European Journal of Wildlife Research

, Volume 58, Issue 4, pp 733–742 | Cite as

Large predators limit herbivore densities in northern forest ecosystems

  • William J. RippleEmail author
  • Robert L. Beschta
Original Paper


There is a lack of scientific consensus about how top-down and bottom-up forces interact to structure terrestrial ecosystems. This is especially true for systems with large carnivore and herbivore species where the effects of predation versus food limitation on herbivores are controversial. Uncertainty exists whether top-down forces driven by large carnivores are common, and if so, how their influences vary with predator guild composition and primary productivity. Based on data and information in 42 published studies from over a 50-year time span, we analyzed the composition of large predator guilds and prey densities across a productivity gradient in boreal and temperate forests of North America and Eurasia. We found that predation by large mammalian carnivores, especially sympatric gray wolves (Canis lupus) and bears (Ursus spp.), apparently limits densities of large mammalian herbivores. We found that cervid densities, measured in deer equivalents, averaged nearly six times greater in areas without wolves compared to areas with wolves. In areas with wolves, herbivore density increased only slightly with increasing productivity. These predator effects are consistent with the exploitation ecosystems hypothesis and appear to occur across a broad range of net primary productivities. Results are also consistent with theory on trophic cascades, suggesting widespread and top-down forcing by large carnivores on large herbivores in forest biomes across the northern hemisphere. These findings have important conservation implications involving not only the management of large carnivores but also that of large herbivores and plant communities.


Large carnivores Wolves Bears Cervids Productivity Trophic cascades Herbivory 



We thank J. Hollenbeck, J. Leniham, C. Melis, L. Painter, and A. Souther for providing assistance on this project.


  1. Abrams PA (2000) The evolution of predator–prey interactions: theory and evidence. Annu Rev Ecol Syst 31:79–105CrossRefGoogle Scholar
  2. Angelstam P, Wikberg PE, Danilov P, Faber WE, Nygren K (2000) Effects of moose density on timber quality and biodiversity restoration in Sweden, Finland, and Russian Karelia. Alces 36:133–145Google Scholar
  3. Aunapuu M, Dahlgren J, Oksanen T, Grellmann D, Oksanen L, Olofsson J, Rammul U, Schneider M, Johansen G, Hyden HO (2008) Spatial patterns and dynamic responses of arctic food webs corroborate the exploitation ecosystems hypothesis. Am Nat 171:249–262CrossRefPubMedGoogle Scholar
  4. Bachelet D, Lenihan JM, Daly C, Neilson RP, Ojima DS, Parton WJ (2001) MC1: a dynamic vegetation model for estimating the distribution of vegetation and associated ecosystem fluxes of carbon, nutrients, and water. USDA General Technical Report PNW-GTR-508, 95 ppGoogle Scholar
  5. Ballard WB, Whitman JS, Gardner CL (1987) Ecology of an exploited wolf population in south central Alaska. Wildl Monogr 98:1–54Google Scholar
  6. Baskin LM (1994) Population ecology of the moose in the Russian Southern Taiga. Alces 30:51–55Google Scholar
  7. Behrend DF, Mattfeld GF, Tierson WC, Wiley JE III (1970) Deer density control for comprehensive forest management. J For 68:695–700Google Scholar
  8. Berger J (2005) Hunting by carnivores and humans: does functional redundancy occur and does it matter? In: Ray JC, Redford KH, Steneck RS, Berger J (eds) Large carnivores and the conservation of biodiversity. Island Press, Washington, DC, pp 315–341Google Scholar
  9. Bergerud AT, Elliot JP (1985) Dynamics of caribou and wolves in northern British Columbia. Can J Zool 64:1515–1529CrossRefGoogle Scholar
  10. Bergerud AT, Wyatt W, Snider B (1983) The role of wolf predation in limiting moose population. J Wildl Manage 47:977–988CrossRefGoogle Scholar
  11. Beschta RL, Ripple WJ (2007) Wolves, elk, and aspen in the winter range of Jasper National Park, Canada. Can J For Res 37:1873–1885CrossRefGoogle Scholar
  12. Beschta RL, Ripple WJ (2009) Large predators and trophic cascades in terrestrial ecosystems of the western United States. Biol Conserv 142:2401–2414CrossRefGoogle Scholar
  13. Beschta RL, Ripple WJ (2010) Mexican wolves, elk, and aspen in Arizona: is there a trophic cascade? For Ecol Manage 260:915–922CrossRefGoogle Scholar
  14. Beschta RL, Ripple WJ (2011) The role of large predators in maintaining riparian plant communities and river morphology. Geomorphology, in press. doi: 10.1016/j.geomorph.2011.04.042
  15. Borer ET, Seabloom EW, Shurnin JB, Anderson KE, Blanchette CA, Broitman B, Cooper SD, Halpern BS (2005) What determines the strength of a trophic cascade? Ecology 86:528–537CrossRefGoogle Scholar
  16. Cairns AL, Telfer ES (1980) Habitat use by four sympatric ungulates in boreal mixedwood forest. J Wildl Manage 47:977–988Google Scholar
  17. Carbyn LN (1983) Wolf predation on elk in Riding Mountain National Park, Manitoba. J Wildl Manage 47:963–976CrossRefGoogle Scholar
  18. Ceballos G, Ehrlich PR (2002) Mammal population losses and the extinction crisis. Science 296:904–907CrossRefPubMedGoogle Scholar
  19. Cederlund G, Markgren G (1987) The development of the Swedish moose population, 1970–1983. Swedish Wildl Res Suppl 1:55–61Google Scholar
  20. Côté SD, Rooney TP, Tremblay J, Dussault C, Waller DM (2004) Ecological impacts of deer overabundance. Annu Rev Eco Evol Syst 35:113–147CrossRefGoogle Scholar
  21. Crête M (1987) The impact of sport hunting on North American moose. Swedish Wildl Res Suppl 1:553–563Google Scholar
  22. Crête M (1989) Approximation of K carrying capacity for moose in eastern Quebec. Can J Zool 7:373–380CrossRefGoogle Scholar
  23. Crête M (1999) The distribution of deer biomass supports the hypothesis of exploitation ecosystems. Ecol Lett 2:223CrossRefGoogle Scholar
  24. Crête M, Manseau M (1996) Natural regulation of cervidae along a 1000 km latitudinal gradient: change in trophic dominance. Evol Ecol 10:51–62CrossRefGoogle Scholar
  25. Crête M, Manseau M (1996) Natural regulation of cervidae along a 1000 km latitudinal gradient: change in trophic dominance. Evol Ecol 10:51–62CrossRefGoogle Scholar
  26. Daly C, Bachelet D, Lenihan JM, Neilson RP, Parton W, Ojima D (2000) Dynamic simulation of tree-grass interactions for global change studies. Ecol Appl 10:449–469Google Scholar
  27. Detling JK (1998) Mammalian herbivores: ecosystem-level effects in two grassland national parks. Wildl Soc Bull 26:438–448Google Scholar
  28. Diamond JM (1983) Ecology: laboratory, field and natural experiments. Nature 304:586–587CrossRefGoogle Scholar
  29. Dixon RK, Solomon AM, Brown S, Houghton RA, Trexier MC, Wisniewski J (1994) Carbon pools and flux of global forest ecosystems. Science 263:185–190CrossRefPubMedGoogle Scholar
  30. 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, Soulé ME, Virtanen R, Wardle DA (2011) Trophic downgrading of planet earth. Science 333:301–306CrossRefPubMedGoogle Scholar
  31. Filonov C (1980) Predator–prey problems in nature reserves of the European part of the USSR. J Wildl Manage 44:389–396CrossRefGoogle Scholar
  32. Frelich LE, Lorimer CG (1985) Current and predicted long-term effects of deer browsing in hemlock forests in Michigan, USA. Biol Conserv 34:99–120CrossRefGoogle Scholar
  33. Fretwell SD (1977) The regulation of plant communities by food chains exploiting them. Perspect Biol Med 20:169–185CrossRefGoogle Scholar
  34. Fuller TK (1989) Population dynamics of wolves in north-central Minnesota. Wildl Monogr 105:1–141Google Scholar
  35. Fuller TK, Keith LB (1980) Wolf population dynamics and prey relationships in northeastern Alberta. J Wildl Manage 44:583–602CrossRefGoogle Scholar
  36. Gasaway WC, Stephenson RO, Davis JL, Shepherd PEK, Burris OE (1983) Interrelationships of wolves, prey, and man in interior Alaska. Wildl Monogr 8413Google Scholar
  37. Gasaway WC, Stephenson RO, Davis JL, Shepherd PEK, Burris OE (1983b) Interrelationships of wolves, prey, and man in interior Alaska. Wildl Monogr 84:1–50Google Scholar
  38. Gasaway WC, Boertje RD, Grangaard DV, Kelleyhouse DG, Stephenson RO, Larson DG (1992) The role of predation in limiting moose at low densities in Alaska and Yukon and implications for conservation. Wildl Monogr 120:1–59Google Scholar
  39. Haber GC (1977) Socio-ecological dynamics of wolves and prey in a subarctic ecosystem. PhD thesis, University of British ColumbiaGoogle Scholar
  40. Hairston NG, Smith FE, Slobodkin LB (1960) Community structure, population control, and competition. Am Nat 94:421–425CrossRefGoogle Scholar
  41. Halaj J, Wise DH (2001) Terrestrial trophic cascades: how much do they trickle? Am Nat 157:262–281CrossRefPubMedGoogle Scholar
  42. Hayes RD, Harestad AS (2000) Wolf functional response and regulation of moose in the Yukon. Can J Zool 78:60–66CrossRefGoogle Scholar
  43. Heuze P, Schnitzler A, Klein F (2005) Is browsing the major factor of silver fir decline in the Vosges Mountains of France? For Ecol Manage 217:219–228CrossRefGoogle Scholar
  44. Hough AF (1949) Deer and rabbit browsing and available winter forage in Allegheny hardwood forests. J Wildl Manage 13:135–141CrossRefGoogle Scholar
  45. Jędrzejewski B, Jędrzejewski W (1998) Predation in vertebrate communities: the Białowieża primeval forest as a case study. Springer, New YorkCrossRefGoogle Scholar
  46. Kochetkov VV (2002) Factors determining moose population dynamics in the Central Forest Reserve. Alces Suppl 2:57–51Google Scholar
  47. Kojola I, Huitu O, Toppinen K, Heikura K, Heikkinen S, Ronkainen S (2004) Predation on European wild forest reindeer (Rangifer tarandus) by wolves (Canus lupus) in Finland. J Zool 263:229–235CrossRefGoogle Scholar
  48. Kolenosky GB (1972) Wolf predation on wintering deer in East-Central Ontario. J Wildl Manage 36:357–369CrossRefGoogle Scholar
  49. Laliberte AS, Ripple WJ (2004) Range contractions of North American carnivores and ungulates. BioScience 54:123–138CrossRefGoogle Scholar
  50. Larsen DG (1982) Moose inventory in the southwest Yukon. Alces 18:142–167Google Scholar
  51. Laundré JW, Hernández L, Altendorf KB (2001) Wolves, elk, and bison: reestablishing the “landscape of fear” in Yellowstone National Park, USA. Can J Zool 79:1401–1409CrossRefGoogle Scholar
  52. Leopold A, Sowls LK, Spencer DL (1947) A survey of over-populated deer ranges in the United States. J Wildl Manage 11:162–183CrossRefGoogle Scholar
  53. Luyssaert S, Schulze ED, Börner A, Knohl A, Hessenmöller D, Law BE, Ciais P, Grace J (2008) Old growth forests as global carbon sinks. Nature 455:213–215CrossRefPubMedGoogle Scholar
  54. Mech LD, Boitani L (eds) (2003) Wolves: behavior, ecology, and conservation. University of Chicago Press, ChicagoGoogle Scholar
  55. Mech LD, Peterson RO (2003) Wolf-prey relations. In: Mech LD, Boitani L (eds) Wolves: behavior, ecology, and conservation. University of Chicago Press, Chicago, pp 131–160CrossRefGoogle Scholar
  56. Melis C, Jędrzejewsk B, Apollonio M, Bartoń KA, Jędrzejewski 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. Global Ecol Biogeogr 18:724–734CrossRefGoogle Scholar
  57. Messier F (1994) Ungulate population models with predation: a case study with the North American moose. Ecology 75:478–488CrossRefGoogle Scholar
  58. Michael ED (1992) Impact of deer browsing on regeneration of balsam fir in Canaan Valley, West Virginia. North J Appl For 9:89–90Google Scholar
  59. Miller B, Dugelby B, Foreman D, Martinez del Rio CR, Noss M, Phillips R, Reading ME, Terborgh Soulé J, Willcox L (2001) The importance of large carnivores to healthy ecosystems. Endang Spec Update 18:202–210Google Scholar
  60. Oksanen L (1992) Evolution of exploitation ecosystems. I. Predation, foraging ecology and population dynamics in herbivores. Evol Ecol 6:15–33CrossRefGoogle Scholar
  61. Oksanen L, Fretwell SD, Arruda J, Niemela P (1981) Exploitation ecosystems in gradients of primary productivity. Am Nat 118:240–261CrossRefGoogle Scholar
  62. Orians GH, Cochran PA, Duffield JW, Fuller TK, Gutierrez RJ, Hanemann WM (1997) Wolves, bears, and their prey in Alaska: biological and social challenges in wildlife management. National Academy Press, Washington, DC, p 207Google Scholar
  63. Palmer SCF, Truscott AM (2003) Seasonal habitat use and browsing by deer in Caledonian pinewoods. For Ecol Manage 174:149–166CrossRefGoogle Scholar
  64. Peterson RO (2007) The wolves of Isle Royale: a broken balance. University of Michigan Press, Ann Arbor, p 192Google Scholar
  65. Peterson RO, Woolington JD, Bailey TN (1984) Wolves of the Kenai Peninsula, Alaska. Wildlife Monogr 88Google Scholar
  66. Peterson RO, Vucetich JA, Page RE, Chouinard A (2003) Temporal and spatial aspects of predator–prey dynamics. Alces 39:215–232Google Scholar
  67. Pimlott DH, Shannon JA, Kolenosky GB (1969) The ecology of the timber wolf in Algonquin Provincial Park. Ontario Department of Lands and Forests Research Paper Wildlife No. 87, Toronto, Ontario, CanadaGoogle Scholar
  68. Polis GA (1999) Why are parts of the world green? Multiple factors control productivity and the distribution of green biomass. Oikos 86:3–15CrossRefGoogle Scholar
  69. Potvin F (1988) Wolf movements and population dynamics in Papineau–Labelle reserve, Quebec. Can J Zool 66:1266–1273CrossRefGoogle Scholar
  70. Ray JC, Redford KH, Steneck RS, Berger J (eds) (2005) Large carnivores and the conservation of biodiversity. Island Press, Washington, DC, p 526Google Scholar
  71. Ripple WJ, Beschta RL (2004) Wolves and the ecology of fear: can predation risk structure ecosystems? BioScience 54:755–766CrossRefGoogle Scholar
  72. Ripple WJ, Beschta RL (2005) Linking wolves to plants: Aldo Leopold on trophic cascades. BioScience 55:613–621CrossRefGoogle Scholar
  73. Ripple WJ, Beschta RL (2006) Linking a cougar decline, trophic cascade, and catastrophic regime shift in Zion National Park. Biol Conserv 133:397–408CrossRefGoogle Scholar
  74. Ripple WJ, Beschta RL (2008) Trophic cascades involving cougar, mule deer, and black oaks in Yosemite National Park. Biol Conserv 141:1249–1256CrossRefGoogle Scholar
  75. Ripple WJ, Beschta RL (2012) Trophic cascades in yellowstone: the first 15 years after wolf reintroduction. Biol Conserv 145:205–213CrossRefGoogle Scholar
  76. Ripple WJ, Rooney TP, Beschta RL (2010) Large predators, deer, and trophic cascades in the mid-latitudes. In: Terborgh J, Estes JA (eds) Trophic cascades. Island Press, Washington, DC, pp 141–161Google Scholar
  77. Ritchie EG, Elmhagen B, Glen AS, Letnic M, Ludwig G, McDonald RA (2012) Ecosystem restoration with teeth: what role for predators? Trends Ecol Evol, in press. doi: 10.1016/j.tree.2012.01.001
  78. Schmitz OJ, Hamback PA, Beckerman AP (2000) Trophic cascades in terrestrial systems: a review of the effects of carnivore removal on plants. Am Nat 155:141–153CrossRefPubMedGoogle Scholar
  79. Schreiner EG, Krueger KA, Houston DB, Happe PJ (1996) Understory patch dynamics and ungulate herbivory in old-growth forests of Olympic National Park, Washington. Can J For Res 26:255–265CrossRefGoogle Scholar
  80. Sergio F, Caro T, Brown D, Clucas B, Hunter J, Ketchum J, McHugh K, Hiraldo F (2008) Top predators as conservation tools: ecological rationale, assumptions and efficacy. Annu Rev Ecol Syst 39:1–19CrossRefGoogle Scholar
  81. Singer FJ, Dalle-Molle J (1985) The Denali ungulate-predator system. Alces 21:339–358Google Scholar
  82. Soulé ME, Estes JE, Berger J, del Rio CM (2003) Ecological effectiveness: conservation goals for interactive species. Conserv Biol 17:1238–1250CrossRefGoogle Scholar
  83. Stoeckeler JH, Strothmann RO, Krefting LW (1957) Effect of deer browsing on reproduction in the northern hardwood-hemlock type in northeastern Wisconsin. J Wildl Manage 21:75–80CrossRefGoogle Scholar
  84. Suominen O (1999) Impact of cervid browsing and grazing on the terrestrial gastropod fauna in the boreal forests of Fennoscandia. Ecography 22:651–658CrossRefGoogle Scholar
  85. Terborgh J, Estes JA (eds) (2010) Trophic cascades: predators, prey, and the changing dynamics of nature. Island Press, Washington, DC, p 464Google Scholar
  86. Terborgh J, Estes JA, Paquet P, Ralls K, Boyd-Heigher D, Miller BJ, Noss RF (1999) The role of top carnivores in regulating terrestrial ecosystems. In: Soulé M, Terborgh J (eds) Continental conservation: scientific foundations of regional reserve networks. Island Press, Washington, DC, pp 39–64Google Scholar
  87. Terborgh J, Lopez L, Nunez P, Rao M, Shahabuddin G, Riveros M, Ascanio R, Lambert TD, Adler GH, Balbas L (2001) Ecological meltdown in predator-free forest fragments. Science 294:1923–1926CrossRefPubMedGoogle Scholar
  88. Trumbull VL, Zielinski EJ, Aharrah EC (1989) The impact of deer browsing on the Allegheny forest type. North J Appl For 6:162–165Google Scholar
  89. Van Ballenberghe V (1987) Effects of predation on moose numbers: a review of recent North American studies. Swedish Wildl Res Suppl 1:431–460Google Scholar
  90. Welsh DA, Morrison KP, Oswald K, Thomas ER (1980) Winter utilization of habitat by moose in relation to forest harvesting. Proc North Am Moose Conf Workshop 16:398–428Google Scholar
  91. Wilmers CC, Post ES, Peterson RO, Vucetich JA (2006) Disease mediated switch from top-down to bottom-up control exacerbates climatic effects on moose population dynamics. Ecol Lett 9:383–389CrossRefPubMedGoogle Scholar
  92. Woodroffe R, Ginsberg JR (2005) King of beasts? Evidence for guild redundancy among large mammalian carnivores. In: Ray JC, Redford KH, Steneck RS, Berger J (eds) Large carnivores and the conservation of biodiversity. Island Press, Washington, DC, pp 154–175Google Scholar
  93. Zager P, Beecham J (2006) The role of American black bears and brown bears as predators on ungulates in North America. Ursus 17:95–108CrossRefGoogle Scholar

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© Springer-Verlag 2012

Authors and Affiliations

  1. 1.Department of Forest Ecosystems and Society, College of ForestryOregon State UniversityCorvallisUSA

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