Red deer allocate vigilance differently in response to spatio-temporal patterns of risk from human hunters and wolves
Nathan J. Proudman
A C D , Marcin Churski A , Jakub W. Bubnicki A , Jan-Åke Nilsson B and Dries P. J. Kuijper A
+ Author Affiliations
- Author Affiliations
A Mammal Research Institute, Polish Academy of Sciences, ul. Stoczek 1 17-230 Białowieża, Poland.
B Department of Biology, Lund University, Sölvegatan 37, 223 62 Lund, Sweden.
C Present address: Oklahoma State University, 008C Agricultural Hall, Stillwater, OK 74078, USA.
D Corresponding author. Email: nathan.j.proudman@okstate.edu
Wildlife Research 48(2) 163-174 https://doi.org/10.1071/WR20059
Submitted: 15 April 2020 Accepted: 25 July 2020 Published: 11 November 2020
Abstract
Context: Ungulate prey can use increased vigilance to reduce their risk of predation, but little is known of the combined and interactive risk effects from humans and wolves in determining ungulate behaviour across time and space. Understanding the interplay between these risk effects is increasingly important, considering the recolonisation of several large carnivores to more human-dominated landscapes in Europe.
Aim: The aim of the present study was to assess the vigilance behaviour expressed by red deer (Cervus elaphus) in response to both humans and wolves in the Polish Białowieża Forest.
Methods: Using a camera-trap transect, the effect of distance to human settlements, hunting season, patterns of space use by wolves (Canis lupus), canopy openness, canopy height, time of day, as well as sex/age of individuals, on the vigilance behaviour observed in red deer was studied using a model-selection approach.
Key results: We did not find a clear effect of patterns of space use by wolves or distance to human settlements on red deer vigilance behaviour at the landscape scale. However, red deer showed increased vigilance during the hunting season and during the day outside of protected areas and reserves, because disturbance from human hunters is highest. Conversely, we also found that red deer were more vigilant at night within more protected areas, which is likely to be explained by the increased activity of wolves because human activity is strictly limited.
Conclusions: Our study showed that vigilance behaviour of red deer in Białowieża Primeval Forest is more driven by human hunting than by the frequency of wolf presence at a landscape scale. This could be explained by the higher temporal and spatial predictability of human hunting activities than wolf risk. We found that patterns of wolf space use, as opposed to the omnipresent fear effects from humans, had only localised effects by increasing vigilance levels during night hours in non-hunting areas of the forest. The reverse was observed outside of protected reserves. Understanding how prey species respond to this new combination of risk from natural predators and humans, is increasingly important in a landscape where human risk is becoming ever more potent and carnivores recolonise.
Keywords: behaviour, ecology, modelling, predator–prey interactions, statistical modelling, wildlife management, stress response, natural resource management, ethology.
References
Altendorf, K. B., Laundré, J. W., González, C. A. L., and Brown, J. S. (2001). Assessing effects of predation risk on foraging behaviour of mule deer. Journal of Mammalogy 82, 430–439.| Assessing effects of predation risk on foraging behaviour of mule deer.Crossref | GoogleScholarGoogle Scholar |
Atickem, A., Loe, L. E., and Stenseth, N. C. (2014). Individual heterogeneity in use of human shields by mountain nyala. Ethology 120, 715–725.
Bednekoff, P. A., and Ritter, R. (1994). Vigilance in Nxai Pan Springbok, Antidorcas marsupialis. Behaviour 129, 1–11.
| Vigilance in Nxai Pan Springbok, Antidorcas marsupialis.Crossref | GoogleScholarGoogle Scholar |
Benhaiem, S., Delon, M., Lourtet, B., Cargnelutti, B., Aulagnier, S., Hewison, A. J. M., Morellet, N., and Verheyden, H. (2008). Hunting increases vigilance levels in roe deer and modifies feeding site selection. Animal Behaviour 76, 611–618.
| Hunting increases vigilance levels in roe deer and modifies feeding site selection.Crossref | GoogleScholarGoogle Scholar |
Berger, J. (2007). Fear, human shields and the redistribution of prey and predators in protected areas. Biology Letters 3, 620–623.
| Fear, human shields and the redistribution of prey and predators in protected areas.Crossref | GoogleScholarGoogle Scholar | 17925272PubMed |
Blanchard, P., and Fritz, H. (2007). Induced or routine vigilance while foraging. Oikos 116, 1603–1608.
| Induced or routine vigilance while foraging.Crossref | GoogleScholarGoogle Scholar |
Bonnot, N., Morellet, N., Verheyden, H., Cargnelutti, B., Lourtet, B., Klein, F., and Hewison, A. J. M. (2013). Habitat use under predation risk: hunting, roads and human dwellings influence the spatial behaviour of roe deer. European Journal of Wildlife Research 59, 185–193.
| Habitat use under predation risk: hunting, roads and human dwellings influence the spatial behaviour of roe deer.Crossref | GoogleScholarGoogle Scholar |
Bonnot, N. C., Couriot, O., Berger, A., Cagnacci, F., Ciuti, S., De Groeve, J., Gehr, B., Heurich, M., Kjellander, P., Kröschel, M., and Morellet, N. (2020). Fear of the dark? Contrasting impacts of humans vs lynx on diel activity of roe deer across Europe. Journal of Animal Ecology 89, 132–145.
| Fear of the dark? Contrasting impacts of humans vs lynx on diel activity of roe deer across Europe.Crossref | GoogleScholarGoogle Scholar | 31799691PubMed |
Brown, J. S., and Kotler, B. P. (2004). Hazardous duty pay and the foraging cost of predation. Ecology Letters 7, 999–1014.
| Hazardous duty pay and the foraging cost of predation.Crossref | GoogleScholarGoogle Scholar |
Bubnicki, J. W., Churski, M., and Kuijper, D. P. J. (2016). Trapper: an open source web‐based application to manage camera trapping projects. Methods in Ecology and Evolution 7, 1209–1216.
| Trapper: an open source web‐based application to manage camera trapping projects.Crossref | GoogleScholarGoogle Scholar |
Bubnicki, J. W., Churski, M., Schmidt, K., Diserens, T., and Kuijper, D. P. J. (2019). Linking spatial patterns of terrestrial herbivore community structure to trophic interactions. eLife 8, e44937.
| Linking spatial patterns of terrestrial herbivore community structure to trophic interactions.Crossref | GoogleScholarGoogle Scholar | 31577225PubMed |
Burger, J., Safina, C., and Gochfeld, M. (2000). Factors affecting vigilance in springbok: importance of vegetative cover, location in herd, and herd size. Acta Ethologica 2, 97–104.
| Factors affecting vigilance in springbok: importance of vegetative cover, location in herd, and herd size.Crossref | GoogleScholarGoogle Scholar |
Burnham, K. P., and Anderson, D. R. (2002). ‘Model Selection and Multimodel Inference: a Practical Information-theoretic Approach.’ (Springer: New York, NY, USA.)
Chapron, G., Kaczensky, P., Linnell, J. D., von Arx, M., Huber, D., Andrén, H., López-Bao, J. V., Adamec, M., Álvares, F., Anders, O., Balčiauskas, L., Balys, V., Bedő, P., Bego, F., Blanco, J. C., Breitenmoser, U., Brøseth, H., Bufka, L., Bunikyte, R., Ciucci, P., Dutsov, A., Engleder, T., Fuxjäger, C., Groff, C., Holmala, K., Hoxha, B., Iliopoulos, Y., Ionescu, O., Jeremić, J., Jerina, K., Kluth, G., Knauer, F., Kojola, I., Kos, I., Krofel, M., Kubala, J., Kunovac, S., Kusak, J., Kutal, M., Liberg, O., Majić, A., Männil, P., Manz, R., Marboutin, E., Marucco, F., Melovski, D., Mersini, K., Mertzanis, Y., Mysłajek, R. W., Nowak, S., Odden, J., Ozolins, J., Palomero, G., Paunović, M., Persson, J., Potočnik, H., Quenette, P. Y., Rauer, G., Reinhardt, I., Rigg, R., Ryser, A., Salvatori, V., Skrbinšek, T., Stojanov, A., Swenson, J. E., Szemethy, L., Trajçe, A., Tsingarska-Sedefcheva, E., Váňa, M., Veeroja, R., Wabakken, P., Wölfl, M., Wölfl, S., Zimmermann, F., Zlatanova, D., and Boitani, L. (2014). Recovery of large carnivores in Europe’s modern human-dominated landscapes. Science 346, 1517–1519.
| Recovery of large carnivores in Europe’s modern human-dominated landscapes.Crossref | GoogleScholarGoogle Scholar | 25525247PubMed |
Childress, M. J., and Lung, M. A. (2003). Predation risk, gender and the group size effect: does elk vigilance depend upon the behaviour of conspecifics? Animal Behaviour 66, 389–398.
| Predation risk, gender and the group size effect: does elk vigilance depend upon the behaviour of conspecifics?Crossref | GoogleScholarGoogle Scholar |
Ciuti, S., Northrup, J. M., Muhly, T. B., Simi, S., Musiani, M., Pitt, J. A., and Boyce, M. S. (2012). Effects of humans on behaviour of wildlife exceed those of natural predators in a landscape of fear. PLoS One 7, e50611.
| Effects of humans on behaviour of wildlife exceed those of natural predators in a landscape of fear.Crossref | GoogleScholarGoogle Scholar | 23226330PubMed |
Clinchy, M., Zanette, L. Y., Roberts, D., Suraci, J. P., Buesching, C. D., Newman, C., and Macdonald, D. W. (2016). Fear of the human ‘super predator’ far exceeds the fear of large carnivores in a model mesocarnivore. Behavioral Ecology 2, 1826–1832.
| Fear of the human ‘super predator’ far exceeds the fear of large carnivores in a model mesocarnivore.Crossref | GoogleScholarGoogle Scholar |
Creel, S., Winnie, J., Maxwell, B., Hamlin, K., and Creel, M. (2005). Elk alter habitat selection as an antipredator response to wolves. Ecology 86, 3387–3397.
| Elk alter habitat selection as an antipredator response to wolves.Crossref | GoogleScholarGoogle Scholar |
Creel, S., Winnie, J. A., Christianson, D., and Liley, S. (2008). Time and space in general models of antipredator response: tests with wolves and elk. Animal Behaviour 76, 1139–1146.
| Time and space in general models of antipredator response: tests with wolves and elk.Crossref | GoogleScholarGoogle Scholar |
Cromsigt, J. P. G. M., Kuijper, D. P. J., Adam, M., Beschta, R. L., Churski, M., Eycott, A., Kerley, G. I. H., Mysterud, A., Schmidt, K., and West, K. (2013). Hunting for fear: innovating management of human–wildlife conflicts. Journal of Applied Ecology 50, 544–549.
| Hunting for fear: innovating management of human–wildlife conflicts.Crossref | GoogleScholarGoogle Scholar |
Cullen, A. C., and Frey, C. H. (1999). ‘Probabilistic Techniques in Exposure Assessment: a Handbook for Dealing with Variability and Uncertainty in Models and Inputs.’ (Eds A. C. Cullen and C. H. Frey.) (Springer: New York, NY, USA.)
Delignette-Muller, M. L., and Dutang, C. (2015). fitdistrplus: an R package for fitting distributions. Journal of Statistical Software 64, 1–34.
| fitdistrplus: an R package for fitting distributions.Crossref | GoogleScholarGoogle Scholar |
Dröge, E., Creel, S., Becker, M. S., and M’soka, J. (2017). Risky times and risky places interact to affect prey behaviour. Nature Ecology & Evolution 1, 1123–1128.
| Risky times and risky places interact to affect prey behaviour.Crossref | GoogleScholarGoogle Scholar |
Eccard, J. A., Meißner, J. K., and Heurich, M. (2017). European roe deer increase vigilance when faced with immediate predation risk by Eurasian lynx. Ethology 123, 30–40.
| European roe deer increase vigilance when faced with immediate predation risk by Eurasian lynx.Crossref | GoogleScholarGoogle Scholar |
Faliński, J. B. (Ed.) (1986). ‘Vegetation Dynamics in Temperate Lowland Primeval Forests: Ecological Studies in Bialowieza Forest.’ (Dr. W. Junk Publishers: Dordrecht, The Netherlands.)
Fischhoff, I. R., Sundaresan, S. R., Cordingley, J., and Rubenstein, D. I. (2007). Habitat use and movements of plains zebra (Equus burchelli) in response to predation danger from lions. Behavioural Ecology 18, 725–729.
| Habitat use and movements of plains zebra (Equus burchelli) in response to predation danger from lions.Crossref | GoogleScholarGoogle Scholar |
Fortin, D., Beyer, H. L., Boyce, M. S., Smith, D. W., Duchesne, T., and Mao, J. S. (2005). Wolves influence elk movements: behaviour shapes a trophic cascade in Yellowstone National Park. Ecology 86, 1320–1330.
| Wolves influence elk movements: behaviour shapes a trophic cascade in Yellowstone National Park.Crossref | GoogleScholarGoogle Scholar |
Garel, M., Cugnasse, J.-M., Maillard, D., Gaillard, J.-M., Hewison, A. J. M., and Dubray, D. (2007). Selective harvesting and habitat loss produce long-term life history changes in a mouflon population. Ecological Applications 17, 1607–1618.
| Selective harvesting and habitat loss produce long-term life history changes in a mouflon population.Crossref | GoogleScholarGoogle Scholar | 17913127PubMed |
Gaynor, K. M., Brown, J. S., Middleton, A. D., Power, M. E., and Brashares, J. S. (2019). Landscapes of fear: spatial patterns of risk perception and response. Trends in Ecology & Evolution 34, 355–368.
| Landscapes of fear: spatial patterns of risk perception and response.Crossref | GoogleScholarGoogle Scholar |
Gervasi, V., Sand, H., Zimmermann, B., Mattisson, J., Wabakken, P., and Linnell, J. D. (2013). Decomposing risk: landscape structure and wolf behavior generate different predation patterns in two sympatric ungulates. Ecological Applications 23, 1722–1734.
| Decomposing risk: landscape structure and wolf behavior generate different predation patterns in two sympatric ungulates.Crossref | GoogleScholarGoogle Scholar | 24261051PubMed |
Gochfeld, M.,, and Burger, J. (1994). Vigilance in African mammals: differences among mothers, other females, and males. Behaviour 131, 153–169.
| Vigilance in African mammals: differences among mothers, other females, and males.Crossref | GoogleScholarGoogle Scholar |
Goldsmith, A. E. (1990). Vigilance behaviour of pronghorns in different habitats. Journal of Mammalogy 71, 460–462.
| Vigilance behaviour of pronghorns in different habitats.Crossref | GoogleScholarGoogle Scholar |
Gulsby, W. D., Cherry, M. J., Johnson, J. T., Conner, L. M., and Miller, K. V. (2018). Behavioural response of white-tailed deer to coyote predation risk. Ecosphere 9, e02141.
| Behavioural response of white-tailed deer to coyote predation risk.Crossref | GoogleScholarGoogle Scholar |
Halofsky, J. S., and Ripple, W. J. (2008). Fine-scale predation risk on elk after wolf reintroduction in Yellowstone National Park, USA. Oecologia 155, 869–877.
| Fine-scale predation risk on elk after wolf reintroduction in Yellowstone National Park, USA.Crossref | GoogleScholarGoogle Scholar | 18224339PubMed |
Hamilton, W. D. (1971). Geometry for the selfish herd. Journal of Theoretical Biology 31, 295–311.
| Geometry for the selfish herd.Crossref | GoogleScholarGoogle Scholar | 5104951PubMed |
Hebblewhite, M., and Merrill, E. H. (2007). Multiscale wolf predation risk for elk: does migration reduce risk? Oecologia 152, 377–387.
| Multiscale wolf predation risk for elk: does migration reduce risk?Crossref | GoogleScholarGoogle Scholar | 17287955PubMed |
Hebblewhite, M., White, C. A., Nietvelt, C. G., McKenzie, J. A., Hurd, T. E., Fryxell, J. M., Bayley, S. E., and Paquet, P. C. (2005a). Human activity mediates a trophic cascade caused by wolves. Ecology 86, 2135–2144.
| Human activity mediates a trophic cascade caused by wolves.Crossref | GoogleScholarGoogle Scholar |
Hebblewhite, M., Merrill, E. H., and McDonald, T. L. (2005b). Spatial decomposition of predation risk using resource selection functions: an example in a wolf–elk predator–prey system. Oikos 111, 101–111.
| Spatial decomposition of predation risk using resource selection functions: an example in a wolf–elk predator–prey system.Crossref | GoogleScholarGoogle Scholar |
Hunter, L. T. B., and Skinner, J. D. (1998). Vigilance behaviour in African ungulates: the role of predation pressure. Behaviour 135, 195–211.
| Vigilance behaviour in African ungulates: the role of predation pressure.Crossref | GoogleScholarGoogle Scholar |
Hurvich, C. M., and Tsai, C.-L. (1989). Regression and time series model selection in small samples. Biometrika 76, 297–307.
| Regression and time series model selection in small samples.Crossref | GoogleScholarGoogle Scholar |
Husseman, J. S., Murray, D. L., Power, G., Mack, C., Wenger, C. R., and Quigley, H. (2003). Assessing differential prey selection patterns between two sympatric large carnivores. Oikos 101, 591–601.
| Assessing differential prey selection patterns between two sympatric large carnivores.Crossref | GoogleScholarGoogle Scholar |
James, G., Witten, D., Hastie, T., and Tibshirani, R. (2013). ‘An Introduction to Statistical Learning.’ (Series Eds G. Casella, S. Fienberg, and I. Olkin.) (Springer: New York, NY, USA.)
Jayakody, S., Sibbald, A. M., Gordon, I. J., and Lambin, X. (2008). Red deer Cervus elephus vigilance behaviour differs with habitat and type of human disturbance. Wildlife Biology 14, 81–91.
| Red deer Cervus elephus vigilance behaviour differs with habitat and type of human disturbance.Crossref | GoogleScholarGoogle Scholar |
Jędrzejewska, B., and Jędrzejewski, W. (1998). Diets and hunting habits of predators. In ‘Predation in Vertebrate Communities, Ecological Studies’. (Eds B. Jędrzejewska, and W. Jędrzejewski.) pp. 180–323. (Springer: Berlin, Germany.)
Jędrzejewska, B., Jędrzejewski, W., Bunevich, A. N., Miłkowski, L., and Krasiński, Z. A. (1997). Factors shaping population densities and increase rates of ungulates in Bialowieza Primeval Forest (Poland and Belarus) in the 19th and 20th centuries. Acta Theriologica 42, 399–451.
| Factors shaping population densities and increase rates of ungulates in Bialowieza Primeval Forest (Poland and Belarus) in the 19th and 20th centuries.Crossref | GoogleScholarGoogle Scholar |
Jędrzejewski, W., Spaedtke, H., Kamler, J. F., Jędrzejewska, B., and Stenkewitz, U. (2006). Group size dynamics of red deer in BiaÅ‚owieża Primeval Forest, Poland. The Journal of Wildlife Management 70, 1054–1059.
| Group size dynamics of red deer in Białowieża Primeval Forest, Poland.Crossref | GoogleScholarGoogle Scholar |
Jędrzejewski, W., Niedziałkowska, M., Hayward, M. W., Goszczyński, J., Jędrzejewska, B., Borowik, T., Bartoń, K. A., Nowak, S., Harmuszkiewicz, J., Juszczyk, A., Kałamarz, T., Kloch, A., Koniuch, J., Kotiuk, K., Mysłajek, R. W., Nędzyńska, M., Olczyk, A., Teleon, M., and Wojtulewicz, M. (2012). Prey choice and diet of wolves related to ungulate communities and wolf subpopulations in Poland. Journal of Mammalogy 93, 1480–1492.
| Prey choice and diet of wolves related to ungulate communities and wolf subpopulations in Poland.Crossref | GoogleScholarGoogle Scholar |
Kamler, J. F., Jędrzejewski, W., and Jędrzejewska, B. (2007a). Survival and cause-specific mortality of red deer Cervus Elaphus in Białowieża National Park, Poland. Wildlife Biology 13, 48–52.
| Survival and cause-specific mortality of red deer Cervus Elaphus in Białowieża National Park, Poland.Crossref | GoogleScholarGoogle Scholar |
Kamler, J. F., Jędrzejewska, B., and Jędrzejewski, W. (2007b). Activity patterns of red deer in Białowieża National Park, Poland. Journal of Mammalogy 88, 508–514.
| Activity patterns of red deer in Białowieża National Park, Poland.Crossref | GoogleScholarGoogle Scholar |
Kohl, M. T., Stahler, D. R., Metz, M. C., Forester, J. D., Kauffman, M. J., Varley, N., White, P. J., Smith, D. W., and MacNulty, D. R. (2018). Diel predator activity drives a dynamic landscape of fear. Ecological Monographs 88, 638–652.
| Diel predator activity drives a dynamic landscape of fear.Crossref | GoogleScholarGoogle Scholar |
Kuijper, D. P. J., de Kleine, C., Churski, M., van Hooft, P., Bubnicki, J., and Jędrzejewska, B. (2013). Landscape of fear in Europe: wolves affect spatial patterns of ungulate browsing in Białowieża Primeval Forest, Poland. Ecography 36, 1263–1275.
| Landscape of fear in Europe: wolves affect spatial patterns of ungulate browsing in Białowieża Primeval Forest, Poland.Crossref | GoogleScholarGoogle Scholar |
Kuijper, D. P. J., Bubnicki, J. W., Churski, M., Mols, B., and van Hooft, P. (2015). Context dependence of risk effects: wolves and tree logs create patches of fear in an old-growth forest. Behavioural Ecology 26, 1558–1568.
| Context dependence of risk effects: wolves and tree logs create patches of fear in an old-growth forest.Crossref | GoogleScholarGoogle Scholar |
Kuijper, D. P. J., Sahlén, E., Elmhagen, B., Chamaillé-Jammes, S., Sand, H., Lone, K., and Cromsigt, J. P. G. M. (2016). Paws without claws? Ecological effects of large carnivores in anthropogenic landscapes. Proceedings. Biological Sciences 283, 20161625.
| Paws without claws? Ecological effects of large carnivores in anthropogenic landscapes.Crossref | GoogleScholarGoogle Scholar |
Lashley, M. A., Chitwood, M. C., Biggerstaff, M. T., Morina, D. L., Moorman, C. E., and DePerno, C. S. (2014). White-tailed deer vigilance: the influence of social and environmental factors. PLoS One 9, e90652.
| White-tailed deer vigilance: the influence of social and environmental factors.Crossref | GoogleScholarGoogle Scholar | 24819253PubMed |
Laundré, J. W., Hernández, L., and Altendorf, K. B. (2001). Wolves, elk, and bison: reestablishing the ‘landscape of fear’ in Yellowstone National Park, U.S.A. Canadian Journal of Zoology 79, 1401–1409.
Liley, S., and Creel, S. (2008). What best explains vigilance in elk: characteristics of prey, predators, or the environment? Behavioural Ecology 19, 245–254.
| What best explains vigilance in elk: characteristics of prey, predators, or the environment?Crossref | GoogleScholarGoogle Scholar |
Lima, S. L. (1995). Back to the basics of anti-predatory vigilance: the group-size effect. Animal Behaviour 49, 11–20.
| Back to the basics of anti-predatory vigilance: the group-size effect.Crossref | GoogleScholarGoogle Scholar |
Lima, S. L. (1998). Nonlethal effects in the ecology of predator–prey interactions. Bioscience 48, 25–34.
| Nonlethal effects in the ecology of predator–prey interactions.Crossref | GoogleScholarGoogle Scholar |
Lima, S. L. (2002). Putting predators back into behavioural predator–prey interactions. Trends in Ecology & Evolution 17, 70–75.
| Putting predators back into behavioural predator–prey interactions.Crossref | GoogleScholarGoogle Scholar |
Lone, K., Loe, L. E., Gobakken, T., Linnell, J. D. C., Odden, J., Remmen, J., and Mysterud, A. (2014). Living and dying in a multi-predator landscape of fear: roe deer are squeezed by contrasting pattern of predation risk imposed by lynx and humans. Oikos 123, 641–651.
Manor, R., and Saltz, D. (2003). Impact of human nuisance disturbance on vigilance and group size of a social ungulate. Ecological Applications 13, 1830–1834.
| Impact of human nuisance disturbance on vigilance and group size of a social ungulate.Crossref | GoogleScholarGoogle Scholar |
McNamara, J. M., and Houston, A. I. (1992). Evolutionarily stable levels of vigilance as a function of group size. Animal Behaviour 43, 641–658.
| Evolutionarily stable levels of vigilance as a function of group size.Crossref | GoogleScholarGoogle Scholar |
Mech, L. D. (2017). Where can wolves live and how can we live with them? Biological Conservation 210, 310–317.
| Where can wolves live and how can we live with them?Crossref | GoogleScholarGoogle Scholar |
Milner, J. M., Bonenfant, C., and Mysterud, A. (2011). Hunting bambi: evaluating the basis for selective harvesting of juveniles. European Journal of Wildlife Research 57, 565–574.
| Hunting bambi: evaluating the basis for selective harvesting of juveniles.Crossref | GoogleScholarGoogle Scholar |
Muhly, T. B., Semeniuk, C., Massolo, A., Hickman, L., and Musiani, M. (2011). Human activity helps prey win the predator–prey space race. PLoS One 6, e17050.
| Human activity helps prey win the predator–prey space race.Crossref | GoogleScholarGoogle Scholar | 21399682PubMed |
Mysterud, A. (1996). Bed-site selection by adult roe deer Capreolus capreolus in southern Norway during summer. Wildlife Biology 2, 101–106.
| Bed-site selection by adult roe deer Capreolus capreolus in southern Norway during summer.Crossref | GoogleScholarGoogle Scholar |
Mysterud, A., and Østbye, E. (1999). Cover as a habitat element for temperate ungulates: effects on habitat selection and demography. Wildlife Society Bulletin (1973–2006) 27, 385–394.
National Oceanic and Atmospheric Administration (NOAA) (2018). NOAA Sunrise/Sunset Calculator. Available at https://www.esrl.noaa.gov/gmd/grad/solcalc/sunrise.html [verified 1 February 2018].
Neteler, M., Bowman, M. H., Landa, M., and Metz, M. (2012). GRASS GIS: a multi-purpose open source GIS. Environmental Modelling & Software 31, 124–130.
| GRASS GIS: a multi-purpose open source GIS.Crossref | GoogleScholarGoogle Scholar |
Okarma, H. (1995). The trophic ecology of wolves and their predatory role in ungulate communities of forest ecosystems in Europe. Acta Theriologica 40, 335–386.
| The trophic ecology of wolves and their predatory role in ungulate communities of forest ecosystems in Europe.Crossref | GoogleScholarGoogle Scholar |
Okarma, H., Jędrzejewski, W., Schmidt, K., Kowalczyk, R., and Jędrzejewska, B. (1997). Predation of Eurasian lynx on roe deer and red deer. Acta Ethologica 42, 203–224.
Olsson, O., Wirtberg, J., Andersson, M., and Wirtberg, I. (1997). Wolf Canis lupus predation on moose Alces alces and roe deer Capreolus capreolus in south-central Scandinavia. Wildlife Biology 3, 13–25.
| Wolf Canis lupus predation on moose Alces alces and roe deer Capreolus capreolus in south-central Scandinavia.Crossref | GoogleScholarGoogle Scholar |
Padié, S., Morellet, N., Hewison, A. J. M., Martin, J.-L., Bonnot, N., Cargnelutti, B., and Chamaillé‐Jammes, S. (2015). Roe deer at risk: teasing apart habitat selection and landscape constraints in risk exposure at multiple scales. Oikos 124, 1536–1546.
| Roe deer at risk: teasing apart habitat selection and landscape constraints in risk exposure at multiple scales.Crossref | GoogleScholarGoogle Scholar |
Preisser, E. L., Bolnick, D. I., and Benard, M. F. (2005). Scared to death? The effects of intimidation and consumption in predator–prey interactions. Ecology 86, 501–509.
| Scared to death? The effects of intimidation and consumption in predator–prey interactions.Crossref | GoogleScholarGoogle Scholar |
Proffitt, K. M., Grigg, J. L., Hamlin, K. L., and Garrott, R. A. (2009). Contrasting effects of wolves and human hunters on elk behavioural responses to predation risk. The Journal of Wildlife Management 73, 345–356.
| Contrasting effects of wolves and human hunters on elk behavioural responses to predation risk.Crossref | GoogleScholarGoogle Scholar |
Proudman, N. J. (2018). A landscape of fear: behavioural responses in red deer (Cervus elaphus) to risk effects posed by wolves (Canis lupus) and human hunters in a European primeval forest. M.Sc. Thesis, Lund University, Lund, Sweden.
R Core Team (2019). ‘R: a Language and Environment for Statistical Computing.’ (R Foundation for Statistical Computing: Vienna, Austria.) Available at https://www.R-project.org/ [verified 31 August 2020].
Rigby, R. A., and Stasinopoulos, D. M. (2001). The GAMLSS project: a flexible approach to statistical modelling. In ‘New Trends in Statistical Modelling: Proceedings of the 16th International Workshop on Statistical Modelling’. (Eds B. Klein, and L. Korsholm.) pp. 249–256. (University of Southern Denmark: Odense, Denmark.)
Rigby, R. A., and Stasinopoulos, D. M. (2005). Generalized additive models for location, scale and shape (with discussion). Journal of the Royal Statistical Society. Series C, Applied Statistics 54, 507–554.
| Generalized additive models for location, scale and shape (with discussion).Crossref | GoogleScholarGoogle Scholar |
Ripple, W. J., and Beschta, R. L. (2007). Restoring Yellowstone’s aspen with wolves. Biological Conservation 138, 514–519.
| Restoring Yellowstone’s aspen with wolves.Crossref | GoogleScholarGoogle Scholar |
Rogala, J. K., Hebblewhite, M., Whittington, J., White, C. A., Coleshill, J., and Musiani, M. (2011). Human activity differentially redistributes large mammals in the Canadian Rockies National Parks. Ecology and Society 16, 16.
| Human activity differentially redistributes large mammals in the Canadian Rockies National Parks.Crossref | GoogleScholarGoogle Scholar |
Sand, H., Wabakken, P., Zimmermann, B., Johansson, Ö., Pedersen, H. C., and Liberg, O. (2008). Summer kill rates and predation pattern in a wolf–moose system: can we rely on winter estimates? Oecologia 156, 53–64.
| Summer kill rates and predation pattern in a wolf–moose system: can we rely on winter estimates?Crossref | GoogleScholarGoogle Scholar | 18270746PubMed |
Schmidt, K. (2008). Factors shaping the Eurasian lynx (Lynx lynx) population in the north-eastern Poland. Nature Conservation 65, 3–15.
Schmidt, K., Jędrzejewski, W., Okarma, H., and Kowalczyk, R. (2009). Spatial interactions between grey wolves and Eurasian lynx in Białowieża Primeval Forest, Poland. Ecological Research 24, 207–214.
| Spatial interactions between grey wolves and Eurasian lynx in Białowieża Primeval Forest, Poland.Crossref | GoogleScholarGoogle Scholar |
Schuttler, S. G., Parsons, A. W., Forrester, T. D., Baker, M. C., McShea, W. J., Costello, R., and Kays, R. (2017). Deer on the lookout: how hunting, hiking and coyotes affect white‐tailed deer vigilance. Journal of Zoology 301, 320–327.
| Deer on the lookout: how hunting, hiking and coyotes affect white‐tailed deer vigilance.Crossref | GoogleScholarGoogle Scholar |
Shannon, G., Cordes, L. S., Hardy, A. R., Angeloni, L. M., and Crooks, K. R. (2014). Behavioral responses associated with a human-mediated predator shelter. PLoS One 9, e94630.
| Behavioral responses associated with a human-mediated predator shelter.Crossref | GoogleScholarGoogle Scholar | 24718624PubMed |
Sönnichsen, L., Bokje, M., Marchal, J., Hofer, H., Jędrzejewska, B., Kramer-Schadt, S., and Ortmann, S. (2013). Behavioural responses of European roe deer to temporal variation in predation risk. Ethology 119, 233–243.
| Behavioural responses of European roe deer to temporal variation in predation risk.Crossref | GoogleScholarGoogle Scholar |
Stankowich, T. (2008). Ungulate flight responses to human disturbance: a review and meta-analysis. Biological Conservation 141, 2159–2173.
| Ungulate flight responses to human disturbance: a review and meta-analysis.Crossref | GoogleScholarGoogle Scholar |
Theuerkauf, J., Jȩdrzejewski, W., Schmidt, K., and Gula, R. (2003a). Spatiotemporal segregation of wolves from humans in the Białowieża Forest (Poland). The Journal of Wildlife Management 67, 706–716.
| Spatiotemporal segregation of wolves from humans in the Białowieża Forest (Poland).Crossref | GoogleScholarGoogle Scholar |
Theuerkauf, J., Jȩdrzejewski , W., Schmidt, K., Okarma, H., Ruczyński, I., Śniezko, S., and Gula, R. (2003b). Daily patterns and duration of wolf activity in the Białowieza Forest, Poland. Journal of Mammalogy 84, 243–253.
| Daily patterns and duration of wolf activity in the Białowieza Forest, Poland.Crossref | GoogleScholarGoogle Scholar |
van Doormaal, N., Ohashi, H., Koike, S., and Kaji, K. (2015). Influence of human activities on the activity patterns of Japanese sika deer (Cervus nippon) and wild boar (Sus scrofa) in central Japan. European Journal of Wildlife Research 61, 517–527.
| Influence of human activities on the activity patterns of Japanese sika deer (Cervus nippon) and wild boar (Sus scrofa) in central Japan.Crossref | GoogleScholarGoogle Scholar |
Werner, E. E., and Peacor, S. D. (2003). A review of trait-mediated indirect interactions in ecological communities. Ecology 84, 1083–1100.
| A review of trait-mediated indirect interactions in ecological communities.Crossref | GoogleScholarGoogle Scholar |
Winnie, J., and Creel, S. (2007). Sex-specific behavioural responses of elk to spatial and temporal variation in the threat of wolf predation. Animal Behaviour 73, 215–225.
| Sex-specific behavioural responses of elk to spatial and temporal variation in the threat of wolf predation.Crossref | GoogleScholarGoogle Scholar |
Zbyryt, A., Bubnicki, J. W., Kuijper, D. P. J., Dehnhard, M., Churski, M., and Schmidt, K. (2018). Do wild ungulates experience higher stress with humans than with large carnivores? Behavioral Ecology 29, 19–30.
| Do wild ungulates experience higher stress with humans than with large carnivores?Crossref | GoogleScholarGoogle Scholar |
