The wolf is a generalist-opportunistic predator that displays diverse and remarkably adaptable feeding strategies across its range with local adaptations to certain prey species depending on their availability and vulnerability. The multi-prey system of the Slovak Carpathians supports important portion of the European wolf population; however, it has been markedly understudied. We evaluated winter diet composition and prey selection of Slovak wolves based on 321 scat samples collected between September-April within four different study areas during 2015-2017. The winter diet of wolves in the Slovak Carpathians was characterized by a 98% occurrence of wild large-sized and medium-sized ungulates with red deer occurring in wolf scats most often, consistent with their highest density among other wild ungulates. However, by comparing the consumption with availability of wild prey, we found that wolves in fact selected for wild boar especially in areas with higher altitudinal range, while selected for red deer in areas with low altitudinal range where this prey species was more spatially predictable. Although wolves showed the potential to switch between red deer and wild boar when their density increases, we found that this variation can be rather linked to changing prey vulnerability, which is dependent on particular environmental conditions at local scale such as topography and snow accumulation. The present study provides valuable insights into the winter foraging ecology of Slovak wolves in a multi-prey system of the Carpathians and allows for practical implications in the management of the rapidly increasing populations of wild ungulates across Europe.

BIOPOLIS Program in Genomics Biodiversity and Land Planning CIBIO Vairão Portugal

CIBIO Centro de Investigação em Biodiversidade e Recursos Genéticos InBIO Laboratório Associado Universidade do Porto Porto Portugal

Department of Biology and Ecology Faculty of Natural Sciences Matej Bel University in Banská Bystrica Banská Bystrica Slovakia

Diana Carpathian Wildlife Research Banská Bystrica Slovakia

Faculty of Ecology and Environmental Studies Technical University in Zvolen Technical University in Zvolen Zvolen Slovakia

Faculty of Forestry Department of Applied Zoology and Wildlife Management Technical University in Zvolen Zvolen Slovakia

Institute of Vertebrate Biology Academy of Sciences of the Czech Republic Brno Czech Republic

Little Fox The Centre of Natural Sciences Research and Environmental Education Staré Hory Slovakia

Muráň Plateau National Park State Nature Conservancy of the Slovak Republic Revúca Slovakia

National Forest Centre Zvolen Slovakia

Poloniny National Park State Nature Conservancy of the Slovakia Republic Stakčín Slovakia

University of Veterinary Sciences Brno Brno Czech Republic

See more in PubMed

Ripple WJ, Estes JA, Beschta RL, Wilmers CC, Ritchie EG, Hebblewhite M, et al.. Status and ecological effects of the world’s largest carnivores. Science. 2014. Jan 10;343(6167). doi: 10.1126/science.1241484 PubMed DOI

Beschta RL, Ripple WJ. Riparian vegetation recovery in Yellowstone: the first two decades after wolf reintroduction. Biol Conserv. 2016. Jun 1;198: 93–103.

Boyce MS. Wolves for Yellowstone: dynamics in time and space. J Mammal. 2018. Oct 10;99(5): 1021–31.

Becker MS, Garrott RA, White PJ, Gower CN, Bergman EJ, Jaffe R. Wolf prey selection in an elk-bison system: choice or circumstance? Terrestrial Ecology. 2008. Jan 1;3: 305–337.

Peterson RO, Ciucci P. The wolf as a carnivore. In: Mech LD, Boitani L, editors. Wolves: behavior, ecology, and conservation. University of Chicago Press; 2003. pp. 104–130.

Sin T, Gazzola A, Chiriac S, Rîșnoveanu G. Wolf diet and prey selection in the South-Eastern Carpathian Mountains, Romania. PloS One. 2019. Nov 21;14(11): e0225424. doi: 10.1371/journal.pone.0225424 PubMed DOI PMC

Zlatanova D, Ahmed A, Valasseva A, Genov P. Adaptive diet strategy of the wolf (Canis lupus L.) in Europe: a review. Acta Zool Bulg. 2014. Dec 1;66(4): 439–452.

Newsome TM, Boitani L, Chapron G, Ciucci P, Dickman CR, Dellinger JA. Food habits of the world’s grey wolves. Mamm Rev. 2016. Oct;46(4): 255–69.

Olsson O, Wirtberg J, Andersson M, Wirtberg I. Wolf Canis lupus predation on moose Alces alces and roe deer Capreolus capreolus in south-central Scandinavia. Wildlife Biol. 1997. Mar;3(3/4): 13–25.

Jędrzejewski W, Niedziałkowska M, Hayward MW, Goszczyński J, Jędrzejewska B, Borowik T, et al.. Prey choice and diet of wolves related to ungulate communities and wolf subpopulations in Poland. J Mammal. 2012. Dec 17;93(6): 1480–1492.

Lagos L, Bárcena F. Spatial variability in wolf diet and prey selection in Galicia (NW Spain). Mamm Res. 2018. Apr;63(2): 125–139.

Petridou M, Youlatos D, Lazarou Y, Selinides K, Pylidis C, Giannakopoulos A, et al.. Wolf diet and livestock selection in central Greece. Mammalia. 2019. Nov 1;83(6): 530–8.

Figueiredo AM, Valente AM, Barros T, Carvalho J, Silva DA, Fonseca C, et al.. What does the wolf eat? Assessing the diet of the endangered Iberian wolf (Canis lupus signatus) in northeast Portugal. PLoS One. 2020. Mar 31;15(3): e0230433. doi: 10.1371/journal.pone.0230433 PubMed DOI PMC

Migli DE, Youlatos D, Iliopoulos YI. Winter food habits of wolves in central Greece. J Biol Res (Thessalon). 2005;4: 217–20.

Pimenta V, Barroso I, Boitani L, Beja P. Wolf predation on cattle in Portugal: Assessing the effects of husbandry systems. Biol Conserv. 2017. Mar 1;207: 17–26.

Kaczensky P, Chapron G, von Arx M, Huber D, Andrén H, et al.. Status, management and distribution of large carnivores–bear, lynx, wolf and wolverine–in Europe. Report to the EU Commission; 2013. pp. 272.

Sillero-Zubiri C, Laurenson MK. Interactions between carnivores and local communities: conflict or co-existence? Conserv Biol Ser—Cambridge. 2001. Jan 1: 282–312.

Graham K, Beckerman AP, Thirgood S. Human–predator–prey conflicts: ecological correlates, prey losses and patterns of management. Biol Conserv. 2005. Mar 1;122(2): 159–171.

Jȩdrzejewski W, Jȩdrzejewska B, Okarma H, Schmidt K, Zub K, Musiani M. Prey selection and predation by wolves in Białowieża Primeval Forest, Poland. J Mammal. 2000. Feb 1;81(1): 197–212.

Freitas J, Lagos L, Álvares F. Horses as prey of wolves: worldwide patterns and management implications. CDPNews. 2021;23: 1–11.

Octenjak D, Pađen L, Šilić V, Reljić S, Vukičević TT, Kusak J. Wolf diet and prey selection in Croatia. Mamm Res. 2020. Oct;65(4): 647–54.

Martins I, Krofel M, Mota PG, Álvares F. Consumption of Carnivores by Wolves: A Worldwide Analysis of Patterns and Drivers. Diversity. 2020. Dec;12(12): 470.

Yearsley JM. Optimal diet selection, frequency dependence and prey renewal. Theor Popul Biol. 2003. Sep 1;64(2): 129–139. doi: 10.1016/s0040-5809(03)00070-4 PubMed DOI

Mattioli L, Capitani C, Gazzola A, Scandura M, Apollonio M. Prey selection and dietary response by wolves in a high-density multi-species ungulate community. Eur J Wildl Res. 2011. Aug;57(4): 909–22.

Sand H, Eklund A, Zimmermann B, Wikenros C, Wabakken P. Prey selection of Scandinavian wolves: single large or several small? PLoS One. 2016. Dec 28;11(12): e0168062. doi: 10.1371/journal.pone.0168062 PubMed DOI PMC

Murdoch WW. Switching in general predators: experiments on predator specificity and stability of prey populations. Ecol Monogr. 1969. Sep;39(4): 335–54.

Oaten A, Murdoch WW. Functional response and stability in predator-prey systems. Am Nat. 1975. May 1;109(967): 289–98.

Fryxell JM, Lundberg P. Diet choice and predator—prey dynamics. Evol Ecol. 1994. Jul;8(4): 407–21.

Mech LD, Peterson RO. Wolf-prey relations. In: Mech LD, Boitani L, editors. Wolves: behaviour, ecology, and conservation. University of Chicago Press; 2003. pp. 131–160.

Huggard DJ. Prey selectivity of wolves in Banff National Park. I. Prey species. Can J Zool. 1993. Jan 1;71(1): 130–139.

Stephens DW, Krebs JR. Foraging theory (Vol. 1). Princeton University Press; 1986.

Lind J, Cresswell W. Determining the fitness consequences of antipredation behavior. Behav Ecol. 2005. Sep 1;16(5): 945–56.

Bergman EJ, Garrott RA, Creel S, Borkowski JJ, Jaffe R, Watson FG. Assessment of prey vulnerability through analysis of wolf movements and kill sites. Ecol Appl. 2006. Feb;16(1): 273–284. doi: 10.1890/04-1532 PubMed DOI

Hegel TM, Mysterud A, Ergon T, Loe LE, Huettmann F, Stenseth NC. Seasonal effects of Pacific‐based climate on recruitment in a predator‐limited large herbivore. J Anim Ecol. 2010. Mar;79(2): 471–482. doi: 10.1111/j.1365-2656.2009.01647.x PubMed DOI

Jedrzejewska B, Jedrzejewski W. Predation in vertebrate communities: the Bialowieza Primeval Forest as a case study. Springer; 1998. Jul 20.

Gervasi V, Sand H, Zimmermann B, Mattisson J, Wabakken P, Linnell JD. Decomposing risk: landscape structure and wolf behavior generate different predation patterns in two sympatric ungulates. Ecol Appl. 2013. Oct;23(7): 1722–1734. doi: 10.1890/12-1615.1 PubMed DOI

Wikenros C, Sand H, Wabakken P, Liberg O, Pedersen HC. Wolf predation on moose and roe deer: chase distances and outcome of encounters. Acta Theriol (Warsz). 2009. Sep;54(3): 207–218.

Klare U, Kamler JF, Macdonald DW. A comparison and critique of different scat‐analysis methods for determining carnivore diet. Mamm Rev. 2011. Oct;41(4): 294–312.

Ruehe F, Buschmann I, Wameling A. Two models for assessing the prey mass of European ungulates from wolf scats. Acta Theriol (Warsz). 2003. Dec 1;48(4): 527–37.

Hindrikson M, Remm J, Pilot M, Godinho R, Stronen AV, Baltrūnaité L, et al.. Wolf population genetics in Europe: a systematic review, meta‐analysis and suggestions for conservation and management. Biol Rev. 2017. Aug;92(3): 1601–1629. doi: 10.1111/brv.12298 PubMed DOI

Gula R, Hausknecht R, Kuehn R. Evidence of wolf dispersal in anthropogenic habitats of the Polish Carpathian Mountains. Biodivers Conserv. 2009. Jul;18(8): 2173–84.

Voskár J. Present problems of wolf preservation in Czechoslovakia. Acta Zool Fennica. 1983; 174 (1): 287–8

Chapron G, Kaczensky P, Linnell JD, Von Arx M, Huber D, Andrén H, et al.. Recovery of large carnivores in Europe’s modern human-dominated landscapes. Science. 2014. Dec 19;346(6216): 1517–19. doi: 10.1126/science.1257553 PubMed DOI

Antal V, Boroš M, Ciberej J, Dóczy J, Finďo S, Kaštier P, et al. [Wolf (Canis lupus) care program in Slovakia]. State Nature Conservancy of the Slovak Republic; 2016. Slovakian.

Ministry of Agriculture and Rural Development of the Slovak Republic. [Wolf hunting quota in the 2020/2021 hunting season]. 2020 October 27 [cited 2022 Feb 23]. Available from: https://www.mpsr.sk/kvota-lovu-vlka-draveho-v-polovnickej-sezone-2020-2021/799-37-799-16019%20. In Slovak.

Kaczensky P. Large carnivore depredation on livestock in Europe. Ursus. 1999. Jan 1: 59–71.

Finďo S. [Food ecology of the wolf (Canis lupus) in the Slovak Carpathians]. In: Urban P, editor. [Research and conservation of mammals in Slovakia]. State Nature Conservancy of the Slovak Republic, Banská Bystrica. 2002. pp. 43–55. In Slovak.

Rigg R, Gorman M. Spring-autumn diet of wolves (Canis lupus) in Slovakia and a review of wolf prey selection. Oecologia Montana. 2004. Dec 1;13(1–2): 30–41.

Levins R. Evolution in changing environments: some theoretical explorations (No. 2). Princeton University Press; 1968.

Fischer LA, Gates CC. Competition potential between sympatric woodland caribou and wood bison in southwestern Yukon, Canada. Can J Zool. 2005. Sep 1;83(9): 1162–73.

Roever CL, Boyce MS, Stenhouse GB. Grizzly bears and forestry: II: grizzly bear habitat selection and conflicts with road placement. For Ecol Manage. 2008. Sep 5;256(6): 1262–9.

Smolko P, Kropil R, Pataky T, Veselovská A, Merrill E. Why do migrants move downhill? The effects of increasing predation and density on red deer altitudinal migration in temperate Carpathian forests. Mamm Res. 2018. Jul;63(3): 297–305.

Jędrzejewski W, Jędrzejewska B, Okarma H, Ruprecht AL. Wolf predation and snow cover as mortality factors in the ungulate community of the Bialowieża National Park, Poland. Oecologia. 1992. Apr 1;90(1): 27–36. doi: 10.1007/BF00317805 PubMed DOI

Gula R. Influence of snow cover on wolf Canis lupus predation patterns in Bieszczady Mountains, Poland. Wildlife Biol. 2004. Mar;10(1): 17–23.

Corine Land Cover. European Union, Copernicus Land Monitoring Service, European Environment Agency (EEA); 2018.

Szalai S, Auer I, Hiebl J, Milkovich J, Radim T, Stepanek P, et al. Climate of the Greater Carpathian Region. Final Technical Report. www.carpatclim-eu.org. CARPATCLIM Database © European Commission–JRC; 2003.

Peifer H. About the EEA reference grid. European Environment Agency; 2011. Available from: https://www.eea.europa.eu/data-and-maps/data/eea-reference-grids-2/about-theeea-reference-grid/eea_reference_grid_v1.pdf.

Trites AW, Joy R. Dietary analysis from fecal samples: how many scats are enough?. J Mammal. 2005. Aug 22;86(4): 704–12.

Gable TD, Windels SK, Bruggink JG. The problems with pooling poop: confronting sampling method biases in wolf (Canis lupus) diet studies. Can J Zool. 2017;95(11): 843–851.

Ciucci P, Boitani L, Pelliccioni ER, Rocco M, Guy I. A comparison of scat-analysis methods to assess the diet of the wolf Canis lupus. Wildlife Biol. 1996. Mar;2(3): 37–48.

Jędrzejewski W, Schmidt K, Theuerkauf J, Jędrzejewska B, Selva N, Zub K, et al.. Kill rates and predation by wolves on ungulate populations in Białowieża Primeval Forest (Poland). Ecology. 2002. May;83(5): 1341–56.

Spaulding R, Krausman PR, Ballard WB. Observer bias and analysis of gray wolf diets from scats. Wildl Soc Bull. 2000. Dec 1: 947–50.

Ciucci P, Tosoni E, Boitani L. Assessment of the point-frame method to quantify wolf Canis lupus diet by scat analysis. Wildlife Biol. 2004. Jun;10(1): 149–53.

Teerink BJ. Hair of West European mammals: atlas and identification key. Cambridge University Press; 2003.

Keller A. [Determination of the mammals of Switzerland by their pelage: V. Carnivora, VI. Artiodactyla]. Revue Suisse de Zoologie. 1978; 88(3): 803–20. In French.

Debrot S, Fivaz G, Mermod C, Weber JM. [Atlas of the hairs of European mammals]. Institut de Zoologie, Université de Neuchâtel, Peseux, Switzerland; 1982. pp. 208. In French.

Félix GA, Piovezan U, Quadros J, Juliano RS, Alves FV, Fioravanti MC. Thricology for identifying mammal species and breeds: Its use in research and agriculture. Archivos de Zootecnia. 2014. Jan 28;63(241): 107–16.

Valente AM, Rocha RG, Lozano E, Ferreira JP, Fonseca C. [Atlas of Hairs of Iberian Terrestrial Mammals]. Universidade de Aveiro, Edições Afrontamento; 2015. In Portuguese.

Smolko P, Veselovská A, Kropil R. Seasonal dynamics of forage for red deer in temperate forests: importance of the habitat properties, stand development stage and overstorey dynamics. Wildlife Biol. 2018. Feb; 2018(1).

Smolko P, Garaj P, Lebocký T, Bútora Ľ, Pataky T, Jaňáková Z, et al.. Soil nutrients and deer density affect antler size of the Carpathian red deer. Mamm Biol. 2022. Jan 8:1–12.

Esri Inc. ArcMap (version 10.5.1). Software. Redlands, CA; 2016.

Dytham C. Choosing and using statistics: a biologist’s guide. John Wiley & Sons, West Sussex, UK; 2011. Aug 2.

Ruprecht AL. Food of the Barn owl, Tyto alba guttata (CL Br.) from Kujawy. Acta Ornithol. 1979;16(19).

Krebs CJ. Ecological methodology (No. QH541. 15. S72. K74 1999.). New York: Harper and Row; 1989.

Jacobs J. Quantitative measurement of food selection. Oecologia. 1974. Dec;14(4):413–417. doi: 10.1007/BF00384581 PubMed DOI

Kusak J, Skrbinšek AM, Huber D. Home ranges, movements, and activity of wolves (Canis lupus) in the Dalmatian part of Dinarids, Croatia. Eur J Wildl Res. 2005. Dec;51(4): 254–262.

Anderson DR, Burnham KP. Avoiding pitfalls when using information-theoretic methods. J Wildl Manage. 2002. Jul 1: 912–18.

Boyce MS, Vernier PR, Nielsen SE, Schmiegelow FK. Evaluating resource selection functions. Ecol Modell. 2002. Nov 30;157(2–3): 281–300.

R Core Team. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria; 2020.

Ansorge H, Kluth G, Hahne S. Feeding ecology of wolves Canis lupus returning to Germany. Acta Theriol (Warsz). 2006. Mar;51(1): 99–106.

Gazzola A, Avanzinelli E, Bertelli I, Tolosano A, Bertotto P, Musso R, et al.. The role of the wolf in shaping a multi‐species ungulate community in the Italian western Alps. Ital J Zool (Modena). 2007. Sep 1;74(3): 297–307.

Mattioli L, Apollonio M, Mazzarone V, Centofanti E. Wolf food habits and wild ungulate availability in the Foreste Casentinesi National Park, Italy. Acta Theriol (Warsz). 1995;40(4): 387–402.

Capitani CL, Bertelli I, Varuzza P, Scandura M, Apollonio M. A comparative analysis of wolf (Canis lupus) diet in three different Italian ecosystems. Mamm Biol. 2004. Jan;69(1): 1–10.

Gazzola A, Bertelli I, Avanzinelli E, Tolosano A, Bertotto P, Apollonio M. Predation by wolves (Canis lupus) on wild and domestic ungulates of the western Alps, Italy. J Zool. 2005. Jun;266(2): 205–13.

Cuesta L, Barcena F, Palacios F, Reig S. The trophic ecology of the Iberian Wolf (Canis lupus signatus Cabrera, 1907). A new analysis of stomach’s data. Mammalia. 1991;55(2).

Barja I. Prey and prey-age preference by the Iberian wolf Canis lupus signatus in a multiple-prey ecosystem. Wildlife Biol. 2009. Jun;15(2): 147–54.

Nowak S, Mysłajek RW, Kłosińska A, Gabryś G. Diet and prey selection of wolves (Canis lupus) recolonising Western and Central Poland. Mamm Biol. 2011. Nov 1;76(6): 709–15.

Corradini A. Wolf (Canis Lupus) in Romania: Winter Feeding Ecology and Spatial Interaction with Lynx (Lynx Lynx). Master’s Thesis, University of Florence, Firenze, Italy, 2015.

Meriggi A, Lovari S. A review of wolf predation in southern Europe: does the wolf prefer wild prey to livestock?. J Appl Ecol. 1996. Dec 1: 1561–71.

Śmietana W, Klimek A. Diet of wolves in the Bieszczady Mountains, Poland. Acta Theriol (Warsz). 1993;38(3): 245–51.

Meriggi A, Brangi A, Schenone L, Signorelli D, Milanesi P. Changes of wolf (Canis lupus) diet in Italy in relation to the increase of wild ungulate abundance. Ethol Ecol Evol. 2011. Jul 1;23(3): 195–210.

Wagner C, Holzapfel M, Kluth G, Reinhardt I, Ansorge H. Wolf (Canis lupus) feeding habits during the first eight years of its occurrence in Germany. Mamm Biol. 2012. May;77(3): 196–203.

Andersone Ž. Beaver: A New Prey of Wolves in Latvia?. In: Busher PE, Dzięciołowski RM, editors. Beaver Protection, Management, and Utilization in Europe and North America. Springer, Boston, MA. 1999. Jun 30: 103–108.

Okarma H. The trophic ecology of wolves and their predatory role in ungulate communities of forest ecosystems in Europe. Acta Theriol (Warsz). 1995;40(4): 335–86.

Nowak S, Mysłajek RW, Jędrzejewska B. Patterns of wolf Canis lupus predation on wild and domestic ungulates in the Western Carpathian Mountains (S Poland). Acta Theriol (Warsz). 2005. Jun;50(2): 263–76.

Mori E, Benatti L, Lovari S, Ferretti F. What does the wild boar mean to the wolf?. European Journal of Wildlife Research. 2017. Feb;63(1): 1–5.

Mattioli L, Capitani C, Avanzinelli E, Bertelli I, Gazzola A, Apollonio M. Predation by wolves (Canis lupus) on roe deer (Capreolus capreolus) in north-eastern Apennine, Italy. J Zool. 2004. Nov;264(3): 249–58.

Blackburn JK, Asher V, Stokke S, Hunter DL, Alexander KA. Dances with anthrax: wolves (Canis lupus) kill anthrax bacteremic plains bison (Bison bison bison) in southwestern Montana. Journal of Wildlife Diseases. 2014. Apr;50(2): 393–6. doi: 10.7589/2013-08-204 PubMed DOI

Tanner E, White A, Acevedo P, Balseiro A, Marcos J, Gortázar C. Wolves contribute to disease control in a multi-host system. Sci Rep. 2019. May 28;9(1): 1–2. PubMed PMC

Ministry of Agriculture and Rural Development of the Slovak Republic. [Let’s stop ASF together]. 2022 Feb 23 [cited 23 February 2022]. Available from: https://www.mpsr.sk/africky-mor-osipanych-aktualny-vyvoj-situacie/424–14648/. In Slovak.

Finďo S, Rigg R, Skuban M. The wolf in Slovakia. Perspectives of wolves in Central Europe. 2008: 15–24.

Kuijper DP, De Kleine C, Churski M, Van Hooft P, Bubnicki J, Jędrzejewska B. Landscape of fear in Europe: wolves affect spatial patterns of ungulate browsing in Białowieża Primeval Forest, Poland. Ecography. 2013. Dec;36(12): 1263–75.

Widén A, Clinchy M, Felton AM, Hofmeester TR, Kuijper DP, Singh NJ, et al.. Playbacks of predator vocalizations reduce crop damage by ungulates. Agric Ecosyst Environ. 2022. Apr 15;328: 107853.

Linnell JD, Cretois B, Nilsen EB, Rolandsen CM, Solberg EJ, Veiberg V, et al.. The challenges and opportunities of coexisting with wild ungulates in the human-dominated landscapes of Europe’s Anthropocene. Biol Conserv. 2020. Apr 1;244: 108500.