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      How climate impacts the composition of wolf‐killed elk in northern Yellowstone National Park

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          Abstract

          1. While the functional response of predators is commonly measured, recent work has revealed that the age and sex composition of prey killed is often a better predictor of prey population dynamics because the reproductive value of adult females is usually higher than that of males or juveniles.

          2. Climate is often an important mediating factor in determining the composition of predator kills, but we currently lack a mechanistic understanding of how the multiple facets of climate interact with prey abundance and demography to influence the composition of predator kills.

          3. Over 20 winters, we monitored 17 wolf packs in Yellowstone National Park and recorded the sex, age and nutritional condition of kills of their dominant prey—elk—in both early and late winter periods when elk are in relatively good and relatively poor condition, respectively.

          4. Nutritional condition (as indicated by per cent marrow fat) of wolf‐killed elk varied markedly with summer plant productivity, snow water equivalent (SWE) and winter period. Moreover, marrow was poorer for wolf‐killed bulls and especially for calves than it was for cows.

          5. Wolf prey composition was influenced by a complex set of climatic and endogenous variables. In early winter, poor plant growth in either year t or − 1, or relatively low elk abundance, increased the odds of wolves killing bulls relative to cows. Calves were most likely to get killed when elk abundance was high and when the forage productivity they experienced in utero was poor. In late winter, low SWE and a relatively large elk population increased the odds of wolves killing calves relative to cows, whereas low SWE and poor vegetation productivity 1 year prior together increased the likelihood of wolves killing a bull instead of a cow.

          6. Since climate has a strong influence on whether wolves prey on cows (who, depending on their age, are the key reproductive components of the population) or lower reproductive value of calves and bulls, our results suggest that climate can drive wolf predation to be more or less additive from year to year.

          Abstract

          The impacts of large carnivores on their prey are importantly impacted by the composition of the prey killed. Here the authors show that climate and elk population size interact in important ways to determine when wolves kill cows, bulls or calves.

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          Most cited references34

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          Patterns of predation in a diverse predator-prey system.

          There are many cases where animal populations are affected by predators and resources in terrestrial ecosystems, but the factors that determine when one or the other predominates remain poorly understood. Here we show, using 40 years of data from the highly diverse mammal community of the Serengeti ecosystem, East Africa, that the primary cause of mortality for adults of a particular species is determined by two factors--the species diversity of both the predators and prey and the body size of that prey species relative to other prey and predators. Small ungulates in Serengeti are exposed to more predators, owing to opportunistic predation, than are larger ungulates; they also suffer greater predation rates, and experience strong predation pressure. A threshold occurs at prey body sizes of approximately 150 kg, above which ungulate species have few natural predators and exhibit food limitation. Thus, biodiversity allows both predation (top-down) and resource limitation (bottom-up) to act simultaneously to affect herbivore populations. This result may apply generally in systems where there is a diversity of predators and prey.
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            Trophic facilitation by introduced top predators: grey wolf subsidies to scavengers in Yellowstone National Park

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              Animal migration amid shifting patterns of phenology and predation: lessons from a Yellowstone elk herd.

              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.
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                Author and article information

                Contributors
                cwilmers@ucsc.edu
                Journal
                J Anim Ecol
                J Anim Ecol
                10.1111/(ISSN)1365-2656
                JANE
                The Journal of Animal Ecology
                John Wiley and Sons Inc. (Hoboken )
                0021-8790
                1365-2656
                27 March 2020
                June 2020
                : 89
                : 6 ( doiID: 10.1111/jane.v89.6 )
                : 1511-1519
                Affiliations
                [ 1 ] Center for Integrated Spatial Research Environmental Studies Department University of California Santa Cruz CA USA
                [ 2 ] Yellowstone Center for Resources Yellowstone National Park WY USA
                [ 3 ] Wildlife Biology Program Department of Ecosystem and Conservation Sciences University of Montana Missoula MT USA
                [ 4 ] Department of Wildland Resources and Ecology Center Utah State University Logan UT USA
                Author notes
                [*] [* ] Correspondence

                Christopher C. Wilmers

                Email: cwilmers@ 123456ucsc.edu

                Author information
                https://orcid.org/0000-0003-2063-1478
                https://orcid.org/0000-0001-8218-1455
                Article
                JANE13200
                10.1111/1365-2656.13200
                7317765
                32145069
                bc883c5e-5953-4d87-ba69-619155bc8b0c
                © 2020 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society

                This is an open access article under the terms of the http://creativecommons.org/licenses/by/4.0/ License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

                History
                : 08 July 2019
                : 04 February 2020
                Page count
                Figures: 4, Tables: 0, Pages: 9, Words: 6931
                Funding
                Funded by: Yellowstone Park Foundation
                Funded by: Yellowstone Forever
                Funded by: National Science Foundation , open-funder-registry 10.13039/100000001;
                Award ID: DEB‐0613730
                Award ID: DEB‐1245373
                Funded by: Yellowstone National Park
                Funded by: US National Park Service , open-funder-registry 10.13039/100007516;
                Categories
                Research Article
                Community Ecology
                Custom metadata
                2.0
                June 2020
                Converter:WILEY_ML3GV2_TO_JATSPMC version:5.8.4 mode:remove_FC converted:26.06.2020

                Ecology
                age structure,canis lupus,climate change,predator–prey dynamics,prey selection
                Ecology
                age structure, canis lupus, climate change, predator–prey dynamics, prey selection

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