In Central Europe, protected areas are too small to ensure survival of populations of large carnivores. In the surrounding areas, these species are often persecuted due to competition with game hunters. Therefore, understanding how predation intensity varies spatio-temporally across areas with different levels of protection is fundamental. We investigated the predation patterns of Eurasian lynx (Lynx lynx) on roe deer (Capreolus capreolus) and red deer (Cervus elaphus) in both protected areas and multi-use landscapes of the Bohemian Forest Ecosystem. Based on 359 roe and red deer killed by 10 GPS-collared lynx, we calculated the species-specific annual kill rates and tested for effects of season and lynx age, sex and reproductive status. Because roe and red deer in the study area concentrate in unprotected lowlands during winter, we modeled spatial distribution of kills separately for summer and winter and calculated-the probability of a deer killed by lynx and-the expected number of kills for areas with different levels of protection. Significantly more roe deer (46.05-74.71/year/individual lynx) were killed than red deer (1.57-9.63/year/individual lynx), more deer were killed in winter than in summer, and lynx family groups had higher annual kill rates than adult male, single adult female and subadult female lynx. In winter the probability of a deer killed and the expected number of kills were higher outside the most protected part of the study area than inside; in summer, this probability did not differ between areas, and the expected number of kills was slightly larger inside than outside the most protected part of the study area. This indicates that the intensity of lynx predation in the unprotected part of the Bohemian Forest Ecosystem increases in winter, thus mitigation of conflicts in these areas should be included as a priority in the lynx conservation strategy.

Department of Research and Documentation Bavarian Forest National Park Freyunger Str 2 94481 Grafenau Germany

Department of Research and Documentation Bavarian Forest National Park Freyunger Str 2 94481 Grafenau Germany; Wildlife Ecology and Management University of Freiburg Faculty of Environment and Natural Resources Tennenbacher Straße 4 79106 Freiburg Germany

Department of Statistics LMU Munich Ludwigstraße 33 80539 Munich Germany

Faculty of Forestry and Wood Sciences Czech University of Life Sciences Prague Kamýcká 1176 16521 Prague 6 Czech Republic; Department of Research and Nature Protection Šumava National Park Administration Sušická 399 34192 Kašperské Hory Czech Republic

Zobrazit více v PubMed

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; 346(6216): 1517–1519. 10.1126/science.1257553 PubMed DOI

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; 343: 10.1126/science.1241484 PubMed DOI

Kaczensky P, Chapron G, von Arx M. Status, management and distribution of large carnivores–bear, lynx, wolf & wolverine–in Europe. Report to the European Commission 2013: 72pp.

Müller J, Wölfl M, Wölfl S, Müller DW, Hothorn T, Heurich M. Protected areas shape the spatial distribution of a European lynx population more than 20 years after reintroduction. Biological Conservation 2014; 177: 210–217.

Araújo MB, Alagador D, Cabeza M, Nogués‐Bravo D, Thuiller W. Climate change threatens European conservation areas. Ecology Letters 2011; 14(5): 484–492. 10.1111/j.1461-0248.2011.01610.x PubMed DOI PMC

Carroll C, Noss RF, Paquet PC, Schumaker NH. Extinction debt of protected areas in developing landscapes. Conservation Biology 2004; 18:1110–1120. 10.1111/j.1523-1739.2004.00083.x DOI

EEA European Environment Agency. Protected areas in Europe–an overview. In: EEA Report No 5/2012 Kopenhagen 2012. ISBN 978-92-9213-329-0 ISSN 1725-9177 pdf doi = 10.2800/55955

Ordiz A, Bischof R, Swenson JE. Saving large carnivores, but losing the apex predator? Biological Conservation 2013; 168:128–133. 10.1016/j.biocon.2013.09.024 DOI

Andrén H, Linnell JD, Liberg O, Andersen R, Danell A, Karlsson J, et al. Survival rates and causes of mortality in Eurasian lynx (Lynx lynx) in multi-use landscapes. Biological Conservation 2006; 131(1): 23–32.

Linnell JDC, Swenson JE, Anderson R. Predators and people: conservation of large carnivores is possible at high human densities if management policy is favourable. Animal Conservation 2001; 4: 345–349.

Liberg O, Chapron G, Wabakken P, Pedersen HC, Thompson Hobbs N, Sand H. Shoot, shovel and shut up: cryptic poaching slows restoration of a large carnivore in Europe. Proceedings of the Royal Society B-Biological Sciences 2012; 279: 910–915. 10.1098/rspb.2011.1275 PubMed DOI PMC

Treves A, Karanth KU. Human-carnivore conflict and perspectives on carnivore management worldwide. Conservation Biology 2003; 17:1491–1499. 10.1111/j.1523-1739.2003.00059.x DOI

Breitenmoser U. Large predators in the Alps: The fall and rise of man's competitors. Biological Conservation 1998; 83: 279–289.

Gervasi V, Nilsen EBE, Odden J, Bouyer Y, Linnell JDC. The spatio‐temporal distribution of wild and domestic ungulates modulates lynx kill rates in a multi‐use landscape. Journal of Zoology 2013; 292:9 10.1111/jzo.12088 DOI

Sunquist M, Sunquist F. Wild cats of the world University of Chicago Press; 2002. 452 pp.

Odden J, Herfindal I, Linnell JD, Andersen R. Vulnerability of domestic sheep to lynx depredation in relation to roe deer density. Journal of Wildlife Management 2008; 72(1): 276–282.

Moa PF, Herfindal I, Linnell JDC, Overskaug K, Kvam T, Andersen R. Does the spatiotemporal distribution of livestock influence forage patch selection in Eurasian lynx Lynx lynx? Wildlife Biology 2006; 12(1): 63–70.

Breitenmoser U, Ryser A, Molinari-Jobin A, Zimmermann F, Haller H, Molinari P, et al. The changing impact of predation as a source of conflict between hunters and reintroduced lynx in Switzerland In: Macdonald DW, Loveridge AJ, editors. Biology and Conservation of Wild Felids. Oxford University Press, Oxford; 2010. pp.493–506.

Červený J, Koubek P, Bufka L. Eurasian lynx Lynx lynx and its chance for survival in central Europe: the case of the Czech Republic. Acta Zoologica Lituanica 2002; 12: 362–366.

Andersen R, Karlsen J, Austmo L. Selectivity of Eurasian lynx Lynx lynx and recreational hunters for age, sex and body condition in roe deer Capreolus capreolus . Wildlife Biology 2007; 4:467–474.

Nowicki P. Food habits and diet of the lynx (Lynx lynx) in Europe. Journal of Wildlife Research 1997; 2:161–166.

Jedrzejewski W, Schmidt K, Milkowski L, Jedrzejewska B, Okarma H. Foraging by lynx and its role in ungulate mortality: the local (Bialowieza Forest) and the Palaearctic viewpoints. Acta Theriologica 1993; 38:384–403.

Okarma H, Jedrzejewski W, Schmidt K, Kowalczyk R, Jedrzejewska B. Predation of Eurasian lynx on roe deer and red deer in Bialowieza Primeval Forest, Poland. Acta Theriologica 1997; 42:203–224.

Molinari-Jobin A, Molinari P, Breitenmoser-Würsten C, Breitenmoser U. Significance of lynx Lynx lynx predation for roe deer Capreolus capreolus and chamois Rupicapra rupicapra mortality in the Swiss Jura Mountains. Wildlife Biology 2002; 8:109–115.

Jobin A, Molinari P, Breitenmoser U. Prey spectrum, prey preference and consumption rates of Eurasian lynx in the Swiss Jura Mountains. Acta Theriologica 2000; 45:243–252.

Mattisson J, Odden J, Nilsen EB, Linnell JDC, Persson J, Andrén H. Factors affecting Eurasian lynx kill rates on semi-domestic reindeer in northern Scandinavia: Can ecological research contribute to the development of a fair compensation system? Biological Conservation 2011; 144:3009–3017. 10.1016/j.biocon.2011.09.004 DOI

Nilsen E, Linnell J, Odden J, Andersen R. Climate, season, and social status modulate the functional response of an efficient stalking predator: the Eurasian lynx. Journal of Animal Ecology 2009; 78:741–751. 10.1111/j.1365-2656.2007.0 PubMed DOI

Odden J, Linnell JDC, Andersen R. Diet of Eurasian lynx, Lynx lynx, in the boreal forest of south-eastern Norway: the relative importance of livestock and hares at low roe deer density. European Journal of Wildlife Research 2006; 52:237–244.

Krofel M, Jerina K, Kljun F, Kos I, Potočnik H, Ražen N, et al. Comparing patterns of human harvest and predation by Eurasian lynx Lynx lynx on European roe deer Capreolus capreolus in a temperate forest. European Journal of Wildlife Research 2014; 60:11–21. 10.1007/s10344-013-0745-4 DOI

Breitenmoser U, Haller H. Patterns of predation by reintroduced European lynx in the Swiss Alps. Journal of Wildlife Management 1993; 57(1): 135–144.

Breitenmoser U, Haller A. Zur Nahrungsökologie des Luchses Lynx lynx in den schweizerischen Nordalpen. Zeitschrift für Säugetierkunde 1987; 52: 168–191.

Molinari-Jobin A, Zimmermann F, Ryser A, Molinari P, Haller H, Breitenmoser-Würsten Ch, et al. Variation in diet, prey selectivity and home-range size of Eurasian lynx Lynx lynx in Switzerland. Wildlife Biology 2007; 13:393–405.

Krofel M, Kos I, Jerina K. The noble cats and the big bad scavengers: effects of dominant scavengers on solitary predators. Behavioral Ecology and Sociobiology 2012; 66: 1297–1304.

Mattisson J, Persson J, Andrén H, Segerström P. Temporal and spatial interactions between an obligate predator, the Eurasian lynx (Lynx lynx), and a facultative scavenger, the wolverine (Gulo gulo). Canadian Journal of Zoology 2011; 89:79–89. 10.1139/Z10-097 DOI

Andrén H, Liberg O. Large impact of Eurasian lynx predation on roe deer population dynamics. PLoS ONE 2015; 10(3): e0120570 10.1371/journal.pone.0120570 PubMed DOI PMC

Laundré JW, Hernández L, Ripple WJ. The landscape of fear: ecological implications of being afraid. Open Ecology Journal 2010; 3:1–7.

Mysterud A, Loe LE, Zimmermann B, Bischof R, Veiberg V, Meisingset E. Partial migration in expanding red deer populations at northern latitudes–a role for density dependence? Oikos 2011; 120(12): 1817–1825.

Luccarini S, Mauri L, Ciuti S, Lamberti P, Apollonio M. Red deer (Cervus elaphus) spatial use in the Italian Alps: home range patterns, seasonal migrations, and effects of snow and winter feeding. Ethology Ecology & Evolution 2006; 18(2): 127–145.

Kjellander P, Hewison AJM, Liberg O, Angibault JM, Bideau E, Cargnelutti B. Experimental evidence for density-dependence of home-range size in roe deer (Capreolus capreolus L.): a comparison of two long-term studies. Oecologia 2004; 139(3): 478–485. PubMed

Mysterud A. Seasonal migration pattern and home range of roe deer (Capreolus capreolus) in an altitudinal gradient in southern Norway. Journal of Zoology (London) 1999; 247: 479–486.

Cagnacci F, Focardi S, Heurich M, Stache A, Hewison A, Morellet N, et al. Partial migration in roe deer: migratory and resident tactics are end points of a behavioural gradient determined by ecological factors. OIKOS 2011; 120: 1790–1802.

Mysterud A. Ungulate migration, plant phenology, and large carnivores: The times they are a-changin'. Ecology 2013; 94:1257–1261. 10.2307/23436143 PubMed DOI

De Vries MF. Large herbivores and the design of large-scale nature reserves in Western Europe. Conservation Biology 1995; 9: 25–33. 10.1046/j.1523-1739.1995.09010025.x DOI

Mayer K, Belotti E, Bufka L, Heurich M. Dietary patterns of the Eurasian lynx (Lynx lynx) in the Bohemian Forest. Säugertierkundliche Informationen 2012; 45:447–453.

Röder J, Bässler C, Brandl R, Dvořak L. Arthropod species richness in the Norway Spruce (Picea abies (L.) Karst.) canopy along an elevation gradient. Forest Ecology and Management 2010; 259:1513–1521.

Heurich M, Neufanger M. Die Wälder des Nationalparks Bayerischer Wald; 2005. 179pp.

Wölfl M, Bufka L, Červený J, Koubek P, Heurich M, Habel H, et al. Distribution and status of lynx in the border region between Czech Republic, Germany and Austria. Acta Theriologica 2001; 46:181–194.

Festetics A. Das ehemalige und gegenwärtige Vorkommen des Luchses, Lynx lynx (Linné, 1758) in Europa und seine Wiederansiedlung in einigen europäischen Ländern. Säugetierkundliche Mitteilungen 1981; 29: 21–77.

Koubek P, Červený J. Lynx in the Czech and Slovak Republics. Acta Scientiarum Naturalium Academiae Scientiarum Bohemicae BRNO XXX Nova Ser; 1996:1–78.

Weingarth K, Heibl C, Knauer F, Zimmermann F, Bufka L, Heurich M. First estimation of Eurasian lynx (Lynx lynx) abundance and density using digital cameras and capture–recapture techniques in a German national park. Animal Biodiversity and Conservation 2012; 35:197–207.

Siebert U, Strauß E, Gräber R, Neubauer D. Rehwildzählung im NP Bayerischer Wald; 2013. Unpublished report. 16pp.

Heurich M, Brand TTG, Kaandorp MY, Šustr P, Müller J, Reineking B. Country, cover or protection: What shapes the distribution of red deer and roe deer in the Bohemian Forest Ecosystem? PlosOne 2015. (in press). PubMed PMC

Cagnacci F, Focardi S, Ghisla A, van Moorter B, Gurarie E, Heurich M, et al. How many routes lead to migration? Re-establishing the link between definitions, methods and movement ecology. 2015. (Submitted to Animal Ecology). PubMed

Möst L, Hothorn T, Müller J, Heurich M. Creating a landscape of management: Unintended effects on the variation of browsing pressure in a national park. Forest Ecology and Management 2014; 338: 46–56.

Gerner J, Heurich M, Günther S, Schraml U. Red deer at a crossroads—an analysis of communication strategies concerning wildlife management in the ‘Bayerischer Wald’ National Park, Germany. Journal of Nature Conservation 2011; 19(5): 319–326.

Ludwig M, Grüninger F, Rothfuß E, Heurich M. Discourse analysis as an instrument to reveal the pivotal role of the media in local acceptance or rejection of a wildlife management project. Erdkunde Archive for Scientific Geography 2012; 66(2):143–156.

Heurich M. Beruecksichtigung von Tierschutzaspekten beim Fang und der Markierung von Wildtieren. In: Erhard M, editor. 12. Internationale Fachtagung zu Fragen von Verhaltenskunde, Tierhaltung und Tierschutz; 2011 May 5–7; München. Deutsche Veterinärmedizinische Gesellschaft. Fachgruppe Ethologie und Tierhaltung. ISBN: 978-3-86345-018-2; 2011: 142–157.

Podolski I, Belotti E, Bufka L, Reulen H, Heurich M. Seasonal and daily activity patterns of free-living Eurasian lynx Lynx lynx in relation to availability of kills. Wildlife Biology 2013; 19(1):69–77.

Heurich M, Hilger A, Küchenhoff H, Andrén H, Bufka L, Krofel M, et al. Activity patterns of Eurasian lynx are modulated by light regime and individual traits over a wide latitudinal range. PlosOne 2014; 9(12), e114143. PubMed PMC

Krofel M, Skrbinšek T, Kos I. Use of GPS location clusters analysis to study predation, feeding, and maternal behavior of the Eurasian lynx. Ecological Research 2013; 28:1–42.

Kleinbaum DG, Klein M. Survival Analysis; 2012. 10.1007/978-1-4419-6646-9 DOI

Kalbfleisch J, Prentice R. The statistical analysis of failure time data, 2nd ed.; 2011. 462pp.

Hosmer DW Jr, Lemeshow S, May S. Applied Survival Analysis: Regression Modeling of Time to Event Data (Google eBook); 2011. 416pp.

Wienke A. Frailty models in survival analysis CRC Press, Boca Raton; 2010. 324pp.

Bretz F, Hothorn T, Westfall P. Multiple Comparisons Using R CRC Press, Boca Raton; 2010. 208 pp.

Efron B. Bootstrap methods: another look at the jackknife. Annals of Statistics 1979; 7:1–26.

R Core Team. R: A language and environment for statistical computing Vienna, Austria: R Foundation for Statistical Computing; 2014. http://www.R-project.org/

Therneau T. A Package for Survival Analysis in S. version 2.38; 2015. http://cran.r-project.org/package=survival

Wickham H. The Split-apply-combine strategy for data analysis. Journal of Statistical Software 2011; 40(1): 1–29. URL http://www.jstatsoft.org/v40/i01/.

Hothorn T, Bretz F, Westfall P. Simultaneous Inference in General Parametric Models. Biometrical Journal 2008; 50(3): 346–363. 10.1002/bimj.200810425 PubMed DOI

Wood SN. Generalized Additive Models: An Introduction with R. Chapman & Hall/CRC, Boca Raton; 2006. ISBN 978–1584884743.

EEA European Environment Agency. Corine Land Cover 2000 (CLC2000); 2009. Available: http://www.eea.europa.eu/data-and-maps/data/corine-land-cover-2000-raster-2. Accessed 28 November 2012.

Ersson L. Estimating lynx kill rate on reindeer using GPS-locations and lynx movement pattern. MSc thesis, Swedish University of Agricultural Sciences, Grimsö, Sweden; 2011. 19 pp.

Stahl P, Vandel JM, Herrenschmidt V, Migot P. Predation on livestock by an expanding reintroduced lynx population: long‐term trend and spatial variability. Journal of Applied Ecology 2001; 38(3): 674–687.

Weingarth K, Zeppenfeld T, Heibl C, Heurich M, Bufka L, Daniszová K, et al. Hide & seek–extended camera-trap session lengths and autumn provide best parameters for estimating lynx densities in mountainous areas. Biodiversity and Conservation 2015. (in press).

Powell RA, Zimmerman JW, Seaman DE. Ecology and behavior of North American black bears: home ranges, habitat, and social organization Chapman & Hall, London; 1997.

Litvaitis J, Sherburne J, Bissonette J. Bobcat habitat use and home range size in relation to prey density. Journal of Wildlife Management 1986; 50(1):110–117.

Belotti E, Kreisinger J, Romportl D, Heurich M, Bufka L. Eurasian lynx hunting red deer: is there an influence of a winter enclosure system? European Journal of Wildlife Research 2014; 60(3): 441–457.

Basille M, Herfindal I, Santin–Janin H, Linnell JDC, Odden J, Andersen R, et al. What shapes Eurasian lynx distribution in human–dominated landscapes: selecting prey or avoiding people? Ecography 2009; 32(4): 683–691. 10.1111/j.1600-0587.2009.05712.x DOI

Niedzialkowska M, Jedrzejewski W, Myslajek RW, Nowak S, Jedrzejewska B, Schmidt K. Environmental correlates of Eurasian lynx occurrence in Poland—Large scale census and GISmapping. Biological Conservation 2006; 133: 63–69.

Bunnefeld N, Linnell JDC, Odden J, Van Duijn MAJ, Andersen R. Risk taking by Eurasian lynx (Lynx lynx) in a human–dominated landscape: effects of sex and reproductive status. Journal of Zoology 2006; 270: 31–39

Selva N, Jędrzejewska B, Jędrzejewski W, Wajrak A. Factors affecting carcass use by a guild of scavengers in European temperate woodland. Canadian Journal of Zoology 2005; 83(12): 1590–1601.

Selva N, Jedrzejewska B, Jedrzejewski W, Wajrak A. Scavenging on European bison carcasses in Bialowieza primeval forest (eastern Poland). Ecoscience 2003; 10(3): 303–311.

Schmidt K, Jędrzejewki W, Okarma H. Spatial organization and social relations in the Eurasian lynx population in Białowieża Primeval Forest, Poland. Acta Theriologica 1997; 42: 289–312.

Ray RR, Seibold H, Heurich M. Invertebrates outcompete vertebrate facultative scavengers in simulated lynx kills in the Bavarian Forest National Park, Germany. Animal Biodiversity and Conservation 2014; 37(1): 77–88.

Krofel M, Huber D, Kos I. Diet of Eurasian lynx Lynx lynx in the northern Dinaric Mountains (Slovenia and Croatia). Acta Theriologica 2011; 56(4): 315–322.

Hebblewhite M. Predation by wolves interacts with the North Pacific Oscillation (NPO) on a western North American elk population. Journal of Animal Ecology 2005; 74:226–233. 10.1111/j.1365-2656.2004.00909.x DOI

Post E, Peterson R, Stenseth N, McLaren B. Ecosystem consequences of wolf behavioural response to climate. Nature 1999; 88:905–907.

Okarma H. The physical condition of red deer falling a prey to the wolf and lynx and harvested in the Carpathian Mountains. Acta Theriologica 1984; 29(23): 283–290.

Schmidt K. Variation in daily activity of the free-living Eurasian lynx (Lynx lynx) in Bialowieza Primeval Forest, Poland. Journal of Zoology (London) 1999; 249: 417–425.

Jedrzejewski W, Schmidt K, Okarma H, Kowalczyk R. Movement pattern and home range use by the Eurasian lynx in Białowieża Primeval Forest (Poland). Annales Zoologici Fennici 2002; 39: 29–41.

Hartová-Nentvichová M, Šálek M, Červený J, Koubek P. Variation in the diet of the red fox (Vulpes vulpes) in mountain habitats: Effects of altitude and season. Mammalian Biology 2010; 75(4): 334–340.

Anděra M, Červený J. Atlas of distribution of the mammals of the Šumava Mts. region (SW-Bohemia). Acta Scientiarum Naturalium Brno 1994; 28 (2–3): 111 pp.

Pulliainen E, Lindgren E, Tunkkari PS. Influence of food availability and reproductive status on the diet and body condition of the European lynx in Finland. Acta Theriologica 1995; 40(2): 181–196.

Wilson RP, Culik BM. The cost of a hot meal: facultative specific dynamic action may ensure temperature homeostasis in post-ingestive endotherms. Comparative Biochemistry and Physiology Part A: Physiology 1991; 100(1): 151–154. PubMed

Zatloukal V. The Šumava National Park—history, mission and basic data.—In: Neuhäuslová Z, editor. The map of potential natural vegetation of the Šumava National Park Explanatory text. Silva Gabreta 2001; Supplementum 1: 77–81.

Kamler JF, Jedrzejewski W, Jedrzejewska B. Survival and cause specific mortality of red deer Cervus elaphus in Bialowieza National Park, Poland. Wildlife Biology 2007; 13:48–52.

Melis C, Nilsen EB, Panzacchi M, Linnell JDC, Odden J. Roe deer face competing risks between predators along a gradient in abundance. Ecosphere 2013; 4: e111 (11 pp).

Melis C, Jedrzejewska B, Apollonio M, Barton KA, Jedrzejewski W, Linnell JDC, et al. Predation has a greater impact in less productive environments: Variation in roe deer Capreolus capreolus population density across Europe. Global Ecology and Biogeography 2009; 18: 724–734.

Heurich M, Möst L, Schauberger G. Survival and causes of death of European Roe Deer before and after Eurasian Lynx reintroduction in the Bavarian Forest National Park. European Journal of Wildlife Research 2012; 58:567–578. 10.1007/s10344-011-0606-y DOI

Hansen AJ, DeFries R. Ecological mechanisms linking protected areas to surrounding lands. Ecological Applications 2007; 17(4): 974–988. PubMed

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

Spatial variation in red deer density in a transboundary forest ecosystem

Timing and synchrony of birth in Eurasian lynx across Europe

Demography of a Eurasian lynx (Lynx lynx) population within a strictly protected area in Central Europe

Habitat selection by Eurasian lynx (Lynx lynx) is primarily driven by avoidance of human activity during day and prey availability during night