Microscale Den-Site Selection of Grizzly Bears in Southwestern Yukon

SHORT COMMUNICATION N Libal et al.

Microscale den-site selection of grizzly bears in southwestern Yukon

Nathan S. Libal1,4, Jerrold L. Belant1, Ramona based on spatial and temporal food availability, while Maraj2, Bruce D. Leopold1, Guiming Wang3, and selecting for security at smaller spatial scales (Ciar- Shelley Marshall2 niello et al. 2007). Multi-scale selection of denning habitat has been demonstrated in other carnivore 1Carnivore Ecology Laboratory, Forest and Wildlife species, including striped skunk (Mephitis mephitis) Research Center, Mississippi State University, Missis- and Canada lynx (Lynx canadensis) (Hwang et al. sippi State, MS 39762, USA 2007, Squires et al. 2008). Similar to grizzly bears, 2Environment Yukon, Government of Yukon, White- resource availability and security were important horse, YK Y1A2C6, Canada variables at large and small scales, respectively 3 Department of Wildlife, Fisheries, and Aquaculture, (Hwang et al. 2007, Squires et al. 2008). Mississippi State University, Mississippi State, MS Denning is a critical component of northern 39762, USA grizzly bear ecology, allowing bears to survive during winter when food is largely unavailable and provid- Abstract: Over-winter denning is a critical compo- ing a secure location for parturition (Nelson 1973, nent of grizzly bear (Ursus arctos) fitness. Conse- Harding 1976). At larger spatial scales, grizzly bears quently, identifying and protecting denning habitat in northern, mountainous environments appear to is important for grizzly bear management. We select den sites at higher elevations (reported ele- evaluated small-scale den-site selection by compar- vations vary, but are at moderate to high elevations ing grizzly bear den sites (n 5 21) in the Southern within respective study areas; Pearson 1975, Vroom Lakes region, Yukon, Canada to random locations et al. 1980, Libal et al. 2011). Higher elevations may within 100 m of dens. We measured indices of be selected in part for increased snow cover, pro- structural stability (slope at den opening, tree and viding improved thermal insulation (Vroom et al. shrub cover, percent soil ,2 mm), and security 1980). Where available, steep terrain is also charac- (horizontal cover) at each den site and random teristic of northern grizzly bear denning habitat location, and used mixed model analysis of variance (Schoen et al. 1987). Steep terrain may be selected to determine selection. Our results indicated selec- for isolation and security (Schoen et al. 1987, Groff tion for both steeper slopes and horizontal cover, et al. 1998), particularly by adult females (Libal et al. suggesting that stability and security are important 2011). components of den-site selection at small spatial At smaller spatial scales, dens are frequently scales. constructed in areas where tree roots, rocks, or other structures provide stability (Vroom et al. 1980, Key words: den-site selection, grizzly bear, micro- Ciarniello et al. 2005). Dens also are constructed scale, Ursus arctos, Yukon commonly on moderately steep slopes (20–40u), Ursus 23(2):226–230 (2012) which may contribute to ease of excavation, den stability, and optimal thermal properties (Pearson 1975, Vroom et al. 1980, Ciarniello et al. 2005). While empirical evidence is lacking, cover also may Habitat selection usually occurs at multiple spatial be an important characteristic of Ursus spp. bed and scales, where an animal first selects a home range den locations (Kusak and Huber 1998, Martorello and then decides which habitats within the home and Pelton 2003, Ordiz et al. 2011). Cover may range to use for different purposes (e.g., foraging, reduce predation or infanticide risk and human- mating; Johnson 1980). Multi-scale habitat selection derived risk, which have all been hypothesized to has been reported for grizzly bears (Ursus arctos), influence grizzly bear resource selection (Wielgus where selection varied between study areas and within and Bunnell 1995, Libal et al. 2011, Ordiz et al. and among individual home ranges (McLoughlin et al. 2011). 2002, Ciarniello et al. 2007). At large spatial extents To better understand den-site selection at smaller grizzly bears appeared food-limited, selecting habitats spatial scales, we investigated denning habitat at a microscale (within 100 m radius surrounding dens). [email protected] Because average home range for this population has

226 SHORT COMMUNICATION N Libal et al. 227 not been quantified, we were unable to evaluate through visual identification of bears, we used DNA selection at larger spatial scales. At microscale, we analysis of hairs (Woods et al. 1999). If neither predicted that excavated material would be predom- sightings nor DNA analysis could be used, we inantly sand, silt, and clay, because these particles conservatively classified dens that were above tree are more cohesive than larger soil structures (e.g., line as grizzly bear and rejected dens below tree line. gravels, cobbles). To further provide structural Based on differences between denning elevations of stability, we predicted that den openings would be sympatric grizzly bears and black bears (U. amer- located on steeper slopes and that surrounding icanus) in mountainous regions (Miller 1990, Aune vegetation would be predominantly trees and shrubs 1994), dens below tree line could have been either because these vegetation types have greater rooting grizzly bear or American black bear dens. Although depth than herbaceous vegetation. We also predicted using this elevation restriction likely excluded some greater horizontal cover at the den than surrounding grizzly bear dens at lower elevations, we believe it areas, which is an indicator of greater security. ensured no black bear dens were mistakenly included in our data set, given current literature and knowledge of our study area. Study area We measured microscale characteristics at the den The study area encompassed approximately site and 1 random point within 100 m of the den 6,800 km2 of the Southern Lakes region of Yukon, opening (Martorello and Pelton 2003, Waller et al. Canada (60u239000 N, 134u039000 W). Mean annual 2012). Random points were representative of areas temperature is about 22uC, with mean January and near known denning sites and were chosen using a July temperatures of 223u and 13uC, respectively random number generator to select a bearing (1– (Smith et al. 2004). Study area elevations range from 360u) and distance (1–100 m). At each location we about 600–2,600 m. The region is in the Coast estimated percent sand, silt, and clay; horizontal Mountains, within the rain shadow of the St. Elias cover; canopy cover of trees and shrubs; and slope Range and receives 200–325 mm annual precipita- steepness to describe den sites. To quantify percent tion, 50–75% as snow (Smith et al. 2004). White sand, silt, and clay, a 30 x 30 x 30 cm soil core was spruce (Picea glauca) and lodgepole pine (Pinus taken from the center of the excavation fan. From contorta) are common tree species, with white spruce soil cores, percent sand, silt, and clay was estimated occurring from valley bottoms to tree line on mesic visually and recorded as the percentage of particles slopes and lodgepole pine dominating flat, xeric ,2 mm (Ministry of Environment, Lands, and Parks areas (Smith et al. 2004). Grizzly bear population and Ministry of Forests 1998). We used a 0.5 x 1.0 m density is less than other areas of Yukon (Ramona cover board (divided into two 0.25-m2 squares; Maraj, unpublished data). In 1989 the population modified from Nudds [1977]) to estimate horizontal was estimated at 82–139 bears (13–22 bears/ cover. The cover board was placed at the den 1,000 km2; Larsen and Markel 1989). entrance or the random point. We recorded horizontal cover readings at 1 m above the ground to approximate a bear’s walking height, 11.3 m from Methods the board, taking our first reading from a random We identified grizzly bear dens using multiple direction (1–360u) and taking 3 subsequent readings methods. During April–May 2010 and 2011 we used 90u from the previous. We visually estimated tree fixed-wing aircraft to fly transects over the study and shrub cover as the percent canopy cover of all area and identify grizzly bear dens during den applicable species in an 11.3-m radius circle centered emergence. To locate additional dens, we used data on the den opening and random point. We excluded from radiocollared individuals in a concurrent vegetation data for dens old enough that vegetation grizzly bear study conducted by Environment cover may have changed since den excavation (e.g., Yukon. We also conducted ground surveys during dens with vegetative growth on excavated material). summer 2010 and 2011. While the use of air and Slope was estimated for a 3-m line centered on the ground surveys to locate dens may have resulted in den opening or random point using a clinometer. sampling bias, there were too few collared bears to We compared attributes of den sites to random rely solely on marked bears for den location. When points using mixed model analysis of variance den sites could not be attributed to grizzly bears (paired comparison test; PROC MIXED, SAS

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Table 1. Habitat attributes of grizzly bear den and random (100 m from dens) locations, Southern Lakes, Yukon, Canada 2010–11. Den Random Variable n Meana SE Range n Meana SE Range Slope (u) 21 30.9A 1.8 15.0–45.0 21 19.3B 2.3 2.0–40.0 Soil ,2mm(%) 20 60.5A 3.2 35.0–80.0 20 53.8A 5.9 0.0–100.0 Horizontal cover (%) 18 62.3A 5.6 18.0–100.0 18 45.8B 8.0 3.0–100.0 Tree and shrub cover (%) 18 60.6A 5.7 16.0–90.0 18 65.4A 4.9 32.0–100.0 aMeans not sharing a letter within rows differed (P , 0.10).

Institute Inc, Cary, North Carolina, USA). Because note that observed selection for greater horizontal of low statistical power due to low sample sizes, we cover may also reflect greater plant cover, and thus accepted an increased probability of committing a greater stability. Type I error by setting a 5 0.10 a priori (Grosbois We observed selection for greater slope, which was et al. 2008). To meet method assumptions, variables also consistent with our hypothesis. Mean value and with non-normal distributions were transformed range of slope were similar to other studies (Pearson using square root and arcsine transformations before 1975, Vroom et al. 1980, Ciarniello et al. 2005), and analyses. All means and standard errors from may represent the range of slopes providing stable analyses are reported in untransformed units. and thermally optimal conditions for an excavated den (Pearson 1975, Vroom et al. 1980, Ciarniello et al. 2005). Observed percentage soil sample ,2mm Results and tree and shrub cover did not differ from We located 21 grizzly bear dens (20 excavated, 1 random, and therefore did not support our hypoth- natural cave), 7 from aircraft, 11 by ground surveys, eses. Thus, we believe these were not accurate and 3 using coordinates from 2 GPS-collared adult metrics for stability at this scale. It is also possible female bears (1 individual was located at dens in that these metrics did not accurately reflect soil consecutive winters). Slope at the den opening was stability. Because of the placement and size of our about 60% steeper (F 5 26.05; 1, 20 df; P , 0.001) soil sample, as well as time elapsed between den than at random locations (Table 1). Horizontal excavation and soil sampling, our samples may not cover at the den was 36% greater (F 5 5.00; 1, 17 have accurately reflected the soil conditions during df; P 5 0.039) than random locations. Percentage of den excavation. In addition, within our 100-m radius soil sample ,2mm(F 5 1.26; 1, 19 df; P 5 0.275) surrounding dens, tree and shrub cover was relative- and tree and shrub cover (F1,17 5 0.68, P 5 0.421) ly homogenous and thus may not have been selected did not differ between dens and random locations. for at this spatial scale. Our results suggest that security and stability may influence den-site selection at small spatial scales. Discussion Which of these factors has a greater influence on We observed selection for greater horizontal den-site selection may depend on individual circum- cover, consistent with our hypothesis. Previous stances (e.g., bear age or sex, local environmental research has suggested that security is an important conditions). Management for grizzly bear denning component of grizzly bear habitat selection at habitat in the Southern Lakes region should include smaller spatial scales (Ciarniello et al. 2007). Risk protection of heavily vegetated, steeper slopes. avoidance may influence bear den- and bed-site Our data did not allow partitioning of dens by age selection, at least for some age–sex classes (Schoen or gender classes of bears. Variation in den-site et al. 1987, Groff et al. 1998, Kusak and Huber 1998, selection between adult male and female grizzly Martorello and Pelton 2003, Libal et al. 2011, Ordiz bears has been observed (Schoen et al. 1987, et al. 2011). Our results support risk avoidance being Haroldson et al. 2002, Libal et al. 2011). Adult an important factor at small spatial scales, which is female bears may spatially segregate from adult male consistent with den-site selection of other mamma- bears while denning to reduce risk to their repro- lian carnivores (striped skunk; Hwang et al. 2007, ductive investment (Libal et al. 2011). The 3 dens for Canada lynx; Squires et al. 2008). It is important to which we could attribute use by gender or age class

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were occupied by adult females, with 2 of these dens GROSBOIS, V., O. GIMENEZ, AND J.M. GAILLARD. 2008. exhibiting maximum observed horizontal vegetation Assessing the impact of climate variation on survival in cover. Future research should incorporate age and vertebrate populations. Biological Reviews 83:357–399. gender class data to investigate possible segregation. HARDING, L.E. 1976. Den-site characteristics of arctic Future research should evaluate selection at larger coastal grizzly bears (Ursus arctos L.) on Richards Island, Northwest Territories, Canada. Canadian Jour- scales (e.g., individual home ranges) to better nal of Zoology 54:1357–1363. understand grizzly bear den-site selection at multiple HAROLDSON, M.A., M.A. TERNENT, K.A. GUNTHER, AND spatial scales. Denning is primarily a mechanism for C.C. SCHWARTZ. 2002. Grizzly bear denning chronology energy conservation (Nelson 1973, Harding 1976), and movements in the Greater Yellowstone Ecosystem. with denning bears exhibiting substantial body mass Ursus 13:29–37. loss (Hilderbrand et al. 2000, Belant et al. 2006). HILDERBRAND, G.V., C.C. SCHWARTZ, C.T. ROBBINS, AND Grizzly bears in northern, mountainous regions T.A. HANLEY. 2000. Effect of hibernation and repro- typically select dens at higher elevation sites with ductive status on body mass and condition of coastal deep snow packs (Pearson 1975, Vroom et al. 1980, brown bears. Journal of Wildlife Management 64:178– Libal et al. 2011). Because deeper snow provides 183. greater insulation in the den, selection for these sites HWANG, Y.T., S. LARIVIE` RE, AND F. MESSIER. 2007. Local- may reduce the energetic cost of thermoregulation and landscape-level den selection of striped skunks on the Canadian prairies. Canadian Journal of Zoology during denning. Thus, we suggest that selection for 85:33–39. thermal cover may occur at large spatial scales and JOHNSON, D.H. 1980. The comparison of usage and avail- should be evaluated in future studies. ability measurements for evaluating resource preference. Ecology 61:65–71. KUSAK, J., AND D. HUBER, D. 1998. Brown bear habitat Acknowledgments quality in Gorski Kotar, Croatia. Ursus 10:281–291. We thank Environment Yukon; the Arctic Insti- LARSEN, D.G., AND R.L. MARKEL. 1989. A preliminary tute of North America; and the Department of estimate of grizzly bear abundance in the southwest Wildlife, Fisheries, and Aquaculture and Forest and Yukon. Yukon Fish & Wildlife Branch, Whitehorse, Wildlife Research Center at Mississippi State Uni- Yukon, Canada. versity for financial support and equipment. We also LIBAL, N.S., J.L. BELANT, B.D. LEOPOLD,G.WANG, AND P.A. OWEN. 2011. Despotism and risk of infanticide thank everyone who assisted with field work, in influence grizzly bear den-site selection. PLoS ONE particular the pilots of Alpine Aviation and Horizon 6:e24133. doi:10.1371/journal.pone.0024133. Helicopters, and K. Everatt. Thanks also to T. Jung MARTORELLO, D.A., AND M.R. PELTON. 2003. Microhabitat for equipment and helpful suggestions. characteristics of American black bear nest dens. Ursus 14:21–26. MCLOUGHLIN, P.D., R.L. CASE, R.J. GAU, H.D. CLUFF,R. Literature cited MULDERS, AND F. MESSIER. 2002. Hierarchical habitat AUNE, K.E. 1994. Comparative ecology of black and selection by barren-ground grizzly bears in the central grizzly bears on the Rocky Mountain Front, Montana. Canadian arctic. Oecologia 132:102–108. International Conference on Bear Research and Man- MILLER, S.D. 1990. Denning ecology of brown bears in agement 9(1):365–374. southcentral Alaska and comparisons with a sympatric BELANT, J.L., K. KIELLAND,E.H.FOLLMANN, AND L.G. black bear population. International Conference on ADAMS. 2006. Interspecific resource partitioning by Bear Research and Management 8:279–287. sympatric ursids. Ecological Applications 16:2333–2343. MINISTRY OF ENVIRONMENT,LANDS, AND PARKS AND MINIS- CIARNIELLO, L.M., M.S. BOYCE, D.C. HEARD, AND D.R. TRY OF FORESTS. 1998. Field manual for describing SEIP. 2005. Denning behavior and den site selection of terrestrial ecosystems. Second edition. Land manage- grizzly bears along the Parsnip River, British Columbia, ment handbook 25. Resource Inventory Branch, Minis- Canada. Ursus 16:47–58. try of Environment, Lands, and Parks and Ministry of ———, ———, D.R. SEIP, AND D.C. HEARD. 2007. Grizzly Forests, Victoria, British Columbia, Canada. bear habitat selection is scale dependent. Ecological NELSON, R.A. 1973. Winter sleep in the black bear: A Applications 17:1424–1440. physiologic and metabolic marvel. Mayo Clinic Pro- GROFF, C., A. CALIARI,E.DORIGATTI, AND A. GOZZI. 1998. ceedings 48:733–737. Selection of denning caves by brown bears in Trentino, NUDDS, T.D. 1977. Quantifying the vegetative structure of Italy. Ursus 10:275–279. wildlife cover. Wildlife Society Bulletin 5:113–117.

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