• Hardin, G. The competitive exclusion principle. Science 131, 1292–1298 (1960).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Barnosky, A. D. et al. Approaching a state shift in Earth’s biosphere. Nature 486, 52–58 (2012).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Di Marco, M. & Santini, L. Human pressures predict species’ geographic range size better than biological traits. Glob. Change Biol. 21, 2169–2178 (2015).

    Article 

    Google Scholar
     

  • Estes, J. A. et al. Trophic downgrading of planet earth. Science 333, 301–306 (2011).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Global Assessment Report on Biodiversity and Ecosystem Services of the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services (IPBES, 2019); https://www.ipbes.net/global-assessment

  • Van Der Putten, W. H., Macel, M. & Visser, M. E. Predicting species distribution and abundance responses to climate change: why it is essential to include biotic interactions across trophic levels. Philos. Trans. R. Soc. B Biol. Sci. 365, 2025–2034 (2010).

    Article 

    Google Scholar
     

  • Wilson, M. W. et al. Ecological impacts of human-induced animal behaviour change. Ecol. Lett. 23, 1522–1536 (2020).

    Article 
    PubMed 

    Google Scholar
     

  • Roemer, G. W., Donlan, C. J. & Courchamp, F. Golden eagles, feral pigs, and insular carnivores: how exotic species turn native predators into prey. Proc. Natl. Acad. Sci. USA. 99, 791–796 (2002).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Cimatti, M. et al. Large carnivore expansion in Europe is associated with human population density and land cover changes. Divers. Distrib. 27, 602–617 (2021).

    Article 

    Google Scholar
     

  • Chapron, G. et al. Recovery of large carnivores in Europe’s modern human-dominated landscapes. Science 346, 1517–1519 (2014).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Levi, T. & Wilmers, C. C. Wolves-coyotes-foxes: a cascade among carnivores. Ecology 93, 921–929 (2012).

    Article 
    PubMed 

    Google Scholar
     

  • Trouwborst, A., Krofel, M. & Linnell, J. D. C. Legal implications of range expansions in a terrestrial carnivore: the case of the golden jackal (Canis aureus) in Europe. Biodivers. Conserv. 24, 2593–2610 (2015).

    Article 

    Google Scholar
     

  • Lanszki, J., Hayward, M. W., Ranc, N. & Zalewski, A. Dietary flexibility promotes range expansion: the case of golden jackals in Eurasia. J. Biogeogr. https://doi.org/10.1111/jbi.14372 (2022).

  • Giannatos, G., Marinos, Y., Maragou, P. & Catsadorakis, G. The status of the golden jackal (Canis aureus L.) in Greece. Belgian J. Zool. 135, 145–149 (2005).


    Google Scholar
     

  • Krofel, M., Giannatos, G., Ćirović, D., Stoyanov, S. & Newsome, T. M. Golden jackal expansion in Europe: a case of mesopredator release triggered by continent-wide wolf persectution?. Hystrix 28, 9–15 (2017).


    Google Scholar
     

  • Sommer, R. & Benecke, N. Late-Pleistocene and early Holocene history of the canid fauna of Europe (Canidae). Mamm. Biol. Z. Säugetierkd. 70, 227–241 (2005).

    Article 

    Google Scholar
     

  • Tóth, T., Krecsák, L., Szűcs, E., Heltai, M. & Huszár, G. Records of the golden jackal (Canis aureus Linnaeus, 1758) in Hungary from 1800th until 2007, based on a literature survey. North. West. J. Zool. 5, 386–405 (2009).

  • Krofel, M. et al. New mesocarnivore at the doorstep of central Europe: historic development of golden jackal (Canis aureus) population in Slovenia. Mamm. Res. 68, 329–339 (2023).

    Article 

    Google Scholar
     

  • Ranc, N. et al. Distribution of large carnivores in Europe 2012–2016: Distribution map for golden jackal (Canis aureus). Zenodo https://doi.org/10.5281/zenodo.6382216 (2022).

  • Hatlauf, J., Bayer, K., Trouwborst, A. & Hackländer, K. New rules or old concepts? The golden jackal (Canis aureus) and its legal status in central Europe. Eur. J. Wildl. Res. 67, 25 (2021).

    Article 

    Google Scholar
     

  • Gorman, J. Rise of the golden jackal. The New York Times (14 January 2019).

  • Newsome, T. M. et al. Top predators constrain mesopredator distributions. Nat. Commun. 8, 15469 (2017).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Prugh, L. R. et al. The rise of the mesopredator. Bioscience 59, 779–791 (2009).

    Article 

    Google Scholar
     

  • Crooks, K. R. & Soule, M. E. Mesopredator release and avifaunal extinctions in a fragmented system. Nature 400, 563–566 (1999).

    Article 
    CAS 

    Google Scholar
     

  • Cunze, S. & Klimpel, S. From the Balkan towards western Europe: range expansion of the golden jackal (Canis aureus)—a climatic niche modeling approach. Ecol. Evol. 12, e9141 (2022).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Šálek, M. et al. Population densities and habitat use of the golden jackal (Canis aureus) in farmlands across the Balkan Peninsula. Eur. J. Wildl. Res. 60, 193–200 (2014).

    Article 

    Google Scholar
     

  • Augustin, N. H., Mugglestone, M. A. & Buckland, S. T. An autologistic model for spatial distribution of wildlife. J. Appl. Ecol. 33, 339–347 (1996).

    Article 

    Google Scholar
     

  • Berger, J. Fear, human shields and the redistribution of prey and predators in protected areas. Biol. Lett. 3, 620–623 (2007).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Kaczensky, P. et al. Distribution of large carnivores in Europe 2012–2016: distribution maps for Brown bear, Eurasian lynx, grey wolf, and Wolverine. Dryad https://doi.org/10.5061/dryad.pc866t1p3 (2021).

  • Fenton, S. et al. Movement, space-use and resource preferences of European golden jackals in human-dominated landscapes: insights from a telemetry study. Mamm. Biol. 101, 619–630 (2021).

    Article 

    Google Scholar
     

  • Yusefi, G. H. et al. Habitat use and population genetics of golden jackals in Iran: insights from a generalist species in a highly heterogeneous landscape. J. Zool. Syst. Evol. Res. 59, 1503–1515 (2021).

    Article 

    Google Scholar
     

  • Ćirović, D., Penezić, A. & Krofel, M. Jackals as cleaners: ecosystem services provided by a mesocarnivore in human-dominated landscapes. Biol. Conserv. 199, 51–55 (2016).

    Article 

    Google Scholar
     

  • Berger, K. M., Gese, E. M. & Berger, J. Indirect effects and traditional trophic cascades: a test involving wolves, coyotes, and pronghorn. Ecology 89, 818–828 (2008).

    Article 
    PubMed 

    Google Scholar
     

  • Sivy, K. J., Pozzanghera, C. B., Grace, J. B. & Prugh, L. R. Fatal attraction? Intraguild facilitation and suppression among predators. Am. Nat. 190, 663–679 (2017).

    Article 
    PubMed 

    Google Scholar
     

  • Newsome, T. M. & Ripple, W. J. A continental scale trophic cascade from wolves through coyotes to foxes. J. Anim. Ecol. 84, 49–59 (2015).

    Article 
    PubMed 

    Google Scholar
     

  • Prugh, L. R. et al. Fear of large carnivores amplifies human-caused mortality for mesopredators. Science 380, 754–758 (2023).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Moll, R. J. et al. Humans and urban development mediate the sympatry of competing carnivores. Urban Ecosyst. 21, 765–778 (2018).

    Article 

    Google Scholar
     

  • Rutkowski, R. et al. A European concern? Genetic structure and expansion of golden jackals (Canis aureus) in Europe and the Caucasus. PLoS ONE 10, 11 (2015).

    Article 

    Google Scholar
     

  • Lanszki, J., Schally, G., Heltai, M. & Ranc, N. Golden jackal expansion in Europe: first telemetry evidence of a natal dispersal. Mamm. Biol. 88, 81–84 (2018).

    Article 

    Google Scholar
     

  • Slatyer, R. A., Umbers, K. D. L. & Arnold, P. A. Ecological responses to variation in seasonal snow cover. Conserv. Biol. 36, e13727 (2022).

    Article 
    PubMed 

    Google Scholar
     

  • Walther, G. et al. Ecological response to recent climate change. Nature 416, 389–395 (2002).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • IPCC 2023 Climate Change 2023: Synthesis Report (eds Core Writing Team, Lee, H. & Romero, J.) (IPCC, 2023).

  • Männil, P. & Ranc, N. Golden jackal (Canis aureus) in Estonia: development of a thriving population in the boreal ecoregion. Mamm. Res. 67, 245–250 (2022).

    Article 

    Google Scholar
     

  • Ripple, W. J. et al. Status and ecological effects of the world’s largest carnivores. Science 343, 80 (2014).

    Article 

    Google Scholar
     

  • Di Bernardi, C. et al. Continuing recovery of wolves in Europe. PLOS Sustain. Transform. 4, e0000158 (2025).

    Article 

    Google Scholar
     

  • Frýbová, Š. et al. Bold wolf behaviour: definitions and analysis of reported past cases across Europe (2025).

  • Chapron, G., Epstein, Y., Bruskotter, J. T. & López-bao, J. V. Europeans support large carnivore recovery while opposing both further population growth and hunting. Nat. Ecol. Evol. 10, 28–33 (2026).

    Article 
    PubMed 

    Google Scholar
     

  • Steyaert, S. M. J. G. et al. Human shields mediate sexual conflict in a top predator. Proc. R. Soc. B Biol. Sci. 283 (2016).

  • Lanszki, J., Hayward, M. W. & Nagyapáti, N. Feeding responses of the golden jackal after reduction of anthropogenic food subsidies. PLoS One 13, 1–18 (2018).

    Article 

    Google Scholar
     

  • Krofel, M., Hočevar, L., Fležar, U., Topličanec, I. & Oliveira, T. Golden jackal as a new kleptoparasite for Eurasian lynx in Europe. Glob. Ecol. Conserv. 36, e02116 (2022).


    Google Scholar
     

  • Raichev, E. G. Effect of shooting on the structure of population of golden jackal (Canis аureus L.) in Sarnena Sredna Gora mountain. Agric. Sci. Technol. 3, 276–280 (2011).


    Google Scholar
     

  • Minnie, L., Gaylard, A. & Kerley, G. I. H. Compensatory life-history responses of a mesopredator may undermine carnivore management efforts. J. Appl. Ecol. 53, 379–387 (2016).

    Article 

    Google Scholar
     

  • Berger, K. M. Carnivore-livestock conflicts: Effects of subsidized predator control and economic correlates on the sheep industry. Conserv. Biol. 20, 751–761 (2006).

    Article 
    PubMed 

    Google Scholar
     

  • Raynor, J. L., Grainger, C. A. & Parker, D. P. Wolves make roadways safer, generating large economic returns to predator conservation. Proc. Natl. Acad. Sci. USA. 118, 1–10 (2021).

    Article 

    Google Scholar
     

  • Gehrt, S. D., Muntz, E. M., Wilson, E. C., Power, J. W. B. & Newsome, S. D. Severe environmental conditions create severe conflicts: A novel ecological pathway to extreme coyote attacks on humans. J. Appl. Ecol. 60, 353–364 (2023).

    Article 
    CAS 

    Google Scholar
     

  • Soulé, M. E., Estes, J. A. & Berger, J. & Martínez del Rio, C. Ecological effectiveness: conservation goals for interactive species. Conserv. Biol. 17, 1238–1250 (2003).

    Article 

    Google Scholar
     

  • Krofel, M., Treves, A., Ripple, W. J., Chapron, G. & Lopez-Bao, J. V. Hunting carnivores at outsized risk. Science 350, 518 (2015).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • van Eeden, L. M. et al. Carnivore conservation needs evidence-based livestock protection. PLoS Biol. 16, 1–8 (2018).


    Google Scholar
     

  • Gese, E. M. Territorial defense by coyotes (Canis latrans) in Yellowstone National Park, Wyoming: Who, how, where, when, and why. Can. J. Zool. 79, 980–987 (2001).

    Article 

    Google Scholar
     

  • Jaeger, M. M., Pandit, R. K. & Haque, E. Seasonal differences in territorial behavior by golden jackals in Bangladesh: howling versus confrontation. J. Mamm. 77, 768–775 (1996).

    Article 

    Google Scholar
     

  • Harrington, F. H. & Mech, L. D. Wolf howling and territory maintanence. Behaviour 68, 207–249 (1979).

    Article 

    Google Scholar
     

  • Szabó, L., Heltai, M., Lanszki, J. & Szucs, E. An indigenous predator, the golden jackal (Canis aureus L. 1758) spreading like an invasive species in Hungary. Bull. USAMV-CN 63–64, 230–235 (2007).


    Google Scholar
     

  • Smith, B. R., Bru, E., Faur, M. & Kershenbaum, A. Acoustic localisation as a tool to aid monitoring of golden jackals (Canis aureus). Ecol. Evol. 15 (2025).

  • Johnson, D. H. The comparison of usage and availability measurements for evaluating resource preference. Ecology 61, 65–71 (1980).

    Article 

    Google Scholar
     

  • Ranc, N., Krofel, M. & Cirovic, D. IUCN Red List Mapping for the Regional Assessment of the Golden Jackal (Canis aureus) in Europe (IUCN, 2018).

  • Lamut, S. The Optimization of the Acoustic Method for Monitoring the Golden Jackal Populations. MSc thesis, Ljubljana Univ. (2020).

  • Giannatos, G. Conservation Action Plan for the Golden Jackal Canis aureus L. (WWF Greece, 2004).

  • Dietz, A. J., Wohner, C. & Kuenzer, C. European snow cover characteristics between 2000 and 2011 derived from improved modis daily snow cover products. Remote Sens. 4, 2432–2454 (2012).

    Article 

    Google Scholar
     

  • Corine Land Cover, Copernicus Land Monitoring Service (European Environment Agency, 2012).

  • Szabó, L., Heltai, M., Szucs, E., Lanszki, J. & Lehoczki, R. Expansion range of the golden jackal in Hungary between 1997 and 2006. Mammalia 73, 307–311 (2009).

    Article 

    Google Scholar
     

  • Water & Wetness, Copernicus Land Monitoring Service (European Environment Agency, 2015).

  • Imperviousness, Copernicus Land Monitoring Service (European Environment Agency, 2015).

  • Root-Gutteridge, H. et al. Not afraid of the big bad wolf: calls from large predators do not silence mesopredators. Wildlife Biol. 2024, e01226 (2024).

    Article 

    Google Scholar
     

  • Guisan, A. & Thuiller, W. Predicting species distribution: offering more than simple habitat models. Ecol. Lett. 8, 993–1009 (2005).

    Article 
    PubMed 

    Google Scholar
     

  • Elith, J. & Leathwick, J. R. Species distribution models: ecological explanation and prediction across space and time. Annu. Rev. Ecol. Evol. Syst. 40, 677–697 (2009).

    Article 

    Google Scholar
     

  • Di Marco, M., Pacifici, M., Maiorano, L. & Rondinini, C. Drivers of change in the realised climatic niche of terrestrial mammals. Ecography 44, 1180–1190 (2021).

    Article 

    Google Scholar
     

  • Gallien, L., Douzet, R., Pratte, S., Zimmermann, N. E. & Thuiller, W. Invasive species distribution models—how violating the equilibrium assumption can create new insights. Glob. Ecol. Biogeogr. 21, 1126–1136 (2012).

    Article 

    Google Scholar
     

  • Krofel, M. Survey of golden jackals (Canis aureus L.) in Northern Dalmatia, Croatia: preliminary results. Natura Croatica 17, 259–264 (2008).


    Google Scholar
     

  • Harrington, F. H. & Mech, L. D. An analysis of howling response parameters useful for wolf pack censusing. J. Wildl. Manage. 46, 686–693 (1982).

    Article 

    Google Scholar
     

  • Forester, J. D., Im, H. K. & Rathouz, P. J. Accounting for animal movement in estimation of resource selection functions: sampling and data analysis. Ecology 90, 3554–3565 (2009).

    Article 
    PubMed 

    Google Scholar
     

  • Thieurmel, B. & Elmarhraoui, A. suncalc: Compute Sun Position, Sunlight Phases, Moon Position and Lunar Phase. R version 0.5.0 https://CRAN.R-project.org/package=suncalc (2019).

  • MacKenzie, D. I. et al. Estimating site occupancy rates when detection probabilities are less than one. Ecology 83, 2248–2255 (2002).

    Article 

    Google Scholar
     

  • Fithian, W. & Hastie, T. Finite-sample equivalence in statistical models for presence-only data. Ann. Appl. Stat. 7, 1917–1939 (2013).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Warton, D. I., Renner, I. W. & Ramp, D. Model-based control of observer bias for the analysis of presence-only data in ecology. PLoS ONE 8, e79168 (2013).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Brooks, M. E. et al. glmmTMB balances speed and flexibility among packages for zero-inflated generalized linear mixed modeling. R J. 9, 378–400 (2017).

    Article 

    Google Scholar
     

  • Bardos, D. C., Guillera-Arroita, G. & Wintle, B. A. Valid auto-models for spatially autocorrelated occupancy and abundance data. Methods Ecol. Evol. 6, 1137–1149 (2015).

    Article 

    Google Scholar
     

  • Bivand, R. S., Pebesma, E. & Gomez-Rubio, V. Applied Spatial Data Analysis with R (Springer, 2013).

  • Dormann, C. F. et al. Methods to account for spatial autocorrelation in the analysis of species distributional data: a review. Ecography 30, 609–628 (2007).

    Article 

    Google Scholar
     

  • Schwarz, G. Estimating the dimension of a model. Ann. Stat. 6, 461–44 (1978).

    Article 

    Google Scholar
     

  • Brewer, M. J., Butler, A. & Cooksley, S. L. The relative performance of AIC, AICC and BIC in the presence of unobserved heterogeneity. Methods Ecol. Evol. 7, 679–692 (2016).

    Article 

    Google Scholar
     

  • Roberts, D. R. et al. Cross-validation strategies for data with temporal, spatial, hierarchical, or phylogenetic structure. Ecography 40, 913–929 (2017).

    Article 

    Google Scholar
     

  • Fielding, A. H. & Bell, J. F. A review of methods for the assessment of prediction errors in conservation presence/absence models. Environ. Conserv. 24, 38–49 (1997).

    Article 

    Google Scholar
     

  • Hirzel, A. H., Le Lay, G., Helfer, V., Randin, C. & Guisan, A. Evaluating the ability of habitat suitability models to predict species presences. Ecol. Modell. 199, 142–152 (2006).

    Article 

    Google Scholar
     

  • Boyce, M. S., Vernier, P. R., Nielsen, S. E. & Schmiegelow, F. K. A. Evaluating resource selection functions. Ecol. Modell. 157, 281–300 (2002).

    Article 

    Google Scholar
     

  • Csányi, S., Márton, M., Kiss, K., Köteles, P. & Schally, G. Vadgazdálkodási Adattár—2019/2020 (Országos Vadgazdálkodási Adattár, 2020).

  • Burnham, K. P. & Anderson, D. R. Multimodel inference: understanding AIC and BIC in model selection. Sociol. Methods Res. 33, 261–304 (2004).

    Article 

    Google Scholar
     

  • Elith, J., Kearney, M. & Phillips, S. The art of modelling range-shifting species. Methods Ecol. Evol. 1, 330–342 (2010).

    Article 

    Google Scholar
     

  • R Core Team R: A Language and Environment for Statistical Computing (R Foundation for Statistical Computing, 2016).

  • QGIS Development Team Quantum GIS Geographic Information System (Open Source Geospatial Foundation Project, 2016).

  • Ranc, N. et al. Human shielding from wolves facilitates jackal expansion across Europe—dataset and code. Zenodo https://doi.org/10.5281/zenodo.18771950 (2026).