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Brown bear body mass and growth in northern and southern Europe

  • Population Ecology
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Abstract

We tested six hypotheses to explain expected geographical differences in body masses of 1,771 brown bears (Ursus arctos) from northern and southern Europe (Sweden and Norway compared with Slovenia and Croatia): Bergmann’s rule, the fasting endurance hypothesis, and the dietary meat hypothesis, which predicted larger bears in the north; and hypotheses stressing the role of high primary productivity, high population density, low seasonality, and length of the growing season, which predicted larger bears in the south. Although brown bear populations in North America vary greatly in body mass, we found no significant difference in body mass between the two European populations using a new analytical approach incorporating modeled age-standardized body masses in linear models, when correcting for sex and season. The greater variation in North America may be due primarily to the presence of large bears that feed on salmon (Oncorhynchus spp.), which does not occur in Europe. Asymptotic body masses were 115 ± 9 (SE) kg in spring and 141 ± 9 kg in autumn for southern females, 248 ± 25 and 243 ± 24 kg for southern males, 96 ± 2 and 158 ± 4 kg for northern females, and 201 ± 4 and 273 ± 6 kg for northern males, respectively. Northern bears gained more body mass before hibernation and lost more during hibernation than southern bears, probably because hibernation was twice as long in the north. Northern bears gained and southern bears lost mass during the spring, perhaps due to the greater availability and use of protein-rich food in spring in the north. As reproductive success in bears is correlated with adult female body mass in interpopulation comparisons, brown bears may have relatively similar reproductive rates throughout Europe, although minimum age at primiparity and litter interval are lower in the south.

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References

  • Adamič M (1997) The analysis of key sources of mortality of the brown bear (Ursus arctos L.) in Slovenia in the last 6-year period (1.4.1991–31.3.1997). Research reports, UNI Ljubljana, Biotechnical Faculty, Ljubljana (in Slovene with English summary). For Wood Sci Technol 53:5–28

  • Almăsan HA, Vasiliu GD (1967) Zur Kenntnis des Rumänischen Karpatenbär. Acta Theriol 12:47–66

    Google Scholar 

  • Atkinson SN, Ramsay MA (1995) The effects of prolonged fasting on the body composition and reproductive success of female polar bears (Ursus maritimus). Funct Ecol 9:559–567

    Article  Google Scholar 

  • Bergmann C (1847) Über die Verhaltnisse der Wärmökonomie der Thiere zu ihrer Grösse. Goettinger Studien 3:595–708

    Google Scholar 

  • Blanchard BM (1987) Size and growth patterns of the Yellowstone grizzly bear. Proc Int Conf Bear Res Manage 7:99–107

    Google Scholar 

  • Boyce MS (1978) Climatic variability and body size variation in the muskrat (Ondatra zibethicus) in North America. Oecologia 36:1–19

    Article  Google Scholar 

  • Burnham KP, Anderson DR (2002) Model selection and multimodel inference: a practical information-theoretic approach. Springer, Berlin

  • Cicnjak L, Huber D, Roth HU, Ruff RL, Vinovrski Z (1987) Food habits of brown bears in Plitvice Lakes National Park, Yugoslavia. Proc Int Conf Bear Res Manage 7:221–226

    Google Scholar 

  • Clutton-Brock TH, Guinness FE, Albon SD (1982) Red deer: behaviour and ecology of two sexes. Edinburgh University Press, Edinburgh

    Google Scholar 

  • Dahle B, Sørensen OJ, Wedul EH, Swenson JE, Sandegren F (1998) The diet of brown bears Ursus arctos in central Scandinavia: effect of access to free-ranging domestic sheep Ovis aries. Wildlife Biol 4:147–158

    Google Scholar 

  • Dahle B, Swenson JE (2003) Home ranges in adult Scandinavian brown bears Ursus arctos: effect of population density, mass, sex, reproductive status and habitat type. J Zool 260:329–335

    Article  Google Scholar 

  • Derocher AE, Stirling I (1994) Age-specific reproductive performance of female polar bears (Ursus maritimus). J Zool London 234:527–536

    Article  Google Scholar 

  • Derocher AE, Stirling I (1996) Aspects of survival in juvenile polar bears. Can J Zool 74:1246–1252

    Google Scholar 

  • Derocher AE, Stirling I (1998) Geographic variation in growth of polar bears (Ursus maritimus). J Zool London 245:65–72

    Google Scholar 

  • Derocher AE, Wiig Ø (2002) Postnatal growth in body length and mass of polar bears (Ursus maritimus) at Svalbard. J Zool London 256:343–349

    Google Scholar 

  • Elgmork K, Kaasa J (1992) Food habits and foraging of the brown bear (Ursus arctos) in central south Norway. Ecography 15:101–110

    Article  Google Scholar 

  • Ferguson SH, McLoughlin PD (2000) Effect of energy availability, seasonality, and geographic range on brown bear life history. Ecography 23:193–200

    Article  Google Scholar 

  • Friebe A, Swenson JE, Sandegren F (2001) Denning chronology of female brown bears in central Sweden. Ursus 12:37–46

    Google Scholar 

  • Frković A, Huber D, Kusak J (2001) Brown bear litter sizes in Croatia. Ursus 12:103–105

    Google Scholar 

  • Fujita R (2000) Bait-hunting for brown bear in Sweden: temporal and spatial occurrence and potential effects on the population. Undergraduate thesis, Swedish University of Agricultural Sciences, Uppsala

  • Große C (1999) Ants—an important food for brown bears (Ursus arctos) in Slovenia? Diploma thesis, Phillips-University of Marburg, Germany

  • Große C, Kaczensky P, Knauer F (2003) Ants: a food source sought by Slovenian brown bears (Ursus arctos)? Can J Zool 81:1996–2005

    Article  Google Scholar 

  • Hell P (1992) Biometrie und Bewirtshaftung der Population des Braunbären im Slowakischen Teil der Westkarpaten. In: F. Bourlière (President of the Scientific Committee) Management and restoration of small and relictual bears populations. Museum d’Historie Naturelle, Grenoble, France, pp 143–153

  • Hilderbrand GV, Jenkins SG, Schwartz CC, Hanley TA, Robbins CT (1999b) Effect of seasonal differences in dietary meat intake on changes in body mass and composition in wild and captive brown bears. Can J Zool 77:1623–1630

    Article  Google Scholar 

  • Hilderbrand GV, Schwartz CC, Robbins CT, Hanley TA (2000) Effect of hibernation and reproductive status on body mass and condition of coastal brown bears. J Wildlife Manage 64:178–183

    Article  Google Scholar 

  • Hilderbrand GV, Schwartz CC, Robbins CT, Jacoby ME, Hanley TA, Arthur SM, Servheen C (1999a) Importance of dietary meat, particularly salmon, to body size, population productivity, and conservation of North American brown bears. Can J Zool 77:132–138

    Article  Google Scholar 

  • Huber D, Frković A (1993) Brown bear management in Croatia. Proc Cong Int Union Game Biol 21(1):287–292

    Google Scholar 

  • Huber D, Roth HU (1993) Movements of European brown bears in Croatia. Acta Theriol 38:151–159

    Google Scholar 

  • Huber D, Roth HU (1997) Denning of brown bears in Croatia. Proc Int Conf Bear Res Manage 9(2):79–83

    Google Scholar 

  • Insightful Corp. (2003) S-PLUS 6.2. Insightful Corp., Seattle, WA. http://www.insightful.com/support/splus62win/. Cited 16 March 2007

  • Kingsley MCS, Nagy JA, Reynolds HV (1988) Growth in length and weight of northern brown bears: differences between sexes and populations. Can J Zool 66:981–986

    Google Scholar 

  • Kojola I, Laitala HM (2001) Body size variation of brown bear in Finland. Ann Zool Fennici 38:173–178

    Google Scholar 

  • Laws RM (1956) Growth and sexual maturity in aquatic mammals. Nature 178:193–194

    Article  Google Scholar 

  • Lindstedt SL, Boyce MS (1984) Seasonality, fasting endurance, and body size in mammals. Am Nat 125:873–878

    Article  Google Scholar 

  • Mahoney SP, Virgl JA, Mawhinney K (2001) Potential mechanisms of phenotypic divergence in body size between Newfoundland and mainland black bear populations. Can J Zool 79:1650–1660

    Article  Google Scholar 

  • Manchi S, Swenson JE (2005) Denning behavior of male Scandinavian brown bears (Ursus arctos). Wildlife Biol 11:123–132

    Article  Google Scholar 

  • Matson GM, Van Daele L, Goodwin E, Aumiller L, Reynolds H, Hristienko H (1993) A laboratory manual for cementum age determination of Alaska brown bear PM1 teeth. Alaska Department of Fish and Game/Matson’s Laboratory, Milltown, MT

  • Mayr E (1956) Geographical character gradients and climatic adaptation. Evolution 10:105–108

    Article  Google Scholar 

  • McLellan B (1994) Density-dependent population regulation of brown bears. In: Taylor M (ed) Density-dependent population regulation in black, brown, and polar bears. Int Conf Bear Res Manage Monogr Ser 3:15–24

  • McNab BK (1971) On the ecological significance of the Bergmann’s rule. Ecology 52:845–854

    Article  Google Scholar 

  • McNab BK (1989) Basal rate of metabolism, body size and food habits in the order Carnivora. In: Gittleman JL (ed) Carnivore behaviour, ecology and evolution. Cornell University Press, Ithaca, NY, pp 335–354

  • Naves J, Palomero G (1993) Brown bear hibernation ecology in the Cantabrian Mountains. In: Naves J, Palomero G (eds) El oso pardo (Ursus arctos) en España. Instituto Nacional para la Conservacion de la Naturaleza, Madrid (in Spanish with English summary), pp 147–181

  • Noyce KV, Garshelis DL (1998) Spring weight changes in black bears in northcentral Minnesota: the negative foraging period revisited. Ursus 10:521–531

    Google Scholar 

  • Opseth O (1998) Brown bear (Ursus arctos) diet and predation on moose (Alces alces) calves in the southern taiga zone in Sweden. Cand Sci Thesis, Norwegian University of Science and Technology, Trondheim

  • Persson IL, Wikan S, Swenson JE, Mysterud I (2001) The diet of the brown bear in the Pasvik, Valley, northeastern Norway. Wildlife Biol 7:27–37

    Google Scholar 

  • Peters RH (1983) The ecological implications of body size. Cambridge University Press, Cambridge

    Google Scholar 

  • Pritchard GT, Robbins CT (1990) Digestive and metabolic efficiencies of grizzly and black bears. Can J Zool 68:1645–1651

    Article  Google Scholar 

  • Rode KD, Robbins CT, Shipley LA (2001) Constraints on herbivory by grizzly bears. Oecologia 128:62–71

    Article  Google Scholar 

  • Rogers L (1976) Effects of mast and berry crop failure on survival, growth, and reproductive success of black bears. Tans North Am Wildlife Nat Resour Conf 41:431–438

    Google Scholar 

  • Rosenzweig ML (1968) Net primary productivity of terrestrial communities: prediction from climatological data. Am Nat 102:67–74

    Article  Google Scholar 

  • Roth HU, Boscagli G, Gentile L (1992) Movements, activity and hibernation of brown bears in the Abruzzo National Park as revealed by radiotelemetry. In: Bourlière F (ed) Management and restoration of small and relictual bears populations. Museum d’Historie Naturelle, Grenoble, France

  • Samson C, Huot J (1995) Reproductive biology of female black bears in relation to body mass in early winter. J Mammal 76:68–77

    Article  Google Scholar 

  • Sæther BE, Engen S, Swenson JE, Bakke Ø, Sandegren F (1998) Assessing the viability of Scandinavian brown bear, Ursus arctos, populations: the effects of uncertain parameter estimates. Oikos 83:403–416

    Article  Google Scholar 

  • Sadleir RMFS (1969) The ecology of reproduction in wild and domestic mammals. Methuen, London

  • Salvatori V, Okarma H, Ionescu O, Dovhanych Y, Find’o S, Boitani L (2002) Hunting legislation in the Carpathian Mountains: implications for the conservation and management of large carnivores. Wildlife Biol 8:3–10

    Google Scholar 

  • Schwartz CC, Franzmann AW (1991) Interrelationship of black bears to moose and forest succession in the northern coniferous forest. Wildlife Monogr 113

  • Schwartz CC, Miller SD, Haroldson MA (2003) Grizzly bear. In: Feldhamer GA, Thompson BC, Chapman JA (eds) Wild mammals of North America: biology, management, and conservation, 2nd edn. John Hopkins University Press, Baltimore, MD, pp 556–586

  • Simonič A (1994) The legal protection of the brown bear in Slovene territory—past and present, and some suggestions for the future. In: Adamič M (ed) Rjavi medved v deželah Alpe-Adria. Gozdarski inštitut Slovenije, Ministrostvo za kmetijstvo in gozdarstvo, Republike Slovenije. Ljubljana, Slovenia, pp 43–75

    Google Scholar 

  • SPSS Inc. (2004) SPSS Release 13.01. SPSS Inc., Chicago, IL. http://www.spss.com/. Cited 16 March 2007

  • Stearns SC (1992) The evolution of life histories. Oxford University Press, Oxford

    Google Scholar 

  • Stearns SC, Koella J (1986) The evolution of phenotypic plasticity in life history traits: predictions from norms of reaction for age- and size-at-maturity. Evolution 34:893–913

    Article  Google Scholar 

  • Stringham SF (1990a) Black bear reproductive rate relative to body weight in hunted populations. Proc Int Conf Bear Res Manage 8:425–432

    Google Scholar 

  • Stringham SF (1990b) Grizzly bear reproductive rate relative to body size. Proc Int Conf Bear Res Manage 8:433–443

    Google Scholar 

  • Swenson JE, Dahle B, Sandegren F (2001) Intraspecific predation in Scandinavian brown bears older than cubs-of-the-year. Ursus 12:81–91

    Google Scholar 

  • Swenson JE, Sandegren F (2000) Conservation of European brown bear populations: experiences from Scandinavia. In: Layna JF, Heredia B, Palomero G, Doadrio I (eds) La conservación del oso pardo en Europa: un reto de cara al siglo XXI (Serie encuentros I). Fundación Biodiversidad, Ministerio de medio ambiente, Madrid, Spain, pp 111–116

  • Swenson JE, Sandegren F, Bjärvall, Söderberg A., Wabakken P, Franzén R (1994) Size, trend, distribution and conservation of the brown bear Ursus arctos population in Sweden. Biol Conserv 70:9–17

    Article  Google Scholar 

  • Swenson JE, Sandegren F, Söderberg A, Franzén R (1995) Estimating the total weight of Scandinavian brown bears from field-dressed and slaughter weights. Wildlife Biol 1:177–179

    Google Scholar 

  • Von Bertalanffy L (1938) A quantitative theory of organic growth (Inquiries on growth laws II). Human Biol 10:181–213

    Google Scholar 

  • Zedrosser A, Dahle B, Swenson JE (2006) Population density and food conditions determine adult female size in brown bears. J Mammal 87:510–518

    Article  Google Scholar 

  • Zedrosser A, Rauer G, Kruckenhauser L (2004) Early primiparity in brown bears. Acta Theriol 49:427–432

    Google Scholar 

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Acknowledgments

We especially thank Steinar Engen, who designed our model and gave us statistical advice, Petra Kaczensky for providing data on denning dates in Slovenia, and Andrew Derocher and Nigel Yoccoz for useful comments on earlier versions of the manuscript. The Scandinavian Brown Bear Research Project has been supported by the Swedish Environmental Protection Agency, Norwegian Directorate for Nature Management, Swedish Association for Hunting and Wildlife Management, WWF-Sweden, Research Council of Norway, and several private foundations. The bear research in Slovenia was funded by the Ministry of Education, Science and Sports and by the Ministry of Agriculture and Forestry. Teeth (P1) for ageing have been extracted by the district wildlife officers of the State Forest Service during the course of regular inspections of dead brown bears. The studies of brown bears in Croatia have been supported by the Croatian Ministry for Science and Technology, the National Geographic Society, the International Association for Bear Research, and Management (Bevins Fund), as well as by local forestry offices.

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Correspondence to Jon E. Swenson.

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Communicated by Jean-Michel Gaillard.

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Swenson, J.E., Adamič, M., Huber, D. et al. Brown bear body mass and growth in northern and southern Europe. Oecologia 153, 37–47 (2007). https://doi.org/10.1007/s00442-007-0715-1

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