Alces30_25.pdf EFFECTS O F BLACK BEAR PREDATION ON CARIBOU--A REVIEW Warren B. Ballard New Brunswick Cooperative Wildlife Research Unit, Faculty of Forestry, P. 0. Box 44555, University of New Brunswick, Fredericton E3B 6C2 ABSTRACT: I reviewed available literature concerning black bear (Ursus americanus) predation on caribou (Rangifer tarandus) in an effort to gain insight on the possible impacts of black bear predation on a potential re-introduction of woodland caribou (Rangifer tarandus caribou) to north-central Minnesota. Several case histories were reviewed and inferences were drawn from several black bear- moose (Alces alces) studies. I concluded that black bear predation on woodland caribou i n the proposed re-introduction area would likely be a secondary source of caribou mortality and that between 6-30% of the calves and 0-5% adults might be killed annually by black bears. ALCES VOL. 30 ( 1 994) pp.25-35 Predation has been identified as a major limiting factor in many populations of ungu- lates. The majority of these losses generally occur among juvenile age classes. Because predation can limit ungulate populations, it can potentially impact re-introductions of ungulate species into areas from which they have been extirpated. The North Central Caribou Corporation is in the process of com- pleting plans for a potential re-introduction of woodland caribou (Rangifer tarandus cari- bou) into north-central Minnesota. Predation, particularly by wolves (Canis lupus) (Bergerud and Elliot 1986), has been identified as a potential factor which could result in the fail- ure of this re-introduction. However, other predator species also occur in the area. Black bears (Ursus americanus) have re- cently been identified as important predators of juvenile ungulates in several areas of North America (Schlegel 1976, Franzmann et al. 1980, Adams et al. 1988). Black bears are moderately abundant within the proposed re- introduction area (i.e., 159 to 244/1,000 km2 [Rogers 1987]), and predation by them could be a significant factor in the success or failure of the re-introduction. Because of the above concerns the North Central Caribou Corpora- tion desired additional information concern- ing the potential of black bears to impact the woodland caribou re-introduction to Minne- sota. The purpose of this report is to review information concerning black bear predation on caribou in general, and on woodland cari- bou in particular. METHODS I reviewed literature concerning black bear predation on ungulates in North America and relied heavily on the review papers by Truett et al. (1989) and Ballard (1992), and on the bibliography on Rangifer tarandus com- piled by Kreeger and Fleming ( 1 991). I contacted a number of individuals seeking unpublished information on the subject; how- ever, this report is not an exhaustive review of black bear predation on caribou. I may have omitted some studies concerning woodland caribou; however, I believe the information presented is representative of the state of knowledge concerning woodland caribou- black bear relationships. CASE HISTORY STUDIES Most studies on bear-caribou relation- ships concern barren ground caribou and griz- zly bears (Ursus arctos) in tundra ecosystems (Page 1976, Garner and Reynolds 1986, Bergerud and Page 1987, Adams et ul. 1988, Whitten et al. 1992). In all cases, predation by wolves and grizzly bears was found to be a significant cause of calf caribou mortality. BLACK BEAR PREDATION ON CARIBOU - BALLARD ALCES VOL. 30 ( 1 994) Although these studies did not involve black death were found for 11 cases; coyotes, black bears, some inferences can be drawn concern- bears, and golden eagles were thought to be ing the timing of mortality and the potential responsible for 11, 3, and 1 mortalities, re- significance of black bear predation on wood- spectively. Yearling and adult female caribou land caribou. Also, there have been several experienced high annual survival between studies which have concerned black bear pre- 1987 and 1992 (>90%) and no cases of preda- dation on woodland caribou. tion were observed during 11,915 caribou- days of monitoring. Black bear feeding sta- Newfoundland tions were in operation during 1989 and CrCte Mahoney et al. (1 990) provides the larg- et al. (1 99 1) concluded that the stations could est data set concerning the impacts of black have reduced bear predation rates. bears on woodland caribou. Between 1979 and 1984 they determined causes of mortality of 220 radio-collared newborn caribou calves within three woodland caribou populations in Newfoundland. Twenty-three percent of the calves died during their first year of life with predation accounting for 78% of the deaths. Predation by lynx (Lynx canadensis) and black bears was each responsible for 35% of the mortalities. Of 52 mortalities, 23 (44%) oc- curred within 2 weeks and 62% occurred within4 weeks of birth (Mahoney etal. 1990). There were no differences in vulnerability of calves by sex or weight. Wolf predation was not a factor because they no longer occur in Newfoundland (Mahoney et al. 1990). No estimates of black bear density were provided and the three caribou populations totaled 17,713 + 15% individuals (Mahoney et al. 1990). The observed total mortality rate of 23% for calves aged 1 1 2 months was not sufficient to limit population growth during the study period. Grand-Jardins Park, Quebec Efforts have been underway to restore caribou to Grand-Jardins Park since 1969 when 41 fawn and yearling caribou were released into the Park (H. Jolicoeur, unpubl. data). However, only 12 of the animals sur- vived or stayed in the area. No caribou were released in 1970 but an additional 25 fawns and yearlings were released in 1971. The latter animals remained in the area apparently because they formed an association with the 12 surviving adult animals from the 1970 release. An additional 12 adults were released in 1972. This herd has increased at an average rate of 5% since 1970 (Cantin 1991) and currently numbers 125 animals (H. Jolicoeur, unpubl. data). Wolf densities within this area were re- duced during 1979 and 1980 and are currently 10 wolves per 1000 km2 (H. Jolicoeur, unpubl. data). Black bear density was estimated at 220 per 1000 km2. The meningeal worm [Parelaphostrongylus tenuis) was not present Gaspesie Park, Quebec in this area (H. Jolicoeur, pers. commun.). Crete and Desrosiers reported On Crete (pers. commun.) suggested that the ex- the status of the Gaspesie Park caribou herd istence of escape habitats (open landscapes) which was declining due to low recruitment. such as alpine tundra or large bogs could play Twelve of 13 calves born to radio-collared a key role in allowing introduced caribou to caribou during 1988 were lost during the increase where bears and wolves exist. summer (majority in July); predation by black - ~ ~ - bears a n d o r coyotes (Canis latrans) was sus- Southeastern British Columbia pected as the cause of these deaths. Newborn Seip (1992) reported on the status of two calves were also radio-tagged in 1989 and woodland caribou herds in southeastern Brit- 1990 with 16 of 25 calves dying during the ish Columbia and sought to identify limiting first summer of life. Indices on the cause of factors. Woodland caribou populations de- ALCES VOL. 30 (1994) BALLARD - BLACK BEAR PREDATION ON CARIBOU clined or disappeared from this area during avoid predators by calving or living on islands the 1900's. Over-hunting was blamed in (D. R. Seip, B. C. Ministry of Forests, Burnaby, many cases, but other populations without pers. commun.). hunting exhibited similar declines. In a non- migratory caribou population the distribution of wolves, grizzly and black bears, and cari- bou overlapped while in a migratory popula- tion wolf and moose (Alcesalces) distribution did not overlap that of caribou during sum- mer. No mention was made of bear distribu- tions but I assume they probably overlapped. Within the non-migratory Quesnel Lake cari- bou population wolves and bears (both griz- zly and black bears) accounted for 55 and 15%, respectively, of the adult mortality. About half of the calves died of unknown causes during the calving period. Seip (1992) suggested that calf survival was related to wolf abundance during summer and that wolf predation was driving the caribou population towards extinction. Within the migratory Wells Gray Park caribou population, Seip (1992) estimated that the adult caribou mortality rate was 8% with most of the mortality being attributed to bear predation (apparently both species). Ap- proximately 40% of the adult female caribou had surviving calves and the population was stable or slowly increasing. Seip (1992) con- cluded that the differences between the two populations were due to differences in wolf predation and overlaps in distribution of alter- nate prey (i.e., moose) and wolves. Seip (1 99 1) concluded that forest-dwell- ing caribou in Ontario, Saskatchewan, Al- berta, and the non-mountainous regions of British Columbia had declined or were elimi- nated due to wolf predation and human har- vest. Increased wolf densities following the range expansion of ungulates such as moose, elk and deer has resulted in greater predation pressure on resident caribou. Caribou persist in large numbers in northern or high elevation areas where predation and hunting pressures are reduced. In southern areas they continue to persist in low numbers where they can Baxter State Park, Maine During 1989, 12 woodland caribou were released in Baxter S t a t e Park, Maine (McCollough and Connery 1990). Within 6 months of the release 10 of the 12 died; at least five died or "were predisposed" to meningeal worm, one died from abomasal ulcers, one from either a fatal accident or black bear predation, and 3 died apparently from black bear predation. Black bear density was estimated at about 232/1,000 krn2, coyote density at 183/1,000 km2, and no wolves were present. The authors pointed out that two major factors (i.e., disease and predation) suggested by Bergerud and Mercer (1989) as being responsible for both the decline of cari- bou in North America and failure of recent re- introductions were also responsible for the re- introduction failure in Maine. EFFECTS OF BEAR PREDATION ON CARIBOU Prior to the use of radio telemetry on neonates, bears were largely thought of as scavengers of ungulates (Jonkel 1978). Since Schlegel's (1976) study of elk (Cervus elaphus) calves, black bears have been iden- tified as significant predators of neonatal elk (Schlegel 1976), moose (Franzmann et al. 1980, Ballard 1992), deer (Odocoileus sp.) (Wilton 1983, Conger and Giusti 1992), and caribou (Mahoney et al. 1990). Wilton (1983) summarized observations of black bears prey- ing on ungulates in North America and con- cluded that they should be considered effec- tive predators of ungulates throughout their range. Black bears are not considered effec- tive predators on adult moose (Ballard 1992), but Mercer (1986, cited in McCollough and Connery 1990) and Seip (1991) suggest that black bears may account for the majority of adult caribou mortality. Also, V. Crichton (Manitoba Dep. of Nat. Resources, Winni- BLACK BEAR PREDATION ON CARIBOU - BALLARD ALCES VOL. 30 (1994) peg, pers. commun.) reports an increasing incidence of anecdotal reports of black bears preying on adult moose in Manitoba and On- tario (Austin et al. 1994). In other areas scattered anecdotal accounts of black bears preying on adult caribou exist, but black bears are not considered significant predators of adult caribou (V. Crichton, pers. commun; L. G. Adams, U . S. National Park Service, An- chorage, Alas., pers. commun.). Regardless, black bear predation is a significant source of mortality to neonates in many ungulate populations. Predation has been reported in many stud- ies to be the most significant mortality factor affecting caribou populations (Miller and Broughton 1974, Bergerud 1980). Most mor- tality of caribou neonates due to predation occurs during the first month of life (Bergerud 1971, 1980; Miller and Broughton 1974; Jakimchuck 1979; Miller 1987; Adams et al. 1988, Mahoney et al. 1990, Whitten et al. 1992; ). Generally the most abundant preda- tor species constitutes the largest source of mortality (Truett et al. 1989). However, this is not always true, particularly with regard to black bear predation. Ballard et al. (1990) examined the causes of mortality to neonate moose in southcentral Alaska where black bears outnumbered grizzly bears and wolves by factors of 3.2 and 32, respectively. Grizzly bears killed 52% of the calves followed by 9% for black bears and 7% by wolves. Thus occurrence of predator species, their relative predation efficiency and density, and prey density probably affect which predator spe- cies are the most significant cause of mortal- ity. Neonatal moose are most vulnerable to bear predation during their first 6 weeks of life (Ballard et al. 198 1). Subsequently their increased mobility appears to make them in- creasingly less vulnerable to bear predation (Ballard etal. 1980). Newborn caribou calves also suffer their highest mortality during the first 1-2 months of life and become less vul- nerable to at least bear predation with increas- ing age and mobility (Truett et al. 1989, Adams et al. 1993). Predation by black bears has accounted for the deaths of from 2 to 50% of radio- collared moose calves in various areas of North America (Table 1). Ballard (1992) concluded that black bears were a significant source of mortality to moose calves when they out number grizzly bears and wolves by factors of 10 and 30, respectively, or their densities were >20011,000 km2. Although only based on two studies from the Kenai Peninsula, Alaska, Ballard (1992) also con- cluded that black bear kill rates (functional response) appeared to be dependent upon the densities of moose calves while the percent of the moose population killed (functional and numerical response) was not dependent on moose densities. Based upon the aforemen- tioned studies the importance of black bear predation as an ungulate limiting factor is dependent upon the density of black bears in relation to the number and density of other predator and prey species. It is not known whether black bear predation is a learned phenomena, or whether it has occurred all along. It may not have been detected earlier because many food habits studies have relied on scat analyses which would underestimate the importance of ungulates (Ballard and Larsen 1987). POTENTIAL EFFECTS OF BLACK BEARS ON WOODLAND CARIBOU RE-INTRODUCTIONS IN THE BOUNDARY WATERS CANOE AREA WILDERNESS Bergerud and Mercer (1989) have sug- gested that even in the absence of deer (the source for P. tenuis) when wolf densities exceed 1011,000 km2, caribou re-introduc- tions will fail. Bergerud and Elliot (1986) indicated that in general, caribou populations can not maintain their numbers when wolf densities are26.5/1,000 km2 in the absence of ALCES VOL. 30 (1994) BALLARD - BLACK BEAR PREDATION ON CARIBOU escape habitat. The increasing caribou popu- lation at Grand-Jardins Park, QuCbec appear to fit their prediction in that the caribou herd has been increasing at 5% annually (Cantin 1991), wolf and black bear densities have been estimated at 10 and 22011,000 km2, respectively (H. Jolicoeur, unpubl. data), and adequate escape cover exists (M. Crete, pers. commun.). Wolf densities within the pro- posed Minnesota caribou reintroduction area have been estimated at 16 to 2011,000 km2 (Nelson and Mech 1992). Nelson and Mech (1 992) acknowledged that their reported wolf densities were well above the threshold re- ported by Bergerud and Elliot (1986) and Bergerud and Mercer (1989), but that caribou coexisted in other areas with higher wolf densities: Spatsizi Provincial Park, British Columbia (16 wolves/1,000 km2), Lake Ni- pigon, Ontario (1 0- 14 wolves/1,000 km2), and Pukaskwa Park, Ontario (14 wolves/1,000 km2). None of the aforementioned authors mentioned bear predation. Bergerud et al. (1983) mentioned that the caribou population at Pukaskwa Park num- bered 20-30 animals and that wolves seldom preyed upon them. Black bears also occur in the area, but their densities are unknown (G. Eason, Ontario Ministry of Natural Resources, Wawa, pers. commun.). The caribou in this area have apparently declined, and only a few remained in the area by 1992 (G. Eason, pers. commun.). Causes for this apparent reintro- duction failure are unknown. Bergerud etal. (1 990) suggested that cari- bou successfully calved on islands to escape predation by wolves. Nelson and Mech (1992) also suggested that islands at Lake Nipigon, Ontario provided escape habitat for caribou from wolf predation. V. Crichton (pers. comm.) indicates that woodland caribou on the east side of Lake Winnipeg, Manitoba traditionally calve on islands in large lakes and on occasion within bogs. These herds are composed of 40-100 individuals. Prior to 1978 few black bears were observed on the islands, but since 1978 numerous black bears have been observed, and the caribou popula- tion has apparently remained stable. It is unknown whether black bears are preying on caribou calves, but the recent occurrence of bears on the islands suggests that they may be preying on caribou calves. Hair from adult caribou has been found in bear feces during July, but it is not known whether these obser- vations represent carrion feeding or actual predation. Caribou persist at low densities (about 0.02/km2) where wolf density is high (161 1,000 km2) at Spatsizi Provincial Park, British Columbia (A. T. Bergerud, Univ. of Victoria, British Columbia, pers. comm.). Both black and grizzly bears occur in the area, but their importance as predators has not been studied. However, if they prey on caribou the magni- tude of such predation would be secondary to that by wolves. Pitt and Jordan (1992) examined the use of islands by black bears within the Boundary Waters Canoe Area by examining usage of bait stations at islands with and without hu- man camp sites. They found that black bears appeared to be more abundant on islands with camp sites than those without camp sites. Although their study results were limited be- cause they could only place bait stations within 1 km of camp sites (this limited the size of the island which could be studied--W. Pitt, Utah State Univ., Logan, pers. commun.) their re- sults have a direct bearing on bear usage of island sites. Islands with camp sites appeared to be occupied less frequently by wolves but more frequently by bears (W. Pitt, pers. commun.). Apparently bears were attracted to the islands because of the presence of food associated with humans. Because woodland caribou use islands as escape habitat during calving, the attraction of bears to island sites occupied by campers will increase the fre- quency of bear-caribou interactions. This may result in increased bear predation be- cause of the presence of human camp sites. Table 1.Causes of mortality and survival rates of radio collared moose calves to November in relation to observed kill rates and predator densities in North America (modified from Ballard 1992). h South central Alaska Kenai Peninsula, AK + cr Areas Area 1-3 Area 1 Area 4 Areas 1947 Bum 1969 Bum Southwest Eastcentral Saskat- New New- f2 '3 Pooled Yukon* Alaska chewan Bmnswick foundland V, Years 1977,1978 1979 1984 1977-84 1977,1978 1981,1982 1983,1985 1984 1982 1983,1985 1983-88 Causes of mortality (%) Grizzly bear 4 1.9 42.9 52.2 44.0 6.4 2.7 41.9 51.5 Black bear W 0 Grizzly & black Wolf 1.6 6.5 2.5 6.4 1.4 17.9 15.2 Unknown predation 2.4 Unknown causes 3.2 Surviving (%) 46.0 42.9 17.4 39.4 44.6 48.5 18.8 18.2 50.0 8 1.8 70.0 Density (no./1000km2) Grizzly bear 24 10 28 24-28 12-28 12-28 16 16 0 0 0 Black bear 0 0 90 0-90 205 258 1 6W 8-11 200-400 mod.? 570 ALCES VOL. 30 (1994) S i 4 4 3 3 'J? 3 09 - 2 '-? N 'J? 3 09 3 2 5 BALLARD - BLACK BEAR PREDATION ON CARIBOU BLACK BEAR PREDATION ON CARIBOU - BALLARD ALCES VOL. 30 ( 1 994) However, if wolves are more abundant on islands not occupied by humans, then caribou may be subjected to high levels of wolf preda- tion on islands without human camp sites. Using the wolf densities reported by Nel- son and Mech (1992), it appears that the area considered for caribou reintroduction exceeds the threshold level of wolf densities that Bergerud and Mercer (1989) indicated were necessary for caribou to survive. Assuming that black bear densities are similar to those reported by Rogers (1987) for the Superior National Forest (1 59 to 244 bears/1,000 km2), bear densities approach the level suggested by Ballard (I 992) for black bears to be a signifi- cant source of mortality. Based upon both the reported bear, wolf, and deer (i.e., <386/1,000 km2, Pitt and Jordan 1991) densities within the proposed reintroduction area and reviewed literature, black bears might kill between 16 to 30% of the calves produced by adult wood- land caribou, and may kill 0-5% of the adults annually. The accuracy of these estimates would depend upon the magnitude of wolf predation. In any case, the evidence suggests that in the presence of wolves, black bear predation will be a secondary source of mor- tality which could be additive to other sources of mortality. The success or failure of the caribou re- introduction may depend on the timing and numbers of caribou released during the initial reintroduction. If black bear predation is, in part, a learned phenomenon then it would appear that introduction of large numbers of caribou at the initial stages of the project may allow caribou to establish a foot-hold before bears and wolves learn of their presence. It may also be advisable to establish diversion- ary feeding stations as was done by Crete et al. (1991) and Boertje et al. (1992) which may reduce black bear kill rates. At low caribou numbers the increased survival of just a few individuals can make the difference between success and failure of a caribou transplant. Although my analysis is based upon reviewed literature, I point out that the interactions between caribou and black bears, particularly woodland caribou, are poorly understood as suggested by the paucity of studies cited in this review. ACKNOWLEDGEMENTS This review was funded by the North Central Caribou Corporation. I thank A. T. Bergerud, A. F. Cunning, V. Crichton, G. J. Forbes, S. Mahoney, W. C. Pitt, D. Seip, H. R. Timmermann, and the North Central Caribou Corporation for critically reviewing early drafts of this manuscript. H. Jolicoeur pro- vided unpublished data on the caribou popu- lation in Grands-Jardins Park, QuCbec. J. L. 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