ALCES VOL. 44, 2008 SEIP - INTERACTIONS AMONG CARIBOU, WOLVES, AND MOOSE 1 MOUNTAIN CARIBOU INTERACTIONS WITH WOLVES AND MOOSE IN CENTRAL BRITISH COLUMBIA Dale R. Seip British Columbia Ministry of Forests and Range, 1011 Fourth Ave., Prince George, B.C. V2L 3H9, Canada, e-mail: Dale.Seip@gov.bc.ca ABSTRACT: Mountain caribou (Rangifer tarandus caribou) populations in south-eastern British Columbia are declining over most of their range and are listed as Threatened. Predation has been documented as the major cause of declining caribou numbers. Excessive predation by wolves (Canis lupus) has been related to increased moose (Alces alces) numbers. The increase in moose appears to be the result of a natural colonization process that has been enhanced by human-caused habitat change. Options to reduce the rate of predation include reducing wolves, reducing moose, and reducing the amount of early seral habitat that supports moose. Current management includes population control of moose and wolves. Monitoring and assessment of these approaches will guide the future management strategy used to maintain mountain caribou in south-eastern British Columbia. ALCES VOL. 44: 1-5 (2008) Key words: Alces alces, British Columbia, Canis lupus, habitat, management, mountain caribou, moose, Rangifer tarandus caribou, wolves Mountain caribou are an ecotype of woodland caribou (Rangifer tarandus caribou) that live in the mountains of south-eastern British Columbia, extending into northern Idaho (Heard and Vagt 1998). The moun- tains inhabited by these caribou experience extremely high snowfall during the winter which prevents the caribou from cratering through the snow to obtain terrestrial forage. Consequently, mountain caribou winter in upper elevation subalpine forests (Terry et al. 2000, Apps et al. 2001, Johnson et al. 2004) and feed almost exclusively on arboreal lichens (Bryoria spp.). Historically, mountain caribou were abun- dant and widely distributed throughout the mountains in south-eastern British Columbia, but over the past century their numbers and distribution have greatly declined (Spalding 2000). Population declines and range contrac- tion have continued in recent years (Wittmer et al. 2005). For example, the George Mountain herd just east of Prince George was observed to be declining over the past few decades and was completely extirpated by 2004 (Seip, unpublished data). As a result of these de- clines, mountain caribou are nationally listed as Threatened and the subject of ongoing recovery efforts (Mountain Caribou Technical Advisory Committee (MCTAC) 2002). Initially, management concern for moun- tain caribou focused on protecting old-growth subalpine forests that provide the arboreal lichens that they rely on during winter. In response, a large amount of subalpine forest habitat has been protected from logging to provide caribou winter range (MCTAC 2002). However, research over the past few decades has indicated that the primary cause of declin- ing mountain caribou herds is excessive levels of predation on both calves and adults during the summer (Seip 1992, Kinley and Apps 2001, Wittmer et al. 2005). Grizzly bears (Ursus arctos), black bears (U. americanus), wolves (Canis lupus), and cougars (Felis concolor) can all be major predators. Bear predation is important throughout the range of mountain caribou, whereas cougar preda- INTERACTIONS AMONG CARIBOU, WOLVES, AND MOOSE - SEIP ALCES VOL. 44, 2008 2 tion is more significant in the southeastern parts of the province, and wolf predation is more significant in the central portion of the province (Wittmer et al. 2005). Winter survival of mountain caribou is quite high because they have minimal expo- sure to predators during this time. Bears are hibernating, and wolves are found at lower el- evations where they are sustained primarily by moose (Alces alces) (Seip 1992). Conversely, moose and wolves often move to higher el- evations in summer which increases both the spatial overlap of wolves and caribou and the predation of adult and calf caribou. The key question is why is predation by native predators resulting in decline of caribou now, given that these animals co-existed in the past? It is unlikely that grizzly bear populations have increased above historic levels, rather, they have probably been reduced in many areas due to hunting. In contrast, wolf popula- tions have probably increased in response to increasing moose populations. Wolves exhibit a strong numerical response to increasing prey availability (Messier 1995), and the number of moose in central British Columbia has greatly increased over the past century. Today, the density of moose in central British Columbia is greater than 1 moose/km2 (Walker et al. 2006). There is some debate as to whether moose were completely absent from this region (Hatter 1950, Peterson 1955), or were sparse and scattered in the early 1900s (Spalding 1990). Regardless, moose began to increase in number in the early 1900s, and their distribution spread contiguously across central British Columbia. For example, in the mountains immediately east of Prince George, the colonization was recorded by local settlers: "In 1911 Ernest Jenson left his native Denmark and came to Canada. A few months later, he moved to the Dome Creek area … In 1912 and 1913, he was hired on as a hunter for the Grand Pacific Railway…. Although moose were present in the area since about 1900, Ernest spent a whole year in the forests before seeing one. The moose were working their way south at about eight to ten miles per year. It wasn’t long before they were in great abundance… Caribou were in abundance and it was common to see small herds in every valley" (Boudreau 1998). Early settlers also reported that the arrival of moose coincided with the decline in caribou, although the interaction among moose, wolves, and caribou was not obvious to them: "I asked him what he thought the reason was for the decline of the caribou popula- tions… Do you think the moose had any bearing on it? I asked. I can’t see how. They don’t even eat the same kind of food, and yet the caribou did start going down hill shortly after the moose arrived. Probably the wolves were more responsible than anything else. I mean they sure killed a lot of them" (Boudreau 1998). It was similarly reported in other parts of their range that decline in mountain caribou numbers coincided with an increase in moose numbers (Edwards 1956, Bergerud and Elliot 1986). It appears that the decline in mountain caribou and their distribution in central Brit- ish Columbia are primarily due to excessive predation, and that this predation is related to high wolf numbers that are sustained by the increased numbers of moose (Bergerud and Elliot 1986, Seip 1992). The cause of the increased number and distribution of moose is probably a combination of natural and human-caused factors. The initial increase and spread of moose during the early 1900s ALCES VOL. 44, 2008 SEIP - INTERACTIONS AMONG CARIBOU, WOLVES, AND MOOSE 3 may have been related to a warming period following the end of the Little Ice Age (Luck- man 2000). Subsequent climatic warming in central British Columbia over the past century (B.C. Ministry of Environment 2002) would have further encouraged increased moose abundance and distribution as severe winters became less common. Consequently, at least part of the increase in moose numbers is prob- ably related to climate change. However, there is also reason to believe that the increase in moose and the associated decline in mountain caribou are, in part, re- lated to human-caused habitat change. The forest ecosystems in and adjacent to traditional range of mountain caribou had a very low frequency of natural disturbance resulting in a landscape dominated by mature and old forests (Seip 1998, Delong 2007). Under the natural disturbance regime, these landscapes would provide little early seral habitat at low elevations that is preferred by moose as winter range. However, forest harvesting and human settlement has greatly increased the amount of early seral habitat, thereby increasing habitat suitability for moose within much of the range of mountain caribou. Such habitat change appears to have benefited moose at the expense of mountain caribou. An analogous situation for mountain caribou exists in south-eastern British Co- lumbia where cougars (Felis concolor) are a major predator of caribou (Kinley and Apps 2001). Climatic and landscape changes that benefit deer (Odocoileus spp.) and elk (Cervus elaphus) habitat have increased these ungulate populations that, in turn, sustain high cougar numbers resulting in increased predation of caribou. MANAGEMENT IMPLICATIONS Recently completed recovery plans for mountain caribou identify the need to reduce the rate of predation to achieve recovery (MCTAC 2002, Hart and Cariboo Mountains Recovery Implementation Group (HCMRIG) 2005). These plans recognise that management actions could occur at different trophic levels to reduce predation, and include: 1. Directly reduce wolf numbers by killing wolves or implementing reproductive control. This approach would have the most direct and immediate effect on reduc- ing wolf numbers and wolf predation on caribou. However, wolf control is very controversial, and the management ac- tion would have to be used as an ongoing program because wolves would quickly recover following the cessation of any control program if moose habitat and moose numbers were not reduced. 2. Reduce wolf numbers by reducing the number of moose which provide the primary prey of the wolf population. Moose populations could be reduced by liberalizing hunting regulations, which is somewhat less controversial than predator control, but still a significant concern to hunters and First Nations. Also, moose reduction would have to be an ongoing program because the moose population will soon recover if good quality habitat is available. The potential for enhanced moose hunting to be a long-term solution is compromised by an ongoing decline in the number of hunters in British Columbia, as well as the feasibility of hunters being able to adequately reduce moose to very low densities in inaccessible areas. There is concern that in the short term, wolves de- prived of moose will temporarily increase their predation on caribou. A short-term wolf control program may be required to reduce wolf numbers to a level compatible with reduced moose numbers. 3. Reduce the rate of forest harvesting, pre- scribed burning, and other practices which create early seral habitat. Reducing early seral habitat on the landscape should reduce INTERACTIONS AMONG CARIBOU, WOLVES, AND MOOSE - SEIP ALCES VOL. 44, 2008 4 moose numbers and lead to lower wolf numbers. This approach is favoured by environmental groups, but further reduc- tions in forest harvesting concerns the for- est industry (HCMRIG 2005). Although much of the upper elevation habitat that is used by caribou is already protected from forest harvesting, reducing the number of moose and wolves would require an ad- ditional reduction in the rate of logging in adjacent valley bottoms, even though caribou seldom use those low elevation areas in central British Columbia. If the latter approach was implemented, there would be a substantial lag time before the existing cut blocks regenerated to a stage where they no longer provided good moose habitat. Consequently, moose reduction and/or wolf control for several decades may be required until cut blocks mature beyond good moose habitat. Some environmental groups accept use of interim predator management if it is used as a temporary tool while habitat recovers, but oppose predator management (reduction) if it is used as a permanent solution in place of habitat management (HCMRIG 2005). The degree that forest harvesting would have to be reduced to adequately reduce wolf numbers is unclear. The HCMRIG (2005) recommended that moose habitat suitability in areas adjacent to mountain caribou habitat should not exceed the amount that would oc- cur under a natural disturbance regime in that area. However, given that moose are a recent colonizer to mountain caribou range, it may be that even under the natural disturbance regime moose and wolf numbers would be too great to allow caribou to co-exist. If so, there may be no possibility of using habitat management to maintain self-sustaining caribou herds in some or all areas, and ongoing predator-prey management would be required to maintain caribou populations. In practice, these three approaches are not mutually exclusive, and a management strategy could include some combination of reducing the amount of early seral habitat, increasing the moose harvest, and some direct wolf control or reduction. A similar set of options also applies to managing the predator- prey interactions among caribou, deer, and cougars in south-eastern British Columbia. At this time, the final management approach to maintain mountain caribou in British Co- lumbia is still evolving. In the meantime, a variety of these man- agement approaches have been implemented, including reproductive control of wolves in the Quesnel Highlands, moose reduction in the Parsnip watershed, and moose reduction in the Revelstoke area. Limiting the creation of early seral habitat is an important manage- ment consideration in parks where there is no competing concern for timber supply and production. Research projects are in place to monitor all these management programs, and the resulting information will be used to refine the future application of these manage- ment practices. REFERENCES Apps, C. D., B. N. McLeLLan, T. A. KinLey, and J. P. FLaa. 2001. Scale-dependent habitat selection by mountain caribou, Columbia Mountains, British Columbia. Journal of Wildlife Management 65: 65-77. Bergerud, A. T., and J. P. ELLiot. 1986. Dy- namics of caribou and wolves in northern British Columbia. Canadian Journal of Zoology 64: 1515-1529. Boudreau, J. 1998. Crazy Man’s Creek. Caitlin Press Inc., Prince George, British Columbia, Canada. British CoLumbia Ministry Of Environment. 2002. Indicators of climate change for British Columbia, 2002. http://www.env. gov.bc.ca/air/climate/indicat/pdf/indcc. pdf (accessed January 2008) DeLong, S. C. 2007. Implementation of natural disturbance-based management in northern British Columbia. Forestry http://www.env.gov.bc.ca/air/climate/indicat/pdf/indcc.pdf http://www.env.gov.bc.ca/air/climate/indicat/pdf/indcc.pdf http://www.env.gov.bc.ca/air/climate/indicat/pdf/indcc.pdf ALCES VOL. 44, 2008 SEIP - INTERACTIONS AMONG CARIBOU, WOLVES, AND MOOSE 5 Chronicles 83: 338-346. Edwards, R. Y. 1956. Snow depths and ungulate abundance in the mountains of western Canada. Journal of Wildlife Management 20: 159-168. (HCMRIG) Hart And Cariboo Mountains Recovery ImpLementation Group. 2005. Recovery implementation plan for threatened woodland caribou (Rangifer tarandus caribou) in the Hart and Cariboo Mountains recovery area, British Colum- bia, Canada. Unpublished report. http:// www.centralbccaribou.ca/crg/24/rap (ac- cessed January 2008) Hatter, J. 1950. The moose of central British Columbia. Ph.D. Thesis, State College of Washington, Pullman, Washington, USA. Heard, D. C., and K. L. Vagt. 1998. Caribou in British Columbia: a 1996 status report. Rangifer Special Issue No.10: 117-123. Johnson, C. J., D. R. Seip, and M. S. Boyce. 2004. A quantitative approach to conser- vation planning: using resource selection functions to identify important habitats for mountain caribou. Journal of Applied Ecology 41: 238-251. KinLey, T. A., and C. D. Apps. 2001. Mortality patterns in a sub-population of endangered Mountain Caribou. Wildlife Society Bul- letin 29: 158-164. Luckman, B. H. 2000. The little ice age in the Canadian Rockies. Geomorphology 32:357-384. Messier, F. 1995. On the functional and numerical response of wolves to chang- ing prey density. in L. N. Carbyn, S. H. Fritts, and D. R. Seip, editors. Ecology and Conservation of Wolves in a Chang- ing World. University of Alberta Press, Edmonton, Alberta, Canada. (MCTAC) Mountain Caribou TechnicaL Ad- visory Committee. 2002. A strategy for the recovery of Mountain Caribou in British Columbia. British Columbia Ministry of Water, Land and Air Protection, Victoria, British Columbia, Canada. Peterson, R. L. 1955. North American Moose. University of Toronto Press, Toronto, Ontario, Canada. Seip, D. R. 1992. Factors limiting woodland caribou populations and their inter- relationships with wolves and moose in southeastern British Columbia. Canadian Journal of Zoology 70: 1494-1503. _____. 1998. Ecosystem management and the conservation of caribou habitat in British Columbia. Rangifer, Special Issue No. 10: 203-211. SpaLding, D. J. 1990. The early history of moose (Alces alces) distribution and relative abundance in British Columbia. Contributions to Natural Science, Royal British Columbia Museum, Victoria, Brit- ish Columbia, Canada. _____. 2000. The early history of Woodland Caribou (Rangifer tarandus caribou) in British Columbia. Wildlife Bulletin No. B-100. British Columbia Ministry of Environment, Lands and Parks, Wildlife Branch, Victoria, British Columbia. Terry, E. L., B. N. McLeLLan, and G. S. Watts. 2000. Winter habitat ecology of mountain caribou in relation to forest management. Journal of Applied Ecology 37: 589-602. WaLker, A. B. D., D. C. Heard, V. MicheLf- eLder, and G. S. Watts. 2006. Moose density and composition around Prince George, British Columbia. Final Report for Forests for Tomorrow. Project No. 2914000. British Columbia Ministry of Environment, Prince George, British Columbia, Canada. Wittmer, H. U., B. N. McLeLLan, D. R. Seip, J. A. Young, T. A. KinLey, G. S. Watts, and D. HamiLton. 2005. Population dynamics of the endangered mountain ecotype of woodland caribou (Rangifer tarandus caribou) in British Columbia, Canada. Canadian Journal of Zoology 83: 407-418. http://www.centralbccaribou.ca/crg/24/rap http://www.centralbccaribou.ca/crg/24/rap