Relationships among food availability, harvest, and human–bear conflict at landscape scales in ,

Martyn E. Obbard1,9, Eric J. Howe1, Linda L. Wall2,10, Brad Allison3, Ron Black4, Peter Davis5, Linda Dix-Gibson6, Michael Gatt7, and Michael N. Hall8

1Wildlife Research and Monitoring Section, Ontario Ministry of Natural Resources, DNA Building, Trent University, 2140 East Bank Drive, Peterborough, ON K9J 7B8, Canada 2Bear Wise Program, Ontario Ministry of Natural Resources, 5520 Highway 101 East, South Porcupine, ON P0N 1H0, Canada 3Regional Resources Unit, Northwest Region, Ontario Ministry of Natural Resources, 435 James Street, S. Suite 221, Thunder Bay, ON P7E 6S8, Canada 4Ontario Ministry of Natural Resources, 7 Bay Street, Parry Sound, ON P2A 1S4, Canada 5Regional Resources Unit, Northeast Region, Ontario Ministry of Natural Resources, 5520 Highway 101 East, South Porcupine, ON P0N 1H0, Canada 6Natural Resources Information Section, Ontario Ministry of Natural Resources, 1350 High Falls Road, Bracebridge, ON P1L 1W9, Canada 7Regional Resources Unit, Southern Region, Ontario Ministry of Natural Resources, 300 Water Street, 3rd Floor North, Peterborough, ON K9J 8M5, Canada 8Ontario Ministry of Natural Resources, 3767 Highway 69S, Sudbury, ON P3G 1E7, Canada

Abstract: Managers of American black bears (Ursus americanus) must maintain populations to ensure viability and opportunities for sport harvest, and minimize human–bear conflict (HBC). Harvest is a cost-effective management tool in most jurisdictions, and intuitively it seems that with fewer bears, there should be fewer conflicts. Therefore, managers may attempt to achieve both objectives by manipulating the harvest. Further, because data describing harvest and HBC are frequently collected, managers sometimes infer changes in population status from trends in harvest and HBC. However, evidence that larger harvests reduce HBC is lacking, and changes in harvest metrics and the frequency of HBC may be independent of bear density. Understanding relationships among food availability, hunter effort, harvest, and HBC could help managers avoid making invalid inferences about population status from data describing harvest and HBC, and evaluate whether management actions are having intended results. We investigated relationships among food availability, HBC, and harvest at landscape scales in Ontario, Canada, 2004–2011. We hypothesized that HBC and harvest would be negatively correlated with food availability; that HBC would be negatively correlated with prior harvest; and that harvest would be positively correlated with number of hunters. We used Spearman rank correlation to test hypotheses. Human–bear conflict was negatively correlated with food availability across Ontario, and in the 2 administrative regions where food availability varied synchronously. Total harvest and the proportion of females in the harvest were negatively correlated with food availability across Ontario and in one region. Human–bear conflict was not correlated with prior harvests, providing no evidence that larger harvests reduced subsequent HBC. Given the variation in natural foods, harvest is unlikely to prevent elevated levels of HBC in years of food shortage unless it maintains bears at low densities—an objective that might conflict with maintaining viable populations and providing opportunities for sport harvest.

Key words: , food availability, harvest, human–bear conflict, Ontario, Ursus americanus

DOI: 10.2192/URSUS-D-13-00018.1 Ursus 25(2):98–110 (2014)

9email: [email protected] 10Present address: 203 Belvidere Street, Winnipeg, MB R3J 2H1, Canada

98 FACTORS AFFECTING HUMAN–BEAR CONFLICT N Obbard et al. 99

During the late 1980s, American black bears (Ursus long-term targets with regard to levels of HBC, americanus) were managed for sustainable yield in sustainability of harvests, and population status. most jurisdictions with healthy populations (Miller During the 1990s, data describing HBC in 1990), though populations in a few U.S. jurisdictions Ontario, Canada, were not maintained in a stan- were still recovering from past over-harvest. Since dardized manner across the province, and food then, minimizing conflicts with humans has become an availability was measured over only a portion of increasingly important aspect of black bear manage- black bear range (Poulin et al. 2003). Controversy ment (Hristienko and McDonald 2007, Spencer et al. surrounding the effects of the 1999 cancellation of 2007). Most management agencies have initiated the spring hunting season on the frequency of HBC programs to educate the public about how to minimize spurred the Ontario Ministry of Natural Resources human–bear conflict (HBC), but their effectiveness is (OMNR) to appoint an independent Nuisance Bear difficult to measure because of multiple influences on Review Committee in 2002 (Howe et al. 2010). That the frequency of, and reporting rate for, human–bear committee recommended standardizing collection of interactions (Garshelis 1989, Gore et al. 2006, Spencer data on HBC, and the expansion of food availability et al. 2007, Howe et al. 2010). Sport harvest can be a surveys to the entire province (Poulin et al. 2003). cost-effective means of removing animals from the Both of these recommendations were implemented in population, so it is intuitive and convenient for 2004. managers to assume that harvest could be used to This study describes the first analysis of relation- limit or reduce HBC by reducing bear density and ships among food availability, HBC, and harvest at distribution. However, evidence demonstrating the landscape scales in Ontario. Our objectives were (1) effectiveness of harvest management for reducing to assess whether food availability varied synchro- HBC is lacking (Garshelis 1989, Treves et al. 2010). nously over large areas; (2) to test for trends in Wildlife managers frequently infer trends in bear harvest and measures of HBC; (3) to test for negative populations from changes in harvest metrics (Gar- correlations between food availability and HBC; (4) shelis 1991, Garshelis and Hristienko 2006, Hris- to test for negative correlations between food tienko and McDonald 2007), and typically evaluate availability and each of total harvest, hunter success the efficacy of education programs designed to rate, and the sex and age ratio of harvests; (5) to test prevent HBC from trends in the number of conflicts for negative correlations between harvest and subse- reported by the public (Schirokauer and Boyd 1998, quent HBC; and (6) to test for positive correlations Gore et al. 2006, Hristienko and McDonald 2007, between hunter effort and harvest. An understanding Spencer et al. 2007). However, both harvest metrics of spatial synchrony in annual variation in food and measures of HBC vary among years in response availability was necessary to assess the validity of to changes in the availability of natural foods (Gilbert pooling data over large areas to test for correlations at et al. 1978; Garshelis 1989; McDonald et al. 1994; landscape scales. Similarly, trends in HBC and Noyce and Garshelis 1997; Ryan et al. 2004, 2007; harvest were potentially relevant to the interpretation Howe et al. 2010). This variation obscures trends and of results of other tests. Objectives 3 and 4 address the may also confound tests for relationships among bear hypothesis that when natural foods are scarce, black population size, hunting pressure, harvest, and HBC, bears are more prone to HBC and more vulnerable to which makes it difficult for managers to assess harvest as they seek out alternative foods. We whether management actions are having the desired expected that food shortages would affect the effect unless concurrent data on food availability are vulnerability of females to harvest more strongly than available to provide insights (Garshelis 1991, Gore vulnerability of males because subadult females et al. 2006, Garshelis and Noyce 2008, Treves et al. typically restrict their movements more than subadult 2010). Knowledge of relationships among food males do (i.e., subadult males are vulnerable to availability, hunter effort, harvest, and HBC would harvest even when natural foods are abundant) and enable us to explain fluctuations or trends in HBC and adult females must meet the energetic requirements of harvest metrics. This could help managers and other gestation and lactation. Objective 5 was intended to stakeholders avoid drawing incorrect inferences about test the assumption (appealing from a management trends in the bear population from HBC or harvest perspective) that harvesting liberally reduces HBC. data, and facilitate evaluations of whether manage- Objective 6 tests whether harvest increased with ment actions are approaching or meeting identified hunting pressure.

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of Ontario’s human population lived in SR (Statis- tics Canada 2007, 2012). Human populations in SR increased from 2001 to 2006 and again from 2006 to 2011, with some of the largest increases occurring in areas with the highest bear densities (Statistics Canada 2007, 2012; Howe et al. 2013). Black bear À 2 harvest densities in SR (x2004–2011 5 2.6/100 km ) were .4 times those in NER or NWR (Dix-Gibson 2012).

Data collection Ontario Ministry of Natural Resources District staff measured food availability annually (2004– 2011) by ranking production of - and - producing species (or groups of species within a genus) on a 5-point scale by visual observation, similar to the approach of Noyce and Coy (1990). Different sets of species were monitored in each of 3 Forest Regions within the greater study area (Boreal Forests, and Great Lakes–St. Lawrence Forests east Fig. 1. Map of Ontario, Canada. The hatched area and west of the Great Lakes; Fig. 1; Table 1). We was permanently occupied by black bears (Scheick et al. 2011). Heavy black lines show the Ontario selected these species based on their prevalence in the border and boundaries between Northwest (NWR), respective Forest Region, and with reference to Northeast (NER), and Southern (SR) administrative studies of black bear feeding habits in Ontario and regions of the Ontario Ministry of Natural Resources. similar habitat elsewhere (Rowe 1972, Obbard 1987, Thin black lines show District administrative bound- Rogers and Allen 1987, Noyce and Garshelis 1997, aries (Districts lie entirely within a Region). Heavy grey lines show boundaries between Rowe’s (1972) Romain et al. 2013; Table 1). All were native to Forest Regions, with Boreal Forests across most of Ontario, except feral domestic apples (Malus pumila) northern Ontario, and Great Lakes–St. Lawrence present in eastern and western Great Lakes–St. transitional forests south of Boreal Forests and east Lawrence Forests, and European mountain-ash and west of the Great Lakes. (Sorbus aucuparia), which contributed to scores for mountain-ash (Sorbus spp.) where present. Our Bear Methods Food Index (BFI) was the average of rankings across Study area all species used by bears in the respective Forest The study area was Ontario, Canada. Most data Region. When calculating the BFI, we first calculat- describing food availability, HBC, and harvest were ed the mean rank for each species across multiple from contiguously forested Boreal and Great Lakes– observations within Districts and years (where St. Lawrence Forest Regions that are permanently available). We then calculated District-specific an- occupied by black bears (Fig. 1); but we collected nual BFI values as the average rank across species, some data on food availability, and recorded occa- and finally we calculated regional and provincial sional occurrences of HBC and harvest, in fragment- BFI values as the mean BFI across Districts within ed habitat used by a small number of bears. The those geographic areas. OMNR divides the province into 3 administrative Staff at many District offices ranked food Regions: Northeast (NER), Northwest (NWR), and availability at multiple locations within their District Southern (SR). Regions are further subdivided into each year; in other cases, rankings reflected avail- administrative Districts (Fig. 1). From 2004 to 2011, ability within entire Districts, as assessed by one or NER and NWR were characterized by contiguous more observers. Food availability was not measured black bear habitat, low black bear harvest densities in all Districts in all years. The mean (SD) number of À 2 (x2004–2011 5 0.6/100 km ), and low and stable Districts that measured food availability annually in human population densities (Natural Resources each Region was 6.5 (2.6) of a possible 9 Districts in Canada 2006; Statistics Canada 2007, 2012). Most NER, 4.0 (1.3) of 7 Districts in NWR, and 5.3 (1.6) of

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Table 1. Species and groups of species monitored and included in the Bear Food Index, in each of eastern and western Great Lakes St–Lawrence (GLSL) Forests and Boreal Forests (Rowe 1972) in Ontario, Canada, 2004– 2011. GLSL GLSL Common name Species East West Boreal Oaks Quercus rubra, Q. alba, Q. velutina, Q. macrocarpa xx x Cherries Prunus serotina, P. pensylvanica, P. virginiana xx x Mountain ash Sorbus americana, S. decora, S. aucuparia xx x Apples Malus pumila xx Beaked Corylus cornuta xx x Juneberry–serviceberry– Amelanchier alnifolia, A. canadensis, A. arborea, xx x Saskatoonberry–shadbush A. amabilis, A. humilis, A. laevis, A. bartramiana Elderberry Sambucus nigra canadensis, S. racemosa xx x Blackberry Rubus occidentalis x Lowbush cranberry macrocarpon xx Highbush cranberry Viburnum opulus var. americanum xx x Gooseberry–currants Ribes spp. x x Wild rose Rosa acicularis, R. blanda, R. palustris, R. woodsii xx x Nannyberry Viburnum lentago x Hawthorn Crataegus spp. x x X Wild raisin Viburnum lantanoides x Wild grape Parthenocissus quinquefolia x Sumac Toxicodendron radicans xx Dogwoods (excluding bunchberry) Cornus sericea, C. alternifolia, C. rugosa, C. racemosa xx X Bunchberry Cornus canadensis xX Raspberry Rubus idaeus xx X Strawberry Fragaria virginiana xx x Vaccinium angustifolium, V. myrtilloides xx x Sarsaparilla Aralia hispida, A. nudicaulis, A. racemosa xx x

8 Districts in SR. The mean (SD) number of annual December of the same year. A reminder was mailed to observations of the BFI across Districts was 34.5 hunters who had not replied. During 2004, the reply (18.7) in NER, 7.0 (SD 3.1) in NWR, and 34.8 (SD rate for the postcard survey was 45% (Dix-Gibson 4.6) in SR. In the case of SR, 95% of observations 2012). Completion and submission of the postcard were from the 5 primarily forested Districts with survey became mandatory in 2005; since then, the contiguous black bear populations, and 5% of reply rate has varied between 60% and 70% (Dix- observations were from the 3 Districts at the Gibson 2012). Information hunters were requested to southeastern fringe of black bear range in Ontario. provide included the Wildlife Management Unit in During 2004, hunting seasons for black bear were which they hunted; whether they hunted using bait; open from 15 August to 15 October in NER and in 17 whether they killed a bear and if so, its sex; and whether of 27 Wildlife Management Units in NWR, from 15 they enclosed a premolar tooth for age determination. August to 31 October in 10 Wildlife Management Total resident harvest and resident-hunter success rates Units in NWR, and from 1 September or the Tuesday were calculated by assuming returned surveys were following Labor Day through 30 November in SR. representative of all licensed hunters. Non-residents of From 2005 to 2011, seasons ran from 15 August to 31 Ontariowererequiredtoreporttheirblackbear October in NER and NWR, and from 1 September or hunting activity and harvest on a Non-Resident Black the Tuesday following Labor Day through 30 Bear Hunting License Validation Certificate obtained November in SR. During 2004, a postcard survey either from an operator licensed to provide bear was mailed to resident hunters, which they were asked hunting services or from an OMNR office. Completion to voluntarily complete. During 2005, the survey was of these certificates was mandatory and reply rates attached to the hunting license, and from 2006 to 2011 were 100% (Dix-Gibson 2012). the survey was provided in the annual Hunting During 2004, OMNR initiated standardized, Regulations Summary. Hunters were required to province-wide recording of the number of reported complete and return the survey to OMNR by 15 occurrences of human–bear conflict (hereafter,

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‘‘occurrences’’) via a toll-free reporting line available We used correlations between calendar year and to the public 24 hours/day, 7 days/week. The number each of occurrences, traps set, total harvest, and the of traps set to capture bears at the site of occurrences number of hunters to test for temporal trends (a 5 was recorded beginning in 2005. Traps were set only 0.05, 2-sided). For all other correlations we used when deemed necessary by OMNR staff to prevent 1-tailed tests because our hypotheses included the property damage or if there was a risk to human expected type of relationship (positive or negative). safety after investigation by OMNR staff. We crossed the BFI with occurrences, traps set, total harvest, hunter success rate, the proportion of Data analyses females in the harvest, and the proportion of adult We assessed synchrony in the direction of change females in the harvest to address objectives 3 and 4. in BFI scores among years within Regions using For objective 5, we would have preferred to cross À Buonaccorsi’s average measure of agreement (A ), occurrences and traps set with harvest in the which is equivalent to the average proportion of time previous year, and the 2- and 3-year moving average that pairs of series agree in their change in direction of prior harvests. However, we were concerned (Buonaccorsi et al. 2001). When assessing synchro- about potential confounding effects of temporally ny, we used data only from Districts that measured autocorrelated BFI data when correlating number of food availability in .5 of the 8 years in our time occurrences and traps set with harvest in only the series. Data from 7 of 9 Districts in NER, and 5 of 7 previous year. Soft mast production varied synchro- Districts in each of NWR and SR met this criterion. nously and exhibited negative autocorrelations with We assessed synchrony among Regions by calculat- production in the previous year over much of NER and SR from 1998 through 2009 (Howe et al. 2012). ing the simple measure of agreement (Aij) between pairs of Regional BFI time series (Buonaccorsi et al. Our BFI included all of the species studied by Howe 2001). We deemed fluctuations in food availability et al. (2012), and exhibited similar negative autocor- À relations at a 1-year lag, both province-wide (r 5 synchronous where A or Aij was .0.75. Because our sample sizes were small (i.e., 7 or 8 20.79; P 5 0.048, 2-tailed) and within NER (r 5 annual observations of each variable), and the BFI 20.93; P 5 0.007, 2-tailed). In our data from NER was a rank variable, we used Spearman rank and across Ontario, years with poor food, high levels correlation to test for trends and for relationships of HBC, and large harvest tended to be followed by years with better food, lower levels of HBC, and among food, HBC, and harvest. We performed all smaller harvest. This tendency could have led to correlations on data from the entire province, and spurious negative correlations between HBC and separately within each of the 3 administrative harvest in the previous year if both HBC and harvest regions. Sudbury District in NER consistently had were responding similarly to variation in food avail- more occurrences than other Districts, which we ability. Therefore, we crossed measures of HBC only attributed to unique landscape characteristics where with the 2- and 3-year average of prior harvests in the interspersion of suburban development and NER and Ontario. We addressed objective 6 using blueberry (Vaccinium spp.) barrens associated with correlations between total harvest and the number of soils acidified by past smelting activity (Winterhalder hunters. We manually calculated measures of agree- 1996) brought large numbers of bears into close ment in the direction of change in food availability proximity with humans during the fruiting season across years. We performed all correlations in R for . We also attributed the high level version 3.0.2 (R Development Core Team 2013). We of occurrences to a long-term research study of calculated the null distribution of the Spearman problem activity by black bears near Sudbury, where Rank correlation statistic (r) as the pre-computed researchers actively solicited reports of problem exact distribution for n , 22 following van de Wiel bears near developed areas (Landriault et al. 2009). and Di Bucchianico (2001) as implemented in the To control for the influence of data from Sudbury ‘‘pspearman’’ package (Savicky 2009). District, we duplicated occurrence data for NER and the province excluding occurrences from there, and tested for trends and correlations with food avail- Results ability and prior harvest using both the complete and Pairs of District-specific BFI values agreed in their the censored data sets. change in direction from one year to the next 77.5%

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Table 2. Mean, standard deviation (SD), and coefficient of variation (CV) of annual numbers of occurrences of human–bear conflict, traps set at the site of human–bear conflict, total harvest, hunter success rate, and the percent females in the harvest in Ontario, Canada, and within each of 3 administrative regions within Ontario, 2004–2011. NER = Northeast Region, NWR = Northwest Region, SR = Southern Region. Hunter success rate Percent females in Occurrences Traps set Total harvest (%) harvest Spatial extent Àx SD CV Àx SD CV Àx SD CV Àx SD CV Àx SD CV Ontario 9,807 2,321 0.24 1,084 325 0.30 5,613 381 0.07 30.0 2.3 0.08 36.1 1.8 0.05 Ontarioa 7,694 1,897 0.25 NER 4,953 1,688 0.34 552 201 0.36 2,593 367 0.14 31.5 3.6 0.11 35.3 2.7 0.08 NERa 2,840 1,064 0.37 NWR 1,830 341 0.19 393 101 0.26 1,679 167 0.10 36.9 3.7 0.10 37.6 1.9 0.05 SR 3,024 766 0.25 139 64 0.46 1,340 224 0.17 18.3 1.7 0.09 35.7 4.0 0.11 aExcluding data from Sudbury District. of the time in NER and 85% of the time in SR Fig. 2). Correlations were stronger when data for (indicating synchrony), but only 54% of the time in Sudbury District were excluded (for Ontario, r 5 NWR. Most cases of disagreement in NER occurred 20.88, P 5 0.004; and for NER, r 5 20.83, P 5 during 2010, when late spring frosts (which can kill 0.008; Fig. 2). The number of traps set was also and hence reduce fruit production; Usui negatively correlated with the BFI at the provincial et al. 2005) of varying severity occurred in different scale (r 5 20.86, P 5 0.012) and for both NER (r 5 Districts within the Region. Regional BFI scores 20.89, P 5 0.006) and SR (r 5 20.75, P 5 0.033; agreed in their direction of change 100% of the time Fig. 2). when NER and SR were compared, but only 43% of Total harvest was negatively correlated with food the time when NER or SR were compared with availability for Ontario (r 5 20.90, P 5 0.002) and NWR. for NER (r 5 20.83, P 5 0.008; Fig. 3). Hunter On average, nearly 10,000 occurrences of HBC success rate was negatively correlated with the BFI were reported, .1,000 traps were set, and nearly in NER (r 5 20.83, P 5 0.008). The proportion of 20,000 licensed hunters harvested approximately females in the harvest was negatively correlated with 5,600 bears (of which 36% were females), annually the BFI in Ontario (r 5 20.86, P 5 0.005), and in during the study period (Table 2). The Northeast NER (r 5 20.81, P 5 0.008; Fig. 3), though there Region had the greatest numbers of occurrences and were no significant correlations with the proportion bears harvested (Table 2). Hunter success rates were of adult females in the harvest. noticeably lower in SR than in other Regions There were no significant correlations between (Table 2). Most hunters in NER and NWR were occurrences or traps set and the 2- or 3-year moving non-residents who targeted bears and hunted over averages of prior harvests at any spatial extent bait, whereas a greater proportion of hunters in SR (Table S1). There were no significant correlations were residents who hunted bears opportunistically between either occurrences or traps set and total while hunting white-tailed deer (Odocoileus virginia- harvest in the previous year in NWR or SR (Table nus) or moose (Alces alces; Dix-Gibson 2012). S1). We did not test for correlations between total The number of hunters in NWR declined during harvest in the previous year and measures of HBC in the study (r 5 20.88, P 5 0.007), though there was Ontario or NER because food data were temporally no significant trend in total harvest (r 5 20.69, P 5 autocorrelated. 0.069). There were no other significant trends across Total harvest was significantly positively correlat- years in measures of HBC (occurrences, traps set) or ed with the number of hunters in SR (r 5 0.74, P 5 harvest metrics (total harvest, or the no. of hunters) 0.023), but not in other Regions or at the provincial at either the provincial scale or within other Regions scale. (Table S1). The number of occurrences was significantly negatively correlated with the BFI at the provincial Discussion scale (r 5 20.79, P 5 0.014), and in both NER (r 5 Synchronous variation in food availability for 20.67, P 5 0.042) and SR (r 5 20.98, P , 0.001; black bears over a vast landscape, including much of

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Fig. 3. Relationships between the bear food index Fig. 2. Relationships between the bear food index and each of total harvest (top panel) and the percent and each of occurrences of human–bear conflict (top of harvested bears that were females (bottom panel) panel) and traps set at the site of occurrences in Ontario, Canada (squares), and in the Ontario (bottom panel) in Ontario, Canada (squares), and in Ministry of Natural Resources’ Northeast Region the Ontario Ministry of Natural Resources’ Northeast (circles), 2004–2011. Lines show least-squares Region (circles) and Southern Region (triangles), linear fits. 2004–2011. Lines show least-squares linear fits. Closed symbols and dashed lines show data exclud- ing Sudbury District. areas of Minnesota, USA (Garshelis and Noyce 2008), and Yukon, Canada (Krebs et al. 2010). northeastern and southern Ontario, enabled us to Because black bear behavior, reproduction, vulner- detect significant negative correlations between food ability to harvest, and propensity to come into availability and each of HBC and harvest at conflict with humans are closely linked to variation landscape scales. No significant relationships were in food supply (Garshelis and Pelton 1981; Rogers detected in NWR, where the BFI did not vary 1987; Noyce and Garshelis 1997; Costello et al. 2003; synchronously among Districts (BFI data from Ryan et al. 2004, 2007; Obbard and Howe 2008; NWR were sparse compared with other Regions). Howe et al. 2010), an understanding of spatiotem- Significant correlations at the provincial scale were poral patterns in forage production is likely to be likely a result of the consistent patterns of variation important to other studies of black bears at in the BFI and measures of HBC and harvest that landscape scales. prevailed in NER and SR. Our BFI was largely Variation in berry production is at least partially based on annual rankings for soft-mast–producing attributable to variation in weather conditions species, which were previously shown to vary during flowering and fruiting (Laine and Henttonen synchronously within and among species over 1983; Sela˚s 1997, 2000; Usui et al. 2005; Krebs et al. subsets of our study area (Obbard et al. 2003, Howe 2009; Holden et al. 2012; Howe et al. 2012). Wide- et al. 2012, Romain et al. 2013). Similar synchrony in spread droughts helped synchronize berry produc- soft mast production was also observed over large tion within and among species over much of NER

Ursus 25(2):98–110 (2014) FACTORS AFFECTING HUMAN–BEAR CONFLICT N Obbard et al. 105 and SR from 1998 to 2009 (Howe et al. 2012). Major management policy (Garshelis and Noyce 2008, climatic oscillations, such as the North Atlantic Gore and Knuth 2009, Howe et al. 2010). However, Oscillation and the El Nin˜o Southern Oscillation, during our study, management policy in Ontario was influence weather conditions similarly over large constant. Serious attacks on humans in Ontario areas, so they could promote spatially synchronous occurred in September 2005 and May 2010. Provin- variation in fruit production (Hurrell et al. 2003, cially, occurrences were large in 2005 (the third- Stenseth et al. 2003). The effects of the North highest in our time series), but this coincided with the Atlantic Oscillation and El Nin˜o Southern Oscilla- second-poorest year for bear foods. During 2006, tion differ qualitatively within Ontario, notably occurrences were low, coinciding with a high BFI along the boundary between Canada’s Continental score, so it seemed there was no lag effect of and Atlantic climatic regions, which roughly bisects publicity surrounding the 2005 attack. Similarly, NWR into eastern and western halves (Bonsal et al. occurrences were about average during 2010 and 2001). This may explain the lack of spatial synchrony there were fewer occurrences during 2011 than any in BFI data, and lack of significant relationships other year, so the highly publicized attack in May between the BFI and HBC or harvest in NWR 2010 apparently had little effect on the reporting rate though it seems likely that the sparse BFI data from for HBC. Furthermore, traps were set by OMNR NWR was also a contributing factor. The influences staff only where deemed necessary after a site of major climatic oscillations on local weather are investigation, thus providing a measure of HBC that somewhat predictable (Hurrell et al. 2003). If we was less dependent on the reporting rate, and could identify relationships between climate indices relationships between the BFI and the number of measured during the winter and berry production traps set were similar to those between the BFI and the following growing season, we would have an occurrences. Though we cannot infer causal rela- opportunity to predict food shortages and associated tionships from correlation analyses, our results increases in vulnerability to harvest and HBC, as corroborate previous studies that demonstrated demonstrated by Zack et al. (2003). For example, negative relationships between the frequency of Holden et al. (2012) recently showed that maximum conflict between humans and American black bears snow-water equivalent can influence production of and fruit or nut production by preferred forage serviceberries (Amelanchier spp.), which are an species (Jorgensen et al. 1978, Peine 2001, Ryan et al. important food for black and brown bears (U. 2007, Garshelis and Noyce 2008, Howe et al. 2010, arctos) in western North America. The potential to Baruch-Mordo et al. 2014). Where humans coexist predict food supply for bears from winter weather with established bear populations, annual variation variables or climate indices warrants additional in the frequency of HBC is apparently well-explained research. by variation in food supply for bears. However, we Although measures of HBC and harvest metrics acknowledge that this might not be true where bear varied among years, they did not change direction- populations are recovering or expanding their range ally over time except for the decline in the number of (Cotton 2008), or where human development or hunters in NWR. This decline was attributable to a recreational use is expanding quickly into occupied .20% decline in the number of non-resident hunters bear habitat (Singer and Bratton 1980, Miller 1990, in NWR after 2008 (Dix-Gibson 2012), which was Schirokauer and Boyd 1998). likely an effect of the economic downturn in the Total harvest and the proportion of females in the . Our sample sizes (n 5 7 or 8 yr) harvest were also negatively correlated with food limited our ability to test for trends while accounting supply in NER and at the provincial scale. Appar- for variation associated with changes in food supply, ently bears, particularly female bears, were more but with longer time series multivariate approaches vulnerable to harvest when food was scarce, as could improve our ability to detect trends in previously observed in Minnesota and Virginia fluctuating data. where baiting was also a common hunting method In our study, the frequency of HBC was negatively (Noyce and Garshelis 1997, Ryan et al. 2004). correlated with food availability in 2 of 3 Regions Increased vulnerability to harvest likely resulted (NER, SR), and across all of Ontario. The reporting from increases in home range size and movement rate for HBC, and therefore our occurrence data, rates during poor food years (Garshelis and Pelton could vary with changes in media coverage and 1981, Pelchat and Ruff 1986, Rogers 1987, Garshelis

Ursus 25(2):98–110 (2014) 106 FACTORS AFFECTING HUMAN–BEAR CONFLICT N Obbard et al. and Noyce 2008), and increased attraction of bears populations to very low densities. This might be at to hunters’ baits (Noyce and Garshelis 1997, Ryan odds with the objectives of maintaining viable et al. 2004). In the case of female bears, food stress populations and providing sustainable sport-harvest associated with the need to gain considerable body opportunities over the long term. A better strategy mass in order to produce cubs the following winter, for management agencies would be to develop a or with the energetic costs of lactation, may have thorough understanding of causes of HBC in their exacerbated these effects. However, despite detecting jurisdiction (Baruch-Mordo et al. 2008), promote an increase in the proportion of females in the programs that focus on practical solutions that deny harvest, we did not detect an increase in the bears access to the many kinds of anthropogenic proportion of adult females in the harvest during attractants, and develop educational programs that poor food years in our study. In Ontario, sex is encourage the public to accept responsibility for reported for approximately 83% of bears harvested, their role in the human–bear conflict dyad. whereas only approximately 40% of premolar teeth In SR, total harvest was a function of the number are submitted for aging (Dix-Gibson 2012). Because of hunters rather than food supply. This could reflect ages are known for only about 40% of females an increase in hunting pressure in SR since 2008, harvested, increasing the proportion of teeth sub- the greater diversity of autumn foods in SR, or mitted might enable us to tease out any effects of differences in hunting methods among Regions. food supply on adult females. Although we did not detect significant trends in the We found no significant correlations between number of hunters or total harvest, both were harvest and subsequent HBC. Although it may be highest during the last 3 years of the study in SR, intuitive to assume that harvesting more bears which would have contributed to the significant should reduce HBC, empirical support for this correlation. Black bears in the boreal forest of assumption is lacking despite considerable research Ontario rely heavily on the of a few (Garshelis 1989, Treves and Karanth 2003, Huygens species (blueberry, bunchberry [Cornus canadensis], et al. 2004, Tavss 2005, Treves 2009, Howe et al. raspberry [Rubus sp.], mountain-ash, pin cherry 2010, Treves et al. 2010). Based on a survey of North [Prunus pensylvanica], and beaked hazel [Corylus American jurisdictions regarding harvest manage- cornuta]) during the hunting season (Romain et al. ment regimes and trends in HBC, Hristienko and 2013). Fruit production by these species, and McDonald (2007) argued that more liberal hunting consumption by black bears, varies considerably regimes would reduce HBC. However, they present- among years (Usui et al. 2005, Romain et al. 2013). ed no quantitative data demonstrating that liberal Furthermore, fruit production by many of these harvesting anywhere actually reduced the frequency species is adversely affected by late spring frosts and of HBC; liberal hunting regimes may simply by dry or cool summer weather conditions (Usui promote greater public acceptance of bears, thus et al. 2005, Howe et al. 2012, Romain et al. 2013). resulting in fewer complaints (Treves and Karanth Should several of these crops fail in the same year, 2003, Howe et al. 2010). bears would likely be strongly attracted to hunters’ We acknowledge that our short time series and the baits. Conversely, in SR oak (Quercus spp.) and beech lack of significant trends in either total harvest or (Fagus grandifolia) may provide more abundant HBC may limit the scope of inference from our data natural food during the hunting season when with respect to effects of harvest on the frequency of available, and bears have more opportunity to exploit HBC. However, given the degree to which natural alternative foods such as feral apples, agricultural food supply for black bears varies among years crops, fruit trees, beehives, garbage, and other (Costello et al. 2003, Usui et al. 2005, Romain et al. anthropogenic food sources. Therefore, the vulnera- 2013, Koenig and Knops 2014), and the importance bility of bears to harvest in SR might be less sensitive of bottom-up processes to bear demography (McLel- to poor production by the species included in our BFI. lan 2011), it seems likely that bears at a range of Differences in hunting methods among Regions may population densities will attempt to access anthro- also be relevant. Most non-residents hunted over bait pogenic food sources and come into conflict with through the services of an outfitter, but many humans when natural foods are scarce. We suggest residents hunt bears opportunistically without the that reducing the associated risk of HBC through use of baits, and the proportion of bears taken by harvest would require high harvest levels that reduce non-resident hunters was larger in NER (66%) than in

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SR (23%; Dix-Gibson 2012). Therefore, the success of Acknowledgments hunters in SR may be less dependent on the We thank the many staff members of the Ontario attractiveness of bait than success in NER. Ministry of Natural Resources who submitted The large number of correlations we performed wildlife food survey reports over the years. We also increased the chance of Type I errors arising in our thank the many outfitters and hunters who submit- study. We did not correct significance thresholds for ted harvest reports. Funding for the study was multiple comparisons because the power of tests was provided by Wildlife Research and Monitoring already low because of small samples, and we Section, Ontario Ministry of Natural Resources. acknowledge that significant results could have Comments from the Associate Editor and 3 anon- occurred randomly. However, we suggest that given ymous reviewers helped us improve an earlier the consistency of the correlations among the BFI, version of the manuscript. HBC, and harvest metrics where food varied syn- chronously, it is unlikely that these were randomly occurring Type I errors rather than real patterns. Literature cited BARUCH-MORDO, S., S.W. BRECK, K.R. WILSON, AND D.M. THEOBALD. 2008. Spatiotemporal distribution of black bear–human conflicts in Colorado, USA. Journal of Management implications Wildlife Management 72:1853–1862. Rank indices of food availability, such as our BFI, ———, K.R. WILSON, D.L. LEWIS,J.BRODERICK, J.S. can be obtained at low cost and have proven useful MAO, AND S.W. BRECK. 2014. Stochasticity in natural in explaining annual variation in black bear harvest forage production affects use of urban areas by black metrics and in the frequency of HBC (Noyce and bears: Implications to management of human–bear Garshelis 1997; Ryan et al. 2004, 2007; Garshelis and conflicts. PLoS ONE 9:e85122. Noyce 2008; Howe et al. 2010). We recommend that BONSAL, B.R., A. SHABBAR, AND K. HIGUCHI. 2001. Impacts food availability surveys be continued or expanded of low frequency variability modes on Canadian winter temperature. International Journal of Climatology in Ontario, and implemented by other jurisdictions 21:95–108. that manage black bears but do not currently BUONACCORSI, J.P., J.S. ELKINTON, S.R. EVANS, AND A.M. measure food availability. LIEBHOLD. 2001. Measuring and testing for spatial This study was made possible by the implementation synchrony. Ecology 82:1668–1679. of province-wide, standardized collection of food avail- COSTELLO, C.M., D.E. JONES, R.M. INMAN, K.H. INMAN, ability and HBC data in 2004. Consistent management B.C. THOMPSON, AND H.B. QUIGLEY. 2003. Relationship from 2004 to 2011 reduced the potential for confound- of variable mast production to American black bear ing effects of management regime changes on harvest reproductive parameters in New Mexico. Ursus 14:1–16. metrics or the reporting rate for HBC. We did not COTTON, W. 2008. Resolving conflicts between humans and augment the time series with data from more recent the threatened Louisiana black bear. Human–Wildlife Conflicts 2:151–152. years because in 2012 the agency’s response to reports DIX-GIBSON, L. 2012. Black bear provincial, regional and of HBC changed. Site visits were not conducted as black bear ecological zone harvest summaries, 1987– frequently, and bears were not trapped and relocated 2011. Unpublished report. Southern Science and except under exceptional circumstances. We suspect Information Section, Ontario Ministry of Natural that the public responded to this change with a Resources, Bracebridge, Ontario, Canada. reduction in the reporting rate for HBC. The number GARSHELIS,D.L.1989.Nuisancebearactivityandmanage- of reported occurrences dropped sharply during 2012 ment in Minnesota. Pages 169–180 in M. Bromley, editor. (to 5,800 province-wide) despite widespread soft-mast Bear–people conflicts. Proceedings of a symposium on failures, and dropped again during 2013 (to 2,419). management strategies. Northwest Territories Depart- Similarly, in Minnesota, calls about HBC to the ment of Renewable Resources, Yellowknife, Northwest managing agency decreased and remained low after Territories, Canada. ———. 1991. Monitoring effects of harvest on black bear the agency stopped trapping and removing bears from populations in North America: A review and evaluation the site of HBC (Garshelis and Noyce 2008). The of techniques. Eastern Workshop on Black Bear potential effects of changes in the management regime Research and Management 10:120–144. on the reporting rate for HBC should be considered ———, AND H. HRISTIENKO. 2006. State and provincial when drawing inferences from changes in the number estimates of American black bear numbers versus of incidences of HBC reported by the public. assessments of population trend. Ursus 17:1–7.

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Supplemental material set against number of hunters; and for proportion Table S1. Spearman rank correlations for occurrenc- (Prop) of adult females (ad F) in the harvest and mean es, traps set, total harvest, and number of hunters set age of harvest set against bear food index. Spatial against year and bear food index (BFI); for occurrences extents are Northeast Region (NER), Northwest Region and traps set against harvest in year prior, or 2- and 3- (NWR), Southern Region (SR), or all of Ontario (Prov). year moving average of prior harvest; for total harvest Data are for Ontario, Canada, 2004–2011.

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