No long-term effect of black bear removal on elk calf recruitment in the southern Appalachians

In 2001 and 2002, 52 elk (Cervus canadensis; 21 males, 31 females), originally obtained from Elk Island National Park, Alberta, Canada, were transported and released into Cataloochee Valley in the northeastern portion of Great Smoky Mountains National Park (GRSM, Park), North Carolina, USA. The annual population growth rate (λ) was negative (0.996, 95% CI = 0.945–1.047) and predation by black bears (Ursus americanus) on elk calves was identified as an important determinant of population growth. From 2006 to 2008, 49 bears from the primary elk calving area (i.e., Cataloochee Valley) were trapped and translocated about 70 km to the southwestern portion of the Park just prior to elk calving. Per capita recruitment (i.e., the number of calves produced per adult female that survive to 1 year of age) increased from 0.306 prior to bear translocation (2001–2005) to 0.544 during years when bears were translocated (2006–2008) and λ increased to 1.118 (95% CI = 1.096–1.140). Our objective was to determine whether per capita calf recruitment rates after bear removal (2009–2019) at Cataloochee were similar to the higher rates estimated during bear removal (i.e., long-term response) or if they returned to rates before bear removal (i.e., short-term response), and how those rates compared with recruitment from portions of our study area where bears were not relocated. We documented 419 potential elk calving events and monitored 129 yearling and adult elk from 2001 to 2019. Known-fate models based on radio-telemetry and observational data supported calf recruitment returning to pre-2006 levels at Cataloochee (short-term response); recruitment of Cataloochee elk before and after bear relocation was lower (0.184) than during bear relocation (0.492). Recruitment rates of elk outside the removal area during the bear relocation period (0.478) were similar to before and after rates (0.420). In the Cataloochee Valley, cause-specific annual calf mortality rates due to predation by bears were 0.319 before, 0.120 during, and 0.306 after bear relocation. In contrast, the cause-specific annual mortality rate of calves in areas where bears were not relocated was 0.033 after the bear relocation period, with no bear predation on calves before or during bear relocation. The mean annual population growth rate for all monitored elk was 1.062 (95% CI = 0.979–1.140) after bear relocation based on the recruitment and survival data. Even though the effects of bear removal were temporary, the relocations were effective in achieving a short-term increase in elk recruitment, which was important for the reintroduction program given that the elk population was small and vulnerable to extirpation.     

Demographics of an Experimentally Released Population of Elk in Great Smoky Mountains National Park

ABSTRACT We assessed the potential for reestablishing elk (Cervus elaphus) in Great Smoky Mountains National Park (GSMNP), USA, by estimating vital rates of experimentally released animals from 2001 to 2006. Annual survival rates for calves ranged from 0.333 to 1.0 and averaged 0.592. Annual survival for subadult and adult elk (i.e., ≥ 1 yr of age) ranged from 0.690 to 0.933, depending on age and sex. We used those and other vital rates to model projected population growth and viability using a stochastic individual-based model. The annual growth rate (λ) of the modeled population over a 25-year period averaged 0.996 and declined from 1.059 the first year to 0.990 at year 25. The modeled population failed to attain a positive 25-year mean growth rate in 46.0% of the projections. Poor calf recruitment was an important determinant of low population growth. Predation by black bears (Ursus americanus) was the dominant calf mortality factor. Most of the variance of growth projections was due to demographic variation resulting from the small population size (n = 61). Management actions such as predator control may help increase calf recruitment, but our projections suggest that the GSMNP elk population may be at risk for some time because of high demographic variation.     

Population viability analysis to identify management priorities for reintroduced Elk in the Cumberland Mountains,Tennessee

We used an individual-based population model to perform a viability analysis to simulate population growth (λ) of 167 elk (Cervus elaphus manitobensis; 71 male and 96 female) released in the Cumberland Mountains, Tennessee, to estimate sustainability (i.e., λ > 1.0) and identify the most appropriate options for managing elk restoration. We transported elk from Elk Island National Park, Alberta, Canada, and from Land Between the Lakes, Kentucky, and reintroduced them beginning in December 2000 and ending in February 2003. We estimated annual survival rates for 156 radio-collared elk from December 2000 until November 2004. We used data from a nearby elk herd in Great Smoky Mountains National Park to simulate pessimistic and optimistic recruitment and performed population viability analyses to evaluate sustainability over a 25-year period. Annual survival averaged 0.799 (Total SE = 0.023). The primary identifiable sources of mortality were poaching, disease from meningeal worm (Parelaphostrongylus tenuis), and accidents (environmental causes and unintentional harvest). Population growth given pessimistic recruitment rates averaged 0.895 over 25 years (0.955 in year 1 to 0.880 in year 25); population growth was not sustainable in 100% of the runs. With the most optimistic estimates of recruitment, mean λ increased to 0.967 (1.038 in year 1 to 0.956 in year 25) with 99.6% of the runs failing to be sustainable. We suggest that further translocation efforts to increase herd size will be ineffective unless survival rates are increased in the Cumberland Mountains. © 2011 The Wildlife Society.     

Summer elk calf survival in a partially migratory population

Decomposing variation in juvenile recruitment is a key component of understanding population dynamics for partially migratory ungulates. We investigated reproductive parameters of adult female elk (Cervus canadensis) with calves at heel, and survivorship, cause-specific mortality, and intrinsic and extrinsic factors affecting risk of mortality for calves in a partially migratory elk population from 2013–2016 in Alberta, Canada. Elk calves born to resident mothers had 45% lower survivorship on average compared to migrant calves (0.24 vs. 0.69) and nearly twice the mortality rate (0.37 vs. 0.19) from bears (Ursus spp.), the dominant source of mortality. Contrary to our predictions, we found that increasing levels of maternal ingesta-free body fat were associated with increasing risk of calf mortality, indicating predation may have overwhelmed nutritional effects. We found no evidence that timing of calf birth or birth weight differed between migratory tactics or influenced mortality risk. We found that as percentage of cut forest increased, risk of calf mortality marginally decreased, which benefited migrant elk that were exposed to more clear-cuts compared to residents. Exposure to bear predation risk was unimportant during the hiding phase (≤10 days after birth) for either migratory tactic, presumably because neonatal hiding behavior reduced vulnerability. In contrast, bear predation risk was important for mortality risk after 10 days in age, especially for resident elk calves, which were exposed to higher bear predation risk compared to migrants. We conclude that relative differences in bear predation between migratory tactics are contributing to the dynamics of partial migration in this population through additive effects on calf mortality. Thus, wildlife managers should anticipate that recovering grizzly bear (U. arctos) populations may substantially lower elk recruitment through effects on summer calf survival, especially in areas with diverse carnivore assemblages.     

Implications of chronic wasting disease,cougar predation,and reduced recruitment for elk management

Emerging diseases and expanding carnivore populations may have profound implications for ungulate harvest management and population regulation. To better understand effects of chronic wasting disease (CWD) and cougar (Puma concolor) predation, we studied mortality and recruitment of elk (Cervus elaphus) at Wind Cave National Park (WICA) during 2005–2009. We marked 202 elk (83 subadult M and 119 subadult and ad F) with Global Positioning System (GPS) collars, observed 28 deaths during 74,220 days of monitoring, and investigated 42 additional deaths of unmarked elk found dead. Survival rates were similar for males and females and averaged 0.863 (SE = 0.025) annually. Leading causes of mortality included hunting (0.065, SE = 0.019), CWD (0.034, SE = 0.012), and cougar predation (0.029, SE = 0.012). Marked elk killed by hunters and cougars typically were in good physical condition and not infected with CWD. Effects of mortality on population growth were exacerbated by low rates of pregnancy (subadults = 9.5%, SE = 6.6%; ad = 76.9%, SE = 4.2%) and perinatal survival (0.49, SE = 0.085 from 1 Feb to 1 Sep). Chronic wasting disease, increased predation, and reduced recruitment reduced the rate of increase for elk at WICA to approximately λ = 1.00 (SE = 0.027) during the past decade. Lower rates of increase are mitigating effects of elk on park vegetation, other wildlife, and neighboring lands and will facilitate population control, but may reduce opportunities for elk hunting outside the park. © 2011 The Wildlife Society     

Dynamics of Newly Established Elk Populations

Abstract: The dynamics of newly established elk (Cervus elaphus) populations can provide insights about maximum sustainable rates of reproduction, survival, and increase. However, data used to estimate rates of increase typically have been limited to counts and rarely have included complementary estimates of vital rates. Complexities of population dynamics cannot be understood without considering population processes as well as population states. We estimated pregnancy rates, survival rates, age ratios, and sex ratios for reintroduced elk at Theodore Roosevelt National Park, North Dakota, USA; combined vital rates in a population projection model; and compared model projections with observed elk numbers and population ratios. Pregnancy rates in January (early in the second trimester of pregnancy) averaged 54.1% (SE = 5.4%) for subadults and 91.0% (SE = 1.7%) for adults, and 91.6% of pregnancies resulted in recruitment at 8 months. Annual survival rates of adult females averaged 0.96 (95% CI = 0.94-0.98) with hunting included and 0.99 (95% CI = 0.97-0.99) with hunting excluded from calculations. Our fitted model explained 99.8% of past variation in population estimates and represents a useful new tool for short-term management planning. Although we found no evidence of temporal variation in vital rates, variation in population composition caused substantial variation in projected rates of increase (Λ = 1.20-1.36). Restoring documented hunter harvests and removals of elk by the National Park Service led to a potential rate of Λ = 1.26. Greater rates of increase substantiated elsewhere were within the expected range of chance variation, given our model and estimates of vital rates. Rates of increase realized by small elk populations are too variable to support inferences about habitat quality or density dependence.     

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.     

Estimating survival in the Apennine brown bear accounting for uncertainty in age classification

For most rare and elusive species, estimating age-specific survival is a challenging task, although it is an important requirement to understand the drivers of population dynamics, and to inform conservation actions. Apennine brown bears Ursus arctos marsicanus are a small, isolated population under a severe risk of extinction, for which the main demographic mechanisms underlying population dynamics are still unknown, and population trends have not been formally assessed. We present a 12-year analysis of their survival rates using non-invasive genetic sampling data collected through four different sampling techniques. By using multi-event capture–recapture models, we estimated survival probabilities for two broadly defined age classes (cubs and older individuals), even though the age of the majority of sampled bears was unknown. We also applied the Pradel model to provide a preliminary assessment of population trend during the study period. Survival was different between cubs [ϕ = 0.51, 95% CI (0.22, 0.79)], adult males [ϕ = 0.85, 95% CI (0.76, 0.91)] and adult females [ϕ = 0.92, 95% CI (0.87, 0.95)], no temporal variation in survival emerged, suggesting that bear survival remained substantially stable throughout the study period. The Pradel analysis of population trend yielded an estimate of λ = 1.009 [SE = 0.018; 95% CI (0.974, 1.046)]. Our results indicate that, despite the status of full legal protection, the basically stable demography of this relict population is compatible with the observed lack of range expansion, and that a relatively high cub mortality could be among the main factors depressing recruitment and hence population growth.     

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.     

Grizzly bear predation links the loss of native trout to the demography of migratory elk in Yellowstone

The loss of aquatic subsidies such as spawning salmonids is known to threaten a number of terrestrial predators, but the effects on alternative prey species are poorly understood. At the heart of the Greater Yellowstone ecosystem, an invasion of lake trout has driven a dramatic decline of native cutthroat trout that migrate up the shallow tributaries of Yellowstone Lake to spawn each spring. We explore whether this decline has amplified the effect of a generalist consumer, the grizzly bear, on populations of migratory elk that summer inside Yellowstone National Park (YNP). Recent studies of bear diets and elk populations indicate that the decline in cutthroat trout has contributed to increased predation by grizzly bears on the calves of migratory elk. Additionally, a demographic model that incorporates the increase in predation suggests that the magnitude of this diet shift has been sufficient to reduce elk calf recruitment (4–16%) and population growth (2–11%). The disruption of this aquatic–terrestrial linkage could permanently alter native species interactions in YNP. Although many recent ecological changes in YNP have been attributed to the recovery of large carnivores—particularly wolves—our work highlights a growing role of human impacts on the foraging behaviour of grizzly bears.     

Trumpeter Swan Abundance and Growth Rates in Yellowstone National Park

ABSTRACT Decreasing abundance of resident, nonmigratory trumpeter swans (Cygnus buccinator) in Yellowstone National Park (YNP), USA, raised concern that this population, which helped facilitate the restoration of the species across North America, may disappear. We quantified trends in abundance of resident and migratory trumpeter swans in YNP from 1967 to 2007 and investigated the potential mechanisms for declining population trends, including cessation of the supplemental feeding program and relocation programs outside of YNP, density dependence, and annual variations in environmental conditions. Estimated abundance of resident trumpeter swans in YNP ranged from 59 individuals in 1968 to 10 individuals in 2007. Using log-linear modeling, the best approximating model chosen from an a priori set of competing models estimated the annual growth rate (r) of resident swans from 1967 to 2007 was −0.036 (95% CI =−0.042 to −0.030, Akaike wt [w_i] = 0.44). A competing model provided evidence that decreases in abundance became more dramatic after supplemental feeding of grain outside of YNP was terminated in winter 1992–1993 (r̂_1967–1992 = −0.027, 95% CI = −0.039 to −0.015; r̂_1993–2007 = −0.053, 95% CI = −0.029 to −0.080; w_i = 0.42). There was little evidence of density-dependent effects on the resident population growth rates (βYNP_pop = 0.006, 95% CI = −0.017 to 0.007), but rates were lower following severe winters, wetter springs, and warmer summers. Our results indicate that the YNP population of trumpeter swans is decreasing and may act as a sink to surrounding populations. Thus, population levels of YNP trumpeter swans may depend on management outside the Park and we recommend the National Park Service collaborate with surrounding agencies in managing trumpeter swans throughout the Tri-state region where more productive habitats may exist.     

Estimating grizzly and black bear population abundance and trend in Banff National Park using noninvasive genetic sampling

We evaluated the potential of two noninvasive genetic sampling methods, hair traps and bear rub surveys, to estimate population abundance and trend of grizzly (Ursus arctos) and black bear (U. americanus) populations in Banff National Park, Alberta, Canada. Using Huggins closed population mark-recapture models, we obtained the first precise abundance estimates for grizzly bears (N=?73.5, 95% CI?=?64-94 in 2006; N=?50.4, 95% CI?=?49-59 in 2008) and black bears (N=?62.6, 95% CI?=?51-89 in 2006; N=?81.8, 95% CI?=?72-102 in 2008) in the Bow Valley. Hair traps had high detection rates for female grizzlies, and male and female black bears, but extremely low detection rates for male grizzlies. Conversely, bear rubs had high detection rates for male and female grizzlies, but low rates for black bears. We estimated realized population growth rates, lambda, for grizzly bear males (λ=?0.93, 95% CI?=?0.74-1.17) and females (λ=?0.90, 95% CI?=?0.67-1.20) using Pradel open population models with three years of bear rub data. Lambda estimates are supported by abundance estimates from combined hair trap/bear rub closed population models and are consistent with a system that is likely driven by high levels of human-caused mortality. Our results suggest that bear rub surveys would provide an efficient and powerful means to inventory and monitor grizzly bear populations in the Central Canadian Rocky Mountains.     

Grizzly Bear Density in Glacier National Park,Montana

Abstract: We present the first rigorous estimate of grizzly bear (Ursus arctos) population density and distribution in and around Glacier National Park (GNP), Montana, USA. We used genetic analysis to identify individual bears from hair samples collected via 2 concurrent sampling methods: 1) systematically distributed, baited, barbed-wire hair traps and 2) unbaited bear rub trees found along trails. We used Huggins closed mixture models in Program MARK to estimate total population size and developed a method to account for heterogeneity caused by unequal access to rub trees. We corrected our estimate for lack of geographic closure using a new method that utilizes information from radiocollared bears and the distribution of bears captured with DNA sampling. Adjusted for closure, the average number of grizzly bears in our study area was 240.7 (95% CI = 202–303) in 1998 and 240.6 (95% CI = 205–304) in 2000. Average grizzly bear density was 30 bears/1,000 km², with 2.4 times more bears detected per hair trap inside than outside GNP. We provide baseline information important for managing one of the few remaining populations of grizzlies in the contiguous United States.     

Reproduction and Survival of Yellowstone Bison

ABSTRACT The conservation of bison (Bison bison) from near extinction to >4,000 animals in Yellowstone National Park has led to conflict regarding overabundance and potential transmission of brucellosis (Brucella abortus) to cattle. We estimated survival and birth rates from 53 radiocollared adult female bison during 1995–2001, and we used calf:adult (C:A) ratios to estimate reproduction with the combined effects of pregnancy, fetal loss, and neonatal mortality during 1970–1997. Annual survival of adult females was high (0.92; 95% CI = 0.87-0.95) and constant. Birth rates differed by brucellosis status and age. Birth rates were 0.40 calves per female (95% CI = 0.15-0.65) for brucellosis-positive 3 year olds, 0.63 (95% CI = 0.39-0.87) for individuals testing negative, and 0.10 (95% CI = 0.00-0.24) for individuals contracting brucellosis that birth year (sero-converters). Birth rates were 0.64 (95% CI = 0.52-0.76) for brucellosis-positive individuals ≥4 years old, 0.81 (95% CI = 0.73-0.89) for brucellosis-negative individuals, and 0.22 (95% CI = 0.00-0.46) for sero-converters. Spring C:A ratios were negatively correlated with snow pack (β = −0.01 to −0.03, R² = 0.26-0.60, P < 0.05). Growth rate was highly elastic to adult survival (0.51), and juvenile survival (0.36) was 3 times more elastic than fecundity (0.12). Simulations suggested brucellosis eradication via vaccination would result in increased birth rates and a 29% increase in population growth (γ = 1.09), possibly leading to more bison movements outside the park. Our results will help park managers evaluate bison population dynamics and explore consequences of management actions and disease control programs.     

Assessing Population Viability of Black Bears using Spatial Capture-Recapture Models

The Central Georgia Bear Population (CGP) is the least abundant and most isolated of Georgia's 3 American black bear (Ursus americanus) populations. Beginning in 2011, changes to regulations governing harvest of the CGP resulted in an increase in female bear harvest, creating concern that future harvest could be an important influence on population viability. Hence, our objective was to assess viability of the CGP under various levels of female mortality. During 2012–2016, we used barbed-wire hair snares to collect bear hair samples from within the range of the CGP in Georgia, USA. We used microsatellite genotyping to identify individual bears and created robust-design, spatial detection histories for all female bears detected. We fit open population spatial capture-recapture (SCR) models to the detection histories in a Bayesian framework. We used the Widely Applicable Information Criterion (WAIC) to rank models that varied with respect to sources of variation in detection probability, survival, and per capita recruitment, and used the model with the lowest WAIC to forecast dynamics of the CGP 50 years into the future under various levels of female mortality. We assessed the 50-year extinction probability under a continuation of mortality levels documented during 2012–2016, and under incremental increases in female mortality above this baseline. The top model included density-dependent per capita recruitment, annual variation in detection probability, and a trap-level behavioral response. Abundance increased from 106 (95% CI = 86–132) females in 2012 to 136 (95% CI = 113–161) females in 2013 and remained relatively stable thereafter. Annual female survival was 0.75 (95% CI = 0.69–0.82) and did not vary among years. The per capita recruitment rate decreased over time as density increased, and was 0.49 (95% CI = 0.33–0.66) during the first time interval and 0.29 (95% CI = 0.20–0.38) during the final time interval. Annual growth rate () was 1.28 (95% CI = 1.07–1.52) between 2012 and 2013 but decreased throughout the study, ending at 1.04 (95% CI = 0.93–1.17). Forecasts indicated continuation of the female mortality levels experienced from 2012–2016 were sustainable over 50 years, with the estimated extinction risk being <0.001%. Increasing annual harvest by 5 females introduced a negligible increase in the 50-year probability of extinction, but harvesting an additional 10 females/year caused extinction risk to rise to 1.15%. We recommend that harvest regulations are structured such that mortality rates remain at current levels or do not increase by more than an annual average of 5 females above levels observed during our study. Furthermore, we recommend that managers continue to monitor the population so that harvest regulations and population models can be refined over time. © 2020 The Wildlife Society.     

Development of a Sightability Model for Low-Density Elk Populations in Ontario,Canada

ABSTRACT The status of recolonizing elk (Cervus elaphus) populations in Ontario, Canada, is unclear and there is a need for effective population survey methods that can be applied locally. We sought to develop a sightability model that could account for both low densities of elk and dense forest cover in elk-release areas in Ontario. We corrected winter aerial survey counts for sightability based on radiocollared animals known to be within observable distance of the aircraft. The multivariate model with the highest Akaike's Information Criterion corrected for sample size weight (w_i = 0.427) revealed that elk group size, elk activity, dominant tree type, percent canopy cover, and percent conifer cover were significant predictors of elk sightability. The group-size effect indicated that odds of sighting an elk increased by 1.353 (95% CI = 0.874-3.689) for every additional elk. Standing elk were 5.033 (95% CI = 0.936-15.541) times more likely to be observed than were resting elk, and those located in conifer cover were 0.013 (95% CI = 0.001-0.278) times less likely to be sighted than elk in deciduous cover. Furthermore, elk located in >50% canopy cover and >50% conifer cover were 0.041 (95% CI = 0.003-0.619) times and 0.484 (95% CI = 0.024-9.721) times less likely to be sighted than elk in more open habitat, respectively. During model validation, observers detected 79% (113/143) of known elk in any given area, and population and sightability model predictions (±90% CI) overlapped with the population estimate, implying that our predictive model was robust. Unsurprisingly, large groups of elk in open habitat increased model precision, which highlights difficulties of counting Ontario elk in their northern range. We conclude that our model provided increased reliability for estimating elk numbers in Ontario compared to existing methods, and that the estimator may be useful in other areas where elk density is low and sightability is poor due to dense forest cover.     

GRIZZLY BEAR DEMOGRAPHICS IN AND AROUND BANFF NATIONAL PARK AND KANANASKIS COUNTRY,ALBERTA

Abstract: The area in and around Banff National Park (BNP) in southwestern Alberta, Canada, is 1 of the most heavily used and developed areas where grizzly bears (Ursus arctos) still exist. During 1994–2002, we radiomarked and monitored 37 female and 34 male bears in this area to estimate rates of survival, reproduction, and population growth. Annual survival rates of bears other than dependent young averaged 95% for females and 81–85% for males. Although this area was largely unhunted, humans caused 75% of female mortality and 86% of male mortality. Females produced their first surviving litter at 6–12 years of age (x̄ = 8.4 years). Litters averaged 1.84 cubs spaced at 4.4-year intervals. Adult (≥6-years-old) females produced 0.24 female cubs per year and were expected to produce an average of 1.7 female cubs in their lifetime, based on rates of reproduction and survival. Cub survival was 79%, yearling survival was 91%, and survival through independence at 2.5–5.5 years of age was 72%, as no dependent young older than yearlings died. Although this is the slowest-reproducing grizzly bear population yet studied, high rates of survival seem to have enabled positive population growth (Λ = 1.04, 95% CI = 0.99–1.09), based on analyses using Leslie matrices. Current management practices, instituted in the late 1980s, focus on alleviating human-caused bear mortality. If the 1970–1980s style of management had continued, we estimated that an average of 1 more radiomarked female would have been killed each year, reducing female survival to the point that the population would have declined.     

Impact of Spatial and Temporal Variation in Calf Survival on the Growth of Elk Populations

Abstract: The realized impact of a vital rate on population growth (λ) is determined by both the relative influence of the vital rate on λ (elasticity) and its magnitude of variability. We estimated mean survival and reproductive rates in elk (Cervus elaphus) and spatial and temporal variation in these rates from 37 sources located primarily across the Rocky Mountain region and northwestern United States. We removed sampling variance from estimates of process variance both within and across vital-rate data sets using the variance discounting method developed by White (2000). Deterministic elasticities calculated from a population matrix model parameterized with these mean vital rates ranked adult female survival (e_Scow = 0.869) much higher than calf survival (e_Scalf = 0.131). However, process variance in calf survival was >11 times greater than process variance in female survival across data sets and 10 times greater on average within studies. We conducted Life-Stage Simulation Analysis to incorporate both vital-rate elasticity patterns and empirical estimates of variability to identify those vital rates most influential in elk population dynamics. The overwhelming magnitude of variation in calf survival explained 75% of the variation in the population growth rates generated from 1,000 matrix replicates, compared to just 16% of the variation in λ explained by variation in female survival. Variation in calf survival greatly impacts elk population growth and calls into question the utility of classical elasticity analysis alone for guiding elk management. These results also suggest that the majority of interannual variability that wildlife managers document in late-winter and spring elk surveys is attributable to variation in calf survival over the previous year and less influenced by variation in the harvest of females during the preceding autumn. To meet elk population size objectives, managers should consider the inherent variation in calf survival, and its apparent sensitivity to management, in addition to female harvest.     

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     

Grizzly bear population vital rates and trend in the Northern Continental Divide Ecosystem,Montana

We estimated grizzly bear (Ursus arctos) population vital rates and trend for the Northern Continental Divide Ecosystem (NCDE), Montana, between 2004 and 2009 by following radio-collared females and observing their fate and reproductive performance. Our estimates of dependent cub and yearling survival were 0.612 (95% CI = 0.300–0.818) and 0.682 (95% CI = 0.258–0.898). Our estimates of subadult and adult female survival were 0.852 (95% CI = 0.628–0.951) and 0.952 (95% CI = 0.892–0.980). From visual observations, we estimated a mean litter size of 2.00 cubs/litter. Accounting for cub mortality prior to the first observations of litters in spring, our adjusted mean litter size was 2.27 cubs/litter. We estimated the probabilities of females transitioning from one reproductive state to another between years. Using the stable state probability of 0.322 (95% CI = 0.262–0.382) for females with cub litters, our adjusted fecundity estimate (m_x) was 0.367 (95% CI = 0.273–0.461). Using our derived rates, we estimated that the population grew at a mean annual rate of approximately 3% (λ = 1.0306, 95% CI = 0.928–1.102), and 71.5% of 10,000 Monte Carlo simulations produced estimates of λ > 1.0. Our results indicate an increasing population trend of grizzly bears in the NCDE. Coupled with concurrent studies of population size, we estimate that over 1,000 grizzly bears reside in and adjacent to this recovery area. We suggest that monitoring of population trend and other vital rates using radioed females be continued. © 2011 The Wildlife Society.