Patterns of Mortality and Factors Influencing Survival of a Recently Restored Elk Population in Ontario,Canada

This study examined patterns of mortality and determinants of survival among elk recently restored to four sites in Ontario, Canada (1998–2005). We predicted that: (1) elk located in release sites closer to the core of their historic range would have higher survival; (2) survival would increase as an animal's time and experience on the landscape increased; and (3) survival rates would decline as animals moved farther away from the release site. During the study, 443 elk were radiocollared and released; 218 mortalities were documented. Predation by wolves was the most important proximate cause of mortality, followed by death due to injuries from translocation and/or capture myopathy, accidents, emaciation, poaching, and Parelaphostrongylus tenuis infection. Overall, annual survival of elk across Ontario ranged from 0.45 (0.37–0.53) to 0.81 (0.66–0.90), with rates being lowest in the years immediately following release and highest in the final years of the study; this pattern was due to high initial mortality from translocation injuries and/or capture myopathy and possibly lack of familiarity with novel habitat. Model‐averaged hazards further support this finding, as the most important factor influencing elk survival was the length of holding period, with elk released after limited holding being less likely to survive than those held for longer periods. Our results suggest that mortalities caused by capture myopathy and transportation‐related injuries are important sources of risk for translocated elk. The method of introduction to the novel landscape and behavior in the first year should be accommodated via soft‐release and appropriate release areas.     

A Seventy-Year History of Trends in Yellowstone's Northern Elk Herd

ABSTRACT We analyzed counts of northern Yellowstone elk (Cervus elaphus) in Yellowstone National Park, Wyoming, USA, over 70 years to evaluate the effects of changing management on population trends. Population reduction efforts and hunter harvests during 1932–1968 removed 71,330 elk and decreased estimated abundance from 16,000 to 6,000 elk. Abundance increased to approximately 17,000 elk (λ = 1.19) when removals ceased and harvests were very small during 1969–1975. Moderate to liberal hunter harvests of antlerless elk outside the Park during 1976–2004 removed a relatively consistent proportion (26 ± 0.1 [SD]%) of females that migrated outside the park, mostly from prime-age (3–15 yr) classes with high reproductive value. Substantial winterkill was infrequent (1989, 1997), but it significantly reduced calf survival when it occurred. Wolves (Canis lupus) were reintroduced in 1995–1996 and rapidly increased in abundance (λ = 1.23) and distribution. Estimated wolf kill of elk now exceeds hunter harvest, but has a smaller effect on population dynamics because wolves concentrate on calves and older females (>14 yr) with low reproductive value. During 1995–2004, estimated abundance decreased from 23,000 to 12,000 elk. The recent ratio of wolves to elk is relatively low compared to the estimated equilibrium ratio, suggesting that the wolf population may yet increase in the future. Thus, reduction of harvests of prime-aged female elk to decrease removals of animals with high reproductive value and increase adult female survival appears essential. We analyzed the relative impact of removals by hunters and by wolves using Fisher's (1930) reproductive value and found that the impact of hunters is far more important than that by wolves, a finding of broad significance.     

Temporal variation in site fidelity: scale-dependent effects of forage abundance and predation risk in a non-migratory large herbivore

Large herbivores are typically confronted by considerable spatial and temporal variation in forage abundance and predation risk. Although animals can employ a range of behaviours to balance these limiting factors, scale-dependent movement patterns are expected to be an effective strategy to reduce predation risk and optimise foraging opportunities. We tested this prediction by quantifying site fidelity of global positioning system-collared, non-migratory female elk (Cervus canadensis manitobensis) across multiple nested temporal scales using a long-established elk–wolf (Canis lupus) system in Manitoba, Canada. Using a hierarchical analytical approach, we determined the combined effect of forage abundance and predation risk on variation in site fidelity within four seasons across four nested temporal scales: monthly, biweekly, weekly, daily. Site fidelity of female elk was positively related to forage-rich habitat across all seasons and most temporal scales. At the biweekly, weekly and daily scales, elk became increasingly attached to low forage habitat when risk was high (e.g. when wolves were close or pack sizes were large), which supports the notion that predator-avoidance movements lead to a trade-off between energetic requirements and safety. Unexpectedly, predation risk at the monthly scale increased fidelity, which may indicate that elk use multiple behavioural responses (e.g. movement, vigilance, and aggregation) simultaneously to dilute predation risk, especially at longer temporal scales. Our study clearly shows that forage abundance and predation risk are important scale-dependent determinants of variation in site fidelity of non-migratory female elk and that their combined effect is most apparent at short temporal scales. Insight into the scale-dependent behavioural responses of ungulate populations to limiting factors such as predation risk and forage variability is essential to infer the fitness costs incurred.     

Bark Beetle-Affected Forests Provide Elk Only a Marginal Refuge from Hunters

For nearly 2 decades, the forests of the Rocky Mountains in the United States experienced a bark-beetle (Dendoctronus ponderosae) epidemic. The number of dead and falling trees from this epidemic likely will affect how elk (Cervus canadensis) and hunters use the forest and their interactions. Downed trees potentially create a component of refuge habitat that could affect the effectiveness of hunting to regulate abundance of growing elk populations. We evaluated how forests affected by bark beetles in south-central Wyoming, USA, influenced resource selection of 50 female elk and 374 hunters in 2012–2016, and interactions between elk and hunters. We employed global positioning system (GPS) technology on elk and hunters, and developed a satellite-derived land classification specifically depicting beetle-affected forests. We tested the predictions that elk would increase use of beetle-killed areas from summer to the hunting season, and that hunters would avoid beetle-killed areas regardless of elk use. Elk increased use of beetle-killed areas during hunting seasons as did hunters during the archery season; however, during the rifle season, hunters avoided beetle-killed areas. Nevertheless, during the rifle season, areas of beetle-kill with a high probability of elk occurrence dampened the aversion hunters had towards beetle-killed areas. Therefore, in contrast to our expectations, forests that have been altered by the bark-beetle epidemic may only function marginally as a refuge for elk. Our study area was at the beginning of the tree-fall phase (i.e., ~3–7 yr after peak infestation) of the bark-beetle epidemic; thus, future research efforts should focus on how a continued increase in downed trees will influence interactions between elk and hunters. At current levels of tree fall, however, beetle-kill should not limit the ability of managers to regulate elk herds through harvest. © 2020 The Wildlife Society.     

Neonatal mortality of elk driven by climate, predator phenology and predator community composition

1.?Understanding the interaction among predators and between predation and climate is critical to understanding the mechanisms for compensatory mortality. We used data from 1999 radio-marked neonatal elk (Cervus elaphus) calves from 12 populations in the north-western United States to test for effects of predation on neonatal survival, and whether predation interacted with climate to render mortality compensatory. 2.?Weibull survival models with a random effect for each population were fit as a function of the number of predator species in a community (3-5), seven indices of climatic variability, sex, birth date, birth weight, and all interactions between climate and predators. Cumulative incidence functions (CIF) were used to test whether the effects of individual species of predators were additive or compensatory. 3.?Neonatal elk survival to 3 months declined following hotter previous summers and increased with higher May precipitation, especially in areas with wolves and/or grizzly bears. Mortality hazards were significantly lower in systems with only coyotes (Canis latrans), cougars (Puma concolor) and black bears (Ursus americanus) compared to higher mortality hazards experienced with gray wolves (Canis lupus) and grizzly bears (Ursus horribilis). 4.?In systems with wolves and grizzly bears, mortality by cougars decreased, and predation by bears was the dominant cause of neonatal mortality. Only bear predation appeared additive and occurred earlier than other predators, which may render later mortality by other predators compensatory as calves age. Wolf predation was low and most likely a compensatory source of mortality for neonatal elk calves. 5.?Functional redundancy and interspecific competition among predators may combine with the effects of climate on vulnerability to predation to drive compensatory mortality of neonatal elk calves. The exception was the evidence for additive bear predation. These results suggest that effects of predation by recovering wolves on neonatal elk survival, a contentious issue for management of elk populations, may be less important than the composition of the predator community. Future studies would benefit by synthesizing overwinter calf and adult-survival data sets, ideally from experimental studies, to test the roles of predation in annual compensatory and additive mortality of elk.     

Effects of wolf pack size and winter conditions on elk mortality

Elk (Cervus canadensis) are high-profile game animals for many states in the western United States, yet over the past several decades some populations have experienced a persistent and broad-scale decline in recruitment. Over this same period, gray wolves (Canis lupus) have become an integral component of many western landscapes and agencies are increasingly challenged to maximize hunting opportunities of ungulates via predator management while simultaneously ensuring wolf conservation. To better understand the implications of predator management on elk populations, we monitored survival of 1,244 adult female elk and 806 6-month-old calves from 29 populations distributed throughout Idaho, USA, from 2004 to 2016. We developed predictive models of mortality that related mortality risk to wolf pack size, winter conditions, and individual-level characteristics. Annual mortality rates (excluding harvest) for adult females and calves were 0.09 and 0.40, respectively. Calf mortality was predicted best with a model that included additive effects of chest girth at time of capture, mean size of surrounding wolf packs, and snow depth. Adult female mortality was predicted best with a model that included female age, mean size of surrounding wolf packs, and snow depth. Based on a sensitivity analysis, chest girth had the largest effect on risk of mortality for calves followed by pack size and snow depth. Other than the effect of senescence in the oldest (>15 yr) individuals, pack size and snow depth had the largest effect on risk of mortality for adult females. We estimated cause-specific mortality and predation was the dominant cause of known-fate mortalities for adult females (35% mountain lion [Puma concolor] and 32% wolf) and calves (45% mountain lion and 28% wolf), whereas malnutrition accounted for 9% and 10% of adult female and calf mortalities, respectively. Wolves preferentially selected smaller calves and older adult females, whereas mountain lions showed little preference for calf size or age class of adult females. Our study indicates managers can increase elk survival by reducing wolf pack sizes on surrounding winter ranges, especially in areas where, or during years when, snow is deep. Additionally, managers interested in improving over-winter calf survival can implement actions to increase the size of calves entering winter by increasing the nutritional quality of summer and early fall forage resources. Although our study was prompted by management questions related to wolves, mountain lions killed more elk than wolves and differences in selection of individual elk indicate mountain lions may have comparably more of an effect on elk population dynamics. Although we were unable to relate changes in mountain lion populations to elk survival in our study, future research should seek a better understanding of multi-predator systems, including how management of one predator affect others and ultimately how these interactions affect elk survival. © 2019 The Wildlife Society     

Influences on Release-Site Fidelity of Translocated Elk

Several eastern states are considering the restoration of free‐ranging elk populations via translocation from western populations. Optimal habitat immediately surrounding release sites has been found to enhance elk reintroduction success in western states. Little information exists, however, to aid eastern managers in identifying release sites with the highest chance of restoration success. We monitored the movements of 415 translocated elk released at three sites in southeastern Kentucky to identify landscape characteristics that enhance release‐site fidelity. The distance elk moved after release differed among sites (F_2,322 = 4.63, p = 0.01), age classes (F_2,322 = 4.37, p = 0.01), and time intervals (F_2,322 = 40.74, p < 0.001). At 6 and 12 months post‐release, adults (15.81 ± 17.32 and 16.38 ± 20.29) and yearlings (13.91 ± 16.44 and 14.61 ± 21.11) moved farther than calves (8.06 ± 14.03 and 9.37 ± 14.40). The release site with the highest fidelity was privately owned, 15% open, and had the highest amount of edge compared with the other release sites. The two remaining sites contained large amounts of expansive openland or forest cover with lower amounts of edge. Additionally, both sites were publicly owned and experienced a higher degree of human‐generated disturbance compared with the site to which elk were most faithful. When selecting release sites, managers should avoid areas dominated by a single cover type with little interspersion of other habitats. Rather, areas with high levels of open‐forest edge (approximately 5.0 km/km²) and limited‐human disturbance will likely enhance release‐site fidelity and promote restoration success.     

Contrasting Effects of Wolves and Human Hunters on Elk Behavioral Responses to Predation Risk

ABSTRACT Prey behavioral responses to predation risk in wolf-ungulate-plant systems are of interest to wildlife managers. Using Global Positioning System data collected from telemetry-collared elk (Cervus elaphus) and wolves (Canis lupus), we evaluated elk behavioral responses to spatial and temporal variation in wolf- and human-predation risk on a winter range in the Greater Yellowstone Area, USA. We found elk changed grouping patterns and increased movement rates as predation risk increased and that these behavioral changes were habitat dependent. Elk behavioral responses to wolf- and human-predation risk were similar; however, responses to human-predation risk were stronger than responses to wolf-predation risk. These results suggest that predation risk from wolves or human hunters may result in elk spending more time on private rangelands away from public-land winter ranges, which may exacerbate problems of landowner tolerance of elk on livestock pastures. However, increased movement and changing grouping patterns on winter ranges may also disperse elk grazing impacts and lessen elk impacts on any one area.     

Evaluating postintervention survival of free‐ranging odontocete cetaceans

Until recently, few data were available for evaluating postintervention survival of free‐ranging cetaceans receiving aid from humans through: rescue from stranding, with rehabilitation and release; rescue, rehabilitation and release of debilitated or entangled individuals that had not beached; rescue of entangled animals with immediate release; and rescue, transport, and release of out‐of‐habitat animals. Advances in medical diagnosis, husbandry and therapy have improved survival of rehabilitation cases, and advances in radio‐telemetry have improved postrelease monitoring. In total, 69 cases (1986–2010) were evaluated, involving 10 species of odontocete cetaceans with release data. Findings suggested a success criterion of surviving at least six weeks postrelease is useful in evaluating intervention strategies. No species had better success than others. Stranded beached cetaceans were less successful than free‐swimming rescued animals. Rehabilitated animals were less successful than those released without rehabilitation. Mass stranded dolphins fared better than single stranded animals. Old age, diminished hearing ability, and lack of maternal care were factors in several unsuccessful cases. Success is not clearly related to rehabilitation duration. Retaining healthy individuals from mass strandings until all animals are ready for release may reduce success for some. Transport durations for unsuccessful cases were greater than for successful cases.     

Differential risk effects of wolves on wild versus domestic prey have consequences for conservation

Predators play integral roles in shaping ecosystems through cascading effects to prey and vegetation. Such effects occur when prey species alter their behavior to avoid predators, a phenomenon called the risk effects of predators. Risk effects of wild predators such as wolves are well documented for wild prey, but not for free ranging domestic animals such as cattle despite their importance for ecosystem function and conservation. We compared risk effects of satellite‐collared wolves (n = 16) on habitat selection by global‐positioning‐system‐collared elk (n = 10) and cattle (n = 31). We calculated resource selection functions (RSFs) in periods before, during and after wolf visits in elk home ranges or cattle pastures. The habitat variables tested included: distance to roads and trails, terrain ruggedness, food‐quality and distance to forest. When wolves were present, elk stayed closer to forest cover and selected less for high‐quality‐food habitat. Thus, the risk effects of wolf presence on elk produced a change in the tradeoff between food and cover selection. Cattle responded by avoiding high‐quality‐food habitat and selecting areas closer to roads and trails (where people likely provided security), but these effects manifested only after wolves had left. Artificial selection in cattle may have attenuated natural anti‐predator behaviors. The effects of predators on ecosystems are likely different when mediated through risk effects on domestic compared to wild animals. Furthermore, predator control in response to livestock predation, an important conservation issue, may produce broad ecosystem effects triggered by decrease of an important predator species. Conservation planners should consider these effects where domestic herbivores are dominant species in the ecosystem.     

Survival of Colonizing Wolves in the Northern Rocky Mountains of the United States, 1982–2004

Abstract: After roughly a 60-year absence, wolves (Canis lupus) immigrated (1979) and were reintroduced (1995-1996) into the northern Rocky Mountains (NRM), USA, where wolves are protected under the Endangered Species Act. The wolf recovery goal is to restore an equitably distributed metapopulation of ≥30 breeding pairs and 300 wolves in Montana, Idaho, and Wyoming, while minimizing damage to livestock; ultimately, the objective is to establish state-managed conservation programs for wolf populations in NRM. Previously, wolves were eradicated from the NRM because of excessive human killing. We used Andersen–Gill hazard models to assess biological, habitat, and anthropogenic factors contributing to current wolf mortality risk and whether federal protection was adequate to provide acceptably low hazards. We radiocollared 711 wolves in Idaho, Montana, and Wyoming (e.g., NRM region of the United States) from 1982 to 2004 and recorded 363 mortalities. Overall, annual survival rate of wolves in the recovery areas was 0.750 (95% CI = 0.728-0.772), which is generally considered adequate for wolf population sustainability and thereby allowed the NRM wolf population to increase. Contrary to our prediction, wolf mortality risk was higher in the northwest Montana (NWMT) recovery area, likely due to less abundant public land being secure wolf habitat compared to other recovery areas. In contrast, lower hazards in the Greater Yellowstone Area (GYA) and central Idaho (CID) likely were due to larger core areas that offered stronger wolf protection. We also found that wolves collared for damage management purposes (targeted sample) had substantially lower survival than those collared for monitoring purposes (representative sample) because most mortality was due to human factors (e.g., illegal take, control). This difference in survival underscores the importance of human-caused mortality in this recovering NRM population. Other factors contributing to increased mortality risk were pup and yearling age class, or dispersing status, which was related to younger age cohorts. When we included habitat variables in our analysis, we found that wolves having abundant agricultural and private land as well as livestock in their territory had higher mortality risk. Wolf survival was higher in areas with increased wolf density, implying that secure core habitat, particularly in GYA and CID, is important for wolf protection. We failed to detect changes in wolf hazards according to either gender or season. Maintaining wolves in NWMT will require greater attention to human harvest, conflict resolution, and illegal mortality than in either CID or GYA; however, if human access increases in the future in either of the latter 2 areas hazards to wolves also may increase. Indeed, because overall suitable habitat is more fragmented and the NRM has higher human access than many places where wolves roam freely and are subject to harvest (e.g., Canada and AK), monitoring of wolf vital rates, along with concomitant conservation and management strategies directed at wolves, their habitat, and humans, will be important for ensuring long-term viability of wolves in the region.     

Density‐independent predation affects migrants and residents equally in a declining partially migratory elk population

Migration is expected to benefit individuals through exposure to higher quality forage and reducing predation rates more than non‐migratory conspecifics. Previous studies of partially migratory ungulates (with migrant and resident individuals) have focused on bottom–up factors regulating resident and migrant segments, yet differential predation between strategies could also be a density‐dependent regulatory mechanism. Our study tested for density‐dependence in mortality, as well as mechanisms of ­bottom–up or top–down regulation in the resident and migrant portions of the partially migratory Ya Ha Tinda elk population. We tested for density dependence in adult female and juvenile survival rates, and then discriminated between predator‐ and food‐regulation hypotheses by testing for density‐dependence amongst mortality causes for adult female elk. Notably, the population declined almost 70% from near previously published estimates of carrying capacity over 10 years, providing ideal conditions to test for density dependence. In contrast to predictions, we found only weak support for density dependence in adult survival and juvenile survival. We also found few differences between migrant and resident elk in adult or juvenile survival, though juvenile survival differences were biologically significant. Predation by humans and grizzly bears was density dependent, but similar between migratory strategies. Predation by wolves was the leading known cause of mortality, yet remained constant with declining elk density equally for both migrant and resident elk, indicating wolf predation was density‐independent. Instead of being strongly regulated by food or predation, we found adult female survival was driven by density‐independent predation and climatic factors. The few differences between migratory strategies suggest equivalent fitness payoffs for migrants and residents. This population is being limited by density‐independent predation leading to declines of both migratory strategies. Our results challenge classical predator–prey theory, and call for better integration between predator–prey and migration theory.     

Influence of Group Size on the Success of Wolves Hunting Bison

An intriguing aspect of social foraging behaviour is that large groups are often no better at capturing prey than are small groups, a pattern that has been attributed to diminished cooperation (i.e., free riding) in large groups. Although this suggests the formation of large groups is unrelated to prey capture, little is known about cooperation in large groups that hunt hard-to-catch prey. Here, we used direct observations of Yellowstone wolves (Canis lupus) hunting their most formidable prey, bison (Bison bison), to test the hypothesis that large groups are more cooperative when hunting difficult prey. We quantified the relationship between capture success and wolf group size, and compared it to previously reported results for Yellowstone wolves hunting elk (Cervus elaphus), a prey that was, on average, 3 times easier to capture than bison. Whereas improvement in elk capture success levelled off at 2–6 wolves, bison capture success levelled off at 9–13 wolves with evidence that it continued to increase beyond 13 wolves. These results are consistent with the hypothesis that hunters in large groups are more cooperative when hunting more formidable prey. Improved ability to capture formidable prey could therefore promote the formation and maintenance of large predator groups, particularly among predators that specialize on such prey.     

How Size and Condition Influence Survival and Cause-Specific Mortality of Female Elk

The size of animal populations fluctuates with number of births, rate of immigration, rate of emigration, and number of deaths. For many ungulate populations, adult female survival is the most important factor influencing population growth. Therefore, increased understanding of survival and causes of mortality for adult females is fundamental for conservation and management. The objectives of our study were to quantify survival rates of female elk (Cervus canadensis) and determine cause-specific mortality. We predicted that hunter harvest would be the leading cause of mortality. Further, we predicted that hunters would harvest animals that were in prime age (2–9 yr) and in better condition than elk predated by mountain lions (Puma concolor). From 2015 to 2017, we captured 376 female elk in central Utah, USA. We assessed body size and condition of captured elk, fitted each animal with a global positioning system-collar, and determined cause of death when we received mortality signals. We estimated survival using Kaplan-Meier estimates and Cox proportional hazard models within an Akaike's Information Criterion model selection framework to identify covariates that influenced survival. We analyzed differences in size and condition measurements between harvested elk and predated elk using analysis of variance tests. Our best model indicated consistent survival across years; mean survival was 78.3 ± 3.5% (SE) including hunter harvest and 95.5 ± 1.7% without hunter harvest. In decreasing order of importance, elk mortality occurred from hunter harvest (21.2%), mountain lion predation (3.7%), depredation removal (0.5%), automobile collision (0.3%), disease (0.3%), complications during calving (0.3%), and those characterized as undetermined (1.3%). Neck circumference and body length were negatively associated with survival, suggesting that larger animals in good condition had lower survival as a result of hunter harvest. Individuals that died because of cougar predation were smaller and had less loin muscle than the average animal. Hunters removed large, healthy, prime-aged females, individuals that likely have a greater effect on population growth than elk lost to other predators. If the proportion of larger, healthy females in the population begins to decline, hunting practices may require adjustment because hunters may be removing individuals with the greatest reproductive value. © 2021 The Wildlife Society.     

Evaluating Indirect Effects of Hunting on Mule Deer Spatial Behavior

Mule deer (Odocoileus hemionus) are widely hunted throughout western North America and are experiencing population declines across much of their range. Consequently, understanding the direct and indirect effects of hunting is important for management of mule deer populations. Managers can influence deer mortality rates through changes in hunting season length or authorized tag numbers. Little is known, however, about how hunting can affect site fidelity patterns and subsequent habitat use and movement patterns of mule deer. Understanding these patterns is especially important for adult females because changes in behavior may influence their ability to acquire resources and ultimately affect their productivity. Between 2008 and 2013, we obtained global positioning system locations for 42 adult female deer at the Starkey Experimental Forest and Range in northeast Oregon, USA, during 5-day control and treatment periods in which hunters were absent (pre-hunt), present but not actively hunting (scout and post-hunt), and actively hunting male mule deer (hunt) on the landscape. We estimated summer home ranges and 5-day use areas during pre-hunt and hunt periods and calculated overlap metrics across home ranges and use areas to assess site fidelity within and across years. We used step selection functions to evaluate whether female mule deer responded to human hunters by adjusting fine-scale habitat selection and movement patterns during the hunting season compared to the pre-hunt period. Mule deer maintained site fidelity despite disturbance by hunters with 72 ± 4% (SE) within-year overlap between summer home ranges and hunt use areas and 54 ± 7% inter-annual overlap among pre-hunt use areas and 56 ± 7% among hunt use areas. Mule deer diurnal movement rates, when hunters are active on the landscape, were higher during the hunting period versus pre-hunt or scout periods. In contrast, nocturnal movement rates, when hunters are inactive on the landscape, were similar between hunting and non-hunting periods. Additionally, during the hunt, female mule deer hourly movements increased in areas with high greenness values, indicating that mule deer spent less time in areas with more vegetative productivity. Female mule deer maintained consistent habitat selection patterns before and during hunts, selecting areas that offered more forest canopy cover and high levels of vegetative productivity. Our results indicate that deer at Starkey are adopting behavioral strategies in response to hunters by increasing their movement rates and selecting habitat in well-established ranges. Therefore, considering site fidelity behavior in management planning could provide important information about the spatial behavior of animals and potential energetic costs incurred, especially by non-target animals during hunting season. © 2020 The Wildlife Society.     

Are migrant and resident elk (Cervus elaphus) exposed to similar forage and predation risk on their sympatric winter range?

Partially migratory populations, where one portion of a population conducts seasonal migrations (migrants) while the other remains on a single range (residents), are common in ungulates. Studies that assess trade-offs between migratory strategies typically compare the amount of predation risk and forage resources migrants and residents are exposed to only while on separate ranges and assume both groups intermix completely while on sympatric ranges. Here we provide one of the first tests of this assumption by comparing the amount of overlap between home ranges of GPS-collared migrant and resident elk and fine-scale exposure to wolf predation risk and forage biomass at telemetry locations on a sympatric winter range in west-central Alberta, Canada. Overlap between migrant and resident home ranges increased throughout the winter, and both groups were generally intermixed and exposed to equal forage biomass. During the day, both migrants and residents avoided predation risk by remaining in areas far from timber with high human activity, which wolves avoided. However, at night wolves moved onto the grasslands close to humans and away from timber. Resident elk were consistently closer to areas of human activity and further from timber than migrants, possibly because of a habituation to humans. As a result, resident elk were exposed to higher night-time predation risk than migrants. Our study does not support the assumption that migrant and resident elk are exposed to equal predation risk on their sympatric range when human presence alters predation risk dynamics and habituation to humans is unequal between migratory strategies.     

Postrelease movement and habitat selection of translocated pine martens Martes martes

Monitoring postrelease establishment and movement of animals is important in evaluating conservation translocations. We translocated 39 wild pine martens Martes martes (19 females, 20 males) from Scotland to Wales. We released them into forested areas with no conspecifics in 2015, followed by a second release in 2016, alongside the previously released animals. We used radio‐tracking to describe postrelease movement and habitat selection. Six martens (15%) were not re‐encountered during the tracking period, of which four undertook long‐distance dispersal. For the remaining individuals, we characterized two phases of movement, “exploration” followed by “settlement,” that differed between releases. In the first release, martens remained in exploration phase for a mean of 14.5 days (SE = 3.9 days) and settled at a mean distance of 8.7 km (SE = 1.8 km) from release sites, whereas martens released in year two, alongside resident conspecifics, traveled away from release sites at a faster rate, settling sooner, at a mean of 6.6 days (SE = 1.8 days), but further, at a mean distance of 14.0 km (SE = 1.7 km) from release sites. Animals released in year one did not exhibit habitat preferences overall but within forests they favored recently felled areas, whereas animals released in year two showed strong selection for forested habitat but did not discriminate between forest types. The presence of conspecifics appeared influential for settlement and site fidelity of translocated martens and was associated with more rapid but more distant dispersal of the later cohort. Releases of animals in close proximity appeared to promote site fidelity and rapid establishment of ranges in the recipient environment.     

Correlates of Mortality in an Exploited Wolf Population

Abstract We investigated the influence of habitat use on risk of death from hunting and trapping of 55 radiocollared gray wolves (Canis lupus) from an exploited insular population in Southeast Alaska, USA. We compared mortality rates for resident and nonresident wolves and used Cox proportional hazards regression to relate habitat composition within 100-m circular buffers around radiolocations to risk of death of resident and nonresident wolves. In addition, we included covariates representing distances to roads, logged stands, and lakes and streams in those analyses. We also compiled harvest data from 31 harvest units within the study area to compare densities of roads and distances from human settlements with rates of harvest. During our study 39 wolves died, of which 18 were harvested legally, 16 were killed illegally, and 5 died from natural causes. Legal and illegal harvest accounted for >87% of the mortality of radiocollared resident and nonresident wolves. Mean annual survival was 0.54 (SE = 0.17) for all wolves. Annual survival was 0.65 (SE = 0.17) for resident wolves and 0.34 (SE = 0.17) for nonresidents. Very few (19%) nonresident wolves survived to colonize vacant territories or join existing wolf packs. Roads, muskegs, and distances from lakes and streams were covariates positively associated with death of resident wolves. Clear-cuts were positively associated with risk of death of nonresident wolves. Rate of harvest increased with density of roads; however, road densities >0.9 km/km² had little additional effect on harvest rates. Harvest rates decreased with ocean distances from nearest towns or settlements. Roads clearly increased risk of death for wolves from hunting and trapping and contributed to unsustainable rates of harvest. Wildlife managers should consider effects of roads and other habitat features on harvest of wolves when developing harvest recommendations. They should expect substantial illegal harvest where wolf habitat is accessible to humans. Moreover, high rates of mortality of nonresident wolves exposed to legal and illegal harvest may reduce or delay successful dispersal, potentially affecting linkages between small disjunct wolf populations or population segments. We conclude that a combination of conservative harvest regulations and large roadless reserves likely are the most effective measures for conserving wolves where risks from human-caused mortality are high.     

Postrelease Dispersal of Reintroduced Elk (Cervus elaphus) in Ontario,Canada

We studied 2 years of postrelease telemetry data of elk (Cervus elaphus) translocated to their historic range limit in Ontario, Canada and sought to determine if postrelease movements were related to behavior, demography of released animals, or site–specific attributes such as length of holding period. During 1998–2004 we radio‐tracked 341 elk in 10 release groups via ground and aerial telemetry and monitored movement patterns relative to gender, age, and pre‐release holding period (4–112 days). We found that elk that were held for short periods prior to release (4–11 days) moved longer distances than those subject to extended conditioning (17–112 days), suggesting that an extended conditioning period is beneficial from the standpoint of promoting philopatry. When all elk were pooled by sex and age class, male calves remained in closer proximity (8.0 ± 13.2 km) to release sites than adult females (19.1 ± 20.6 km), adult males (19.7 ± 15.1 km), and female calves (14.4 ± 20.4 km). Most calves dispersed in a southeasterly direction whereas adults tended to travel southwest. Our results reveal that elk movement characteristics are influenced by factors such as release protocol and group demographics; these findings provide further insight regarding appropriate release methods for restoring natural populations near their historical range limit.     

Demographic balancing of migrant and resident elk in a partially migratory population through forage–predation tradeoffs

Partial migration is widespread in ungulates, yet few studies have assessed demographic mechanisms for how these alternative strategies are maintained in populations. Over the past two decades the number of resident individuals of the Ya Ha Tinda elk herd near Banff National Park has been increasing proportionally despite an overall population decline. We compared demographic rates of migrant and resident elk to test for demographic mechanisms partial migration. We determined adult female survival for 132 elk, pregnancy rates for 150 female elk, and elk calf survival for 79 calves. Population vital rates were combined in Leslie‐matrix models to estimate demographic fitness, which we defined as the migration strategy‐specific population growth rate. We also tested for differences in factors influencing risk of mortality between migratory strategies for adult females using Cox‐proportional hazards regression and time‐varying covariates of exposure to forage biomass, wolf predation risk, and group size. Despite higher pregnancy rates and winter calf weights associated with higher forage quality, survival of migrant adult females and calves were lower than resident elk. Resident elk traded high quality food to reduce predation risk by selecting areas close to human activity, and by living in group sizes 20% larger than migrants. Thus, residents experienced higher adult female survival and calf survival, but lower pregnancy and calf weights. Cause‐specific mortality of migrants was dominated by wolf and grizzly bear mortality, whereas resident mortality was dominated by human hunting. Demographic differences translated into slightly higher (2–3%), but non‐significant, resident population growth rate compared to migrant elk, suggesting demographic balancing between resident strategies during our study. Despite statistical equivalence, our results are also consistent with slow long‐term declines in migrants because of high predation because of higher wolf‐caused mortality in migrants. These results emphasize that migrants and residents will make different tradeoffs between forage and risk may affect the demographic balance of partially migratory populations, which may explain recent declines in migratory behavior in many ungulate populations around the world.