Effects of predator treatments,individual traits,and environment on moose survival in Alaska

We studied moose (Alces alces) survival, physical condition, and abundance in a 3-predator system in western Interior Alaska, USA, during 2001–2007. Our objective was to quantify the effects of predator treatments on moose population dynamics by investigating changes in survival while evaluating the contribution of potentially confounding covariates. In May 2003 and 2004, we reduced black bear (Ursus americanus) and brown bear (U. arctos) numbers by translocating bears ≥240 km from the study area. Aircraft-assisted take reduced wolf (Canis lupus) numbers markedly in the study area during 2004–2007. We estimated black bears were reduced by approximately 96% by June 2004 and recovered to within 27% of untreated numbers by May 2007. Brown bears were reduced approximately 50% by June 2004. Late-winter wolf numbers were reduced by 75% by 2005 and likely remained at these levels through 2007. In addition to predator treatments, moose hunting closures during 2004–2007 reduced harvests of male moose by 60% in the study area. Predator treatments resulted in increased calf survival rates during summer (primarily from reduced black bear predation) and autumn (primarily from reduced wolf predation). Predator treatments had little influence on survival of moose calves during winter; instead, calf survival was influenced by snow depth and possibly temperature. Increased survival of moose calves during summer and autumn combined with relatively constant winter survival in most years led to a corresponding increase in annual survival of calves following predator treatments. Nonpredation mortalities of calves increased following predator treatments; however, this increase provided little compensation to the decrease in predation mortalities resulting from treatments. Thus, predator-induced calf mortality was primarily additive. Summer survival of moose calves was positively related to calf mass (β > 0.07, SE = 0.073) during treated years and lower (β = −0.82, SE = 0.247) for twins than singletons during all years. Following predator treatments, survival of yearling moose increased 8.7% for females and 21.4% for males during summer and 2.2% for females and 15.6% for males during autumn. Annual survival of adult (≥2 yr old) female moose also increased in treated years and was negatively (β = −0.21, SE = 0.078) related to age. Moose density increased 45%, from 0.38 moose/km² in 2001 to 0.55 moose/km² in 2007, which resulted from annual increases in overall survival of moose, not increases in reproductive rates. Indices of nutritional status remained constant throughout our study despite increased moose density. This information can be used by wildlife managers and policymakers to better understand the outcomes of predator treatments in Alaska and similar environments. © 2011 The Wildlife Society.     

Habitat use by black bears in relation to conspecifics and competitors

Sympatric black bears (Ursus americanus) and brown bears (Ursus arctos) are common in many boreal systems; however, few predator assemblages are known to coexist on a single seasonally abundant large prey item. In lowland southwestern interior Alaska, black bears and brown bears are considered the primary cause of moose (Alces alces) calf mortality during the first 6 weeks of life. The objective of this study was to document habitat use of global-positioning system (GPS)-collared black bears during peak and non-peak seasons of black bear-induced and brown bear-induced moose calf mortality within southwestern interior Alaska, in spring 2002. We compared habitats of GPS-collared black bears to those of presumably uncollared black bears and brown bears at their moose calf mortality sites. Results from this study suggest that GPS-collared black bears use similar habitat as conspecifics more than expected during the peak period of black bear predation on moose calves, whereas they use habitat in proportion to home range availability during the peak in brown bear predation on moose calves. Sex-specific Ivlev's electivity indices describe greater than expected use of mixed-deciduous forest and needleleaf forest by male GPS-collared black bears during the peak of moose calf predation, whereas females have a tendency to use these habitats less than expected. Juvenile GPS-collared black bears largely use the same habitat as other sympatric predators during the peak of moose calf predation, whereas during the non-peak period juveniles use opposite habitats as adult GPS-collared black bears. The outcome of this study offers possible explanations (e.g., sex, age) for spatial overlap or segregation in one member of a complex predator guild in relation to a seasonal pulse of preferred prey.     

Managing for Elevated Yield of Moose in Interior Alaska

ABSTRACT Given recent actions to increase sustained yield of moose (Alces alces) in Alaska, USA, we examined factors affecting yield and moose demographics and discussed related management. Prior studies concluded that yield and density of moose remain low in much of Interior Alaska and Yukon, Canada, despite high moose reproductive rates, because of predation from lightly harvested grizzly (Ursus arctos) and black bear (U. americanus) and wolf (Canis lupus) populations. Our study area, Game Management Unit (GMU) 20A, was also in Interior Alaska, but we describe elevated yield and density of moose. Prior to our study, a wolf control program (1976–1982) helped reverse a decline in the moose population. Subsequent to 1975, moose numbers continued a 28-year, 7-fold increase through the initial 8 years of our study (λ_B1 = 1.05 during 1996–2004, peak density = 1,299 moose/1,000 km²). During these initial 8 hunting seasons, reported harvest was composed primarily of males ( = 88%). Total harvest averaged 5% of the prehunt population and 57 moose/1,000 km², the highest sustained harvest-density recorded in Interior Alaska for similar-sized areas. In contrast, sustained total harvests of <10 moose/1,000 km² existed among low-density, predator-limited moose populations in Interior Alaska (≤417 moose/1,000 km²). During the final 3 years of our study (2004–2006), moose numbers declined (λ_B2 = 0.96) as intended using liberal harvests of female and male moose ( = 47%) that averaged 7% of the prehunt population and 97 moose/1,000 km². We intentionally reduced high densities in the central half of GMU 20A (up to 1,741 moose/1,000 km² in Nov) because moose were reproducing at the lowest rate measured among wild, noninsular North American populations. Calf survival was uniquely high in GMU 20A compared with 7 similar radiocollaring studies in Alaska and Yukon. Low predation was the proximate factor that allowed moose in GMU 20A to increase in density and sustain elevated yields. Bears killed only 9% of the modeled postcalving moose population annually in GMU 20A during 1996–2004, in contrast to 18–27% in 3 studies of low-density moose populations. Thus, outside GMU 20A, higher bear predation rates can create challenges for those desiring rapid increases in sustained yield of moose. Wolves killed 8–15% of the 4 postcalving moose populations annually (10% in GMU 20A), hunters killed 2–6%, and other factors killed 1–6%. Annually during the increase phase in GMU 20A, calf moose constituted 75% of the predator-killed moose and predators killed 4 times more moose than hunters killed. Wolf predation on calves remained largely additive at the high moose densities studied in GMU 20A. Sustainable harvest-densities of moose can be increased several-fold in most areas of Interior Alaska where moose density and moose: predator ratios are lower than in GMU 20A and nutritional status is higher. Steps include 1) reducing predation sufficient to allow the moose population to grow, and 2) initiating harvest of female moose to halt population growth and maximize harvest after density-dependent moose nutritional indices reach or approach the thresholds we previously published.     

Influence of Predator Harvest,Biological Factors,and Landscape on Elk Calf Survival in Idaho

ABSTRACT We evaluated survival of elk (Cervus elaphus) calves on 2 contrasting study areas in north-central Idaho, USA, from 1997 to 2004. Recruitment was modest (>30 calves:100 F [calves of either sex: F elk 1 yr old]) and stable on the South Fork study area and low (<20 calves:100 F) and declining on the Lochsa study area. The primary proximate cause of calf mortality on both study areas was predation by black bears (Ursus americanus) and mountain lions (Puma concolor). We experimentally manipulated populations of black bears and mountain lions on a portion of each study area. Black bear harvest (harvest density/600km²) initially doubled on the Lochsa treatment after manipulating season bag limits. Mountain lion harvest also increased by 60% but varied widely during the manipulation period. Harvest seasons were closed for black bears and mountain lions on the treatment portion of the South Fork study area. Using the Andersen—Gill formulation (A-G) of the Cox proportional hazards model, we examined effects of landscape structure, predator harvest levels, and biological factors on summer calf survival. We used Akaike's Information Criterion (AIC_c) and multimodel inference to assess some potentially useful predictive factors relative to calf survival. We generated risk ratios for both the best models and for model-averaged coefficients. Our models predicted that calf survival was influenced by biological factors, landscape surrounding calf locations, and predator harvest levels. The model that best explained mortality risk to calves on the Lochsa included black bear harvest (harvest density/600 km²), estimated birth mass of calves, and percentage of shrub cover surrounding calf locations. Incorporating a shrub X time interaction allowed us to correct for nonproportionality and detect that effect of shrub cover was only influential during the first 14 days of a calf's life. Model-averaging indicated that estimated birth mass of calves and black bear harvest were twice as important as the next variables, but age of calves at capture was also influential in calf survival. The model that best explained mortality risk to calves on the South Fork included black bear harvest, age of calves at capture, and gender of calves. Model-averaging indicated that age at capture and black bear harvest were twice as important as the next variable, forest with 33–66% canopy cover (Canopy 33–66). Risk to calves decreased when calves occupied areas with more of this forest cover type. Model-averaging also indicated that increased mountain lion harvest lowered calf mortality risk 4% for every 1-unit increase in lion harvest (harvest density/600 km²) but was lower (<25%) in importance compared to age at capture and black bear harvest. Our results suggest that levels of predator harvest, and presumably predator density, resource limitations expressed through calf birth mass, and habitat structure had substantial effects on calf survival. Our results can be generalized to other areas where managers are dealing with low calf elk recruitment. However, because factors vary spatially, a single management strategy applied in different areas will probably not have the same effect on calf survival.     

Extensive Predator Space Use Can Limit the Efficacy of a Control Program

ABSTRACT Reduced to small isolated groups by anthropogenic habitat losses or habitat modifications, populations of many endangered species are sensitive to additive sources of mortality, such as predation. Predator control is often one of the first measures considered when predators threaten survival of a population. Unfortunately, predator ecology is often overlooked because relevant data are difficult to obtain. For example, the endangered Gaspésie caribou (Rangifer tarandus caribou) has benefited from 2 periods of predator control that targeted black bears (Ursus americanus) and coyotes (Canis latrans) in an attempt to reduce predation on caribou calves. Despite a high trapping effort, the number of predators removed has remained stable over time. To assess impact of predator movements on efficacy of a control program, we studied space use of 24 black bears and 16 coyotes over 3 years in and around the Gaspésie Conservation Park, Quebec, Canada, using Global Positioning System radiocollars. Annual home ranges of black bears averaged 260 km² and 10 individuals frequented area used by caribou. Annual home ranges of resident coyotes averaged 121 km², whereas dispersing coyotes covered >2,600 km². Coyotes were generally located at lower altitudes than caribou. However, because coyotes undertook long-distance excursions, they overlapped areas used by caribou. Simulations based on observed patterns showed that 314 bears and 102 coyotes potentially shared part of their home range with areas used by female caribou during the calving period. Despite low densities of both predator species, extensive movement and use of nonexclusive territories seem to allow predators to rapidly occupy removal areas, demonstrating the need for recurrent predator removals. Our results underscore the necessity of considering complementary and alternative solutions to predator control to assure long-term protection of endangered species.     

Modeling female brown bear kill rates on moose calves using global positioning satellite data

The recent development in Global Positioning System (GPS) techniques has started a new era in predation studies. Estimates of kill rates based on animal movements and GPS relocation clusters have proven to be valid in several obligatory carnivores. The main focus has been to obtain accurate mean predation estimates for the management of wildlife populations. We present a model to estimate individual kill rates of moose calves by adult female brown bears in Sweden, based on spatiotemporal clustering of 30,889 bear GPS relocations and 71 moose calves verified killed during 714 field investigations in 2004–2006. In this virtually single-predator single large prey system, the omnivorous brown bear is an efficient predator on moose calves up to 4 weeks of age. The top model set only included models with cluster radii of 30 m or 50 m, indicating very high kill-site fidelity. The best model included a cluster radius of 30 m and number of periods of bear activity at the kill site as a single covariate. The mean estimated individual kill rate of 7.6 ± 0.71 (n = 18, ± SE) moose calves per calving season is comparable to the estimate of 6.8 from a previous study of radio-tracked moose in our study area, though at a lower moose/bear ratio. The mean annual kill rates varied from 6.1 to 9.4 calves per bear. The estimated individual kill rates ranged from 2 to 15 calves per season, indicating a large individual variation in hunting skills and possibly effort. Predation and livestock depredation represent a core conflict between humans and carnivores in rural Scandinavia. Accurate predation estimates represent an important step in quantifying costs of carnivores and reducing human–carnivore conflicts. Our technique may be applied in the exploration of predation mechanisms and predator–prey interactions, and contribute to the old and global debate of problem individuals in livestock depredation. © 2012 The Wildlife Society.     

Mobility of moose—comparing the effects of wolf predation risk,reproductive status,and seasonality

In a predator–prey system, prey species may adapt to the presence of predators with behavioral changes such as increased vigilance, shifting habitats, or changes in their mobility. In North America, moose (Alces alces) have shown behavioral adaptations to presence of predators, but such antipredator behavioral responses have not yet been found in Scandinavian moose in response to the recolonization of wolves (Canis lupus). We studied travel speed and direction of movement of GPS‐collared female moose (n = 26) in relation to spatiotemporal differences in wolf predation risk, reproductive status, and time of year. Travel speed was highest during the calving (May–July) and postcalving (August–October) seasons and was lower for females with calves than females without calves. Similarly, time of year and reproductive status affected the direction of movement, as more concentrated movement was observed for females with calves at heel, during the calving season. We did not find support for that wolf predation risk was an important factor affecting moose travel speed or direction of movement. Likely causal factors for the weak effect of wolf predation risk on mobility of moose include high moose‐to‐wolf ratio and intensive hunter harvest of the moose population during the past century.     

Conservation of wildlife populations: factoring in incremental disturbance

Progressive anthropogenic disturbance can alter ecosystem organization potentially causing shifts from one stable state to another. This potential for ecosystem shifts must be considered when establishing targets and objectives for conservation. We ask whether a predator–prey system response to incremental anthropogenic disturbance might shift along a disturbance gradient and, if it does, whether any disturbance thresholds are evident for this system. Development of linear corridors in forested areas increases wolf predation effectiveness, while high density of development provides a safe‐haven for their prey. If wolves limit moose population growth, then wolves and moose should respond inversely to land cover disturbance. Using general linear model analysis, we test how the rate of change in moose (Alces alces) density and wolf (Canis lupus) harvest density are influenced by the rate of change in land cover and proportion of land cover disturbed within a 300,000 km² area in the boreal forest of Alberta, Canada. Using logistic regression, we test how the direction of change in moose density is influenced by measures of land cover change. In response to incremental land cover disturbance, moose declines occurred where <43% of land cover was disturbed; in such landscapes, there were high rates of increase in linear disturbance and wolf density increased. By contrast, moose increases occurred where >43% of land cover was disturbed and wolf density declined. Wolves and moose appeared to respond inversely to incremental disturbance with the balance between moose decline and wolf increase shifting at about 43% of land cover disturbed. Conservation decisions require quantification of disturbance rates and their relationships to predator–prey systems because ecosystem responses to anthropogenic disturbance shift across disturbance gradients.     

Disturbance-Mediated Apparent Competition Decouples in a Northern Boreal Caribou Range

The most widely reported threat to boreal and mountain populations of woodland caribou (Rangifer tarandus caribou; caribou) involves habitat- or disturbance-mediated apparent competition (DMAC). With DMAC, natural and anthropogenic disturbances that increase the abundance of deciduous-browsing cervids (e.g., moose [Alces alces], deer [Odocoileus spp.]) are thought to promote predator (especially wolf [Canis lupus]) numbers, which heightens predation risk to caribou. We know most about the effects of DMAC on caribou where the species is under threat by anthropogenic activities in relatively productive southern boreal and mountain systems. Yet, >60% of extant boreal caribou range in North America consists of northern shield and taiga ecoregions of low productivity where caribou may compete with only 1 ungulate species (moose) in the context of DMAC. In this environment, we know very little of how DMAC acts as a limiting factor to caribou. In Saskatchewan, Canada, from 2014–2018, using a combination of vegetation sampling, aerial surveys, and telemetry data (n = 38 wolves), we searched for evidence of DMAC (trends in data consistent with the hypothesis) in an 87,193-km² section of the Western Boreal Shield, a poorly productive but pristine region (0.18% of land cover classed as an anthropogenic feature) with a historically high fire-return interval (47% of stands aged <40 years). Despite the high levels of disturbance, moose density was relatively low (47 moose/1,000 km²), likely because of the scarcity of deciduous or mixed-wood stands and low abundance of deciduous browse in the young conifer stands that dominated the landscape. In contrast, boreal caribou density was relatively high for the species (37 caribou/1,000 km²). Wolf density (3.1 wolves/1,000 km²) and pack sizes ( = 4.0 wolves/pack) were low and resident (established) territories were large ( = 4,360 km²; 100% minimum convex polygon). The low density of wolves mirrored the low (standardized) ungulate biomass index (UBI; moose + boreal caribou) of the study area (0.36 UBI/km²). We conclude that wolf and hence caribou populations were not responding in accordance with the outcomes generally predicted by DMAC in our study area because the requisite strong, positive response to fire of deciduous-browse and alternate-prey abundance was lacking. As a limiting factor to caribou, DMAC is likely modulated at a macroecological scale by factors such as net primary productivity, a corollary to the general hypothesis that we advance here (i.e., primary productivity hypothesis of DMAC). We caution against managing for caribou based on the presumption of DMAC where the mechanism does not apply, which may include much of boreal caribou range in the north. © 2020 The Wildlife Society.     

Science and Values Influencing Predator Control for Alaska Moose Management

ABSTRACT We encourage informed and transparent decision-making processes concerning the recently expanded programs in Alaska, USA, to reduce predation on moose (Alces alces). The decision whether to implement predator control ultimately concerns what society should value; therefore, policymakers, not objective biologists, play a leadership role. From a management and scientific standpoint, biological support for these predator-control programs requires convincing evidence that 1) predators kill substantial numbers of moose that would otherwise mostly live and be available for harvest, 2) low predation can facilitate reliably higher harvests of moose, 3) given less predation, habitats can sustain more moose and be protected from too many moose, and 4) sustainable populations of Alaska's brown bears (Ursus arctos), black bears (Ursus americanus), and wolves (Canis lupus) will exist in and out of control areas. We reviewed 10 moose mortality studies, 36 case histories, 10 manipulative studies, 15 moose nutrition studies, and 3 recent successful uses of nutrition-based management to harvest excess female moose. Results of these studies support application of long-term, substantial predator control for increasing yield of moose in these simple systems where moose are a primary prey of 3 effective predators. We found no substantive, contradictory results in these systems. However, to identify and administer feasible moose population objectives, recently established moose nutritional indices must be monitored, and regulatory bodies must accept nutrition-based management. In addition, the efficacy of techniques to reduce bear predation requires further study. Predicting precise results of predator control on subsequent harvest of moose will continue to be problematic because of a diversity of changing interactions among biological, environmental, and practical factors. In Alaska, the governor has the prerogative to influence regulations on predator control by appointing members to the Board of Game. At least annually, the Board of Game hears a wide spectrum of public opinions opposing and favoring predator control. We summarized these opinions as well as the societal and cultural values and expectations that are often the primary basis for debates. Advocates on both sides of the debate suggest they hold the higher conservation ethic, and both sides provide biased science. We recommend a more constructive and credible dialogue that focuses openly on values rather than on biased science and fabricated conspiracies. To be credible and to add substance in this divisive political arena, biologists must be well informed and provide complete information in an unbiased and respectful manner without exaggeration.     

Predator densities and white-tailed deer fawn survival

Predation is the dominant source of mortality for white-tailed deer (Odocoileus virginianus) <6 months old throughout North America. Yet, few white-tailed deer fawn survival studies have occurred in areas with 4 predator species or have considered concurrent densities of deer and predator species. We monitored survival and cause-specific mortality from birth to 6 months for 100 neonatal fawns during 2013–2015 in the Upper Peninsula of Michigan, USA, while simultaneously estimating population densities of deer, American black bear (Ursus americanus), coyote (Canis latrans), bobcat (Lynx rufus), and gray wolf (Canis lupus). We estimated fawn predation risk in response to sex, birth mass, and date of birth. Six-month fawn survival pooled among years was 36%, and fawn mortality risk was not related to birth mass, date of birth, or sex. Estimated mean annual deer and predator densities were 334 fawns/100 km², 25.9 black bear/100 km², 23.8 coyotes/100 km², 3.8 bobcat/100 km², and 2.8 wolves/100 km². Despite lower estimated per-individual kill rates, coyotes and black bears were the leading sources of fawn mortality because they had greater densities relative to bobcats and wolves. Our results indicate that the presence of more predator species in a system is not entirely additive in its effect on fawn survival. © The Wildlife Society, 2019     

Moose-wolf dynamics and the natural regulation of moose populations

Summary In southwestern Québec, non-harvested moose populations stabilize at a density of 0.40 animal·km^-2. In an attempt to test whether or not moose were regulated by predators, we investigated wolf predation near this equilibrium density (0.37) and at 2 lower densities (0.23, 0.17). Scat analysis in summer and feeding observations in winter indicated a greater use of alternative food resources by wolves at lower moose densities. Each wolf pack killed on average 5.3, 1.8, 1.1 moose·100 days in the area of 0.37, 0.23, and 0.17 moose·km^-2, respectively. Consumption of moose per wolf was 2.8, 1.7, and 1.6 kg/day, respectively. January wolf densities were estimated at 1.38, 0.82, and 0.36 animals·100 km^-2, respectively. Year-long predation rates proved to be density-dependent, increasing with moose density from 6.1 to 19.3% of the postnatal populations. We conclude that moose populations in southwestern Québec are regulated largely by predators (wolves and maybe black bears) at a density where competition for forage produces no detrimental effect. We support the concept that wolf predation can have an important regulatory effect at low moose densities but also a depensatory (inversely density-dependent) effect at higher densities.     

Invading white-tailed deer change wolf–caribou dynamics in northeastern Alberta

Human-caused habitat change has been implicated in current woodland caribou (Rangifer tarandus caribou) population declines across North America. Increased early seral habitat associated with industrial footprint can result in an increase in ungulate densities and subsequently those of their predator, wolves (Canis lupus). Higher wolf densities can result in increased encounters between wolves and caribou and consequently higher caribou mortality. We contrasted changes in moose (Alces alces) and deer (Odocoileus spp.) densities and assessed their effects on wolf–caribou dynamics in northeastern Alberta, Canada, pre (1994–1997) versus post (2005–2009) major industrial expansion in the region. Observable white-tailed deer (O. virginianus) increased 17.5-fold but moose remained unchanged. Wolf numbers also increased from approximately 6–11.5/1,000 km². Coincident with these changes, spatial overlap between wolf pack territories and caribou range was high relative to the mid-1990s. The high number of wolf locations in caribou range suggests that forays were not merely exploratory, but rather represented hunting forays and denning locations. Scat analysis indicated that wolf consumption of moose declined substantively during this time period, whereas use of deer increased markedly and deer replaced moose as the primary prey of wolves. Caribou increased 10-fold in the diet of wolves and caribou population trends in the region changed from stable to declining. Wolf use of beaver (Castor canadensis) increased since the mid-1990s. We suggest that recent declines in woodland caribou populations in the southerly extent of their range have occurred because high deer densities resulted in a numeric response by wolves and consequently higher incidental predation on caribou. Our results indicate that management actions to conserve caribou must now include deer in primary prey and wolf reduction programs. © 2010 The Wildlife Society     

The non-impact of hunting on moose <Emphasis Type="Italic">Alces alces</Emphasis> movement,diurnal activity,and activity range

Previous studies on moose Alces alces have suggested that interactions with humans may trigger anti-predator behaviors and generate a demographical cost. Therefore, we hypothesized that disturbances from small and big game hunting may have negative effects on moose movements, diurnal activity, and activity range. Using location data from 64 moose equipped with GPS collars from three populations (Low Alpine, Inland, Coastal) with different temporal human presence and spatial accessibility, we evaluated the impact of hunting on moose activity rhythms. On average, female moose in the low human population density (Low Alpine) area (<0.5/km²) had significantly lower movement rates during moose hunting season, but variation in movement rates among individuals were higher compared with female moose in regions with denser human populations (6–24/km²). We found no evidence that reproductive status influenced female moose sensitivity to disturbance. As expected, females used smaller activity ranges and were less active nocturnally than males. The high within-group variation suggests that current hunting disturbance levels do not alter moose population behavior in general. Our data indicate that alterations in movement were related to rutting activity, not human disturbance induced by hunting. In line with behavioral theory, our study suggests that some individuals were more sensitive to hunting disturbance than the general population. Our work suggests that individual moose may perceive human predation risk to be similar to other predation risks.     

Summer kill rates and predation pattern in a wolf–moose system: can we rely on winter estimates?

So far the vast majority of studies on large carnivore predation, including kill rates and consumption, have been based on winter studies. Because large carnivores relying on ungulates as prey often show a preference for juveniles, kill rates may be both higher and more variable during the summer season than during the rest of the year leading to serious underestimates of the total annual predation rate. This study is the first to present detailed empirical data on kill rates and prey selection in a wolf–moose system during summer (June–September) as obtained by applying modern Global Positioning System-collar techniques on individual wolves (Canis lupus) in Scandinavia. Moose (Alces alces) was the dominant prey species both by number (74.4%) and biomass (95.6%); 89.9% of all moose killed were juveniles, representing 76.0% of the biomass consumed by wolves. Kill rate in terms of the kilogram biomass/kilogram wolf per day averaged 0.20 (range: 0.07–0.32) among wolf territories and was above, or well above, the daily minimum food requirements in most territories. The average number of days between moose kills across wolf territories and study periods was 1.71 days, but increased with time and size of growing moose calves during summer. Over the entire summer (June–September, 122 days), a group (from two to nine) of wolves killed a total of 66 (confidence interval 95%; 56–81) moose. Incorporation of body growth functions of moose calves and yearlings and wolf pups over the summer period showed that wolves adjusted their kill rate on moose, so the amount of biomass/kilogram wolf was relatively constant or increased. The kill rate was much higher (94–116%) than estimated from the winter period. As a consequence, projecting winter kill rates to obtain annual estimates of predation in similar predator–prey systems may result in a significant underestimation of the total number of prey killed. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Moose calf mortality in central Ontario,Canada

Although some populations remain stable, moose (Alces alces) density and distribution have been declining in many areas along the southern edge of their North American distribution. During 2006–2009, we deployed 99 vaginal implant transmitters (VITs) in 86 adult female moose in central Ontario, Canada to assist in locating and radiocollaring neonatal moose calves. We monitored radiocollared calves to estimate calf survival and assess the relative importance of specific causes of death. Calves in the western portion of our study area (WMU49) were exposed to a 6-day general hunting season, whereas calves in the eastern portion of our study area (Algonquin Provincial Park [APP]) were not exposed to hunting. Annual survival for 87 collared calves was greater in the protected area than the harvested area (72.4 ± 6.8% and 55.8 ± 8.3%, respectively) and averaged 63.7 ± 7.1% overall. Predation by wolves (Canis sp.) and American black bears (Ursus americanus) was the dominant cause of death but occurred predominately in APP, whereas other natural mortality agents were 4× more common in WMU49. Only 16% of the collared calves in WMU49 were harvested each year despite a high proportion (approx. 50%) of accessible, public land. Most natural mortality occurred prior to the autumn hunting season such that reductions in natural mortality had little potential to compensate for calf harvest. Overall, calf survival in our study area was moderate to high and our findings suggest predator control or further restrictions of calf hunting in this area is not justified. © The Wildlife Society, 2013     

Home ranges of brown bears on the Kamchatka peninsula and Sakhalin Island

Territorial activity was studied using satellite tracking of four brown bears (Ursus arctos) in Kamchatka in 2005–2006 and three brown bears on Sakhalin in 2011–2012. The size of annual home ranges was 6.09–27.58 km² for females and 153.12 km² for males. The size of the nuclear zone of the annual home ranges did not exceed 1.68 km². Seasonal home ranges were largest in August-September and smallest in May. The home ranges of two females in Kamchatka were significantly overlapped, the lesser degree of overlap was noted for two females on Sakhalin. The nature of the use of the study area by bears, essentially depends on the seasonal distribution of food, in particular salmon.     

Encounter frequencies between GPS-collared wolves (Canis lupus) and moose (Alces alces) in a Scandinavian wolf territory

Over 6,000 GPS fixes from two wolves (Canis lupus) and 30,000 GPS fixes from five moose (Alces alces) in a wolf territory in southern Scandinavia were used to assess the static and dynamic interactions between predator and prey individuals. Our results showed that wolves were closer to some of the moose when inside their home ranges than expected if they had moved independently of each other, and we also found a higher number of close encounters (<500 m) than expected. This suggests that the wolves were actively seeking the individual moose within their territory. Furthermore, the wolves showed a preference for moving on gravel forest roads, which may be used as convenient travel routes when patrolling the territory and seeking areas where the moose are. However, due to the particularly large size of the wolf territory combined with relatively high moose densities, the wolves generally spent a very small proportion of their time inside the home range of each individual moose, and the frequency of encounters between the wolves and any particular moose was very low. We suggest that the high moose:wolf ratio in this large Scandinavian wolf territory compared to that typically occurring in North America, results in a relatively low encounter frequency and a low predation risk for individual moose, as the predation pressure is spread over a high number of prey individuals.     

Molecular identification of Taenia spp. in wolves (Canis lupus), brown bears (Ursus arctos) and cervids from North Europe and Alaska

Taenia tapeworms of Finnish and Swedish wolves (Canis lupus) and Finnish brown bears (Ursus arctos), and muscle cysticerci of Svalbard reindeer (Rangifer tarandus platyrhynchus), Alaskan Grant's caribou (Rangifer tarandus granti) and Alaskan moose (Alces americanus) were identified on the basis of the nucleotide sequence of a 396 bp region of the mitochondrial cytochrome c oxidase subunit 1 gene. Two species were found from wolves: Taenia hydatigena and Taenia krabbei. The cysticerci of reindeer, caribou and one moose also represented T. krabbei. Most of the cysticercal specimens from Alaskan moose, however, belonged to an unknown T. krabbei-like species, which had been reported previously from Eurasian elks (Alces alces) from Finland. Strobilate stages from two bears belonged to this species as well. The present results suggest that this novel Taenia sp. has a Holarctic distribution and uses Alces spp. as intermediate and ursids as final hosts.     

Black bear movements and habitat use during a critical period for moose calves

In sub-Arctic and north-temperate ecosystems, opportunistic carnivores, such as black bears (Ursus americanus) and brown bears (Ursus arctos), are active on the landscape for a shorter period annually than sympatric gray wolves (Canis lupus). Therefore, bear movement patterns and habitat use might be expected to be more deliberate and of greater consequence, in terms of energy acquisition, than those of predators not undergoing hibernation. Habitat choices concerning feeding, bedding, and denning grounds made by black bears therefore should reflect seasonal abundance and distribution of vegetation and key prey items as these are sites where bears remain and forage for prolonged periods of time. We recorded the movement patterns of 6 GPS-collared black bears from den emergence to onset of moose (Alces alces) parturition in 2003. Over approximately 3 weeks prior to parturition, results from average distance calculations suggest that black bears moved closer to probable moose calving-site habitat. Additionally, the seasonal habitat use by black bears surrounding dens reflected the same trend for areas where cow moose gave birth in spring 2003, with a propensity to use needleleaf forest more than any other habitat.