Migration is a striking behavioral strategy by which many animals enhance resource acquisition while reducing predation risk. Historically, the demographic benefits of such movements made migration common, but in many taxa the phenomenon is considered globally threatened. Here we describe a long-term decline in the productivity of elk (Cervus elaphus) that migrate through intact wilderness areas to protected summer ranges inside Yellowstone National Park, USA. We attribute this decline to a long-term reduction in the demographic benefits that ungulates typically gain from migration. Among migratory elk, we observed a 21-year, 70% reduction in recruitment and a 4-year, 19% depression in their pregnancy rate largely caused by infrequent reproduction of females that were young or lactating. In contrast, among resident elk, we have recently observed increasing recruitment and a high rate of pregnancy. Landscape-level changes in habitat quality and predation appear to be responsible for the declining productivity of Yellowstone migrants. From 1989 to 2009, migratory elk experienced an increasing rate and shorter duration of green-up coincident with warmer spring-summer temperatures and reduced spring precipitation, also consistent with observations of an unusually severe drought in the region. Migrants are also now exposed to four times as many grizzly bears (Ursus arctos) and wolves (Canis lupus) as resident elk. Both of these restored predators consume migratory elk calves at high rates in the Yellowstone wilderness but are maintained at low densities via lethal management and human disturbance in the year-round habitats of resident elk. Our findings suggest that large-carnivore recovery and drought, operating simultaneously along an elevation gradient, have disproportionately influenced the demography of migratory elk. Many migratory animals travel large geographic distances between their seasonal ranges. Changes in land use and climate that disparately influence such seasonal ranges may alter the ecological basis of migratory behavior, representing an important challenge for, and a powerful lens into, the ecology and conservation of migratory taxa.

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...

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.

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Elk (Cervus elaphus) in the western United States are an economically and socially valuable wildlife species. They have featured species status for federal land management planning; hence, considerable modeling focused on habitat evaluation and land management planning has been undertaken for elk. The extent to which these and other habitat models for large ungulates account for influences of nutritional resources varies greatly, probably because of varying recognition of the importance of nutrition and uncertainty about how to measure and model nutrition. Our primary goals were to 1) develop greater understanding of how habitat conditions influence foraging dynamics and nutrition of elk in summer and autumn; and 2) illustrate an ecological framework for evaluating and predicting nutritional resources so that nutritional needs of elk can be integrated within landscape‐scale plans, population models, and habitat evaluation models. We evaluated foraging responses of elk to clearcut logging and commercial thinning, forest succession, and season across ecological site potentials. We also identified the extent to which plant communities satisfied nutritional requirements of lactating female elk and their calves. Our study was conducted in the temperate rainforests of the Pacific Northwest on industrial and public timberlands. We evaluated relations between habitat conditions and elk nutrition in plant communities representing a range in stand age and ecological conditions at 3 study areas, 1 near the Canadian border in the north Cascades Mountains (Nooksack), 1 in the Coast range southwest of Olympia, Washington (Willapa Hills), and the third in the central Cascades near Springfield, Oregon (Springfield), from late June to November, 2000–2002. In 98–143 macroplots per study area, we measured forage abundance by plant species, digestible energy content by plant life‐form group, and forest overstory. In a subset of these macroplots (∼30 per study area), we held 4 tame lactating elk with calves in electrified pens (n = 15–25 adult elk per year), and sampled activity budgets, dietary composition, forage selection, and other measures of foraging behavior; dietary digestible energy (DE; kcal/g) and protein (DP; %) levels; and intake rates of these nutrients. In 15 of these pens, we held elk for extended periods (13–21 days) to monitor changes in body fat of adults and growth of calves. We developed equations to predict dietary DE and DP and per‐minute intake rates of each in a nutrition prediction model that reflected vegetation attributes and ecological site influences. Total abundance of forage in the western hemlock series after clearcut logging in low to moderate elevations (≤1,000 m) ranged from a peak of 3,000–4,500 kg/ha in 5‐ to 10‐year‐old stands to 100–300 kg/ha in 20‐ to 50‐year‐old stands with only moderate increases through late succession. Patterns were similar in higher elevation forests (1,000–1,800 m), although peaks and troughs in forage abundance developed more slowly. Deciduous shrubs, forbs, an...

We developed new, and validated existing, indices of nutritional condition for live and dead mule deer (Odocoileus hemionus). Live animal indices included a body condition score (BCS), thickness of subcutaneous fat and selected muscles using ultrasonography, and body mass. Dead animal indices included femur, metatarsal, and mandible marrow fat, 3 kidney fat indices, and 2 carcass scoring methods. We used 21 female deer and 4 castrates (1‐11 yr old) varying widely in nutritional condition (2‐28% ingesta‐free body fat). Deer were euthanized and homogenized for chemical analysis of fat, protein, water, and ash content. Estimates of fat and gross energy (GE) were regressed against each condition indicator using regression. Subcutaneous fat thickness, a rump BCS, and rLIVINDEX (an arithmetic combination of subcutaneous fat thickness and the rump BCS) were most related to condition for live animals (r2 ≥ 0.87, P < 0.001) whereas the Kistner score and kidney fat were most related to fat and GE for dead animals (r2 ≥ 0.77, P < 0.001). We also evaluated range of usefulness and sensitivity to small changes in body condition for all models. In general, indices with moderate or highly curvilinear statistical relations to body fat or those based on only one fat depot or a small number of ranking scores will have limitations in their use. Our results identify robust tools for a variety of research and monitoring designs useful for evaluating nutrition's effect on mule deer populations.

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.

Each spring, migratory herbivores around the world track or ‘surf’ green waves of newly emergent vegetation to distant summer or wet‐season ranges. This foraging tactic may help explain the great abundance of migratory herbivores on many seasonal landscapes. However, the underlying fitness benefits of this life‐history strategy remain poorly understood. A fundamental prediction of the green‐wave hypothesis is that migratory herbivores obtain fitness benefits from surfing waves of newly emergent vegetation more closely than their resident counterparts. Here we evaluate whether this behavior increases body‐fat levels – a critically important correlate of reproduction and survival for most ungulates – in elk Cervus elaphus of the Greater Yellowstone Ecosystem. Using satellite imagery and GPS tracking data, we found evidence that migrants (n = 23) indeed surfed the green wave, occupying sites 12.7 days closer to peak green‐up than residents (n = 16). Importantly, individual variation in surfing may help account for up to 6 kg of variation in autumn body‐fat levels. Our findings point to a pathway for anthropogenic changes to the green wave (e.g. climate change) or migrants’ ability to surf it (e.g. development) to impact migratory populations. To explore this possibility, we evaluated potential population‐level consequences of constrained surfing with a heuristic model. If green‐wave surfing deteriorates by 5–15 days from observed, our model predicts up to a 20% decrease in pregnancy rates, a 2.5% decrease in population growth, and a 30% decrease in abundance over 50 years. By linking green‐wave surfing to fitness and illustrating potential effects on population growth, our study provides new insights into the evolution of migratory behavior and the prospects for the persistence of migratory ungulate populations in a changing world.