Abstract
Conservation strategies for populations of woodland caribou Rangifer tarandus caribou frequently emphasize the importance of predator–prey relationships and the availability of lichen-rich late seral forests, yet the importance of summer diet and forage availability to woodland caribou survival is poorly understood. In a recent article, Wittmer et al. (Can J Zool 83:407–418, 2005b) concluded that woodland caribou in British Columbia were declining as a consequence of increased predation that was facilitated by habitat alteration. Their conclusion is consistent with the findings of other authors who have suggested that predation is the most important proximal factor limiting woodland caribou populations (Bergerud and Elliot in Can J Zool 64:1515–1529, 1986; Edmonds in Can J Zool 66:817–826, 1988; Rettie and Messier in Can J Zool 76:251–259, 1998; Hayes et al. in Wildl Monogr 152:1–35, 2003). Wittmer et al. (Can J Zool 83:407–418, 2005b) presented three alternative, contrasting hypotheses for caribou decline that differed in terms of predicted differences in instantaneous rates of increase, pregnancy rates, causes of mortality, and seasonal vulnerability to mortality (Table 1, p 258). These authors rejected the hypotheses that food or an interaction between food and predation was responsible for observed declines in caribou populations; however, the use of pregnancy rate, mortality season and cause of mortality to contrast the alternative hypotheses is problematic. We argue here that the data employed in their study were insufficient to properly evaluate a predation-sensitive foraging hypothesis for caribou decline. Empirical data on seasonal forage availability and quality and plane of nutrition of caribou would be required to test the competing hypotheses. We suggest that methodological limitations in studies of woodland caribou population dynamics prohibit proper evaluation of the mechanism of caribou population declines and fail to elucidate potential interactions between top-down and bottom-up effects on populations.
Similar content being viewed by others
References
Adams LG (2005) Effects of maternal characteristics and climatic variation on birth masses of Alaskan caribou. J Mammal 86:506–513
Adams LG, Dale BW, Mech LD (1995a) Wolf predation on caribou calves in Denali National Park, Alaska. In: Carbyn LN, Fritts SH, Seip DR (eds) Ecology and conservation of Wolves in a changing world. Canadian Circumpolar Institute Occasional Publication 35, University of Alberta, Edmonton
Adams LG, Singer FJ, Dale BW (1995b) Caribou calf mortality in Denali National Park, Alaska. J Wildl Manage 59:584–594
Ahti T, Hepburn RL (1967) Preliminary studies on woodland caribou range, especially on lichen stands, in Ontario. Technical Series, Research Report (Wildlife) No. 74. Department of Lands and Forests, Ontario
Albon SD, Langvatn R (1992) Plant phenology and the benefits of migration in a temperate ungulate. Oikos 65:502–513
Apps CD, McLellan BN, Kinley TA, Flaa JP (2001) Scale-dependent habitat selection by mountain caribou, Columbia Mountains, British Columbia. J Wildl Manage 65:65–77
Bergerud AT (1972) Food habits of Newfoundland caribou. J Wildl Manage 36:913–923
Bergerud AT (1974) Decline of caribou in North America following settlement. J Wildl Manage 38:757–770
Bergerud AT, Elliot JP (1986) Dynamics of caribou and wolves in northern British Columbia. Can J Zool 64:1515–1529
Bergerud AT, Butler HE, Miller DR (1984) Antipredator tactics of calving caribou: dispersion in mountains. Can J Zool 62:1566–1575
Boertje RD (1990) Diet quality and intake requirements of adult female caribou of the Denali herd, Alaska. J Appl Ecol 27:420–434
Brown GS, Mallory FF (2007) A review of ungulate nutrition and the role of top-down and bottom-up forces in woodland caribou population dynamics. Technical Bulletin No. 934. National Council for Air and Stream Improvement, Research Triangle Park, NC
Cameron RD, Smith WT, Fancy SG, Gerhart KL, White RG (1993) Calving success of female caribou in relationto body weight. Can J Zool 71:480–486
Cook JC (1990) Habitat, nutrition, and population ecology of two transplanted bighorn sheep populations in southcentral Wyoming. PhD thesis, University of Wyoming, Laramie
Cook JG (2002) Nutrition and food. In: Toweill DE, Thomas JW (eds) North American elk: ecology and management. Smithsonian Institution, Washington D.C.
Cook RC, Cook JG, Murray DL, Zager P, Johnson BK, Gratson MW (2001a) Development of predictive models of nutritional condition for Rocky Mountain elk. J Wildl Manage 65:973–987
Cook RC, Cook JG, Murray DL, Zager P, Johnson BK, Gratson MW (2001b) Nutritional condition models for elk: which are the most sensitive, accurate, and precise? J Wildl Manage 65:988–997
Cook RC, Cook JG, Mech LD (2004a) Nutritional condition of northern Yellowstone elk. J Mammal 85:714–722
Cook JG, Johnson BK, Cook RC, Riggs RA, Delcurto T, Bryant LD, Irwin LL (2004b) Effects of summer-autumn nutrition and parturition date on reproduction and survival of elk. Wildl Monogr 155:1–61
Courtois R (2003) La conservation du caribou dans un contexte de perte d’habitat et de fragmentation du milieu. Thèse de doctorat, Université du Quebec à Rimouski, Rimouski Couturier S, Brunelle J, Vandal J, St-Martin G (1990) Changes in the population dynamics of the George River caribou herd, 1976–87. Arctic 43:9–20
Crête M, Desrosiers A (1995) Range expansion of coyotes, Canis latrans, threatens a remnant herd of caribou, Rangifer tarandus, in southeastern Québec. Can Field-Nat 109:227–235
Crête M, Huot J (1993) Regulation of a large herd of migratory caribou: summer nutrition affects calf growth and body reserves of dams. Can J Zool 71:2291–2296
Crête M, Manseau M (1996) Natural regulation of cervidae along a 1,000 km latitudinal gradient: change in trophic dominance. Evol Ecol 10:51–62
Dauphiné TC (1971) Physical variables as an index to condition in barren-ground caribou. Trans N E Fish Wildl Conf 28:91–108
Dauphiné TC (1976) Biology of the Kaminuriak population of barren-ground caribou. Part 4. Growth, reproduction and energy reserves. Report Series Number 38, Canadian Wildlife Service, Canada
Edmonds EJ (1988) Population status, distribution, and movements of woodland caribou in west central Alberta. Can J Zool 66:817–826
Ferguson SH, Bergerud AT, Ferguson R (1988) Predation and habitat selection in the persistence of a remnant caribou population. Oecologia 76:236–245
Gaillard JM, Festa-Bianchet M, Yoccoz NG, Loison A, Toigo C (2000) Termporal variation in fitness components and population dynamics of large herbivores. Annu Rev Ecol Syst 31:367–393
Gerhart KL, Russell DE, Wetering D, White RG, Cameron RD (1997a) Pregnancy of caribou (Rangifer tarandus): evidence for lactational infertility. J Zool 242:17–30
Gerhart KL, White RG, Cameron RD, Russell DE, Wetering D (1997b) Pregnancy rate as an indicator of nutritional status in Rangifer: implications of lactational infertility. Rangifer 17:21–24
Gratton C, Denno RF (2003) Seasonal shift from bottom-up to top-down impact in phytophagous insect populations. Oecologia 134:487–495
Gustine DD, Parker KL, Lay RJ, Gillingham MP, Heard DC (2006) Calf survival of woodland caribou in a multi-predator ecosystem. Wildl Monogr 165:1–32
Harder JD, Kirkpatrick RL (1994) Physiological methods in wildlife research. In: Bookhout TA (ed) Research and management techniques for wildlife and habitats, 5th edn. The Wildlife Society, Bethesda
Hayes RD, Farnell R, Ward RMP, Carey J, Dehn M, Kuzyk GW, Baer AM, Gardner CL, O’Donoghue M (2003) Experimental reduction of wolves in the Yukon: ungulate responses and management implications. Wildl Monogr 152:1–35
Hearn BJ, Luttich SN, Crête M, Berger MB (1990) Survival of radio-collared caribou (Rangifer tarandus caribou) from the George River herd, Nouveau-Quebec–Labrador. Can J Zool 68:276–283
Holter JG, Hayes HH (1977) Growth in white-tailed deer fawns fed varying energy and constant protein. J Wildl Manage 41:506–510
Hunter MD, Price PW (1992) Playing chutes and ladders: heterogeneity and the relative roles of bottom-up and top-down forces in natural communities. Ecology 73:724–732
Jiang L, Morin PJ (2005) Predator diet breadth influences the relative importance of bottom-up and top-down control of prey biomass and diversity. Am Nat 165:350–363
Keech MA, Bowyer RT, Ver Hoef JM, Boertje RD, Dale BW, Stephenson TR (2000) Life-history consequences of maternal condition in Alaskan moose. J Wildl Manage 64:450–462
Keith LB, Cary JR, Rongstad OJ, Brittingham MC (1984) Demography and ecology of a declining snowshoe hare population. Wildl Monogr 90:1–43
Kinley TA, Apps CD (2001) Mortality patterns in a subpopulation of endangered mountain caribou. Wildl Soc Bull 29:158–164
Kistner TP, Trainer CE, Hartmann NA (1980) A field technique for evaluating physical condition of deer. Wildl Soc Bull 8:11–17
Kotler BP, Gross JE, Mitchell WA (1994) Applying patch use to assess aspects of foraging behavior in Nubian ibex. J Wildl Manage 58:299–307
Leibold MA (1989) Resource edibility and effects of predators and productivity on the outcome of trophic interactions. Am Nat 134:922–949
Lima SL, Dill LM (1990) Behavioural decisions made under the risk of predation: a review and prospectus. Can J Zool 68:619–640
McLoughlin PD, Dzus E, Wynes B, Boutin S (2003) Declines in populations of woodland caribou. J Wildl Manage 67:755–761
Mech LD, Delgiudice GD (1985) Limitations to the marrow-fat technique as an indicator of body condition. Wildl Soc Bull 13:204–206
Menge BA (1992) Community regulation: under what conditions are bottom-up factors important on rocky shores? Ecology 73:755–765
Olofsson J (2006) Short- and long-term effects of changes in reindeer grazing pressure on tundra heath vegetation. J Ecol 94:431–440
Parker KL, Gillingham MP, Hanley TA, Robbins CT (1999) Energy and protein balance of free-ranging black-tailed deer in a natural forest environment. Wildl Monogr 143:1–48
Pimlott DH, Shannon JA, Kolenosky GB (1969) The ecology of the timber wolf in Algonquin Provincial Park. Research Report No. 87. Ontario Department of Lands and Forests, Toronto
Rettie WJ, Messier F (1998) Dynamics of woodland caribou populations at the southern limit of their range in Saskatchewan. Can J Zool 76:251–259
Reimers E, Klein DR, Sørumgård R (1983) Calving, growth rate, and body size of Norwegian reindeer on different ranges. Arctic Alpine Res 15:107–118
Rominger EM, Oldemeyer JL (1990) Early-winter diet of woodland caribou in relation to snow accumulation, Selkirk Mountains, British Columbia, Canada. Can J Zool 68:2691–2694
Schaefer JA, Veitch AM, Harrington FH, Brown WK, Theberge JB, Luttich SN (1999) Demography of decline of the red wine mountains caribou herd. J Wildl Manage 63:580–587
Singer FJ, Harting A, Symonds KK, Coughenour MB (1997) Density dependence, compensation, and environmental effects on elk calf mortality in Yellowstone National Park. J Wildl Manage 61:12–25
Skogland T (1983) The effects of density-dependent resource limitation on the size of wild reindeer. Oecologia 60:156–168
Skogland T (1986) Density-dependent food limitation and maximal production in wild reindeer herds. J Wildl Manage 50:314–319
Skogland T (1991) What are the effects of predators on large ungulate populations? Oikos 61:156–168
Thomas DC, Gray DR (2002) Updated status report on “forest-dwelling” woodland caribou, caribou du bois (Rangifer tarandus caribou). Committee on the Status of Endangered Wildlife in Canada, Ottawa
Tollefson T (2007) The influence of summer and autumn forage quality on body condition and reproduction in lactating mule deer and their fawns (Odocoileus hemionus). PhD thesis, Washington State University, Pullman
Tveraa T, Fauchald P, Yoccoz NG, Ims RA, Aanes R, Høgda KA (2007) What regulate and limit reindeer populations in Norway? Oikos 116:706–715
Van der Wal R (2006) Do herbivores cause habitat degradation or vegetation state transition? Evidence from the tundra. Oikos 114:177–186
Verme LJ (1969) Reproductive patterns of white-tailed deer related to nutritional plane. J Wildl Manage 33:881–887
Verme LJ, Ozoga JJ (1980a) Influence of protein-energy intake on deer fawns in autumn. J Wildl Manage 44:305–314
Verme LJ, Ozoga JJ (1980b) Effects of diet on growth and lipogenesis in deer fawns. J Wildl Manage 44:315–324
Whitten KR, Garner GW, Mauer FJ, Harris RB (1992) Productivity and early calf survival in the porcupine caribou herd. J Wildl Manage 56:201–212
Wittmer HU, McLellan BN, Hovey FW (2006) Factors influencing variation in site fidelity of woodland caribou (Rangifer tarandus caribou) in southeastern British Columbia. Can J Zool 84:537–545
Wittmer HU, McLellan BN, Seip DR, Young JA, Kinley TA, Watts GS, Hamilton D (2005a) Population dynamics of the endangered mountain ecotype of woodland caribou (Rangifer tarandus caribou) in British Columbia, Canada. Can J Zool 83:407–418
Wittmer HU, Sinclair ARE, McLellan BN (2005b) The role of predation in the decline and extirpation of woodland caribou. Oecologia 144:257–267
Author information
Authors and Affiliations
Corresponding author
Additional information
Communicated by John Fryxell.
An erratum to this article can be found at http://dx.doi.org/10.1007/s00442-007-0881-1
Rights and permissions
About this article
Cite this article
Brown, G.S., Landriault, L., Sleep, D.J.H. et al. Comment arising from a paper by Wittmer et al.: hypothesis testing for top-down and bottom-up effects in woodland caribou population dynamics. Oecologia 154, 485–492 (2007). https://doi.org/10.1007/s00442-007-0855-3
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s00442-007-0855-3