We conducted a 3-year study (May 2003-Apr 2006 of mortality of northern Yellowstone elk (Cervus elaphus) calves to determine the cause for the recruitment decline (i.e., 33 calves to 13 calves/100 adult F) following the restoration of wolves (Canis lupus). We captured, fit with radiotransmitters, and evaluated blood characteristics and disease antibody seroprevalence in 151 calves 6 days old (68M:83F). Concentrations (x, SE) of potential condition indicators were as follows: thyroxine (T4; 13.8 lg/dL, 0.43), serum urea nitrogen (SUN; 17.4 mg/dL, 0.57), c-glutamyltransferase (GGT; 66.4 IU/L, 4.36), gamma globulins (GG; 1.5 g/dL, 0.07), and insulin-like growth factor-1 (IGF-1; 253.6 ng/mL, 9.59). Seroprevalences were as follows: brucellosis (Brucella abortus; 3%), bovine-respiratory syncytial virus (3%), bovine-viral-diarrhea virus type 1 (25%), infectious-bovine rhinotracheitis (58%), and bovine parainfluenza-3 (32%). Serum urea nitrogen, GGT, GG, and IGF-1 varied with year; T4, SUN, and GG varied with age (P 0.01); and SUN varied by capture area (P ¼ 0.02). Annual survival was 0.22 (SE¼0.035, n¼149) and varied by calving area but not year. Neonates captured in the Stephens Creek/Mammoth area of Yellowstone National Park, USA, had annual survival rates .33 higher (0.54) than those captured in the Lamar Valley area (0.17), likely due to the higher predator density in Lamar Valley. Summer survival (20 weeks after radiotagging) was 0.29 (SE ¼ 0.05, n ¼ 116), and calving area, absolute deviation from median birth date, and GG were important predictors of summer survival. Survival during winter (Nov-Apr) was 0.90 (SE ¼ 0.05, n ¼ 42), and it did not vary by calving area or year. Sixty-nine percent (n ¼ 104) of calves died within the first year of life, 24% (n ¼ 36) survived their first year, and 7% (n ¼ 11) had unknown fates. Grizzly bears (Ursus arctos) and black bears (Ursus americanus) accounted for 58-60% (n ¼ 60-62) of deaths, and wolves accounted for 14-17% (n ¼ 15-18). Summer predation (95% of summer deaths) increased, and winter malnutrition (0% of winter deaths) decreased, compared with a similar study during 1987-1990 (72% and 58%, respectively). Physiological factors (e.g., low levels of GG) may predispose calves to predation. Also, the increase in bear numbers since wolf restoration and spatial components finer than the northern range should be considered when trying to determine the causes of the northern Yellowstone elk decline. This is the first study to document the predation impacts from reintroduced wolves on elk calf mortality in an ecosystem already containing established populations of 4 other major predators (i. para determinar las causas del descenso del reclutamiento (de 33 a 13 crías /100 hembras adultas) tras la restauración del lobo (Canis lupus). Hemos capturado, marcado con radiotransmisores y evaluado las características de la sangre y la seroprevalencia de los anticuerpos a enfermedades de 151 crías 6 días (68M:83H). Las concentraciones (x, SE) de los indicadores del estado pote...

Ecological theory predicts that the diffuse risk cues generated by wide-ranging, active predators should induce prey behavioural responses but not major, population- or community-level consequences. We evaluated the non-consumptive effects (NCEs) of an active predator, the grey wolf (Canis lupus), by simultaneously tracking wolves and the behaviour, body fat, and pregnancy of elk (Cervus elaphus), their primary prey in the Greater Yellowstone Ecosystem. When wolves approached within 1 km, elk increased their rates of movement, displacement and vigilance. Even in high-risk areas, however, these encounters occurred only once every 9 days. Ultimately, despite 20-fold variation in the frequency of encounters between wolves and individual elk, the risk of predation was not associated with elk body fat or pregnancy. Our findings suggest that the ecological consequences of actively hunting large carnivores, such as the wolf, are more likely transmitted by consumptive effects on prey survival than NCEs on prey behaviour.

A “landscape of fear” (LOF) is a map that describes continuous spatial variation in an animal's perception of predation risk. The relief on this map reflects, for example, places that an animal avoids to minimize risk. Although the LOF concept is a potentially unifying theme in ecology that is often invoked to explain the ecological and conservation significance of fear, little is known about the daily dynamics of an LOF. Despite theory and data to the contrary, investigators often assume, implicitly or explicitly, that an LOF is a static consequence of a predator's mere presence within an ecosystem. We tested the prediction that an LOF in a large‐scale, free‐living system is a highly dynamic map with “peaks” and “valleys” that alternate across the diel (24‐h) cycle in response to daily lulls in predator activity. We did so with extensive data from the case study of Yellowstone elk (Cervus elaphus) and wolves (Canis lupus) that was the original basis for the LOF concept. We quantified the elk LOF, defined here as spatial allocation of time away from risky places and times, across nearly 1,000‐km2 of northern Yellowstone National Park and found that it fluctuated with the crepuscular activity pattern of wolves, enabling elk to use risky places during wolf downtimes. This may help explain evidence that wolf predation risk has no effect on elk stress levels, body condition, pregnancy, or herbivory. The ability of free‐living animals to adaptively allocate habitat use across periods of high and low predator activity within the diel cycle is an underappreciated aspect of animal behavior that helps explain why strong antipredator responses may trigger weak ecological effects, and why an LOF may have less conceptual and practical importance than direct killing.

While many wildlife species are threatened, some populations have recovered from previous overexploitation, and data linking these population increases with disease dynamics are limited. We present data suggesting that free-ranging elk (Cervus elaphus) are a maintenance host for Brucella abortus in new areas of the Greater Yellowstone Ecosystem (GYE). Brucellosis seroprevalence in free-ranging elk increased from 0-7% in 1991-1992 to 8-20% in 2006-2007 in four of six herd units around the GYE. These levels of brucellosis are comparable to some herd units where elk are artificially aggregated on supplemental feeding grounds. There are several possible mechanisms for this increase that we evaluated using statistical and population modeling approaches. Simulations of an age-structured population model suggest that the observed levels of seroprevalence are unlikely to be sustained by dispersal from supplemental feeding areas with relatively high seroprevalence or an older age structure. Increases in brucellosis seroprevalence and the total elk population size in areas with feeding grounds have not been statistically detectable. Meanwhile, the rate of seroprevalence increase outside the feeding grounds was related to the population size and density of each herd unit. Therefore, the data suggest that enhanced elk-to-elk transmission in free-ranging populations may be occurring due to larger winter elk aggregations. Elk populations inside and outside of the GYE that traditionally did not maintain brucellosis may now be at risk due to recent population increases. In particular, some neighboring populations of Montana elk were 5-9 times larger in 2007 than in the 1970s, with some aggregations comparable to the Wyoming feeding-ground populations. Addressing the unintended consequences of these increasing populations is complicated by limited hunter access to private lands, which places many ungulate populations out of administrative control. Agency-landowner hunting access partnerships and the protection of large predators are two management strategies that may be used to target high ungulate densities in private refuges and reduce the current and future burden of disease.

Variation in vital rates of an unharvested elk (Cervus elaphus) population was studied using telemetry for 7 consecutive years, 1991–1998. We found pronounced senescence in survival rates, but no evidence for reproductive senescence. Prime-age females (<10 years old) experienced very high annual survival rates (mean = 0.97, SE = 0.02), with lower survival rates for senescent animals ([Formula: see text]10 years old; mean = 0.79, SE = 0.06). There was evidence that the severity of snowpack conditions had little effect on survival of prime-age animals except during the most extreme winter, while survival of senescent animals was progressively depressed as the severity of snowpack conditions increased. Reproductive rates remained essentially constant, near their biological maxima (mean = 0.91, SE = 0.02). Annual re cruitment was highly variable. Snowpack had a pronounced effect on recruitment (r2 = 0.91), the most severe snowpack conditions resulting in the virtual elimination of a juvenile cohort. Population estimates and recruitment rates obtained during this investigation and historic data collected from 1965 to 1980 support the premise that the population has been maintained in a dynamic equilibrium for at least three decades despite the stochastic effects of climate variation on vital rates. We conclude that the population is resource-limited, with variation about the equilibrium caused primarily by variable recruitment driven by stochastic annual snowpack.

Summary1. Well-informed management of harvested species requires understanding how changing ecological conditions affect demography and population dynamics, information that is lacking for many species. We have limited understanding of the relative influence of carnivores, harvest, weather and forage availability on elk Cervus elaphus demography, despite the ecological and economic importance of this species. We assessed adult female survival, a key vital rate for population dynamics, from 2746 radio-collared elk in 45 populations across western North America that experience wide variation in carnivore assemblage, harvest, weather and habitat conditions. 2. Proportional hazard analysis revealed that 'baseline' (i.e. not related to human factors) mortality was higher with very high winter precipitation, particularly in populations sympatric with wolves Canis lupus. Mortality may increase via nutritional stress and heightened vulnerability to predation in snowy winters. Baseline mortality was unrelated to puma Puma concolor presence, forest cover or summer forage productivity. 3. Cause-specific mortality analyses showed that wolves and all carnivore species combined had additive effects on baseline elk mortality, but only reduced survival by <2%. When human factors were included, 'total' adult mortality was solely related to harvest; the influence of native carnivores was compensatory. Annual total mortality rates were lowest in populations sympatric with both pumas and wolves because managers reduced female harvest in areas with abundant or diverse carnivores. ‡Present address: Natural Resources Canada, 506 Burnside Road W, Victoria, BC V8Z 1M5, Canada *Correspondence author. E-mail: jedediah.brodie@gmail.com †Authorship alphabetical after B. Johnson.© 2013 This article is a US Government work and is in the public domain in the USA. 2013, 50, 295-305 doi: 10.1111/1365-2664.12044 4. Mortality from native carnivores peaked in late winter and early spring, while harvest-induced mortality peaked in autumn. The strong peak in harvest-induced mortality during the autumn hunting season decreased as the number of native carnivore species increased. 5. Synthesis and applications. Elevated baseline adult female elk mortality from wolves in years with high winter precipitation could affect elk abundance as winters across the western US become drier and wolves recolonize portions of the region. In the absence of human harvest, wolves had additive, although limited, effects on mortality. However, human harvest, and its apparent use by managers to offset predation, primarily controls overall variation in adult female mortality. Altering harvest quotas is thus a strong tool for offsetting impacts of carnivore recolonization and shifting weather patterns on elk across western North America. Journal of Applied Ecology

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Non‐native species can have adverse impacts on native species. Predicting the potential extent of distributional expansion and abundance of an invading non‐native species can inform appropriate conservation and management actions. Non‐native mountain goats (Oreamnos americanus) in the greater Yellowstone area (GYA) have substantial potential to occupy similar habitats to native Rocky Mountain bighorn sheep (Ovis canadensis canadensis). To understand the potential for expansion of mountain goats in the GYA, this study evaluated detection‐nondetection data derived from ground‐based occupancy surveys of viewsheds partitioned into a grid of 100 × 100 m sampling units. Surveys were conducted over three summer seasons (2011–2013) in two study areas with well‐established mountain goat populations. Relationships between scale‐specific habitat covariates and mountain goat selection were evaluated to model occupancy and detection probabilities based on mountain goat detections in 505 of the 53,098 sampling units surveyed. Habitat selection was most strongly associated with terrain covariates, including mean slope and slope variance, at a spatial scale of 500 × 500 m, and canopy cover, heat load, and normalized difference vegetation index at a spatial scale of 100 × 100 m. These results provide new insight into multi‐scale patterns of mountain goat habitat selection, as well as evidence that mean slope and slope variance are more informative terrain covariates than distance to escape terrain, which has been commonly used in published mountain goat habitat models. The model predicted 9,035 km2 of suitable habitat within the GYA, of which 57% is currently un‐colonized. Seventy‐five percent of all bighorn observations recorded in the GYA fall within predicted suitable mountain goat habitat. We also estimated that the GYA might have the potential to support 5,331–8,854 mountain goats when all predicted habitat is occupied, or approximately 2.5–4.2 times the most recent abundance estimate of 2,354.