SPACE USE PATTERNS OF MOOSE (ALCES ALCES) IN RELATION TO FOREST COVER IN SOUTHEASTERN ONTARIO, CANADA

A thesis submitted to the Committee on Graduate Studies in partial fulfillment of the requirements for the degree of Master of Science in the facult y of Arts and Science

TRENT UNIVERSITY Peterborough, Ontario, Ca nada Copyright by Karen Hussey, 2009 Environmental and Life Sciences Graduate Program

January 2010

Space use patterns of moose (Alces alces) in relation to forest cover in southeastern Ontario, Canada.

ABSTRACT I investigated habitat use relative to distance to cover of moose (Alces alces) in Algonquin Provincial Park, Ontario, Canada to validate assumptions in the Ontario Ministry of Natural Resources (OMNR) habitat suitability model for moose. I compared distances to cover of 21 GPS-collared female moose to random points within seasonal home ranges to determine selection and calculate several distance -to-cover parameters. At the population level moose showed no selection for proximity to cover in the summer and marginal selection in the winter. When considering four habitat types, moose generally showed no selection for proximity to cover in either season while in any of the habitat groups. Considerable variation in selection for cover existed within and among individuals, with many individuals selecting proximity to cover in one year and avoiding or showing no selection in other years. I identified several areas where the existing habitat suitability model could be improved. I recommend further testing of an adapted model that 1) redefines stand age according to the most recent harvest date instead of the age of the oldest trees, 2) adopts a broader list of cover types defined as cover in the growing season, and 3) increases the distance moose are assumed to travel through open areas in the dormant season from 200 to 300 meters.

Keywords: Alces alces, Algonquin Provincial Park, cover, distance, habitat, habitat suitability model, home range, moose, Ontario, ungulate

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ACKNOWLEDGMENTS I am indebted to many who have provided me with assistance and support throughout the project. First of all I would like to thank my supervisors Brent Patterson of the Ontario Ministry of Natural Resources (OMNR) and Dennis Murray of Trent University for the great opportunity to be a part of the moose project and for their financial and academic contributions. Id also like to thank my third committee member, Bruce Pond, of the OMNR for his thoughtful input and supportive demeanor. Funders of the project included the Natural Sciences and Engineering Research Council of Canada (NSERC), Ontario Federation of Anglers and Hunters (OFAH), Canada Foundation for Innovation (CFI), OMNR Wildlife Research & Development Section, and Ontario Parks (Algonquin Provincial Park). Id especially like to thank the Algonquin Forestry Authority (AFA) for providing the critical funding allowing me to finish my degree. I am deeply indebted to several office extras who generously shared their time and expertise during numerous and impromptu occasions: Raul Ponce-Hernandez, Joe Nocera, Jeff Bowman, and especially Kevin Middel and Colin Garroway, for without their technical help, I would still be processing my sea of data. Phil Elkie of OMNR provided assistance with technical aspects of the model and Joe Yaraskavitch, Keith Fletcher, and Gord Cumming of AFA answered all the forestry-related questions I could throw at them. Id especially like to thank Linda Cardwell who is a pillar of support for all of us graduate students and tirelessly holds the Environmental and Life Sciences Department together. There are many who have assisted with moose captures and/or field work: Andy Silver, Stacey Lowe, Ken Mills, John Benson, Kevin Downing, Kiira Siitari, Josh Sayers,

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Tom Habib, Mike Ward, and especially Karen Loveless (aka Kwolf) who ran the field component until I arrived and was a wonderful field mentor and friend from the beginning. Lastly, Id like to thank my friends and family for all their support and especially my husband, Travis Hussey, who sacrificed so much to join me on this journey. I cant imagine completing this endeavor without all your love, support, patience, and tasty dinners. Thank you, Travis, from the bottom of my heart.

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TABLE OF CONTENTS Page ABSTRACT. ..ii ACKNOWLEDGEMENTS... iii TABLE OF CONTENTS...v LIST OF TABLES... .. vi LIST OF FIGURES..... .. viii CHAPTER 1: INTRODUCTION AND LITERATURE REVIEW. 1 CHAPTER 2: METHODS.. .. 5 2.1 Study area.5 2.2 Field methods....... 7 2.3 Habitat data and validation...... 8 2.4 Analyses... 11 2.4.1 Selection of cover..... 11 Population level selection . .... 11 Home range level selection.... 14 Year effect..15 2.4.2 Distance parameters...... 15 2.4.3 Model validation... 18 Model description.. 18 Validation method 1...20 Validation method 2. ..21 Validation method 3...22 CHAPTER 3: RESULTS...... 22 3.1 Selection of cover year effect... 23 3.2 Selection of cover home range level.... 24 3.3 Population level seasonal results (selection & distance parameters).... 26 3.4 Population level habitat results (selection & distance parameters).. 27 3.5 Model validation.. 38 3.5.1 Validation method 1......38 3.5.2 Validation method 2..... 41 3.5.3 Validation method 3.. ... 45 CHAPTER 4: DISCUSSION 46 4.1 Selection of cover............ 46 4.1.1 Prediction 1... 46 4.1.2 Prediction 2... 48 4.1.3 Prediction 3... 49 4.2 Model validation.......... 49 4.3 Model recommendations. 54 4.4 Implications of variability in habitat studies. ...... 55 LITERATURE CITED.. 59 APPENDIX A: delineation of habitat groups, seral stages, and cover values...........69 APPENDIX B: selection ratio normality plots.......... 72 v

LIST OF TABLES

Page Table 2.1: The Ontario Ministry of Natural Resources habitat suitability models description of cover categories (Naylor et al. 1999) and my corresponding habitat groups.. 20 Distance-to-cover assumptions in the Ontario Ministry of Natural Resources habitat suitability model for moose and expected 95 percentile distances from moose locations to distance categories ........ 20 Mean temperature and snow depths ( SD) for the analysis periods taken from Algonquin Park East Gate weather station (Environment Canada 2008). Historic normals are averaged from 1971-2000. Year effect analyses were performed using growing season and midwinter season. Asterisks indicate snow depths known to influence moose movement........24

Table 2.2:

Table 3.1:

Table 3.2a: Results of habitat group level mature conifer cover selection ANOVAs. Asterisks indicate marginal significance. ........ 29 Table 3.2b: Results of habitat group level selection of the lesser cover categories associated with presapling and sapling in the dormant season. Asterisks indicate marginal significance. Sapling plus refers to the combination of any sapling or mature forest. ........ 29 Table 3.3a: Moose median observed and expected distances to cover plus 95% confidence intervals (in meters) by season and habitat group for all home ranges, those where cover was selected, and those where cover was avoided. N refers to the number of home ranges in each group. %S = % home ranges showing selection, %NS = % showing no selection, and %A = % showing avoidance . . 30 Table 3.3b: Moose observed and expected ninety-five percentile distances plus 95% confidence intervals (in meters) to cover by season and habitat for all home ranges, those where cover was selected, and those where cover was avoided. N refers to the number of home ranges in each group. %S = % home ranges showing selection, %NS = % showing no selection, and %A = % showing avoidance....................................... 31

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Table 3.4a: Moose observed and expected median distances plus 95% confidence intervals (in meters) to lesser cover categories associated with presapling and sapling in the dormant season for all home ranges, those where cover was selected, and those where cover was avoided. N refers to the number of home ranges in each group. Sapling plus refers to the combination of any sapling or mature forest. . 32 Table 3.4b: Moose observed and expected ninety-five percentile distances plus 95% confidence intervals (in meters) to lesser cover categories associated with presapling and sapling in the dormant season for all home ranges, those where cover was selected, and those where cover was avoided. N refers to the number of home ranges in each group. Sapling plus refers to the combination of any sapling or mature forest....... 32 Table 3.5: Breakdown of distance assumption violations for the dormant season. The mature conifer assumption refers to points greater than 1600 meters from cover. Mature forest assumption refers to points greater than 400 meters from all mature habitat groups (hardwood, mixed, and cover). Sapling plus assumption refers to points greater than 200 meters from saplings and all mature habitat groups combined. N refers to the number of animals associated with each result. * The sum of distance assumption violations 1 - 3 may exceed 100% in a given habitat group because some points violated more than one assumption.. 45

APPENDIX A Table 1: Forest Resource Inventory Landscape Guide Forest Units (LGFU) included in the three forest types used in this study... 69 Delineation of seral stages for my study groups in relation to the Forest Resource Inventory Landscape Guide Forest Units (LGFUs) that make up each study group. LGFU descriptions are located in Appendix A Table 1 70 Delineation of cover values for my mature study groups in relation to the Forest Resource Inventory Landscape Guide Forest Units (LGFUs) that make up each study group. LGFU descriptions are located in Appendix A, Table 1. Cover values apply to forest stands classified in the FRI as immatur e, mature, or old... 71

Table 2:

Table 3:

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LIST OF FIGURES

Page Figure 2.1: Study area- Algonquin Provincial Park in southeastern Ontario, Canada ..... 7

Figure 2.2a: Observed and expected distributions of distance -to-cover from hardwood locations for Moose7 in the dormant season of 2006-2007. Preference zone edge can be estimated as the general area where the expected values begin to outnumber the observed values...... 17 Figure 2.2b: Linear regression for Moose7 in the dormant season of 2006-2007 when in hardwood stands. The preference zone is the area within 394 meters of cover. A Y value of 0.301 is equal to the selection ratio of one .. 18 Figure 3.1a: Variation of selection behavior at the home range level for mature conifer cover. S = selection, A = avoidance, and NS = no selection. N refers to the number of home ranges in each category. 25 Figure 3.1b: Variation of selection behaviour at the home range level for the lesser cover categories associated with presapling and sapling forest in the dormant season (mature forest and sapling plus). S = selection, A = avoidance, and NS = no selection. N refers to the number of home ranges in each category. ..... 26 Figure 3.2a: Moose observed (o) and expected (e) average median distances to cover ( 95% CI) of home ranges where selection of cover occurred for each season and habitat group. %S is the percent of all home ranges where cover was selected and n is the number of home ranges where cover was selected 33 Figure 3.2b: Moose observed (o) and expected (e) average 95 percentile distances to cover ( 95% CI) of home ranges where selection of cover occurred for each season and habitat group. %S is the percent of all home ranges where cover was selected and n is the number of home ranges where cover was selected ... 34 Figure 3.2c: Moose average preference zones for cover ( 95% CI) for each season and habitat group. %S is the percent of all home ranges where cover was selected and n is the number of home ranges where cover was selected... .... 35

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Figure 3.3a: Moose observed (o) and expected (e) average median distances ( 95% CI) to lesser cover categories associated with presapling and sapling forest in the dormant season for home ranges where selection of cover occurred. Sapling plus refers to the combination of any sapling or mature forest. %S is the percent of all home ranges where cover was selected and n is the number of home ranges where cover was selected.... 36 Figure 3.3b: Moose observed (o) and expected (e) average 95 percentile distances ( 95% CI) to lesser cover categories associated with presapling and sapling forest in the dormant season for home ranges where selection of cover occurred. Sapling plus refers to the combination of any sapling or mature forest. %S is the percent of all home ranges where cover was selected and n is the number of home ranges where cover was selected .... 37 Figure 3.3c: Moose average preference zones ( 95% CI) in relation to lesser cover categories associated with presapling and sapling in the dormant season. Sapling plus refers to the combination of any sapling or mature forest. %S is the percent of all home ranges where cover was selected and n is the number of home ranges where cover was selected ... . 38 Figure 3.4a: Map of the growing season range calculated in OMNRs moose habitat suitability model and the seasonal moose locations falling in and outside of the available habitat in the southwestern portion of Algonquin Provincial Park, Ontario, Canada. Distance assumption violations occurred in 49% of moose locations.......... 40 Figure 3.4b: Map of the dormant season range calculated in OMNRs moose habitat suitability model and the seasonal moose locations falling in and outside of the available habitat in the southwestern portion of Algonquin Provincial Park, Ontario, Canada. Distance assumption violations occurred in 0.1% of moose locations. 41 Figure 3.5a: Map of the growing season range created from my adapted model and the seasonal moose locations falling in and outside of the available habitat in the southwestern portion of Algonquin Provincial Park, Ontario, Canada. Distance assumption violations occurred in 3% of moose locations... 42

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Figure 3.5b: Map of the dormant season range created from my adapted model and the seasonal moose locations falling in and outside of the available habitat in the southwestern portion of Algonquin Provincial Park, Ontario, Canada. Distance assumption violations occurred in 12% of moose locations... 43 APPENDIX B Figure 1: Distance to cover selection ratio normality plots for moose in Algonquin Park by year-season for year affect analysis. 72 Distance to cover selection ratio normality plots for moose in Algonquin Park by season.. 73 Distance to cover selection ratio normality plots for moose in Algonquin Park by habitat group-season 74 Distance to cover selection ratio normality plots for moose in Algonquin Park for lesser cover categories 75

Figure 2:

Figure 3:

Figure 4:

CHAPTER 1: INTRODUCTION & LITERATURE REVIEW Because cover provides protection from harsh environmental conditions and concealment from predators, it is a critical habitat component for species of many taxa, including insects (Sih 1990) , fish (Gilliam and Fraser 1987), amphibians (Holomuzki 1986) , birds (Radford et al. 2005), and mammals ( Holmes 1984, Kotler and Blaustein 1995). How ever, these protective areas are often deficient in other high quality resources (typically food) needed to adequately sustain individuals and/or facilitate reproduction (Lima and Dill 1990, Mysterud and stbye 1995, Moody et al. 1996, Dussault et al 2004, 2005, Sinclair et al. 2006, p. 69). Consequently individuals must make trade -offs to maximize fitness and often this involves deciding how far to stray from cover to acquire resources. For moose and other ungulates, cover can be important in all seasons for reasons including protection from predation, and relief from deep snow, heat, or extreme cold and wind (Peek 1997, pp. 368-372). Moose are susceptible to predation from wolves , bears and humans (Van Ballenberghe and Ballard 1994). To hide from predators, ungulates need lateral cover which is comprised of dense low and mid-level vegetation that breaks up the shape of their bodies making them harder to detect by coursing predators (Timmermann and McNicol 1988, Mysterud and Ostbye 1999, Altendorf et al. 2001, White and Berger 2001). Concealment cover is presumed to be especially critical for moose calves because of their high susceptibility to predation, with various studies reporting neonate predation-induced mortality to be between 30 and 70% (Franzmann et al. 1980, Ballard et al. 1981, Larsen et al. 1989, Osborne et al. 1991, Gasaway et al. 1992, Garner 1994).

Deep snow is energetically costly for any ungulate species and although moose are well-adapted for this condition, their movements can still be limited by snow depth and condition, especially for calves. As snow depth increases, movement becomes more costly and eventually moose need to shift to habitats dominated by mature conifers with less snow accumulation (Coady 1974, Timmermann and McNicol 1988, Courtois et al. 2002). Depths of 70 cm impede moose movement and at 90-100 cm moose are confined to areas with a dense coniferous canopy (Coady 1974). Peterson and Allen (1974) found that more calves on Isle Royale were killed when snow depths exceeded 76 cm and Loveless (2009) found that wolves in Algonquin Park consumed the most moose biomass when snow depth was highest. In addition to lower snow depths, dense coniferous canopies provide softer snow often making travel easier for moose. In a mild winter, Peek (1971) found that moose moved to denser canopies at a snow depth of only 30 cm because snow hardness under open canopies made movement more costly. Thermal stress is presumed to be a common condition for moose though nearly always due to heat, not cold (Schwartz and Renecker 1997, p. 468). Moose have many physiological adaptations for low temperatures, including their insulative pelage and large size which helps by conserving heat and reducing energy needs in the winter. At the northern edge of their distribution, moose may be constrained not by cold temperatures but by the absence of forest (Kelsall and Telfer 1974). Lower critical temperatures have not been well-tested, although adult moose have been reported to show no visual sign of distress at -40C (Schwartz and Renecker 1997, p. 469). Evidence suggests cold stress can occur more commonly in calves (at -30C, Renecker et al. 1978) and in late winter as a result of tick-induced hair loss (Blyth and Hudson 1987, Glines and Samuel 1989).

However, the southern range periphery is thought to be determined largely by temperature as moose are reported to be heat-stressed in the winter at 5C, and in the summer at 14C, with a panting threshold at 20C (Renecker and Hudson 1986). To avoid heat stress moose may seek mature stands with coniferous trees to reduce exposure to solar radiation (Schwab and Pitt 1991, Dussault et al. 2004, but see Lowe 2009). Because forest stands providing high quality cover for moose are usually dominated by closed coniferous canopies, they offer food resources at a lower quantity and quality than other forested habitat. Moose generally prefer hardwood browse over conifer species (Crte 1989, Conrad 2000) possibly because conifers have elevated levels of secondary compounds, such as tannins, that can affect palatability and nutrient absorption (Robbins et al. 1987, Bryant et al. 1991). Additionally, the closed canopy of cover habitat allows less sunlight to penetrate and less vegetation is able to grow in the midstory, diminishing browse quantity. Therefore moose need to leave cover in order to better meet their nutritional requirements. The concept that moose movements are constrained by the abundance and juxtaposition of cover has been well-documented (Coady 1974, Crte 1977, Telfer 1978, Welsh et al.1980, Hamilton et al. 1980, Thompson and Vukelich 1981, Eastman and Retcey 1987, Peek et al. 1987, Courtois and Beaumont 2002, Dussault et al. 2006). In Ontario, Hamilton et al. (1980) found declining trends in winter browse use with increasing distance to cover. Although they did not find an upper distance limit from cover to feeding locations , 95% of browse use was recorded within 80 m of forested cover. Thompson and Vukelich (1981) found that although exceptional distances of 400 meters were found in early winter, average distance to forested cover was 27 meters.

Habitat suitability models for moose commonly include a component related to availability or proximity to cover (Allen et al. 1987, Puttock et al. 1996, Naylor et al. 1999, Dussault et al. 2006), yet these components are often assumed without empirical validation, especially in the geographic region in which it is intended to be applied. This is the case for moose habitat models in the temperate forests of the Great Lakes Region. Two models were created in 1987 by the United States Fish and Wildlife Service (USFWS) (Allen et al. 1987). The models operate at different scales with Model I estimating carrying capacity at the fine scale of 600 ha (the assumed size of a moose home range) and Model II estimating carrying capacity at a coa rser scale of approximately 9000 ha (an area presumed large enough to support a population) (Allen et al. 1987). In 1999 the Ontario Ministry of Natural Resources (OMNR) created a new model based upon components of Model I to aid in forest planning in the Great Lakes St. Lawrence Forest (Naylor et al. 1999). The distance to cover as sumptions in this model as well as Model I upon which it was based have never been empirically validated, although a few partial validations of Model II have occurred (Allen et al.1991, Naylor et al. 1992, Puttock et al. 1996, Rempel et al. 1997, Koitzsch 2002). However these validations were based on winter aerial survey data or in the la tter case, harvest data, so have limited interpretation value. Harvest data are spatially coarse and subject to biases (Koitzsch 2002) and aerial surveys provide only a snapshot of moose behaviour because they reflect only the winter season and the particular conditions on the day of flight: snow depth, snow hardness, temperature, etc. In contrast, GPS collars provide concise, frequent, and consistent location data for all seasons, daily periods, and weather conditions and therefore are a better tool for model validation.

The purpose of this study wa s to determine if moose select areas closer to cover and if the selection depends upon season or habitat. I also was interested in determining specific distance parameters: are moose constrained to certain distances from cover, how close do the y prefer to stay to cover, and do these values differ by season or habitat type the moose is in? The second goal of my study was to validate the distance to cover assumptions used in OMNRs habitat suitability model for moose. I expected to find that: 1) overall, moose would select areas close to cover, 2) cover in the dormant season would be more important than in the growing season, and 3) moose would be closer to cover when they were in early successional stands than when in mature forest, especially during the dormant season.

CHAPTER 2: METHODS 2.1 Study Area This study took place in a 2545 km2 portion on the western side of Algonquin Provincial Park located in southeastern Ontario, Canada in the Great Lakes- St. Lawrence forest region (45 North, 78 West, Figure 2.1). The 7600 km2 park is comprised of shade -tolerant upland hardwoods on poorly drained glacial till, and is intersperse d with lakes, wetlands, and mixed and conifer stands in the lower areas (Crins et al. 2008). Elevation ranges from 150 - 590 meters above sea level (Friends of Algonquin Park, 2005) and dominant tree species include sugar maple (Acer saccharum), American beech (Fagus grandifolia ), eastern hemlock (Tsuga canadensis), yellow birch (Betula alleghaniensis), and red maple (Acer rubrum) (Crins et al. 2008, Appendix A, Table 1). Timber harvesting occurs in 78% of the park (56% with water and non-harvestable areas

within the harvesting zones excluded) and is comprised largely of selective and shelterwood cuts with clearcuts consisting of < 5% (Cumming 2009). Highway 60, a major 2-lane route, bisects the southern portion of the park. Recreational human use (camping, boating, fishing, hiking) is focused on the major lakes and the few side roads within the highway corridor. Interior (logging) roads are not open to public vehicles, though backcountry access is available via canoe routes. Although Aboriginals harvest moose on the east side of the park, my study area in the western side was not subject to moose harvest. During the study period, moose population estimates for Wildlife Management Unit 51, the area comprising the majority of the park and the entire study area , increased from 2100 in 2006 to 3100 in 2009, producing a density of 0.29/km2 and 0.43/km2 , respectively (Steinberg and Francis 2006, Steinberg, 2009).

Figure 2.1. Study area- Algonquin Provincial Park in southeastern Ontario, Canada.

2.2 Field Methods A total of 21 adult female moose (mean age : 4 2 SD years, age range: 1-7) were equippedwith Lotek 3300L store-on-board GPS collars (Lotek Wireless, Newmarket, ON, Canada) in January 2006 and February 2007. Animals were captured via helicopter by net-gunning in 2006 (Bighorn Helicopters Inc., Cranbrook, BC, Canada) , and by darting in 2007 (Heli-horizon Inc., Quebec City, QC, Canada ). The anaesthetic applied in 2007 included a mixture of carfentanil (Wildlife Pharmaceuticals Inc., Ft. Collins, Colorado, USA) at approximately 0.0070 mg/kg combined with xylazine hydrochloride at approximately 0.2 mg/kg. This drug combination was reversed with naltrexone at 7

approximately 0.7 mg/kg. Radio collars recorded fixes at a two-hour interval and were deployed for approximately two years. Survival was monitored usually once or twice per month by fixed-wing aircraft and mortalities were investigated from the ground to determine cause of death. Collars were removed in March 2008 by Heli-horizon Inc. using the same drugging method described above. Field methods were approved by the Trent University and Ontario Ministry of Natural Resources Animal Care Committees under the guidelines of the Canadian Council on Animal Care. GPS collar positional accuracy was believed to be similar to results of an accuracy study done with the same collar model in the same study area (Maxie 2009). Approximately 1150 fixes were collected in open, hardwood, conifer, and mixed coniferhardwood habitats with an average 3D fix error of 14 2 m SD, 2D fix error of 34 4 m SD, and 3D fixes comprising 83.5% of all locations.

2.3 Habitat Data and Validation Habitat analysis was conducted using a Forest Resource Inventory (FRI) map of the study area. The FRI is a digital database created from visual interpretation of 1:15,840 aerial photos, calibrated by ground data, and updated with fire and harvest events every five years. It provides stand-level detail including species composition, age, height, and stocking (OMNR 2008) with a minimum stand size of 4 ha in the study area. The current FRI for the study area was interpreted from photos taken in 1989. Recent timber harvest spatial data including harvest type and date was provided by the Algonquin Forestry Authority (Huntsville, Ontario, Canada) and used to determine forest age.

In 2007, we evaluated species composition accuracy of the FRI in the study area by comparison with plot-based field estimates of species composition (Maxie et al. in press). In our study area, 25 standard forest units and 8 non-productive forest units were derived from the FRI (Elkie et al. 2007). We collapsed similar units into eight study groups, including six forest types, wetlands and water. Poor agreement between the FRI and field data led us to further collapse forest types to four. These four forest types (hemlock, conifer, hardwood, and mixed conifer-hardwood) yielded an overall accuracy level of 77%. According to our ground data, hemlock stands were misclassified as hardwood 50% of the time (Maxie et al. in press). Because of poor map accuracy, I was unable to include the full spectrum of standard forest units in my analyses and model validation, and although hemlock may be an important source of cover for moose (Forbes and Theberge 1993, Naylor et al, 1999), I merged it into the hardwood group because of the frequency of misclassification as hardwood. The resulting habitat groups in the analysis consist of three forest types (conifer, hardwood, and mixed) and three seral groups (presapling, sapling, and mature) yielding an accuracy of 91%. Appendix A, Table 1 illustrates how FRI-based standard forest units were combined into the three forest types. Seral stage refers to periods of forest succession and is defined according to age and FRI-based forest unit. Because some types of forest mature faster than others, the age at which a forest advances to the next stage depends on the specific forest type. Appendix A, Table 2 illustrates how seral stages were calculated for the habitat groups in relation to the FRI forest units that make up each study group. I designed the seral stage age cutoffs to reflect those of the majority of FRI forest units within each study group. However,

I strayed from the FRI in the way in which I calculated age. The FRI, and consequently OMNRs moos e model, defines age based on the oldest trees but I defined age according to the most recent harvest. Although clear-cutting was not used in this study area, 30-35% of the total volume is typically removed in each cut (AFA 2005). Thus, factors important to moose such as browse availability and canopy closure more closely reflect younger seral stages than older seral stages. Field observations during our habitat accuracy exercise supported this belief. Consequently, I have slightly adapted the seral stage definitions found in Holloway et al. (2004). I defined the presapling seral stage as the youngest categor y of forest where vegetation includes herbaceous plants, shrubs, and tree seedlings <3 meters in height and possibly some mature trees as well. The sapling seral stage was defined as the second category of development where trees begin to dominate and sublethal competition begins to occur. Mean tree height of the new generation of trees is three meters. My mature seral group combined the FRI seral groups of immature, mature, and old and was defined as forest older than sapling where canopies become more closed, understories more sparse, and lethal competition occurs. I combined the three forest species types (hardwood, mixed, and conifer) in the presapling and sapling seral groups yielding five study groups for my analyses: presapling, sapling, mature hardwood, mature mixed, and mature conifer. Though constrained to only a few habitat groups by poor map accuracy, I delineated habitat groups to best validate the distance-to-cover assumptions in OMNRs moose habitat suitability model (Naylor et al. 1999). In the study area, mature hardwood was the most dominant habitat group at 25% followed by presapling (19%) , sapling (18%) , mature

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mixed (11%), and mature conifer (5%). Non-forested areas comprised 22% consisting almos t entirely of water and wetland, and were excluded from analysis. Though some recent moose studies measured a single level of cover (Dussault et al. 2006, Lowe 2009), I considered a gradient of cover in my analyses. I will refer to mature conifer as cover because it represents the highest quality cover of my habitat groups in both seasons. I additionally considered two lesser cover categories in the dormant season, mature forest, and sapling plus, with the latter term referring to any forest at sapling stage or older. These lesser cover categories are explained more thoroughly in the model validation section of analysis methods.

2.4 Analyses 2.4.1 Selection of Cover Population Level Selection Selection of proximity to cover, which I will refer to as selection of cover, was assessed using a type III study design in which use and availability are measured separately for each individual instead of as a group (Thomas and Taylor 1990, Manly et al. 2002). A Euclidean distance approach was used with distance of forested animal locations from cover as use and distance from random forested locations within home ranges from cover as available (Conner and Plowman 2001). Spatial analyses were performed using ArcInfo (Environmental Systems Research Institute Inc. 1999, 2006) and home ranges were created in Home Range Tools for ArcGIS (Rodgers et al. 2007). Seasonal home ranges were created using 99% fixed kernels and a referenced bandwidth. A fixed kernel density estimator was used instead of an adaptive kernel model

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because as Kenward and Hodder (1996) suggested, the adaptive model widened the kernels in the outlying regions of the home range, effectively over-expanding the utilization distribution and leading to an overly liberal area of availability for my analysis. The referenced bandw idth smoothing parameter (Worton 1995) was chosen because it smoothed all home ranges in a consistent way so that they were comparable. In contrast, t he least squares cross-validation method (Bowman 1984, Rudemo 1982) failed to calculate a smoothing parameter for some home ranges and defaulted to a smaller smoothing parameter that resulted in highly under-smoothed utilization distributions compared to the rest. Biased cross-validation (Scott and Terrell 1987), the plug-in approach (Wand and Jones 1995) and Brownian bridge method (Horne et al. 2007) each produced overly conservative (under-smoothed) home ranges for my purpose, effectively excluding areas that were likely available to non-territorial and highly mobile species such as moose. Separate home ranges were estimated for each individual during each season for each year (Rettie and Messier 2000). Seasons were based on browse availability and define d according to the OMNRs moose habitat suitability model (HSM) with a growing season from May 16th to September 30th and a dormant season from October 1st to May 15th (Naylor et al. 1999). Selection was determined for cover (mature conifer) in both seasons and for the lesser cover categories (mature forest and sapling plus) in the dormant season. To represent availability, random locations were generated within each home range using Hawths Analysis Tools (Beyer 2004). Locations were stratified in a 1:1 ratio by habitat group according to the animals proportional use in each group. For example,

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if in a given home range, an animal had 400 locations in mature hardwood, then 400 random locations were created in mature hardwood within that home range. The distance from each random location to the nearest element of each of three levels of cover was determined by creating Euclidean distance rasters (10-meter cell size) in ArcGIS Spatial Analyst Tools and extracting distance values at the random points. These va lues were averaged for each seasonal home range and for each habitat group within each home range , producing the expected values under the null hypothesis of no selection. The same procedure was applied using the moose locations to calculate the average observed distances to cover. Selection ratios of observed mean distances to expected mean distances were calculated for each home range and for each habitat group within each home range. These ratios were used to determine selection of cover, with ratios significantly less than one indicating selection and significantly greater than one indicating avoidance (Conner and Plowman 2001, Conner et al. 2003). Selection of cover was determined for each season and for each habitat group within season using selection ratios as the dependent variable in ANOVAs (Statistica 7.0, StatsSoft Inc 2001). The expected value of the null hypothesis was converted from one to zero by subtracting a constant of one from the ratio data and consequently a n ANOVA intercept significantly different than zero would lead to a rejection of the null hypothesis (Conner and Plowman 2001). Individual animal was used as a blocking variable to assess variation in individuals and account for multiple measures of the same animal because seasonal selection ratios were calculated separately for 2-3 consecutive years (Chamberlain and Leopold 2000). Individual seasonal home ranges were removed from habitat-level analysis if they had fewer than 20 locations in that habitat. This helped

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avoid spurious selection ratios resulting from insufficient sample sizes and ensure that adequate numbers of locations existed for bootstrapping at the home range level (described below) (Stine 1985, Boos and Brownie 1989, Zhang et al. 1991) .

Home Range Level Selection Because I was interested not only in population level behaviour but also in variation within the population, I tested selection of the three types of cover on a home range basis, both for the entire home range and for each habitat group within the home range. To determine selection for each home range , I bootstrapped the means of both the observed and expected distances (Gillingham and Parker 2008) 1000 times to create 95% confidence intervals using the statistical package R (R Development Core Team 2006). Each of the 1000 replications used 95% of the sample size and was sampled with replacement. Bootstrapping was chosen over the parametric standard error of the mean because the distance data were not normally distributed and some home ranges had small sample sizes (approaching as low as n = 20). Selection of cover was inferred to have occurred in a home range if the confidence intervals around the means of observed and expected distances did not overlap and the observed mean was lower than the expected mean. Because approximately 450 comparisons were made, experiment-wise error likely resulted in approximately 5% or 23 false positive s, underestimating the outcome of no selection by 5% and over-estimating the outcomes of selection and avoidance by 2.5% each. I considered this when interpreting results.

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Year Effect Environmental conditions affecting habitat use such as temperature and snow depth can vary annually so I tested for a year effect for both dormant and growing seasons using ANOVA. Because radio-collars were deployed in the middle of the first dormant season and removed in the middle of the third dormant season, the three years represented different periods and werent comparable. Accordingly, to test for a year effect in the dormant season I created separate home ranges and random points to calculate selection ratios based only on the overlapping mid-winter period, February 2nd March 6th. The dependent variable in the ANOVAs was selection ratio and individual animal was again used as the blocking factor.

2.4.2 Distance Parameters After selection was assessed, three distance parameters were calculated on a home range basis and summarized at the population level: median distance, 95-percentile distance, and for home ranges where selection occurred, the preference zone was determined. Distance parameters were calculated with all habitats combined as well as with each habitat separate. Ninety-five percentile distances were calculated to indicate how far animals strayed from cover barring temporary excursions and anomalous movements. Preference zone is described below. The preference zone was defined as the area within which moose preferred to be from cover. Visually the edge of this zone is the point in the distance distribution where the number of expected locations exceeds the number observed locations (See Figure

15

2.2a for example). To calculate the preference zone edge, observed and expected distance-to-cover values for each home range were binned into 100-meter sections. In each distance bin, selection ratios were formed using the count of observed locations over expected locations with ratios above one indicating selection for the given distance category and ratios below one indicating avoidance. These selection ratios and the midpoint of their associated distance bins were used in a linear regression as the dependent and independent variables, respectively. However, because the relationship between selection ratio and distance followed a negative exponential distribution, I logtransformed the selection ratios after adding a constant of one. Because selection ratios created from a very small number of locations may be overly influential outliers, I weighted the regression by the total number of observed and expected locations in each distance category. I then solved all the linear regression equations for a selection ratio of one (y value of 0.301) to find the edge of the preference zone for each home range where selection for cover occurred (See Figure 2.2b for example). Determining the preference zone is a new approach and may be more useful than traditional measures (such as means) for management and habitat modeling because it describes actual animal behaviour. Unlike a simple mean distance, the preference zone incorporates both use and expe cted distributions and is a more valid measure of resource selection.

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Observed Mean = 357 Expected Mean = 435 240 220 200 180 Number of observations 160 140 120 100 80 60 40 20 0
100 200 300 400 500 600 700 800 900 1000 1100 1200 1300 1400

Observed Expected

Distance to cover (meters)

Figure 2.2a. Observed and expected distributions of distance-to-cover from hardwood locations for Moose7 in the dormant season of 2006-2007. Preference zone edge can be estimated as the general area where the expected values begin to outnumber the observed values.

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Figure 2.2b. Linear regression for Moose7 in the dormant season of 2006-2007 when in hardwood stands. The preference zone is the area within 394 meters of cover. A Y value of 0.301 is equal to the selection ratio of one.

2.4.3 Model Validation Model Description The second goal of the study was to validate the distance to cover parameters in OMNRs habitat suitability model (HSM) (Naylor et al. 1999). In the growing season the HSM delineates two types of forest stands : those which provide thermal cover and those which do not, and it assumes that moose are confined to the area within 1500 meters of thermal cover. In the dormant season, the model delineates four types of forest providing

18

various degrees of cover: late winter cover, early winter cover, lateral cover, and no cover. It assumes that moose are always constrained to areas within 1600 meters of late winter cover. Additionally, when moose are in a stand providing only lateral cover, they are assumed to also be constrained to areas within 400 meters of early or late winter cover. Three assumptions exist when moose are in a stand that provides no cover. In addition to the assumptions above, the y are assumed to also be constrained to areas within 200 meters of any level of cover (lateral, early winter, or late winter). HSM descriptions of these levels of cover and my corresponding habitat groups are given in Table 2.1 and a summary of the distance assumptions is given in Table 2.2. I assigned my habitat groups to the models cover categories in the way that most closely reflected the majority of FRI forest units within each group (See Appendix A, Table 3). I will refer to the 1600-meter assumption as mature conifer, the 400-meter assumption as mature forest, and the 200-meter assumption as sapling plus. (These terms refer respectively to distance 1, distance 2, and distance 3 in the HSM documentation.) The HSM uses the distance assumptions to create grids indicating the range of available forested habitat for moose in each season. The full model carrying capacity map is a product of three sub-model carrying capacity maps: dormant season, growing season, and aquatic feeding. The dormant and gr owing season maps are a product of their seasonspecific range and forage grids. The range grids I am validating in this study directly affect the model output because areas that fall outside of the available range result in a carrying capacity of zero in the sub-model and subsequently in the final model. I assessed the validity of the distance to cover parameters using three methods.

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Season Dormant

Cover Category No cover Lateral cover Early winter cover Late winter cover Thermal cover

Description Height < 3 m Height between 3 and 6 m Height > 6 m with some conifer cover Height > 10 m and conifer or coniferdominated mixedwood Height = 10 m and lowland conifer, lowland mixed, or lowland hardwood

Habitat group Presapling Sapling Mature hardwood and mixed Cover (mature conifer) Cover (mature conifer)

Growing

Table 2.1. The Ontario Ministry of Natural Resources habitat suitability models description of cover categories (Naylor et al. 1999) and my corresponding habitat groups.

Season Dormant

Habitat moose is in Presapling Sapling Mature hardwood or mixed Growing Non-thermal cover forest

Mature conifer 1600 m 1600 m 1600 m 1500 m

Mature forest 400 m 400 m ---

Sapling plus 200 m ----

Table 2.2. Distance-to-cover assumptions in the Ontario Ministry of Natural Resources habitat suitability model for moose and expected 95 percentile distances from moose locations to distance categories.

Validation Method 1 The first validation method simply involved running the HSM and plotting the moose locations over the seasonal range outputs from the model in ArcGIS. Each range output is a binary surface of hexagonal parcels with each hectare parcel indicating that it is either available habitat (parcel value of 1) or unavailable (parcel value of 0). I calculated the percent of all forested moose locations that fell within the forested area 20

deemed unavailable. This helped to determine if the models distance parameters were overly conservative (i.e., if moose were willing to travel further from cover than assumed). To understand which specific assumptions may be overly conservative, I further calculated the percent of moose locations that fell within the assumed unavailable area for each habitat group. Inferential statistics could not be used here to compare habitat groups because of the lack of variance but the percentage measures were still able to indicate effect size.

Validation Method 2 Beyond this first simple approach, I was limited in my ability to validate the HSM directly as it is based upon the full spectrum of FRI forest units and low map accuracy forced me to use combined habitat groups . However, I was able to evaluate how well my habitat groups performed in the model. As a second validation method, I substituted the HSMs levels of cover with my corresponding habitat groups (Table 2.1) to determine the available and unavailable areas of my study area. As in the first method, I calculated the percent of all forested moose locations that fell within the unavailable area and further calculated the percent of locations in each habitat group that fell within that area. I was also able to quantify which of the three distance assumptions were violated for each location using the distance values extracted from the Euclidean distance rasters (see Analyses: Population Level Selection). Again, inferential statistics could not be used to compare habitat groups because of the lack of variance but the percentage measures were still able to indicate effect size.

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Validation Method 3 The two approaches described above were only able to detect instances where the model may be too conservative. To determine if any of the models distance parameters are overly liberal (i.e., if moose arent willing to travel as far from cover as assumed), I compared the distance assumptions to the 95 percentile of the distribution of distances that moose were located from cover. Unlike the first two validation methods that used pooled moose locations, this comparison operated at the individual seasonal home range level. The validation for the growing season was simple since there is only one distance assumption (1500 m from cover). Because the dormant season has multiple assumptions depending on the habitat type the animal is in, I calculated 95 percentile distances to three groups: my cover group (mature conifer), the three mature forest groups combined (mature forest), and the sapling and three mature forest groups combined (sapling plus). The 95 percentiles were compared to the HSM distance assumptions (Table 2.2).

CHAPTER 3: RESULTS A total of 94 home ranges (21 individual animals) comprised of 58 dormant season and 36 growing season ranges were calculated for the analyses. Growing season analyses included 19 individuals because one moose died and one collars data were censored from analyses for the growing seasons due to poor fix success. Seasonal home range sizes were quite variable, averaging 50 km2 ( 44 SD) in the 7.5-month dormant season and 38 km2 ( 23 SD) in the 4.5-month growing season. Overall GPS collar fix

22

success was 91% but many of the missed fixes originated from the one collar that malfunctioned during the summers. With this collar removed, fix success averaged 94% 12% SD.

3.1 Selection of Cover - Year Effect Environmental conditions were similar in the growing seasons but differed in the dormant seasons with the second dormant season having less snow depth than the first and third (Table 3.1). Home ranges were larger during the second dormant season but this increased use of space did not affect distance to cover. Selection of cover was not significantly affected by year in either the growing or dormant season (F1,16 = 2.83, p = 0.11 and F2,34 = 1.484, p = 0.24, respectively). All ANOVA assumptions were met. Normality plots are depicted in Appendix B: Figure 1.

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Season Growing 2006 Growing 2007

Historic Normal

Dates
May 16 - Sept 30

Dormant 2005-2006 Dormant 2006-2007 Dormant 2007-2008

Historic Normal

Oct 1 - May 15

Midwinter 2006 Midwinter 2007 Midwinter 2008

Historic Normal

Feb 2 - Mar 6

Mean Daily Max. Temp (C) 22 ( 5.6) 23 ( 4.8) 21 4 ( 9.3) 4 ( 9.2) 4 ( 9.6) 4 -5 ( 4.7) -7 ( 5.4) -4 ( 5.1) -4

Mean Daily Min. Temp (C) 8 ( 5.0) 7 ( 4.7) 9 -8 ( 10.1) -8 ( 10.4) -10 ( 10.5) -7 -19 ( 9.0) -21 ( 7.2) -19 ( 9.3) -16

Mean Temp (C) 15 ( 4.8) 15 ( 4.2) 15 -2 ( 9.2) -2 ( 9.3) -3 ( 9.6) -2 -12 ( 6.2) -14 ( 5.6) -11 ( 6.8) -10

Mean Snow Depth (cm) ---32 ( 27.9) 12 (11.7) 34 ( 12.2) 28 72 ( 7.4) 27 ( 5.1) 35 ( 4.5) 65

Table 3.1. Mean temperature and snow depths ( SD) for the analysis periods taken from Algonquin Park East Gate weather station (Environment Canada 2008). Historic normals are averaged from 1971-2000. Year effect analyses were performed using growing season and midwinter season. Asterisks indicate snow depths known to influence moose movement.

3.2 Selection of Cover - Home Range Level At the home range level, bootstrapping revealed great variability in selection of mature conifer cover in each habitat group of each season (Figure 3.1). In the growing season the number of home ranges where cover was selected was greater than or equal to the number of home ranges where cover was avoided for each habitat group; in the dormant season the number of home ranges indicating selection always exceeded the number of home ranges indicating avoidance (Figure 3.1a). Selection of the lesser cover categories in the dormant season (mature forest and sapling plus) revealed less variation. The number of home ranges where cover was selected always outnumbered those where cover was avoided but in each case, the most common outcome, no 24

selection, comprised 50% of home ranges (Figure 3.1b). Because approximately 450 comparisons were made, experiment-wise error likely resulted in 5% or 23 false positives, potentially underestimating the outcome of no selection by 5% and overestimating the outcomes of selection and avoidance by 2.5% each. However, this adjustment for experiment-wise error does not mask the trends in variance of selection.

70% 60% 50% 40% 30% % S 20% 10% 0% Mature Mixed Mature Hardwood Combined habitats Presapling Combined habitats Mature Hardwood Mature Mixed Presapling Sapling Sapling % NS % A

n = 36 n = 36 n = 31 n = 15 n = 22 n = 58 n = 54 n = 49 n = 26 n = 40 Growing Season Dormant Season

Figure 3.1a. Variation of selection behaviour at the home range level for mature conifer cover. S = selection, A = avoidance, and NS = no selection. N refers to the number of home ranges in each category.

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60% 50% 40% 30% 20% 10% 0% Sapling plus Presapling n = 26 Mature forest Presapling n = 26 Mature forest Sapling n = 40 %S % NS %A

Figure 3.1b. Variation of selection behaviour at the home range level for the lesser cover categories associated with presapling and sapling forest in the dormant season (mature forest and sapling plus). S = selection, A = avoidance, and NS = no selection. N refers to the number of home ranges in each category.

3.3 Population Level Seasonal Results In the growing season the intercept of the ANOVA indicated no selection for proximity of cover, meaning that at the population level, moose tended to be the same distance from cover within their home ranges as expected by chance (F1,17 = 0.908, p = 0.354, selection ratio = 1.00 0.23 SD) (Table 3.2a). However, moose ID was a significant source of variation (F18,17 = 7.339, p < 0.01) and bootstrapping revealed that in only 19% of home ranges did moose show no selection with moose in 42% of home ranges showing selection, and 39% showing avoidance. The average median distance animals were from cover was 470 meters and 95% of the locations were within 1144 26

meters of cover. Those figures for just the moose that selected cover (n = 15 of 36) were 321 and 991 meters, respectively (Table 3.3). The average preference zone edge was 621 meters. In the dormant season the intercept of the ANOVA indicated a marginally significant trend in selection of cove r at the population level (F1,37 = 3.176, p = 0.083, 0.93 0.21 SD) (Table 3.2a), with moose ID also being marginally significant (F20,37 = 1.715, p = 0.076). The average median distance animals were from cover was 427 meters and 95% of the locations were within 1135 meters. When just considering those moose that selected cover (n = 36 of 58), those figures were 304 and 1014 meters, respectively (Table 3.3). The average preference zone edge was 532 meters. All ANOVA assumptions were met. Normality plots associated with these ANOVAs are shown in Appendix B, Figure 2.

3.4 Population Level- Habitat Results During the growing season, moose as a whole did not significantly select or avoid proximity to cover when they were in any of the habitat groups, although there was a marginally significant avoidance when moose were in saplings (F1,10 = 3.747, p = 0.082, 1.407 1.04 SD ) (Table 3.2a). Distance parameters for all home ranges, those selecting cover, and those avoiding cover are presented in Table 3.3. Figures 3.2a-c illustrate distance parameters for home ranges showing selection for cover. Depending upon the habitat group, selection of cover occurred in 32 to 46% of growing season home ranges. In the dormant season, moose as a whole did not significantly select or avoid proximity to cover when they were in any of the habitat groups, although there was

27

marginally significant selection in mature hardwoods (F1,34 = 3.919, p = 0.056, 0.937 0.29 SD) (Table 3.2a). The preference zone edge was significantly lower in mature hardwood than in presapling stands (Figure 3.2c). The proportion of dormant season home ranges in which selection of cover occurred ranged from 38 to 48% depending upon the habitat group. For every habitat group in this season, proportionately more individuals selected cover and distance parameters were consistently lower, indicating that cover is more important in the dormant season than the growing season. All ANOVA assumptions were met. Normality plots at the habitat group level for both seasons are shown in Appendix B, Figure 3. In the dormant season I also assessed selection and distance parameters of the two lesser categories of cover, mature forest and sapling plus. When in young forest, moose as a whole did not select areas closer to any lesser type of cover (Table 3.2b). Distance parameters for all home ranges, those selecting cover, and those avoiding cover are presented in Table 3.4a-b. Figures 3.3a-c illustrate distance parameters for home ranges that showed selection for cover. All ANOVA assumptions were met. Normality plots are shown in Appendix B, Figure 4.

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A. Selection Ratio SD 1.003 0.234 0.989 0.325 1.010 0.321 0.999 0.280 1.407 1.042 0.934 0.212 0.937 0.288 0.901 0.287 1.065 0.456 1.031 0.486

Season Habitat Group Growing Combined Hardwood Mixed Presapling Sapling* Dormant Combined* Hardwood* Mixed Presapling Sapling

ANOVA results F1,17 = 0.908, p = 0.354 F1,16 = 0.153, p = 0.700 F1,14 = 0.122, p = 0.732 F1,7 = 1.305, p = 0.291 F1,10 = 3.747, p = 0.082 F1,37 = 3.176, p = 0.083 F1,34 = 3.919, p = 0.056 F1,30 = 2.002, p = 0.167 F1,16 = 0.219, p = 0.646 F1,23 = 1.814, p = 0.191

B. Selection Ratio SD 0.955 0.107 1.018 0.093 0.935 0.105

Habitat Presapling Presapling Sapling

Cover category ANOVA results Sapling plus F1,16 = 0.007, p = 0.936 Mature forest F1,16 = 0.484, p = 0.497 Mature forest F1,24 = 2.645, p = 0.117

Table 3.2 Results of habitat group level mature conifer cover selection ANOVAs (A) . and selection of the lesser cover categories associated with presapling and sapling in the dormant season. Asterisks indicate marginal significance. Sapling plus refers to the combination of any sapling or mature forest (B).

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All home ranges % S 42 46 35 40 32 62 48 39 38 40 % NS 19 29 39 20 36 14 31 41 31 30 % A 39 26 26 40 32 24 20 20 31 30 Observed Median 470 407 448 589 531 427 335 350 565 503 CI 82 66 82 160 168 81 56 56 99 136 Expected Median 446 393 395 597 497 448 343 376 546 461 CI 63 40 47 148 165 72 29 29 83 118

Home ranges selecting cover Observed Median 321 246 277 561 323 304 217 212 436 242 CI 43 54 65 260 95 50 32 40 104 99 Expected Median 408 362 395 700 456 396 327 347 648 374 CI 56 60 90 153 96 54 41 44 151 131

Home ranges avoiding cover Observed Median 545 596 638 558 695 749 584 581 822 832 CI 94 79 191 125 217 235 164 150 174 314 Expected Median 411 380 354 443 458 601 351 444 480 523 CI 68 68 103 117 239 248 40 76 121 319

Season

Growing Season

Habitat Group Combined habitats Mature Hardwood Mature Mixed Presapling Sapling Combined habitats Mature Hardwood Mature Mixed Presapling Sapling

n 36 36 31 15 22 58 54 49 26 40

Dormant Season

Table 3.3a. Moose median observed and expected distances to cover plus 95% confidence intervals (in meters) by season and habitat group for all home ranges, those where cover was selected, and those where cover was avoided. N refers to the number of home ranges in each group. %S = % home ranges showing selection, %NS = % showing no selection, and %A = % showing avoidance.

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Season

Habitat Group Combined habitats Mature Hardwood Mature Mixed Presapling Sapling Combined habitats Mature Hardwood

n 36 36 31 15 22 58 54 49 26 40

% S 42 46 35 40 32 62 48 39 38 40

% NS 19 29 39 20 36 14 31 41 31 30

% A 39 26 26 40 32 24 20 20 31 30

All home ranges Observed Expected 95% 1144 1000 932 1135 1171 1135 943 911 1062 1132 CI 154 120 118 196 291 117 81 94 134 216 95% 1225 1079 1044 1328 945 1247 1054 1061 1223 1219 CI 149 98 122 298 269 116 72 88 154 224

Home ranges selecting cover Observed Expected 95% 991 856 759 1132 943 1014 760 785 1059 837 CI 143 179 86 284 379 130 78 152 263 314 95% 1152 1067 1127 1535 1243 1203 988 1048 1394 1134 CI 113 151 143 385 206 141 97 140 264 367

Home ranges avoiding cover Observed Expected 95% 1149 1188 1034 970 1364 1496 1223 1197 1136 1383 CI 190 213 246 172 421 272 118 175 179 438 95% 1106 1060 995 926 1013 1451 1107 1111 1050 1309 CI 136 196 235 222 454 276 146 213 177 461

Growing Season

Dormant Season

Mature Mixed Presapling Sapling

Table 3.3b. Moose observed and expected ninety-five percentile distances plus 95% confidence intervals (in meters) to cover by season and habitat for all home ranges, those where cover was selected, and those where cover was avoided. N refers to the number of home ranges in each group. %S = % home ranges showing selection, %NS = % showing no selection, and %A = % showing avoidance.

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All home ranges Habitat Group Presapling Presapling Sapling Cover Category Sapling plus Mature forest Mature forest % S 35 35 30 % NS 50 50 50 % A 15 15 20 Observed Median 115 189 147 CI 16 37 38 Expected Median 123 183 149 CI 16 32 37

Home ranges selecting cover Observed Median 119 194 131 CI 29 54 60 Expected Median 157 224 194 CI 20 48 83

Home ranges avoiding cover Observed Median 131 274 262 CI 63 148 73 Expected Median 93 178 173 CI 49 118 64

n 26 26 40

Table 3.4a. Moose observed and expected median distances plus 95% confidence intervals (in meters) to lesser cover categories associated with presapling and sapling in the dormant season for all home ranges, those where cover was selected, and those where cover was avoided. N refers to the number of home ranges in each group. Sapling plus refers to the combination of any sapling or mature forest.

All home ranges Habitat Group Presapling Presapling Sapling Cover Category Sapling plus n 26 % S 35 35 30 % NS 50 50 50 % A 15 15 20 Observed 95% 310 432 396 CI 33 56 88 Expected 95% 361 461 412 CI 45 61 85

Home ranges selecting cover Observed 95% 312 435 409 CI 42 87 158 Expected 95% 456 521 541 CI 62 87 179

Home ranges avoiding cover Observed 95% 341 495 654 CI 161 212 189 Expected 95% 293 513 541 CI 163 254 136

Mature forest 26 Mature forest 40

Table 3.4b. Moose observed and expected ninety-five percentile distances plus 95% confidence intervals (in meters) to lesser cover categories associated with presapling and sapling in the dormant season for all home ranges, those where cover was selected, and those where cover was avoided. N refers to the number of home ranges in each group. Sapling plus refers to the combination of any sapling or mature forest.

32

Figure 3.2a. Moose observed (o) and expected (e) average median distances to cover ( 95% CI) of home ranges where selection of cover occurred for each season and habitat group. %S is the percent of all home ranges where cover was selected and n is the number of home ranges where cover was selected.

33

Figure 3.2b. Moose observed (o) and expected (e) average 95 percentile distances to cover ( 95% CI) of home ranges where selection of cover occurred for each season and habitat group. %S is the percent of all home ranges where cover was selected and n is the number of home ranges where cover was selected.

34

1800 1600 1400 1200 Meters 1000 800 600 400 200 0 Combined habitats % S = 42 n = 15 Hardwood % S = 46 n = 15 Mixed % S = 35 n = 11 Growing Season Presapling % S = 40 n=6 Sapling % S = 32 n=7 Combined habitats % S = 62 n =36 Hardwood % S = 48 n = 26 Mixed % S = 39 n = 19 Dormant Season Presapling % S = 38 n = 10 Sapling % S = 40 n = 16

Figure 3.2c. Moose average preference zones for cover ( 95% CI) for each season and habitat group. %S is the percent of all home ranges where cover was selected and n is the number of home ranges where cover was selected.

35

Figure 3.3a. Moose observed (o) and expected (e) average median distances ( 95% CI) to lesser cover categories associated with presapling and sapling forest in the dormant season for home ranges where selection of cover occurred. Sapling plus refers to the combination of any sapling or mature forest. %S is the percent of all home ranges where cover was selected and n is the number of home ranges where cover was selected.

36

Figure 3.3b. Moose observe d (o) and expected (e) average 95 percentile distances ( 95% CI) to lesser cover categories associated with presapling and sapling forest in the dormant season for home ranges where selection of cover occurred. Sapling plus refers to the combination of any sapling or mature forest. %S is the percent of all home ranges where cover was selected and n is the number of home ranges where cover was selected.

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350 300 Meters 250 200 150 100 50 0 Sapling plus Presapling % S = 35 n=9 Mature forest Presapling % S = 35 n=9 Mature forest Sapling % S = 30 n = 12

Figure 3.3c. Moose preference zone edge ( 95% CI) in relation to lesser cover categories associa ted with presapling and sapling in the dormant season. Sapling plus refers to the combination of any sapling or mature forest. %S is the percent of all home ranges where cover was selected and n is the number of home ranges where cover was selected.

3.5 Model Validation 3.5.1 Validation Method 1 For the first validation approach I simply plotted the moose locations on top of the habitat availability range grids from the HSM. The growing season range grid designated 40% of the study area, mainly in the greater Highway 60 corridor, as unavailable habitat. The large unavailable area is a result of the limited forest types that the model assumes to provide thermal cover. Specifically, thermal cover as defined by the HSM model 38

composed less than 2% of the study area and 49% of all forested moose locations fell in the unavailable area, including one animals entire home range (Figure 3.4a). To find out if violations occurred more often in any particular habitat group, I compared the proportion of points that violated the assumptions to points that did not for each habitat group. If moose were equally likely to violate the HSM assumptions in any given habitat, the n I would expect approximately the same proportion of locations (49% 5%) within each habitat group to fall within area deemed unavailable. This was the case for all habitat groups except for the sapling group where 63% of the locations were in violation of the distance assumptions , a magnitude of 14 percentage points more than the expected 49%. These violations occurred in locations of all 12 of the moose that used the habitat group. In all other habitat groups, violation proportions were within 5 percentage points of the expected 49% and therefore less likely to be of particular interest. The dormant season range grid designated only 1% of the study area as unavailable and virtually no forested moose locations (0.1%) occurred within this limited area (Figure 3.4b).

39

Figure 3.4a. Map of the growing season range calculated in OMNRs moose habit at suitability model and the seasonal moose locations falling in and outside of the available habitat in the southwestern portion of Algonquin Provincial Park, Ontario, Canada. Distance assumption violations occurred in 49% of moose locations.

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Figure 3.4b. Map of the dormant season range calculated in OMNRs moose habitat suitability model and the seasonal moose locations falling in and outside of the available habitat in the southwestern portion of Algonquin Provincial Park, Ontario, Canada. Distance assumption violations occurred in 0.1% of moose locations.

3.5.2 Validation Method 2 The second method of validation was independent of the HSM; I substituted my habitat groups into the models groups that represent various degrees of cover: no cover, lateral cover, early winter cover, and late winter cover (Table 2.1). My adapted grids designated 8% of the study area as unavailable in the growing season and 10% as unavailable in the dormant season (Figures 3.5a -b). 41

Figure 3.5a. Map of the growing season range created from my adapted model and the seasonal moose locations falling in and outside of the available habitat in the southwestern portion of Algonquin Provincial Park, Ontario, Canada. Distance assumption violations occurred in 3% of moose locations.

42

Figure 3.5b. Map of the dormant season range created from my adapted model and the seasonal moose locations falling in and outside of the available habitat in the southwestern portion of Algonquin Provincial Park, Ontario, Canada. Distance assumption violations occurred in 12% of moose locations.

Distance assumption violations in the growing season were minimal with only 3% of forested moose locations falling within the area assumed to be unavailable. To find out if these violations occurred more often in any particular habitat group, I compared the proportion of points that violated the assumptions to points that did not for each habitat group and expected the same proportion of locations (3%) within each habitat group to fall within the area deemed unavailable. Instead, 19% of sapling locations were in 43

violation of the distance assumption, though these violations only occurred among three of 12 animals. Violations occurring in the other habitat groups were all minor at 1%, 1%, and 3% for presapling, mature hardwood, and mature mixed, respectively and therefore not of particular interest. In the dormant season distance assumption violations occurred in 12% of forested moose locations. Again, if violations occurred equally in the habitat groups, I would expect the same proportion of locations (12%) within each habitat group to fall within the area deemed unavailable. In contrast 41 and 31% of the locations in presapling and sapling, respectively, were in violation and locations in mature hardwood and mature mixed forest were only in violation 0.3 and 0.5%, respectively. The vast majority of violations that occurred in young forest were violations of the distance assumptions of lesser cover: i.e., the animals were further than 200 meters fr om all types of forest providing cover (sapling plus) or further than 400 meters from all types of mature forest (Table 3.5).

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Habitat group Presapling Sapling Mature hardwood Mature mixed

n 10 16 20 19

% of points in violation 41% 31% 0% 1%

n 10 9 0 5

Mature conifer

Assumption Violations* Mature Sapling forest plus n n 0.03% 2 47% 10 77% 48% 4 74% 8 -----100% 5 ----

n 10 ----

Table 3.5. Breakdown of distance assumption violations for the dormant season. The mature conifer assumption refers to points greater than 1600 meters from cover. Mature forest assumption refers to points greater than 400 meters from all mature habitat groups (hardwood, mixed, and cover). Sapling plus assumption refers to points greater than 200 meters from saplings and all mature habitat groups combined. N refers to the number of animals associated with each result. * The sum of distance assumption violations 1 - 3 may exceed 100% in a given habitat group because some points violated more than one assumption.

3.5.3 Validation Method 3 The third validation method compared the distance assumptions to the 95 percentile distances of moose locations from cover (see Tables 3.3b and 3.4b). In all instances, moose at the population level did not significantly select areas closer to cover, meaning the 95 percentile distances merely reflect what was available within the home ranges. Therefore, these distances dont necessarily reflect the true limits for the moose population as a whole, but they at least represent a minimum threshold of the distance moose are willing to travel from cover. All of the 95 percentile distances were below the model assumptions except for presapling stands in the dormant season, where 95% of the

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moose locations were within 310 (33) meters from any type of cover (sapling plus) instead of the HSM-assumed 200 meters.

CHAPTER 4: DISCUSSION 4.1 Selection of Cover 4.1.1 Prediction 1: Overall, moose will select areas close to cover Overall, I found that cover s election by moose in Algonquin was not as strong as I had hypothesized but I did note a high degree of variation in selection ratios among home ranges. If distance to cover is not an important factor for moose, then I would expect individuals in the majority of home ranges to show no selection. Instead the populationlevel result of no selection was caused by more of a bimodal distribution of selection behaviour , with the number of home ranges where cover was selected cancelled out by home ranges where cover was avoided. This was most evident in the selection of mature conifer cover (Figure 3.1a). The variance in selection behaviour was not due to a year effect nor was it fully explained by an animal effect, as several animals switched their selection behaviour during the study. In the dormant season, 65% of individuals switched selection behaviour during the three years and 24% switched behaviour during the two growing season years. I suspect that some of this variation could be explained by calf presence, with encumbered females staying closer to cover while unencumbered moose are able to forage more freely away from cover. The belief t hat calf presence influenc es cow movements with respect to cover is fairly well documented in the literature. In southwestern Montana, bulls and cows without calves made greater use of open areas than cows with calves (Peek 1962). In northeastern Ontario, cows with calves made

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much more use of areas abundant with cover and forage than open areas (Thompson and Vukelich 1981) . In Quebec, Dussault et al. (2005) found cows with calves to have a higher preference for 10-year old mixed and deciduous stands and mature conifer stands, habitats that provide the most concealment cover, indicating a focus on protection from predators. However, to my knowledge, no study has fully examined habitat selection in relation to specific distance to cover and the needs of calves. These distance parameters would be valuable for modeling and management purposes. Factors causing moose to choose areas far from cover are less apparent but it may be a behavioura l response to those selecting cover. Moose, like most ungulates are polygynous and tend to sexually segregate during many parts of the year (Bleich et al. 1997, Bowyer 1984, Bowyer et al. 1996, 2001, Kie and Bowyer 1999, Miller and Litvaitis 1992, Miquelle et al. 1992). In a winter habitat management study, Bowyer et al. (2001) crushed areas of old-growth willow to provide more accessible forage and found the crushed areas to be beneficial only to males, as females with calves were deterred by the lack of cover. The authors consequently recommended that the sexes be treated as separate species in terms of habitat management and emphasized the importance of predation risk in management intended to aid females. Kie and Bowyer (1999) had the same recommendation for white-tailed deer. My study revealed variation in selection of cover even within females. However, this same mechanism may have been at work as Dussault et al. (2005) found habitat use of unencumbered females to more closely resemble that of males than that of females with calves. This spatial segregation may be a behavioural response of two potential processes. Moose may be employing a density-dependent foraging strategy, in which unencumbered

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individuals choose to forage in lower quality feeding areas (areas far from cover) to avoid intraspecific competition with the encumbered cows. A second factor affecting this spatial segregation could be a predator avoidance strategy, in which individuals avoid aggregating to reduce chances of detection by predators, a common strategy among ungulates of forested habitats where concealment options exist (Kie 1999).

4.1.2 Prediction 2: Cover will be more important in the dormant season than in the growing season I hypothesized that moose would show greater select ion for cover in the dormant season than the growing season and found evidence supporting this. In the dormant season the percent of home ranges where cover was selected was higher than in the growing season. In addition all median and 95-percentile distances were lower in the dormant season than in the growing season. These results are not surprising because the difference between cover and non-cover is more distinct when dec iduous foliage is absent. In the dormant season, deciduous trees provide considerably less concealment cover and thermal protection than they do in the growing season. Additionally, movement in the dormant season through stands dominated by deciduous species become s more energetically costly as snow accumulates. These phenomena are compounded as winter progresses and ungulates are weakened by a less nutritious diet (Schwartz et al. 1987, 1988, Renecker and Hudson 1989, Schwartz 1992), making proximity to cover in the dormant season even more important relative to the growing season.

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4.1.3 Prediction 3: Moose will be closer to cover when in early successional stands than when in mature stands, especially during the dormant season. At the habitat level in the dormant season, I expected that distance to cover would be least important to moose when they were in mature stands , especially mature mixed stands, since mature forest most closely resembles cover. Also a large body of literature supports the belief that browsing in open areas is focused at edges with stands providing cover (Neu et al. 1974, Bangs et al. 1985, Allen et al. 1991, Dussault et al. 2006). However, my data showed more support for the opposite effect, with selection ratios and distance parameters tending to be lower in mature forest than in young forest. Perhaps young forest provided such a high quality feeding area in my study that they were important to moose regardless of distance to protective cover. This apparent nonselection for proximity to cover in young forest could also be an artifact of the spatial patterns in my study area. Young stands (i.e. timber harvests) were typically greater than 500 m from cover, possibly too far to influence space use within those habitats. Repeatable studies in other locations would help clarify these patterns and mechanisms.

4.2 Model Validation All three of my validation approaches indicated that the model could be improved, with the first approach illustrating this most clearly. In Algonquin the biggest apparent weakness in the full HSM is the growing season thermal cover assumptions that designated a large area, mostly in the greater Highway 60 corridor, as unavailable to moose. This unavailable designation gets carried through the growing season sub-model and further to the full model, resulting in a predicted carrying capacity in that area of zero

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moose/km2 . The nearly 50% of moose locations, including all locations of an individual, that fell in this unavailable area provides strong evidence that the model is flawed. Indeed, a kriged map based on OMNRs winter aerial surveys predicted the area to have a density between 0.4 and 0.5 moose/km2 (Loveless 2009) and the two other sub-models, dormant season carrying capacity and aquatic feeding carrying capacity, predicted this area to be capable of supporting between 0.6 and 0.8 moose/km2. The estimates of these two sub-models better agree with aerial survey data than the growing season sub-model, especially considering that estimations of carrying capacity (in the HSM) will be somewhat higher than estimates of density (from aerial surveys). It is apparent in our study area that the parameters related to thermal cover are too conservative, biasing low the estimate of available growing season area , growing season carrying capacity, and finally total carrying capacity. There are a few possible reasons why the thermal cover assumptions in the growing season are flawed. One possibility is that moose are willing to feed much further from thermal cover than the 1500 m presently assumed. Although this could be true, the HSMs unavailable area is occupied by entire home ranges of a relatively high density of moose that have no access to thermal cover as defined by the model. Hence it is more likely that either 1) moose dont need thermal cover at all (Lowe 2009) or 2) that they are using other habitats as thermal cover. It is unlikely that moose dont need thermal cover because during the study period, ambient summer temperature exceeded the presumed critical temperature for moose 65% of the time and the panting threshold 31% of the time (Lowe 2009). The most plausible answer is that the forest types that comprise thermal cover in the model are too restrictive. The model assumes that thermal cover is provided

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only by four types of forest: cedar, lowland conifer, lowland mixedwood, and black spruce. These forest types comprise less than 2% of the study area , and therefore leave large areas designated as unavailable , but used as demonstrated by the study animals. When I used my adapted definition of thermal cover (all mature conifers, comprising 5% of the study area) and applied the same distance assumption, I found that moose only rarely violated the assumption (3% of the time), an improvement over the 50% of violations of the original model. Cedar, lowland conifer, lowland mixedwood, and black spruce may indeed provide better quality thermal cover than the collective group of mature conifers, but if this is the case, it doesnt appear that moose are constrained by a lack of these habitat classes. Perhaps moose make a trade-off by using lower quality but more widely available thermal cover so they can access browse across more of the landscape. Alternatively, moose may be thermo-regulating in the highly abundant wetlands in the study area. This possibility should be investigated further. If moose can utilize wetlands for thermal relief as well as the select forest stands, then distance to wetlands should be factored into the growing season range grid. During the dormant season, the models range grid deeme d almost the entire study area as available compared to the 10% unavailable of my adapted range grid. There are two differences between the off the shelf dormant season model and my adapted version leading to this discrepancy. The first difference is in the definition of late winter cover. The models designation of late winter cover includes five types of conifer forest: cedar, spruce-fir, hemlock, and two types of white pine, covering 8.5% of the study area. Hemlock was the most available type of this group making up nearly 6% of the study area. My designation of late winter cover comprised 5% of the study area and included all

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eleven types of conifer available in the study area except hemlock because of its unfortunate misclassification problem with hardwood (Maxie et al. in press). The second difference between the dormant season model and my adapted version was in the way stand age was determined, relating directly to how seral stages were defined. Because the FRI, and therefore the HSM, defines a stands age as the age of the oldest trees and I defined stand age as the time since last harvest, I ended up with a much larger presapling and sapling group than the model. All dormant season violations of my adapted model (12% of total moose locations) occurred in these two habitat groups and nearly all of them were related to the distance moose were from the lesser cover categories: >200 meters from sapling or mature forest, or >400 meters from mature forest. These violations could indicate that I was incorrect in assuming that time since last harvest was a better way to determine seral stage and what I called presapling and sapling more closely reflected the characteristics of mature forest. However, if the majority of my presapling and sapling study groups really belonged in mature forest, the n I would expect the selection ratios, medians, 95 percentiles, and preference zones to have resembled those of mature forest. Instead the selection ratios and distance parameters for the young seral groups were consistently higher than the mature groups indicating that 1) my method of calculating forest age is biologically appropriate for moose and 2) that the assumed distances moose are willing to travel through young seral forests may be too conservative. Perhaps at least in my study area, these young forests provide such a high forage value that moose are willing to travel further from cover to access them than assumed.

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Finally, although 95 percentile distances have the potential to determine thresholds and validate distance assumptions, I was unable to take full advantage of my third validation technique because proximity to cover was not selected at the population level, meaning that the 95 percentile values merely reflected what was available collectively within home ranges. However the se values do provide minimum distance thresholds indicating that animals are at least willing to travel those distances from cover. All of the 95 percentiles were below the model assumptions except for presaplings in the dormant season, where the distance to the sapling plus cover category extended approximately 100 meters beyond the assumed 200 meters. Accordingly, I recommend that parameter be extended to 300 meters. Although selection for proximity to cover was not established at the population level, it was present in 42% of growing season home ranges and 62% of dormant season home ranges. Of this subset, the observed distances were consistently lower than those of all home ranges combined and expected dista nces were relatively similar to those of all home ranges combined (Table 3.3). This indicates that the subset of home ranges where selection occurred was a result of differences in 3rd order level of selection behaviour and not merely differences in availability within home ranges. Because of this variability of selection behaviour, it may be appropriate to consider adding a second set of assumptions to the model that reflects the needs of animals requiring proximity to cover. More research is needed, but if I speculated correctly that much of the variation of selection behaviour is explained by calf presence/absence, then the model should be adapted to reflect the needs of cows with calves. Parts of the model do this already as the growing season browse component is based upon the needs of lactating cows (Allen et al. 1987).

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Because calf recruitment is an important component of population stability, habitat models and timber cut placements should explicitly address the special needs of encumbered cows. Models failing to do so will be ineffective at protecting and predicting population levels into the future. One approach for model adaptation is to add a third category to the range grids to designate the proportion of available habitat that is best suited for cows with calves. Preference zone would be the ideal parameter to use here. This category of the range grid could be given a slightly higher value than the broader category of available habitat needed for unencumbered moose. However, t his type of adaptation would be easiest to implement in a habitat suitability index that predicts habitat quality rather than the more specific prediction of carrying capacity as in the model I evaluated. Another option is to keep the range grid for encumbered cows separate from the original range grid that is used to predict density. The encumbered cow range grid could be used in the model in areas where moose numbers are below target or calf mortality rates are high in order to determine if habitat for encumbered cows is a limiting factor.

4.3 Model Recommendations In summary, I recommend testing an adapted habitat suitability model in which 1) stand age is determined according to timber harvest dates, 2) growing season thermal cover is expanded to include all conifer types, and 3) the dormant season distance threshold for travelling through areas of no cover is increased from 200 m to 300 m. I also emphasize the need for further research in two areas: 1) the role of wetlands as a source of thermal regulation and its role in determining available growing season range

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and 2) the cover requirements of cows with calves. If cows with calves are found to have a greater need for proximity to cover, I recommend a second set of distance parameters be added to the model to accommodate those needs. These model adaptations should be tested in other study areas before being implemented, as conditions in Algonquin Provincial Park are not representative of the entire Great Lakes Saint Lawrence forest region. I recommend testing the adapted model in an area with different anthropogenic and ecologic characteristics- specifically on a harvested population in an area with more fragmentation, human disturbance, private land, and different timber harvesting regimes. If possible the area should also have a more sparse spatial arrangement of cover that allows for the distance assumptions to be fully tested. The new study areas should have access to an FRI map based on recent photos as timber harvest information is not reported for most private land in Ontario, so the older the photo, the less-reliable the forest age. These adaptations and further research will improve our ability to manage healthy moose populations.

4.4 Implications of Variability in Habitat Studies My study revealed great variation in selection behaviour , both among and within individuals. Because individuals are unique in terms of their genetic make-up, physiology, and behaviour (Lomnicki 1988, DeAngelis and Gross 1992), inferences at the population level may be hard to ascertain, especially for complex behaviour such as habitat selection. Gillingham and Parker (2008) for instance, found that after completing a global resource selection model, there were no individuals in the study that fit the model. Further, the level of variation within individuals led to misleading conclusions

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about the general habitat selection portrayed in the global model High individual . variability affecting population inferences of habitat use has also been reported by others (Holmes et al. 1978, Ehlinger 1990, Tyler and Rose 1994, Volkl 1994, Compton et al. 2002, Marques 2004) but I believe the phenomenon is even more widespread, as some analyses commonly used in habitat studies (described by Thomas and Taylor 1990, and Manly et al. 1993) are not designed to detect individual variability. Thomas and Taylor (1990) and later Manly et al. (1993) described categories of study designs for habitat analyses that differ in terms of the level of which use and availability data are collected. My study used a type III design in which use and availability are determined separately for each individual. This design allows for the most thorough investigation of individual variability. A type II study design determines use at an individual level but availability at the population level. Individual variability can still be determined in this design but sometimes to a lesser extent. If the type II design is used in a study of 2nd order level of selection (selection of home range, Johnson 1980), then variation of selection at an individual level can be fully measured. However, when a type II study design is combined with a third order level of selection such as selection of feeding sites (Johnson 1980) , then individual variability can not be fully addressed. One can compare measures of use within individuals but because availability is not determined on an individual basis, comparison of selection may not be informative. A type I study design does not allow for any measure of individual variability since availability is defined at the population level and use data from all individuals are pooled. Type I study designs tend to be the most cost-effective because radio-marking

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individuals is replaced with cheaper methods such as track counts. However, when using this design, population level statements should be made with caution as they may be flawed in systems with high individual variability. My study also revealed great variation within individuals, which leads to a similar discussion as above but at a scale less commonly discussed. Aebischer et al. (1993) emphasized the importance of using individual as the sampling unit in habitat studies. This is an improvement over older techniques that use radio location as the sampling unit and pool all locations across animals (Smith et al. 1982, Byers et al. 1984). These older techniques are problematic because radio locations are likely serially correlated and individuals are liable to display different behaviour, both of which result in an inflated Type I error rate. Additionally, results will be biased towards the behaviour of animals represented by more locations. For these reasons, treating individual as the sampling unit is often superior to using animal locations, especially for habitat selection studies of species that dont move in tight social groups . However, depending upon the studys research goals, individual may not be the appropriate sampling unit. Using individual as the sampling unit assumes that an animals habitat needs are fixed, when in fact habitat requirements are dynamic and vary with abiotic conditions (such as weather and seasons) and biotic conditions (such as health, pregnancy, and provision of young). Measures important to habitat studies such as use, availability, and preference are likely to vary in these time periods . In situations such as this, it may be more appropriate to use animal-period as the sampling unit. This technique introduces some degree of Type I error because multiple animal-periods of the same animal are not entirely independent. However the inflated error rate can be

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mitigated using various techniques such as blocking for individual. Pooling all animalperiods of the same individual under conditions when animal-periods are different is inappropriate in the same manner as pooling all animals when individuals are different. More importantly, pooling animal-periods may mask critical habitat needs in the same way that my population level analyses masked the bimodal nature of selection behaviour. I therefore recommend that researchers give critical thought to the factors affecting habitat selection of their study species and the implications that might have on their research goals before defining a sampling unit.

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Appendix A Table 1. Forest Resource Inventory Landscape Guide Forest Units (LGFU) included in the three forest types used in this study.
Cover type Dominant species 1 Mh Mh - Be He - Mh - By Mh - Bw - Po By - Mh Or - Ms Po - Ms - Bw - Bf Bw - Ms - Mh - Po - Ce Po - Bw Po - Bw Po - Bw - Pj Ab - Ms - Ce Bf - Sw - Sb - Ce Sb Pw - Po - Bw - Pr Pw Sb - Pj - Bw Sb - La - Ce Ce Pw - Pr Pw - Po - Bw Pr % Landcover 75.3% 49.2% 9.7% 7.4% 4.9% 3.9% 0.1% 18.6% 7.1% 4.0% 3.7% 3.1% 0.2%

LGFU Total HDSL1 HDLS2

Description Hardwoods Tolerant hardwood selection Mid-tolerant hardwood Eastern hemlock Tolerant hardwood shelterwood Yellow birch Red oak Mixedwoods Rich upland mixedwood Rich upland mixedwood White birch Poplar Poor upland mixedwood Lowland mixedwood Conifers Spruce/Fir mix Pure black spruce White pine/Red pine White pine shelterwood Upland black spruce, mixed Lowland conifer, mixed Lowland conifer, mixed White pine/Red pine mix White pine/Red pine mix Red pine

Hardwood

HE HDUS BY OAK Total MWR MWUS

Mixed

BW PO MWD LMWM Total SF SB PWST PWUS4

0.5%
6.2% 2.2% 1.1% 0.6% 0.5% 0.5% 0.3% 0.3% 0.3% 0.2% 0.1%

Conifer

SP1 LC CE PWUSC PWUSH PR

1 Species Codes: Ab black ash; Be American beech; Bf balsam fir; Bw white birch; By yellow birch; Ce cedar; He eastern hemlock; La larch; Mh hard maple; Ms soft maple; Or red oak; Pj jack pine; Po poplar; Pr red pine; Pw white pine; Sb black spruce; Sw white spruce

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Table 2: Delineation of seral stages for my study groups in relation to the Forest Resource Inventory Landscape Guide Forest Units (LGFUs) that make up each study group. LGFU descriptions are located in Appendix A Table 1. Max age Presapling 14 14 14 14 14 14 14 9 9 14 9 9 9 14 14 14 9 14 14 9 14 14 14 14 14 Max age Sapling 39 39 34 44 34 39 34 29 29 34 29 24 29 34 29 29 24 34 34 24 29 34 34 34 39 Min age Mature 40 40 35 45 35 40 35 30 30 35 30 25 30 35 30 30 25 35 35 25 30 35 35 35 40

Cover type

Hardwood

Mixed

Conifer

LGFU Total HDSL1 HDLS2 HE HDUS BY OAK Total MWR MWUS BW PO MWD LMWM Total SF SB PWST PWUS4 SP1 LC CE PWUSC PWUSH PR

% Landcover 75.3% 49.2% 9.7% 7.4% 4.9% 3.9% 0.1% 18.6% 7.1% 4.0% 3.7% 3.1% 0.2% 0.5% 6.2% 2.2% 1.1% 0.6% 0.5% 0.5% 0.3% 0.3% 0.3% 0.2% 0.1%

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Table 3: Delineation of cover values for my mature study groups in relation to the Forest Resource Inventory Landscape Guide Forest Units (LGFUs) that make up each study group. LGFU descriptions are located in Appendix A. Cover values apply to forest stands classified in the FRI as immature, mature, or old. % Study Area Landcover 75.3% 49.2% 9.7% 7.4% 4.9% 3.9% 0.1% 18.6% 7.1% 4.0% 3.7% 3.1% 0.2% 0.5% 6.2% 2.2% 1.1% 0.6% 0.5% 0.5% 0.3% 0.3% 0.3% 0.2% 0.1% Dormant Cover Value 1 2 2 1 3 2 2 1 2 2 2 2 1 2 2 3 3 2 2 3 2 2 3 3 2 2 Growing Cover Value 2 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 0 0 0 1 1 0 0 0

Cover type

Mature Hardwood

Mature Mixed

Mature Conifer

LGFU Group Values HDSL1 HDLS2 HE HDUS BY OAK Group Values MWR MWUS BW PO MWD LMWM Group Values SF SB PWST PWUS4 SP1 LC CE PWUSC PWUSH PR

1. Dormant season cover value codes: 1 = lateral cover, 2 = early winter cover, and 3 = late winter cover. 2. Growing season cover value codes: 0 = no cover, 1 = thermal cover.

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Appendix B
Normal Probability Plot of SelectionRatio (Mid-winter Years)
2.0 Expected Normal Value 1.5 1.0 0.5 0.0 -0.5 -1.0 -1.5 -2.0 0.2 0.4 0.6 0.8 1.0 1.2 1.4 1.6 1.8 2.0 0.2 0.4 0.6 0.8 1.0 1.2 1.4 1.6 1.8 2.0

K-S p > 0.20

K-S p > 0.20

Year: MW06 2.0 Expected Normal Value 1.5 1.0 0.5 0.0 -0.5 -1.0 -1.5 -2.0 0.2 0.4 0.6 0.8 1.0 1.2 1.4 1.6 1.8 2.0

Year: MW07

K-S p > 0.20

Year: MW08

Categ. Normal P-Plot: Selection Ratio for Growing Season Years 2.0 K-S p > 0.20 1.5 1.0 Expected Normal Value 0.5 0.0 -0.5 -1.0 -1.5 -2.0 -0.4 -0.2 K-S p > 0.20

0.0

0.2

0.4

0.6

0.8

1.0 -0.4 -0.2

0.0

0.2

0.4

0.6

0.8

1.0

Gyear: 6

Gyear: 7

Figure 1. Distance to cover selection ratio normality plots for moose in Algonquin Park by year-season for year affect analysis. 72

Growing Season 2.5 K-S p > 0.20 2.0 1.5 1.0 0.5 0.0 -0.5 -1.0 -1.5 -2.0 -2.5 0.4

Expected Normal Value

0.6

0.8

1.0

1.2 Observed Value

1.4

1.6

1.8

2.0

Dormant Season 3 K-S p < 0.20 2

Expected Normal Value

-1

-2

-3 0.4

0.6

0.8

1.0 Observed Value

1.2

1.4

1.6

Figure 2. Distance to cover selection ratio normality plots for moose in Algonquin Park by season.

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Dormant Season Mature Hardwood 3 2 1 0 -1 -2 -3 0.2 3

Dormant Season Mature Mixed

K-S p < 0.20

Expected Normal Value

Expected Normal Value

2 1 0 -1 -2 -3 0.2

K-S p > 0.20

0.4

0.6

0.8

1.0

1.2

1.4

1.6

1.8

0.4

0.6

0.8

1.0

1.2

1.4

1.6

1.8

2.0

Observed Value Dormant Season Presapling 2.5 2.0 Expected Normal Value 1.5 1.0 0.5 0.0 -0.5 -1.0 -1.5 -2.0 -2.5 0.2 0.4 0.6 0.8 1.0 1.2 1.4 1.6 1.8 2.0 2.2 2.4 2.6 Observed Value
Growing Season Mature Hardwood 2.5 2.0 Expected Normal Value 1.5 1.0 0.5 0.0 -0.5 -1.0 -1.5 -2.0 -2.5 0.2 0.4 0.6 0.8 1.0 1.2 1.4 1.6 1.8 2.0 2.5

Observed Value Dormant Season Sapling 2.5

K-S p > 0.20

Expected Normal Value

2.0 1.5 1.0 0.5 0.0 -0.5 -1.0 -1.5 -2.0

K-S p > 0.20

-2.5 0.0 0.2 0.4 0.6 0.8 1.0 1.2 1.4 1.6 1.8 2.0 2.2 2.4 Observed Value
Growing Season Mature Mixed

K-S p > 0.20

Expected Normal Value

2.0 1.5 1.0 0.5 0.0 -0.5 -1.0 -1.5 -2.0 -2.5 0.4

K-S p > 0.20

0.6

0.8

1.0

1.2

1.4

1.6

1.8

2.0

Observed Value Growing Season Presapling 2.0 1.5 Expected Normal Value 1.0 0.5 0.0 -0.5 -1.0 -1.5 -2.0 0.4 0.6 0.8 1.0 1.2 1.4 1.6 1.8 2.5

Observed Value Growing Season Sapling

K-S p > 0.20

Expected Normal Value

2.0 1.5 1.0 0.5 0.0 -0.5 -1.0 -1.5 -2.0 -2.5 0.0

K-S p < 0.10

0.5

1.0

1.5

2.0

2.5 3.0 3.5 4.0 4.5

5.0

5.5

Observed Value

Observed Value

Figure 3. Distance to cover selection ratio normality plots for moose in Algonquin Park by habitat group-season. 74

Presapling Distance 2 2.5 2.0 Expected Normal Value 1.5 1.0 0.5 0.0 -0.5 -1.0 -1.5 -2.0 -2.5 0.6 0.8 1.0 1.2 Observed Value Sapling Distance 2 2.5 2.0 Expected Normal Value 1.5 1.0 0.5 0.0 -0.5 -1.0 -1.5 -2.0 -2.5 0.2 0.4 0.6 0.8 1.0 1.2 1.4 1.6 1.8 2.0 1.4 1.6 1.8 2.5

Presapling Distance 3

K-S p > 0.20

Expected Normal Value

2.0 1.5 1.0 0.5 0.0 -0.5 -1.0 -1.5 -2.0 -2.5 0.4

K-S p > 0.20

0.6

0.8

1.0

1.2

1.4

1.6

1.8

2.0

Observed Value

K-S p > 0.20

Observed Value

Figure 4. Distance to cover selection ratio normality plots for moose in Algonquin Park for lesser cover categories.

75