Human activity and associated landscape modifications alter the movements of animals with consequences for populations and ecosystems worldwide. Species performing long-distance movements are thought to be particularly sensitive to human impact. Despite the increasing anthropogenic pressure, it remains challenging to understand and predict animals' responses to human activity. Here we address this knowledge gap using 1206 Global Positioning System movement trajectories of 815 individuals from 14 red deer (Cervus elaphus) and 14 elk (Cervus canadensis) populations spanning wide environmental gradients, namely the latitudinal range from the Alps to Scandinavia in Europe, and the Greater Yellowstone Ecosystem in North America. We measured individual-level movements relative to the environmental context, or movement expression, using the standardized metric Intensity of Use, reflecting both the directionality and extent of movements. We expected movement expression to be affected by resource (Normalized Difference Vegetation Index, NDVI) predictability and topography, but those factors to be superseded by human impact. Red deer and elk movement expression varied along a continuum, from highly segmented trajectories over relatively small areas (high intensity of use), to directed transitions through restricted corridors (low intensity of use). Human activity (Human Footprint Index, HFI) was the strongest driver of movement expression, with a steep increase in Intensity of Use as HFI increased, but only until a threshold was reached. After exceeding this level of impact, the Intensity of Use remained unchanged. These results indicate the overall sensitivity of Cervus movement expression to human activity and suggest a limitation of plastic responses under high human pressure, despite the species also occurring in human-dominated landscapes. Our work represents the first comparison of metric-based movement expression across widely distributed populations of a deer genus, contributing to the understanding and prediction of animals' responses to human activity.

Animal Ecology Unit Research and Innovation Centre Fondazione Edmund Mach San Michele all'Adige Italy

Chair of Wildlife Ecology and Management Albert Ludwigs University Freiburg Freiburg Germany

Department for Conservation Biodiversity and Wildlife Management Bavarian State Institute of Forestry Freising Germany

Department of Biology and Biotechnologies Charles Darwin University of Rome La Sapienza Rome Italy

Department of Biosciences Centre for Ecological and Evolutionary Synthesis University of Oslo Oslo Norway

Department of Civil Environmental and Mechanical Engineering University of Trento Trento Italy

Department of Ecoscience Wildlife Ecology Aarhus University Aarhus Denmark

Department of Environmental Science Policy and Management University of California California Berkeley USA

Department of Nature Management University of Applied Sciences of Western Switzerland Jussy Switzerland

Department of Visitor Management and National Park Monitoring Bavarian Forest National Park Grafenau Germany

Haub School of Environment and Natural Resources University of Wyoming Wyoming Laramie USA

Idaho Department of Fish and Game Idaho Boise USA

Idaho Department of Fish and Game Idaho Coeur d'Alene USA

Inland Norway University of Applied Science Institute for Forest and Wildlife Management Koppang Norway

Institute for Biodiversity and Ecosystem Dynamics University of Amsterdam Amsterdam The Netherlands

LTSER ZA PYRénées GARonne Auzeville Tolosane France

Montana Department of Fish Wildlife and Parks Montana Bozeman USA

Montana Department of Fish Wildlife and Parks Montana Helena USA

National Park Service Grand Teton National Park Wyoming Moose USA

Natural and Agricultural Environmental Studies Department Service Public de Wallonie Gembloux Belgium

NBFC National Biodiversity Future Center Palermo Italy

Office Français de la Biodiversité DRAS Montfort Birieux France

School of Business Innovation and Sustainability Halmstad University Halmstad Sweden

School of Computing University of Wyoming Wyoming Laramie USA

Stelvio National Park Ersaf Lombardia Bormio Italy

Šumava National Park Vimperk Czech Republic

U S Geological Survey Wyoming Cooperative Fish and Wildlife Research Unit Department of Zoology and Physiology University of Wyoming Wyoming Laramie USA

Université de Toulouse INRAE CEFS Castanet Tolosan France

US Fish and Wildlife Service National Elk Refuge Wyoming Jackson USA

Wildlife Sciences Faculty of Forest Sciences and Forest Ecology University of Goettingen Goettingen Germany

Wyoming Game and Fish Department Wyoming Cody USA

Wyoming Game and Fish Department Wyoming Jackson USA

Comment In

Glob Chang Biol. 2023 Oct;29(20):5741-5743. doi: 10.1111/gcb.16893. PubMed

See more in PubMed

Aikens, E. O., Mysterud, A., Merkle, J. A., Cagnacci, F., Rivrud, I. M., Hebblewhite, M., Hurley, M. A., Peters, W., Bergen, S., De Groeve, J., Dwinnell, S. P. H., Gehr, B., Heurich, M., Hewison, A. J. M., Jarnemo, A., Kjellander, P., Kröschel, M., Licoppe, A., Linnell, J. D. C., … Kauffman, M. J. (2020). Wave-like patterns of plant phenology determine ungulate movement tactics. Current Biology, 30(17), 3444-3449.e4. https://doi.org/10.1016/j.cub.2020.06.032

Aikens, E. O., Wyckoff, T. B., Sawyer, H., & Kauffman, M. J. (2022). Industrial energy development decouples ungulate migration from the green wave. Nature Ecology & Evolution, 1-9, 1733-1741. https://doi.org/10.1038/s41559-022-01887-9

Almeida, P. J. A. L., Vieira, M. V., Kajin, M., Forero-Medina, G., & Cerqueira, R. (2010). Indices of movement behaviour: Conceptual background, effects of scale and location errors. Zoologia (Curitiba), 27(5), 674-680. https://doi.org/10.1590/S1984-46702010000500002

Armstrong, J. B., Takimoto, G. T., Schindler, D. E., Hayes, M. M., & Kauffman, M. J. (2016). Resource waves: Phenological diversity enhances foraging opportunities for mobile consumers. Ecology. 97, 1099-1112. https://doi.org/10.1890/15-0554

Ball, A., John, P., Ball, J. P., Nordengren, C., & Wallin, K. (2001). Partial migration by large ungulates: Characteristics of seasonal moose Alces Aalces ranges in Northern Sweden partial migration by large ungulates: Characteristics of seasonal moose Alces Ranges in Northern Sweden. Academia, 7(3), 39-47. https://doi.org/10.2981/wlb.2001.007

Barker, K. J., Mitchell, M. S., Proffitt, K. M., & DeVoe, J. D. (2019). Land management alters traditional nutritional benefits of migration for elk. Journal of Wildlife Management, 83(1), 167-174. https://doi.org/10.1002/jwmg.21564

Bates, D., Mächler, M., Bolker, B., & Walker, S. (2015). Fitting linear mixed-effects models using lme4. Journal of Statistical Software, 67(1), 1-48. https://doi.org/10.18637/jss.v067.i01

Bauer, S., & Hoye, B. J. (2014). Migratory animals couple biodiversity and ecosystem functioning worldwide. Science, 344(6179), 1242552. https://doi.org/10.1126/science.1242552

Benítez-López, A., Alkemade, R., & Verweij, P. A. (2010). The impacts of roads and other infrastructure on mammal and bird populations: A meta-analysis. Biological Conservation, 143(6), 1307-1316. https://doi.org/10.1016/j.biocon.2010.02.009

Berger, J. (2007). Fear, human shields and the redistribution of prey and predators in protected areas. Biology Letters, 3(6), 620-623. https://doi.org/10.1098/rsbl.2007.0415

Beyer, H. L., Gurarie, E., Börger, L., Panzacchi, M., Basille, M., Herfindal, I., Van Moorter, B., Lele, S. R., & Matthiopoulos, J. (2016). “You shall not pass!”: Quantifying barrier permeability and proximity avoidance by animals. Journal of Animal Ecology, 85(1), 43-53. https://doi.org/10.1111/1365-2656.12275

Bischof, R., Loe, L. E., Meisingset, E. L., Zimmermann, B., van Moorter, B., & Mysterud, A. (2012). A migratory northern ungulate in the pursuit of spring: Jumping or surfing the green wave? American Naturalist, 180(4), 407-424. https://doi.org/10.1086/667590

Bond, N. (2019). Hydrologic indices for daily time series data. Package ‘Hydrostats.’ https://github.com/nickbond/hydrostats

Cagnacci, F. (2023). Memory pays off. Nature Ecology & Evolution, 7(3), 316-317. https://doi.org/10.1038/s41559-022-01968-9

Cagnacci, F., Focardi, S., Ghisla, A., van Moorter, B., Merrill, E. H., Gurarie, E., Heurich, M., Mysterud, A., Linnell, J., Panzacchi, M., May, R., Nygård, T., Rolandsen, C., & Hebblewhite, M. (2016). How many routes lead to migration? Comparison of methods to assess and characterize migratory movements. Journal of Animal Ecology, 85(1), 54-68. https://doi.org/10.1111/1365-2656.12449

Cagnacci, F., Focardi, S., Heurich, M., Stache, A., Hewison, A. J. M., Morellet, N., Kjellander, P., Linnell, J. D. C., Mysterud, A., Neteler, M., Delucchi, L., Ossi, F., & Urbano, F. (2011). Partial migration in roe deer: Migratory and resident tactics are end points of a behavioural gradient determined by ecological factors. Oikos, 120(12), 1790-1802. https://doi.org/10.1111/j.1600-0706.2011.19441.x

Calenge, C. (2006). The package adehabitat for the R software: A tool for the analysis of space and habitat use by animals. Ecological Modelling, 197, 516-519.

Colwell, R. K. (1974). Predictability, constancy, and contingency of periodic phenomena. Ecology, 55(5), 1148-1153. https://doi.org/10.2307/1940366

Corradini, A., Randles, M., Pedrotti, L., van Loon, E., Passoni, G., Oberosler, V., Rovero, F., Tattoni, C., Ciolli, M., & Cagnacci, F. (2021). Effects of cumulated outdoor activity on wildlife habitat use. Biological Conservation, 253, 108818. https://doi.org/10.1016/j.biocon.2020.108818

Couriot, O., Hewison, A. J. M., Saïd, S., Cagnacci, F., Chamaillé-Jammes, S., Linnell, J. D. C., Mysterud, A., Peters, W., Urbano, F., Heurich, M., Kjellander, P., Nicoloso, S., Berger, A., Sustr, P., Kroeschel, M., Soennichsen, L., Sandfort, R., Gehr, B., & Morellet, N. (2018). Truly sedentary? The multi-range tactic as a response to resource heterogeneity and unpredictability in a large herbivore. Oecologia, 187(1), 47-60. https://doi.org/10.1007/s00442-018-4131-5

Creel, S., Merkle, J., Mweetwa, T., Becker, M. S., Mwape, H., Simpamba, T., & Simukonda, C. (2020). Hidden Markov models reveal a clear human footprint on the movements of highly mobile African wild dogs. Scientific Reports, 10(1), 17908. https://doi.org/10.1038/s41598-020-74329-w

D'Amico, M., Périquet, S., Román, J., & Revilla, E. (2016). Road avoidance responses determine the impact of heterogeneous road networks at a regional scale. Journal of Applied Ecology, 53(1), 181-190. https://doi.org/10.1111/1365-2664.12572

Doherty, T. S., Hays, G. C., & Driscoll, D. A. (2021). Human disturbance causes widespread disruption of animal movement. Nature Ecology & Evolution, 5, 513-519. https://doi.org/10.1038/s41559-020-01380-1

Dormann, C. F., Elith, J., Bacher, S., Buchmann, C., Carl, G., Carré, G., Marquéz, J. R. G., Gruber, B., Lafourcade, B., Leitão, P. J., Münkemüller, T., McClean, C., Osborne, P. E., Reineking, B., Schröder, B., Skidmore, A. K., Zurell, D., & Lautenbach, S. (2012). Collinearity: A review of methods to deal with it and a simulation study evaluating their performance. Ecography, 36(1), 27-46. https://doi.org/10.1111/j.1600-0587.2012.07348.x

Eggeman, S. L., Hebblewhite, M., Bohm, H., Whittington, J., & Merrill, E. H. (2016). Behavioural flexibility in migratory behaviour in a long-lived large herbivore. Journal of Animal Ecology, 85(3), 785-797. https://doi.org/10.1111/1365-2656.12495

English, A. K., Chauvenet, A. L. M., Safi, K., & Pettorelli, N. (2012). Reassessing the determinants of breeding synchrony in ungulates. PLoS One, 7(7), 1-7. https://doi.org/10.1371/journal.pone.0041444

European Environment Agency (EEA). (2020). European Digital Elevation Model (EU-DEM), version 1.1.

Fahrig, L. (2007). Non-optimal animal movement in human-altered landscapes. Functional Ecology, 21(6), 1003-1015. https://doi.org/10.1111/j.1365-2435.2007.01326.x

Fong, Y., Huang, Y., Gilbert, P. B., & Permar, S. R. (2017). Chngpt: Threshold regression model estimation and inference. BMC Bioinformatics, 18(1), 1-7. https://doi.org/10.1186/s12859-017-1863-x

Harris, G., Thirgood, S., Hopcraft, J., Cromsight, J., & Berger, J. (2009). Global decline in aggregated migrations of large terrestrial mammals. Endangered Species Research, 7(1), 55-76. https://doi.org/10.3354/esr00173

Hijmans, R. J. (2023). Geographic data analysis and modeling. https://cran.r-project.org/package=raster

Hu, P., Shao, Y., Xu, J., Wang, T., Li, Y., Liu, H., Rong, M., Su, W., Chen, B., Cui, S., Cui, X., Yang, F., Tamate, H., & Xing, X. (2019). Genome-wide study on genetic diversity and phylogeny of five species in the genus Cervus. BMC Genomics, 20(1), 1-13. https://doi.org/10.1186/s12864-019-5785-z

Ito, T. Y., Lhagvasuren, B., Tsunekawa, A., Shinoda, M., Takatsuki, S., Buuveibaatar, B., & Chimeddorj, B. (2013). Fragmentation of the habitat of wild ungulates by anthropogenic barriers in Mongolia. PLoS One, 8(2), e56995. https://doi.org/10.1371/journal.pone.0056995

Jesmer, B. R., Merkle, J. A., Goheen, J. R., Aikens, E. O., Beck, J. L., Courtemanch, A. B., Hurley, M. A., McWhirter, D. E., Miyasaki, H. M., Monteith, K. L., & Kauffman, M. J. (2018). Is ungulate migration culturally transmitted? Evidence of social learning from translocated animals. Science, 361(6406), 1023-1025. https://doi.org/10.1126/science.aat0985

Johnson, P. C. D. (2014). Extension of Nakagawa & Schielzeth's R2 GLMM to random slopes models. Methods in Ecology and Evolution, 5(9), 944-946. https://doi.org/10.1111/2041-210X.12225

Joly, K., Gurarie, E., Sorum, M. S., Kaczensky, P., Cameron, M. D., Jakes, A. F., Borg, B. L., Nandintsetseg, D., Hopcraft, J. G. C., Buuveibaatar, B., Jones, P. F., Mueller, T., Walzer, C., Olson, K. A., Payne, J. C., Yadamsuren, A., & Hebblewhite, M. (2019). Longest terrestrial migrations and movements around the world. Scientific Reports, 9(1), 15333. https://doi.org/10.1038/s41598-019-51884-5

Jones, J. D., Kauffman, M. J., Monteith, K. L., Scurlock, B. M., Albeke, S. E., & Cross, P. C. (2014). Supplemental feeding alters migration of a temperate ungulate. Ecological Applications, 24(7), 1769-1779. https://doi.org/10.1890/13-2092.1

Kauffman, M. J., Cagnacci, F., Chamaillé-Jammes, S., Hebblewhite, M., Hopcraft, J. G. C., Merkle, J. A., Mueller, T., Mysterud, A., Peters, W., Roettger, C., Steingisser, A., Meacham, J. E., Abera, K., Adamczewski, J., Aikens, E. O., Bartlam-Brooks, H., Bennitt, E., Berger, J., Boyd, C., … Zuther, S. (2021). Mapping out a future for ungulate migrations. Science, 372(6542), 566-569. https://doi.org/10.1126/science.abf0998

Koch, A., & Heipke, C. (2001). Quality assessment of digital surface models derived from the shuttle radar topography Mission (SRTM). IGARSS 2001. Scanning the present and resolving the future. Proceedings. IEEE 2001 International Geoscience and Remote Sensing Symposium (cat. no. 01CH37217). 6 (January 2000), 2863-2865. https://doi.org/10.1109/IGARSS.2001.978187

Linnell, J. D. C., Trouwborst, A., Boitani, L., Kaczensky, P., Huber, D., Reljic, S., Kusak, J., Majic, A., Skrbinsek, T., Potocnik, H., Hayward, M. W., Milner-Gulland, E. J., Buuveibaatar, B., Olson, K. A., Badamjav, L., Bischof, R., Zuther, S., & Breitenmoser, U. (2016). Border security fencing and wildlife: The end of the transboundary paradigm in Eurasia? PLoS Biology, 14(6), e1002483. https://doi.org/10.1371/journal.pbio.1002483

Long, J. A. (2020). Jtools: Analysis and presentation of social scientific data. https://cran.r-project.org/package=jtools

Main, M. T., Davis, R. A., Blake, D., Mills, H., & Doherty, T. S. (2020). Human impact overrides bioclimatic drivers of red fox home range size globally. Diversity and Distributions, 26(9), 1083-1092. https://doi.org/10.1111/ddi.13115

Manly, B. F. J., McDonald, L. L., Thomas, D. L., McDonald, T. L., & Erickson, W. P. (2002). Resource selection by animals-statistical design and analysis for field studies.

Martin, J., Tolon, V., Morellet, N., Santin-Janin, H., Licoppe, A., Fischer, C., Bombois, J., Patthey, P., Pesenti, E., Chenesseau, D., & Saïd, S. (2018). Common drivers of seasonal movements on the migration-Residency behavior continuum in a large herbivore. Scientific Reports, 8(1), 7631. https://doi.org/10.1038/s41598-018-25777-y

McInturff, A., Xu, W., Wilkinson, C. E., Dejid, N., & Brashares, J. S. (2020). Fence ecology: Frameworks for understanding the ecological effects of fences. Bioscience, 70(11), 971-985. https://doi.org/10.1093/biosci/biaa103

Meisingset, E. L., Loe, L. E., Brekkum, Ø., Van Moorter, B., & Mysterud, A. (2013). Red deer habitat selection and movements in relation to roads. The Journal of Wildlife Management, 77(1), 181-191. https://doi.org/10.1002/jwmg.469

Mendgen, P., Dejid, N., Olson, K., Buuveibaatar, B., Calabrese, J. M., Chimeddorj, B., Dalannast, M., Fagan, W. F., Leimgruber, P., & Müller, T. (2023). Nomadic ungulate movements under threat: Declining mobility of Mongolian Gazelles in the Eastern Steppe. https://doi.org/10.1101/2023.02.05.526430

Merkle, J. A., Monteith, K. L., Aikens, E. O., Hayes, M. M., Hersey, K. R., Middleton, A. D., Oates, B. A., Sawyer, H., Scurlock, B. M., & Kauffman, M. J. (2016). Large herbivores surf waves of green-up during spring. Proceedings of the Royal Society B: Biological Sciences, 283(1833), 20160456. https://doi.org/10.1098/rspb.2016.0456

Middleton, A. D., Kauffman, M. J., Mcwhirter, D. E., Cook, J. G., Cook, R. C., Nelson, A. A., Jimenez, M. D., & Klaver, R. W. (2013). Rejoinder: Challenge and opportunity in the study of ungulate migration amid environmental change. Ecology, 94(6), 1280-1286. https://doi.org/10.1890/12-2142.1

Morelle, K., Lehaire, F., & Lejeune, P. (2013). Spatio-temporal patterns of wildlife-vehicle collisions in a region with a high-density road network. Nature Conservation, 5, 53-73. https://doi.org/10.3897/natureconservation.5.4634

Morrison, T. A., Merkle, J. A., Hopcraft, J. G. C., Aikens, E. O., Beck, J. L., Boone, R. B., Courtemanch, A. B., Dwinnell, S. P., Fairbanks, W. S., Griffith, B., Middleton, A. D., Monteith, K. L., Oates, B., Riotte-Lambert, L., Sawyer, H., Smith, K. T., Stabach, J. A., Taylor, K. L., & Kauffman, M. J. (2021). Drivers of site fidelity in ungulates. Journal of Animal Ecology, 90, 955-966. https://doi.org/10.1111/1365-2656.13425

Mueller, T., Olson, K. A., Dressler, G., Leimgruber, P., Fuller, T. K., Nicolson, C., Novaro, A. J., Bolgeri, M. J., Wattles, D., DeStefano, S., Calabrese, J. M., & Fagan, W. F. (2011). How landscape dynamics link individual- to population-level movement patterns: A multispecies comparison of ungulate relocation data. Global Ecology and Biogeography, 20(5), 683-694. https://doi.org/10.1111/j.1466-8238.2010.00638.x

Mysterud, A., Loe, L., & Zimmermann, B. (2011). Partial migration in expanding red deer populations at northern latitudes-A role for density dependence? Oikos, 120, 1817-1825. https://doi.org/10.1111/j.1600-0706.2010.19439.x

Nandintsetseg, D., Bracis, C., Leimgruber, P., Kaczensky, P., Buuveibaatar, B., Lkhagvasuren, B., Chimeddorj, B., Enkhtuvshin, S., Horning, N., Ito, T. Y., Olson, K., Payne, J., Walzer, C., Shinoda, M., Stabach, J., Songer, M., & Mueller, T. (2019). Variability in nomadism: Environmental gradients modulate the movement behaviors of dryland ungulates. Ecosphere, 10(11), e02924. https://doi.org/10.1002/ecs2.2924

Nathan, R., Getz, W. M., Revilla, E., Holyoak, M., Kadmon, R., Saltz, D., & Smouse, P. E. (2008). A movement ecology paradigm for unifying organismal movement research. Proceedings of the National Academy of Sciences of the United States of America, 105(49), 19052-19059. https://doi.org/10.1073/pnas.0800375105

Nickel, B. A., Suraci, J. P., Allen, M. L., & Wilmers, C. C. (2020). Human presence and human footprint have non-equivalent effects on wildlife spatiotemporal habitat use. Biological Conservation, 241(August 2019), 108383. https://doi.org/10.1016/j.biocon.2019.108383

Olson, K. A., Fuller, T. K., Mueller, T., Murray, M. G., Nicolson, C., Odonkhuu, D., Bolortsetseg, S., & Schaller, G. B. (2010). Annual movements of Mongolian gazelles: Nomads in the eastern steppe. Journal of Arid Environments, 74(11), 1435-1442. https://doi.org/10.1016/j.jaridenv.2010.05.022

Ossi, F., Gaillard, J. M., Hebblewhite, M., Morellet, N., Ranc, N., Sandfort, R., Kroeschel, M., Kjellander, P., Mysterud, A., Linnell, J. D. C., Heurich, M., Soennichsen, L., Sustr, P., Berger, A., Rocca, M., Urbano, F., & Cagnacci, F. (2017). Plastic response by a small cervid to supplemental feeding in winter across a wide environmental gradient. Ecosphere, 8(1). https://doi.org/10.1002/ecs2.1629

Panzacchi, M., Van Moorter, B., Jordhøy, P., & Strand, O. (2013). Learning from the past to predict the future: Using archaeological findings and GPS data to quantify reindeer sensitivity to anthropogenic disturbance in Norway. Landscape Ecology, 28(5), 847-859. https://doi.org/10.1007/s10980-012-9793-5

Panzacchi, M., Van Moorter, B., & Strand, O. (2013). A road in the middle of one of the last wild reindeer migration routes in Norway: Crossing behaviour and threats to conservation. Rangifer, 33, 15-26. https://brage.nina.no/nina-xmlui/handle/11250/2480558

Passoni, G., Coulson, T., Ranc, N., Corradini, A., Hewison, A. J. M., Ciuti, S., Gehr, B., Heurich, M., Brieger, F., Sandfort, R., Mysterud, A., Balkenhol, N., & Cagnacci, F. (2021). Roads constrain movement across behavioural processes in a partially migratory ungulate. Movement Ecology, 9(1), 1-12. https://doi.org/10.1186/s40462-021-00292-4

Peters, W., Hebblewhite, M., Mysterud, A., Eacker, D., Hewison, A. J. M., Linnell, J. D. C., Focardi, S., Urbano, F., De Groeve, J., Gehr, B., Heurich, M., Jarnemo, A., Kjellander, P., Kröschel, M., Morellet, N., Pedrotti, L., Reinecke, H., Sandfort, R., Sönnichsen, L., … Cagnacci, F. (2019). Large herbivore migration plasticity along environmental gradients in Europe: Life-history traits modulate forage effects. Oikos, 128(3), 416-429. https://doi.org/10.1111/oik.05588

Peters, W., Hebblewhite, M., Mysterud, A., Spitz, D., Focardi, S., Urbano, F., Morellet, N., Heurich, M., Kjellander, P., Linnell, J. D. C., & Cagnacci, F. (2017). Migration in geographic and ecological space by a large herbivore. Ecological Monographs, 87(2), 297-320. https://doi.org/10.1002/ecm.1250

Prokopenko, C. M., Boyce, M. S., & Avgar, T. (2017). Extent-dependent habitat selection in a migratory large herbivore: Road avoidance across scales. Landscape Ecology, 32(2), 313-325. https://doi.org/10.1007/s10980-016-0451-1

R Core Team. (2023). R: A language and environment for statistical computing. R Foundation for Statistical Computing. https://www.r-project.org/

Ranc, N., Moorcroft, P. R., Ossi, F., & Cagnacci, F. (2021). Experimental evidence of memory-based foraging decisions in a large wild mammal. Proceedings of the National Academy of Sciences of the United States of America, 118(15), 1-9. https://doi.org/10.1073/pnas.2014856118

Rickbeil, G. J. M., Merkle, J. A., Anderson, G., Atwood, M. P., Beckmann, J. P., Cole, E. K., Courtemanch, A. B., Dewey, S., Gustine, D. D., Kauffman, M. J., McWhirter, D. E., Mong, T., Proffitt, K., White, P. J., & Middleton, A. D. (2019). Plasticity in elk migration timing is a response to changing environmental conditions. Global Change Biology, 25(7), 2368-2381. https://doi.org/10.1111/gcb.14629

Rutz, C., Loretto, M. C., Bates, A. E., Davidson, S. C., Duarte, C. M., Jetz, W., Johnson, M., Kato, A., Kays, R., Mueller, T., Primack, R. B., Ropert-Coudert, Y., Tucker, M. A., Wikelski, M., & Cagnacci, F. (2020). COVID-19 lockdown allows researchers to quantify the effects of human activity on wildlife. Nature Ecology and Evolution, 4, 1156-1159. https://doi.org/10.1038/s41559-020-1237-z

Salvatori, M., Oberosler, V., Rinaldi, M., Franceschini, A., Truschi, S., Pedrini, P., & Rovero, F. (2023). Crowded mountains: Long-term effects of human outdoor recreation on a community of wild mammals monitored with systematic camera trapping. Ambio, 52, 1085-1097. https://doi.org/10.1007/s13280-022-01825-w

Sawyer, H., Korfanta, N. M., Nielson, R. M., Monteith, K. L., & Strickland, D. (2017). Mule deer and energy development-Long-term trends of habituation and abundance. Global Change Biology, 23(11), 4521-4529. https://doi.org/10.1111/gcb.13711

Signer, J., Fieberg, J., & Avgar, T. (2019). Animal movement tools (amt): R package for managing tracking data and conducting habitat selection analyses. Ecology and Evolution, 9(2), 880-890. https://doi.org/10.1002/ece3.4823

Teitelbaum, C. S., Fagan, W. F., Fleming, C. H., Dressler, G., Calabrese, J. M., Leimgruber, P., & Mueller, T. (2015). How far to go? Determinants of migration distance in land mammals. Ecology Letters, 18(6), 545-552. https://doi.org/10.1111/ele.12435

Tucker, M. A., Böhning-Gaese, K., Fagan, W. F., Fryxell, J. M., Van Moorter, B., Alberts, S. C., Ali, A. H., Allen, A. M., Attias, N., Avgar, T., Bartlam-Brooks, H., Bayarbaatar, B., Belant, J. L., Bertassoni, A., Beyer, D., Bidner, L., van Beest, F. M., Blake, S., Blaum, N., … Mueller, T. (2018). Moving in the Anthropocene: Global reductions in terrestrial mammalian movements. Science, 359(6374), 466-469. https://doi.org/10.1126/science.aam9712

Urbano, F., Cagnacci, F., & Euromammals Collaborative Initiative. (2021). Data management and sharing for collaborative science: Lessons learnt from the Euromammals initiative. Frontiers in Ecology and Evolution, 9(September), 1-13. https://doi.org/10.3389/fevo.2021.727023

Venter, O., Sanderson, E. W., Magrach, A., Allan, J. R., Beher, J., Jones, K. R., Possingham, H. P., Laurance, W. F., Wood, P., Fekete, B. M., Levy, M. A., & Watson, J. E. (2018). Last of the Wild Project, Version 3 (LWP-3): 1993 Human Footprint, 2018 Release. https://doi.org/10.7927/H4H9938Z

Venter, O., Sanderson, E. W., Magrach, A., Allan, J. R., Beher, J., Jones, K. R., Possingham, H. P., Laurance, W. F., Wood, P., Fekete, B. M., Levy, M. A., & Watson, J. E. M. (2016). Global terrestrial human footprint maps for 1993 and 2009. Scientific Data, 3(1), 160067. https://doi.org/10.1038/sdata.2016.67

Vuolo, F., Mattiuzzi, M., Klisch, A., & Atzberger, C. (2012). Data service platform for MODIS vegetation indices time series processing at BOKU Vienna: Current status and future perspectives. Earth Resources and Environmental Remote Sensing/GIS Applications III, 8538(October), 85380A. https://doi.org/10.1117/12.974857

Wyckoff, T. B., Sawyer, H., Albeke, S. E., Garman, S. L., & Kauffman, M. J. (2018). Evaluating the influence of energy and residential development on the migratory behavior of mule deer. Ecosphere, 9(2). https://doi.org/10.1002/ecs2.2113

Xu, W., Barker, K., Shawler, A., Van Scoyoc, A., Smith, J. A., Mueller, T., Sawyer, H., Andreozzi, C., Bidder, O. R., Karandikar, H., Mumme, S., Templin, E., & Middleton, A. D. (2021). The plasticity of ungulate migration in a changing world. Ecology, 102, 1-14. https://doi.org/10.1002/ecy.3293

Xu, W., Dejid, N., Herrmann, V., Sawyer, H., & Middleton, A. D. (2021). Barrier behaviour analysis (BaBA) reveals extensive effects of fencing on wide-ranging ungulates. Journal of Applied Ecology, 58, 690-698. https://doi.org/10.1111/1365-2664.13806

Zimmermann, B., Nelson, L., Wabakken, P., Sand, H., & Liberg, O. (2014). Behavioral responses of wolves to roads: Scale-dependent ambivalence. Behavioral Ecology, 25(6), 1353-1364. https://doi.org/10.1093/beheco/aru134

Ecological and intrinsic drivers of foraging parameters of Eurasian lynx at a continental scale