When death comes: linking predator-prey activity patterns to timing of mortality to understand predation risk.

The assumption that activity and foraging are risky for prey underlies many predator-prey theories and has led to the use of predator-prey activity overlap as a proxy of predation risk. However, the simultaneous measures of prey and predator activity along with timing of predation required to test this assumption have not been available. Here, we used accelerometry data on snowshoe hares (Lepus americanus) and Canada lynx (Lynx canadensis) to determine activity patterns of prey and predators and match these to precise timing of predation. Surprisingly we found that lynx kills of hares were as likely to occur during the day when hares were inactive as at night when hares were active. We also found that activity rates of hares were not related to the chance of predation at daily and weekly scales, whereas lynx activity rates positively affected the diel pattern of lynx predation on hares and their weekly kill rates of hares. Our findings suggest that predator-prey diel activity overlap may not always be a good proxy of predation risk, and highlight a need for examining the link between predation and spatio-temporal behaviour of predator and prey to improve our understanding of how predator-prey behavioural interactions drive predation risk.

Does coat colour influence survival? A test in a cyclic population of snowshoe hares.

Some mammal species inhabiting high-latitude biomes have evolved a seasonal moulting pattern that improves camouflage via white coats in winter and brown coats in summer. In many high-latitude and high-altitude areas, the duration and depth of snow cover has been substantially reduced in the last five decades. This reduction in depth and duration of snow cover may create a mismatch between coat colour and colour of the background environment, and potentially reduce the survival rate of species that depend on crypsis. We used long-term (1977-2020) field data and capture-mark-recapture models to test the hypothesis that whiteness of the coat influences winter apparent survival in a cyclic population of snowshoe hares (Lepus americanus) at Kluane, Yukon, Canada. Whiteness of the snowshoe hare coat in autumn declined during this study, and snowshoe hares with a greater proportion of whiteness in their coats in autumn survived better during winter. However, whiteness of the coat in spring did not affect subsequent summer survival. These results are consistent with the hypothesis that the timing of coat colour change in autumn can reduce overwinter survival. Because declines in cyclic snowshoe hare populations are strongly affected by low winter survival, the timing of coat colour change may adversely affect snowshoe hare population dynamics as climate change continues.

Balancing food acquisition and predation risk drives demographic changes in snowshoe hare population cycles.

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.

Rapidly declining body size in an insectivorous bat is associated with increased precipitation and decreased survival.

Reduced food availability is implicated in declines in avian aerial insectivores, but the effect of nutritional stress on mammalian aerial insectivores is unclear. Unlike birds, insectivorous bats provision their young through lactation, which might protect nursing juveniles when prey availability is low but could increase the energetic burden on lactating females. We analyzed a 15-year capture-mark-recapture data set from 5312 individual little brown myotis (Myotis lucifugus) captured at 11 maternity colonies in northwestern Canada, to test the hypothesis that nutritional stress is impacting these mammalian aerial insectivores. We used long-bone (forearm [FA]) length as a proxy for relative access to nutrition during development, and body mass as a proxy for access to nutrition prior to capture. Average FA length and body mass both decreased significantly over the study period in adult females and juveniles, suggesting decreased access to nutrition. Effect sizes were very small, similar to those reported for declining body size in avian aerial insectivores. Declines in juvenile body mass were only observed in individuals captured in late summer when they were foraging independently, supporting our hypothesis that lactation provides some protection to nursing young during periods of nutritional stress. Potential drivers of the decline in bat size include one or both of (1) declining insect (prey) abundance, and (2) declining prey availability. Echolocating insectivorous bats cannot forage effectively during rainfall, which is increasing in our study area. The body mass of captured adult females and juveniles in our study was lower, on average, after periods of high rainfall, and higher after warmer-than-average periods. Finally, survival models revealed a positive association between FA length and survival, suggesting a fitness consequence to declines in body size. Our study area has not yet been impacted by bat white-nose syndrome (WNS), but research elsewhere has suggested that fatter bats are more likely to survive infection. We found evidence for WNS-independent shifts in the body size of little brown myotis, which can inform studies investigating population responses to WNS. More broadly, the cumulative effects of multiple stressors (e.g., disease, nutritional stress, climate change, and other pressures) on mammalian aerial insectivores require urgent attention.

Snow depth influences harvest of a boreal ungulate more than socio-economic factors: Implications for food security in a changing climate.

Climate change may affect the ability of hunters to harvest wildlife and, hence, threaten food security of local people. However, few studies have investigated the relative influence of environmental conditions on wildlife harvest rates. We harnessed a 24-year dataset of harvest dates for a boreal ungulate in a region where climate change is having pronounced impacts on snow depth, precipitation, and temperatures to investigate the effect of weather on harvest rates. We used generalized linear models and a model selection framework to examine the influence of weather covariates (snow depth, mean daily temperature, precipitation) and socio-economic factors (gasoline and red meat prices, employment rates, and moose [Alces americanus] harvest) on harvest rates of bison (Bison bison) in Yukon, Canada, at two temporal scales: annual and daily. At an annual scale, snow depth was the only covariate that was important in explaining bison harvest. No socioeconomic variables improved our model beyond the null. At the daily scale, snow depth and mean daily temperature influenced bison harvest rates, with a 1 SD increase resulting in a 14 % and 9 % increase in daily harvest rates, respectively. Increased snow depth facilitates ease of travel in remote, roadless areas by snowmobile to locate bison and truncates movements of bison, resulting in increased harvest rates. Decreased snow depth due to climate change will impact hunter access to boreal ungulates and food security for northern people. More broadly, our data suggests that in some socioecological systems, environmental covariates have a greater influence on wildlife harvest rates than socioeconomic factors and need to be considered in future studies to better understand and predict harvest rates.

Coat color mismatch improves survival of a keystone boreal herbivore: Energetic advantages exceed lost camouflage.

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.

Diet and landscape characteristics drive spatial patterns of mercury accumulation in a high-latitude terrestrial carnivore.

Limited information exists on mercury concentrations and environmental drivers of mercury bioaccumulation in high latitude terrestrial carnivores. Spatial patterns of mercury concentrations in wolverine (Gulo gulo, n = 419) were assessed across a 1,600,000 km2 study area in relation to landscape, climate, diet and biological factors in Arctic and boreal biomes of western Canada. Hydrogen stable isotope ratios were measured in wolverine hair from a subset of 80 animals to assess the spatial scale for characterizing environmental conditions of their habitat. Habitat characteristics were determined using GIS methods and raster datasets at two scales, the collection location point and a 150 km radius buffer, which was selected based on results of a correlation analysis between hydrogen stable isotopes in precipitation and wolverine hair. Total mercury concentrations in wolverine muscle ranged >2 orders of magnitude from 0.01 to 5.72 µg/g dry weight and varied geographically, with the highest concentrations in the Northwest Territories followed by Nunavut and Yukon. Regression models at both spatial scales indicated diet (based on nitrogen stable isotope ratios) was the strongest explanatory variable of mercury concentrations in wolverine, with smaller though statistically significant contributions from landscape variables (soil organic carbon, percent cover of wet area, percent cover of perennial snow-ice) and distance to the Arctic Ocean coast. The carbon and nitrogen stable isotope ratios of wolverine muscle suggested greater mercury bioaccumulation could be associated with feeding on marine biota in coastal habitats. Landscape variables identified in the modelling may reflect habitat conditions which support enhanced methylmercury transfer to terrestrial biota. Spatially-explicit estimates of wet atmospheric deposition were positively correlated with wolverine mercury concentrations but this variable was not selected in the final regression models. These landscape patterns provide a basis for further research on underlying processes enhancing methylmercury uptake in high latitude terrestrial food webs.

Isotopic evidence for bioaccumulation of aerosol lead in fish and wildlife of western Canada.

Lead (Pb) is a toxic element which is released as a result of anthropogenic activities, and Pb stable isotope ratios provide a means to distinguish sources and transport pathways in receiving environments. In this study, isotopes of bioaccumulated Pb (204Pb, 206Pb, 207Pb, 208Pb) were examined for diverse terrestrial and aquatic biota from three areas in western Canada: (a) otter, marten, gulls, terns, and wood frogs in the Alberta Oil Sands Region (AOSR), (b) fish, plankton, and gulls of Great Slave Lake (Yellowknife, Northwest Territories), and (c) wolverine from the Yukon. Aquatic and terrestrial biota from different habitats and a broad geographic area showed a remarkable similarity in their Pb isotope composition (grand mean ± 1 standard deviation: 206Pb/207Pb = 1.189 ± 0.007, 208Pb/207Pb = 2.435 ± 0.009, n = 116). Comparisons with Pb isotope ratios of local sources and environmental receptors showed that values in biota were most similar to those of atmospheric Pb, either measured in local aerosols influenced by industrial activities in the AOSR or in lichens (an aerosol proxy) near Yellowknife and in the Yukon. Biotic Pb isotope ratios were different from those of local geogenic Pb. Although the Pb isotope measurements could not unambiguously identify the specific anthropogenic sources of atmospheric Pb in biota, initial evidence points to the importance of fossil fuels currently used in transportation and power generation. Further research should characterize bioavailable chemical species of Pb in aerosols and important emission sources in western Canada.

High prevalence, intensity, and genetic diversity of Trichinella spp. in wolverine (Gulo gulo) from Yukon, Canada.

BACKGROUND: Species of Trichinella are globally important foodborne parasites infecting a number of domestic and wild vertebrates, including humans. Free-ranging carnivores can act as sentinel species for detection of Trichinella spp. Knowledge of the epidemiology of these parasites may help prevent Trichinella spp. infections in northern Canadian animals and people. Previous research on Trichinella spp. in wildlife from Yukon did not identify risk factors associated with infection, or the diversity and identity of species of Trichinella in regional circulation, based on geographically extensive sampling with large sample sizes. METHODS: In a cross-sectional study, we determined the prevalence, infection intensity, risk factors, and species or genotypes of Trichinella in wolverine (Gulo gulo) in two regions of Yukon, Canada, from 2013-2017. A double separatory funnel digestion method followed by mutiplex PCR and PCR-RFLP were used to recover and identify species of Trichinella, respectively. RESULTS: We found larvae of Trichinella in the tongues of 78% (95% CI 73-82) of 338 wolverine sampled. The odds of adult (≥ 2 years) and yearling (1 year) wolverine being Trichinella spp.-positive were four and two times higher, respectively, compared to juveniles (<1 year). The odds of Trichinella spp. presence were three times higher in wolverine from southeast than northwest Yukon. The mean intensity of infection was 22.6 ± 39 (SD, range 0.1-295) larvae per gram. Trichinella T6 was the predominant genotype (76%), followed by T. nativa (8%); mixed infections with Trichinella T6 and T. nativa (12%) were observed. In addition, T. spiralis was detected in one wolverine. Out of 22 isolates initially identified as T. nativa in multiplex PCR, 14 were analyzed by PCR-RFLP to distinguish them from T. chanchalensis, a recently discovered cryptic species, which cannot be distinguished from the T. nativa on multiplex PCR. Ten isolates were identified either as T. chanchalensis alone (n = 7), or mixed infection with T. chanchalensis and T. nativa (n = 2) or T. chanchalensis and Trichinella T6 (n = 1)]. CONCLUSIONS: Wolverine hosted high prevalence, high larval intensity, and multiple species of Trichinella, likely due to their scavenging habits, apex position in the food chain, and wide home range. Wolverine (especially adult males) should be considered as a sentinel species for surveys for Trichinella spp. across their distributional range.

Hiding in plain sight: discovery and phylogeography of a cryptic species of Trichinella (Nematoda: Trichinellidae) in wolverine (Gulo gulo).

Understanding parasite diversity and distribution is essential in managing the potential impact of parasitic diseases in animals and people. Imperfect diagnostic methods, however, may conceal cryptic species. Here, we report the discovery and phylogeography of a previously unrecognized species of Trichinella in wolverine (Gulo gulo) from northwestern Canada that was indistinguishable from T. nativa using the standard multiplex PCR assay based on the expansion segment 5 (ESV) of ribosomal DNA. The novel genotype, designated as T13, was discovered when sequencing the mitochondrial genome. Phylogenetic analyses of the mitochondrial genome and of 15 concatenated single copy orthologs of nuclear DNA indicated a common ancestor for the encapsulated clade is shared by a subclade containing Trichinella spiralis and Trichinella nelsoni, and a subclade containing T13 and remaining taxa: T12 + (T2 + T6) + [(T5 + T9) + (T3 + T8)]. Of 95 individual hosts from 12 species of mammalian carnivores from northwestern Canada from which larvae were identified as T. nativa on multiplex PCR, only wolverines were infected with T13 (14 of 42 individuals). These infections were single or mixed with T. nativa and/or T6. Visual examination and motility testing confirmed that T13 is encapsulated and likely freeze-tolerant. We developed a new Polymerase Chain Reaction-Restriction Fragment Length Polymorphism which unequivocally distinguishes between T13 and T. nativa. We propose Trichinella chanchalensis n. sp. for T13, based on significant genetic divergence from other species of Trichinella and broad-based sampling of the Trichinella genome. Exploration of Alaskan and Siberian isolates may contribute to further resolution of a phylogeographically complex history for species of Trichinella across Beringia, including Trichinella chanchalensis n. sp. (T13).

HEALTH STATUS OF REINTRODUCED WOOD BISON ( BISON BISON ATHABASCAE): ASSESSING THE CONSERVATION VALUE OF AN ISOLATED POPULATION IN NORTHWESTERN CANADA.

A central goal for reintroduced populations of threatened wood bison ( Bison bison athabascae) is to maintain them free of diseases of concern, particularly bovine tuberculosis (caused by Mycobacterium bovis) and brucellosis (caused by Brucella abortus). A wood bison population in southwestern Yukon, Canada was reintroduced into the wild in 1988, but no health assessment has been done since then. To provide an initial assessment of the health status and, hence, the conservation value of this population, we serologically tested 31 wood bison (approximately 3% of the population) for pathogens of interest and obtained histopathology results for select tissues. We found no evidence of exposure to M. bovis or Brucella spp., but antibodies were present to bovine parainfluenza virus 3, bovine coronavirus, Leptospira interrogans, and Neospora caninum, with seroprevalences of 87, 7, 61, and 7% of the tested animals, respectively. Reintroduced wood bison in southwestern Yukon may be of high value for wood bison recovery because it is a large and geographically isolated population with no bacteriologic, histopathologic, or serologic evidence of exposure to Brucella spp. or M. bovis.

Impact of rewilding, species introductions and climate change on the structure and function of the Yukon boreal forest ecosystem.

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.

Efficacy of a Butorphanol, Azaperone, and Medetomidine Combination for Helicopter-Based Immobilization of Bison ( Bison bison).

Decreased access to potent narcotics for wildlife applications has stimulated the need to explore alternative drug combinations for ungulate immobilizations. A combination of butorphanol, azaperone, and medetomidine (BAM) has been used for some ungulate species, but information on its use in bison ( Bison bison) is limited. We conducted field trials using BAM, in conjunction with atipamezole and naltrexone as antagonists, for reversible field immobilization of bison during ground- and helicopter-based operations. We compared times to induction and recovery, vital rates (rectal temperature and respiration rate), and the quality of induction, immobilization, and recovery between ground- and helicopter-based immobilizations of bison. Overall, 15 of 21 bison were induced with the volume we used (mean±SD=3.4±0.6 mL); two other animals darted from a helicopter required a full second dose, and four others (two darted from the ground and two from a helicopter) required a supplemental partial dose to achieve induction. All immobilizations achieved a sufficient plane of anesthesia to permit minor invasive procedures (e.g., skin biopsy and blood sampling). All animals recovered, and most (17 of 21) were reversed in ≤5 min. The mean time to induction was 10.8±7.3 min while that for recovery was 5.0±2.1 min. We found few differences in vital rates or the quality of immobilizations between ground- and helicopter-based captures. The drug combination provided good immobilization and was reliably reversed; however, inconsistent inductions at the doses we used may limit its use in field immobilizations of bison, particularly those animals being darted from a helicopter.

Tongue has higher larval burden of Trichinella spp. than diaphragm in wolverines (Gulo gulo).

Trichinella is an important zoonotic parasite found in a range of wildlife species harvested for food and fur in Canada. We compared larval intensity from tongue and diaphragm, the best predilection sites in other animal species, from naturally infected, wild wolverines (Gulo gulo) (n = 95). Muscle larvae of Trichinella spp. were recovered by the pepsin/HCl artificial digestion method (gold standard) using double separatory funnels, and species were identified using multiplex PCR. Prevalence was 83% (79/95). Of those positive for Trichinella spp. (n = 79), 76 (96.2%) were detected in both tissues, 2 (2.5%) were positive only on diaphragm, and 1 (1.3%) only on tongue. A total of 62 of 79 wolverines (78.5%) had higher larval burden in tongue than in diaphragm, whereas 17 wolverines (21.5%) had higher larval burden in diaphragm. The predilection site (higher larval burden) of Trichinella spp. larvae did not vary significantly between juvenile and adult wolverines (P = 0.2), between male and female wolverines (P = 0.9), and among wolverines classified as having low and high larval intensities overall (P = 0.2). Trichinella T6 was the predominant genotype (63 of 79; 80%), followed by T. nativa (T2) (6 of 79; 8%). Mixed infections of T2 and T6 were observed in 9 of 79 (12%) wolverines. Larval intensity of Trichinella T6 was higher in tongues than diaphragms. No statement can be made for T2 due to insufficient T2 positive samples. In conclusion, tongues are a better site for sampling than diaphragms in future surveys of Trichinella larval intensity in wolverines; however, either tissue is suitable for prevalence studies.

Spatial variation of mercury bioaccumulation in bats of Canada linked to atmospheric mercury deposition.

Wildlife are exposed to neurotoxic mercury at locations distant from anthropogenic emission sources because of long-range atmospheric transport of this metal. In this study, mercury bioaccumulation in insectivorous bat species (Mammalia: Chiroptera) was investigated on a broad geographic scale in Canada. Fur was analyzed (n=1178) for total mercury from 43 locations spanning 20° latitude and 77° longitude. Total mercury and methylmercury concentrations in fur were positively correlated with concentrations in internal tissues (brain, liver, kidney) for a small subset (n=21) of little brown bats (Myotis lucifugus) and big brown bats (Eptesicus fuscus), validating the use of fur to indicate internal mercury exposure. Brain methylmercury concentrations were approximately 10% of total mercury concentrations in fur. Three bat species were mainly collected (little brown bats, big brown bats, and northern long-eared bats [M. septentrionalis]), with little brown bats having lower total mercury concentrations in their fur than the other two species at sites where both species were sampled. On average, juvenile bats had lower total mercury concentrations than adults but no differences were found between males and females of a species. Combining our dataset with previously published data for eastern Canada, median total mercury concentrations in fur of little brown bats ranged from 0.88-12.78µg/g among 11 provinces and territories. Highest concentrations were found in eastern Canada where bats are most endangered from introduced disease. Model estimates of atmospheric mercury deposition indicated that eastern Canada was exposed to greater mercury deposition than central and western sites. Further, mean total mercury concentrations in fur of adult little brown bats were positively correlated with site-specific estimates of atmospheric mercury deposition. This study provides the largest geographic coverage of mercury measurements in bats to date and indicates that atmospheric mercury deposition is important in determining spatial patterns of mercury accumulation in a mammalian species.