European Journal of Wildlife Research

, Volume 60, Issue 1, pp 11–21 | Cite as

Comparing patterns of human harvest and predation by Eurasian lynx Lynx lynx on European roe deer Capreolus capreolus in a temperate forest

  • Miha KrofelEmail author
  • Klemen Jerina
  • Franc Kljun
  • Ivan Kos
  • Hubert Potočnik
  • Nina Ražen
  • Petra Zor
  • Anamarija Žagar
Original Paper


Human harvest is the most important mortality factor for wild ungulates in Europe and can affect several aspects of ungulate biology. There is a growing concern about possible negative side effects of human harvest. To better understand the differences between human and natural mortality, we compared the extent, age and sex structure, nutritional condition, spatial and temporal distribution of human harvest, and natural predation by the Eurasian lynx Lynx lynx on the European roe deer Capreolus capreolus, the most abundant wild ungulate in Europe. Compared to the human harvest, lynx were less likely to kill fawns and yearlings than adults, and among adult deer, lynx were more likely to kill females. The proportion of roe deer with fat-depleted bone marrow was higher among lynx prey than among harvested animals. Average lynx kill rate was estimated to 47.8 roe deer per year, and lynx predation was considerably lower than the human harvest in the same area. While human harvest increased with higher roe deer density, lynx predation was similar across the gradient of roe deer densities. Comparison with other countries indicated that differences between human harvest and natural mortality of ungulates vary considerably in different parts of Europe. Variation in hunting practices and, even more importantly, carnivore predation may have an important role in buffering unwanted side effects of harvest of wild ungulates.


Dinaric Mountains Hunting Mortality Slovenia Ungulates Wildlife management 

Supplementary material

10344_2013_745_MOESM1_ESM.doc (47 kb)
ESM 1 (DOC 47 kb)


  1. Adamič M, Jerina K (2010) Ungulates and their management in Slovenia. In: Apollonio M, Andersen R, Putman R (eds) European ungulates and their management in the 21st century. Cambridge University Press, Cambridge, pp 507–526Google Scholar
  2. Andersen R, Linnell JDC, Solberg EJ (2006) The future role of large carnivores on terrestrial trophic interactions: the northern temperate view. In: Danell K, Bergström R, Duncan P, Pastor J (eds) Large herbivore ecology, ecosystem dynamics and conservation. Cambridge University Press, Cambridge, pp 413–448CrossRefGoogle Scholar
  3. Andersen R, Karlsen J, Austmo LB, Odden J, Linnell JDC, Gaillard JM (2007) Selectivity of Eurasian lynx Lynx lynx and recreational hunters for age, sex and body condition in roe deer Capreolus capreolus. Wildl Biol 13:467–474CrossRefGoogle Scholar
  4. Andersen R, Lund E, Solberg EJ, Saether BE (2010) Ungulates and their management in Norway. In: Apollonio M, Andersen R, Putman R (eds) European ungulates and their management in the 21st century. Cambridge University Press, Cambridge, pp 14–36Google Scholar
  5. Apollonio M, Andersen R, Putman R (2010) Present status and future challenges for European ungulate management. In: Apollonio M, Andersen R, Putman R (eds) European ungulates and their management in the 21st century. Cambridge University Press, Cambridge, pp 578–604Google Scholar
  6. Apollonio M, Putman R, Grignolio S, Bartoš L (2011) Hunting seasons in relation to biological breeding seasons and the implications for the control or regulation of ungulate populations. In: Putman RJ, Apollonio M, Andersen R (eds) Ungulate management in Europe: problems and practices. Cambridge University Press, Cambridge, pp 80–105CrossRefGoogle Scholar
  7. Beschta RL, Ripple WJ (2009) Large predators and trophic cascades in terrestrial ecosystems of the western United States. Biol Conserv 142:2401–2414. doi: 10.1016/j.biocon.2009.06.015 CrossRefGoogle Scholar
  8. Beyer HL (2012) Geospatial modelling environment (Version Accessed 14 February 2013
  9. Bischof R, Fujita R, Zedrosser A, Soderberg A, Swenson JE (2008) Hunting patterns, ban on baiting, and harvest demographics of brown bears in Sweden. J Wildl Manag 72:79–88. doi: 10.2193/2007-149 CrossRefGoogle Scholar
  10. Breitenmoser U, Breitenmoser-Würsten C (2008) Der Luchs: Ein Grossraubtier in der Kulturlandschaft. Salm Verlag, Wohlen and BernGoogle Scholar
  11. Breitenmoser-Würsten C, Zimmermann F, Stahl P, Vandel JM, Molinari-Jobin A, Molinari P, Capt S, Breitenmoser U (2007) Spatial and social stability of a Eurasian lynx Lynx lynx population: an assessment of 10 years of observation in the Jura Mountains. Wildl Biol 13:365–380CrossRefGoogle Scholar
  12. Brown JS, Kotler BP (2007) Foraging and the ecology of fear. In: Stephens DW, Brown JS, Ydenberg RC (eds) Foraging: behavior and ecology. The University of Chicago Press, Chicago, pp 437–480Google Scholar
  13. Burnham KP, Anderson DR (2002) Model selection and multimodel inference: a practical information-theoretic approach, 2nd edn. Springer, New YorkGoogle Scholar
  14. Ciuti S, Muhly TB, Paton DG, McDevitt AD, Musiani M, Boyce MS (2012) Human selection of elk behavioural traits in a landscape of fear. Proc R Soc B Biol Sci 279:4407–4416. doi: 10.1098/rspb.2012.1483 CrossRefGoogle Scholar
  15. Coltman DW, O’Donoghue P, Jorgenson JT, Hogg JT, Strobeck C, Festa-Bianchet M (2003) Undesirable evolutionary consequences of trophy hunting. Nature 426:655–658PubMedCrossRefGoogle Scholar
  16. Estes JA, Terborgh J, Brashares JS, Power ME, Berger J, Bond WJ, Carpenter SR, Essington TE, Holt RD, Jackson JBC, Marquis RJ, Oksanen L, Oksanen T, Paine RT, Pikitch EK, Ripple WJ, Sandin SA, Scheffer M, Schoener TW, Shurin JB, Sinclair ARE, Soule ME, Virtanen R, Wardle DA (2011) Trophic downgrading of planet Earth. Science 333:301–306. doi: 10.1126/science.1205106 PubMedCrossRefGoogle Scholar
  17. Festa-Bianchet M (2003) Exploitative wildlife management as a selective pressure for the life-history evolution of large mammals. In: Festa-Bianchet M, Apollonio M (eds) Animal behavior and wildlife conservation. Island Press, Washington, pp 191–207Google Scholar
  18. Franzmann AW, Arneson PD (1976) Marrow fat in Alaskan moose femurs in relation to mortality factors. J Wildl Manag 40:336–339CrossRefGoogle Scholar
  19. Gervasi V, Nilsen EB, Sand H, Panzacchi M, Rauset GR, Pedersen HC, Kindberg J, Wabakken P, Zimmermann B, Odden J, Liberg O, Swenson JE, Linnell JDC (2011) Predicting the potential demographic impact of predators on their prey: a comparative analysis of two carnivore–ungulate systems in Scandinavia. J Anim Ecol 81:443–454. doi: 10.1111/j.1365-2656.2011.01928.x PubMedCrossRefGoogle Scholar
  20. Ginsberg JR, Milner-Gulland EJ (1994) Sex-biased harvesting and population dynamics in ungulates: implications for conservation and sustainable use. Conserv Biol 8:157–166CrossRefGoogle Scholar
  21. Gossow H, Honsig-Erlenburg P (1985) Several predation aspects of red deer-specialized lynx. 17th IUGB Congress, Brussels, BruseljGoogle Scholar
  22. Haglund B (1966) Winter habits of the lynx (Lynx lynx L.) and wolverine (Gulo gulo L.) as revealed by tracking in the snow. Viltrevy 4:81–229Google Scholar
  23. Harris RB, Wall WA, Allendorf FW (2002) Genetic consequences of hunting: what do we know and what should we do? Wildl Soc B 30:634–643Google Scholar
  24. Heurich M, Möst L, Schauberger G, Reulen H, Sustr P, Hothorn T (2012) Survival and causes of death of European roe deer before and after Eurasian lynx reintroduction in the Bavarian Forest National Park. Eur J Wildl Res 58:567–578. doi: 10.1007/s10344-011-0606-y CrossRefGoogle Scholar
  25. Hewison AJM, Vincent JP, Angibault JM, Delorme D, Van Laere G, Gaillard JM (1999) Tests of estimation of age from tooth wear on roe deer of known age: variation within and among populations. Can J Zool 77:58–67CrossRefGoogle Scholar
  26. Husseman JS, Murray DL, Power G, Mack C, Wenger CR, Quigley H (2003) Assessing differential prey selection patterns between two sympatric large carnivores. Oikos 101:591–601Google Scholar
  27. Jedrzejewska B, Jedrzejewski W (1998) Predation in vertebrate communities: the Białowieża Primeval Forest as a case study. Springer, HeidelbergCrossRefGoogle Scholar
  28. Jedrzejewski W, Schmidt K, Milkowski L, Jedrzejewska B, Okarma H (1993) Foraging by lynx and its role in ungulate mortality: the local (Bialowieza Forest) and the Palearctic viewpoints. Acta Theriol 38:385–403Google Scholar
  29. Jedrzejewski W, Apollonio M, Jedrzejewska B, Kojola I (2011) Ungulate–large carnivore relationships in Europe. In: Putman RJ, Apollonio M, Andersen R (eds) Ungulate management in Europe: problems and practices. Cambridge University Press, Cambridge, pp 284–318CrossRefGoogle Scholar
  30. Jelenko I, Jerina K, Pokorny B (2010) Impact of environmental factors on the appearance and distribution of dental fluorosis in roe deer (Capreolus capreolus L.) in eastern Slovenia. Zbornik gozdarstva in lesarstva 92:21–32Google Scholar
  31. Klevezal GA (1996) Recording structures of mammals: determination of age and reconstruction of life history. A. A. Balkema, BrookfieldGoogle Scholar
  32. Kos I, Potočnik H, Skrbinšek T, Skrbinšek Majić A, Jonozovič M, Krofel M (2005) Lynx in Slovenia, 2nd edn. University of Ljubljana, LjubljanaGoogle Scholar
  33. Kos I, Koren I, Potočnik H, Krofel M (2012) Status and distribution of Eurasian lynx (Lynx lynx) in Slovenia from 2005 to 2009. Acta Biol Slov 55:49–63Google Scholar
  34. Krofel M (2012) Predation-related interspecific interactions in Eurasian lynx (Lynx lynx) in northern Dinaric Mountains. Doctorate thesis. University of Ljubljana, LjubljanaGoogle Scholar
  35. Krofel M, Huber D, Kos I (2011) Diet of Eurasian lynx Lynx lynx in the northern Dinaric Mountains (Slovenia and Croatia): importance of edible dormouse Glis glis as alternative prey. Acta Theriol 56:315–322. doi: 10.1007/s13364-011-0032-2 CrossRefGoogle Scholar
  36. Krofel M, Jonozovič M, Jerina K (2012) Demography and mortality patterns of removed brown bears in a heavily exploited population. Ursus 23:91–103. doi: 10.2192/URSUS-D-10-00013.1 CrossRefGoogle Scholar
  37. Krofel M, Skrbinšek T, Kos I (2013) Use of GPS location clusters analysis to study predation, feeding, and maternal behavior of the Eurasian lynx. Ecol Res 28:103–116. doi: 10.1007/s11284-012-1005-x CrossRefGoogle Scholar
  38. Langvatn R, Loison A (1999) Consequences of harvesting on age structure, sex ratio and population dynamics of red deer Cervus elaphus in central Norway. Wildl Biol 5:213–223Google Scholar
  39. Linnell JDC, Zachos FE (2011) Status and distribution patterns of European ungulates: genetics, population history and conservation. In: Putman R, Apollonio M, Andersen R (eds) Ungulate management in Europe: problems and practices. Cambridge University Press, Cambridge, pp 12–53CrossRefGoogle Scholar
  40. Melis C, Jedrzejewska B, Apollonio M, Barton KA, Jedrzejewski W, Linnell JDC, Kojola I, Kusak J, Adamic M, Ciuti S, Delehan I, Dykyy I, Krapinec K, Mattioli L, Sagaydak A, Samchuk N, Schmidt K, Shkvyrya M, Sidorovich VE, Zawadzka B, Zhyla S (2009) Predation has a greater impact in less productive environments: variation in roe deer, Capreolus capreolus, population density across Europe. Glob Ecol Biogeogr 18:724–734. doi: 10.1111/j.1466-8238.2009.00480.x CrossRefGoogle Scholar
  41. Melis C, Basille M, Herfindal I, Linnell JDC, Odden J, Gaillard JM, Hogda KA, Andersen R (2010) Roe deer population growth and lynx predation along a gradient of environmental productivity and climate in Norway. Ecoscience 17:166–174. doi: 10.2980/17-2-3314 CrossRefGoogle Scholar
  42. Metz MC, Smith DW, Vucetich JA, Stahler DR, Peterson RO (2012) Seasonal patterns of predation for gray wolves in the multi-prey system of Yellowstone National Park. J Anim Ecol 81:553–563. doi: 10.1111/j.1365-2656.2011.01945.x PubMedCrossRefGoogle Scholar
  43. Milner JM, Nilsen EB, Andreassen HP (2007) Demographic side effects of selective hunting in ungulates and carnivores. Conserv Biol 21:36–47. doi: 10.1111/j.1523-1739.2006.00591.x PubMedCrossRefGoogle Scholar
  44. Moa PF, Herfindal I, Linnell JDC, Overskaug K, Kvam T, Andersen R (2006) Does the spatiotemporal distribution of livestock influence forage patch selection in Eurasian lynx Lynx lynx? Wildl Biol 12:63–70CrossRefGoogle Scholar
  45. Muggeo VMR (2003) Estimating regression models with unknown break-points. Stat Med 22:3055–3071. doi: 10.1002/sim.1545 PubMedCrossRefGoogle Scholar
  46. Mysterud A (2010) Still walking on the wild side? Management actions as steps towards ‘semi-domestication’ of hunted ungulates. J Appl Ecol 47:920–925. doi: 10.1111/j.1365-2664.2010.01836.x CrossRefGoogle Scholar
  47. Neiland KA (1970) Weight of dried marrow as indicator of fat in caribou femurs. J Wildl Manag 34:904–907CrossRefGoogle Scholar
  48. Nilsen EB, Linnell JDC, Odden J, Andersen R (2009) Climate, season, and social status modulate the functional response of an efficient stalking predator: the Eurasian lynx. J Anim Ecol 78(4):741–751. doi: 10.1111/j.1365-2656.2009.01547.x PubMedCrossRefGoogle Scholar
  49. Okarma H (1984) The physical condition of red deer falling a prey to the wolf and lynx and harvested in the Carpathian Mountains. Acta Theriol 29:283–290Google Scholar
  50. Okarma H (1995) The trophic ecology of wolves and their predatory role in ungulate communities of forest ecosystems in Europe. Acta Theriol 40:335–386Google Scholar
  51. Pedersen VA, Linnell JDC, Andersen R, Andren H, Linden M, Segerstrom P (1999) Winter lynx Lynx lynx predation on semi-domestic reindeer Rangifer tarandus in northern Sweden. Wildl Biol 5:203–211Google Scholar
  52. Perko D, Orožen Adamič M (1998) Slovenija: pokrajine in ljudje. Mladinska knjiga, LjubljanaGoogle Scholar
  53. Poličnik H, Pokorny B, Varljen Bužan E, Kryštufek B (2011) Spreminjanje spola uplenjenih/izločenih parkljarjev - le čemu? Kontrola z analizo genskega lokusa SRY. Lovec 94:75–76Google Scholar
  54. Preisser EL, Bolnick DI, Benard MF (2005) Scared to death? The effects of intimidation and consumption in predator–prey interactions. Ecology 86(2):501–509. doi: 10.1890/04-0719 CrossRefGoogle Scholar
  55. Proaktor G, Coulson T, Milner-Gulland EJ (2007) Evolutionary responses to harvesting in ungulates. J Anim Ecol 76:669–678. doi: 10.1111/j.1365-2656.2007.01244.x PubMedCrossRefGoogle Scholar
  56. Putman R (2011) A review of the various legal and administrative systems governing management of large herbivores in Europe. In: Putman R, Apollonio M, Andersen R (eds) Ungulate management in Europe: problems and practices. Cambridge University Press, Cambridge, pp 54–79CrossRefGoogle Scholar
  57. Ransom AB (1965) Kidney and marrow fat as indicators of white-tailed deer condition. J Wildl Manag 29:397–398CrossRefGoogle Scholar
  58. Ratcliffe PR (1980) Bone marrow fat as an indicator of condition in roe deer. Acta Theriol 25:333–340Google Scholar
  59. Ruth TK, Buotte PC, Quigley HB (2010) Comparing ground telemetry and global positioning system methods to determine cougar kill rates. J Wildl Manag 74(5):1122–1133. doi: 10.2193/2009-058 CrossRefGoogle Scholar
  60. Sand H, Wikenros C, Ahlqvist P, Strømseth TH, Wabakken P (2012) Comparing body condition of moose (Alces alces) selected by wolves (Canis lupus) and human hunters: consequences for the extent of compensatory mortality. Can J Zool 90:403–412. doi: 10.1139/z2012-007 CrossRefGoogle Scholar
  61. Schmidt K (2008) Behavioural and spatial adaptation of the Eurasian lynx to a decline in prey availability. Acta Theriol 53:1–16CrossRefGoogle Scholar
  62. Slovenia Forest Service (2007) Dolgoročni načrt za IV Notranjsko lovsko upravljavsko območje za obdobje 2007–2016. Slovenia Forest Service, PostojnaGoogle Scholar
  63. Smole I (2012) The importance of large herbivore carcasses for vertebrate scavengers in Dinaric forests of Slovenia. Diploma thesis. University of Ljubljana, LjubljanaGoogle Scholar
  64. Stephens DW, Brown JS, Ydenberg RC (2007) Foraging: behavior and ecology. The University of Chicago Press, ChicagoCrossRefGoogle Scholar
  65. Stergar M, Borkovič D, Hiršelj J, Kavčič I, Krofel M, Mrakič M, Troha R, Videmšek U, Vrčon B, Jerina K (2012) Estimation of wild ungulate densities with a combined method of pellet group counting and removal data. Biotechnical Faculty. University of Ljubljana, LjubljanaGoogle Scholar
  66. Temple SA (1987) Do predators always capture substandard individuals disproportionately from prey populations? Ecology 68:669–674CrossRefGoogle Scholar
  67. Wilmers CC, Crabtree RL, Smith DW, Murphy KM, Getz WM (2003) Trophic facilitation by introduced top predators: grey wolf subsidies to scavengers in Yellowstone National Park. J Anim Ecol 72(6):909–916CrossRefGoogle Scholar
  68. Wright GJ, Peterson RO, Smith DW, Lemke TO (2006) Selection of northern Yellowstone elk by gray wolves and hunters. J Wildl Manag 70(4):1070–1078CrossRefGoogle Scholar
  69. Zar JH (1999) Biostatistical analysis. Prentice Hall, New JerseyGoogle Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2013

Authors and Affiliations

  • Miha Krofel
    • 1
    • 2
    Email author
  • Klemen Jerina
    • 1
  • Franc Kljun
    • 2
  • Ivan Kos
    • 2
  • Hubert Potočnik
    • 2
  • Nina Ražen
    • 2
  • Petra Zor
    • 2
  • Anamarija Žagar
    • 2
  1. 1.Department of Forestry and Renewable Forest Resources, Biotechnical FacultyUniversity of LjubljanaLjubljanaSlovenia
  2. 2.Department of Biology, Biotechnical FacultyUniversity of LjubljanaLjubljanaSlovenia

Personalised recommendations