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

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

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     

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     

Behavioral responses of male elk to hunting risk

Prey respond to predation risk with a range of behavioral tactics that can vary based on space use and hunting mode of the predator. Unlike other predators, human hunters are often more spatially and temporally restricted, which creates a period of short-duration, high-intensity predation risk for prey. Consequently, identifying the roles different hunting modes (i.e., archery and rifle), hunts for targeted and non-targeted species, and landscape features play in altering spatial and temporal responses of prey to predation risk by humans is important for effective management of harvested populations. From 2009 to 2016, we used a large-scale experiment including 50 animal-years of location data from 38 unique male elk (Cervus canadensis) to quantify changes in movement and resource selection in response to hunters during 3 separate 5-day controlled hunts for antlered males (elk archery, deer [Odocoileus spp.] rifle, and elk rifle) at the Starkey Experimental Forest and Range in northeast Oregon, USA. We evaluated competing hypotheses regarding elk responses to varying levels of prey risk posed by the different hunt types. We predicted that the strength of elk behavioral responses would increase with perceived hunter lethality (i.e., weak response to elk archery but similar response to elk and deer rifle hunts) and that prey response would be closely associated with hunter activity within the diel cycle (greater during diurnal than nocturnal hours) and across hunting seasons. Elk responses were strongest during diurnal hours when hunters were active on the landscape and were generally more pronounced during both rifle hunts than during the archery hunt (supporting our perceived lethality hypothesis). Male elk avoided open roads across all periods except during nocturnal hours of the breeding season and alternated between avoidance of areas with high canopy cover during nocturnal hours and selection during diurnal hours. In combination these patterns led to distinct distributional changes of male elk from pre-hunt to hunt periods. Patterns of male elk selection highlight the importance of managing for heterogeneous landscapes to meet a variety of habitat, harvest, hunter satisfaction, and escapement objectives.     

Lifetime Sex-Specific Moose Mortality During an Intentional Population Reduction

Where elevated harvest of ungulates is a priority, managers benefit by understanding how various sources of mortality affect the age and sex structure and trend of ungulate populations. Prior studies reported a long period (1997–2014) of moose (Alces alces gigas) nutritional stress from overabundance in our study area, an intentional 31% reduction in moose numbers using liberal harvests of females (2004–2012), and low bear (Ursus spp.) predation and high moose harvest densities relative to other largely roadless systems with moose, bears, and wolves (Canis lupus). In this paper, we detailed management findings after describing causes and rates of mortality from 226 female and 164 male moose radio-collared at 9 months of age (1997–2008) and followed through life (1997–2019) and throughout the population reduction. We listened for mortality signals on radio-collars 1–2 times/month when snow cover was complete and 2–4 times/month when snow cover was incomplete. Upon hearing a mortality signal, we investigated mortality sites usually within 24 hours via helicopter. Excluding hunter-caused mortality, we estimated 28% annual mortality for male yearlings versus 17% for female yearlings, then low annual mortality rates (0–4%) to 84 months of age for males and 96 months of age for females, and gradually increasing annual mortality rates thereafter. Most (83%) male moose ≥24 months of age died from hunters; minor causes included wolves (8%), malnutrition or disease (5%), grizzly bears (U. arctos; 2%), and accidents (2%). Most female moose ≥24 months of age died from wolves (37%) or hunters (33%); minor causes included malnutrition or disease (15%), grizzly bears (10%), and accidents (5%). The proportion of radio-collared females killed by hunters varied depending on numbers of permits issued to hunters; the kill rate of females ≥24 months of age was 58% during the initial 4 years of the 9-year reduction, moderated at 29% during the final 5 years of the reduction, and was only 7% for all other study years. We attributed 32% of hunter kills to illegal harvest and unrecovered hunter kills. Hunters played a key role in the intentional population reduction by harvesting prime-age and near prime-age male and female moose that rarely died from other sources of mortality compared with calf, yearling, and older moose. Restricting general season hunters to primarily harvesting prime-age and older male moose with antler spreads ≥127 cm did not appreciably reduce harvest of adult males. Male moose 2.0–5.3 years of age rarely died from non-hunter causes and were largely harvested at older, prime ages (5.3–8.3 yr of age). Yearling moose of both sexes died primarily from wolves, with wolves selecting more for males. By using liberal harvests of female moose to reduce the population, managers improved moose nutrition and reproduction, met mandates for elevated harvests, and may have avoided a reoccurrence of a previous precipitous decline in moose numbers that was initiated by overabundance and extreme snow depths. © 2019 The Wildlife Society.     

Influence of Landscape Characteristics on Hunter Space Use and Success

Sport hunting of ungulates is a predominant recreational pursuit and the primary tool for managing their populations in North America and beyond, given its influence on ungulate distributions, social organization, and population performance. Similarly, land management, such as motorized vehicle access, influences ungulate distributions during and outside hunting seasons. Although research on ungulate responses to hunting and land use is widespread, knowledge gaps persist about space use of hunters and what landscape features discriminate among hunt types and between successful and unsuccessful hunters. We used telemetry location data from hunters (n = 341) to estimate space use from 2008–2013 during 3 types of controlled, 5-day hunts for antlered mule deer (Odocoileus hemionus) and elk (Cervus canadensis) in northeastern Oregon, USA: archery elk, rifle deer, and rifle elk. To evaluate space use, we developed utilization distributions for each hunter, created core areas (50% contours) for groups of hunters, and derived several metrics of space-use overlap between successful and unsuccessful hunters. We also modeled predictors of space use using resource utilization functions with beta regression and stepwise model building. Hunter space use was compressed, with even the largest core area (unsuccessful rifle elk hunters) encompassing <16% (1,178 ha) of the area. We found strong similarities in space use of rifle hunters compared to archers, and core areas of successful hunters were markedly smaller than those of unsuccessful hunters (e.g., = 104 ha vs. 681 ha, respectively, for archers). Percentage cover and distance from open roads were the most consistent covariates in the 6 final models (successful vs. unsuccessful for each of 3 hunts) but with different signs. For example, predicted use of archery and rifle elk hunters increased with cover but decreased for rifle deer hunters. Although the same covariates were in the final models for unsuccessful and successful rifle elk hunters, their negligible spatial overlap suggested they sought those features in different locales, a pattern also documented for rifle deer hunters. Our models performed well (Spearman's rank correlation coefficients = 0.99 for 5 of 6 models), reflecting their utility for managing hunters and landscapes. Our results suggest that strategic management of open roads and forest cover can benefit managers seeking to balance hunter opportunity and satisfaction with harvest objectives, especially for species of special concern such as mule deer, and that differences in space use among hunter groups should be accounted for in hunting season designs. © 2021 The Wildlife Society. This article has been contributed to by US Government employees and their work is in the public domain in the USA.     

Ecosystem Scale Declines in Elk Recruitment and Population Growth with Wolf Colonization: A Before-After-Control-Impact Approach

The reintroduction of wolves (Canis lupus) to Yellowstone provided the unusual opportunity for a quasi-experimental test of the effects of wolf predation on their primary prey (elk – Cervus elaphus) in a system where top-down, bottom-up, and abiotic forces on prey population dynamics were closely and consistently monitored before and after reintroduction. Here, we examined data from 33 years for 12 elk population segments spread across southwestern Montana and northwestern Wyoming in a large scale before-after-control-impact analysis of the effects of wolves on elk recruitment and population dynamics. Recruitment, as measured by the midwinter juvenile∶female ratio, was a strong determinant of elk dynamics, and declined by 35% in elk herds colonized by wolves as annual population growth shifted from increasing to decreasing. Negative effects of population density and winter severity on recruitment, long recognized as important for elk dynamics, were detected in uncolonized elk herds and in wolf-colonized elk herds prior to wolf colonization, but not after wolf colonization. Growing season precipitation and harvest had no detectable effect on recruitment in either wolf treatment or colonization period, although harvest rates of juveniles∶females declined by 37% in wolf-colonized herds. Even if it is assumed that mortality due to predation is completely additive, liberal estimates of wolf predation rates on juvenile elk could explain no more than 52% of the total decline in juvenile∶female ratios in wolf-colonized herds, after accounting for the effects of other limiting factors. Collectively, these long-term, large-scale patterns align well with prior studies that have reported substantial decrease in elk numbers immediately after wolf recolonization, relatively weak additive effects of direct wolf predation on elk survival, and decreased reproduction and recruitment with exposure to predation risk from wolves.     

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.     

Population and Harvest Dynamics of Midcontinent Sandhill Cranes

Sandhill cranes (Antigone canadensis) inhabiting the midcontinent of North America have been hunted since the 1960s under management goals of maintaining abundance, retaining geographic distribution, and maximizing sustainable harvest. Some biologists have raised concerns regarding harvest sustainability because sandhill cranes have lower reproductive rates than other game birds. We summarized demographic information in an age-structured matrix model to better understand population dynamics and harvest. Population indices and recovered harvest since the early 1980s suggest midcontinent sandhill cranes have experienced an average long-term annual growth of 0.9%; meanwhile, harvest has increased 1.8% annually. Adult survival and recruitment rates estimated from field data required modest adjustments (1–3%) so that model-derived growth rates matched growth estimated from a long-term survey (0.887 adult survival and 0.199 females/breeding female). Considering 0.9% long-term annual growth, sandhill cranes could be harvested at a rate of 6.6% if harvest was additive to natural mortality (assumed to be 0.05) or 11.3% if harvest mortality compensated for natural mortality. Life-history characteristics for long-lived organisms and demographic evidence suggested that hunter harvest was primarily additive. Differential harvest rates of segments of sandhill cranes in the midcontinent population derived from differential exposure to hunting suggested potentially unsustainable harvest for greater sandhill cranes (A. c. tabida) from 2 breeding segments. Overall, demographic evidence suggests that the harvest of sandhill cranes in the midcontinent population has been managed sustainably. Monitoring activities that reduce nuisance variation and estimate vital and harvest rates by subspecies would support continued management of sandhill cranes that are of interest to hunters and bird watchers. Published 2020. This article is a U.S. Government work and is in the public domain in the USA.     

Survival,cause-specific mortality,and population growth of white-tailed deer in western Virginia

Understanding the role of recruitment in population dynamics of white-tailed deer (Odocoileus virginianus) is important for management. In the central Appalachian Mountains, deer are part of a largely forested ecosystem that supports 3 carnivore species thought to be capable of influencing white-tailed deer recruitment: black bears (Urus americanus), coyotes (Canis latrans), and bobcats (Lynx rufus). Yet little is known about predation, how other environmental factors influence recruitment, or the importance of neonate survival to white-tailed deer population performance in the region. Our objectives were to identify causes of mortality for neonates, analyze effects of landscape attributes on survival of neonates, estimate survival rates for neonates and adult female white-tailed deer, and to model population growth trends based on current vital rates and hypothetical harvest and neonate survival scenarios. During 2019–2020, we captured 57 neonate deer in Bath County, Virginia, USA, by monitoring 38 pregnant females equipped with global positioning system collars and vaginal implant transmitters and by conducting transect searches for recently born neonates. We observed 37 neonate mortalities and identified cause of death using field and genetic evidence. Mortalities included 28 predation events and 9 deaths from other causes (e.g., abandonment, malnutrition, disease). Black bears accounted for 48.6% of neonate mortalities, and 64.2% of predation events (n = 18), followed by bobcats (n = 5) and coyotes (n = 3). Annual survival for adult female deer was 0.871 and neonate survival to 12 weeks old was 0.310. Elevation was a significant predictor of neonate survival; mortality risk increased 20% for every 100-m increase in elevation. Models of annual population growth using observed vital rates predicted an increasing population (λ = 1.10). A 10% increase in female harvest would still result in a potential population increase of 2% (λ = 1.02), but a 20% increase in harvest rate would result in a potential 7% decline (λ = 0.93). Neonate survival was higher near fertile valley bottoms and lower along forested ridges characterized by shallow, infertile soils and limited edge or early successional forests. While predation, largely influenced by black bears, was the leading cause of neonate mortality and contributed to low neonate survival, we observed little evidence of population decline, and suggest there is opportunity for a modest increase in harvest of female deer.     

Effects of hunter access and habitat security on elk habitat selection in landscapes with a public and private land matrix

Traditional elk habitat management on public land has focused on providing security habitat for bull elk during the hunting season to provide for both adequate hunter opportunity and bull survival. This paradigm has given less consideration to adult female elk habitat use, patterns of adjacent land ownership, and hunter access. This paradigm also was developed when elk population sizes were much smaller in many areas. In many Rocky Mountain states, the focus of elk population management has recently shifted to reducing or maintaining elk population sizes, necessitating a better understanding of the implications of security habitat management, as well as patterns of adjacent land ownership and hunter access, on adult female elk. We addressed this need by testing the hypotheses that during the hunting season: 1) adult female elk selection for areas prohibiting or limiting hunter access is stronger than elk selection for publicly owned and managed elk security habitat, 2) these effects occur during the archery hunting period and intensify during the rifle hunting period, and 3) the effects of hunter access on selection are consistent among herds that occupy landscapes characterized by a matrix of public and private lands. We used global position system locations collected from 82 females in 2 different Greater Yellowstone Ecosystem (GYE) elk herds to evaluate effects of hunter access, security habitat as defined by the Hillis paradigm, and other landscape attributes on adult female elk resource selection during the pre-hunting, archery, rifle, and post-hunting periods. We found that female elk selection for areas restricting public hunting access was stronger than selection for security habitat in both study areas, and that the density of roads open to motorized use was the strongest predictor of elk distribution. Increases in selection for areas that restricted hunting access occurred during the rifle hunting period, and we did not find consistent evidence these movements were triggered by the archery hunting period. Our results provide evidence that in landscapes characterized by a matrix of public and privately owned lands, traditional concepts of elk security habitat need to be expanded to also include areas that restrict hunter access to plan for elk population management that is regulated through adult female harvest. Future efforts should investigate whether elk use of areas that restrict hunter access are flexible behavioral responses to hunting risk, or if these behaviors are passed from generation to generation such that a learned pattern of private land use becomes the normal movement pattern rather than a short-term behavioral response. Published 2013. This article is a U.S. Government work and is in the public domain in the USA.     

Effects of early horn growth on reproduction and hunting mortality in female chamois

1.?Environmental conditions during early development can affect the growth patterns of vertebrates, influencing future survival and reproduction. In long-lived mammals, females that experience poor environmental conditions early in life may delay primiparity. In female bovids, annual horn growth increments may provide a record of age-specific reproduction and body growth. Horn length, however, may also be a criterion used by hunters in selecting animals to harvest, possibly leading to artificial selection. 2.?We studied three populations of chamois (Rupicapra rupicapra) in the western Alps to explore the relationships between female horn length and early growth, age of primiparity and age-specific reproduction. We also compared the risk of harvest to reproductive status and horn length. 3.?Early horn growth was positively correlated with body mass in pre-reproductive females and with reproduction in very young and senescent adults. Females with strong early horn growth attained primiparity at an earlier age than those with weak early growth. Horn length did not affect hunter selection, but we found a strong hunter preference for nonlactating females. 4.?Our research highlights the persistent effects of early development on reproductive performance in mammals. Moderate sport harvests are unlikely to affect the evolution of phenotypic traits and reproductive strategies in female chamois. A policy of penalizing hunters that harvest lactating females, however, may increase the harvest of 2-year-old females, which have high reproductive potential.     

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

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

Chukar Seasonal Survival and Probable Causes of Mortality

ABSTRACT Chukars (Alectoris chukar) have been introduced throughout the world. Despite this widespread distribution, limited information regarding seasonal survival, probable causes of mortality, and other basic life history characteristics is available to manage this harvested species. We estimated the probable cause of mortality for chukars with radiotransmitters by examining evidence at kill sites. We used model selection to evaluate influences of seasonal effects (fall raptor migration, peak migration, and reproductive period), demographic effects (age and sex), radio weight, and year on survival of chukars in western Utah, USA, by using a known-fate model in Program MARK and 2 years of telemetry data. We captured and randomly fitted 125 individual chukars with 2 different-sized radiotransmitters (97 F, 20 M, 8 sex undetermined). Model selection results showed our top 3 models accounted for 99% of Akaike's Information Criterion weight, and each one had seasonal and year effects. Two-week survival estimates were lower during peak raptor migration in both years and significantly (P < 0.05) so in year 2 (2-week S = 0.87, 95% CI = 0.77–0.94) compared with other year 2 intervals (2-week S > 0.91). Annual survival was lower in 2005 (S = 0.03, 95% CI = 0.01–0.09) compared with 2006 (S = 0.19, 95% CI = 0.12–0.31). We documented 95 deaths and classified 56% unknown, 33% avian predation, 8% hunter harvest, and 3% mammalian predation. Our research suggests that predation on chukars is substantial during the peak fall raptor migratory period and that the hunting take under current regulations is relatively small and likely compensatory.     

Interaction effects between weather and space use on harvesting effort and patterns in red deer

Most cervid populations in Europe and North America are managed through selective harvesting, often with age‐ and sex‐specific quotas, with a large influence on the population growth rate. Less well understood is how prevailing weather affects harvesting selectivity and off‐take indirectly through changes in individual animal and hunter behavior. The behavior and movement patterns of hunters and their prey are expected to be influenced by weather conditions. Furthermore, habitat characteristics like habitat openness are also known to affect movement patterns and harvesting vulnerability, but how much such processes affect harvest composition has not been quantified. We use harvest data from red deer (Cervus elaphus) to investigate how weather and habitat characteristics affect behavioral decisions of red deer and their hunters throughout the hunting season. More specifically, we look at how sex and age class, temperature, precipitation, moon phase, and day of week affect the probability of being harvested on farmland (open habitat), hunter effort, and the overall harvest numbers. Moon phase and day of week were the strongest predictors of hunter effort and harvest numbers, with higher effort during full moon and weekends, and higher numbers during full moon. In general, the effect of fall weather conditions and habitat characteristics on harvest effort and numbers varied through the season. Yearlings showed the highest variation in the probability of being harvested on farmland through the season, but there was no effect of sex. Our study is among the first to highlight that weather may affect harvesting patterns and off‐take indirectly through animal and hunter behavior, but the interaction effects of weather and space use on hunter behavior are complicated, and seem less important than hunter preference and quotas in determining hunter selection and harvest off‐take. The consideration of hunter behavior is therefore key when forming management rules for sustainable harvesting.     

Survival and cause-specific mortality rates of female sika deer in eastern Hokkaido, Japan

Survival and cause-specific mortality rates of female sika deer (Cervus nippon) were studied using radio telemetry in eastern Hokkaido, Japan. We captured and radio-collared 18 female deer, and monitored their survival from April 1993 to May 1996. Estimated annual survival rate for adult females was 0.779 (95% confidence interval was 0.609–0.997). The harvest mortality rate of adult females was higher than the natural mortality rate. Experimental female hunting during 1994–1996 contributed to an increase in the mortality rate for females and was useful in the control of the sika deer population.     

Influence of habitat features and hunter behavior on white-tailed deer harvest

Sport hunting may help in controlling cervid populations over large areas. As with natural predators, several environmental factors can influence sport harvest. A better understanding of the environmental variables that limit the efficiency of sport hunting could provide guidelines for more efficient wildlife management using hunting. We studied white-tailed deer (Odocoileus virginianus) hunting on a high deer density island where hunting was the sole form of predation. Our objective was to study the behavior of sport hunters and determine the habitat characteristics (e.g., abundance of deer forage, visibility of the deer from the hunter's point of view, and accessibility of the territory to hunters) that are associated with a successful harvest. We collected movements and harvest site location data from 477 hunters equipped with handheld Global Positioning System (GPS) units. Harvest sites were visited and characterized, along with a paired random site, to determine the environmental conditions associated with a successful hunt. We also developed a model to predict the daily number of deer seen by hunters considering weather conditions, hunter characteristics (e.g., age, experience), and date of hunting. We used the mean number of deer seen per hunter per day as a relative index of local density in each hunted territory. At both the site and landscape scales, the combination of visibility and access had a positive effect on the distribution of harvested deer. Habitat types with less visual obstruction from vegetation enabled hunters to see more deer in a given day. At the site scale, harvested deer were located in areas with a lower density of access routes compared to areas where hunters travelled throughout the day. Using an innovative approach of studying hunter behavior with GPS technology, digital maps, and questionnaires, we highlighted the factors associated with hunter success. Our study suggests that habitat characteristics could be modified to increase harvest by improving accessibility and visibility near roads. Creating openings in mature and regenerating forest near access roads could make sport hunting a more efficient management tool, but the potential impact of increased forage availability in forest openings should not be overlooked. © 2012 The Wildlife Society.     

Elk survival and mortality causes in the Blue Mountains of Washington

We studied survival of elk (Cervus elaphus) ≥1 yr old and quantified mortality sources in the Blue Mountains of Washington, 2003–2006, following a period of extensive poaching. The population was managed under a spike-only general hunting season, with limited permits for larger males and for females. We radiomarked 190 elk (82 males and 39 females >1 yr old and 65 males 11 months old), most with rumen transmitters and neck radiocollars; 60 elk only received rumen transmitters. We estimated annual survival using known fate models and explored survival differences among sex and age classes and in 2 potentially different vulnerability zones for males. We found little support for differences in survival between younger (2–3-yr old) and older (≥4-yr old) branch-antlered males or zone differences for yearling males. A model with zone differences for branch-antlered males was the second ranked model and accounted for 14% of the available model weight. From the best-supported models, we estimated annual survival for yearling males at 0.41 (95% CI: 0.29–0.53). We estimated pooled adult female survival at 0.80 (95% CI: 0.64–0.93); when an age-class effect was included, point estimates were higher for prime-aged females (2–11 yr: S = 0.81 [0.70–0.88]) than for older females (≥12 yr: S = 0.72 [0.56–0.83]), but confidence intervals broadly overlapped. Only 1 of 7 models with a female age effect on survival was among the competitive models. For branch-antlered males, survival ranged 0.80–0.85, depending on whether zone variation was modeled. We recorded 78 deaths of radiomarked elk. Human-caused deaths (n = 55) predominated among causes and most were of yearling males killed during state-sanctioned hunts (n = 28). Most subadult male deaths were from tribal hunting (n = 5), and most mature males died from natural causes (n = 6) and tribal hunting (n = 5). We detected few illegal kills (n = 4). Our results suggest that increased enforcement effectively reduced poaching, that unreported tribal harvest was not a trivial source of mortality, and that spike-only general seasons were effective in recruiting branch-antlered males. © 2011 The Wildlife Society.     

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

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

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.