in the subnivean space. In the first winter, all bank voles survived well on all plots, whereas in the second winter voles on almost all plots went extinct except for those receiving supplemental food. Survival was highest on the feeding treatment in both winters, whereas improving habitat structure had no effect. We conclude that food limitation was a key factor in causing winter population crashes. [ABSTRACT FROM AUTHOR]

Chronic activation of the stress axis caused by long-term uncontrollable and unpredictable factors in the environment has been regarded as causing maladaptive and/or pathological effects, both by those studying animals in the laboratory and in nature. While pathology may apply to the former, I argue that it does not apply to the latter., Our thinking on the role of chronic stress in animals in nature has been heavily influenced by biomedical research, but much less so by the ecological and evolutionary context within which animals actually function. I argue that when such stressors occur (e.g. periods of high predation risk, food limitation, prolonged severe weather, social conflict, etc.), although the animal may be chronically stressed, its responses are adaptive and continue to promote fitness., Chronic stressors in nature can be subdivided into whether they are reactive (direct physiological challenges threatening homeostasis and not requiring cognitive processing - for example, food limitation) or anticipatory (perceived to be threatening and requiring cognitive processing - for example, high predation risk). For anticipatory stressors, their impact on the animal should not be based on their absolute duration (they may be acute), but rather by the duration of their physiological consequences., The anticipatory stressor of persistent high predation risk does not elicit chronic stress in all prey classes. Cyclic snowshoe hare and arctic ground squirrels exhibit evidence of chronic stress when predator numbers are high, but cyclic vole and noncyclic elk populations do not. I suggest that chronic stress has evolved to benefit the fitness of the former and not the later, with the key factors being lifespan and life history. I propose that chronic stress evolves in a species only if it is adaptive. [ABSTRACT FROM AUTHOR]

Rapid, effective and enduring responses of physiology and behaviour to perturbations of the environment are key to robustness of an organism (ability to resist perturbations) and resilience (ability to resist and recover quickly from perturbations) so that the normal life cycle can be resumed quickly., Perturbations of the environment can be labile (i.e. eventually subside) or permanent such as when human activity changes the environment in the long term, for example, deforestation, urbanization, etc., Hormonal responses to labile perturbation factors ( LPFs) allow organisms to cope during the perturbation and then return to the normal life cycle. These hormonal responses are called stress responses especially in cases when major changes in physiology and behaviour occur (emergency life-history stage)., Permanent perturbations require more than just temporary acclimation resulting in changes in range, adaptation or in some cases local extinction. Perturbations can be abiotic, biotic and social, but these are not mutually exclusive., Here I focus on the effects of abiotic perturbation factors and their effects on the hypothalamo-pituitary-adrenal/interrenal axis in vertebrates. There is a great need for more field investigations of responses of free-living populations to perturbations of the environment, especially now that it appears the frequency and intensity of these events is increasing. However, such studies will require a high degree of opportunism on the part of the investigators to take advantage of unpredictable events when they occur. [ABSTRACT FROM AUTHOR]

Our goal in this review is to discuss how measures beyond simple quantification of total glucocorticoid levels are needed in comparative studies of stress. We need to measure corticosteroid binding globulin - CBG - and further downstream performance metrics to properly evaluate the significance and impact of stress in wild populations. We briefly cover the current literature, discuss methods that may enable detection of chronic stress and point to directions for future research to continue to clarify this field., CBG may regulate access of hormones to tissues, but disagreement remains as to the functional importance of total vs. free vs. bound hormone in the plasma. Here we focus on recent studies providing solid evidence supporting the free hormone hypothesis. These studies unequivocally indicate that the biologically active fraction consists only of the portion that is free (not total levels in the blood). We also present the 'reservoir hormone hypothesis', indicating the relevance of CBG-bound hormone in the plasma., The vast majority of physiological studies on stress in natural populations only measure blood corticosterone or its faecal metabolites. However, downstream metrics (e.g. immune function, oxidative stress and body mass changes) can assess the physiological impact of the changes in corticosterone and CBG and ultimately of the adaptiveness of these changes. Here we will discuss some of the most promising factors that can be measured downstream of the plasma and the rationale for their measurement, how to do it, and an introduction to the evidence., Although we draw on biomedical findings for some of this insight, we recognized that findings from a few mammalian laboratory species (primarily rats and mice) may not apply to the vast diversity of vertebrate species from fish to birds. In large measure we are still in the natural history phase of the accumulation of fundamental knowledge in terms of trying to extract general principles in the stress biology of natural populations. Our review outlines what is needed to lay the foundation for these principles. [ABSTRACT FROM AUTHOR]

Glucocorticoids ( GC) are integral to the stress response of vertebrates to environmental challenges. Their impact is immediate and widespread, and prolonged exposure can result in major activational and organizational changes. The vertebrate body has two mechanisms to limit GC impact: first, a rapid negative feedback system to turn off their release, and second, a protein - corticosteroid-binding globulin ( CBG) - to prevent them being free in the blood. Species used in biomedical research have CBG levels that normally bind 90-95% of blood GC. This evidence was the basis for the 'Free Hormone Hypothesis,' which posits that only unbound, free hormone is available for use by tissues and is biologically active. High levels of free GC typically occur only under conditions of chronic stress and are associated with major suppression of key body functions. The hypothesis proposes that the primary role of CBG is to render GCs unavailable, thereby preventing tissue exposure., From a field study in southern Ontario on northern ( Glaucomys sabrinus) and southern ( Glaucomys volans) flying squirrels, we found virtually no CBG binding capacity in their plasma, resulting in only about 10% of cortisol being bound throughout the year. However, neither species showed any evidence of being physiologically compromised. This presents a major challenge to the potential consequences of chronically high levels of free GC., To assess the generality of this finding, we carried out a phylogenetic comparison of 91 vertebrate species in which both GC and CBG levels were measured. In 93%, CBG levels were sufficient to bind about 90% of circulating GCs. This evidence is thus concordant with that from biomedical research. However, both flying squirrel species and four species of New World monkeys were extreme outliers. These two groups had the highest GC concentrations, but relatively little binding capacity (10% or less of their GC was bound)., An ancestor state reconstruction of the proportion of GCs bound indicated that the flying squirrels and New World monkeys evolved this character state from ancestors that followed the common 90% bound pattern. It also indicated that the original vertebrates - the earliest fishes - have most of their GC in the free state and little or no steroid-binding protein., We thus demonstrate a dichotomous pattern with respect to CBG: a dominant branch, where high levels of CBG bind most of the GC, applies to the majority of vertebrates; and a secondary branch, where low levels of CBG bind almost none of the GC, applies to a very small subset. For the latter, the critical unknown is how these species mitigate the impact of the high free GC levels and how such a dramatic trait shift could evolve. [ABSTRACT FROM AUTHOR]

Glucocorticoids hormones ( GCs) are intuitively important for mediation of age-dependent vertebrate life-history transitions through their effects on ontogeny alongside underpinning variation in life-history traits and trade-offs in vertebrates. These concepts largely derive from the ability of GCs to alter energy allocation, physiology and behaviour that influences key life-history traits involving age-specific life-history transitions, reproduction and survival., Studies across vertebrates have shown that the neuroendocrine stress axis plays a role in the developmental processes that lead up to age-specific early life-history transitions. While environmental sensitivity of the stress axis allows for it to modulate the timing of these transitions within species, little is known as to how variation in stress axis function has been adapted to produce interspecific variation in the timing of life-history transitions., Our assessment of the literature confirms that of previous reviews that there is only equivocal evidence for correlative or direct functional relationships between GCs and variation in reproduction and survival. We conclude that the relationships between GCs and life-history traits are complex and general patterns cannot be easily discerned with current research approaches and experimental designs., We identify several future research directions including: (i) integration of proximate and ultimate measures, including longitudinal studies that measure effects of GCs on more than one life-history trait or in multiple environmental contexts, to test explicit hypotheses about how GCs and life-history variation are related and (ii) the measurement of additional factors that modulate the effects of GCs on life-history traits (e.g. GC receptors and binding protein levels) to better infer neurendocrine stress axis actions., Conceptual models of HPA/I axis actions, such as allostatic load and reactive scope, to some extent explicitly predict the role of GCs in a life-history context, but are descriptive in nature. We propose that GC effects on life-history transitions, survival probabilities and fecundity can be modelled in existing quantitative demographic frameworks to improve our understanding of how GC variation influences life-history evolution and GC-mediated effects on population dynamics [ABSTRACT FROM AUTHOR]

Biomedical researchers have long appreciated that maternal stressors can induce preparative and adaptive programming in offspring via exposure to maternal Glucocorticoids ( GCs). However, few ecologists are aware of the capacity for maternal GC exposure to translate ecological and environmental stressors into preparative and adaptive programmed offspring responses in free-living systems. We review a growing body of experimental work indicating that circulating maternal GCs link ecological stressors with adaptive programming of the stress axis. Throughout, we emphasise that natural and human-induced ecological stressors play a fundamental role in programming the capacity of individuals, populations and communities to respond to both predictable and unpredictable ecological change via translating maternal adversity into responsive programming of the vertebrate stress axis., To encourage rigorous testing of this paradigm in a broad range of ecological systems, we introduce the principal extrinsic stressors with a recognised potential to alter maternal circulating GC levels. We then review from the biomedical literature regarding the underlying physiological and epigenetic mechanisms of stress-induced programming of individual phenotypes to predict how variation in ecological stressors can produce individual variation in stress axis management., To appreciate the potential evolutionary inertia (i.e. adaptive value) of maternally programmed individual variation, we review key recent studies in free-living systems that test its adaptive function, and then discuss how variation in stress-axis programming may scale up to influence populations and ecological communities., Given the huge potential of this field, it is encouraging that ecologists are beginning to examine how and why maternal GCs translate ecological and environmental stressors into preparative stress axis programming in free-living systems. [ABSTRACT FROM AUTHOR]

The ability to store energy endogenously is an important ecological mechanism that allows animals to buffer predictable and unpredictable variation in allostatic load. The secretion of glucocorticoids, which reflects changes in allostatic load, is suggested to play a major role in the adjustment of endogenous stores to these varying conditions., Although crucially important, the relationship between allostatic load and energy stores remains largely unexplored. Two contrasting hypotheses describe how stores may be adjusted: animals may use low allostatic loads to increase stores to a maximum possible ( 'fat and fit'), or they can attain a lean physique due to fitness advantages of a low body mass ( 'lean and fit')., We compiled observational and experimental data available for a long-lived seabird to examine the relationship between glucocorticoids and stored energy at two life history stages (incubation and chick-rearing). Data were collected across multiple years and colonies in the North Pacific, thereby reflecting the wide range of environmental conditions birds' encounter in the marine environment. During experimental manipulations, allostatic load was minimized by supplementing food to free-living birds., We found that the relationship between allostatic load and energy stores was clearly curvilinear at both life history stages. Observational data suggested that energy stores remained relatively stable under low allostatic load and decreased under high loads. Experimental data showed that birds did not maximize energy stores under favourable conditions but maintained energy stores below a physiologically attainable level., Energy stores remained consistently lower during chick-rearing compared to incubation across the wide range of variations in allostatic load suggesting that stage-specific trade-offs limit the accumulation of energy during favourable environmental conditions. Secretion of glucocorticoids did not appear to mediate this shift in energy stores between the life history stages., Overall, results of this study support the ' lean and fit' hypothesis. We conclude that increased energy stores may not necessarily reflect better environmental conditions experienced by individuals or predict their higher fitness. A major advantage of adopting a lean physique when environmental conditions allow may be the avoidance of additional energetic costs for moving a heavy body. In breeding seabirds, this advantage may be more important during chick-rearing. In the focal species, the secretion of glucocorticoids might be involved in regulation of energy stores within a life history stage but does not appear to mediate an adaptive shift in energy stores between the incubating and chick-rearing stages of reproduction. [ABSTRACT FROM AUTHOR]

Predator-induced stress has been used to exemplify the concept of stress for close to a century because almost everyone can imagine the terror of fleeing for one's life from a lion or a tiger. Yet, because it has been assumed to be acute and transitory, predator-induced stress has not been much studied by either comparative physiologists or population ecologists, until relatively recently., The focus in biomedical research has always been on chronic stress in humans, which most comparative physiologists would agree results from ' sustained psychological stress - linked to mere thoughts' rather than ' acute physical crises' (like surviving a predator attack) or ' chronic physical challenges' (such as a shortage of food). Population ecologists have traditionally focused solely on the acute physical crisis of surviving a direct predator attack rather than whether the risk of such an attack may have a sustained effect on other demographic processes (e.g. the birth rate)., Demographic experiments have now demonstrated that exposure to predators or predator cues can have sustained effects that extend to affecting birth and survival in free-living animals, and a subset of these have documented associated physiological stress effects. These and similar results have prompted some authors to speak of an 'ecology of fear', but others object that 'the cognitive and emotional aspects of avoiding predation remain unknown'., Recent biomedical studies on animals in the laboratory have demonstrated that exposure to predators or predator cues can induce ' sustained psychological stress' that is directly comparable to chronic stress in humans, and this has now in fact become one of the most common stressors used in studies of the animal model of post-traumatic stress disorder (PTSD)., We review these recent findings and suggest ways the laboratory techniques developed to measure the 'neural circuitry of fear' could be adapted for use on free-living animals in the field, in order to: (i) test whether predator risk induces ' sustained psychological stress' in wild animals, comparable to chronic stress in humans and (ii) directly investigate 'the cognitive and emotional aspects of avoiding predation' and hence the 'ecology of fear'. [ABSTRACT FROM AUTHOR]

Many aspects of the social environment affect hypothalamic-pituitary-adrenal ( HPA) axis function and increase circulating glucocorticoid concentrations. In this review, we examine the relationships between the social environment and the function of the HPA axis in vertebrates., First, we explore the effects of the social environment on glucocorticoid secretion in territorial (primarily non-social) species, with an emphasis on the effects of variation in population density, as modified by environmental factors such as predation risk and food availability. In general, high population density or frequent territorial intrusions are associated with increased glucocorticoid secretion in a wide range of taxa, including mammals, birds, fish and reptiles, although there is considerable variability across species., Second, we consider the effects of social interactions and dominance rank on glucocorticoid secretion in social species, mostly in birds and mammals. We review studies that have detected an association between social status and glucocorticoid levels - sometimes with higher glucocorticoid levels in low-ranking individuals, and sometimes with higher glucocorticoid levels in dominant individuals. The relationship between dominance and glucocorticoid levels varies among species, populations and years, in a manner that depends on the stability of the social hierarchy, environmental conditions, the type of breeding system, and the manner in which high rank is obtained and maintained., Finally, we discuss the concept of allostasis and consider interactions between social effects and other environmental factors, noting that there is relatively little research on these interactions to date. For both non-social and social species, we identify priorities of future research. These priorities include more complete descriptions of HPA function that move beyond measurements of basal glucocorticoid concentrations (which will generally require field experiments), to studies that examine organizational effects of social stressors, that directly test the relationship between HPA function and fitness, and that examine how glucocorticoid responses affect population dynamics., Although several lines of evidence suggest that glucocorticoid responses can affect the fitness of individuals and therefore can alter the dynamics of populations, the effect of glucocorticoid responses on population dynamics remains essentially unstudied. [ABSTRACT FROM AUTHOR]

Disturbances, or stressors, are pervasive in nature and profoundly influence fitness of individuals and, ultimately, species extinction risk. Glucocorticoid ( GC) hormones affect multiple physiological and behavioural traits regulating the responses of vertebrates to stressors. Current understanding of what influences GC stress response variation among species is poor. However, identifying environmental causes of species-level variation in the GC stress response is important to understanding macroevolutionary patterns in this key physiological system and potentially offers a metric for evaluating species risk to global change., We tested whether GC stress response of 22 reptile and 66 bird species exposed to a uniform capture stress protocol could be predicted from a GC stressor space model. This model predicts that species-level variation in GCs results from multivariate selection from environmental gradients and species-specific metabolic sensitivities. Thus, these multivariate selection interactions delimit the cumulative selection intensity causing GC stress response variation among vertebrates., Using phylogenetic generalized least square regression analysis, we ranked models incorporating putative variables pertaining to the corticosterone stress response ( CSR) for reptiles and birds separately. The top-ranked models explained between 14% and 33% of species variation in the CSR. Models indicated that combinations of variables including increasing body mass, net primary productivity and latitude influenced GC stress responsiveness in reptiles and birds., We conclude that GC stress response variation among species represents, in part, evolution to species-specific traits and large-scale environmental variation. Further improvement to predicting species variation in GC stress response could be achieved by expanding our stressor space model to integrate microevolutionary and ecological processes., There is a pressing need to identify physiological traits that predict species responses to environmental change. Conceptually, the use of plasma GC levels as a species-specific marker for environmental tolerance could be plausible. However, we identify three major knowledge gaps and ensuing research areas to aid understanding of the utility of variation in GC stress response as a metric for inferring how species might respond to global change. [ABSTRACT FROM AUTHOR]

Summary 1. At high latitudes, evolutionary adaptations focus on those that maximize survival, with hibernation being a major one used by many smaller mammals. Typically, mammalian hibernators overwinter in sites that are ≈0°C. However, in arctic regions, such sites do not exist, necessitating hibernation at sites well below 0°C. Lipid, the normal fuel of most hibernators, may not provide sufficient glucose needed by certain tissues to permit survival, with muscle breakdown being required. Critical to enhancing muscle stores are high concentrations of anabolic androgens prior to hibernation when the gonads are inactive. 2. We compare and contrast androgen levels in arctic ground squirrels (AGS) ( Urocitellus parryii Richardson) from the Yukon and Columbian ground squirrel (CGS) ( U. columbianus Ord) from southern Alberta. 3. In males, changes in testes mass over the active season were similar between AGS and CGS. In contrast, during the breeding and the nonbreeding, pre-hibernation periods, androgen levels in AGS were 6-10 and 20-25 times, respectively, those of CGS. From the breeding to the pre-hibernation periods, androgen levels declined 41% in AGS, but 86% in CGS. In females, androgen levels in AGS were high throughout the active season and, prior to hibernation, were 24 times those in CGS. 4. In pre-hibernating AGS, we determined the source of these high androgen levels from two studies. First, using a hormonal challenge protocol to probe the hypothalamic-pituitary-adrenal axis, we found that androgen levels in both females and males decreased ≥10% in response to suppression by dexamethasone (an artificial glucocorticoid that inhibits ACTH release) and increased ≥18% in response to direct adrenal stimulation by adrenocorticorticotropic homone (ACTH). Second, by sequential hormonal injections in males of gonadotropin-releasing hormone (GnRH) and of ACTH coupled with gonadectomy (GDX), followed by adrenalectomy (ADX), we found that GnRH had no effect, whereas ACTH stimulated androgen levels by >40%, both before and after GDX. After ADX, levels fell by 80%. Thus, the adrenals, not the gonads, are the source of these androgens. 5. We hypothesize that pre-hibernating AGS have evolved a solution seen in no other known mammal: exploiting the benefits of high adrenal androgen levels prior to hibernation to build muscle that is then catabolized overwinter. The unknown is how AGS have mitigated the costs of these high androgen levels. [ABSTRACT FROM AUTHOR]

The study of adult neurogenesis has had an explosion of fruitful growth. Yet numerous uncertainties and challenges persist. Our review begins with a survey of species that show evidence of adult neurogenesis. We then discuss how neurogenesis varies across brain regions and point out that regional specializations can indicate functional adaptations. Lifespan and aging are key life-history traits. Whereas 'adult neurogenesis' is the common term in the literature, it does not reflect the reality of neurogenesis being primarily a 'juvenile' phenomenon. We discuss the sharp decline with age as a universal trait of neurogenesis with inevitable functional consequences. Finally, the main body of the review focuses on the function of neurogenesis in birds and mammals. Selected examples illustrate how our understanding of avian and mammalian neurogenesis can complement each other. It is clear that although the two phyla have some common features, the function of adult neurogenesis may not be similar between them and filling the gaps will help us understand neurogenesis as an evolutionarily conserved trait to meet particular ecological pressures. [ABSTRACT FROM AUTHOR]

Maternal effects may be a major factor influencing the demography of populations. In mammals, the transmission of stress hormones between mother and offspring may play an important role in these effects. Laboratory studies have shown that stressors during pregnancy and lactation result in lifelong programming of the offspring phenotype. However, the relevance of these studies to free-living mammals is unclear. The 10-year snowshoe hare (Lepus americanus) cycle is intimately linked to fluctuating predation pressure and predation risk. The enigma of these cycles is the lack of population growth following the decline phase, when the predators have virtually all disappeared and the food supply is ample. We have shown that a predator-induced increase in maternal stress hormone levels resulted in a decline in reproduction. Here we examine population and hormone changes over a four-year period from the increase (2005) to the decline (2008). We report (1) that an index of maternal stress (fecal corticosteroid metabolite [FCM] concentrations) fluctuates in synchrony with predator density during the breeding season; (2) that maternal FCM levels are echoed in their offspring, and this occurs at a population-wide level; and (3) that higher maternal FCM levels at birth are correlated with an increased responsiveness of the hypothalamic-pituitary-adrenal (HPA) axis in their progeny. Our results show an intergenerational inheritance of stress hormones in a free-ranging population of mammals. We propose that the lack of recovery of reproductive rates during the early low phase of the hare cycle may be the result of the impacts of intergenerational, maternally inherited stress hormones caused by high predation risk during the decline phase. [ABSTRACT FROM AUTHOR]

Ecology Letters (2010) 13: 980–988 Predator-induced changes in physiology and behaviour may negatively affect a prey’s birth rate. Evidence of such indirect predator effects on prey demography remains scarce in birds and mammals despite invertebrate and aquatic studies that suggest ignoring such effects risks profoundly underestimating the total impact of predators. We report the first experimental demonstration of indirect predator effects on the annual ‘birth’ rate resulting from negative effects on the size of subsequent clutches laid by birds. We manipulated the probability of nest predation and measured the size of subsequent clutches and multiple indices of the mother’s physiological condition, while controlling for food availability, date and stage of breeding. Females subject to frequent experimental nest predation laid smaller subsequent clutches and were in poorer physiological condition, particularly regarding non-resource-based indices (e.g. oxidative stress and glucocorticoid mobilization) consistent with both a response to the threat of predation and an increased cost of egg production. [ABSTRACT FROM AUTHOR]

1. Prey responses to high predation risk can be morphological or behavioural and ultimately come at the cost of survival, growth, body condition, or reproduction. These sub-lethal predator effects have been shown to be mediated by physiological stress. We tested the hypothesis that elevated glucocorticoid concentrations directly cause a decline in reproduction in individual free-ranging female snowshoe hares, Lepus americanus. We measured the cortisol concentration from each dam (using a faecal analysis enzyme immunoassay) and her reproductive output (litter size, offspring birth mass, offspring right hind foot (RHF) length) 30 h after birth. 2. In a natural monitoring study, we monitored hares during the first and second litter from the population peak (2006) to the second year of the decline (2008). We found that faecal cortisol metabolite (FCM) concentration in dams decreased 52% from the first to the second litter. From the first to the second litter, litter size increased 122%, offspring body mass increased 130%, and offspring RHF length increased 112%. Dam FCM concentrations were inversely related to litter size ( r2 = 0·19), to offspring birth mass ( r2 = 0·32), and to offspring RHF length ( r2 = 0·64). 3. In an experimental manipulation, we assigned wild-caught, pregnant hares to a control and a stressed group and held them in pens. Hares in the stressed group were exposed to a dog 1–2 min every other day before parturition to simulate high predation risk. At parturition, unsuccessful-stressed dams (those that failed to give birth to live young) and stressed dams had 837% and 214%, respectively, higher FCM concentrations than control dams. Of those females that gave birth, litter size was similar between control and stressed dams. However, offspring from stressed dams were 37% lighter and 16% smaller than offspring from control dams. Increasing FCM concentration in dams caused the decline of offspring body mass ( r2 = 0·57) and RHF ( r2 = 0·52). 4. This is the first study in a free-ranging population of mammals to show that elevated, predator-induced, glucocorticoid concentrations in individual dams caused a decline in their reproductive output measured both by number and quality of offspring. Thus, we provide evidence that any stressor, not just predation, which increases glucocorticoid concentrations will result in a decrease in reproductive output. [ABSTRACT FROM AUTHOR]

1. Across the vast boreal forests of North America, no population cycles in Clethrionomys species occur. In Eurasia, by contrast, some Clethrionomys populations of the same species undergo regular 3–5-year cycles. We examined the effects of nutrients, food, competitors, predators and climate on population limitation in the northern red-backed vole ( Clethrionomys rutilus Pallas) in the south-western Yukon to determine why this difference occurs. 2. From 1986 to 1996 we added food, reduced large mammal predators and excluded snowshoe hares ( Lepus americanus Erxleben) from large plots and found that none of these manipulations affected red-backed vole abundance. Adding nutrients as nitrogen, phosphorus and potassium (NPK) fertilizer had a slight negative effect, probably acting through a reduction in dwarf shrub productivity caused by competition from grasses. 3. We monitored weasel populations directly through trapping and indirectly through snow tracking. Predation by these vole specialists was irrelevant as a limiting factor most of the time because voles in this area do not reach the densities needed to sustain weasel populations. Other boreal forest mammal and bird predators did not focus on red-backed voles. However, when red-backed vole populations increased in the forest and Microtus voles also increased in the meadows, weasel populations increased and may have temporarily depressed red-backed voles in winter. 4. We monitored one major potential food, white spruce seeds, but seed fall was not related to population changes in red-backed voles, even after mast years. 5. We assessed the impact of weather variables, and the average depth of the snow pack during winter (October–March) was correlated directly with vole demography, having both direct effects in that year and delayed effects in the following year. 6. Our long-term trapping data (1973–96) indicate that Clethrionomys populations fluctuated, with peaks following hare peaks by 2–3 years. 7. We propose that the key variable limiting these vole populations is overwinter survival, and this is a function of overwinter food from berries produced during the previous summer by dwarf shrubs. These shrubs may be stimulated by abundant moisture from winter snows or by periodic fertilization from large quantities of pellets produced at snowshoe hare peaks. [ABSTRACT FROM AUTHOR]

1. Overwinter mass loss can reduce energetic requirements in mammals (Dehnel's phenomenon). Alternatively, mass loss can result from food limitation or high predation risk. 2. We use data from fertilizer, food-supplementation and predator-exclusion experiments in the Yukon during a population cycle from 1986 to 1996 to test the causes of overwinter mass loss by snowshoe hares ( Lepus americanus). In all years, some hares on control sites gained mass overwinter. During the increase phase the majority gained mass, but in all other phases the majority lost mass. 3. Snowshoe hares weighing < 1000 g in autumn always gained mass overwinter, as did the majority that weighed 1000–1400 g. Hares weighing > 1800 g in autumn usually lost mass. 4. Snowshoe hares on the predator-exclosure + food site gained mass overwinter in all years. Hares on the food-supplementation sites lost mass during the decline but gained mass in all other phases. Fertilization had little effect on mass dynamics. 5. Snowshoe hares were more likely to lose mass during winters with low survival rates. Snowshoe hares on the predator-exclosure treatments were more likely to gain mass than were hares on control sites. 6. Overwinter mass loss was correlated with maximum snow depth. At equivalent snow depths, hares on food-supplemented areas lost 98 g (± 14·6 SE) less on average than hares on the controls and predator-exclosure treatment. 7. Bone-marrow fat was related to body mass and cause of death. Small hares had the lowest marrow fat. Hares killed by humans had higher marrow fat than those killed by predators; hares that simply died had the lowest marrow fat. Hares on food-supplemented sites had the highest kidney and marrow fat. 8. Overwinter-mass loss for snowshoe hares is explained interactively by winter conditions, food supply, predation risk and autumn mass. Some snowshoe hares lost mass overwinter in all years and on all treatments, suggesting that reducing body mass may facilitate survival, especially in cases where foraging costs are high energetically or increase predation risk. [ABSTRACT FROM AUTHOR]

We investigated the effect of local environment on the demography and population dynamics of arctic ground squirrels (Spermophilus parryii plesius) by comparing reproduction, survival, and population trends of squirrels living in low elevation boreal forest and high elevation alpine tundra sites in southwestern Yukon Territory, Canada. Contrary to the trend for most birds and mammals, reproduction was significantly lower at the lower elevation and females living at higher elevation did not delay the age at which they first reproduced. Even though survival in the boreal forest was lower in summer than in the alpine, it was higher over winter so annual adult female survival was similar between sites.Sensitivity analysis of model parameters revealed that in the forest, population growth rate (λ) was most sensitive to small changes in adult active season survival whereas for the alpine population,λ was most sensitive to changes in juvenile winter survival. In their respective habitats, these parameters also showed high year to year variation and thus contributed greatly to the population trends observed. Even though ground squirrels persisted in the boreal forest, the measured demographic rates indicate the forest was sink habitat (λ<1) and may have relied on nearby grassy meadows for immigrants. In contrast, the alpine habitat maintained a ground squirrel population in the absence of immigration (λ=1).The variation in demographic rates between ground squirrels living at high and low elevation may arise from phenotypic responses of squirrels to different habitat structure. Arctic ground squirrels rely on sight to detect predators from a safe distance, and the boreal forest, with its lower visibility and higher predator density, appears to be suboptimal habitat. [ABSTRACT FROM AUTHOR]

In ecological risk assessment, a key necessity is to understand how contaminants known to have negative impact on laboratory mammals affect the population demography of mammals living in their natural environment. We examined the demography of six local populations of the short-tailed shrew (Blarina brevicauda) living in eastern deciduous forest palustrine habitat along the Housatonic River (MA, USA) on soils contaminated with a range of polychlorinated biphenyl (PCB) concentrations (1.5-38.3 ppm). The objective of the study was to assess whether PCBs adversely affect the population demography of these small mammals living in their natural environment. Blarina were selected for study because they would be expected to readily bioaccumulate PCBs from the soil. Populations were intensively live trapped on 1-ha grids from spring to autumn 2001. There was no relationship between any demographic parameter and PCB soil concentrations. Densities were high (usually exceeding 20/ha, and on two grids exceeded 60/ha in summer); survival was good (typically 60-75% per 30 d); and sex ratio, reproduction rates, growth rates, and body mass were within the ranges reported in the literature. Thus, these shrew populations showed no detectable impact on their population demography from living on PCB-contaminated sites. [ABSTRACT FROM AUTHOR]

Reproductive traits of voles vary with the phases of the population density fluctuations. We sought to determine whether the source of this variation resides in the individuals or in their environment. Overwintering field voles (Microtus agrestis) from two cyclic out-of-phase populations (increase and peak phases) were sampled in early spring and bred in the laboratory for two generations under standardised conditions with ambient light and temperature. Monitoring of the source populations by capture-mark-recapture showed large differences in reproductive performance. In the increase area, reproduction started six weeks earlier, the probability of maturation of young-of-the-year was more than ten times higher during mid-summer, and reproduction continued nearly two months later in the autumn than in the peak area. These differences were not found to be associated with a difference in age structure of overwintered animals between the two areas (assessed by the distribution of eye lens masses from autopsy samples). Although the population differences in reproductive traits were to some degree also present among the overwintered animals in the laboratory, we found no difference in reproductive traits in the laboratory-born generations. There was a strongly declining seasonal trend in probability of sexual maturation both in the field and in the laboratory under ambient light conditions. However, in the field there were large population differences in the steepness of the seasonal decline that were not seen under the standardised laboratory conditions. We conclude that seasonal decline in maturation rates is governed by change in photoperiod, but that the population level variation in the shape of the decline is caused by a direct response to the environment and not due to variation in any intrinsic state of the individuals reflecting the environment experienced by the previous generation(s). [ABSTRACT FROM AUTHOR]

Assesses the physiological stress response of male arctic ground squirrels, Spermophilus parryii plesius. Percent of male population during breeding season; Biology of arctic squirrels; Immunological competence; Blood parameters and immune response; Factors affecting changes in testosterone levels.

The meadow vole, Microtus pennsylvanicus , is the most widely distributed Microtus species in North America. Across its range, it shows marked demographic differences, experiences a large range of climatic conditions, and varies considerably in body size and life-history characteristics. To study the genetic basis of the geographic variation in size and life history of this species, we subjected three populations, one from central Canada and two from eastern Canada, to quantitative genetic analysis in the lab. We studied the variance and covariance of several size and growth variables as well as age and size at maturity by means of population crosses, full-sib analysis, and parent-offspring regressions. We found that the phenotypic differences among these populations are almost entirely due to environmental effects. However, within populations, additive genetic and maternal effects explain most of the variation. We discuss possible explanations for the lack of genetic differences among the populations and speculate that a similar reaction norm is maintained in all populations through heterogeneity in the temporal or spatial environment that the populations experience. The heterogeneity may be mediated through population density fluctuations, climatic variation, or variation in site productivity. Thus, we hypothesize that M. pennsylvanicus has evolved to be the best in all possible worlds rather than in one actual world. This study highlights the crucial importance of maternal and environmental effects on the size, growth, and life history of small rodents. [ABSTRACT FROM AUTHOR]

Provides information on a study which compared the effects of 3 inhalant anesthetics and manual restraint on induction and recovery time, stress response, and hematological parameters in Arctic ground squirrels. Research methods; Results and discussion; Conclusion.

Examines the impact of predator-induced stress on snowshow hare population dynamics. Effects to physiology and demography; Population cycles of snowshoe hares; Signs of chronic stress in hares.

Examines the cycles of population found among small mammals. What the cycles characterize; Fluctuation of populations of some small mammals; Variation of the amplitude of cycles; Potential of these small mammals; Information on the low phases; Contrasting population factors.

We tested the hypothesis that low doses of gamma radiation have beneficial, hormetic effects on the stress axis (the hypothalamic-pituitary-adrenocortical axis) of free-ranging meadow vole populations (Microtus pennsylvanicus). Voles were exposed to chronic gamma radiation from a 137Cs field irradiator. In isolated populations, voles received one of three treatments over a four-year period: Controls (0.19-0.42 µGy/h -- levels that were 2-5x above background levels [0.1 µGy/h] and live-trapped in all years -- 1982-1985), low doses (22.6 µGy/h - 50-200x background, live-trapped from November 1982-April 1985), or high doses (3,840 µGy/h - 40,000x background, live-trapped from November 1983-April 1985). Voles exposed to a low dose had levels of free and total corticosterone that were significantly higher than those in the control or high-dose groups. Differences in response to radiation between the sexes were apparent for maximum corticosterone-binding capacity, with females exposed to low doses having higher binding capacity than control or high-dose females, whereas males exposed to low doses had lower binding capacity than control or high-dose males. Low-dose voles had higher counts of neutrophils than either the controls or high-dose voles; hematocrit was greater in the controls than in irradiated voles. These results indicate that voles display a hormetic response to radiation, wherein low doses of an otherwise harmful agent produce a beneficial effect. The stimulation of the stress axis resulting in the increased secretion of glucocorticoids, which may protect against the excessive actions of the immune and inflammatory responses, may be a key mechanism producing this effect. [ABSTRACT FROM AUTHOR]

Summary1. We examined the effects of food and predators on population limitation in the arctic ground squirrel (Spermophilus parryii plesius Richardson) in the boreal forest of the south-western Yukon. We focused on ground squirrel reproduction and overwinter survival. 2. Squirrel populations were monitored by live-trapping and radio-telemetry from 1993 to the spring of 1996 on four control and four experimental areas (one predator exclosure treatment, two food addition treatments, and one predator exclosure plus food addition treatment). 3. Predator exclusion increased body condition, percentage lactating, percentage weaning litters, litter size, and doubled population density relative to controls, but had no effect on juvenile growth rate, overwinter survival, or juvenile emergence date. 4. Food addition advanced juvenile emergence date and increased adult body condition, percentage lactating, percentage weaning litters, litter size, population density relative to controls (4–7 fold), but had no effect on juvenile growth rate or overwinter survival. 5. Predator exclusion combined with food addition increased adult body condition, percentage lactating, percentage weaning litters, and population density relative to controls (19-fold). 6. We conclude that arctic ground squirrels in the boreal forest are limited by an interaction between food and predation, acting primarily through changes in reproduction, and that their impact on density was multiplicative. [ABSTRACT FROM AUTHOR]