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Peers, Michael; Majchrzak, Yasmine; Menzies, Allyson; Studd, Emily; Bastille-Rousseau, Guillaume; Boonstra, Rudy; Humphries, Murray; Jung, Thomas; Kenney, Alice; Krebs, Charles; Murray, Dennis; Boutin, Stan 2021-06-16 <p style="text-indent:36.0pt;">Canada lynx (<i>Lynx canadensis</i>) and snowshoe hares (<i>Lepus americanus</i>) form a keystone predator-prey cycle that has large impacts on the North-American boreal forest vertebrate community. Snowshoe hares and lynx are both well-suited for snowy winters, but climate change associated shifts in snow conditions could lower hare survival and alter cyclic dynamics. Using detailed monitoring of snowshoe hare cause-specific mortality, behaviour, and prevailing weather, we demonstrate that hare mortality risk is strongly influenced by variation in snow conditions. Although predation risk from lynx was largely unaffected by snow conditions, coyote (<i>Canis latrans</i>) predation increased in shallow snow. Maximum snow depth in our study area has decreased 33% over the last two decades and predictions based on prolonged shallow snow indicate future hare survival could resemble that seen during population declines. Our results indicate that climate change could disrupt cyclic dynamics in the boreal forest.</p>
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Lamb, Clayton T.; Mowat, Garth; Gilbert, Sophie L.; McLellan, Bruce N.; Nielsen, Scott E.; Boutin, Stan 2018-08-25 Brown bears are known to use rubbing behavior as a means of chemical communication, but the function of this signaling is unclear. One hypothesis that has gained support is that male bears rub to communicate dominance to other males. We tested the communication of dominance hypothesis in a low-density brown bear population in southeast British Columbia. We contrasted rubbing rates for male and female bears during and after the breeding season using ten years of DNA-mark-recapture data for 643 individuals. Here we demonstrate that male brown bears rub 60% more during the breeding than the non-breeding season, while female rubbing had no seasonal trends. Per capita rub rates by males were, on average, 2.7 times higher than females. Our results suggest that the function of rubbing in the Rocky Mountains may not only be to communicate dominance, but also to self-advertise for mate attraction. We propose that the role of chemical communication in this species may be density-dependent, where the need to self-advertise for mating is inversely related to population density and communicating for dominance increases with population density. We suggest that future endeavors to elucidate the function of rubbing should sample the behavior across a range of population densities using camera trap and genotypic data.
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Studd, Emily K.; Landry-Cuerrier, Manuelle; Menzies, Allyson K.; Boutin, Stan; McAdam, Andrew G.; Lane, Jeffrey E.; Humphries, Murray M. 2018-12-28 1. The miniaturization and affordability of new technology is driving a biologging revolution in wildlife ecology with use of animal-borne data logging devices. Among many new biologging technologies, accelerometers are emerging as key tools for continuously recording animal behavior. Yet a critical, but under-acknowledged consideration in biologging is the trade-off between sampling rate and sampling duration, created by battery- (or memory-) related sampling constraints. This is especially acute among small animals, causing most researchers to sample at high rates for very limited durations.Here, we show that high accuracy in behavioral classification is achievable when pairing low frequency acceleration recordings with temperature. 2. We conducted 84 hours of direct behavioral observations on 67 free-ranging red squirrels (200-300 g) that were fitted with accelerometers (2 g) recording tri-axial acceleration and temperature at 1 Hz. We then used a random forest algorithm and a manually-created decision tree, with variable sampling window lengths, to associate observed behavior with logger recorded acceleration and temperature. Finally, we assessed the accuracy of these different classifications using an additional 60 hours of behavioral observations, not used in the initial classification. 3. The accuracy of the manually-created decision tree classification using observational data varied from 70.6% to 91.6% depending on the complexity of the tree, with increasing accuracy as complexity decreased. Short duration behavior like running had lower accuracy than long duration behavior like feeding. The random forest algorithm offered similarly high overall accuracy, but the manual decision tree afforded the flexibility to create a hierarchical tree, and to adjust sampling window length for behavioral states with varying durations. 4. Low frequency biologging of acceleration and temperature allows accurate behavioral classification of small animals over multi-month sampling durations. Nevertheless, low sampling rates impose several important limitations, especially related to assessing the classification accuracy of short duration behavior.
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Peers, Michael; Konkolics, Sean; Majchrzak, Yasmine; Menzies, Allyson; Studd, Emily; Boonstra, Rudy; Boutin, Stan; Lamb, Clayton 2021-07-23 <p>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 days) relative to herbivore carcasses (1.1 days), and carnivore carcasses persisted on the landscape for over a month longer (48.4 days and 5.5 days, respectively). The longer persistence times were due to the reduction in scavenging by carnivores such as Canada lynx (<i>Lynx canadensis</i>). 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.</p>
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Hendrix, Jack G; Fisher, David; Martinig, April; Boutin, Stan; Dantzer, Ben; Lane, Jeffrey; McAdam, Andrew 2021-03-06 <p>1) Juvenile survival to first breeding is a key life history stage for all taxa. Survival through this period can be particularly challenging when it can coincide with harsh environmental conditions such as a winter climate or food scarcity, leading to highly variable cohort survival. However, the small size and dispersive nature of juveniles generally makes studying their survival more difficult. 2) In territorial species, a key life history event is the acquisition of a territory. A territory is expected to enhance survival, but how it does so is not often identified. We tested how the timing of territory acquisition influenced the winter survival of juvenile North American red squirrels Tamiasciurus hudsonicus, hereafter red squirrels, and how the timing of this event mediated sources of mortality. We hypothesized that securing a territory prior to when food resources become available would reduce juvenile susceptibility to predation and climatic factors over winter. 3) Using 27 years of data on the survival of individually-marked juvenile red squirrels, we tested how the timing of territory acquisition influenced survival, whether the population density of red squirrel predators and mean temperature over winter were related to individual survival probability, and if territory ownership mediated these effects. 4) Juvenile red squirrel survival was lower in years of high predator abundance and in colder winters. Autumn territory owners were less susceptible to lynx Lynx canadensis, and possibly mustelid Mustela and Martes spp., predation. Autumn territory owners had lower survival in colder winters, but surprisingly non-owners had higher survival in cold winters. 5) Our results show how the timing of a life history event like territory acquisition can directly affect survival and also mediate the effects of biotic and abiotic factors later in life. This engenders a better understanding of the fitness consequences of the timing of key life history events.</p>
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Broadley, Kate; Burton, Cole; Boutin, Stan; Avgar, Tal 2020-11-20 <p>Camera-traps (CTs) are an increasingly popular tool for wildlife survey and monitoring. Estimating relative abundance in unmarked species is often done using detection rate as an index of relative abundance, which assumes a positive linear relationship with true abundance. This assumption may be violated if movement behavior varies with density, but the degree to which movement is density-dependent across taxa is unclear. The potential confounding of population-level relative abundance indices by movement depends on how regularly, and by what magnitude, movement rate and home-range size vary with density. We conducted a systematic review and meta-analysis to quantify relationships between movement rate, home range size, and density, across terrestrial mammalian taxa. We then simulated animal movements and CT sampling to test the effect of contrasting movement scenarios on CT detection rates. Overall, movement rate and home range size were negatively correlated with density and positively correlated with one another. The strength of the relationships varied significantly between taxa and populations.  In simulations, detection rates were related to true abundance but underestimated change, particularly for slower moving species with small home ranges. In situations where animal space use changes markedly with density, we estimate that up to thirty percent of a true change in abundance may be missed due to the confounding effect of movement, making trend estimation more difficult. The common assumption that movement remains constant across densities is therefore violated across a wide range of mammal species. When studying unmarked species using CT detection rates, researchers and managers should consider that such indices of relative abundance reflect both density and movement. Practitioners interpreting changes in detection rates should be aware that observed differences may be biased low relative to true changes in abundance, and that further information on animal movement may be required to make robust inferences on population trends.</p>
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Lamb, Clayton; Mowat, Garth; Reid, Aaron; Smit, Laura; Proctor, Michael; McLellan, Bruce N.; Nielsen, Scott E.; Boutin, Stan 2020-04-06 Human activities have dramatic effects on the distribution and abundance of wildlife. Increased road densities and human presence in wilderness areas have elevated human-caused mortality of grizzly bears and reduced bears' use. Management agencies frequently attempt to reduce human-caused mortality by managing road density and thus human access, but the effectiveness of these actions is rarely assessed. We combined systematic, DNA-based mark–recapture techniques with spatially explicit capture–recapture models to estimate population size of a threatened grizzly bear population (Kettle–Granby), following management actions to recover this population. We tested the effects of habitat and road density on grizzly bear population density. We tested both a linear and threshold-based road density metric and investigated the effect of current access management (closing roads to the public). We documented an c. 50% increase in bear density since 1997 suggesting increased landscape and species conservation from management agencies played a significant role in that increase. However, bear density was lower where road densities exceeded 0.6 km/km2 and higher where motorised vehicle access had been restricted. The highest bear densities were in areas with large tracts of few or no roads and high habitat quality. Access management bolstered bear density in small areas by 27%. Synthesis and applications. Our spatially explicit capture–recapture analysis demonstrates that population recovery is possible in a multi-use landscape when management actions target priority areas. We suggest that road density is a useful surrogate for the negative effects of human land use on grizzly bear populations, but spatial configuration of roads must still be considered. Reducing roads will increase grizzly bear density, but restricting vehicle access can also achieve this goal. We demonstrate that a policy target of reducing human access by managing road density below 0.6 km/km2, while ensuring areas of high habitat quality have no roads, is a reasonable compromise between the need for road access and population recovery goals. Targeting closures to areas of highest habitat quality would benefit grizzly bear population recovery the most.
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Lamb, Clayton T.; Mowat, Garth; McLellan, Bruce N.; Nielsen, Scott E.; Boutin, Stan 2016-09-29 Habitat choice is an evolutionary product of animals experiencing increased fitness when preferentially occupying high-quality habitat. However, an ecological trap (ET) can occur when an animal is presented with novel conditions and the animal's assessment of habitat quality is poorly matched to its resulting fitness. We tested for an ET for grizzly (brown) bears using demographic and movement data collected in an area with rich food resources and concentrated human settlement. We derived measures of habitat attractiveness from occurrence models of bear food resources and estimated demographic parameters using DNA mark–recapture information collected over 8 years (2006–2013). We then paired this information with grizzly bear mortality records to investigate kill and movement rates. Our results demonstrate that a valley high in both berry resources and human density was more attractive than surrounding areas, and bears occupying this region faced 17% lower apparent survival. Despite lower fitness, we detected a net flow of bears into the ET, which contributed to a study-wide population decline. This work highlights the presence and pervasiveness of an ET for an apex omnivore that lacks the evolutionary cues, under human-induced rapid ecological change, to assess trade-offs between food resources and human-caused mortality, which results in maladaptive habitat selection.
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Studd, Emily K.; Boutin, Stan; McAdam, Andrew G.; Krebs, Charles J.; Humphries, Murray M. 2015-08-04 Neonatal reproductive failure should occur when energetic costs of parental investment outweigh fitness benefits. However, little is known about the drivers of neonatal reproductive failure in free ranging species experiencing continuous natural variation in predator abundance and in the energetic and fitness costs and benefits associated with parental investment. Long-term comprehensive studies are required to better understand how biotic, abiotic, and life history conditions interact to drive occurrences of reproductive failure in the wild. Using 24 years (1987-2011) of reproductive data from a northern boreal population of North American red squirrels in southwestern Yukon, we examined the effects of predator abundance, energetics (resource availability, ambient temperature and litter size), and fitness benefits (probability of overwinter juvenile survival and maternal age) on occurrences of neonatal reproductive failure (494/2670 reproductive attempts; 18.5%). Neonatal reproductive failure was driven by a combination of predator abundance, and the energetic and fitness costs and benefits of parental investment. The abundance of mustelids and maternal age were positively related to the occurrence of neonatal reproductive failure. High energy costs associated with a combination of low resource availability and cold ambient temperatures or large litters, corresponded to increased occurrences of neonatal reproductive failure. However, the strength of these relationships was influenced by variation in juvenile overwinter survival (i.e. fitness benefits). We provide evidence that predation pressure is an important driver of neonatal reproductive failure. In addition, we found a trade-off occurs between resource-dependent energetic and fitness costs and benefits of raising the current litter to independence
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Fisher, David N.; Boutin, Stan; Dantzer, Ben; Humphries, Murray M.; Lane, Jeffrey E.; McAdam, Andrew G. 2022-04-26 Individuals often interact more closely with some members of the population (e.g. offspring, siblings or group members) than they do with other individuals. This structuring of interactions can lead to multilevel natural selection, where traits expressed at the group-level influence fitness alongside individual-level traits. Such multilevel selection can alter evolutionary trajectories, yet is rarely quantified in the wild, especially for species that do not interact in clearly demarcated groups. We quantified multilevel natural selection on two traits, postnatal growth rate and birth date, in a population of North American red squirrels (Tamiasciurus hudsonicus). The strongest level of selection was typically within-acoustic social neighbourhoods (within 130m of the nest), where growing faster and being born earlier than nearby litters was key, while selection on growth rate was also apparent both within-litters and within-study areas. Higher population densities increased the strength of selection for earlier breeding, but did not influence selection on growth rates. Females experienced especially strong selection on growth rate at the within-litter level, possibly linked to the biased bequeathal of the maternal territory to daughters. Our results demonstrate the importance of considering multilevel and sex-specific selection in wild species, including those that are territorial and sexually monomorphic.
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Serrouya, Robert; Paetkau, David; McLellan, Bruce N.; Boutin, Stan; Jenkins, Deborah A.; Campbell, Mitch 2012-03-02 Identifying conservation units below the species level is becoming increasingly important, particularly when limited resources necessitate prioritization for conservation among such units. This problem is exemplified with caribou, a mammal with a circum-Arctic distribution that is exposed to a broad spectrum of ecological conditions, but is also declining in many parts of its range. We used microsatellite markers to evaluate the suitability of existing intra-specific taxonomic designations to act as population units for conservation, and contrasted this with landscape features that were independent of taxonomy. We also quantified the relationship between genetic differentiation and subpopulation size, a factor that has been under-represented in landscape genetic research. Our dataset included three subspecies and three ecotypes of caribou that varied in population size by five orders of magnitude. Our results indicated that genetic structure did not correspond to existing taxonomic designation, particularly at the level of ecotype. Instead, we found that major valleys and population size were the strongest factors associated with substructure. There was a negative exponential relationship between population size and FST between pairs of adjacent subpopulations, suggesting that genetic drift was the mechanism causing the structure among the smallest subpopulations. A genetic assignment test revealed that movement among subpopulations was a fraction of the level needed to stabilize smaller subpopulations, indicating little chance for demographic rescue. Such results may be broadly applicable to landscape genetic studies, because population size and corresponding rates of drift have the potential to confound interpretations of landscape effects on population structure.
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DeMars, Craig; Boutin, Stan; Serrouya, Robert; Gilbert, Sophie; Kelly, Allicia; Larter, Nicholas; Hervieux, Dave 2021-10-07 <p>As global climate change progresses, wildlife management will benefit from knowledge of demographic responses to climatic variation, particularly for species already endangered by other stressors. In Canada, climate change is expected to increasingly impact populations of threatened woodland caribou (<i>Rangifer tarandus caribou</i>) and much focus has been placed on how a warming climate has potentially facilitated the northward expansion of apparent competitors and novel predators. Climate change, however, may also exert more direct effects on caribou populations that are not mediated by predation. These effects include meteorological changes that influence resource availability and energy expenditure. Research on other ungulates suggests that climatic variation may have minimal impact on low-density populations such as woodland caribou because per-capita resources may remain sufficient even in “bad” years. We evaluated this prediction using demographic data from 21 populations in western Canada that were monitored for various intervals between 1994 and 2015. We specifically assessed whether juvenile recruitment and adult female survival were correlated with annual variation in meteorological metrics and plant phenology. Against expectations, we found that both vital rates appeared to be influenced by annual climatic variation. Juvenile recruitment was primarily correlated with variation in phenological conditions in the year prior to birth. Adult female survival was more strongly correlated with meteorological conditions and declined during colder, more variable winters. These responses may be influenced by the life history of woodland caribou, which reside in low-productivity refugia where small climatic changes may result in changes to resources that are sufficient to elicit strong demographic effects. Across all models, explained variation in vital rates was low, suggesting that other factors had greater influence on caribou demography. Nonetheless, given the declining trajectories of many woodland caribou populations, our results highlight the increased relevance of recovery actions when adverse climatic conditions are likely to negatively affect caribou demography.</p>
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Menzies, Allyson; Studd, Emily; Majchrzak, Yasmine; Peers, Michael; Boutin, Stan; Dantzer, Ben; Lane, Jeffrey; McAdam, Andrew; Humphries, Murray 2020-07-31 <ol> <li style="margin-bottom:12px;">Organisms survive environmental variation by combining homeostatic regulation of critical states with allostatic variation of other traits, and species differences in these responses can contribute to coexistence in temporally-variable environments.</li> <li style="margin-bottom:12px;">In this paper, we simultaneously record variation in three functional traits – body temperature (Tb), heart rate, and activity - in relation to three forms of environmental variation – air temperature (Ta), photoperiod, and experimentally-manipulated resource levels – in free-ranging snowshoe hares and North American red squirrels to characterize distinctions in homeotherm responses to the extreme conditions of northern boreal winters.</li> <li style="margin-bottom:12px;">Hares and squirrels differed in the level and precision of Tb regulation, but also in the allostatic pathways necessary to maintain thermal homeostasis. Hares demonstrated a stronger metabolic pathway (through heart rate variation reflective of the thermogenesis), while squirrels demonstrated a stronger behavioral pathway (through activity variation that minimizes cold exposure).</li> <li style="margin-bottom:12px;">As intermediate-sized, winter-active homeotherms, hares and squirrels share many functional attributes, yet, through the integrated monitoring of multiple functional traits in response to shared environmental variation, our study reveals many pairwise species differences in homeostatic and allostatic traits, that both define and are defined by the natural history, functional niches, and coexistence of sympatric species.</li> </ol> https://creativecommons.org/publicdomain/zero/1.0/
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Fletcher, Quinn E.; Speakman, John R.; Boutin, Stan; Lane, Jeffrey E.; McAdam, Andrew G.; Gorrell, Jamieson C.; Coltman, David W.; Humphries, Murray M. 2015-06-13 1. Energy expenditure is a trait of central importance in ecological and evolutionary theory. We examined the correlates of, the strength of selection on, and the heritability of, daily energy expenditure (DEE; kJ/day) during lactation in free-ranging North American red squirrels (Tamiasciurus hudsonicus). 2. Over seven years, lactating squirrels with greater DEE had higher annual reproductive success (ARS; standardized selection gradient: β’ = 0.47; top 12% of published estimates). Surprisingly, positive fecundity selection on lactation DEE for increased ARS did not result because lactation DEE was correlated with typical measures of reproductive performance and/or investment. 3. We found no evidence of costs of elevated lactation DEE acting through female survival, subsequent year lactation DEE, or subsequent year reproduction. 4. Lactation DEE was not significantly repeatable and heritability was not significantly different from zero. 5. Elevated lactation DEE enhances ARS through a link between DEE and an unidentified measure of maternal or environmental quality, but there is limited evolutionary potential for lactation DEE to respond to our documented selection.
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DeMars, Craig; Boutin, Stan 2020-03-20 Rapid landscape alteration associated with human activity is currently challenging the evolved dynamical stability of many predator-prey systems by forcing species to behaviorally respond to novel environmental stimuli. In many forested systems, linear features (LFs) such as roads, pipelines and resource exploration lines (i.e. seismic lines) are a ubiquitous form of landscape alteration that have been implicated in altering predator-prey dynamics. One hypothesized effect is that LFs facilitate predator movement into and within prey refugia, thereby increasing predator-prey spatial overlap. We evaluated this hypothesis in a large mammal system, focusing on the interactions between boreal woodland caribou (Rangifer tarandus caribou) and their two main predators, wolves (Canis lupus) and black bears (Ursus americanus), during the calving season of caribou. In this system, LFs extend into and occur within peatlands (i.e. bogs and nutrient-poor fens), a habitat type highly used by caribou due to its refugia effects. Using resource selection analyses, we found that LFs increased predator selection of peatlands. Female caribou appeared to respond by avoiding LFs and areas with high LF density. However, in our study area most caribou cannot completely avoid exposure to LFs and variation in female response had demographic effects. In particular, increasing proportional use of LFs by females negatively impacted survival of their neonate calves. Collectively, these results demonstrate how LFs can reduce the efficacy of prey refugia. Mitigating such effects will require limiting or restoring LFs within prey refugia, though the effectiveness of mitigation efforts will depend upon spatial scale, which in turn will be influenced by the life history traits of predator and prey.
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Dickie, Melanie; Serrouya, Robert; McNay, R. Scott; Boutin, Stan 2017-06-21 Predation by grey wolves Canis lupus has been identified as an important cause of boreal woodland caribou Rangifer tarandus caribou mortality, and it has been hypothesized that wolf use of human-created linear features such as seismic lines, pipelines and roads increases movement, resulting in higher kill rates. We tested if wolves select linear features and whether movement rates increased while travelling on linear features in north-eastern Alberta and north-western Saskatchewan using 5-min GPS (Global Positioning System) locations from twenty-two wolves in six packs. Wolves selected all but two linear feature classes, with the magnitude of selection depending on feature class and season. Wolves travelled two to three times faster on linear features compared to the natural forest. Increased average daily travelling speed while on linear features and increased proportion of steps spent travelling on linear features increased net daily movement rates, suggesting that wolf use of linear features can increase their search rate. Synthesis and applications. Our findings that wolves move faster and farther on human-created linear features can inform mitigation strategies intended to decrease predation on woodland caribou, a threatened species. Of the features that can realistically be restored, mitigation strategies such as silviculture and linear deactivation (i.e. tree-felling and fencing) should prioritize conventional seismic lines (i.e. cleared lines used for traditional oil and gas exploration) and pipelines, as they were selected by wolves and increased travelling speed, before low-impact seismic lines.
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Fletcher, Quinn E.; Selman, Colin; Boutin, Stan; McAdam, Andrew G.; Woods, Sarah B.; Seo, Arnold Y.; Leeuwenburgh, Christiaan; Speakman, John R.; Humphries, Murray M. 2012-11-08 A central principle in life-history theory is that reproductive effort negatively affects survival. Costs of reproduction are thought to be physiologically-based, but the underlying mechanisms remain poorly understood. Using female North American red squirrels (Tamiasciurus hudsonicus), we test the hypothesis that energetic investment in reproduction overwhelms investment in antioxidant protection, leading to oxidative damage. In support of this hypothesis we found that the highest levels of plasma protein oxidative damage in squirrels occurred during the energetically-demanding period of lactation. Moreover, plasma protein oxidative damage was also elevated in squirrels that expended the most energy and had the lowest antioxidant protection. Finally, we found that squirrels that were food-supplemented during lactation and winter had increased antioxidant protection and reduced plasma protein oxidative damage providing the first experimental evidence in the wild that access to abundant resources can reduce this physiological cost.
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Peers, Michael; Konkolics, Sean; Lamb, Clayton; Majchrzak, Yasmine; Menzies, Allyson; Studd, Emily; Boonstra, Rudy; Kenney, Alice; Krebs, Charles; Martinig, April Robin; McCulloch, Baily; Silva, Joseph; Garland, Laura; Boutin, Stan 2020-10-09 <p>1. 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.</p> <p>2. 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 (<i>Lepus americanus</i>), influence scavenging communities in the northern boreal forest. We further examined the impact of habitat and temperature on scavenging dynamics.</p> <p>3. 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.</p> <p>4. 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 (<i>Lynx canadensis</i>). 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.</p> <p>5. 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.</p>
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Krebs, Charles; Boonstra, Rudy; Boutin, Stan; Krebs, Charles J. 2018-06-07 1. 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. 2. 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. 3. 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. 4. 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. 5. 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.
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McFarlane, S. Eryn; Boutin, Stan; Humphries, Murray M.; McAdam, Andrew G.; Gorrell, Jamieson C.; Coltman, David W. 2015-01-21 A trait must genetically correlate with fitness in order to evolve in response to natural selection, but theory suggests that strong directional selection should erode additive genetic variance in fitness and limit future evolutionary potential. Balancing selection has been proposed as a mechanism that could maintain genetic variance if fitness components trade off with one another and has been invoked to account for empirical observations of higher levels of additive genetic variance in fitness components than would be expected from mutation–selection balance. Here, we used a long-term study of an individually marked population of North American red squirrels (Tamiasciurus hudsonicus) to look for evidence of (1) additive genetic variance in lifetime reproductive success and (2) fitness trade-offs between fitness components, such as male and female fitness or fitness in high- and low-resource environments. “Animal model” analyses of a multigenerational pedigree revealed modest maternal effects on fitness, but very low levels of additive genetic variance in lifetime reproductive success overall as well as fitness measures within each sex and environment. It therefore appears that there are very low levels of direct genetic variance in fitness and fitness components in red squirrels to facilitate contemporary adaptation in this population.

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