Behavioral Ecology and Sociobiology

, Volume 66, Issue 2, pp 295–303 | Cite as

The causes and consequences of partial prey consumption by wolves preying on moose

  • John A. VucetichEmail author
  • Leah M. Vucetich
  • Rolf O. Peterson
Original Paper


For a wide range of taxa, partial prey consumption (PPC) is a frequent occurrence. PPC may arise from physiological constraints to gut capacity or digestive rate. Alternatively, PPC may represent an optimal foraging strategy. Assessments that clearly distinguish between these causes are rare and have been conducted only for invertebrate species that are ambush predators with extra-intestinal digestion (e.g., wolf spiders). We present the first strong test for the cause of PPC in a cursorial vertebrate predator with intestinal digestion: wolves (Canis lupus) feeding on moose (Alces alces). Previous theoretical assessments indicate that if PPC represents an optimal foraging strategy and is not caused by physiological limitations, then mean carcass utilization is negatively correlated with mean kill rate and the utilization of individual carcasses is uncorrelated with time between kills. Wolves exhibit exactly this pattern. We explore how the typical portrayal of PPC by wolves has been not only misleading but also detrimental to conservation by promoting negative attitudes toward wolves.


Alces alces Canis lupus Numerical response Optimal foraging Predator–prey dynamics 



We thank the US National Science Foundation (DEB-0918247) and the US National Park Service for financial support.

Ethical Standards

This work complies with the current Michigan Technological University Institutional Animal Care and Use Committee guidelines, which are guided by the US federal regulations and ethical principles, intended to ensure the humane care and use of animals in research.

Conflict of Interest Statement

The authors declare that they have no conflict of interest.


  1. Abrams PA (1982) Functional responses of optimal foragers. Am Nat 120:382–390CrossRefGoogle Scholar
  2. Bayliss P, Choquenot D (2002) The numerical response: rate of increase and food limitation in herbivores and predators. Philos T Roy Soc Lond B 357:1233–1248CrossRefGoogle Scholar
  3. Bjärvall A, Nilson E (1976) Surplus-killing of reindeer by wolves. J Mamm 57:585CrossRefGoogle Scholar
  4. Boitani L (2003) Wolf conservation and recovery. In: Mech LD, Boitani L (eds) Wolves: behavior, ecology, and conservation. University of Chicago Press, Chicago, pp 317–340Google Scholar
  5. Burnham KP, Anderson DR (2010) Model selection and multi-model inference: a practical information-theoretic approach. Springer, New YorkGoogle Scholar
  6. Calder WA (1984) Size, function, and life history. Harvard University Press, CambridgeGoogle Scholar
  7. Carbyn LN (1983) Wolf predation on elk in riding mountain national park, Manitoba. J Wildlife Manage 47:963–976CrossRefGoogle Scholar
  8. Charnov EL (1976) Optimal foraging, the marginal value theorem. Theor Popul Biol 9:129–136Google Scholar
  9. Chen X, Dickman CR, Thompson MB (2004) Selective consumption by predators of different body regions of prey: is rate of energy intake important? J Zool 264:189–196CrossRefGoogle Scholar
  10. Chew MK, Laubichler MD (2003) Natural enemies: metaphor or misconception? Science 301:52–53PubMedCrossRefGoogle Scholar
  11. Conover RJ (1966) Factors affecting the assimilation of organic matter by zooplankton and the question of superfluous feeding. Limnol Oceanogr 11:346–354CrossRefGoogle Scholar
  12. Cook RM, Cockrell BJ (1978) Predator ingestion rate and its bearing on feeding time and the theory of optimal diets. J Anim Ecol 47:529–547CrossRefGoogle Scholar
  13. Cosner C, DeAngelis DL, Ault JS, Olson DB (1999) Effects of spatial grouping on the functional response of predators. Theor Popul Biol 56:65–75PubMedCrossRefGoogle Scholar
  14. DelGiudice GD (1998) Surplus killing of white-tailed deer by wolves in north-central Minnesota. J Mammal 79:227–235CrossRefGoogle Scholar
  15. Ehrlinge S, Bergsten B, Kristiansson B (1974) The stoat and its prey: hunting behavior and escape reactions. Fauna Flora 69:203–211Google Scholar
  16. Eide SH, Ballard WB (1982) Apparent case of surplus killing of caribou by gray wolves. Can Field-Nat 96:87–88Google Scholar
  17. Fuller TK, Mech LD, Cochrane JF (2003) Wolf population dynamics. In: Mech LD, Boitani L (eds) Wolves: behavior, ecology, and conservation. University of Chicago Press, Chicago, pp 161–191Google Scholar
  18. Gillisa DM, Pikitchb EK, Petermanc RM (1995) Dynamic discarding decisions: foraging theory for high-grading in a trawl fishery. Behav Ecol 6:146–154CrossRefGoogle Scholar
  19. Griffiths D (1982) Test of alternative models of prey consumption by predators, using antlion larvae. J Anim Ecol 51:363–373CrossRefGoogle Scholar
  20. Haberl W (1998) Prey handling times and partial prey consumption in five species of European shrews (Soricidae, Insectivora). Pak J Biol Sci 1:53–54CrossRefGoogle Scholar
  21. Holling CS (1966) The functional response of invertebrate. Predators to prey density. Mem Entomol Soc Can 48:1–86CrossRefGoogle Scholar
  22. Johnson DM, Akre BG, Crowley PH (1975) Modeling arthropod predation: wasteful killing by damselfly naiads. Ecology 56:1080–1093Google Scholar
  23. Jost C, Devulder G, Vucetich JA, Peterson RO, Arditi R (2005) The wolves of Isle Royale display scale-invariant satiation and density dependent predation on moose. J Anim Ecol 74:809–816CrossRefGoogle Scholar
  24. Juliano SA (1989) Queuing models of predation and the importance of contingent behavioural choices for optimal foragers. Anim Behav 38:757–770CrossRefGoogle Scholar
  25. Krebs CJ, Boutin S, Boonstra R (2001) Ecosystem dynamics of the boreal forest: the Kluane project. Oxford University Press, OxfordGoogle Scholar
  26. Kruuk H (1972) Surplus killing by carnivores. J Zool 166:233–244CrossRefGoogle Scholar
  27. Lang A, Klarenbeg AJ (1997) Experiments on the foraging behaviour of the hunting spider Pisaura mirabilis (Araneae: Pisauridae): utilization of single prey items. Eur J Entomol 94:453–459Google Scholar
  28. Loiterton SJ, Magrath RD (1996) Substrate type affects partial prey consumption by larvae of the antlion, Myrmeleon acer (Neuroptera: Myrmeleontidae). Aust J Zool 44:589–597CrossRefGoogle Scholar
  29. Lounibos LP, Makhni S, Alto B, Kesavaraju B (2008) Surplus killing by predatory larvae of Corethrella appendiculata: prepupal timing and site-specific attack on mosquito prey. J Insect Behav 21:47–54PubMedCrossRefGoogle Scholar
  30. Lucas JR (1985) Partial prey consumption by antlion larvae. Anim Behav 33:945–958CrossRefGoogle Scholar
  31. Lucas JR, Grafen A (1985) Partial prey consumption by ambush predators. J Theor Biol 113:455–473CrossRefGoogle Scholar
  32. Maupin JL, Riechert SE (2001) Superfluous killing in spiders: a consequence of adaptation to food-limited environments? Behav Ecol 12:569–576CrossRefGoogle Scholar
  33. McCay CM (1949) Nutrition of the dog. Comstock, IthacaGoogle Scholar
  34. Mech LD (1966) The wolves of Isle Royale. Natl. Park Service Sci. Monogr. Ser. No. 7. US Government Printing Office, Washington, DCGoogle Scholar
  35. Mech LD, Frenzel LD (eds) (1971) Ecological studies of the timber wolf on northeastern Minnesota. USDA Forest Service Research Paper NC-52. North Central Forest Experiment Station, St. Paul, 62 ppGoogle Scholar
  36. Mech LD, Peterson RO (2003) Wolf–prey relationships. In: Mech LD, Boitani L (eds) Wolves: behavior, ecology, and conservation. University of Chicago Press, Chicago, pp 131–160Google Scholar
  37. Mech LD, Adams LG, Meier TJ, Burch JW, Dale BW (1998) The wolves of Denali. University of Minnesota Press, MinneapolisGoogle Scholar
  38. Metz JAJ, Sabelis MW, Kuchlein JH (1988) Sources of variation in predation rates at high prey densities: an analytic model and a mite example. Exp Appl Acarol 5:187–205CrossRefGoogle Scholar
  39. Miller FL, Gunn A, Broughton E (1985) Surplus killing as exemplified by wolf predation on newborn caribou. Can J Zool 63:295–300CrossRefGoogle Scholar
  40. Mittler J (1997) What happens when predators do not completely consume their prey? Theor Popul Biol 51:238–251PubMedCrossRefGoogle Scholar
  41. Nakamura K (1974) A model of the functional response of a predator to prey density involving the hunger effect. Oecologia 16:265–278CrossRefGoogle Scholar
  42. Nakamura K (1977) A model for the functional response of a predator to varying prey densities; based on the feeding ecology of wolf spiders. Bull Nat Inst Agric Sci 31:29–89Google Scholar
  43. Oksanen T, Oksanen L, Fretwell SD (1985) Surplus killing in the hunting strategy of small predators. Am Nat 126:328–346CrossRefGoogle Scholar
  44. Patterson BR (1994) Surplus killing of white-tailed deer, Odocoileus virginianus, by coyotes, Canis latrans, in Nova Scotia. Can Field-Nat 108:484–487Google Scholar
  45. Peterson RO (1977) Wolf ecology and prey relationships on Isle Royale. National Park Service Scientific Monograph Series Number 11. US Government Printing Office, Washington, DCGoogle Scholar
  46. Peterson RO, Allen DL (1974) Snow conditions as a parameter in moose–wolf relationships. Nat Can 101:481–492Google Scholar
  47. Peterson RO, Ciucci P (2003) The wolf as a carnivore. In: Mech LD, Boitani L (eds) Wolves: behavior, ecology, and conservation. University of Chicago Press, Chicago, pp 104–130Google Scholar
  48. Peterson RO, Page RE (1988) The rise and fall of Isle Royale wolves, 1975–1986. J Mammal 69:89–99CrossRefGoogle Scholar
  49. Peterson RO, Thomas NJ, Thurber JM, Vucetich JA, Waite TA (1998) Population limitation and the wolves of Isle Royale. J Mammal 79:487–841CrossRefGoogle Scholar
  50. Pimlott DH, Shannon JA, Kolenosky GB (1969) The ecology of the timber wolf in Algonquin Provincial park, Ontario. Research Report (Wildlife), No. 87. Ontario Department of Lands and Forests, Toronto, 92 ppGoogle Scholar
  51. Pollard SD (1989) Constraints affecting partial prey consumption by a crab spider, Diea sp. indet. (Araneae, Thomisidae). Oecologia 81:392–396Google Scholar
  52. Post E, Peterson RO, Stenseth NC, McLaren BE (1999) Ecosystem consequences of wolf behavioural response to climate. Nature 401:905–907CrossRefGoogle Scholar
  53. Rathje WL (1984) The garbage decade. Amer Behav Sci 28:9–29CrossRefGoogle Scholar
  54. Reynolds PE, Reynolds HV, Shideler R (2002) Predation and multiple kills of muskoxen by grizzly bears. Ursus 13:79–84Google Scholar
  55. Samu F (1993) Wolf spider feeding strategies: optimality of prey consumption in Pardosa hortensis. Oecologia 94:139–145CrossRefGoogle Scholar
  56. Sand H, Zimmermann B, Wabakken P, Andrèn H, Pedersen HC (2005) Using GPS technology and GIS cluster analyses to estimate kill rates in wolf-ungulate ecosystems. Wildl Soc Bull 33:914–925CrossRefGoogle Scholar
  57. Sih A (1980) Optimal foraging: partial consumption of prey. Am Nat 116:281–290CrossRefGoogle Scholar
  58. Vucetich JA, Peterson RO (2004a) The influence of prey consumption and demographic stochasticity on population growth rate of Isle Royale wolves (Canis lupus). Oikos 107:309–320CrossRefGoogle Scholar
  59. Vucetich JA, Peterson RO (2004b) The influence of top-down, bottom-up, and abiotic factors on the moose (Alces alces) population of Isle Royale. Proc R Soc Lond B 271:183–189CrossRefGoogle Scholar
  60. Vucetich JA, Peterson RO, Schaefer CL (2002) The effect of prey and predator densities on wolf predation. Ecology 83:3003–3013CrossRefGoogle Scholar
  61. Vucetich JA, Peterson RO, Waite TA (2004) Raven scavenging favours group foraging in wolves. Anim Behav 67:1117–1126CrossRefGoogle Scholar
  62. Vucetich JA, Hebblewhite M, Smith DW, Peterson RO (2011) Predicting prey population dynamics from kill rate, predation rate and predator–prey ratios in three wolf-ungulate systems. J Animal Ecol. doi: 10.1111/j.1365-2656.2011.01855
  63. Young SP, Goldman EA (1944) The wolves of North America. American Wildlands Institute, Washington, DCGoogle Scholar

Copyright information

© Springer-Verlag 2011

Authors and Affiliations

  • John A. Vucetich
    • 1
    Email author
  • Leah M. Vucetich
    • 1
  • Rolf O. Peterson
    • 1
  1. 1.School of Forest Resources and Environmental ScienceMichigan Technological UniversityHoughtonUSA

Personalised recommendations