Demographic data show many populations of Rocky Mountain (Cervus elaphus nelsoni) and Roosevelt (Cervus elaphus roosevelti) elk have been declining over the last few decades. Recent work suggests that forage quality and associated animal nutritional condition, particularly in late summer and early autumn, influence reproduction and survival in elk. Therefore, we estimated seasonal nutritional condition of 861 female elk in 2,114 capture events from 21 herds in Washington, Oregon, Wyoming, Colorado, and South Dakota from 1998 to 2007. We estimated ingesta-free body fat and body mass, and determined age, pregnancy status, and lactation status. We obtained estimates for most herds in both late winter-early spring (late Feb-early Apr) and in autumn (Nov-early Dec) to identify changes in nutritional condition of individuals across seasons.Body fat levels of lactating females in autumn were consistently lower than their non-lactating counterparts, and herd averages of lactating elk ranged from 5.5% to 12.4%. These levels were 30-75% of those documented for captive lactating elk fed high-quality diets during summer and autumn. Body fat levels were generally lowest in the coastal and inland northwest regions and highest along the west-slope of the northern Cascades. Adult females in most herds lost an average of 30.7 kg (range: 5-62 kg), or about 13% (range: 2.6-25%) of their autumn mass during winter, indicating nutritional deficiencies. However, we found no significant relationships between spring body fat or change in body fat over winter with winter weather, region, or herd, despite markedly different winter weather among herds and regions. Instead, body fat levels in spring were primarily a function of fat levels the previous autumn. Thinner females in autumn lost less body fat and body mass over winter than did fatter females, a compensatory response, but still ended the season with less body fat than the fatter elk.Body fat levels of lactating females in autumn varied among herds but were unrelated to their body fat levels the previous spring. Within herds, thinner females exhibited a compensatory response during summer and accrued more fat than their fatter counterparts over summer, resulting in similar body fat levels among lactating elk in autumn despite considerable differences in their fat levels the previous spring. Level of body fat achieved by lactating females in autumn varied 2-fold among herds, undoubtedly because of differences in summer nutrition. Thus, summer nutrition set limits to rates of body fat accrual of lactating females that in turn limited body condition across the annual cycle.Pregnancy rates of 2-to 14-year-old females ranged from 68% to 100% in coastal populations of Washington, 69% to 98% in Cascade populations of Washington and Oregon, 84% to 94% in inland northwestern populations of Washington and Oregon, and 78% to 93% in Rocky Mountain populations. We found evidence of late breeding, even in herds with comparatively high pregnancy rates. Mean body mass of calves (n ¼ 242) in 3 popul...

Because they do not require sacrificing animals, body condition scores (BCS), thickness of rump fat (MAXFAT), and other similar predictors of body fat have advanced estimating nutritional condition of ungulates and their use has proliferated in North America in the last decade. However, initial testing of these predictors was too limited to assess their reliability among diverse habitats, ecotypes, subspecies, and populations across the continent. With data collected from mule deer (Odocoileus hemionus), elk (Cervus elaphus), and moose (Alces alces) during initial model development and data collected subsequently from free-ranging mule deer and elk herds across much of the western United States, we evaluated reliability across a broader range of conditions than were initially available. First, to more rigorously test reliability of the MAXFAT index, we evaluated its robustness across the 3 species, using an allometric scaling function to adjust for differences in animal size. We then evaluated MAXFAT, rump body condition score (rBCS), rLIVINDEX (an arithmetic combination of MAXFAT and rBCS), and our new allometrically scaled rump-fat thickness index using data from 815 free-ranging female Roosevelt and Rocky Mountain elk (C. e. roosevelti and C. e. nelsoni) from 19 populations encompassing 4 geographic regions and 250 free-ranging female mule deer from 7 populations and 2 regions. We tested for effects of subspecies, geographic region, and captive versus free-ranging existence. Rump-fat thickness, when scaled allometrically with body mass, was related to ingesta-free body fat over a 38-522-kg range of body mass (r 2 5 0.87; P , 0.001), indicating the technique is remarkably robust among at least the 3 cervid species of our analysis. However, we found an underscoring bias with the rBCS for elk that had .12% body fat. This bias translated into a difference between subspecies, because Rocky Mountain elk tended to be fatter than Roosevelt elk in our sample. Effects of observer error with the rBCS also existed for mule deer with moderate to high levels of body fat, and deer body size significantly affected accuracy of the MAXFAT predictor. Our analyses confirm robustness of the rump-fat index for these 3 species but highlight the potential for bias due to differences in body size and to observer error with BCS scoring. We present alternative LIVINDEX equations where potential bias from rBCS and bias due to body size are eliminated or reduced. These modifications improve the accuracy of estimating body fat for projects intended to monitor nutritional status of herds or to evaluate nutrition's influence on population demographics.

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

Raw counts from aerial surveys make no correction for undetected animals and provide no estimate of precision with which to judge the utility of the counts. Sightability modeling and double‐observer (DO) modeling are 2 commonly used approaches to account for detection bias and to estimate precision in aerial surveys. We developed a hybrid DO sightability model (model MH) that uses the strength of each approach to overcome the weakness in the other, for aerial surveys of elk (Cervus elaphus). The hybrid approach uses detection patterns of 2 independent observer pairs in a helicopter and telemetry‐based detections of collared elk groups. Candidate MH models reflected hypotheses about effects of recorded covariates and unmodeled heterogeneity on the separate front‐seat observer pair and back‐seat observer pair detection probabilities. Group size and concealing vegetation cover strongly influenced detection probabilities. The pilot's previous experience participating in aerial surveys influenced detection by the front pair of observers if the elk group was on the pilot's side of the helicopter flight path. In 9 surveys in Mount Rainier National Park, the raw number of elk counted was approximately 80–93% of the abundance estimated by model MH. Uncorrected ratios of bulls per 100 cows generally were low compared to estimates adjusted for detection bias, but ratios of calves per 100 cows were comparable whether based on raw survey counts or adjusted estimates. The hybrid method was an improvement over commonly used alternatives, with improved precision compared to sightability modeling and reduced bias compared to DO modeling. © 2013 The Wildlife Society.

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