Seasonal shifts in habitat selection of a large herbivore and the influence of human activity

Failure to recognize factors contributing to variation in habitat models like resource selection functions (RSFs) can affect their application for projecting probabilities of occurrence, and thereby limit their relevance for conservation and management. We compared seasonal RSFs (2006–2008) for 16 adult female moose (Alces alces) with home ranges located in western Algonquin Provincial Park (APP), Ontario, Canada, to those of 14 adult females located in provincial Wildlife Management Unit (WMU) 49, 40 km west of the protected area. Wildlife and habitat management practices differed between regions: hunting was higher in WMU 49 compared to APP, and APP preserved large tracts of old growth forest rarely found in WMU 49. Seasonal RSFs projected expected similarities in moose resource use between regions (e.g., responses to wetlands and stands of eastern hemlock, Tsuga canadensis [in winter]); however, we also observed differences consistent with the hypothesis that animals, through effects of hunting, would shift habitat use seasonally and in response to roads. We further observed evidence of functional responses in habitat selection due to underlying differences in forestry practices (e.g., responses to stands of old-growth hemlock forest). Given the close proximity and shared biogeographic region between study areas, we believe that observed spatial dynamics in RSFs were ultimately reflective of divergent management strategies between areas and ensuing differences in predation and hunting mortality risk, and functional habitat.     

Survival and cause-specific mortality of moose calves in Northeastern Minnesota

Ungulate reproductive success (calf production and survival) influences population performance. The moose (Alces alces) population in northeastern Minnesota, USA, has declined 65% from 2006 to 2018 but has begun to stabilize. Because causes of this decline were largely unknown, we investigated production, survival, and cause-specific mortality of calves of the global positioning system (GPS)-collared females in this population. In 2013 and 2014, we GPS-collared 74 neonates and monitored them for survival. In 2015 and 2016, we monitored 50 and 35 calving females for signs of neonatal mortality using changes in adult female velocities and assessed seasonal calf survival by aerial surveys. In 2013 and 2014 (pooled), survival to 9 months was 0.34 (95% CI = 0.23–0.52) for collared calves, and in 2015 and 2016 (pooled) survival was 0.35 (95% CI = 0.26–0.48) for uncollared calves. Mortality in all 4 years was high during the first 50 days of life. In 2013 and 2014 (pooled), calving sites were relatively safe for collared neonates; predator-kills occurred a median 17.0 days after departure and a median 1,142 m from calving sites. Predation was the leading cause of death of collared calves (84% of mortalities), with wolves (Canis lupus) accounting for 77% of these. Other forms of mortality for collared and uncollared calves included drowning, infection, vehicle collision, and natural abandonment. We documented higher wolf predation than other recent studies with similar predator communities. Identifying specific causes of calf mortality and understanding their relations to various landscape characteristics and other extrinsic factors should yield insight into mechanisms contributing to the declining moose population in northeastern Minnesota and serve as a basis for ecologically sound management responses. © 2019 The Wildlife Society.     

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.     

Maternal Behavior Indicates Survival and Cause-Specific Mortality of Moose Calves

Continuing research on cause-specific mortality and annual survival of moose (Alces alces) calves in northeastern Minnesota, USA, is important to understanding the long-term trajectory of the population. In 2013 and 2014, we observed global positioning system (GPS)-collared, female moose exhibit a specific behavior (i.e., mortality movement) associated with the death of their GPS-collared neonate. The females made a rapid, long-distance movement (flee), followed by a return to the calf mortality site. We used characteristics of this movement in 2013–2014 (n = 46) to develop models for assessing calf survival, and then evaluated these models using female movement rates (n = 49) in 2015−2016. Using this behavior as an indicator of calf mortality in 2016, we conducted field investigations, leading to evidence of 15 mortalities at a mean age of 30.6 ± 15.5 (SE) days (range = 3–243 days). We launched 21 investigations in response to a mortality movement and they resulted in confirmation of 11 of the 15 calf mortalities. Specific causes of mortality included 9 wolf (Canis lupus)-kills, 3 black bear (Ursus americanus)-kills, 1 unknown predator-kill, and 2 deaths following vehicle collisions. The mean distance females fled after a mortality was 1,873 ± 412 m (range = 126–5,805 m, n = 14). Females that made return visits returned a mean 2.8 ± 0.5 times (range = 1–5, n = 8) to within a mean 106 ± 22 m (range = 34–230 m, n = 8) of the mortality site. Calf survival to 30 days of age was 67 ± 8% (95% CI = 53–84%, n = 36) but declined to 53 ± 8% (95% CI = 39–72%, n = 36) by 3 months of age. We developed 2 population-level movement models to improve the efficacy of using the mortality movement to identify and locate calf mortalities in real time via field investigations. The first approach, a temporal-based model, used a 3-day average movement velocity threshold (118 m/hr) for all females to indicate calf mortality and accurately predicted survival status in 51% (n = 105) of the cases. The second approach, an age-specific model using different thresholds (28–135 m/hr) for females relative to calf age, was 80% (n = 231) accurate. Using movement behavior of females to assess calf mortality yielded important insights into mechanisms influencing the population decline that will inform future management decisions. © 2019 The Wildlife Society     

Sex ratio variation in harvested moose (<Emphasis Type="Italic">Alces alces</Emphasis>) calves: does it reflect population calf sex ratio or selective hunting?

During the last 30 years, the proportion of males in the calf harvest of moose (Alces alces) in Norway has decreased, indicating a decline in proportions of males recruited to the autumn populations. At the same time, the percentages of exclusive calf hunting permits and of calves shot have increased. The change in calf sex ratio may thus simply be the result of hunter preferences for slightly larger (6.2% higher body mass) male calves combined with fewer opportunities for selective hunting due to increasing hunting quotas of calves. We examined this hypothesis by analyzing the variation in sex, number of siblings, carcass mass, date, and location of kill of 16,330 moose calves harvested during 1970–2004. In the presence of hunting selection for larger calves, we predicted larger proportions of male calves to be harvested in populations with large sexual size dimorphism among calves. Similarly, we expected more males to be harvested from twin than single litters because hunters then can more easily compare twins and select the larger calf, which is more often a male. Increasing proportions of single female calves were also expected to occur in the daily harvest as the accumulated number of harvested calves increased and the proportion of calves left in the population decreased. We found no positive relationship between the proportion of male calves and the level of sexual size dimorphism, no clear difference in sex ratio between harvested single and twin calves, and no increase in the proportion of single female calves as the accumulated number of calves in the harvest increased. This suggests that the spatiotemporal variation in the harvest calf sex ratio in Norway most likely reflects differences in population calf sex ratios prior to the hunting season and not varying degrees of hunting selectivity.     

Patterns of hunting mortality in Norwegian moose (Alces alces) populations

We used a simple life table approach to examine the age-specific patterns of harvest mortality in eight Norwegian moose populations during the last 15 years and tried to determine if the observed patterns were caused by hunter selectivity. The general opinion among local managers is that hunters prefer to shoot female moose not in company with calves to keep a high number of reproductive females in the population (and because of the emotional stress involved in leaving the calf/calves without a mother), and relatively large males because of the higher return with respect to meat and trophy. In support of the former view, we found the harvest mortality of adult females to be higher among pre-prime (1–3 years old) than prime-aged age classes (4–7 years old). This is probably because prime-aged females are more fecund and, therefore, more likely to be in company with one or two calves during the hunting season. As the season progressed, however, the selection pressure on barren females decreased, probably due to more productive females becoming ‘legal’ prey as their calf/calves were harvested. In males, we did not find any evidence of strong age-specific hunter selectivity, despite strong age-dependent variation in body mass and antler size. We suggest that this was due to the current strongly female-biased sex ratio in most Norwegian moose populations, which leaves the hunters with few opportunities to be selective within a relatively short and intensive hunting season. The management implications of these findings and to what extent the results are likely to affect the future evolution of life histories in Norwegian moose populations are discussed.Electronic Supplementary Material Supplementary material is available for this article at and accessible for authorised users.     

Moose (Alces alces) calf survival rates in the presence of wolves (Canis lupus) in southeast Norway

Wolves (Canis lupus) are recolonizing Scandinavia and becoming a new limiting factor that should be taken into account in the management of moose (Alces alces). However, there is a lack of empirical estimates of moose survival after wolf recolonization. We investigated the effects of wolf abundance, moose litter size (single/twin calves), and climatic factors on annual and seasonal calf survival rates in a moose population in southeast Norway. We used data that were obtained over 7?years by radio-tracking and regular visual observations of 68 moose cows to determine the presence or absence of calves at heel. Annual and winter calf survival rates were 20–40 percentage points lower in the wolf territory compared with previous estimates of moose calf survival in similar areas that lacked wolves. Cause-specific studies of mortality would further enhance our ability to determine the relative role of various limiting factors. Our study suggests that moose managers should regulate quotas to buffer the lower survival rates after wolf recolonization.     

Divergent estimates of herd‐wide caribou calf survival: Ecological factors and methodological biases

Population monitoring is a critical part of effective wildlife management, but methods are prone to biases that can hinder our ability to accurately track changes in populations through time. Calf survival plays an important role in ungulate population dynamics and can be monitored using telemetry and herd composition surveys. These methods, however, are susceptible to unrepresentative sampling and violations of the assumption of equal detectability, respectively. Here, we capitalized on 55 herd‐wide estimates of woodland caribou (Rangifer tarandus caribou) calf survival in Newfoundland, Canada, using telemetry (n = 1,175 calves) and 249 herd‐wide estimates of calf:cow ratios (C:C) using herd composition surveys to investigate these potential biases. These data included 17 herd‐wide estimates replicated from both methods concurrently (n = 448 calves and n = 17 surveys) which we used to understand which processes and sampling biases contributed to disagreement between estimates of herd‐wide calf survival. We used Cox proportional hazards models to determine whether estimates of calf mortality risk were biased by the date a calf was collared. We also used linear mixed‐effects models to determine whether estimates of C:C ratios were biased by survey date and herd size. We found that calves collared later in the calving season had a higher mortality risk and that C:C tended to be higher for surveys conducted later in the autumn. When we used these relationships to modify estimates of herd‐wide calf survival derived from telemetry and herd composition surveys concurrently, we found that formerly disparate estimates of woodland caribou calf survival now overlapped (within a 95% confidence interval) in a majority of cases. Our case study highlights the potential of under‐appreciated biases to impact our understanding of population dynamics and suggests ways that managers can limit the influence of these biases in the two widely applied methods for estimating herd‐wide survival.     

Effects of Winter Ticks and Internal Parasites on Moose Survival in Vermont,USA

Moose (Alces alces) have experienced considerable declines along the periphery of their range in the northeastern United States. In Vermont, the population declined 45% from 2010 to 2017 despite minimal hunter harvest and adequate habitat. Similarly, nearby populations recently experienced epizootics characterized by >50% mortality. Declines have largely been associated with the effects of winter ticks (Dermacentor albipictus), but uncertainty exists about the effects of environmental and other parasite-related conditions on moose survival. We examined patterns of moose survival among a radio-collared population (n = 127) in Vermont from 2017 to 2019. Our objectives were to estimate causes of mortality and model survival probability as a function of individual and landscape variables for calves (<1 yr) and adults (≥1 yr). Observed adult survival was 90% in 2017, 84% in 2018, and 86% in 2019, and winter calf survival was 60% in 2017, 50% in 2018, and 37% in 2019. Winter tick infestation was the primary cause of mortality (91% of calves, 25% of adults), and 32% of all mortalities had evidence of meningeal worm (Parelaphostrongylus tenuis). Other sources of mortality such as vehicles, harvest, predation, deep snow, and other parasitic infections were negligible. The best supported calf model included sex differences and negative effects of tick engorgement (%/week) and parasite level (roundworm and lungworm). The best supported adult model included the effect of cumulative tick engorgement (cumulative %/week), which negatively affected survival. Our results indicate that winter tick engorgement strongly affects survival, and is probably compounded by the presence of meningeal worm and other parasites. Reduced tick effects may be achieved by decreasing moose density through harvest and managing late winter habitat to minimize tick density. Management of white-tailed deer (Odocoileus virginianus) density may also affect the transmission of meningeal worm. © 2021 The Authors. The Journal of Wildlife Management published by Wiley Periodicals LLC on behalf of The Wildlife Society.     

Pintail and Mallard Survival in California Relative to Habitat,Abundance, and Hunting

ABSTRACT The influence of habitat, waterfowl abundance, and hunting on winter survival of waterfowl is not well understood. We studied late August-March survival of 163 after-hatch-year (AHY) and 128 hatch-year (HY) female mallards (Anas platyrhynchos) radiotagged in Sacramento Valley (SACV) and 885 AHY female northern pintails (A. acuta) radiotagged throughout the Central Valley of California, USA, relative to flooded habitat (HAB), January abundance of each species (JMAL or JPIN), hunter-days (HDY), and a hunting pressure index (HPI) that combined these variables. From EARLY (1987–1994) to LATE (1998–2000), HAB increased 39%, JPIN increased 45%, JMAL increased 53%, HDY increased 21%, duck-hunting season increased from 59 days to 100 days, and the female daily bag limit doubled to 2 for mallards but remained 1 for pintails. Survival (± SE) was greater during LATE versus EARLY for pintails radiotagged in each region (SACV: 93.2 ± 2.1% vs. 87.6 ± 3.0%; Suisun Marsh: 86.6 ± 3.2% vs. 77.0 ± 3.7%; San Joaquin Valley: 86.6 ± 3.1% vs. 76.9 ± 4.1%) but not for SACV mallards (AHY: 70.6 ± 7.2% to 74.4 ± 7.7% vs. 80.1 ± 7.2% to 82.8 ± 5.6%; HY: 48.7 ± 9.1% [1999–2000 only] vs. 63.5 ± 8.8% to 67.6 ± 8.0%). Most pintail (72%) and mallard (91%) deaths were from hunting, and lower HPI and higher JPIN or JMAL were associated with reduced mortality. Increased HAB was associated with reduced winter mortality for pintails but not for SACV mallards. Pintail survival rates that we measured were within the range reported for other North American wintering areas, and during LATE were higher than most, even though our study duration was 68–110 days longer. Winter survival rates of SACV mallards were also within the reported range. However, with higher bag limits and longer seasons, mallard survival during LATE was lower than in most other wintering areas, especially during 1999–2000, when high winds on opening weekend resulted in high hunting mortality. Habitat conservation and favorable agriculture practices helped create a Central Valley wintering environment where natural mortality of mallards and pintails was low and survival varied with hunting mortality. We recommend regulations and habitat management that continue to minimize natural mortality while allowing sustainable harvest at a level that helps maintain strong incentive for management of Central Valley waterfowl habitats, including the large portion that is privately owned.     

Lying in wait: Limiting factors on a low-density ungulate population and the latent traits that can facilitate escape from them

Predation, habitat, hunting, and environmental conditions have all been implicated as regulatory mechanisms in ungulate populations. The low-density equilibrium hypothesis predicts that in low-density populations, predators regulate their prey and that the population will not escape unless predation pressure is eased. We evaluated survival of adult and juvenile moose (Alces alces) in north-central Alaska to determine whether or not the population supported the hypothesis. We instrumented adult male and female moose with radiocollars and used aerial observations to track parturition and subsequent survival of juvenile moose. Generalized linear mixed-effects models were used to assess survival. Adult annual survival rates were high (∼89%), but may be negatively influenced by winter conditions. Migratory status did not affect moose survivorship or productivity. Approximately 60% of the calf crop died before 5 months of age. Productivity was significantly lower in the northern section of the study area where there is less high-quality habitat, suggesting that, even in this low-density population, nutrition could be a limiting factor. It appears that predation on young calves, winter weather, and nutritional constraints may be interacting to limit this population. Latent traits, such as overproduction of calves and migratory behavior, which do not currently enhance fitness, may persist within this population so that individuals with these traits can reap benefits when environmental conditions change.     

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.     

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.     

Wolf-avoidance strategies of moose

Two possible anti-predator strategies of prey were suggested by the distribution of moose at Isle Royale, a 544 km² island with numerous offshore islets. Aerial surveys in winter indicated that moose density on small islets (<1.5 km²) averaged 415% greater than on the main island, with about 11% of the calf population existing on the small islets which comprise less than 2% of the total land area. On small islets and shorelines, protection from wolves (in the form of escape terrain) is provided by nearby water in summer and coniferous cover in winter. Calves on islands appear to have a higher survival rate. In summer, cow moose with calves were seen more commonly in camps with human activity (and no wolves) than away from camps, suggesting that people provided a refuge from wolf predation for this unhunted moose population. These subtle patterns of habitat selection by moose are attributed to the intensive natural selection pressure of wolf predation.     

Predation on Moose Calves by European Brown Bears

Abstract: In North America, brown bears (Ursus arctos) can be a significant predator on moose (Alces alces) calves. Our study in Sweden is the first in which brown bears are the only predator on moose calves. Bears and moose occurred at densities of about 30/1,000 km² and 920/1,000 km², respectively, and bears killed about 26% of the calves. Ninety-two percent of the predation took place when calves were <1 month old. Bear predation was probably additive to other natural mortality, which was about 10% in areas both with and without bears. Females that lost their calves in spring produced more calves the following year (1.54 calves/F) than females that kept their calves (1.11 calves/F), which reduced the net loss of calves due to predation to about 22%.     

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     

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.     

Disentangling Natural From Hunting Mortality in an Intensively Hunted Wild Boar Population

Abstract We assessed age-specific natural mortality (i.e., excluding hunting mortality) and hunting mortality of 1,175 male and 1,076 female wild boar (Sus scrofa) from Chǎteauvillain-Arc en Barrois (eastern France), using a 22-year dataset (1982–2004) and mark-recapture-recovery methods. Overall yearly mortality was >50% for all sex and age-classes. Low survival was mostly due to high hunting mortality; a wild boar had a >40% of chance of being harvested annually, and this risk was as high as 70% for adult males. Natural mortality rates of wild boar were similar for males and females (approx. 0.15). These rates were comparable to rates typical of male ungulates but high for female ungulates. Wild boar survival did not vary across sex and age-classes. Despite high hunting mortality, we did not detect evidence of compensatory mortality. Whereas natural mortality for males was constant over time, female mortality varied annually, independent of fluctuations in mast availability. Female wild boar survival patterns differed from those reported in other ungulates, with high and variable natural mortality. In other ungulates, natural mortality is typically low and stable across a wide range of environmental conditions. These differences may partly reflect high litter sizes for wild boar, which carries high energetic costs. High hunting mortality may induce a high investment of females in reproduction early in life, at the detriment to survival. Despite high hunting mortality, the study population increased. Effective population control of wild boar should target a high harvest rate of piglets and reproductive females.     

Survival and cause-specific mortality of female Rocky Mountain elk exposed to human activity

Animal populations are becoming increasingly exposed to human activity as human populations expand and demand for energy resources (e.g., coal, oil and natural gas) increases. We initiated this study to document survival and cause-specific mortality patterns of female Rocky Mountain elk (Cervus elaphus) exposed to increasing levels of human activity. We fitted 184 females with VHF or GPS collars over 4 years and used the Kaplan–Meier survival estimator to calculate annual survival rates. We used multinomial logistic regression to assess differences in cause-specific mortality and generalized linear mixed models to determine how probability of survival was structured during hunting season; both analyses examined a suite of 5 covariates (i.e., age, year, extent of space use, cover, and human footprint) as potentially influencing cause-specific mortality and survival probability. Annual probability of survival averaged 0.8 (±0.02 SE) over 4 years but averaged 0.91 (±0.03 SE) when harvest mortality was excluded, which was the most significant source of mortality in most years ( [`(x)] = 0.13 ±0.02 \textSE \bar{x} = 0.13 \pm 0.02\,{\text{SE}} ). We found no difference between cause-specific mortality sources relative to elk that survived during the hunting season (χ ₁₀² = 5.79, P = 0.832). The probability of a female surviving during hunting season was negatively influenced by age, year, extent of space use, cover, and human footprint. We found evidence that human activity may have influenced annual rates of natural survival (i.e., exclusive of hunting mortality) and probability of survival during the hunting season. We note that this study occurred largely on privately owned and managed residential and ranch land and focused on female elk; we acknowledge that survival rate and cause-specific patterns of mortality may vary as a function of land ownership (private vs. public), demographic status, and management and harvest practices. While temporal and spatial scales of 1 week may be sufficient to describe patterns of direct mortality during hunting season, broad temporal or spatial scale analyses may be needed to address natural mortality during other seasons.     

Evaluating how hunters see and react to telemetry collars on white-tailed deer

Fates of individuals outfitted with radiotransmitters commonly are used for estimating survival rates in populations of large animals that are hunted. Despite precautions, this practice may be subject to complex biases associated with hunter reaction to presence of radiotransmitters. To assess this potential bias we conducted an experiment using artificial deer (i.e., decoys) to measure hunters' abilities to see deer and determine if deer seen were wearing radiocollars. We used logistic regression to quantify probabilities that seeing deer and subsequently seeing radiocollars might be influenced by distance, percent visual obstruction, body orientation, hunter experience, and antler characteristics of deer. Additionally, we evaluated how experience and antler characteristics of deer might influence a hunter's decision to harvest a radiocollared deer. We found that 25.8% of the potentially observable collared deer (n = 663) were subsequently observed by hunters. Odds of observing deer and radiocollars increased 95% and 230%, respectively, for each additional log(yr) of hunting experience. Willingness to harvest radiocollared deer increased 89% for each additional log(yr) of hunting experience and 144% for large-antlered deer relative to antlerless deer. When hunting is an important source of mortality, analysts need to understand how potential biases associated with observing deer are associated with hunters' reactions to and subsequent decisions to harvest radiocollared animals. Our study suggested that presence of radiocollars may influence a deer's potential risk of being harvested and in turn bias telemetry-based estimates of survival, given that hunting mortality is the largest component of total mortality in hunted deer populations. Collar-based telemetry is used nearly universally by wildlife managers and researchers throughout North America and elsewhere to estimate and monitor the survival of big game populations that are managed through hunting. Our findings demonstrate that these estimates are likely subject to complex and systematic biases that managers should consider when evaluating future population-level effects of managed hunting. © 2011 The Wildlife Society