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Summer kill rates and predation pattern in a wolf–moose system: can we rely on winter estimates?

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Abstract

So far the vast majority of studies on large carnivore predation, including kill rates and consumption, have been based on winter studies. Because large carnivores relying on ungulates as prey often show a preference for juveniles, kill rates may be both higher and more variable during the summer season than during the rest of the year leading to serious underestimates of the total annual predation rate. This study is the first to present detailed empirical data on kill rates and prey selection in a wolf–moose system during summer (June–September) as obtained by applying modern Global Positioning System-collar techniques on individual wolves (Canis lupus) in Scandinavia. Moose (Alces alces) was the dominant prey species both by number (74.4%) and biomass (95.6%); 89.9% of all moose killed were juveniles, representing 76.0% of the biomass consumed by wolves. Kill rate in terms of the kilogram biomass/kilogram wolf per day averaged 0.20 (range: 0.07–0.32) among wolf territories and was above, or well above, the daily minimum food requirements in most territories. The average number of days between moose kills across wolf territories and study periods was 1.71 days, but increased with time and size of growing moose calves during summer. Over the entire summer (June–September, 122 days), a group (from two to nine) of wolves killed a total of 66 (confidence interval 95%; 56–81) moose. Incorporation of body growth functions of moose calves and yearlings and wolf pups over the summer period showed that wolves adjusted their kill rate on moose, so the amount of biomass/kilogram wolf was relatively constant or increased. The kill rate was much higher (94–116%) than estimated from the winter period. As a consequence, projecting winter kill rates to obtain annual estimates of predation in similar predator–prey systems may result in a significant underestimation of the total number of prey killed.

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References

  • Alfredéen AC (2006) Denning behaviour and movement pattern during summer of wolves Canus lupus on the Scandinavian Peninsula. Thesis no. 164, Department of Conservation Biology, Swedish University of Agricultural Sciences

  • Anderson CR, Lindzey FG (2003) Estimating cougar predation rates from GPS location clusters. J Wildl Manage 67:307–316

    Article  Google Scholar 

  • Arnemo JM, Ahlqvist P, Segerström P (2004) Biomedical protocol for free-ranging wolves (Canis lupus): capture, anaesthesia, surgery, yagging, sampling and necropsy procedures. http://skandulv.nina.no/skandulv%20new/Publikasjoner/publikasjoner.htm

  • Ballard WB, Whitman JS, Gardner CL (1987) Ecology of an exploited wolf population in south-central Alaska. Wildl Monogr 98:1–54

    Google Scholar 

  • Ballard WB, Ayres LA, Krausman PR, Reed DJ, Fancy SG (1997) Ecology of wolves in relation to a migratory caribou herd in northwest Alaska. Wildl Monogr 135:47

    Google Scholar 

  • Cederlund GN, Sand HKG (1991) Population dynamics and yield of a moose population without predators. Alces 27:31–40

    Google Scholar 

  • Dale BW, Adams LG, Bowyer RT (1994) Functional response of wolves preying on barren-ground caribou in a multi-prey ecosystem. J Anim Ecol 63:644–652

    Article  Google Scholar 

  • Dale BW, Adams LG, Bowyer TB (1995) Winter wolf predation in a multiple ungulate prey system, gates of the Arctic National Park, Alaska. In: Carbyn LN, Fritts SH, Seip DR (eds) Ecology and conservation of wolves in a changing world. Canadian Circumpolar Institute, University of Alberta, Edmonton, pp 223–230

    Google Scholar 

  • Eberhardt LL, Garrot RA, Smith DW, White PJ, Peterson RO (2003) Assessing the impact of wolves on ungulate prey. Ecol Appl 13:776–783

    Article  Google Scholar 

  • Floyd TJ, Mech LD, Jordan PA (1978) Relating wolf scat content to prey consumed. J Wildl Manage 42:528–532

    Article  Google Scholar 

  • Fritts SH, Mech LD (1981) Dynamics, movements and feeding ecology of a newly protected wolf population in north-western Minnesota. Wildl Monogr 80:79

    Google Scholar 

  • Fuller TK (1989) Population dynamics of wolves in north-central Minnesota. Wildl Monogr 105:41

    Google Scholar 

  • Hayes RD, Baer AM, Wotschikowsky U, Harestad AS (2000) Kill rate by wolves on moose in the Yukon. Can J Zool 78:49–59

    Article  Google Scholar 

  • Hebblewhite M, Pletcher DH, Paquet PC, Lessard RB, Callaghan CJ (2003) Development and application of a ratio estimator to estimate wolf kill rates and variance in a multiple-prey system. Wildl Soc Bull 31:933–946

    Google Scholar 

  • Holling CS (1959) The components of predation as revealed by a study of small-mammal predation of the European sawfly. Can Entomol 91:293–320

    Article  Google Scholar 

  • Huggard DJ (1993) Effects of snow depth on predation and scavenging by gray wolves. J Wildl Manage 57:382–388

    Article  Google Scholar 

  • Jedrzejewski J, Schmidt K, Theuerkauf J, Jedrzejewska B, Selva N, Zub K, Szymura L (2002) Kill rates and predation by wolves on ungulate populations in Bialowieza primeval forest (Poland). Ecology 83:1341–1356

    Article  Google Scholar 

  • Knappwost U (2006) Territorial variation in wolves diet? A comparison of 11 territories in Sweden. Diploma thesis, School of Forest Science and Resource Management, München, Germany

  • Kolenosky GB (1972) Wolf predation on wintering deer in east-central Ontario. J Wildl Manage 36:357–369

    Article  Google Scholar 

  • Kunkel KE, Mech LD (1994) Wolf and bear predation on white-tailed deer and fawns in north-eastern Minnesota. Can J Zool 72:1557–1565

    Article  Google Scholar 

  • Kunkel KE, Pletcher DH (2000) Habitat factors affecting vulnerability of moose to predation by wolves in south-eastern British Columbia. Can J Zool 78:150–157

    Article  Google Scholar 

  • Markgren G (1969) Reproduction of moose in Sweden. Swed Wildl 6:127–299

    Google Scholar 

  • Mech LD (1966) The wolves of Isle Royale. US National Parks Fauna series no. 7, Washington, DC

  • Mech LD (1971) Wolves, coyotes and dogs. In: Nelson M (ed) Symposium on the white-tailed deer in Minnesota. Minnesota Department of Natural Resources, St Paul, pp 19–22

    Google Scholar 

  • Mech LD (1977) Population trends and winter deer consumption in a Minnesota wolf pack. In: Phillips RL, Jonkel C (eds) Proceedings of the 1975 predator symposium. Montana forest and conservation experiment station, University of Montana, Missoula, pp 55–83

    Google Scholar 

  • Mech LD (1988) The Arctic wolf: living with the pack, 2nd edn. Voyageur, Stillwater

    Google Scholar 

  • Mech LD, Peterson RO (2003) Wolf–prey relations. In: Mech LD, Boitani L (eds) Wolves: behaviour, ecology and conservation. University of Chicago Press, Chicago, pp 104–130

    Google Scholar 

  • Mech LD, Adams LG, Meier TJ, Burch JW, Dale BW (1998) The wolves of Denali. University of Minnesota Press, Minneapolis

    Google Scholar 

  • Messier F (1994) Ungulate population models with predation: a case study with North American moose. Ecology 75:478–488

    Article  Google Scholar 

  • Messier F, Crête M (1985) Moose–wolf dynamics and the natural regulation of moose populations. Oecologia 65:503–512

    Article  Google Scholar 

  • Müller S (2006) Diet composition of wolves (Canis lupus) on the Scandinavian Peninsula determined by scat analyses. Diploma thesis, School of Forest Science and Resource Management, München, Germany

  • Nelson ME, Mech LD (1981) Deer social organisation and wolf predation in northeastern Minnesota. Wildl Monogr 77:53

    Google Scholar 

  • Nelson ME, Mech LD (1986) Mortality of white-tailed deer in northeastern Minnesota. J Wildl Manage 50:691–698

    Article  Google Scholar 

  • Olsson O, Wirtberg J, Andersson M, Wirtberg I (1997) Wolf Canis lupus predation on moose Alces alces and roe deer Capreolus capreolus in south-central Scandinavia. Wildl Biol 3:13–25

    Google Scholar 

  • Peterson RO (1977) Wolf ecology and prey relationships on Isle Royale. US Nat Park Serv Sci Monogr 11:1–210

    Google Scholar 

  • Peterson RO, Allen DL (1974) Snow conditions as a parameter in moose-wolf relationships. Nat Can 101:481–492

    Google Scholar 

  • 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 104–130

    Google Scholar 

  • Peterson RO, Woolington JD, Bailey TN (1984) Wolves of the Kenai Peninsula, Alaska. Wildl Monogr 88:1–52

    Google Scholar 

  • Peterson RO, Thomas NJ, Thurber JM, Vucetich JA, Waite TA (1998) Population limitation and the wolves of Isle Royale. J Mammal 79:828–841

    Article  Google Scholar 

  • Promberger C (1992) Wölfe und Scavenger (English abstract). Diplomarbeit, Ludwig Maximilians Universität, Munchen

  • Rühe F, Buschmann I, Wameling A (2003) Two models for assessing the prey mass of European ungulates from wolf scats. Acta Theriol 48:527–537

    Google Scholar 

  • Saether BE, Andersen R, Hjeljord O, Heim M (1996) Ecological correlates of regional variation in life history of the moose Alces alces. Ecology 77:1493–1500

    Article  Google Scholar 

  • Sand H (1996) Life history strategies in moose (Alces alces): geographical and temporal variation in body growth and reproduction. PhD thesis, Swedish University of Agricultural Sciences, Uppsala

  • Sand H, Zimmermann B, Wabakken P, Andrèn H (2005) Using GPS-technology and GIS-cluster analyses to estimate kill rates in wolf-ungulate ecosystems. Wildl Soc Bull 33:914–925

    Article  Google Scholar 

  • Sand H, Wikenros C, Wabakken P, Liberg O (2006a). Effects of hunting group size, snow depth and age on the success of wolves hunting moose. Anim Behav 72:781–789

    Article  Google Scholar 

  • Sand H, Wikenros C, Wabakken P, Liberg O (2006b) Cross-continental differences in patterns of predation: will naïve moose in Scandinavian ever learn? Proc R Soc Lond B 273:1–7

    Article  Google Scholar 

  • Schmidt PA, Mech LD (1997) Wolf pack size and food acquisition. Am Nat 150:513–517

    Article  PubMed  CAS  Google Scholar 

  • Smith DW, Drummer TD, Murphy KM, Guernsey DS, Evans SB (2004) Winter prey selection and estimation of wolf kill rates in Yellowstone National Park 1995–2000. J Wildl Manage 68:153–166

    Article  CAS  Google Scholar 

  • Solberg EJ, Sand H, Linnell JD, Brainerd SM, Andersen R, Odden J, Brøseth H, Swenson JE, Strand O, Wabakken P (2003) The effects of large carnivores on wild ungulates in Norway: implications for ecological processes, harvest and hunting methods (in Norwegian with English summary). NINA Fagrapport 63, Trondheim, Norway

  • Solberg EJ, Grotan V, Rolandsen CM, Brøseth H, Brainerd S (2005) Changes-in-sex ratio as an estimator of population size for Norwegian moose Alces alces. Wildl Biol 11:163–172

    Article  Google Scholar 

  • Solomon ME (1949) The natural control of animal populations. J Anim Ecol 18:1–135

    Article  Google Scholar 

  • Statistics Norway (2003) Statistisk Sentralbyrå Befolkning. Publikasjoner, Oslo

    Google Scholar 

  • Swedish National Atlas (1991) Befolkningen. Svensk National Atlas, Stockholm

    Google Scholar 

  • Swenson JE, Sandegren F, Söderberg A (1998) Geographic expansion of an increasing brown bear population: evidence for pre-saturation dispersal. J Anim Ecol 67:819–826

    Article  Google Scholar 

  • Swenson JE, Dahle B, Busk H, Opseth O, Johansen T, Wallin K, Cederlund G (2006) Predation on moose calves by European brown bears. J Wildl Manage 71:1993–1997

    Article  Google Scholar 

  • Taylor RJ (1984) Predation. Chapman and Hall, New York

    Google Scholar 

  • Thurber JM, Peterson RO (1993) Effects of population density and pack size on the foraging ecology of gray wolves. J Mammal 74:879–889

    Article  Google Scholar 

  • Tremblay JP, Jolicoeur H, Lemleux R (2001) Summer food habits of grey wolves in the boreal forest of the Lac Jacques-Cartier highlands, Quebec. Alces 37:1 –12

    Google Scholar 

  • Wabakken P, Sand H, Liberg O, Bjärvall A (2001) The recovery, distribution, and population dynamics of wolves on the Scandinavian peninsula, 1978–1998. Can J Zool 79:710–725

    Article  Google Scholar 

  • Wabakken P, Aronson Å, Strømseth TH, Sand H, Svensson L, Kojola I (2006) The wolf in Scandinavia: status report of the 2005–2006 winter (in Norwegian with English summary). Oppdragsrapport no. 3–2006, Høgskolen i Hedmark

  • Wam HK, Hjeljord O (2003) Wolf predation on moose––a case study using hunter observations. Alces 30:263–270

    Google Scholar 

  • Weaver JL (1993) Refining the equation for interpreting prey occurrence in gray wolf scats. J Wildl Manage 57:534–538

    Article  Google Scholar 

  • Vucetich J, Peterson RO, Schaefer CL (2002) The effect of prey and predator densities on wolf predation. Ecology 83:9003–3013

    Article  Google Scholar 

  • Zimmermann B, Storaas T, Wabakken P, Nicolaysen K, Steinset OK, Dötterer M, Gundersen H, Andreassen HP (2001) GPS collars with remote download facilities, for studying the economics of moose hunting and moose-wolf interactions. In: Sibbald AM, Gordon IJ (eds) Tracking animals with GPS. Macaulay Institute, Aberdeen, pp 33–38

    Google Scholar 

  • Zimmermann B, Wabakken P, Sand H, Pedersen HC, Liberg O (2007) Wolf movement patterns: a key to estimation of kill rate? J Wildl Manage 71:1177–11

    Article  Google Scholar 

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Acknowledgements

We are indebted to J. M. Arnemo, P. Ahlqvist and P. Segerström, who captured and handled the wolves. C. Wikenros, A. Blixgard, S. E. Bredvold, A. Bye, M. Dötterer, J. Enerud, F. Holen, P. Kirkeby, P. Larsson, E. Maartmann, B. Olsson, H. Rønning, R. Skyrud, T. H. Strømseth, O-K. Steinset carried out excellent fieldwork. The study was supported by the Swedish Environmental Protection Agency, the Swedish Association for Hunting and Wildlife Management, World Wildlife Fund for Nature (Sweden), Swedish University of Agricultural Sciences, Norwegian Directorate for Nature, Management, Norwegian Research Council, Norwegian Institute for Nature Research, Hedmark University College, County Governors of Hedmark and Värmland, Borregaard Skoger, Glommen Skogeierforening, Norskog, Norges Skogeierforbund, Olle and Signhild Engkvists Stiftelser, Carl Tryggers Stiftelse, Swedish Carnivore Association, and Stor-Elvdal, Åmot, Åsnes and Trysil municipalities. All research presented complies with the current laws of the country in which the experiments were performed.

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Correspondence to Håkan Sand.

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Communicated by Janne Sundell.

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Sand, H., Wabakken, P., Zimmermann, B. et al. Summer kill rates and predation pattern in a wolf–moose system: can we rely on winter estimates?. Oecologia 156, 53–64 (2008). https://doi.org/10.1007/s00442-008-0969-2

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