Northeastern Naturalist 830 M.D. Jones, A.N. Tri, J.W. Edwards, and H. Spiker 22001155 NORTHEASTERN NATURALIST 2V2(o4l). :2823,0 N–8o4. 14 Home-Range Dynamics of Female Ursus americanus (Pallas) (American Black Bear) in a Recovering Population in Western Maryland Michael D. Jones1, Andrew N. Tri1,2,*,, John W. Edwards1, and Harry Spiker3 Abstract - Western Maryland’s population of Ursus americanus (American Black Bear; hereafter Black Bear) was nearly extirpated by the 1950s but recovered to 326 individuals by 2005. A knowledge gap currently exists regarding home-range dynamics of this recovering population. One of the most basic questions that managers wish to understand is how much space these Black Bears are using. To provide this information, we examined the home-range dynamics of 18 adult female Black Bears in western Maryland from 2006 to 2007 using GPS collars. We predicted that home-range estimates in our study population would be similar to that of surrounding states because Black Bear populations have been recovering for the past 50 years throughout Appalachia. Fixed-kernel estimates for spring, summer, and fall home ranges were 8.9 km2, 15.4 km2, and 20.7 km2, respectively. Fall and summer home ranges were similar, and both were larger (P < 0.10) than spring home ranges. Solitary females had spring home ranges 6.9 times larger than females with cubs, but ranges for all females were similar during other seasons. Home-range fidelity among seasons was high. As predicted, home-range sizes were comparable to those from other Appalachian states. With our results, managers can better understand space use of Black Bears in this recovering population. Introduction The home range of an animal is most simply described as “that area traversed by the individual in its normal activities of food gathering, mating, and caring for young” (Burt 1943:352). Estimating the size and spatial distribution of Ursus americanus (Pallas) (American Black Bear; hereafter Black Bear) home ranges is potential very useful to researchers and wildlife managers. Female Black Bears can be territorial and their home ranges are affected by the distribution of food sources (Garshelis and Pelton 1981, Jonkel and Cowan 1971, Lindzey and Meslow 1977, Rudis and Tansey 1995). Therefore, female Black Bear home ranges are often indicative of the overall habitat quality of an area (Ford 1983, Koehler and Pierce 2003). The size and spatial distribution of home ranges change seasonally as Black Bears shift to the most abundant and nutritious food sources available (Garshelis and Pelton 1981, Rogers 1987). Moreover, because females are territorial, higher Black Bear density may cause a reduction in average home-range size (Oli et al. 1Division of Forestry and Natural Resources, West Virginia University, Morgantown, WV 26506. 2Current address - Forest Wildlife and Populations Research Group, Minnesota Department of Natural Resources, Grand Rapids, MN 55744. 3Game Mammal Section Leader, Maryland Department of Natural Resources – Wildlife and Heritage Service, Mt. Nebo WMA, Oakland, MD 21550. *Corresponding author - andrew.tri@state.mn.us. Manuscript Editor: James Cardoza Northeastern Naturalist Vol. 22, No. 4 M.D. Jones, A.N. Tri, J.W. Edwards, and H. Spiker 2015 831 2002, Young and Ruff 1982). There is evidence that 2 factors reduce the territoriality of female Black Bears: genetic relatedness and abundant food sources (Elowe 1984, Reynolds and Beecham 1980, Rogers 1987). Female Black Bears generally exhibit some degree of natal philopatry (Costello 2010, Lee and Vaughan 2003, Reynolds and Beecham 1980), so there is often more overlap between the home ranges of sows and their female offspring than between sows and unrelated females (Moyer et al. 2006). Female Black Bears also have been shown to be less territorial around highly abundant food sources (e.g., garbage dumps) where resource competition is lower (Rogers 1987, Young and Ruff 1982). Black Bear home-range size varies widely across North America, with reported female home ranges from 0.9 km2 in Louisiana (Leigh 2007) to 294.8 km2 in Manitoba (Pacas and Paquet 1994). In the Appalachian region, female (both adult and subadult) home-range estimates vary from 5.0 to 49.0 km2 (Table 1). Only 2 studies have analyzed Black Bear home-range dynamics in Maryland (Table 1; Dateo 1997, Webster 1994), both of which occurred in the same study area as our research. The Black Bear population in western Maryland has increased dramatically since the 1990s, while the habitat has been altered by human development. Due to these changes, we expected current home-range dynamics to differ from those in the previous Maryland studies. We used GPS telemetry to examine seasonal home-range sizes, shifts, and overlap of female Black Bears in western Maryland. Field-Site Description We conducted our study in Garrett County, MD, which is the westernmost county in the state. Garrett County encompasses 1722 km2 and is bordered by Pennsylvania to the north and West Virginia to the south and west. Maryland’s Black Bear population occurs at its highest densities in Garrett County and adjacent Allegany County to the east, which is the only area in the state where Black Bear hunting is currently permitted (Spiker 2011). Elevations on the study area range from 292 to 1028 m. Table 1. Home-range estimates for female Ursus americanus (American Black Bear) in the Appalachian Region. Some of these estimates include subadult female bears. If separate adult female estimates were recorded in literature source, we used those. n = sample size within each study. MCP = minimum convex polygon. Mean home-range State Author size (km²) n Estimator Maryland Webster 1994 41 3 MCP Maryland Dateo 1994 36 5 MCP North Carolina Brody 1984 17 11 MCP North Carolina Jones and Pelton 2003 8 13 MCP Pennsylvania Alt et al. 1980 41 12 Bivariate normal Tennessee Quigley 1982 5 10 MCP Virginia Garner 1986 22 25 MCP Virginia Higgins 1997 7 27 Fixed kernel Virginia Olfenbuttel 2005 30 76 Fixed kernel West Virginia Brown 1980 49 8 Bivariate normal West Virginia Kraus 1990 26 15 MCP Northeastern Naturalist 832 M.D. Jones, A.N. Tri, J.W. Edwards, and H. Spiker 2015 Vol. 22, No. 4 The human population density in the area is relatively low at 18 persons/km2 (US Census Bureau 2010). Approximately 22% of the area is public land (GCPC 2008), including several large contiguous public areas. The 221-km2 Savage River State Forest, which is open to Black Bear hunting, is the largest public area in the county and makes up almost 13% of the study area. The majority of the study area is forested, with approximately 68% of the total area covered in deciduous forests. Five different forest-type groups occur on the area: (1) (Oak/hickory (dominated by Quercus spp. and Carya spp.) is the most common, making up 54% of all forested land in Garrett County; (2) Northern hardwood (dominated by Acer rubrum L. [Red Maple], A. saccharum Marsh. [Sugar Maple], Fagus grandifolia Ehrh. [American Beech], Betula alleghaniensis Britton [Yellow Birch], and Prunus serotina Ehrh. [Black Cherry]) is also common, making up 33% of forests; (3) Elm/Ash/Red Maple (dominated by Ulmus spp., Fraxinus spp., and Red Maple); (4) White/Red Pine (dominated by Pinus strobus L. [White Pine] and P. resinosa Aiton [Red Pine]); and (5) Spruce/Fir ( dominated by Picea spp. and Abies spp.). The latter 3 types all account for ≤7% of the total forested land (USFS 1999). The understory vegetation in these forest-type groups includes Kalmia latifolia L. (Mountain Laurel), Rhododendron spp. (rhododendron), Amelanchier arborea (F. Michx.) Fernald (Serviceberry), Cornus spp. (dogwood), and Corylus spp. (hazelnut), which are important sources of food and thick cover for Black Bears. Methods Black Bear GPS data collection Maryland Department of Natural Resources (MDNR) captured Black Bears in Garrett County during 2006–2007 using barrel traps, spring-activated foot snares, and running with hounds (Table 2; H.A. Spiker, MDNR, Oakland, MD, pers. comm.). Black Bears were immobilized using an injection of ketamine (4.4 mg/ kg) and xylazine (2.2 mg/kg). Females with neck circumferences of >48 cm were fitted with Lotek Model 3300S GPS collars weighing 285 g (Lotek Wireless, Inc. Newmarket, ON, Canada). Black Bears with neck circumferences <48 cm do not allow for the proper positioning of the collar antenna, which reduces the accuracy of the GPS coordinates. All individuals were marked with a metal ear tag (Self- Piercing “Round Post” Ear Tag – Style 56; National Band and Tag, Newport, KY) with a unique identification number. We recorded morphometric measurements as well as sex and reproductive status for each captured Black Bear. A premolar was extracted from each Black Bear and used to age the individual using cementum annuli techniques (Matson’s Laboratory LLC, Milltown, MT). We programmed the GPS collars to record a waypoint every 4 hours, and the battery life was approximately 1 year. Each waypoint recorded latitude and longitude at the current position, along with the date and time. Each collar also emitted a VHF signal to locate the Black Bear in the event of a GPS failure. If a collar remained stationary for >12 hrs, the collar would emit a unique VHF “mortality signal.” We then located these collars to determine the cause of the lack of movement. We recorded any mortalities Northeastern Naturalist Vol. 22, No. 4 M.D. Jones, A.N. Tri, J.W. Edwards, and H. Spiker 2015 833 or slipped collars (i.e., collar was removed by the individual) and recovered data from the collar if possible. For Black Bears remaining collared during the denning season, we located den sites using the VHF signal. We serviced the collars and downloaded the GPS data during MDNR’s annual den checks. Home-range analysis We estimated seasonal home ranges using GPS data. These estimates represented a sample of all adult female bears (3.5 years or older). We estimated seasonal home ranges based on 3 seasons: spring (den emergence–2 June), summer (3 June– 30 August), and fall (1 September–den entry). The 2 June division represents the approximate separation date between adult females and their yearlings (Lee and Vaughan 2004, Schwartz and Franzmann 1992) and the approximate shift in diet to soft mast for the region. This date also was prior to the peak breeding season for the mid-Appalachian region (Echols 2000, Ryan 1997). We chose the 1 September division to approximate the timing of the annual shift from soft mast to hard mast as the primary food source for Black Bears in the Appalachian region (Powell et al. 1997). We identified den emergence using the first sustained movement from the den site and defined time of den entry as the date when fall movement ceased or became drastically reduced. Den emergence ranged from 19 March to 10 May and den entry ranged from 29 November to 21 January. We used 95% minimum convex polygon (MCP) and 95% fixed-kernel density estimates for seasonal home-range estimates. We used the MCP method to facilitate comparison with earlier Black Bear home-range studies, specifically the 2 studies conducted in western Maryland (Dateo 1997, Webster 1994). We chose the 95% Table 2. Seasonal tracking of adult female Ursus americanus (American Black Bear) fit with GPS collars in Garrett County, MD, 2006–2007. x indicates locations were available during the above season for a given bear; – indicates no locations were available during this season for a given bear. Bear Year Spring Summer Fall A 2006 x x x B 2006 x – – C 2006 x x – D 2006 x x – E 2006 – – x F 2006 – x – G 2006 x x x H 2006 x x – I 2006 x x – D 2007 – x – J 2007 x x x G 2007 x – – E 2007 – x x K 2007 – – x L 2007 – x x M 2007 – x x B 2007 x x – N 2007 x x – Northeastern Naturalist 834 M.D. Jones, A.N. Tri, J.W. Edwards, and H. Spiker 2015 Vol. 22, No. 4 fixed-kernel density method to create a more informative and reliable home-range estimate (Seaman et al. 2000). We used the 95% level for both methods to exclude potential outliers that represented occasional travel outside of the defined home range. We used Geospatial Modeling Environment (Beyer 2011) in ArcGIS 10 for the fixed-kernel and MCP estimates. For seasonal home ranges, we only included Black Bears that had at least 30 locations for a given season (Girard et al. 2002). We calculated the appropriate fixed-kernel smoothing parameter (h) using least-squares cross-validation (LSCV), likelihood cross-validation (CVh) reference smoothing parameter (href), and proportions of href (0.4, 0.6, 0.8). We used a Kruskal-Wallis test to compare seasonal home ranges. If the Kruskal-Wallis statistic was significant, we used a pairwise Wilcoxon rank-sum test to compare medians of the groups. We used a Wilcoxon rank-sum test to examine home range differences between females with cubs and solitary females (i.e., Black Bears without cubs). Due to the small sample sizes, we used α = 0.10 to determine significance when comparing home-range sizes. To quantify seasonal shifts in home range, we used ArcGIS to create centroids for each seasonal home-range polygon and measured the distances between centroids. We also measured percent overlap of seasonal home ranges within and among individuals. We used Kruskal-Wallis tests to detect differences in home-range shifts between seasons as well as percent overlap among individuals. Results We included 7843 GPS locations from 18 adult female Black Bears in our home-range analyses, excluding all locations recorded during the denning period (Table 2). We tracked 9 Black Bears during 2006 and 9 during 2007. Five Black Bears were tracked for multiple years, although not always continuously; when bears were tracked for the same season among years, we randomly selected one season of data to preserve sample independence. We did not have enough data to estimate annual home-range sizes. We estimated 8 spring, 12 summer, and 6 fall home ranges (Jones 2012). The number of locations recorded for an individual Black Bear during a year was 56–498 (Table 3). Among fixed-kernel estimates, LSCV failed to select a smoothing factor for most Black Bears due to the large number of duplicate locations associated with GPS data. CVh led to obvious undersmoothing of most home ranges, causing highly fragmented polygons. The reference bandwidth resulted in oversmoothing, with polygons extending far beyond the extent of Black Bear locations. After Table 3. Home-range estimates for adult female Ursus americanus (American Black Bear) in Garrett County, MD, 2006–2007. 95% fixed kernel (km²) 95% MCP (km²) n No. locations Mean SE Range Mean SE Range Spring 8 124–333 8.93 3.63 0.07–35.54 6.73 1.31 0.05–34.79 Summer 12 81–369 15.40 2.74 3.72–48.01 10.57 1.60 4.14–27.75 Fall 6 56–498 20.66 7.47 5.18–80.95 14.18 3.86 3.03–39.35 Northeastern Naturalist Vol. 22, No. 4 M.D. Jones, A.N. Tri, J.W. Edwards, and H. Spiker 2015 835 visually comparing different proportions of href (0.4, 0.6, 0.8), we determined that 0.8href produced the most biologically relevant home-range polygons with minimal undersmoothing or oversmoothing, consistent with the results of Worton (1995). We used this bandwidth-selection method for all reported fixed-ke rnel estimates. Seasonal home-range size Mean spring, summer, and fall home-range sizes were 8.9 km2, 15.4 km2, and 20.7 km2, respectively (Table 3). Home-range size differed among seasons (H = 5.97, P = 0.07). Fall home ranges were larger than spring home ranges (P = 0.10), but not different than summer home ranges (P = 0.75). Summer home ranges also were larger than spring home ranges (P = 0.07). Reproductive status Among 33 seasonal home-range estimates of female Black Bears, 20 (60.6%) were from females with cubs: 14 (70.0%) were tracked during 2006 and 6 (30.0%) during 2007. Solitary females (n = 14) had larger spring and summer home ranges than females with cubs, but only spring home ranges were significantly different (W = 21, P = 0.02; Table 4). Mean spring home ranges of solitary females were 6.9 times larger than those of females with cubs. Home-range fidelity and overlap We monitored 11 Black Bears for consecutive seasons, allowing us to measure home-range fidelity. Home-range centroids shifted 0.22–2.40 km (mean = 0.85 km) between any 2 seasons. While mean home-range shift was slightly higher from summer to fall (0.88 km) than from spring to summer (0.83 km), no differences were detected (W = 28, P = 0.91). Reproductive status of Black Bears did not affect seasonal home-range shifts (W = 32.5, P = 0.97). Individual summer home ranges overlapped 55.2–100.0% (mean = 79.26%) of spring home ranges (Table 5). Similar overlap occurred with summer and fall home ranges, where individual fall ranges contained 30.9–98.3% (mean = 73.78%) of summer home ranges. We tracked 5 Black Bears over multiple years within the same season, allowing us to examine interannual differences in seasonal home ranges. One Black Bear had a spring home range 9.9 times larger in 2006 than 2007, even though Table 4. Seasonal 95% fixed-kernel home-range estimates based on reproductive status for adult female Ursus americanus (American Black Bear) in Garrett County, MD, 2006–2007. * = statistically different at α = 0.10. Solitary = females without cubs. 95% fixed kernel (km²) Season Reproductive status n Mean SE Range Spring Solitary 3 22.20* 8.09 7.60–35.54 With Cubs 7 3.24* 0.69 0.07–5.30 Summer Solitary 6 19.40 3.00 8.57–28.06 With Cubs 11 14.28 3.81 3.72–48.01 Fall Solitary 5 12.91 3.72 5.18–25.75 With Cubs 5 25.79 14.01 6.23–80.95 Northeastern Naturalist 836 M.D. Jones, A.N. Tri, J.W. Edwards, and H. Spiker 2015 Vol. 22, No. 4 that Black Bear had cubs in 2006 and not in 2007. Her 2006 spring home range encompassed the perimeter of her 2007 spring home range. Of the 3 Black Bears tracked over multiple summers, summer home-range size changed among years by 32.7–292.4%. The larger summer home range for each Black Bear overlapped their smaller summer home range by 71.5–97.9%. For the Black Bear tracked during 2 fall seasons, the home range from 2007, when she was solitary, overlapped her 2006 fall home range, when she had cubs, by 89.9%. Discussion Our MCP estimates of Black Bear spring, summer, and fall home ranges in western Maryland were smaller than others in our study region (Table 1; Dateo 1997, Webster 1994). A number of factors may have influenced differences between our MCP estimates and those from the 2 previous Maryland Black Bear studies. First, the difference in number of locations used to estimate home ranges may account for some of the discrepancy. While Dateo (1997) and Webster (1994) located each Black Bear approximately 3 times per week, our GPS collars collected locations at a much higher rate. MCP estimates are heavily influenced by the number of locations (Anderson 1982), so our estimates would be expected to differ somewhat from previous studies. Given the magnitude of difference among our estimates and those of Dateo (1997) and Webster (1994), the disparity in home-range estimates likely extends beyond sample-size differences and is more a result of biological, behavioral, and landscape changes occurring since those studies. The Black Bear population in the study area has increased rapidly in recent decades, with a 438% increase in Black Bear density from 1991 to 2011 (Jones 2012). This increase likely influenced differences between previous home-range estimates and those of our study. Female Black Bears are generally territorial, so the increase in Black Bear density could result in a reduction in home-range size (Oli et al. 2002, Young and Ruff 1982). Higher Black Bear density likely would increase intraspecific interactions for resources, causing female Black Bears to more aggressively defend resources within their home ranges. Limitations on the size of an area that a female Black Bear could successfully defend may have an indirect negative effect on overall home-range size. Recent habitat and land-use changes in western Maryland may also have influenced home-range size of female Black Bears. Garrett County is a popular vacation destination, and the construction of businesses and vacation homes has Table 5. Overlap of seasonal home ranges among adult female Ursus americanus (American Black Bear) in Garrett County, MD, 2006–2007. Area of overlap (km²) % overlap n Mean SE Range Mean SE Range Spring 4 1.70 0.11 1.59–1.82 50.11 21.87 4.47–100.00 Summer 6 10.27 3.01 1.31–22.18 41.24 12.00 15.29–97.90 Fall 2 4.77 0.00 4.77–4.77 12.21 6.32 5.89–18.52 Northeastern Naturalist Vol. 22, No. 4 M.D. Jones, A.N. Tri, J.W. Edwards, and H. Spiker 2015 837 increased rapidly in certain parts of the county. From 1990 to 2005, the human population in Garrett County increased only 6%, whereas the number of housing units rose 33% (GCPC 2008). Development can affect Black Bear home-range size by reducing the amount and connectivity of forest cover, and by increasing anthropogenic food sources. Urbanization tends to reduce the area of forested Black Bear habitat and to fragment remaining habitat and travel corridors, which may result in a decrease in home-range size. A potentially positive result of increased human presence is the availability of anthropogenic food sources. Anthropogenic food sources may supplement a Black Bear’s diet and allow them to secure adequate food resources within a smaller home range. The interspersion of forests and developed areas, common in some parts of Garrett County, could be considered high-quality Black Bear habitat in terms of cover and food availability. Previous studies have shown that Black Bear home ranges are generally smaller in higher-quality habitat (Garshelis and Pelton 1981, Koehler and Pierce 2003, Lindzey and Meslow 1977, Moyer et al. 2007). Our estimates of Black Bear spring home ranges were smaller than summer and fall home ranges (P < 0.10), which was consistent with previous Maryland Black Bear studies (Dateo 1997, Webster 1994), and those from Pennsylvania (Alt et al. 1980) and Virginia (Kasbohm et al. 1998). One explanation for the size difference was the relatively low activity levels of Black Bears immediately following den emergence. Tøien et al. (2011) found that Black Bears maintained reduced metabolic rates for up to 3 weeks after den emergence, even though body temperature returned to normal rather quickly. Garshelis and Pelton (1980) reported low spring activity levels in Black Bears in the Great Smoky Mountains National Park. They suggested that, because Black Bears largely relied on herbaceous material in spring, the low nutritional value of spring diets limited the energy Black Bears could expend traveling. Another possible explanation for this finding was that herbaceous material is easily available to Black Bears in the spring and they did not have to travel far to find food. Black Bears with cubs had smaller spring home ranges than solitary females, supporting the findings of previous studies in the region (Alt et al. 1980, Dateo 1997, Kasbohm et al. 1998, Webster 1994) and some from other parts of the United States (Moyer et al. 2007, Smith and Pelton 1990), but differing from others (Alt et al. 1980, Olfenbuttal 2005, Reynolds and Beecham 1980). Spring home ranges among females with cubs may be constrained in size by the limited mobility of their cubs immediately after den emergence. Cubs become much more mobile and independent during the summer, which may explain why we found no difference between summer or fall home ranges of solitary Black Bears and Black Bears with cubs. It should be noted that we only calculated 3 spring home ranges for solitary females. Our results may have been different with larger sample sizes or if we included comparisons of homerange sizes for individual females during years with and without cubs. Female Black Bears demonstrated strong home-range fidelity, which was reflected in minimal centroid shifts and high degrees of overlap between seasons within years. Our findings were similar to home-range shifts observed in Pennsylvania (Alt Northeastern Naturalist 838 M.D. Jones, A.N. Tri, J.W. Edwards, and H. Spiker 2015 Vol. 22, No. 4 et al. 1980) and Virginia (Olfenbuttel 2005). Although remaining relatively constant for most Black Bears, some individuals did show more marked changes in size and/ or shape of seasonal home ranges. We surmise that changes in home-range size and shape were influenced by resource availability (Powell et al. 1997), but we did not have a sufficient data to assess this relation. Despite varying differences in seasonal home-range sizes among years, most Black Bears showed extensive overlap. This result may be explained by variation in the quality, quantity, and spatial distribution of food resources on the landscape. Black Bears can adjust their home ranges to take advantage of the most abundant and nutritious foods available in a given season (Garshelis and Pelton 1981). In contrast to our extensive overlap, 83% of Black Bears in the Great Smoky Mountains National Park showed large shifts in seasonal home ranges, especially from summer to fall when they were seeking areas with more oak mast (Garshelis and Pelton 1981). MDNR fall hard-mast surveys for Garrett County classified 2007 as a mast failure, while the 2006 crop was rated as average (MDNR, Mt. Nebo, MD, unpubl. data). An absence of mast within a Black Bear’s current home range may require a shift to a new area to find an alternate food source. However, the availability of anthropogenic food sources may have helped buffer the effects of variation in natural food availability, resulting in high home-range overlap. The presence of refuse, bird feed, and other supplemental foods may provide enough energy to reduce the need for Black Bears to substantially shift home ranges among seasons or years. Due to the cost of GPS collars and technological issues with the collars, our sample sizes were small, which may have influenced our findings. It is possible that our pooling of data and unequal sample sizes among seasons and reproductive statuses reduced our ability to detect differences or similarities in home-range size. We also did not have adequate data to examine home-range size variation on an individual level. For these reasons, caution should be used when making inferences beyond our study population. Although our results are useful for managers, future home-range studies in the area would be helpful to provide additional data where our study had gaps. Managers and researchers often focus on female Black Bears because their survival and reproductive output strongly influence population dynamics. To best manage the female segment of a Black Bear population, it is essential to identify the spatial scale at which individual Black Bears select habitat. Our results will help managers better understand how management decisions (e.g., habitat management, recreational hunting) will impact the Black Bear population and at which spatial scale Black Bear management and research should be conducted. Our home-range estimates provide insight into female Black Bear movements that can help MDNR develop expectations for female movements based on season and reproductive status. 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