White-tailed deer population dynamics in a multipredator landscape shaped by humans.
Canis lupus
Odocoileus virginianus
Puma concolor
agriculture
carnivore
population dynamics
predator–prey interactions
survival analysis
timber harvest
ungulate
Journal
Ecological applications : a publication of the Ecological Society of America
ISSN: 1051-0761
Titre abrégé: Ecol Appl
Pays: United States
ID NLM: 9889808
Informations de publication
Date de publication:
18 Jun 2024
18 Jun 2024
Historique:
revised:
23
02
2024
received:
01
09
2023
accepted:
22
04
2024
medline:
19
6
2024
pubmed:
19
6
2024
entrez:
19
6
2024
Statut:
aheadofprint
Résumé
Large terrestrial mammals increasingly rely on human-modified landscapes as anthropogenic footprints expand. Land management activities such as timber harvest, agriculture, and roads can influence prey population dynamics by altering forage resources and predation risk via changes in habitat, but these effects are not well understood in regions with diverse and changing predator guilds. In northeastern Washington state, USA, white-tailed deer (Odocoileus virginianus) are vulnerable to multiple carnivores, including recently returned gray wolves (Canis lupus), within a highly human-modified landscape. To understand the factors governing predator-prey dynamics in a human context, we radio-collared 280 white-tailed deer, 33 bobcats (Lynx rufus), 50 cougars (Puma concolor), 28 coyotes (C. latrans), and 14 wolves between 2016 and 2021. We first estimated deer vital rates and used a stage-structured matrix model to estimate their population growth rate. During the study, we observed a stable to declining deer population (lambda = 0.97, 95% confidence interval: 0.88, 1.05), with 74% of Monte Carlo simulations indicating population decrease and 26% of simulations indicating population increase. We then fit Cox proportional hazard models to evaluate how predator exposure, use of human-modified landscapes, and winter severity influenced deer survival and used these relationships to evaluate impacts on overall population growth. We found that the population growth rate was dually influenced by a negative direct effect of apex predators and a positive effect of timber harvest and agricultural areas. Cougars had a stronger effect on deer population dynamics than wolves, and mesopredators had little influence on the deer population growth rate. Areas of recent timber harvest had 55% more forage biomass than older forests, but horizontal visibility did not differ, suggesting that timber harvest did not influence predation risk. Although proximity to roads did not affect the overall population growth rate, vehicle collisions caused a substantial proportion of deer mortalities, and reducing these collisions could be a win-win for deer and humans. The influence of apex predators and forage indicates a dual limitation by top-down and bottom-up factors in this highly human-modified system, suggesting that a reduction in apex predators would intensify density-dependent regulation of the deer population owing to limited forage availability.
Types de publication
Journal Article
Langues
eng
Sous-ensembles de citation
IM
Pagination
e3003Subventions
Organisme : University of Washington College of the Environment Hall Genetics Grant
Organisme : Washington Department of Fish and Wildlife Aquatic Lands Enhancement Account
ID : 19-135
Organisme : Washington State Legislature
Organisme : National Science Foundation
ID : DEB-1652420
Organisme : Rocky Mountain Elk Foundation
ID : WA190025
Informations de copyright
© 2024 The Ecological Society of America.
Références
Anderson, D. R., and K. P. Burnham. 2002. “Avoiding Pitfalls when Using Information‐Theoretic Methods.” Journal of Wildlife Management 66: 912–918.
Ballard, W. B. 2011. “Predator‐Prey Relationships.” In Biology and Management of White‐Tailed Deer, edited by D. G. Hewitt, 251–286. Boca Raton: CRC Press.
Ballard, W. B., D. Lutz, T. W. Keegan, L. H. Carpenter, and J. C. deVos Jr. 2001. “Deer‐Predator Relationships: A Review of Recent North American Studies with Emphasis on Mule and Black‐Tailed Deer.” Wildlife Society Bulletin 29: 99–115.
Ballard, W. B., H. A. Whitlaw, B. F. Wakeling, R. L. Brown, J. C. deVos Jr, and M. C. Wallace. 2000. “Survival of Female Elk in Northern Arizona.” Journal of Wildlife Management 64: 500–504.
Barker, K. J., E. Cole, A. Courtemanch, S. Dewey, D. Gustine, K. Mills, J. Stephenson, B. Wise, and A. D. Middleton. 2023. “Large Carnivores Avoid Humans while Prioritizing Prey Acquisition in Anthropogenic Areas.” Journal of Animal Ecology 92: 889–900.
Barker, K. J., M. S. Mitchell, K. M. Proffitt, and J. D. DeVoe. 2019. “Land Management Alters Traditional Nutritional Benefits of Migration for Elk.” Journal of Wildlife Management 83: 167–174.
Bassing, S. B. 2022. “Spatial and Temporal Patterns of Predator‐Prey Interactions in a Large Mammal Community in Eastern Washington.” PhD diss., University of Washington.
Beausoleil, R. A., L. S. Welfelt, I. N. Keren, B. N. Kertson, B. T. Maletzke, and G. M. Koehler. 2021. “Long‐Term Evaluation of Cougar Density and Application of Risk Analysis for Harvest Management.” Journal of Wildlife Management 85: 462–473.
Berger, J. 2007. “Fear, Human Shields and the Redistribution of Prey and Predators in Protected Areas.” Biology Letters 3: 620–623.
Berger, K. M., and M. M. Conner. 2008. “Recolonizing Wolves and Mesopredator Suppression of Coyotes: Impacts on Pronghorn Population Dynamics.” Ecological Applications 18: 599–612.
Bergman, E. J., C. J. Bishop, D. J. Freddy, G. C. White, and P. F. Doherty. 2014. “Habitat Management Influences Overwinter Survival of Mule Deer Fawns in Colorado.” Journal of Wildlife Management 78: 448–455.
Berry, S. L., L. A. Shipley, R. A. Long, and C. Loggers. 2019. “Differences in Dietary Niche and Foraging Behavior of Sympatric Mule and White‐Tailed Deer.” Ecosphere 10: e02815.
Bishop, C. J., G. C. White, D. J. Freddy, B. E. Watkins, and T. R. Stephenson. 2009. “Effect of Enhanced Nutrition on Mule Deer Population Rate of Change.” Wildlife Monographs 172: 1–28.
Boyce, M. S., C. J. Johnson, E. H. Merrill, S. E. Nielsen, E. J. Solberg, and B. vanMoorter. 2016. “Can Habitat Selection Predict Abundance.” Journal of Animal Ecology 85: 11–20.
Boyce, M. S., P. R. Vernier, S. E. Nielsen, and F. K. A. Schmiegelow. 2002. “Evaluating Resource Selection Functions.” Ecological Modelling 157: 281–300.
Carstensen, M., G. D. Delgiudice, B. A. Sampson, and D. W. Kuehn. 2009. “Survival, Birth Characteristics, and Cause‐Specific Mortality of White‐Tailed Deer Neonates.” Journal of Wildlife Management 73: 175–183.
Carter, N. H., and J. D. C. Linnell. 2016. “Co‐Adaptation Is Key to Coexisting with Large Carnivores.” Trends in Ecology & Evolution 31: 575–578.
Chamberlain, S. 2018. “Rphylopic: Get Silhouettes of Organisms from PhyloPic.” R Package Version 0.2.0. https://www.phylopic.org.
Clark, T. J., and M. Hebblewhite. 2021. “Predator Control May Not Increase Ungulate Populations in the Future: A Formal Meta‐Analysis.” Journal of Applied Ecology 58: 812–824.
Clover, M. R. 1954. “A Portable Deer Trap and Catch‐Net.” California Fish & Game 40: 367–373.
Conover, M. R. 2019. “Numbers of Human Fatalities, Injuries, and Illnesses in the United States Due to Wildlife.” Human‐Wildlife Interactions 13: 264–276.
Cook, J. G., R. C. Cook, R. W. Davis, and L. L. Irwin. 2016. “Nutritional Ecology of Elk during Summer and Autumn in the Pacific Northwest.” Wildlife Monographs 195: 1–81.
Cook, R. C., J. G. Cook, D. J. Vales, B. K. Johnson, S. M. McCorquodale, L. A. Shipley, R. A. Riggs, et al. 2013. “Regional and Seasonal Patterns of Nutritional Condition and Reproduction in Elk.” Wildlife Monographs 184: 1–45.
Côté, S. D., and M. Festa‐Bianchet. 2001. “Birthdate, Mass and Survival in Mountain Goat Kids: Effects of Maternal Characteristics and Forage Quality.” Oecologia 127: 230–238.
Cox, D. R. 1972. “Regression Models and Life‐Tables.” Journal of the Royal Statistical Society: Series B (Methodological) 34: 187–202.
Cunningham, C. A., T. A. Nuñez, Y. Hentati, B. Sullender, C. Breen, T. R. Ganz, S. E. S. Kreling, et al. 2022. “Permanent Daylight Savings Time Would Reduce Deer‐Vehicle Collisions.” Current Biology 32: 4982–4988.
Daly, C., M. Halbleib, J. I. Smith, W. P. Gibson, M. K. Doggett, G. H. Taylor, J. Curtis, and P. P. Pasteris. 2008. “Physiographically Sensitive Mapping of Climatological Temperature and Precipitation across the Conterminous United States.” International Journal of Climatology 28: 2031–2064.
Dellinger, J. A., C. R. Shores, A. Craig, M. R. Heithaus, W. J. Ripple, and A. J. Wirsing. 2019. “Habitat Use of Sympatric Prey Suggest Divergent Anti‐Predator Responses to Recolonizing Gray Wolves.” Oecologia 189: 487–500.
DeMars, C. A., and S. Boutin. 2018. “Nowhere to Hide: Effects of Linear Features on Predator‐Prey Dynamics in a Large Mammal System.” Journal of Animal Ecology 87: 274–284.
DeYoung, C. 2011. “Population Dynamics.” In Biology and Management of White‐Tailed Deer, edited by D. G. Hewitt, 147–180. Boca Raton: CRC Press.
Donadio, E., and S. W. Buskirk. 2016. “Linking Predation Risk, Ungulate Antipredator Responses, and Patterns of Vegetation in the High Andes.” Journal of Mammalogy 97: 966–977.
Dormann, C. F., J. Elith, S. Bacher, C. Buchmann, G. Carl, G. Carré, J. R. García Marquéz, et al. 2013. “Collinearity: A Review of Methods to Deal with it and a Simulation Study Evaluating their Performance.” Ecography 36: 27–46.
Duquette, J. F., J. L. Belant, D. E. Beyer, and N. J. Svoboda. 2012. “Comparison of Pregnancy Detection Methods in Live White‐Tailed Deer.” Wildlife Society Bulletin 36: 115–118.
Eacker, D. R., M. Hebblewhite, K. M. Proffitt, B. S. Jimenez, M. S. Mitchell, and H. S. Robinson. 2016. “Annual Elk Calf Survival in a Multiple Carnivore System.” Journal of Wildlife Management 80: 1345–1359.
Eberhardt, L. L. 2002. “A Paradigm for Population Analysis of Long‐Lived Vertebrates.” Ecology 83: 2841–2854.
Elbroch, L. M. 2003. Mammal Tracks & Sign: A Guide to North American Species, 1st ed. Mechanicsburg: Stackpole Books.
Elbroch, L. M., P. E. Lendrum, J. Newby, H. Quigley, and D. J. Thompson. 2015. “Recolonizing Wolves Influence the Realized Niche of Resident Cougars.” Zoological Studies 54: 1–11.
Elbroch, L. M., and H. U. Wittmer. 2012. “Table Scraps: Inter‐Trophic Food Provisioning by Pumas.” Biology Letters 8: 776–779.
Elbroch, L. M., and H. U. Wittmer. 2013. “The Effects of Puma Prey Selection and Specialization on Less Abundant Prey in Patagonia.” Journal of Mammalogy 94: 259–268.
Elbroch, M., and C. McFarland. 2019. Mammal Tracks & Sign: A Guide to North American Species, 2nd ed. Guilford: Stackpole Books.
Estes, J. A., J. Terborgh, J. S. Brashares, M. E. Power, J. Berger, W. J. Bond, S. R. Carpenter, et al. 2011. “Trophic Downgrading of Planet Earth.” Science 333: 301–306.
Fieberg, J., and G. D. DelGiudice. 2009. “What Time Is it? Choice of Time Origin and Scale in Extended Proportional Hazards Models.” Ecology 90: 1687–1697.
Fischer, J., and D. B. Lindenmayer. 2007. “Landscape Modification and Habitat Fragmentation: A Synthesis.” Global Ecology and Biogeography 16: 265–280.
Forrester, T. D., and H. U. Wittmer. 2013. “A Review of the Population Dynamics of Mule Deer and Black‐Tailed Deer (Odocoileus hemionus) in North America.” Mammal Review 43: 292–308.
Frame, P. F., and T. J. Meier. 2007. “Field‐Assessed Injury to Wolves Captured in Rubber‐Padded Traps.” Journal of Wildlife Management 71: 2074–2076.
Gaillard, J.‐M., M. Festa‐Bianchet, and N. G. Yoccoz. 1998. “Population Dynamics of Large Herbivores: Variable Recruitment with Constant Adult Survival.” Trends in Ecology & Evolution 13: 58–63.
Ganz, T. R., M. T. DeVivo, B. N. Kertson, R. Roussin, L. Satterfield, A. J. Wirsing, and L. R. Prugh. 2022. “Interactive Effects of Wildfires, Season and Predator Activity Shape Mule Deer Movements.” Journal of Animal Ecology 91: 2273–2288.
Ganz, T. R., M. T. DeVivo, E. M. Reese, and L. R. Prugh. 2022. “Wildlife Whodunnit: Forensic Identification of Predators to Inform Wildlife Management and Conservation.” Wildlife Society Bulletin 47: e1386.
Gervasi, V., E. B. Nilsen, H. Sand, M. Panzacchi, G. R. Rauset, H. C. Pedersen, J. Kindberg, et al. 2012. “Predicting the Potential Demographic Impact of Predators on their Prey: A Comparative Analysis of Two Carnivore‐Ungulate Systems in Scandinavia.” Journal of Animal Ecology 81: 443–454.
Gilbert, S. L., M. S. Lindberg, K. J. Hundertmark, and D. K. Person. 2014. “Dead before Detection: Addressing the Effects of Left Truncation on Survival Estimation and Ecological Inference for Neonates.” Methods in Ecology and Evolution 5: 992–1001.
Gilbert, S. L., K. J. Sivy, C. B. Pozzanghera, A. DuBour, K. Overduijn, M. M. Smith, J. Zhou, J. M. Little, and L. R. Prugh. 2017. “Socioeconomic Benefits of Large Carnivore Recolonization through Reduced Wildlife‐Vehicle Collisions.” Conservation Letters 10: 431–439.
Griffin, K. A., M. Hebblewhite, H. S. Robinson, P. Zager, S. M. Barber‐Meyer, D. Christianson, S. Creel, et al. 2011. “Neonatal Mortality of Elk Driven by Climate, Predator Phenology and Predator Community Composition.” Journal of Animal Ecology 80: 1246–1257.
Guiden, P. W., S. L. Bartel, N. W. Byer, A. A. Shipley, and J. L. Orrock. 2019. “Predator–Prey Interactions in the Anthropocene: Reconciling Multiple Aspects of Novelty.” Trends in Ecology & Evolution 34: 616–627.
Hansen, M. C., P. V. Potapov, R. Moore, M. Hancher, S. A. A. Turubanova, A. Tyukavina, D. Thau, S. V. Stehman, S. J. Goetz, and T. R. Loveland. 2013. “High‐Resolution Global Maps of 21st‐Century Forest Cover Change.” Science 342: 850–853.
Hawkins, R. E., L. D. Martoglio, and G. G. Montgomery. 1968. “Cannon‐Netting Deer.” Journal of Wildlife Management 32: 191–195.
Hayes, T. A., N. J. DeCesare, C. J. Peterson, C. J. Bishop, and M. S. Mitchell. 2022. “Trade‐Offs in Forest Disturbance Management for Plant Communities and Ungulates.” Forest Ecology and Management 506: 119972.
Hebblewhite, M., and E. H. Merrill. 2007. “Multiscale Wolf Predation Risk for Elk: Does Migration Reduce Risk.” Oecologia 152: 377–387.
Hijmans, R. J., J. Van Etten, J. Cheng, M. Mattiuzzi, M. Sumner, J. A. Greenberg, O. P. Lamigueiro, A. Bevan, E. B. Racine, and A. Shortridge. 2015. “Package ‘Raster’.” R Package Version 3.6‐14. http://CRAN.R-project.org/package=raster.
Hoenes, B., S. Hansen, R. Harris, and J. Nelson. 2016. Wildlife Program 2015–2017 Ungulate Assessment. Olympia: Washington Department of Fish and Wildlife https://wdfw.wa.gov/publications/01948.
Huijser, M. P., P. McGowan, A. Hardy, A. Kociolek, A. P. Clevenger, D. Smith, and R. Ament. 2008. “Wildlife‐Vehicle Collision Reduction Study: Report to Congress.” U.S. Department of Transportation, Federal Highway Administration. https://www.fhwa.dot.gov/publications/research/safety/08034/.
Hull, I. T., L. A. Shipley, S. L. Berry, C. Loggers, and T. R. Johnson. 2020. “Effects of Fuel Reduction Timber Harvests on Forage Resources for Deer in Northeastern Washington.” Forest Ecology and Management 458: 117757.
Ivlev, V. S., and D. Scott. 1961. Experimental Ecology of the Feeding of Fishes. New Haven: Yale University Press.
Jensen, A. J., C. J. Marneweck, J. C. Kilgo, and D. S. Jachowski. 2022. “Coyote Diet in North American: Geographic and Ecological Patterns during Range Expansion.” Mammal Review 52: 480–496.
Jessup, D. A. 1982. “The Use of the Helicopter in the Capture of Free‐Roaming Wildlife.” In Chemical Immobilization of North American Wildlife, edited by L. Nielsen, J. C. Haight, and M. E. Fouler, 289–303. Milwaukee: The Wisconsin Humane Society Inc.
Johnson, B. K., P. K. Coe, and R. L. Green. 2013. “Abiotic, Bottom‐Up, and Top‐Down Influences on Recruitment of Rocky Mountain Elk in Oregon: A Retrospective Analysis.” Journal of Wildlife Management 77: 102–116.
Johnson, B. K., T. McCoy, C. O. Kochanny, and R. C. Cook. 2006. “Evaluation of Vaginal Implant Transmitters in Elk (Cervus elaphus nelsonii).” Journal of Zoo and Wildlife Medicine 37: 301–305.
Johnson, D. H. 1980. “The Comparison of Usage and Availability Measurements for Evaluating Resource Preference.” Ecology 61: 65–71.
Kaplan, E. L., and P. Meier. 1958. “Nonparametric Estimation from Incomplete Observations.” Journal of the American Statistical Association 53: 457–481.
Kauffman, M. J., N. Varley, D. W. Smith, D. R. Stahler, D. R. MacNulty, and M. S. Boyce. 2007. “Landscape Heterogeneity Shapes Predation in a Newly Restored Predator‐Prey System.” Ecology Letters 10: 690–700.
Kendall, B. E., M. Fujiwara, J. Diaz‐Lopez, S. Schneider, J. Voigt, and S. Wiesner. 2019. “Persistent Problems in the Construction of Matrix Population Models.” Ecological Modelling 406: 33–43.
Kennedy, C. M., J. R. Oakleaf, D. M. Theobald, S. Baruch‐Mordo, and J. Kiesecker. 2019. “Managing the Middle: A Shift in Conservation Priorities Based on the Global Human Modification Gradient.” Global Change Biology 25: 811–826.
Kertson, B. N., R. D. Spencer, J. M. Marzluff, J. Hepinstall‐Cymerman, and C. E. Grue. 2011. “Cougar Space Use and Movements in the Wildland–Urban Landscape of Western Washington.” Ecological Applications 21: 2866–2881.
Kuijper, D. P. J., E. Sahlén, B. Elmhagen, S. Chamaillé‐Jammes, H. Sand, K. Lone, and J. P. G. M. Cromsigt. 2016. “Paws without Claws? Ecological Effects of Large Carnivores in Anthropogenic Landscapes.” Proceedings of the Royal Society B: Biological Sciences 283: 20161625.
Latham, A. D. M., M. C. Latham, N. A. Mccutchen, and S. Boutin. 2011. “Invading White‐Tailed Deer Change Wolf‐Caribou Dynamics in Northeastern Alberta.” Journal of Wildlife Management 75: 204–212.
Lima, S. L., and L. M. Dill. 1990. “Behavioral Decisions Made under the Risk of Predation: A Review and Prospectus.” Canadian Journal of Zoology 68: 619–640.
Manly, B. F. J., L. L. McDonald, D. L. Thomas, T. L. McDonald, and W. P. Erickson. 2002. Resource Selection by Animals, 2nd ed. Dordrecht: Kluwer Academic Publishers.
Marshall, K. N., A. C. Stier, J. F. Samhouri, R. P. Kelley, and E. J. Ward. 2016. “Conservation Challenges of Predator Recovery.” Conservation Letters 9: 70–78.
McCullough, D. R. 1999. “Density Dependence and Life‐History Strategies of Ungulates.” Journal of Mammalogy 80: 1130–1146.
McShea, W. J. 2012. “Ecology and Management of White‐Tailed Deer in a Changing World: Deer and Eastern Forests.” Annals of the New York Academy of Sciences 1249: 45–56.
Melis, C., B. Jędrzejewska, M. Apollonio, K. A. Bartoń, W. Jędrzejewski, J. D. C. Linnell, I. Kojola, et al. 2009. “Predation Has a Greater Impact in Less Productive Environments: Variation in Roe Deer, (Capreolus capreolus), Population Density across Europe.” Global Ecology and Biogeography 18: 724–734.
Merems, J. L., L. A. Shipley, T. Levi, J. Ruprecht, D. A. Clark, M. J. Wisdom, N. J. Jackson, K. M. Stewart, and R. A. Long. 2020. “Nutritional‐Landscape Models Link Habitat Use to Condition of Mule Deer (Odocoileus hemionus).” Frontiers in Ecology and Evolution 8: 98.
Miller, B. F., D. A. Osborn, W. R. Lance, M. B. Howze, R. J. Warren, and K. V. Miller. 2009. “Butorphanol‐Azaperone‐Medetomidine for Immobilization of Captive White‐Tailed Deer.” Journal of Wildlife Diseases 45: 457–467.
Moen, A. N. 1978. “Seasonal Changes in Heart Rates, Activity, Metabolism, and Forage Intake of White‐Tailed Deer.” Journal of Wildlife Management 42: 715–738.
Montgomery, R. A., R. J. Moll, E. Say‐Sallaz, M. Valeix, and L. R. Prugh. 2019. “A Tendency to Simplify Complex Systems.” Biological Conservation 233: 1–11.
Monzingo, D. S., J. G. Cook, R. C. Cook, J. S. Horne, and L. A. Shipley. 2023. “Influences of Succession and Biogeoclimate on Forage Resources for Elk in Northern Idaho.” Northwest Science 96: 94–116.
Monzingo, D. S., L. A. Shipley, R. C. Cook, and J. G. Cook. 2022. “Factors Influencing Predictions of Understory Vegetation Biomass from Visual Cover Estimates.” Wildlife Society Bulletin 46: e1300.
MTBS Project. 2022. “MTBS Data Access: Fire Level Geospatial Data.” USDA Forest Service/U.S. Geological Survey. https://mtbs.gov/direct-download.
Muff, S., E. B. Nilsen, R. B. O'Hara, and C. R. Nater. 2022. “Rewriting the Results Section in the Language of Evidence.” Trends in Ecology & Evolution 37: 203–210.
Muhly, T. B., C. Semeniuk, A. Massolo, L. Hickman, and M. Musiani. 2011. “Human Activity Helps Prey Win the Predator‐Prey Space Race.” PLoS One 6: e17050.
Murray, D. L. 2006. “On Improving Telemetry‐Based Survival Estimation.” Journal of Wildlife Management 70: 1530–1543.
Neilson, E. W., and S. Boutin. 2017. “Human Disturbance Alters the Predation Rate of Moose in the Athabasca Oil Sands.” Ecosphere 8: e01913.
Newsome, T. M., L. Boitani, G. Chapron, P. Ciucci, C. R. Dickman, J. A. Dellinger, J. V. López‐Bao, et al. 2016. “Food Habits of the World's Grey Wolves.” Mammal Review 46: 255–269.
Northrup, J. M., C. R. Anderson, and G. Wittemyer. 2014. “Effects of Helicopter Capture and Handling on Movement Behavior of Mule Deer.” Journal of Wildlife Management 78: 731–738.
Oregon State University Libraries & Press and Institute for Natural Resources. 2021. “Oregon Explorer.” http://oregonexplorer.info/.
Pierce, B. M., R. T. Bowyer, and V. C. Bleich. 2004. “Habitat Selection by Mule Deer: Forage Benefits or Risk of Predation.” Journal of Wildlife Management 68: 533–541.
Pollock, K. H., S. R. Winterstein, C. M. Bunck, and P. D. Curtis. 1989. “Survival Analysis in Telemetry Studies: The Staggered Entry Design.” Journal of Wildlife Management 53: 7–15.
Popp, J. N., J. Hamr, J. L. Larkin, and F. F. Mallory. 2018. “Black Bear (Ursus americanus) and Wolf (Canis spp.) Summer Diet Composition and Ungulate Prey Selectivity in Ontario, Canada.” Mammal Research 63: 433–441.
Proffitt, K. M., R. Garrott, J. A. Gude, M. Hebblewhite, B. Jimenez, J. T. Paterson, and J. Rotella. 2020. “Integrated Carnivore‐Ungulate Management: A Case Study in West‐Central Montana.” Wildlife Monographs 206: 1–28.
Prugh, L. R., and S. M. Arthur. 2015. “Optimal Predator Management for Mountain Sheep Conservation Depends on the Strength of Mesopredator Release.” Oikos 124: 1241–1250.
Prugh, L. R., C. X. Cunningham, R. M. Windell, B. N. Kertson, T. R. Ganz, S. L. Walker, and A. J. Wirsing. 2023. “Fear of Large Carnivores Amplifies Human‐Caused Mortality for Mesopredators.” Science 380: 754–758.
Prugh, L. R., and K. J. Sivy. 2020. “Enemies with Benefits: Integrating Positive and Negative Interactions among Terrestrial Carnivores.” Ecology Letters 23: 902–918.
Prugh, L. R., C. J. Stoner, C. W. Epps, W. T. Bean, W. J. Ripple, A. S. Laliberte, and J. S. Brashares. 2009. “The Rise of the Mesopredator.” Bioscience 59: 779–791.
R Core Team. 2021. R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing. Vienna: R Foundation for Statistical Computing http://www.r-project.org/.
Raynor, J. L., C. A. Grainger, and D. P. Parker. 2021. “Wolves Make Roadways Safer, Generating Large Economic Returns to Predator Conservation.” Proceedings of the National Academy of Sciences of the United States of America 118(22): e2023251118.
Rearden, S. N., R. G. Anthony, and B. K. Johnson. 2011. “Birth‐Site Selection and Predation Risk of Rocky Mountain Elk.” Journal of Mammalogy 92: 1118–1126.
Rice, C. G. 2016. “Development of a System for Remotely Monitoring Vaginal Implant Transmitters and Fawn Survival.” Wildlife Biology 22: 22–28.
Rowland, M. M., and C. D. Vojta. 2013. A Technical Guide for Monitoring Wildlife Habitat. Washington: Department of Agriculture, Forest Service, Pacific Northwest Research Station https://www.fs.usda.gov/research/treesearch/45213.
Ruth, T. K., and K. Murphy. 2010. “Cougar‐Prey Relationships.” In Cougar: Ecology and Conservation, edited by M. Hornocker and S. Negri, 138–162. Chicago: The University of Chicago Press.
Shuman, R. M., M. J. Cherry, T. N. Simoneaux, E. A. Dutoit, J. C. Kilgo, M. J. Chamberlain, and K. V. Miller. 2017. “Survival of White‐Tailed Deer Neonates in Louisiana.” Journal of Wildlife Management 81: 834–845.
Sikes, R. S., and The Animal Care and Use Committee of the American Society of Mammalogists. 2016. “2016 Guidelines of the American Society of Mammalogists for the Use of Wild Mammals in Research and Education.” Journal of Mammalogy 97: 663–688.
Stonehouse, K. F., C. R. Anderson, Jr., and M. E. Peterson. 2016. Approaches to Field Investigations of Cause‐Specific Mortality in Mule Deer (Odocoileus hemionus). Fort Collins: Colorado Parks and Wildlife.
Therneau, T. 2022. “A Package for Survival Analysis in R.” R Package Version 3.4‐0. https://CRAN.R-project.org/package=survival.
Therneau, T. M., and P. M. Grambsch. 2000. “The Cox Model.” In Modeling Survival Data: Extending the Cox Model. New York: Springer.
Toledo, D. P., L. B. Abbott, and J. E. Herrick. 2008. “Cover Pole Design for Easy Transport, Assembly, and Field Use.” Journal of Wildlife Management 72: 564–567.
Tollefson, T. N., L. A. Shipley, W. L. Myers, and N. Dasgupta. 2011. “Forage Quality's Influence on Mule Deer Fawns.” Journal of Wildlife Management 75: 919–928.
Tollefson, T. N., L. A. Shipley, W. L. Myers, D. H. Keisler, and N. Dasgupta. 2010. “Influence of Summer and Autumn Nutrition on Body Condition and Reproduction in Lactating Mule Deer.” Journal of Wildlife Management 74: 974–986.
Ulappa, A. C., L. A. Shipley, R. C. Cook, J. G. Cook, and M. E. Swanson. 2020. “Silvicultural Herbicides and Forest Succession Influence Understory Vegetation and Nutritional Ecology of Black‐Tailed Deer in Managed Forests.” Forest Ecology and Management 470: 118216.
USDA National Agricultural Statistics Service Cropland Data Layer. 2019. “Published Crop‐Specific Data Layer.” https://nassgeodata.gmu.edu/CropScape.
van deKerk, M., B. R. McMillan, K. R. Hersey, A. Roug, and R. T. Larsen. 2020. “Effect of Net‐Gun Capture on Survival of Mule Deer.” Journal of Wildlife Management 84: 813–820.
Wagoner, S. J., L. A. Shipley, R. C. Cook, and L. Hardesty. 2013. “Spring Cattle Grazing and Mule Deer Nutrition in a Bluebunch Wheatgrass Community.” Journal of Wildlife Management 77: 897–907.
Wang, G., N. T. Hobbs, S. Twombly, R. B. Boone, A. W. Illius, I. J. Gordon, and J. E. Gross. 2009. “Density Dependence in Northern Ungulates: Interactions with Predation and Resources.” Population Ecology 51: 123–132.
Washington Department of Fish and Wildlife. 2014. Livestock Injury and Mortality Investigations: A Reference Guide for WDFW Field Personnel. Olympia: The Washington Department of Fish and Wildlife https://wdfw.wa.gov/publications/01581.
Washington Department of Fish and Wildlife. 2019. Washington Department of Fish and Wildlife Standard Wolf Foot‐Hold Capture, Immobilization, and Handling Procedures. Olympia: Washington Department of Fish and Wildlife.
Washington Department of Fish and Wildlife, Confederated Tribes of the Colville Reservation, Spokane Tribe of Indians, USDA—APHIS Wildlife Services, and U.S. Fish and Wildlife Service. 2022. Washington Gray Wolf Conservation and Management 2021 Annual Report. Ellensburg: Washington Department of Fish and Wildlife https://wdfw.wa.gov/publications/02317.
White, C. G., P. Zager, and M. W. Gratson. 2010. “Influence of Predator Harvest, Biological Factors, and Landscape on Elk Calf Survival in Idaho.” Journal of Wildlife Management 74: 355–369.
White, M., F. F. Knowlton, and W. C. Glazener. 1972. “Effects of Dam‐Newborn Fawn Behavior on Capture and Mortality.” Journal of Wildlife Management 36: 897–906.
Wickstrom, M. L., C. T. Robbins, T. A. Hanley, D. E. Spalinger, and S. M. Parish. 1984. “Food Intake and Foraging Energetics of Elk and Mule Deer.” Journal of Wildlife Management 48: 1285–1301.
Wiles, G. J., and G. E. Hayes. 2011. Wolf Conservation and Management Plan for Washington. Olympia: Washington Department of Fish and Wildlife https://wdfw.wa.gov/publications/00001.
Williams, C. L., K. Blejwas, J. J. Johnston, and M. M. Jaeger. 2003. “A Coyote in Sheep's Clothing: Predator Identification from Saliva.” Wildlife Society Bulletin 31: 926–932.
Wirsing, A. J., M. R. Heithaus, J. S. Brown, B. P. Kotler, and O. J. Schmitz. 2021. “The Context Dependence of Non‐Consumptive Predator Effects.” Ecology Letters 24: 113–129.
Wittmer, H. U., B. N. Mclellan, R. Serrouya, and C. D. Apps. 2007. “Changes in Landscape Composition Influence the Decline of a Threatened Woodland Caribou Population.” Journal of Animal Ecology 76: 568–579.