GRIZZLY BEAR HABITAT SELECTION: Along the Parsnip River, British Columbia Prepared for: British Columbia Ministry of Forests Dr. Dale R. Seip, Wildlife Ecologist Prince George Forest Region 1011 - 4th Avenue, Prince George, BC, V2L 3H9 Report Date: 22 March 2002 Prepared by: Lana M. Ciarniello Mark S. Boyce Hawthorne Beyer Department of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E9, Canada Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests Abstract Resource Selection Functions (RSF) were estimated to map the relative probability of occurrence of habitats for 49 radiocollared grizzly bears along the Parsnip River, British Columbia, Canada between April 1998 and February 2002. Twenty-two of these bears ranged in the plateau, a landscape heavily manipulated by humans. In contrast, 27 bears ranged in a relatively pristine mountainous landscape. Bears were monitored with VHF radiotelemetry twice per week using a fixed-wing aircraft. Population level RSF models were estimated at the landscape scale for the mountains and the plateau. In both landscapes greenness was the most significant predictor of bear occurrence followed by warm, southwest facing aspects (hillshade). Mountain bears selected for sub-alpine/balsam fir and non- forested mountain (e.g., alpine grasslands or avalanche chutes) greater than their reference habitat of white spruce. Surprisingly, at the landscape scale we could not detect significant selection for any of the landcover types by plateau bears. However, they selected against deciduous mixed forest, balsam fir, pine and nonforested plateau categories. Overall the mountain bears were more affected by covariates than plateau bears, with the proportion of variance explained in the mountains being greater. Therefore, the mountain model did a better job of predicting spatial patterns in the probability of use by grizzly bears than the plateau model. Resource Selection Ratios were examined for model covariates that could not be GIS referenced as well as to uncover mechanisms tha t would aid in increasing the predictive capability of the plateau model. Common to mountain and plateau bears was a significant selection for the ESSFwk2 biogeoclimatic zone. In the mountains, the AT zone was used in proportion to its availability. On the plateau, the SBSwk1 zone was highly selected by plateau bears but avoided by mountain bears. In both landscapes, bears selected areas at a straight-line distance of 5 to 11 km from a road far greater than expected by chance. The Limited Entry Hunt and Problem Wildlife databases were examined to determine areas of mortality risk for grizzly bears. Seventy-three percent of grizzly bears were killed within 1 km of a road. In order of importance, kills occurred most frequently near Highway 97, then secondary logging roads, primary trunk roads, and decommissioned roads. Bears who resided within the plateau were subjected to differential mortality risks than from bears residing in the mountains. We suggest that plateau bear occurrence is a combination of habitat related attributes and avoidance of high-risk areas. GIS layers that reflect active landscape change are currently being developed. In addition, more information on plateau bear habitat selection is required to adequately predict occurrence by grizzly bears in human- manipulated landscapes. Key words: British Columbia, forestry, grizzly bear, habitat availability, habitat selection, mountains, plateau, radiotelemetry, Geographic Information Systems, Resource Selection Function, Ursus arctos. Ciarniello, Boyce & Beyer ii Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests Table of Contents 1.0 INTRODUCTION .................................................................................................................... 1 2.0 STUDY AREA ......................................................................................................................... 1 3.0 METHODS ............................................................................................................................... 4 3.1 Capture and radiotelemetry monitoring ................................................................................ 4 3.1.1 Mountains and Plateau: a definition.............................................................................. 4 3.2 Resource Selection Function Models.................................................................................... 4 3.2.1 Mapping Procedures and Covariates ............................................................................. 5 3.2.2 Habitat Covariates.......................................................................................................... 5 3.2.3 Forest Age Classes ......................................................................................................... 6 3.2.4 Hillshade ........................................................................................................................ 6 3.2.5 Greenness....................................................................................................................... 6 3.2.6 Distance to the Nearest Road and Road Type ............................................................... 6 3.2.7 Data Analysis ................................................................................................................. 6 3.2.8 Model Selection............................................................................................................. 7 3.3 Resource Selection Ratios..................................................................................................... 7 3.3.1 Limited Entry Hunt and Problem Wildlife Data............................................................ 8 4.0 RESULTS ................................................................................................................................. 8 4.1 Habitat Model Results........................................................................................................... 9 4.1.1 Model 1. The Mountains ................................................................................................ 9 4.1.2 Model 2. The Plateau................................................................................................... 11 4.1.3 Comparing the Mountain and Plateau Models............................................................. 13 4.2 Resource Selection Ratios................................................................................................... 14 4.2.1 Age Classes.................................................................................................................. 14 4.2.2 Distance to the nearest road ......................................................................................... 15 4.2.3 Road Type.................................................................................................................... 17 4.2.4 Biogeoclimatic Zone .................................................................................................... 18 4.2.3 Limited Entry Hunt and Problem Wildlife Kills.......................................................... 19 5.0 DISCUSSION......................................................................................................................... 21 6.0 ACKNOWLEDGEMENTS.................................................................................................... 22 7.0 LITERATURE CITED........................................................................................................... 23 Ciarniello, Boyce & Beyer iii Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests List of Figures Figure 1. Map of the Parsnip Grizzly Bear Project Study Area Boundary. All analysis refers to the study area ................................................................................................................3 Figure 2. Map of the Relative Probability of Use Across the Mountains. Darker areas represent an increased probability of use (i.e., higher RSF values) ................................................ 11 Figure 3. Map of the Re lative Probability of Use Across the Plateau. Darker areas represent an increased probability of use (i.e., higher RSF values) .....................................................13 Figure 4. Ratio of Used Versus Available Landscape Age Classes by Mountain and Plateau Grizzly Bears, 1998 to 2001 ............................................................................................15 Figure 5. Grizzly Bear Locations Versus Distance to the Nearest Road for both Landscapes, 1998 to 2001.....................................................................................................................15 Figure 6. Grizzly Bear Locations Versus Distance to Nearest Road for the Mountainous Landscape, 1998 to 2001 .................................................................................................16 Figure 7. Grizzly Bear Locations Versus Distance to Nearest Road for the Plateau Landscape, 1998 to 2001.....................................................................................................................16 Figure 8. Grizzly Bear Locations Versus Type of Road for the Mountainous Landscape, 1998 to 2001..............................................................................................................................17 Figure 9. Grizzly Bear Locations Versus Type of Road for the Plateau Landscape, 1998 to 2001 ................................................................................................................................18 Figure 10. Grizzly Bear Locations of Biogeoclimatic Zones in Mountainous and Plateau Landscapes, 1998 to 2001................................................................................................ 19 Figure 11. Grizzly Bear Kills in Relationship to Distance to the Closest Road in Mountainous and Plateau Landscapes, 1998 to 2001 ............................................................................20 Figure 12. Grizzly Bear Kills in Relationship to Distance to the Closest Road in Mountainous and Plateau Landscapes, 1998 to 2001 ............................................................................20 List of Tables Table 1. Resource selection function coefficients, standard errors, and P-values based on Wald statistics for the mountain portion of the Parsnip River study area ....................................10 Table 2. Resource selection function coefficients, standard errors, and P-values based on Wald statistics for the plateau portion of the Parsnip River study area........................................12 Ciarniello, Boyce & Beyer iv Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests 1.0 INTRODUCTION The management of grizzly bears (Ursus arctos) and their habitat is a high profile conservation issue in British Columbia. Because grizzly bears are sensitive to human activity, there is considerable concern about the rapidly expanding development of forest and mineral resources in western Canada. Intense public concern regarding B.C.’s grizzly management practices occurs at the international, national, provincial and local level. In 1991 the Committee on the Status of Endangered Wildlife in Canada listed the grizzly bear as vulnerable. In B.C., grizzly bears are a blue-listed species (vulnerable). The B.C. Forest Practices Code requires that the needs of red and blue listed species be addressed during forest management activities. Forest companies that have, or are trying to obtain 3rd party environmental certification for their products must implement acceptable practices to protect threatened and endangered species within their operating area. Developing an understanding of the nature of habitat loss and fragmentation will be crucial to the development of sound management practices for grizzly bears. Habitats for grizzly bears can be modeled statistically using resource selection functions (RSF; Manly et al. 1993) facilitated by applications of geographical information systems (GIS; Mace et al. 1996, 2000). RSF offers a statistically rigorous way to quantify bear use of the landscape through a use versus availability analysis using multiple logistic regression. In this paper we will examine grizzly bear use of the study area by developing Resource Selection Functions for two adjacent landscapes, one mountainous and one plateau. Further we characterize habitat use by calculating resource selection ratios, which are the proportion used o ver the proportion available. The objectives of this report were to predict areas with higher probabilities of grizzly bear occurrences as well as to examine whether or not these results were consistent with our knowledge of grizzly bear use of the study area. 2.0 STUDY AREA The study area was approximately 17,500-km2 centred along the Parsnip River, British Columbia, north by northeast of Prince George, and encompassed the Parsnip River and its tributaries including the Missinka, Hominka, Table, Anzac and Chuchinka river drainages, but also extended past the Crooked River drainage in the west, and the Wolverine River drainage in the east (Figure 1). The core study area was within the Arctic watershed. To the south, the boundary extended past the Salmo n River, which was known to support some small salmon runs, however, salmon was not known to be a food source and study bears lacked this high-quality, concentrated, and predictable food supply that is important to bears along Pacific watersheds (Hilderbrand et al. 1999). The study area depicted in Figure 1 was derived by manually connecting the outside locations of radiocollared bears gathered during 1998-2001, excluding outliers in the western portion (i.e, bears who went to Vanderhoof district or Fort St. James). The rationale for limiting our western boundary was that we did not have a representative sample of bears using this region. Consequently, this would affect the results of a use versus availability analysis. Two distinct topographical areas were represented: the plateau which contains rolling hills and flat valleys, and the west and east slopes of the Hart Ranges of the Rocky Mountains with steepsided bowls, avalanche chutes and upper elevation valleys. The plateau was within the Nechako Lowland and the McGregor Plateau Ecosections of the Fraser Basin Ecoregion. The mountainous habitat was primarily within the Hart Ranges and the Hart Foothills Ecosections of Ciarniello, Boyce & Beyer 1 Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests the Central Canadian Rocky Mountains Ecoregion. Three biogeoclimatic zones were contained within the study area. The sub-boreal spruce zone (SBS) is the montane zone and occurred primarily in the plateau and some lower-elevation areas in the mountains (e.g., along major rivers). Plateau subzones included moist cool, wet cool, dry warm, and dry cool. Mountainous subzones included very wet cool and very cool. The Engelmann spruce – subalpine fir zone (ESSF) occurred above the sub-boreal spruce zone and predominated in the mountainous portion of the study area. The subzone on the west side of the Rocky Mountains was wet and cool, while the eastern slopes were moist very cold. The alpine-tundra zone (AT) occurred in high-elevation habitats beginning at approximately 1,400 m. The AT zone typically consisted of small shrubs or krummholtz, heath communities, barren rock, or alpine snow and ice. Less than 1 percent of the study area was rock and ice at elevations over 2,400 m. Elevations ranged from 750 m in the plateau to 2,500 m in the mountains. Lower elevation valley bottoms of the Missinka, Hominka, Table, and Anzac Rivers drain from the mountains into the plateau and were dominated by a mix of white spruce and subalpine fir, with the proportion of subalpine fir becoming progressively greater with increasing elevation. High elevation mountain habitats were classified as subalpine parkland, which as elevation increased become subalpine fir krumholtz tree formations. Subalpine grassland slopes and avalanche chutes were comprised of glacier lily (Erythronium grandiflorum), Indian helabore (Veratrum viride), and arrow- leaved groundsel (Senecio triangularis). The highest elevation alpine areas were alpine tundra communities, barren rock, or ice/snow. Large burns within the mountains had abundant huckleberries (Vaccinium membranaceum), blueberries (Vaccinium myrtilloides), and Canadian buffalo-berry (Shepherdia canadensis). The mountainous portion of the study area occurred in a bottleneck of the Rocky Mountains (Hart Ranges) and may be important in providing connectivity between the southern and northern Rocky Mountains. White spruce (Picea glauca) forests existed mainly in the wetter portion of the plateau, while lodgepole pine (Pinus contorta) occurred primarily in the dryer portions. Most of the plateau habitats were a mix of white spruce and pine or spruce and subalpine fir (Abies lasiocarpa). Although rare, small pockets of subalpine fir forests also existed on the plateau. Black spruce bogs (Picea mariana ) occurred in lower elevation wet areas. Mixed wood forests were lar gely the result of older cutblocks and consisted of regenerations of balsam poplar (Populus balsamifera), trembling aspen (Populus tremuloides), and paper birch ( Betula papyrifera). Interior Douglas fir (Pseudotsuga menziesii spp.), western hemlock (Tsuga heterophylla), and larch (Larix spp.) also occurred in small portions on the plateau and lower elevation mountain valley bottoms. Forestry was the predominant industry in the study area and the plateau contained the majority of forestry activities. Howe ver, logging activities were expanding up the 4 main river valleys (Missinka, Hominka, Table and Anzac Rivers) leading into mountainous areas. The majority of logging on the plateau had taken place within the last 45 years, resulting in a mosaic of forest habitats in various successional stages. The mountainous area had experienced much less industrial development, although all major watersheds had logging roads along the valley bottom and varying proportions of previous harvesting at lower elevations. However, upper elevations and the back ends of most watersheds were undeveloped wilderness. Ciarniello, Boyce & Beyer 2 Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests Other potential disturbances to grizzly bears included the town of Bear Lake, Highway 97, a railway line through the mountains for coal extraction, two sawmills, two logging camps, and various consumptive and non-consumptive recreational activities, such as hunting, fishing, snowmobiling and hiking. With the exception of two small provincial parks, there were no protected areas within the study area. The location of the Parsnip Grizzly Bear Project provided a number of opportunities to better understand grizzly bear movements and habitat requirements: 1) the study area ranged from wilderness mountain habitat to plateau habitat that had extensive road access and for est harvesting activities. Prior to this study, little was known about grizzly bear habitat use on the sub-boreal plateau; 2) the majority of the area was in the Arctic watershed so bears did not have access to spawning salmon; and 3) the area occurred in a bottleneck of the Rocky Mountains (Hart Ranges) and may be important in providing connectivity between the southern and northern Rocky Mountains. Figure 1. Map of the Parsnip Grizzly Bear Project Study Area Boundary. All analysis refers to the study area. Study Area Ciarniello, Boyce & Beyer 3 Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests 3.0 METHODS 3.1 Capture and radiotelemetry monitoring We captured 49 grizzly bears and fitted them with VHF radiocollars between August 1997 and October 2001 using a combination of aerial darting, leg snares, and culvert traps. Study bears were monitored by fixed-wing aircraft twice per week or as often as weather permitted. Once the bear was located a Universal Transverse Mercator (UTM) coordinates (x, y grid system) was taken with a hand-held 12 Channel Garmin GPS unit. If a position was taken in latitude/longitude it was converted to UTM (NAD 83) using the Geographic Calculator (Blue Marble Geographics). A biologist onboard the aircraft recorded the bear number, collar frequency, date, time, habitat type and a location confidence code on a telemetry data form. Confidence codes ranged from 1 (absolute certainty), to 2 (difficult to pinpoint due to terrain or the bear moving but generally certain), to 3 (heard but unable to pinpoint). Only confidence codes of 1 and 2 were used in this analysis. 3.1.1 Mountains and Plateau: a definition The Parsnip River provided a natural divide between the plateau landscape and the mountainous landscape. We digitized a line down the middle of the River based on the Landsat TM imagery using Geographic Information System (GIS; Arc View). Bears that had > 50% of their locations on the east side of the Parsnip River were classified as mountain bears, while bears that resided primarily on the west side of the River were referred to as plateau bears. Likewise, random points that fell west of the Parsnip River were identified as ‘mountain’ while those that fell on the east side of the river were identified as ‘plateau’. Thus, a bear that traveled between the two landscapes would have some plateau and some mo untain locations. 3.2 Resource Selection Function Models Resource Selection Functions reflecting the relative probability of use of a habitat type were estimated using logistic regression. We employed a Design 2 (Manly et al. 1993), third-order selection (Johnson 1983) study design at the population level because the entire study boundary extent was likely occupied by grizzly bears. With this design, data from individual animals were pooled and use and availability was censused for the entire study area. Bear locations gathered during 1998, 1999, 2000, and 2001 along with over 33,800 randomly generated computer points were overlaid on map images using a Geographic Information System (GIS; Arc Info). Only locations for which we were confident of the position of the animal were used in analysis. Repeat locations, for example multiple den-site locations, were removed from analysis. In addition, if a family group was collared (e.g., GF35, GM36 and GF37) only the locations of the mother were included. However, if an offspring split from its mother for a time (e.g., GM36 in 2001) its locations were considered independent until it returned to its mother, when locations were again removed from analysis. Ciarniello, Boyce & Beyer 4 Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests 3.2.1 Mapping Procedures and Covariates Digital elevation maps were built from terrain resources inventory map (TRIM; British Columbia Ministry of Land, Air and Water Protection) images and were used to obtain elevation, slope, and aspect data for bear use and random locations. Four Landsat 5 TM images were obtained from Spatial Mapping (on behalf of Canadian Forest Products Ltd.). These four images were all collected within one month (mid- August to mid-September 1999) and were mosaiced together to form a continuous image of the study area. Erdas Imagine was used for all image processing. Forest Cover Maps (FCM) and road networks were obtained from the BC Ministry of Forests, Canfor East, and Canfor West. These map layers were used to obtain the habitat type, stand age, distance to roads, and type of road information. Vegetation Resources Inventory data was obtained from Canfor West and applied to areas where it existed. 3.2.2 Habitat Covariates The availability of landcover types for each bear was characterized from forest cover maps. These maps provided the following primary tree species classifications: AC AT BA BL DH DR E EP ES FD HW PA PL SB SW LA LT LW Cottonwood Aspen Subalpine fir Balsam fir Douglas hemlock Red alder Birch species general Birch Engelmann Spruce Douglas fir Western Hemlock White bark pine Pine Black spruce Spruce white Alpine larch Tamarack Western Larch To minimize the number of landcover classes tree species were combined into 7 primary landcover categories: ACATEP (cottonwood, aspen, and common paper birch), BBABL (subalpine and balsam fir), FDHWL (Douglas fir, western hemlock, larch species), SSW (spruce species and white spruce), SB (black spruce), PL (pine species), NPF (non-forested mountain) and NFP (non-forested plateau). Non-forested plateau included those habitats of noncommercial value, such as non-commercial brush, non-productive brush, and non-productive forest. Thus, the majority of non-forested plateau habitats were those related to riparian areas. Non-forested mountain habitat also included those habitats of non-commercial value. However, in the mountains these landcover types consisted largely of alpine or meadow habitats as described by the non-forest or non-productive descriptor columns provided in the database. Because the landcover types added up to 100% we withheld white spruce as reference habitat (i.e. coefficients for those variables were not included in the models). We did this because white spruce occured in abundance in both the mountain and plateau landscapes. Ciarniello, Boyce & Beyer 5 Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests 3.2.3 Forest Age Classes Forest cover maps provide a projected age for all commercial forest types. From these projected ages forests were classified into the following categories: young (0 to 45 years and all not sufficiently restocked areas), mature (46 to 99 years), old (>100 years). However, habitats of non-commercial value do not contain age information. Therefore, if an age class was not assigned we went to the Non-Productive Descriptors and Non-Commercial Descriptors to gain information on these landcove r types. From these, we added the age classes of alpine for all locations with a descriptor of A (alpine) or AF (alpine forest). Finally, an ‘other’ category was used to classify the remaining locations without ages. Other included habitats of noncommercial brush, non-productive brush, swamps, and meadows. Mature forest was used as a reference category to which all other categories were compared because it commonly occurred in both the mountain and plateau landscapes. Therefore, a coefficient was not obtained for the mature age class. 3.2.4 Hillshade Aspect and slope were used to calculate hillshade values. Aspect was set at 225° while degree slope was set at 45° . Southwest slopes receive high hillshade values while northeast slopes correspond to low hillshade values. In the transformation, negative coefficients indicate selection for cooler, northeast aspects, while positive coefficients reflect selection for warmer southwest aspects. 3.2.5 Greenness A tasseled-cap transformation of Landsat Thematic Mapper (TM) satellite images using Erdas Imagine was used to calculate greenness. We used images from mid-August to mid-September with a 30 x 30 m pixel size. The higher the greenness scores the more green vegetation present. 3.2.6 Distance to the Nearest Road and Road Type Road network data from FCM, TRIM, Canfor East and Canfor West was amalgamated and used to query both distance to the nearest road and the type of road. For each bear and random location the straight- line distance to the nearest road was calculated in GIS. The type of road was categorized into 4 classes: highway, primary trunk logging roads, secondary logging road, and deactivated. The highway category referred to Highway 97. Primary trunk roads included any main stem road used to transport logs to the point of the Highway. An example of a primary trunk road would be the Anzac 2000 road or the 100 road. Secondary logging roads stem off of primary trunk roads and were used to access the cutblock. Decommissioned roads were at one time secondary logging roads that have not had a drainage ditch or some other type of obstruction (e.g., piling up stumps on road) put in place by the logging company that made the site less accessible. 3.2.7 Data Analysis Prior to model construction a correlation matrix was examined for all model variables. Because all correlations were less than 0.7 we concluded that colinearity was not a serious concern (Sokal Ciarniello, Boyce & Beyer 6 Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests and Rohlf 2000). The following log-linear equation was assumed to characterize the influence of covariates on relative use, w(x): w(x) = exp(β 1x1 + β2x2 + β3x3…βpx p ) (1) where β i are selection coefficients for each covariate, x i, for i = 1,2,. . .p, estimated using logistic regression. Bear locations were the used sites, 1, while random sites were assigned 0. Negative β coefficients imply avoidance, while positive coefficients imply selection. We constructed one RSF model for the entire 2001 study area. Next, to examine the relative contribution of the habitats across the study area, we constructed separate RSF models for mountain bears and for plateau bears. Models were estimated using SATISTICA (StatSoft 1999). 3.2.8 Model Selection Initially, model selection was based on Akaike Information Criteria (AIC; Burnham and Anderson 1998). However, to examine whether the large home range size and lower density of plateau bears was a function of the different landcover types, we wanted the same coefficients in both the plateau and mountain models. Therefore, to make the models comparable we did not necessarily choose the individual model with the lowest AIC score but rather the best models in which each variable occurred within both landscapes. Model estimates for the mountains and the plateau were interfaced with GIS to create maps of relative probability of grizzly bear use across the study area. 3.3 Resource Selection Ratios We also provided Resource Selection Ratios (Manly et al. 1993), which is the ratio of the proportion used to the proportion available. In this population- level design (i.e., all animals pooled), use for all animals (wi) is characterized by the following form: w( x ) = (Ui / U + ) /( Ai / A+ ) (2) Where (Ui / U + ) refers to the proportion used of bear use locations, while ( Ai / A+ ) refers to the proportion available for randomly generated locations. If use is proportional to availability (i.e., no selection) then the number is approximately 1. If use is greater than availability (i.e., numbers greater than 1) there is said to be selection for the covariate, whereas use less than availability (i.e., a number less than 1) implies avoidance. There were two covariates that we wanted to examine that could not be used in the logistic regression RSF model because currently they cannot be applied back to the GIS. These were biogeoclimatic zone, subzone and variant as well as the Limited Entry Hunt data for the study area. Ciarniello, Boyce & Beyer 7 Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests 3.3.1 Limited Entry Hunt and Problem Wildlife Data The grizzly bear limited entry hunt and problem wildlife database was obtained through the Ministry of Water, Land and Air Protection for 1977 to 2001. Locations were plotted in Arc View and only locations within the study area were considered. Some animals appeared in the database twice and therefore one location was removed. All locations pertaining to the grizzly bears killed in association with the landfill closure in Mackenzie were left in the database. There were 13 grizzly bear kills that occurred in the database that were in a burn off the ‘700 road.’ This was a known guide-outfitter hunting area which was known to contain many ATV trails. However, because we could not obtain information on the trail network, we excluded these 13 locations from analysis. 4.0 RESULTS Of the 49 bears captured, 22 were plateau bears (12 female; 10 male), while 27 were mountain bears (18 female; 9 male). In 2000 and 2001, we also monitored a plateau female and plateau male that were captured as part of another study (Peace-Williston Compensation Program). From 1998 to 2001, a total of 2,307 bear locations met the criteria for analysis on 49 grizzly bears (58P: 367M in 1998; 133P: 465M in 1999; 238P: 383M in 2000; 358P: 305M in 2001). Of these 2,307 locations, 1,520 were from the mountains while 787 were from the plateau. Mountain bears have significantly smaller home ranges than plateau bears (Ciarniello et al. 2001). For all years combined, male grizzly bears that live on the plateau have 5 times the home range size of males that live in the mountains. Furthermore, the home range size of plateau males is likely underrepresented because their necks were often larger than their heads allowing them to slip their radiocollars. In addition, with the exception some subadult males, we were frequently unable to locate plateau males. Due to intense search efforts for missing bears we believe they moved extended distances outside the study area boundary. Females that live in the plateau have 6 times the home range sizes of mountain females (Ciarniello et al. unpublished). Project capture success was also higher in the mountains than the plateau. For example, despite significant effort we could not catch any female plateau bears in 1998. A DNA census conducted in the spring of 2000 yielded a density estimate of 12 bears per 1000 km2 in the plateau, and 49 bears per 1000 km2 in the mountains (Mowat et al. 2002). Compared to other DNA-based population estimates in interior British Columbia, grizzly bear density in the mountains is high (McLellan 1989; Hovey and McLellan 1996), while the density in the plateau is low (Mowat and Strobeck 2000). We examined movements of bears to evaluate barriers between the mountains and the plateau. On the plateau, both male and female bears regularly crossed Highway 97 as well as the Parsnip River. In addition, 3 male mountain bears moved between the mountains and the plateau. Furthermore, male and female bears moved between the west and east slopes of the Rocky Mountains. In November 2000, one female family group moved a straight- line distance of 40.5 km from the plateau to the mountains for denning. These bears moved back to the plateau after den emergence the next spring. These have been the only bears monitored to spend spring, summer and fall in the plateau and den in the mountains. Based on the movements of bears there Ciarniello, Boyce & Beyer 8 Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests does not appear to be significant barriers to movement within the study area and bears are free to select habitats within the mountains or the plateau (see Fretwell and Lucus 1970). 4.1 Habitat Model Results A number of differences existed between the mountain and plateau habitats. These differences occurred in available habitats, length of foraging and denning seasons, as well as consumptive and non-consumptive human use. In addition, due to the uneven sample sizes (i.e., 1,520 mountain locations versus 787 plateau locations), and diverse bear use selection patterns, we felt that generating separate models with the same covariates and then comparing the results would be a better categorization of relative bear use of the landscape than one model for the entire landscape. Figure 2 provides a map of the relative probability of use across the mountainous landscape, while Figure 3 provides the relative probability of use across the plateau. Higher probability of use by grizzly bears is represented by darker areas. The following sections will examine each of the model coefficients by their respective landscapes. 4.1.1 Model 1. The Mountains The most significant predictor of mountain bear landscape use was greenness followed by hillshade (Table 1). Therefore, mountain bears were more likely to be found on warm aspects that contain a high green herbaceous phytomass. Elevation was found to be negative, simply meaning that use was higher on the sides of mountains as opposed to the tops or ridges. Mountain bears exhibited selection for sub-alpine fir and balsam fir (BBABL) as well as nonforested mountain (NF_M) landcover types. These forest t ypes were normally mid-to-high elevation habitats that consist of krumholtz trees, slide alder, and avalanche chutes. The upper limit of the sub-alpine parkland was classified as BBABL habitat. These findings are consistent with radio-telemetry observations of habitat types recorded by the biologist. In addition, visual rates on telemetry flights for mountain bears were over double those of plateau bears, which were related to the ease of viewing the ground in open mountain habitats Mountain bears selected for alpine and young age classes in reference to mature age class. This was not surprising because mountain bears were often located in alpine landscapes. The selection for young-age landscapes within the mountains was a reflection of bears moving to burns that produced an abundance of blueberries. In addition, cutblocks that were difficult to access by ground occurred in the northeastern portion of the mountains (upper Sukunka River area). In the spring of 2001, one mountain male bear was often located within these young-age (i.e., 0-45 years) cutblocks. Mountain bears were seldom located in the low elevation valley bottoms, which consisted primarily of old age forest stands. In relationship to mature forests, mountain bears selected against old age forests. However, site visits to old-growth forest habitat in the mountains revealed well-used travel routes that were largely associated with adult male locations. Further investigation of old- growth habitat type is required to better interpret this coefficient. Mountain bears also selected against ‘other’ landcover types. This was not surprising because in the mountains the ‘other’ habitat category largely refers to rock or talus terrain. Mountain bears were located a greater distance to roads than random (Table 1). Because the coefficient for distance is positive, the greater the distance from roads the more likely there are to Ciarniello, Boyce & Beyer 9 Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests be bears. For mountain bears with roads within or adjacent to their home range a general avoidance of roads has also been noted on telemetry flights. In addition, all of the cutblocks used by mountain bears have had to be accessed by helicopter due to decommissioning of the road. During 2000 and 2001, active logging operations and new cutblocks have been recorded during aerial telemetry flights within some of the mountain bear home ranges. However, we have never recorded a mountain bear near active logging activity or on a logging road. Table 1. Resource selection function coefficients, standard errors, and P-values based on Wald statistics for the mountain portion of the Parsnip River study area. These estimates for P-values using the Wald Statistics are conservative given our study design (Boyce et al. 2002). Variables GREENNESS ELEVATION HILLSHADE DISTANCE RD ACATEP BBABL PL NF_M Categorical Age Classes AGE Alpine AGE Old AGE Other AGE Young AGE Mature ß Standard Error P-Value +.036455 -.001322 +.003188 +.000087 -.036463 +.013488 -.026964 +.019632 .001958 .000174 .000586 .000009 .008612 .001730 .006791 .002438 <.00001 <.00001 <.00001 <.00001 .000023 <.00001 .000072 <.00001 +.511529 -.404497 -1.967851 +.084712 -------- .238955 .139493 .299779 .189942 -------- .032299 .003734 <.00001 .655603 -------- Deviation Explained = 22.52%, AIC = 7729.14 Ciarniello, Boyce & Beyer 10 Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests Figure 2. Map of the Relative Probability of Use Across the Mountains. Darker areas represent an increased probability of use (i.e., higher RSF values). 4.1.2 Model 2. The Plateau Plateau bears also exhibited a selection for higher greenness scores and positive hillshade values. The selection for lower elevation areas within the plateau was due to a lack of use by bears of the two small pockets of alpine habitats that existed within the plateau. One of these areas is a small ridge west of the Williston Reservoir. This ridge was quite different than the alpine habitat within the mountains but was classified as alpine within the forest cover database. The second area was in the southeastern portion of the plateau by the divide to Arctic Lake. This area was similar to lower elevation alpine habitat found within the mountains. The absence of bear use in this area may be due to the lack of study bears rather than an avoidance of this habitat type. Remarkably, in the plateau, we were unable to detect significant positive selection for landcover types (Table 2). Rather, bears significantly avoided the deciduous, pine, and subalpine fir Ciarniello, Boyce & Beyer 11 Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests landcover types. It must be mentioned that this avoidance is relative to the withheld habitat types of white spruce, black spruce, and Douglas fir. In a previous analysis, selection for spruce dominated forests was found to be weakly positive (Ciarniello et al. unpublished), while selection for Douglas fir habitats was positive at the de n-site selection scale. Subalpine firdominated habitats occurred in small pockets on the plateau, whereas they dominate within the mountains. We have recorded a male denning in higher elevation subalpine fir habitat in the plateau. Pine-dominated habitats are typically much dryer than spruce-dominated habitats. Therefore they lack the abundance of herbaceous vegetation suitable for grazing that our habitat plots have revealed important for bears (Ciarniello et al. 2001). In 1998 and 1999, avoidance of pinehabitats was recorded on telemetry flights. However, in 2000 an abundant berry crop resulted in more bears using the dryer pine habitats during berry season. Therefore, use of pine forests was dependent upon the productivity of the berry crop. Contrary to primary landcover types, there was selection for all age classes in relation to the mature forest reference category, however, only the ‘other’ age class was significantly selected. In the plateau, ‘other’ referred primarily to riparian areas, bogs and fens for which age information was not available. Selection for wet areas adjacent to rivers and ponds was consistent with aerial telemetry findings. These riparian areas were also known to contain important forage for bears during all seasons . We have provided a resource selection ratio for age classes for both the mountains and plateau under section 4.2.1. Distance to roads is a positive beta, meaning the further from a road the more likely there are to be bears, however, this coefficient is not significant. This selection was likely a function of the density of roads within the plateau – in the plateau it may be difficult to be far from a roaded area while also meeting nutritional requirements. We were unable to apply type of road to this analysis because this information is currently being incorporated into the GIS so it may be referenced, however, this information appears to be important to more adequately assess relative bear use of the plateau (please refer to Section 4.2.3). Table 2. Resource selection function coefficients, standard errors, and conservative P-values based on Wald statistics for the plateau portion of the Parsnip River study area. Variables ß Standard Error P-Value GREENNESS ELEVATION HILLSHADE DISTANCE RD ACATEP BBABL PL NF_P Categorical Age Classes AGE Old AGE Other AGE Young AGE Mature Ciarniello, Boyce & Beyer +.01984 -.00128 +.00410 +.00003 -.00351 -.00628 -.01387 -.03952 .003779 .000310 .001963 .000031 .001706 .002070 .001634 .004599 <.00001 .000035 .036699 .265928 .039685 .002423 <.00001 <.00001 +.03650 +3.57140 +.16013 -------- .106596 .454416 .117943 -------- .732075 <.00001 .174569 -------- 12 Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests Figure 3. Map of the Relative Probability of Use Across the Plateau. Darker areas represent an increased probability of use (i.e., higher RSF values). 4.1.3 Comparing the Mountain and Plateau Models Eleven of the 15 variables measured were significant predictors of mountain bear landscape use (Table 2), while 8 of variables were significant predictors of plateau-bear use of the landscape (Table 3). Common to both RSF models was selection for higher greenness scores and southwest facing aspects. Southwest aspects are snow free earlier, and remain snow free longer, than cooler aspects, which is an important consideration for food availability in northern environments. Also common to both models was a selection for lower elevation habitats, however, for mountain bears this was a reflection of use of mid-elevation habitats, while for Ciarniello, Boyce & Beyer 13 Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests plateau bears this reflected an avoidance of the small pockets of alpine habitat that occurred in two locations within the plateau landscape. Overall, mountain bears were more affected by the covariates in the RSF models than plateau bears. The mountain model does a better job of predicting spatial patterns in the probability of use by grizzly bears than the plateau model. Furthermore, the density of bears is higher in the mountains than the plateau (Mowat et al. 2002). We examined resource selection ratios to determine the type of information that may be required to improve the plateau model. 4.2 Resource Selection Ratios Resource selection ratios are similar to Resource Selection Functions (Manly et al. 1993). Their main advantage is that they allow one to view the selection pattern in a graphical format. In addition, at this stage of analysis some information has not yet been created as a spatial explicit GIS layer and therefore cannot be used to create GIS maps. This section provides selection ratios for some of the model covariates, information not contained in the models, as well as further information on road types. A ratio of 1 indicates use in proportion to availability (no selection), whereas greater than 1 indicates selection, and less than 1 indicates avoidance. The ratio can be thought of as an odds ratio, so the higher the number the more it is selected. 4.2.1 Age Classes We included a ratio of the forest age class to explain a limitation in the Resource Selection Function models presented above (Figure 4). In ratio form, the histogram depicts the information contained within the models. For example, one can clearly see that alpine age classes were highly selected for by mountain bears, whereas plateau bears selected for ‘other’ and young age classes. There are two things to note on this histogram. The first is that the selection for young age classes now appears significant. This is because a ratio is a direct proportion, whereas the RSF models used deviance coding and referenced the age classes to mature. Therefore, although the beta value was positive, in relation to mature age classes there was not a significant selection for young habitat, however, in ratio comparison there appears to be selection for this age class. Complete avoidance was also a problem for the RSF models and can be seen when examining the urban category. Random points fell within non-forest descriptors that were labeled urban, however, we did not have any bear use points in urban. Therefore, we had to eliminate all random urban points from RSF analysis. In the ratio depicted in Figure 4, these also come out as 0 because there was no bear use. However, complete avoidance of an area is an extremely important feature of grizzly bear habitat use. Ciarniello, Boyce & Beyer 14 Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests Figure 4. Ratio of Used Versus Available Landscape Age Classes by Mountain and Plateau Grizzly Bears, 1998 to 2001. Ratio of Used Versus AvailableForest Age Classes 3.5 3 Ratio 2.5 2 Mountain Plateau 1.5 1 0.5 0 Alpine Mature Old Other Urban Young Age Class 4.2.2 Distance to the nearest road Distance to the nearest road is provided for both landscapes combined (Figure 5), for the mountains (Figure 6), and for the Plateau (Figure 7). For the study area, there was an overall avoidance of roads by grizzly bears until approximately 2.5 km. There was neither selection nor avoidance associated with roads from 2.5 to 4.5 km. Overall, grizzly bear use of the landscape increased with further distances from roads. Figure 5. Grizzly Bear Locations Versus Distance to the Nearest Road for both Landscapes, 1998 to 2001. 11.5 11 10.5 10 9.5 9 8.5 8 7.5 7 6.5 5 4.5 4 3.5 3 2.5 2 1.5 1 12 11 10 9 8 7 6 5 4 3 2 1 0 0.5 Ratio Bear Locations vs. Distance to Nearest Road (Study Area) Distance to Road (KM) Ciarniello, Boyce & Beyer 15 Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests For the mountains the selection ratio provides a depiction of avoidance of areas within 1 kilometer of roads (Figure 6). At 1.5 kilometers use began to increase and was in proportion with availability at approximately 2.5 km. At 5 kilometers from the road use was greater than availability and remained so until 11.5 km. This reveals an overall avoidance of road networks by mountain bears. Figure 6. Grizzly Bear Locations Versus Distance to Nearest Road for the Mountainous Landscape, 1998 to 2001. Grizzly Bear Locations versus Distance to Nearest Road (Mtns. Only) 8 7 Ratio 6 5 4 3 2 11.5 11 10.5 10 9.5 9 8.5 8 7.5 7 6 6.5 5.5 5 4 4.5 3.5 3 2.5 2 1 1.5 0 0.5 1 Distance to Nearest Road Figure 7. Grizzly Bear Locations Versus Distance to Nearest Road for the Plateau Landscape, 1998 to 2001. Grizzly Bear Locations versus Distance to Closest Road (Plateau Only) 6 5 Ratio 4 3 2 1 0 0.5 1 1.5 2 2.5 3 3.5 4 4.5 5 6.5 7 7.5 Distance to Road (KM) Ciarniello, Boyce & Beyer 16 Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests For the plateau use was in proportion to availability within 500 m of the road, which was likely a function of plateau bears being attracted to the new growth along roadsides during the spring foraging season. Also, plateau grizzly bears have been recorded to travel along deactivated logging roads on a number of occasions. From 1 to 5 km from the road selection fluctuated slightly above and below (avoidance) 1. At 6.5 km use became increasingly greater than availability. An interesting note is that this is very similar to the pattern observed in the mountains, where at approximately 5 km use became greater than availability and remained so up to 11 km from the road. Therefore, in either landscape the further from a road the more likely there are to be grizzly bears. 4.2.3 Road Type We also examined the types of roads that were closest to the bear’s locations (Figure 8 and Figure 9). Mountain bear locations were closer to roads that had been decommissioned while they most strongly avoided the highway. Possibly the majority of the collared sample did not actively avoid the highway but simply did not move out of the mountainous habitat. Therefore, road types other than highway were more applicable for mountain bears that resided in the interior mountain. As mentioned, all of the cutblocks we visited in the mountains to conduct onsite habitat investigations were classified as decommissioned or largely inaccessible. Figure 8. Grizzly Bear Locations Versus Type of Road for the Mountainous Landscape, 1998 to 2001. Mountain Bear Ratio of Nearest Road Type 6 5 Ratio 4 3 2 1 0 Highway Logging Road Secondary Logging Decommissioned Type of Roads Ciarniello, Boyce & Beyer 17 Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests Figure 9. Grizzly Bear Locations Versus Type of Road for the Plateau Landscape, 1998 to 2001. Plateau Bear Ratio of Nearest Road Type 1.6 1.4 1.2 Ratio 1 0.8 0.6 0.4 0.2 0 Highway Logging Road Secondary Logging Type of Road Plateau bear locations were closer to primary logging roads than the highway or secondary logging roads. However, the plateau landscape results for type of road should be viewed with caution because there were no roads listed as deactivated within the database. However, our onsite knowledge of the land base reveals that there were places on the plateau that had been deactivated making some inaccessible or ATV-only accessible. For example, on several occasions we investigated a known area of deactivated road networks by Summit Lake. In the GIS database these roads were classified either as logging roads or secondary logging roads. Therefore, for this analysis to accurately reflect the use of areas associated with various types of road, the databases must be updated and reclassified. Thus, the selection for logging road and use in proportion to secondary logging road may not be accurate. 4.2.4 Biogeoclimatic Zone We also examined the biogeoclimatic zones in which the locations occurred (Figure 10). For mountain and plateau bears there was a selection for the ESSFwk2 biogeoclimatic zone. Meidinger and Pojar (1991) considered this biogeoclimatic zone to be year round high-quality grizzly bear habitat. The ESSFwk2 zone occurred in a relatively small proportion on the plateau but bears were located within this zone greater than its availability. The majority of this use occurred along the mountain ridge west of McLeod Lake. The Alpine Tundra (AT) biogeoclimatic zone occurred in very small proportions on the plateau and there was no use recorded by bears in this zone. However, this may be due to a lack of study animals in the area where this zone occurs (i.e., by the divide of Arctic Lake). In the mountains use of the AT zone was in proportion to its availability. Meidinger and Pojar (1991) state that Ciarniello, Boyce & Beyer 18 Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests the AT alpine zone offers important habitats for grizzly bears, particularly where the soils allow easy digging. Figure 10. Grizzly Bear Locations of Biogeoclimatic Zones in Mountainous and Plateau Landscapes, 1998 to 2001. Ratio of Used Versus Available Biogeoclimatic Zones 2.5 Ratio 2 1.5 1 Mountain Plateau 0.5 SBSwk2 SBSwk1 SBSvk SBSmk2 SBSmk1 SBSmh SBSdw3 ESSFwk2 ESSFmv3 ESSFmv2 BWBSwk1 BWBSmw1 AT p 0 Biogeoclimatic Zone 4.2.3 Limited Entry Hunt and Problem Wildlife Kills The plateau model did not perform well in predicting bear use of the landscape (deviation explained was 3.5%). We believe that this may be because the majority of the covariates in the model were habitat related, whereas bear use in the plateau may be dictated by risk of mortality and avoidance of active resource extraction activities rather than driven by habitat related variables. To increase the predictive capability of the plateau model we are currently developing a risk layer for bears that will be used to reflect avoidance of areas that based on habitat variables alone may predict bear use. For example, it is possible that plateau bears were actively avoiding areas of high mortality risk due to human access. To investigate this hypothesis we are examining the Limited Entry Hunt (LEH) and Problem Wildlife (PW) database (Figure 11). Figure 11 depicts the number of grizzly bears killed as a function of the distance to the nearest road. Seventeen percent (n=54) of the bears killed within the study area were within 0-100 m of a road, 32% (n=99) within 0-200 m, 39% (n=121) within 0-300m, 43% (n=133) within 0-400 m, and 58% (180) within 0-500 m of the closest road. 73% (n=227) occurred within 1 km of a road. Hunters are only required to report their kill site to within the nearest kilometer. Thus, it is likely that the majority of these kills occurred closer to a road than 1 km. Overall, the majority of grizzly bears killed within the study area occurred closer to roads than random and the frequency decreased as the distance to the road increased (Figure 11). The majority of the kills occurring furthest from roads occurred at a fly in guide outfitting camp within the mountains. A few of Ciarniello, Boyce & Beyer 19 Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests the bears in the database had been hit by a train along the railway used to access coal from two mines within the mountains. Figure 11. Grizzly Bear Kills in Relationship to Distance to the Closest Road in Mountainous and Plateau Landscapes, 1998 to 2001. Grizzly Bears Killed 1977 - 2001 in the PGBP Study Area 250 Number of Bears Killed 200 150 100 50 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 Distance to Road (KMs) We also examined the types of roads closest to the mortality locations (Figure 12). The majority of grizzly bears killed occurred near the highway, followed by secondary logging roads, primary trunk roads, and decommissioned logging roads (Figure 12). Deactivating roads may offer an increased advantage to grizzly bear survival. Figure 12. Grizzly Bear Kills in Relationship to Distance to the Closest Road in Mountainous and Plateau Landscapes, 1998 to 2001. Grizzly Bear Mortality - Relationship to Nearest Road Type 1.2 1 Ratio 0.8 0.6 0.4 0.2 0 Highway Primary Trunk Secondary Logging Deactivated Road Type Ciarniello, Boyce & Beyer 20 Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests 5.0 DISCUSSION The location of the Parsnip Grizzly Bear Study provided a number of special opportunities to better understand grizzly bear habitat requirements. In particular, the study area ranged from wilderness mountain habitat to plateau habitat that had extensive road access and forest harvesting activities. Prior to this study, little was known about habitat use of grizzlies on the sub-boreal plateau. Similar to results of other studies, we thought that most of the bears would den in the mountains, move to lower, snow- free habitat in the plateau at den emergence and then move back to the mountains as foods became available, resulting in a population of migrating bears (McLellan and Hovey 2001). However, we found that mountain bears tended to live solely in the mountains and plateau bears on the plateau. During our 4 years of monitoring an average of 19 grizzly bears per year only 1 large adult male frequently traveled between the mountains and plateau, although he primarily lived in the mountains. One male dispersed from the mountains to the plateau and another occasionally traveled between landscapes. However, both were killed in the limited entry hunt. Only 1 female, accompanied by 2, 3- yr-old offspring, moved from the plateau to the mountains for denning. This family group returned to the plateau at den emergence. The density of bears within the two landscapes also differed markedly (Mowat et al. 2002). In a previous analysis we examined the link between bear density and landscape structure by investigating the changes in the RSF models as a result of applying the mean values of the different habitat types as well as the average density of roads from one landscape to the other (Ciarniello et al. in review). Although the probability of use by grizzly bears accounted for by the model is higher in the mountains than the plateau we could explain only 5 to 6 % of the difference in density between the 2 areas by examining the habitat variables and/or the distance to roads. Therefore, we were unable to support the hypothesis that 4- fold differences in bear density between the mountains and the plateau were attributable to differences in the respective habitats or road density. However, such a conclusion would have a high probability of a Type II error, i.e., a difference might exist that we are unable to detect for whatever reason. Because there is a four- fold difference in grizzly bear density as identified by the DNA census, and we could only account for a 5-6% difference when switching the road and habitat attributes,we concluded that we did not captured the mechanism behind the difference in population density between the 2 areas. We believe that more information on plateau bears is required to sort out habitat selection by these bears. We state this because we have over twice the number of locations in the mountains than we have in the plateau, plateau bears were more prone to slipping their collars and therefore we do not have a full year of data on a number of plateau bears, and the habitat model has trouble predicting bear use of the plateau landscape. In addition, uncovering the mechanisms of bear use on the plateau will allow us to hypothesize about future use of the mountain landscape by bears as it becomes increasingly fragmented. We present a further hypothesis that selection by plateau bears may be a combination of habitat related attributes and avoidance of high-risk areas. To adequately predict use of landscapes with a myriad of human activities an avoidance/risk layer must be developed that can be interfaced with GIS and applied back to the landscape. We are currently developing a risk layer that will be applied to RSF models and the GIS database. The analysis presented in this paper reveals that bears were selecting for habitats further from roads than was available. Therefore, a model based mainly on habitat-related Ciarniello, Boyce & Beyer 21 Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests covariates would predict the animal to be in an area, however, the risk of mortality likely outweighs the benefits of foraging close to roads. Based on RSFs characterizing female grizzly bear habitat use in Montana, Mace et al. (1999) recommended management to reduce road density. Although our results indicate a significant influence of roads, the magnitude of response is unlikely to account for the observed differences in bear density between the mountains and the plateau (Ciarniello et al. in review). Clearly the extensive road network on the plateau subjects bears to higher risk of human- induced mortality than in the mountains by providing access for human activities. However, adding layers will require more locations to have the statistical power to accurately predict relative use of the plateau landscape. Another possible explanation for our results might be that bears have been avoiding areas that are being harvested for timber. A limitation with the data obtained from forest cover maps is that the information is not updated yearly. Therefore, map layers do not reflect current manipulation of the landscape, which may have an influence on the distribution and dens ity of bears. Our next step will be to update GIS layers with timber harvesting patterns by year to account for present timber harvesting, and to build new RSF models. Mortality and current land-use data will be added to the model to explicitly consider risks in areas of concentrated human activity associated with living in the plateau. Addition of these variables should better predict relative use of the plateau landscape as well as explaining the 4-fold differences in grizzly bear densities found between the mountains and the plateau. This information will help land managers in government and forest industry develop land- use practices that are compatible with the conservation of grizzly bears and their habitat. 6.0 ACKNOWLEDGEMENTS The Parsnip Grizzly Bear Project is supported by Forest Renewal British Columbia (FRBC), Canadian Forest Products Ltd. (Canfor), and the Pas Lumber. Additional support was provided by: BC Ministry of Environment, Lands and Parks, BC Ministry of Forests, BC Conservation Foundation, Spruce City Wildlife Association, Natural Science and Engineering Research Council of Canada scholarship to L. Ciarniello, and the University of Alberta. This project report was initiated by Dale Seip, BC Ministry of Forests, and supported by the Crown Land Use Planning Enhancement Fund. We thank our funding agencies. We thank John Paczkowski for work on the project and for the1998 data. Elena Jones was the field technician and aided in some of the resource selection ratio analysis. The success of the Parsnip Project would not have been possible without the assistance of Dale Seip, Doug Heard, Glen Watts, Carla Wainwright, Ian Ross, Charles Mamo, Doug Wilson, Kerry Deschamps, Andrew De Vries, and Joe Kavanagh. We also thank Greg Altoft, Pie rre Bock, Ken Knight, Chris Norman, and Biker Mike for helicopter skills. Fixed-wing airplane pilots Larry Frey and Eric Stier kept us safe on telemetry flights. We also thank Mari Woods of the McLeod Lake Landfill Study (Peace-Williston Compensation Program) for providing data on 2 of their study animals, which have been included in our 2000 and 2001 databases. Last but certainly not least, we thank all the study bears. Ciarniello, Boyce & Beyer 22 Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests 7.0 LITERATURE CITED Burnham, K. P., and D. R. Anderson. 1998. Model selection and inference: a practical information theoretic approach. Springer-Verlag, New York, New York, USA. Ciarniello, LM, J. Paczkowski, D. Heard, I. Ross and D. Seip. 2001. Parsnip Grizzly Bear Population and Habitat Project: Progress Report for 2000. Unpub lished report for Canadian Forest Products Ltd. And BC Ministry of Forests, Prince George, BC, Canada. 55pp. Ciarniello, L.M., M.S. Boyce, D.R. Seip, D.C. Heard. In review. Grizzly bear habitat selection in mountain and plateau landscapes along the Parsnip River, BC. Ursus. Committee on the Status of Endangered Wildlife in Canada (COSEWIC). 1996. Canadian species at risk. Queen’s Printer, Ottawa, Ontario, Canada. Fretwell, S. D., and H. L. Lucus, Jr. 1970. On territorial behavior and other factors influencing habitat distribution in birds: I. Theoretical development. Acta Biotheoretica 19:16-36. Hilderbrand, G.V., Schwartz, C.C., Robbins, C.T., Jacoby, M.E., Hanley, T.A., Arthur, S.M. and C. Servheen. 1999. The importance of meat, particularly salmon, to body size, population productivity, and conservation of North American brown bears. Can. J. Zool. 77:132-138. Johnson, D. H. 1980. The comparison of usage and availability measurements for evaluating resource preference. Ecology 61:65-71. Mace, R. D., J. S Waller, T. L. Manley, L. J. Lyon, and H. Zuuring. 1996. Relationships among grizzly bears, roads and habitat in the Swan Mountains, Montana. Journal of Applied Ecology 33:1295-1404. Mace, R. D., J. S. Waller, T. L. Manley, K. Ake, and W. T. Wit tinger. 1999. Landscape evaluation of grizzly bear habitat in western Montana. Conservation Biology 13:367-377. Manly, B. F. J., L. L. McDonald, and D. L. Thomas. 1993. Resource selection by animals. Chapman and Hall, London, United Kingdom. McLellan, B. N. 1989. Dynamics of a grizzly bear population during a period of industrial resource development, I. Density and age-sex composition. Canadian Journal of Zoology 67:1856-1860. McLellan, B. N., and F. W. Hovey. 2001. Habitats selected by grizzly bears in multiple use landscapes. Journal of Wildlife Management 65:92-99. Meidinger, D., and J. Pojar. 1991. Ecosystems of British Columbia. Special Report Series 6. Ministry of Forests, Victoria, British Columbia, Canada. Mowat, G., K. G. Poole, D. R. Seip, D. C. Heard, R. Smith, and D. W. Paetkau. 2002. Grizzly and Black Bear Densities in Interior British Columbia. Forest Renewal BC Workplan No. OPM 01103, Activity No. 720869 and Common Land Information Base Project No. CLB1029. Mowat, G., and C. Strobeck. 2000. Estimating population size of grizzly bears using hair capture, DNA profiling, and mark-recapture analysis. Journal of Wildlife Management 64:183-193. Ciarniello, Boyce & Beyer 23 Habitat Selection by Mountain and Plateau Grizzly Bears Ministry of Forests Nielsen, S. E., and M.S. Boyce, G.B. Stenhouse, R.H.M. Munro. In press. Modeling grizzly bear habitats in the Yellowhead ecosystem of Alberta: taking autocorrelation seriously. Ursus. Sokal, R. R., and F. J. Rohlf. 2000. Biometry, 3rd edition. W.H. Freeman and Company, New York, New York, USA. Ciarniello, Boyce & Beyer 24
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